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SMITHSONIAN
MISCELLANEOUS COLLECTIONS

ARE: INCRE 2
“oe OWE VIO AES
Gres CM nck Alo) 0,

““ EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES,
AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN ””__sMITHSON

(PUBLICATION 3529)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1939

The Lord Gaftimore (Press

BALTIMORE, MD., U. 8 As

Me, oY poet ys

ADVERTISEMENT

The Smithsonian Miscellaneous Collections series contains, since the
suspension in 1916 of the Smithsonian Contributions to Knowledge,
all the publications of the Institution except the Annual Report, the
annual pamphlet describing the Institution’s field-work, and occasional .
publications of a special nature. As the name of the series implies, its
scope is not limited, and the volumes thus far issued relate to nearly
every branch of science. Papers in the fields of biology, geology,
anthropology, and astrophysics have predominated.

C. G. ABBor,
Secretary of the Smithsonian Institution.

(iii)

to

6.

NY

Io.

Teles

CONTENTS

STRONG, WILLIAM DUNCAN; Kipper, ALFRED, II; and PAaAuvt,
A. J. DREXEL, Jr. Preliminary report on the Smithsonian
Institution-Harvard University archeological expedition to
northwestern Honduras, 1936. 129 pp., 16 pls., 32 figs.,
Jan. 17, 1938. (Publ. 3445.)

Jounston, Eart S. Plant growth in relation to wave-length
balance. 18 pp., 4 pls., Jan. 12, 1938. (Publ. 3446.)

RESSER, CHARLES ELMER. Middle Cambrian fossils from Pend
@redleseake, Idaho» 12 pp. 1 pli, Jan. 3; 19038. (Publ:
3447.)

ScumiTt, JoHN B. The feeding mechanism of adult Lepidoptera.
28 pp., 12 figs., Jan. 10, 1938. (Publ. 3448.)

StTrrtinc, M. W. Three pictographic autobiographies of Sitting
Bull. 57 pp., 46 pls., July 22, 1938. (Publ. 3482.)

Snoperass, R. E. Evolution of the Annelida, Onychophora, and
Arthropoda. 159 pp., 54 figs., Aug. 23, 1938. (Publ. 3483.)

WebEL, Watpo R. The direct-historical approach in Pawnee
archeology. 21 pp., 6 pls., Oct. 19, 1938. (Publ. 3484.)

BusHNELL, Davin I., Jr. Drawings by George Gibbs in the Far
Northwest, 1849-1851. 28 pp., 18 pls., 4 figs., Dec. 30, 1938. -
(Publ. 3485.)

Deicnan, H. G. A new nuthatch from Yunnan. 2 pp., Oct. 10,
1938. (Publ. 3486.)

RESSER, CHARLES ELMER. Fourth contribution to nomenclature
of Cambrian fossils. 43 pp., Dec. 17, 1938. (Publ. 3487.)

WEINTRAUB, Rosert L. An assay method for growth-promoting
substances utilizing straight growth of the Avena coleoptile.
10) pps tpl, Wee, 31, 1938. (Publ. 3486.)

Resser, CHARLES’ Ermer. The Spence shale and its fauna.
29 pp., 6 pls., Jan. 20, 1939. (Publ. 3490.)

(v)

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"SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER 1.

_ PRELIMINARY REPORT ON THE
_ SMITHSONIAN INSTITUTION-HARVARD UNIVERSITY
ARCHEOLOGICAL EXPEDITION TO NORTHWESTERN
HONDURAS, 1936

(WiTH 16 PLATEs)

BY

WILLIAM DUNCAN STRONG
Anthropologist, Bureau of American Ethnology

ALFRED KIDDER II
Peabody Museum, Harvard University
AND

A. J. DREXEL PAUL, JR.
Peabody Museum, Harvard University

(PUBLICATION 3445)

Ae CITY OF WASHINGTON
PueESHeD BY THE SMITHSONIAN INSTITUTION
JANUARY 17, 1938

(‘Apasse) “5 “y Aq sunuied e wos)

VaEIa0 V1 ‘2 ALIS ‘(AdAL GIOAVW
‘ASVA ANOYHDAIOd VOFOA V NO SAYNDIA IWNOISSSD0O¥d

L “Id ‘bk “ON ‘26 “10A SNOILO31100 SNOANVTISZOSIN NVINOSHLINS

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 97, NUMBER 1

PRELIMINARY REPORT ON THE
SMITHSONIAN INSTITUTION-HARVARD UNIVERSITY
ARCHEOLOGICAL EXPEDITION TO NORTHWESTERN

HONDURAS, 1936

(WiTH 16 PLATEs)

BY
WILLIAM DUNCAN STRONG
Anthropologist, Bureau of American Ethnology
ALFRED KIDDER II

Peabody Museum, Harvard University
AND

ARE DINE MEIS PAU ike
Peabody Museum, Harvard University

ZU
4s THSO
Nig Lungity

(PUBLICATION 3445)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 17, 1938

The Lord Baltimore Press

BALTIMORE, MD., U. & A.

CONTENTS

PAGE
iifirrayAIBVS AVOUT Ps, drags cies eed LAOS RIE OIE MGU Gd Oo Mad ab on oucs onsen I
EMaeieo COP Tap CUSEELUM Ors a tuenehet e oai oie airs el eeiaocrerensie aieneioheron eters eeiece creel 2
Bahnnicrandm lines cibackorounds... cs «= cece ane se eee cree cece e era 8
Banivehistomc contacts, im northwestern Honduras); qqs-sse seco seee ae 19
NGchem O2tGalmexp OnatONSe cera acre se vic sisi he ett aie Soe ee ic Tce ore e oS 27
hianiaiel EconmelLeiviete erase srotes.cke aides SE Rhee cl BRUT aa eae ee RRS 27
HNECTC Oe RT ease Bos coe Ne cai ols Sear aT Seon TOS Tare pear eek aie Rewehitee omeestrcs 27

NE ASMAVIC DAG ME rE T ATC reer cco ia e asd Sco tecetspere o OmHeeOe naire ot arcte eae 34

ite IPAS GRIS. 6 A's oes o 5 Ai eR GeO On RO Gs SOO ra Geos oma nea 35

(Me TEMSILC SME IRR ee cre Mr ec Gat tera Rcc stent eee Deanne 37

Wiliam Gomeya ctiam RIVES oasis clare + varie eeiea cic. e cece treieeescrereets 39
[brow OF CSU Ol Sam MRR tees serait als Oe La ed oe oe rae re 39
SaTltAMNitcie Cail aera Aceh nis ees ee rene scram ee eee 45
Binyarde losmiierios (akin IT ya. of ont ews soc senses Seine eee 62
@Cheresitesue an ween Aen renee arc ta ceeisele oes ave asia eRe eee ee ee 76
INonthmendmotelalcemVioy Gamera css eke colar Oe cere sta Stein eee 76
INSHACATC ATIC PA Ottaitalleanams caine wiles s/eur eieis sis owveisele else miclentetevemsiars ts 80

Een Bei lye eee APE ie Sebcici ane tsb chs aoe sua eters Eien Saarare exe imianocala Reererey go

SC CMR Pe TEP eee or acai oy so RET Cs obi Ba acl cone sslehchea rer ncbiate QI

Stee 2m ee mete eR ES heey aR emir lar a Sina pe es ce ae em 94

Site este ae area aster acetic on etches ancl chenahs Stee (ake Wager ete ofS eee ree 99

Causeway, didencanaly near Jatal: ins. o.'sh2'% neve s <jate stole Maver 100
Pyramids and stone statues near Los Naranjos.................-. 102
Excavations on the northern border of Los Naranjos.............. 105
M@hevolderhionzonvatweoseNatanyjossaeacsn sees ce cease ee ee III

@fhenmesitesme cert Re AE ETA Ce ain ieee cS Brae aris SiN aCe waate cee ea 115
Sunumany and tentative: COnClISIONSs 6 l.).:c sie sss ce ves nse oialetslejnre wialele dic e's 118
NPitera hah cmc ite cummeen reer strate al sie euetchs ioe ielenstio Sista cve dais er closely suse genoa te tetous ones cisind.« 125
ares AeA ICAL CO MERU LEVEE SI Hye avast yeh cL 3) i sou faints! 2) re) eats sia eidhorrepeie a urerorevay mate wines S wens 127

ili

WAKER Eb

10.
ie
12.
13.
IA.
TS
16.

Cee taco ated

10.

Hie
12.
12:
14.

15.
16.

ILLUSTRATIONS

PLATES
PAGE

(Frontispiece) Processional figures on a Yojoa Polychrome vase of
Mayoid' type: site. a Geibas i etn Seles eerie eee eee title
Narious) Ghameleconr and: Ulan Rivetrcttesmsycrcrtieie tertile tien ttre 130
INIACOMSHERAS: . fos cities oc 2 sicuste lhe lotta inlets ence oto ielsureterocsars terion
INacomsherdsmandaattitacts sieerreeriee ect icr tie ieiericte italian eet
Upper Ulua Polychrome pottery, types, Maas) Plores = .1.- ie vis ee a aire
Upper Ulua Polychrome pottery types, Las Flores..................--
Ulua Polychrome, Bold Geometric pottery types, Santa Rita and Naranjo
hain ee oe NG och lee soi oo EPO eae Te dine lonbare eines Cee teeta
Ulua Polychrome, Mayoid pottery types, Santa Rita..................
Ulua Bichrome sherds, from deepest level at Santa Rita..............
Playa de los Muertos Bichrome sherds....... sr sharltats Banal catecoharerele ap aeaeets
Playa de los Muertos Bichrome sherds and figurines..................
Wojoa: Polychrome’ vessels, Mavyordity pes «ic aj). ccm eieteei sine ete
Yojoa Polychrome vessels, Bold Animalistic and other types...........
Yojoa Polychrome vessels, Bold Animalistic and other types...........
Rarlyceramic types: at eaake YO @disels ale os a's cters rs ercherora(->yayeraeyoe eres
Mos Naranjos,: lealke "YO Gains. 2 clemisscicracn ehalecetricieie olepe foletalct ye ersictoroiamy sitet

TEXT FIGURES

PAGE
Map of northwestern Hlondttras’. 5 ..ict) sis om 3s 5 oa elas cloeinnn seals 4
Map of the-region aroutid: NAC. amass aneeiosorass Sorte e ere ee ee 7
Sketchimap.of the suins of Waco. accra © oe oes poe eee 30
North wall of cross-section trench through mound 3, Naco............ 31
Map of the lower Ulua’ and ChamelecomRiverss 22.2001. sss 42 eee 40
West wall of excavation 1, Santa Rita (farm 17)............ Pee 47
Hollow figurines, whistles, and “candelario”, from the Ulua Poly-
Chinemerperiod, Satitay Ritan (farina 17) ie cictetrsts cr ete peter nee eee 52
Ulua Polychrome, Bold Geometric tripod dish, excavation 2, Santa Rita
(eaten 7a ta oy sevarn sree ts ceemne a pataite lc} sha crneere) aleve mans kegenstniee sNeneia gates tee 54
Unusual Ulua Polychrome, Bold Geometric dish, excavation 2, Santa
Riba MCHA OL7, tert yams: ce or eee ers + ues eisisia te se eine eee et lero 55
Ulua Polychrome, Bold Geometric bowl, excavation 2, Santa Rita
(Getame I) ee eens see titer tonnes Sse aintelomie ae +e Oeste aint ae errr rae 56
Ulua Polychrome bowl, excavation 2, Santa Rita (farm 17)........... 57
Ulua Polychrome bowl, excavation 2, Santa Rita (farm 17)........... 58
Ulua Polychrome, Mayoid vase, excavation 2, Santa Rita (farm 17)... 59
Inside design from Ulua Polychrome, Lower Mayoid vases (pl. 8, a, b),
excavation x, cantar ieita. (farm 17) a... trices ec = oe atelier 60
Ulua Polychrome Bowl, excavation 2, Santa Rita (farm 17).......... 61

North wall of excavation I, showing stratification, Playa de los Muertos
(Eaten eUT ha dechas Pe iexe sks p orsiate are puniohers taba Gch yates ete oro ter eee ee AEN eee

1V

10.
20.
21
22.
2a
24.
25.
20.
27.
28.
20.
30.
Bie

32.

Led ILLUSTRATIONS Vv
PAGE
Outlines of vessels of the Playa de los Muertos culture obtained by
Weartathty ee EGPEHOE. ...2.chowieatas Aiclsnrsin te eis cine © emitter oe ie taliers «i abalmnata'e 70
Outlines of vessels of the Playa de los Muertos culture obtained by
Woascothyall a OpenOeracysctitrelenrra ets a trerrnere eicieysieetelet iat teteelralelered 71
Polychrome vase of Ulua Mower Mayoid types s--.--.6+2ee ces oss 6 77
Sketch map of archeological sites around the north end of Lake Yojoa.. 78
Yojoa Polychrome bowl, Bold Animalistic type, Aguacate............ 82
Yojoa Polychrome pot, Bold Animalistic type, Aguatal............... 84
Yojoa Polychrome bowl, Bold Animalistic type, Aguacate............ 85
Yojoa Polychrome tripod dish, Bold Animalistic type, Aguacate...... 86
Yojoa Polychrome bowl, Bold Animalistic type, Aguacate............ 86
Outline of Yojoa Polychrome pot showing “ vestigial” spout, Aguacate. 87
NWojoam oly chrome: cooking spot, Asuatalsssasei cee cee citeieiercll te 88
Yojoa Polychrome bowl, Mayoid type, Aguacate...................-- &9
Yojoa Polychrome bowl, Mayoid (?) type, Aguacate................. go
Yojoa Polychrome vase, Mayoid type, Aguacate......... ............ QI
Cross-section of west wall, site 1, Los Naranjos, showing house floor,
burial, and superimposed cultural horizons.....................-.-- 107
Cross-section of excavation A, near site 1, Los Naranjos, showing

ShraniedMcuitidicl ahOrtZOlsemcmren acer i atreee oer cs. Sree coterie eee 113

s ; ws M " nit ohm ¥)

Vek Wik ,

( ‘ge roa h as ae
"aly eae 3

PREEIMINARY “REPORT ON THE
SMITHSONIAN INSTITUTION-HARVARD UNIVERSITY
ARCHEOLOGICAL EXPEDITION TO NORTHWESTERN

HONDURAS, 1936

By WILLIAM DUNCAN STRONG
Anthropologist, Bureau of American Ethnology;

ALFRED KIDDER, II
Peabody Museum, Harvard University ;
AND

Ae). DREXEL “PAUL. JR:
Peabody Museum, Harvard University

INTRODUCTION

The present paper presents in tentative and outline form certain
major results of the Smithsonian Institution-Harvard University
Archeological Expedition to northwestern Honduras in 1936. The ex-
pedition personnel included the senior author as leader and represen-
tative of the Bureau of American Ethnology, Smithsonian Institution,
and the two other authors as representatives of the Peabody Museum
of Harvard University. Mrs. Strong and Mrs. Kidder accompanied
the expedition in the field and performed invaluable services in cata-
loging and caring for the collections. The purpose of the expedition
was twofold: to extend the work carried on by the Smithsonian Insti-
tution in northeastern Honduras in 1933, and to follow up earlier work
on the Ulua River so successfully inaugurated, under the auspices of
the Peabody Museum, by George Byron Gordon and by Mrs. Dorothy
Hughes Popenoe. The discoveries of Mrs. Popenoe at Playa de los
Muertos in 1928 and 1929 opened new vistas in Honduras archeology,
and her untimely death was a sad blow to science and to all who were
fortunate enough to know her. In a sense our work was merely a con-
tinuation of that which she had so ably begun. The original suggestion
for the present expedition came from Dr. Wilson Popenoe, and it was
due to him that the successful cooperative effort was launched and
completed.

The expedition received cordial support from the government of the
Republic of Honduras, and our warmest thanks are extended to the

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97, No. 1

bo

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

officials in Tegucigalpa and elsewhere who not only furthered the
cause of science but put us deeply in their debt for many personal
courtesies. Similarly, the officials of the United Fruit Company, both
in the United States and in Honduras, furnished very material aid
in ways too numerous to mention. Without this generous assistance
our results would have been tremendously curtailed. Considerations
of space prevent listing the many people who aided us in our work, but
we cannot forbear mentioning Mr. Walter S. Turnbull, and Mr. Regi-
nald Hamer. To them and to many other persons in Honduras we are
deeply grateful. Later, in the final report, it will be possible to
acknowledge more adequately our appreciation of the many cour-
tesies, both official and personal, which we received on every hand.

Our choice of this particular area for excavation was based on
numerous geographic, historic, ethnographic, and archeological con-
siderations. For this reason we have devoted considerable space to
these important factors. Viewed against this background, it is hoped
that a condensed account of our excavations may have value. In due
course a final report will be prepared by the senior author for publi-
cation by the Peabody Museum. As this may not appear for some
time, it seems advisable to make our major results available without
undue delay. Most of this important area still awaits adequate exca-
vation, and it is our hope that these notes and sketch maps may be of
value to future scientific explorers and excavators.

BRIEF GEOGRAPHIC SETTING

The general area covered by this report includes the eastern por-
tion of the Department of Cortes, the western border of the Depart-
ment of Yoro, and certain places on the eastern border of the De-
partment of Santa Barbara, all within the Republic of Honduras.’

From the archeological standpoint, however, modern political
boundaries are of minor importance compared to factors of terrain
and drainage, which conditioned aboriginal human occupation no less
than they do that of the present inhabitants. Of primary importance

*See map, fig. 1. The best general maps of Honduras at present are the
“Carta General de la Republica de Honduras, America Central, of the Pan-
American Institute of Geography and History, 1933”, and the “ Mapa General
de la Republica de Honduras, Levantado por el Prof. Jesus Aguilar Paz, 1933.”
These maps are far superior to any of their predecessors. Many contradictions
still exist, however, owing to the present inadequacy of cartographic exploration
in much of Honduras. The present map (fig. 1) is primarily based on that of
Direaz,

NOs HONDURAS—STRONG, KIDDER, AND PAUL 3

is the fact that at this point occurs one of the easiest passages across
the Central American Isthmus from Tehauntepec to Panama.”

From the mouth of the Ulua River, where it enters the Gulf of
Honduras, a series of elevated valleys extend up the Rio Blanco to
Lake Yojoa, over the plateau of Siguatepeque, across the Plains of
Comayagua, and down the valley of the Goascaran River into the
Gulf of Fonseca and the Pacific. It can hardly be coincidental that it 1s
at this point that the higher aboriginal cultures of the Pacific High-
land extend north to the Caribbean Sea, in marked contrast to the lower
cultures of the remainder of the Atlantic Lowland region in Honduras.
The present archeological reconnaissance covers the northern half of
this natural transition area between the Pacific Highland and the
Atlantic Lowland regions.

If we include the valley of the Chamelecon River, which at no very
distant time emptied into the Gulf of Honduras through the Ulua
River, this entire area from Lake Yojoa north may be termed the
Ulua drainage. The lower portions of the Ulua and Chamelecon Riv-
ers flow through the Plain of Sula, a rich alluvial valley, down to their
respective mouths in the great mangrove swamps extending along the
Gulf of Honduras from Puerto Cortez almost to Tela. Because of
these swamps and their shallow, silted-up channels, neither river
offers much inducement to modern navigation, whereas such impedi-
ments were probably of small import to the numerous trading canoes
of pre-Conquest times. Above the mangrove swamps, which extend
some 20 kilometers upstream, is the rich valley floor that today is cov-
ered with banana plantations. Formerly the valley supported a rich
tropical flora, described by Gordon (1898) and others, but at present,
except for isolated remnants in swamps and low areas, the great
mahogany, ceiba, and other trees, have been replaced by the ubiqui-
tous banana. To the northwest the Ulua valley is hemmed in by the
great Mountains of Omoa, which reach a height of 7,000 to 8,000 feet.
To the east occur the Mountains of Mico Quemado and Tiburon. Be-
tween these two ranges the Ulua-Chamelecon valley reaches a breadth
of some 45 kilometers, terminating about 75 kilometers in a direct
southwest line from the mouth of the Ulua at Potrerillos, where the

? This has been pointed out time and again in the voluminous literature referring
to the much-talked-of but never completed transoceanic railroad across Honduras.

See Squier, 1858 and 1870, and Wells, 1857. Although perhaps unduly opti-
mistic on some points, Squier’s various reports remain the best general geographic
descriptions of Honduras.

Wells gives a detailed and delightful picture of Honduran life in the middle
of the last century. So far as the remote interior is concerned, much of his
description holds good today.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

CARIBBEAN SEA

KILOMETERS
ae Sianes

-

1
NICARAGUA

AA
- =
°

Lc, SIGUATEPEQUE|

Q
3
x Sy
. \
4 \

‘Sie § ZT ie /

Soro a / TEGUCIGALPA
aN MO
LNTHB U CA \ Dee

a
LA ESPERANZA 2
Vl PAA &

& MARCALA \

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GUAJIQUIRO

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EGS) i

Map of northwestern Honduras (based on the General Map of the
Republic of Honduras, by Prof. Jesus Aguilar Paz, 1933).

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 5

bordering mountains converge and the Ulua splits up into its three
main branches (maps, figs. 1, 5). These are, from north to south, the
Comayagua, the Rio Blanco, and the Ulua proper. The Chamelecon,
after running parallel to the Ulua for some 50 kilometers above its
mouth, turns north into mountains, where, as a rapid mountain stream,
it extends almost as far southwest as Copan. Similarly, the three
branches which form the Ulua are rapid, clear streams, in marked con-
trast to the slow moving, muddy lower Ulua and Chamelecon.

Owing to its configuration and the mountainous character of its
terrain, Honduras has a wide variety of climates and seasons. In
general, however, the dry season in the region we are considering be-
gins in December or January and extends until June or early July.
The temperature, which is pleasantly low in the early part of the dry
season, increases as the wet season approaches. The rainy season is
cooler, but more unpleasant, owing to rain, wind, and great humidity.
Moisture brought by the northeast trade winds is deposited when they
hit the high mountains bordering the Ulua valley. Thus, despite their
relatively short courses, the Ulua-Chamelecon tributaries at times
carry a tremendous volume of water. These rivers rise to their great-
est heights about October and flood the valleys. A smaller rise occurs
in the late Spring. As the rivers spread over the lower valleys, they
deposit the sediment brought down from the mountains, and in this
way the valley has been aggraded or built up. As proved by human
occupation levels buried in situ 6 meters or more deep along the pres-
ent channels, this building-up process has been relatively rapid and
continuous.” .

As the present rivers shift their channels across the valley floor, they
thus expose in their steep banks the various human occupation strata of
past centuries, which elsewhere in the valley are inaccessible, because
of depth and lack of surface indication. The majority of the sites
investigated along the Ulua by the present expedition were of this
type.

Whereas the lower Ulua valley was formerly covered with a luxuri-
ant rain forest, the sites which we worked on the upper Chamelecon

*Yde, 1936, p. 39, in our opinion, exaggerates the difficulties due to depth of
deposit facing the archeologist in this region. Nevertheless, there is no doubt
that the earliest human remains in the lower Ulua valley may be buried at
inaccessible depths. However, as the present report demonstrates, it is possible
to obtain stratification. Gordon, 1898, shows the manner in which the rivers
cut and shift their channels as well as the dangers of re-deposition which must
- be borne in mind by the archeologist. Neither Mrs. Popenoe nor the present
writers encountered cultural remains at the extreme depths mentioned by Gordon
and Yde.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

in the vicinity of Naco (map, fig. 2) are located on clear, rapid streams.
These streams are bordered by narrow strips of tropical forest, but
back from these are steep hills or elevated rocky plains covered with
pines and oaks. There is no reason to believe that the environment here
was different in aboriginal times. It is a region admirably adapted to
maize. Abundant food supplies possessed by the numerous Indian
pueblos, as well as the occurrence of gold in the surrounding moun-
tains, early attracted the Spaniards to these mountain valleys. The
climate appears to be much more healthful than that of the lower
river valleys.

The northern end of Lake Yojoa, where other excavations were
carried on, offers a similar environment. This marks our most south-
erly working point as well as the limit of the Ulua drainage in this
direction, since Lake Yojoa in part drains through an underground
channel into the Rio Blanco. To the south, it is said to drain into the
Santa Barbara River by means of the Jaitique River and by sub-
terranean channels, principally the Rio Sacapa and the Rio Salala.’
We did not investigate the southern end of the lake.

Lake Yojoa is located in a small mountain valley or bolson at an
altitude of some 2,050 feet. The auto road from the north coast to
Tegucigalpa utilizes the lake as a water connection by means of auto-
mobile ferries. To reach Jaral, the little town on the north shore, one
leaves the low banana country at Potrerillos and climbs through rocky
hills covered with oak, pine, and scrub, following the Rio Blanco River,
which is crossed only once at the little town of that name. Just before
the road reaches the lake, the grade increases sharply and then drops
down onto the small triangular plain bordering the lake (map, fig. 20).
This bush-covered plain is bounded on the east by low pine-covered
hills, and on the north by high (5,000-6,000 feet) volcanic mountains.
We suspect that this plain originally supported a heavy rain forest,
but both the aboriginal and modern inhabitants have long practised
milpa-type farming here, and today there exist few remnants of the
original forest. With the exception of open, rolling, pine-covered hills
on the northeast shore near Agua Azul, the remainder of the lake
is bordered by steep slopes covered with rain forest. At the southern
extremity of the lake, there is a considerable belt of low, swampy land,
most of which is overflowed when the lake is full. Beyond the water-

* Published reports and maps of Lake Yojoa are utterly inadequate. Squier,
1858, pp. 96-104, and 1860, pp. 58-63, is still the authority. The lake has been
studied from time to time by Honduran and American engineers, but we know
of no up-to-date maps or reports. Amory Edwards, and Squier, 1860, describe
io outlets.

STRONG, KIDDER, AND PAUL

HONDURAS

I

NO.

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8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97
shed to the south in the vicinity of Taulebé and San José, there are
fertile plains and valleys. These are separated from the Comayagua
valley by the high plateau of Siguatepeque.

In general, therefore, it can be said that the sites investigated by
us in the Ulua drainage occupy two main environmental regions: First,
those on the lower Comayagua and Ulua Rivers, located in the rain
forests of the broad, alluvial river valleys, and second, mountain val-
leys, as at Naco and at Lake Yojoa, where the elevation was consider-
ably greater, the rain forest limited to the borders of stream or lake,
and the more open pine and oak association close at hand.

ETHNIC AND LINGUISTIC BACKGROUND

At the present time there are no obvious, aboriginal remnants of
population in the part of Honduras considered in this report. It is
true that the present population of the region is in considerable part
made up of assimilated Indian groups, but the language is Spanish
and the culture Latin American. Isolated groups of Jicaque Indians
are reported as still living in the more remote parts of the Depart-
ment of Yoro.’

It is possible that Lenca-speaking Indians may still be found in
our region, and groups of Chorti Maya occur in the departments to
the south and west, but, as yet, ethnographic and linguistic research
in Honduras has received little attention. If we desire to connect the
archeological remains with historic Indian groups, it is therefore neces-
sary to turn back the pages of history and consider the region at the
time of the Spanish Conquest.

Early sources on northwestern Honduras are fairly numerous, in-
cluding Cortez, Bernal Diaz, Palacio, Las Casas, Torquemada, Mar-
roquin, Montejo, Palaez, Pedraza, Espino, and the historians Her-
rera, Oviedo, and Gomara, but, with the exception of the first three, the
grains of ethnography to be gleaned from the works of these writers
seem rather scant. In a later paper other sources will be considered,
but for the present we shall confine ourselves to the most important
primary sources and more recent general studies.

As was the case in regard to the geography of Honduras, one must
still consult E. G. Squier’s “ States of Central America”, 1858, re-
garding the ethnography of Honduras. Similarly, H. H. Bancroft’s

* Described by Habel, 1880, p. 17. In June 1933 the senior author was told by
Mr. Acley, then American Consul at Tegucigalpa, of a very primitive group of
Jicaque Indians he had visited that year near the town of Morale, in the Sierra
de la Flor of the Department of Yoro, near the Olancho line. From photographs,
they appeared very similar to those described by Habel.

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 9

“Native Races’ (1882) is a treasure house of ethnographic source
material, and the same author’s “‘ History of Central America” (1883)
not only indicates the sources but also the major trends of native and
European contact in the period of conquest. More recently Cyrus
Thomas and John R. Swanton (1911) have presented in brief form
the salient facts regarding the distribution of Indian languages in
Honduras as part of their study of the languages of Mexico and Cen-
tral America. Preeminent in this field, however, is the voluminous
and detailed work of Walter Lehmann, who in a preliminary report
in I910, and in “ Zentral Amerika”’ (1920), his given us a wealth
of data based on close examination of the sources as well as personal
linguistic work in the field. In addition to an intensive study of origi-
nal sources and present-day Indian dialects, Lehmann has also in-
cluded many sweeping theoretical generalizations. The latter, however,
concern us less in the present study than do the specific references to
linguistic and cultural distributions in Honduras at the time of the
Conquest.

In general, the linguistic maps of Thomas and Swanton (1910),
and Lehmann (1920) agree as regards the distribution of native
languages in the Ulua drainage. Both place Chol, Chorti, and other
Maya-speaking groups to the west of the Ulua River proper. Follow-
ing Lehmann, we find that the Lenca occupied a large area around
Lake Yojoa, extending north almost to the junction of the Comayagua
and the Ulua. From here to the coast the valley of the Ulua and
Chamelecon Rivers was Jicaque territory. To the west, Lenca and
Jicaque territory bordered on that of the Chorti and Chol Maya,
Copan being one center of Chorti occupation. The Lenca and Jicaque
demesnes extended east to that of the Paya who with the Sumu, oc-
cupied the northeastern corner of Honduras.

To the south, peoples of Lenca speech extended to the Pacific coast.
To the west they were bordered by the Pipil of Salvador along the
Lempa River, and to the east by various Chiapanecan and Matagalpan
groups (see linguistic maps, Thomas and Swanton, 1911, and Leh-
mann, 1920).

Thus it appears that all of the territory investigated by the present
expedition was occupied by Jicaque and Lenca-speaking peoples at the
time of the Conquest—with one very important exception. This was
the occurrence in the same region of various Nahuatl-speaking pueblos
along what appear to have been trade routes extending from south-
ern Mexico and from the Pipil (Nahuatl) territory in Salvador. Leh-

* Linguistic and tribal distributions in Honduras have already been discussed
in some detail; see Strong, 1935, pp. 7-19 and 140-172.

Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

mann indicates one such line of late Nahuatl influence and settlement
which crossed the Chamelecon River in the vicinity of Naco and ex-
tended east to the Nahuatl pueblos mentioned by Cortez, located a
short distance south of Trujillo (Lehmann, 1920, vol. 2, p. 629 and
map). Both Cortez and Bernal Diaz in their accounts of the traverse
from Mexico to Honduras indicate the importance and vogue of
these trade routes and mention the many pueblos engaged in trade
which they visited (Bernal Diaz, 1916). Recent Nahuatl settlements
would be thus expectable in the upper Chamelecon valley near Naco
and probably elsewhere in the Ulua valley proper. These Nahuatl
influences from southern Mexico were apparently quite recent, but
the Pipil occupation of Salvador was much older. This is clearly
indicated by Palacio (1860, pp. 21, 31, and 65), who points out the
acquisition of the Pipil tongue by many neighboring groups originally
of different linguistic affiliation. Moreover, Pipil cultural influences
were obviously very active in southwestern Honduras at the time of
which Palacio writes (1. e., 1576).

The name of the Ulua River was apparently derived from the Ulba
language, which Palacio ascribes to Honduras (1860, p. 21). Both
Squier (Palacio, 1860, p. 114) and Lehmann (1910, p. 747; 1920,
vol. 2, p. 624) concur in this identification. The extension of the term
Ulba, Ulua, or Ulvan to the Sumu is explained by Lehmann on the
grounds that the Sumu, Jicaque and Matagalpan languages (includ-
ing the Cacaopera and Lislique) are all basically related. This seems
quite probable but has not yet been satisfactorily demonstrated. Since
the Jicaque lived along the Ulua river, it is most probable that Palacio
referred to them as the Ulba. Both Squier (Palacio, 1860, p. 114)
and Lehmann (1920, p. 624) regard Palacio’s designation “‘ Chontal ”
as a general term for non-Pipil-speaking peoples. According to Leh-
mann, the term would include the Lenca with the Jicaque. Specifically,
Palacio seems to refer to the Lenca when he speaks of the Taulepa.
This is the old name for Lake Yojoa (Taulebé, according to Squier,
1860). Inthe Lenca language, Taulepa means “‘ House of the Jaguars ”
(Lehmann, 1910, p. 747; 1920, vol. 2, p. 624). The jaguar was of
special importance in Lenca mythology. Lehmann is convinced that
the region around Lake Yojoa and the entire central portion of
Honduras was occupied by the Lenca (the Taulepa of Palacio), and
that the valley of the lower Ulua and the adjacent Department of
Yoro was primarily occupied by Jicaque groups (the Ulba of Palacio).
During the seventeenth century, the names Lenca and Jicaque were
often confused, but, as indicated by the work of Thomas and Swanton,
the general regions assigned to these groups by Lehmann seem
accurate.

NO} = HONDURAS—STRONG, KIDDER, AND PAUL ret

It is therefore apparent that our archeological investigations were
made in a contact area between advanced Mayan and Nahuatl peoples
to the west, and Lenca, Jicaque, and other less advanced groups to the
east. As Lehmann points out (1920, vol. 2, p. 625, and map), Maya
influence, as indicated primarily by archeological objects, extended
well into Lenca territory, including all the region west of a line drawn
from the junction of the Ulua and Comayagua Rivers southeast to the
Gulf of Fonseca. Similar influences were also present in the lower
Ulua valley and in Salvador. Moreover, Palacio (1860) clearly
indicates that cultural influences from the Nahuatl Pipil of Salvador
extended east well into Lenca territory during early historic times.
Whether Lehmann’s assumption that the language of the Lenca is re-
lated, on the one hand, with the Jicaque and the Paya, and on the
other, with the Cacaopera, Matagalpa, Sumu, Ulua, Miskito, Rama-
Guatuso, etc. (1920, vol. 2, p. 637), be accepted or not, there is little
doubt that the majority of these languages are affliated with major
linguistic stocks to the south. There is at least a strong probability
that the majority of these languages are in some degree related to the
Chibchan linguistic stock centering in northern South America. As
has been pointed out elsewhere (Strong, 1935, pp. 170-172), the scant
ethnological information on certain of these groups likewise points to
a southern derivation. On the other hand, Chol and Chorti Maya and
Nahautl (Pipil and Aztecan) linguistic connections are clearly with
the north. Thus the Ulua-Yojoa region comprised an important
buffer area between two sets of cultural traditions and linguistic
stocks, the one derived from Mexico and northern Central America,
the other from southern Central America and, eventually, from
South America. A very complex archeological situation is therefore
expectable. It is, however, a situation that, when clearly understood,
is certain to throw much new light on the ultimate derivation and
development of the higher civilizations of the New World.

Since the historic occupants of our particular region were the
Lenca and Jicaque Indians, we are particularly concerned with what-
ever ethnological information has survived concerning their cultural
status at the time of the Conquest. Regarding the Jicaque, little is
on record but for the Lenca, or at least their near neighbors and cul-
tural kin, we have the brief but excellent account of Palacio. Speak-
ing of the plain of Jiboa in the province of San Miguel, Salvador,
he says that here the Indians begin to speak a new language, called the
Chontal. He states that they “are a very rude people, but had
anciently a great reputation for valor among their neighbors.’ His
description of the customs observed prior to 1576 at Micla, (Mita),

2

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

apparently a cultural center representing a blending of Pipil and
Lenca ceremonies and customs, is so important that we quote it in
full, following Squier’s translation (Palacio, 1860, pp. 65-89).

Three leagues distant, is the village of Micla, which anciently the Pipil Indians
of this district held in great veneration; it was here they came to make their
offerings and sacrifices, as did also the Chontals, and other neighboring Indians
of different languages. Their modes of sacrificing differed in some respects from
those of other parts. They had cues or temples, and teupas or priests of high
authority, of which there are still many signs and traces.

Besides their cazique or secular lord, they had a kind of pope, called Tecti,
who dressed in a long blue robe, and wore on his head a diadem, or sometimes a
mitre embroidered with many colors, at the crown of which rose a cluster of
very beautiful feathers, taken from a bird, called in this country, Quetzal.
This pontiff carried in his hand a staff, which resembled the crook of a bishop,
and he was obeyed in all spiritual things. After him, next in sacerdotal authority,
was the tehu a matlini, who was the ablest diviner and the man best versed in
their ancient books and in their arts. He it was who made auguries and foretold
future events. After him were four priests called teupixquis, who went dressed
in long robes, falling to their feet, each of different color, black, red, green and
yellow. These were the counsellers of the pontiff, and directed all the super-
stitous ceremonies and follies of their religion. Their was also a kind of mayor-
domo, who had charge of the sacred jewels and the instruments of sacrifice. He
also opened the breasts of the victims of sacrifice, and tore out their hearts,
and performed such other personal services as were requisite. Besides all these
there were other functionaries, who played on the drums, trumpets and other
instruments used in convoking the people to the sacrifices.

ELECTION OF THE POPE AND PRIESTS

When the chief priest died, he was buried in his own house, seated in a
painted chair, and all the people mourned for him for fifteen days, with loud
cries and lamentations. They also fasted during this period; but when this was
over, the cazique and the wife, man or diviner selected a new pontiff by lot.
It was requisite that he should be one of the four priests above mentioned. When
the choice was made, they had great feasts and dances, and he who was chosen
drew blood from his tongue and private parts, and offered it in sacrifice to the
idols. He also named his successor in the priesthood, who was required to be
a son of the deceased pontiff, if he had left one, if not, the son of some other
priest. He filled also the other offices which at any time became vacant in the
teupas, or temples. They adored the rising sun, and had two idols, one repre-
senting a man, whom they called Quetzalcoatl, and the other a woman named
Itzqueye. All their sacrifices were made to them, and they had a calendar,
with days specially set apart for each one, and on these the sacrifices were made.

SACRIFICES

Each year they had two principal and very solemn sacrifices; one at the com-
mencement of summer, and the other at the beginning of winter. These were
made in the interior of the sacred place or temple, and were of boys between
the ages of six and twelve years, bastards, born among themselves.

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 13

MODE OF THESE SACRIFICES

They sounded their trumpets and drums for one day and night before the
sacrifice, and when the people were assembled, the four priests came out from
the temple, with four small braziers in which they burnt copal and caoutchouc ;
and the four together, turning in the direction of the rising sun, bent their knees
to it, offering incense, and reciting words of invocation. After this they separated,
and did the same in the direction of the four cardinal points, south, east, north and
west, preaching and explaining their rites and ceremonies. When the sermon
was finished, they retired within four houses or chapels which were built at the
four corners of the temple, and there rested for a little while. They next went
to the house of the high priest, which was close to the temple, and took thence
the boy who was to be sacrificed, and conducted him four times around the court
of the temple, dancing and singing. When the ceremony was finished, the high
priest came out of his house, with the second priest and mayordomo, and
ascended the steps of the temple, accompanied by the cazique and principal
Indians, who, however, stopped at the door of the sanctuary. The four priests
next seized the victim by his extremities, and the mayordomo coming out, with
little bells on his wrists and ankles, opened the left breast of the boy, tore out
his heart and handed it to the high priest, who put it into a little embroidered
purse, which he closed. The priests received the blood of the victim in four
jicaras, which are vessels made from the shell of a certain kind of fruit (the
calabash), and, descending one after another into the court, sprinkled it, with
their right hands, in the direction of the cardinal points. If any blood remained
over, they returned it to the high priest, who put it back, with the purse
containing the heart, into the body of the victim, which was interred in the
temple itself. This was the kind of sacrifice made at the opening of the two
seasons of the year.

The high priest, his second, and the four priests were accustomed to meet to
ascertain, by sorcery and enchantment, if they should make war, or if their foes
were coming to attack them; and if it appeared that such an event was to take
place, they called together the cazique and war chief, and advised them of the
approach of their enemies, and whether they should go to meet them. The
cazique then assembled the soldiers, and all went out to battle. If he was
victorious, he despatched a messenger to the high priest, advising him of the date
of the occurrence, and on this information the diviner ascertained to which of the
gods sacrifice was to be made. If to Quetzalcoatl, the ceremonies lasted fifteen
days; if to Itzqueye, five days, and on each day they sacrificed a prisoner. These
sacrifices were made as follows: All those who had taken part in the war,
returned home in order, singing and dancing, and bringing with them those who
were to be sacrificed, decorated with feathers and chalchiuites on their wrists
and ankles, and with strings of cacao beans around their necks, the captains
themselves conducting them in their midst. The pontiff and priests, at the head
of the people, went out to meet the victors, with music and dancing; and when
they encountered them, the captains delivered over the victims to be sacrificed,
to the high priest; after which all went together to the court of their teupa,
where they kept up the dancing night and day, for the periods above named. In
the middle of this court was placed a block or bench of stone, on which the victim
was stretched, the four priests holding him by the feet and hands. The sacrificer
then came forward, loaded with plumes and bells, with a knife of flint, with

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

which he opened the breast of the victim, and took out his heart, and tossed it
in the air in the direction of the four cardinal points, and finally threw it aloft
directly in the middle of the court, in this way soliciting the divinity to accept
the sacrifice, in return for the victory. This sacrifice was public to all the
Indians, great and small.

During this period, the soldiers returning from the war, could not cohabit with
their wives, but were obliged to sleep in certain calpules or barracks, which
were given up to them for the occasion, by the young men who were learning
the art of war. During the day they went to the houses of their women to eat
and drink, and from thence to their plantations, always however, leaving a
company to guard the town. The men sacrificed blood drawn from their private
parts, and he who had most wounds in these was reputed to be most valiant. The
women sacrificed blood drawn from their tongues and ears, and they sacrificed
their entire bodies, taking up the blood with cotton and offering it to their idols—
the men to Quetzalcoatl, and the women to Itzqueye.

Their superstitious ceremonies, at the time of planting their fields, were as
follows: They put in little cups of calabash the seeds which they had selected
for the purpose, and placed them before the altar of their idols. They next
dug a trench in the ground, in which they planted the seeds regularly, covering
them with earth; and over all they placed a large brazier, full of burning coals,
on which they sprinkled copal and caoutchouc. The four priests then drew blood
from their ears and nose, receiving it in certain large reeds, which they burnt
before their idols. At other times they drew blood from their tongues and
private members, and petitioned their gods to prosper the fruits of the earth,
and give them abundant harvests. The high priest, in sacrificing, drew blood
from the same parts, and with it anointed the feet and hands of the idols, invoking
the demon, who spoke with him, and told him what kind of weather would follow,
all of which was communicated to the people by the four priests, who always
concluded by ordering the men to have connection with their wives, and then
proceed to plant their fields. And such was the sacrifice of planting.

We come now to their sacrifices for hunting and fishing. They took a living
deer to the courtyard of the cue or temple which they had outside of the town,
where they strangled and skinned him, collecting all his blood in a vase, and
cutting in small pieces the liver, lungs and stomach. These were put aside, with
the heart, head and feet. They next cut up and cooked the deer by itself, and the
blood by itself, and while these were cooking they had their dances. Next the
high priest and his assistant took the head by the ears, and each of the four priests
one of the feet, and the mayordomo put the heart in a brazier and burned it,
with copal and caoutchouc, as incense to the idol of the god who was held to be
protector of hunting and fishing. When the dance was finished, the head and
feet were scorched in the fire before the idol, as an offering, and afterwards taken
to the house of the high-priest and eaten. The flesh and blood were then eaten
before the idol; and the same was done with all the animals which they offered
in sacrifice. When they sacrificed fish, the entrails were burnt before the idol.

When a woman was in travail, the midwives made her confess her sins; but
if this was not sufficient to hasten the birth, they made her husband do the same;
and finally, if the woman admitted illicit connection with any other man, they
went to his house and took his clothes and placed them beneath her; if this
failed, as a last resort, the husband sacrificed blood from his tongue and ears.
When the child was born, if a boy, they put in his hands a bow and arrows;

No. I HONDURAS—STRONG, KIDDER, AND PAUL | 15

if a girl, a spindle of cotton; and the mother made a streak of soot mixed with
water on the right foot of the child, which they believed would prevent it, when
grown up, from being lost in the woods. At the end of twelve days, the child
was taken to the priest, green branches being scattered under the feet of the
bearers. The priest gave it the name of its grandfather or grandmother, as the
case might be, and they presented it with cacao and fowls, which were the
offerings made to the priest. When it was taken back to the house, the mother
carried it to a river and bathed it, offering to the stream, cacao and copal, that
it should never do evil to the child.

As regards the rites for the dead; if the defunct were a cazique or captain,
or the wife or child of either, all the people mourned for four days and nights.
At the rising of the sun on the fifth day, the high priest announced that the soul
of the dead was with the gods, and that it was useless to mourn any longer.
They buried the dead man dressed in all of his riches, in a sitting posture, and
in his own house. Their manner of mourning during the four days and nights
resembled a mitote, in which they chanted the lineage and deeds of the dead.
If he were a cazique who died, the high priest and all the people, immediately
recognized as his successor his son or daughter; or, if he had neither, his
brother or nearest relative.

On such occasions they had great feasts, dances and sacrifices, and the new
chief entertained in his house all the priests and captains. If a common man died,
only his children and relatives mourned; and if a woman lost her child, she
reserved her milk for four days, without giving it to another; for they believed,
if she failed in this, the dead child would do the living one some injury. This
sacrifice they called navitia.

It was the office of the cacique to order the plantings, and direct the marriages.
They always married when young; and when the affair was arranged, and the
affanced groom met his future father-in-law he turned aside, as also did the
afhanced bride when she met her future mother-in-law. They did this, because
the devil had made them believe that such encounters would prevent their having
children. Marriages were celebrated in this wise: the male relatives of the
woman sought the bridegroom and made him bathe in a river; and the female
relatives of the woman did the same with the bride; they then wrapped each of
them in a new, white cloth, and took them to the house of the bride, where they
tied them up naked, in their garments. The relatives of the young man then
made presents to the bride of cloths, cottons, fowls and cacao, while those of
the bride gave presents of the same kind to the bridegroom; after which they
all feasted together. At these marriages the high priest and cazique were obliged
to be present.

Concerning relationship: They have a tree painted, with seven branches, which
represent the seven degrees of relationship in direct descent, within which no
person might marry, excepting those who had distinguished themselves in war,
but even these might not marry within three degrees of blood. In respect of the
line collateral, they made use of another tree with four branches, which repre-
sented the four degrees within which no one could marry.

Aside from other laws which these Indians possessed in common throughout
the province, those of this nation have the following as inviolable:

Whoever contemned or ridiculed the sacrifices to the idols, or the ceremonies
connected therewith, was condemned to death.

Whoever had connection with a strange woman, was condemned to death.

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Those who had carnal intercourse with relatives, within the degrees above
proscribed, both suffered death.

He who spoke libidinously with a married woman, or who made improper
signs to her, was banished and his property confiscated.

Whoever had commerce with a strange slave (one not his own ?) was himself
reduced to slavery, unless pardoned by the high priest for services in war.

Whoever wounded another, if the wound were serious, suffered death therefor.

Whoever violated a virgin was sacrificed.

Whoever lied was severely whipped; and if it were in any matter concerning
war, he was enslaved.

Those of the people who were not soldiers cultivated the plantations of the
cazique, pontiff and priests; and also gave a part of their own crops for the
support of the warriors.

This is what I have been able to gather concerning the manners and custom of
this people.

Near this place, is a high rocky hill from which flow two streams of water,
close to each other, one hot and the other cold. Here too is found an abundance
of spices, which the Indians use in their drink and food; and an earth which
resembles copperas, and which it must be judging from its effects. With this
they make a dye.

From here to the borders of the province of Chiquimula de la Sierra, the
country is for the most part high, of good temperature, abounding in pasturage,
and adapted for the support of cattle, and the cultivation of all kinds of grains.

In the portion of this province which lies in the direction of Gracios 4 Dios in
Honduras, are the Chontal Indians. While there, complaint was made to me
against a cazique of a place called Gotera, who since the time of his paganism
had had his private member split open, as was the custom anciently, among the
most valiant. In 1563, certain idolatrous Indians of another village called Cezori,
got together in a neighboring forest.where one of them performed the same
operation; and afterwards they circumcised four boys of twelve years of age,
in the Jewish manner, offering the blood to an idol of stone of a cylindrical form,
with a double visage and many eyes, called Icelaca. They say that he is the
god which knows the present and the past, and sees all things. Both his faces
were anointed with blood, and they sacrificed to him deer, fowls, rabbits, peppers,
and other things which they used in ancient times.

Torquemada (1723, lib. 3, cap. 41, vol. 1, p. 330) has recorded
a Lenca myth which, he says, was told him by the old people. Ac-
cording to them, 200 years before this time, there came to Cerquin
(Lehmann states, 1920, vol. 2, p. 636, that this was probably Corquin
in the Department of Gracias, Honduras), a lady, white as a Castilian,
whose name was Comicahual, meaning “ jaguar that flies”, so named
because she was very wise and versed in supernatural arts. These
Indians held the jaguar in high esteem. She made her abode in
Calcoquin, which was the most fertile land in the province. Here
there were stone “lions”? which they worshiped, and a large three-
pointed stone which had on each point three grotesque faces. Some
said that Comicahual carried it there through the air and by its virtue

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 17,

won battles, thus extending her realm. Some said that she had three
supernaturally conceived sons, others said they were her brothers.
When she grew old, she distributed her territories among them with
advice concerning the good treatment of her subjects. She then com-
manded that her bed be taken out of the house. Lightning flashed
and thunder roared. The people saw a beautiful bird flying across
the sky and, as they never saw the lady again, they believed she was
the bird and thus went to the sky. The sons (or brothers) divided
the realm and governed it well. The people were courageous and
warlike. They had been taught religion and enchantments by the
Lady Comicahual. Among the many idols which they adored, there
was one called the Great Father and another called the Great Mother.
To these idols they prayed for their well being. Other gods were
introduced, to whom they prayed for food, property, riches, and
that their lands might prosper, and produce abundantly. And, “ for
many years these superstitions and deceits of Satan lasted among the
old people.” Lehmann (1920, vol. 2, p. 637) is inclined to identify
the Calcoquin of Torquemada with the Icelaca of Palacio. He goes
on to show that similar rites, presumably originating with the Pipil as
indicated by Palacio, extended as far north as the Bay Islands in the
Caribbean. This evidence, derived from Salcedo, has already been
cited elsewhere (Strong, 1935, pp. 14, 15) and need not be repeated
here. Sufficient for our present purposes is the fact that elaborate
but basically similar cult observations extended from Salvador north
beyond the mouth of the Ulua River and that many of these at the
time of the Conquest seem to have originated in Pipil territories.
Only the results of scientific archeology can show whether this his-
toric Salvadorean center was actually primary or was derived from
still earlier sources of cultural development. This will be discussed
in relation to the results of our own archeological excavations.

Linguistically, the Lenca and the Jicaque have since been studied
by various travelers. This material has been summed up and amplified
by Lehmann (1920, vol. 2, pp. 649-722). From the ethnographic
standpoint, recent work on the Lenca and Jicaque has been pitifully
inadequate. Habel (1880) describes various Jicaque he met in the
Department of Yoro as follows:

The Xicaques differ in the form of their bodies from all the other tribes of
Central America. Their stature, on the average, being equal to that of Europeans,
is greater than that of the other tribes. Their skin is of a lighter color, and
their features resemble more closely those of the Caucasians, having a more
pleasant and intelligent expression than any other tribe of this region known to

me. Both sexes wear a kind of apron made of the inner bark of the Caoutchouc
tree. That of the women reaches around the waist and the ends hang down

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

from the hips to the knees. These two flaps are attached to the body by a strap
of the same material fastened around the waist. By another narrower strap, tied
around the head, they secure the long black hair, parted in front, floating down
to the shoulders.

According to Habel, the Jicaque had but recently been gathered
into permanent settlements through the splendid efforts of a Spanish
missionary. He adds that they were improvident, did not cultivate
the soil nor raise any large domesticated animals. They had formerly
been permitted to sell themselves into practical slavery, but this prac-
tice had then been stopped. They traded in sarsaparilla and tobacco.
Habel goes on to discuss the physical and other characteristics of the
still numerous Paya, who appeared to him to be much darker in pig-
mentation than the Jicaque. We have previously indicated that quite
primitive groups of Jicaque survive at the present time.

According to Otis T. Mason (1889), the Lenca of Honduras had
an ingenious method of straightening lance shafts. A pole about
16 feet long was suspended vertically from the limb of a tree by a
lariat attached by half hitches to both ends of the pole. At the lower
end, the lariat was attached to a rock weighing around 50 pounds, the
shaft being thus held straight while seasoning. He goes on to de-
scribe a variant of the musical bow used by the Lenca which was
called a “ bumbum.” This strung bow had a small gourd on the back
of the bend which was attached to the bow cord by another string
running at right angles. The bow was rested on a half gourd inverted
on the ground, which gave added resonance while playing (Mason,
1889). Apparently this instrument was not confined to the Lenca,
for Habel (1880, p. 31) describes an identical instrument used at
about the same time by the Pipil of the Balsam Coast of Salvador.
Here it was strung with wire and called the “ carimba.’”” The melody
was produced by strumming the wires with a stick and cupping the
hand over the gourd. Quite possibly this represents a variant of the
musical bow, or it may be a historic borrowing from the African
marimba so popular in Central America at present. Whether it is
primarily of New World or of African origin, we cannot say.

In June 1936 the junior and senior authors of the present report
were grounded by an airplane accident at the town of San Pedro
Sula in Honduras. While waiting for a track car, we were enter-
tained by a small native boy who, with a short stick, strummed dole-
fully on the identical instrument described by Mason. In this case
the bow string was of wire and the bow rested on an empty carton
instead of a gourd.

NOs E HONDURAS—STRONG, KIDDER, AND PAUL 19

Squier (1859, pp. 603-619) gives a brief but vivid picture of a
fiesta at Comayagua in which Indians from the nearby mountains
performed dances accompanied by much ceremonial drinking and
native ritual. The deer and the ocelot were the symbols of the two
main dancing groups. Their musical instruments consisted of flutes,
the Panspipe, the marimba, and a covered pot with a string drawn
through the bottom. At this fiesta, the Indians, the majority of whom
were probably Lenca, visited the numerous ruined towns in the
vicinity of Comayagua that had been occupied at the time of the
Conquest. He also described an extremely isolated village of the
Guajiquero Lenca and gives an amusing account of the difficulties
involved in securing linguistic or ethnographic information from the
Indians. As anyone knows who has attempted work with Honduras
Indians, the repression of almost half a millennium combined with
linguistic barriers is not an easy thing to overcome. However, it is
obvious from Squier’s account that a wealth of native custom and
belief still survives among the more isolated groups.

Such survivals, combined with the extremely haphazard nature
of previous research among the living Indians, indicate that there
is much more information available in Honduras for the trained
ethnologist and linguist than has been generally realized.

EARLY HISTORIC CONTACTS IN NORTHWESTERN HONDURAS

The fourth voyage of Columbus gives us our first historic glimpse
of conditions on the Honduras mainland. Having visited the Bay
Islands, Columbus landed at Punta de Caxinas (the Cape of Hon-
duras) on August 14, 1502. The chroniclers of this voyage give a
brief but vivid picture of the advanced agricultural life and the
thriving coastal trade then existing on the north coast of Honduras.’

In 1524 Gil Gonzalez named Puerto Caballos (later to become
Puerto Cortez) and established a settlement at San Gil de Buena
Vista. From this base he sailed down the coast and marched over-
land into the Olancho valley, where he met and defeated a force under
Hernando de Soto that had been exploring this region from Nica-
ragua. Returning to Puerto Caballos he was informed of the arrival
of a Spanish fleet under Cristobal de Olid.

It is of interest that again Honduras becomes a buffer area and
battleground between two earlier established southern and northern

* Pertinent historical and ethnographic information regarding the Bay Islands
and the adjacent mainland have been given elsewhere (Strong, 1935, pp. 7-19).
The following historical résumé of the Ulua region is primarily condensed from
Bancroft, History of Central America, vols. 1 and 2, 1883. Other sources are
cited as they occur.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

centers, the one in Panama under Pedrarias, the other in Mexico
under Hernando Cortez. Not content with the rich spoil of the Aztec
Empire, Cortez had already cast covetous eyes to the south where
rumor painted the golden glories of Hibueras or Honduras. For
this reason he despatched a trusted lieutenant with a fleet to conquer
the province. Having already reached an agreement with Velasquez,
Governor of Cuba and the rival of Cortez, Olid, in 1524, established
the settlement of Triunfo de la Cruz east of Puerto Caballos and
withdrew his allegiance from Cortez. The latter countered by dis-
patching another fleet under Las Casas, which proceeded from
Mexico to the Bay of Honduras. Olid promptly attacked Las Casas.
As Bancroft says, “it was an original spectacle in these parts, Span-
iards fighting Spaniards, in regular naval engagement; and as the
hissing projectiles flew out from the smoke over the still waters,
followed now and then by a crash, the noise reverberating over the
forest-clad hills, the dusky spectators should have been exceedingly
grateful for this free exhibition of the wisdom and power of Euro-
pean civilization that had come so far to instruct them in such a
fashion.”

Although the honors of battle, if any, went to Las Casas, a tropical
storm wrecked his fleet and he was forced to surrender. Along with
Gil Gonzales, who had also been captured by Olid, Las Casas was
taken inland to Olid’s headquarters newly established in the large
Indian town of Naco (see maps, figs. 1, 2). Here, although they
were treated as guests by their captor, the two captives plotted against
Olid, and eventually cut his throat with a table knife. Crawling away
into hiding, Olid sent for a priest. The latter being followed, Olid
was dragged into the plaza at Naco and publicly beheaded. Las Casas
returned to Mexico by an overland route through Guatemala. Even
today, over 400 years later, a tradition still persists among the un-
lettered inhabitants of present-day Naco that here “the king was
killed’ after being dragged in from his hiding place at El Salto, the
falls of the beautiful little Naco river. Here, as elsewhere in the
New World, European civilization was ushered in with blood and
treachery.

Meanwhile Cortez had had no word from his latest Honduras
venture. Despite the advice of his other lieutenants, he decided to
leave Mexico and proceed overland to Honduras.”

“The best sources on this amazing expedition are given in Maudslay’s trans-
lation of Bernal Diaz, “ The True History of the Conquest of New Spain, vol. 5”,
1916. The pertinent letters of Cortez to the Emperor Charles V are also included
in this volume.

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 21

It is of interest that before starting, Cortez obtained maps from
the Indians of the Vera Cruz region showing the entire area between
that point and Panama. It is apparent that he was traveling along
well-known aboriginal trade routes throughout most of his journey,
and he mentions that nearly all the towns he stopped in were full
of traders. The details of his Yucatan traverse do not particularly
concern us here until he arrived at Nito near the Gulf of Dulce. Here
he found the diseased, starving remnants of Gonzales’ colony. Mak-
ing an expedition up the Gulf of Dulce, Cortez captured a well-
provisioned pueblo and obtained supplies for continuing his journey.
From Nito, Cortez proceeded by sea to the vicinity of Puerto Caballos,
where he established a settlement. He sent Sandoval overland to
Naco. After crossing the Motagua River and visiting several pueblos,
Sandoval’s force arrived at Naco. The town had been recently de-
serted by its native inhabitants but contained abundant provisions and
even salt ; and here the Spaniards settled themselves, in the words of
Bernal Diaz, ‘‘as though we were going to stay there forever.” Ina
later section on the excavations at Naco, we will give more details
regarding native conditions in the vicinity of Naco at the time of
Sandoval’s visit.

Regarding the probable linguistic affiliations of the natives of Naco
and the adjacent pueblos, it is significant that Lehmann lists three
pueblos mentioned by Bernal Diaz “in the neighborhood of Naco ”
as having Nahuatl names.’

Similarly, Cortez states :

When I first arrived at this pueblo (San Andrés), I heard from the Spaniards
who had come from Naco that the natives of that pueblo and of the neighboring
pueblos, were somewhat disturbed, and had left their houses for the hills and
forests, and that although some of them had been reasoned with they refused to
be pacified from fear of the treatment that they had received at the hands of
the followers of Gil Gonzalez and Cristobal de Olid. I wrote the Captain in
charge there and told him to do all that he could to capture some of the natives
by whatever means he could devise, and to send them to me so that I could
speak to them and reassure them. This he did, and he sent me certain natives
whom he had captured during an expedition which he had undertaken, and I
talked to them and gained their confidence, and let them talk with some of the
native Chiefs from Mexico, whom I had brought with me. These Chiefs told
them who I was, what I had done in their country, and what good treatment
they had received from me when once we were friends, and how they were
protected and governed in justice—they and their property, their wives and
children—and the punishment that those received who rebelled against the
service of your Majesty, and many other things which they told them. After this,

Cee eee

*Lehmann, 1920, vol. 2, p. 1018. Also see Nahuatl distributions on linguistic
map.

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

they regained confidence, although they still told me that they had some fear
that they were not being told the truth, for those captains who had come in
advance of me had told them the same things and more to the same effect, and
that they had lied to them and had carried off their women when they had sent
them to make bread, and that the men who accompanied them had been forced
to carry loads, and they believed that I would do the same. Nevertheless, with
the assurances which the Mexicans and the Interpreter (Marina, a Mexican
woman) whom I had with me gave them, and seeing those of my company
happy and well treated, they were somewhat reassured. I sent them off to speak
to the Chiefs and people of the pueblos, and in a few days the Captain at Naco .
wrote me that some of the neighboring pueblos had become peaceful, particularly
the chief pueblos which are: Naco, where the Spaniards are residing, Quimiztlan,
Sula and Tholoma (Cheloma)—the smallest of these had more than two thousand
houses—and other villages which were subject to them; and that the envoys
said the whole country would soon be at peace, for they had sent messengers to
pacify the people, telling them of my arrival among them and all that I had said
to them, and also what they had heard from the natives of Mexico; they added
that they greatly desired that I would go to Naco, as my arrival there would give
confidence to the people. This I would have done with good will, had it not
been very necessary for me to continue my journey in order to arrange that
which I shall explain to your Majesty in the following chapter.”

From the foregoing it seems quite possible that the people of Naco
spoke a Nahuatl dialect understandable to the Aztec caciques and
to Dona Marina, Cortez’ famous Mexican Indian woman interpreter.
Had the temporary captives from Naco been Jicaque, Lenca or Maya,
this would not have been possible. It is also possible that certain
Nahuatl dialects served as a lingua franca in the area, due to the
obviously extensive trade connections then in existence with Mexico
and to the extent of Pipil influence exerted from Salvador. How-
ever, elsewhere Cortez mentions linguistic difficulties when entirely
alien languages were encountered by his men, but this does not seem
to have been the case here.

Cortez then proceeded by sea to the newly founded town of Tru-
jillo. His settlement near Puerto Caballos was soon abandoned, owing
to sickness and lack of food, in favor of Naco. A number of large
and rich pueblos in this vicinity were gradually conciliated by San-
doval, but the inhabitants of Naco, owing to the severe treatment
they had received from Olid, refused to return to their homes. While
Sandoval was at Naco, the caciques of two pueblos named Quespan
and Talchinalchapa came to him to report the depredations of some
other Spaniards who had arrived from the South.” These were seized
and proved to be a party under Garro from Nicaragua that had been

* Bernal Diaz, 1916, vol. 5, p. 407, from the fifth letter of Hernando Cortez to
the Emperor Charles V; also see p. 60.
1 Bernal Diaz, 1916, vol. 5, p. 66.

NON HONDURAS—STRONG, KIDDER, AND PAUL 23

sent to claim the lands to the north for Pedrarias. They were well
equipped with arms and horses and had handsome Nicaraguan Indian
women with them. Sandoval sent them under guard to Cortez at
Trujillo. Bernal Diaz, who was with this overland party, describes
the difficulties and the Indian fights they encountered. Unless one
has actually traveled through these mountainous, tropical countries,
it is impossible to appreciate how truly amazing such early Spanish
journeys were. Even today an overland trip from Naco into Nicara-
gua would be an expedition not to be undertaken lightly. Yet in the
time of Cortez, Spanish adventurers seem to have already traversed
the Central American cordilleras from end to end. Cortez returned
Garro to Nicaragua with messages of good will and mining supplies.
For some time Cortez toyed with the idea of adding Olancho and
Nicaragua to his conquests and even went so far as to start a road
from Trujillo to Nicaragua! However, a mission from Mexico
arrived with bad news, and the road is still unbuilt.

Hearing that his holdings in Mexico had been seized by enemies,
Cortez determined to return at once. Before departing he ordered
Luis Marin with a number of discontented colonists from Trujillo
to proceed to Naco where there was abundant good land. Saavedra,
who was then campaigning in Olancho, was to remain as Governor
of Honduras. After a hard trip Marin arrived at Naco, and the next
day, in company with Sandoval, set out on the overland trip through
hostile Guatemala to Mexico. Meanwhile Cortez, who, strange to say,
appears to have been a very bad and timorous sailor, had been driven
back by storms. Messengers were sent to Sandoval ordering him
to stop and settle. This was a great blow to the overland party, for
they desired above all else to return to Mexico. Sandoval hurried to
Trujillo to plead with Cortez, that he set sail and let the overland
party proceed. Under Marin the latter went “to some pueblos
called Maniani and thence to another pueblo named Acalteca, where
at that time there were many houses.”

Despite Sandoval’s plea, Cortez still refused to sail. Sandoval was
dispatched to Olancho where he drove out Rojas, a lieutenant of
Pedrarias. On Sandoval’s return, Cortez sent orders to Marin to
proceed, and he ordered Godoy, who was forming a settlement at
Puerto Caballos, to go to Naco with all his people. Finally, in 1526,
Cortez set out for Cuba and, eventually, Mexico.

Finally, Pedro de Alvarado, having received orders from Cortez to
proceed from Guatemala to Honduras, began his march. Marin,

* Bernal Diaz, 1916, vol. 5, p. 86.

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

desperate and without orders, sent a party of Io men through Olancho
to go to Trujillo. According to Bernal Diaz, they got as far as the
gold-working region on the Guayape River, when they learned of
Cortez’ departure. Receiving orders from Saavedra to return, they
did so, and, Bernal Diaz remembers, they threw stones at the country
as they left. They met Marin at the pueblo of Acalteca and then
proceeded to another pueblo called Maniani, where they encountered
six of Alvarado’s soldiers. In two days’ marching they reached
Alvarado “near the town called Chuluteca Malcala.” This was
probably the site of Tegucigalpa on the Choluteca River. From here
the combined parties proceeded toward Guatemala after a difficult
crossing of the Lempa River, which was then in flood.

Years later, Bernal Diaz (1916, vol. 5, pp. 328, 329) thus recalled
the country of Naco and of the Ulua River, as it was when he first
saw it and as it soon became:

and what I state I know, for when I came with Cortés on the expedition to
Honduras I was present in Trujillo, which was called by the Indian name of
Guaimura, and I was at Naco and the Rio de Pichin, and that of Balama, and
that of Ulua, and in nearly all of the pueblos of that neighborhood, and it was
thickly peopled and at peace (and the people were living) in their houses with
their wives and children; but as soon as those bad governors came they destroyed
them to such an extent, that in the year fifteen hundred and fifty one, when I
passed through there on my return from Castile, two Caciques who had known
me in the old days, told me with tears in their eyes of all their misfortunes and
the treatment (they had received), and I was shocked to see the country in such
a condition.

The details of this tragic and complex period in Honduran history
cannot be considered here. The withdrawal of Cortez threw the new
colony into turmoil and the starving colonists engaged in every form
of intrigue. Coming from Guatemala, Pedro de Alvarado took over
the governorship and set about pacifying the country. He built the
town of San Juan at Puerto Caballos and founded San Pedro. For
the Indians this was an even more tragic period. According to Ban-
croft (vol. 7, pp. 233-234) Indian slaves were kidnapped and sold in
Honduras by the shipload. In the vicinity of Trujillo where there had
been villages of from 600 to 3,000 houses, there were not more than
180 Indians left in 1547. Those not enslaved or killed had fled to the
mountains. At Naco, where there had originally been a population
of about 10,000, there were, in 1536, only 45 remaining. At La Haga,
a coastal town some g leagues from Trujillo, there had been about
goo houses, but of the entire population, only the daughter of the
Cacique remained. The cruelty toward the natives was even greater
than in Guatemala. In 1539, when Alvarado returned from Spain

IORI HONDURAS—STRONG, KIDDER, AND PAUL 25

and transported the materials for building a fleet across the isthmus,
the entire remaining Indian population fled. These evils were pre-
sented in full detail by Bartolome de Las Casas, and the new laws
resulting from his famous publication at least gave nominal protection
to the oppressed natives.

In answer to a petition from Trujillo, the Emperor appointed
Francisco de Montejo, the former governor of Yucatan, as ruler
of Honduras. Only a handful of starving Spanish colonists remained.
Montejo subdued but did not enslave the Indians of the mountains
near Trujillo. Many Indians returned voluntarily to their homes in
this region. Montejo then visited the town of Gracias a Dios. Here,
owing to the murder of a Spaniard, he arrested and punished the
Lenca Indian ring leaders in the presence of the Caciques of the
district of Cerquin previously referred to. This aroused the opposi-
tion of the famous Lenca leader Lempira, “ Lord of the Mountains.”
Lempira had previously withstood Alvarado and driven off Spanish
attacks under Chavez, and he now opposed Montejo. The great
Indian leader had secured allies from various interior tribes including
several that had formerly been hostile to the Lenca, and was estimated
to have a force of some 30,000 warriors.

According to Lehmann (1920, vol. 2, p. 637), he had united the
men of more than 200 towns and commanded over 2,000 ‘‘ men and
gentlemen of distinction.”

“Lempira, the last of the chiefs of Corquin, made his final stand
against the Spaniards on the mountains of Piriera, which overlooks
the valley of the river Lempa, in the name of which beautiful stream
his own is commemorated.” (Squier, 1858, p. 320.)

Here for 6 months he was besieged by Caceres, a lieutenant of
Montejo, but so greatly were the Spaniards harassed by the Indians
that they were on the point of failure. Siege and assault having
failed, Caceres resorted to treachery. Under a flag of truce Lempira
came to the walls of his stronghold to parley with his enemies and
was shot by a hidden marksman. The Lenca and their allies fled,
and the great conspiracy soon fell to pieces.

Mrs. Popenoe, quoting from a letter from Montejo to the King
of Spain, June 1, 1539, gives the following account of the latter part
of this campaign against the Lenca:

Disturbing news reached Gracias, where Montejo was sojourning with 11

Spanish soldiers. The Indians were preparing stubbornly to resist him. In
Yamala, a nearby village, they were building many houses on a great, very

*D. H. Popenoe, 1936, pp. 559-560. For the original, see Coleccion de Docu-
mentos Inéditos, 1864, vol. 2, pp. 212-266.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

strong rock which they have, and providing them with provisions. The Spanish
chieftain sent a Negro spy, who knew the language of the Indians, to enter the
stronghold and bring back a report. The frightened Negro found there four
houses built very large, and four more larger ones full of corn, and he set fire
to the houses and to the corn. Word came of a great disaster in the valley of
Comayagua. The Indians had risen. One Spaniard had been killed and several
others wounded. Four horses had been lost. Unable longer to withstand the
siege, the Spaniards had fled at night to a neighboring province where the
inhabitants were friendly.

Montejo realized that the time had come for desperate action. Supplies were
brought together, and soldiers were called in from regions where the danger of
rebellion was not imminent. Others who had been wounded but now had
recovered sufficiently to join the colors, augmented the small band which was
placed under the leadership of Alonzo de Caceres, recently returned from the final
campaign against Lempira.

When they arrived at Comayagua they found that the Indians, doubtless
apprised of their approach, with all available supplies would fortify themselves on
big rocks. Cattle which they could not take with them had been killed and
eaten, so that the valley was now in a state of starvation.

Montejo advanced into one part of the valley, Caceres into another, attacking
and capturing a mountain fortress “which was the strongest in that region.”
The last named leader then proceeded to a village, by name Guaxerequi, where
six Christians had recently been killed. There he found another fortress. At
this point he was rejoined by Montejo, who describes the place in his letter.
He says: “and (has) seen (or visited) a great rock, which was the strongest
thing that has been seen, which, if they had time to cut a ridge of mountain,
which they were cutting, would be impossible to capture, for they had on it
water and wood and cultivated fields and many provisions; they had 220 large
houses, and certain temples and places of worship.”

It took the combined forces of Montejo and Caceres four months to conquer
the valley of Comayagua, after which they carried the campaign into Olancho.

Such stories as the above throw much light on the importance of fortified
mountain tops at the time of the Conquest. Although it has been impossible to
place Tenampua (the famous archeological site near Comayagua, first described
by Squier, 1858 and 1869, see map, fig. 1), among the strongholds described in
the early accounts at my disposal, it seems probable that it may have been one
of those captured during the campaign carried out in the Comayagua region by
Francisco de Montejo and his lieutenant, Alonzo de Caceres. It may have been
the formidable Guaxerequi described in Montejo’s letter.

’

In the light of the partially cut “cuchillo” or narrow neck con-
necting Tenampua with the main promontory to the northeast (D. H.
Popenoe, 1936, pp. 562, 563 and map), I am inclined to believe that
this identification of Guaxerequi and Tenampua is indeed very
probable.

It is certain that a complete combing of the sources, combined with
first-hand examination of the available archives in Honduras and
neighboring countries, would yield a considerable mass of informa-
tion on the Lenca and their neighbors, but this is not possible at

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 27

present. All that has been attempted here is to suggest the main trends
of a fascinating historical period and to indicate the probable dis-
tribution of ethnic groups in the region under investigation. We turn
now to outlining the results of direct archeological research.

ARCHEOLOGICAL EXPLORATIONS
CHAMELECON RIVER

Our reconnaissance of the middle Chamelecon River extended from
May 26 until June 17, 1936. It was aimed primarily at Naco but
several other sites were also investigated. Through the courtesy
of the United Fruit Company we lodged comfortably at Manacal
Ranch (map, fig. 2) which is located about a mile south of the town
of Cofradia. Here we obtained horses and mules and were thus able
to work at a number of archeological sites in the general vicinity.
We first visited the San Luis site just above the confluence of the
Naco and Chamelecon Rivers (map, fig. 2). Next we spent 2 weeks
mapping and digging exploratory trenches at Naco. The remainder
of the time was occupied in making stratigraphic sections and maps
at the prehistoric Las Vegas and Tres Piedras mound sites.

NACO

All Honduras records of the Conquest refer to Naco, first as a
thriving Indian town and later as the site of repeated Spanish settle-
ments. The Indian pueblo of Naco was only one of a considerable
group in the vicinity. Montejo states that the original population of
Naco was 10,000 persons (Colleccién de Documentos Inéditos, 1864,
vol. 2, p. 228), an estimate that agrees reasonably well with the previ-
ously cited statement of Cortez that the smallest of the pueblos in that
vicinity had more than 2,000 houses. Las Casas, speaking of Hon-
duras, says: “Tenia Pueblos innumerables, y una vega de treinta
leguas y mas, toda muy poblada . . . la ciudad de Naco que tenia
sobre dos cientas mil animas, y muchos edificios de piedra, en especial
los templos en que adoraban”’ (cited by Bancroft, Native Races, vol.
4, p. 77). When compared to the other authorities, as well as to the
size of the ruins, this would seem to be an extremely exaggerated
estimate. Similarly, his statement (Las Casas, 1822, p. 45) that be-
tween the years 1524 to 1535 more than 2,000,000 Indians perished in
the kingdom of Naco and Honduras, leaving only 2,000 inhabitants in
a territory 100 leagues square, must be taken with a large grain of
salt. Diaz, Montejo, and others give ample proof that the natives of
Honduras were cruelly despoiled and that whole districts were de-

3

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS ~— VOL. 97

populated in the early days of the Conquest. Nevertheless, the
wholesale statistics of Las Casas seem to be those of a crusader
rather than a historian.

Bernal Diaz (1916, vol. 5, pp. 56-59) gives a first-hand picture
of Naco as it was in 1525.

At the hour of Mass we went to Naco. At that time it was a good pueblo,
but we found it had been deserted that very day, and we took up our quarters
in some very large courts where they had beheaded Crist6bal de Olid. The
pueblo was well provisioned with maize and beans and Chili peppers, and we
also found a little salt which was the thing we needed most, and there we settled
ourselves with our baggage as though we were going to stay there forever. In
this pueblo is the best water we have found in New Spain, and a tree which
in the noonday heat, be the sun ever so fierce, appears to refresh the heart
with its shade, and there falls from it a sort of very fine dew which comforts
the head. At that time this pueblo was thickly peopled and in a good situation,
and there was fruit of the Zapotes, both of the red and small kind, and it was
in the neighborhood of other pueblos.

.. . When we arrived at the Pueblo of Naco and had collected maize, beans
and peppers, we captured three chieftains in the maizefields and Sandoval coaxed
them and gave them beads from Castile, and begged them to go and summon
the other caciques and we would do them no harm whatever. They set off as
they were ordered to do, and two caciques came in, but Sandoval could not
induce them to repeople the pueblo, only to bring a little food from time to time;
they did us neither good nor harm, nor we to them, and thus we continued for
the first days. . . . When Sandoval saw that the neighboring Indians and natives
of Naco did not want to come and settle in the pueblo, although he sent to
summon them many times, and that the people of the neighboring pueblos did
not come or take any notice of us, he decided to go himself and manage to make
them come. We went at once to some pueblos called Girimonga and Agula, and
to three other pueblos near Naco, and all of them came to give fealty to His
Majesty. Then we went to Quimistan [Quimistlan in preceding chapter, Quimi-
stan on map] and to other pueblos of the Sierra, and they too came in, so that
all the Indians of that district submitted, and as nothing was demanded of them
beyond what they were inclined to give, their submission did not weigh on them,
and in this manner all was pacified as far up as to where Cortés founded the
town which is now called Puerto de Caballos.

Modern Naco is a small village of perhaps a dozen mud-walled
and thatched houses on the beautiful little Naco River. Permission
to excavate was kindly granted us by the son of the owner, Dr. Paz
Barraona, and by Don Santiago Nolasco, the head man of the village.
Don Santiago and the other residents of Naco were interested specta-
tors or laborers during our work here and the children brought us
many fragmentary specimens from the adjacent river banks. The
heart of the site is still covered by the small but very dense shade
trees mentioned by Bernal Diaz. These shaded our work but made
mapping difficult. Noontime siestas spent under great jungle trees

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 29

bordering the rapid, sparkling, Naco River made us appreciate the
remark of the soldier-historian that “here is the best water we have
found in New Spain.”

It is.not the purpose of the present report to discuss fully the
excavations at each site nor to analyze the archeological findings in
any detail. Instead, a brief summary of significant excavations will
be given, and at least one stratigraphic or horizontal artifact record
at each site will be outlined in an effort to indicate the apparent
trend of local cultural development. This preliminary analysis will
be confirmed or amended in the final report in accord with the full
statistical findings and in relation to all the excavations. Although no
numerical record of artifact or ceramic types is given at this time,
an effort has been made to discuss them quantitatively rather than
selectively. In regard to ceramics, which greatly preponderate over
any other artifact types throughout the entire Ulua drainage, we
have here attempted to suggest the relative proportions of all wares
at each site or in each stratigraphic section discussed. When the very
extensive sherd collections of the expedition have been analyzed and
the data fully presented it will be possible to check this preliminary
analysis against the complete record. In regard to technical names
applied to various soil layers these have been used in a very general
sense. When our soil samples have been fully studied by experts it
may be possible to supplement the cultural record with the detailed
findings of soil analyst and sedimentation expert.

As previously indicated, the ruins around Naco are extensive, and
our detailed survey deals with only the central area. The map (fig. 3)
gives the essential data in regard to mound orientation and elevations.
In general, the Naco mounds are low and rounded, apparently form-
ing the foundations of houses, but the group just northeast of the ball
court differs in this regard. Mound 6 appears to have been the center
of the complex. It is still the highest and was, in all probability,
originally faced with squared stones, forming a square-faced pyramid
with a flat top (fig. 3). A few of the cut stones are still in place.
It has been sadly damaged by the disruptive effects of tree growth.
According to local authorities it has also suffered by an earthquake,
by having its stone facing removed for road foundations, and, about
1902, by treasure-hunting excavations. It is still quite impressive,
however. Mound 6 is flanked by mounds 3, 4, 5, all of which are
exceptionally large. To judge by mound 3, which we cross-sectioned
near its southern end (fig. 4), mounds 3 and possibly 5 were origin-
ally capped by thick white plaster. This had eroded off the steep
sides of mound 5 but was present at the base and over the flattened

30

Curving Mounds 2 and 1

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HONDURAS—STRONG, KIDDER, AND PAUL 31

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32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

top (fig. 4). Mound 3 included an inner structure the nature of
which could not be satisfactorily determined by our one cross trench.
This occurrence of two thin plaster walls running through the heart
of the mound is shown in the illustrations (pl. 1, 2 and text fig. 4).
A small trench in mound 9 (fig. 3) revealed aboriginal refuse and dis-
articulated human remains. Owing to its proximity to certain historic
burials, work here was discontinued.

Horizontal stripping of mound 1 (fig. 3) revealed considerable
portions of the floors of two houses with massed small boulders on
the north side and tumbled adobe blocks on the south side (Strong,
1936, fig. 68). The plastered floors were stained a rich, dark red.
I‘ragments of plaster apparently from the walls showed five succes-
sive layers of red, yellow, red, blue gray, and red indicating the
varying washes used in decorating the interiors of the houses. These
colors were very fresh when uncovered but have since faded slightly.
To judge from our test cuts, these long, low mounds north of the
central pyramid complex consist of rows of house floors. Owing to
the curve of mounds 2 and 14 (fig. 3) they enclose a crescentic area
which may have been the old plaza of Naco. Our excavations,
although very incomplete, indicate that with adequate time a whole
series of house floors could be easily cleared. The earth covering
them is shallow, and the floors are intact. Such work would be of
the greatest value in revealing actual living conditions in aboriginal
Naco. When the potsherds from inside mound 1 were being washed,
we encountered two pieces of European glazed crockery. One of these
(pl. 4, ™) was obviously an early Spanish piece, the other might
possibly have been intrusive from more recent times. Since it was in
these houses that Olid, Bernal Diaz, and other Conquistadores lived,
further excavations here might cast light on early historic as well as
late prehistoric events. Certainly this association of early European
and late Indian ceramics links the prehistoric and the early historic
periods in Honduras. We also cross-sectioned mound 19, which is
located about 30 meters east of mound 17 but beyond the edge of the
map. This mound was about 1 meter high and 15 meters in diameter.
It contained sherds and snail shells to a depth of 35 centimeters but
no structural features of any sort. Here, as elsewhere in the vicinity
of Naco, the underlying soil is hard and gravelly, making excavations
below or beyond the artificially built or accumulated earth structures
extremely difficult.

There remains to be mentioned the ball court. For present pur-
poses the general diagram (fig. 3) and the photographs (pl. 2, fig. 1,
and Strong, 1936, fig. 69) show the main features. Excavations here

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 33

were confined to the southeastern end, owing to the presence in the
other end of modern burials placed here under the impression that
the structure was a colonial church. The discovery of a portion of
one of the ball court rings in the center of the north wall was of
especial interest (pl. 2, fig. 1). A complete ring of very similar type,
said to have been found at Naco by Sr. Roque Hernandez of San
Pedro Sula and presented by him to Mrs. Dorothy H. Popenoe, is
now at Lancetilla. It is possible that this specimen came from one of
the neighboring pueblos, since none of the present Naco inhabitants
remember its discovery and removal. As with the other structures at:
Naco, the ball court will be described in more detail at a later time.

We searched in vain for any large refuse heaps along the Naco
River. Scattered sherds occur where the deeply worn trail leads
down the steep gravelly bank to the river at the village but we found
no thick deposits. The children brought us various fragmentary arti-
facts from along these banks but could show us no concentrated
deposits. We saw little at Naco indicating any great antiquity, but
our impressions were based on only limited study. Naco appears as a
one-culture site, and we obtained no indication of stratigraphic
changes.

Artifacts do not appear to be very abundant at Naco, although
considerable broken pottery occurs in the various mounds dnd scat-
tered along the river bank. The bulk of the ceramic remains here as
elsewhere in the Ulua region are from monochrome cooking vessels
which, so far as present knowledge goes, are rarely distinctive. From
the samples preserved, this ware appears to be primarily dull red in
color ranging from smoke-stained black to gray. For the most part it
is fairly well polished, but a considerable portion has artificially
scratched and roughened surfaces. Sizes are highly variable, but
medium to small vessels seem to predominate. Rims are usually
direct or slightly flaring; broad strap handles, notched flanges
(pl. 3, 2) and projecting lugs occur; and bottoms are either flat or
slightly dimpled. Plain ware legs do not occur in our sample. The
tempering of all the Naco wares is a fine micaceous grit. Particu-
larly significant, though much less abundant, is the Naco painted
ware (pl. 3). Characteristically, this ware has a white slip and
painted, geometric or curvilinear decorations on both sides in red and
black. One sherd (pl. 3, a) of this ware is unusual in showing a
portion of what appears to be a plumed figure. The painted vessels
appear to have been small and flat-bottomed with either direct or
slightly flaring rims. Tripod supports containing rattles are very
common in Naco painted ware (pl. 3, s-w). A strange, four-pointed,

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

bird- or animal-head foot is most the common type (pl. 3, t, w). The
painted ware is not very well made, and the designs are usually
badly eroded. A small proportion of unpainted and a few painted
sherds have either heavily incised or raised geometric designs (pl. 4,
q, S-V, *-z) in the interior. These were apparently made by some
sort of a stamp. In one case the raised design left by the stamp was
smoothed down and its outer border carved for emphasis. Three
plain sherds show finely woven textile designs impressed on their
inner surface (pl. 4, n-p). Sherds with incised designs also occur
(pl. 4, 7). On the whole, Naco ceramics consist of these two wares,
the plain and the painted, but in excavations at mound 1, two intrusive
types occurred. The first of these, consisting of two fragments of
European crockery (pl. 4, m), has already been mentioned. The
second type consisted of three sherds of well-made, highly polished
and painted ware which apparently belong to prehistoric ceramic series
from other earlier sites on the Chamelecon and Ulua Rivers.

Incensario fragments from Naco are of the usual frying pan shape
(pl. 4, @) with the distinctive Naco painted designs. Two candelarios
(pl. 4, w) are crude but unique. They are made of unslipped coarse
pottery and have tripod supports. They represent the only type found
by us at Naco. Spindle whorls are quite common at Naco and are well
decorated with incised designs similar to those painted on pottery
(pl. 4, 7, 7). Undecorated “ bobbins ”’, probably to hold cotton thread,
are even more common (pl. 4,7). The occurrence of spindle whorls,
bobbins and textile-marked pottery bears witness to the importance
of cloth manufacture in aboriginal Naco. No distinctive type of
figurine was noted at Naco. The various pottery heads, ranging from
simple to complex, and the “speak no evil”? monkey, are illustrated
(pl. 4, b-f). Whistles seem rare at this site. Only one specimen
was found (pl. 4, 2) and this animal form suggests Chiriqui, although
the red and black paint design is in the Naco style. The only other
artifacts noted were the ubiquitous obsidian prismatic flake knives
(pl. 4, k), and fragmentary legged metates and manos of lava. At
Naco, as elsewhere in Honduras, there appears to have been an amaz-
ing emphasis on pottery in comparison with any other type of non-
perishable artifact. Textiles and wooden implements have left only
indirect evidences of their probable importance.

LAS VEGAS |

This site, also known locally as “ Potrerito de los Calpullis ”, is
located less than 1 kilometer in a direct line and about 2 kilometers
by trail from Manacal (see map, fig. 2). It is a neatly arranged

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 35

mound group and is one of the few in the Ulua-Chamelecon region
that can be photographed to advantage (see Strong, 1937, fig. 70).
The main features are four large mounds forming a rough square,
with another low mound in the center. The largest mound, to the
north, is about 2 meters in height, 27 meters in length, and 12 meters
in width. The others are slightly smaller, those to the east and south
being rounded rather than rectangular. The eastern mound had a
trench, made by workmen from Manacal, in the east side. The central
mound is about I meter in height with a diameter of 8 meters. It is
connected with the eastern mound by a low neck. The four main
mounds roughly correspond with the cardinal points, but there is no
exact orientation. A low, stone-covered mound is located about
40 meters to the west. The entire group is located on an open strip
of high, flat land, flanked on the east by a deep gully and on the south
and west by the steep river banks. An artificial terrace of river
boulders borders the site to the south. Behind the site rise rolling
pine-covered hills, and between it and the river proper is a densely
wooded flood plain.

No artifacts occur on the surface other than a very few sherds
of plain brown ware. A rounded boulder in the central plaza suggested
an ape’s head somewhat similar to that shown in plate 16, figure 3,
but we were unable to determine whether the stone had been actually
worked. The men who had dug the deep trench in the eastern mound
encountered nothing but stones and broken pottery. Pottery is visible
in the cut to a depth of 2.5 meters. We ran a trench through the
heart of the low central mound reaching a depth of 14 meters in the
center. The upper meter consisted of soil with many large boulders,
stones, and a few pieces of plaster; below this was hard gravel. A
few lava metate and mano fragments and a considerable amount of
plain, brown pot sherds came from the upper meter. The Las Vegas
ceramic remains are predominantly of an unslipped brown ware
indistinguishable from cooking ware at Naco and in Ulua Polychrome
sites. However, a few polished and slipped sherds occur, and some
of these have linear designs in red and black. A few sherds of dull
orange ware with red stripes, a small orange rim with red and black
monkeys, and a hollow round tripod leg were also found.

TRES PIEDRAS

According to our workmen, this site received its modern name
“because it is a place where they catch many fish’’, a puzzling ex-
planation unless one is aware that the name “Tres Piedras”? may
be given to any person or place of particular potency. In a sense

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Tres Piedras is a very miniature Copan since the Chamelecon River
has nicely cross-sectioned it (pl. 2, fig. 4). It is located less than a

kilometer down stream from Las Vegas and on the same or western ~

bank (map, fig. 2).

Originally, the site must have resembled Las Vegas in outward
appearance, having four mounds enclosing a central plaza. In the
photograph (pl. 2, fig. 4) two of these mounds can be seen in cross-
section on the right and left of the cut; the rear mound is visible
to the left of the figures, but the fourth or nearest mound has been
completely washed away except for the many boulders deposited
in the river channel. A fifth mound, likewise cross-sectioned, occurs
to the west. Unlike Las Vegas, the plaza at Tres Piedras had a series
of three plaster floors, the highest at a depth of 1.5 meters below the
present surface, the lowest at a depth of 2 meters. The material
was a thick, white “ mezcla” or plaster. The upper floor appeared
to be flat, but the two lower floors each had one step rising to the
east. From our limited excavations it was impossible to tell how
extensive these floors originally may have been. Along the river bank
they extended for about Io meters, and a considerable amount of
broken plaster was visible elsewhere on this general level and in the
talus deposit at the foot of the bank. It seems probable that the entire
court or plaza between the mounds was once paved, but until adequate
excavations are made here this cannot be proved, nor can the nature
of the steps or mound approaches be determined. Over the plaster to
a depth of three-fourths of a meter is a thick deposit of large river
boulders. These may have rolled down from the mounds or may have
been placed here later to raise the level.

Among the vast quantity of stones deposited in the river bed from
the portion of the site that has been washed away are many that
indicate human workmanship. The most tantalizing of these are a
considerable number of large lava blocks that strongly suggest
sculpture in the round. None, however, are definite enough for
certainty, but they do give an impression of either a dying or a
nascent sculptural drive. The “ape’s head” from Las Vegas is of
this type and may have been transported there from Tres Piedras.
In addition, there are numerous squared blocks of limestone or gray-

green schist, one circular block with abrupt edges, and several thick .

slabs with holes drilled through them. Metate and mano fragments,
as well as lapstones without legs, occur. Stones are particularly
concentrated in the river bed below what was once the position of
the east mound. With them occur large fragments of plaster flooring.
This flooring often contains boulders or shows the molds from which

*

NO.’ I HONDURAS—STRONG, KIDDER, AND PAUL a7

boulders came. Structurally, the Tres Piedras site appears to have
been more pretentious than the majority of sites in the vicinity, and the
remaining half is well worthy of complete excavation.

We made two small stratigraphic cuts, the first west of the central
mound group between that and the outlying mounds to the west. The
second was in the heart of the plaza. Both sections were on the face
of the river bank and each was 5 meters long by 1 meter broad,
extending down to sterile soil. The first excavation yielded some
pottery at 10 centimeters and reached barren soil at about 1.70 meters.
The second cut passed through three plaster floors and reached barren
soil at 2 meters. The first yielded the most potsherds although even
here they were not overly abundant. Gray to red cooking ware was
most abundant in each of the five 30-centimeter levels. A very few
sherds of Mayoid polychrome occurred in all but the bottom level.
Above I meter all fragments of this type were from pots with buff
to orange slips covered with florid, conventionalized, red, purplish,
and black designs. Below 1 meter the same Mayoid types occurred,
but in. association with more realistic designs having human head
panels. At this same level occurred fragments of an excellent Mayoid
vessel with a panel of square, grotesque heads around the rim and,
below this, an intricately carved design. The design had been carved
after firing. Associated with these lower Mayoid types were a few
sherds suggesting orange over buff negative painting; and cooking
ware with dull, dark red line decoration. The second excavation in
the plaza yielded few but similar pot sherds. However, the occurrence
of a small, restorable imitation Ulua marble bowl just above the
upper floor at a depth of 14 meters was significant. Maya carved
ware occurred at this same depth in the first excavation. The three
plaster floors in this second excavation have already been mentioned.

OTHER SITES

There are numerous mounds and other archeological sites in this
region, but time to examine many of them was lacking. Close to
Manacal Ranch is the site of Los Cocos, consisting of a few low earth
and stone mounds that are being rapidly eaten away by the river
(map, fig. 2). There is a deep 30-foot bank at this place. No notable
structural details could be observed. The only pottery we obtained
were some coarse, blackish brown sherds and one heavy, dull orange,
sherd with eroded red and black designs.

Farther upstream, beyond the mouth of the Naco River, is the site
of San Luis (see map, fig. 2). Here in a cut of some 3 meters occur
many river boulders and large amounts of broken pottery. The

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

majority of the pottery is a coarse, brown or buff ware. There are
also a number of heavy dull orange pieces with broad red stripes
and some polish ; as well as a polished red incised piece and a fragment
of a heavy platter with coarse red and black line decoration. In the
talus below this bank were two large, square cut stones of volcanic
origin. A few crumbling human bones were also found in the bank
and on the talus. There are no surface mounds at San Luis, but
broken pottery occurs from just below the present surface to a depth
of about 3 meters. No stratigraphic changes in type occur so far
as our very small pottery sample is concerned, but the site merits much
more careful study than we were able to give it.

As one rides past Cofradia on the way to Naco a few low mounds
are visible to the south of the road just after one has crossed the
Manchagualay River. We did not examine these in detail. Farther
along the Naco road, about 1 kilometer from that village, there is a
small Spanish colonial ruin located in dense bush about 20 meters
north of the road. It is the foundation of a small house made of bricks
and plaster, and the local people have tales concerning a magical
cross of gold that was once found here. As already stated, the ruins
of Naco extend for about 1 kilometer up the Naco River, and there are
said to be numerous mounds across the river from the modern town.’
We visited the site of Quebrada Tostada, about two leagues upstream
from Naco in a hanging valley some 400 feet above El Salto, the wild
and beautiful falls of the Naco River. The main site at Quebrada
Tostada includes 4 or 5 acres of pine- and thorn-covered land. Low
stone and earth mounds are scattered over this area, and we found a
few sherds of coarse brown pottery. Local tradition says that “the
King”, i. e., Olid, fled to El Salto after he was wounded. We cut
our way down the steep, rough gorge to the falls but found no signs
of any settlement there. Our guide, Don Santiago Nolasco, said
that there were many low mounds scattered over the hills and moun-
tain valleys in the general vicinity of Quebrada Tostada, but he knew
of no nearby site comparable in size to the ruins at Naco.

There is another important site in this general vicinity which we
had hoped to visit. This is the Bell Cave, which Blackeston (1910)
located near the headwaters of a small stream flowing into the
Chamelecon River, about 25 miles from the ruins of Naco. Blackes-
ton obtained a considerable number of copper bells and a few other
unusual artifacts at this site. We were told by Sr. Roque Hernandez
of San Pedro Sula that the site was not yet exhausted. Just before
we left Manacal, Sr. Juan Antonio Sarmiento of San Antonio Mahada
offered to guide us to the cave which he said was near his home.

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 39

Unfortunately, we were unable to make the trip. It would be very
important to learn what types of pottery, if any, occur in association
with these copper bells. Spinden (1925, p. 544) has suggested that
the cache formed part of a Toltec trader’s outfit.

ULUA AND COMAYAGUA RIVERS

Our most extensive excavations on the Ulua proper were at Las
Flores Bolsa and at Playa de los Muertos. In addition, ceramic
samples were obtained at various river bank and mound sites between
Naranjo Chino and the mouth of the Comayagua (see map, fig. 5).
Our investigations were for the most part confined to the eastern
bank of the Ulua. In a region as rich in sites as is the Ulua, it
seemed better to confine our efforts to a few promising places rather
than attempt too wide a survey. The depths at which cultural layers
occur necessitated moving dirt on a very large scale for even a
reasonable ‘stratigraphic sample. On the Comayagua River, near |
Santa Rita, we made excavations similar to those at Las Flores Bolsa
and at Playa de los Muertos.

LAS FLORES BOLSA

Las Flores Bolsa is located on the east bank of the Ulua River
just south of the division line between the Las Flores and Naranjo
Chino banana plantations. This was the farthest down-river site
excavated by the expedition (see map, fig. 5). We worked here from
January 20 to February 20, 1936. The site was chosen because of the
fact that examination of the steep river bank from a dugout canoe
revealed several human skeletons one above the other at this place.
We therefore hoped for some sort of stratification. This was also
the exact place where O. P. Swofford found a deformed skull with
filed and inlaid teeth and with a jade bead in its mouth. This skull
has been described as that of a Maya chieftain from Santa Ana (see
Blom, Grosjean, and Cummins, 1933). It should be noted that the
Las Flores site is actually a considerable distance downstream from
Santa Ana (see map, fig. 5). In addition to fragmentary human
bones there was a considerable amount of broken pottery projecting
from the bank and on the small talus at the water’s edge.

We made two deep stratigraphic cuts paralleling the bank and ex-
tending down to the water line. At the time of our work the almost
vertical river bank was 5.25 meters in height. Excavation I was
approximately 10 meters long by 4 meters wide. The top 2 meters
was a recent sand and silt. Cultural debris, mainly broken pottery,

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Naranjo Chino 5

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BASED ON TELA RAILROAD CO. MAP, OCTOBER, 1934.

Fic. 5—Map of the lower Ulua and Chamelecon Rivers.

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 4I

occurred in the heavy clay below this for 2.25 meters; below this was
a sterile light clay loam extending to and below the water level. From
the point where we struck the first artifact (at a depth of 2 meters),
the soil was stripped off in successive layers each 25 centimeters
thick, the first one being designated as P 1 (i. e., pottery level 1)
and so on through the occupation level. In all, 13 burials were en-
countered in this excavation, 10 extended (P 5-9) and 3 bundle
burials (P 4-6). The skeletons were in crumbly condition, and the
skulls were badly distorted by the pressure of the earth. Only two
complete skulls could be saved. Grave gifts were sparse, no complete
pots occurring with any of the burials. One bundle burial had 2 clay
spindle whorls (top of P 4), and another bundle burial (P 6) had 1
copper fishhook, 16 obsidian flake knives with needle sharp points, and
a broken cooking pot containing bird bones. Layers of small adobe
bricks and small baked clay basins near certain burials were the main
structural features encountered. Broken pottery was quite abundant
in this excavation, coming from 10 levels. The types and sequence
represented will be discussed in the final report.

Excavation 2 was 16 meters east of excavation 1. It was roughly
5 meters long, 4 meters wide, and 5.40 meters deep in the deepest
portion. It contained only one extended skeleton (P 3). From the
surface, mixed sand and silt extended down a little less than 2
meters ; here the soil changed to a light clay. This layer of light clay,
without artifacts, extended down slightly more than 1 meter. Be-
neath this was a dark, heavy clay containing artifacts. Artifacts
occurred throughout 7 levels or 1.75 meters. The cultural deposit
sloped down toward the south (i. e., toward the river) so that it
extended to the top of P 8 there, whereas on the north side of the
excavation it terminated on top of P 4. A living level occurred
in P 3 and it is the termination of this which sloped down to P 8 on
the river side suggesting a refuse heap. In absolute level, P 1 in
excavation 2 corresponds to P 4 in excavation 1. Obviously, the
deposition of refuse at this site had been little disturbed by burials.
On the other hand, the occurrence of only three levels on the north
side with what appears to be a dump heap (correlated with level P 3)
on the south suggests that the deposit represented a relatively uni-
form period of no very great duration. A bed of coarse sand occurred
at a depth of 4.25 meters and below this was a light clay loam extending
to and below the water level. Except for the sloping dump on the south
edge, this stratum was devoid of artifacts.

The various ceramic and artifact types from excavation 2 will be
briefly discussed and any obvious stratigraphic changes noted.

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Pottery predominates tremendously over any other form of artifact,
and plain or domestic wares are much more abundant than decorated
wares. In all layers at this site the pottery shows the effects of water
action, and the surfaces of many sherds are eroded. There is, however,
no observable indication of re-deposition. The majority of sherds
from all levels are of unslipped, undecorated wares ranging in color
from a smoked or burned black, through brick red or brown to light
buff. Sizes are highly variable. Pots with constricted and medium
flaring lips are common, as are direct rimmed bowls. Vertical strap
and solid round handles are most abundant. Many of these, from
all levels, have a knob, filleting, or a crude monkey head on the
bend. Bottoms are flat, rounded, dimpled, and annular, the first
three types being most abundant. A few plain, hollow, conical feet
occur. In P 6 and 7, large, thick, highly polished sherds also occur.
Since grit tempering seems practically universal in the Ulua-Yojoa
region, it may be taken for granted unless variants are mentioned.
Domestic (that is coarse or household) ware with painted decoration
is rare at this site. It occurs sparsely in P 3 and 4 where large
vessels with high flaring necks are decorated with rayed circles, cross-
hatches, or lines of dull red or brown paint applied in a splotchy
fashion. Plain incised ware is rare but occurs in P 4 and 5 where
necks are decorated with delicate, wavy, comblike patterns forming
both vertical and horizontal patterns.

With the wares which are both painted and incised we pass out
of the strictly utilitarian class and find several intergrading types.
A striking Las Flores type occurs in levels P 3-5 (pl. 5, a, b, c, d, e).
These sherds are from thick-walled, vertical vases or bowls with high
vertical necks, having a polished red slip, a band of black geometric
designs below the lip and another band of incised design below this
(compare Strong, 1935, pl. 18, fig. 1, b, c, e, for similar Bay Island
types). Another striking and distinctive ware, which occurs com-
monly at Santa Rita (farm 17) (pl. 7, a-d) and rarely at Lake Yojoa
(pl. 14, d), we have here called the Bold Geometric, monkey-handled
type. It is very similar if not identical with Bay Island Polychrome
II ware figured elsewhere (Strong, 1935, fig. 11). This is found
in all levels at excavation 2 but undergoes some change in the two
bottom levels. The typical vessel is large, with an orange slip and
intricate black and red geometric designs around the neck, the body,
and on the handles. The neck design is often of the interlocking textile
type (compare pl. 5, c, and Strong, 1934a, fig. 54, and 1935, fig.11),
and the handle at the bend usually has a monkey head in relief with
modeled or punctate features. At Las Flores, excavation 2, numerous

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 43

sherds of this type have incised as well as painted designs around
the neck. The more or less realistic birds and animals occurring on
vessels of this type from the lower levels at Santa Rita (farm 17)
(pl. 7, b-d) are lacking at Las Flores. In levels P 6-7, vessels of this
type have lower necks, irregular handles, and incised as well as painted
designs.

Polychrome sherds from thin-walled, vertical vases of so-called
Mayoid type occur in all levels in excavation 2. The majority have
florid, conventionalized, all-over designs in red, black, white or purple
on buff, orange, black, or white slips (pl. 5, f, g, h, 1, 7, k, 1, m).

The majority of designs are elaborated and extremely conventional-
ized reptilian, animal, mask, or anthropomorphic forms. They often
cover the entire surface of the vessel and are difficult or impossible
to reconstruct in their entirety from sherds. Crude skeuomorphic
glyph bands occur from P 5-7, as do elaborately modeled projecting
monkey or animal head lugs in the same levels (pl. 5, f, g). ‘In some
cases the designs are outlined with incisions. In the upper levels
several sherds with red and purple spots occur (pl. 5,:7). Bases are
flat, dimpled or annular, and hollow cylindrical as well as solid, thin,
rectangular, tripod legs occur in all levels. None of the isolated
and graceful processional or “ dancing ”’ figures occur in excavation
2, although a few sherds with this type of decoration were found in the
deepest levels of excavation 1.

In addition to polychrome, straight-walled vases, a number of low
bowls or small jars have similar types of designs. In P 1-2 occur
polished red or orange sherds. In P 3 there are fragments of about
six small jars with solid rectangular, tripod feet and eroded black and
red designs. From P 4 to P 7, small tripod jars and low bowls with
an orange slip, and red and black conventional or crudely realistic
designs are common. These are in the Mayoid rather than the Bold
Geometric tradition, though an occasional blending between these
major styles occurs. In some instances incision is used to outline
painted designs. In P 7 was found an unusual, restorable bowl of
thin, polished ware, with an orange slip, and conventional, black
and red, monkey and rosette designs outlined with incisions, a dimple
base, and a low “ vestigial”? spout to one side of the direct rim
(pl. 6, b). Three similar low “ vestigial” spouts occurred in P 2-3
as well; hence they cannot be regarded as strictly early at Las Flores.

From P 5-7 came a few fragments of Mayoid sculptured pottery.
A restorable tripod vessel of this type is painted all over with an
orange wash, except for the carved panel of elaborate Mayoid faces
which apparently had no slip (pl. 6, d). A tiny vessel with a similar

4

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

face panel is brown with no slip. It has an annular base and in shape
rather suggests certain of the Ulua marble bowl types, though the
sculptured design is Mayoid. A third fragment is the rounded
bottom of a bowl with intricate Mayoid design in high but rounded
relief. The slip was originally red but has disappeared except between
the raised designs, and glittering micaceous tempering material shows
on the surface. If it were not for Lothrop’s statement (1936a, p. 142)
that this mold-made appearance is due to delicate carving and the
obscuring effect of the slip, one would be inclined to regard these as
stamped or molded rather than carved. The type will repay much more
detailed analysis than is possible here. From P 5 and 6 come three
small restorable pots of the imitation Ulua marble bowl type (pl. 6,
e, f). There are a few other sherds of this type. No slip is visible
on these pieces, though all are considerably eroded. The association
of Mayoid sculptured ware and imitation Ulua marble bowl pieces
in the same levels may very well be significant.

Incensario fragments occur in every level except P 1 and P 7.
All seem to be of the usual perforated frying pan type with hollow,
round handles. They lack painted decoration and range from light
buff to brown in color. Fragments from P 2 and P 6 are very thin
and delicate, but a fragment from P 3 is thick and crude. Candelarios,
or small incense burners, occur in P 3 and P 5. All are of the un-
slipped, single-hole type. That from P 3 is undecorated, whereas the
two fragments from P 5 have crude linear incision and punctate
ornamentation. Cassava-grinders, or round, handled, disks of coarse
pottery, with one surface ridged with cross-hatched incisions like a
grater, occur from P 3 to P 6. They are most numerous in P 3.
Spindle whorls occurred only once, with burial A I in level P 4.
Of the three, two were plain and one had neatly incised decorations.

Figurines and whistles occur in practically all levels. They show
little change in types from top to bottom. Solid, mold-made figurines
of Mayoid type (like fig. 7, s) occur in P 2, 5, and 6. A portion
of a pottery figurine mold was found in P 5. Modeled figures of thin,
polished, brown pottery occur from P 2 to P 7. Some of these were
originally rather pretentious (pl. 6, a), but nearly all are very frag-
mentary and their original form often cannot be determined. Besides
the human figurines and larger hollow statues, both solid and hollow
animal and bird heads occur in all levels. Many of these were probably
from whistles (like fig. 7, a, c, e). Similarly, many of the human
figures once formed parts of whistles. Strange bulbous animal forms
occur from P 2 to P 7. Some of these were whistles, others were not.
A particularly interesting whistle from P 2 is in the form of a realistic

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 45

frog with a small one on its back (compare Gordon, 1898, pl. 9, 7, 7).
Pottery stamps likewise occur from top to bottom. From P 1 comes
a round, stemmed stamp with a neat monkey design; from P 5 a
rectangular, stemmed stamp with a squirrel design and a butterfly-
shaped stamp with two crude faces; from P 6 an elaborate froglike
stamp with small circles for designs, and, from P 7 a rectangular
stamp with a geometric design.

Compared to the amount of pottery recovered from this excavation,
the total list of other artifacts is pitifully small. P 1 yielded 1 broken
T-shaped drill of obsidian ; P 2, 30 fragments of obsidian flake knives,
t crude obsidian drill, 2 polished pebbles, 1 piece of crudely flaked
quartzite ; P 3, 2 crudely chipped stones, 1 polished pebble, 2 pieces
of baked clay with wattle and daub impressions ; P 4, 1 lump of clay;
P 5, 6 fragments of obsidian knives; P 6, 1 obsidian knife fragment ;
2 quartzite stones, 1 smoothed piece of baked clay, 1 large alligator
(?) bone with 2 perforations, I tapering, cylindrical brick of baked
clay ; and P 7, the butt end of a small celt of hard green stone. This
slim list clearly indicates what a tremendous proportion of the ancient
material culture was perishable. Were it not for the advanced and
abundant ceramic remains in prehistoric Ulua sites, one might reason-
ably, but erroneously, conclude that only a very simple prehistoric
culture had flourished there.

SANTA RITA (FARM 17)

Excavation work was carried on at this site by Dr. and Mrs. Kidder
from the middle of March until the rising water level drove them out
of the excavations on May 20, 1936. Work was also going on at
Lake Yojoa, but all the other members of the expedition spent some
time in the Santa Rita excavations. The site is located on the
Comayagua River just below the little town of Santa Rita (map,
fig. 5). It consists of refuse deposits and living levels exposed in
the steep banks of a flood channel of the river and is only 200 meters
west of the overseer’s house on farm 17 of the Tela Railroad Co.
The main irrigation canal for the lower valley takes out from the
Comayagua just east of the overseer’s house. Thanks to the courtesy
of the Tela Railroad Co. and of the overseer, Mr. John Thompson,
we were able to board comfortably at the farm house and to use its
broad porches for sorting specimens.

The physiographic and cultural evidences revealed by the Santa
Rita excavations are complex and require far more detailed treatment
than is possible here. However, certain very significant correlations

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

between these factors are already apparent, and these can be briefly
outlined. In all, three adjoining excavations were made at this site,
the main stratigraphic cut designated as excavation 1; a northern ex-
tension of this cut resulting from the discovery toward the close
of the work of older and deeper cultural material; and excavation 2
extending through a polychrome refuse heap to the east. For present
purposes we will confine our remarks to excavation I and, to a
lesser extent, to the northern extension. The beginnings of all these
cuts can be seen in the illustration (pl. 2, fig. 3).

A cross-section (fig. 6) of the west end of excavation I shows the
outstanding stratigraphic features. This excavation was originally
5 meters long from east to west, paralleling the cut river bank, and
4.5 meters in width from north to south. Owing to the outward slope
of the bank, the bottom of the excavation was 8 meters in breadth
(fig. 6). When the May floods made further work impossible, we had
reached a depth of 5.20 meters in excavation 1 and 5.40 meters in
the northern extension. In size, the northern extension was less than
one-third of excavation I.

The cross-section along the west wall of excavation 1 (fig. 6) shows
the various soil layers. The 2 upper meters consist of alternating
deposits of dark silt, light silt, and sand. Below this is a thick deposit
of dense clay which terminates at a total depth of 3.80 meters in a
thin bed of sand or sandy silt (level 8, fig. 6). This sand layer has
here a slight dip from the north and thins out near the southern
edge. Beneath this layer is a sandy clay (level 9, fig. 6) which, with
certain minor changes, extends down to the bottom of our excava-
tion. On the extreme southern edge and in the deepest portion is a
deposit of sand and gravel (fig. 6) which ran the length of the exca-
vation and may represent an old stream bed. In the west wall cross-
section the sand layer (level 8, fig. 6) seems to dip toward this sand
and gravel deposit but, on the east wall cross-section, the sand is
much thicker (40 cm) and extends on a level plane to the edge of
the bank at a point 1 meter above the lower sand and gravel deposit.

The first potsherds and other cultural detritus occur in the dense
clay deposit simultaneously with a layer of river boulders (fig. 6).
Throughout this clay deposit polychrome pottery is abundant, as are
other cultural manifestations. The polychrome debris is thickest in
a definite refuse deposit on the southern edge which dips slightly
less than I meter below the main clay and pottery-bearing stratum
(refuse heap, fig. 6). Debris extends down almost to the low sand
and gravel deposit suggesting that it had been dumped over a low
bank at the edge of an old water course. The refuse heap here

HONDURAS—STRONG, KIDDER, AND PAUL 47

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48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

terminates before the east wall of excavation I is reached, but beyond
this point another polychrome refuse heap at the same depth occurs
and it was in this that we made excavation 2. The sand level (level 8,
fig. 6) was sterile of artifacts throughout all of our excavations. In
excavation I, the same was true of the underlying sandy clay (level 9)
except on the southern edge where the overlying polychrome refuse
material dipped, and at the northern edge where a very few mono-
chrome potsherds were encountered (level 9, fig. 6). These latter,
found toward the end of our work, seemed highly significant and
for this reason the northern extension was made extending north
from the northwest corner of excavation 1. Soil layers and pottery
deposits were generally similar in excavation I and in the northern
extension. However, in the northern extension more abundant pot-
sherds differing from the polychrome type were found in the sandy
clay (level 9, fig. 6) beneath the sterile sand stratum (level 8). This
will be discussed subsequently.

Cultural features, other than abundant potsherds and rare artifacts,
were not marked in any of the excavations. River boulders occurred
throughout the upper portions of the main clay stratum (level 7,
fig. 6) in all. What appears to have been a roasting pit or oven is out-
lined in figure 6. Small clay-lined fire pits and small irregular clay
bricks also occurred. Eight burials, all in bad condition, were en-
countered, four in excavation 1 and four in the northern extension.
In all cases these occurred either in the clay stratum (level 1) con-
taining polychrome pottery or just below it and clearly intrusive
into the sand. All were extremely friable and crumbled on exposure
to the air. One of the burials in excavation I was extended and had
notched upper incisors, three were flexed, only one had any grave gift
(a ground stone knife). In the northern extension, one burial con-
sisted merely of an immature skull, jaw, and humerus; one was
extended ; one was flexed; and the last was the extended skeleton of
a new-born child under a large, plain red, two-handled bowl. Excava-
tion 2 yielded no burials.

The succession of ceramic and artifact types from excavation I
and the northern extension will be briefly outlined. This site was
richer in polychrome pottery than any other we dug on the Ulua, but
it should be remembered that even here there was much more plain
than painted ware. The sherds from this site show little erosion
through direct water action and the majority of the painted pieces
are fresh and bright. We will discuss the material according to four
major stratigraphic levels, A (F 1-3, see fig. 6), B (P 4-6), C (P
7-9), and D (P 10-12). As indicated on the diagram (fig. 6) levels

NO. I : HONDURAS—STRONG, KIDDER, AND PAUL 49

A include the upper portion of the clay occupation stratum. The
southern face of the cut, including the uppermost portion of the
refuse heap, had been removed to expose skeleton B I prior to com-
pleting the diagram. Levels B include the lower portion of the clay
occupation level and the upper portion of the southern dump heap.
Levels C, the very bottom of the clay occupation level and the middle
of the dump heap. Levels D include the lowest portion of the dump.
With the exception of this southern polychrome dump heap, most of
the remainder of levels C, and all of levels D, were devoid of artifacts
except in the extreme northern portion where a very few bichrome
sherds were encountered beneath the sand stratum (level 8, fig. 6).
These will be discussed separately.

Levels A contained a large amount of plain cooking ware of a
red brown to blackish gray color. The vessels were fairly large, in-
cluding direct bowls and pots with flaring necks and vertical handles.
These handles are either round or flat in cross-section and, in a few
cases, have a monkey head lug on the bend. Rounded, flat and
dimpled bases and a few conical and round hollow feet occur. There
are also some highly polished thin sherds tan or buff in color. The
upper portion of B contained the same types but in the lower portions
crudely painted ware superseded the plain cooking ware. In C and D
plain cooking ware was very scarce except for a few very thick
gray and brown sherds and some vertical strap handles. A portion of
a thick, plain tray with horizontal handles occurs in C, and a plain
annular base in D. Similar cooking ware but deccrated with blotchy
red or brown designs on neck and body occurs in A. These designs
are usually rayed circles, criss-cross lines, and more or less irregular
blotches. In B this type supersedes the plain ware in the lower levels.
A squat, swollen pot form with flaring neck and vertical handles is
characteristic. These are better made than in A, and the dull red,
criss-cross line decoration on a lighter background sometimes sug-
gests negative painting. This type also predominates in both C and
D where undecorated domestic wares are rare. Plain incised ware
is lacking in all levels. From both A and B levels came a few similar
pot fragments in which the neck of the vessel is also incised with
delicate, wavy, vertical lines and where the handle is replaced by small
tripartite adornos. This variant of the swollen, simply-painted pot is
more numerous in the lower levels, 1. e., C and D. In D there is some
blending of this type with the Bold Geometric, monkey-handled ware.
Three sherds from thick-walled, vertical vases have a slip and painted
designs with a band of heavily incised decoration around the upper

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

body. They all come from A and closely resemble the more numerous
representatives of the type from Las Flores Bolsa (pl. 5, a-e).

The Bold Geometric polychrome type occurs in all of the lower
major levels at farm 17. In A it occurs only in the lower third (i. e.,
level P 3, fig. 6). The vessels of the characteristic swollen olla type
are medium rather than large in size. Textile and geometric designs
are common (figs. 8, 9, compare Strong, 1934a, fig. 54, p. 46), but
conventional birds and animals are lacking. In B similar designs
occur in the upper two-thirds, and a few animal and bird designs occur
in the lower third. In C and D animalistic designs (pl. 7, b-d, and
fig. 10) are common, but geometric and numerous textile motifs also
occur. The Bold Geometric vessels of the lower levels appear to have
been slightly larger and better finished than those from the upper
levels. Characteristic cursive, conventionalized bird, feline, bat, and
reptile designs from the lower levels are illustrated (pl. 7, b-d, and
fig. 10) and their association with geometric motifs indicated. In Da
few Bold Geometric type vessels have incised patterns on the neck,
similar to the squat, painted and incised domestic ware previously
described. Bold Geometric monkey-handled bowls are numerous at
this site and, with the straight-walled Mayoid vases, constitute one
of the two most distinctive ceramic types.

Straight-walled, vertical vases of Mayoid type are represented by
sherds from all four major levels in excavation 1. In A the pre-
dominant, painted decorations are complex over-all designs on white,
black, orange, or yellow backgrounds. Designs are in red, black,
white, purple, and, in one case, blue.

As at Las Flores Bolsa (pl. 5, f-m), the majority of the design
motifs from A are elaborately conventionalized monster animal or
human forms. One large fragment has a conventionalized jaguar
with a row of conventionalized human heads above. An elaborately
modeled and. painted monkey-head lug occurs, as do hollow cylindrical
feet and two annular bases. In B similar types occur, with the addi-
tion of textile designs and the common occurrence of bands of con-
ventionalized heads of several types (compare upper panel, pl. 8, a, b,
and fig. 13). Squat, elaborated human or deity figures (pl. 8, d, and
fig. 13) occur in this horizon and one of these is outlined with carved
lines. An elaborate modeled monkey-head lug and a black monkey
in low relief on a painted bowl came from levels B. One sherd with
blue paint used as a design was noted.

In C, panels containing paired “ dancing figures” occur for the
first time. This unique design motif, on beautifully polished and
painted pottery, has been noted from northern Honduras to Salvador.

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 51

Lehmann (1910, p. 740 and illus. 8, p. 736) believes that copulation,
not dancing, is indicated by this design and supports his view by a
drawing of a Salvadorean example. To us, the latter seems no more
definite than do the Ulua examples here illustrated (pl. 8, a, b,
and fig. 14). In the light of Palacio’s information regarding the cere-
monial importance of the mutilation of male genitalia among Pipil
and Lenca, we rather incline to connect this widespread design with
phallic rather than procreative rites. Undoubtedly, the correlation of
outer dancing figures with a unique design inside such vessels (fig. 14)
is significant. This peculiar, and always slightly variable, inner design
suggests some sort of record. It occurs inside “ dancing figure ” and
certain processional vases and bowls from the Ulua River, Comayagua
River, and Lake Yojoa. We suspect it also occurs inside Salvadorean
vessels. This is an extremely interesting problem which at this time
may only be mentioned in passing. Associated with the “dancers ”
are sherds decorated with isolated, processional figures. Like the
“dancers”, these are usually well proportioned and graceful. The
manner in which they are fitted into the simpler but more beautiful
panels and design areas contrasts markedly with the florid, over-all
designs of the upper levels. With these more realistic figures occur a
variety of conventionalized human head designs (pl. 8, a, b). Simi-
larly, the squat, conventionalized deity or priest figures (pl. 8, d,
and fig. 13) also occur in association with the well-proportioned
“ dancers ” and processional figures. Flat bases are most common in
this level, and tripod feet are usually solid and rectangular or ovoid,
though a few cylindrical feet occur. Lugs and annular bases do not
occur in our sample. Levels D are identical with levels C so far as the
Mayoid cylindrical vase shapes and designs are concerned. As was
true of the Bold Geometric ware there is here also a slight but obvious
development from the realistic to the conventional in painted decora-
tions. It is significant, however, that during the time involved in
these stylistic changes, neither the basic form of the Mayoid straight-
walled vase or the Bold Geometric monkey-handled pot changed in
any very marked degree.

Fragments of Mayoid sculptured ware, as well as some examples
of carved designs, come from B and C. From levels C there are two
fragments from small jars in imitation Ulua marble bowl style (com-
pare pl. 6, e, fy. Here, as at Las Flores, Mayoid sculptured ware
and imitation Ulua marble bowl incised ware are in close association.
At Las Flores, excavation 2, these are in the lowest levels; at Santa
Rita, excavation I, in the two middle levels.

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS — VOL. 97

Fic. 7.—Hollow figurines, whistles, and “ candelario”, from the Ulua Poly-
chrome period, Santa Rita (farm 17). (Specimens in National Museum of
Honduras at Tegucigalpa.)

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 53

Numerous small polychrome jars and vases are represented in
excavation I. Certain of these are Mayoid, others Bold Geometric,
and still others suggest blends between the two (compare figs. IT,
12, 15). Any attempt to clearly delineate these two major Ulua
polychrome styles, or to demonstrate the exact nature of their blend-
ing, would necessitate a far more extensive analysis of design motifs
than is possible here. Considered very generally, however, there
are certain top-to-bottom variations which seem to be significant. In
A, small red bowls with black geometric designs and conventionalized
animal or anthropomorphic designs, either outside or inside, occur.
Some of these are definitely Mayoid in feeling, having circle, diamond,
or feather designs and dimpled bottoms. The majority, however,
seem more closely allied to the Bold Geometric type. In C, an orange
tone is particularly prevalent and numerous pieces show a rather
unique blending of Mayoid and Bold Geometric styles (compare
figs. II, 12, 15). Conventionalized birds, animals, and reptiles occur
both outside and inside open bowls (compare the similar bat designs
on two vessels from approximately the same levels in excavation 2,
where one (fig. 15) has a Mayoid, the other (fig. 10) a Bold Geometric
feeling). Both flat and dimpled bottoms occur in C. In D there are
numerous small flat-bottomed jars of Mayoid type with processional
figures and other elaborate anthropomorphic designs, and open bowls
with Bold Geometric designs on the inside. These less clearly pro-
nounced vessel forms, therefore, seem to recapitulate the tendency
to change from realistic to geometric decoration observed elsewhere.

A few fragments of polished gray ware came from Levels A, B,
and-C. The fragments were from small, slightly pear-shaped bowls
without legs or handles. One fragmentary vessel had a narrow band
of red paint around the inside of the neck. Another interesting feature
in levels C is represented by two very definite spouts of red and brown
polished ware. They are more similar to those from the deep layers
at Playa de los Muertos (pls. 10, 11) than to the “ vestigial” spouts
from Las Flores (pl. 6, b). Strange to say, no Plumbate ware oc-
curred in any of our excavations.

At Santa Rita there are two other distinctive polychrome vessel
types, both of which were lacking in excavation 2 at Las Flores. One
of these, a flat plate on high tripod legs (pl. 8, e, f), may be called a ©
tripod plate. The other, with somewhat higher walls, and either low
(fig. 8) or high (compare pl. 12, f) tripod feet, may be termed a
tripod dish. In excavation 1, tripod dish fragments are lacking in A,
fairly abundant in B, still more numerous in, CG. but-rare in Diy Dhey
characteristically have more or less intricate and geometric, red and

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

black designs on a light red or orange back ground (fig. 8 and pl. 7, ¢).
An unusual vessel of this type from excavation 2, which has loop
handles and an auxiliary annular base, is also figured (fig. 9). In
general, at Santa Rita, these vessels are in the Bold Geometric
style, though elsewhere (as at Lake Yojoa, pl. 12, f) they may be
more Mayoid. Tripod plate fragments or restorable pieces (pl. 8,
e, f) are rare in levels A and B, fairly numerous in C, and rather
abundant in D. Characteristically, the tripod legs are high with
vertical slits and contain rattles. The plates are heavy and flat with
slightly inward-dipping rims (pl. 8, f). The designs are often

[_] veLvow purr RED BROWN J BLACK

Fic. 8.—Ulua Polychrome, Bold Geometric tripod dish, excavation 2, Santa
Rita (farm 17). (Specimen in National Museum of Honduras at Teguci-
galpa.)

intricate, conventionalized serpents (pl. 8, e) in black and dark red on
a lighter red background. Although very involved, such designs are
often very irregular in execution. The style seems rather unique
but is more “ Mayoid”’ than Bold Geometric in feeling.

No incensario fragments came from levels A in excavation I. From
B are nine unpainted incensario fragments, all of the perforated
“frying pan” type. The handles are tubular, except one that is
rather crude and solid. Two handles end in clutching triangular
claws and one has slits down the side. The same number of frag-
ments came from levels C, but half are painted with dull red and
brown stripes or simple geometric polychrome designs. Two frag-
ments came from D, one plain handle has a horizontal slit and another
is painted with red and black. There is some indication here that
painted incensarios may be relatively earlier than unpainted ones.

NO. 1 HONDURAS—STRONG, KIDDER, AND PAUL 55

Candelarios are lacking in levels A. In B two were recovered,
one decorated by an incised bird (compare fig. 7, 7) and one with
incised lines. Both are of the single-hole variety. Levels C yielded
one two-hole candelario decorated with a delicate incised pattern. Six
candelario fragments came from D, one plain two-hole type and five
single-hole specimens. One of the latter was decorated with a nicely
executed textile design unit. There were no fragmentary cassava-
grinders from levels A, but B yielded one, C five, and D seven. One
of the latter is almost restorable. Like the others, it was of coarse gray
pottery, round, with a broken strap handle on the rear and a series

\ “ily el
Lf LTTE FITZY PEER GOOG Gh Uy
aS oe b/ i

fa;

7
MUM

Zi

Yk

Aa

tL

WL
Cy

(7d [f VV. VV A

= TTT i, M,
ar

eS] DULL ORANGE es BLACK DARK RED

Fic. 9 —Unusual Ulua Polychrome, Bold Geometric dish, excavation 2, Santa
Rita (farm 17). (Specimen in National Museum of Honduras at Teguci-

galpa.)

of very rough-edged incisions or graters on the face. Only one plain,
biconical, pottery spindle whorl was recovered. It came from levels D.

Figurines and whistle fragments were, rather strangely, completely
lacking in levels A and D, at excavation 1, though they were fairly
abundant in the two middle horizons, B and C. Excavation 2 yielded
the finest assortment of such modeled pieces, and certain of these, now
in the National Museum of Honduras, are reproduced here from our
field sketches and photographs (fig. 7). All of these came from the
polychrome horizon between pottery levels 8 and 11 in excavation 2,
but, in general, are similar to the fragmentary pieces from levels B
and C in excavation 1. The latter types show no obvious stratigraphic
differences ; fragments of large ornate busts and statues of polished
brown pottery (like pl. 6, a) ; solid, mold-made figurines of Mayoid

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

type (like fig. 7, s) ; hollow faces with beards or ornate head dresses
(like fig. 7, m), bulbous human, animal or composite figures (like
fig. 7, h, r), tubular birds (fig. 7, e), howling dogs, (fig. 7, c), and

<2V

o%< VA.
Re

J x
Sy

G 5 250% X
» . TREE
Up

yp jj

Te

Ml

DARK RED ie] DULL ORANGE Ea BLACK

Fic. 10—Ulua Polychrome, Bold Geometric bowl, excavation 2, Santa Rita
(farm 17). (Specimen in National Museum of Honduras at Tegucigalpa.)

a variety of squatting animals (like fig. 7, a, k, p) all occurred in both
horizons. Some of the smaller human figures were once attached
to whistles, but many are simply figurines, or ornate hollow statutes
whose functions remain conjectural. Only a few exceptional pieces

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 57

show any traces of painted decoration. From C came a fragmentary
animal with a mouthpiece suggesting a spout. An incensario or pot
cover, from P 10-11 in excavation 2, representing a deer similar to
certain figures in the Dresden Codex, is remarkable (pl. 8, c). At
present the distribution of these numerous products of the sculptor’s
art gives little indication of the lines of their development within
the polychrome period on the Ulua. However, comparison with
similar types from earlier horizons and a complete typographical
analysis will be a large and important task.

a

g
fi y

OU

7

yj

y
]

cm.

ee DARK BUFF RED Pea BLACK

Fic. 11—Ulua Polychrome bowl, excavation 2, Santa Rita (farm 17). (Specimen
in National Museum of Honduras at Tegucigalpa.)

Pottery stamps are rare from excavation 1, though they were fairly
numerous in excavation 2. In excavation 1, levels A yielded one
cylindrical, roller stamp with a neat, squatting monkey design; and
one flat, stemmed stamp with a geometric design. Levels B and D
yielded no stamps, but levels C yielded one flat stamp with a con-
ventionalized serpent head design. :

As at Las Flores Bolsa, the disproportion between the abundant
ceramic remains and all other artifact types was enormous in exca-
vation 1, Santa Rita. Levels A produced one large, conical, stone
pestle and one obsidian flake knife; levels B, six pieces of ground-

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

down animal rib-bones and one ground stone knife ; levels C two small,
polished bone needles, two obsidian flake knife fragments and one
ground stone knife ; and levels D yielded, aside from ceramics, nothing
but one small stone celt.

Soil conditions in the northern extension were practically identical
with those in the main portion of excavation 1 (compare fig. 6).
The occurrence of four burials in the northern extension has already
been noted. As in excavation I, the polychrome horizon in the north-

; 7
ELL W
ZTE “fA

ZZ
ZILLI

cm.

DARK BRICK RED wHite fi scack [|_| orance

Fic. 12.—Ulua Polychrome bowl, excavation 2, Santa Rita (farm 17). (Specimen
in National Museum of Honduras at Tegucigalpa.)

ern extension corresponded with the dense clay stratum (level 7,
fig. 6) and was marked by a concentration of river boulders in the
upper levels. In the northern extension the polychrome horizon
(and burials) which were included in pottery levels A and B, termi-
nated abruptly just above the sand layer (level 8, fig. 6). The latter
was sterile and averaged 20 centimeters in thickness throughout this
area. As in excavation I (excepting the polychrome dump heap on
the southern border) the polychrome pottery horizon in excavation 2
terminated abruptly on the sterile sand stratum. However, under this

NOTE HONDURAS—STRONG, KIDDER, AND PAUL

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KAJ BROWNISH DARK RED ORANGE BuFF [| veLLow BuFF ff Brack

Fic. 13.—Ulua Polychrome, Mayoid vase, excavation 2, Santa Rita (farm 17).

(Specimen in National Museum of Honduras at Tegucigalpa.)

5

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

sand in C (P 7) there occurred a considerable number of potsherds
of a different monochrome or bichrome type. These were in a clay
stratum and were mainly concentrated in P 7, although they occurred
very sparingly in D (down to P 12). Only three sherds came from
P 12. This lowest clay stratum was sandier at the bottom of the
excavation.

Fic. 14.—Inside design from Ulua Polychrome, Lower Mayoid vases (pl. 8 a, b),
excavation I, Santa Rita (farm 17).

Space permits only a very brief analysis of the ceramic sequence
in the northern extension. Dividing the 12 pottery levels (P 1-12) as
in the main excavation, A (P 1-3) contained considerable amounts
of undecorated cooking ware; some fragments of straight-walled
Mayoid vases with black slip, florid designs, and solid rectangular
legs; a few small Mayoid bowl fragments; a few fragments of Bold
Geometric ware; and, finally, several sherds from _ thick-walled,
painted and incised vases (Las Flores type, pl. 5, a-e). B (P 4-6)
contained numerous sherds from excellent, thin Mayoid vases with an
orange slip and well-executed bat and thin-line human designs in

No. I HONDURAS—STRONG, KIDDER, AND PAUL 61

black and red (an excellent example from P 12, just above the sand,
is illustrated in pl. 9, t). In addition, B contained a number of non-
descript polychrome pieces. As in the main excavation, the lowest
polychrome types in the northern extension were the best finished
and had the most realistic and artistic designs. The sand level below
P 12 was barren of artifacts.

Below this sand level (layer 8, fig. 6) potsherds were rather numer-
ous in P 7 and occurred in very small quantities down to P 12 (i. e¢.,
through C and D but concentrated in the upper portion of C). All of
these sherds are monochrome or bichrome and not a single example

Ta aa
me Why

ay

Jelly

C VY :
I Or Oe a) | 7

SF MTT

inning: a nas iSite 38

cm.

RED DARK RED erack =[ | Mepium BROWNISH BUFF

Fic. 15.—Ulua Polychrome bowl, excavation 2, Santa Rita (farm 17).
(Specimen in National Museum of Honduras at Tegucigalpa.)

of either Mayoid or Bold Geometric polychrome occurred. The
sample is too limited to define the type adequately but is undoubtedly
significant as indicating a different and earlier ceramic type, here
designated Ulua Bichrome, at this site. The majority of these lower
sherds are monochrome ranging from highly polished red and orange
ware to more numerous coarse brick red or sooty gray sherds. The
highly polished red or orange sherds show examples of flat, heavily
incised lips (pl. 9, 7, n); swollen lips (pl. 9, f); flanges below the
rim; flat bottoms; and small, solid tripod feet (pl. 9, aa, bb). They
are from small vessels for the most part. The paste and tempering of
these pieces is very fine and the ware is light and hard. A direct rim
from a bowl of this type has a light gray polished interior.

A number of the orange sherds (pl. 9, 0, ~, g, 7, 5, u, v-, aa, bb)
are definitely of Usulatan ware (Lothrop, 1933, p. 50). The faded red

62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

or black linear designs on the bright orange background makes them
very hard to distinguish from examples of negative painting since
the slip at present appears to form the design, in contrast to the darker
red or blackish overlay. Several sherds retain the black color of the
original design, whereas in the others this has faded to a brown or
even a dull reddish color. One very coarse potsherd, apparently
from a flat tripod vessel, has a dull white slip on the inside with
broad, criss-cross red lines (pl. 9, cc). Aside from the Usulatan type
sherds this is the only painted fragment. This is similar to the red-on-
white sherds from the old Playa de los Muertos horizon.

Among the heavier, coarser sherds occur examples of low, flaring,
swollen lips; direct rims ; broad, vertical loop handles, smooth rocker
zigzags (pl. 9, e), and both fine and coarse incised decoration (pl. 9,
a,c). At the present stage of preliminary analysis this coarser pottery
shows no very striking differences from the monochrome or domestic
wares associated with the upper polychrome horizons. The polished
orange ware and especially the Usulatan or related painted pieces are
unique so far as this site is concerned. Aside from pottery the only
other artifacts from these levels are a few fragmentary obsidian flakes
(pl. 9, k, m) and a heavy, stemmed, pottery stamp with a geometric
design (pl. 9, 7). The stamp comes from P 8. The nature of the
deposit below the sand level in the northern extension rather suggests
the fringe of a midden whose concentration lay still farther to the
north. Unfortunately, it was impossible to follow up this problem
at the time, owing to the rapidly rising water level. These subsand
layer ceramics at Santa Rita suggest definite affiliations with the oldest
horizon at Playa de los Muertos.

PLAYA DE LOS MUERTOS (FARM ITI)

This important site is located on the east bank of the Ulua River
close to the northwest corner of farm 11 (see map, fig. 5). In this
general vicinity Gordon (1898) carried on extensive excavations in
1895 and 1897, and later, in 1929, Mrs. Dorothy Hughes Popenoe
(1934; also see Vaillant, 1934) isolated the old Playa de los Muertos
culture at this exact spot. For this reason we visited the site on
January 18, 1936, the day after establishing our headquarters at
Progreso. First impressions regarding the possibility of further work
were extremely discouraging. The terriffic flood of the preceding fall
had removed most of the point where Mrs. Popenoe worked, as well as
the entire island just below it (see map, Popenoe, 1934, p. 81). A
small hard-pan or dense clay playa remained, on which we found a
few Playa de los Muertos type potsherds. However, we found none

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 63

im situ on the adjacent steep banks, nor did we note any traces of
burials or of polychrome pottery deposits in the vicinity. It was
apparent that, at most, only a tiny remnant of the area worked by
Mrs. Popenoe remained at the small playa previously mentioned.
This opinion, based on comparison with Mrs. Popenoe’s map, was
verified by Mr. Roberts, overseer at farm 11, who had assisted Mrs.
Popenoe in her work. For this reason we sought other sites, hoping
to encounter elsewhere, the older type of Playa de los Muertos
material in direct relationship to the polychrome horizons.

By the middle of April 1936 it was apparent that we were not
going to find typical old Playa de los Muertos material at any of
our other sites, despite the discovery of polychrome ware super-
imposed on pottery suggesting the Playa de los Muertos culture at
Santa Rita (farm 17). For this reason, while work continued at
Santa Rita, the senior author returned to the Playa de los Muertos
site on April 17.

Subsequent to our first visit to the site, a large levee had been
constructed along the river just east of the main site, and on the levee,
and in the deep borrow pit or trench, we found numerous fragments
of polychrome pottery. This material was concentrated at one place
on the west wall of the borrow trench and here we later excavated
(excavation 2). This site was only 80 meters southeast of the playa
(with the old type sherds on its surface) which marked the eastern
boundary of the grave area worked by Mrs. Popenoe. At this latter
point close examination of the 4-meter bank behind the playa revealed
a few sherds of coarse brown cooking ware and one tiny polychrome
fragment in situ. Here excavation I was commenced.

Excavation 1, which was made on the very top of a point projecting
out onto the clay playa, was L-shaped. The main cut was 2 meters
wide and 6 meters from west to east. To facilitate handling the dirt
from the deep cut, a north to south extension 4 meters long by 14
meters wide and slightly more than 2 meters deep was made from
the east end of the main cut south to a steep bank on that side.
The north wall of excavation 1 is illustrated (fig. 16; also see Strong,
1937, fig. 79), and the position of the shallow north to south L
extension is indicated by the shelf under skeleton 1. The main
east to west trench attained a maximum depth of 6 meters. The soil
layers from top to bottom are well indicated in the diagram (fig. 16).
Potsherds were first encountered at a depth of 80 centimeters at the
west end and 1.30 meters at the east end. Scattered sherds extended
through this layer of gray clay (P 1, fig. 16) to a depth of 1.8 meters,
where the sterile yellow clay began. The majority of these sherds

VOL. 97

SMITHSONIAN MISCELLANEOUS COLLECTIONS

64

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NO. I HONDURAS—STRONG, KIDDER, AND PAUL 65

were from monochrome red to gray cooking ware, but enough poly-
chrome sherds were found to establish the horizon as definitely
belonging to the polychrome period. Sherds were too scarce, however,
to make 30-centimeter levels of value, so that the entire stratum
was designated P 1. Two extended skeletons occurred at the bottom
of this horizon (fig. 16). Each was accompanied by a broken mono-
chrome. cooking pot, but definite polychrome sherds were also found
next to each skeleton. Skeleton 1 had, in addition, an obsidian flake
knife and a perforated pottery labret or ear plug. Below these
skeletons we ran into a layer of yellow clay which was absolutely
sterile. At a depth of 3.35 meters more sherds of a different type
(Playa de los Muertos culture) were encountered coincident with
our passing from the yellow clay into a hard brown clay. Owing to
their abundance, it was now possible to work by 30-centimeter levels,
thus P 2 (pottery level 2) began at this point. The Playa de los
Muertos horizon (P 2-10, fig. 16) sloped down from west to east.
As indicated in the cross-section (fig. 16) at least one and possibly
two definite occupation or house floor levels, marked by black soil,
concentrated charcoal, animal bones, sherds, etc., and a small deposit
of mussel shells, were encountered. Owing to the depth of the deposit
and to lack of time, it was impossible to work out these living levels
beyond the walls of the excavation. No post holes were encountered,
but baked clay with wattle and daub impressions was fairly abundant.
In the west end sterile soil underlay P 6, but in the east end the
occupation strata dipped to the top of P 10, terminating just above
the then level of the river (fig. 16).

It is obvious, both from the direct superimposition of the two
ceramic horizons separated by a barren stratum (fig. 16) and from
the markedly different ceramic content of each, that two distinct cul-
tures are represented at this site. Of these, the lower or Playa de los
Muertos horizon is the older. Since this horizon extended well to the
west prior to the recent flood and since the main concentration of the
upper or polychrome horizon occurred on a similar level 80 meters
to the southeast (excavation 2), it would appear probable that only the
edges of the two occupation levels overlap at excavation 1 (fig. 16).
For the purpose of obtaining direct stratification, we were therefore
extremely fortunate in choosing the place for our main trench.

Excavation 2, in the west wall of the levee borrow-pit, was small
but yielded a considerable amount of polychrome pottery. The
excavation was 5 meters from north to south and slightly less from
east to west. The first potsherd was encountered at a depth of 70
centimeters and the lowest at 2.40 meters. No noticeable changes

66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

in polychrome pottery types were observable in this deposit, and 30-
centimeter levels were not recorded. Abundant polychrome sherds
were scattered throughout a gray to brown clay stratum. The pottery
level contained concentrations of ash, charcoal, and sherds, one
lenticular hearth, numerous small boulders, and abundant sherds.
Below the pottery level an absolutely sterile, brown sandy clay was
encountered. The maximum depth of this excavation was 3 meters.
In absolute level the polychrome horizon at excavation 2 compared
closely with the upper or polychrome horizon (P 1, fig. 16) at
excavation I.

Excavation 3 was made on the northern side of the playa in the
same dense brown clay level where old Playa de los Muertos material
occurred in the main trench. At excavation 3, this level was on the
surface, owing to the removal of the top soil by the river. An excava-
tion 6 meters long (from northeast to southwest) and 1.5 meters
wide was carried down to a depth of about 1 meter. No sherds or
other artifacts were encountered below the surface and, as it was
apparent that we were outside the area of ancient occupation, work
was stopped.

Before describing the artifact content of the various levels at exca-
vation I, it will be well to discuss briefly the material from excavation
2. All the ceramics (other artifacts were extremely rare) from excava-
tion 2 correspond with those from the A (P 1), the upper or poly-
chrome level at excavation 1. These two horizons are actually on the
same level, and since material was scarce in A (excavation 1) and
abundant in excavation 2, the latter must be considered in order to de-
fine the polychrome wares characteristic of the upper horizon. Owing
to the apparent uniformity of all wares exposed in the cut bank at ex-
cavation 2, it was considered as one unit. To check this, however,
material from the very bottom portions was segregated for comparison
with the remainder. This will be discussed after the bulk of the
material has been analyzed.

The domestic ware from excavation 2 is predominantly mono-
chrome, of a dull red color. A much smaller number of sherds have
traces of crude linear designs in brown, dark red, or black. The bulk
of the domestic sherds appear to be from medium large vessels which
were fairly well polished, with openings varying from heavy direct
lips to slightly flaring rims. Vertical, solid, loop, and strap handles
occur frequently. There are two dimpled bases and one partially hol-
lowed, conical foot (from a unique vessel form). Six monochrome
sherds are decorated with well-executed but simple incised geometric
designs.

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 67

Nine rim sherds from finer vessels that were both painted and in-
cised are of the thick Las Flores vertical-walled vase type (compare
pl. 5,a-e). These have a polished slip ranging from dark red to orange,
a band of black geometric designs under the lips and below this another
band of incised design. As indicated earlier, this Las Flores type of
incised and painted ware is also represented on the Bay Islands
(Strong, 1935, pl. 18, b, c, e). The sherds of this type from excava-
tion 2 (farm 11) also have inner and outer design elements that rather
definitely suggest Bay Island Polychrome I pottery. One other sherd
with more delicate painted and incised designs (similar to pl. 5, 1)
indicates the same fusion between the Las Flores painted and incised
vase style and the Mayoid painted style that occurred at Las Flores.
At excavation 2 (farm II), as at Las Flores, the Mayoid polychrome
type of vertical vase is the more numerous. Sherds from these vases
are very similar to those from Las Flores (compare pl. 5, f-m). They
are relatively thick (compared to the vases from the lower levels at
Santa Rita) with elaborate but conventionalized over-all designs in
red and black on yellow buff. Geometric motifs such as crossed circles
are also common. One flat bottom, one low, round, solid, tripod leg,
and one thin, solid, rectangular, tripod leg occur. Two elaborately
sculptured sherds have a curvilinear Mayoid design. One vestigial
spout (identical with pl. 6, b from Las Flores) is from a painted and
incised vessel.

Smaller bowls with black and red designs on light red or orange
are even more common than the Mayoid vase type. Some of these
have conventionalized ‘“ Mayoid” figures but more have geometric
designs such as lines and circles. They are small to medium in size
including direct bowls, small pots, and small vases. One vertical strap
handle, one flat bottom and numerous rounded bottoms occur. In style
these vessels represent a blending between the Mayoid and the Bold
Geometric with the latter style predominant. Tripod plates and dishes
are lacking here as was true at Las Flores (excavation 2).

Bold Geometric ware is fairly common and the large swollen vessel
with broad strap handles occurs (like pl. 7, a). The monkey lug,
however, is absent at this site. The slip of these pieces is a very dark
polished red or orange with geometric designs in black. Animal design
forms are lacking. One sherd of this type has a geometric design in
white paint. Two typical deep dimple bottoms occur. No figurines,
stamps, or whistles were found, but there is a brown pottery foot
from a rather large hollow effigy. Two fragmentary prismatic flakes
of obsidian were the only other artifacts.

68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

The material segregated from the lowest level in excavation 2 con-
tains fragments of all these types and establishes the uniformity of
the deposit. The domestic ware is identical, numerous pieces having
blotchy dull red or brown designs. One well-polished sherd has a
flange outside the neck with a dull red criss-cross design extending
from flange to body. One sherd represents the Las Flores type painted
and incised vertical vase. There are several small bowl fragments with
conventionalized Mayoid and geometric designs, and one typical Bold
Geometric swollen bowl fragment. A sample gathered from the sur-
face of the borrow-pit is similar but contains several “ Mayoid ” verti-
cal vase fragments rather suggesting the Bay Island Polychrome I
type (Strong, 1935, pl. 21 and fig. 21). A portion of a very small
tripod vase with red slip and black line decoration has an outer wall
panel with excellently sculptured Mayoid faces in profile. In general,
all the material from excavation 2, Playa de los Muertos, agrees very
closely with that from excavation 2, Las Flores, and with pottery
levels A and B in excavation 1, Santa Rita.

Returning to excavation 1 at Playa de los Muertos, we will first |
consider the material from P 1, the upper or polychrome horizon
(fig. 16). The fairly abundant domestic ware is identical with that
just described at excavation 2. Two restorable vessels of this type
accompanied the two burials in the lower portion of P 1 (fig. 16).
That with skeleton I is a round-bottomed pot with a low flaring rim
and two vertical round handles. It is of coarse brown, unslipped ware
with triangular incised designs over the lower neck and upper body.
The vessel with skeleton 2 is a polished black vessel with a direct
rim and three small solid legs. In direct association with the coarser
ware throughout P 1, polychrome sherds occurred. The majority of
these are small and some of them are eroded, but their type is definite.
The majority come from small bowls with a red or orange slip. The
lips of these sherds are painted red or black and similar linear designs
occur on the body of the vessels. Ten small sherds are colored buff
to orange and have remnants of complex red and black designs. Two
orange sherds with red lines and large dots suggest the Bold Geometric
ware. Two polychrome sherds are from flat bottoms, and one is a
rounded flat bottom. One large, hollow, cylindrical leg with an orange
slip and red and black designs is from a tripod dish. The leg has a
vertical perforation in the lower portion and holes in the part joining
the body. It originally contained a rattle. This type of vessel (compare
pl. 7, e, f) was lacking in excavation 2. The only other artifact en-
countered was a fragmentary prismatic flake of obsidian. Although

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 69

the polychrome sherd sample from excavation 1 is small, it is very
similar to the material from excavation 2.

Playa de los Muertos culture material is abundant throughout an
average of 2 meters in the lower portion of excavation I (P 2-9
inclusive, fig. 16). Broken pottery comprises the bulk of the collec-
tion, since no complete vessels were recovered by us and other artifact
types were rare. This discovery of undisturbed refuse deposits entirely
pertaining to the old Playa de los Muertos culture is exceptionally
important. Not only does it give a representative and unselected
sample of the culture but it also permits the inclusion of burial ma-
terials obtained by Gordon and Popenoe as definitely pertaining to
the older horizon. Although Gordon in his brief published report
gives no data on relative depths and states that no observable strati-
fication occured (1898, p. 38), it is undoubtedly significant, that the
majority of complete vessels he illustrates (1898, pl. 7, a, }, ¢, d,
e, h, k, n, 0, p, g, 7, S, w) are characteristic of the older Playa de los
Muertos culture. Furthermore, examination of his letters from the
field and the occasional depths he recorded in cataloging, now on file
in the Peabody Museum, indicates that all these complete vessels came
from the lowest portions of his Playa de los Muertos (Largartijo)
excavations. These undoubtedly were from burials of the old Playa
de los Muertos period. The old burials excavated by Mrs. Popenoe
are fully documented (Popenoe, 1934, pp. 65-79). All are from below
4 meters in depth and contain only Playa de los Muertos materials.
Since we found no entire vessels of the Playa de los Muertos culture,
we have included outline sketches of vessels obtained from graves
by Mrs. Popenoe (figs. 17, 18). Thus, each ware or ceramic subtype
of this culture, established on the basis of our potsherd collection,
can be illustrated in its complete form by a vessel from Mrs. Popenoe’s
collection pertaining to the same Playa de los Muertos type or subtype
(also compare Mrs. Popenoe’s illustrations, 1934, figs. 2, 6, 8, II,
12, and 15). The final description of the Playa de los Muertos cul-
tural horizon must include a complete study of the abundant com-
parable Gordon and Popenoe materials, but this is not attempted here.

For present purposes we have grouped our 30-centimeter strati-
graphic levels of Playa de los Muertos culture material (fig. 16, P 2-9
inclusive) into two uneven divisions, an upper (P 2-4 inclusive),
and a lower (P 5-9 inclusive). The lower grouping of levels, which
we may call levels C (P 5-9), yielded almost twice as much material
as did the upper levels, here designated as B (P 2-4), owing to the
fact that level P 5 was unusually rich and overweighted whichever
half it was placed with. This discrepancy can be avoided later when

7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

finer analyses are attempted, but for present purposes this segregation
into a smaller upper and later grouping of Playa de los Muertos cul-
tural materials (B); and a larger, lower, and earlier grouping (C)

F / 7m
Fic. 17.—Outlines of vessels of the Playa de los Muertos culture obtained by
Dorothy H. Popenoe. Not to scale. a, burial 2; b, c, burial 4; d-l, burial 5;
m, burial 7.

must suffice. Even such an arbitrary division suggests certain ceramic
trends within the period that may well be significant.

The ceramic materials from B and C fall into six main wares or
ceramic subtypes based on surface finish or decorations (pls. 10, 11,

NO] I HONDURAS—STRONG, KIDDER, AND PAUL Ji

177

Fic. 18.—Outlines of vessels of the Playa de los Muertos culture obtained by
Dorothy H. Popenoe. Not to scale. a, burial 7-03 6s Gd; €, }, 4; butial 85) g, 4;
k, |, m, burial 11.

72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

and figs. 17, 18). Each of these subtypes share common features of
form and decoration uniting them into a very definite major ceramic
type characteristic of the Playa de los Muertos culture. The most
abundant sherds from both B and C may be described as (1) un-
slipped, rough, brick-red to sooty gray ware. These are often from
large vessels with slightly flaring rims and necks of variable height,
or from smaller vessels with low necks and swollen lips. Broad,
vertical, strap handles are common in both B and C, but the great
majority of handles in C have two or three vertical ridges and corre-
sponding depressions down the outside. Two round, solid, vertical
handles of large size from B and one from C have conical tenons on
the ends for attachment to the body of the vessel. Of the nine basal
fragments of this ware in B, eight are flat and one dimpled ; in C three
of the four basal sherds are flat and are slightly rounded. One spout
of this ware occurs. Decoration is rare, several sherds have incised
lines forming criss-cross designs, and three sherds have a raised ridge
with regular indentations about the greatest diameter of the vessel.
Subtype I seems very similar to the plain or domestic wares character-
istic of the upper or polychrome horizons at Playa de los Muertos, but
the prevalence of flat bottoms seems rather distinctive. Owing to lack
of space, this subtype is not illustrated here except for an outline sketch
(fig. 17,6).

Subtype 2, (pl. 10, a-h), slipped and polished orange-red to brown
ware, 1s almost as abundant as subtype 1 in B, but only about one-
fourth as abundant in C. Shapes in subtype 2 are very similar to
those in subtype 1, but the vessels were somewhat smaller (fig. 17,
g,h, 1, j, and fig. 18, e, g, i,j,k). One spout from this sub-type is from
B and 2 from C. A rather heavy basinlike bowl or vase (fig. 17, k, l,
and fig. 18, c) is rather common, as are direct bowls (fig. 18, g, /).
Handles seem rare but a few vertical strap handles occur. Fluted
sherds (pl. 10, f) are fairly abundant in B but do not occur in our
sample of this ware from C. Fluted fragments are usually from the
body portion of rather small rounded or swollen bowls (like fig. 18, m,
in shape). The fluting varies in width and is either vertical or diagonal.
Incised lines often set off the fluted portions. Incised and modeled
sherds of this type occur in B, but only incising in C. Several frag-
ments from B have intricate and well-executed geometric and curvi-
linear incised designs (pls. 10, 11). In C basinlike bowls with heavy
incised designs are represented. In B a hand in high relief is the best
modeled piece. Broad, flattened lips with deeply incised decoration (pl.
10, h, 7 represents the type) are very common in B and fairly com-
mon in C. Usually the entire rim is flattened and decorated, the rim ex-

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 73

tending farther out on two sides, forming a handle or a definite tab
(pl. 10, h, +).

Subtype 3 is a dark gray to black, highly polished ware (pl. Io,
i-n; figs. 17, m; 18, m). This is a very distinctive slipped ware with
such a high polish and so much fire clouding that certain pieces have an
almost purplish color. The forms are very similar to those in subtypes
2 and 4, and fluted sherds and flat, heavily incised rims (pl. 10, f, h, 1)
are common to all. Fragments from basinlike bowls are common
(pl. 10, 7, m) and the incisions on such pieces are sometimes so deep
as to suggest a series of outer flanges. There are no handles of this
ware, but one small, solid, cylindrical foot occurred in B. This is the
only foot noted in the entire Playa de los Muertos culture horizon.
Material of subtypes 2 and 3 are about equal in amount but subtype 3
is more abundant in C than in B. This suggests that subtype 3 is
generally earlier than subtype 2.

Subtype 4 is a slate-gray to. buff, highly polished ware (pl. 10, o-s;
fig. 17, d, e, f, k; fig. 18, a, b, f). In amount this ware is about equal
to the two preceding subtypes and is slightly more abundant in C
than in B. The majority of pieces appear to have had a light-colored
slip and a subsequent high polishing that gives them almost a glazed
appearance. The paste is exceptionally fine, and the pottery very hard.
Irregular dark firing clouds are very common (pl. 10, 7). In general
the shapes are similar to those already discussed, but small bowls
with low, slightly flaring and swollen lips are common (pl. 10, 0, q, 5).
Several sherds have ridges, tabs, and-human features in relief, and a
number of spouts of this ware occur in C. An unusual flaring, trumpet-
like neck has been figured elsewhere (Strong, 1937, fig. 76, upper
left). A fragment similar to this was found in the older deposits at
Lake Yojoa by Mr. Rittenhouse and erroneously restored as a trumpet.
Flat bottoms are common, but a few rounded bottoms occur. Handles
are rare.

Subtype 5 may be designated as a ware with a chalky white wash
(pl. 11, a-e). It is relatively rare in both levels but somewhat more
abundant in C than in B. It should be noted that the majority of the
figurines from B are of this ware (pl. 11, g, r). The majority of
sherds come from heavy, direct bowls or from pots with low necks
and slightly flaring lips. One extremely broad, vertical strap handle
occurs, as well as two large spouts (pl. 11, a). The figurines and a
few sherds of this type with painted designs will be discussed later.

The sixth ceramic subtype from this horizon is comprised of various
painted wares. Painted pottery is relatively very rare in the Playa
de los Muertos culture horizon, yet forms an important and varied

74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

type. It was about equally divided between B and C, perhaps indicating
that it was more abundant in later times since the smaller upper section
yielded an equal amount of painted sherds. Red and black painted
sherds constitute one type (pl. 11, f, g, k). Some of those sherds have
alternate areas of black and red sometimes separated by incised lines
(pl. 11, g). In other cases these red and black areas are very irregular,
and the colors form irregular blotches rather than controlled designs.
Numerous flattened and incised rims of the very dark subtype 3 have
flecks or small areas of red paint (pl. 11, f), others have red on black,
or black on red, painted areas. The under side of such flattened
rims are usually black. Several fluted fragments have black and red
painted areas. A few small vertical handles and one very small
horizontal handle (pl. 11, g, k) occur in this red and black ware.
The red and black painted sherds are more numerous in B than in C.
Red on buff sherds are the next most abundant type (pl. 11, 1, 7, 0).
The majority of red on buff sherds come from C, but the type is rep-
resented in B. Most of these sherds come from small vessels with
a red band on the inside and outside of the rim (pl. 11, 7). The
lower portions and the bottoms of these vessels were often red, and
the remainder, except for red rims, was buff. Several have incisions
in the red area showing the underlying buff. One rim sherd from a
direct bowl, polished red on the outside, has faint red linear designs
on the inside with the lighter buff showing through them like negative
painting. A very well-modeled and painted red and buff lug comes
from B (pl. 11, 0). Several large sherds of coarse unslipped brown
or buff ware have red bands on lip, neck, or rim (pl. 11, 4). Two
sherds in B and four sherds in C have a dull white slip with red
lines or bands on the outside and in one case on the inside (pl. 11, m, /).
They are from large vessels with low flaring lips and include one
broad, vertical strap handle. Three sherds from B and one sherd
from C have irregular white designs on a red painted background.
These are from large, coarse vessels. In two cases the lip has a band
of white inside and out ; in one there are broad, irregular vertical lines
extending down the rim, and in another there are blotchy white de-
signs on the inner surface.

One polished, dull red rim sherd (pl. 11, 2) has vertical lines of
dull gray paint extending down the neck. This suggests negative
painting, owing to the fading of the gray paint. It has already been
stated that several of the other painted sherds with faded linear
designs imitate negative painting. In passing it may be said that
although Usulatan ware is not present in our Playa de los Muertos
culture ceramic sample, it does occur at the site (Vaillant, 1934, p. 90).

NOs HONDURAS—STRONG, KIDDER, AND PAUL 75

Gordon obtained an excellent complete vessel of this ware at a depth
of “26 feet’? (Peabody Museum C 1054), and we found sherds of
Usulatan ware in the lowest horizon at Santa Rita (farm 17). It is
probably one of the components of this early ceramic complex. In
concluding this brief description, it is interesting to note that whereas
the polished and incised ware of the Playa de los Muertos culture is
of an advanced and mature type, the very small percentage of painted
wares is highly variable and suggests an early, experimental interest
in this technique.

Of the remaining artifact types, the figurine fragments are perhaps
the most interesting, From B come four fragments, all from hollow
figurines and all but one with a polished white slip (pl. 11,v,7). Three
of these, all with a polished white slip, represent a bulbous, seated
type that is much conventionalized (pl. 11, v, r). The fourth is an
unusually well-modeled face of polished brown ware (pl. 11, p).
From C come Io figurine fragments, all of which are solid and all
but one without any slip. Three represent female torsos (pl. 11, t,
u, v). Although very simple, they have a primitive naturalism that is
rather pleasing. Six fragments of solid figurines are more frag-
mentary but suggest similar types. Gordon (1808, pl. 10, d, f, g)
shows complete examples of this solid, naturalistic type. The last
fragment is also solid but has a dull, polished white slip like those from
the upper level (pl. 11, s). Our sample is too small for certainty, but
there is at least a hint that the hollow, slipped figurines were later,
being preceded by the solid, naturalistic, hand-modeled type.

Artifacts of materials other than clay were very rare. B yielded
one small jade bead, four fragmentary prismatic flakes of obsidian,
two retouched pieces of obsidian, two polishing stones (one stained
red), one piece of pink chalk (?), and two irregular flakes of hard
stone. C yielded one prismatic flake of obsidian, one small rectangular
muller, and several battered hammer-stones. As is true of the later
horizons, the proportion of perishable to nonperishable artifacts, other
than pottery was very low in the Playa de los Muertos culture. It is
interesting to note that animal bones were unusually abundant in this
horizon, suggesting a considerable dependence on hunting. Over a
dozen fragments of baked clay retain the impress of wattle and daub
house construction. When the present midden material is considered
in relation to the much more elaborate grave materials obtained by
Mrs. Popenoe (1934) and by Gordon (1898), a reasonably complete
record of the period is available.

6

76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

OTHER SITES

In addition to the sites intensively worked, numerous other mounds
or refuse deposits were also examined. The majority of these are
shown on the map (fig. 5). Mound groups are abundant on both sides
of the Ulua River but to date have yielded relatively little material.
Aside from superficial examination and the gathering of small sample
sherd collections, we did no work at such sites. In general, the mounds
on the present valley floor appear to be relatively recent and yield
ceramics that are inferior to those of the earlier polychrome periods.
However, until adequate work has been accomplished at such sites,
the linkage between definitely historical sites, such as Naco, and the
deeply buried, earlier polychrome periods will be obscure. Our failure
to excavate mounds was due to lack of time, not of interest. Small
excavations were made at two polychrome sites, one below Naranjo
Chino and the other on farm 15 (see map, fig. 5). These yielded
splendid polychrome sherds apparently contemporaneous with the
lower levels at Santa Rita (farm 17). On farm I0 we excavated
a shallow polychrome deposit containing pottery identical with that
from the late polychrome horizon at the nearby Playa de los Muertos
site (see map, fig. 5). It is probable that earlier polychrome deposits
occur here also as indicated by the excellent, realistic Mayoid vase
(fig. 19) which is reported as having come from farm 10. We have
illustrated this specimen, which is in a private collection near Trujillo,
because it is a splendid example of the best Mayoid tradition in early
Ulua-Yojoa polychrome wares (compare pls. 1 and 8, a, b, also fig.
30). It is also unique in showing ceremonial drinking among the
Maya.

Nortu Enp or LAKE YOjoA

On February 22 and 23, 1936, Mrs. Strong, Dr. Wilson Popenoe,
and the senior author stopped over at Jaral and visited sites east and
west of that town where commercial excavations had been carried on
(see map, fig. 20). At Aguacate we obtained a considerable amount
of broken but restorable pottery that had been discarded by these
earlier diggers. On March 9, Mr. Paul, Mrs. Strong, and the senior
author returned to Jaral and remained there until April 6, carrying on
excavations at various polychrome sites in the hope ef obtaining at
least a partial scientific record prior to their entire destruction by pot
hunters. Our work was interrupted by “ Holy Week” during which
period neither work nor travel was practicable. Later, on May 26,
Mr. Paul returned for a week in order to carry on deeper excavations,

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 77

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Fic. 19.—Polychrome vase of Ulua Lower Mayoid type, said to have been
found downstream from Playa de los Muertos on the Ulua River bank. (From a
private collection near Trujillo.)

VOL. 97

SMITHSONIAN MISCELLANEOUS COLLECTIONS

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NO. I HONDURAS—STRONG, KIDDER, AND PAUL 79

seeking an earlier type of culture at the Los Naranjos site. In this
venture he was successful despite the very limited time available.

Only within the last 4 or 5 years has the occurrence of prolific
mounds around the north end of Lake Yojoa become a matter of sci-
entific knowledge. The first of these were dug up by local farmers
and, later with the permission of the Honduras government, J. B.
Edwards carried on extensive excavations in the region. In 1934 Mrs.
Doris Zemurray Stone ™ visited Los Naranjos and published a brief
report.

In 1935 Frans Blom, Dr. Jens Yde, and Prentiss Andrews spent 4
days around Jaral in the course of the Tulane University-Danish Na-
tional Museum Expedition.” They explored around the various sites,
worked with Mr. Edwards, and from him obtained collections of the
polychrome ware.

On our second visit we rented a dilapidated house in Jaral and
boarded at the “‘ Grand Hotel Rats’ Nest ’’, as it was fittingly christened
by Yde and his companions. Our genial host, “ El Chino Alejandro ”,
however, made us as comfortable as his limited resources permitted.
In Jaral we were also greatly aided by Capt. Evalyn Bush, and in the
field by our two main workers, Paco Ramirez, of Dos Caminos, and
Miguel Hernandez, of El Eden. Information furnished by Mr. J. B.
Edwards proved very useful throughout our work.

The major geographic characteristics of Lake Yojoa have already
been touched upon. Outstanding archeological features of the plain
at the northern end are, first of all, the great mound group and frag-
mentary stone statues at Los Naranjos (great mound group, map,
fig. 20) ; next the long earth mound or causeway, with its parallel
ditch, just east of the road to Jaral (map, fig. 20) ; and finally the
series of “ancient cemeteries”, or low house mounds containing
burials, about 3 miles east of Jaral near the lake shore (map, fig. 20).
In the following account we will attempt to present very briefly the
major characteristics of certain of these features, each of which merits
at least a full season’s work and a complete report. Our primary aim
was to determine the nature and association of the major ceramic
wares present at such sites and, if possible, to determine whether strati-
fication of culture horizons might be present. The apparent richness

** 1934, pp. 123-128. Mrs. Stone, p. 128, mentions the occurrence of gold at
this site. To the best of our knowledge, no metals of any kind have ever been
found there.

* See Yde, 1935, and 1036. The earlier report (fig. 5) shows four typical
Yojoa vessels; the upper has the “dancing figures’? and is Mayoid in type.
The three lower vessels are in the local Bold Animalistic style.

8o SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

and complexity of Lake Yojoa ceramics makes any brief description
extremely difficult. However, since the majority of Yojoa collections
in various museums are highly selective, even a preliminary account
of the manner in which the various types of vessels and artifacts occur
in situ, should have value. Since complete or restorable pottery ves-
sels are more abundant in Lake Yojoa sites than in those previously
described, we will discuss them in this preliminary report, leaving
detailed potsherd analysis for a later time.

AGUACATE AND AGUATAL

Modern place names around the north end of the lake are usually
derived from certain trees that mark favorable areas for milpa farm-
ing. Aguacate and Aguatal (map, fig. 20) are so named, and it was
here that the first finds of Yojoa polychrome pottery were made. Both
sites have been sadly looted, and though we visited them on our first
trip to the lake, we were unable to find any mounds or promising areas
sufficiently undisturbed to merit scientific excavation. Probably a
very large proportion of Yojoa polychrome vessels in collections inside
and outside Honduras, have come from these sites which appear to
have been exceptionally prolific. We reached them after a long walk
along the trail to Dos Caminos (map, fig. 20), then cutting south
through the generally low but very dense bush. Each site consists
of a large (Aguacate is the larger) irregular area covered with low
mounds ranging from barely visible eminences to some 2 meters high.
Originally, the mounds were covered with rocks, many of large size,
but at the time of our visit both areas were entirely covered with
shallow, irregular burrowings and piles of rich black dirt and stones
which obscured almost all natural contours. The dense bush added
to the difficulty. The excavations ranged from 30 centimeters to about
2.5 meters in depth and, starting in what originally seem to have
been mounds, run labyrinthian courses wherever the machete-wielding
“huaqueros”’ believed they were in mixed soil. Potsherds were
abundant, polychrome pieces seeming to predominate over plain or
cooking ware fragments, and a number of splendid and only slightly
broken vessels lay about, indicating that they had been carelessly
excavated and then abandoned in favor of harder and more resistant
complete vessels. Numerous three-legged metate fragments of various
sizes, roller pestles, rectangular mullers, and two large polished celts
were noted. There were no human bones in sight, but our guide said
that small fragments were occasionally encountered in association with
complete vessels. Among the great number of rough, volcanic rocks

No. I HONDURAS—STRONG, KIDDER, AND PAUL 81

that once formed these mounds we noted a few that appeared to be
artificially squared or smoothed, and in one or two cases the disturbed
rocks appeared to have once formed part of some simple artificial
structure.

Although the occurrence of some domestic pottery and broken arti-
facts, such as metates, suggests human habitation at these sites, the
predominance of elaborately painted sherds and the reported occur-
rence of very numerous deposits of complete polychrome vessels sug-
gests a burial ground wherein the human bones had vanished owing
to the damp, very humous soil. In the light of our later excavations at
La Ceiba and Los Naranjos, it seems probable that both habitations
and burials formerly occurred here, with the latter coming to be pre-
dominant before the sites were finally abandoned. Except that Agua-
cate covers a larger area than Aguatal, and that the rather closely
adjacent sites have been given different names by modern farmers,
the two seem to be identical in types of pottery represented, in the
nature of the mounds, and in the complete manner in which they have
both been looted.

An analysis of all the vessels from, or reported to be from, these
two sites would probably run the complex gamut of Lake Yojoa
polychrome ware. Sherds of almost every Yojoa polychrome type
were actually present in the old excavation pits, confirming the reports
of various of the diggers that the majority of these vessel types occur
in association. Our necessarily brief discussion of ceramics from these
two sites is based, first, on 14 broken but restorable vessels which we
ourselves picked up in the diggings at Aguacate on our first visit. Al-
though these abandoned pieces may not represent the finest types from
the site, they are definite as to site provenience and probably generally
representative. Next, we were able to acquire a number of complete
vessels obtained by local diggers at Aguacate and Aguatal, and in
some cases to sketch and photograph other vessels from these sites
which were not acquired (for example, fig. 30). Complete vessels
thus obtained were sent to the National Museum of Honduras at
Tegucigalpa, and our present illustrations were made from field photo-
graphs and sketches. Some of these latter vessels, reported to be from
Aguacate and Aguatal, but not excavated in our presence, may have
come from La Ceiba. However, we talked directly to the excavator
of each, and there is strong probability that the majority did come
either from the place designated or from one of the adjacent south-
eastern sites (see ‘“‘ ancient cemeteries’, map, fig. 20).

The 14 restored vessels we obtained at Aguacate fall into five major
types in regard to form: first, straight-walled or only slightly flaring

82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

vases or jars, with or without tripod feet ; second, tripod dishes ; third,
open bowls with dimpled bases ; fourth, two-handled pots with dimpled
bases ; and, fifth, bichrome or monochrome pots with direct or slightly
flaring rims and two or four vertical handles. The finest of the first
type is a vertical-walled vase with three slightly hollow, low, cylindri-
cal feet. In form and coloring it is almost identical with one illustrated
(pl. 1). It has the identical step and scroll design pattern on the lip,
but the body design consists of two pairs of interlocked horned or

(ice DULL ORANGE W/Y/ DARK DULL ORANGE

Fig. 21.—Yojoa Polychrome bowl, Bold Animalistic type, Aguacate. (Specimen
in the National Museum of Honduras at Tegucigalpa.)

A DARK RED

plumed serpents with spearlike flames coming from their nostrils
(similar to pl. 13, d). A low, flat-bottomed vase with thickened lips
and paneled walls is too badly eroded to make out the design. A third
piece of this general type has slighty flaring, straight walls and a
dimpled base. Around the neck is a band of skeuomorphic glyphs,
dark red and black on an orange background. On the orange body
of the vessel there are two conventionalized parrots in dark red and
black. Tripod dishes, the second type, are here represented by only
one example (pl. 14, c). This is of medium size with hollow, cylindri-

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 83

cal legs containing rattles. The body color is dark orange with panels
of geometric and conventional designs in red and black.

The third type, bowls with dimpled bases, includes five vessels ;
the finest of these is of thin ware with a light orange background and
elaborate design in dark red and black on the outside. Around the
neck is a series of plumed Mayoid faces conventionally but exquisitely
executed ; these are identical with those on a very similar vessel from
Aguatal (pl. 12, c). The body has complex, human figures in the
same elaborate style, but erosion prevents a clear understanding of
the design. A smaller thicker bowl with a buff background has purple
around the lips and on the body, enclosing buff circles in which are
crude, conventionalized Mayoid faces. Around the neck is a buff
band with black, skeuomorphic glyphs. A heavy bowl has a white slip
with massive, dark red, dull orange and black panels, bands, and
designs. On the sides are two heavy monkeys squatting in profile.
One has a forward-sweeping plume similar to those on the priestly
figures, the other has a backward-sweeping plume and a long tongue.
A thick but well-executed bowl is light orange with dark orange and
black designs. On the rim, these are geometric, but in two circular
areas on the side are ornate birds, evidently the Muscovy duck, with
strange, wrapped objects on their backs. The last open bowl is light
orange with two extremely ornate black and purple birds. It has iso-
lated black stepped scrolls outside the lip (like pl. 1).

There are two vessels of the fourth type, i. e., bowls with two verti-
cal strap handles and dimpled bottoms. One of these is light orange
with a low straight neck and black and red geometric designs. On the
sides these form two highly conventionalized monkeys whose raised
faces with indented eyes project like lugs (compare fig. 22 and pl.
E2aiG) i

The second vessel of this type has a low neck and more swollen
body (like fig. 26). It is orange in color with a band of red and black
geometric designs around the neck, a band of curvilinear designs
around the shoulders, and three ornate concentric diamond designs
down the body (fig. 26 had similar but more elaborate designs).

The three vessels of the fifth type, monochrome or bichrome pots
with direct or slightly flaring rims and two or four vertical handles,
suggest domestic or cooking ware. The largest of these has a round
bottom, low lip, and four solid, vertical, loop handles. It is a dull,
slightly polished red with smoke stains on the bottom. A smaller but
heavier vessel is similar as to handles and base. It is lower, however,
is dark red, and has more polish. The third vessel is dull buff with

84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

vertical bands of dark red (like fig. 27). Unlike the above, it has a
dimpled base and only two vertical loop handles.

A very brief analysis of other vessels reported to be from Aguacate
and Aguatal, probably including some from La Ceiba as well, will
bring out the major types represented here. The majority of the
straight-walled vases from these sites bear Mayoid designs, very
often identical with those on similar vessels from the Ulua or Sal-

V7

CUM

Ss put. BLACK ~=V RED BROWN faa DULL BUFF

Fic. 22——Yojoa Polychrome pot, Bold Animalistic type, Aguatal. (Specimen in
the National Museum of Honduras at Tegucigalpa.)

vador regions (see pls. 12, b; 13, f, and fig. 30). Since these vases
are the ones mostly highly valued by collectors, they are apt to pre-
ponderate in purchased collections, disproportionately to their actual
occurrence in the field. An exceptionally fine vase of this type, said
to come from Aguacate, is illustrated (fig. 30). There are three
design units ; two are seated priests, and the third is a monkey shown
against a black medallion. The two priest designs are similar (fig. 30)
except that the one not illustrated holds a five-branched scepter. This

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 85

vessel was sketched and photographed in a private collection at Jaral.
The tripod dish appears to be rare, but tripod plates, with either high,
hollow legs (like pl. 12, f) or low, hollow feet (pl. 14, c) containing
rattles, are rather common. The majority of these have conventional
designs of the Bold Animalistic type, but Mayoid designs do occur
(pl. 12, f), including skeuomorphic glyphs and “ dancing figures ”
associated with textile designs. Yojoa vessels of this sort appear to
be somewhat more variable in size than are those from the Ulua.
Small, dull brown vessels, with or without low, solid, tripod feet and
decorated in the imitation Ulua marble bowl technique of incising,

[__]vertow sure A stack 77 purptish rep

Fic. 23—Yojoa Polychrome bowl, Bold Animalistic type, Aguacate. (Specimen
in the National Museum of Honduras at Tegucigalpa.)

also occur (pl. 14, e). A few of these vessels, with lugs suggesting
the Ulua marble bowl technique, have sculptured designs that appear
more Mayoid. Particularly noteworthy at Aguacate are a few flat-
bottomed dishes of highly polished dark brown ware having unique
carved designs (pl. 14, f). These conventionalized designs are carved
through the slip and, owing to the light color of the paste, stand out
clearly. This seems to be a ware and decorative technique distinct
from either the imitation Ulua marble vases or the Maya sculptured
vessels. Open bowls vary in size and, as a rule, have two main types
of design: Mayoid (often against a dark background) (pl. 12, ¢, e,

86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Oo 5
oe

cm.

BLACK DESIGN ON DULL ORANGE

Fic. 24.—Yojoa Polychrome tripod dish, Bold Animalistic type, Aguacate.
(Specimen in the National Museum of Honduras at Tegucigalpa.)

[__]vettow sure = sack RED

Fic. 25.—Yojoa Polychrome bowl, Bold Animalistic type, Aguacate. (Specimen
in the National Museum of Honduras at Tegucigalpa.)

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 87

and figs. 28, 29); or a combination of Bold Animalistic and geo-
metric motifs including highly conventionalized birds (pl. 14, a, b;
figs. 21, 24), monkeys, (pl. 13, a, b, c, and figs. 22, 23), alligators
(fig. 25), peccary, and “dancing” jaguars (pl. 12, d). Although
somewhat similar animal motifs occur on true Maya wares, these
Yojoa forms are generally distinctive and are usually associated with
other designs suggesting the Bold Geometric style of the Ulua. Mon-

cm.

Fic. 26.—Outline of Yojoa Polychrome pot showing “ vestigial’’ spout, Aguacate.
(From a private collection at Jaral.)

key designs occur commonly on the two-handled pots with dimpled
bases (pl. 13, a, c, and fig. 22). The range of Lake Yojoa monkey
designs is extremely wide and interesting. The Bold Geometric swollen
vessel with monkey lug handles is not overly common at Aguacate or
other Lake Yojoa sites but does occur (pl. 14, d). Such Yojoa vessels
are smaller than the majority of those from the Ulua and often have
vestigial lugs and less striking red and black designs. These vestigial
handle-lugs are also very common on the dull buff cooking vessels
with dull red stripes (fig. 27). Another two-handled straight-necked

88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

type of pot from Aguacate is decorated with intricate dark purple
designs on orange. One such vessel has a panel of isolated orange heads
on purple around the shoulder and an intricate purple “mask” de-
sign on the lower body and neck. The two lower design areas are
separated by an ornate concentric diamond design. Figure 26 from
Aguacate had a very similar design and shape except for the rather
unusual but significant vestigial spout. Another vessel form, the an-

cm.

[_] out Burr DULL RED

Fic. 27.—Yojoa Polychrome cooking pot, Aguatal. (Specimen in the National
Museum of Honduras at Tegucigalpa.)

nular-based “ salad bowl” type (pl. 14, g), also occurs at Aguacate.
Cooking pots of dull buff color with vertical dull red stripes and two
handles (fig. 27) and four-handled, polished red pots are far more
common at Aguacate than any of the selected collections would indi-
cate, for the looters usually throw these away.

No brief description can do justice to the diversity of Aguacate,
Aguatal, and other Yojoa polychrome forms and decorative elements,
yet it must be remembered that all of these come from the same sites

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 89

and from depths that are rarely as much as 2 meters. Moreover,
despite the occurrence of at least two distinct major styles, the Mayoid
and the local Animalistic or Bold Geometric, the composition and
even the colors of both are similar. There is, moreover, a great over-
lapping of design elements. At Santa Rita (farm 17) the typical
Mayoid and the Bold Geometric polychrome vessels, exclusive of the
numerous intermediate types where they blend, seem more distinct
than do the two major types at Lake Yojoa. Moreover, despite the
great richness of color and design, the bulk of Lake Yojoa polychrome

eS

8.

cm.

RED - ORANGE fee BLACK tes] BUFF

Fic. 28.—Yojoa Polychrome bowl, Mayoid type, Aguacate. (Specimen in the
National Museum of Honduras at Tegucigalpa.)

pottery appears to be technically inferior to either the Mayoid or
the Bold Geometric ware at Santa Rita or other old polychrome sites
on the Ulua. Occasional Yojoa pieces, and these are the ones eagerly
acquired by collectors, have a fine hard paste and fast colors, but
for every one of these, the looters discarded or ruined hundreds of
pieces that were crumbly in texture, with faded or eroded paints.
Had the peoples of early polychrome times on the Ulua had the arche-
ological generosity to bury complete vessels with their dead, as did
their contemporaries on Lake Yojoa, this comparison would be more
obvious than it is at present. Analysis of paste, form, size, color, and

go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

design, and the intercorrelation of these factors in the Aguacate-
Aguatal collections must await future publication, but the foregoing,
very brief, description may give some idea of their richness and the
manner of their occurrence insofar as the looters have not destroyed
all such evidence.

is

ark

cm

[] putt oRANGE MM siack (]] sure

DULL BROWNISH BLACK WY RED

Fic. 29—Yojoa Polychrome bowl, Mayoid (?) type, Aguacate. (Specimen in
the National Museum of Honduras at Tegucigalpa.)

LA CEIBA

The first time we visited La Ceiba we walked in from Jaral by
the trail to Dos Caminos and came back along the lake shore (map,
fig. 20). The latter was an especially hard trip through the black mud
and dense jungle of the lake shore. It was enlivened, however, by the
profusion of orchids, animal tracks, and land and water birds we

NO I HONDURAS—STRONG, KIDDER, AND PAUL g!i

encountered. Later we always rowed down in a cranky old boat,
forcing our way in to shore through the massed water hyacinths
(see fig. 71, Strong, 1937) to land at the old stone steps (see map,
fig. 20). These roughly laid tiers of unworked stone extend several

FETT EL TOS TOT E!

B

D>

Yl Ma“

V7], vickt BROWN DARK BROWN [MMMM crack = [__] Licht vetow Burr

Fic. 30—Yojoa Polychrome vase, Mayoid type, Aguacate.
(From a private collection at Jaral.)

meters from the shore up over the barrier of volcanic rocks that lie
just beyond. They appear to be artificially laid and of native origin.

SITE I

Our first excavations were made in one of a series of low, rock-
covered mounds about 1 kilometer southeast of the stone steps (site I,

7

g2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

map, fig. 20). En route to the site one passes two sharp mounds about
2 meters high that have been deeply pitted by pot hunters. There are
a large number of such mounds in the general vicinity. The mound
selected, mound 1, when cleared, was 10.75 meters long from north
to south, 10.5 meters wide from east to west, and about half a meter
high. It had not been pitted. A trench 1 meter wide and 7 meters
long was run from the eastern edge to just beyond the center of
the mound. The soil at and above ground level was a rich black humus
full of various-sized volcanic rocks. At a depth of slightly less than
I meter below ground level, both stones and artifacts ran out, and
a sterile, yellow clay was encountered. Above this, the black soil was
flecked with the same yellow clay, indicating disturbance. The bulk
of the sherds and charcoal occurred in the black soil below the natural
ground level, the slightly raised mound area consisting mainly of
rocks. No definite house floors, burials, or other structural features
were encountered.

Potsherds were the most numerous artifacts from mound 1. No
complete pots were encountered. The sherds were predominantly
from unslipped brown cooking vessels with very heavy and solid
handles which were vertical or round in cross-section. Coarse buff
ware with dull red stripes and similar handles was also abundant.
Polychrome sherds were few in number, mostly from dull orange
bowls and tripod plates with geometric red and black designs. Legged
metate fragments, fragmentary roller pestles, rectangular manos, half
an ovoid sandstone bowl (1 m deep), and three prismatic flakes of
obsidian were the only other artifacts. Mound 1 was apparently a
habitation site judging by its contents. In all probability there were
burials in the mound somewhere which led to the site having been
covered with rocks, but this is problematical.

Less than a meter south of mound 1 were two small rock piles which
may once have formed one mound, since the space between appeared
to have been pitted long ago. An L trench which sectioned both
portions revealed similar soil conditions to those in mound 1, except
that the mixed earth extended to a greater depth. Artifacts were
more numerous, as were also huge stones that had to be moved with
crowbars. Five whole or restorable pots were found. The first, at
1.10 meters depth, was a tiny two-handled pot with its original red
and black design almost entirely eroded. The second, 1.40 meters deep,
was a larger, swollen-bodied red vessel with two vertical strap handles
having knobs at the bend of each. A black geometric design on the
neck was almost entirely eroded. Both these vessels were lying on
their sides. Nearby, at a depth of 1.30 meters, was a rather high-walled,

NO: I HONDURAS—STRONG, KIDDER, AND PAUL 93

dull orange open bowl with a simple geometric design in red around
the neck. It is chiefly remarkable because it has wavy, vertical lines
down the body that appeared to have been executed in negative paint-
ing. Actually, these seem to have been formed by the disappearance
of the dark red paint that once covered them. A particularly inter-
esting little tripod vase came from a depth of 1.65 meters. Like the
above, it was in an upright position. This vessel has a light orange slip,
two broad black lines inside and below the lip and, on the outer wall,
three prancing jaguars with raised heads and open mouths, vividly
executed in purplish red and black. A black and red geometric panel
separates the three identical figures (the design on pl. 12, d, is similar
but less realistic). Among the many sherds from about the depth of
I meter we found enough fragments to restore an interesting bowl
representing a bird with head, wings, and tail projecting (similar to
but more elaborate than pl. 14, 4). The basic color is light orange,
and the rim, lip, bird head and tail, as well as the median portion,
have dark brown, dark red, and white designs. All but the first of
these vessels were broken when found, and only two were upright.
It is possible that they had been placed with burials, the bones of which
had disappeared, but it seems more probable that they had been dis-
carded with the abundant sherds and other broken artifacts. Two
more partially complete vessels were so soft that they crumbled to
bits when we tried to remove them.

Sherds were more abundant in mound 2 and of better quality.
Cooking ware was abundant and similar to that in mound 1, but there
were also present a large number of fragments from large, straight-
walled vases of Mayoid type. The majority of these had heavy dark
red and black designs on buff or orange, with hollow rectangular or
cylindrical tripod feet. A rim with a painted twilled basketry design
and several with typical Ulua conventionalized heads (on thin, hard,
polished ware) occur. Skeuomorphic glyph bands also occur, and
there is one painted “ bird ” head lug. A number of excellently painted
tripod dish fragments and a number of very large dark red sherds
with typical Ulua Bold Geometric designs are noteworthy. These
include two broad straphandles with monkey face lugs. Two figurine
heads were encountered at a depth of 1.20 meters. One of these is
solid, suggesting a pouting “ baby face ” with hair but no head dress ;
the other is hollow with an elaborate head dress, having a raised St.
Andrew’s cross above the forehead. Both are rather badly eroded.
A few roller pestles, cylindrical manos, double-ended hammerstones,
two crude obsidian side scrapers, one quartzite side scraper, and a
few broken prismatic flakes of obsidian complete the artifact list. The

94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

occurrence of much charcoal and small amounts of animal bone bears
out the evidence of the broken pottery in suggesting that this, too,
was primarily a habitation site. Our workers, accustomed to under-
cutting and burrowing in general, in their zeal, ruined our trench
profiles at mound 2, and we decided to try another site.

Attention may be called in passing to a small collection obtained from
a rock mound similar to and very close to site 1, at La Ceiba. These
objects were dug up by a local pot hunter who heard us working and
visited us. They included an excellent tetrapod dish (pl. 12, f) with
conventionalized Mayoid designs, and feet representing an alligator’s
head and containing rattles. The colors are dark red and black on a
yellow background. A small tripod dish with low, solid feet had a
textile knot design with three pairs of crudely executed “ dancing
figures.” These two vessels are of interest since they have Mayoid
designs on a vessel form usually decorated in the Bold Geometric or
Bold Animalistic style. One large broken whistle of unslipped brown
pottery was unusually interesting since it represented a tusked monster
almost identical to one found on the Ulua at Santa Rita (pl. 13, e, cf.
fig. 7, p). There were also a number of Mayoid figurine and bulbous
animal whistles, including howling dogs, similar to those from the
Ulua. The same mound had also yielded a rectangular and an ovoid
bark beater, excellently made of polished gray stone.

SITE 2

This excavation was on the southern border of the area intensively
dug over 2 years earlier by J. B. Edwards. The rise or mound
selected was less than I meter west of the remains of his headquarters
shack (La Ceiba, site 2, map, fig. 20). From this point north there
are a great number of irregular excavations both in mounds and in
the areas between. There are numerous mounds in this immediate
vicinity, and all of them are badly pitted. According to our men a
very great number of pots came from this general area. The small
rise or mound which we selected for work was not more than 30
centimeters high and had three irregular pot holes on its surface. It
sloped slightly from the volcanic dyke on the west, extended about
18 meters to the east, and was 13 meters from north to south. Its
surface was very irregular, owing to numerous volcanic rocks and to
the old dirt heaps. We completed an east to west cross trench I meter
wide through the center of the mound, but, finding that there were no
regular structural details to be observed in this fashion, we carried
out various extensions to the north and south. In cross-section the
“mound ” showed a top layer of darker soil averaging ten centimeters

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 95

in thickness. Below this was mixed brown earth containing flecks of
yellow clay and innumerable large volcanic boulders. At an average
depth of 80 centimeters sterile yellow clay was encountered with still
more volcanic boulders, many of such great size that they could not
be moved even with crowbars. Aside from various sherds and broken
artifacts throughout the brown earth there were no definite floors
or other evidences of artificial structure except for five groups of
complete pots evidently marking graves. It proved impossible to
penetrate far into the sterile clay owing to the innumerable great
boulders which apparently formed part of the natural volcanic dyke.

Pottery deposit, or grave, I occurred just north of our cross trench
on the edge of the mound. Here, at a depth of from I to 1.25 meters
in the mixed soil just above the yellow clay and under a large number
of great volcanic slabs, we found four pottery vessels (see Strong,
1937, figs. 75, 77). Three were very close together (Strong, 1937,
fig. 77), and the fourth, an incense burner (not shown in the illus-
tration), was 80 centimeters away. Three vessels were intact, but
the fourth and finest (pl. 1) was broken by another bowl which had
been forcibly nested in it. The broken vase, when restored, (pl. 1)
was unusually interesting, since it depicted a processional group of
priests calling to mind the description of Palacio (see p. 12). The first
figure (pl. 1) is the high priest with the ceremonial staff ; behind him is
an assistant. The latter either holds a copal container or has removed
the high priests’ bustle with one hand and is reaching back with the
other for one of the two objects carried by the third priest. These are
probably incensarios, but they could possibly be obsidian mirrors or
some other ceremonial objects. The three priests are followed by two
musicians playing on wind instruments of an unusual type. From the
attitudes of the figures, it would seem that the procession had just come
to a halt prior to the performance of some rite. Further description of
this vase is made unnecessary by the illustrations. The three other
vessels are comparatively simple. The bowl nested in the broken vase
(Strong, 1937, fig. 77 and 75, lower center) has a simple but striking
black design on a cream-white background. Red and black designs
occur on the lip, there is a black band inside the rim, and the under
and inner slip is a dull orange. The small two-handled bowl (Strong,
1937, fig. 75, lower left) is unusually interesting since it is of the
Bold Animalistic type with geometric designs around the neck, a
cursive and conventionalized, twice repeated animal and circle de-
sign on the body, two handles with definite nodes on the bend, and
a deep dimpled bottom. It has a bright orange slip with designs in
black, dark red, and white. Thus, although the processional vase is

96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

definitely Mayoid in form and decoration, this accompanying vessel
is indubitably Bold Animalistic in type. The incensario (Strong, 1937,
fig. 75, upper) is very crudely made of coarse buff pottery with dull
red bands around handle and rim. From their distribution it would
seem that these vessels had been laid around a skeleton, all traces of
which had disappeared.

Pottery deposit, or grave, 2 occurred in our cross trench near the
center of the mound at a depth of only 15 centimeters. It consisted
of two vessels upright and side by side. The larger of these (pl. 13, a)
has a bright buff slip with geometric designs in black, dark red, and
bright red on neck, body, and handles. On the central body it has
an extremely conventionalized monkey face with a miniature body.
It is an unusually conventionalized piece of the local Lake Yojoa
Bold Animalistic style. The second vessel is smaller with swollen
body, slightly flaring neck, two vertical strap handles, and a small,
cross-incised node on each side of its greatest diameter. It is one-
color bright red and, like its companion piece, very fresh in appear-
ance. Both vessels have dimpled bases. The larger pot contained one
small ovoid bead of grayish jadeite or diorite, and the smaller pot a
larger, cylindrical bead of greenish gray jadeite. The latter bead has
a groove around one end and both have complete biconodont perfora-
tions. The shallowness of the deposit may indicate relative recency
and the extremely conventionalized type of Bold Animalistic design
on the larger vessel appears to be late (pl. 13, a). The fact that
each vessel contained a stone bead suggests deposition with the dead,
although here again all trace of human remains had disappeared.

One meter east of deposit 2 in the cross trench, at a depth of 35
centimeters, there occurred two restorable little jars of chocolate-
brown ware in close association with a larger restorable pot (pottery
deposit 3). One of these little straight-walled jars has three low tripod
feet and is decorated with an incised diamond and dot design. The
other is flat-bottomed, has two vertical lugs and a carved or sculptured
design in low relief. The lugs and form strongly suggest the small |
pottery imitations of Ulua marble bowls, but the partially restored
sculptured design seems more Mayoid. The other small jar also sug-
gests the imitation Ulua marble bowl type (similar to pl. 14, e). The
third restorable vessel is a typical, two-handled, local Bold Animalistic
pot with a striking, heavy black and red monkey design (similar to pl.
13, c, and fig. 22). The broken condition of these vessels makes it
uncertain whether or not they represent a grave offering. However
the association of types at this depth is interesting.

NOL HONDURAS—STRONG, KIDDER, AND PAUL . 97

Pottery deposit, or grave, 4 occurred under a mass of great rocks
and consisted of three nested pots at a depth of 72 centimeters. It
was located 2.5 meters east of deposit 3 in the cross trench. The upper
vessel (pl. 13, d) was inverted over an upright, smaller, two-handled
bowl, and also contained a very crude, unslipped, and slightly shoe-
shaped vessel with horizontal, solid, round handles. Inside the latter
was one cylindrical, thin pottery bead. The upper vessel is a striking
open bowl (pl. 13, d) one-half of the surface of which is eroded.
The original slip is dull orange, but the entire outside was covered
with black, against which a thrice repeated dull orange, dark orange,
and red serpent design stands out. The serpent, with bulrushlike
flames darting from its nostrils, is definitely Nicoyan in style. A band
of conventionalized serpent heads circle the outside of the rim and
two black bands the inside. The small two-handled pot is even more
eroded. It has a light orange slip, two conventionalized red and black
alligator designs (similar to fig. 25) and other geometric designs on
the body. The vertical strap handles have definite nodes. The coarse
brown slightly shoe-shaped pot is very badly eroded and lacks all
surface finish. Despite the lack of skeletal remains, this pottery de-
posit has all the earmarks of a funerary offering. It is particularly
interesting since it contained only the local Bold Animalistic type of
pottery in association with a shoe-shaped vessel.

Pottery deposit 5 consisted of several vessels uncovered in the
northwest quadrant of the mound. They occurred over a triangular
area 2 by 3 meters in extent and may or may not have represented
one or more grave offerings. No human remains were found here or
elsewhere at La Ceiba. The first vessel was a small, straight-walled
bowl with a band of red frets against a brighter background around
the neck. The entire middle portion of the outer body is black but
much of the surface is eroded. It was found in an upright position
at a depth of 45 centimeters. The next is a small, swollen pot with
slightly flaring lips and small, solid, rectangular, tripod legs. It is
badly eroded but has traces of black and dull buff circular designs
on a dull orange background. It was found in an upright position
at a depth of 30 centimeters. The third vessel is a large, badly eroded
bowl found upright and wedged in among great rocks at a depth of
‘50 centimeters. Traces remain of an intricate but conventionalized
dark red and light orange design against a black background. Like
many of the Yojoa pots it has two black bands inside, below the lip.
At a depth of 32 centimeters a badly eroded straight-walled vase with
solid, rectangular tripod support was found upright, covered with a
broken bowl. The vase had only traces of black paint on the outer

98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

surface, but the bowl was slightly better preserved. It had an original
orange slip, a band of small red horizontal chevrons outside the lip
and two circular panels surrounded by a black background on the out-
side. The design inside these circles was gone. These vessels had
evidently been broken up by roots. Nearby, at a depth of 60 centi-
meters, another upright bowl was encountered. It was badly eroded
over its entire surface and crumbled to pieces when exposed. It seems
probable that the original nature and finish of the individual pieces
has more to do with their state of preservation than does their relative
age or depth.

About 8 meters northeast of the mound or rise described above,
and only a short distance north of the remains of Edward’s “ casita ”’,
was a rough stone cairn formed by about a dozen large stone slabs
lying in rather orderly fashion. We commenced a trench at this point
but soon ran into innumerable great boulders, laid in no particular
sort of order. The trench yielded nothing but potsherds, fragments
of bulbous whistles, and a few mano and metate fragments. The
other mounds in this vicinity appeared to be similar to the one we
cross-trenched. Many of them were higher, but all had been so badly
pitted that further excavation seemed useless.

Space is lacking to describe the potsherds from these two excava-
tions. Elaborate polychrome types were abundant, a number of Ulua
types such as rows of conventionalized heads and imitation textile and
basketry designs occurred; several sherds of brown engraved ware
were noted; and a number of large handles having monkey faces in
relief on the bend, from red-on-buff cooking vessels, closely approxi-
mate the Ulua Bold Geometric style. A few heavy, coarse sherds with
rough incisions suggest graters, and a number of ground-down disks
of polychrome pottery occur. The more localized Yojoa Animalistic
and Mayoid polychrome types are generally the same as those described
in the complete vessels and in the Aguacate ceramic material. Com-
plete figurines are lacking, but a brown ware fragment, from a depth
of 30 centimeters, depicts a woman’s breasts supported by a bar or
pendant as in certain Maya stone sculptures. A few bulbous bird
and animal whistle fragments are present. Heavy volcanic stone
metates, both with and without tripod supports, were fairly numerous,
and both roller pestles and small rectangular manos occur. An ovoid
wedge or chisel, 10 centimeters in length, of hard gray-green stone is
interesting. From a depth of 20 centimeters comes a flat slab of hard
gray stone with a sharp, ground-down edge. Numerous prismatic
flakes of obsidian, a few crude obsidian and flint side scrapers, and
some red ochre, were also found. Round stone balls were fairly

NO; t HONDURAS—STRONG, KIDDER, AND PAUL 99

numerous at the site. The occurrence of two jadeite beads has already
been noted. On the whole, nonceramic artifacts are more abundant
in Lake Yojoa than in Ulua Polychrome sites.

SITE 3

About one-third of a kilometer north of site 2 we briefly investigated
what appears to be a quite different type of mound. To reach it one
proceeds through the extremely dense bush past a great number of
low, pitted, rock mounds (La Ceiba, site 3, map, fig. 20). Despite
its relative proximity to the lake we doubt if we could have found
it without the aid of Paco. The mound in question we called the
“ cut-stone mound ”’, because of the occurrence there of several large
slabs which appeared to have been worked. The main structure is
a rectangular platform-mound, 2.80 meters in height, with a north
to south length of about 20 meters, and a breadth of approximately
10 meters. The walls of this mound rise sharply, and the top, which
measures roughly 14 by 6 meters, is rather flat. The south end, which
faces the lake, has a more gradual slope, but the north end and east
and west sides rise abruptly. This platform-mound is set upon a low
circular rise, or mound, which has an estimated diameter of almost 40
meters. It was impossible to clear this entire area with the time and
men available; hence these measurements are merely approximations.

An excavation had been made near the center of the’ platform-
mound which reached down to subsoil, a depth of exactly 2.80 meters.
When cleared, the walls of this pit proved to be of brown soil con-
taining, especially near the bottom, some potsherds and charcoal. The
very bottom of the pit reached sterile yellow clay. No large rocks
occurred in the walls of the pit, but we found a few just under the
surface elsewhere on the platform. The local man who had dug the
pit told us that he had found nothing. To the south, where the
platform-mound rises from the low irregular substructure, we en-
countered a row of boulders which seemed to form a lower border.
Ten meters farther south, still on the sloping substructure, we un-
covered a number of large, flat slabs, several of which appeared to
have been more or less ground into shape. These were immediately
adjacent to an old excavation containing other slightly worked, flat
slabs. Our workers told us that four small pots had been found in
this pit. Aside from being laid flat, none of these large slabs ap-
peared to be in any particular arrangement. Two meters farther
south on the outer edge of the substructure, we encountered a row
of boulders and smoothed slabs laid end to end just under the surface.
These slabs and boulders formed a definite border to the substructure

100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

which we followed for 8 meters, paralleling the south face of the
inner platform-mound.

Approximately 20 meters south of the “cut-stone mound” is _
another, lower mound and, running west-northwest of this, is a row
of regularly aligned boulders barely projecting above the surface of
the ground. These extend for about 20 meters and then take a 90°-
turn to the south. We lacked time for further investigation, but it
is apparent that both the “ cut-stone mound ” and its neighbors repre-
sent a structural unit of some sort, the nature of which may only be
determined by adequate clearing and excavation. The rough boulder
and slab structures are similar to those at Agua Azul, to be mentioned
later. Similarly, we encountered very few polychrome sherds in our
brief work around the “cut-stone mound”’, the majority being of
coarse, plain ware.

‘ ’

CAUSEWAY AND “ CANAL” NEAR JARAL

With the exception of the concentration of burial and other mounds
near Aguacate and La Ceiba, the great bush-covered plain east of
the Jaral-Potrerillos road appears to be without noticeable archeologi-
cal sites. At present this is the area where most of the scattered
milpa farming takes place, the soil being reported as very fertile.
Just to the east of this road we discovered a great causeway and
“canal” which separates the ancient ceremonial center near Los
Naranjos from the main agricultural area and the burial sites farther
to the east (see map, fig. 20). Following up local stories of “ an ancient
canal to drain the lake ’’, we visited El Eden and found that the story
had a basis in fact (Strong, 1937, fig. 73). Later, guided by Miguel,
we followed the entire length of this structure from where it enters
the lake to its northeastern termination on the Rio Blanco (map,
fig. 20), an estimated distance of 5 kilometers. With the exception of
perhaps 300 meters at El Eden which are cleared, the remainder of
the structure is covered with dense bush, and we had to cut our way
through. It took us about 5 hours to make the trip.

The structure, which appears to be continuous, consists of a large,
flat-topped causeway on the west, bordered by a definite borrow-pit
or “canal” on the east. It enters the lake about one half a kilometer
east of Jaral. Here the borrow-pit is 25 meters wide and the mound
to the west about 8 meters wide and .75 meter high. To the east of \
the borrow-pit is a rise of about 1 meter. Where the mound crosses
the trail from Jaral to Dos Caminos (map, fig. 20) it is about 14 meters
wide and 2-3 meters in height. The ditch is not visible at this point.
‘About three-fourths of a kilometer farther north the mound is 5

NOL TE HONDURAS—STRONG, KIDDER, AND PAUL IOI

meters across and 4 meters high. The borrow-pit or ditch is 25 to
30 meters wide and is flanked by high ground to the east. At El
Eden, where it crosses the road to the cemetery, the mound is 21 meters
wide, 3.5 to 4 meters high on the ditch side, and 2-3 meters high on
the west side. The ditch here is 9 meters wide across the bottom.
About 100 meters north of this road is an apparently intentional break
in the mound wall about 16 meters wide (Strong, 1937, fig. 73).
About 22 meters farther on is another smaller break, perhaps worn
through by an old road. With these exceptions the mound or causeway
appears to be continuous throughout its entire length, though the
poor visibility due to the dense bush prevented our perceiving all
details as we cut our way through. About half a kilometer north of
El Eden the mound, now definitely turned to the west, crosses the
road, where it shows in cross-section on the east side. The ditch
here is not marked. Beyond the road both mound and ditch again
become very definite with fairly steep walls. Here, as elsewhere, the
mound has a flat top. Both terminate in a series of mysterious, deep,
dry gorges which mark the underground course of the Rio Blanco.
Miguel pointed out another series of deep pits or small gorges just
south of here extending to the west (map, fig. 20), which he said
marked the course of another underground stream called the Jutosa.
At the time of our visit (April 4) no water was visible in either stream
at this point, but during certain seasons the water level is said to rise
to a considerable height.

It is certain that any clear understanding of the function of this
interesting causeway and “canal” will depend on an equally clear
understanding of the nature and recent history of these mysterious,
semisubterranean streams. Hidden amidst almost impenetrable bush
and marked by precipitous canyons and yawning sink-holes, the solu-
tion of the problem of the Rio Blanco, which apparently drains Lake
Yojoa by some subterranean passage, is not one to be lightly attempted.
There is probably some connection between the past level of this
stream and the “canal” in question. If the river level was at one
time higher than at present, the “ canal’ would have served to irrigate
a large portion of the lower plain. Strange to say, local tradition
reverses this explanation and claims that the ancient Indians sought
to drain the lake! Since returning to Washington, the senior author
has also heard a story that a canal was dug in this vicinity about 1880
by a commercial company with some similar end in view. We have
as yet been unable to secure more definite information in this regard.
We are unable to state positively that the causeway and ditch are
not of historic origin, but, to say the least, this seems highly improbable.

102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

From the slope of the terrain we would estimate that the northern end
of the “ canal ”’ is at least 50 to 60 feet higher than its southern termina-
tion on the lake shore, hence any attempt to “ drain the lake’ would
be absurd. This same factor, however, would favor the theory of a
great central irrigation ditch, should geologists determine that the
water level of the Rio Blanco was once considerably higher than at
present.

There is another possible explanation which emphasizes the con-
tinuous mound or causeway and accounts for the ditch or “ canal ”
as merely a borrow-pit. A glance at the sketch-map (fig. 20) will show
that the causeway might well have been a ceremonial or defensive
structure enclosing the great mound group west of Jaral, since it
extends from the steep, encircling mountains all the way to the lake.
A flat area is thus entirely enclosed and in the center of this rise
the great mounds of Los Naranjos (fig. 20). Here we must leave
the problem, the true answer to which must depend on the cartographer,
the geologist, and the adequate excavation of the archeologist.

PyRAMIDS AND STONE STATUES NEAR Los NARANJOS

The dominant archeological feature on the north shore of Lake
Yojoa is the extremely impressive group of great mounds, or pyra-
mids, located about 20 minutes’ easy walk west of Jaral (see map,
fig. 20). This site was first described by Mrs. Doris Zemurray Stone
(1934) as the southernmost known Maya city and designated Los
Naranjos, after the little modern village to the west. J. B. Edwards
has made what appears to be an excellent sketch map of this site
based on his own explorations. He very kindly furnished us with
a copy of this. So far as our own sketch map (fig. 20) is concerned,
we have located and numbered these mounds in general accordance
with Mr. Edwards’ map, omitting, however, mounds 6 and 7, slightly
east of the main group, which we did not examine ourselves. Since
the Los Naranjos mounds or pyramids cover a large area and are
all covered with dense forest or bush, the preparation of an accurate,
surveyed map would be a considerable task. Until this is accomplished
Mr. Edwards’ map is the best available and, so far as our own limited
explorations went, seems generally accurate. We have not reproduced
it here, however, since it was primarily made for Frans Blom and will
probably appear in connection with publications of Tulane University.

The Los Naranjos mounds or pyramids are of great size, as indi-
cated by our photograph of one of the smaller terraced mounds
(mound 1, pl. 16, fig. 4). Yde has overdone it, however, when he shows

NON I HONDURAS—STRONG, KIDDER, AND PAUL 103

a photograph (1935, fig. 4) of the sharp, natural hills behind Los Nar-
anjos with the caption, “ View of the Mounds at Jaral.” His photo-
graph, probably taken from mound 1 (pl. 16, fig. 4), overlooks the
great mounds to the south which, however, are shrouded in jungle
and do not show in the picture. As a result, the reader might easily
assume that the natural hills which do show are the pyramids. Per-
haps the translation should have been “ View over the Pyramids ”,
rather than “ View of the Pyramids.” Mound 1, (pl. 16, fig. 4)
appears to be terraced, and we estimated its height at some 6 meters.
Mound 4 (fig. 20) is much larger and higher, perhaps 8 to 10 meters.
It is terraced and has a number of smaller mounds forming a court
on the top. There has been considerable digging here, probably by
road workers seeking paving stones, as well as by pot hunters. Sherds
seemed to be scarce on the surface. The other mounds appear to be
smaller than mound 4, but several of them are terraced and all are
worthy of careful mapping and investigation. Owing to the dense
bush, it is impossible to make adequate observations without a great
deal of clearing. Since we lacked facilities for this type of work or
for any large scale excavation, we limited our own activities to smaller
sites on the norther border of the great group (site 1, Los Naranjos,
map, fig. 20).

The occurrence of a number of fragmentary stone statues at the
Los Naranjos site is particularly interesting (Stone, 1934; Yde, 1935
and 1936; and Strong, 1937). These have all been removed from
their original sites, probably by road workers, who have undoubtedly
broken up and carried off others. Those which we located were lying
in the great plaza between the Los Naranjos mounds at three places
near the Jaral-Los Naranjos trail (map, fig. 20). Probably these
had once been placed on top of mound 4, or one of its neighbors, and
later tumbled down by the road workers. One statue represents the
body of a man or ape, with one hand resting on the hip, the other
crossing the body and resting on the shoulder (pl. 16, fig. 3). Feet,
arms, and the head had all been broken off long ago. The material
is a hard, gray, volcanic stone, and the body at present is I meter high
and 50 centimeters wide across the shoulders. The neck break at
present measures 23.5 centimeters from front to back. The body has
a primitive simplicity and grace despite its solidity. Aside from two
parallel incised lines on the back, suggesting a belt, there are no other
notable features. Mrs. Stone describes a similar mutilated figure with
a string of beads around the neck (1934, p. 126) ; hence there must
be at least two of these figures. Next to this stone body was a large
grotesque head (pl. 16, fig. 3) which evidently belongs to the body,

104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

although the uniform patination indicates that the original break oc-
curred long ago. On the head the outline of breakage at the neck is
very similar to that on the body. It measures 22.5 centimeters from
front to back. The apelike head is disproportionately large for the
body and distinctively prognathous. Anthropomorphic characteristics
are the elongated and, presumably, decorated ears and a row of incised
circles down the back of the head. Next to the other anthropomorphic
torso with beads around the neck described by Mrs. Stone (1934, p.
126) there was a similar head. We would be inclined to regard this
as identical with the one here figured (pl. 16, fig. 3), but Mrs. Stone
does not mention the macrocephaly, which is so outstanding in the
head here figured, and states that it had circular ear plugs. From her
description of it as “thick lipped” and “soft nosed” with eroded
features, the two, if not the same object, must have been similar.
We saw only one head and Yde and his party none.

Mrs. Stone (1934, p. 126) calls the anthropomorphic statue a
“stela ”, and stresses the position of the hands as representing “ with-
out a question of doubt, the Mayan sign for submission.” Yde (1936,
figs. 3, 4, and pp. 27-29) also figures this statue and apparently con-
curs with the interpretation of Mrs. Stone. In our opinion, neither
the body nor the head are Mayan. Rather they appear to us as
closely related to that widespread, and probably older, “ Chorotegan ”
style of stone statue which occurs commonly in Costa Rica, Nicaragua,
the highlands of Guatemala, and, rarely, on the Ulua (Lothrop, 1921).
The archaic simplicity of the torso, plus the crudity and simian
characteristics of the head, seem totally non-Mayan in feeling and
technique. The position of the hands alone suggests a definite Maya
convention, which may be relatively early but was certainly in vogue
at a late period at Chichen Itza (see Tozzer, 1930, pp. 155-158).
However, the same position of the arms, as well as crossed arms with
the hands on the shoulders, occurs on a number of simian stone
statues from Costa Rica, now in the United States National Museum.
These statues, and others figured by Lothrop (1921) seem much
closer to the Lake Yojoa stone figures than do the highly ornamented
and definitely stylized Mayan bas reliefs or vase paintings. If the
coincidence of hand position is not accidental in regard to the two
types, it may well have some historic significance. The relationship
of the Lake Yojoa stone carvings to the southern, ‘“ Chorotegan ”,
type is even more forcibly demonstrated by another anthropomorphic,
cylindrical, stone carving from Los Naranjos (pl. 16, fig. 1, and Yde,
1936, fig. 6). This type is identical with the rather common, anthropo-

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 105

,

morphic, giant “roller pestles’
1935, p. 148).

With the anthropomorphic head and body, we also found the stone
serpent head (pl. 16, fig. 2) referred to by Mrs. Stone (1934, p. 126)
and Yde (1936, p. 29). By some strange mistake, Yde (1936, fig. 28)
figures a side view of the “ submissive figure” which he designates
as the serpent’s head. This piece is 80 centimeters long and 37 centi-
meters wide across the broken base. It, too, is of hard gray volcanic
stone. A short distance west of mound 4, we noted a cylindrical stone,
g5 centimeters long, smooth on one end and broken on the other, which
apparently had once formed the base of the serpent head. It would be
interesting to know whether the stone serpent on the “ Islita’’, men-
tioned by Yde (1936, p. 30), was of the same type. He refers to a
photograph of it in his article (1936, p. 30), but there is none. The
style of this Los Naranjos serpent head carving (pl. 16, fig. 2) is
very well executed and distinctive, but we cannot definitely place it.
It would be extremely interesting to know whether it pertains to the
same period as do the anthropomorphic statues.

We did not see the various, undecorated, great stone slabs described
and figured as “ stelae”” and “altars” by Mrs. Stone (1934, p. 127)
and Yde (1936, p. 29, fig. 5). In the light of general distribution,
however, we would be prone to relate these to similar erect stone
slabs at Plan Grande in the Bay Islands and elsewhere on the main-
land of northeastern Honduras (compare Strong, 1935, pl. 33 and
pp. 160, 161) rather than to true hieroglyphic stelae of the Maya.
As Yde (1936, p. 29) points out, the flat rock with irregular carved
grooves on its surface in the plaza of Los Naranjos is very similar
to others occurring at Tenampua (compare D. H. Popenoe, 1936, pl. 5,
fig. 1). The adjacent flat rock with depressions suggesting three shal-
low bed-rock mortars seems more unique in this area. In a later report,
it will be possible to publish adequate photographs of these interesting
statues and carvings, but this cannot be done here. When the great
site of Los Naranjos has been cleared, and excavations on a scale
worthy of its size and importance have been commenced, more sta-
tues will undoubtedly come to light. It should then be possible to -
correlate them with their exact cultural horizons and thus end the
unsatisfactory speculation which must always center about disas-
sociated art objects.

of northeastern Honduras (Strong,

EXCAVATIONS ON THE NORTHERN BORDER OF LOS NARANJOS

Just north of mound 1 is a bushy field where a considerable amount
of digging has been done in the last 3 years. We chose this place for

106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

work because Mr. Edwards reported deep, and possibly stratified,
burials here, and M. K. Rittenhouse reported the finding of two pots
of the old Playa de los Muertos type (pl. 15, a, b) amidst similar
sherds at a depth of less than a meter. The surface of the field was
irregular, but definite mounds were hard to find in the dense, low bush.
However, ticks of all sorts were not. We selected and cleared a roughly
circular mound, 18 meters wide from north to south, 21 meters wide,
and 50 centimeters high, located about 100 meters north of the western
end of mound 1 (map, fig. 20). East of the center line of the mound
we dug a north-to-south trench 2 meters wide and 12 meters long.
The west wall of the trench was later extended 3.5 meters north, and
two western side trenches were dug well beyond the center of the
mound.

A small portion of the long western wall of this cut is shown (pl.
16, fig. 5, and text fig. 31). A layer of dark, humous soil occurred
just below the surface on the entire mound (fig. 31). Just below this,
in the thick deposit of yellow-brown mixed soil we cross-sectioned the
entire floor of a house (see house floor, fig. 31) composed of black,
burned soil containing many sherds, metate fragments, and refuse.
Beyond the edge of the diagram here shown (fig. 31), this occupation
level or floor dipped, forming a level area for about 5 meters, then
rose to the ground level, extending on into what appeared to be another
floor or occupation level beyond the limits of the excavation. The
same type of occupation level also occurred at the surface on the
south end of our main trench. Our western cross trenches showed
that the central floor area extended 2 meters to the west, where it
again rose to the surface. No post holes occurred in our cross-section
of the central house, but one was found extending below the occupa-
tion level at the north end of the central trench. No special fireplaces
were noted, but charcoal was abundant. Judging from our trenches,
there are numerous house floors in this vicinity, on or just below the
present level of the ground. These contain the finest Yojoa poly-
chrome and associated cooking ware sherds, along with other refuse.
Here, as at Naco, an expedition engaged in other than exploratory,
stratigraphic work, could easily clear entire house floors and work
out the features in detail. During the first part of our work the
trenches were taken down below the occupation area into the sterile,
yellow clay and gravel stratum (fig. 31) which occurred at an average
depth of about 1.3 meters below the surface. Polychrome sherds,
stones, charcoal, burned clay, fragments of pumice, and broken arti-
facts occurred throughout the yellow-brown soil. The mixed soil level
became darker just above the sterile layer (pl. 16, fig. 5, and text fig.

107

STRONG, KIDDER, AND PAUL

HONDURAS

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108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

31). Below the house floors, or occupation levels, we encountered
several burials (see P 2, fig. 31) of the polychrome period. For the
present we will confine ourselves to a brief discussion of features and
artifacts from this upper or polychrome level, later discussing the
materials below the sterile stratum. All vessels and important arti-
facts were photographed im situ, but these, like our complete cross-
sections and ground plans, cannot be presented here.

At the extreme south end of the main trench, at a depth of 94
centimeters, we found a small orange bowl decorated with crude red
alligators and black scrolls. It was tipped on one side. No bones
were present. A deposit of three vessels occurred near the north
end of the main trench at a depth of 1 meter. There were two super-
imposed bowls, one with a cream-white slip on which were three dark
red and orange designs of Mayoid type, probably conventionalized
serpents’ heads; the other was orange with a much conventionalized,
seated Mayoid figure. These two bowls were inverted. Next to them
was an upright, small, but striking, effigy bowl, modeled to represent
a frog. It was brown and unslipped. This was probably a grave,
occurring at the base of a refuse heap. However, no bone was found.
About 2.50 meters from the south end of the trench near the west
wall, at a depth of 1.45 meters, we found an interesting upright bowl
(pl. 14, d). This was small but of the typical Santa Rita Bold Geo-
metric olla shape and color (compare pl. 14, d, and pl. 7, a). The
present vessel has the same light orange slip, with similar red semi-
circles inside the lips as do the monkey-handled ollas, but the geo-
metric and conventionalized designs are in dull red with no black.
The base has the marked dimple, and the handles have the lugs of the
Ulua Bold Geometric olla type.

Slightly north of this pot, at the junction of one of the western
cross trenches, five vessels were uncovered at depths of from I to
1.45 meters. This immediate area had been badly disturbed by arma-
dillo burrowings (see grave P 2, fig. 30, and pl. 16, fig. 5), but all
the vessels were evidently part of one grave offering. In between them
were found a few small, crumbling fragments of human bone and
three caps from human molar teeth, thus proving for the first time
the presence of a burial. As can be seen from the photograph (pl.
16, fig. 5) and diagram (fig. 30) this burial occurred under the
southern edge of a house floor, Only one vessel shows in the cross-
section diagram, the others being just east of the trench wall. This
bowl was inverted. It was fluted and had a splotchy, light orange
slip, with crude red linear and geometric designs. On the inside a
slightly darker orange wash has been added, leaving thin vertical

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 109

stripes of the lighter slip, thus suggesting negative painting. The bowl
is blackened by fire on one side of the bottom, suggesting that, despite
its thinness and fine, hard paste, it might have been used for cooking.
Just east of this vessel, at a depth of 1 meter, was a crumbly red bowl
containing a fragile little tripod vase with black and red decorations.
Both these vessels, despite our greatest care, crumbled into tiny frag-
ments when removed. At a depth of 1.45 meters, 30 centimeters to the
south of these, was an upright incensario containing a considerable
amount of charcoal. It had nine perforations and a solid, rectangular
handle which had been completely hollowed out from the top. The
incensario had a dirty, cream-colored slip, with both upper and lower
edges outlined in red. The fifth bowl (pl. 13, 0) was slightly to the
north, at a depth of 1.45 meters, in an upright position. It has an
orange slip, with a band of white below the rim and three white
bands down the sides, dividing the outer surfaces into three panels
(pl. 13, 5). On the white bands are unusual geometric and curvilinear
designs in dark red and, on the sides, orange. In each of the three
panels occurs a most interesting prancing monkey, done in dark red.
The bottom of the bowl is flat.

In the south wall of the southerly, east to west extension trench,
at a depth of 64 centimeters, was an upright, two-handled, cooking
pot, 22 centimeters high and of a dull yellow color. Inside this large
vessel was an inverted polychrome bowl with a yellow slip, a row of
red conventionalized Mayoid designs outside the lip, and three big
black circles on the sides. The designs were badly eroded, but the
bowl was intact. The outer vessel barely held together while being
uncovered and photographed, but the moment we touched it, to re-
move it, it fell into over a hundred small pieces. Close to these two
vessels was a dull cylindrical stone bead. On the north wall of the
other extension trench, 1.25 meters deep, occurred a small open bowl
of rough gray unslipped ware. At a depth of about I meter, near
the west wall of the main trench, we found restorable fragments of
a vertical walled vase with solid, rectangular tripod feet. It has a
rich orange slip, divided into three parts on the sides by dark red
and black linear designs. Red and black lines circle top and bottom,
and each panel contains a well-executed seated Mayoid figure, with
elaborate headdress, bustle, and outstretched hand, done with fine
lines. Later, when this site was reopened to dig through the sterile
layer searching for deeper cultural material, a small ‘salad bowl”
type vessel (pl. 14, g) with an annular base, was found nearby right
side up at a depth of 1.10 meters. This bowl is interesting because
of its shape and because of the darker orange wash through which

IIo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

horizontal and vertical lines of the lighter underslip stand out. More
complicated curvilinear designs of this type occur on the inside of
the bowl. This rather peculiar type of negative painting is well shown
in the photograph (pl. 14, g).

The foregoing account, in conjunction with that of our excavations
at La Ceiba, gives an idea of the manner in which vessels representing
the various types of polychrome ware occur in the smaller Lake Yojoa
mounds. Later, in connection with a site at El Eden, we will discuss
the present slender evidence regarding the apparent vertical distribu-
tion of Yojoa polychrome pottery types in these relatively shallow
sites. Although traces of human bones occurred with only one of
our burials, there seems good reason to believe that the majority of
these pottery caches were once with skeletons, all traces of which have
now disappeared. It is further indicated that these low mounds also
served as places of habitation during the polychrome period, and that
burials occurred beneath the house floors. Probably, as at La Ceiba
and Aguacate, many of these mounds were used, or came to be used,
almost entirely for burial purposes, and it is in these that the great
masses of rocks occur. Others, like the site we are discussing, served
primarily for habitation, but burials also took place under and near
the houses. Such habitation mounds seem to have relatively few large
rocks. There remains briefly to sketch in the rest of the artifact con-
tent of the polychrome horizon at this site, and then to describe the
occurrence of a deeper, older, cultural horizon which was encountered
at the very end of our stay at Jaral.

The sherds from this one Los Naranjos mound site present a wide
variety of Yojoa polychrome types. In addition to those already men-
tioned among the entire vessels are Mayoid pieces with incising as
well as painting; Bold Animalistic sherds; heavy Bold Geometric
sherds; polished brown carved fragments; heavy gray or buff sherds
painted only on the flat upper surface with bright black, red, and
orange designs (one of these is flat with a small annular base) ; un-
slipped brown grater fragments ; and two cylindrical spouts of coarse
brown pottery. The latter may have been carried in by the natives
from older deposits since we have seen no Lake Yojoa polychrome
vessels with this type of spout. One candelario fragment of coarse
brown ware has three compartments and simple incised designs. There
are two spindle whorls, one of plain brown ware, the other a ground-
down, painted sherd. All the above come from depths ranging from
the surface to 1.45 meters in depth. A complete Mayoid figure form-
ing a whistle comes from a depth of 65 centimeters, and a bird whistle
from 1.10 meters. In addition, there are numerous fragments includ-

NO. I HONDURAS—STRONG, KIDDER, AND PAUL II!

ing broken but ornate, hollow, Mayoid effigies of fairly large size.
Figurines are varied. Solid and hollow figurines with square Mayoid
headdresses (like fig. 7, b, i, s; also see Gordon, 1808, pl. 9, 1, n, v,
s),as well as solid heads with pouting faces and simpler hair dresses,
all come from depths of 1 to 1.45 meters. This latter type of simple,
well modeled, solid figurine also occurs in polychrome deposits on
the Ulua. Several of the Los Naranjos figurines are extremely crude,
solid lumps of baked clay with grotesque, punctate faces or filleted
“ coffee-bean ” eyes. These have a decided “Archaic ” appearance but
occur in the same horizon with the polychrome pottery and ornate
figurines. In addition to the figurines this deposit yielded a consider-
able number of filleted or modeled fragments of baked clay. Many
of these are quite complex but their original form is uncertain. Ground
stone artifacts are fairly abundant, including flat ovoid lapstones ; both
flat and tripod rectangular metates (the majority with a broad grind-
ing groove) ; cylindrical roller pestles (including some that taper at
both ends) ; hammerstones ; one small rock mortar ; two small, sharp,
jadeite celts (1-1.10 m deep) ; and one brown stone bead. Chipped
stone artifacts are simple but relatively abundant. Large and small,
fragmentary, prismatic flakes of obsidian occur. There are numbers
of crude obsidian flakes, evidently used as scrapers, and a few flakes
of other stone. At a depth of I.10 meters we found the only definite
projectile point encountered, a planoconvex, obsidian dart point with
a slight, tapering stem. Even this brief summary indicates that these
small Lake Yojoa sites are far more prolific in nonceramic artifacts
than are sites on the Ulua or the Chamelecon Rivers.

Tue Oper Horizon at Los NARANJOS

Our main work at Lake Yojoa was terminated by the advent of
“Holy Week.” It was then necessary to move on to Playa de los
Muertos on the Ulua, and then to Naco if we were to complete the
survey we had outlined. We had determined the general association
of polychrome wares at Lake Yojoa but had not found any marked
stratification of cultures, nor had we encountered the old type of Playa
de los Muertos ware (pl. 15, a, D) discovered in the vicinity of Los
Naranjos by Mr. Rittenhouse. Despite his statement that it had been
found here at depths of less than a meter, we had so far been unsuc-
cessful in locating any remains other than those of the polychrome
period. It was obviously necessary to go deeper and penetrate below
the sterile yellow clay stratum encountered at our Los Naranjos site.

Therefore, in May, when the rest of the expedition went to Naco,
Mr. Paul returned to Jaral for this purpose. Efforts to locate the

112 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

exact Rittenhouse site through his former workmen again proved un-
successful. Mr. Paul therefore sank a small test trench (B) 6 meters
southwest of the mound we had excavated (fig. 20). Passing through
the polychrome horizon, which was removed in 30-centimeter levels,
he again encountered the sterile layer of yellow clay and gravel which
here averaged about 50 centimeters in thickness. Digging through this,
he encountered a brownish black clay which contained a small amount
of cultural admixture. All potsherds from this lower occupation level
proved to be of a crude, monochrome type. This lower cultural hori-
zon averaged about 65 centimeters in thickness, dipping toward the
east end of the trench, and terminating in a very hard yellow clay
which appeared to be absolutely sterile. To check these results he
dug another test pit (A) 8 meters to the southeast of the same mound.
Here he again encountered the same soil and cultural conditions (fig.
32), the brownish black clay below the sterile clay yielding only coarse
monochrome potsherds and a few other artifacts. He then returned
to our former excavation and sank a pit next to our old test cut into
the sterile stratum (fig. 31). Only 25 centimeters below the lowest
level of our former excavation he ran through this sterile layer into
the darker clay, obtaining monochrome potsherds and a small mano.
It is evident, therefore, that this direct superimposition of two cul-
tural horizons, separated by a sterile stratum of yellow clay and gravel,
extends over a considerable area. The same strata vary slightly in
thickness and absolute level at the different pits (compare figs. 31 and
32), but the sequence is the same in all. Material from the lower
cultural horizon is likewise uniform and may be discussed as a unit.

The deeper ceramic remains, some 700 sherds, are extremely crude
(pl. 15, c-w). They are all of a crumbling ware, tempered with finely
ground stone or sand. The apparent similarity in texture between
this older ware and the poorer grade of Yojoa Polychrome, especially
the cooking ware, suggests that both were made of inferior, local
clays. This point may be determined later by microscopic analysis.
All sherds from the old horizon seem to have come from small vessels.
The thickest sherd is 1.4 centimeters, the thinnest .4 centimeter, and
the majority average about .7 centimeter. Some are badly waterworn
(pl. 15, r), and the majority have one or both faces considerably
eroded. Of the 51 rim sherds, the great majority have low, slightly
flaring lips (pl. 15, c,d, f, g). A small proportion of lips are swollen,
and there are a few vertical and a few direct rims. Two sherds from
the same vessel, seem to be parts of an annular base, but the remain-
ing 30 basal sherds are all from small, flat-bottomed vessels (pl. 15 s,
u, v, w). There are no spouts, handles, lugs, or feet in the present

NO: I HONDURAS—STRONG, KIDDER, AND PAUL 113

sample. Only 12 sherds show definite traces of slip or paint. The
others range in color from a dull buff, through dull red, to a grayish
black. Despite the obvious erosion on many sherds the majority do
not appear to have ever been slipped or painted, though we cannot
be positive of this. The painted sherds include eight that have a faded

ems
Layer |

Monochrome
Layer |

Nery: Hard: Yellow: i

Clay: Gi no, >, mixture ):

YI
~— Sterile Areas

x — Approximate location of potsherds
oO 4

1 meter

Fic. 32.—Cross-section of excavation A, near site 1, Los Naranjos, showing
stratified cultural horizons.

red or pinkish slip (pl. 15, 7) ; two with a dull white slip or wash;
and two that have definite areas painted a very dull red and black on
the inner surface. Other decorative efforts are scant. A few sherds
have raised ridges below the rim (pl. 15, 2, /), one has such a ridge
with regular indentations, and a few sherds have simple, linear de-
signs incised on the outer rim or body. If the present sample is
at all adequate, this, the oldest known Lake Yojoa pottery, appears

114 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

to be the most primitive ceramic type yet encountered in Honduras.
Technically, since a few sherds are painted, we should designate this
ware as Yojoa bichrome. Actually, the great majority of sherds are
unpainted and all of them are definitely inferior in both texture and
finish to either the Playa de los Muertos Bichrome or the Santa Rita
(farm 17) Bichrome wares. For this reason we have tentatively
designated it as Yojoa “‘ Monochrome ”’, subject to change when the
results of larger excavations yielding adequate ceramic samples are
at hand.

Equally puzzling is the relationship of this Yojoa ‘‘ Monochrome ”
to the two vessels excavated by Mr. Rittenhouse in this immediate
vicinity. Both these vessels are well modeled, incised, and painted red
and buff (pl. 15, a, b). Both originally had spouts. They are un-
doubtedly closely allied, if not identical, with the old Playa de los
Muertos Bichrome (pls. 10, 11, and figs. 17, 18). Yet not one of
our sherds from the deep stratum is positively of this type. We have
no reason to doubt that the general location of the two Rittenhouse
vessels was substantially as reported. It is apparent, therefore, that
our Yojoa Monochrome is either a strangely isolated sample of crude,
domestic ware, actually pertaining to the Playa de los Muertos Bi-
chrome, or else that both Yojoa Polychrome, Playa de los Muertos
Bichrome, and a new type, Yojoa Monochrome, all occur in shallow
deposits on the northern borders of Los Naranjos.

Other artifacts from the older horizon at Los Naranjos included
three figurine fragments (pl. 15, e, 7). All are of solid, baked clay,
and none are slipped. The crudely modeled little head (pl. 15, e)
has a knot of hair on the back, and the body (pl. 15, 7) has broad
modeled and grooved buttocks which have been smoked black. Head
and body are from different figurines. The third figurine torso is
also of coarse, dull buff pottery. It is somewhat similar to the old
Playa de los Muertos horizon figurine torsos (pl. 11, f-v), but is not
so well modeled and has no breasts. Ground stone artifacts from this
older Yojoa horizon include one small rectangular mano with ground
sides and battered ends; one small rectangular stone (pl. 15, 1) of
unknown use; and a fragment of ground sandstone (pl. 15, ¢) which
may be froma simple metate, although it has uneven grinding surfaces
on the two sides. There are no true prismatic flake fragments from
this horizon. There are, however, several irregular flakes of obsidian
(pl. 15, m), and one irregular prismatic flake with a side point which
also shows a use retouch. There is one rather large, gray flint side
scraper anda flint flake. Here again, definite conclusions are precluded

NO. I HONDURAS—STRONG, KIDDER, AND PAUL II5

by the small size of the present sample, but both the figurine and
artifact fragments, like the pottery, show unique types.

There can be no doubt that, just below the elaborate polychrome
horizon at Los Naranjos, there occurs another cultural level which
appears to be surprisingly primitive. When it is remembered that
our deepest excavations at Los Naranjos were slightly less than 3
meters, it can be seen that here is an area where deep excavations may
yet furnish evidence regarding the truly simple cultures of Central
America. On the Ulua, where we conducted our largest and deepest
excavations, we were eventually stopped by reaching the water level.
At Lake Yojoa this was not the case. A larger expedition, with
adequate time and equipment, providing it is not led too far astray
by the richer polychrome deposits, should be able to work out a most
important sequence of human occupation in this immediate region.

OTHER SITES

Seeking for a deep Yojoa polychrome refuse heap suitable for
stratigraphic analysis, we conducted a small excavation about 1 kilo-
meter northeast of El Eden (see site 2, near that village, map, fig.
20). Miguel had brought us a number of polychrome sherds from this
place, his sample including a dark brown and highly polished tripod
bowl fragment with delicate geometric incisions on the body, an in-
censario fragment with rather elaborate geometric painted designs,
and a small whistle shaped like a turtle. This sherd deposit was located
in the abrupt face of a steep bank terminating a small, densely wooded
arroyo. This arroyo led down toward one of the deep sink-holes which
here mark the course of the Rio Blanco. The region is a maze of small,
abrupt canyons or sink-holes, and is covered by unbelievably dense
bush.

We dug a trench, 3.5 meters long and 1 meter wide, along the face
of the bank, encountering our first potsherd at a depth of 40 centi-
meters. From this point down all artifacts were segregated according
to horizontal levels averaging 30 centimeters in depth. Unfortunately,
only three levels were encountered when we ran into sterile yellow
clay. The exposed surface of the bank below this point appeared
devoid of any human detritus. The upper 30-centimeter level con-
tained a number of well-executed fragments of polychrome ware
with highly conventionalized Mayoid designs. A tripod leg proved
to belong to the dark brown and incised dish fragment secured earlier
by Miguel. A number of heavier polychrome sherds had geometric
designs in red and black, suggesting the Bold Geometric Ulua type,

116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

but no definite Bold Animalistic sherds occurred. Crudely painted
and occasionally incised sherds of domestic ware were fairly abundant
in all levels. The middle layer yielded one conventionalized Mayoid
sherd similar to the above, and a considerable number of red and black
or orange sherds with designs suggesting the Bold Geometric. Defin-
itive Bold Animalistic designs were lacking. The bottom level was
similar but lacked both definitive Mayoid or Bold Animalistic designs,
although several badly eroded polychrome sherds may have been of
these types. To sum up, the El Eden polychrome site proved negative
so far as any obvious stratification of ceramic types was concerned.
The absence of definite Bold Animalistic type sherds is interesting
but hardly significant, owing to the small sherd sample.

At excavation B, Los Naranjos (near site 1, map, fig. 20), a simi-
lar stratigraphic excavation was made. Here, again, only three 30-
centimeter levels of polychrome sherds were obtained, the depth of
the upper Yojoa Polychrome deposit being similar to excavation A
(fig. 32). The top level contained rim fragments of small bowls,
many of which had thickened lips. These sherds have conventional
and rather massive red and black designs on orange and, in one case,
white, backgrounds. Three basal fragments, one flat, one dimpled,
and one annular, occurred. The latter is a dark brown, almost black,
overfired piece. The middle level contained sherds with similar, con-
ventionalized Mayoid designs, and a few with well-executed and iso-
lated serpent motifs. A few sherds from this level are of the Bold
Animalistic type. Basal fragments include two flat and two dimpled
bottoms. The lowest sherd level included a bowl fragment with an
elaborate open-winged bat (like pl. 3, b; Gordon, 1898). Several
rim sherds from vertical-walled Mayoid vases have rows of typical
Ulua conventionalized faces, and a fragment of a tripod plate has
a similar design motif. In addition, there are a number of dull orange
sherds with more conventionalized black and red geometric designs.
From this lowest level comes a splendid Mayoid vase (pl. 12, a)
encountered in a broken condition at a depth of 1.25 meters in the
original excavation B test pit. This vase, with a definite rim, marked
entasis, and a flat bottom, has an orange slip with complex anthropo-
morphic and glyph designs in brownish yellow, purplish red, and
black. Thus, although the evidence is slender, there is some sugges-
tion that the Lake Yojoa Polychrome wares exhibit the same trend
from the more realistic to the conventional as was true of Ulua
Polychrome pottery decoration. The occurrence at La Ceiba of both
an extremely conventionalized Bold Animalistic vase (pl. 13, @)
at a depth of only 15 centimeters, and a splendid, realistic Mayoid

NO. I HONDURAS—STRONG, KIDDER, AND PAUL II7

_vase (pl. 1) at a depth of 1.25 meters furthers this possibility. That
both the more realistic and the more conventional aspects of the
Mayoid and the Bold Animalistic tradition occur in the shallow Lake
Yojoa Polychrome sites is certain. Their exact interrelationship, how-
ever, remains to be demonstrated.

Two other sites on the north end of Lake Yojoa may be briefly
mentioned. The first of these is a little island, called merely, La
Islita. It is near the shore between Jaral and Agua Azul (see map,
fig. 1). Yde (1936, p. 30) describes a stone serpent head from this
place which, subsequent to their visit, was reported to have been
smashed by natives looking for treasure. He refers to, but does not
reproduce, a photograph of this statue. Our guide brazenly showed
us a simple cylindrical statue, apparently anthropomorphic and about
I meter tall, the head and face of which had been completely smashed
by him in a futile search for treasure! Only a carved ear remained
and the rounded pediment which was like the simplest statue at Los
Naranjos (pl. 16, 1). He did not know of the stone serpent head
but claimed a similar anthropomorphic statue had been taken from
the island to Tegucigalpa. Another man told us of a stone serpent
head that had been found on this island, but said that local people

had thrown it in the lake! The island is very steep and densely wooded.
- On top of one of the hills are a number of low, irregular mounds,
some of which are covered with rock slabs. Our guide had dug pits
in several of these and claimed to have found a few pieces of painted
pottery. The soil of the mounds is a red clay. Aside from a few
coarse brown sherds we saw no pottery at the site. There are said
to be other mounds of a similar nature on the island, which we did
not see. It is sincerely to be hoped that this interesting site may be
scientifically worked by archeologists before it is completely ruined.

The other site is a group of three impressive mounds located in
the open pine and savannah country about 2 kilometers northwest
of the ranch house at Agua Azul (see map, fig. 1). The largest mound,
to the north, is conical with a flattened top. It is approximately 7
meters high by 9 meters across, and is flanked to the west and south
by a terrace edged with straight walls of large boulders about 1 meter
in height. The west wall is about 5 meters from the edge of the mound
and the south wall about 6 meters from it. The south wall is com-
posed of several thicknesses of stone and terminates just east of the
center of the mound. The west wall is only one stone thick and termi-
nates just beyond the north edge of the mound. About Io meters
south of the edge of this terrace are two more parallel mounds. The
mound to the east has an approximate length of 7 meters, and a

118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

height of 5 meters. The mound to the west is smaller and steeper,
being about 4 meters long and perhaps 3 meters high. Local men from
Siguatepeque have excavated a small hole in the terrace south of
the big mound and a large trench, 5 meters wide and nearly 6 meters
deep, on the west side of the same mound. The earth wall of this
trench shows successive curving layers of black charcoal, suggesting
that the mound had been built up at different times and the remains
of fires on the top had been scattered down the sides. The small pit
on the terrace showed nothing. We were unable to find any potsherds,
either in the cut or on the surface. According to local report the origi-
nal diggers encountered nothing but a very little broken pottery. This
is a striking mound group and, as already mentioned, seems similar
in some ways to the “ cut stone mound ” which is buried in the dense
bush near La Ceiba (site 3, map, fig. 20).

This concludes the list of sites visited by us around the north end
of Lake Yojoa. We have since heard that local pot hunters have
opened up a new series of ancient cemeteries between La Ceiba and
Agua Azul. Other sites are reported in the mountains to the north
(see map, fig. 20), at Saucé, and elsewhere around the lake, but we
lacked time to visit these.

SUMMARY AND TENTATIVE CONCLUSIONS

The present reconnaisance of the Ulua-Yojoa region opens promis-
ing vistas. It reveals incomplete but considerable sequences of local
development, and it demonstrates that the interplay of northern and
southern cultural forces, so strongly suggested by linguistic, eth-
nographic, and historic sources, is very definitely reflected in the arche-
ological record.

Since ceramic remains constitute the most abundant and helpful
guides in attaining any understanding of the development of the pre-
historic cultures of northwestern Honduras, we may preface our brief
summary by a table showing the sequence and groupings of Ulua-
Yojoa ceramic types as known at present (table 1). Of these the
Naco Polychrome is definitely historic and represents, apparently, the
late Nahuatl occupation of the region. Spinden, Tozzer, Mason, and
Vaillant, who have examined this material, state that it appears to be
related to certain late prehistoric wares of Mexico. Naco Polychrome
pottery will probably be found at other sites occupied or influenced
by these intrusive Nahuatl peoples. It may occur at Tenampua (com-
pare Popenoe, 1936, p. 572 and fig. 2). In the same way that the
occurrence of Spanish crockery in association with Naco Polychrome
sherds connects the site with the historic period, so the occurrence of

NO. I HONDURAS—STRONG, KIDDER, AND PAUL II9Q

simply decorated Ulua Polychrome sherds (pl. 3, 0) in Naco refuse
mounds indicates that other, local cultures were contemporaneous in
the region. Little attention has as yet been paid to the historic and late
prehistoric cultures of the Jicaque and other local inhabitants of
northwestern Honduras.

TABLE 1.—Apparent Sequence of Ceramic Types in Northwestern Honduras.

Ulua Yojoa
Historic: Naco Polychrome
Prehistoric: (surface mounds?)
(late) Ulua Polychrome (including) Yojoa Polychrome (including)
Upper Upper
; Bold
Mayoid p Mayoid Bold Animalistic
Geometric
Lower Lower
(to)
Ulua Bichrome (Santa Rita) (here?)
Playa de los Muertos Bichrome Yojoa ‘‘ Monochrome ”
(P.d.1.M.) (Los Naranjos)
(early) (or here?)

TaBe 2—Probable Correlation between present Ulua Polychrome Classification
(Table 1) and those of Gordon (1898) and Vaillant (1927)

Upper Mayoid | Gordon’s B | Vaillant’s III and IV
(Wilttaw Poliy.chworiey /s) siete =i elererereseleneus

Upper Bold Gordon’s C | Vaillant’s V
Geometric

Ul IPolichrome secre ceteleterciensieverare
Sie ecs a, oy Lower Bold Gordon’s C | Vaillant’s V

ts Mayoid| Gordon’s A | Vaillant’s I and II
Geometric

The prehistoric polychrome wares of the Ulua have been classified
on typological grounds by Gordon (1898), and by Vaillant (1927).
In general their classifications seem to accord with our stratigraphic
findings as above (table 2). Gordon clearly distinguished the Bold
Geometric as Type C, and the Mayoid as Types A and B, but has
nothing to say regarding sequence. Vaillant makes a mistake when
he assumes, on stylistic grounds, that the Bold Geometric (Ulua Poly-
chrome V) developed out of the Mayoid style and was therefore later.
Strong makes the same mistake in regard to the related Bay Island

120 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Polychrome I (Upper Mayoid, plus southern influences) and the iden-
tical Bay Island Polychrome II (Upper Bold Geometric) (1935,
p- 145). Actually the Mayoid and the Bold Geometric have been
shown to be parallel developments; thus Vaillant’s Ulua Polychrome .
V is contemporaneous with his Ulua Polychrome I and II (his
Ulua Polychrome IV contains both Upper Mayoid and Upper Bold
Geometric constituents), and Strong’s Bay Island Polychrome I
is, in all probability, contemporaneous with his Bay Island Polychrome
II. Vaillant (1927, p. 266) was careful to point out the entirely tenta-
tive nature of his assumed sequences. Further, in regard to the Ulua
Polychrome V (and Salvador Polychrome VI) he states: “ There is
a strong suspicion of the same non-Maya factors influencing both
these styles. The source of the influence is not discoverable in Maya
districts, and one thinks vaguely of the south and east, of Nicaragua,
eastern Honduras, and Costa Rica to locate a source” (1927, p. 170).

Recently Tschopik (1937), in a brief but valuable analysis of tex-
tile motifs on Gordon’s Ulua Polychrome pottery, has independently
pointed out this stylistic dichotomy. He groups Vaillant’s Ulua Poly-
chrome I-IV as Ware A [Mayoid], and the latter’s Ulua Poly-
chrome V as Ware B [Bold Geometric]. Tschopik points out that
there are consistent differences in both form and decoration between
the two, and that A is Mayoid, whereas B has a definite relationship
in form and decoration with ceramic types from Salvador, Nicaragua,
and Costa Rica. He, too, repeats the theory that naturalistic designs
are apt to be earlier than geometric, suggesting that Ware A is earlier
than Ware B, thus falling into the same error as Vaillant and Strong.

At Santa Rita these two major styles (Mayoid and Bold Geometric)
are intermixed throughout almost 4 meters of Ulua Polychrome de-
posits. Although they blend in certain intermediate types of vessels,
each style in general keeps to its own particular genius, and each shows
a parallel development from a finer and somewhat more realistic dec-
oration in the lower levels, to a more conventionalized and geometric
decoration in the upper levels. Thus, the Lower Mayoid has priestly,
processional, and “ dancing ” figures in open panels, whereas the Up-
per Mayoid has florid, conventionalized, over-all designs, geometric
motifs and, often, animal head lugs. The Lower Bold Geometric has
intricate linear and geometric designs with remarkable, cursive ani-
mals or birds in open panels, whereas the Upper Bold Geometric be-
comes simpler, drops the animals, but retains textile and geometric
designs. At Las Flores, also, both the Mayoid and the Bold Geometric
styles occur in the same excavation, but here both are of the upper
and later, conventionalized type. It is worth noting that the only

NO. I HONDURAS—STRONG, KIDDER, AND PAUL I21

metal object recovered in any of our excavations, a small copper fish-
hook, came from these levels at Las Flores. The Ulua Polychrome
horizon overlying the deep stratum at Playa de los Muertos is also
late. Here only the Upper Mayoid and Upper Bold Geometric occur.
It is undoubtedly significant that the typical, swollen, monkey-handled
olla form of the Bold Geometric tradition (pl. 7, a-d), and the vertical-
walled vase form of the Mayoid tradition (pl. 8, a-b), both persist
practically unchanged throughout the entire Ulua Polychrome series.
This occurrence argues rather strongly against any very considerable
time period being assigned to the Ulua Polychrome period.

The polychrome wares of Lake Yojoa are closely related to those
of the Ulua. Not only does Yojoa Polychrome ware contain a large
number of forms and motifs identical with those of the Ulua Poly-
chrome, but it also manifests a very similar division into two major
stylistic traditions. In general, however, Ulua and Yojoa Polychrome
ware vessels are distinguishable. The Yojoa Mayoid type, as well
as the Bold Animalistic type, finds many close parallels in polychrome
. vessels from eastern Salvador (see Vaillant, 1927, figs. 35-40). It
seems strange that no Plumbate ware whatsoever was recovered in any
of our excavations, either on the Ulua or at Lake Yojoa. The Bold
Animalistic type from Lake Yojoa differs from the Ulua Bold Geo-
metric in the relative rarity of monkey-handled ollas and the prevalence
of bird, monkey, alligator, and other animal design motifs. Regard-
ing the internal development of Yojoa Polychrome ware decoration,
there is some very slight evidence that it parallels the trend of the Ulua
Polychrome series from better executed realistic, to conventional
and geometric design. However, the 14 meters of Yojoa Polychrome
deposits so far investigated have not as yet yielded very satisfactory
evidence in this regard. The fact that Ulua Polychrome deposits oc-
cur throughout 3 to 4 meters of alluvial and cultural deposition,
whereas the known Yojoa Polychrome refuse deposits are less than
2 meters in depth, is undoubtedly significant. In our opinion, how-
ever, this discrepancy is probably due to the very different physio-
graphic conditions in the two regions, rather than to differences in
time.

Of the three wares that have been stratigraphically established as
earlier than the Ulua-Yojoa Polychrome series, the Playa de los
Muertos Bichrome (table 1, and pls. 10, 11) is the most clearly
defined. This is the type D of Gordon (1898). Vaillant (1934) has
pointed out that this horizon contains a majority of traits, mainly
ceramic, that are characteristic of the QO complex. It is undoubtedly
significant that, whereas Playa de los Muertos Bichrome ceramics

122 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

represent an advanced pottery type so far as texture, surface finish,
modeling, and incising are concerned, they appear to mark an experi-
mental and inept stage in the use of surface painting. Especially char-
acteristic of this horizon are highly polished, modeled and spouted
forms ; flat-bottomed, vertical-walled vases; low dishes with flaring
incised walls or everted, flat, incised lips or both; and solid female
figurines, which may or may not have a white slip. There is con-
siderable resemblance between the simple but effective modeling of
these figurines (pl. 11, ¢, u, v) and the stone statue of a man or
ape at Los Naranjos, Lake Yojoa (pl. 16, fig. 3). These traits,
plus the occurrence of jadeite artifacts and the varied experiments
with painted decoration, all indicate that here was an early and potent
cultural manifestation of more than local significance. In so far as
data are available (R. E. Smith, 1936, and Uaxactun sample sherd
collections), we see considerable resemblance between this Playa de
los Muertos Bichrome ware and the two earliest stratigraphic periods
at the old Maya city of Uaxactun. These have been termed Mamon
and Chicanel, and both precede the Maya Polychrome period.

The determination of the northern and the southern extent of the
Playa de los Muertos horizon is one of the important problems in
Middle American archeology. Even more important is the determina-
tion of the simpler ceramic horizons from which it developed. In
Honduras we have as yet no clues to this earlier period unless the
so-called Yojoa “ Monochrome” (pl. 15, c-w) is as truly primitive
as it superficially appears, and can be demonstrated as stratigraphically
earlier than the developed Playa de los Muertos culture. The later
break, between the Playa de los Muertos Bichrome and the Ulua
Polychrome, is in part bridged by the deepest cultural horizon at
Santa Rita containing Ulua Bichrome ware. The most outstanding
feature of the Ulua Bichrome ceramics is the presence of Usulatan
ware sherds. According to Lothrop this is “the earliest painted
pottery now known from Central America”, and, although it occurs
occasionally in the form of trade pieces at Old Empire Maya sites,
it seems to center in Lenca territory in eastern Salvador (1933, pp.
47-51). There is rather close resemblance between our Usulatan
sherds with short, solid legs (pl. 9) and the early Chukumuk pottery
from Lake Atitlan in the highlands of Guatemala (Lothrop, 1933,
p. 49). Similarly, the tetrapod Usulatan bowl recovered by Gordon at
a depth of “ 26 feet”? in his Playa de los Muertos excavations is of
an identical type. Thus, there is a linkage in this regard between the
deep horizons at Playa de los Muertos and at Santa Rita, despite the
fact that our own sample of Playa de los Muertos Bichrome ceramics

No. I HONDURAS—STRONG, KIDDER, AND PAUL 123

contains no definite Usulatan ware. In addition, this clear linkage
between early Ulua and early Guatemalan highland cultural horizons
is of great interest. We have assumed that Ulua Bichrome is some-
what later than the Playa de los Muertos Bichrome on stylistic grounds
and because the sterile area separating the former from the Ulua
Polychrome is thin compared to that separating the Playa de los
Muertos Bichrome from the overlying Ulua Polychrome (compare
fig. 6, and fig. 16). This, however, is at best a dubious procedure, since
we do not as yet know the physiographic nature of either sterile
stratum. Moreover, it must be remembered that only the Upper May-
oid and Bold Geometric Ulua Polychrome types occur in the over-
lying cultural stratum at Playa de los Muertos, whereas both these
and the earlier Lower Ulua Polychrome wares occur above the Ulua
Bichrome at Santa Rita. These details, like the cultural and temporal
placing of the puzzling Yojoa “ Monochrome” ceramics and the
“Chorotegan ” stone statues at Los Naranjos, must await further
excavation.

Tracing the relationship of the native cultures of northwestern
Honduras backward from the known historic, we have already veri-
fied the presence of a late Nahuatl migration from Mexico through
the finds made at Naco. Similarly, in the Ulua Polychrome period
we find two interlocked but distinct styles occurring in the same sites,
the Mayoid and the Bold Geometric, which at Santa Rita persist and
develop simultaneously over a considerable period. Lake Yojoa Poly-
chrome is also composed of a Mayoid and a so-called Bold Animalistic
tradition. This original fission and subsequent parallelism of both Ulua
and Yojoa Polychrome ceramic development has obvious sociological
as well as archeological implications. At both Ulua River and Lake
Yojoa Polychrome sites one of these styles is Mayoid and the other
is of southern origin. For linguistic and ethnographic reasons previ-
ously discussed, it seems highly probable that the Bold Geometric
element of the Ulua Polychrome was contributed by Jicaque peoples,
whereas the very similar Bold Animalistic element in Yojoa Poly-
chrome was due to the related Lenca. Since the Mayoid element com-
prises about one half of the Ulua and Yojoa Polychrome ceramic re-
mains, it can hardly be explained as due solely to trade or indirect
influence. It seems far more logical to assume that intermixed Maya,
Jicaque, and Lenca peoples were living together at these sites and
that perhaps the pottery-makers of each ethnic group clung to their
own art styles over a considerable period. The quite remarkable
florescence and the high and complex artistic attainments of the Ulua

124 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

and the Yojoa Polychrome periods are in all probability the direct
results of this cultural and physical amalgamation.

We have at present no means for dating the exact period repre-
sented by these Lenca and Jicaque styles which apparently stem from
Nicaraguan and Nicoyan culture centers to the south. On the other
hand, there is in the nearby Maya city of Copan a series of dated
monuments ranging from 9: 11.0.0.0 (stela 3) .to 9: 17. 12.0.0. (stela
C) (or, roughly, according to the Goodman-Thompson-Martinez
correlation, between 650 and 800 A. D.), in association with which
there occur pottery vessels (Vaillant, 1927, and Lothrop, 1933, p. 66
and 1936 b, p. 69). We have attempted to correlate our Ulua and
Lake Yojoa Polychrome series with Vaillant’s classification of Copan
wares, but owing to the selective nature of the Copan collections, as
well as the paucity of illustrative material, this has proved imprac-
ticable for the present. The Copan ceramic series in the Peabody
Museum, as a whole, seems quite distinct from the Ulua-Yojoa Poly-
chrome wares, although numerous similarities do exist. Vaillant points
out the occurrence of Ulua Polychrome sherds at mound 36 in Copan,
a point we were able to verify for ourselves at the site, but there is
reason to believe that these deposits are later than the Copan series or
perhaps intrusive. According to Vaillant (1927, p. 271) the trend
of the Ulua Polychrome wares “suggests the years after the fall
of Copan.” If this is the case, it may serve to point out when the
Maya Old Empire dispersal into Salvador and northwestern Honduras
took place and how their developed polychrome wares came to be
grafted on to those of the Lenca, Jicaque, and, probably, the Pipil,
with whom the various Maya groups settled. When adequate strati-
graphic studies of the entire range of Copan ceramics have been made
and correlated with the ceremonial series from the stelae vaults,
described by Vaillant, there is reason to believe that the Ulua-Yojoa
Polychrome series may also be approximately dated.

Such excavations should also throw light on the origin or deriva-
tion of the southern Mayoid Polychrome ceramic tradition. Did it
arise from a groundwork similar to the Playa de los Muertos cul-
ture in the Peten, perhaps at Uaxactun or Holmul, spread from there
to Copan, and thence to Salvador and the Ulua? Or are there inter-
mediate stages between the developed Polychrome and the Playa de
los Muertos horizons present but as yet unknown in Honduras, at
Copan, or in Salvador? An even more basic problem concerns the
suggested relationship between the ceramics in the oldest horizons
at Uaxactun in the Peten and Chukumuk in the Guatemalan high-
lands, with the Playa de los Muertos Bichrome and Ulua Bichrome

NO. I HONDURAS—STRONG, KIDDER, AND PAUL 125

wares respectively. Lothrop (1933, p. 62) believes that the elements
shared in common by the earliest known cultural horizons in the high-
lands and the Atlantic lowlands of Guatemala were derived, not from
one another, but from a parent culture to the south. When the
Uaxactun materials are available, the role of the southern Playa de
los Muertos culture as a donor or a recipient may be tested. These
are questions for the future but, thanks to the growing scientific
vogue of the shovel, they are questions that may soon be answered.

LITERATURE CITED

Bancrort, H. H.
1882. The native races of the Pacific States, vols. 1-5. San Francisco.
1883-7. History of Central America, vols. 1-3. (Vols. 6, 7, and 8 of
Bancroft’s Works. San Francisco, 1882-1890.)
BLACKESTON, R. H.
1910. Recent discoveries in Honduras. Amer. Anthrop., n. s., vol. 12, no. 4,
Pp. 536-541.
Brom, Frans, Grosjean, S. S., and Cummins, Harotp
1933. A Maya skull from the Uloa Valley, Republic of Honduras. Tulane
Univ. Studies in Middle America, Publ. 5, no. 1. New Orleans.
CoLteccION DE Documentos INEpIrTos relativos al descubrimiento, conquista
y colonizacién de las posesiones Espafiolas en America y Oceania,
tomos 1-41. Madrid, 1854-84.
Cummins, Harotp. See Biom, FRANs, GrosyJEAN, and CUMMINS.
Diaz pEL CAsTILLo, BERNAL
1908-16. The true history of the conquest of New Spain, vols. 1-5. Edited
and published in Mexico by Genaro Garcia. Translated by A. P.
Maudsley. Hakluyt Soc. Publ., ser. 2, vols. 23-25, 30, and 40.
London.
Gorpon, GEORGE Byron
1898. Researches in the Uloa Valley, Honduras. Mem. Peabody Mus.
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GrosjJEAN, S. S. See Biom, Frans, GrosJEAN, and CUMMINS.
Haset, S.
1880, The sculptures of Santa Lucia Cosumalwhuapa in Guatemala. Smith-
sonian Contr. Knowl., vol. 22, art. 3, pp. I-90. Washington.
pE Las CASAS, BARTHELEMI
1922. (Euvres de Don Barthelemi de Las Casas, évéque de Chiapa. Edited
by J. A. Llorente. Vols. 1-2. Paris.
LEHMANN, WALTER
1910. Erbebnisse einer Forschungsreise in Mittelamerika und Mexico, 1907-
‘1909. Zeitschr. Ethnol., vol. 42, pp. 687-749. Berlin.
1920. Zentral Amerika. Vols. 1-2. Berlin.
LotnHrop, S. K.
1921. The stone statues of Nicaragua. Amer. Anthrop., n. s., vol. 23, no. 3,
PP. 311-319.
1933. Atitlan. An archaeological study of ancient remains on the borders
of Lake Atitlan, Guatemala. Carnegie Inst. of Washington. Publ.

444.

126 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

1936a. Sculptured pottery of the Maya and Pipil. Maya Research (Mexico
and Central America), vol. 3, no. 2, pp. 140-152. New Orleans.
1936b. Zacualpa. A study of ancient Quiche artifacts. Carnegie Inst. of
Washington, Publ. 472.
Mason, Oris T.
1889 a. How to straighten a spear shaft. Amer. Anthrop., ser. 1, vol. 2,
no. 2, p. 158.
1889 b. Music in Honduras. Amer. Anthrop., ser. 1, vol. 2, no. 2, p. 158.
pE Paracio, D1irco GARCIA
1860. Carta dirijida al Rey de Espana. Afio. 1576. (Original Spanish with
English translation.) Squier's “Collection of rare and original
documents and relations”, no. 1. New York.
PorENOE, DororHy H.
1934. Some excavations at Playa de los Muertos, Ulua River, Honduras.
Maya Research, vol. 1, no. 2, pp. 61-86. New York.
1936. The ruins of Tenampua, Honduras. Ann. Rep. Smithsonian Inst.
1935, PP. 559-572.
SPINDEN, HERBERT J.
1925. The Chorotegan culture area. Compt. Rend. 21st Congr. Internat.
Amer., Goteborg, 1924, pp. 529-545. Goteborg.
SguieErR, EPHRAIM GEORGE
1858. The States of Central America. New York.
1859. A visit to the Guajiquero Indians. Harper’s New Month. Mag.,
vol. 19, no. 113, pp. 603-619. New York.
1860. Some account of the Lake of Yojoa or Taulebé, in Honduras, Central
America. Journ. Roy. Geogr. Soc., vol. 13, pp. 58-63. London.
1869. Tongues from tombs, or the stories that graves tell. Frank Leslie’s
Illustr. Newsp. New York.
1870. Honduras. Descriptive, historical, and statistical. London.
SmitH, H. E.
1936. Preliminary shape analysis of the Uaxactun pottery. (Typed script,
with illustrations reproduced photographically.) Guatemala.
Stone, Doris ZEMURRAY
1934. A new southernmost Maya City. Maya Research, vol. I, no. 2, pp.
125-132. New York.
Stronc, WILLIAM DUNCAN
1934a. Hunting ancient ruins in northeastern Honduras. Explorations and
Field-Work of the Smithsonian Inst., 1933, pp. 44-48.
1934 b. An archeological cruise among the Bay Islands of Honduras. Ibid.,
PP. 49-50.
1935. Archeological investigations in the Bay Islands, Spanish Honduras.
Smithsonian Misc. Coll., vol. 92, no. 14, pp. 1-176.
1937. Archeological explorations in northwestern Honduras. Explorations
and Field-Work of the Smithsonian Inst., 1936, pp. 75-82.
Tuomas, Cyrus, and Swanton, JoHN R.
1911. Indian languages of Mexico and Central America and their geo-
graphical distribution. Bur. Amer. Ethnol. Bull. 44.
DE TORQUEMADA, JUAN
1723. Los veinte i un libros rituales i Monarchia Indiana, con el origen y
guerras, de los Indios Occidentales. Madrid.

NO. 1 HONDURAS—STRONG, KIDDER, AND PAUL 127

Tozzer, ALFRED M.
1930. Maya and Toltec figures at Chichen Itza. Proc. 23d Internat. Congr.

Amer., New York, 1928, pp. 155-164. New York.
TscHopIk, Harry, JR.
1937. Textile motifs from Uloa Valley pottery. MS., Peabody Mus.,
Harvard Univ.

VAILLANT, G. C.
1927. The chronological significance of Maya ceramics. Manuscript thesis

submitted in partial fulfillment of the requirements for the degree
of Doctor of Philosophy, Harvard Univ.
1934. The archaeological setting of the Playa de los Muertos culture. Maya
Research, vol. 1, no. 2, pp. 87-100. New York.
WELLs, Wo. V.
1857. Explorations and adventures in Honduras. New York.

YodE, JENS
1935. Forelgbig Beretning on Nationalmuseets og Tulane Universitetets

Ekspedition til Mellemamerika 1935. Sertryk Geogr. Tidsskr.,
vol. 38, nos. 3-4, pp. 137-155.

1936. A preliminary report of the Tulane University-Danish National Mu-
seum expedition to Central America. Maya Research (Mexico and
Central America), vol. 3, no. 1, pp. 25-37. New Orleans.

EXPLANATION OF PLATES
PLATE I

Processional figures on a Yojoa Polychrome vase, Mayoid type. Site 2, La
Ceiba (13.2 cm high).

PLATE 2
Various Chamelecon and Ulua River sites

Fragment of ball court ring im situ, Naco.

BIG. 1.

Fic. 2. Thin plaster walls, heart of mound 3, Naco.

Fic. 3. Excavation 1 and start of excavation 2 (right) at Santa Rita (farm 17).
Fic. 4. Site at Tres Piedras, showing mounds and plaza cross-sectioned by

Chamelecon River.
PLATE 3
Naco sherds

a, c-w, Naco Polychrome; b, Ulua Polychrome sherds found at Naco.

PLATE 4
Naco sherds and artifacts

a, incensario; b-g, figurine fragments; h, whistle; 7, j, spindle whorls;
k, obsidian flake knives; J, clay bobbins; m, Spanish colonial crockery ;
n-p, textile-marked sherds; gq, s, t, u, v, x, y, 2, sherds with molded or
carved designs; r, incised sherd; zw, “ candelarios.”

128 SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. 97

PLATE 5
Upper Ulua Polychrome pottery types, Las Flores
a-e, Las Flores painted and incised vase sherds; f-m, Upper Mayoid type
sherds; n, sculptured or molded Mayoid sherd.
PLATE 6
Upper Ulua Polychrome pottery types, Las Flores

a, human effigy (16.5 cm) ; b, vessel with “vestigial” spout (14 cm); ¢,
Upper Mayoid jar (7 cm); d, sculptured or molded Mayoid jar (6.5 cm) ;

e (5 cm), f (6.5 cm), jars of imitation Ulua marble bowl type.
PLATE 7
Ulua Polychrome, Bold Geometric pottery types, Santa Rita
a, Bold Geometric olla (12 cm) ; b (22 cm); ¢ (30 cm) ; d Lower Bold Geo-
metric olla type (30 cm) ; e, Bold Geometric tripod dish (7.5 cm) ; f, Bold
Geometric tripod dish (10 cm). (a-e, Santa Rita; f, Naranjo Chino.)
PLaTeE 8
Ulua Polychrome, Mayoid pottery types, Santa Rita
a, Lower Mayoid type vase (20 cm); b, Lower Mayoid type vase (20 cm) ;
deer effigy pot cover (19.5 cm); e, Mayoid type vase (17.8 cm) ; e, f, tripod
plate, type uncertain (14.5 cm).
PLATE 9
Ulua Bichrome sherds, deepest level, Santa Rita

a-j, various sherds; k, obsidian scraper; /, pottery stamp; m, obsidian flake
knife fragment; o-s, u-z, aa, bb, Usulatan ware sherds; ¢, Lower Mayoid
type sherds from just above sterile sand layer; cc, red-on-white sherd.

PLATE 10

Playa de los Muertos Bichrome sherds

a-h, polished orange-red to brown; i-n, polished dark gray to black; o-s,
polished slate-gray to buff. (Lower cultural horizon, Playa de los Muertos.)

PLATE II

Playa de los Muertos sherds and figurines

a-e, sherds with chalky white wash; f, g, k, red and black; i, 7, 0, red on
buff ; 4, unslipped brown and red; 1, m, red on white; n, gray on dull red;

p, polished brown face; q, r, s, polished figurines with white slip; ¢, 4, v,
solid brown figurines (lower cultural horizon, Playa de los Muertos).

NOI HONDURAS—STRONG, KIDDER, AND PAUL I29

PLATE 12
Yojoa Polychrome vessels, Mayoid types
a, excavation B, Los Naranios (25 cm) ; b, Aguacate (16.3 cm) ; ¢ (10.7 cm) ;
d (10.5 cm), Aguatal; e, Aguacate (9.5 cm) ; f, La Ceiba (12 cm).
PLATE 13
Yojoa Polychrome vessels

Bold Animalistic type: a, La Ceiba (15 cm); b, site 1, Los Naranjos (12.5
cm) ; c, Aguatal (10 cm) ; d, La Ceiba (11 cm); e, Effigy (type ?, compare
fig. 7, p, p. 52), La Ceiba (11 cm); f, Mayoid type, Aguatal (11.5 cm).

PLATE I4

Yojoa Polychrome vessels

a (10.7 cm); b (10.2 cm), Bold Animalistic type, Aguacate; c (8 cm); d (10
cm), Bold Geometric or Bold Animalistic types, Aguacate and Los Naranjos,
site 1; e, imitation Ulua marble bowl type, Aguacate (6.7 cm); f, carved
brown ware, Aguacate (7.3 cm); g, bowl with negative painting, Los
Naranjos, site 1, (6 cm); h, bird-shaped pot, Aguatal (7 cm).

PLATE I5

Early ceramic types at Lake Yojoa

a (11 cm); b (12cm), Playa de los Muertos Bichrome type (?), Los Naranjos
(exact provenience uncertain) ; c, d, f-i, k, 1, o-s, u-w, Yojoa “ Monochrome”
sherds; e, j, figurine fragments ; m, obsidian flake; n, t, ground stone artifacts.
(c-w, lower cultural horizon at Los Naranjos, site 1, and excavations A
and B.)

PLATE 16

Los Naranjos, Lake Yojoa

Fic. 1. Crude anthropomorphic statue.

Fic. 2. Stone serpent head.

Fic. 3. Stone torso and head.

Fic. 4. Mound 1, from the north near site 1.

Fic. 5. Section of trench at site 1, showing house floor and burial.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO eo iis O-line enn

VARIOUS CHAMELECON AND ULUA RIVER SITES

1, Ball court at Naco; 2, mound structure at Naco; 3, excavation at
Santa Rita (farm 17); 4, Tres Piedras site.
9

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97, NO. 1, PL. 3

U

NACcO SHERDS

b, Ulua Polychrome sherds at Naco.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO Oia INO)-mdige ant

NACO SHERDS AND ARTIFACTS

m, Spanish colonial sherd at Naco.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97, NO. 1, PL. 5

UPPER ULUA POLYCHROME POTTERY TYPES, LAS FLORES

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO Siig NOs bene

ae

UPPER ULUA POLYCHROME POTTERY TYPES, LAS FLORES

SMITHSONIAN MISCELLANEOUS COLLECTIONS Wolta Sry INE ap Ss 7

ULUA POLYCHROME, BOLD GEOMETRIC POTTERY TYPES. SANTA RITA
f, Naranjo Chino.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOTE OT INO medline

bh Riana eat ts

Si oe
Rttimisisieeet ce eon oe

ULUA POLYCHROME, MAYOID POTTERY TYPES, SANTA RITA

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOPR 7peNOw dpe Pleo

ULUA BICHROME SHERDS, DEEPEST LEVEL, SANTA RITA
t, Lower Mayoid sherds on sterile sand layer above Ulua Bichrome horizon.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO Eom NOmr leant O

p

PLAYA DE LOS MUERTOS BICHROME SHERDS

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLO INOm ainda

PLAYA DE LOS MUERTOS BICHROME SHERDS AND FIGURINES

SMITHSONIAN MISCELLANEOUS COLLECTIONS WAIL 75 INOS qo Velbe WE

YOJOA POLYCHROME VESSELS, MAYOID TYPES

SMITHSONIAN MISCELLANEOUS COLLECTIONS WAKES 765 INO i lel Gs as)

YOJOA POLYCHROME VESSELS, BOLD ANIMALISTIC TYPES

c, uncertain type; f, Mayoid type.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOM Si aaNOe dig Plea 4:

A

YOJOA POLYCHROME VESSELS, VARIOUS TYPES

a—d, Bold Animalistic and Bold Geometric; ¢, imitation Ulua marble bowl.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VON 975 INOj dy Pl. 45

EARLY CERAMIC TYPES AT LAKE YOJOA

a, b, Playa de los Muertos Bichrome (?) ; c-w, Yojoa ‘“ Monochrome.”

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO EO NOluiti ier iG

iS a

“~*
hk

LOS NARANJOS, LAKE YOJOA

I, crude human statue; 2, stone serpent head; 3, stone torso and head;
4, mound 1; 5, section of trench, site 1.

A ‘SMITHSONIAN, MISCELLANEOUS. COLLECTIONS
mri. _ VOLUME 97, NUMBER 2 is

_ WAVE- LENGTH BALANCE |

: . “(Wire Four PLATS)
Loss - ae uBy
OAM teh) BAR Se JOHNSTON
es Assistant Director, Division of Radiation and Organisms_
; _ Smithsonian Institution
“ (PUBLICATION 3446)
, i
| _ GITY OF WASHINGTON
‘PUBLISHED BY THE SMITHSONIAN INSTITUTION
. JANUARY 12, 1938

PLANT GROWTH IN RELATION TO

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER 2

PLANT GROWTH IN RELATION TO
WAVE-LENGTH BALANCE

(WiTH Four PLATES)

BY
EARL S. JOHNSTON

Assistant Director, Division of Radiation and Organisms
Smithsonian Institution

(PUBLICATION 3446)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 12, 1938

The Lord Galtimore Hress

BALTIMORE, MD., U. & A.

PLANT GROWTH IN RELATION TO WAVE-LENGTH
BALANCE

By EARL S. JOHNSTON
Assistant Director, Division of Radiation and Organisms, Smithsonian Institution

(With Four PLATEs)

INTRODUCTION

There can be little doubt that wave-length distribution exerts an
enormous influence on the growth of plants. Numerous experiments
show that stem elongation is greatly retarded under blue light, whereas
an acceleration takes place in the red and near infrared regions.
Chlorophyll production takes place better toward the red than toward
the blue end of the visible spectrum. Phototropic sensitivity is great-
est in the blue and zero in the red. For equal amounts of energy
falling on the leaf, two maximal regions of CO, absorption have been
found—one in the red, the other in the blue. It thus appears that a
wave-length region best suited to a given plant process may be entirely
without effect upon another.

In plant nutrition studies, experiments have shown that there is a
general balance in the proportionate amounts of mineral elements of
a nutrient solution that brings about a favorable growth response in
plants. Although there may be considerable latitude in the ratio of
-amounts of elements in such a solution, it may be said that a balanced
condition exists.

In a somewhat analogous manner, it is possible to think of the light
requirements of plants as a balanced condition of intensities of dif-
ferent wave lengths which bring about good plant growth. Both light
intensity and wave-length distribution vary to a considerable extent
over the earth’s surface. Likewise the character of the vegetation
varies. Since plants have been growing on the earth for countless
ages, it is reasonable to assume that their physiology is adjusted best
to sunlight. Although there is experimental evidence to show that
different processes go on better in some wave-length regions of the
spectrum than in others, yet the best growth, when all the processes
are considered simultaneously, apparently takes place in the natural

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97, No. 2

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

light of the sun. A direct experimental comparison between sunlight
and artificial light is, of course, difficult to make because of the great
number of variables entering into the problem.

Numerous experiments have been made for the purpose of growing
plants under artificial illumination. The object of many such experi-
ments was to find a satisfactory artificial light which could be operated
economically on a commercial scale. In other experiments the technical
and scientific aspects were the main objectives. So far as is known,
there is no available light source which is like that of the sun in its
wave-length distribution. Plants have been grown fairly successfully
in a few instances under well-controlled laboratory conditions, but
the problem is by no means solved. It may even be found that plants
can be grown normally under greatly reduced intensities of light pro-
vided a proper proportion between the intensities of its component
wave lengths is worked out.

The purpose of the present report is to discuss briefly some pre-
liminary experiments dealing with the question of a wave-length
balance of artificial light.

EXPERIMENTATION

In the experiments herein described, plants were grown between
two different light sources. Three or more types of lights could be
used, but for this preliminary survey it was thought best to limit the
wave-length distribution to two types. All the experiments were con-
ducted in a small room (approximately 15 x 10 ft. x 8 ft. high) the
walls and ceilings of which were painted a flat black to minimize
scattered light effects. Both temperature and humidity were auto-
matically controlled. The plants were grown in I-quart jars contain-
ing nutrient solution. Each culture was placed on a small rotating
table and usually grown for 3 weeks with a daily light period of 12
to 18 hours. By constantly rotating the plants (3.4 r.p.m.) on an axis
parallel to their stems, the phototropism of these stems was reduced
to zero. The leaves in some experiments showed phototropic response.
The wave-length distribution depended upon the light source. The
intensity was regulated largely by the distance the culture was placed
from the light.

In an earlier paper Johnston (1932) found that the excess of near
infrared of the Mazda lamp caused a distinct yellowing of tomato
leaves. If this region of the spectrum was not actually destructive to
chlorophyll,.it was of little or no benefit to its formation. It would
thus appear that more nearly normal color could be obtained by re-

NO. 2 PLANT GROWTH AND WAVE-LENGTH BALANCE—JOHNSTON 3

ducing the infrared or by increasing the intensity of the rest of the
spectrum. An experiment was therefore planned in which this was
partially accomplished by building up the: blue end of the spectrum.

Experiment 1—Two 1,000-watt projection Mazda lamps (115 v.)
were placed I meter apart. Surrounding each lamp was a clear Pyrex
thermos bottle blank fitted with a water inlet and outlet. The radiation
of each lamp was thus filtered through 5 mm of water. The constant
flow of water through this jacket was a great aid in maintaining a
constant temperature condition in the room, since a great deal of heat
was thus removed.

A copper sulphate (sp. gr. 1.08, about 8 percent) filter (6 cm thick)
was placed in front of one of the lamps. This was the added blue
light source. The individual rotating plant cultures were located
at positions relative to these two light sources which gave the intensity
values expressed as watts/cm’ in table 1.

TABLE 1.—Radiation intensities and plant data from experiment 1

Light intensity

watts/cm2 Stem Total
Culture —----- tO ht. dry wt.

no. White Blue Total cm gram
Ler stevens ose .0396 .0006 .0402 5.0 .086

Pig pee re eer .0285 .OO10 .0295 aT .102

BERS eras existe .0166 .OO14 .0180 6.1 .043

AMR cr ee Sai 8 .0064 .0027 OOO 6.3 025

See eyelets .0046 - .0054 .O100 3.8 O19

Marglobe tomato seeds were sprouted between moist filter paper
at a temperature of 25° C. for 3 days. The sprouted seeds were then
transferred to a germination net, and after about a week of growth
five similar seedlings were selected and set out in quart jars, one per
jar, and placed on the five small rotating tables. After 2 weeks of
growth these plants were measured and dried in an oven at 103° C.
to obtain the dry weight. These data are also shown in table 1.

Because of the meagerness of data, no definite conclusions can be
drawn. The first three plants were heavier than similar ones grown
in the north and south laboratory windows. Although the total in-
tensity of no. I was greatest, yet maximum dry weight occurred in
plant no. 2. Here the added blue radiation was about 3 percent of
the total as compared to 1.5 percent in plant no. 1, which was yellow-
green in color. Plant no. 2 was a light green when compared to plants
3, 4, and 5, whose percentage of added blue to total radiation were
respectively 8, 30, and 54.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Experiment 2—The next experiment was very similar to that just
described. Here again individual variation was too great to draw any
accurate conclusions.

Experiment 3.—In the next experiment three duplicate sets of
tomato plants were grown under three different sets of light condi-
tions. In front of one lamp a Corning heat-absorbing 212 percent
red filter was placed. In front of the other lamp a filter jar containing
a M/2 CuSO, solution was placed. Both filters cut off at 6040 A, the
CuSO, solution transmitting light of shorter wave length and the
Corning filter transmitting light of longer wave length. Two duplicate
sets of cultures were placed between these filtered light sources. A
third set was located to the rear of the blue filter light in such a position
that the plants received only the full Mazda spectrum. Intensities
were measured at the beginning and at the end of the experiment.
These average values, together with the plant data for 3 weeks’
growth are given in table 2.

TABLE 2.—Fadiation intensities and average plant data from experiment 3

aes A Average data per plant
Radiation intensity

watts/cm? Stem Root Total

Culture (oo ht. length dry wt.
nos. Red Blue White Total cm cm gram
Tecate es ane .0055 .0022 aR .0057 6.3 38 .026
Sandys sere .0028 .OOIL ese .0039 7.4 50 .038
iS Ehitel Wonoaccce Synane eee .0056 .0056 8.0 44 .028

The greatest amount of dry weight was produced by cultures 3 and
4, although the total light intensity was less than under the other two
conditions of growth. Here the blue radiation was about 28 percent
of the total. Although these data are meager, there is an indication
that considerable differences in growth are obtained by manipulating
the wave-length distribution as well as the total intensity.

Experiment 4.—In the next experiment wave-length distribution
was further restricted by using neon and mercury grids as light
sources. These were constructed in our laboratory by Mr. L. B.
Clark. In order to increase the intensities a mirror was placed back
of each. Three duplicate cultures were placed between these two
light sources, each culture jar containing three tomato seedlings.
This increased the number of plants per treatment to six. Because
of reflections in the mirrors some red light came from the blue side
and some blue light came from the red side of the cultures. As will
be seen in table 3, the intensity of radiation was considerably less
than in the earlier experiments. The plants were grown for 26 days

NO. 2 PLANT GROWTH AND WAVE-LENGTH BALANCE—JOHNSTON 5

and then harvested. The average stem height and total dry weight
per plant for each of the three light conditions appear in the same table.

Although the stems of plants in group 1-2 were thicker than those
in the other groups, their leaves were quite yellow. Here again yel-
lowing is associated with energy distribution where the greatest
amount is found in the red end of the spectrum. The plants in group
5-6 had the best color, even though the total amount of energy was
about half that of group 1-2. These leaves had flat smooth surfaces,
while those in group I-2 were quite pointed and curled. The general
appearance of these plants is shown in plate 1. After another experi-
ment with these lamps it was definitely indicated that the plants were
getting insufficient illumination.

Experiment 5.—To increase the radiation, a General Electric 400-
watt high-pressure mercury lamp was substituted for the mercury

TABLE 3.—Radiation intensities and average plant data from experiment 4

Radiation intensity Average data
at beginning of experiment per plant
watts/cm2 ——

Stem Total

Culture Neon Mercury ht. dry wt.
no. (red) (blue) Total cm gram
Deie.s ais atee .0002 .00002 .00025 6.6 .029
De Nein hse .0OOI9 .00002 .0002I 6.6 .029
Ber Reet aoieor .00008 .00002 .OOOLO 3.6 .OI4
AY isa ae .00008 .00002 .0OO1O 3.6 O14
Dicaswsien et ace ies’ .00005 .00007 .OOOI2 3.4 O13
eran .00005 .00007 00012 3.4 .013

grid and four instead of two transformers were used with the neon
lamp. The daily light period was increased from 12 hours to 18
hours. Because of the marked decrease in the life of the neon lamp
under these forced conditions, the experiment was discontinued at
the end of 20 days. In this exploratory experiment no accurate in-
tensity measurements were made. However, general improvement in
growth was noted.

Experiment 6.—To increase further the light intensity, a 1,000-
watt, 110-volt projection lamp housed in a water jacket as noted
earlier (experiment 1) was substituted for the neon grid lamp. Three
plants per quart culture jar were used and the cultures run in dupli-
cate so far as the light relations were concerned. Throughout all
the previous experiments the plants were grown in a three-salt
nutrient solution similar to that used by Johnston and Dore (1929).
In this experiment cultures 2, 4, and 6 had (NH,)2SO, added to the
former solution which contained Ca(NO;).2, MgSO,, and KH,PO,

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

and traces of Mn, B. Iron was added as FeSO, to all cultures from
time to time as conditions demanded. Because the Mazda lamp was
run at about its voltage limit its life was short, and replacements were
necessary every 6 or 7 days. The plants were grown for 3 weeks with
a daily light period of 18 hours. The added heat from the lamps
caused a slight daily temperature fluctuation. The average maximum
was 24° C. and the average minimum 21.5°C. This resulted in a
change in humidity which averaged 57 and 51 percent for the dark
and light periods respectively. As found in previous experiments, a
temperature fluctuation is beneficial to the tomato plant. Better
growth was obtained by subjecting the plants to a lower dark period

TasLe 4.—Radiation measurements at beginning of experiment 6

Foot-candles with

Watts/cm?2 small G. E. meter
Culture a M@M@V[{T_-
nos. Mazda Mercury Total Mazda Mercury Total
Titanic 2etetiyan« .0404 .0O13 .0417 2,800 * 200 3,000
A euil Mecowooc .0172 .0031 .0203 1,200 600 1,800
iG amduOaere cir .0005 .0067 .0132 550 1,000 1,550

TABLE 5.—Plant data from experiment 6 expressed as averages per plant

Dry wt.
Stem Green wt. gram

Culture ht. grams ——_—_—_——.
no, cm Tops Tops Roots Total

Tris a.aleyevevstaneieiete 175 6.6 520 .139 .668

Di scisterevoyegevegerone 21.1 7.8 .671 174 845
Bivona ees 18.9 5.6 403 .OOI 554
Alwaatagniue aerate 20.9 6.1 .469 .074 543
Reese ete 23.5 5.0 384 .074 458

OA oss 23.8 5.4 371 .057 .428

temperature (about 3° C. lower) than by maintaining a constant
temperature during the dark and light periods.

The intensity measurements which were made at the beginning of
the experiment are presented in table 4.

After 3 weeks of growth the plants were photographed (pl. 2) and
harvested. Data giving average stem height, green weight of tops, and
dry weight of tops and roots are given in table 5.

Both the illustrations and plant data show that this group of plants
was normal in appearance and comparable to good greenhouse plants.
It was by far the best we have grown under the 100 percent artificial
conditions of our laboratory. In an earlier publication, Johnston
(1932) reported that tomato plants exposed to an intense illumination
from a Mazda lamp grew very well but soon became yellow in color.

NO. 2 PLANT GROWTH AND WAVE-LENGTH BALANCE—JOHNSTON a

The near infrared radiation was apparently destructive to chlorophyll
or inhibited its formation. This again appeared to be the case for the
three plants in culture 1. However, one of the most interesting ob-
servations made in this experiment was that the color of the plants in
culture 2, which received the same radiation intensity as those of
no. I, was much greener. This color difference is seen to some extent
in plate 2 as differences in light and dark tones of the plants in the
upper and lower figures. This was also true for the plants in cul-
tures 4 and 6, as compared with cultures 3 and 5 respectively, which
were grown under similar light conditions. All the plants grown in
nutrient solution to which (NH,).,SO, had been added were greener
than the corresponding ones without this additional nitrogen. This
observation suggests the influence of the type of radiation on the up-
take of mineral nutrients. This same solution without the (NH,).SO,
has been used in growing tomato plants in the greenhouse but the
characteristic chlorotic effects were not noted until the plants were
grown under Mazda lamps.

The percentages of added mercury radiation to total illumination
were 3, 15, and 50 respectively for cultures 1-2, 3-4, 5-6. The green
color of the leaves was deeper where this percentage was larger. A
more striking color difference occurred, however, between the plants
in cultures with and without the (NH,).SO,.

The average total dry weight per plant for each of the three light
conditions 1-2, 3-4, 5-6 was .757, .549, and .443 gram respectively.
Under these three light conditions the efficiency in the production of
dry weight per watt/cm* was 18, 27, and 34 respectively. Although
the total intensity of 5-6 was about a third that of I-2, on the basis
of efficiency in producing dry weight per unit energy, the less intense
radiation was about double that of the more intense.

One other factor in addition to wave-length distribution must be
recognized in an experiment of this type. One lamp (Mazda) gave
practically continuous illumination; the other (mercury, 60-cycle), a
fluctuating illumination varying from a minimum considerably below
the average to a maximum much greater than the average as de-
termined by the thermocouple and photoelectric cell. McAlister
(1937) clearly shows that a change in efficiency of carbon dioxide
assimilation occurs with frequency of intermittency of illumination.
Although it may be comparatively safe to compare the different cul-
tures in any one experiment since the “ flicker” effect is doubtless
the same, it is impossible to compare results of experiments in which
the light is continuous with those in which it is intermittent or with
those in which it is half continuous and half intermittent.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. 97

Experiment 7.—An experiment varying a little from the one just
described was next performed. In this, five cultures of tomato seed-
lings were placed around the Mazda lamp at positions which gave
them approximately equal light intensities from this lamp. A sixth
culture was located at a position where the Mazda intensity was about

half that of the other cultures.

lamps for each culture of these plants are best seen in table 6.
After 18 days the plants were harvested and their dry weights
determined. The plant data appear in tables 7 and 8.

TABLE 6.—Radiation measurements* from experiment 7

Culture
no. Mazda
Lote ye eee 0234
Dede A Net ae 0241
EB onahcde alors ee 0241
Alsrotwretatsgute vate 0240
sere terstalot eta sai 0231
Oana O1I2

Watts/cm?
Hg
.0097
.oo18
.0013
.0013

.0039
.0090

Total

0331
.0259
.0254
.0253
.0270
-0202

Foot-candles with
small G. E. meter

Mazda
1,300
1,300
1,400
1,400
1,300
600

Hg
2,000
300
200
200
600
1,900

The intensities of each of the two

Total

3,300
1,600
1,600
1,700
1,900
2,500

1 Since the original Mazda lamp was replaced after 6 days, these measurements were made
on the second lamp on the 1oth day of the experiment.

Plant

Av

Avy.
Av.

TABLE 7.—Stem height (cm) data from experiment 7

Culture number

3
19.0
15.5
19.0
17.8

2.2

15.6

Dry wt.
Roots
-199

-044
.060

.004
.099

4
18.5
22.3
20.0

20.3
1.6

18.7

5
22.2
22.5

~ 20.6

21.8
1.9

19.9

19.8
19.5
18.5

19.3
1.8

17.5

Total
.QI12
355
.451
531
681

I 2
Da Grae Pae TTA rche eaten ee eerie 16.5 16.2
Be eae iar ents Le oto eo Rinteey 15.8 18.0
By is Siva Us bey Wi aE PERS recent ae 14.5 10.4
Auhits tathanvieSts waiter ersctorainete 15.6 17.9
Original hiss oseesic wae eeear 2:3 iT
DOVES iol Mites ao sonodamoec mae 15.8
TasLe 8.—Average green and dry weights (grams) of plants from
experiment 7
Culture Green wt.
no. Tops Tops
Diacwahae’s cote 6.1 1s
DRAM 4.2 Pair
QURASEA scorsrere 4.8 391
ASA nb geees 5.8 .407
ee ciate peepee 6.6 582
Game cetaoas 6.3 .610

145

755

NO.2 PLANT GROWTH AND WAVE-LENGTH BALANCE—JOHNSTON 9

During the experiment water was added and fresh nutrient solution
renewed as required. Because of frequent stopping of rotating table
no. I, these plants were slightly burned. These plants had the shortest
internodes. Plants in culture 6 had next to the shortest internodes
and were the best green. The leaves of plants in cultures 2, 3, 4, and
5 were slightly chlorotic. In order of their dry weights, plants in
cultures 1, 6, and 5 were the best. It is interesting to compare the
total dry weight per unit total energy with the percentage of energy
received from the mercury lamp (table 9).

TABLE 9.—Comparison of dry weight efficiency with amount of radiation from
the mercury lamp

Cultures ; I 2 3 4 5 6
Ratio total dry wt. to. watts/em™...........: 27 OS aE Oue 2 ONE2 52s 7d
Percentage radiation from mercury lamp.... 29 7 5 5 14 45

Plants in cultures 6, 1, and 5 produced the greatest amount of dry
weight per watt/cm’. These same cultures in the order given received
the largest percentages of radiant energy rich in the blue. Total
energy (table 6) was greatest for culture 1 and least for culture 6.
Cultures 2, 3, and 4 were practically equal. Thus, plants of culture 1
had the greatest total dry weight, and those of 6 were second. How-
ever, for greatest efficiency in the production of dry weight, plants
in culture 6 were much better than those in culture 1. This is evi-
dently related to the greater percentage of shorter wave length in
the one case than in the other. When light intensity as measured
by the foot-candle meter is considered, plants of culture 1 are shown
as receiving the greatest amount of light and those of culture 2 the
next greatest amount.

By consulting the table of stem heights it will be noted that the
average height at harvest for plants in culture 1 was less than any of
the other groups although the average original height was greatest.
The least average stem elongation shown by this group may be cor-
related with the greatest amount of total energy received by these
plants. . But little difference in stem height is seen between plants
of the other cultures. Likewise there is but little difference in total
energy received by these same cultures. Other observations bear out
this same point that an intense light retards stem elongation more
than a less intense one. Although the shorter wave lengths have a
greater retarding effect, this difference between plants of cultures 6
and I must have been offset by the differences in total radiation
intensities.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

Experiment §—In the last experiment of this series, the same types
of lamps were used. Also each culture contained three tomato plants.
The first four cultures (nos. I, 2, 3, 4) were arranged around the
mercury lamp at approximately equal distances. The other two (nos.
5 and 6), together with those numbered 3 and 4, were located about
equal distances from the water-cooled 115-watt Mazda projection
lamp. The intensity measurements taken at the beginning of the
experiment are shown in table Io.

TABLE 10.—Kadiation measurements in experiment 8

Intensity measurements

Watts/cm? Foot-candles

Culture aaa LER Pe a ag oo ee
no. Mazda Hg Total Mazda Hg Total
Taree e .0O7I .0048 .O11Q 350 700 1,050

DB ierees .007I .0047 .O118 350 700 I,050

iB Npaaa ate .O140 .0046 .0186 800 700 1,500

AUN ero letere .O144 .0045 .0189 800 700 1,500
Cisenten .0137 .0O1O .O147 900 200 1,100
Oxenee .O146 .00OI0 * .0156 900 200 1,100

At the end of three weeks the plants were photographed and har-
vested. Since each culture of three plants was duplicated, the average
of the six plants is shown in table 11.

TABLE 11.—Plant data from experiment 8 expressed as averages per plant

Plant data expressed as averages per plant
po

Total
transpira-
Final Green wt. tional Dry weight (g)

Culture stem ht. No. of of tops water loss
nos. (cm) leaves (zg) (ml) Tops Roots Total
eae EUS 6 a5 157 .240 .042 282
Brandl Ae ee 20:0 7 6.1 251 437 .0907 534
amd Olean sO 5 2.6 08 .140 .021 161

Plants of cultures 3-4 were best in general appearance and had the
thickest stems. Those of cultures 5-6 were lightest green. Plants
with longest roots were found in cultures 5-6; those with shortest
roots occurred in cultures 1-2.

The general appearance of the cultures about the two lamps in this
experiment may be seen in plate 3, and the appearance of the tops
and roots of the plants at the end of the experiment is seen in plate 4.

It will be recalled that in experiment 6 the ratio of dry weight to
total energy increased with the percentage of added radiation from
the mercury lamp. Also in experiment 7, table 9, the three cultures

NO. 2 PLANT GROWTH AND WAVE-LENGTH BALANCE—JOHNSTON II

in which the greatest dry weight was produced per watt/cm* were the
same three cultures which received the greatest percentage of radia-
tion from the mercury lamp. In experiment 8, however, an exception
occurred. The dry weight efficiencies for the three groups of cultures
I-2, 3-4, 5-6, were 23.8, 28.6, 10.7 respectively, while the percentages
of total radiation attributed to the mercury lamp were 40, 25, and 7
for these same cultures in the order given. It is not clear from the
data at hand why this exception occurred.

In order to compare all these data which are fairly comparable,
table 12 has been constructed. Since two types of solutions were used
in experiment 6, cultures 2, 4, and 6 were selected as their solutions

TABLE 12.—Average dry weight production per unit total radiation (Mazda plus
mercury lamp) in relation to percentage of radiation from the mercury lamp

Percentage
Experiment Culture Dry wt. per radiation from
number number watt/cm? mercury lamp
FP APRS SAILS eis misters ie 6 a7 An 45
Be Ae a eral Sante neta Baiaicleis 3 227 25
OPE cere tna oe tisl es veais, sre wes 6 32.4 51
SEs Ano Saiaatelanes Gesiheceteais I 28.2 40
Tees Mart Sale oNe asta arbre tlw ah I 27.6 20
Oye cree rere cra lereibin s Gis sre one 4 26.8 15
Phe Seid Na ier he Distancia a. Ss 5 25.2 14
Sareea Serene ceeeve etoile cvereiare 4 24.4 24
PPE PERS ASS OMe it oso o2 4 21.0 5
(Detehevaterapaeyersie Srexd toy </b. oe ore wisi tox 2 20.3 3
Be at oP ena a EN oe ai ate 2 19.4 40
Nase eran nay crevie aie" tr etsy Saevs (32% 3 17.8 5
isiatate es WN ofrecer s ueter sis) ecaisters 2 Nei a
SPE ert fa aiiand sigla rane bie er csakene 6 11.0 6
OB eretavsnrusteitera cia a cise: clbisue ents 5 10.3 7

were similar to those used in experiments 7 and 8. All plant values
given in this table are the averages of three plants. There is a slight
difference in the duration of the three experiments which should be
kept in mind in making this comparison. In these experiments, 6, 7,
and 8, the plants were grown for 20, 18, and 21 days respectively.
The data showing dry weight produced per unit total energy in
table 12 have been arranged from greatest to least value. The cor-
responding values showing the percentages of total radiation that are
obtained from the mercury lamp fall roughly into two groups. The
first eight values are high (14 to 51 percent). The remaining seven
with the exception of culture 2 in experiment 8 are low (3 to 7
percent). Although there is no regular decrease in these percentage
values with the decrease in dry weight per unit total radiation, there

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

appears to be a general decrease in dry weight efficiency with illumi-
nation containing less of the shorter wave lengths found in the
mercury lamp.

DISCUSSION

Plants have been grown by Harvey (1922), Hendricks and Harvey
(1924), and others under Mazda lamps. Davis and Hoagland (1928),
Arthur, Guthrie and Newell (1930), Garner and Allard (1931),
Steinberg and Garner (1936), and others have conducted numerous
experiments in which good growth was obtained with Mazda lamps
for various lengths of daily light and dark periods. Many other in-
vestigators both in Europe and in this country have shown that plants
may be grown in artificial light whose wave-length distribution is
continuous from blue-violet to red. Other investigators have de-
termined the growth of plants in different portions of the spectrum.
Here it was necessary to use glass or liquid filters. Others, like
Roodenburg (1932), have used gaseous discharge lamps such as
neon. Most of these experiments indicate the necessity of the full
visible spectrum for normal growth. Popp’s (1926) results indicate
that the blue-violet end of the spectrum is necessary for normal
growth although the ultraviolet may not be indispensable. Shirley
(1929) states that “ The entire visible and ultra-violet solar spectrum
is more efficient for the growth of the plants studied than any portion
of it used; the blue region of the spectrum is more efficient than the
red region.” Schappelle (1936) concluded that white light is best for
normal plant response. Either end of the visible spectrum without
the other causes abnormal growth. Infrared, between 0.8 » and 2.0 pw
was ineffective in bringing about fruiting of Marchantia, while red
and blue lights were of approximately equal effectiveness.

Arthur and Stewart (1935) made a comparison of the growth of
buckwheat plants under Mazda, neon, sodium, and mercury vapor
lamps. For short periods of 8 to 10 days the sodium lamp was found
to be most efficient in the production of dry weight. No relation was
found between the absorption bands of chlorophyll and the emission
bands of the various lamps. These gaseous discharge lamps produced
plants with greener leaves than the Mazda lamps. Later Arthur
and Harvill (1937) show that the sodium lamp alone is not ideal for
the continuous growth of plants over long periods of time. If, how-
ever, the continuous exposure from the sodium lamps is supplemented
by an exposure of 2 hours per day from an 85-watt capillary mercury
vapor lamp, excellent leaf color and flowering could be produced in

NO.2 PLANT GROWTH AND WAVE-LENGTH BALANCE—JOHNSTON 13

such plants as begonia, gardenia, cotton, geranium, buckwheat, and
snapdragon. Although this light source was not satisfactory for the
tomato plant, the authors point out that other wave bands of light
may be found which should be added or subtracted for the best
growth of some plants such as the tomato.

Dastur and Mehta (1935) determined the rate of photosynthesis in
approximately equal intensities of red, blue, and white light. Photo-
synthetic activity was greatest in the white light, intermediate in the
red light, and least in the blue light. They state that both the red and
blue regions are necessary for normal photosynthesis.

Equally interesting are the results of Hoover’s (1937) investigation
on determining the rate of CO, absorption as a function of wave
length on the basis of equal incident energy. The principal maximum
occurred at 6500 A in the red, and a secondary maximum came at
4400 A in the blue. The greater transmission and reflection of radia-
tion in the green region decreased the effectiveness in that portion
of the spectrum. The limits of CO, absorption were placed between
7200 A and 7500 A in the red, and below 3650 A in the blue end of
the spectrum.

Dastur and Solomon (1937) show the importance of the blue-
violet end of the spectrum in photosynthesis in a series of experi-
ments in which plants are grown in the light of a carbon arc, in
“mixed” light where the gas-filled electric lamp light has super-
imposed upon it a beam of blue-violet light, and in the light of the
gas-filled electric lamp alone. The “mixed” light was composed of
two beams originating in a single source (1,000-watt flood lamp) and
reflected to the plant by mirrors. One beam was passed through a
copper sulphate filter which limited the wave-length band to the
region 4200A to 4720A. These beams (white and blue) were
reunited in the proportion I: 1 on an intensity basis. Plants grown
in these three lights showed greatest photosynthetic activity in the
carbon arc light, intermediate in the “ mixed” light, and least in the
gas-filled electric bulb light. This follows the order of richness in
blue-violet light of the three sources.

From the foregoing discussion it would appear that plants can be
grown in artificial light, but for more or less normal growth the light
should include those wave lengths found in the visible solar spectrum.
An increase in intensity or the absence of a given portion of this
spectrum brings about abnormal growth responses. Undoubtedly,
the more nearly the artificial light resembles sunlight in its energy
distribution, the more nearly normal are the plant growth responses.

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

In the experiments reported in the present paper, a method for
mixing artificial lights was used, but one quite different from that
used by the above-mentioned investigators. A beginning was made
by using two light sources, one rich in red, the other rich in blue
light. By locating the plants on small rotating tables at different
distances from these light sources, practically any intensity ratio of
the two could be obtained. With this general scheme the number of
lights could be increased, thus making it possible to study the effects
of any given mixture of restricted wave-length regions on the growth
of plants. With each added light, however, the interpretation of
data becomes more difficult. By using this method it is difficult to
grow many duplicate individuals at one time, especially if they grow
large. This objection may be met in part by repeating an experiment
often enough to obtain more reliable statistical data.

The first two experiments with the Mazda light vs. the Mazda light
filtered through a CuSO, solution were mostly exploratory in nature.
There is some indication that the greatest dry weight produced is
associated with wave-length distribution and not entirely correlated
with intensity of radiation. Although the data of experiment 3 are
meager, a considerable difference in growth was obtained between
plants receiving different amounts of red (wave lengths longer than
6040 A) and blue (wave lengths shorter than 6040 A). The dry
weight increase for the plants receiving red-blue light in the ratio
72:28 was about 40 percent over those receiving white light and
those receiving a mixture in the proportion 96: 4, although the total
intensities of these two cultures were over 40 percent greater.

An attempt was made to change the type of red and blue light by
‘the use of neon and mercury grids. In these experiments (nos. 4 and
5) it was found that the intensity of radiation was too low for good
growth. This made it impossible to draw any definite conclusion
regarding the proportion of red to blue that gave best growth. Yellow-
ing or lack of greenness was associated with those light mixtures
predominant in red.

In order to obtain lights of higher intensities, one rich in red, the
other rich in blue, the water-jacketed projection Mazda lamp used in
experiments I and 2 and the 400-watt high-pressure mercury lamp
used in experiment 5 were employed. With this combination of lights
very good growth was obtained under 100 percent artificial conditions.
Because of this good growth and the increased number of plants per
treatment, more weight can be attached to the data from experiments
6, 7, and 8, than to the earlier ones. Where light and not carbon

NO.2 PLANT GROWTH AND WAVE-LENGTH BALANCE—JOHNSTON I5

dioxide is the limiting factor, the-dry weight increases with increased
illumination. Hoover, Johnston, and Brackett (1933), working with
wheat plants, found that in normal air CO, became limiting at a light
intensity of about 0.05 to 0.06 watts/cm’. In none of these experi-
ments with the tomato plant was the intensity greater than these
values. Although the two plants may not behave exactly alike, it is
reasonable to suppose they are similar enough to assume that at no
time was CO, the limiting growth factor. In order to accentuate
_ growth differences due to wave-length mixtures and minimize the
effect of intensity on dry weight production, the dry weight data
were divided by watts/cm*. This dry weight efficiency of comparable
cultures in the last three experiments was used as a criterion of the
effect short-wave (blue) radiation added to that of longer wave length
had on plant growth. It would appear from the data given in table
12 that a greater amount of dry weight is produced with a Mazda
light by enriching it with blue from a mercury lamp to the extent of
14 to 51 percent under the conditions of these experiments. Care
should be exercised in drawing any far-reaching conclusions, for with
a change in quality or wave-length distribution of the Mazda or other
source rich in red, changes undoubtedly will be necessary in other
portions of the spectrum. Although for good growth plants very
probably tolerate a rather wide range in wave-length distribution, yet
it would appear that the more neariy this distribution in artificial
light approaches that of sunlight the better will the plants grow.

SUMMARY

Emphasis is placed on the importance of quality or wave-length
distribution of light in affecting plant growth. A method and several
experiments are described in which plants were grown in “ mixed ”
lights. By placing the plants on small rotating tables between two
light sources, one rich in red, the other rich in blue, the proportion
of each type of radiation falling on each culture was varied by the
position of the culture with reference to the light sources.

As found in previous experiments, yellowing of leaves occurred
in light rich in near infrared. Since this trouble could be corrected
to a considerable extent by the type of nutrient solution used, it
indicates the importance of wave-length distribution on the uptake
of mineral nutrients.

Excellent growth under entirely artificial conditions was obtained
with plants grown between a 1,000-watt, water-jacketed, projection
Mazda lamp and a 400-watt, high-pressure mercury lamp. The posi-

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

tions of the plants for good growth were such that from 14 to 51
percent of the total radiation falling on them came from the mercury
lamp. In several cases better growth was attained in one mixture of
wave-lengths than in another where the total intensity was higher.
However, the relatively high growth efficiency may in part be due to
an intermittency effect occurring in gaseous discharge tubes such as
the mercury lamp here used.

LITERATURE CITED

ArtHurR, Joun M., GutuHrir, Joun D., and NEweEL, Joun M.
1930. Some effects of artificial climates on the growth and chemical com-
position of plants. Amer. Journ. Bot., vol. 17, pp. 416-482.
ARTHUR, JoHN M., and Harvitt, Epwarp K.
1937. Plant growth under continuous illumination from sodium vapor lamps
supplemented by mercury arc lamps. Contr. Boyce Thompson Inst.,
vol. 8, no. 5, pp. 433-443.
ARTHUR, JOHN M., and Stewart, W. D.
1935. Relative growth and dry weight production of plant tissue under
Mazda, neon, sodium and mercury vapor lamps. Contr. Boyce
Thompson Inst., vol. 7, no. 2, pp. 119-130.
Dastur, R. H., and Menta, R. J.
1935. The study of the effect of blue-violet rays on photosynthesis. Ann.
Bot., vol. 49, no. 196, pp. 809-821.

Dastur, R. H., and Sotomon, S.
1937. A study of the effect of blue-violet rays on the formation of carbo-
hydrates in leaves. Ann. Bot., n. s., vol. I, no. I, pp. 147-152.
Davis, A. R., and Hoacianp, D. R.
1928. An apparatus for the growth of plants in a controlled environment.
Plant Physiol., vol. 3, no. 3, pp. 277-292.
GARNER, W. W., and ALLarp, H. A.
1931. Effect of abnormally long and short alternations of light and darkness
on growth and development of plants. Journ. Agr. Res., vol. 42,
no. 10, pp. 645-651.

Harvey, R. B.
1922. Growth of plants in artificial light. Bot. Gaz., vol. 74, no. 4, pp.
447-451.

HEnprIcKs, Esten, and Harvey, R. B.
1924. Growth of plants in artificial light required for blooming. Bot. Gaz.,
vol. 77, Pp. 330-334.
Hoover, W. H.
1937. The dependence of carbon dioxide assimilation in a higher plant on
wave length of radiation. Smithsonian Misc. Coll., vol. 95, no. 21,
pp. I-13.
Hoover, W. H., Jounston, Eart S., and Brackett, F. S.
1933. Carbon dioxide assimilation in a higher plant. Smithsonian Misc.
Coll., vol. 87, no. 16, pp. I-19.

NO.2 PLANT GROWTH AND WAVE-LENGTH BALANCE—JOHNSTON E7

JOHNSTON, Eart S.
1932. The functions of radiation in the physiology of plants. II. Some
effects of near infra-red radiation on plants. Smithsonian Misc.
Coll., vol. 87, no. 14, pp. I-15.
JoHNSTON, EArt S., and Dore, W. H.
1929. The influence of boron on the chemical composition and growth of
the tomato plant. Plant Physiol., vol. 4, pp. 31-62.
McAttsTER, E. D.
1937. Time course of photosynthesis for a higher plant. Smithsonian Misc.
Coll., vol. 95, no. 24, pp. I-17.
Popp, HENry WILLIAM.
1926. A physiological study of the effect of light of various ranges of
wave length on the growth of plants. Amer. Journ. Bot., vol. 13,
pp. 706-736
Roopensure, J. W. M.
1932. Kunstlichtcultuur. II. Over de noodzakelijke van planten en neonbe-
lichting bij bloemcultures. Med. Wagen. (Nederland), vol. 36,
no. 2, pp. I-37.
SCHAPPELLE, N. A.
1929. Effect of narrow ranges of wave-lengths of radiant energy, and other
factors, on the reproductive growth of long-day and short-day
plants. Cornell Univ. Agr. Exp. Stat. Mem. 185, pp. 1-33.
SHIRLEY, Harpy L.
1929. The influence of light intensity and light quality upon the growth
of plants. Amer. Journ. Bot., vol. 16, pp. 354-390.
STEINBERG, Ropert A., and GARNER, W. W.
1936. Response of certain plants to length of day and temperature under
controlled conditions. Journ. Agr. Res., vol. 52, no. 12, pp. 943-960.

EXPLANATION OF PLATES

PLATE I

Tomato plants grown for 26 days under the following intensities (watts/cm?).
Daily illumination was 12 hours.

Culture Neon Mercury
Le mecehenensncucneret ola velciedelchare ce te -00023 -O0002
Bi eset eae ac vetiane Gu Manaxe .00008 .00002
RR Tercera eae ete .00005 .00007
PLATE 2

Tomato plants grown for 21 days under the following intensities (watts/cm’).
Daily illumination was 18 hours.

Mazda Mercury
Culture (water-cooled) (400 watt)
TetATAG a2 arcoe te betctn acre ravenna re .0404 .0013
Be AN Are aye sys hale cine ster eae .0172 .0031
Sera Olesen araysvamitareronaare .0065 .0067

The darker green leaves in cultures 2, 4, and 6, due to the added (NH,)SOu,,
appear in the illustrations as a deeper shade than those in cultures 1, 3, and 5.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

PLATE 3

General arrangement of cultures in experiment 8 on rotating tables placed
about the two light sources. The Mazda lamp encased in a water jacket is on
the left and the 400-watt mercury lamp on the right. The small rotating tables
turned at the rate of 3.4 r.pm. This prevented phototropic curvature of the
stems but not of the leaves which, although turgid, appear wilted.

PLATE 4

Tomato plants grown for 21 days under the following intensities (watts/cm’).
Daily illumination was 18 hours.

Mazda Mercury
Culture (water-cooled) (400 watts)
Dosen ciate outers eee eae .0071 .0048
DN te Aa aN hen PRED tes .0071 .0047
Biidvenanie oa din apa rene .O140 .0046
Ale lie Set Re she Narcan 0144 0045
Ba SR cr a en aiken .0137 .OO10

Gis State aekoe vo cremieere .0146 .OO10

(‘41 pue F sased aas ‘uoreurldxe 10,7)

SdNVT]1 NOAN OGNV
AYNDYAW NSEaMLAG NMOUD ‘pb LNAWIYNSadxXy NI SLNV1I1d OLVNOL AO SAONVYVAddY IWHANAD

14a ‘o> «ma 'ye CIMA CAIALI ABAMAN AQHMBAIWIVIIMACIIAL AIWIAIOCW TTIAIC

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97, NO. 2, PL. 2

GENERAL APPEARANCE OF PLANTS IN EXPERIMENT 6, GROWN BETWEEN
MAZDA AND MERCURY LAMPS

(For explanation, see pages 5 and 17.)

‘QT puke OF Sased as ‘uoreueldxo 30,
3 I A

SA9YHNOS LHSIT OML AHL WOUA SAONVLSIG
IN3e8Ss4I0 LY GS90V1d ‘SSA1IESVL DONILVLOY NO ‘8 LNAWIYSsdxXy NI SAYNLAIND AO LNAWAONVYYY IWYANAD

ao em 9 bs eprat t+ Rr *7A ra ee pees ee ee ee ee a ree ee oe erry oe

(‘gI pue OF sased 90s ‘uo1yzeue[dxa 104, )

8 LNAWIedadxZ AO NOISNADNOD SHL LV SLNV1d 40 FAONVYVAddyY IWYANAD

+ “Id ‘2 "ON ‘L6 “10A SNOILO3Z1II00 SNOSNVIISOSIN NVINOSHLIWS

pias

SRS
i vay

Ne

BY |
CHARLES ELMER RESSER

- Curator, Division of Invertebrate Paleontology,
‘ic Ue S. National Museum

(PuBLicaTIon 3447)

; GITY OF WASHINGTON
~ PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 3, 1938

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER 3

MIDDLE CAMBRIAN FOSSILS FROM
PEND: OREILLE LAKE, IDAHO

(WITH ONE PLATE)

BY
GHARLES ELMER RESSER

Curator, Division of Invertebrate Paleontology,
U. S. National Museum

(PUBLICATION 3447)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 3, 1938

The Lord Battimore Press

BALTIMORE, MD., U. 8. A.

MIDDLE CAMBRIAN FOSSILS FROM PEND OREILLE
LAKE, IDAHO

By CHARLES ELMER RESSER

Curator, Division of Stratigraphic Paleontology
U. S. National Museum

(WitH ONE PLATE)
INTRODUCTION

From 1921 to 1924 Dr. Edward Sampson, of Princeton University,
then a member of the United States Geological Survey, examined the
Pend Oreille mining district which surrounds the southern part of
Pend Oreille Lake, Bonner County, Idaho. A fossiliferous Middle
Cambrian series crops out in several of the fault blocks into which
the district is divided. This was the first Cambrian outcrop discovered
in that part of North America. Subsequently, other Cambrian areas
were found in the northwestern United States and the adjacent por-
tions of Canada, in the extensive area previously thought barren of
Cambrian strata except for the occurrence in the Lewis and Clark
Range, west-central Montana. These occurrences were briefly dis-
cussed in 1934.’ Since the Pend Oreille Lake area is isolated from
other Cambrian outcrops and the stratigraphic succession is clearly
determined, description of the faunas is desirable.

Dr. Sampson published a brief summary* of his findings in the
district and described the stratigraphy, naming three Cambrian
formations.

BELTIAN

Before discussing the Middle Cambrian formations a few words
descriptive of the underlying Beltian strata are in order. Five Beltian
formations, totaling more than 30,000 feet, are described beneath the
Middle Cambrian. This enormous thickness of sediments consists of
argillaceous sandstone, fine-grained massive sandstone, and hetero-
geneous beds of quartzite, sandstone, and argillite, with or without a

* Resser, C. E., Recent discoveries of Cambrian beds in the northwestern United
States. Smithsonian Misc. Coll., vol. 92, no. 10, 1934.

* Sampson, Edward, Geology and silver ore deposits of the Pend Oreille Dis-
trict, Idaho. Idaho Bur. Mines Geol., Pamphlet 31 (mimeographed), 1928.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97, No. 3

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

calcareous content. Cross-bedding, sun cracks, ripple marks and the
other usual Beltian features characterize the series, but contemporane-
ous igneous rocks are evidently lacking, which is also the case in the
Beltian strata of nearby western Montana.

MIDDLE CAMBRIAN

The Middle Cambrian series of the Pend Oreille region begins with
a quartzite, which is followed in turn by argillaceous shale and calcare-
ous formations. No mention is made of younger strata, and the pub-
lished structure sections indicate that the Cambrian is not overlain
by other beds.

Gold Creek quartzite —The Gold Creek quartzite, which is estimated
to average 400 feet in thickness, is easily distinguished from the
Beltian by its coarser grain. Some of the conglomeratic beds contain
pebbles up to 3 inches in diameter and cross-bedding is a characteristic
feature. Outcrops are conspicuous because of the resistance of the
rock to weathering. Unfortunately, the contact of the Gold Creek with
the Beltian rocks is not clearly exposed so that essential history is
lacking.

Rennie shale-—This formation is only 50 to 75 feet thick and con-
sists of soft olive argillaceous shale, sometimes micaceous. Because
it is so easily eroded, the Rennie shale seldom crops out. In fact, the
fossils here described were collected from the bed of a brook.

The published summary fails to mention the fossiliferous limestone
nodules present in the Rennie shale. They evidently are about the same
size and shape as similar nodules obtainable from most Cambrian
shales in the Cordilleran region. Internally, however, these nodules
are peculiar as they consist of an odd mixture of brown and blue lime-
stone. Both sorts occur as distinct masses, sometimes sharply angular,
but more frequently irregular in shape, and the change from one to
the other is abrupt. The brown limestone, which looks like a fine
sandstone, is evidently rather pure calcium carbonate, judging from
its rapid effervescence, and, since this portion of the rock does not
scratch steel, it is assumed to be free from silica or sand. On the
other hand, the blue limestone masses contain sand grains or silica,
even though they also effervesce freely. Fossils are absent from the
brown masses but are very abundant in the blue portions. Strangely,
there are but two species of trilobites in these nodules, the abundant
Vanuxemella idahoensis and rare examples of Albertella sampson.
Neither species has been recognized in the larger shale fauna, although
elsewhere these trilobite forms are found together.

NO. 3 MIDDLE CAMBRIAN FOSSILS FROM IDAHO—RESSER 3

The shale has the species listed below. Besides the names given in
this list there is a poor specimen that seems to be Eocystites. A few
imperfect specimens of Obolus and a fragment of a shell similar to
Westonia ella represent the brachiopods.

Elrathia sampsoni Resser Margaretia angustata Resser
Elrathia longiceps Resser Schistometopus typicalis Resser
Glossopleura intermedia Resser Urotheca sampsoni Resser

Hyolithes idahoensis Resser

Lakeview limestone —Where unaltered, two rock types characterize
this conspicuous and commercially valuable formation. One type con-
sists of cliff-forming massive beds which vary from nearly pure lime-
stone to nearly pure dolomite. The other beds are shaly, containing
thin-bedded, highly fossiliferous limestone. Sampson does not state
what relative position the two types hold with respect to each other,
but the thin-bedded and shaly material probably forms the lower por-
tion of the formation. Metamorphism caused different degrees of
alteration, some of the beds becoming a crystalline marble.

Black crystalline limestone from the shaly beds yields an abundant
fauna which is listed below.

Acrothele speciosa Resser Clavaspidella minor Resser
Acrotreta mitens Resser Elrathia idahoensis Resser
Agnostus bonnerensis Resser Iphidella cf. pannula (White)
Alokistocare noduliferum Resser Lingulella idahoensis Resser
Alokistocare natale Resser Pagetia fossula Resser
Alokistocare nactum Resser Oryctocephalus walcotti Resser
Alokistocare notatum Resser Utia curio Walcott
Alokistocare normale Resser Zacanthoides sampsoni Resser

Alokistocare nothwm Resser

A small collection of altered rock, presumably from the Lakeview
formation, contains a pygidium of Glossopleura. Another lot of im-
pure dark blue limestone is especially interesting because it contains,
among other fossils, a species of Tonkinella, unfortunately too poorly
preserved to illustrate. This is not the Tonkinella-like form described
below as the pygidium possibly belonging to Utia.

RELATIONSHIP OF THE FAUNAS

It has already been pointed out that the faunas of the Rennie shale,
both in the limestone nodules and in the shale, have no species in com-
mon. Nevertheless, both must be regarded as one fauna, since else-
where they occur together. Neither have any species been found com-
mon between the Rennie shale and the Lakeview limestone. These
faunas also are elsewhere found intermingled. From these facts it

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

seems that these faunas represent faunal subzones or possibly facies
developments, but for purposes of correlation it is necessary to treat
the faunas of the Rennie and Lakeview formations as a unit.

The fossils in the Rennie shale are clearly related to those in the
Stephen formation.’ Margaretia, Elrathia, Glossopleura, and the par-
ticular form of Hyolithes are definite relatives of species in the
Stephen. On the other hand Vanusxemella and Albertella are more
characteristic of the older Ptarmigan formation of the Canadian Rock-
ies. The Lakeview is also related to the Stephen, particularly by the
Agnostus, Oryctocephalus, and Zacanthoides. The numerous species
of Alokistocare are found more commonly in other Middle Cambrian
formations than in the Stephen.

Close connection exists between the Lakeview and the Spence shale *
of southern Idaho. Pagetia and the rare trilobite Utia curio indicate
that these two formations are identical in age. The other genera,
both in the Lakeview and in the Rennie, occur in the Spence also.

DESCRIPTION OF THE FOSSIES

The identifiable material is described and illustrated as completely
as possible. In order to avoid unnecessary printing, locality numbers
are given with the descriptions and in the plate legend. A full descrip-
tion of the two localities is given below.

Locality 37m: Middle Cambrian, Rennie shale; headwaters North
Gold Creek, south side of Packsaddle Mountain, east of Pend Oreille
Lake, Idaho.

Locality 37n: Middle Cambrian, Lakeview limestone ; cement mine
just north of Lakeview, Pend Oreille Lake, Idaho.

MARGARETIA Walcott, 1931
MARGARETIA ANGUSTATA, n. sp.
Plate 1, figs 2

A number of narrow flexible tubes have a surface roughened by
elongate depressions typical of Margaretia. Compared with the geno-
type, M. dorus, as well as species in process of publication, M. angus-
tata is considerably smaller in size, averaging less than one-fourth the
diameter of the smaller specimens of the other species.

Locality 37m.

Holotype —U.S.N.M. no. 95019.

* Walcott, C. D., Mount Stephen rocks and fossils. Canadian Alpine Journ.,

vol. I, no. 2, 1908.
Walcott, C. D., Smithsonian Misc. Coll., vol. 53, no. 1, p. 8, 1908.

NO. 3 MIDDLE CAMBRIAN FOSSILS FROM IDAHO—RESSER 5

UROTHECA Matthew, 1899
UROTHECA SAMPSONI, n. sp.

Plate 1, fig. 1

Long, slender tubes abundant in the Rennie shale are referable to
this genus. The illustrated specimen appears to have a carina but is
merely broken in the middle. Faint annulations seem to occur on some
individuals.

Locality 37m.

Holotype —U.S.N.M. no. 95020.

HYOLITHES Lichwald, 1840
HYOLITHES IDAHOENSIS, n. sp.
Plate 1, figs. 57, 58

A species of Hyolithes occurs in the shale; unfortunately, most of
the specimens are poorly preserved. The species is evidently related
to H. carinata but is larger, the carina is less pronounced, and the
operculum has wider wings.

Locality 37m.

Cotypes—U.S.N.M. no. 95021.

LINGULELLA Salter, 1866
LINGULELLA IDAHOENSIS, n. sp.
Plate 1, fig. 18

This shell is nearest to L. isse in shape, but it is a smaller brachio-
pod. It is possible that this brachiopod is one of the Middle Cambrian
forms now included in L. desiderata.

Locality 37n.

Holotype —U.S.N.M. no. 95022.

ACROTHELE Linnarsson, 1876
ACROTHELE SPECIOSA, n. sp.
Plate 1, figs. 6, 7

This form is most like A. colleni, from which it differs in having
weaker ribs and growth lines, but more particularly in the narrowness
of the false area.

Locality 37n.

Cotypes—U.S.N.M. no. 95023.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

ACROTRETA Kutorga, 1847
ACROTRETA NITENS, n. sp.
Plate 1, figs. 3-5

The generic reference is not certain, for this species differs from all
described forms of Acrotreta. Recently, similarly constructed species
have been found in both Lower and Middle Cambrian collections. The
illustrations present clearly the characteristics of the species.

Locality 37n.

Cotypes—U.S.N.M. no. 95024.

AGNOSTUS Brongniart, 1822
AGNOSTUS BONNERENSIS, n. sp.

Plate 1, figs. 16, 17

This agnostid is a typical form of the Cordilleran Middle Cambrian.
The characteristic features place it between A. montis Matthew of the
Stephen formation and A. interstrictus White from the Wheeler shale
of Utah. A. bonnerensis has also been compared with the undescribed
species in the Spence shale fauna, from which it differs in possessing
axial furrows on the pygidium.

Locality 37n.

Holotype and paratypes —U.S.N.M. no. 95025.

PAGETIA Walcott, 1916
PAGETIA FOSSULA, n. sp.
Plate 1, figs. 8-11

P. fossula is similar to P. clytia from the Spence shale, but differs
in having a median furrow like P. bootes. The pygidium has short
axial spines.

Locality 37n.

Cotypes.—U.S.N.M. no. 95026.

ALBERTELLA Walcott, 1908
ALBERTELLA SAMPSONI, n. sp.
Plate 1, figs. 24-26

The glabella of this species is long and is not expanded much in
front. The pygidium is wide, much like A. helena and has a rather
wide concave border, with a nearly straight posterior margin. The
spines diverge more than average.

Locality 37m.

Holotype and paratypes —U.S.N.M. no. 95027.

NO. 3 MIDDLE CAMBRIAN FOSSILS FROM IDAHO—RESSER i

ALOKISTOCARE Lorenz, 1906
ALOKISTOCARE NORMALE, n. sp.
Plate 1, fig. 14

This species is much like A. subcoronatum except for its larger size.
Also the furrows, eyelines, and distribution of relief in the brim are
different.

Locality 37n.

Holotype —U.S.N.M. no. 95028.

ALOKISTOCARE NODULIFERUM, n. sp.
Plate 1, figs. 52, 54

This species has a wide brim, the test is finely and closely granu-
lated, the brim is striated beneath the test, and two nodes are situated
in the dorsal furrow a short distance forward of the occipital furrow.

Locality 37n.

Holotype and paratype—U.S.N.M. no. 95029.

ALOKISTOCARE NATALE, n. sp.
Plate 1, fig. 53

A, natale is nearest like A. noduliferum. It has a brim of about the
same size, and other proportions are similar. Small nodes are also
present in the rear portion of the dorsal furrow. However, the sur-
face of A. natale, which is finely granulated, has, in addition, scattered
larger granules.

Locality 37n.

Holotype—vU.S.N.M. no. 95030.

ALOKISTOCARE NACTUM, n. sp.
Plate 1, figs. 41, 42

A. nactum is characterized by a medium brim, on which a rather
wide flat rim is differentiated by its upturned position.

Locality 37n.

Holotype and paratype —U.S.N.M. no. 95032.

ALOKISTOCARE NOTHUM, n. sp.
Plate 1, figs. 51, 55

This species is more normal than A. nactum, which it resembles.
A medium swelling causes the rim to be less even in width throughout.

Locality 37n.

Holotype and paratype-—U.S.N.M. no. 95031.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

ALOKISTOCARE NOTATUM, n. sp.
Plate 1, fig. 43

This species departs considerably from the norm of the genus be-
cause of its convexity. The test is finely granulated, and only a nar-
row rim is differentiated by the upturned edge.

Locality 37n.

Holotype.—U.S.N.M. no. 95033.

ELRATHIA Walcott, 1924
ELRATHIA IDAHOENSIS, n. sp.
Plate 1, figs. 36-40
A fullness in all parts of the cranidium characterizes this species.
It is typical of the genus in all respects. The thorax has about 15
segments.

Locality 37n.
Holotype and paratypes —U.S.N.M. no. 95034.

ELRATHIA SAMPSONI, n. sp.
Plate 1, figs. 31, 35, 58

Several cranidia of various sizes are illustrated, thus presenting the
characteristics of the species. Compared with E. idahoensis, the spe-
cies is somewhat narrower at the eyes, and the glabella is also tapered
more.

Locality 37m.

Holotype and paratypes —U.S.N.M. no. 95035.

ELRATHIA LONGICEPS, n. sp.
Plate 1, fig. 50
Compared with E. sampsont, this species has a longer glabella and
relatively shorter brim; also, the brim is divided more nearly equally
between the rim and preglabellar area.

Locality 37m.
Holotype —U.S.N.M. no. 950306.

GLOSSOPLEURA Poulsen, 1927
GLOSSOPLEURA INTERMEDIA, n. sp.
Plate 1, fig. 56

Several pygidia, libragenes, and two incomplete hypostomata, but
no cranidia, were found in the shale collections. One pygidium is

NOD 5 MIDDLE CAMBRIAN FOSSILS FROM IDAHO—RESSER 9g

figured. It is more like G. boccar than the Spence shale form because
the doublure is not so wide. Fusion is carried nearly to the extinction
of the rib furrows.

Locality 37m.

Holotype.—U.S.N.M. no. 95037.

ORYCTOCEPHALUS Walcott, 1886
ORYCTOCEPHALUS WALCOTTI, n. sp.

Plate 1, figs. 22, 23

A small fragmentary granulated cranidium, with a typical Orycto-
cephalus glabella is tentatively referred to the species.

The pygidium is nearest like O. reynoldsi, differing in having heavier
spines and more clearly impressed pleural grooves.

Locality 37n.

Holotype and paratype —U.S.N.M. no. 95038.

CLAVASPIDELLA Poulsen, 1927
CLAVASPIDELLA MINOR, n. sp.
Plate 1, figs. 45, 49

A number of specimens in the Lakeview limestone evidently belong
to Clavaspidella. This species is much smaller than any other thus
far described ; also, both the pygidial axis and the eye lobes are long.

Locality 37n.

Holotype and paratypes —U.S.N.M. no. 95039.

UTIA Walcott, 1924
UTIA CURIO Walcott

Plate 1, figs. 19-21

Specific differences are not apparent between the Utia of the Lake-
view limestone and of the Spence shale, consequently the Idaho form
is identified as U. curio.

A peculiar small pygidium characterized by its radiating furrows and
grooves is tentatively assigned to the species.

Locality 37n.

Plesiotypes —U.S.N.M. no. 95041.

VANUXEMELLA Walcott, 1908
VANUXEMELLA IDAHOENSIS, n. sp.
Plate 1, figs. 13-15
This species has stronger pygidial furrows than V. nortia, and also
larger rear spines. Comparison with the Montana species, VY. con-

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

tracta, shows that V. idahoensis has the rear spines set wider apart,
giving the entire pygidium a wider aspect. ”. 1idahoensis has incom-
pletely fused pleural furrows.

Locality 37m.

Cotypes—U.S.N.M. no. 95042.

SCHISTOMETOPUS, n. gen.

Diagnosis —Glabella long, occupying nearly the entire cranidial
length ; tapered slightly. There are four pairs of glabellar furrows and
the occipital furrow. Fixigenes about half width of glabella. Anterior
suture slightly divergent. Posterolateral limbs rather short. Eyes
small, situated slightly behind the midpoint. Eyelines curved back-
ward, arising opposite anterior pair of glabellar furrows. Brim con-
sists of rim only. Two deep furrows run forward from the anterior
angles of the dorsal furrow and separate a central thickened portion
from the two flat lateral portions of the rim.

Genotype.—sS. typicalis, new species.

Name.—oyxworos = divided : peroros = forehead.

SCHISTOMETOPUS TYPICALIS, n. sp.

Plate a, issn

During preparation the important rim was injured because the speci-
men was thought to be an Elrathia. Fortunately, enough of the rim
remains to show its features. It will be observed that the glabella of
S. typicalis is like that of Elrathia sampsoni because of the four sets
of furrows.

Locality 37m.

Holotype-—vU.S.N.M. no. 95040.

ZACANTHOIDES Walcott, 1888
ZACANTHOIDES SAMPSONI, n. sp.
Plate 1, figs. 27-30
A small form of Zacanthoides is present in the Lakeview limestone.
The pygidium has spines of nearly equal length, and the thorax has 8
or g segments. Glabellar furrows are short and shallow.

Locality 37n.
Holotype and paratypes —U.S.N.M. no. 95043.

NO. 3 MIDDLE CAMBRIAN FOSSILS FROM IDAHO—RESSER

EXPLANATION OF PLATE

Bice. Urotheca ‘sampsons, Mews SPECIES» se. o 7 seo osien (seeder

Portion of a specimen. Holotype, X 2.

Fic. 2. Margaretia angustata, new species.........-..-.-.:------

Ten

ire tic 4

Portion of the holotype, X 2. Top portion is of the outside;

lower portion, the impression.

Bircsg-5 Acrotreta mitens, Ne we SPeCleSaa-- «esas somes

3, 4. Ventral valves, X 4.
5. Dorsal valve, X 4.

res. (0.17. Acratnele speciosa, MEW SPECIES... <4 22 =.) 4200020000 0%

Two ventral valves, X 2.

EGS uo-Dheiogena fossula, New) SPECles--- eee sero -

8, 11. Cranidia, X 4.
9, 10. Pygidia and a good cranidium, X 4.

Fic. 12. Schistometopus typicalis, new specieS................06.

Holotype cranidium.

Fics. 13-15. Vanuxemella idahoensis, new species..........+++++.

13. A cranidium.
14. Pygidium showing the spines.
15. Cranidia and pygidium.

Fics. 16, 17. Agnostus bonnerensis, new species..............0--

16. A good cranidium, X 4.
17. Holotype pygidium, x 4.

Fic. 18. Lingulella idahoensts, new SpeCi€S.............0200 000s

Interior of a ventral valve, X 4.

EAGSueTO-21., (Upia. Cirle. WWialCOttes macs in eriosoeciie cis oeeinor elses

19, 20. Cranidia.
21. Pygidium referred to the species, X 4.

Fics. 22, 23. Oryctocephalus walcotti, new species...............
22. Cranidium referred to the species, X 4.

23. Holotype pygidium.

Figs. 24-26. Albertella sampsoni, new species.............000005

24, 25. Two cranidia.
26. Holotype pygidium. ©

Fies. 27-30. Zacanthoides sampsoni, new specieS............++-+-

27. A small cranidium, X 4.
28. The holotype.

29. A large pygidium.

30. A small cranidium, X 2.

Bese 3i-25. Elratiia sampsonm, new SpeCieS..5-.«-...22-- ..) 4-5:

31, 32, 35. Cranidia of various sizes.
33. Holotype cranidium (see also fig. 58).
34. Libragene.

Fics. 36-40. Elratiia idahoensis, new species............-...+---

36-38. Cranidia of several sizes.
39. Holotype cranidium, X 2.
40. Entire individual.

etter 6

on

wiaaratahey it 6

un

TZ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Fics. 41, 42. Alokistocare nactum, new SPpe€CieS.............0cccceeevees

41. Cranidium with libragene, > 2.
42. Holotype cranidium, X 2.

Fic. 43. Aloktstocare. notatum, tlew SpecieS..........0..0cescucccccecece

Holotype cranidium.

Fic. 44. Alokistocare normale, new species..............-00+-+ee eee reee

Holotype cranidium.

Fics. 45-49. Clavaspidella minor, new species..............0eeeeecceees

45. A small cranidium, > 2.
46. Holotype cranidium, X 2.
47. Libragene, X 2.

48. A pygidium, X 4.

49. A pygidium, X 2.

BiG. 50, lrathiavlongiceps newaspecics=: yee eee eee eee

Holotype cranidium.

Figs. 51, 55. Alokistocare nothum, new species.............eeeeencceces

51. Holotype cranidium, > 2.
55. Partially exfoliated cranidium, X 2.

Fics. 52, 54. Alokistocare noduliferum, new specieS............eeeceeeee

52. Holotype cranidium.
54. A small cranidium.

HIG. 53.1 -4okistocare nade: newaspecicsee ren ee en een ene eee

Holotype cranidium, X 2.

Fic. 56. Glossopleura intermedia, new SpecieS..............ceececcccecs

Holotype pygidium.

Fics. '57, 58: Hyolithes tdahoensts: new. Species: /1..). cde tne aes eae eee

57. Operculum, and several tubes.

58. A large example, and cranidia of Elrathia sampsoni.

Figs. I, 2, 12-15, 24-26, 31-35, 56, 58, are from Loc. 37m.
Figs. 3-11, 16-24, 27-30, 36-55, are from Loc. 37n.

SMITHSONIAN MISCELLANEOUS COLLECTIONS WOKE E75 NOs 5 TE, Al

MIDDLE CAMBRIAN FOSSILS FROM PEND OREILLE LAKE, IDAHO

(For explanation, see page ir.)

VOLUME ” "NUMBER 4

BY
JOHN B. SCHMITT

_ Assistant. Entomologist, New Jersey Agricultural
iat Experiment Station. New Brunswick

(PUBLICATION 3448)

_ GITY OF WASHINGTON
"PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 10, 1938

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER 4

THE FEEDING MECHANISM OF
AULT ERY OP DeRA

BY
JOHN B. SGHMITT

Assistant Entomologist, New Jersey Agricultural
Experiment Station, New Brunswick

(PUBLICATION 3448)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 10, 1938

The Lord Baltimore Press

BALTIMORE, MD., U. & As

TBE PEEDING MECHANISM OF ADULT LEPIDOPTERA

By JOHN BSCHMITT

Assistant Entomologist, New Jersey Agricultural
Experiment Station, New Brunswick

CONTENTS

PAGE
NsIELOMUCELOM Fr pecy ae tttne hatuel ce Selon re erie iene crsiemniowsctretaleverstorsiens Sersiac orate I
i Generalstructune of the head and mouthparts. (/. 42.2 ).i0%2-<00-.-% 2
RO DT OUOSCIS! (ES. ni oud adie chucege rem eo yerpee a ohare tele aster de eh uale 5
Structunevand musculature. easiness sterseis aiels clusinie ait eating oe 5
Mechanism) of cotline aid extensions 7. a.1shs.cae suentaleic/slere cian eats 8
Comparative structure in lepidopterous families................... 8
MAMIPm EMG SUC KAIIS 5 PITTING es staat us ec etre tiave Sere x adel ata ge ee eee eee 16
Generalized@structure: thet ccc te sen cee ab one he ele i ee ae 17
ie CH LEPIdOPLEN atiarcwer = vcassumteirey ete wis week a ae etree 18
Mie Mamma stadas ss, sacvers een ara Sareea Seance lata es aU Ree cha eee 20
Comparative structure in lepidopterous families................2.. 20
IM Ae lehieteallairarnieyse ecrsevsaciereeetice eyes cee tae estilo sire oer ee eee 24
SOUS ISMUCTIcL i VM eerste sect poorer es ae atatd Yoda ae ore ara oe eeu OR es 26
mbireviations sed) on the wisures.: «<tc cacs-s nc ted cc amod cee a cue 27
PSHE ECTIC GSE cerns sunt EN Grate aye GPa Were one A rar ies etree ce Mean 28

INTRODUCTION

The mechanism of the feeding apparatus of moths and butterflies has
been studied by a number of anatomists since Reamur and Latreille,
but the exact means by which the proboscis is extended has not been
determined, and it is this problem with which this paper is chiefly con-
cerned. The morphology of the sucking pump has also engaged the
writer’s attention, as have various other parts of the lepidopterous
head.

The literature is not extensive and (as the general information on
the head is contained in most textbooks) there is little need except for
historical purposes to review the contributions previous to the work of
Burgess (1880) who was the first worker to describe correctly the
muscles within the proboscis. Kirbach (1883) wrote on the sucking
pump of Vanessa io and also on the muscles within the proboscis. In
1890 Burgess published further information on the structure of the
head of the milkweed butterfly, followed by Kellogg (1893) on the
same subject. In 1895 Kellogg showed that the pilifers are labral lobes

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97, No. 4

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

and not mandibles. Berlese’s “ Gli Insetti’’ (1910) contains some in-
formation on the maxillary musculature. Tillyard (1923) demon-
strated that the maxillary lobes forming the proboscis are probably
the galeae. Weber (1924) has contributed to our knowledge of the
occipital area and the cervix of certain species. Snodgrass (1935) de-
scribed the sucking pump of a sphingid.

This study was made possible only through the invaluable instruc-
tion and encouragement of R. E. Snodgrass, of the Bureau of Ento-
mology and Plant Quarantine of the United States Department of
Agriculture, and I am therefore especially indebted to him. I am also
indebted to Dr. T. J. Headlee, of Rutgers University, and to Dr. E. N.
Cory, of the University of Maryland, for their cooperation while at
their respective institutions. I also appreciate the aid of Dr. A. B.
Klots, of the City College of New York, in determining microlepi-
doptera. This study formed the larger part of a thesis submitted in
partial fulfillment of the requirements for the degree of Doctor of
Philosophy at Rutgers University.

I. GENERAL STRUCTURE OF THE HEAD AND MOUTHPARTS

The cranium of the lepidopterous head is a relatively simple struc-
ture showing very few sutures. The clypeus forms an elongate an-
terior area and is not marked off from the frons. A suture extends
on each side from the invagination of the anterior arm of the tentorium
to the antenna fossa in the butterflies and most of the higher moths,
but is usually absent in the more generalized groups. Whether this
suture is a true frontal suture is questionable, for, as will be seen later,
its internal ridge seems to have been developed secondarily for the
purpose of bracing the cranium against the pull of the antenna muscles,
which originate on the anterior arms of the tentorium. The internal
ridge of this suture will be called the antennal ridge (fig. 1, A, AR).
The parietals are large, and in the higher Lepidoptera their size is
further increased by the great development of the. compound eyes.
The ventral and anterior ends of each parietal are recurved mesally,
thereby providing between them a recess for the maxillae and the
labium (fig. 12, B). Posteriorly and dorsally, the parietals merge
with the occiput, there being no limiting suture. The postoccipital
suture has a well-developed internal ridge and is itself usually evident
externally. It limits the dorsal part of the posterior edge of the occi-
put, the ventral part being limited by the much lengthened hypostomal
sutures. The invaginations of the posterior arms of the tentorium are
located in the ends of the postoccipital suture, and since most of each
hypostomal suture lies in the same dorsoventral line as the lateral part

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCHMITT 3

of the postoccipital suture, the posterior tentorial pits appear to be
“higher ” in the lepidopterous head than they are in most other insects.
Internally, this part of the hypostomal suture is marked by a well-de-
veloped ridge, on which are inserted the ventral intersegmental muscles
from the thorax. The postocciput and the posterior part of the hypo-
stoma are either poorly developed or entirely membranous.

The ventral areas of the parietals are not marked off from the sub-
genal areas by sutures, so it may be said that the pleurostomal and
anterior part of the hypostomal sutures are nonexistent. Since the in-
vaginations of the anterior tentorial arms of pterygote insects are
always found in either the pleurostomal or the epistomal sutures, it
might be supposed that the furrow extending ventrally from each
anterior tentorial pit is the pleurostomal suture. Such, however, is not
the case. This deep infolding is the line along which the clypeus and
the parietal have been brought into juxtaposition, so that the true
pleurostomal suture would necessarily be within the infolded area. In
some groups, as in the Tineidae and the Pyralidae, this infolding is
not pronounced, but in the butterflies it is extremely well developed.
From a practical viewpoint, these infolded ridges are continuous with
the ridges that brace the floor of a sucking pump, and will be de-
scribed later.

The cephalic endoskeleton, or tentorium, of moths and butterflies
presents practically the same structure throughout the order (fig.1 A).
The anterior arms of the tentorium are well developed and are the most
important part. They are without dorsal arms, and the antennal
muscles arise directly on them. The anterior arms are attached to the
posterior bridge, close to the invaginations of the posterior arms. In
many cases the actual posterior tentorial pits are really large open
depressions, so that when seen from the inside of the head the anterior
arms and the tentorial bridge appear to have separate invaginations.
The posterior bridge is always small and poorly developed, and no
muscles actually arise on its span.

The only muscles arising on the anterior arms of the tentorium are
the antennal muscles and two pairs of muscles affecting the extension
of the proboscis, which will be described later. In the butterflies and
in moths having functional mouth parts the tentorial arms are often
provided with large flanges and ridges, to allow greater attachment
surface. In moths having degenerate or obsolete mouthparts the ten-
torial arms are often bulging and thin-walled, especially in the anterior
halves.

The foramen magnum is sharply constricted near the invaginations
of the posterior tentorial arms, although the degree of constriction

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

varies considerably throughout the order. However, the posterior
bridge of the tentorium is always short.

The mouth parts of adult Lepidoptera consist of the maxillae, the
labium, the labrum, and the hypopharynx. The maxillae, as is well
known, are the most important, their galeae forming the long suc-
torial proboscis in those which have functional feeding mechanisms.
Various degrees of degeneracy may be found, until the point is
reached, as in the males of Thyridopteryx, where the maxillae are no
longer recognizable as distinct appendages. The hypopharynx of
moths and butterflies is incorporated in the floor of the sucking pump

Fic. 1.—Structural details of the head and feeding mechanism of Argynnis
and Danaus.

A, right half of head, mesal view, of Argynnis cybelle, showing endoskeleton
and floor of sucking pump (SP) formed by hypopharynx (Hphy). B, left
half of clypeus and base of left maxilla attached to parietal part of head of
Danaus menippe, anterior view.

and will be described under that heading. The labrum, like the hypo-
pharynx, is really a part of the cranium, but since it plays a part in
the mechanics of feeding it may be described as a mouthpart. The
lateral lobes of the labrum, called the pilifers, bear against the pro-
boscis base, and in some butterflies the labrum is sufficiently flexible
to move as a unit with the proboscis base. In such cases (fig. 1 B)
the proboscis base is provided with a knob that fits against the pilifer
under its fringe of setae. In many moths, however, the labrum does
not have this function. The labium is evident only as a small triangu-
lar area bearing the three-segmented labial palpi. These palpi are
usually so placed that the coiled proboscis can be clasped between
them and be almost completely hidden from view. The labial palpi

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCH MITT 5

are capable of some motion, each palpus having usually one or two
muscles at its base, which enable the palpi to clasp the coiled probos-
cis or to release it.

The mouth cavity, or preoral cavity, is defined by Snodgrass (1935)
as “an external space bounded anteriorly by the epipharyngeal wall
of the labrum and clypeus, posteriorly by the labium, and laterally by
the mandibles and the maxillae.” The hypopharynx is described as
lying in this cavity as a tonguelike lobe. The cibarium is that part of
the preoral cavity which is anterior to the hypopharynx ; that is, the an-
terior surface of the hypopharynx forms its “floor.” The salivary
meatus is the portion of the preoral cavity which is posterior to the
hypopharynx, i. e., enclosed between the hypopharynx and the an-
terior surface of the prementum. The median salivary duct pours its
secretions into this cavity from a small pocket called the salivarium,
between the labium and the hypopharynx.

In the Lepidoptera most of the cibarium is incorporated with a part
of the pharynx in the sucking pump, as will be demonstrated later.
There is a small portion of the epipharyngeal surface which is not a
part of the sucking pump, and this part is usually applied against the
proboscis base. The salivary meatus is practically nonexistent, as the
hypopharynx has completely lost its lobular character, and there is no
protrusion of a prementum beyond the salivarium. The hypopharynx,
in fact, forms most of the “ floor” of the sucking pump (fig. 1 A) as
a single well-sclerotized piece.

Il. THE PROBOSCIS

Savigny long ago discovered that the proboscis of Lepidoptera is
derived from the maxillae or rather from one pair of the lobes of
the maxillae. The cardo and the stipes are usually quite distinct, and
form no part of the proboscis as such; hence the term proboscis
should be reserved for the conveying structure itself. Tillyard (1923)
has produced evidence that the proboscis is derived from the galeae.

Structure and musculature-—Each half or unit of the proboscis is
therefore a tube, the lumen of which is continuous with the body
cavity through the stipes. Each proboscis unit is rendered flexible by
a series of fine rings separated by membrane, as described by Burgess.
These rings are absent in nonfunctional proboscides. In the butterflies
and higher moths the rings are made up of many small flat circles of
hard cuticula, like small beads set in rows. The food channel is also
lined with rings, similar but having only about one-third the width
of the outer rings. Muscles passing obliquely between the rings were

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

described by Burgess as effecting the coiling of the proboscis, a find-

ing verified by Berlese and later writers. ‘
The cardo in functional maxillae is usually a small flat sclerite just

anterior to the labial palpi. The stipes varies in shape throughout the

Fic. 2.—Pressure-producing mechanisms of the maxillae.

A, base of right maxilla of a swallowtail butterfly, ventral view (morpho-
logically posterior), showing at A: a cross-section through the line ab, giving
appearance of stipes when proboscis is coiled. B, cross-section of stipes of
Catocala sp., showing appearance of pressure chamber (PC) formed by stipes.
C, same of Arcyonis alope. D, same of Hemaris thysbe. E, same of Danaus
menippe. EF, same of Pieris rapae. G, base of proboscis of Danaus menippe,
lateral view, showing insertion of posterior tentorial proboscis extensor (ftp).
H, cross-section of stipes of Hemaris thysbe near insertion of posterior tentorial
proboscis extensor (ptp). I, cross-section of stipes of Atrytone zabulon, showing
appearance of pressure chamber.

families above the Tineidae, but when functional always presents
fundamentally the same structures. The proximal portion in cross-
section always has a median flat sclerite continuous with a tubular
lateral part (fig. 2). This tubular part fits into the recurved ventral
and anterior ends of the parietal, the lateral edge of the stipes being

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCH MITT 7

continuous with the parietal. In some cases the curved lateral part of
the proximal portion of the stipes is membranous, as in Cercyonia
alope (fig. 2C). In others the lateral part is heavily sclerotized and
the tubular half is modified so that in effect there are two tubes set
side by side with membrane between. This arrangement is found in
the Pieridae (fig. 2F). The distal portion of the stipes is also re-
cessed under the parietal. The mesal surfaces of the maxillae bear
against each other or against a small projection of the labium.

In addition to the muscles within the proboscis mentioned above
there are three pairs of maxillary muscles inserting on each stipes
and originating within the cranium. Two of these muscles originate

Fic. 3—Proboscis extensor musculature.

A, left half of head of Danaus menippe, showing interior by removal of eye.
B, right half of head, mesal view, of Thyridopteryx ephemeraeformis (male), as
exposed by median sagittal cut. C, left half of head of Desmia funeralis, showing
interior by removal of eye.

on the anterior arm of the tentorium, and the third originates on the
anterior part of the gena. Of the tentorial muscles, one arises on the
lateral surface of the anterior arm and inserts on the distal part of
the stipes, on the median flat sclerite. It is therefore called the anterior
tentorial proboscis extensor (fig. 3 A, atp). The second muscle origi-
nates on the mesal surface of the anterior arm of the tentorium and
inserts near the distal point of the stipes. Its origin on the tentorium
is always posterior to that of the anterior tentorial proboscis extensor,
so that the paths of these muscles cross within the head. This second
muscle is called the posterior tentorial proboscis extensor (fig. 3 A,
ptp). The genal muscle originates on the anterior part of the gena
and inserts on the flat mesal sclerite of the stipes. It is called the
cranial proboscis extensor (fig. 3 A, cp).

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Mechanism of coiling and extension.—It should now be possible to
understand the functions of these muscles. A study of figure 2 shows
that by their contraction, the anterior tentorial proboscis extensors
draw the tubular part of the stipes up against the recurved end of the
gena. (This action is shown diagrammatically in fig. 2A.) It will be
noticed that there is a valve arrangement between the tubular part of
the stipes and the flat sclerite on which the muscles are inserted. As
the muscles draw the stipes upward, the valve (vlv) closes, with the
result that the tubular part becomes a closed cylinder. Thus pressure
is exerted against the blood within the stipes cylinder as it is forced
against the recurved flange of the gena. The stipes cylinder forms a
closed point at its proximal end, and therefore the blood displaced as
the pressure continues must move outward through the stipes, toward
the proboscis. The stipital ridge is enlarged at the proximal end of the
stipes and thus practically covers the lumen of the proboscis unit. The
posterior tentorial proboscis extensor is inserted on this ridge, and con-
traction of this muscle not only creates pressure on the blood enclosed
within the stipes, but also moves the base of the proboscis unit upward,
which effects a tight seal with the functional mouth (fig. 2 G,H;9D).

The blood displaced from the stipes is thus forced out into the lumen
of each tightly coiled proboscis unit, thereby causing the proboscis to
unroll. The diagonal muscles within each proboscis unit, described by
Burgess, cause the proboscis to coil. That blood pressure might be the
agency for uncoiling the proboscis was first suggested to the writer by
R. E. Snodgrass, who, in his “ Principles of Insect Morphology ”’
points out the mechanical analogy of such a mechanism with the toy
paper snake which a child uncoils by blowing into it. The uncoiling
action of one proboscis unit is shown diagrammatically in figure 4.

The mechanism described above is the simplest which the writer has
seen. In many moths and butterflies the stipital cylinder is further
modified, but the principle is invariably the same, as may be seen in
figure 2. The musculature concerned in the extension of the proboscis
seems to be fundamentally the three pairs of muscles described, but
in a large number of insects one or two pairs may be absent. However,
functional maxillae always have at least two pairs.

Comparative structure in lepidopterous families—The maxillae of
a number of species representing the more important families were
examined, primarily to determine the fundamental musculature of the
lepidopterous maxilla. Moths having degenerate or obsolete mouth-
parts were also studied and, indeed, proved to be one of the most in-
teresting phases of this investigation. To expose the proboscis ex-
tensor musculature, a simple procedure is first to make a complete

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCH MITT 9

median sagittal cut, and then, using either half of the head, to remove
the compound eye, the brain, suboesophageal ganglion, and the suck-
ing pump.

Tineidae: In the females of the common Yucca moth, Pronuba
yuccasella, all three pairs of proboscis extensors are present, very
much as described above. There is a single muscle at the base of
each maxillary palpus and each maxillary tentacle. The common
clothes moth, Tineola, lacks the cranial proboscis extensors, but other-
wise its musculature is complete. In certain other Tineidae, determined

Fic. 4.—Diagram of the action of the proboscis extensor muscles of the right
halt of the head, mesal view.

A, proboscis coiled. B, proboscis extended by blood forced into it by com-
pression of stipes (see fig. 2).

to family only, there are no tentorial muscles but only the cranial
proboscis extensors, a very unusual condition.

Coleophoridae: A number of coleophorids, determined to family
only, were found to possess both pairs of tentorial proboscis exten-
sors, but to lack the cranial proboscis extensors.

Limacodidae: A single representative, Euclea cloris indeterminia,
was studied. This moth has practically no proboscis, but only two
very small lobes, each a remnant of a proboscis unit. A single pair of
tentorial proboscis extensors is all that is left of the maxillary mus-
culature.

Oecophoridae: A species of the genus Agonopterix, with a well-
developed proboscis, was also examined. All three pairs of extensors

Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

are well developed, and strands of the anterior proboscis extensor
have migrated onto the clypeus, so that there appears to be a fourth
pair of muscles.

Pyralidae: In the pyralids examined, the cranial proboscis exten-
sor is often absent. In the common wax moth, Galleria mellonella,
both tentorial muscles are clearly evident and apparently able to act.
The proboscis does not appear to be functional, and it is probable
that the proboscis extensors serve only to move the maxillary palpi.
The lesser wax moth, Achroia grisella Fab., has only remnants of the
tentorial extensors, the proboscis being evident only as two short
lobes, apparently nonfunctional. Two other pyralids, Nomophila
noctuella and Ephestia kuehniella, have the proboscis well developed.
The musculature is complete and the cranial muscle is especially well
developed.

In figure 3 C the head of a pyralid, Desmia funeralis, is represented
with the left eye removed. In this case all three proboscis extensor
muscles are present, although the anterior tentorial proboscis extensor
is very small. The cranial proboscis extensor is remarkably large, but
since it originates on the ocular ridge, a relatively thin structure, it
may be doubted whether it exerts much force. It may also be seen
that a large lobe has been formed on each anterior tentorial arm in
order to accommodate a very large antennal muscle, thereby depriving
the proboscis extensor muscles of their usual position. Such a sacri-
fice of feeding structures for nonfeeding structures may be found in
many moths.

Tortricidae: The tortricids usually possess all three pairs of probos-
cis extensors. Figure 10 B represents the head of the common codling
moth, Carpocapsa pomonella, as seen when opened by a median sagit-
tal cut. The sucking pump is shown in place, but the tentorial exten-
sors may be seen just below the pump.

Psychidae: The male of the common bag-worm moth, Thyridop-
teryx ephemeraeformis, was studied in this group. These moths were
found to have an extremely degenerate proboscis (fig. 3 B) repre-
sented only by two large lobes. Each lobe has a single proboscis ex-
tensor muscle, arising on the anterior arm. The antennal muscles,
however, are by far the largest muscles in the head.

Sphingidae: The feeding mechanisms found in this family are gen-
erally very well developed. The musculature of the head of one spe-
cies of sphingid, Sphinx convolvuli, has already been described to some
extent by Berlese (1910). The proboscis muculature which he found
homologizes thus : his no. 190 1s the cranial proboscis extensor, his no.
171 is the anterior tentorial proboscis extensor, and his no. 172 is the

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCH MITT II

posterior tentorial proboscis extensor. However, Berlese apparently
did not examine the insertions of these muscles and makes no refer-
ence to their functions. In discussing the possible mode of extension
of the proboscis Berlese followed the suggestion of Burgess, that it is
unrolled by its own elasticity.

The proboscis musculature of one. species of sphinx moth, Hemaris
thysbe, may be taken as generally typical of the family. In this moth
(fig. 10 D) the tentorial muscles are equally well developed and are
well spaced on the anterior arms of the tentorium. The cranial probos-
cis extensor is moderately developed and in general the whole ar-
rangement is well balanced.

Yet, in some sphingids, there are decidedly inferior proboscis ex-
tension mechanisms. For example, the sucking pump in Smerinthus
geminatus is so large that there is very little space left for the probos-
cis musculature, and the brain also is reduced in size and displaced
posteriorly (fig. 10C). In this moth there is no cranial proboscis
extensor, and only one tentorial muscle, which appears to be the
posterior tentorial proboscis extensor judging by its insertion. An-
other sphingid, Darapsa pholus (fig. 6B), has an extremely large
cranial proboscis extensor.

Geometridae: The feeding mechanisms of members of this family
are weak or degenerate. The proboscis musculature of Haematopis
grataria is shown in fig. 9g C. All three stipital muscles are present, but
very weak, especially the tentorial muscles. The cranial proboscis
extensor is also very short. In the little green geometrids (sub-
family Hemitheinae) the proboscis musculature resembles that of
Haematopis.

The geometrid Ennomos subsignarius is typical of further degen-
eracy in this family. Only the tentorial muscles are present and they
are very weak (fig. 5 B). Each anterior arm is very thin-walled and
is considerably enlarged. This enlargement, of course, “ lightens ”
the head by replacing blood volume with air and is of interest in view
of the remarkable flights of this insect. Caberodes confusaria re-
sembles Ennomos in this respect but has a functional cranial probos-
cis extensor.

The adults of the spring cankerworm, Paleacrita vernata, differ, as is
well known, in that the males are winged and the females wingless.
However, there is practically no difference between the sexes in the
proboscis and its musculature. The proboscis itself, in both cases, is
represented by two small lobes. Only a pair of tentorial proboscis ex-
tensors can be found, and they are very weak.

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Noctuidae: In this family the proboscis and its musculature are
very well developed. The cranial proboscis extensor here reaches a
remarkable size and importance, in comparison with the tentorial pro-
boscis muscles. This is of interest in view of the fact that the cranial
proboscis extensor is the muscle most frequently absent in cases where
there is not a full complement of proboscis muscles. The proboscis
musculature of a large moth, Catocala nubilis, is shown in figure 6 A
and is in general typical of the family.

Arctiidae: All examined members of this family have degenerate
feeding mechanisms. Figure 6 C shows a section through the head of
a typical species, Apantesis virgo. Both A. virgo and A. vittata have

Fic. 5.—Proboscis extensor musculature and the sucking pump.

A, right half of head, mesal view, of Haematopis ‘grataria, as exposed by
median sagittal cut. B, same of Ennomos subsignarius.

all three pairs of proboscis extensors. The posterior tentorial probos-
cis extensor originates well back on the anterior arm of the tentorium.
Estigmene acraea and Utetheisa bella show about the same conditions.

The members of the genus Haploa exhibit the strongest proboscis
musculature seen in this family, and there can be but little doubt that
the proboscis is functional. The members of the genus Diacrisia, on
the other hand, have the most degenerate feeding mechanisms seen
in this family. In D. virginica the anterior tentorial proboscis exten-
sor has been lost and the posterior muscle is very weak. The cranial
proboscis extensor is still evident.

In Jsia isabella all three pairs of proboscis muscles are present and
apparently functional. The anterior tentorial proboscis extensor, how-
ever, has migrated to the antennal ridge, above the anterior arm.

Saturnoidea: The degeneracy of the feeding mechanism in the
giant silk moths is so complete that there are few traces left of the

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCH MITT 13

proboscis musculature. In the large moth Samia cecropia, for ex-
ample, the proboscis is represented by two small shapeless lobes (fig.
11 B, Prb), associated with which there is a single pair of tentorial
muscles. The position of these muscles suggests that they may be the
posterior tentorial proboscis extensors.

Fic. 6.—Proboscis extensor musculature.

A, left half of head of Catocala nubilis, showing interior by removal of eye.
B, same of Darapsa pholus. C, right half of head, mesal view, Apantesis virga,
as exposed by median sagittal cut. D, same of Malacosoma americana.

The tentorium of this moth has developed a peculiar secondary
function. In addition to having the anterior part of the anterior arms
thin-walled and bulging, displacing blood with air, the posterior part
of the anterior arms is tubular and curved to provide a sort of cradle
for the brain and suboesophageal ganglion. This is done by having
each arm pass between the brain and the optic lobe on its side. If it
were not for this the brain would be supported only by the optic lobes.

In other saturniids remnants of one or both pairs of the tentorial
proboscis muscles can be found, but the cranial proboscis extensor is

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

invariably absent. In Basilona imperalis (Ceratocampidae) the pro-
boscis still has a tubular shape and shows a food channel. Both ten-
torial muscles are present, but no cranial muscle.

In all saturniids examined, the position of the tentorium in the
head—that is, the length of the hypostomal area below the posterior
tentorial pits—suggests that the ancestors of these moths had power-
ful feeding mechanisms with the tentorial muscles well developed. The
antennal muscles in these moths are always large and powerful, and it
may be that the great enlargement of the anterior part of the anterior
arms results from the need for a large base for these muscles rather
than from an effort to lighten the head. However, the latter purpose
is unmistakably served.

Lasiocampidae: The adult of the eastern tent caterpillar, Mala-
cosoma americana, was studied as an example of a lasiocampid, but its
resemblance to a saturniid was so complete that no new information
was obtained. The antennal, proboscis, and sucking pump musculature
is illustrated in figure 6 D.

Bombycidae: Similar conditions were found in the common silk
moth, Bombyx mort (fig. 7 A). The proboscis is represented by two
shapeless lobes. Both pairs of tentorial muscles are present.

Papilionoidea: Representatives of five families of butterflies were
studied, and some interesting differences were found. In the Nym-
phalidae and Danaidae the three pairs of proboscis extensors are most
perfectly preserved. Figure 11 shows the proboscis extensor of Danaus
menippe, known as the “ Monarch butterfly.” All three pairs of
muscles are well developed. The musculature of Vanessa atalanta, the
red admiral butterfly, closely resembles it, as does also the mourning
cloak butterfly, Aglais antiopia, and the great spangled frittillary,
Argynnis cybela. In certain other Nymphalidae, however, the cranial
proboscis extensor has been lost. Such butterflies include the viceroy,
Basilarchia archippus, and the common grayling, Cercyonis alope. A
few species of Lycaenidae were also examined. In Lycaenopsis argio-
lus, the common blue, and Everes comyntas, the tailed blue, the mus-
culature consists of the familiar three pairs. In Chrysophanus hypo-
phleas the cranial muscle has been lost, but both tentorial muscles are
well developed.

Thus, in these two families nothing unusual was found. In the
Papilionidae and the Pieridae, however, no trace was found of the
cranial proboscis extensor, but instead there was a remarkable migra-
tion of part of the anterior tentorial proboscis extensor. Inserting on
each stipes with the anterior tentorial proboscis extensor but arising
on the clypeus, between the anterior tentorial arms, there is a single

>

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCH MITT 15

large muscle. This is illustrated in figure 7 B, showing the muscula-
ture of Pieris rapae. The migrant is marked atps, and it really passes
laterad of the posterior tentorial proboscis extensor, its strands insert-
ing with those of the anterior tentorial proboscis extensor which origi-
nate on the anterior arm. Papilio polyxenes, P. troilus, and P. mar-
cellus show similar conditions. Sometimes there is a continuous band
of muscle from the clypeus to the anterior arm of the tentorium.

For

Fic. 7—Proboscis extensor musculature.

A, right half of head, mesal view, of Bombyx mori, as exposed by median
sagittal cut. B, same of Pieris rapae. C, left half of head of Pieris rapae,
showing interior by removal of eye. D, right half of head, mesal view, of
Epargyreus tityrus, as exposed by median sagittal cut.

In the Pieridae this modification is more complete, with the “ mi-
grant” atp. originating higher on the clypeus than in the Papilionidae.
In Pieris protodice, the checkered white, this muscle originates just
anterior of the antennal socket. The musculature of Pieris rapae is
illustrated in figure 7 B. Other pierids examined included Colias eury-
theme and Anthocharis genutia.

By eliminating the cranial proboscis extensor, the Papilionidae and
Pieridae seem to have considerably narrowed the parietal area, or per-

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

haps it should be said, increased the extent of the compound eye, yet
without sacrificing muscular power. In effect, this muscle is here lo-
cated between the anterior arm of the tentorium and the anterolateral
surface of the sucking pump, thus utilizing what might be charac-
terized as “ waste space.” At the same time, the development of this
muscle is necessarily limited by the sucking pump and its muscles.

A fourth pair of muscles should now be described. This pair con-
sists of one muscle located in each proboscis unit, arising on the stipi-
tal ridge and inserting in the proboscis base, and called the proboscis
base muscle (PBm). In direction it is a continuation of the posterior
tentorial proboscis extensor. Its position in the head of Papilio is
indicated in figure 9 B, PBm, also in figure 2 G.

Hesperiidae: Only two species of skippers have been examined:
Epargyreus tityrus and Atrytone zabulon. In this family the probos-
cis extensors are short but very well developed. The anterior arms are
greatly broadened to provide greater attachment surface, while the
cranial proboscis extensors in Epargyreus have invaded the antennal
ridge to secure greater attachment surface.

The remarkable simplicity of the stipital tube in Atrytone is well
worthy of note. In figure 2 I it is represented in cross-section under
compression. In this type, closure of the pressure chamber is effected
directly by the cranial proboscis extensor, as it presses the mem-
branous stipital ridge against the recurved flange of the parietal. The
membranous fold labeled F2 becomes much larger as it approaches
the base of the proboscis unit, at which point its outer portion is firmly
sclerotized, while its inner lateral section (that is, its morphologically
lateral section) remains membranous.

1 PHE SUCKING (PUMP.

Among the orders of insects equipped with sucking pumps, the
Hemiptera and the Diptera have received considerable study. Snod-
grass (1935) has shown that the pump of the cicada is prepharyngeal
in origin and evolved almost entirely from the preoral cibarium. Simi-
larly, Jobling (1929) and Snodgrass (1935) have demonstrated that
the sucking pump in Diptera is derived from the cibarium. In respect
to the sucking pump of the Hymenoptera, Snodgrass (1935) states
that ‘‘ while the morphology of the organ is not entirely clear,... .
judging from the musculature, it includes without doubt the pharynx
and the buccal cavity and perhaps the cibarium.” It is, therefore, of in-
terest to determine to what extent the sucking pump of Lepidoptera
is preoral in derivation.

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCH MITT 17

Generalized structure-—In the more generalized insects there is
usually a large preoral cavity bounded anteriorly by the epipharyngeal
wall of the labrum and clypeus, laterally by the mandibles and the
maxillae, and posteriorly by the labium. The hypopharynx is sus-
pended between these organs (fig. 8 A, Prc) and thus divides the pre-
oral cavity into an anterior food meatus (fm), having the anterior
wall of the hypopharynx for its floor, and a posterior salivary meatus
(sm) enclosed between the posterior wall of the hypopharynx and
the anterior surface of the prementum.

The food meatus, of course, is not part of the stomodaeum, but
simply space enclosed by certain mouthparts. It leads to the true
mouth which marks the beginning of the alimentary canal. A portion

Fic. 8—Comparison of the orthopteroid head with the lepidopterous head.

A, diagram of orthopteroid head (from Snodgrass). B, right half of head,
mesal view, of Danaus menippe, as exposed by median sagittal cut.

of the food meatus just before the mouth is used to hold food before
swallowing and is therefore known as the cibarium (fig. 8 A, Cb).
The dilators of the cibarium always arise on the clypeus.

The part of the stomodaeum just inside the mouth is termed the
buccal cavity (BuC). Beyond the buccal cavity, extending to the
cerebral nerve connectives, we may distinguish the pharynx (Phy).
The dilators of the buccal cavity (dlbc) arise on the clypeus, but the
dilators of the pharynx (dlphy) arise on the frons. The frontal gan-
glion (Fr Gng) lies on the anterior wall of the stomodaeum between
the buccal cavity and the pharynx. The connectives of the frontal
ganglion (Fr Con) always pass laterad of the dilators of the pharynx.
From figure 8 A it may be seen that the pharyngeal dilators are thus
encircled by two nerve rings, outside of which they cannot migrate.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Pump of Lepidoptera——Snodgrass (1935) has shown that the suck-
ing pump of moths and butterflies includes at least the buccopharyn-
geal region of,the stomodaeum. This is evidenced by the fact that the
dilator muscles of the pump are inserted both before and behind the
connectives of the frontal ganglion, which lies on the dorsal wall of
the pump. The sucking pump of a butterfly, Danaus menippe, is il-
lustrated in figure 8 B showing the dilators of the true pharynx in-
serted on the posterior portion of the pump. Whether the cibarium or
any portion of the food meatus is also incorporated in the lepidopterous
sucking pump has therefore been an open question.

The labrum (Lr) of moths and butterflies is usually described as a
narrow transverse band at the lower edge of the clypeal region, bear-
ing the pilifers (P/f) on its lateral extremities (fig. 1B). In orthop-
teroid insects there is a pair of muscles, the compressors of the labrum
(cplr), originating on the anterior wall of the labrum and inserting
on the epipharyngeal wall. If the small lobe between the pilifers is the
labrum, as it appears to be, this pair of muscles exists in the Lepidop-
tera (fig. 8 B and 9 B), and the cibarium then necessarily forms part
of the anterior section of the pump.

However, there is certain other evidence that the cibarium is in-
cluded in the pump, based on the structure of the floor of the pump.
At the base of the salivary meatus in many generalized insects there
is a small cuplike depression or pocket into which the median salivary
duct pours its secretions. This pocket is known as the salivarium
(fig. 8 A, Slv). Itis supplied with three pairs of muscles, a dorsal pair
(rs) arising on the suspensorial sclerites of the hypopharynx, and two
ventral pairs, arising on the prementum. In the Lepidoptera, only the
dorsal pair of muscles, arising on the hypopharynx, may be found.
Their point of origin is on the floor of the sucking pump (fig. 9 A, B)
showing that the anterior part of the floor is derived from the hypo-
pharynx and therefore that this portion of the sucking pump belongs
to the cibarium.

In orthopteroid insects the hypopharynx has a pair of retractors
(fig. 8 A, rhphy) originating on the tentorium. In a geometrid moth,
Haematopis grataria (fig. 5 A), a pair of muscles was found insert-
ing on the floor of the pump and originating on the anterior arms of
the tentorium. Since the ventral dilators of the true pharynx in
orthopteroid insects pass between the circumoesophageal connectives,
they could not possibly migrate from the tentorial bridge to the an-
terior arms. Hence, this pair of muscles in Haematopis must repre-
sent the retractors of the hypopharynx, and although they may have
migrated beyond the limits of hypopharynx, their presence, neverthe-

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCH MITT 19

less, is evidence that the hypopharynx is incorporated in the sucking
pump. In an oecophorid, Agonopterix sp., a similar pair of retractors
occurs.

The highest development of the sucking pump is to be found in the
Sphingidae, the Noctuidae, and especially in the butterflies. Dilation
of the pump is produced by the muscles originating on the wall of the
head ; contraction in the lower moths is produced by the intrinsic elas-
ticity of the pump itself, but in the above-named groups, bands of

Fic. 9.—Various structural details of the head.

A, cross-section of sucking pump of Danaus menippe. B, mesal view of right
half of head and base of right proboscis unit of Papilio sp. C, ventral view of
salivarium of Lycaenopsis argiolus. D, base of proboscis and sucking pump of
Papilio sp. as seen from right side. E, left half of head of Haematopsis grataria,
showing interior by removal of eye.

muscles passing around the pump are mainly responsible. The “ floor ”
of the pump is heavily sclerotized and well braced to withstand the pull
of the dilating muscles. Figure 1 shows the pump in a specimen
cleared in KOH, with a portion of the dorsal wall of the pump re-
moved. The infolded ridge on each side between the parietal and the
clypeus can be seen passing under the floor of the pump (Hphyk),
thus providing support. Morphologically, these ridges are more diffi-
cult to trace. Figure 9 D shows the bracing arrangement of the pump
of a butterfly, Papilio sp. The infolded ridge between the clypeus

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

and the parietal can be seen to merge with the hypopharyngeal ridge,
which is apparently formed jointly by the epipharynx and the hypo-
pharynx. In effect, the hypopharynx has contributed the median sur-
faces of each ridge and the area in the floor of the pump between the
ridges. The dorsal dilators (rs) of the salivarium usually arise on
these ridges, indicating that at least that much is hypopharynx.

The dorsal wall of the pump varies greatly throughout the order
in the particular arrangement of its dilating muscles, but shows in-
teresting consistency in the relative development of the true dilators
of the pharynx and the dilators of the cibarium. Moths beginning
with the Tineidae were examined, but no means was found whereby
muscles which might be dilators of the buccal cavity could be dif-
ferentiated from dilators of the cibarium. Therefore, any dilator
muscles not included in the frontal complex (1. e., encircled by the
connectives of the frontal ganglion, hence true pharyngeal dilators)
are labeled as dilators of the cibarium. With very few exceptions, the
true pharyngeal dilators are restricted to the posterior part of the suck-
ing pump. It does not follow, of course, that the portion of the pump
derived from the pharynx is necessarily limited to this area; it merely
shows the extent to which the dilators of each part have contributed to
the musculature of the pump.

Mechanism.—In figure 12 A the complete musculature of the pump
of Danaus menippe is indicated. The muscles compressing the pump
are shown in cross-section in figure 9 A also. These muscles are
arranged in two groups, transverse pump muscles (tpm) and longi-
tudinal pump muscles (/pm), with two layers in each group. Figure
g A was drawn from a hand-cut section of the pump imbedded in
paraffin. Focusing through the section showed that fibers of the trans-
verse pump muscles passed directly into the dilating muscles, indicating
a possible origin of the compressor muscles from the dilators.

At the anterior end of the pump, a group of transverse pump muscles
are often arranged in a distinct group, forming what Burgess (1880)
called the “ oral valve” (fig. 8B, OVm). Its purpose is believed to
be to prevent the imbibed juices from escaping when the pump is
emptied. This arrangement was found to be especially well developed
in the butterflies and in the Sphingidae.

Comparative structure in lepidopterous families —A number of un-
determined tineids were examined, including the common clothes
moth, Tincola biselliella. In this family the axis of the sucking pump,
that is, a straight line from the anterior to the posterior end of the
pump, is practically perpendicular to the longitudinal body axis. Such
a pump is illustrated in figure 10 A. A single pair of pharyngeal

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCH MITT 21

dilators is encircled by the frontal connectives. The remainder of the
pump dilators form four or more pairs of well-developed short muscles.
The pump musculature of the Yucca moth is poorly developed although
the pump floor is fairly well sclerotized.

In a coleophorid (fig. 10 A) practically the same type of pump was
observed, except that the dilators of the cibarium were grouped an-

Fic. 10.—The sucking pump.

A, right half of head, mesal view, of Coleophora coruscipennella, as exposed
by median sagittal cut. B, same of Carpocapsa pomonella. C, same of Smerinthus
geminatus. D, same of Hemaris thysbe.

teriorly into a large median band of muscles, with a large band placed
laterally on the pump. An oecophorid, Agonopterix sp., has a similar
and well-developed pump. Males of the common bagworm moth, Thy-
ridopteryx ephemeraeformis, possess a very degenerate pump, with
the muscles evident but very weak.

In the Tortricidae the axis of the sucking pump is inclined to a more
horizontal position. In the codling moth, Carpocapsa pomonella (fig.
10 B), there is a single large pair of pharyngeal dilators, and the dorsal

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

pump wall itself is well supplied with muscles. Archips offers nothing
unusual in either pump or proboscis.

Excellent development of the sucking pump may be found in such
pyralids as Desmia funeralis and Nomophila noctuella. The cibarial
dilators are strong and well spaced, although the pharyngeal dilators
are limited to a single pair. In other pyralids, such as the common
wax moth, Galleria mellonella, and the lesser wax moth, Achroia
grisella, the pump is relatively weak, especially in the last-named
species. Other species of pyralids were studied, but nothing unusual
was found.

Sphingidae: Snodgrass (1935) has described the sucking pump of
a Sphinx moth. In Hemaris thysbe the structure of the pump is
typical of this family (fig. 10 D). The pair of pharyngeal dilators is
large and set close together, and with the cibarial dilators, provide the
pump with powerful suction. In one species, Smerinthus geminatus
(fig. 10 C), this development of the sucking pump has reached such
a point that little space is left for the brain and the suboesophageal
ganglion. The proboscis extensor musculature is also reduced to a
single pair of extensors. In fact, the anterior arms of the tentorium
are curved laterally in order to accommodate the expanded pump. In
Darapsa pholus the pump is of more moderate proportions, although
quite well developed. In this species there are two pairs of pharyngeal
dilators. A mouth valve or oral valve is common in this family. The
dorsal muscles of the salivarium are also easily found in the sphingidae.

Geometridae: In this family the sucking pump is generally weak.
Figure 5 B shows the head of a typical geometrid, Ennomos subsig-
narius. There are three pairs of pharyngeal dilators and three pairs
of cibarial dilators, but all are relatively thin muscles. The dorsal
salivarium muscles are also evident, although very small. Caberodes
confusaris shows about the same pump as Ennomos. The sucking
pump of both the males and the females of the spring cankerworm
moth, Paleacrita vernata, is very weak, although provided with four
pairs of dilators.

The sucking pump of Haematopts grataria, in addition to possessing
a pair of hypopharyngeal retractors (fig. 5 A; rhphy), is of interest
because of its unusual formation. There are three pairs of pharyngeal
dilators, the posterior pair originating posterior to the antennae and
passing between the antennal nerves (fig. 9 E). There are also two
pairs of cibarial dilators.

Noctuidae: Members of this family possess well-developed sucking
pumps, of which that of Heliothis obsoleta (fig. 11 A) is typical.
Laterally, the pump is provided with a sheet of fibers on each side, the

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCH MITT 23

posterior bundle of which is shown by the position of the frontal con-
nective to be derived from the pharyngeal dilators. In addition to
these sheets of muscle, there are two pairs of dilators on the anterior
part of the pump. The dorsal salivarium muscles are well developed.
The sucking pump of Autographa falcifera resembles that of Heliothis.

Arctiidae: Moths of this family are provided with poorly-developed
feeding mechanisms. Figure 6C illustrates the head of an arctiid,
Apantesis virgo. The pump muscles are mere strands, and the floor
of the pump is but weakly sclerotized. In /sia isabella the pump and

Fic. 11.—The sucking pump.

A, right half of head, mesal view, of Heliothis obsoleta, as exposed by median
sagittal cut. B, same of Samia cecropia. C, head of Epargyreus tityrus, as seen
with dorsal wall removed.

proboscis are weak but apparently functional. In the genus Haploa
the pump is relatively strong. Diacrisia virginica shows the most
degenerate condition observed in this family, the pump dilators being
mere strands. Yet, in many arctiids the dorsal salivarium muscles are
present and probably functional.

Saturniidae: In this family the sucking pump is extremely weak.
Figure 11 B illustrates the sucking pump of Samua cecropia. There
is a single pair of pharyngeal dilators, still recognizable by means of
the frontal ganglion. Laterally, there are two pairs of muscles which
might be functional. No salivarium muscles could be found. The

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

sucking pumps of other saturniids have about the same development
as in Samua.

Bombycidae: The well-known silk-moth, Bombyx mori, also has
a very feeble sucking pump (fig. 7A). The remnants of only two
pairs of muscles are present, one pair being pharyngeal dilators.

Ceratocampidae: The sucking pump of Basilona imperalis is very
weak and in general much as in the saturniids. There are two pairs of
pharyngeal dilators.

Lasiocampidae: The adult of the eastern tent caterpillar, Malaco-
soma americana, was studied as an example of this family (fig. 6 D).
The pump has a single pair of pharyngeal dilators and three pairs of
cibarial dilators, but all are mere threads.

Hesperuidae: The skippers have well-developed sucking pumps, not
unlike those of the butterflies. The head of Epargyreus tityrus is
illustrated in figures 11 C and 7D. There is only a single pair of
pharyngeal dilators, most of the contraction being provided by the
anterior muscles.

Papilionoidea: The swallowtail butterflies have a large sheet of
muscle on each side of the pump, as well as a pair of median muscles
and a pair of pharyngeal dilators. In the family Pieridae the pump
very much resembles that in Papilionidae, except that two pairs of
pharyngeal dilators are usually present. The Nymphalidae and Danai-
dae show one or two pairs of pharyngeal dilators ; in Argynnis cybele
the dilators of the pharynx originate as two pairs but insert practically
as one. The sucking pump of Danaus menippe is illustrated in figures
8 Band 12A.

However, throughout the families of the Lepidoptera it is probable
that these pairs of pharyngeal dilators do not represent original pairs of
muscles immediately homologous with the dilators of the pharynx of
such insects as Dissostetra. For example, in Dissosteira there is a pair
of retractors of the mouth angles encircled by the frontal connectives,
but it is improbable that any of the muscles encircled by the frontal
connectives in Haematopsis are exactly homologous with the retractors
of the mouth angles (see fig. 8 A, rao).

IV. THE LABIUM

The structure and limits of the labium in adult Lepidoptera have
been previously described by other writers, most recently by Snodgrass
(1935). In figure 12 B the labium of Hemaris thysbe is illustrated.
In this case the labium is limited to a median strip passing to the base
of the proboscis, and a small area around each labial palpus. Pos-
teriorly, the labium is supported by a hypostomal bridge (HBr.).

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCH MITT 25

In the yucca moth, Pronuba yuccasella, there is a small paired fleshy
lobe at the distal end of the labium. The Oriental fruit moth, Grapho-
litha molesta, also has a pair of minute lobes at the tip of the labium,
but it is improbable that these lobes have any significance.

The labium of many moths and butterflies possesses a strong ventral
ridge at the distal end (figs. 11 A and 9 C, Keel). A possible function
of this ridge is to serve as a bearing surface for the proboscis base.

Fic. 12.—Various structural details of the head.

A, head of Danaus menippe, as seen with dorsal wall removed. B, labium and
basal part of maxillae, ventral view, of Hemaris thysbe. C, labial palpus mus-
culature of Papilio glaucus. D, same of Danaus menippe. E, same of Pieris
rapae.

The musculature of the labium is limited to the palpi muscles. Ber-
lese and Burgess have figured palpus muscles arising on the tentorium,
but apparently did so by mistaking proboscis extensors for palpi
muscles. In every moth and butterfly examined by the writer there
were never more than two pairs of palpus muscles, and these arise
either on the labium itself or on the hypostomal bridge. The articula-
tion of the labial palpus with the head is so formed that little or no
blood passes out into the palpus. If the palpus of a live butterfly is
snipped off, it will be found that the walls of both the first and the
second joints are barely moist inside. This, of course, greatly lightens
the palpi.

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

The palpus musculature of Danaus menippe is illustrated in figure
12D. Each palpus is supplied with two muscles, a levator (lplp) and
a depressor (dplp). Nymphalidae usually have a pair of muscles for
each palpus, although the depressor is often very weak. In the Papili-
onidae and the Pieridae there is no depressor muscle, but the levator
is always well developed (fig. 12 C, E). In this case the levator arises
either on the sclerotized median plate (MP) of the labium, or on the
hypostomal bridge.

The presence or absence of labial palpus muscles is extremely
variable in the other families. Pronuba yuccasella has no palpus
muscles, nor does an Agonopterix sp. Each palpus of Galleria mel-
lonella has a single large muscle. Geometridae show only a single
palpus muscle or none at all. In the Arctiidae there is only a single
muscle, usually arising on the hypostomal bridge. Sphingidae have
either one or two pairs of palpus muscles, commonly only one. Satur-
niids and other extremely degenerate groups usually lack any palpus
musculature, and the trembling motion of the palpi sometimes seen in
this family is usually caused by the remnants of the proboscis exten-
sors. However, Basilona imperalis has a single muscle in each palpus,
as does also Malacosoma americana.

V. SUMMARY

I. The coiled proboscis of Lepidoptera is extended by means of
blood pressure created in the stipes of each maxilla. This pressure is
caused by three pairs of muscles, which by their contraction press the
stipes against the head wall. Two pairs of these muscles arise on the
anterior arms of the tentorium and the third pair arises on the gena.

2. The sucking pump is a compound organ, derived from the
pharynx, the buccal cavity, and the cibarium. This is evidenced by
these facts: (1) true pharyngeal dilators are inserted only in the
posterior part of the pump; (2) muscles homologous with the com-
pressors of the labrum are present in some Lepidoptera; and (3) the
dorsal salivarium muscles arise on the pump floor, showing that the
hypopharynx forms at least the anterior part of the floor.

3. There is no labial musculature except that of the palpi. There
are generally two pairs of palpus muscles, but in many families only
one pair, or none at all, may be found.

4. The area posterior to the labial palpi is bounded by the hypos-
toma, the hypostomal ridge offering an insertion for the ventral inter-
segmental muscles. A hypostomal bridge is sometimes present.

5. The anterior arms of the tentorium are well developed but lack
dorsal arms. The posterior tentorial bridge is short and weak. The

NO. 4 FEEDING MECHANISM OF LEPIDOPTERA—SCHMITT 27
great length of the hypostoma in Lepidoptera elevates the tentorium
to a higher position in the head, with respect to other cephalic struc-
tures, than is common.

6. The antennal muscles arise on the anterior arms of the tentorium
and vary in number from one to five pairs. They are always well
developed, sometimes at the expense of other head structures and, in
moths with obsolete feeding structures, are often the only functional

muscles within the head.

ABBREVIATIONS USED ON THE FIGURES

Ant, antenna.

ant mcl, antennal muscle.

Ant Nv, antennal nerve.

AR,.antennal ridge.

AT, anterior tentorial arms.

at, invagination of anterior arm.

atp, anterior tentorial proboscis ex-
tensor.

BuC, buccal cavity.

Br, brain.

Cd, cardo.

Clp, clypeus.

cp, cranial proboscis extensor.

Cu, cervix.

dlbc, dilator of buccal cavity:

dlcb, dilator of cibarium.

dplp, depressor of palpus.

dlphy, dilator of pharynx.

eye:

fm, food meatus.

Fr, frons.

Fr Con, frontal connective.

Fr Gng, frontal ganglion.

HBr, hypostomal bridge.

Hphy, hypopharynx.

HphyR, hypopharyngeal ridge.

HR, hypostomal ridge.

hs, hypostomal suture.

Hst, hypostoma.

Lb, labium.

Lb Pip, labial palpus.

Iplp, levator of palpus.

lpm, lateral pump muscle.

Lrm, labrum.

MP, median plate.

Mth, mouth.

Nv, nerve.

Oc, occiput.

OcR, occipital ridge.

ocs, occipital suture.

Oe, oesophagus.

OVm, oral valve muscle.

PBm, proboscis base muscles.

PC, pressure chamber.

Phy, pharynx.

Pif, pilifer.

Plp, palpus.

Poc, postocciput.

PoR, postoccipital ridge.

pos, postoccipital suture.

Prb, proboscis.

Prb Ext, proboscis extensor.

Pritl, parietal.

PT, posterior tentorial arms.

pt, invaginations of posterior arms.

ptp, posterior tentorial proboscis ex-
tensor.

rao, retractor of mouth angles.

rhphy, retractor of hypopharynx.

rm(RNv), recurrent nerve.

Is, anterior salivarium muscle.

2s, 3S, posterior salivarium muscle.

SID, salivary duct.

Slv, salivarium.

sm, salivary meatus.

Soe Gung, suboesophageal ganglion.

SP, sucking pump.

SR, stipital ridge.

St, stipes.

thm, transverse pump muscle.

Tr, trachea.

ulv, valve.

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

REFERENCES

BERLESE, A.
1910. Gli insetti, vol. 1. Milan.
Burcess, E.
1880a. The structure and action of a butterfly’s trunk. Amer. Nat., vol. 14,
PP. 313-319.
1880 b. Contribution to the anatomy of the milkweed butterfly. Anniv.
Mem. Boston Soc. Nat. Hist.
JoBLING, B.
1929. A comparative study of the structure of the head and mouthparts in
the Streblidae (Diptera Pupipara). Parasitology, vol. 21, pp. 417-
444.
KeEttoce, V. L.
1893. The sclerites of the head of Danaus archippus. Kansas Univ. Quart.,
vol. 2, pp. 51-59.
1895. The mouthparts of the Lepidoptera. Amer. Nat., vol. 29, pp. 546-556.
KirBacH, P.
1883. Uber die Mundwerkzeuge der Schmetterlinge. Zool. Anz., vol. 6, pp.
553-558.
Snoperass, R. E.
1932. Evolution of the insect head and the organs of feeding. Ann. Rep.
Smithsonian Inst. for 1931, pp. 443-489.
1935. Principles of insect morphology. McGraw-Hill Book Co., New York.
WimeESONO, IRS J).
1923. On the mouth-parts of the Micropterygoidea. Trans. Ent. Soc. Lon-
don, vol. 71, pp. 181-206, 12 figs.
WEBER, H.
1924. Das Thorakalskelett der Lepidopteren. Ein Beitrag zur vergleichenden
Morphologie des Insektenthorax. Zeitschr. Anat. und Entwick.,
vol. 73, pp. 277-331, 9 figs.

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- "SMITHSONIAN MISCELLANEOUS COLLECTIONS
; _ VOLUME 97, NUMBER $

THREE PICTOGRAPHIC AUTOBIOGRAPHIES
«OF SITTING BULL

(WirTH 46 PLaTEs)

Ne Sea ek
SUES iain OR 4S My, WeeSTIRLING
2 - ~ Chief, Bureau of American Ethnology

(PUBLICATION 3482)
CITY OF WASHINGTON
- PUBLISHED BY THE SMITHSONIAN INSTITUTION
JULY 22, 1938

o

Photograph by Barry

SITTING BULL

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER 5

THREE PICTOGRAPHIC AUTOBIOGRAPHIES
OF SI LEING BUEL

(WiTH 46 PLATES)

BY
M. W. STIRLING

Chief, Bureau of American Ethnology

(PUBLICATION 3482)

GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JULY 22, 1938

The Lord Baltimore Presa

BALTIMORE, MD., U. & A.

ike Es PICTOGRAPEIC AUPOBIOGRAPHIES OF
SITTING BULE

By M. W. STIRLING
Chief, Bureau of American Ethnology

(WitH 46 PLATES)

INTRODUCTION

The name of Sitting Bull will probably always remain as the best
known of any American Indian. Whether or not this preeminent place
is deserved, it is a fact that more has been printed about him than
any other Indian and his name has most intrigued the popular imagi-
nation. Part of this notoriety resulted from the fact that he was a
prominent and influential Indian. during a crucial period in the
history of his tribe and partly from the fact that during the latter
years of his life he was exploited, both in this country and abroad,
in a manner calculated to bring his achievements in a highly colored
manner before the general public.’

It is not the purpose of this introduction to outline the career of
Sitting Bull. This has been adequately done by a number of biogra-
phers, and the interested reader is referred to the attached selected
bibliography.

There is no question concerning the fact that Sitting Bull was a
great man, in spite of the fact that many of his contemporaries at-
tempted to belittle his character. It is true, however, that the promi-
nence he later achieved in the popular mind was partly due to
circumstances which gave unusual publicity to his career.

In view of his status with the government, Sitting Bull was always
reluctant to speak with white men regarding his personal adventures.
However, in keeping with the custom of his people, he was proud
of his war exploits and kept a careful record of them.

*In an unpublished manuscript on Sioux names by the famous scout and
interpreter, E. H. Allison, the author says of the Indian name of Sitting Bull,
Tatanka Yotanka; “Sitting Bull’s totem was a bull standing in a defiant attitude,
which clearly expressed the meaning of his name, ‘The Bull in Possession,’
‘The Conquering Bull, ‘The Bull of Occupation,’ ‘The Sitting Bull.’ ”

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97, No. 5

I. THE KIMBALL PICTOGRAPHIC RECORD

When Sitting Bull recounted his honors at the dance following
the Sioux victory over the Crow in 1870, Frank Grouard, who was
present, states that at this time Sitting Bull was entitled to 63 coups.’
About this same time Sitting Bull made his pictographic record after
the usual manner of the Plains Indians, representing the feats which
entitled him to special credit among the Indians. This set of drawings
he gave to his adopted brother, Jumping Bull, who placed with them
a pictographic record of his own. While these drawings were in the
possession of Jumping Bull, Four Horns copied 55 of them, includ-
ing 40 from the record of Sitting Bull and the remainder from that
of Jumping Bull. In some manner, not yet explained, these copies
fell into the hands of another Indian who brought them to Fort
Buford, where they eventually came into the possession of Assistant
Surgeon James Kimball in August 1870. The fate of the original
drawings from which this set was made is not known, although
Sitting Bull stated that they were still in the possession of Jumping
Bull as late as 1881. According to Col. H. M. Morrow, his father,
also Col. H. M. Morrow, who was with Dr. Kimball, procured an
identical set at the same time. These copies were both drawn on
roster sheets of the Thirty-first United States Infantry. The copy
retained by the Morrow family was destroyed in San Francisco in
the great fire of 1906. The copy obtained by Dr. Kimball was de-
posited by him, together with explanations of the pictures obtained
at the time from other Indians, with the Medical Director’s Office,
Department of Dakota, on March 14, 1871. The same year they
were transferred to the Army Medical Museum in Washington,
DiC. On May 15, 1915, Dr. D:. S. Lamb of the Army Medical
Museum transferred them to the archives of the Bureau of American
Ethnology, where they are at the present time.

From time to time this pictographic record has attracted consider-
able attention, but it has been reproduced only in part, and the
supporting documents concerning it have never before been published.

Although the name of Sitting Bull had already become well known
to the whites, he did not become a figure of outstanding national
interest until after the annihilation of General Custer and the Seventh
Cavalry, June 25, 1876, in which battle Sitting Bull participated.

*De Barthe, 1894, p. 105.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

As news concerning the details of the Custer defeat slowly filtered
in from the northern plains, newspaper men realized that they had
the biggest news story since the Civil War. The fact that the colorful
Custer, who had become something of a national idol, was the central
figure in the tragic affair, made the story ideal from the standpoint
of the journalists. In search of material which could be tied in with
the Custer fight, a Washington correspondent learned of the existence
of the copy of Sitting Bull’s autobiography, then in the Army Medical
“Museum. Sitting Bull was known to have participated in the battle.
Here then was the perfect nucleus for a follow-up story. On July 6,
1876, the New York Herald published a highly colored account of
the pictographic record, neglecting to note that the autobiography
was not the original handiwork of Sitting Bull. The record was cited
as proof of. Sitting Bull’s cruelty, lust for battle and vainglorious
boastfulness. This story was a huge journalistic success. It was
copied and revamped by newspapers and magazines throughout the
United States. Sitting Bull, who heretofore in the public mind had
been but one of a group of hostile chiefs resisting the westward ad-
vance of the whites, now became Public Enemy Number I and a
character of outstanding interest.

Apparently, an introduction written by Dr. Kimball formerly ac-
companied the explanatory index and the Williamson letter of verifi-
cation which are now with the pictographic record.

The news release from Washington of July 6, 1876, as published
by the New York Herald says:

Among the many ghastly souvenirs preserved at the Army Medical Museum
of this city is an autobiography of Sitting Bull, gotten up in the highest style
of the art of savage picture history, and telling, in fifty-five drawings or
sketches, the story of his life down to 1870. Each picture is. rudely outlined
in ink, the men, horses and other objects being such as children would make.
Many of them are partly filled in with red and blue colors as if Sitting Bull
had at some time got possession of one of the red and blue pencils so well
known in newspaper offices, and with it elaborated his pictorial efforts. Blood
or a wound is indicated by a red blotch with streamers falling down from it.
The blue is used generally in indicating the white man’s pantaloons. Each
picture is made on a sheet of paper eight by ten inches, and is pasted into a
book of blank leaves, such as are used for a scrap book. By holding the sheets
up to the light it is seen that they are the muster-roll blanks of the 31st United
States Infantry, of which Col. de Trobriand was the commandant. The papers
probably fell into Sitting Bull’s hands at the evacuation of a camp, or, as is
more likely, were stolen by him during a visit to some of our outposts. Sitting
Bull is not-at all modest in committing to posterity the story of his great deeds.
Whether it be the scalping of a soldier in battle or the sly theft of a mule, he

brags equally of his prowess in his curious autobiography. This literary work,
which is now likely to be famous, fell into the hands of Assistant Surgeon

NO. 5 SITTING BULL—-STIRLING 5

James C. Kimball, of the army, in the month of August, 1870, while he was
stationed at Fort Buford, Dakota Territory. He had the pictures translated,
and sent them, with the translation and an index, to the Curator of the Army
Medical Museum, Washington, Surgeon George A. Otis, United States Army,
who has filed them in book shape, among the archives of the Museum. The
introduction, written by Dr. Kimball goes, on to say that the autobiography
contains a description of the principal adventures in the life of Sitting Bull,
who is an Unk-pa-pa chief. It was sketched by himself in the picture language,
in common use with the Indians. Since the establishment of Fort Buford, in
1866, Sitting Bull, at the head of from sixty to seventy warriors, had been
the terror of mail-carriers, wood-choppers and small parties in the vicinity of
the post and from 100 to 200 miles from it either way, up and down the
Missouri River. During the time from 1866 to 1870, when the autobiography
was written, this band had several times captured and destroyed the mail and
had stolen and run off over 200 head of cattle and killed near a score of white
men in the immediate vicinity of the fort. The Unk-pa-pas are a tribe of the
great Sioux Nation, living in the Yellowstone and Powder River countries.

The book was brought into Fort Buford by a Yanktonnais Sioux, and offered
for sale and purchased for $1.50 worth of provisions. The Indian gave con-
flicting statements regarding the manner in which he came into possession of
the book, exciting suspicion that he had stolen it from Sitting Bull, who in his
turn, undoubtedly stole the book in blank from the whites.

In an article over the name of Porte Crayon published in the
supplement to Harper’s Weekly of July 29, 1876, the editor says:

About the year 1870 a collection of MS. drawings, put up in book form, ©
bearing the autograph of Sitting Bull and exhibiting a record of his exploits
and adventures, was brought into Fort Buford by a Yanktonnais Sioux and
sold for a dollar and fifty cents worth of provisions. When cross-questioned
regarding the ownership of the book, the Indian shuffled and prevaricated so
as to confirm the belief that he had stolen it from Sitting Bull himself. The
authenticity of the work, with its general historical accuracy, is confirmed by
Assistant Surgeon James C. Kimball, U.S.A., who, with the aid of interpreters,
Indians, and others versed in the picture-language of the Northwestern tribes,
wrote a detailed explanation of the scenes represented, accompanied by a brief
sketch of the warrior-artist’s life. The book was then forwarded to the Superin-
tendent of the Army Medical Museum at Washington, who placed it in the
hands of the present editor.

The series consists of fifty-five designs, drawn on the blank side of printed
rosters of the Thirty-first United States Infantry, of uniform size (about eight
by ten inches), clearly outlined with a pen and a brown ink resembling sepia.
There is no attempt at shading, but the outlines are filled in with flat tints,
very crudely laid on, with red and blue chalk, yellow ochre, green, and the
same brown ink or pigment used in the outlines. The coloring, which is quite
appropriate in the dress and trappings of the human figures, is rather florid
in the animals. Thus while there seems to be great care in showing the
characteristic spots and markings of the horses and mules, the sorrels are
represented with bright yellow, the grays with blue, the bays red, and the
browns and blacks with the aforesaid brown ink.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

This coloring, however, serves to impart life and meaning to the designs,
to relieve the groupings from confusion, and is sometimes so arranged as to
produce quite an artistic effect of chiaro-oscuro. It may be further noted that
there is no attempt at foreshortening, the objects and figures being all shown
in flat profile, and without exception, all looking and moving in the same
direction, that is, from right to left.

Of all the objects presented by the artist, the figure of the buffalo bull is
elaborated with the most intelligent and loving minuteness. The horses and
mules are drawn with a free and well-assured hand, with a tendency to manner-
ism, relieved somewhat by distinctive character in color, markings, and details.
He is least happy in his delineations of the human figure, draperies, and
accoutrements, although in some scenes his attitudes are spirited and his
costumes sufficiently marked to enable us to identify the sex and country of
those who have had the honor to sit for their portraits to this distinguished
limner.

The information in the two foregoing newspaper accounts con-
cerning the manner in which the pictographic record was obtained
at Fort Buford presumably was obtained from Dr. Kimball’s now
missing introduction. The article published by Harper’s Weekly
reproduces 11 of the drawings with a rather detailed description of
the set based partly on the Kimball index and partly upon specula-
tion by the editor.

As already indicated, the pictures are drawn on the reverse side
of loose-leaf: roster pages of the Thirty-first United States Infantry.
The numbers were subsequently placed on them arbitrarily without
regard to the actual chronology of the events described.*

In 1881 the pictures, together with the Kimball index, were for-
warded through Col. George S. Andrews to Rev. John P. Williamson,
missionary with the Sioux, who showed them to Sitting Bull for
purposes of verification. The results obtained from this interview
are explained in the following letter :

Fort RANDALL, DAKOTA TER.
Dec. 12, 1881.
Cot. Gro. L. ANDREWS,

25 U. S. Infantry,
Commanding Post,

Sir:

I have the honor to state that in connection with Capt. G. Lawson, I inter-
viewed Sitting Bull in regard to the supposed “Hyeroglyphic Autobiography”
of himself, contained in pictured sketches, numbered 1 to 55, obtained by Jas. C.
Kimball, Ass’t. Surgeon, U.S.A., in the year 1870.

* Vestal, in describing them, has placed them in what he considers to be the
order in which the different feats took place.

NO. 5 SITTING BULL—STIRLING ih

Sittting Bull immediately recognized the pictures as scenes from his early
life, with the exception of Nos. 39 to 51, and 53 and 54, which he said were
not his, but were adventures of his brother Jumping Bull.

As to the scenes from his own life, he says these are all true scenes, and
he drew a similar set many years ago and gave them to his brother Jumping
Bull. He saw his brother last summer and understood from him that he still had
them. He thinks therefore that this set must be a copy of the one he made,
and has been drawn off by some Indian, he does not know by whom. He
could tell perhaps by seeing his brother who is at Standing Rock.

Sitting Bull verified in the main the Index accompanying the pictures.

No. 1 he says was his first feat, accomplished when he was fourteen years
of age.

No. 10 he says was a Ree, who drops his gun and bow from fear. He was
struck (for “coup”) but not killed (no blood is shown). The scalp at the
horses bridle, here and elsewhere, not being intended to represent the scalp
of the enemy drawn.

No. 55 he says is not completed—should have his “name” (as he calls the
sitting buffalo).

As to the particular history of each event recorded, we found Sitting Bull
rather reserved, especially in regard to Scenes Nos. 11 to 26, and we could
see that any narration he gave of the several events was colored by the circum-
stances of his present situation. And I would suggest that if a more full account
of his war deeds is desired, a better time to secure it would be at some future
date when his status is definitely determined.

Yours Respectfully,
(signed) JoHN P. WILLIAMSON,
Missionary.

In reproducing the drawings, the explanation of each is given
exactly as written in the Kimball index in 1870. It should be borne
in mind that these interpretations were furnished by Indians familiar
with the career of Sitting Bull but not by the Sioux warrior himself.

For purposes of comparison these explanations are supplemented
by the interpretations published by Vestal.*

No. 54 is missing from the set. This picture was one of the Jump-
ing Bull series and represented an episode in the famous battle of
1870 between the Sioux and the Crow. The Kimball description
says “Sitting Bull at the head of his band charges into a camp of
Crows and kills thirty of them. (This happened in the winter of
1869-70. )”’

“Vestal says, “For Sitting Bull’s interpretations of these drawings, given in
1885, I am indebted to Mr. Seth C. Jones, Secretary, Municipal Art Com-
mission, Rochester, N. Y.”

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 1

“Sitting Bull, a young man without reputation and therefore wear-
ing no feather, engages in his first battle and charges his enemy, a
Crow Indian who is in the act of drawing his bow, rides him down
and strikes him with a ‘coup’ stick. Sitting Bull’s autograph—a
buffalo bull sitting on his haunches—is inscribed over him. His
shield suspended in front has on it the figure of an eagle which
he considers his ‘medicine’-—in the Indian sense of the term.” ”
KIMBALL.

®*See Williamson letter, p. 7. For detailed circumstances of this exploit,
see Vestal, 1932, p. 13.

“1846. On Red Water. The boy Sitting Bull, as yet an unfledged
warrior, is shown on horseback, charging an enemy whom he strikes
with a coup stick. On his blue shield a black bird is painted, and
four black-tipped eagle feathers flutter from the edges of the shield.”
—VESTAL.

No, 2

“Sitting Bull wearing a war bonnet is leader of a war party who
takes a party of Crows consisting of three women and a man, so
completely by surprise that the man has not time to draw his arrows
from the quiver. Sitting Bull kills one woman with his lance and
captures another, the man meanwhile endeavoring to drag him from
his horse, from which it is supposed he is forced to desist by others
of the war-party. The fate only of Sitting Bull and his victims is
given in this history.”—K1MBALL.

“1858. Rainy Butte. This picture commemorates the capture of
three Crow women, at the time when Sitting Bull’s father was killed.
Sitting Bull carries the lance made for him by his parents, and wears
a bonnet with horns and a long trail of eagle feathers. A Crow |
warrior is represented as trying to arrest his charge.”—VESTAL.

NO. § SITTING BULL—STIRLING 9

No. 3

“Sitting Bull pursuing his enemy, a Crow Indian whom he strikes
with his lance.”—KIMBALL.

“1856. On Yellowstone River. Sitting Bull counts coup with his
lance on a mounted Crow warrior who carries a shield and a gun.
As required by the obligations belonging to his shield, Sitting Bull
wears his hair in a knot like a horn on his forehead.”—VEsTAL.

No. 4

“Lances a Crow woman.”—KIMBALL.

“1860. Sitting Bull counts coup on a Crow woman riding a mule.
She turns to fend off his lance as he strikes at her. This happened
when the Sioux encountered Crow hunters among the buffalo herds
and Makes-the-Enemy killed two Crow women.”—VESTAL.

TO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 5

“T ances a Crow Indian.”—-KIMBALL.

“1853(?). Sitting Bull unhorses a Crow warrior with his lance.
The story is well known, but no eye-witnesses now live, and the
date and place are uncertain.’”’—VESTAL.

No. 6

“Sitting Bull twice wounded, and unhorsed. His enemy, a Crow,
at length killed by a shot in the abdomen and his scalp taken and
hung on Sitting Bull’s bridle.” “—KimBa tt.

** Regarding the mention of scalps in this and succeeding pictures, see William-
son letter, p. 7.

“7856. On Porcupine Creek. Sitting Bull, shown wearing his
Strong Heart bonnet and sash, crouches behind his shield and shoots
a Crow chief through the belly, at the same time being wounded
in the foot. Flame and smoke pour from the guns, and the wounds
bleed freely. Sitting Bull’s black war horse awaits its master in the
background.”’—VESTAL.

NO. 5 SITTING BULL—STIRLING er

Nos37

“In an engagement with the Crows, Sitting Bull mortally wounds
one of the enemy and dropping his lance rides up and strikes him
with his whip. The lines and dashes in the picture represent the
arrows and bullets that were flying in the air during the combat.”—
KIMBALL.

“7861. Sitting Bull, amid a hail of enemy bullets, wounds a
Crow warrior with his lance, then drops it and strikes him over the
head with the heavy notched wooden handle of his quirt, which is
decorated with a dangling kit-fox skin—the insignia of his Warrior
Society. The Crow carries a quiver, and bleeds freely.”—-VEsTAL.

No. 8

“Counts ‘coup’ on a Gros Ventre de Prairie, by striking him with
his lance. Gros Ventre distinguished from Crow by manner of
wearing the hair.” —KIMBALL.

“1857. On the Missouri River. Winter. Sitting Bull, armed with
a gun and wearing his Strong Heart and white blanket coat, strikes
with his lance the Hohe lad whom he is to save and to adopt as his
brother, named Jumping Bull, or Little Assiniboin.”—VeEsTAL.

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 9

“Lances a Crow Indian.”—KiIMBALL.

“1858. Near Rainy Butte. Sitting Bull lances and kills a Crow
warrior, the slayer of his father in that very fight.’ “—VEsSTAL.

*’ For details see Vestal, p. 44.

No. 10

“A Crow Indian attempts to seize Sitting Bull’s horse by the
bridle. Sitting Bull knocks him down with a ‘coup’ stick, takes his

99 6

scalp and hangs it to his bridle.” “—KIMBALL.

* Sitting Bull corrected this interpretation saying that his opponent is a Ree
who drops his gun and bow from fear. The Ree was struck for coup but not
killed (no blood is shown). See p. 7.

“1859. Near Fort Berthold. A Ree enemy grabs the bridle of
Sitting Bull’s horse. Sitting Bull kills him, and takes his gun and
bow.” —VESTAL.

,

10

12

NO. 5 SITTING BULL——-STIRLING 13

No» 17

“Sitting Bull with his brother mounted behind him kill a white

man—a soldier.” “—KIMBALL.

“When Williamson showed these pictures to Sitting Bull for verification,
he found him unwilling to go into detail concerning his war exploits involving
white opponents.

“1868. In a skirmish with white men Sitting Bull rescues his
unhorsed companion Jumping Bull, takes him up behind, and charges
a white man armed with a rifle. Jumping Bull, being armed with
a long lance, is able to strike the white man first. Sitting Bull has
to be content with the second couwp.”’—VESTAL.

No. 12

“Counts ‘coup’ on a white man by striking him with a ‘coup’ stick.”
—KIMBALL.

“1868. Sitting Bull strikes a white man. This happened on the
same warpath as the deed recorded in Fig. 11. Circling Hawk, now
living, was leader of this war party.”—VESTAL.

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 13

“In a warm engagement with the whites, as shown by the bullets
flying about, Sitting Bull shoots an arrow through the body of a
soldier who turns and fires wounding Sitting Bull in the hip.”—
KIMBALL.

“1864. Near White Butte, on the Little Missouri River. Under
heavy fire, Sitting Bull charges a white soldier. Though transfixed
by an arrow from behind, and bleeding copiously from mouth and
wounds, the brave soldier turns and shoots Sitting Bull through the
buttocks, causing great loss of blood.” —VeEsrTat.

’ This episode is described in detail by Vestal. See Vestal, 1932, p. 64.

No. 14

“Sitting Bull counts ‘coup’ on a white man by striking him with
his bow. Sitting Bull wears a jacket and bandanna handkerchief
taken from some of his victims.”—K1MBALL.

“7867-68 (winter). On the Montana Trail. Sitting Bull counts
coup on a white man. In this affair Sitting Bull counted nine coups.
This picture is followed by eight others showing the other coups
struck. But as the drawings differ only in the details of the dress
and persons of the white men, they have not been given here. Several
of the white men were represented as having hair on their bodies—
a thing considered loathsome by the Sioux.”—VESTAL.

14

16

NO.

5 SITTING BULL—STIRLING

No. 15

“Sitting Bull counting ‘coup’ on a white man.’”—KIMBALL.

No. 16

“Sitting Bull counting ‘coup’ on a white man.’’—KIMBALL.

10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 17

“Sitting Bull counting ‘coup’ on a white man.’”—KIMBALL.

No. 18

“Sitting Bull counting ‘coup’ on a white man.”—KIMBALL.

17

18

KB

20

NO.

5 SITTING BULL—STIRLING

No. 19

“Sitting Bull counting ‘coup’ on a white man.’”—KIMBALL.

No. 20

“Sitting Bull counting ‘coup’ on a white man.’’—KIMBALL.

17

18

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 21

“Sitting Bull counting ‘coup’ on a white man.’”—IKIMBALL.

No. 22

“Sitting Bull counting ‘coup’ on a white man.’’—KIMBALL.

21

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22

24

NO. 5 SITTING BULL——STIRLING 19

No. 23

“Sitting Bull shoots a frontiersman wearing a buckskin shirt,
takes his scalp which he hangs on his own bridle and captures his
horse. Sitting Bull wears a blanket.”—KIMBALL.

“1863. Near Fort Totten, in the Devil’s Lake country. Sitting
Bull, wearing a red blanket, chases a mounted white man in a fringed
buckskin coat, and shoots him between the shoulders. This was
Sitting Bull’s first white victim.’”—VESTAL.

No. 24

“Sitting Bull strikes a white soldier with his ‘coup’ stick, takes
his scalp and his mule. Wears a.war shirt.”—KIMBALL.

“1863, June. The skirmish with General H. H. Sibley’s wagon-
train on the Missouri River, near the mouth of Apple Creek. Sitting
Bull, facing a heavy fire, as shown by flying bullets, charges a mule-
skinner armed with a blacksnake whip, counts coup on him, and
makes off with a saddled mule.”—VEsTAL.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 25

“Counts ‘coup’ on a soldier, mounted with overcoat on, gun slung
across his back, by riding up and striking with his riding whip.”—
KIMBALL.

“1867. On the Montana Trail. Sitting Bull overtakes a white man
wearing an overcoat and armed with a rifle. Sitting Bull carries
only a quirt, with which he strikes the fugitive. On his head Sitting
Bull wears a bandanna taken from some enemy.’”—VESTAL.

No. 26

“Kills a white man and takes his scalp.”—KIMBALL.

“1867. On the Niobrara River near the Missouri. Sitting Bull
shoots a white man armed with a sawed-off shotgun. Sitting Bull
carries a revolver, and is riding a rawhide saddle, made by his uncle.”
—VESTAL.

26

NO. 5 SITTING BULL——STIRLING Za

No. 27
“Captures a mule and a scalp.”—KIMBALL.
“1865. North of the Black Hills. In a skirmish with the troops

under Colonel N. Cole, of the Powder River Expedition, Sitting Bull
runs off a slow pack-mule.”—VESTAL,

No. 28
“In a warm engagement, captures a horse and a scalp.”—KIMBALL.
“7865. On the Montana Trail. Under heavy fire from the soldiers,

Sitting Bull captures a buckskin mare. He afterward gave her to
his sister.”—VESTAL.

ho
to

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 29

“Steals a mule.”—KIMBALL.

“November 6, 1867. Fort Buford. In an attack on the woodcutters
from the post, one soldier was killed, one wounded. Sitting Bull

captures a fine brown Army mule with a black spot on the withers,
off side. He gave the mule to his sister.” —VESTAL.

No. 30

“Captures two horses in action.”—K1IMBALL.

“1864. Under fire, Sitting Bull takes from the soldiers a chestnut

and a buckskin horse. The buckskin he trained to run buffalo, and

then gave it to his sister. These horses were captured in the Badlands
from General Sully’s troops.”—VEsTAL.

30

32

NO. 5 SITTING BULL—STIRLING

to
ow

Nove3t

“Steals a horse.”—KIMBALL.

“7865. On the Montana Trail. Sitting Bull steals a fast buckskin
war horse. He gave it to his adopted brother, Jumping Bull.’—
VESTAL.

No. 32

“Steals and runs off a drove of horses from the Crows.’—
KIMBALL.

“1863-64 (winter). Sitting Bull brings home nine Crow ponies:
five bays, two blacks, one buckskin mule, and a little white mare.
The mare he presented to his favorite sister, Pretty Plume.’—
VESTAL.

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 33

“In an engagement captures a government horse, and mule, and a
scalp.” —KIMBALL.

“1860. Amid a shower of bullets, which fill the air, Sitting Bull,
riding his famous war horse Blackie, runs off two animals from a
Crow camp. One of them is a branded Army mule, picked up or
stolen by the Crows. These animals Sitting Bull gave for Brown
Eyes, the girl who became his fourth wife.”’—VEsTAL.

No. 34.

“Steals a horse.” —K1MBALL.

“1866. On the Montana Trail. Sitting Bull takes a horse with
a split ear from white men.”—VeESsTAL.

34

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39

36

NO. 5 SITTING BULL—-STIRLING 2

cn

No. 35

“Captures three horses and a scalp.”—KIMBALL.

“1866. Wearing his Strong Heart bonnet, and riding Blackie,
Sitting Bull captures three Crow ponies; one bay, one black, one
mouse-colored.’”—VESTAL.

No. 36

“Steals a drove of horses from the Crows.’’-—KIMBALL.

“1862. Sitting Bull runs off a bunch of Crow ponies. Sitting Bull
was such a noted horse-stealer that the old men say nobody can
remember all his raids. Chief Charging Thunder stated that to his
own knowledge Sitting Bull took horses from the Crows twenty
times, sometimes as many as thirty head at a time.” —VESTAL.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 37
“Steals a government horse.’”—KIMBALL.
“1865. Wearing beaded leggins and a fur cap with earflaps,

Sitting Bull runs off a horse belonging to the Powder River Expedi-
tion.” —VESTAL.

No. 38
“Steals a drove of horses from the Crows.’’—KiIMBALL.
“1859-60. Wearing his Strong Heart bonnet, Sitting Bull runs off

seven Crow ponies: two white, two black, one bay, one buckskin,
and one mouse-colored.”—VESTAL.

38

40

NO. 5 SITTING BULL——-STIRLING 27

No. 39

“In an engagement captures a mule. Sitting Bull first appears here
as Chief of the Band of Strong Hearts, to which dignity his prowess
has raised him. The insignia of his rank—a bow having on one end
a lance head—he carries in his hand.” “—KIMBALL.

* This drawing actually depicts a feat of Jumping Bull the adopted son of
Sitting Bull. See Williamson letter, p. 7.

No. 40

“Sitting Bull, Chief of the Band of Strong Hearts, captures two
horses in an engagement in which his horse is wounded in the
shoulder.” “—KIMBALL.

* This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

No. 41

99 11

“Captures a horse in a fight.” “—KIMBALL.

“ This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

No. 42

99 12

“Steals a mule. KIMBALL.

“This drawing actually represents an exploit of J umping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

41

42

44

NO. 5 SITTING BULL—STIRLING 29

No. 43

“Captures two horses in a fight in which his horse is wounded in

99 13

the leg.” “—KIMBALL,

* This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

No. 44

yyoi4

“Mounted on a government horse, captures a white man.
KIMBALL.

“This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 45

99 15

“Steals two horses.” “—KIMBALL.

* This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

No. 46

“Captures four mules in a fight in which his horse is wounded in
the hip.” ““—KIMBALL.

*® This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

48

NO. 5 SITTING BULL—-STIRLING 31

No. 47
“Counts ‘coup’ on white man.” “—KIMBALL,

™ This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

No. 48

9) 18

“Counts ‘coup’ on white man.” “—-KIMBALL.

*® This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 49

99 19

“Steals a government horse. KIMBALL.

” This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

No. 50

“Fastens his horse to his lance driven into the earth and in a hand
to hand fight kills a white man with his own gun. The black marks
show the ground fought and trampled over.” “—KIMBALL.

This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

52

NO. 5 SITTING BULL—STIRLING 33

No. 51

“A fort into which his enemies the Crows, have retreated and from
which they maintain a hot fire through which Sitting Bull charges
the fort.” "—KIMBALL.

“This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

This apparently represents the feat of Jumping Bull performed at Spoon
Horn Butte, where he drew the fire of the Crows in order to empty their guns
before the Sioux charged. This exploit is described in Vestal, 1932, p. 116.

No. 52

“In a fight with the Crows, Sitting Bull kills and scalps one Indian,
and counts ‘coup’ on another who fired at him barely missing him.” *
—KIMBALL.

“For details concerning this exploit see Vestal, 1932, chap. 16.

“1869. Near the Big Dry. An incident of the battle in which the
thirty Crows were killed. Sitting Bull, wearing a horned bonnet and
beaded leggins, charges the rocky barrier (indicated by the circle),
and counts coup upon a Crow, who fires in his face, but misses. The
air is full of flying lead.”—VESTAL.

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

No. 53

2

“Steals a drove of mules.” *“—KIMBALL.

“This drawing actually represents an exploit of Jumping Bull. It is errone-
ously attributed to Sitting Bull by Kimball. (See p. 7.)

No. 55

“Kills one Crow and counts ‘coup’ on two others, who run from
him disgracefully.” “—KIMBALL.

* Sitting Bull stated to Williamson that this picture is incomplete. It should
carry his “name” glyph. (See p. 7.)

“This, the last of the series, is incomplete, and lacks the picture
of the seated buffalo, which should identify Sitting Bull. However,
the shield is enough to serve that purpose. Sitting Bull himself
explained that this unfinished sketch represented a fight with the
Crows in which he killed one and counted coup on two others, who
ran from him disgracefully. The date and place of this fight are
unknown.”—VESTAL.

09

i

_ <ict a |

II. THE SMITH PICTOGRAPHIC RECORD

In June 1923, through the generosity of Mr. Robert A. Smith,
the Bureau of American Ethnology archives were enriched by an-
other Sitting Bull document of even greater interest. This consists
of a later Sitting Bull pictographic autobiography drawn by the
great Sioux warrior himself. Although it contains drawings of only
22 exploits, it is well documented, and the explanations of the draw-
ings were given by Sitting Bull at the time the pictures were made.

The drawings were made with a pencil on the pages of an army
ledger book, and colored by means of water-color paint. The human
figures are rather crudely drawn in the usual Plains Indian style,
“but Sitting Bull shows his individuality even in the field of art, by
the manner in which the horses are depicted. Departing from the
general Plains Indian style of representing horses in a slender and
much conventionalized fashion, he draws his horses: realistically and
in a well rounded manner. The various horses shown are so consci-
entiously delineated that some of them can be recognized from
descriptions of Sitting Bull’s favorite mounts given by Vestal and
others.”

In the Four Horns copy of Sitting Bull’s autobiography of 1870,
the warrior is always identified by his name glyph in the form of a
seated buffalo. At the time the present picture record was made,
Sitting Bull had learned to write his name, and his signature accom-
panies each drawing in the place of the buffalo.

The following letter from Mr. Smith accompanied the book of
pictures and the documents concerning them.

*®Bob Davis, the well known newspaper writer,.informed the author that
in an interview in 1931, he learned that Rudolph Cronau was sent to America
by the Gartenlaube, a weekly periodical published in Leipzig, to cover the Indian
wars as illustrator. In 1881 the artist made the acquaintance of Sitting Bull
shortly after his surrender at Fort Buford. Sitting Bull was much interested
in watching Cronau make his sketches and wishing to gain the friendship
of the Sioux leader, Cronau spent some time in teaching him to draw. This
very probably accounts for the sophisticated and un-Indian appearance of the
horses drawn by Sitting Bull.

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Rosert A. SMITH
430 S. Garden Street
South Bellingham, Wash.
June 20, 1923.
SMITHSONIAN INSTITUTION
Washington, D. C.

GENTLEMEN :

I am sending with this a book of paintings by Sitting Bull, with interpre-
tation of same, letters from Wallace Tear, Lieut. 25th Infantry U.S.A. to my
father, General John C. Smith, which explain themselves. Lieut. Tear was a
soldier in the 96th Regt. Infantry—Illinois U.S.V. 1860-65.

My father was Captain, Major, Lieutenant Colonel, Brevet-Colonel 96th
Regt., and Brevet Brig. General. At the close of the war he was able to get
a commission in the Regular Army for Tear and did him some favors after-
wards, hence this History of Bull.

All the people mentioned are gone and when I pass on there will be none
that will be interested, so I would like to have this book where perhaps it might
interest someone—sometime.

I am, Gentlemen,

Respectfully,
Rosert A. SMITH
430 South Gardner Street
South Bellingham
Washington

Two letters of explanation from Lieutenant Tear addressed to
Gen. John C. Smith were with the pictographic record.

Fort RanpDALL, D. T.
August 10, 1882
DEAR GENERAL:

Yours of 31th ult., with photographs of yourself and wife recd. Many thanks.
Mrs. Smith looks younger than she did 20 years ago—fact—you don’t look
very old yourself. I may be a little prejudiced in this matter as I am “passing
off” as a young man yet.

I have Sitting Bull’s description of the paintings, taken down when the
pictures were made. Am copying them for you, and will send them next mail.
Intended to send them with the book, but I had to send the- book to keep it
from being stolen. I came near losing it. Some tourists wished to look at it
and then borrowed it for a while to show to some friends.- I only got it back
“by a scratch.” They had hidden it with the intention of carrying it off. As
soon as I got my hands on it, I put it into the mail.

Bull is very diffident about giving any incidents of his fights with the whites.
I have tried to have him give me a detailed description of the Custer fight
but he seems rather timid. Once in speaking of the affair he said: “I did not
hunt Custer. I thought I had a right to protect my own women and children.
If he (Custer) had taken our village he would have killed our women and
children. It was a fair fight.”

I will try and get him to make a picture of some portion of the fight.

NO. 5 SITTING BULL—STIRLING a7.

I will try and think of some trinket that would please him as a gift from

you—something that will cost but littl—I can’t think of anything at present.

Am quite well. Have been out in the field a good deal this summer and am
quite busy. I have the luck to be alone with the Co. most of the time.

Love to all. Of course you will be elected. If you need my vote I will come

home on election day.

Yours truly,

W. Tear (signed)

Fort RANDALL, D. T.
August 16, 1882
DEAR GENERAL:

I send you inclosed Sitting Bull’s interpretation of his paintings recently sent
you.

I furnished the book which contains the paintings and from time to time
saw him at work on them. These notes were taken down by me, after the
paintings were completed, in Sitting Bull’s tipi in the same routine as given
by himself (thro an interpreter of course) Bull having the picture before him
while giving a description of the fight. It was impossible to locate the scenes
with any definiteness; ‘““The Land of the Sioux’; “The Land of the Crows”
and “a long way from the Missouri” being the usual location given. In talking
of his life Bull uses his name instead of the pronoun “l’’; that is he speaks in the
third person. In these notes you must understand that it is Sitting Bull speaking.
Bull made these pictures for me to show his gratitude for blankets and clothing
furnished his children last winter before the Government supply of clothing for
his band arrived.

I am endeavoring to get him to complete his history up to the present time,
and if successful you shall have it.

Bull says he is 43 years old. I think he is nearer 50. These scenes of his
life of course comprise his life from the time that he was able to ride a horse
and handle a bow. The scars of the wounds he speaks of are visible now.

Regards to your family.

Yours truly,
W. TeEAR (Signed)
Lieut. 25th Infantry
Gen. J. C. SmitH
Chicago, III.
250 W. Van Buren Street

The list of explanations with the picture record are written in
the hand of Lieutenant Tear on nine pages of foolscap paper. These
explanations are here placed opposite the pictures described, exactly
as written.

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. o

Assinniboine Chief taken prisoner by Sitting Bull in a fight between
Sioux and Assinniboines. 100 Sioux—whole tribe of Assinniboines
engaged—about 27 years ago, when Sitting Bull was about 16 years
old—Land of the Sioux. Kept the chief prisoner for while and then
gave him the horse he (“Bull”) rode and the bonnet he (“Bull”)
wore in the fight and then sent him to his people with a good heart.

No. I

Fight with Assinniboines—14o Sioux—43 Assinniboines—23 As-
sinniboines killed—8 Sioux killed—2o0 Sioux wounded—“Bull” 18
years old—In land of Sioux. Bull took several prisoners. Didn’t
kill prisoners. Kept them many days. Gave them ponies and sent
them home. Assinniboines were hunting in Land of the Sioux.

NO. 5 SITTING BULL——-STIRLING 39

No. 2

Scene in same fight.

No. 3

Fight with Assinniboines. “Bull” 24 years old. 50 Sioux—20o
Assinniboines. No Sioux killed. 5 Assinniboines killed. “Bull” kills
warrior. “Bull” 24 years old.

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 4

Assinniboine woman taken prisoner by “Bull” in a fight. “Bull”
16 years old.

No. 5

Same fight. 5 women captured by “Bull.”

NO. 5 SITTING BULL—STIRLING 41

No. 6

Same fight. Woman captured by “Bull.” The warrior toutches
[sic] woman with his lance and she becomes a prisoner. Warrior
never strikes a woman in a fight except to save his own life. These
women were kept with the Sioux a short time and then sent back to
their own people except 3 who married Sioux warriors and remained ;
one of them here now. No one killed in this fight. Assinniboines
passing thro Land of the Sioux.

Now?

Fight with the Crows. Crow Indian killed by “Bull.” 30 Sioux
warriors—200 Crows, men, women and children. 14 Crows killed.
No Sioux killed. Crows were travelling with their camps. “Bull”
20 years old. On the Little Missouri river, “Crows always fighting
the Sioux—tried to make friends with them but they were always
doing something bad.”—Butt.

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 8

Fight with Assinniboines. Warrior killed by “Bull.” 350 Sioux
run upon 10 Assinniboines and killed 2. 1 Sioux killed and 2
wounded. Land of the Sioux, a little above the forks of the Missouri
near mouth of Yellow Stone. “Bull” 25 years old.

No. 9
Scene in same fight described in No. 7. “Bull” kills Crow Indian.

SITTING BULL—STIRLING 43

NO. 5

No. 10

Fight with Crows. “Bull” killed 2 men and captured 2 women.
“Bull” 25 years old. Crows were stealing ponies. Let women go
home with presents for Crow chiefs to try and make friends.

No. II

Fight with Crows. 100 Sioux—whole tribe of Crows. Bull killed
Crow Chief. 3 Crows killed. 1 Sioux warrior and I woman killed.
Land of the Sioux—a little above Tongue River. Bull 22 years.

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 12

Same fight. “Took long time to kill these people. Here is where
I got wounded in leg and got off of horse and killed this man. No
prisoners in that fight. This is ‘Stand and Kill’ Crow Chief. Had
guns in this fight. The Sioux used to take the Crows prisoners
and give them good clothes and feed them up and give them good
ponies and then send them back so they could tell a good story of
the Sioux to their people.’ (‘‘Bull’s” description of fight.)

No. 13

Fight with Assinniboines. “Bull” takes 2 prisoners. “Bull” 30
years old. 300 Sioux—20 Assinniboines—2 Assinniboines killed—
no Sioux killed. On the big fork of the Missouri. “Bull” took one
prisoner, “Jumping Bull,” home to his (Bull’s) tipi (wigwam) gave
him his (“Bull’s”) horse and war bonnet. Jumping Bulls’ father
was a Chief. Jumping Bull is now at Standing Rock (Fort Yates,
D. T.) with my people. They call him my son.

li

13

15

NO. 5 SITTING BULL-—STIRLING 45

No. 14

Fight with the Rees. 16 Sioux. 100 Rees. Sioux were fighting
and retreating. Sioux turned and chased Rees. This Ree Indian,
Chief “Bull Head,” fell down dead. “Bull” took him prisoner and
he came to life again. This is the only Ree caught in the fight. No
Sioux killed. In the land of the Rees. “Bull” sent this prisoner home
with presents. Made peace with Rees, and peace with Assinniboines.
“Bull” 33 years old.

No. 15

Fight with Gen. Miles’ Scouts and Crow Indians. “Bull” kills
“Brave Indian,” one of Gen. Miles Scouts. About three years ago—
the time Gen. Miles was out after the Sioux near the Queens’ land
(Canada) “Brave Indian” was away ahead of the soldiers and was
following up the Sioux too close. “Bull” turned and killed “Brave
Indian.” One Cheyenne Indian (scout) also killed. Sioux did not
fight soldiers—wanted to get away from soldiers. The scouts and
Crows killed 5 Sioux before they got to Canada. Gen. Miles’ Scouts
seemed to be from every Indian nation. The Sioux run away.

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 16

Fight with Flat Heads. “Bull” wounded in left arm and side by
arrow. “Bull” killed Flat Head. 15 Sioux, young men, went on
war path. Flat Heads killed them all. Sioux then went out with
300 warriors. 40 Sioux attacked the camp of the Flat Heads; the
main body of Sioux being hid back from the camp; the Flat Heads
chased the 40 Sioux back through the main force of Sioux. The
Sioux charged and killed 33 Flat Heads. 4 Sioux killed—good many
wounded. 7 years ago. Bull 36 years old. Near Muscle Shell river.

Nomw17

Crow Indian killed by “Bull.” 200 Sioux run upon 7 Crows hunt-
ing in Land of the Sioux and killed them all. Crows had guns. Sioux
had nothing but bows and lances. Crows were crossing Missouri—
(river). A few years ago. (“‘Bull” gave his age at the time of this
fight but my notes are defaced at this point so that I am uncertain
as to how old he said he was.—TEar.)

Pe

>»

19

NO. 5 SITTING BULL—STIRLING 47

No. 18

Fight with Crows. 200 Sioux—whole tribe of Crows. “Bull”
kills Crow warrior. 5 Crows killed in the fight. No Sioux killed.
“Bull” 33 years old. (Chicken Hawk, skinned and skin stuffed, worn
as ornament where picture of bird is seen near Bull’s head.)

No. 19

Fight with Crows. 200 Sioux—whole tribe of Crows. “Bull”
kills warrior. 7 Crows killed. No Sioux killed. “Bull” dressed in
war bonnet trimmed with eagle feather. “Bull” 24 years old. Near
mouth of Tongue River.

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 20
Fight with Assinniboines. 100 Sioux—6o Assinniboines. “Bull”
kills warrior. 3 Assinniboines killed. 1 Sioux wounded. Near big
fork of the Missouri. “Bull” 29 years old.

No. 21

Fight with Assinniboines. 320 Sioux attacked big winter camp
of Assinniboines. Bull kills warrior. 5 Assinniboines killed. No
Sioux killed. Did not get in to their (Assinniboines’) camp—there
were too many and fought too well. Sioux run off after killing these

men.

s LLL

nw

21

ny va ae ee eee ty

Ill. THE PETTINGER PICTOGRAPHIC RECORD

During the month of February, 1938, a news release appeared
concerning the Sitting Bull autobiographies in the Bureau of Ameri-
can Ethnology. The following letter came as a result :

OsWEGO, OREGON
March 7, 1938
SMITHSONIAN INSTITUTION
Washington, D. C.

GENTLEMEN :

My uncle the late Dan’l L. Pratt of Seattle was Post Trader at Fort Randall,
Dak. Ter. in 1882, 56 years ago and knew Sitting Bull and his band very well.

Sitting Bull sketched for him 13 pictures of himself on horseback showing
him in action against the Crows, Gros Ventres etc.—each one is drawn on
paper 103 by 84 inches and marked in print—D. L. Pratt, Post Trader, Fort
Randall, 188-. The horses are very well drawn—in Indian style—some in colors.
These pictures came to me in book form bound in oil cloth. I now have them
with the affidavit of Mr. Pratt in a large walnut frame—they are quite im-
pressive.

Hoping this information will be interesting,

I am
Yours truly,
/s/ Mrs. G. H. PETTINGER
Oswego, Ore.

The writer communicated at once with Mrs. Pettinger, who very
kindly forwarded the pictures to the Bureau of American Ethnology
so that they could be included with this publication. They belong to
the George Howard Pettinger Collection, which contains several
other very interesting Sitting Bull items, including the tomahawk
surrendered by Sitting Bull to Lieutenant Ogle on the occasion of
Sitting Bull’s surrender to the Commanding Officer at Fort Buford,
Dakota Territory. The writer wishes here to express his deep ap-
preciation to Mr. Pettinger for making available this interesting
addition to the Sitting Bull record.

These pictures, like those of the Smith autobiography, were made
at Fort Randall in 1882 and were probably painted from the same
paint box, as the shades of the colors used are identical in the two
sets. The Pettinger drawings are exactly the same in style but lack
the signature. In the Pettinger drawings, Nos. 2 and 5 are unique
in that full-face figures are shown. It seems probable that some
pictures are missing from the series as originally drawn, for in two
instances the descriptive sequence appears to refer to a preceding
episode which is not shown.

49

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. I

“Winter” scene. Killing a Gros Ventres Indian—1oo Sioux on
war path killed 3 Gros Ventres whom they found hunting.”

* This is the same episode as that shown in No. 21 of the Smith autobiography.
The horse is blue-black in color. Sitting Bull is wearing black army trousers
with a red stripe. His victim wears a green blanket coat.

No. 2

1860. Sitting Bull killing a Crow Indian. One hundred Sioux
chased thirty Crows all night. Caught them in the morning and
killed them all.”

* This is probably the fight that took place in the winter of 1869. See Vestal,
i atl

Ce Bo. Ccelie> GaLLGenr Bear OF En, fe~
Pra frie, Filler! 3S Gye0 lewtiee thane Clog
7

=e ;
CL Atttetiees
file sep

PRATT, JR
ThADE

Qui htt thie Lat, bina. Le 2égtl— Chtyae hia
le Che TEE, Cnc Mclod Phew BE

L. PRATT; JR.

As

L. PRATT, JR

+
\

NO. 5 SITTING BULL—STIRLING 51

Noi 3

Sitting Bull killing a Crow Indian. 40 Sioux against 7 Crows.
Sitting Bull killed 3 Crows in this fight.

No. 4
Same fight.

on
ty

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 5
Sitting Bull killing a Crow Indian.*
* Sitting Bull is here shown wearing the same costume depicted in No. 2.

These two pictures are unique in that in three instances the men pictured are
shown in full face instead of profile.

No. 6
Killing a Flathead Indian in a battle.”

* Sitting Bull is wearing a shirt covered with green spots.

Jr

ATT

Aedlan Gs Glee Mth Bite te Ce Kllbe Ae,

j .
e ee ee Cla oh Gul lt Chee Btrey

Wen
" WH

NO. 5 SITTING BULL——STIRLING 53

No: 7
Killing a Flathead and receiving an arrow wound in left side.”

* This is the same episode as depicted as No. 16 of the Smith series. This
battle with the Flatheads and the particular incident here shown is described
by Vestal, p. 125.

No. 8
200 Sioux in a fight with 50 Chippeways. Sitting Bull killed one.*

* Sitting Bull is mounted on a yellow horse, black nose and ear tips. His
shield is painted blue. In all of the other pictures in which the shield is shown,
the color is green.

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

No. 9

Jumping Bull, a Gros Ventres, captured in a fight by Bull—Sitting
Bull took a fancy to him and presented him with his war bonnet and
horse and permitted him to be free—made chief after this occur-
rence.”

“ This represents the same episode as pictured in No. 13 of the Smith record
and No. 5 of the Kimball record.

No. 10

1880. Killing a Crow Indian Scout who belonged to General
Miles’ command. Sitting Bull wears a war bonnet which once be-
longed to Crazy Horse.”

“This represents the same episode shown in No. 15 of the Smith record.
The horse in each instance is colored a light purple, probably a roan.

3 me " :
\
*

‘Post
=
‘

. liltmy a re Davee | bose Wes Le OF be wee
pee %
ee ee dag oi

72

10

dima fi gle — ae a Crow tu
EL So
ep Mie

D, L. PRATT, JR.

/1

11

12

NOI 5 SITTING BULL—STIRLING 55

No. 11

Same fight. Killing a Crow who dismounted and fought desperately
and wounded Bull in two places.”

3 This does not refer to the same fight pictured in No. 10. It is the same
episode, evidently, as shown in No. 12 of the Smith record and No. 4 of the
Kimball record. It was in this encounter that Sitting Bull received the wound
in his foot which crippled him for the rest of his life.

No. 12
Same.”

® This, evidently, does not refer to the same fight as depicted in No. 11.
Apparently, it represents the capture of an Assiniboin woman, probably on the
occasion shown in Nos. 5 and 6 of the Smith record.

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Nos r
Same fight. Killed two.”

% Sitting Bull is shown wearing a shirt spotted with red. His opponent is
wearing a shirt spotted with green. This may represent the fight between the
Sioux and the Assiniboines shown in Nos. 1 and 2 of the Smith record.

2

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13

BIBLIOGRAPHY

Jounston, W. FLETCHER
1891. The Red Record of the Sioux Life of Sitting Bull and History of
the War of 1890-91. Edgewood Publishing Company, Philadelphia.
De BarTHE, JOE
1894. The Life and Adventures of Frank Grouard. Combe Printing Co.,
St. Joseph, Mo.
’ Mooney, JAMES .
1897. The Ghost-Dance Religion and the Sioux Outbreak of 1890. 14th
Ann. Rep., Bur. Ethnol., pt. 2.
McLauGHLIN, JAMES
1910. My Friend, the Indian. Houghton Mifflin Co., Boston.
VESTAL, STANLEY
1932. Sitting Bull, Champion of the Sioux. Houghton Mifflin Co., Boston
& New York.

57

On IAN ‘MNISCELLANEOUS COLLECTIONS
Be ere a VOLUME s 97 NUMBER 6 kien

st
¥ it

BY
-R, E. SNODGRASS
Bureau of. Entomology and Plant Quarantine
ATS: Department of Agriculture

(PUBLICATION 3483)

CITY OF WASHINGTON
"PUBLISHED BY THE SMITHSONIAN INSTITUTION
AUGUST 23, 1938

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER 6

EVOLUTION OF THE ANNELIDA, ONYCHOPHORA,
AND ARTHROPODA

BY
R. E. SNODGRASS

Bureau of Entomology and Plant Quarantine
U. S. Department of Agriculture

(PUBLICATION 3483)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
AUGUST 23, 1938

The Lord Waltimore Press

BALTIMORE, MD., U. S. A.

EVOLUTION OF THE ANNELIDA, ONYCHOPHORA,
AND ARTHROPODA

By R. E. SNODGRASS
Bureau of Entomology and Plant Quarantine, U. S. Department of Agriculture

CONTENTS
PAGE
fee nypothietical-annelid AncestOrs 3. se eiSee ae cides obs os eenen es I
II. The mesoderm and the beginning of metamerism..................00: 9
her Development of the annelid nervous system. ... 2.602.000.0600 cees 21
NAV mrcl cl il tar aatarie incl seca avese a chavo rctc fovetars Cte vere re ont oh nlstonnte oi cca Tak utvaiere ee every rato 26
iheateloblastics OG. postlatvalesomitess 14456 eieeiitee ones 26
hesprostomiunmandeitsrappendapessmenmacs ae seedoo es eee 32
Mthesbodyeanduitsrappendasesmectmsntirmscceas coe ales cee eters 34
Meme rVOUS ASV Stellen «trees tina tetes ciel ae ae Sa ame sean res 39
IRE Mey Shweta rsterererarcrcsstoney oot teens tA Naites s1ct Soap rath voney hake SESE SP aS 45
ihe mephridiayand: theycenitalyductsaeeremiace se cee cenneeeeeee 45
Witeeibiee @) ny. chophio tans. <p se rercic foetstaera So cmacieiere Sehe iol ave Sao tia oe erorets 50
iBarlyastages ofrdevelopinentsa meres sc cine con tier oh econ doses t 52
OME TVOUSM SY SOM metysteoreeieys es secre aie trae reece ers Loe oeereee 55
AN GS: RESET, oe HORE SOS ROI TENG TOTO EISIC BCE ROR ICR IE ener © oar 62
Later history of the mesoderm and the coelomic sacs............ 62
MPhensomaticamtiscul attendees ce artic co clenaiceeiseininae oer 64
ie sepmentalmappendagesiac cine cries steelers wires cies erroiearsion 67
MEE tees IL ATO My WOMANS 5 6 thes vic cee A eactiates 2 chore sess horace ne Oe 70
shemcircilarony, Systems cents ete es Gels ooo kon a oh eee 70
BIRETG ale Poa ICRA ts ys tae scanete Sede 4n/ clade vanes scetare. Seda s alewal-e cae lave wie erecete 72
MH erODeansOr LeproaiuctiOtiase.< (5 -mirers sioc = a6 cserio ioe ola eared 74
Wale LICUPAT CLO POC an ceytee cenit ialereras ts secuntcrotorers oie sist, Socaie onic eiauka arenes 76
Barly emiryonie development .(05.,6<e/s is ee leicod Ss cmale sae URIS oe 80
Primary (And WSECONGALY \SOMULES se, oars ds = evelsie’e sale aleve Utewaade o 82
The cephalic segmentation and the development of the brain...... 89
MC IMGUTOlNtHe s EAC eacse = aise et cree oa foils ais a alerts co abate Mae 107
Coclomiclorezans oteadimlh arthropods see. eres cie oem 126
SE em taltell (OUCES ture Sete tee obetaics *, S auity since ova oie cactus sbsyelsi@laetelamieye i & 131
Wem VAGR CNEL. CONCINSIONS aj./3 oocs.00 oi dc ac0 cu alc ao.obu de oc tc eenace o0n8 132
TRXENPETREVAYOSY | S'S SO nor cRCRCERS Io Ree OE ES TE Cee EE ae I oe RE oe a 149

I. THE HYPOTHETICAL ANNELID ANCESTORS

Among the simplest of the metazoic animals that lead an active,
free existence is the planula larva of the Coelenterata. The planula
develops into a polyp or a medusa because it is a young coelenterate,
but, so far as its structure goes, it contains the fundamental building

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97, No. 6

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

elements that, with the appropriate hereditary influences, might be
fashioned into a flatworm, an annelid, an arthropod, a mollusk, or
a vertebrate.

The typical planula is a minute oval or elongate creature (fig.
1 A, C) consisting of an outer layer of ectoderm cells, and an inner
mass of endoderm cells. The planula, therefore, represents the
gastrula stage of embryonic development, though it may have no
enteric cavity and no blastopore. Its motor mechanism is a covering

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Fic. 1.—The coelenterate planula, and two methods of endoderm formation.

A, planula of Sympodium corraloides (from Kowalevsky and Marion, 1883).
B, blastula of Carmarina fungiformis, showing differentiation of endoderm from
ectoderm by delamination of blastoderm cells (from Metschnikoff, 1882). C-F,
formation of endoderm by internal proliferation from posterior pole of planula
(from Hatschek, 1888, after Claus).

Blc, blastocoele; Bld, blastoderm; Ecd, ectoderm; End, endoderm.

of vibratile cilia. The embryology of the planula is very simple. The
cleavage of the coelenterate egg produces a morula, and the morula
becomes a blastula. In the succeeding planula stage the inner endo-
dermal cell mass is formed, but it is not certain that gastrulation
takes place by simple invagination in any of the coelenterates. With
some forms the endoderm arises as an inward migration of scattered
cells from the blastoderm; in others the blastomeres divide regularly
each into an outer ectoderm cell and an inner endoderm cell (fig. 1 B) ;
but the most common method of endoderm formation is the internal

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 5

proliferation of cells from the posterior pole of the blastula (D, E, F),
and this last process suggests that it is an embryonic modification
of gastrulation by invagination. When, shortly, the planula settles
to the bottom of the water and becomes attached preparatory to its
metamorphosis into a polyp, a stomach cavity appears in the endo-
derm, and a mouth cavity breaks through at the free pole.

The development of the planula shows clearly that there is in
ontogeny no fixed method for the formation even of so important
an organ as the stomach. The effective thing in embryonic develop-
ment is the inherited organizing property resident in the egg that
converts a mass of cells, however formed, into a definite functional
structure. The same principle, as we shall see later, applies also to
the development of the annelids and the arthropods, for in these
animals there is so much apparent irregularity in the formation of
the germ layers that attempts to interpret all observed facts in terms
of cell genealogy lead only to confusion. Ontogeny and phylogeny,
therefore, while they produce the same end results, may follow quite
different methods of procedure. In phylogeny we must visualize the
successive stages in the evolution of an animal as free-living adult
forms, each structurally adapted for performing the functions of an
independent animal.

If the coelenterate planula were an adult animal instead of a
temporary larval form, or if it had to maintain itself for any con-
siderable length of time, it almost certainly would have a stomach
cavity and a mouth. Thus modified, as it is later in its own develop-
ment, the planula would be an independent, motile gastrula, having
a stomach in the form of an open pocket of the blastoderm for the
retention of food particles (fig. 2 A). An animal of this simple type
of structure, we must suppose, was the actual ancestor of the polyp
and medusa forms of the Coelenterata ; but equally well it might have
been the progenitor of the annelids, and through the latter of the
arthropods. Various writers on phylogeny have proposed an origin of
the segmented worms direct from a coelenterate polyp, but it should
be recognized as a fundamental principle in evolution that a special-
ized type of animal does not give rise to another specialized type—if
two forms are related, they are related through some simple common
ancestor. This principle as applied to the coelenterate derivatives is
expressed by Ziegler (1898), who says:

It is to be supposed that the higher animals derived from coelenterate stock took
their origin not from the highly specialized forms of the Coelenterata, such as
the anthozoans and ctenophores, but from a planula-like or gastrula-like ancestral
form of the coelenterates.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

The theoretical planulalike gastrula postulated above as the com-
mon ancestor of the Coelenterata and the Annelida (fig. 2A) pre-
sumably swam habitually in one direction by means of a covering of
cilia, and the mouth, or blastopore, was at the posterior pole where
food particles might be swept into the stomach with the eddy of
currents converging to the rear.

In the ontogenetic development of the annelids, gastrulation
generally takes place by epiboly, which is the overgrowth of the endo-
derm by the ectoderm, and the primary open blastopore is at the
posterior pole of the embryo. There is no reason why this ontogenetic
stage should not represent an early phylogenetic stage, and one iden-
tical with the gastrula ancestor of the Coelenterata (fig. 2 A). With
the further development of the annelid embryo, however, the blasto-
pore elongates forward on the ventral surface of the gastrula (fig.
2F) until its anterior end comes to be near the anterior pole (G);
but, at the same time, the lips of the blastopore grow together from
behind forward, leaving finally only the anterior end open into the
archenteron, and this opening is the primitive mouth (H, Mth).
Secondarily, an anal aperture (An) is formed later at the original
posterior end of the blastopore on the caudal extremity of the embryo.
The endodermal archenteron of the annelid thus becomes a simple
alimentary canal having the oral aperture located ventrally near the
anterior end of the body, and the anal aperture situated terminally
at the posterior end.

If we visualize the change in the position of the blastopore as an
event in the phylogenetic history of the annelids, we must see a corre-
lated change in the habits of the animal. The actively swimming
gastrula (fig. 2 A) in its search for food, we may suppose, took to
brushing over the surfaces of stones or aquatic plants (B), where
food particles were more numerous and more easily obtained. For
this manner of feeding, a ventrally placed blastopore would be a
distinct advantage, or, even more efficient, a blastopore drawn out
lengthwise on the under surface (C). With a form thus modified in
habits and structure, there may easily have developed a creeping
habit, and an adaptation of the ventral cilia for progression on solid
surfaces (D). Finally, then, came a more complete adaptation to
feeding on a subsurface, resulting in an elongate flattened body, and
the establishment of an alimentary canal with a ventral mouth and
a terminal anus (E) produced by the closure of the intermediate
part of the blastoporic slit.

A creeping mode of locomotion may be subserved entirely by a
ciliary coating of the body wall, as is shown in the Platyhelminthes,

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 5

but a creeping animal encounters irregularities and obstructions. A
provision for body movements, therefore, becomes an advantageous
adjunct to the motor mechanism, and such movements can be pro-
duced only by an internal muscular system. Hence, the next stage
in evolution, recorded in both the flat worms and the annelids, was

SK!
Fi de RE UP Cy
" . ee

SRN
Ss

TRESS
SSS

hs

Fic. 2—Hypothetical evolution of a swimming planulalike creature with an
open gastrocoele into a creeping wormlike animal with a simple alimentary
canal, a subapical ventral mouth, and a terminal anus.

A, primitive swimming form with posterior blastopore. B, the same having
acquired the habit of sweeping up food particles from a solid surface. C, blas-
topore elongated forward on surface of contact to accommodate the feeding
habit. D, the same more fully adapted to subsurface feeding. E, final develop-
ment of alimentary canal, with ventral mouth and terminal anus, formed by
closure of intermediate part of blastopore, creeping habit fully established.
F-H, three stages of elongation and closure of the blastopore, ventral view.

AlCnl, alimentary canal; An, anus; Bpr, blastopore; Gc, gastrocoele, or
archenteron; Mth, mouth.

the development of contractile tissue that conferred the power of
diversified adjustive movements on the body itself. Muscles, how-
ever, are not generally automatically active, as are cilia, and hence
the development of muscle tissue is usually accompanied by the
development of a mechanism for its activation. Furthermore, since
a muscular system is a provision for adjustment to external con-
ditions, the source of its stimulus must come from the environment.
The sponges are said to have a primitive contractile tissue that is

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

stimulated directly by environmental changes; in all other animals
there is intimately associated with the contractile muscle tissue a
specifically receptive and conductive nerve tissue, through which
environmental stimuli become effective on the muscles. Finally, the
high metabolic rate of muscular activity creates the need of special
excretory organs for the removal of waste products from the body.

The genesis of contractile and conductive tissues, and their inte-
gration into a neuromuscular system are best seen in the Coelenterata.
Contractility, being a common property of protoplasm, may become
localized and specially developed in a particular part of any cell of
the body in a primitive animal. In the coelenterates fingerlike muscle
processes are produced from the inner ends of cells in both the
ectodermal and the endodermal epithelium, those of the ectoderm
(fig. 3, mp) taking a longitudinal course, those of the endoderm a

Cnb SCl

\ / |
NCI mf mp NC1. SNC1 mf mp
Fic. 3.—Diagram of the ectodermal neural and muscular elements of Hydra.
(From Curtis and Guthrie, 1927.)

Cnb, cnidoblast; Ecd, ectoderm; mf, muscle fiber; mp, muscle process of
epithelial cell; NCJ, neural cell; NSC/, neurosensory cell; SC/, sensory cell;
SL, supporting lamella.

transversely circular course. Fibrils of contractile tissue (mf) be-
come differentiated in these processes. In the hydra, the body of the
muscle cell remains as a part of the epithelial layer, but in some of
the other coelenterates the entire cell may be withdrawn beneath the
surface and converted into a muscle fiber. A primitive nerve cell is
an epithelial cell in which the common protoplasmic properties of
irritability and conductivity are specially developed both in the cell
body and in branching processes given off from the latter, but the
nerve cells become differentiated into superficial receptive cells and
deeper-lying conductive cells. In the hydra the receptive cells (fig. 3,
SCI) and the sensory cells (SNCT7) contained in the ectoderm have
connections, on the one hand, with the surface of the body, and, on
the other, send branches to the strictly neural cells (NCJ), which are
distributed through the inner parts of the ectoderm, and in turn
send branches to the muscle processes of the muscle cells. The endo-

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—S NODGRASS 7

derm of the coelenterates, though its cells have numerous muscle
processes, contains relatively few sensory and neural cells, and fibrous
branches of these cells are but little developed.

The polychaete annelid larva of the trochophore type (fig. 8) has
a muscular system of which the elements appear to be quite analogous
to the ectodermal muscles of the coelenterates, though the system
itself is carried to a higher degree of development. Furthermore,
the larval muscles are parts of a neuromuscular system, since gener-
ally they follow the inner surfaces of nerve tracts in the ectoderm.
The muscle fibers are formed from cells derived directly from the
larval ectoderm, along with numerous small undifferentiated cells
that constitute a loose layer of mesenchyme distributed through the
haemocoele. The fibers are arranged principally in longitudinal and
circular tracts, though some of them extend from the body wall to
the alimentary canal. The endoderm of the larva does not produce
directly either muscular or neural cells. The nervous system of the
polychaete larva, when best developed, consists of longitudinal and
circular strands of ectodermal nerve cells and fibers following the
muscle tracts, and of ganglionic groups of nerve cells developed
particularly in connection with sensory organs on the preoral part
of the body. The larval elaboration of the neuromuscular system is
largely a temporary adaptation to the specialized form and habits of
the trochophore, for most of it is lost when the larva undergoes its
metamorphosis into the definitive worm form; but the preoral part
of the larval nervous system forms the brain of the adult, and some
of the larval muscle fibers are taken over into the definitive muscular
system.

If now we endow our hypothetical annelid ancestor (fig. 2 E) with
a primitive neuromuscular system derived from the ectoderm, and
provide it with a pair of primitive nephridia, it will have reached an
evolutionary stage entirely comparable in structure with that of
an annelid in the ontogenetic stage of the young polychaete larva.
The usual trochopore larva of the Polychaeta (fig. 4 A), however,
leads a purely pelagic life; it floats upright in the water and swims
by means of bands of cilia that encircle the body. Its radial and
circular neuromuscular system appears to be entirely adapted to its
upright position, and many zoologists have regarded the trochophore
as the ancestral form of the annelids as well as of various other
invertebrates. The lateral position of the mouth, however, just
below the principal circle of cilia (Mth), gives us good reason for
suspecting that the shape of the trochophore and the position assumed
in the water are secondary adaptations to a brief swimming existence ;

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

in fact, the later horizontal development of the worm form along
the vertical axis of the larva shows clearly that the trochophoral
position is one quite out of harmony with the general organization
of a worm.

The trochophore, therefore, is to be regarded as a temporary,
specialized larval form in polychaete ontogeny, adapted to a free
pelagic life for the purpose of disseminating the individuals of its
species. The metamorphic alterations that it undergoes at its trans-
formation to the worm are changes of a nature that could not have
been a part of the phylogenetic evolution of any animal. On the

ApPl

Fic. 4——The polychaete trochophore and the crustacean nauplius, two spe-
cialized larval forms of an early ontogenetic stage, having, therefore, primi-
tive characters, but no phylogenetic significance in their shape or general
structure.

A, typical structure of a trochophore, diagrammatic. B, nauplius of a cirriped,
Alcippe lampas Hancock, dorsal surface (from Kuhnert, 1935).

An, anus; 1Ant, first antenna; 2Ant, second antenna; ApGung, apical ganglion;
ApPl, apical plate; CNv, circular nerve; Epsp, episphere; Hpsp, hyposphere;
lh, lateral horn; LNv, longitudinal (radial) nerve; Md, mandible; Ment,
mesenteron; MsT, mesodermal teloblast; Mth, mouth; Nph, nephridium; O,
naupliar ocellus; Patr, paratroch; Proc, proctodaeum; Prtr, prototroch; Stom,
stomodaeum.

other hand, inasmuch as the trochophore is an early ontogenetic stage,
its general organization is primitive, and is repeated in the onto-
genetic development of many other invertebrates besides the annelids.
It should be noted, furthermore, that the trochopore is not a uni-
versal larval form even among the annelids, for most of the archi-
annelids, some of the polychaetes, and all the oligochaetes have a
direct development, in which either there is no suggestion of the
trochophore form, or a remnant of it is preserved from ancestors
that had a typical swimming larva.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 9

The presence of a mesoblastic muscle system and of a mesenchyme,
or parenchymatous layer between the ectoderm and the endoderm,
gives the annelid larva, or the platyhelminth adult, the status of a
triploblastic animal; but the middle layer is here only an elaboration
of elements present also in the so-called diploblastic coelenterates.
The young annelid larva, however, is endowed from its parents with
hereditary influences that will mold its growing tissues into structures
never attained by the coelenterates or flat worms. Particularly affected
are two individualized groups of mesoblast cells, which, though they
may be set apart in the platyhelminths, will give rise in the annelids
to special bands of mesoblastic tissue, known as the mesoderm.
Within the mesoderm will be formed a new body cavity, the coelom,
and from the walls of the latter will be produced a new muscular
system, a more efficient excretory system, a circulatory system, and
various tissues of special functions, to all of which is added an exten-
sion and elaboration of the nervous system. With the formation of
the mesodermal cavities the triploblastic annelid larva becomes a
coelomate animal, but, shortly before the appearance of the coelom,
there takes place a segmentation of the body affecting the ectoderm
and the mesoderm, so that the young annelid worm is almost at once
a segmented and a coelomate animal.

Il. THE MESODERM AND THE BEGINNING OF METAMERISM

In the ontogeny of the articulate animals, the formation of the
coelomic cavities in the mesoderm is so closely associated with the
appearance of body segmentation as to give the impression that the
two are intimately related developmental processes, and since the
segmentation of the mesoderm is usually more conspicuous than the
segmentation of the body, embryologists often describe metamerism
in terms of mesoderm segmentation, as if the formation of “meso-
derm somites” were equivalent to body segmentation. Closer atten-
tion recently given to the sequence of events in the development of
the Polychaeta, however, shows that metamerism begins in the ecto-
derm and the primary ectodermal musculature, and that it secondarily
effects a division of the coelomic mesoblast into segmental sections.
Subsequently, the coelomic cavities are formed in the segmented
mesoderm. That coelomic sacs do not determine metamerism is
shown also by the formation of paired coelomic cavities in the preoral
cephalic mesoderm of the Onychophora and Arthropoda, in which
there is no corresponding external segmentation.

Metamerism, therefore, probably took its origin in a subdivision
of the primary somatic musculature into successive sections (myo-

Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

tomes) to give greater efficiency to body movement. The segmen-
tation of the ectoderm and the mesoderm then followed as a result of
the segmentation of the muscular system. The primitive coelomic
cavities were probably spaces formed’ in the mesoderm for the accu-
mulation of waste products in the body fluid, to be discharged through
primitive nephridial tubules. The coelomic mesoblast, however,
formed also a secondary musculature that reinforced the primary
musculature, and which, in the higher arthropods, has completely
replaced the latter. Evidence that such has been the course of evolution
in the Articulata will be shown in the following discussion of the
early stages in annelid ontogeny; but there still remains the question
as to the origin and nature of the primitive mesoderm, which antedates
metamerism.

A study of the growth and differentiation of the annelid mesoderm
takes us into the later part of larval development, but to obtain light
on the origin of the middle germ layer we must go back to an earlier
ontogenetic stage. During cleavage of the annelid egg most of the
yolk remains consistently in the blastomeres situated on the vege-
tative surface of the blastula (fig. 5 A), with the result that, in the
64-cell stage, there are 8 large, yolk-filled blastomeres at the posterior
pole (B). These cells are designated by embryologists 44, 4B, 4C,
4D, and 4a, 4b, 4c, 4d, since they comprise the so-called macromeres
of the fourth generation and the fourth quartet of micromeres. All
of them at this stage would appear to be endodermal, and at the time
of gastrulation they all become internal, owing to their overgrowth
by the ectoderm. Seven of them, in fact, give rise to purely endo-
dermal progeny, but the 4d cell will form in most cases both endo-
derm and mesoderm. The first cleavage of 4d produces two bilaterally
symmetrical cells, 4d’ and gd? (C), and these cells, in their immedi-
ately following divisions, give rise to a few very small cells (D, end),
usually regarded as endoderm cells, and a pair of large cells (MsT)
that are destined to produce the coelomic mesoblast, and hence con-
stitute the mesodermal teloblasts. (It is perhaps possible that the
small ‘““endoderm”’ cells of this generation are the primary germ cells.)

The common occurrence in the annelids of mesodermal teloblasts
derived from cells closely associated with the endoderm has given
rise to the idea that the coelomic mesoblast is of endodermal origin,
and for this reason it is often called the “endodermal mesoblast” to
distinguish it from the larval mesoblast, which is derived from the
ectoderm. In most animals the mesoderm is, one way or another,
associated in its origin with the endoderm, but among the annelids
there are many cases where its endodermal connection is not evident.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS II

Eloise

nee

Fic. 5.—Late cleavage stages and mesoderm formation in Annelida and
Platyhelminthes.

A, diagram of posterior pole of annelid blastula showing four yolk-filled
“macromeres” of third generation. B, posterior pole of blastula of Arenicola
cristata Stimpson after next cleavage forming fourth quartet of ‘‘micromeres,”
showing differentiation of 4d blastomere (adapted from Child, 1900). C, same,
after cleavage of 4d into 4d* and 4d° (adapted from Child, 1900). D, blastula
of Podarke obscura Ehlers, showing mesodermal teloblasts (MsT) derived
from blastomeres 4d’ and 4d° (C) after separation of small endoderm cells
(from Treadwell, 1901). E, posteroventral view of 40-hour embryo of Podarke
obscura with mouth and anus, showing position of mesoderm bands (Msd) in
body (from Treadwell, 1901). F, optical frontal section of embryo of Capitella
capitata Fabr., showing mesodermal teloblasts and rudiments of mesoderm
bands (from Eisig, 1899). G, optical section of embryo of Dinophilus sp., with
mesoderm bands (Msd) extending forward from teloblasts (from Nelson,
1904). H, optical section of blastula of Planocera inquilina Wheeler (Poly-
cladia) from right side just after division of 4d, producing 4d’ that will form
endoderm, and 4ad* that will form mesoblast (from Surface, 1907). I, same,
later stage seen from posterior pole, showing mesoblast (Msb*) derived from
4d’, and mesoblast (Msb*) derived from second quartet of ectodermal blastomeres
(from Surface, 1907).

3A-3D, 4A-4D, “macromeres” of third and fourth generations; 4a-4d, “micro-
meres” of fourth quartet; An, anus; Bpr, blastopore; Br, brain; 4d’, 4d’,
daughter cells of 4d blastomere; Ecd, ectoderm; End, endoderm; end, endoderm
derived from 4d' and 4d* blastomeres; Msb*, mesoblast derived from second
quartet of micromeres; Msb*, mesoblast derived from 4d cell of fourth quartet ;
Msd, mesoderm; Msnc, mesenchyme; MsT, mesodermal teloblast; Mth, mouth.

,

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

It is claimed by both Kleinenberg (1886) and E. Meyer (1901), for
example, that in the larva of Lopadorhynchus the mesoderm arises
from the ectoderm, and in Capfitella, according to Eisig (1899), the
coelomic mesoblast is produced from blastomeres other than 4d.
Furthermore, the mesoderm of the postlarval somites is said by
Iwanoff (1928) to be formed in many polychaetes directly from the
ectoderm, and the same is probably true in cases of regeneration. The
mesoderm of certain other coelomate invertebrates also may have no
genetic relation to the endoderm, as in the gastropod Paludina, in
which the embryonic mesoblast that gives rise to the usual mesodermal
organs is generated directly from cells of the ventral ectoderm (see
Dautert, 1929).

During larval life, or at the transformation of the larva to the
worm, the annelid mesodermal teloblasts, however formed, proliferate
within the haemocoele two masses of mesoderm cells (fig. 5 E, F, G,
Msd), which eventually take the form of ventrolateral bands extend-
ing forward at least as far as the sides of the mouth (fig. 6 F). These
primary mesoderm bands are solid cell masses; they are never ob-
served at this early stage to contain cavities, and there is no evidence
from annelid embryology to suggest that they represent phylogeneti-
cally a pair of open pouches. Later, with body segmentation, the
bands are broken up into solid segmental blocks (G), and finally the
blocks are excavated by coelomic cavities (H). The nature of the
mesoderm and the primitive function of the coelomic cavities can be
better discussed after we have examined the known facts concerning
the beginning of metamerism, but it should be noted here that the
formation of the mesoderm bands precedes body segmentation.

Metamerism in the polychaete larva becomes first evident as a
subdivision of the body region between the mouth and the pygidium
into a small number of somites (fig. 7 A, J, IJ, III). There is ample
reason to believe, as Iwanoff (1928) claims, that the formation of
these primary somites, or larval segments of ontogeny, represents the
beginning of metamerism in phylogeny, and, as we shall see, the same
phenomenon of direct segmentation in the body of the embryo or
young larva recurs in various arthropods. The primary somites are
thus to be distinguished from the secondary somites later added by
teloblastic growth in a subterminal generative zone, and which will
constitute the major part of the adult animal. The larval somites of
the Polychaeta, Iwanoff shows, are formed approximately simul-
taneously in contrast with the successive, individual generation of
the teloblastic somites. E. Meyer (1901) observes that in Lopa-
dorhynchus metamerism takes place so rapidly as to give the impres-

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 13

sion that a relatively large number of somites are formed all at once,
but Sokolow (1911) says that in Ctenodrilus the intermediate somites
or the more anterior ones are first differentiated and the series then
completed anteriorly and posteriorly. Segmentation may be delayed
until the beginning of metamorphosis, as in Polygordius, or it may
take place while the larva is still in the swimming trochophore stage.

Fic. 6.—Transformation of the annelid blastopore, primary segmentation of
the body, growth and segmentation of the mesoderm bands, and formation of
the coelomic sacs, diagrammatic.

A, blastopore and mesodermal teloblasts at posterior pole of embryo. B,
blastopore elongating forward on ventral surface; rudiments of mesoderm gen-
erated from teloblasts. C, blastopore still more elongate, closing posteriorly ;
mesoderm growing forward. D, blastopore closed posteriorly; mesoderm bands
extended to prostomium. E, blastopore obliterated except for mouth opening
at anterior end; anus formed secondarily at posterior end; mesoderm segmented
following metamerism of body, and extended into prostomium. F, polychaete
trochophore before segmentation. G, same after segmentation, mesoderm cut
into solid segmental blocks. H, same, mesoderm blocks excavated by coelomic
cavities.

AlCnl, alimentary canal; An, anus; Bpr, blastopore; Coel, coelomic cavity ;
Epsp, episphere; Msd, mesoderm; MsT, mesodermal teloblast; Mth, mouth;
Prst, prostomium; Pyg, pygidium; ZG, zone of growth.

In Polynoé, as described by Hacker (1895), seven somites are first
marked out in the body of the trochophore, which is transformed
while still active into a swimming “nectochaete” larva with seven
segments and corresponding chaeta-bearing parapodia. The number
of larval somites is always small, three or four being usual (fig. 7 A,
B, C), the maximum not more than 13. Completion of larval meta-
merism is followed by a pause in development.

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Observations on the beginning of embryonic segmentation in the
annelids differ somewhat as to whether the intersegmental divisions
appear first in the ectoderm or in the mesoderm, but most students
of annelid development find either that the ectoderm and the meso-

Pys

Fic. 7—Examples of primary segmentation in polychaete larvae.

A, Eupomatus uncinatus, trochophore showing primary segmentation of the
mesoderm (from Iwanoff, 1928). B, same, later larval stage, horizontal sec-
tion showing development of chaetal sacs in primary somites, and extension of
posterior part of body (from Iwanoff, 1928). C, Platynereis dumerilii Aud.
& Milne-Edw., nereidogen larva just out of egg, with four primary somites
(from Hempelmann, 1911). D, Capitella capitata Fabr., embryo before seg-
mentation, ventral view (from Eisig, 1899). E, same, embryo with seven
somites and zone of growth formed directly in primary body region (from
Eisig, 1899). F, same, later stage with two additional somites formed from
zone of growth (from Eisig, 1899). ;

E, eye; [-IX, somites; Mth, mouth; Prst, prostomium; Pyg, pygidium; SP,
somatic plate; 7/, tentacle; ZG, zone of growth.

derm are segmented at the same time, or that the first signs of meta-
merism are to be seen in the ectoderm.

In the development of the polychaete Capitella, according to Eisig
(1899), on the sixth day after fertilization of the egg, the cells of the
ventral somatic plates of the larva (fig. 7D, SPl) become arranged
in transverse rows, and on the sixth day seven or eight somites are
already demarked by transverse lines in the ectoderm of the larval

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 15

body region between the mouth and the pygidium (E). On the same
day, however, the mesodermal bands also become divided into seg-
mental sections. At first the ectodermal and mesodermal somites of
Capitella do not entirely correspond, there being several super-
numerary mesodermal divisions in the mouth region, but by the
twelfth or thirteenth day the larva has 13 somites with coincident
limits in both the ectoderm and the mesoderm.

The segmentation of the mesoderm bands as described by E. Meyer
(1901) in Psygmobranchus, Polygordius, and Lopadorhynchus ap-
pears to be determined by elements of the mesenchymatic primary
mesoblast in the form of spindle-shaped cells that penetrate into the
mesoderm bands at the intersegmental lines and cut the bands into a
series of segmental sections. From the penetrating mesenchyme cells
are later formed, according to Meyer, the muscles of the interseg-
mental dissepiments. Similarly in the Serpulidae and Spionidae the
larval segmentation is said by Iwanoff (1928) to be secondarily im-
posed upon the mesoderm bands by metamerism in other parts of the
body, as by the ectodermal segmentation, the ingrowth of the chaetal
sacs (fig. 7 B), the penetration into the mesoderm of mesenchymatous
muscle elements, or by the segmental formation of blood lacunae in
the general mesoderm mass.

The primary larval segments are seldom as fully developed in the
adult worm as are the teloblastic segments, and both the segment
limits and the differentiation of ganglia on the nerve cords may remain
obscure. In the Spionidae, Iwanoff (1928) says, the trochophoral
mesoderm is very weakly developed, the dissepiments are only imper-
fectly formed, often absent, and in some species a segmentation of
the mesoderm in the primary segments is absent even in the adult.
Chlorogogen cells are not developed in the coelomic walls of the larval
segments, and in these segments germ cells are never present.

As a result of body metamerism, the mesoderm bands are divided
each into a series of segmental sections, and these sections, as the
bands themselves, are at first solid blocks of cells (fig.6 G). Later the
coelomic cavities appear as cleavage spaces within the cell blocks (H).
Hence, just as there is no evidence that the primary mesoderm bands
represent primitive sacs, so there is no evidence from ontogeny that
the coelomic cavities of the annelids took their origin as a series of
separate mesodermal pouches. The facts of development suggest only
that the primitive mesoderm bands were continuous tracts of cells,
and that the formation of cavities within them was a secondary
process, subsequent to segmentation.

With the formation of the coelomic cavities in the mesoderm, the
young annelid becomes a coelomate animal. Before the appearance

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

of the coelom, however, it might pass for the ancestor of a flatworm,
for even in the Platyhelminthes there is a teloblastic proliferation of
cells that appear to correspond with the mesoderm cells of the anne-
lids, though the cells thus produced soon disperse and become a part
of the parenchyma. It is in the development and elaboration of the
mesoderm, or teloblastic mesoblast, therefore, that the Coelomata
depart from the Platyhelminthes. Segmentation is a feature super-
imposed upon the mesoderm in the Annelida as a result of body
metamerism, in which apparently the ingrowth of the septal muscles
plays an important part.

The mesoderm of the adult annelid or arthropod gives rise to a
large variety of tissues and organs, but most of the specialized deriva-
tives of the mesoderm are formed in the secondary segments of the
adult animal. The principal products of the primary mesoderm are
muscle and connective tissues, and an epithelial lining of the coelomic
cavities.

According to E. Meyer (1901), the mesodermal myoblasts of the
polychaete larva are not recognizable as such until the mesoderm
bands have become broken up into segmental sections, and the trans-
formation of the myoblasts into functional muscle fibers is not evident
until after the appearance of the coelomic cavities. The myoblasts of
each mesodermal segment, Meyer says, consist of four large cells on
each side, two dorsal and two ventral, lying along the lines of the
larval longitudinal muscles of mesenchymatic origin already present.
The mesoderm fibers finally replace the mesenchyme fibers and be-
come the definitive longitudinal muscles of the worm. The coelomic
myoblasts, Meyer shows, are true epithelial muscle cells that form
muscle processes from their outer surfaces, while the plasmatic bodies
of the cells retain their places for some time in the coelomic walls.
The parts of the coelomic walls not involved in muscle formation be-
come thinner, and finally transform into typical peritoneal epithelium.

The important part that the mesoderm plays in the development of
the coelomate animals is entirely clear; but what the mesoderm be-
comes in the course of evolution does not explain what it was in its
beginning. Most of the theories that have been proposed to account
for the primitive mesoderm, it will be found, are attempts to explain
the functional nature of the coelomic sacs rather than the origin of
the mesoderm itself.

The theory most widely accepted at one time as to the origin of
the mesoderm is the enterocoele theory (Hertwigs, 1882, Sedgwick,
1884), by which the coelomic sacs are explained as diverticula of the
archenteron. In some animals the coelomic sacs are thus formed in

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 17

the embryo, and the enterocoele theory has some plausibility as a
wide generalization, considering the very common early association
of the coelomic mesoderm rudiments with the endoderm; but, as
applied to the annelids and arthropods, the theory must entirely discard
the direct evidence from embryology that the mesoderm first appears
as solid proliferations of cells, which only in a purely hypothetical
manner could be interpreted as representing pouches of the archen-
teron. The only known case of the formation of the mesoderm from
enteric pouches that might be referred to the articulates occurs in
the Tardigrada (see Marcus, 1929), but there is much uncertainty
concerning the relationships of the tardigrades.

A second mesoderm theory is the gonocoele theory, based on the
almost universal association of the germ cells with the coelomic meso-
derm in the coelomate animals. Hatschek (1877, 1894) believed that
the mesodermal teloblasts of the annelid larva are themselves germ
cells, and Rabl (1879, 1889) adopted this view. The gonocoele theory
of the origin of the coelomic sacs, however, was principally elaborated
by E. Meyer (1891, 1901). Meyer contended that the primitive
coelomic sacs were muscular pouches, from the epithelial walls of
which the germ cells are generated, and that, as these gonadial sacs
expanded to increase the reproductive function, they finally preémpted
the haemocoele, and their muscles were transferred to the body wall.
The gonocoele theory loses much of its support now that the old belief
that the germ cells are direct products of the coelomic epithelium
is no longer tenable, and, moreover, it entirely breaks down in view
of the fact that the primary larval somites of the annelids do not
contain germ cells. In the primitive annelids, as will be shown later,
the germ cells probably were located in the zone of undifferentiated
tissue behind the last primary somite. If so, the reproductive function
had nothing to do with the origin of the mesoderm or the formation
of the coelomic sacs.

A third theory, concerned principally with the function of the
coelomic sacs, is the nephrocoele theory (Ziegler, 1898; Faussek, 1899,
Ig01). According to Ziegler, the primitive coelomic cavities were
- open pouches for the accumulation of waste products; they were not
diverticula of the archenteron, but were, perhaps, of the nature of
protonephridia. The nephrocoele theory as modified by Faussek holds
that the excretory coelomic sacs are not primitive structures in a
phylogenetic sense, but that they have been developed for excretory
purposes in the embryo, and are hence purely ontogenetic organs.
Faussek supports his theory with the generalization that the open
metanephridia constitute exits from the coelom, while the closed

2

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

protonephridia serve for removal of waste products from the haemo-
coele. This statement, however, is not entirely true, for in some of
the Polychaeta protonephridia are associated with coelomic sacs, and
the primary larval somites of the annelids do not have metanephridia.
On the other hand, there can be no question that the coelomic fluid
does contain waste products of metabolism.

A fourth theory, that of Kleinenberg (1886), identifies the primi-
tive mesoderm with muscle tissue, and is thus more satisfactory than
the other theories because it deals with the beginning of the meso-
derm as a functional tissue. Kleinenberg attributes the idea of a
muscle origin for the mesoderm to Rabl, who later discarded it, but
the theory rests principally on Kleinenberg’s studies of the develop-
ment of Lopadorhynchus. Kleinenberg claimed that in the larva of
Lopadorhynchus the mesoderm is derived directly from the ectoderm
at the posterior end of the body, and that the ectodermal myoblasts,
and the neuroblasts of the ventral nerve cords, arise from a common
neuromuscular rudiment. The mesoderm bands, or “muscle plates,”
become divided into segmental myotomes consequent on metamerism
of the body, and the myotomes give rise to the body musculature,
including, according to Kleinenberg, the dorsal and ventral longi-
tudinal muscles, the parapodial muscles, and the circular muscles of
the body wall. Then follows a separation of the muscle plates into
parietal and visceral layers in each somite, producing thus the paired
coelomic cavities, the peritoneal linings of which are formed by the
inner cells of the myotomes. Kleinenberg’s theory of the origin of
the mesoderm thus gives to metamerism a mechanical significance,
since it explains body segmentation as an adaptation to more efficient
locomotion. Certainly, when once established, the chief function of
metamerism is effective movement of the body, and to this feature
the segmented annelids owe their superiority over the unsegmented
flatworms. A serious weakness of the muscle theory of the origin
of the coelomic mesoblast, however, is found in the fact that so many
tissues other than muscle are evolved from it. Muscle fiber is a highly
specialized tissue, and it seems hardly likely that epithelial tissue, for
example, would be formed from muscle cells, since ordinarily it is
epithelial tissue that gives rise to muscle fibers and to the various
other specialized tissues of the body. Furthermore, as shown by
Meyer (1901), muscle is not formed from the coelomic mesoblast of
Lopadorhynchus until after the segmentation of the mesoderm bands
and the formation of the coelomic cavities.

The literature of annelid morphology is replete with discussions
on the nature and difference of the “two kinds of mesoblast”; but

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 19

the facts concerning the ontogenetic origin of the annelid mesoblast
apparently can be expressed in the simple statement that mesoblastic
tissue may be formed by internal proliferation from any part of the
blastoderm, and may, therefore, be both “ectodermal” and “endo-
dermal.” The mesoblast of the first three quartets of the blastula
(see Torrey, 1903) gives rise to the so-called larval mesoblast, or
mesenchyme; from the fourth quartet ordinarily arises the coelomic
mesoblast, or mesoderm. That these two groups of mesoblast cells
primarily have the same morphological status is indicated by the fact
that in the Platyhelminthes they do not become differentiated into
separate tissues. Surface (1907), who first followed the divisions of
the 4d cell in a flatworm, shows that in Planocera the 4d blastomere
gives rise to both endoderm and mesoblast as it does in the annelids,
since, of the two cells of the first division, 4d' (fig. 5H) forms the
endoderm (I, End), and 4d? gives rise to two lateral groups of scat-
tered mesoblast cells (I, Msb*), which are at first quite distinct from
the mesoblast of the second quartet (Msb*), though eventually they
intermingle with the latter to form the parenchymatous tissue of the
adult. In Planocera the usual endodermal “macromeres” degenerate
and almost the entire endoderm proceeds from the 4d* cell. Finally,
we may correlate the “double origin” of the annelid mesoblast with
the production of muscle tissue from both the ectoderm and the
endoderm in the Coelenterata.

From the condition in the Platyhelminthes, it becomes evident that
the primitive mesoblast was a parenchymatous mass of undiffer-
entiated cells occupying the haemocoele, which had been proliferated
internally from both the ectoderm and the endoderm. In the unseg-
mented ancestors of the annelids, the ectodermal mesoblast must have
formed a primary somatic muscular system, represented by the larval
musculature of modern annelids, which is derived from the ectodermal
quartets of the blastula. The principal part of the parenchyma, there-
fore, came to be that part of the mesoblast proliferated in the posterior
part of the body, chiefly, or usually, from the 4d cell of the fourth
quartet. The persistent parenchyma thus became the embryonic
middle layer known specifically as the mesoderm.

Since the most important result of metamerism is the production
of a mechanism of movement based on the division of the body into
consecutive motor units, it can scarcely be questioned that meta-
merism had its origin as an adaptation to more effective body move-
ment. Inasmuch as the evidence from embryonic development shows
that metamerism originates ontogenetically in the ectoderm and its
derivatives, and is secondarily imposed upon the mesoderm, we may

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

suppose that it took its inception phylogenetically from an attachment
of the primary (ectodermal) longitudinal somatic muscles at con-
secutive rings on the body wall, and from the accompanying ingrowth
of fibers that formed contractile dissepiments between the myotomes.
The ingrowth of the septal muscles cut the parenchymatous meso-
dermal bands into segmental blocks. This modification and elaboration
of the primitive muscular system, and the consequent segmental
division of the mesoderm bands, give at once the essential quality of
metamerism, and from it there follows as a necessary result the
metamerization of other organs, such as external ectodermal struc-
tures, the ventral nerve cords, and all structures of mesodermal origin.

The coelomic cavities first appear in the annelid embryo or larva
as cleavage spaces in the segmental mesoderm blocks. Since the un-
segmented Platyhelminthes have nephridial organs, it may be assumed
that the primitive annelids possessed simple segmental nephridia in
the form of internally closed tubules extending into the haemocoele.
The primitive coelomic cavities, therefore, were probably spaces
formed in the segmented parenchyma for the accumulation of body
fluid charged with excretory products. The inner cells of the paren-
chyma now formed epithelial walls about the nephric cavities, which
became the coelomic sacs; the outer cells were converted largely into
muscles and connective tissue. The muscle cells gave rise to fibers
that reinforced the somatic musculature, and eventually came to
be its principal constituents. The definitive musculature of modern
annelids, therefore, is a composite of fibers derived from the larval
ectoderm and of fibers formed from the coelomic mesoblast, but in
the Onychophora and the Arthropoda the entire musculature appears
to be now a coelomic product. There is no reason necessarily for
supposing that the primitive mesodermal muscles were functional
elements of the coelomic sacs, for, though in ontogeny the mesoderm
usually takes the form of two-layered bands of cells, within which
the coelomic cavities are formed, it would seem probable that the
primitive mesoderm was a loose parenchymatous tissue. The coelomic
sacs are specifically the epithelial walls formed about the nephric
cavities; the surrounding muscles were probably generated from the
outer undifferentiated cells of the original parenchyma.

With the later development of the teloblastic somites, into which
the germ cells were distributed from their posterior source of pro-
liferation, the reproductive products were discharged into the coelomic
sacs of these somites, which thus became gonocoelic as well as nephro-
coelic in function. Open nephridia or coelomoducts now connected
the coelomic cavities with the exterior and served both as excretory

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 21

and as genital outlets. Finally, in the Onychophora and the Arthrop-
oda, the coelomic sacs have been divided into gonadial compartments
and nephridial compartments, which have become reduced in size and
limited to restricted parts of the body, with the result that the
haemocoele is restored as the functional body cavity.

III]. DEVELOPMENT OF THE ANNELID NERVOUS SYSTEM

The annelids and the arthropods undoubtedly have a closer bond
of union in the structure of the nervous system than in any other
feature of their organization, except metamerism itself. The definitive
central nervous system of the polychaete annelids is developed from
two distinct sources, one located in the prostomium, or episphere of
the trochophoral larva, the other in the somatic region, or hyposphere
of the larva. From the first is produced the brain; from the second,
the ventral nerve cords. The nervous system of the trochophore con-
sists of ganglionic centers in the prostomium connected by circular
and radial nerve tracts, from which trunks proceed into the hypo-
sphere (fig. 4 A). This primary system centering in the prostomium
must represent the primitive neural system of the unsegmented an-
cestors of the annelids, adapted to the structure of the trochophoral
larva, and is probably congenetic in its origin with the nervous system
of the Platyhelminthes. The segmentally ganglionated ventral nerve
cords of the postoral region of the trunk are correlated in their
development with the development of body metamerism; they pertain,
therefore, to a later stage of evolution, and have no homologues in
the unsegmented worms. The definitive connection between the pro-
stomial and somatic parts of the nervous system is established secon-
darily in the ontogeny of the polychaetes, but in the oligochaetes the
two parts are said to be continuous from their inception. The funda-
mental structure of the somatic nervous system of the articulate
animals is an adaptation to the function of regulating the muscular
mechanism of metameric body movement; the prostomial system is
primarily sensory in function, except insofar as it controls the move-
ments of prostomial appendages.

The phylogenetic origin of the articulate nervous system can prob-
ably be interpreted very closely from the development of the neural
elements in the trochophore larva of the polychaete annelids, and must
have been about as follows: The primary neurocytes were probably
sensory cells of the ectoderm closely associated with the primary myo-
cytes, and were thus, at first, both receptive and motor in function.
As the muscular system became elaborated, however, the primary
neurocytes were withdrawn to the inner surface of the ectoderm,

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

while other superficial cells assumed the receptive function and trans-
mitted secondarily the impulses from external stimuli to the first set
of cells, which now became purely motor neurones. Finally, still other
neurocytes gave rise to a subepidermal plexus of fibrous tracts that
formed lines of intercommunication between the scattered motor and
sensory elements, and thus unified and coordinated the entire nervous
system. Then the nerve cells of the prostomial region became aggre-
gated into a number of ganglionic centers, principally associated with
groups of receptive cells in primitive sensory organs, and the nerve

a Sarat

WSS
IW
Ts
I Magid
ByeS }
ay
\ LY,
nm
|

Fic. 8.—The nervous and muscular elements of a young trochophore larva of
Lopadorhynchus, nerve tissue represented in white, muscle tissue in black.
(From E. Meyer, Igor.)

A, aboral surface of larva. B, oral surface.

da, rudiments of so-called dorsal antennae; dv, median dorsal nerve of hypo-
sphere; gSo, ganglion of left ciliary organ; Ks, apical plate; mcl, muscle fibers;
Msd, mesoderm; n, longitudinal nerves (seven pairs in episphere) ; NC/s, nerve
cells; An, circular nerve of prototroch; rno’, rno*, rno*, circular nerves of
episphere; rnu, circular nerve of hyposphere; So, left ciliary organ; so, rudimen-
tary right ciliary organ; Stom, stomodaeum; dn, larval stomodaeal nerve.

tracts assumed definite courses. Thus was evolved the primary
nervous system of the polychaete larva. The prostomial ganglia of
this system later coalesce to form the definitive brain. The somatic
nervous system, subsequently developed in correlation with meta-
merism, took its origin from restricted ventrolateral tracts of the
somatic ectoderm, became connected with the brain, and finally re-
placed the primary system in the body region.

. The most primitive nerve center of the annelids probably is repre-
sented by the apical ganglion of the polychaete trochophore (fig. 4 A,
ApGng) situated beneath the ectodermal apical plate (ApP/), which

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 23

usually bears a tuft of cilia, and with which there may be associated a
pair of small larval tentacles, and sometimes a pair of “eye spots.”
From the apical ganglion, nerves radiate posteriorly (LNv) on the
inner surface of the epidermis, and these longitudinal radial nerves
are connected by bands of circular fibers (CNv), chief of which is
the nerve ring of the prototroch (Prtr). The nerve tracts, both radial
and circular, closely follow the peripheral muscle bands of the larva
(fig. 8), thus attesting that the nervous and contractile elements arose
from common ectodermal neuromuscular rudiments. The nerve tissue
is situated between the muscle fibers and the epidermis, the nerve cells
being scattered individually, or condensed in small ganglionic groups.
The nervous system of the polychaete trochophore is best known
from the elaborate studies of Kleinenberg (1886) and of E. Meyer
(1901) on the larval development of Lopadorhynchus, a small errant
polychaete of the family Phyllodocidae (fig. 13 D) having two pairs
of prostomial tentacles but no palpi. The larva of Lopadorhynchus
is a typical trochophore (fig. 8) with an equatorial band of cilia, the
prototroch, just above the mouth. The apical ciliary organ, however,
does not have the usual form and position; it is transposed to the
anterior ventral surface, and is divided into a well-developed organ
on the left (B, So), and a rudimentary organ on the right (so). The
episphere contains seven pairs of longitudinal nerves (m'-n"), and is
encircled by three nerve rings (rno'-rno*) above that of the prototroch
(Fn). In the hyposphere there is but a single nerve ring (A, rnw).
The largest of the longitudinal nerves are two thick lateroventral
nerve tracts (B, n?), which anteriorly (apically) are continuous with
each other in a wide transverse commissural arch within the episphere,
and posteriorly are extended into the hyposphere as a pair of large
lateral trunks (Vd) that break up into smaller branching nerves.
The neural cells of the Lopadorhynchus larva are described in great
detail by Meyer. In general they lie along the fiber tracts (fig. 8 B,
NCls), where many of them are aggregated into small ganglionic
clumps, particularly in the episphere. In the early stages of develop-
ment, according to both Meyer and Kleinenberg, the neurocytes are
generated from the ectoderm in association with muscle cells, and the
principal neuromuscular rudiments of the episphere represent larval
sensory organs (fig. 16 A), of which the nerve cells (~) form small
ganglionic centers. The scattered neurocytes are probably the gener-
ative cells of the fibers in the nerve tracts. The ganglionic centers of
the larva pertain to the apical ciliary organs, a pair of transient larval
antennae, the two pairs of persistent tentacles, which are dorsal and
ventral in the adult (fig. 13 D), and the nuchal organs, but include

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

also two cell groups of unknown significance situated on the dorsal
surface of the episphere (fig. 8 A, da). From some of the ganglion
cells nerve processes go to the muscles, and from others fibers pene-
trate centrally into the nerve tracts.

Before the beginning of larval metamorphosis, Meyer says, the
production of myocytes ceases in the larval neuromuscular centers,
and during metamorphosis a large part of the larval musculature is

Fic. 9.—Theoretical evolution of the annelid nervous system, diagrammati-
cally following Kleinenberg’s and Meyer’s accounts of the development of the
nervous system in the larva of Lopadorhynchus.

A, early trochophore with diffuse nerve cells (NCls) along the radial and
circular nerve tracts. B, nerve cells of episphere concentrated in incipient
ganglia connected with ectodermal sensory organs along lateral radial nerves;
ventral nerve cords (VNC) developed from ectoderm of hyposphere. C, scat-
tered ganglia of episphere condensed into a cerebral mass (Br); cerebral con-
nectives united with ventral nerve cords. D, generalized adult nervous system;
podial ganglia developed at bases of body appendages. EE, nervous system of
adult polychaete, lateral view.

AlCnl, alimentary canal; An, anus; Br, brain; E, eye; Mth, mouth; NCls,
nerve cells; NO nuchal organ; PdGng, podial ganglion; P/p, palpus; Prst,
prostomium; 7/, tentacle; V NC, ventral nerve cord; ZG, zone of growth.

lost. Many of the primary muscles, however, remain, including those
of the persistent tentacles and nuchal organs, and certain other muscles
of the prostomium. The neural cells of the various ganglionic centers
of the larval episphere, on the other hand, increase in number until
they become so crowded that details of their further development
cannot be followed. The cells thus generated, however, are massed
upon the large lateral nerve trunks of the episphere (fig. 8B, n?)
and their anterior commissure. In this manner there is formed from

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 25

numerous agglomerated centers in the larval episphere (fig. 9 A, B)
a compact cellular and fibrous body of nerve tissue (C, Br), which
becomes the brain of the adult worm (D). Hence, as Kleinenberg
remarks, the developmental history of the brain in Lopadorhynchus
shows how extraordinarily complicated in its origin is the cephalic
ganglion even in the annelids. However, that details in the probable
phylogenetic history of the nervous system are not necessarily reca-
pitulated in ontogeny is shown in many annelids having a direct
development, or one in which the trochophoral stage is passed within
the egg, for in such forms the brain is differentiated from the

Fic. 10.—Median vertical sections of the anterior end of an embryo of the
viviparous polychaete Ctenodrilus branchiatus Sokolow (Cirratulidae), show-
ing extension of the mesoderm into the prostomium, and the direct development
of the brain from the prostomial ectoderm. (From Sokolow, Igrt.)

A, embryo before appearance of coelom, with mesoderm (Msd) extended into
prostomium (PMsd). B, full-grown embryo, with coelom and dissepiments,
coelomic cavity of prostomium (PCoel) continuous with coelomic cavity of
first postoral somite (metastomium), which in the embryo is separated from
second somite by a temporary dissepiment (1Dsp).

Br, brain; Coel, coelom; 1Dsp, first (temporary) dissepiment, behind first
postoral somite; 2Dsp, 3Dsp, second and third (permanent) dissepiments; Ecd,
ectoderm; mcl, muscles; Msd, mesoderm; Mth, mouth; PCoel, prostomial
coelom; PMsd, prostomial mesoderm; Stom, stomodaeum.

prostomial ectoderm as a single, compact mass of neural cells
(fig. 10, Br). ,

The larval innervation of the hyposphere gives way during meta-
morphosis to the definitive body nervous system, consisting of the
ganglionated ventral nerve cords and their peripheral nerves. The
rudiments of this system appear first in the embryo as continuous
strands of neurocytes proliferated in the ventral parts of the ecto-
dermal somatic plate as the median edges of the latter unite to close
the blastopore. The cords later become ganglionated by the segmental
aggregation of their cells. The neuroblasts of the somatic nerve cords,
Meyer believes, represent the nerve cells of a series of primitive
ectodermal sense organs. Though there are no persistent remnants of

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

such sensory organs in the annelids, the so-called “ventral organs” of
the Onychophora, from which the nerve cords are differentiated, sug-
gest that the latter took their origin from ectodermal structures of
some kind.

The final connection between the brain and the ventral nerve cords,
according to Kleinenberg and Meyer, is established by fibers that grow
forward from the first ventral ganglia (fig. 9 B) and unite with the
lateral nerve trunks (fig. 8B, Ydn) extending posteriorly from the
arms (n*) of the cerebral commissure. The union thus formed pro-
duces the stomodaeal (circumoesophageal) connectives, through which
the prostomial and somatic nerve centers are unified in the definitive
nervous system.

The peripheral subcutaneous nervous system of the adult worm is
developed directly from scattered neurocytes of the ectoderm. To
this system Kleinenberg ascribes the parapodial ganglia (fig. 9 D,
PdGng), which, he says, are formed quite independently of the central
system by groups of ectodermal neurocytes situated mesad of the
parapodial bases. Secondarily, the parapodial ganglia send connecting
nerves to the ventral nerve cords.

IV. THE ADULT ANNELID

The final development of the adult polychaete annelid from the
larva depends upon the histogenic activity in the zone of undiffer-
entiated cells situated between the last larval somite and the pygidium
(fig. 11 B, ZG). Within this zone of growth is generated anteriorly
a series of secondary postlarval somites (C, D), which does not repre-
sent an extension of the body, but an expansion of a small part of it,
since the new somites are interpolated between the primarily seg-
mented larval body and the pygidium. The more anterior somites of
the new series, being those first formed, are the first to acquire the
mature structure. The teloblastic growth-process is the same whether
the larva is a typical trochophore (fig. 12 B, D), or one more nearly
resembling the adult worm (FE, F, G), but in the first case a greater
degree of metamorphosis accompanies the formation of the new
somites. Hence, though we may eliminate the trochophore from our
concept of the primitive annelid, we cannot dismiss the secondary
formation of the teloblastic somites as a purely ontogenetic process—
it must be explained in terms of phylogeny.

THE TELOBLASTIC, OR POSTLARVAL, SOMITES

The zone of growth, as described by Lillie (1906) in Arenicola
cristata, is a mass of large clear mesodermal and ectodermal cells,

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS = 27

which are frequently to be seen in the process of mitosis. Posteriorly
the growing zone is sharply defined from the pygidial region, but
anteriorly it passes by gradual transition into the more fully differen-
tiated region in front. Its ectodermal cells, Lillie says, must be derived
from the last transverse row of cells in the ectodermal somatic plate
produced from the 2d cell of the embryo. The space between the
ectoderm and the endoderm is filled with a mass of mesoderm cells
very probably generated from the mesodermal teloblasts. Anteriorly
the mesoderm of the growing zone is shut off by a roughly defined

Fic, 11.—Diagrams illustrating the direct primary segmentation of the body
of a larval polychaete, and the growth of the worm by successive addition of
secondary teloblastic somites generated in the subterminal zone of growth.

A, larva with unsegmented soma and mesoderm bands. B, larval soma and
mesoderm directly segmented. C, D, successive formation, from subterminal
zone of growth, of teloblastic somites interpolated between primary larval
somites and terminal pygidium.

AlCnl, alimentary canal; Coel, coelomic cavity; E, eye; J-JII/, primary larval
somites; JV-IX, secondary teloblastic somites; Msd, mesoderm; MsT, meso-
dermal teloblast; P/p, palpus; PMsd, prostomial mesoderm; Prst, prostomium ;
Pyg, pygidium; Soma, body region between prostomium and pygidium in which
somites are formed; Tl, tentacle; ZG, zone of growth at end of soma.

transverse partition from the coelomic cavity of the somite before it.
The first evidence of new somite formation is the appearance of an
irregular space in the mesodermal mass of the zone of growth, which
enlarges upward around the alimentary canal and becomes the coelomic
cavity of the new somite. (Arenicola has a dorsal mesentery but
none beneath the alimentary canal.) The anterior coelomic wall
is pressed against the preceding partition and becomes the posterior
lamella of the dissepiment thus formed. Longitudinal muscle fibers
make their appearance at an early period in the somatic layers of
the mesoderm, but the circular muscles, Lillie claims, appear much
later, and evidently, as described by Meyer (1901), are derived from

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

the inner surface of the ectoderm. According to Iwanoff (1928) the
mesoderm of the postlarval somites is formed in Polygordius, Aricia,
Arenicola, and the Oligochaeta from the mesodermal teloblasts that
generate the larval bands of mesoderm, but in the rest of the Poly-
chaeta the postlarval mesoderm is proliferated from the ectoderm of
the zone of growth.

The pygidial region posterior to the zone of growth retains its
primitive characters throughout the course of development, and is
carried continuously backward as the number of somites increases.
When the definitive number of somites has been formed, the growing
zone loses its distinctive features and becomes indistinguishable as
such. Structurally the secondary, or teloblastic, somites are modeled
according to the general plan of the primary somites before them;
but, though they may differ in various structural details from the
latter, they have one distinctive feature, which is that they alone
contain the germ cells. Germinal centers (“‘gonads”) may occur in
all the teloblastic segments, but in most of the polychaetes they are
limited to a definite part of the body (the epitoke), and in the
oligochaetes they are usually restricted to a few segments.

The ancestral annelids necessarily were reproductive as adults
in all their evolutionary stages, but phylogenetic forms recapitulated
in ontogeny are generally not reproductive. Hence, it is difficult to
study the evolution of the reproductive system from ontogenetic
development. The germ cells of the annelids usually are not recog-
nizable as such in the larva, and little is known of their embryonic
origin. It is claimed by Malaquin (1925), however, that in the
serpulid Salmacina dysteri the sex elements first appear as differen-
tiated cells in the gastrula, and that later (Malaquin, 1924) these
cells become localized immediately before the zone of growth in the
posterior segments of the young larva, where they lie ventral to
the rectum, and are distinguishable from the surrounding cells by
their large, clear, spherical nuclei containing numerous small chro-
matic masses. In the oligochaete Pachydrilus, Penners (1930) claims
the germ cells arise directly from the mesodermal teloblasts, and are
the first cells formed by the latter. The germ cells, as shown also by
Penners and Stablein (1930) in Tubificidae, appear prior to the for-
mation of the definitive gonad somites, and migrate in the haemocoele
to these somites, where they penetrate the mesoderm and finally take
their definitive positions in the dissepiments. It seems highly probable,
therefore, that the primitive annelids, at a phylogenetic stage before
the teloblastic somites were formed, carried the germ cells in the
undifferentiated posterior part of the body behind the last primary

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS = 29

somite. From this point the germ cells must have been distributed to
the secondary somites when the latter began to be developed during
the course of evolution. Hence, primarily, the entire series of telo-
blastic somites would appear to have been genital segments.

Fic. 12—Examples of the growth of larval Archiannelida and Polychaeta
by proliferation in a subterminal zone of growth of teloblastic segments added
to the primary larval body.

A, larva of Eupomatus uncinatus with series of teloblastic, or “postlarval,”
segments (7Segs) interpolated between the three primary larval somites (J,
II, III, see fig. 7 A, B) and the terminal pygidium (simplified from Iwanoff,
1928). B, larva of Polygordius neapolitanus Fraipont during metamorphosis,
with series of teloblastic segments added to the trochophoral body, which is
itself unsegmented and contains no primary mesoderm (from Woltereck, 1905).
C, half-grown young of Nerilla antennata Schmidt (from Schlieper, 1925).
D, larva of Lopadorhynchus brevis Grube with series of teloblastic segments
(from Kleinenberg, 1886). E-G, growth stages of “nereidogen” larva of Platy-
nereis dumerilii Aud. & Milne-Edw. (from Hempelmann, 1911, see also fig. 7 C,
first stage larva).

AnCir, anal cirrus; Cirl, Cirll, tentacular cirri of first two somites, united
in peristomium; J-X/, somites; Papd, parapodium; Perst, peristomium; Plp,
palpus; Prst, prostomium; Pyg, pygidium; T/, prostomial tentacle; TSegs,
teloblastic segments.

A condition similar at least to that which we should expect to find
in the primitive annelids is seen in the archiannelid Dinophilus (fig.
13 A). The body of Dinophilus consists of six or seven somites
clearly defined externally between the prostomium and the pygidium,
but there are no coelomic cavities in the diffuse mesoderm of the

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

somites anterior to the last one, though each of these somites has a
pair of simple protonephridia. In the terminal somite are formed
the reproductive organs, which, in the female, consist of a delicate
gonadial sac, either single or double, extending forward in the body,
and opening posteriorly by a median pore, at least at the time of egg
laying. The gonadial sac appears to represent the coelom of the last
segment, though, as Iwanoff (1928) points out, it may be simply a
space accommodating the germ cells in the undifferentiated tissue
near the end of the body. Hence, the apparent last somite is either
a single teloblastic genital somite, or a region corresponding with
the zone of growth of the polymerous annelids.

A concrete example of the secondary distribution of the germ cells
in a polymerous annelid is given by Malaquin (1924 a) in his study
of the development of Salmacina dysteri. The germ cells, as we have
seen, are first localized in the growing zone of the young larva. When
the formation of the postlarval segments begins, Malaquin says, the
germ cells multiply, and three, four, or five of the resulting gono-
cytes become adherent to the outer wall of each new coelomic sac.
Thus the germ cells, proliferated from a constant source, are dis-
tributed to the newly forming somites, and are extracoelomic both in
their origin and in their secondary segmental positions. After a
period of inactivity the segmentally distributed gonocytes begin to
multiply in the coelomic walls, and here form the small masses of
germinal cells ensheathed in peritoneal folds that are known as the
“gonads.”

If, now, the ontogenetic facts of annelid growth are given a phylo-
genetic significance, it becomes evident, as claimed by Iwanoff (1928),
that the extension of the worm by the teloblastic generation of new
somites, in which are apportioned groups of the multiplying germ
cells, was primarily a means of amplifying the reproductive function.
In the course of evolution it gave rise to a type of animal from which
have been derived the modern Annelida, the Onychophora, and the
Arthropoda.

The teloblastic genital segments are in many respects mechanical
improvements over the primary segments; their muscular equipment
is stronger, the parapodia better constructed for locomotion, the
dissepiments usually more complete, and the nephridia more efficient
for excretory purposes. Hence, the whole worm is clearly a stronger
and a more active animal by reason of the addition of the well-
organized reproductive somites. At the bottom of the water the
creeping worm is better able to force its way under stones or into
crevices, or to burrow into sand or mud; but at the breeding season

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 31

its new powers of locomotion come into effective service, for now
many modern species that habitually live at the bottom rise to the
surface in swarms of energetically swimming individuals, both males
and females, and here discharge the matured gametes.

That the genital segments may be of no special physiological
importance to the worm, except for carrying, maturing, and distrib-

Fic. 13—Examples of annelid types.

A, B, Dinophilus, a very simple archiannelid, perhaps a primitive form, lack-
ing teloblastic somites, tentacles, cirri, chaetae, parapodia, and coelomic sacs;
with five pairs of protonephridia, reproductive organs in posterior part of body
(A, D. gyrociliatus Schmidt, adult female; B, adult male, from Shearer, 1912).
C, Nerilla, an archiannelid with polychaete characters, perhaps a degenerate
form, having a coelom, open metanephridia, and direct development (fig. 12 C)
(N. antennata Schmidt, from Goodrich, 1912). D, Lopadorhynchus, an errant
polychaete (Phyllodocidae), having typical trochophoral development (fig. 8)
with metamorphosis (fig. 12 D) producing long series of teloblastic somites
(L. uncinatus Fauvel).

An, anus; AnCir, anal cirrus; Ch, chaetae; Cir, cirrus; J, IJ, first two
somites; Mth, mouth; Nph, nephridium; Ov, ovary; Papd, parapodium; Phy,
pharynx; Plp, palpus; Prst, prostomium; S/G/d, salivary gland; TJ, tentacles.

uting the reproductive elements, is shown by the various ways in
which the annelids can dispose of these segments without otherwise
impairing their functional integrity. There is the well-known case
of the palolo worms, Eunice fucata and E. viridis, for example,
which live in crevices of rocks at the bottom of the water, and at the
time for spawning detach the rear parts of their bodies, already

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

loaded with the mature generative elements. The reproductive tail-
ends (epitokes) then actively swim to the surface, where myriads
of them congregate to liberate the gametes. In their accustomed
haunts the anterior nonreproductive sections (atokes) regenerate the
discarded epitokes in preparation for next year’s consignment to the
breeding grounds. Various other species of Polychaeta have similar
habits. The Syllidae are famous for the many forms of schizo-
genesis, strobilation, and budding that take place among them, but
here the detached piece, either before or after separation, generates
a new head and becomes a complete worm except perhaps for the
lack of an alimentary canal and a few other unimportant structures.
Again, in some of the Ctenodrilidae the worm breaks up by con-
striction into several pieces of a few segments each, and the middle
pieces regenerate both a head and a tail.

The periodic fragmentation of the body for reproductive purposes,
however, cannot lead to anything in the way of constructive evolution,
and, with the annelids in general, the tendency has been to integrate
the entire series of somites into a mechanical and physiological unit,
in which the reproductive cells are assigned to definite segments. In
the Arthropoda, though the body may still be composed of freely
movable segments, the process of integration has been carried so far,
and the various organs so interdependently distributed, that fission
becomes impossible without fatal results. It would seem, therefore,
that the teloblastic somites, first added apparently for reproductive
efficiency, have been found so useful in other ways that they have
come to constitute not only the largest part of the body in all the
articulate animals, but its most important part, except for the primary
sensory and nervous elements contained in the head.

A structural differentiation between groups of somites, forming
distinct body regions, or tagmata, has taken place in many of the
polychaetes, particularly in the Sedentaria, and is a characteristic
feature of all the Arthropoda. The zone of growth, therefore, which
presumably at first gave rise to a series of identical somites, has
acquired the remarkable faculty of differential activity, producing
successively, at definite segment intervals, two or more series of
somites having often a strongly contrasting structure, while minor
differences may be distributed throughout the entire series of
segments.

THE PROSTOMIUM AND ITS APPENDAGES

The annelid prostomium is the part of the trunk that is not invaded
by the blastopore as the latter elongates forward on the ventral sur-

No.6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 33

face of the embryo (fig. 6D, Prst); in the adult it is reduced to a
small lobe overhanging the mouth (fig. 14, Prst). Appendages of
the prostomium are best developed in the errant polychaetes, where
typically they include a pair of anterior tentacles, or “antennae” (T1),
with frequently a median tentacle between them, and a pair of more
posterior and ventral palpi (Plp). The prostomial appendages are
clearly not equivalent to the parapodia of the postoral body somites,
but they have the same development in the larva as the parapodial
cirri (cf. fig. 16, A and B). Since the prostomium usually contains
the brain and bears the apical sense organs, it constitutes the “head”’
-of the worm. In the absence of prostomial appendages and sense

__ Fic. 14.—Head and anterior body segments of Nereis virens Sars. A, dorsal;
B, ventral.

Cirl, CirlI, tentacular cirri of first and second somites united in peristomium ;
E, eye; III, IV, third and fourth somites; Papd, parapodium; Perst, peristomium
(somites J and JJ); Plp, palpus; Prst, prostomium; 7/, prostomial tentacle.

organs, however, the brain may be secondarily withdrawn into the
body, as in the earthworms (fig. 17 C, D, Br).

The prostomium is not affected by the process of metamerism that
cuts the postoral body region into a series of somites. Since the
mesoderm bands of the larva do not proceed anterior to the mouth
(fig. 6 F). the larval prostomium does not contain mesoderm; but
in later stages the mesoderm of the first somite may be extended
into the prostomium (fig. 10 A) and give rise to a cephalic coelom
and peritoneum (B, PCoel). Ordinarily the cephalic mesoderm is
not segmented, but according to Binard and Jeener (1928) there is
present in the prostomium of the spionid Scolelepis fuliginosa a pair
of distinct coelomic sacs, which are continuous with the cavities of
the palpi, and have no connection with the coelomic sacs of the first

-postoral somite. This fact, the authors point out, gives a new argu-

3

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

ment in favor of the homology of the polychaete palpi with the
tentaclelike antennae of the Onychophora; but evidently it does not
prove their further contention (1929) that the palpi are appendages
of a secondarily “cephalized’’ somite, since it must first be demon-
strated that coelomic cavities may not pertain to the preoral mesoderm
itself.

THE BODY AND ITS APPENDAGES

The body of the annelid is the segmented part of the trunk posterior
to the acronal prostomium, including the region of the true somites,
the zone of growth, and the pygidium; but the term soma, in a
restricted sense, would apply literally only to the region of the somites
between the prostomium and the zone of growth or the pygidium.
In the Polychaeta the first two somites are generally united with each
other in a double segment known as the peristomium, (fig. 14, Perst),
the tentaclelike cirri of which (Cir/, CirI/) take an anterior position
closely associated with the prostomium. The “cephalization” of the
anterior segments in the polychaetes, therefore, contrasts with that
in the arthropods, since, with the latter, the first stage of cephalization
is a union of the first somite with the prostomium. In the oligochaetes,
however, the first somite and the prostomium may unite to form a
composite head as in the arthropods.

The fundamental demarcation of the annelid somites is the attach-
ment of the longitudinal muscle fibers of the body wall and the
muscles of the dissepiments on transverse circular grooves of the
integument; but the coelomic sacs when present are strictly intra-
segmental, and most of the ectodermal and mesodermal organs are
segmentally repeated. The locomotor mechanism of the annelids
consists primarily of the somatic musculature and the regulating
nerve ganglia, which give movement to the body wall, but it usually
includes external adjuncts in the form of bristles or chaetae, and,
in the Polychaeta, lobelike segmental appendages, the parapodia. The
annelid body musculature should be the basis of the derived arthropod
musculature, but there is reason to doubt that the polychaete para-
podia are prototypes of the arthropod legs.

The somatic musculature of the annelids includes the muscles of
the body wall, the muscles of the chaetal sacs, and the muscles of
the parapodia. The muscle fibers, with possibly rare exceptions, are
of the nonstriated type. The musculature of the body wall is of a
very simple pattern, so far as the arrangement of the fibers is con-
cerned, but it may attain a strong development in the rapacious
polychaetes and the burrowing oligochaetes. The longitudinal muscles

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 35

can produce only contraction or lateral undulatory movements of the
body ; the circular muscles are constrictors producing peristaltic waves
of body compression, and longitudinal extension of the body by the
creation of internal pressure. The arthropod type of body mechanism,
involving intersegmental movement of integumental plates, can be
derived from the intrasegmental annelid mechanism only by the
establishment of new intersegmental divisions.

The polychaete somatic musculature is well developed in the
Nereidae, of which Nereis virens may be taken as an example (fig.
15). The outermost layers of body wall muscles consist of fine cir-
cular fibers closely adherent to the integument (A, D, 1). Internal
to these there may be bands of oblique fibers (D, 2) crossing each
other in opposite directions. The largest of the somatic muscles,
however, are four thick bundles of longitudinal fibers (A, 3, 4) lying
internal to the others, two dorsal and two ventral, the fibers of which
are attached on deeply inflected intersegmented folds of the integu-
ment (D, isf). The longitudinal muscles of the terrestrial oligochaetes
are continuous in a thick layer around the entire circumference of
each somite, except where they are interrupted by the intrusion of
the four chaetal sacs. Besides the muscles of the body wall there is
in Nereis a double series of paired, obliquely transverse ventral
muscles, one pair anterior and the other posterior in each segment
(D, 5, 6), which extend outward and upward from the median ventral
fold of the body wall (A) to the lateral intersegmental folds between
the parapodial bases. The intersegmental folds give attachment also
to the transverse or radial muscles of the intercoelomic dissepiments.
Most of the other muscles of the body pertain to the chaetal sacs and
the parapodia, and will be described in connection with the parapodia.

A typical polychaete parapodium is a lateral outgrowth of the body
wall (fig. 15 A, Papd), flattened antero-posteriorly, and usually
divided into a dorsal lobe and a ventral lobe, which again may be
subdivided into secondary lobules. Each major lobe bears distally a
fan-shaped group of long chaetae (B, Ch), and on its base a slender
cirrus (dCir, vCir). The chaetae arise from the inner walls of chaetal
sacs (C, chS), from each of which a long rod, the acicula (Acic),
extends inward to give attachment to protractor and retractor muscles.

The larval rudiments of the parapodia represent the cirri and the
chaetal sacs, and are differentiated as cellular bodies within the ecto-
derm. The rudiments of the cirri, as described by Kleinenberg
(1886) and by Meyer (1901) in the larva of Lopadorhynchus (fig.
16 B, dcR, vcR), consist each of an outer layer of myoblasts (m)
and an inner core of sensory nerve cells (7). The cirri in their origin,

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

“ AN i
RAINS po
\S AY j

f NY . SESS

eae =

pS

K——

BIZ.

vCir

Fic. 15.—The polychaete locomotor mechanism: parapodia, and somatic and
parapodial muscles of Nereis virens Sars.

A, transverse section of a body segment, posterior view, somewhat diagram-
matic. B, a parapodium and its muscles, posterior view. C, chaetal apparatus
of a parapodium. D, muscles of right half of a body segment, inner view.

Acic, acicula; AlCnl, alimentary canal; Ch, chaetae; chS, chaetal sac; Cir,
cirrus; dCh, dorsal chaetae; dCir, dorsal cirrus; Dsp, intersegmental dissepi-
ment; DV, dorsal blood vessel; isf, intersegmental fold of integument; Papd,
parapodium; vCh, ventral chaetae; vCir, ventral cirrus; VNC, ventral: nerve
cord.

I, circular muscles of body wall; 2, oblique muscles of body wall; 3, dorsal
longitudinal muscles; 4, ventral longitudinal muscles; 5, 6, anterior and pos-
terior lateroventral, obliquely transverse muscles; 7, 8, dorsal motors of
parapodium; 9, 10, ventral motors of parapodium; 17, intrinsic muscle of para-
podium between dorsal and ventral lobes; 12, protractors of dorsal acicula and
chaetal sac; 13, r4, retractor and protractor of dorsal chaetae; 15, retractor
of dorsal chaetal sac and acicula; 16, protractors of ventral acicula and chaetal
sac; 17, 18, retractor and protractor of ventral chaetae; 19, retractor of ventral
chaetal sac.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 37

therefore, resemble the tentacular rudiments of the prostomium (A),
and later they grow out as tentaclelike processes. The bristle sacs
are formed as ectodermal cell masses between the cirri (B, chS), the
outer cells of which become myoblasts, while some of the inner cells
enlarge and produce the chaetae ; a lumen then appears in the cell mass,
and the latter becomes an open eversible sac from which the chaetae
protrude. Finally the cirri and the chaetal pouches are carried out-
ward on an outgrowth of the body wall that becomes the principal
part of the appendage. The mature parapodia of Lopadorhynchus
are not of typical form in that each consists of a single lobe (fig. 16 C)
with both chaetal sacs at its extremity.

In some of the polychaetes, particularly in the Sedentaria, ee
are two rows of podial organs on each side of the body (fig. 16 F),
those of one series, the notopodia (dPd), being situated dorso-
laterally, those of the other, the neuropodia (vPd), ventrolaterally.
Each organ includes a cirrus (Cir) and a chaetal sac (chS), and
is innervated separately from the corresponding podial ganglion
(PdGng). In the Oligochaeta the podial organs are represented only
by the chaetae, which usually are arranged in two separated rows on
each side of the body. It is possible, therefore, that the usual two-
branched parapodium of the Polychaeta (fig. 15 B) has been formed
by the union of a notopodium and a neuropodium. Furthermore, the
double composition of each notopodium and neuropodium suggests
that the primitive polychaetes had dorsolateral and ventrolateral rows
of cirri, and between them on each side two series of chaetal sacs.
On the peristomial segments of adult polychaetes generally only the
cirri are present (fig. 14 A, CirJ, CirIJ), but on the rest of the body
segments the chaetae-bearing lobes are usually the more important
podial elements.

The musculature of a parapodium is somewhat complex: it includes
extrinsic muscles that move the appendage as a whole, and intrinsic
muscles concerned principally with the movement of the chaetae. In
Nereis virens there are four extrinsic muscles for each parapodium,
two dorsal (fig. 15 D, 7, 8), and two ventral (9, zo). The dorsal
muscles arise anteriorly and posteriorly on the body wall, but cross
each other obliquely to opposite margins of the parapodial base. The
ventral muscles take their origins on the median infold of the ventral
wall of the body segment (A), and extend laterally and dorsally,
above the ventral longitudinal body muscles (4), to the anterior and
posterior margins of the base of the parapodium. If the dorsal and
ventral muscles inserted anteriorly act in opposition to those inserted
posteriorly, the parapodium is moved anteriorly and posteriorly on

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

the vertical axis of its base, and this is the usual motion of the
appendage; but the latter can also be lifted and depressed, and the
up-and-down motion evidently results from an antagonistic action

>” we
PdGng VNC 4 (4

Fic. 16—Development, structure, and innervation of the polychaete appendages.

A, section through larval rudiment of persistent dorsal tentacle (t/R) of
trochophore of Lopadorhynchus (from E. Meyer, 1901). B, transverse sec-
tion of larva of Lopadorhynchus through rudiments of a pair of chaetal sacs
(chS) and associated cirri (from Meyer, 1901). C, parapodium of adult Lopa-
dorhynchus. D, structure of the armature of a parapodium of Myzostomum
asteriae Marinzeller, diagrammatic (from Stummer-Traunfels, 1903). E, a
myzostomid, ventral view, showing parapodia. F, diagrammatic section of an
amphinomid, Hermodice carunculata Pallas, showing widely separated notopodia
(dPd) and neuropodia (vPd) and their innervation (from Storch, 1913). G,
section of Nereis virens Sars, showing innervation of parapodia (from Hamaker,
1808).

a, lateral nerve from podial ganglion; Acic, acicula; acmcls, acicular muscles;
b, notopodial ganglion; Brn, branchia; c, neuropodial ganglion; Ch, chaeta or
chaetae; chS, chaetal sac; Coel, coelom; dCir, dorsal cirrus; dcR, dorsal cirrus
rudiment; dPd, notopodium; Ecd, ectoderm; Gung, ventral ganglion; m, pri-
mary muscle cell; , primary neural cell; NCls, nerve cells; Nv, nerve; Pafd,
parapodium; PdGng, podial ganglion; t/R, rudiment of tentacle; vCir, ventral
cirrus; vck, ventral cirrus rudiment; VNC, ventral nerve cord; vPd, neuro-
podium; 3, dorsal muscles; 4, ventral muscles.

between the dorsal and ventral muscles. The intrinsic muscles of the
parapodium include protractors and retractors of the chaetal sacs. The
principal protractors (B, C, 12, 76) converge from the parapodial
walls upon the inner ends of the acicular processes of the sacs (B,

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 39

C, D), but the sacs themselves are eversible by muscles in their own
walls (C, 14, 78). A retractor (75, 19) arising within the para-
podium is inserted on the distal part of each chaetal sac, and a muscle
(13, 17) from the acicula, attached on the base of the sac, opposes
the muscles (74, 78) that evert the sac itself.

The parapodia are subject to numerous structural modifications in
the different groups of Polychaeta, and among the specialized types
the small leglike parapodia of the Myzostomidae (fig. 16E) are of
particular interest because of their resemblance to the legs of Ony-
chophora. Each myzostomid appendage, as described by Stummer-
Traunfels (1903), contains a deep apical pouch (D, chS), from the
inner end of which a large hooked process (Ch) projects outward,
while from its distal wall a thick rod (Acic) extends inward and
gives attachment to protractor muscles (acmcls) and muscles inserted
on the base of the hook. It is evident that the hook is a single, greatly
enlarged chaeta, and the internal arm an acicula. The myzostomid
“leg,” therefore, is only a modified parapodium adapted for clinging
to the crinoid hosts on which the Myzostomidae live, and has only a
superficial likeness to the appendages of Onychophora (fig. 31).

THE NERVOUS SYSTEM

The central nervous system of the Polychaeta, as shown in the
larval development, is produced from separate prostomial and somatic
rudiments, which secondarily become united (fig. 9); in the Oligo-
chaeta the two parts are said to be continuous from their inception.
The definitive brain, whether formed from discrete ganglionic centers,
as in Lopadorhynchus, or from a single generative zone of the pro-
stomial ectoderm (fig. 10 A, B, Br), is always a compact organ,
though it is generally bilobed (fig. 17 A) or differentiated into several
consecutive parts (fig. 18C). The ventral nerve cords in the more
primitive condition found in most of the archiannelids and in various
polychaete and oligochaete families are entirely separate, except for
their connection by commissures (fig. 19 A,-B), and in such cases
the nerve tissue usually preserves a close contact with the ectoderm
from which it is derived (C). More commonly, however, the paired
ganglia of the cords are united in single median ganglia (fig. 17 C, D),
giving the cords themselves a median position; but even in such cases
the ganglia of one or more pairs carried by the divergent anterior
ends of the cords may remain widely separated. The first pair of
united ganglia on the cords constitutes the so-called ‘‘suboesophageal
ganglion,” but it is evident that this ganglion does not belong always

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

J Prac

eoo------

|

Fic. 17—Nervous system of Annelida.

A, anterior nervous system of Nereis virens Sars, diagrammatic, showing
nerves of cerebral and suboesophageal ganglia (adapted from Hamaker, 1808).
B, anterior nervous system of an amphinomid, Hermodice carunculata Pallas,
showing podial nerves (PdNv) from brain connecting the podial ganglia, and
divergence of ventral nerve cords (VNC) through several somites around
stomodaeum (from Gustafson, 1930). C, anterior neryous system of Lumbricus
terrestris Linn., lateral view, showing retraction of brain into third somite
(simplified from Hess, 1925). D, same, dorsal view (from Hess, 1925).

Br, brain; ComII, ComV, commissures of second and fifth somites; Gugl-
GngVI, central ganglion of first to sixth somites; J-VJ//, first to eighth somites ;
Mth, mouth; PdGng, podial ganglion of second peristomial cirri; PdGngl,
PdGngVIII, podial ganglia of first and eighth somites; PdNv, podial nerve;
Perst, peristomium; P/p, palpus; Prst, prostomium; SoeGng, suboesophageal
ganglion; StCon, stomodaeal nerve connective; StGng, stomodaeal ganglion;
Stom, stomodaeum; JJ, tentacle.

Nerves of Nereis (fig. A): a, nerve to stomodaeum; b, tentacle nerve; c, d,
nerves to muscles and prostomial integument; e, nerve to proboscis; f, palpus
nerve; g, tegumentary nerve; h, i, ocular nerves; 7, nerve to nuchal organ; k,
commissural ganglion; /, m, nerves to proboscis; , connective between peri-
stomial ganglia; 0, nerve to second peristomial ganglion (podial ganglion,
PdGng); p, nerve to proboscis; g, 7, nerves to muscles and integument of
peristomium.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 4I

to the same somite, and, furthermore, it sometimes contains the
ganglia of more than one somite. In the polychaete family Amphinom-
idae there is, in addition to the median nerve cords, a pair of lateral
cords extending posteriorly from the brain (fig. 17 B, PdNv), which
unite the series of podial ganglia (PdGng) lying at the bases of the
parapodia (see Storch, 1912, 1913, Gustafson, 1930). The tetra-
neurous structure is regarded by Storch as representing the more
primitive condition of the annelid nervous system, though Gustafson
contends that it is probably secondary. According to a theory pro-
posed by Jeener (1928) the lateral line system represents a primitive
series of neuromuscular sensory organs, from which there has been
preserved and developed in the Sedentaria the sensorial elements, in
the Errantia the ganglionic elements, and in the Oligochaeta the
muscular elements.

The annelid brain in its simplest form probably consists of a
homogeneous mass of neurocytes aggregated upon a fibrous com-
missure continuous on each side with the stomodaeal connectives, and
through the latter with the ventral nerve cords (fig. 19 A, B, Br).
With higher development, however, specialized groups of cells appear
in the cortex, and specific tracts of fibers are individualized in the
neuropile. A very simple brain structure occurs in the archiannelid
Polygordius (fig. 18 A), in which, according to Hanstrom (1929),
a pair of glomerulous association centers (PlpGlm) receive the roots
of the palpal nerves and are connected by a palpal commissure
(PlpCom). The peripheral sense cells of the palpi form ganglionlike
masses (SCls) at the bases of the appendages. Two posterior lobes
of the brain (NL) are connected with the nuchal organs, but eyes
and anterior tentacles are absent in Polygordius.

In the active polychaetes, in which cephalic tentacles, palpi, eyes,
and nuchal organs are well developed, the brain takes on a more
complex form and may acquire a high degree of differentiation in its
internal organization. Particularly conspicuous are the paired cellular
and fibrous masses known as corpora pedunculata. Each of these
bodies consists of a cap of small chromatic cells lying in the upper
anterior part of the cortex (fig. 18B, E, Gb), and of a stalk, or
pedunculus (Ped), composed of the neurites of the cap cells, which
penetrate the central part of the brain. Within the stalks the terminals
of the neurites (B, d) form synaptic associations between fibers from
all other parts of the brain and from the ventral nerve cords (a, b,c).
A simple development of the corpora pedunculata is shown by Han-
strom (1927) to occur in the Hesionidae (fig. 18 D), in which the
caps consist each of a single globulus of cells, and the stalks are

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

connected by a fibrous commissure. In most of the other errant poly-
chaetes the corpora pedunculata are more highly developed, and the
cap cells become segregated into two or three distinct globuli (E, F).
On the other hand, in the sedentary polychaetes with reduced cephalic

Fic. 18.—Structure of the brain in Archiannelida and Polychaeta.

A, outline of head and diagram of simple brain structure in Polygordius (from
Hanstr6m, 1929). 8B, horizontal section of brain of Sthenelais picta Verrill
(Aphroditidae), diagrammatic (from Hanstrém, 1927). C, brain of Eunice
punctata Risso (Eunicidae), showing high degree of external differentiation
(from Heider, 1925). D, transverse section of brain of Podarke obscura Ehlers
(Hesionidae) through corpora pedunculata (from Hanstrom, 1927). E, hori-
zontal section of brain of Nereis virens Sars (Nereidae) showing internal
structure (from Hanstr6m, 1927). F, diagram of a corpus pedunculatum of
Nereis virens (from Hanstrom, 1027).

a, sensory fibers of palpus nerve; Za, 2a, 3a, nerves of lateral and median
prostomial tentacles; b, c, fibers of stomodaeal connectives; d, axons of glob-
uli cells of corpus pedunculatum; E, eye; f, central neurocytes in anterior
part of brain; fb, forebrain; Gb, globulus of corpus pedunculatum ; hb, hindbrain;
mb, midbrain; NL, nuchal lobe; NCom, nuchal commissure; OpCom, optic
commissure; OpNv, optic nerve; Ped, pedunculus of corpus pedunculatum;
Plp, palpus; PlpCom, palpal commissure; P/pGlm, palpal glomeruli; PlpNv,
palpal nerve; SC/s, sense cells of palpus; StCon, stomodaeal connective.

sense organs, the corpora pedunculata are correspondingly reduced
or are vestigial, and in the Oligochaeta they are absent.

The development of the corpora pedunculata in the Polychaeta is
clearly correlated with the development of the prostomial sense organs,
but the particular relationships of the bodies are with the sensory
nerves of the palpi. It is shown by Hanstrom (1927, 1928, 1929) in

NO.6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 43

the Hesionidae (fig. 18D), the Aphroditidae (B), the Nereidae
(E, F) and other errant families, that the roots of the palpal nerves
are closely associated in glomerulous bodies with the stalks of the
corpora pedunculata, which fact, Hanstrom points out, clearly sug-
gests that the corpora pedunculata had their inception as association
centers for the sensory nerves of the palpi. Much importance attaches
to a study of the corpora pedunculata in connection with annelid and
arthropod phylogeny, because bodies very similar in position, struc-
ture, and variations are characteristic features also of the brain of
Onychophora and Arthropoda.

The relative positions of the principal internal structures of the
polychaete brain, it should be noted for later comparison with the
onychophoran and arthropod brain, are as follows: Anteriorly and
dorsally are the corpora pedunculata (fig. 18B, D, E); closely
associated with the stalks of the latter are the palpal glomeruli (A, B,
F, PlpGlm), and the glomeruli are connected by a palpal commissure ;
behind the corpora pedunculata is the optic commissure (D, IE,
OpCom) ; and in the posterior part of the brain are the nerve centers
of the nuchal organs and a nuchal commissure (E, NCom). The
stomodaeal connectives attach to the ventral surface of the brain.

The number of nerves given off from the brain is highly variable
according to the development of prostomial sense organs. In Poly-
gordius (fig. 18 A) there is but a single pair of cerebral nerves,
which innervate the tentaclelike palpi, while in such forms as Nereis
(fig. 17 A) an elaborate innervation of the prostomial walls, the sense
organs, and the stomodaeum proceeds from the brain.

The principal stomodaeal nerves of the Polychaeta arise in some
families from the first ganglia of the ventral nerve cords, or from the
brain connectives near these ganglia, while in others they come from
the upper parts of the connectives or from the back of the brain.
It is contended by Hanstrom (1927, 1928), therefore, that in the
second case the primitive first ganglia of the cords have been drawn
forward and united with the brain, forming thus in certain polychaete
families a posterior part of the definitive brain corresponding with
the tritocerebral lobes of the arthropod brain, which always have
connections at least with the stomodaeal (stomatogastric) system
of nerves.

The stomodaeal innervation of the Polychaeta is most elaborate in
those forms that have a large and eversible stomodaeal proboscis, and
in such cases the innervation of the organ may be derived from so
many sources that the evidence adduced in favor of Hanstrom’s theory
is not convincing. In Nereis, for example, Hamaker (1898) describes

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Fic 19.—Examples of generalized structure in the annelid nervous system,
and the structure of the annelid eye.

A, the “rope-ladder” type of nervous system in an oligochaete, Aecolosoma
tenebrarum (from Brace, 1901). 3B, same in an archiannelid, Dinophilus conk-
lint Nelson (from Nelson, 1907). CC, cross-section of ventral body wall of
Aeolosoma tenebrarum, showing nerve cords not separated from epidermis (from
Brace, 1901). D, vertical section of an eye of primitive structure in a chaetopterid,
Rangania sagittaria Claparéde (from Hesse, 1899). FE, diagram of a typical
annelid retinal cell (based on Pflugfelder, 1932). F, cross-section of optic rods
of retina of Heteronereis sp. (from Pflugfelder, 1932). G, vertical section of
eye of a nereid, Lycastis sp. (from Pflugfelder, 1932).

a, outer layer of epidermis over eye; b, inner layer of epidermis forming
ocular vesicle (see fig. 28 C, D, E); Br, brain; c, optic rod of sensory retinal
cell; CB, crystalline body; Com, nerve commissure; Cor, cornea; Ct, cuticula;
d, cell body of sensory retinal cell; e, striated border of retinal optic rod; Epd,
epidermis; Ln, lens; Mth, mouth; nf, nerve fiber; nfb/, neurofibrillae; Nu,
nucleus; Nv, nerve trunk; Ret, retina (including sensory and supporting cells) ;
StCom, stomodaeal connective; V NC, ventral nerve cord.

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 45

five pairs of stomodaeal nerves, two pairs of which proceed from the
anterior part of the brain (fig. 17 A, a, c), a third pair (/) from
small ganglia on the upper ends of the stomodaeal connectives, a
fourth (m) from the ganglia of the first peristomial cirri, and a
fifth (p) from the suboesophageal ganglion. In the earthworm,
Lumbricus, the stomodaeal innervation arises from the connectives
between the brain and the first ganglia of the cords (fig. 17 C, StGng).
Other examples would only show further inconsistencies i the origin
of the nerves that supply the annelid stomodaeum. We can, therefore,
most readily agree with Gustafson (1930), who concludes that no
homology exists between the stomodaeal nervous system of the Anne-
lida and that of the Arthropoda. Gustafson points out, furthermore,
in reference to Hanstrém’s theory of transposed ganglia, that there
is no concrete evidence of the transfer of a pair of ventral ganglia to
the brain in any of the annelids, whereas in the arthropods there is
conclusive proof that the tritocerebral ganglia have been secondarily
united with the brain. In the higher arthropods, moreover, the ganglia
of the stomodaeal nervous system are derived directly from the
ectodermal wall of the stomodaeum itself, and their definitive nerve
connections with the central system appear to be secondary.

THE EYES

Light-receptive organs in the form of eyes are widely present in
the Polychaeta. The polychaete type of eye is fundamentally a
vesicular ingrowth of the integument (fig. 28C, D, E), the retinal
cells being epithelial cells of the vesicle wall converted into primary
sense cells by the extension of their inner ends as nerve fibers. In the
simpler forms of eyes the cuticula may form a mere plug in the cavity
of the retinal sac (fig. 19 D), but usually the ingrown part of the
cuticula is enlarged and becomes a lenslike body, either connected with
the surface by a cuticular strand, or entirely shut in by the union of
the lips of the retinal sac (G, Ln). The outer ends of the retinal
cells form optic rods, converging upon the inner surface of the lens
(E, G, c); which contain the distal parts of the neural fibrillae (E,
nfbl), but the apposed surfaces of adjacent rods do not form rhab-
doms (F), as they do in the Arthropoda.

THE NEPHRIDIA AND THE GENITAL DUCTS

The most primitive excretory organs of the annelids are the proto-
nephridia of trochophore larvae. These are minute tubes, one or two
pairs, extending from the body wall into the haemocoele, where they

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

end blindly, but may be branched ; each tube or each branch terminates
with a cell that sends a long vibratile flagellum into the lumen of the
tube. The larval protonephridia are apparently of ectodermal origin,
being said to be formed from primary nephroblasts derived from
cells of the third quartet of blastomeres ; their structure is essentially
that of the “flame cell’? tubes of the excretory canals of the Platy-
helminthes. Since the larval nephridia are present before the coelomic
sacs are formed, they lie within the primary body cavity, which later
becomes the haemocoele.

A type of closed nephridium resembling the larval nephridia, and
therefore often called a protonephridium, occurs in the five pregenital
somites of the archiannelid Dinophilus (fig. 13 A), and in the post-
larval somites of several families of Polychaeta. The closed nephridia
of the adult worm, however, are more highly developed excretory
structures than the larval organs, and usually have a more complex
end apparatus of tube-cells (solenocytes), which contain long fila-
ments resembling the flagella of the larval nephridia, but said to be
nonmotile. The nephridial canal has a simple structure, and its lumen
is ciliated. These nephridia project into the coelomic cavities, but,
inasmuch as they are ensheathed in folds of the peritoneum, they lie
morphologically in the haemocoele. Because of the similarity of their
structure to that of the larval nephridia, the closed nephridia of the
adult are supposed also to be of ectodermal origin, but their develop-
ment apparently has not been studied.

The usual adult excretory organ, occurring in most Archiannelida
and Polychaeta, and in all Oligochaeta, is of the type called a meta-
nephridium. The characteristic feature of a metanephridium is the
presence of an inner opening, or nephrostome, by which the nephridial
canal communicates with the coelom. Solenocytes in this case are
absent. The nephrostome may be a simple ciliated aperture, as in
the archiannelids, but more commonly it has the form of a wide, open,
ciliated funnel. Unless coelomic dissepiments are absent, the nephro-
stome always lies in the anterior lamella of the dissepiment before
the somite in which the canal opens to the exterior. The canal thus
appears to traverse the coelomic cavity behind the funnel, but morpho-
logically it is extracoelomic, since it is ensheathed in a peritoneal fold
produced from the posterior lamella of the dissepiment bearing its
funnel. A closed nephridium is without doubt strictly an excretory
organ, but an open nephridium may serve both for the removal of
excretory products and for the discharge of the gametes from the
coelom.

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 47

The reproductive elements of the annelids are liberated in various
ways. In some of the Archiannelida and Polychaeta there is no
anatomical provision for the discharge of the sex products from the
coelomic sacs, and in such cases the gametes escape by a rupture of
the body wall or by fission of the rear part of the body. With certain
polychaetes having closed nephridia, a funnel-shaped structure is
developed in the genital somites on the anterior surface of the septum,
which at maturity opens into the canal of the nephridium, and serves
as an outlet for the gametes; but again in others the funnel, though
present, is a mere “ciliated organ” of the coelomic peritoneum, not
known to acquire an opening. Special genital ducts with an internal
funnel and an external aperture are present in only a few Polychaeta,
as in some of the Capitellidae, but they are characteristic features of
the genital segments of Oligochaeta and Hirudinea. In most of the
Polychaeta the nephridial funnels serve for the discharge of the
gametes.

The relationship of the various types of annelid excretory organs
and genital ducts to one another is difficult to understand. According
to the well-known theory of Goodrich (1898-1900), nephridia and
genital ducts, or coelomoducts, originally formed two separate series
of segmental organs, and are still retained as such in Oligochaeta,
Hirudinea, and certain Capitellidae. In the majority of the Poly-
chaeta, however, Goodrich claimed, the genital funnel has lost its own
duct and its funnel has united with the mouth of the nephridium,
intermediate stages being suggested in some forms where there is a
partial fusion between the funnel and the nephrostome.

The study of the development of the open nephridia has given rise
to much difference of opinion as to the origin of the nephridial
rudiments. The earlier investigators, such as Hatschek and Vejdov-
sky, regarded the nephridial funnels and canals as mesodermal
structures, but Whitman (1886) claimed that the nephridia of the
leech Clepsine are entirely of ectodermal origin. Wilson (1889), in
his work on the development of Lumbricus, described the nephridial
canals as being apparently ectodermal structures, developed from
continuous rudiments formed from the second and third rows of
ectodermal cells of the germ band, though he admitted they might
be mesodermal; the funnels, however, he said are derived separately
from the anterior walls of the coelomic septa. Staff (1910) asserted
also that the nephridial canals are ectodermal products in Criodrilus,
but are formed from only the second row of cells in the germ band;
and Tannreuther (1915) claimed the nephridia of Bdellodrilus have
the same origin, though he did not follow their complete development.

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

On the contrary, nearly all other investigators have stoutly main-
tained that both the funnels and the canals are mesodermal, though
some regard these two parts as derived from separate rudiments. In
this class may be mentioned E. Meyer (1887, Psygmobranchus),
Bergh (1888, Criodrilus, 1890, Lumbricus, 1899, IRhynchelmis),
Burger (1891, Nephelis, 1894, Hirudo, Aulastomum), Michel (1808,
Allolobophora), Lillie (1906, Arenicola), Bychowsky (1921, Clep-
sine), Penners (1924, Tubifex), and A. Meyer (1929, Tubifex).
Only Bergh is insistent that the entire nephridium is mesodermal;
most of the others admit that a terminal part, perhaps including the
reservoir, may be formed from the ectoderm.

According to Lillie, the nephridia of the polychaete Arenicola
cristata are gradually differentiated in the somatic mesoderm, starting
from the posterior angles between the septa and the body wall, but
the mesoderm in early stages of somite formation presents no cell
boundaries. The lumen of each organ appears as a minute intra-
cellular canal, which from its inception opens through the dissepi-
ment into the preceding coelomic cavity. Later, as the nephridial
cells divide, the lumen becomes intercellular, and finally it opens
posteriorly through the ectoderm. Lillie says, however, that there is
no invagination of the ectoderm, and no specific evidence that the
reservoir is an ectodermal formation.

Those writers who claim that the nephridia of the Oligochaeta
and Hirudinea are of mesodermal origin agree essentially with Bergh
that each organ is formed from a single cell of the anterior lamella
of an intersegmental septum. According to A. Meyer (1929), for
example, the nephridioblasts of Tubifex are early differentiated from
the other cells of the septa by their large size (fig. 20 A, Npbl). By
successive divisions of the nephridioblast a column of cells is formed
that pushes backward within a sheath of ordinary epithelial cells
derived from the posterior lamella of the septum (B-E). The young
nephridium extends in a space between the somatopleure and the
longitudinal muscles, and is thus extracoelomic. The lumen appears
first as an intracellular canal, which later becomes intercellular by a
radial division of the cells; it is ciliated from an early stage. Pos-
teriorly the canal ends against an epidermal cell (G), through which
it eventually opens to the exterior, and from which is later gener-
ated the reservoir. The coelomic funnel is formed by the original
teloblast, the nucleus of which divides into four nuclei, one taking
a position in the dorsal lip of the funnel, the other three in the ventral
lip (H, I). According to Bergh (1899) only the lower lip of the
funnel in Rhynchelmis is derived from the nephridioblast, the upper

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 49

lip being formed from a neighboring group of septal cells. In
Clepsine, Bychowsky (1921) says, the first division of the nephridio-
blast is in the plane of the dissepiment, and gives rise to an anterior
cell that forms the funnel and the adjacent part of the canal, and a
posterior cell that generates the rest of the canal. The latter opens
finally to the exterior through an ectodermal invagination. Bergh

Fic. 20.—Successive early stages in the development of the posterior nephridia
of the oligochaete Tubifex rivulorum Lam. (From A. Meyer, 1929.)

A, a primary nephridioblast developed from a cell of the anterior lamella of
a dissepiment. B, proliferation of nephridial cells by transverse division of
the nephridioblast. C-F, successive extensions of the nephridial canal within
a peritoneal sheath derived from the posterior lamella of the dissepiment; the
canal acquires first an intracellular lumen. G, the canal still more elongate and
looped upon itself, attached posteriorly to an epidermal cell, through which the
lumen penetrates to the exterior, and which later forms the nephridial bladder.
H, I, two stages in the final development of the nephrostome in the primary
nephridioblast by radial division of the nucleus.

Dsp, dissepiment ; Lum, lumen; NCnl, nephridial canal; Npbl, nephridioblast ;
Npr, nephropore; Nst, nephrostome; PSh, peritoneal sheath,

claims that there is no ectodermal element in the nephridium of
Criodrilus, Rhynchelmis, or Lumbricus.

It thus appears to be now well established that the metanephridia
of the annelids in general are structures of the nature of coelomo-
ducts, formed principally as outgrowths of the posterior walls of the
coelomic sacs, but perhaps including a terminal part of variable extent
derived from the ectoderm. They are extracoelomic, inasmuch as
each nephridial canal is invested in a fold of the coelomic peritoneum.
The nephridial organs have always been important subjects in dis-

4

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

cussions of relationships between the Annelida, the Onychophora,
and the Arthropoda. The onychophoran nephridia, however, are
developed as simple diverticula of the ventral walls of the coelomic
sacs, which connect with short ectodermal ingrowths of the same
segments situated mesad of the leg bases, and the nephridial organs
of the arthropods most probably have had the same genesis as the
onychophoran organs. Hence, it is possible that the coelomic exits
have had an independent origin in the higher Annelida on the one
hand, and in the common ancestors of the Onychophora and Arthrop-
oda on the other.

V. THE ONYCHOPHORA

Somewhere from a generalized annelid stock there must have
branched off in remote pre-Cambrian time the ancestors of the group
of animals that includes the modern Onychophora (fig. 21 A), the
Cambrian Aysheaia (B), and the pre-Cambrian Xenusion (C). The
primitive onychophorons undoubtedly were segmented, wormlike
creatures, in which coelomic sacs and the basic features of the annelid
muscular and nervous systems had long been established, and in
which the body had been lengthened by the addition of a series of
reproductive somites generated from the posterior zone of growth.
A distinctive feature of the Protonychophora, however, was the pos-
session of movable locomotor appendages having the form of small
lobelike outgrowths of the body wall along the lateroventral lines of
the segments. The ancestors of the lobopod Onychophora, and the
ancestors of the chaetopod Annelida, therefore, probably constituted
two divergent branches from a generalized annelid stock. The primi-
tive chaetopods were creeping worms that progressed by the usual
vermiform movements of the body, produced by the body musculature
with the aid of integumental chaetae. The primitive onychophorons
became distinguished as walking worms, a character well expressed
in the name Peripatus (Guilding, 1826) given to the first-described
modern form. The walking habit led to the adaptation of the modern
Onychophora to life on land, but the older forms, such as Aysheaia
and Nenusion, may have been inhabitants of the ocean.

A typical onychophoron is a slender wormlike creature with a pair
of tentacular antennae at the anterior end of the trunk, and a double
row of short, conical, lateroventral legs along the length of the body
(fig. 21 A). The trunk is cylindrical or somewhat depressed, blunt
anteriorly, and tapering posteriorly. The rough integument is closely
ringed, but there is no external sign of segmentation except for the
series of appendages. The animal has no distinct head; the anterior

NO.6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS- 51

part of the trunk, however, forms a cephalic lobe (D, E) bearing
the antennae, a pair of small dorsal eyes (E, E) just behind the
antennal bases, and on the ventral surface the mouth (D). ._The
mouth, which is a triangular opening into the stomodaeum, is sunken
into a preoral cavity surrounded by an integumental circumoral fold

enor
Ls

Ne—c cee cocccc cee

<)
|
°
°
°
°
°
°
°
°
sl
\

:

eoccoccee
cc ococcess

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;
ol
e
°
o
°
°
°
°
Oy

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SU mee eog® 2770

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~eoe

Fic. 21.—Onychophora, ancient and modern.

A, Peripatoides novae-zealandiae Hutton. B, Aysheaia pedunculata Walcott
(1911), of Middle Cambrian, British Columbia, “conjectural restoration” (from
Hutchinson, 1930). C, Xenusion auerswaldi Pompeckj (1927), of Algonkian,
proterozoic pre-Cambrian (from Heymons, 1928, broken lines hypothetically
completing lacking parts). D, Peripatoides novae-zealandiae, anterior part of
trunk, ventral view. E, same, head and anterior part of body, lateral view.
F, same, right jaw, dorsal view, with muscles.

a-d, jaw muscles; Ant, antenna; Ap, apodeme of jaw muscles; cof, circumoral
fold; E, eye; J, jaw; IL, first leg; Lm, labrum; OP, oral papilla.

(cof). Within the preoral cavity is a small anterior labral lobe
(Lm), and a pair of flat, two-hooked jaws (J) that converge pos-
teriorly at the sides of the mouth. On the sides of the head, laterad
of the mouth, is a pair of oral papillae (E, D, Op) that give vent
to a pair of large, many-branched slime glands widely spread in the

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

body cavity (fig. 32 A, SlmGld). The following appendages are
the legs, varying in number with different species from a minimum
of 13 pairs to an average of perhaps 25 or 30 pairs, though some
species have 40 or more. Behind the last legs the body tapers to a
terminal cone on which is situated the anus. The genital aperture
in each sex is a median ventral opening lying either between the legs
of the last pair, or behind the last pair present in species having one
or two of the posterior pairs of legs absent.

EARLY STAGES OF DEVELOPMENT

Were it not for the evidence of annelid relationships shown in the
adult structure of the Onychophora, we should have little reason for
believing that the onychophorons are descended from Annelida, for
in their ontogeny we encounter none of the familiar early phases of
development so characteristic of the annelids. Most of the Ony-
chophora are viviparous, the embryos developing to maturity in
uterine chambers of the oviducts (fig. 32 A, Utrs); only a few
species are known to be oviparous. Eggs supplied with a large quantity
of deutoplasm complete their development from their own store of
yolk, but the embryos of viviparous species with small eggs receive
nourishment from the uterine walls, and in some cases a placentalike
growth of the blastoderm forms a large vesicular trophoblast applied
to the walls of the uterus.

The early stages of onychophoran development are so variable in
different species that it is impossible to give any general account of
the processes of cleavage and germ-layer formation. Cleavage in
some species with small eggs is holoblastic, producing first a solid
morula and then a hollow blastula (see Sclater, 1888). Contrary to
what we might expect, however, gastrulation in such cases does not
take place by invagination. In Peripatus imthurni, as described by
Sclater, an internal proliferation of cells proceeds from a definite
point on the blastula, and the cells thus produced become differen-
tiated into endoderm and mesoderm. A similar method of endoderm-
mesoderm formation is described by Kennel (1888) in Peripatus
edwardsi, there being here a blastoporic depression of the blastoderm
from which an internal proliferation gives rise to endoderm and to
ventrolateral bands of mesoderm. With eggs having much yolk,
meroblastic cleavage is the rule. The egg nucleus divides within the
yolk, and the cleavage nuclei enclosed in small masses of cytoplasm
migrate to the surface and form a blastoderm. In Peripatoides novae-
zealandiaec, however, according to Sheldon (1888), the blastoderm

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—S NODGRASS 53

lies beneath a superficial layer of yolk, the early embryo in this case
being a sac not only containing yolk, but also surrounded by it. The
outer yolk is later absorbed. In this species the manner of germ-
layer formation has not been definitely determined, but the endoderm
cells appear within the yolk, and the mesoderm takes the form of
two widely separated bands along the sides of the embryo, in which
the coelomic sacs are formed.

Fic. 22.—Early developmental stages of Onychophora.

A-D, successive embryonic stages of Peripatopsis capensis Grube, showing
elongation and closure of the blastopore except at oral and anal extremities,
and forward growth and segmentation of mesoderm bands (from Balfour,
1883). E, young embryo of Eoperipatus weldoni Evans, ventral view, mouth
covered by external yolk (from Evans, 1902). F, young embryo of Pert-
patopsis moseleyi Wood-Mason with open blastopore (from Bouvier, 1905).
G, cross-section of embyro of Peripatopsis capensis through open blastopore
(from Balfour, 1883). H, cross-section of embryo of Eoperipatus weldoni,
blastopore covered with yolk (from Evans, 1902).

An, anus; Bpr, blastopore; Coel, coelomic cavity; 2Coel, 3Coel, second and
third coelomic cavities; Ecd, ectoderm; End, endoderm; Gc, gastrocoele; He,
haemocoele; Msd, mesoderm; MsT, mesodermal teloblast (“primitive streak’) ;
Mth, mouth; Y, internal yolk; y, external yolk.

In Peripatopsis capensis, Sedgwick (1885) says, cleavage is complete
but unequal, the blastomeres being differentiated into four small, dark
ectodermal cells at the animal pole of the egg, and four large, clear
endodermal cells at the vegetative pole. Subsequent divisions proceed
in each group separately. The endoderm cells soon draw together
into the center of the egg, and are here overgrown by the ectoderm
until completely enclosed by the latter, except at one point where

54. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

the endoderm remains exposed on the surface. A cavity now
appears in the endodermal mass, and opens externally where the
endoderm is not covered by the ectoderm. The opening is the blasto-
pore (fig. 22 A, Bpr). With the growth of the embryo, the blastopore
lengthens to an elongate slit on the ventral surface (B). The first
observations on the development of Peripatopsis capensis were made
by Balfour (1883), who believed that the mesoderm arises in the form
of paired coelomic pouches along the edges of the elongate blastopore
where the ectoderm and endoderm are confluent. From the subse-
quent work of Sedgwick, however, it appears that the mesoderm in
P. capensis is generated from an opaque area of the blastoderm
situated behind the posterior end of the blastopore (A, B, C, MsT).
From this area, or ‘“‘primitive streak,” there takes place an internal
proliferation of cells, which, migrating forward in each side of the
embryo, produce two ventrolateral mesoderm bands along the margins
of the blastopore (B). The bands then break up into sections that
mark the primitive somites of the embryo, and later are excavated by
the coelomic cavities (G, Coel). The elongate blastopore finally closes
by the fusion of its lips, except at the two ends, which become the
primary mouth and the primary anus (D, Mth, An).

The development of the endoderm of Eoperipatus weldoni, as
described by Evans (1902), is again different from that of Peri-
patopsis capensis. ““The endodermal elements,’ Evans says, “are
derived from the lips of the blastopore and travel inward along the
outer layers of the yolk, which is at first devoid of nuclei.” Here,
evidently, is a process suggesting invagination. On the surface of
the yolk the endoderm cells form a complete investing layer, but later
some of them invade the yolk, probably bringing about its partial
digestion, and then again most of these cells return to the surface,
where they reconstruct a permanent endodermal sac containing the
yolk (fig. 22 H, End). A few endodermal cells, however, remain
within the yolk. The mesoderm of Eoperipatus weldom, according
to Evans, is formed in the same way as described by Sedgwick for
Peripatopsis capensis, that is, from a proliferating area of the blasto-
derm situated immediately behind the blastopore (E, MsT).

Considering the various processes by which the organization of
the onychophoron is accomplished in the embryo, it would appear
that the manner of development has little significance. In extreme
cases the assembling of the germ layers seems to be almost haphazard.
Sheldon (1888) observes of Peripatoides novae-zealandiae that the
embryo might be said to be formed “by a process of crystallizing out
in situ from a mass of yolk, which is a protoplasmic reticulum con-

NO. 6 ANNELIDA, @NYCHOPHORA, AND ARTHROPODA—SNODGRASS 55

taining nuclei.” Among the early developmental phases of the Ony-
chophora, however, we cannot fail to note two important likenesses
to annelid development. The first is the elongation of the blastopore
on the ventral surface of the embryo as it occurs in Pertpatopsis
capensis (fig. 22 A, B), followed by the closure of its median part
(C), finally leaving only the persistent oral and anal apertures at the
two extremities (D). We have here evidently a condition even more
generalized than in the annelids, in which the anus is usually a secon-
dary perforation. The second suggestion of annelid development,
shown in several onychophoran species, is the forward growth of the
mesoderm as bands of cells generated from a proliferating area of
the blastoderm situated behind the blastopore (fig. 22 A, B, C,
E, MsT). The mesoderm is, therefore, a teloblastic product, though
it is not possible to identify in the generative area a primary pair of
teloblastomeres. It would appear, however, that the onychophoran
mesoderm may not be entirely of teloblastic origin, for Sedgwick
(1887) finds that the forwardly growing bands in Peripatopsis
capensis are augmented by cells proliferated from the lips of the
blastopore along the lines where ectoderm and endoderm meet. The
later development of the mesoderm is unquestionably a strictly
homologous process in both the Annelida and the Onychophora, for
in the latter, as in the annelids, the primarily solid mesoderm bands
are first segmented corresponding with the body somites (fig. 22 B),
and then excavated by coelomic cavities (C, G).

Beyond the early stages of cleavage and germ-layer formation the
course of onychophoran ontogeny is well standardized and gives a
good basis for comparison of the Onychophora with the Annelida on
the one hand, and with the Arthropoda on the other. It will be found,
however, that many of the irregular earlier processes of onychophoran
development are duplicated among the Arthropoda.

THE NERVOUS SYSTEM

The onychophoran nervous system includes a brain situated in the
head above’and before the decurved anterior end of the stomodaeum
(fig. 32 A, Br), and two long, widely separated nerve cords (NC)
extending from the brain to the posterior end of the body, where they
appear to be continuous in an arc above the rectum. The cords are
connected by numerous ventral commissures (Com), and they give
off in each segment a series of dorsal nerves against the body wall
(fig. 24 .B) and ventral nerves that go downward to the legs and
other ventral parts. Opposite the legs the nerve cords are slightly

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

thickened, but they have no differentiated ganglia, since the neuro-
cytes are scattered along their lengths.. The brain, on the other hand,
is a well-developed, bilobed cerebral body (fig. 25 A, B) extending
horizontally forward from the anterior ends of the nerve cords (C).
It bears anteriorly the large antennal nerves (AntNv), laterally a
pair of small optic lobes supporting the eyes (£), and ventrally a
pair of small pear-shaped bodies (B. C. rVO). Numerous other

Fic. 23.—Development of the onychophoran head and anterior body region
as shown in three embryonic stages of Peripatus edwardsi Blanchard, ventral
view. (From Kennel, 1888.)

A, young embryo with large prostomial cephalic lobes, postoral jaw appen-
dages (J) and oral papillae (OP) resembling legs. B, older embryo with
prostomial antennal rudiments, jaws approaching mouth and surrounded by
circumoral fold (cof), ventral organs (VO) becoming differentiated. C, still
older embryo; jaws with definitive form, retracted into preoral cavity, ventral
organs more distinct; head region composed of procephalic lobes, jaw somite,
and somite of oral papillae.

a3V O, anterior ventral organ of papillar somite; cof, circumoral fold; J, jaw;
L, leg; Lm, labrum; Mth, mouth; OP, oral papilla; PrC, preoral mouth cavity ;
Prst, prostomium ; p3/’O, posterior ventral organ of papillar somite; S/O, orifice
of salivary gland; zr’ O, ventral organ of preoral cephalic lobe; 2VO, ventral
organ of jaw somite; 3/O, ventral organ of somite of oral papillae (subdi-
vided into anterior and posterior parts) ; 4/O, 5VO, ventral organs of first and
second leg somites.

small nerves are given off from the brain (fig. 24 A), among which
are anterior ventral nerves that go to the mouth and the circumoral
fold, a dorsal median nerve (f) that turns downward and posteriorly
on the dorsal surface of the stomodaeum, a pair of posterior stomo-
daeal nerves (7), and the nerves of the jaws (j), which arise from
the nerve cords just behind the brain.

The entire central nervous system of the Onychophora is developed
in the embryo from a series of paired ventral thickenings of the
ectoderm known as the “ventral organs” (fig. 23 B, C, VO), which

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS = 57

correspond with the embryonic somites, except that the first pair
(1VO) lies on the preoral head region. Whether these thickenings
represent primitive organs or are merely embryonic structures is
open to question, but they suggest the paired tubercles on what may
be the ventral surface of Xenusion (fig. 21C). From the inner
surfaces of the ventral organs of the body are differentiated the

Fic. 24.—Nervous system of the head and of a body segment of Peripatus
tholloni Bouvier. (From Fedorow, 1926, 1920.)

A, diagram of brain and anterior parts of nerve cords, with bases of nerves,
dorsal view. B, nerve cord and peripheral nerves of left side of a body segment,
lateral view.

@, sensory antennal nerve; aip, anterior interpedal nerve; Ant, antenna; b,
motor nerves of antenna; Br, brain; c, optic nerve; Com, nerve commissure;
d, lateral dorsal nerve; E, eye; e, nerve to circumoral fold; f, median dorsal
nerve; g, nerve to dorsal muscles of head; 4, commissural nerve from f to g;
i, stomodaeal nerve; JCom, first ventral commissure; j, k, nerves of jaw; L,
leg; J, nerves of oral papilla; m, », 0, nerves of first leg segment; Ip, 2), first
and second pedal nerves; r1pp, 2pp, first and second postpedal nerves; pip, pos-
terior interpedal nerve; Iprp, 2prp, first and second prepedal nerves; v, ventral
nerves.

ventral nerve cords; the outer parts are then gradually reduced in
size until finally they disappear as distinct areas of the epidermis.
When the nerve strands become free cords within the body they do
not approach each other or unite as do the nerve cords of most anne-
lids or arthropods; on the contrary they move farther apart until they
take positions along the sides of the body on a level with the leg
bases (fig. 29, NC). The definitive cords, moreover, lie laterad of

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

series of dorsoventral lateral muscles (dvm) attached dorsally and
ventrally on the body wall. A condition thus arises in the Ony-
chophora that has no counterpart in the annelids or arthropods, for
in the latter the nerve cords, even when laterally situated, have no
barrier to a median approximation or union.

The major part of the brain, from which arise the antennal and
optic nerves, is shown by Sedgwick (1888), Kennel (1888), and
Evans (1902) to be generated from the paired ventral organs of

Fic. 25.—Brain of Peripatoides novae-zealandiae Hutton.

A, dorsal surface of brain and anterior parts of nerve cords, showing posterior
antennal commissure and dorsal position of antennal tracts. B, ventral surface
of brain, with remnants of ventral organs. C, lateral view of brain and stomo-
daeal connectives.

AntCom, antennal commissure; AntNv, antennal nerve; AntT, antennal tract;
b, motor nerves of antenna; E, eye; f, median dorsal nerve; 7, stomodaeal
nerves; Com, first ventral commissure; 7, k, nerves of jaw; NC, nerve cord;
OpL, optic lobe; StCon, stomodaeal connective; 1/O, remnant of first ventral
organ.

the head (figs. 23 B, 27 B, 1/0). Evans says that the brain includes
also a pair of anterior “‘archicerebral lobes’”’ belonging to the apical
part of the head, but in his account of the embryonic development
of Eoperipatus weldoni he makes no mention of observing a separate
origin of such lobes, and attributes the entire brain, except a pos-
terior part, to the neural elements derived from the cephalic ventral
organs. The ventral organs of the head, unlike those of the body,
are finally invaginated as vesicles connected with the nerve tissue;
eventually they are reduced, but persist as the small bodies attached
to the ventral side of the brain (fig. 25 B, C, 1VO).

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 59

The small posterior lobes of the brain from which arise the posterior
stomodaeal nerves (fig. 25 B, 7), together with the adjoining parts
of the nerve cords that give off the nerves of the jaws (A, B, C, 7),
are said by Evans to be secondarily added to the antenno-ocular lobes
from the ventral organs of the postoral jaw somite (fig. 23 B, 2VO),
and Kennel clearly shows in a head section (fig. 27 A) the inclusion
in the brain of a mass of neural cells given off from these generative
centers (2VO). The definitive onychophoran brain, therefore, as

Fic. 26.—Internal structure of the brain of Peripatopsis capensis Grube.
(From Holmgren, 1916.)

AntCom, antennal commissure; AntGlm, antennal glomeruli; AntNv, sensory
antennal nerve; AntT, antennal tract; b, motor nerves of antenna; Cc, corpus
centrale; e, nerve to circumoral fold; f, median dorsal nerve; Gb, globuli of
corpus pedunculatum ; 7, stomodaeal nerves; 7, nerve of jaw; OpNuv, optic nerve;
Ped, peduncle of corpus pedunculatum; StCon, stomodaeal connective.

shown by the records of its development, and as claimed by Holmgren
(1916) and by Hanstrom (1928, 1935) from histological evidence,
would appear to be a syncerebrum composed of a prostomial fore-
brain including the ocular and antennal centers, and of a postoral
hindbrain containing the centers of the posterior stomodaeal nerves
and the nerves of the jaw appendages.

A quite different concept of the composition of the onychophoran
brain is deduced by Fedorow (1929) from a study of Peripatus
thollont, in which he attempts to correlate the cerebral nerves with
the nerves of a series of body segments (fig. 24 A, B). Fedorow

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

concludes that the anterior part of the brain, lying before the antennal
commissure and bearing the optic lobes, represents the prostomial
archicerebrum of the annelids, and that the rest of the brain is of
postoral origin, being formed of the united anterior ends of the
nerve cords extended secondarily in front of the stomodaeum. This
alleged postoral part of the definitive brain, Fedorow believes,
includes the ganglionic centers of the antennal somite, and the ganglia
of a reduced premandibular somite that has lost its appendages. The
jaw centers, he contends, are contained in the parts of the nerve cords
immediately behind the brain, from which arise the nerves of the jaw
muscles (7), and which are connected by the first postoral commissure
(r1Com). Fedorow’s elaborate analysis of the brain structure and
nerves would be more convincing if it took into account the embryonic
development of the brain; his results are entirely unsupported by
ontogenetic evidence, and are mostly at variance with observations on
the brain development reported by other investigators.

The internal structure of the onychophoran brain (fig. 26) shows
fundamental characters of the polychaete brain, and contains certain
arthropod features, but it presents also special modifications that are
not found in either the annelids or the arthropods. Corpora peduncu-
lata are well developed, each consisting of a cap of three globuli (Gb)
of small chromatic cells lying in the anterior part of the brain, and
of a large pedunculus (Ped) composed of three confluent groups of
fibers springing from the globuli cells. The sensory antennal nerves
(AntNv) coming into the anterior angles of the brain traverse the
upper part of the cerebrum in distinct antennal tracts (AntT), which
are united posteriorly in a broad antennal commissure (AntCom).
The association centers of the antennal nerve fibers, called by Holm-
gren (1916) and Hanstrom (1928, 1935) the antennal glomeruli
(AntGlm), lie laterad of the anterior ends of the corpora pedunculata,
and are said by Hanstr6m to be closely connected with neurites of
the globuli cells. In this feature, Hanstrom points out, the Ony-
chophora have a distinctly polychaete character in the brain structure,
since the antennal glomeruli of the onychophoran brain evidently
correspond with the palpal glomeruli of the polychaete brain (fig.
18 B, F, PlpGlm). On the other hand, the onychophoran brain shows
arthropodan characters in the presence of a well-developed central
body (Cc) and an antennal commissure (AntCom). But again, the
small optic lobes of the eyes (fig. 25 A, OpL) contain each only a
single ganglionic center, while all arthropods have at least two. The
optic ganglia are connected with the corpora pedunculata and with
the central body.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS- OI

The onychophoran brain thus appears to contain, as Hanstrom
(1935) has shown, a mixture of polychaetous and arthropodan char-
acters. Its origin must be found in the annelid brain; but certain
peculiar features of the onychophoran brain would seem to preclude

Ment mel Stom
\ \

Br
Fic. 27,—Developmental stages of various head structures of Onychophora.
(A, C from Kennel, 1888; B, D, E from Evans, 1902.)

A, cross-section of head of embryo of Peripatus edwardsi Blanchard through
jaws (J), showing groups of brain cells proliferated from ventral organs (2V O)
of jaw somite. B, cross-section of embryonic head of Eoperipatus weldoni
Evans, showing coelomic sacs of antenna embracing the stomodaeum, and gen-
eration of brain (Br) from cephalic ventral organs (IV O). C, section through
anterior part of head of embryo of Peripatus edwardsi, showing antennal coelomic
sac extending into antenna. D, section of embryonic head of Eoperipatus weldoni
with canal (d) from antennal coelom opening mesad of circumoral fold (cof).
E, same, more anterior section, showing antennal coelomic sacs, and a pre-
antennal sac on left side.

Ant, antenna; AntCoel, coelomic sac of antenna; AntNv, antennal nerve;
Br, brain; cof, circumoral fold; Cpr, coelomopore of antennal coelom; £, pit
of developing eye; Ecd, ectoderm; End, endoderm; Hc, haemocoele; J, jaw;
mcl, muscles of stomodaeum, Ment, mesenteron (folded forward on stomo-
daeum); PrntCoel, preantennal coelom; Stom, stomodaeum; 1/0, preoral
cephalic ventral organ; 2/’O, ventral organ of postoral jaw somite; Y, yolk.

the possibility of its having given rise to the arthropod brain. The
superficial position of the antennal nerve tracts (fig. 25 A, C, AntT),
which traverse the forebrain dorsal to the optic lobes (A, OpL), con-
stitutes a condition quite at variance with that in any arthropod, for
in all the Arthropoda the antennal nerves issue from antennal lobes

62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

that lie ventral to the optic lobes, showing that the antennae have
migrated forward beneath the eyes, and not above them as in the
Onychophora. Moreover, in the arthropod brain the antennal glomer-
uli are not immediately connected with the corpora pedunculata. The
onychophoran brain in its modern form, therefore, could not have
given rise to a brain of arthropod structure, and we can assume only
that the two types of cerebral structure have taken their origins
separately from some common progenitor not far removed from a
generalized annelid. Even the inclusion of the nerve centers of the
first postoral somite in the onychophoran brain cannot be taken as
evidence that the Onychophora are ancestral to the Arthropoda, for
in some of the lower members of the second group the first postoral
(tritocerebral) ganglia are not united with the brain.

THE EYES

The eyes of the Onychophora resemble the eyes of annelids in
structure and development. An eye of the annelid-onychophoran
type is formed from an invagination of the body wall (fig. 28 C),
which becomes closed by an approximation or union of its lips (D, E),
thus producing an inner optic vesicle (Op) beneath an outer layer
of epidermis and corneal cuticula (Cor). The cavity of the vesicle
is occupied by a crystalline lens (Ln), probably of a cuticular nature,
and its inner wall becomes the retina (Jeet). In the onychophoran
eye (A), as described by Dakin (1921), the lens is strongly convex
outwardly and rests on the thick retina (Ret). Each retinal cell (B)
is differentiated into a distal cylindrical rod (c) and a basal pigmented
part (d), which contains the nucleus (Nw), and is prolonged proxi-
mally as a nerve fiber (nf) that enters the optic lobe of the brain.
The rods appear to have peripheral striations (e), but, as shown in
cross-section (F), they do not form structures between them corre-
sponding with the rhabdoms of arthropod eyes.

LATER HISTORY OF THE MESODERM AND THE COELOMIC SACS

The mesoderm bands of the Onychophora in their forward growth
(fig. 22 B, C) continue into the head, where they form a pair of
distinct coelomic sacs in the antennal region diverging anteriorly from
the mouth (D). The cephalic coelomic sacs are described by Sedg-
wick (1887) in Peripatopsis capensis, by Kennel (1888) in Peripatus
edwardsi, and by Evans (1902) in Eoperipatus weldoni. The sacs
are at first of large size (fig. 27 B) ; posteriorly their splanchnic walls
embrace the stomodaeum (Stom) and give rise to a part of the

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 63

stomodaeal musculature; anteriorly they extend into the antennae
(C, AntCoel), and thus show their relation to these appendages.
According to Evans, the antennal sacs acquire temporary coelomo-
ducts (D, d) opening ventrally to the exterior (Cpr) within the
circumoral fold (cof). With the increase in the size of the brain,

Fic. 28.—Structure of the onychophoran eye. (A, B, F from Dakin, 1921.)

A, vertical longitudinal section of eye of Peripatoides occidentalis Fletcher,
right half of retina depigmented. B, a retinal cell, differentiated into basal
‘plasmatic part (d) and distal optic rod (c). C, D, E, diagrams of development
of an eye of the vesicular type (see also fig. 19 G). F, tangential section
through optic rods of retina.

a, b, outer and inner layers of corneal epidermis; Br, brain; c, optic rod of
retinal cell; Cor, cornea; Ct, cuticula; d, basal plasmatic part of retinal cell;
e, striated border of optic rod; Epd, epidermis; Ln, lens; mcl, muscle fibers ;
nf, nerve fiber; Nu, nucleus; OpNv, optic nerve; Opl, optic vesicle; Pig,
pigment; Fet, retina.

the antennal sacs become reduced until finally, Evans says, they appear
only as two small spaces situated above the brain in front of the eyes.

A pair of small mesoderm masses observed by Evans in an embryo
of Eoperipatus weldoni, lying above and before the antennal sacs, in
one of which a cavity was present (fig. 27 E, PrntCoel), are regarded
by Evans as representing a pair of preantennal coelomic sacs, possibly

64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

corresponding with a pair of transient rudiments of preantennal
appendages mentioned by Kennel in Peripatus edwardsi.

The coelomic sacs of the body region conform with the series of
postoral somites. The sacs of the jaw somite soon disappear. Those
of the following somites attain a high state of development during the
early embryonic period, leaving thus no doubt that the Onychophora
are descended from typically metameric ancestors. The coelomic
cavities become connected with the exterior by ventral diverticula
from the mesodermal walls of the sacs (fig. 32 C, c) that unite with
ectodermal invaginations (d), and thus form ducts opening on the
mesal aspects of the bases of the legs (D). These outlet ducts of
the coelomic sacs (coelomoducts) probably served primarily in the
early history of the Onychophora for the discharge of excretory
products and the gametes (fig. 34 A); but the coelomic sacs of the
somites anterior to the somite of the definitive genital outlets become
differentiated into dorsal gonadial and ventral nephridial compart-
ments (B, C, a, b). The gonadial compartments eventually disappear
except in a few posterior segments where they unite to form the
gonads; the nephridial compartments are reduced to the form of
delicate vesicles at the inner ends of the coelomoducts (D, b), and
thus persist as end-sacs of the definitive nephridia. In the somite of
the genital outlet the entire coelomic sacs (figs. 32 E, 34 E, a, b) with
their coelomoducts (d) are converted into the lateral genital ducts.
The sacs of the second postoral somite become the salivary glands
that open into the preoral mouth cavity. Derivation products of the
coelomic walls include the entire muscular system, the dorsal pulsating
blood vessel (fig. 29, DV’), and a muscular dorsal diaphragm (DDph)
beneath the blood vessel.

THE SOMATIC MUSCULATURE

The body musculature of the Onychophora is in general similar to
that of the annelids in so far as it consists mostly of flat sheets or
bands of circular, oblique, and longitudinal fibers closely applied to
the integument throughout the length of the animal (fig. 29), but it
includes a series of lateral dorsoventral fibers (dum) along each side
of the body cavity, which have no representatives in annelid muscu-
lature. These lateral muscles divide the body cavity into a median
compartment (mBC) containing the alimentary canal (A/Cnl) and
the slime glands (S/mGld), and lateral compartments (JBC) enclos-
ing the salivary glands (S/G/d), the nephridia (Nph), and the nerve
cords (NC). The muscle fibers are all very slender, and for the

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 65

most part are not closely grouped into bundles forming specific
muscles as in the arthropods. Each fiber is invested in a delicate
sarcolemma, the nuclei are superficial, and the axis is distinctly fibril-
lated but shows no trace of cross striation (see Camerano, 1897).
The following account of the onychophoran body musculature is
based on a study of Peripatoides novae-zealandiae. When the body
is laid open from above there are exposed on each side three sets of

DS DV dm
DDph WA Aad sine
gE ROR Tw Ze

Fic. 29.—Cross-section of middle body region of Peripatoides novae-zealandiae
Hutton, showing position of principal organs, diagrammatic.

AlCnl, alimentary canal; Com, commissure of nerve cords; DDph, dorsal
diaphragm; dm, dorsal muscles; DS, dorsal sinus; DY, dorsal blood vessel;
dvm, dorsoventral lateral muscles; /BC, lateral compartment of body cavity;
mBC, median compartment of body cavity; NC, nerve cord; Nph, nephridium ;
Npr, nephropore, S/Gld, salivary gland; SlmGld, slime gland (reservoir) ;
vm, ventral muscles.

fibers. Dorsally is a broad, thin band of internal dorsal longitudinal
fibers (fig. 30, 1), the more median fibers beginning anteriorly at the
bases of the antennae, the more lateral ones behind the bases of the
oral papillae. Ventrally is a much narrower band of ventral longi-
tudinal fibers (2) lying along the midventral line. Between the dorsal
and ventral longitudinal muscles is a series of flat, closely adjacent,
straplike lateral dorsoventral muscles (fig. 29, dum, fig. 30, 3), begin-
ning anteriorly midway between the oral papillae and the first legs.
When fully exposed, however, these lateral muscles are seen to be

5

66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

nearly semicircular in extent (fig. 29), since they are attached dorsally
high up on the back external to the dorsal muscles, and ventrally along
the midline of the body external to the median ventral muscles.

By removing a section of the lateral muscles and the more lateral
fibers of the dorsal muscles (fig. 30, left), there will be exposed two
flat external laterodorsal longitudinal muscles (4, 5) lying above the
leg base, an external lateroventral longitudinal muscle (6) mesad of
the leg base, two dorsal muscles of the leg (7, 8), and a layer of

Fic. 30.—Muscles of body wall of Peripatoides novae-zealandiae Hutton.
The various muscle layers exposed on right side of three successive segmental
areas.

1, dorsal longitudinal muscles; 2, ventral longitudinal muscles; 3, dorsoventral
lateral muscles; 4, 5, internal and external laterodorsal longitudinal muscles ;
6, lateroventral longitudinal muscles; 7, dorsal promotor of leg; 8, dorsal re-
motor of leg; 9, internal oblique muscles; ro, external oblique muscles (9 and
10, reversed in position between legs) ; 12, ventral promotor of leg; 12, ventral
remotor of leg; 13, circular muscles.

oblique muscles (9, 10). The fibers of the leg muscles penetrate
between the oblique fibers to make attachments on the body wall.
The oblique muscles (fig. 30, 9, 10) lie external to all the other
muscles thus far described. They consist of two thin sheets of fibers
crossing each other at right angles in opposite directions. The fibers
that are internal on the back (9) go from above downward and for-
ward; those that are external dorsally (zo) go downward and
posteriorly. Just above each leg, however, a broad band of the ex-
ternal fibers becomes internal by crossing over a similar band of the
otherwise internal fibers (9) going below the leg from behind. Between

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 67

each two successive legs, therefore, the relation of the two sets of
oblique fibers is reversed. On the venter all the fibers again take the
same relative position that they have on the back. The two sets of
oblique fibers arise on the integument close to the middorsal and
midventral lines, and are hence not continuous from one side to the
other. External to the oblique fibers may be seen the anterior and
posterior ventral muscles of the legs (fig. 30, right, zz, 12).

Finally, outside all the other muscles of the body wall, are the
circular muscles (fig. 30, 13). They consist of extremely fine fibers
closely adherent to the inner surface of the integument, and are
apparently continuous across the middorsal and midventral lines.

A few other body muscles occur in the region of the mouth, and
the jaws have an elaborate musculature quite different from the
musculature of the legs (fig. 21 F).

THE SEGMENTAL APPENDAGES

The appendages of the Onychophora include the antennae, the
jaws, the oral papillae, and the legs. Their rudiments appear in the
embryo as conical outgrowths of the body wall (fig. 23). The an-
tennae arise from the anterior angles of the cephalic lobes (B, Ant)
and retain this position. The jaws, which are the appendages of the
first postoral somite, arise posterior to the mouth (A, J), but later
they migrate mesally and forward (B), and are finally buried in the
preoral mouth cavity (C), where they become reduced to a pair of
double flattened hooks (fig. 21 F) converging in a horizontal plane
beneath the mouth (D, J). The oral papillae are the appendages of
the second postoral somite (fig. 23 A, OP), but in the definitive state
they take a more anterior position at the sides of the mouth (fig. 21 D,
E, OP). The legs retain their primary lateroventral positions (fig.
23 B, C), and show but little variation in their final structure.

The onychophoran appendages in their development give no evi-
dence of having been derived from polychaete parapodia; they have
no cirri or bristle sacs, and nothing suggests that they are composite
organs formed of notopodial and neuropodial elements. The terminal
claws of the onychophoran leg in no way resemble parapodial chaetae,
and the general structure and musculature of the leg has little in
common with a parapodium, except features that adapt each appen-
dage to forward and backward movement on its base. On the other
hand, the segmental appendages of the Onychophora and the Arthrop-
oda have the same manner of origin and growth in the embryo, the
organs in each case being hollow musculated lobes of the body wall,

68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

and it is only in their later development that they assume the structure
characteristic of the adult appendages in each group.

An onychophoran leg (fig. 31 A) is a hollow, conical outgrowth
of the body wall terminating in a small pedal lobe bearing a pair of
decurved claws. The leg integument is thrown into permanent circular
folds, which on the thick basal part of the limb are covered with

oO

°

oO
oO

e205 a2
LoS" 0 0

Fic. 31.—Structure and musculature of an onychophoran leg, Peripatoides
novae-zealandiae Hutton.

A, anterior view of a leg. B, lateral view of distal part of leg. C, horizontal
section of basal part of leg. D, section of more distal part of leg. E, diagram-
matic vertical section of distal part of leg. F, mesal view of leg. G, section
of entire leg in transverse plane of body.

a, b, c, d, distal nontuberculate rings of leg; e, claw-bearing pedal lobe; Nor,
nephropore; Un, claws; 14, transverse muscle of leg base; 15, peripheral muscles
of basal part of leg; 16, anteroposterior septal muscles of leg; 17, flexor muscle
of leg; z7a, flexor of distal leg rings; 18, circular muscles of foot; roa, I9b,
two-branched retractor of claws.

bristle-bearing tubercles. The distal folds, however, form distinct
segmentlike rings (A, B, a, b, c, d) and are devoid of tubercles.
The pedal lobe (e) appears to be a larger terminal ring bearing the
claws (Un).

The leg is movable anteriorly and posteriorly on the obliquely
transverse axis of its base by the four somatic muscles (fig. 30, 7,
8, 11, 12) that converge from the body wall into its basal opening.
These muscles undoubtedly serve principally as promotors and re-

NO.6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 69

motors, but are probably also levators and depressors of the leg as
a whole. Within the leg the fibers of the four somatic muscles spread
out into a thick peripheral layer of intrinsic leg fibers (fig. 31 C, D,
G, 15) attached on the successive rings of the thick basal part of the
appendage. Running through the narrow axial cavity of the leg is
an antero-posterior muscular septum (76), the fibers of which diverge
among those of the peripheral layer to the anterior and posterior walls
of the leg (C, D). The rest of the leg muscles, except a slender
transverse basal muscle (G, 14), are motors of the distal rings and
of the claws. The former include a bundle of fibers (17) arising
mesally in the leg base (G), with its fibers distributed to the ventral
walls of the distal rings (E, G), and a series of strong circular
muscles (18) in the pedal lobe. The claws are provided with a large
two-branched muscle (E, G, 19), the larger branch arising in the
base of the leg (G, roa), the other in the distal part (EF, G, rgb);
the short common terminal part is inserted dorsally between the bases
of the claws. The claw muscle is, therefore, a levator, or extensor,
of the claws and has no antagonist.

It is quite reasonable to suppose that the onychophoran leg is a
prototype of the arthropod limb, but if we look for structural resem-
blances in these two sets of locomotor organs we find few such, if
any at all. The differentiation of the onychophoran leg into a thick
basal part and a slenderer distal part, and the individualization of the
distal rings, on which muscle branches are separately inserted, might
be seen as an incipient segmentation. There is, however, no actual
parallelism between the structure of the onychophoran leg and that
of any arthropod leg, so that all we can say of the former is that it
suggests a mode by which segmentation might arise in an ambulatory
appendage. We may conclude, therefore, that the appendages of the
Onychophora and the appendages of Arthropoda have had a common
origin as lobiform outgrowths of the body wall containing extensions
of the somatic muscles. The common need of a mechanism for an-
terior and posterior movement of each appendage on its base then
brought about a differentiation of the extrinsic parts of the limb
muscles into promotors and remotors, while the parts of the muscles
within the leg were elaborated to give greater efficiency to movements
of the leg itself. The further course of evolution producing segmen-
tation and correlated musculation in the limb evidently has proceeded
independently in the Onychophora and the Arthropoda from a very
primitive common beginning, and has gone much farther in the
Arthropoda than in the Onychophora.

7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

THE RESPIRATORY ORGANS

The Onychophora are provided with numerous fine tubular in-
growths from the body wall, which undoubtedly serve for respiration,
and are therefore termed tracheae, though it is possible that ana-
tomically they are more of the nature of insect tracheoles. The
tubules, which are only one to three microns in diameter, arise in
dense bundles (fig. 32 B, Tra) from small flask-shaped pits (tp) of
the integument, and extend long distances into the body cavity. The
tracheal pits may be very numerous; they occur on all parts of the
body, on the head, and around the mouth, but they are most abundant
on the back, where several may occupy the space of a square milli-
meter. For the most part the pits are irregularly distributed, but in
some species they are arranged in longitudinal rows. The tracheal
bundles issuing from the inner ends of the pits contain large, con-
spicuous nuclei in their basal parts (Nw), which probably pertain to
the matrix cells, but the tubes themselves diverge and extend far
beyond these nuclei. According to Dakin (1920), the tracheal walls
are strengthened by excessively minute but perfect spiral fibers visible
in fresh material. In their distal parts the tracheae are branched and
go to practically all the internal organs, but their final terminations
have not been observed.

Since tracheal invaginations of the body wall are developed for
respiratory purposes in nearly all groups of terrestrial arthropods,
the mere presence of such organs can have no taxonomic significance,
any more than has the presence of gills in diverse groups of aquatic
animals. Inasmuch as invertebrates breathe through the skin in any
case, evaginations or invaginations of the integument are about the
only devices they can develop for improving their respiratory
functions.

THE CIRCULATORY SYSTEM

The blood vascular system of the Onychophora consists only of a
tubular dorsal vessel (fig. 29, DV’) extending the entire length of
the body, said to be open anteriorly and posteriorly. The walls of
the vessel consist of circular muscle fibers, and are perforated dorsally
in each segment by a pair of ostia. The tube is suspended from the
body wall by connective tissue strands, and is supported on a mem-
branous and muscular dorsal diaphragm (DDph). The diaphragm
muscles are fine, regularly transverse fibers medially attached on the
ventral wall of the blood vessel; laterally they penetrate between the
fibers of the dorsal somatic muscles and are apparently attached on

Fic. 32.—Internal structure of Onychophora, and later development of the
coelomic sacs.

A, general view of internal anatomy of Peripatoides novae-szealandiae Hutton,
female, dorsal view; muscles, nephridia, peripheral nerves, and dorsal blood
vessel omitted. B, tracheal pit of Peripatopsis capensis Grube and respiratory
tubules extending inward from it (from Schneider, 1902). C, section of embryo
of Peripatus edwardsi Blanchard, showing constriction of coelomic cavity into
dorsal gonadial compartment (a) and lateral nephridial compartment (0) ; rudi-
ments of coelomoduct (c, d) not yet united (from Kennel, 1888). D, same,
later stage, dorsal compartment (a) of coelomic sac (which later disappears
except in genital somites) and nephridial compartment (b) entirely separated,
coelomoduct (c,d) open to exterior (from Kennel, 1888). E, section of embry-
onic somite of genital outlets, coelomic sacs narrowed but not divided as in
other segments (C, D), continuous from gonads (a) through coelomoducts (d)
to exterior (from Kennel, 1888). KF, male reproductive organs of Peripatopsis
blainvillei Gay-Gervais (from Bouvier, 1902, with accessories omitted). G, sper-
matophore of same (from Bouvier, 1902).

a, dorsal gonadial compartment of coelomic sac; AcGld, genital accessory
gland; AlCnl, alimentary canal; Ant, antenna; AntNv, antennal nerve; D, ne-
phridial compartment of coelomic sac; Br, brain; c, mesodermal component of
coelomoduct; Com, nerve commissures; Ct, cuticula; d, ectodermal component
of coelomoduct; Dej, ductus ejaculatorius; E, eye; Epd, epidermis; Epdm,
epididymis; mcl, muscle; NC, nerve cord; Nu, nucleus; Od, oviduct; Oe,
oesophagus, OP, oral papilla; Ov, ovary; Phy, pharynx; Res, reservoir of
slime gland; Rect, rectum; S/G/d, salivary gland; SlmGld, slime gland; Sphr,
spermatophore; Tes, testis; tp, tracheal pit; Tra, tracheal tubules; Utrs, uteri;
Vd, vas deferens; Vent, ventriculus; VO, ventral “organ”; Vsm, vesicula
seminalis.

(71)

72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

the body wall. Above the diaphragm on each side of the blood vessel
are masses of small individual cells, probably ‘“‘nephrocytes.”’ The
circulatory system of the Onychophora thus resembles that of the
Chilopoda and the Hexapoda in the simplicity of its structure. Since
many of the arthropods, in common with the annelids, have a highly
developed blood vascular system, it would seem probable that the
simpler forms represent reductions from a more elaborate primitive
system such as that of the Annelida.

THE NEPHRIDIA

The nephridialike excretory organs of the Onychophora are paired
segmental structures usually present in all the somites between the
somite of the oral papillae and that of the genital ducts, though they
may differ much in size and in the relative development of their parts.
They lie in the lateral compartments of the definitive body cavity at
the bases of the legs (fig. 29, Nph), and open externally in grooves
on the ventral surfaces of the leg bases (figs. 29, 31 F, 33 A, Npr),
except those of the fourth and fifth pairs, which in most species open
at the bases of the distal rings of the legs (fig. 33 C).

A well-developed onychophoran nephridium consists of five distinct
parts (fig. 33 A): First, beginning externally, is a short outlet duct
(Nd) ; second, a bladderlike enlargement, or reservoir (Bl) ; third,
a tubular canal (Cnl) varying in length and usually coiled; fourth,
a funnel-shaped enlargement of the inner end of the canal (Fun) ;
and fifth, a thin-walled end-sac (ESc). The walls of the funnel (B)
are relatively thick and are histologically different from the rest of
the canal; they are clothed with long vibratile cilia directed toward
the nephridial exit (see Dakin, 1920, Cuénot, 1926, Zilch, 1936).

The funnel and the canal of an adult onychophoran nephridium
are comparable with an entire metanephridium of the annelids; the
end-sac is a remnant of the coelomic sac of the embryonic somite.
The opening of the nephridial funnel into the end-sac, therefore, is
the nephrostome (fig. 33 B, Nst). The canal is developed in the
embryo as an exit duct of the coelomic sac, formed by the union of
a ventral diverticulum of the sac (fig. 32 C, c) with a tubular in-
growth (d) from the ectoderm of the same segment mesad of the
leg rudiment (D). The primitive function of the coelomoducts un-
doubtedly was the discharge of excretory products and, in the
genital segments, of the gametes. Embryonic coelomoducts occur,
according to Evans (1902), in connection with the coelomic sacs of
the antennae (fig. 27D, d), and in all the postoral somites except

No.6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 73

the somite of the jaws. During embryonic development the coelomic
sacs of those segments that eventually contain nephridia become each
constricted into a dorsal section (fig. 32 C, a) and a ventral section
(b), which soon become entirely separate compartments (D). Except
in the genital region the dorsal compartments disappear; in the
nephridial somites the ventral compartments become much reduced,
but they retain their open connections with the coelomoducts, and
persist as the delicate end-sacs of the nephridia (fig. 33, ESc).

Fun Cil

(pte er J
m SEZ os TE RES
LULA" Uf oN

Fic. 33.—Structure of onychophoran nephridia.

A, diagrammatic transverse section of leg and nephridium of mature embryo
of Peripatopsis capensis Grube (from Sedgwick, 1888). B, inner part of nephrid-
ium of Peripatoides sp., showing ciliated funnel (Fun) with nephrostome open-
ing into coelomic end-sac (from Dakin, 1920). C, diagrammatic transverse sec-
aa leg and nephridium of adult Peripatus tholloni Bouvier (from Fedorow,
1926).

Bl, nephridial bladder; Cil, cilia; Cnl, nephridial canal; ESc, coelomic end-
sac of nephridium; Fun, nephridial funnel; NC, nerve cord; Nd, nephridal duct;
Npr, nephropore; Nst, nephrostome.

It is commonly held that the excretory organs of the Onychophora
are homologous with the annelid metanephridia (see Glen, 1919).
The simple development’ of the canals as open ventral diverticula of
the coelomic walls (not of the septa), the direct opening of the canals
to the exterior on the same segment, and the occurrence of embryonic
coelomic ducts in the head, however, are all features distinctive of
the Onychophora. Considering, therefore, that there is little proba-
bility on other grounds that the Onychophora have been derived from
annelids having metanephridia, we may conclude that the open

74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

nephridia of the higher Annelida and the coelomic exits of the Ony-
chophora have been separately acquired and developed in each group.
On the other hand, there can be little doubt that the nephridial organs
of Arthropoda (antennal, maxillary, and coxal glands) are entirely
comparable with the onychophoran nephridia.

THE ORGANS OF REPRODUCTION

In the evolution of specific reproductive organs the Onychophora
are far in advance of any of the polychaete or oligochaete annelids ;
but the development and the definitive structure of the genital organs
are so closely parallel in the Onychophora and the Arthropoda that
we can scarcely question the probability of the genital systems in
these two groups having had a common origin. In fact, it is the
fundamental similarity in the genital system that would appear to
constitute the closest bond of union between the Onychophora and
the Arthropoda, and which most strongly suggests that the two groups
have been derived from a common progenitor. The germinal centers
of the Onychophora, as in the arthropods, are entirely enclosed in
gonadial sacs of coelomic derivation, and the gametes are discharged
through ducts whose lumina are continuous with those of the gonads.
Ani approach to a closed genital system is seen in the Oligochaeta in
the development of coelomic seminal vesicles containing the genital
outlet funnels, and a system as completely closed as that of the Ony-
chophora and Arthropoda is perfected in the Hirudinea; but the
ontogeny of the organs in these several groups shows that there is
no possibility of the onychophoran-arthropod reproductive system
having been evolved from that of the higher annelids.

The primary germ cells of the Onychophora become localized at
an early stage of embryonic development in the median dorsal parts
of the splanchnic walls of one or several posterior pairs of coelomic
sacs (fig. 34 A, Grm). According to Evans (1902) there are four
embryonic genital somites in Eoperipatus weldoni, while Kennel
(1888) says the germ cells of Peripatus edwardsi occur in but
one somite. Whatever the number of genital segments may be in
modern forms, we must suppose that the germ cells once occupied
most of the somites, for the early embryonic relation of the germinal
centers to the coelomic sacs is identical with the adult condition in
the Polychaeta, and undoubtedly means that in the primitive Ony-
chophora the gametes were discharged into the coelomic sacs (A,
Spz), and were liberated from the latter through the coelomoducts
(d). As we have seen, the upper parts of all the coelomic sacs between

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 75

the somite of the oral papillae and the somite of the genital ducts
become constricted from the ventral parts (fig. 32 C, a), and then
separated as independent dorsal compartments (D, a). In the pre-
genital somites the dorsal compartments disappear, but in the defini-
tive genital somites they persist as gonadial sacs containing the
germaria (fig. 34 C, G). The gonadial sacs of each lateral series,

Fic. 34.—Diagrams showing the transformation of the onychophoran coelomic
sacs and coelomoducts into genital organs and nephridia. (From Snodgrass,
1936, based on Sedgwick, 1885, Kennel, 1888, and Evans, 1902.)

A, theoretical primitive stage in which excretory products and the gametes
were discharged from the coelomic sacs through coelomoducts. B, C, D, differ-
entiation and division of the coelomic sacs into dorsal gonadial sacs (a) and
ventral nephric sacs (b), the last finally reduced (D) to end-sacs of the
nephridia. E, gonadial sacs of definitive genital segments united on each side
in a gonadial tube (G) opening through undivided coelomic sac of penultimate
somite.

a, gonadial compartment of primitive coelomic sac; A/Cnl, alimentary canal ;
An, anus; b, nephric compartment of coelomic sac; BC, definitive body cavity
(haemocoele) ; c, nephridial diverticulum of coelomic sac; Coel, coelomic cavity ;
Cpr, coelomopore; d, ectodermal part of coelomoduct; DI, dorsal blood vessel ;
G, gonad; Gdl, lateral gonoduct; G/d, accessory genital gland ; Grm, germarium ;
Msd, mesoderm ; NC, nerve cord; Nph, REAR Spz, spermatozoa; VO,
ventral ‘ ‘organ’ ’ of ectoderm.

however, unite in a continuous tube (E, G), which becomes the
definitive gonad with a germinal band in its ventral wall (C, D, G).
Furthermore, the posterior ends of the gonadial tubes open into the
coelomic sacs of the following somite, and these sacs, which maintain
their integrity, and their continuity with the coelomoducts (fig. 32 E,
a, b, c, d), become the lateral genital ducts (fig. 34 E, Gdl). Eventu-

76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

ally the apertures of the lateral ducts come together on the midline
of the venter, where they are carried inward at the end of an ecto-
dermal invagination that forms a common definitive exit tube, the
ejaculatory duct or median oviduct.

The adult reproductive organs of the Onychophora are strikingly
arthropodan in character. In the male, the testes retain the tubular
embryonic form (fig. 32 F, Tes); each discharges into a seminal
vesicle (Vsm) from which proceeds a long tubular vas deferens
(Vd), the anterior part of which is thrown into an epididymislike
mass of coils (Epdm). The ejaculatory duct (Dej) is usually long
and irregularly looped ; its opening is on the region of the penultimate
somite. Associated with the gonopore is a pair of tubular accessory
glands (AcGl/d), said to be the reduced coelomic sacs of the last
somite (fig. 34, Gld). In the female, the tubular ovaries are
united at their extremities and lie on the dorsal surface of the ali-
mentary canal in the posterior part of the body (fig. 32 A, Ov). The
oviducts (Od) proceed first forward from the posterior ends of the
ovaries, and then turn backward to unite beneath the rectum (Rect)
in a very short terminal atrium, or common oviduct, opening in the
same position as the gonopore of the male. In viviparous species the
intermediate parts of the oviducts are enlarged in a series of uterine
chambers (Utrs) containing the embryos. Sperm receptacles usually
occur on the lateral oviducts near their ovarian ends.

VI, THE ARTHROPODA

The fundamental characters of the arthropods are those of the
Onychophora and the Annelida. The three groups have in common
the following features: (1) The ventral elongation of the blastopore
and the closure of its intermediate part, resulting in the formation
of a tubular enteron with a ventral subapical mouth and a terminal
anus, and in the conversion of the preblastoporic region of the trunk
into a prostomial cephalic lobe; (2) a definitive tripartite alimentary
canal composed of the primitive endodermal enteron, and of a secon-
dary ectodermal stomodaeum and proctodaeum; (3) the differen-
tiation of a part of the mesoblast, originally formed in the posterior
end of the body, into a specific mesoderm taking the form of ventro-
lateral bands that extend forward through the entire length of the
body and penetrate into the prostomium; (4) metamerism of the
somatic ectoderm and mesoderm, involving a segmental repetition of
organs derived from these germ layers; (5) the continuity of the
acronal centers of the primary nervous system with the somatic centers

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS Ti

secondarily developed in connection with metamerism; (6) internal
cleavage of the mesoderm segments to form paired coelomic cavities ;
(7) a somatic muscular system applied against the body wall, con-
sisting primarily of an outer set of constrictor fibers running in
transverse planes, and of an inner set of contractor fibers taking a
longitudinal course, each of which may be variously amplified or
reduced; (8) the development of a blood vascular system from the
mesoderm, composed essentially of a dorsal and a ventral longitudinal
vessel connected by lateral vessels, but often reduced to a dorsal
vessel and more or less well-defined sinuses; (9) the association of
the germ cells with the walls of the coelomic sacs, and their discharge
into the coelom.

The common basic features of organization above enumerated
attest the origin of the Arthropoda, the Onychophora, and the higher
Annelida froma common ancestral form, which itself must necessarily
be visualized as a generalized annelid. It is to be assumed that the
progenitors of the three groups had already acquired a lengthened
body by the addition of secondary genital somites proliferated from
a subterminal zone of growth. Though teloblastic growth does not
appear in the ontogeny of the Onychophora, it is quite as character-
istic of certain arthropods as of the annelids.

The Arthropoda have in common with the Onychophora the follow-
ing nonannelid characters: (1) A chitinous ectodermal cuticula;
(2) segmental ambulatory appendages formed as simple outgrowths
of the body wall, which in their structure and development give no
suggestion of a community of origin with the composite parapodia
of the Polychaeta; (3) segmental excretory organs (antennal, maxil-
lary, and coxal glands) that resemble the nephridia of Onychophora
in being remnants of coelomic sacs connected with the exterior by
simple coelomoducts, but which have neither the anatomical position
nor the development of annelid metanephridia; and (4) closed gona-
dial sacs of coelomic origin, containing the germarial centers in their
walls, and connected with the exterior by a pair of coelomic sacs set
apart to serve as genital ducts. A feature characteristic of both the
Arthropoda and the Onychophora is the restoration of the haemocoele
as the definitive body cavity, resulting from the reduction of the
coelom to the cavities of gonadial and nephridial sacs, but it is not
distinctive of them because an obliteration of the coelom occurs also
in certain annelids.

The small but important assemblage of characters given above as
common to the Onychophora and the Arthropoda would seem to in-
dicate that the two groups have been evolved from the same ancestral

78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

stock, which arose from some generalized nonchaetopodous annelid ;
but since none of the modern annelids has these characters it is
evident that the annelidan progenitors of the Protonychophora-
arthropoda have left no direct descendents. The Arthropoda differ
in so many respects from present-day Onychophora that it is certain
they must have branched off from the common onychophoran-
arthropod trunk before the latter had gone far in the onychophoran
direction. Arthropod forms were highly developed and differentiated
in the early Cambrian period of geological history, and must, therefore,
have had their origin in remote pre-Cambrian times, though in the
rocks of this period there is no specific evidence of their existence.

As an individualized group, the Arthropoda are characterized
particularly by the development of hard plates in the cuticular layer
of the integument, separated by areas of flexibility. In the Mandib-
ulata sclerotization results from the presence of nonchitinous sub-
stances in the otherwise chitinous cuticula; in the Trilobita and
Chelicerata sclerotization may be due to a structural differentiation
of the chitin itself, though apparently little attention has been given
to the chemical composition of the cuticular skeleton in these groups.
Ruser (1933) describes the physical structure of “hard chitin” and
“elastic chitin” in the Ixodidae, but makes no determination of their
chemical nature.

Since the muscles are primarily attached on the body wall, the
differentiation of the latter into hard and flexible areas at once created
a possibility for unlimited development of skeletomuscular mecha-
nisms, and it is through the elaboration of such mechanisms that the
arthropods have attained their exalted position among the articulates,
and their wonderful diversity of structure. It is true, of course, that
some of them, particularly those that have taken up parasitic habits,
have renounced their birthright, and among the latter we find examples
of physical degeneration carried to such an extent that every semblance
of arthropod structure may be lost.

Sclerotization of the integument involved first a complete change
in the mechanism of body movement, for if the rings of flexibility
between segmental plates remained at the primary intersegmental
grooves, on which the longitudinal muscles are attached, there would
be little if any possibility of movement. Hence, each dorsal and
ventral plate includes the primary intersegmental groove in front,
while the areas of flexibility occupy the posterior parts of the seg-
mental regions. The sclerotized parts of the primary intersegmental
grooves, carrying the muscle attachments, thus come to form internal
ridges, or antecostae, on or near the anterior margins of the definitive

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 79

tergal and sternal plates, and the primary intersegmental grooves
become the submarginal antecostal sutures. As a consequence, a new,
secondary type of segmentation has been established, in which the
functional intersegmental rings are the membranous posterior parts
of the primary segments, and the action of the longitudinal muscles
becomes intersegmental instead of intrasegmental. A body mechanism
of this kind is typical of all the arthropods, but still it is by no means
fixed, for innumerable modifications of it have been introduced
in adaptation to the development of special structures for specific
purposes.

The acquisition of an exoskeleton necessarily limits freedom of body
movement, such as that possessed by the highly flexible annelids,
but at the same time it furnishes a mechanism by which movements
may become more specific, since the development of definite hinge
joints becomes possible, and muscles can assume more effective
antagonistic relations to each other. The longitudinal muscles lose
nothing of their efficiency, but their contraction now results in a
telescoping of the body segments. The presence of dorsal and ventral
plates, however, necessarily eliminates the constrictor effect of the
primitive circular or semicircular muscles; the latter, therefore, have
become reduced to lateral tergosternal muscles, the contraction of
which produces a flattening of the body. The primitive mechanism
of dilation and extension by unequal distribution of internal pressure
is still operative; but the potentiality of developing endoskeletal
structures gives the possibility of a new mechanism of expansion, for
the ingrowth of apodemal arms from tergal or sternal areas, on which
primarily compressor muscles are attached, may reverse the position
of such muscles to the extent that they become dilators. A separation
of contiguous plates, however, may be brought about also by the
contraction of intersegmental muscles that have been reversed by
the overlapping of the plates. All these mechanical devices and many
others are variously and often highly developed in the different
arthropod groups, and their elaboration has set the arthropods far
above the annelids and onychophorons in the power of performing
definite and specific acts. Even the wing mechanism of pterygote
insects has been built up from little more than the skeletal parts and
musculature common to the body segments. It should be observed,
however, that although the musculature of the body segments and
the appendages is fairly definite and fixed within the major arthropod
groups, there seems to be no limit to the potential genesis of new
muscles in connection with special organs, such as the male genitalia
of insects, and, furthermore, that the entire body musculature is

80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

subject to adaptive changes, which may be very extensive, as in
certain holometabolous insect larvae.

Sclerotization of the integument has affected not only the wall of
the body, but also the walls of the tubular segmental appendages, and
the latter are jointed by definite rings of flexible membrane interposed
between the resulting limb segments, or podomeres. Hence, the
arthropod limb itself has possibilities of much variety and specificity
of action. As a consequence, while probably the appendages in the
first place were all simple locomotor organs, many of them have been
converted into instruments adapted to various purposes, and those
that still subserve the locomotor function are capable of all the kinds
of mechanical progression except flying known among animals.

Concomitant with the evolution of the skeletomuscular mechanisms,
the nervous system and the sense organs have necessarily acquired a
high state of development, and the elaboration of most intricate in-
stincts has been possible because of the facility with which tools may
be produced and adapted to their ends.

The primitive arthropods, being closely related to the primitive
onychophorons, and together with the latter derived from generalized
annelids, must have been slender, many-segmented, polypodous crea-
tures resembling modern centipedes. They differed from their con-
temporaneous onychophoran relatives in having dorsal and ventral
segmental plates and specifically jointed appendages. The Protarthrop-
oda were early differentiated into primitive trilobites and primitive
mandibulate forms. From the primitive trilobites were evolved the
later Trilobita, Xiphosurida, Eurypterida, and Arachnida, while the
Protomandibulata gave rise to the Crustacea, the Diplopoda, the
Chilopoda, and the Hexapoda.

EARLY EMBRYONIC DEVELOPMENT

The processes of cleavage and germ-layer formation are so variable
among the arthropods that they can have little value in a phylogenetic
study of arthropod relationships. Cleavage, whether total or partial,
results usually in the formation of a superficial blastoderm, and the
embryo appears as a germ band on the ventral side of the egg.
Gastrulation in some of the Crustacea takes place by invagination,
but more commonly both the endoderm and the mesoderm are formed
by delamination or by proliferation from the blastoderm or the germ
band. Manton (1928) gives a precise account of the proliferation of
the germ layers and the primary germ cells from the blastoporic
region in the crustacean //emimysis, the cells of the several groups

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 81

being first differentiated on the surface of the germinal disk. The
first endoderm cells in many of the arthropods scatter through the
yolk as independent trophocytes (vitellophags) and the definitive
enteron may then be formed either by a reassembling of the cells
about the yolk, or by regeneration from intact endodermal rudiments.

The mesoderm in some of the Crustacea, Chilopoda, and Chelicerata
is proliferated forward from a posterior generative zone very much
in the manner of the sonychophoran mesoderm, and suggestive of
the teloblastic origin of the coeloblast in the Annelida. Among the
Crustacea there are in fact a few cases in which the mesoderm takes
its origin, at least in part, from a single pair of teloblastomeres derived
from the endoderm, as in the cirriped Lepas. The mesodermal telo-

Fic. 35——Early stages in the development of a cirriped, Lepas. (Simplified
from Bigelow, 1902.)

A, 8-cell stage, with large yolk-filled posterior cell. B, 30-cell stage, endoderm
surrounded by mesoderm comprising a posterior cell (Msd) of endodermal
origin, and four cells (msd) of ectodermal origin. CC, the posterior mesoblast
cell divided into mesodermal teloblasts (MsT). D, near close of gastrulation,
but with mesoderm cells still exposed.

Bpr, blastopore; Ecd, ectoderm; End, endoderm; Msd, endodermal meso-
derm; msd, ectodermal mesoderm; MsT, mesodermal teloblast; vCl, yolk-
filled cell at vegetative pole of morula.

blasts of Lepas, according to Bigelow (1902), appear in the 32-cell
stage on the posterior lip of the blastopore (fig. 35 C, Mst), and are
produced from a single mesoblast cell (B, Msd) that results from
the division of a primary yolk-filled blastomere (A, vCl) at the
posterior pole of the morula. Four other mesoblast cells, however,
are formed in Lepas from the ectodermal lips of the blastopore
(B, C, msd), and eventually the entire mesoblast sinks into the blasto-
pore (D). A separate destiny of the mesoblast from the two sources,
entoblastic and ectoblastic, has not been distinguished in Lepas, but
it is a point of much interest to note that here the mesoblast completely
surrounds the open blastopore between the ectoderm and the endo-
derm, a part of it being of endodermal and a part of it of ectodermal

6

82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

derivation. It is not difficult, then, to understand from this condition
how, in forms having a closed blastopore, the coelomic mesoblast may
arise from the entire length of the linear blastoporic area, and we
may further see some significance in the statement by Sedgwick
(1887) that in the onychophoron Peripatopsis the mesoderm bands
in their forward growth are augmented by cells derived from the
lips of the blastopore. In the more specialized types of arthropod
development evidence of teloblastic generation of the mesoderm is
entirely lost, or at least obscured, and the whole of the mesoderm
appears to be a direct product of the germ band closely associated
with the endoderm. In its full development the arthropod mesoderm
surrounds the blastopore anteriorly, since in the adult the lateral bands
of the cephalic mesoderm may be continuous from side to side in
front of the mouth.

Segmentation of the mesoderm and the subsequent formation of
coelomic sacs take place in the early embryonic stages of many Crus-
tacea and Arachnida almost as completely as in the Onychophora and
Annelida, but in the myriapods the coelomic sacs are small, and in
the insects they are for the most part represented only by cleavage
spaces in the lateral parts of the mesoderm. In all cases, however,
the walls of the sacs break down, except such parts of them as are
retained in the formation of certain organs of coelomic origin, and
the haemocoele is restored as the definitive body cavity. Probably
all muscle tissue of the arthropods is produced from the coelomic
mesoblast; though some writers have claimed that certain muscles
are produced directly from the ectoderm, the evidence is open to
question and needs closer scrutiny (see Needham, 1937).

PRIMARY AND SECONDARY SOMITES

There is ample reason from arthropod ontogeny for believing that
the arthropods have been derived, as have the annelids, from primi-
tively unsegmented ancestral forms in which metamerism first ap-
peared as a direct subdivision of the primary body region into a small
number of somites, and that the subsequent increase in the number
of somites proceeded secondarily from growth in a subterminal
zone of undifferentiated cells. This dual method of somite produc-
tion is recapitulated in the embryogeny of some of the arthropods,
and teloblastic growth is of frequent occurrence in postembryonic
development.

In the Trilobita it seems very probable, as contended by Iwanoff
(1933) and Schulze (1936), that the so-called head represents the

NO.6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 83

area of primary segmentation, for there is no doubt that the post-
cephalic segments are produced by teloblastic growth. The youngest
trilobite larvae known give no evidence of metamerism (fig. 46 A),
but there soon appears in the glabellar region four pairs of lateral
impressions or transverse grooves that divide the glabella into five
consecutive lobes (fig. 36 A). These depressions produce internal

Fic. 36.—Segmentation and tagmosis of Trilobita.

A-D, four successive stages in larval development of Liostracus linnarssoni
Brogger (from Warburg, 1925). E, Olenellus vermontanus Hall (from Wal-
cott, 1910). F, Olenellus gilberti Meek (from Walcott, 1910). G, Schmidtiel-
lus mickwitzi Schmidt, distal body segments (from Walcott, 1910). H,
Asaphiscus wheeleri Meek, example of a trilobite with distal segments united
in a caudal fan, or pygidium (from Walcott, 1916). I, Agnostis montis Matthew,
example of the group Agnostia having only two free segments between head
and pygidium (from Walcott, 1908).

fg, fixed cheek, or fixigene; fr/, frontal lobe; H, head; lg, free cheek, or
libragene; Pyg, pygidium; sp, spine; Tel, terminal lobe of body, probably the
telson; 7h, thorax; ZG, zone of growth.

ridges or apodemes most probably for muscle attachments, and their
formation, therefore, does not represent the process of segmentation
itself, but unquestionably they mark the primary intersegmental lines
of the segments united in the larval body. The first glabellar division,
known as the frontal lobe (A, fri), is continuous with a pair of
lateral areas (Jg) that become the “free cheeks” of the adult bearing
the compound eyes (fig. 46 E, /Jg). The frontal lobe, therefore, may

84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

be regarded as a part of the eye segment, or acron, and further reasons
for so regarding it will be given later. The other four glabellar lobes
must then represent four primary larval somites, the intersegmental
lines of which should, theoretically, have extended to the lateral
margins of the simple oval body before segmentation in the latter was
suppressed. The postlarval somites of the adult trilobite are generated
teloblastically (fig. 36B, C, D) from a small region of the larva
behind the glabella (A, ZG), and are, therefore, clearly secondary
somites. The definitive segments of the postcephalic series remain
distinct in some of the trilobites to the end of the body (E, F), where
there is a small terminal lobe (E, G, Tel?) that may be the telson;
in others the posterior segments are united in a tail-fan, or pygidium
(H, Pyg), and in the Agnostia (I) only two segments retain their
independence between the head and the pygidium.

The Xiphosurida in the adult stage resemble the Trilobita in so
many respects that we should expect to find an even closer approach
to the trilobite structure in their developmental stages; and, in fact,
it has been shown by Iwanoff (1933) that the primary segmentation
in the embryo of Limulus moluccanus produces four somites (fig.
37 A, I-IV), those of the chelicerae, the pedipalps, and the first two
pairs of legs, which evidently represent the four postacronal head
somites of a trilobite. Because of the large amount of yolk in the
ectoderm, embryonic metamerism appears first in the mesoderm,
which is early divided almost simultaneously into four sections corre-
sponding with the four primary somites. The preoral cephalic region
of L. moluccanus, Iwanoff says, is at first not distinctly differentiated
from the surrounding blastoderm, but later it becomes apparent as a
preoral head segment without appendages, and in an older embryo it
forms a pair of definite cephalic lobes (B, Prc). Behind the fourth
somite there is in the young embryo (A) only an unsegmented tail
piece, but at the base of this region are later generated consecutively
(B) the remaining segments of the adult, which are thus typically
teloblastic in the manner of their formation.

It would thus appear that the primary segmentation of the ancestors
both of the trilobites and the xiphosurids produced only four somites.
These four primary somites, united with one another and with the
cephalic lobe, or acron, constitute the “head” in the Trilobita (fig.
36 H, H); in the Xiphosurida they form the anterior part of the
prosoma, for in this group three following somites and part of a
fourth are combined with the four primitive somites in the anterior
section of the body (fig. 47 E). Moreover, in the Xiphosurida a
union has taken place between all the opisthosomatic somites, so that

No.6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 85

Fic. 37—Embryonic and adult segmentation of Limulus. (A-E from Iwanoff,
1933.)

A, Limulus moluccanus Linn., germ band with mesoderm divided into four
postoral somites, cephalic lobes not yet differentiated from blastoderm. B, same,
embryo with nine pairs of appendages, cephalic lobes (Prc) present. C, first
instar larva, segments of opisthosoma indicated by internal mesoderm bands
before moulting. D, first instar larva before moulting stage. E, second instar
larva. F, Limulus polyphemus Linn., young adult, veytral view, prosomatic
appendages removed, showing radial position of their bases around the central
mouth.

Acr, acron, derived from procephalic lobes of embryo; An, anus; Chi,
chilarium; Chl, chelicera; csp, caudal spine; db/, doublure; dO, dorsal ocellus ;
E, compound eye; fg, fixigene; glb, glabella; /-VJ, first six somites; L, leg;
lg, libragene; Lm, labrum; Mth, mouth; Ofl, genital operculum; Pdp, pedipalp;
Prc, procephalic lobe; vO, ventral ocellus.

86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

there is no intermediate region of free somites as in the Trilobita
and Agnostia (fig. 36 H, I).

The adult structure of Limulus contains evidence of the presence
of 14 postoral somites, the last somite being behind the last gill-
bearing segment (fig. 47 D, XIV) ; but Iwanoff (1933) says that in
the embryo rudiments of three somites appear in the postbranchial
region, giving thus a total of 16 somites anterior to the caudal spine.
The caudal spine of the Xiphosurida is often called the “telson,” but,
as shown by Schulze (1936), a comparison with the subterminal spine
of such trilobites as Mesonacis and Olenellus (fig. 36 E, F, G), which
arises from a segment some distance from the end of the body, sug-
gests that the caudal spine of the xiphosurids may not be a true
terminal structure, and that several primitive somites beyond it may
have been lost.

Studies on the embryogeny of Arachnida have not brought out any
distinction between primary and secondary somites, and the arachnids _
have no postembryonic teloblastic growth. Schulze (1936), however,
has pointed out many features in the adult structure of the arachnids,
especially in the Acarina, that suggest the trilobite type of segmen-
tation. The area of the four primary somites, he shows, is often
evident as a differentiated anterior region of the prosoma, and in the
segmentation and body form of such acarinids as Oxypleurites there
may be seen a striking general resemblance to a mesonacid trilobite.
The arachnid prosoma contains six postacronal somites, and in this
respect, therefore, is intermediate between the trilobite “head” and
the xiphosurid prosoma.

The embryonic development of segmentation in the Crustacea
has been particularly studied by Sollaud (1923) in the palaemonid
Leander, The germ band of Leander is at first V-shaped (fig. 38 A),
its two arms diverging forward on the blastoderm from a posterior
area of proliferation (GD) in the region of the blastopore, whence
also are proliferated forward two corresponding bands of mesoderm.
Each mesoderm band soon becomes divided into four consecutive
parts, which appear as four lobes on the surface (B). The germ
bands themselves gradually become less divergent, and finally their
anterior ends curve mesally and unite by a bridge between their
anterior lobes (C). At the same time the rudiments of three pairs
of appendages appear on the second, third, and fourth lobes, which
are respectively the first antennae (B, D, rAnt), the second antennae
(2Ant), and the mandibles (Md). The first lobes (Prc) have no
appendages, but they give rise to the compound eyes and the optic
ganglia. There now appear in the ectoderm of the young embryo,

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 87

Sollaud says, three transverse grooves which define the first seg-
mentation (D). The most anterior groove runs between the first
and second pairs of antennae, the next between the second antennae
and the mandibles, and the third behind the mandibles. The body of
the embryo is thus divided into an anterior prostomial head segment

Fic. 38—Early embryonic stages of a palaemonid crustacean, showing the
development of the procephalic lobes and the antennules from the unsegmented
prostomial region, and the formation of four primary body somites. (From
Sollaud, 1923.) A, Leander squilla Linn. B-F, L. serratus Pennant.

A, ventral surface of egg showing germinal disk and anterior proliferation
of germ bands. B, early nauplius stage with first appearance of appendages.
C, later stage with germ bands united anteriorly. D, nauplius stage, with
caudal papilla (CdP) differentiated, but circle of ectodermal teloblasts (EcT)
yet incomplete. E, older nauplius embryo with ventral groove (cf) in caudal
papilla. EF, metanauplius stage, with rudiments of first and second maxillae
formed on posterior part of nauplius body before generation of teloblastic
somites has begun.

rAnt, first antenna; 2Ant, second antenna; CdP, caudal papilla; EcT, ecto-
dermal teloblasts; GD, germinal disk; J, JJ, first two somites; Lm, labrum;
Md, mandible; 1Mx, 2M +x, first and second maxillae; Prc, procephalic lobe;
Prst, prostomium; Tel, telson.

(Prst) bearing the procephalic lobes and the first antennae, a second
segment (J) bearing the second antennae, a third segment (//) bear-
ing the mandibles, and a terminal unsegmented piece (CdP), which
is the caudal papilla. The embryo is now in the nauplius stage. The
first segment, bearing the optic lobes and first antennae, Sollaud

88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

claims, is the prostomium, the other two segments being the first and
second true somites (J, JJ). (See also Sollaud, 1933.)

The caudal papilla of the malacostracan embryo (fig. 38D, E,
CdP) projects from the blastoderm. In its distal part is a circle of
undifferentiated cells, ectodermal (EcT) and mesodermal, which are
the teloblasts that will generate the postnaupliar somites. Beyond
the teloblasts is the region of the telson (Tel) containing the anus
(An). In its development the caudal papilla bends forward (F)
beneath the part of the embryo contained in the blastoderm.

When the malacostracan embryo reaches the metanauplius stage
there appear at the base of the caudal papilla the two maxillary somites
and their appendages (fig. 38 F, 1Mx, 2Mx). In a study of the
development of Hemuimysis, Manton (1928) includes the two maxil-
lary somites in the part of the body produced from the teloblasts.
Sollaud (1923), however, asserts that in Leander both maxillary
somites arise from the base of the caudal papilla before the beginning
of activity in the teloblast, and that the first somite of the teloblastic
series is that of the first maxillipeds. According to Sollaud, there-
fore, the four somites of the metanauplius (F), namely, those of the
second antennae, the mandibles, the first maxillae, and the second
maxillae, are primary somites formed directly in the primitive em-
bryonic body between the acronal prostomium and the caudal papilla.
If so, it would seem to be more than a coincidence that the same
number of primary somites occurs in Malacostraca, Xiphosurida,
and Trilobita.

In most of the entomostracan Crustacea the embryo hatches in the
nauplius stage when only three pairs of appendages are present
(fig. 4B). The trunk is not yet distinctly segmented, but it con-
sists of three regions. The first region is a preoral cephalic lobe
bearing a median eye, the first antennae (zAnt), and the labrum;
the second carries anteriorly the second antennae (2Ant) and the
mandibles (J/d), and includes posteriorly the area on which the first
and second maxillae will be formed; the third region is a terminal
unsegmented lobe, the telson, at the base of which is the generative
zone from which will be formed the teloblastic somites. The nauplius,
therefore, represents an ontogenetic stage in which the body region of
the four primary somites is present, though the appendages of the
posterior two of these somites are as yet undeveloped.

The crustacean nauplius has often been likened to the trochophore
larva of the Polychaeta (fig. 4 A), and the two forms are comparable
in so far as each represents an early stage of ontogenetic development.
We cannot suppose, however, that the arthropods and the annelids

NO.6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 89

are separately derived from an ancestral form represented by the
polychaete trochophore, since the adult arthropods have too many
features in common with adult annelids that are not yet present in
the trochophore. The common ancestor of the two groups, therefore,
is to be found in a much later stage of annelid development than that
of the trochophore. The trochophore and the nauplius are specialized
larval forms, adapted in their general shape and structure to a tem-
porary pelagic life; but, since they represent an early stage of phylo-
genetic development, and probably originated as larvae at an early
phylogenetic period of evolution in their respective groups, they
necessarily show primitive characters in their basic organization.

THE CEPHALIC SEGMENTATION AND THE DEVELOPMENT OF THE BRAIN

The question of the number of segments that enters into the com-
position of the arthropod “head” has been widely investigated and
discussed, but with such lack of uniformity in the results as to lead
to the suspicion that interpretation of the observed facts has been too
much influenced by theoretical considerations. The writer believes
that a more literal acceptance of the known facts of embryonic
development in the case of the arthropod head will give a simpler
and more satisfactory concept of the fundamental cephalic structure
than that which has been current for several decades.

In the first place, it should be understood that there is no specific
“arthropod head.” The cephalic structure is a variable combination
of segments, and the number of cephalized segments may be quite
different in different arthropod groups, or even within a single major
group. The more complex types of head, such as occur in the Man-
dibulata, include an anterior procephalic region bearing the labrum,
the eyes, and two pairs of antennae, and a posterior gnathal region
bearing the mandibles, the first and second maxillae, and in some
forms the first maxillipeds or also the second maxillipeds. In the
Trilobita the so-called “head” is a combination of at least four postoral
somites with the prostomial acron, and the “prosoma” of the Chelic-
erata is a similar composite structure, except that it contains six or
eight somites. On the other hand, in many of the Crustacea, the true
head is a primitive structure corresponding with the procephalic part
of the head in other mandibulate groups. However, differences of
opinion as to the number of somites involved in the head composition
pertain chiefly to the procephalic region, since the segments of the
gnathal region are usually distinct in the embryo, and are readily
identified by their appendages.

go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

On the assumption that the Arthropoda and the Onychophora
are derived from generalized annelids, the primary head of the
onychophoran-arthropod ancestors must have been the prostomium.
The prostomium, therefore, constitutes the archicephalon in the series
of articulate animals. In the polychaete annelids the prostomium
(fig. 39 A, Prst) supports two pairs of sensory appendages, the
tentacles (77) and the palpi (P/p), and often a median anterior
tentacle, and bears dorsally the eyes and the nuchal organs, while
between it and the first somite (/) is situated ventrally the mouth
(Mth). The neural elements of the prostomium, probably including
originally a median apical ganglion and several paired ganglia devel-
oped in connection with the sensory organs (fig. 9 B), unite to form
the composite suprastomodaeal nerve mass known as the brain, or
archicerebrum (C, D, Br).

The young arthropod embryo characteristically has at the anterior
end of the body a large cephalic lobe (fig. 39 B, Acr). On this head
lobe are developed the eyes, both simple and compound (£), the
first antennae (rAnt), in some cases a pair of transient preantennal
rudiments (Prnt), and the labrum (Lim). The neural elements of
the embryonic head, which may include an anterior median ganglionic
rudiment and as many as four paired lateral rudiments, soon unite
to form the suprastomodaeal brain. The exact parallelism in structure
and development between the cephalic lobe of the arthropod embryo
and the prostomium of the polychaete worm (A) certainly suggests
a morphological identity between the two organs. In neither is there
ever any external mark of segmentation, or direct evidence of the
confluence of more primitive segments.

Sollaud (1923, 1933), from his study of the development of the
crustacean Leander, contends that the embryonic head region (fig.
38 D, E, Prst) on which are developed the procephalic (ocular) lobes
(Pre) and the first antennae (zAdnt) must represent the annelid
prostomium, since the first intersegmental groove runs behind the
first antennae, and there is no external evidence of segmentation
before it. Moreover, in the procephalic nerve ganglia, he says, only
a slight constriction occurs at an early stage between the ocular, or
protocerebral, parts and the antennal, or deutocerebral, parts. Sollaud
asserts, therefore, that there is no valid reason for the commonly
accepted view that the first antennae are homodynamous with the
following appendages in the sense that they are the appendages of a
primarily postoral somite that has been secondarily incorporated. with
the prostomium. The first antennae of Leander, he shows, remain
uniramous, while almost from the beginning the second antennae

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS QI

(D, E, 2Ant) take on the biramous structure characteristic of the
following somatic appendages. The postoral segment of the second
antennae is thus, according to Sollaud’s interpretation, the first true
somite. The same view is strongly advocated by Holmgren (1916)
and Hanstrom (1928) from a comparative study of the annelid and
arthropod brain, but, as will be shown later, the evidence adduced by
these authors from the brain structure must be qualified by facts
of development.

The principal ground for the generally accepted belief that the
acronal region of the arthropod embryo contains one or more “cepha-

Fic. 39.—Diagrams of cephalization in the Polychaeta and Arthropoda, show-
ing the relation of the annelid prostomium to the arthropod head on the assump-
tion that the first antennae are prostomial appendages.

A, an adult polychaete with prostomial tentacles and palpi, first two somites
united in the peristomium. B, an insect embryo in which the head (acron) is
an archicephalon representing the annelid prostomium, and may bear two pairs
of appendages. C, a theoretical protomandibulate arthropod, in which the head
is a protocephalon (Prtc) composed of the acron and one somite. D, a
chelicerate arthropod, in which the acron is extended laterally and dorsally
over several somites united in the prosoma.

Acr, acron (arthropod prostomium); 1Ant, first antenna (acronal appen-
dage) ; 2Ant, second antenna (appendage of first somite) ; Chl, chelicera (equiva-
lent to second antenna); &, lateral eye; J-VJ, first six somites; Lm, labrum;
Md, mandible; Mth, mouth; 1Mx, 2Mx, first and second maxillae; Pdp, pedi-
palp; Perst, peristomium; P/p, palpus; Prnt, preantenna; Prst, prostomium;
Prtc, protocephalon; TI, tentacle.

lized somites” is the occurrence of temporary coelomic sacs in this
region. However, it has not been shown that the presence of cavities
in the cephalic mesoderm is necessarily indicative of somites, and it
would seem that the burden of proof should be on the positive side
of this question.

The mesoderm bands of the annelids,.as shown in an earlier part
of this paper, extend forward in the sides of the body from their
posterior centers of propagation. In the trochophore larva the meso-
derm is arrested at the mouth, but in the later development of the
worm the bands extend into the prostomium and may here contain a
pair of coelomic cavities. While it is usually observed that the pro-

g2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

stomial coelom of the annelids is a continuation from the coelomic
cavities of the first somite, it is claimed by Binard and Jeener (1928)
that the prostomial cavities of the spionid Scolelepis fuliginosa belong
to a distinct pair of mesodermal sacs associated with the palpi. In the
Onychophora and Arthropoda the mesoderm likewise extends into
the head region at the sides of, or before, the stomodaeum (fig. 41 A),
and is usually excavated by a pair of well-developed coelomic sacs
pertaining to the antennae (C, AntCS) ; but in the arthropods there
may be formed also a pair of sacs pertaining to transitory preantennal
appendages (fig. 42 B, PrntCS), and even a third pair in the labral
region (D, LmCS). The position of the antennal sacs, as that of the
antennal rudiments themselves, is somewhat variable in different
arthropods, both structures being in some cases postoral, in others
adoral, and again preoral; in the Onychophora the antennal sacs are
decidedly preoral, though their posterior mesal ends embrace the
stomodaeum and give rise to some of the stomodaeal muscles. The
preantennal sacs are usually slightly preoral ; the labral sacs lie directly
before the mouth.

When we consider that the forwardly growing mesoderm bands, in
their fullest development, should finally meet in front of the blasto-
pore, it is evident that coelomic cavities formed in the cephalic region
must assume adoral and preoral positions with their axes centering
in the mouth (fig. 40 B). Being thus radial in position, the cephalic
coelomic sacs cannot represent “somites” in the manner of the paired
sacs lying posterior to the mouth, which are transversely opposed
to each other. Hence, the assumption that these anterior sacs
represent ‘“‘cephalized somites” is inconsistent with the anatomical
conditions that arise in the acronal region of the trunk. Moreover,
as we have seen in a study of the annelids, the coelomic sacs them-
selves do not determine metamerism ; the segmentation of the postoral
parts of the mesoderm bands is secondary to metamerization of the
primary somatic muscular system, and the coelomic cavities are later
formed probably for physiological purposes. The coelomic sacs, there-
fore, correspond with the somites in the segmented part of the trunk,
but similar mesodermal cavities might be formed for the accumulation
of waste products in an unsegmented region such as the prostomium.
The usual absence of well-differentiated coelomic sacs in the annelid
prostomium, and the fact that the fullest development of the head
sacs is found in the higher arthropods indicate that the formation of
cavities in the cephalic mesoderm is a secondary accompaniment of
advancing organization in the prostomial lobe; but the temporary

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 93

nature of the head cavities might equally suggest that they are purely
ontogenetic structures, as claimed by Faussek (1899, 1901), for
coelomic cavities in general.

The association of the antennal coelomic sacs with the antennae
and the association of the preantennal sacs with preantennal appen-
dicular rudiments suggest that in a primitive stage there may have

a

Fic. 40.—Diagrams illustrating two theories of the fundamental structure of
the Articulata.

A, the theory of radial structure, based on a supposed origin of the articulates
from a zoantharian polyp, according to which the coelomic sacs represent radial
pouches of the enteron, and the nervous system a circumoral nerve ring, seg-
mentation of the body being determined by the enteric pouches.

B, the theory adopted in this paper, which assumes an origin of the articu-
lates from a creeping wormlike ancestor, based on the facts that, though the
mouth is subapical, the anus is terminal, and that in embryonic development
segmentation precedes the formation of the coelomic sacs, which have no con-
nection with the enteron; the mesoderm, being teloblastic, grows forward, and,
in its fullest development, may surround the mouth anteriorly, and thus give
rise to a secondary radial symmetry in the prostomial region.

Bpr, blastopore; cCom, cerebral commissure; CS, coelomic sac; Mth, mouth;
NR, nerve ring; Prst, prostomium; VNC, ventral nerve cord.

been a pair of appendages in the labral region corresponding with
the labral sacs. Some writers have contended that the labrum itself
represents a pair of united appendages, but since the labrum is imme-
diately preoral, a pair of “labral” appendages in an annelid would arise
from the base of the prostomium. Perhaps, by a long stretch of the
imagination, the labral sacs might better be correlated with a hypo-
thetical pair of primitive apical prostomial tentacles (fig. 4oB),

94. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

possibly represented by the median tentacle of certain Polychaeta
(fig. 13 C), which, having a double nerve root in the brain (fig. 45 B,
C, 1TINv), might be supposed to have had itself a double origin.
However, the possibility of the median polychaete tentacle having
been formed by the union of a pair of apical tentacles is denied by
Binard and Jeener (1928).

The theory here proposed to explain the occurrence of coelomic
sacs in the prostomial region of the articulate animals has no relation
whatever to the theory of Sedgwick (1884), Lameere (1926), and
Binard and Jeener (1928) that the annelids and arthropods are
derived from a coelenterate polyp form, and therefore have funda-
mentally a radial organization (fig. 4o A). A radial structure secon-
darily affects the anterior end of the articulate trunk because of the
subapical position of the mouth (B) ; but the terminal position of the
anus creates a quite different structure at the posterior end.

The term acron (Janet, 1899) is frequently used by students of
arthropod embryology to designate the apical part of the arthropod
head that lies anterior to the first true somite; its exact application,
therefore, differs according to each writer’s interpretation of the
head segmentation. Janet defined the acron as the preantennal part
of the head. As the term is used in the present paper, the arthropod
acron is equivalent to the annelid prostomium, and is represented in
the arthropod embryo by the cephalic lobe (or lobes) bearing the
eyes, the labrum, the preantennae, and the first antennae. The pro-
stomium is primarily the anterior part of the trunk not invaded by
the blastopore (fig. 6D, Prst); the median part of the arthropod
acron is always preoral, but its lateral parts may lap backward and
extend even a considerable distance behind the mouth. The telson
at the posterior end of the trunk is not morphologically equivalent to
the acron. It is traversed by the alimentary canal, and has the anus
at its extremity; it does not contain coelomic sacs, but its represen-
tative in the annelids, the so-called pygidium, may support a pair of
tentaclelike appendages.

The principal reasons for regarding the oculo-antennal region of
the arthropod head, here defined as the acron, as representing a
primarily unsegmented archicephalon corresponding with the annelid
prostomium may be summarized as follows: (1) There is never any
external division of the acronal region into segmental areas; (2) there
is no specific evidence of the cephalization of primarily postoral
somites, except in the case of the tritocerebral somite; (3) the embry-
onic coelomic sacs of the first antennae, the preantennae, and the
labrum are formed directly where they occur in the cephalic meso-

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 95

derm, and give no evidence of having been drawn forward from
behind the mouth; (4) coelomic sacs of the acronal region, so
far as known, are best developed in the higher arthropods, and thus
do not appear to be primitive structures; (5) the protocerebral and
deutocerebral parts of the brain are always connected by preoral
commissures, the only postoral cerebral commissure being that of the
cephalized tritocerebral ganglia; (6) the mouth and labrum are in-
nervated from the tritocerebral ganglia, which would not likely be
the case if several other postoral ganglia preceded the tritocerebral
ganglia; (7) paired appendages, sense organs, and primarily discrete
nerve centers pertain both to the annelid prostomium and to the
arthropod acron; (8) the first antennae of the arthropods never have

Lm LmMsd

/

Fic. 41.—Development of the procephalic mesoderm in Orthoptera. (A, B
from Roonwal, 1937; C from Wiesmann, 1926.)

A, horizontal section of anterior end of 52-hour embryo of Locusta migratoria
Linn. showing cephalic mesoderm extending to labrum anterior to stomodaeum.
B, same of 564-hour embryo, with coelomic cavities in labral mesoderm. C, re-
construction of head of embryo of Carausius morosus Brunner, lateral view,
with developing antennal coelom, and mesoderm extending into clypeolabral
region.

Am, amnion; Ant, antenna; AntCS, antennal coelomic sac; AntMsd, antennal
mesoderm; Br, brain; Lm, labrum; LmCS, coelomic sac of labrum; LmMsd,
labral mesoderm; Msd, mesoderm; Prc, cephalic lobe; Stom, stomodaeum.

the structure or musculature of the following appendages; in the
Crustacea they are never truly biramous.

A brief review of the facts now known concerning the develop-
ment of the procephalic mesoderm and nervous system of the arthro-
pods will show that the facts are not inconsistent with the idea that
both coelomic sacs and multiple nerve centers may be formed directly
in the otherwise unsegmented acronal region, and that the phenomena
of embryonic development pertaining to the head are most easily
understood if they are taken approximately at their face value for
phylogenetic recapitulations.

The cephalic mesoderm of the arthropods is usually continuous
with the mesoderm bands of the anterior somites. In a 52-hour
embryo of Locusta, Roonwal (1937) says, “it is seen that a pair of

96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

mesoderm bands extends upward from the junction of the head-lobe
with the trunk and meet over the stomodaeum” (fig. 41 A, Msd).
The same is true of Carausius (B), as shown by Wiesmann (1926),
but in the crustacean Hemimysis, according to Manton (1928), a
part of the preoral mesoderm has an independent origin from the
germ band. .

Among the Chelicerata the cephalic mesoderm is less developed
or differentiated than in the Mandibulata. In Limulus longispina, as
described by Kishinouye (1893), the first pair of coelomic sacs in
the embryo occupies both the cephalic lobe and the cheliceral somite.
Later these sacs become partially divided by an incomplete septum
into a pair of cephalic sacs and a pair of cheliceral sacs, but the latter
soon disappear. In the scorpion, according to Brauer (1895), the
cephalic coelom is an extension of the coelomic cavities of the chelic-
eral somite, and is never shut off from the latter in a pair of specific
head sacs. Likewise in the Pedipalpida (Thelyphonus) Schimkewitsch
(1906) says the coelomic sacs of the head segment are continuous
with those-of the cheliceral segment. Kishinouye (1894) finds, on
the other hand, in the Araneida (Lycosa and Agelena) a pair of
coelomic sacs in the cephalic lobe that are entirely separate from the
sacs of the cheliceral somite. The cephalic sacs are later divided each
into two parts; the ventral sections disappear, the dorsal sections
elongate upward and form between them the cephalic aorta.

In the Mandibulata coelomic cavities associated with the first
antennae are of common occurrence in the cephalic mesoderm. A
diverticulum from each antennal sac extends into the corresponding
antenna (fig. 41 C, AntCS) and gives rise to the antennal muscula-
ture. The inner dorsal parts of the sacs, as observed by the majority
of investigators (see Wiesmann, 1926, Roonwal, 1937), grow mesally
into the space between the stomodaeum and the brain, where they
extend anteriorly and posteriorly and form the cephalic part of the
aorta, including the anterior end of the tubular aorta proper, and an
open distributing section that extends from beneath the brain to the
clypeal region. The cephalic aorta of the crustacean Hemuimysis,
however, is said by Manton (1928) to be a product of the preantennal
mesoderm.

The presence of preantennal coelomic sacs associated with small
evanescent rudiments of preantennal appendages (figs. 42 A, 43 A,
Prnt) is recorded by Heymons (1901) in Scolopendra (fig. 42 B,
PrntCS), and by Wiesmann (1926) in Carausius (F, PrntCS), and
the occurrence of coelomic cavities in the preantennal mesoderm of
Hemimysis is reported by Manton (1928), though vestiges of pre-

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS = Q7

antennal appendages are not known in the Crustacea. In the diplopod
Platyrrhacus amauros, Pflugfelder (1932a) shows that a pair of
coelomic sacs is formed in the cephalic lobes of the embryo in con-

Lm Prte

aS
PrntCs

Ant “Pat Ma iM
B CS

Fic. 42——Embryonic appendages and coelomic sacs of the procephalic region
of an insect, a chilopod, and a crustacean.

A, head (protocephalon) and two following somites of young embryo of
Carausius morosus Brunner, ventral view (from Wiesmann, 1926). B, length-
wise section through cephalic appendages and coelomic sacs of embryo of
Scolopendra (irom Heymons, 1901). C, lengthwise section through a coelomic
sac of the embryonic labral rudiment of Carausius (from Wiesmann, 1926).
D, cross-section of same through labral coelomic sacs (from Wiesmann, 1926).
E, cross-section through preantennular coelomic sacs of embryo of Hemimysis
lamornae (from Manton, 1928). F, cross-section through preantennal coelomic
sacs of embryo of Carausius (from Wiesmann, 1926).

Am, amnion; Ant, rAnt, first antenna; Lm, labrum; LmCS, labral coelomic
sac; Md, mandible; 1Mx, 2M, first and second maxillae; PntCS, postantennal
coelomic sac; Prut, preantenna (preantennule); PrntCS, preantennal (pre-
antennulary) coelomic sac; Prtc, protocephalon (acron and first somite) ; Stom,
stomodaeum.

nection with the protocerebral lobes of the brain (fig. 44 D, Per),
and a second pair in connection with the deutocerebral lobes (Der).
Hence, if there is any necessary homology between the cavities of
the cephalic mesoderm in different arthropods, the “protocerebral”’

di

98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

sacs of Platyrrhacus should represent the preantennal sacs of Scolo-
pendra, Carausius, and Hemimysis, though there are in the diplopod,
as in the crustacean, no corresponding appendage rudiments. While,
in most cases observed, the preantennal mesoderm is a part of the
general mesoderm, the preantennal mesoderm of Hemumysis is said
by Manton (1928) to have an independent origin from the germ
band just behind the optic lobes. When the arms of the V-shaped
germ band of Hemimysis later come together, the preantennal meso-
derm rudiments are approximated immediately before the mouth. In
their growth, Manton says, they extend posteriorly and embrace the
lateral and dorsal walls of the stomodaeum, their cavities entirely
disappear, and their walls give rise to a part of the stomodaeal
(‘‘stomach’’) muscles, and to the cephalic aorta.

Coelomic sacs of the labral region of the embryonic head were
first described by Wiesmann (1926) in the stick insect, Carausius
morosus, and have since been observed by Mellanby (1936) in the
hemipteron Rhodnius, and by Roonwal (1937) in a grasshopper,
Locusta migratoria. Pflugfelder (1932a) describes in the diplopod
Platyrrhacus a pair of mesodermal cavities in the “clypeus” (fig. 44 D,
Clp), but since these cavities lie immediately before the mouth, they
evidently correspond with those called “labral’’ in the insects. In both
Locusta (fig. 41 A) and Carausius (C) the head mesoderm extends
into the labrum (LmMsd) anterior to the stomodaeum (Stom), and
the cavities formed in it are thus literally preoral in position (fig. 42 C,
LmCS) ; the mesal walls of the labral sacs of Carausius are united
before the mouth (D). In Locusta, Roonwal says, the labral and
stomodaeal mesoderm is loosely continuous prior to the appearance of
the labral cavities (fig. 41 A), but when the sacs are formed the
latter are independent structures (B, LmCS). After the disap-
pearance of the cavities the coelomic cells remain as two bodies of
mesoderm that suggest similar mesodermal masses found in the
labrum of certain other insects in which corresponding cavities are
not known to occur. .

The definitive brain of the mandibulate arthropods consists of an
anterior bilobed part, including the protocerebrum and the deuto-
cerebrum, which innervate respectively the eyes and the first antennae,
and of a pair of posterior lobes, the tritocerebrum, which innervate
the second antennae when these appendages are present. The proto-
deutocerebral lobes are always united above the stomodaeum, and
thus appear to belong to the prostomial part of the head; the trito-
cerebral lobes, on the other hand, are unquestionably derived from
the postoral somite of the second antennae, and are connected by a

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 99

postoral commissure. In many cases the dorsal lobes are developed
in the embryo from a single pair of generative centers in the ecto-
derm, just as are the corresponding lobes of the brain in the Ony-
chophora and in some of the Annelida. Considering, however, that
the annelid brain, as shown in the larva of Lopadorhynchus, has
probably taken its origin from a number of discrete prostomial
ganglionic centers corresponding with the sensory organs of the
prostomium, we should expect that a more primitive condition in the
arthropods would show that the definitive brain of these animals is
likewise a composite structure formed by the union of primarily

Clp—
Jer a—

IGng- =
(SSP)

A

Fic. 43.—Embryonic development of the cephalic appendages and nerve
ganglia of a chilopod, Scolopendra. (From Heymons, 1901.)

A, anterior end of germ band with rudiments of appendages, including pre-
antennal, antennal, mandibular, and maxillary lobes, but no rudiments of post-
antennal (intercalary) appendages, though postantennal (tritocerebral) somite
marked by a pair of ganglia ([Gng). B, same, later stage (antenna removed
on left), showing ganglionic pits (gp) of ectoderm from which ganglia are
developed.

Ant, antenna; C/p, clypeus; Igp, generative pit of optic ganglion; 2gp/, pit
of protocerebral ganglion; 3g, pit of preantennal ganglion; 4g, pit of antennal
ganglion; 59), pit of tritocerebral ganglion; 6gp-Sgp, pits of mandibular and
maxillary ganglia; rGng, tritocerebral ganglion ; IL, first leg; Lm, labrum;
Md, mandible; rMx, 2M x, first and second maxillae; Prnt, preantenna.

separate ganglia. Various studies on the development of the arthropod
brain, in fact, demonstrate its diffuse origin.

The best-known example of the development of the arthropod brain
from diffuse ganglionic centers is that described by Heymons (1901)
in Scolopendra. The embryonic cephalic appendages of Scolopendra
that correspond with cerebral rudiments include the persistent an-
tennae (fig. 43 A, Ant) and a pair of transient preantennae (Prnt),
appendages of the postantennal “intercalary,”’ or tritocerebral, somite
being absent. The definitive brain of Scolopendra, according to Hey-
mons, is formed by the coalescence of an anterior unpaired ganglionic
rudiment and five paired rudiments. The unpaired rudiment arises

I0O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

directly from the ectoderm of the clypeal region of the cephalic lobes
(fig. 43 B, Clp), and appears before any of the appendages except
_ the antennae are formed. The paired rudiments are groups of gan-
glionic cells proliferated from the inner ends of small ectodermal
pits (1gp-5gp). The first of these rudiments to be formed (2g)
lie at the extremities of the median rudiment, and their cells become
closely associated with the latter to produce a cellular mass that be-
comes the lamina dorsalis of the definitive protocerebrum. Laterad
of these rudiments are formed a pair of pits (zgp) that furnish
principally the cells of the definitive frontal lobes of the brain, and
later when the eyes appear give rise also apparently to the small optic
lobes. Following the lateral rudiments of the lamina dorsalis on each
side are formed in series three other cephalic pits, which generate
respectively the primitive ganglionic centers of the preantennae
(39p), of the antennae (4gp), and of the appendageless tritocerebral
somite (5gp). The two series of neurogenic pits are continued pos-
teriorly on the mandibular, the maxillary, and the leg somites.

Heymons regards the median unpaired brain rudiment as the equiv-
alent of the annelid ‘‘archicerebrum,”’ but it would seem rather to
correspond with the ganglion of the apical plate of the polychaete
larva. The two paired rudiments that combine with the median rudi-
ment to form the definitive protocerebrum he refers also to the
“acronal,” or prostomial, part of the head, but the preantennal,
antennal, and tritocerebral rudiments he claims represent postoral
somites. The preantennal ganglia constitute at first a connection
between the protocerebrum and the deutocerebrum, but later they
merge so completely into the brain that their identity is lost in the
definitive brain structure. The deutocerebral lobes formed of the
antennal ganglia lie primarily behind the protocerebrum, but with
the forward migration of the antennae they become transposed to a
position anterior to the protocephalon and take on a conical form
with the antennal nerves issuing from their distal ends. The trito-
cerebral lobes lie beneath the deutocerebral lobes and are continuous
with the stomodaeal connectives.

The claim of Heymons that the preantennal and antennal ganglia
represent postoral somites is not substantiated by any external evi-
dence of segmentation in the corresponding cephalic region of the
scolopendrid embryo, and as represented in Heymons’ figure (fig.
43 B) these ganglia appear to be actually preoral in position. In
none of the arthropods do the true cerebral ganglia have postoral
commissures, but the preoral position of their commissures in the
brain mass, Heymons says, is to be explained ontogenetically by the

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS_ IOI

fact that the commissures are not formed until after the respective
ganglia have taken a preoral position. This proposed explanation,
however, is merely the statement of a fact that can as well be taken as
evidence that the ganglia themselves are morphologically preoral.

In the Diplopoda the embryonic rudiments of the nervous system
appear to be completely double, for no median ganglionic center has
been observed corresponding with the “archicerebral” rudiment
described by Heymons in Scolopendra. Preantennal appendages are
absent so far as known, and the tritocerebral somite always lacks
appendages, as in the Chilopoda. According to Robinson (1907) the
nervous system of a 16-day-old embryo of Archispirostreptus consists
of a pair of trilobed “archicerebral’’ rudiments situated before the
mouth (fig. 44 E, Arc), and of two ganglionated nerve cords pro-
ceeding posteriorly from the latter around the stomodaeum. The
first ganglia of the cords (AntGng), which are distinctly postoral,
Robinson claims are the antennal ganglia. The next pair, she says,
are the ganglia of the tritocerebral somite (TcrGng), which has no
appendages, and the next pair (/dGng) belong to the mandibles.
At a later stage (F), just before hatching, the “antennal ganglia”
(AntGng), to which Robinson says the tritocerebral ganglia are now
joined, still lie behind the mouth and are approximated to the median
line. It is very surprising, however, that antennal ganglia should be
postoral at such a late stage of development, and not yet united with
the protocerebrum, so much so, in fact, that the relation of these
alleged “antennal” ganglia to the antennae becomes questionable.
Robinson gives no evidence of any nerve connection between the
ganglia and the antennae (Ant); hence we might suspect that the
antennae are innervated from the posterior ganglia of the “archi-
cerebral’ groups (Arc), and that the first postoral ganglia are the
tritocerebral ganglia.

Heymons (1897) gives a brief description of the embryo of
Glomeris (fig. 44 C), in which the antennae (Ant) appear as adoral
appendages of the cephalic lobes (Prec), whence presumably they
derive their innervation.

A more detailed account of the cephalic and cerebral segmentation
of a diplopod is given by Pflugfelder (1932a) for Platyrrhacus
amauros, but it only adds to the difficulties of understanding the
development and morphology of the diplopod head. According to
Pflugfelder, the protocerebral and deutocerebral elements of the brain
appear on the surface of the young embryo of Platyrrhacus as a
single pair of preoral cephalic lobes (fig. 44 A, Prc). Just behind
the mouth is the somite of the antennae (Ant), and the latter is

102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Fic. 44—Embryonic segmentation of the head of Diplopoda as interpreted
by different investigators.

A, germ band of Platyrrhacus amauros Attems before invagination (from
Pflugfelder, 1932 a). B, longitudinal section of head of embryo of Platyrrhacus
amauros shortly before hatching (adapted from Pflugfelder, 1932 a). C, young
embryo of Glomeris (from Heymons, 1897). D, longitudinal section of germ
band of Platyrrhacus amauros just before invagination, showing preoral coelomic
sac of clypeal region and sacs of four postoral somites (from Pflugfelder,
1932 a). E, embryo of Archispirostreptus sp., about four days before hatching
(from Robinson, 1907). F, same, one day before hatching (from Robinson,
1907).

Ant, antenna; AntGng, antennal ganglion; AntNv, antennal nerve; Arc,
archicerebrum; Cl/p, clypeus (labrum); Der, deutocerebrum; Gch, gnathochi-
larium; IL, first leg; Md, mandible; MdGng, mandibular ganglion; Mz,
maxilla; 1Mx, 2M-x, first and second maxillae; Pcr, protocerebrum; Pm-x,
postmaxillary appendage, or somite; Prc, procephalic lobe; TcrGng, tritocere-
bral ganglion; VNC, ventral nerve cord.

NO.6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 103

followed directly by the mandibular (Wd) and two maxillary somites
(Mx, Pma), there being no evidence of a tritocerebral somite. In
sections the cephalic lobes show internally a distinct division into a
protocerebral rudiment (D, Pcr)and a deutocerebral rudiment (Der),
each later accompanied by a coelomic sac. It would seem to be in-
ferred from Pflugfelder’s description, though not so stated, that the
primary antennal ganglia lie in the postoral “antennal somite” (Ant),
and yet he says, “das Deutocerebrum tritt sehr frith mit den Antennen
in Verbindung durch den Antennennerv,” and he clearly shows the
antennal nerve connection with the preoral deutocerebrum (B,

aieg Per
F\ _--AntGngy

- 2-lerGne
ze —MdGne

‘ele

_ Fic. 44 G.—Germ band of a diplopod, Archispirostreptus gigas Peters, show-
ing rudiments of appendages and ganglia. (From Silvestri, 1933.)

An, anus; Ant, antenna; AntGng, antennal ganglion; rL, first leg; Md, mandi-
ble; MdGng, mandibular ganglion; Mth, mouth; Pcr, protocerebrum; TcrGng,
tritocerebral ganglion.

AntNv). The anatomical evidence here would seem to show that
the true morphological relations of the antennae are with the deuto-
cerebral ganglia, and we can only suppose, therefore, as in the case
of Robinson’s account of Archispirostreptus, that the postoral so-
called “antennal” ganglia are the tritocerebral ganglia. In any event,
the implication from Pflugfelder’s statements that the antennae are
appendages of a postoral somite, but are finally innervated from the
preoral deutocerebrum gives the impression that there is some error
involved.

The interpretation of the anterior cephalic ganglia of the diplopod
embryo given by Silvestri (1933), illustrated in Archispirostreptus
gigas (fig. 44 G), unquestionably presents the most reasonable view

104. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

that can be taken concerning the homologies of the ganglionic rudi-
ments, since it disposes of the latter in a manner entirely consistent
with the evident facts in other arthropods. According to Silvestri the
ganglia of the antennae (AntGng) are neural masses situated mesad
of the antennal bases, and the first pair of postoral ganglia (TcrGngq)
are the tritocerebral ganglia (ganglia of the intercalary somite). It
should be observed that the antennal ganglia, as shown by Silvestri,
have a preoral position and are not separated from the protocerebral
‘lobes of the brain (Per).

Among the higher arthropods the more primitive stages in the brain
development are generally not shown in embryonic recapitulation, for
the proto-deutocerebral centers are usually proliferated from the ecto-
derm as a unified ganglionic cell mass, just as in the Onychophora
and in many of the Annelida. It is observed by Baden (1936) and
by Roonwal (1937), however, that the brain of the grasshopper
(Melanoplus, Locusta) is formed from five pairs of ganglionic
centers, three of which give rise to the protocerebrum and the optic
lobes, and the other two to the deutocerebrum and the tritocerebrum,
respectively. On the other hand, Nelson (1915) finds that in the
honey bee the lateral surfaces of the primarily undivided cephalic
lobes of the embryo become directly differentiated into three areas
from which are proliferated the neural centers of the protocerebrum,
the deutocerebrum, and the tritocerebrum.

In view of the well-authenticated examples of a diffuse origin of
the cerebral ganglionic centers in the arthropods, the theory of
Holmgren (1916) and of Hanstrém (1928) that the protocerebrum
and the deutocerebrum are secondarily differentiated parts of a primi-
tive, undivided archicerebrum does not appear to be substantiated by
the facts of embryogeny. However, since the definitive brain is
evidently a conglomerate of primitively separate ganglionic centers
in the Annelida as well as in the Arthropoda, the general contention
of these authors is not invalidated, namely, that both the protocerebral
and the deutocerebral parts of the arthropod brain belong to the
preoral prostomial region of the head, and, therefore, together
represent the annelid archicerebrum.

The concept that all coelomic sacs and corresponding nerve centers
represent postoral somites seemed reasonable enough, as applied to
the arthropod head, when only antennal sacs were known; it was
somewhat stretched, though still acceptable, when preantennal sacs
were discovered ; but now that we must add a third pair of cephalic
sacs lying directly before the mouth in the labral region it begins to
look farfetched. The theory here proposed, illustrated at D of

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 105

figure 45, accepts the embryonic facts more literally. It assumes that
the archicephalic nervous system of the arthropods, as that of the
annelids, has been built up from groups of ganglionic cells centering

eTi(Pmnt) TRL ApGng: Beate an ike ee
ae YY I AGng Y a yy
Prst_ YA aij.

Plp (Ant) .

StCon )\/ SoeGng rd-

Bo C D

Fic. 45.—Suggestions of homologies between the prostomium of the annelids
and the acronal region of the arthropod head.

ee)

A, diagram of the anterior segments of a theoretical “lobopod” annelid, with
elemental ganglia corresponding with those of the arthropod cerebrum dis-
tributed on a preoral commissural arch of the nerve cords, and the potential
number of coelomic sacs of the arthropod acron shown in their possible rela-
tion to the prostomial appendages. B, diagram of the fundamental structure of
the prostomial nervous system of a polychaete annelid (from Binard and Jeener,
1928). C, reconstructed frontal section of dorsal fibrillar mass of brain and
nerves arising from it in a sedentary polychaete, Sabellaria spinulosa Leuckart
(from Binard and Jeener, 1928). D, analytical diagram of the relation of the
coelomic sacs of an arthropod to the central nerve ganglia and the associated
appendages.

acCom, anterior cerebral commissure; Acr, acron (prostomium) ; An, anus;
Ant, antenna; ApGng, apical ganglion; Br, brain; CS, coelomic sac; cv, ventral
fibrillar mass of brain; E, lateral eye; rGng, protocerebral ganglion; 2Gng,
preantennal ganglion; 3Gug, antennal ganglion; /, JJ, first and second somites;
[Gng, first somatic (tritocerebral) ganglion; mfd, dorsal fibrillar mass of
brain; Msd, mesoderm; Mth, mouth; Plp, palpus; PlpNv, palpal nerve; Put,
postantennal appendage; Prut, preantenna; Prst, prostomium; rd, dorsal root
of stomodaeal connective; SoeGng, suboesophageal ganglion; StCon, stomodaeal
connective; Tel, telson; r7/, first tentacle; 27/7, second tentacle; r7/Nv, nerve
roots of median tentacle; 27/Nv, nerve of second tentacle; / NC, ventral nerve
cord; ZG, zone of growth; 1, labral coelomic sac; 2, preantennal coelomic sac;
3, antennal coelomic sac.

upon a fibrous commissural tract arched forward around the mouth
and continuous posteriorly with the ventral nerve cords of the somatic
system. The primary cephalic ganglia included a median anterior

106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

ganglion, paired protocerebral and optic ganglia, paired preantennal
ganglia, and paired first antennal ganglia. That these ganglia belong
to the preoral acron (Acr) is shown by the fact that the paired
ganglia are always connected by preoral commissures. The cephalic
mesoderm extends forward from the somatic mesoderm bands, and,
in its fullest development, surrounds the mouth anteriorly; it may
become excavated by cavities corresponding with the first antennae
(3), the preantennae (2), and the labrum (7). The development of
the prostomial nerve ganglia and mesodermal cavities is determined
probably in all cases by external structures (appendages or sense
organs), but the acronal neuromeres and coelomic sacs, because of
their radial position around the mouth, cannot have the same relation
in the body structure as have their postoral counterparts that are
transversely opposed to each other. For a like reason there is no
prostomial metamerism of the muscular system. The same concept
may be applied to the preoral lobe of the polychaete annelids (A),
assuming that potentially the annelid prostomium might have a full
quota of coelomic sacs corresponding with its appendages, which actu-
ally it does not have. The tritocerebral somite of the arthropods thus
represents the first postoral somite of the annelids. The tritocerebral
ganglia are secondarily united with the preoral cerebrum in the Ony-
chophora and in most of the Arthropoda, and always have a postoral
commissure ; the corresponding appendages are the jaws of the Ony-
chophora, the chelicerae of the Chelicerata, and the second antennae
of the Mandibulata.

The definitive arthropod brain more closely resembles the brain of
the Polychaeta than that of the Onychophora. Its principal part is
the protocerebrum, formed of a median apical ganglion and the first
pair of lateral ganglia, with which are connected the optic ganglia.
The preantennal ganglia lose their individuality in the general cerebral
mass. The antennal ganglia form the deutocerebral lobes, but the
latter take a forward position beneath the protocerebrum, with the
result that, in the definitive condition, the antennal nerves arise
anteriorly below the optic lobes. In the Onychophora, on the other
hand, though the antennae are anterior, the brain maintains a hori-
zontal position (fig. 25 A, C) with the antennal commissure behind
the optic region, and the antennal tracts (AntT) traverse the dorsal
part of the brain above the optic lobes. The tritocerebral ganglia are
united with the primary cerebrum in the Onychophora and in nearly
all the Arthropoda, but the union would seem to have taken place
separately in the two groups, since in some of the lower Crustacea
the corresponding centers are independent ganglia on the nerve cords,
as they are in most of the Annelida.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS_ 107

EVOLUTION OF THE HEAD

The prostomial acron does not constitute the definitive head of any
known arthropod; there is always added to the acron at least one
postoral somite, and generally the definitive head includes from four
to six somites. A head composed of the acron and one somite, how-
ever, recurs so frequently, either in the adult stage or in ontogenetic
development, as to suggest that a simple head structure of this kind
(fig. 39 C, Prtc) represents the earliest stage in the evolution of the
more complex types of arthropod head. It may hence be termed the
protocephalon. The best example of a functional protocephalon is to
be seen in the anostracan Branchiopoda (fig. 50 A), in which the
definitive head is a large cephalic lobe (Prtc) bearing the eyes, both
pairs of antennae, and the labrum. The protocephalon is unquestion-
ably the primitive head of all the mandibulate arthropods. There is
no direct evidence, however, that it ever occurred as a specific stage
in the evolution of the Trilobita or the Chelicerata, and hence, in the
ancestors of these groups, and in the protarthropods generally, the
primitive head may have been merely the prostomial acron.

Crampton (1928) applies the term “‘archicephalon” to a supposed
stage in the cephalic evolution of the arthropods when the head con-
sisted of the procephalic region and the mandibular somite. That
such a stage occurred relatively late in the phylogenetic history of
the head, however, is clearly shown in the ontogeny of the Mandibu-
lata, in which the primitive embryonic head is always a cephalic lobe
bearing the first antennae and usually including the second antennal
somite, while the gnathal somites are still a part of the body region.
Antedating this protocephalic stage, however, there must theoretically
have been a truly primitive stage when there was no head structure
other than the prostomium. The prostomium, therefore, which be-
comes the acronal region of the definitive head, is the only stage in
the evolution of the arthropod head that might properly be termed
the “‘archicephalon.”

The trilobite branch of the protarthropods is characterized by a
lateral expansion of the body, produced by an extension of the lateral
margins of the tergal plates into long flat lobes (fig. 36 E, 48 D). The
dorsal surface of the body thus presents a median elevated area
(rhachis) accommodating the alimentary canal, and broad depressed
lateral areas (pleurae). On the under surface the true venter (fig.
48 D, V’) is the area between the leg bases, the areas laterad of the
legs being the ventral doublure (dbl) of the dorsum. The appen-
dages bear long coxal epipodites (Eppd) supporting branchial
lamellae or filaments.

108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

The so-called “head” of an adult trilobite (fig. 36 H, H), as we
have seen, represents the 5-segmented body of the larva (A), the
“body” segments of the adult being formed secondarily of a series
of teloblastic somites generated from a subterminal zone of growth
(ZG). The very young larva (fig. 46 A) presents a broad anterior
acronal region (Acr), and a postacronal region in which are already
differentiated the elevated median glabella (glb), which is the cephalic
part of the rhachis, and the broad lateral areas (fg) that become the
fixed cheeks of the adult (E). When the glabellar impressions appear
(B, C) the glabella is cut into five consecutive divisions, but it is
evident that the first division, or frontal lobe (C, frl), is derived
from the acron, and that the following four divisions represent the
first four postacronal somites (J-[V). With successive stages of
development (B, C, D), the lateral wings of the acron (/g) extend
posteriorly along the sides of the somites and eventually form the
so-called free cheeks of the adult’ (E, /g), on which are located the
compound eyes (E). The cephalic segmentation of the trilobite larva,
therefore, may be represented as at I of figure 46, in which the
intersegmental lines (zs-4s) are theoretically extended to the lateral
margins of the body. A median dorsal ocellus, when present, is
always situated on the glabella, but since it must belong to the acron,
it is placed on the frontal lobe in the diagram (I, dO).

In the mature trilobite head of typical structure (fig. 46 E), the
preocular part of the acronal suture (I, zs) has disappeared, but the
postocular parts become the posterior parts of the sutures known as
the “facial sutures” (fsp), the preocular parts of which (fsa) are
probably secondary lines of cleavage developed to facilitate moulting.
In some forms the facial sutures end on the lateral margins of the
head ; in others they go to the posterior margin (E), and in such cases
the genal spines are continuations of the free cheeks. On the ventral
surface of the head (I) the acronal surface is broadly inflected to
form the doublure (dbl), which carries the labrum (Lm), or “hypo-
stome,” on its preoral margin. The probable dorsal segmentation of
the adult trilobite head, therefore, may be represented diagram-
matically as shown at J of figure 46. The acron (Acr) clearly forms
an extensive part of the mature cephalic structure, since it must in-
clude the frontal lobe of the glabella (frl), the free cheeks (Jg) with
the compound eyes (£), and the doublure (F, db/) with the labrum
(Lm). Furthermore, since the dorsal ocellus often occurs far back
on the glabella (J, dO), we must assume that it is contained in a
median, tongue of the frontal lobe extended posteriorly into the
glabellar somites, because the simple eyes as well as the compound
eyes always belong to the acronal segment.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 109

The antennal appendages of the trilobites, judging from their
filamentous form in contrast with the segmented structure of the
following appendages, evidently represent the first antennae (anten-

<-Acr
glb
te

Fic. 46.—Segmentation of the trilobite “head,’ or prosoma.

A-D, four consecutive stages in the larval development of Blainia gregaria
Walcott, showing gradual posterior extension of lateral wings (free cheeks)
of acron against sides of anterior somites, and division of glabella (C) into four
segmental areas behind frontal lobe of acron (from Lalicker, 1935). E, diagram
of typical trilobite head, dorsal surface. F, diagram of ventral surface of trilobite
head, showing labrum attached to margin of doublure. G, labrum of Paedewmias
transitans Walcott, example of a stalked labrum (from Walcott, 1910). H,
head of Holotrachelus punctillosus, with segmentation obliterated in the large
swollen glabella (from Warburg, 1925). I, diagram of larval trilobite, with
head segmentation indicated. J, head of adult trilobite with probable segmenta-
tion deduced from the larval structure (I).

Acr, acron; dbl, doublure; dO, dorsal ocellus; E, compound eye; fg, fixigene
(fixed cheek) ; frl, frontal lobe (of acron) ; fsa, anterior part of facial suture ;
fsp, posterior part of facial suture; glb, glabella; /-IJV’, cephalic somites; lg,
libragene (free cheek) ; Lm, labrum; p/, palpebral lobe; rs-4s, intersegmental
sutures of head.

nules) of other arthropods; if so, according to the theory here
followed, they should belong to the acron, and perhaps had their
muscle attachments on the frontal lobe. The position of the antennal

I1o SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

bases is not exactly known, but the antennal appendages are generally
represented as arising at the sides of the labrum. The four following
segmented, leglike appendages of the head clearly pertain to the four
postfrontal somites of the dorsal shield.

Henriksen (1926), in his analysis of the segmentation of the trilo-
bite head, convincingly argues that the free cheeks bearing the com-
pound eyes must belong to the “eye segment” (acron), and that the
preocular parts of the facial sutures are secondary lines of cleavage
to facilitate moulting; but the median part of the eye segment he
believes is represented only by the narrow anterior marginal rim of
the dorsal shield before the frontal lobe. Henriksen notes, however,
the anomalous position of the median eye far back on the glabella,
and it is not clear why the reasoning by which he assigns the free
cheeks to the eye segment does not demand that the eye segment
include also the area of the median eye. The antennae, Henriksen
contends, belong to a separate postoral somite, represented dorsally
by the frontal lobe of the glabella. Furthermore, since he believes
that the trilobite head must have the same segmentation as the head
of certain Crustacea, Henriksen concludes that a second antennal
somite has been lost by the trilobites. To the writer this theoretical
elaboration of the trilobite head to give conformity with crustacean
structure appears quite unnecessary, since the trilobites are non-
mandibulate arthropods having no immediate relations with the Crus-
tacea, and their structure clearly leads into that of the Chelicerata.

The Xiphosurida, in the structure of the prosoma, show unmis-
takably their trilobite derivation, for the trilobite head is carried over
into the xiphosurid prosoma with few changes other than the inclusion
of a few extra segments, the loss of the antennae, and a differentiation
of the other appendages.

A comparison of figure 47 A with figure 46 E will show at once
the likeness of the prosomatic carapace of Limulus to the typical head
shield of a trilobite. The segmentation of the xiphosurid prosoma is
evident from the position of the limb bases on the ventral surface
(fig. 47 C), where it is seen that the anterior somites lap forward at
the sides of the labrum from behind the central mouth, while the
posterior somites curve somewhat backward. The chelicerae (Ch)
thus come to have anatomically a preoral position at the sides of the
labrum, though their somite (/) is morphologically postoral, and the
same is true of the pedipalps (Pdp) and the first legs (7). On
the inner surface of the prosomatic carapace the attachments of the
limb muscles (fig. 47 B), as depicted by Benham (1885), follow the
segmentation indicated ventrally by the limb bases. The cheliceral

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS III

Fic. 47—Segmental analysis of Xiphosurida (Limulus polyphemus Linn.).

A, young adult, dorsal surface. B, ventral surface of dorsal carapace, show-
ing muscle attachments and series of dorsal apodemes, or entapophyses (from
Benham, 1885, with dorsal ocelli added). C, ventral surface of prosoma, show-
ing segmentation as indicated by position of leg bases. D, dorsal surface of
opisthosoma, with segmentation indicated. E, theoretical approximate segmen-
tation of prosoma.

Acr, acron; Ap, tergal apodemes (entapophyses) ; Chi, chilarium; Chl, chelic-
era; dbl, doublure (ventrally inflected part of acron); dO, dorsal ocellus; E,
compound eye; fg, fixigene; fs, facial suture; J-X/V, postoral somites; L, leg;
lg, libragene; Lm, labrum; Mth, mouth; Opl, genital operculum; /, external
pits of tergal apodemes; 12a, b, f, dorsal attachments of tergo-sternal muscles
of opercular and gill somites; 20a, b, f, dorsal attachments of anterior muscles
of opercular and gill appendages; 25a-e, 28, dorsal attachments of tergo-coxal
muscles of prosomatic appendages.

T12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

muscles (24) arise near the midline just behind the dorsal eyes; the
muscles of the pedipalps (25a) take their origins farthest forward;
and the muscles of the other appendages (25b-25c) are distributed
on the following areas of the “fixed cheeks.’’ Diagrammatically, there-
fore, we may represent the segmentation of the prosomatic carapace
as given at E of figure 47. The horseshoe-shaped acron (Acr) bear-
ing the eyes encloses the region of the prosomatic somites (J-VJI/),
and sends posteriorly, between the lobes of the anteriorly curved
cheliceral and pedipalp somites, a median tongue bearing the dorsal
ocelli (dO). The structural conformity with the trilobite head (fig.
46 J) is exact, except for the greater number of somites included in
the xiphosurid prosoma.

Students of the embryology of Limulus (Kishinouye, 1893, Iwanoff,
1933) have indicated the segmental divisions of the prosoma as sub-
tending the lateral areas of the carapace bearing the compound eyes.
Branches of the segmental nerves, the “haemal nerves” of Patten and
Redenbaugh (1900), extend into these parts, but, as in the case
of the trilobites, the location of the compound eyes on the lateral plates
of the prosoma is sufficient proof that these plates belong to the eye
segment, or acron. Hence, they cannot be lateral extensions of the
median somites.

The gills of the trilobite legs, borne on coxal epipodites (fig. 48 D,
Eppd), have not been retained on the prosomatic appendages of
Xiphosurida, though an epipodite is present on the fourth leg (E,
Eppd), but gill-bearing epipodites are highly developed on the opistho-
somatic appendages, which are otherwise much reduced.

The prosomatic appendages of Limulus, except the chelicerae, as
shown by Benham (1885), have the typical arthropod coxal muscu-
lature, consisting of dorsal promotor and remotor muscles (fig. 48 F,
I, J), and ventral muscles (K, L). Of the latter, two (33, 34) are
promotors and remotors, but two others (32m, 32n) are united
proximally and evidently function as adductors. The dorsal muscles
arise on the tergal carapace (C). The ventral muscles, however, are
attached on an internal plate, or “entochondrite” (k), suspended in
the ventral part of the body by dorsal muscles (t-s). The same
structure (B) is characteristic of most of the Chelicerata, and a
similar structure occurs in the gnathal segments of many of the
Mandibulata (figs. 50 E, H, 51 B, k). Since the ventral muscles of
the appendages should primarily arise on the ventral body wall, the
“entochondrite” might be supposed to be a sternal derivative, but
Schimkewitsch (1895, 1906) claims that in the Arachnida it is pro-
duced from transformed muscle tissue, In various mandibulate

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS I13

arthropods some of the adductor fibers of the mandibles go con-
tinuously from one appendage to the other.

That the prosoma of Limulus contains at least a part of the eighth
somite is evident from several structural features, but the writer’s

Loo Ss aadan: =

[OOO

co vi / AR mS
Bnd Cxpd ©, Y

ip Tipae 228

Fic. 48.—Structure of Chelicerata and Trilobita.

A, Liobunum sp. (Phalangida), anterior view of body, showing secondary
preoral position of chelicerae above base of labrum. B, same, “endosternite”
of prosoma with adductor leg muscles (suspensory dorsal muscle not shown).
C, Limulus polyphemus Linn. (Xiphosurida), section of prosoma behind third
legs, leg muscles somewhat diagrammatic. D, diagrammatic cross-section of
a trilobite. E, Limulus polyphemus, fourth leg, with coxal epipodite. F, same,
base of a left leg, mesal view, with muscle insertions (from Benham, 1885).

a, dorsal articulation of coxopodite; b, ventral end of coxal axis; Bnd, basen-
dite; Chl, chelicera; Cp, carapace; Cxpd, coxopodite; D, dorsum; dbl, doublure;
dO, dorsal ocellus; E, lateral eye; Eppd, epipodite; 7, tergal promotor muscles
of coxopodite; J, tergal remotors of coxopodite; K, anterior ventral muscles of
coxopodite; k, ligamentous “endosternite’ on which ventral leg muscles are at-
tached; L, posterior ventral muscles of coxopodite; 3L, third leg; Lm, labrum;
Pdp, pedipalp; Tlpd, telopodite; t-s, tergal suspensory muscle of “endosternite” ;
V, venter; 26, 27, dorsal promotor muscles of coxopodite arising on carapace;
25, 28, 29, dorsal remotor muscles arising on carapace; 32m, 32n, anterior and
posterior branches of coxal adductor arising on “endosternite”; 33, 34, ventral
remotor and promotor muscles arising on “endosternite”.

former statement (1936) that the prosoma and opisthosoma of

Limulus are separated between segments VIII and [X is not strictly

correct. The attachment of the muscles and the distribution of the

nerves in this region demonstrate that the dorsal hinge between the

prosomatic carapace and the opisthosomatic carapace lies within the
8

II4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

eighth segment itself, and not behind it, a narrow anterior median
part of this segment being incorporated into the posterior margin of
the prosoma, while lateral parts of it form the anterior lateral lobes
of the opisthosomatic carapace (fig. 47 D, VIII). Six following seg-
ments of the opisthosoma are marked by the six pairs of impressions
bordering the median elevation of the carapace, and by the six pairs
of marginal spines. The intrasegmental division of the body into
movable parts is not an anomalous condition; it occurs between the
thorax and the abdomen of many insects, and is a necessary mechanical
adaptation resulting from the primarily intersegmental attachments
of the longitudinal muscles.

The six pairs of dorsal impressions on the opisthosoma of Limulus
(fig. 47 D) and a pair of similar impressions on the posterior margin
of the prosomatic carapace (p) form internally (B) a double series
of tergal apodemes, the “entapophyses” of Benham (1885), of which
the larger first pair (VIIJJAp) is on the prosoma, and the other six
pairs (IXAp-XIV Ap) are on the opisthosoma. The tergosternal
muscles of the five gill-bearing segments are shown by Benham to have
their dorsal attachments (72b-12f) at the bases of the first five opistho-
somatic apodemes, while the corresponding muscles of the opercular
segment (VJ//) arise at the bases of the corresponding prosomatic
apodemes (12a). On the other hand, while the “external branchial
muscles” of the gill segments have their dorsal attachments (20b-
20f) just laterad of the first five opisthosomatic apodemes, the corre-
sponding muscles of the operculum take their origins also on the
opisthosomatic shield, but more laterally on the anterior lateral lobes.
The muscle attachments, therefore, show that the dorsal part of the
eighth segment has been divided between the prosoma and the opistho-
soma, or, as Benham says, the first pair of tergal apophyses has been
transferred from the opisthosoma to the prosoma. The dorsal longi-
tudinal muscles between the prosoma and the opisthosoma of Limulus
have been condensed into a single large bundle of fibers, the “arthro-
tergal muscle” of Benham, and the attachments of this muscle (fig.
47 B, 78) have extended somewhat anteriorly and posteriorly on the
two body regions to acquire greater efficiency as a flexor of the
opisthosoma.

The innervation of the hinge region of the carapace gives the same
evidence of division within the eighth segment as that furnished by
the musculature. As shown by Patten and Redenbaugh (1900), the
nerves of the genital operculum proceed from the composite ventral
ganglion of the prosoma, while the corresponding somatic nerves
(‘haemal nerves” of segment VJ//) are distributed to the anterior

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS II5

lateral lobes of the opisthosoma. From a comparative study of the
position of the cardio-aortic valve in the Chelicerata, Petrunkewitch
(1922) found that the valve is always between segments ’//I and
IX, and he therefore claimed that segment V/JIJ is included in the
prosoma of Limulus. His contention is but little affected by the
modified view here shown to be in better accord with the facts. The
operculum is anatomically more closely connected with the prosoma,
from which it derives its innervation, than with the opisthososa, and
the partition of the tergum of its segment between the prosoma and
the opisthosoma, as above noted, is but a necessary adaptation to give
intersegmental action to primarily intrasegmental muscles.

The Eurypterida and Arachnida differ from the Xiphosurida in that
the prosoma includes only six somites, and in this respect they are
nearer to the Trilobita, which have only four prosomatic somites.
The eurypterids and arachnids, however, lack the lateral expansions
of the prosomatic carapace characteristic of the trilobites and xipho-
surids, and, judging from the more anterior position of the lateral eyes
(fig. 49 E, E), it seems probable that the acronal element of the
prosoma is less extensive on the marginal areas of the latter, though
medially it must include the region of the dorsal eyes (dO).

In a typical arachnid embryo (fig. 49 A) the somites are regular
transverse sections of the trunk behind the large prostomial acron
(Acr), which is produced laterally into a pair of cephalic lobes.
Ordinarily there are no appendages on the acron, but Jaworowski
(1891) has described a pair of apparent antennal rudiments in a
species of Trochosa (C, b), and Pokrowsky (1899) found two pairs
of transient precheliceral lobes in an embryo of Pholcus opilionides
(B, a, b), the second of which, he says, correspond in position with
the “antennal” rudiments described by Jaworowski. The nature of
these embryonic lobes may be doubtful, but since the trilobites have
well-developed antennae, there is no reason why embryonic vestiges
of acronal appendages might not recur in some chelicerate forms. In
adult Solpugida there is a pair of small appendages (fig. 49 F, Ant?)
arising at the sides of the epistomal lobe, which are movable by
muscles (G, mcl), and are, therefore, suggestive of being antennal
remnants.

The cheliceral somite of the arachnid embryo (fig. 49 A, J) lies
transversely immediately behind the acronal lobes; but in the adult
this somite curves forward around the sides of the labrum from
behind the mouth as in Limulus, so that the chelicerae come to have
a preoral position above the labrum (fig. 48 A, Chl), though usually
they are separated by a median epistomal bar extending downward

116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Fic. 49.—Embryonic and adult structures of Arachnida.

A, embryo of Agelena labyrinthica (from Balfour, 1880). B, embryo of
Pholcus opilionides Schranck with two lateral lobes (a, b) on acron (from
Pokrowsky, 1899). C, embryo of 7rochosa singoriensis Laxm., with possible
antennal rudiments (from Jaworowski, 1891). D, longitudinal section through
anterior end of prosoma of a phalangid (Liobunum), showing anterior tergal
attachments of cheliceral muscles. E,.dorsal surface of phalangid (Liobunum),
legs removed from coxopodites. F, epistomal lobe and labrum of a solpugid,
lateral view, showing movable appendage (Ant?) at side of epistoma. G, same,
longitudinal section, showing muscles of epistomal appendage.

a, possible preantennal rudiment of embryo; Acr, acron (cephalic lobe of
embryo); Ant?, adoral (possibly antennal) appendage of adult solpugid; )D,
possible antennal rudiment of embryo; Ap, apodeme; Chl, chelicera; Chlmcls,
cheliceral muscles; dO, dorsal ocellus; E, lateral eye; Epst, epistoma; GC,
genital chamber; Gtr, gonotreme; //Bnd, IJ] Bnd, basendites of second and
third appendages; JX.S, sternum of ninth somite; IL, first leg; Lm, labrum;
mcl, muscles; Mth, mouth; Pdp, pedipalp; Stom, stomodaeum; [-XV,, postoral
somites.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS I17

to the labrum from the frontal region of the carapace. As in Limulus
again, the chelicerae have only dorsal muscles, which arise on the
anterior part of the carapace (fig. 49 D, Chlmcls).

The ancestors of the modern Mandibulata were represented in the
more generalized members of the Protarthropoda that persisted after
the trilobite-chelicerate branch had been given off from the main stem
(fig. 54). The Protomandibulata undoubtedly retained the primitive
centipedelike form of the protarthropods, but, as shown in the embry-
ology of modern Mandibulata, the head at this stage must have been
a composite protocephalon (fig. 39 C, Prtc) formed by an intimate
union of the first somite (J) with the highly developed prostomial
acron (Acr). It carried, therefore, the eyes (E), the labrum (Lm),
the acronal appendages, or first antennae (rAnt), and the appendages
of the included somite, which became a second pair of antennal organs
(2Ant). The distinctive feature of the early mandibulates, however,
was the presence of a pair of jaws, the mandibles (Md), developed
from the bases of the appendages of the first postcephalic somite.
Probably also the appendages of the following two somites were
reduced in size and modified in a manner suggestive of their future
transformation into maxillae; and perhaps a pair of paragnathal lobes
was developed between the mandibles and the first maxillary appen-
dages, since these structures are not present in the chelicerate branch.

The Crustacea represent the first offshoot from the mandibulate
section of the arthropod stem that has given rise to a specialized group
of modern forms (fig. 54). The wide recurrence among the Crustacea
of cursorial appendages identical in segmentation with the legs of the
trilobites can leave little doubt that the primitive crustaceans were
polypodous walking animals, living on the bottom of the water or on
aquatic plants along the ocean shores, and adapted to life in the water,
as were the trilobites, by the development of branchial organs on exite
lobes of the coxopodites. According to this view, the natatory appen-
dages of swimming or purely pelagic Crustacea are legs that have
been modified secondarily for swimming purposes, just as the gnathal
appendages have been modified for feeding. It is a sound principle
of ecology that pelagic forms in all cases have been derived from
benthonic forms (see Hesse, Allee, and Schmidt, 1937, p. 179), and
the fact that many of the more generalized modern Crustacea are
pelagic is no argument that such forms are ancestral. The frequent
biramous structure of crustacean appendages is entirely a crustacean
feature, since the exopodite is a specially developed outer branch of
the basipodite, and therefore has no counterpart in the Trilobita or
in any other arthropod group.

118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Il III+IV

Prte

Fic. 50.—Cephalic structures of Crustacea in which the protocephalon (acron
and first somite) is either the definitive head, or is united with several follow-
ing somites to form a more extensive syncephalon.

A, Eubranchipus vernalis Hay (Anostraca): protocephalon a distinct head
lobe (Prtc) separate from mandibular somite (//); maxillary somites (J/J/,
IV) united with each other. B, Apus longicaudatus Le Conte (Notostraca),
dorsal view: mandibular and maxillary somites united with protocephalon, maxil-
lary tergum produced in large cephalic carapace (Cp). C, same, ventral view
of head, showing labrum, antennules, mandibles, and maxillae. D, Daphnia
pulex Degeer (Cladocera): head structure as in Apus, body covered by bi-
valved maxillary carapace. E, Eubranchipus vernalis, detached mandibular seg-
ment, anterior view, showing mandibles suspended from tergum, and mandibu-
lar musculature. F, Nebalia bipes Fabr. (Leptostraca) : bivalved carapace has
same composition as in Apus and Daphnia. G, section of head of Nebalia show-
ing muscle attachments of protocephalic appendages. H, mandibles of Nebalia,

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS_ IIQ

The crustacean head is variable in structure according to the number
of somites it contains. The most primitive crustacean head, as already
noted, is a simple protocephalon formed by the union of the trito-
cerebral somite with the prostomial acron. A head of this type occurs
in some of the Branchiopoda, and in all the Malacostraca except
Leptostraca, Amphipoda, and Isopoda.

The best example of a protocephalic head is seen in the anostracan
branchiopods. The head of Eubranchipus, for example (fig. 50 A,
Prtc), is a large cephalic capsule bearing only the eyes, both pairs of
antennae, and the labrum. Behind it is the small but distinct tergum
of the mandibular somite (JJ), which supports the large mandibles
(Md). The next following segment is evidently the two maxillary
somites united (J/7+JV), since it carries the vestigial first and second
maxillae. The muscles of the head appendages, including those of
the eye stalks, the antennules, the second antennae (in the male), and
the labrum, all take their origins on the walls of the head capsule.
The mandibular muscles, on the other hand, arise on the mandibular
tergum (E), except the adductors (KL), which are united on a
median ligament (k) and thus form a zygomatic muscle between the
two jaws.

The head of most of the other Entomostraca and of Leptostraca is
a more extensive structure than that of the Anostraca, since it includes
the mandibular and maxillary somites united with the protocephalon.
The maxillary region of the head is often expanded to form a large
cephalic shield, or bivalved shell, covering the anterior part of the
body. In Apus (fig. 50 B) the region of the protocephalon (Prtc)
forms a distinct anterior part of the definitive head bearing the
eyes dorsally and the antennae and labrum ventrally (C). Behind
the protocephalon the limits of the mandibular tergum (B, JJ) are
clearly marked, but the maxillary terga (III+I1V) are confluent as in

posterior view. I, Porcellio sp. (Isopoda), head, composed of protocephalon
and four following somites (maxillae and maxillipeds removed). J, Orches-
toidea californica Brandt (Amphipoda), head, same composition as in Porcellio,
approximate division between protocephalic ‘and gnathal regions indicated by
broken line (2s). K, Talorchestia longicornis Say (Amphipoda), right mandi-
ble, mesal view.

a, primary (dorsal) articulation of mandible; rAnt, first antenna (antennule) ;
2Ant, second antenna; Bnd, basendite; c, secondary (anterior) articulation of
mandible; Cp, carapace; E, compound eye; /, tergal promotor muscles of man-
dible ; JJ-V J, second to sixth somites ; //T, mandibular tergum; J, tergal remotor
of mandible; k, ligament uniting ventral adductors of mandibles; KL, ventral
adductor muscles of mandible; Lm, labrum; Md, mandible; rMx, 2M~x, first and
second maxillae; MarGld, maxillary gland; 1M xp, first maxilliped; Plp, palpus;
Prtc, protocephalon (acron-+ somite I); 2s, suture between protocephalon and
mandibular somite (B), or theoretical line of division between protocephalic and
gnathal regions of head (J); 3s, suture between mandibular and maxillary
somites; V.Stn, sternum of first maxilliped somite.

120 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Eubranchipus (A). In the Cladocera (D) the general head structure
and composition is the same as in Apus, except for the lateral com-
pression of the maxillary shield, which gives the latter its “bivalved”
form, but the intersegmental lines are lost, and the limits of the proto-
cephalon (Prtc) are marked only by the attachments of the antennal
muscles. The Leptostraca (F) have the cladoceran type of head and
bivalved maxillary shield, but are distinguished by the presence of a
large frontal lobe (s) projecting above the bases of the eye stalks.
Here again the protocephalic area of the composite head is marked
only by the origins of the muscles of the protocephalic appendages
(G), including those of the eye stalks and the two pairs of antennae.
The mandibles of the Leptostraca (Nebalia) retain the palpi (H, Pip),
but their basal structure and musculature is the same as those of
Eubranchipus (E) and other Entomostraca.

The Malacostraca, other than Amphipoda and Isopoda, are com-
monly said to have a “cephalothorax,’ which includes the gnathal
somites and a number of following somites up to a maximum of 12
in all. Most of this composite structure, however, which in its fullest
development is covered by the carapace (fig. 51 C, Cp), is more truly
a gnathothorax, since the true head is always a distinct though small
protocephalic lobe more or less concealed beneath the overhanging
rostrum (r) of the mandibular somite. When the protocephalon is
detached, as shown in the figure (C), it is seen to be a distinct cephalic
structure bearing the stalked eyes, both pairs of antennae, and the
labrum. The typical malacostracan head is thus identical. with the
protocephalic head of the Anostraca (fig. 50 A, Prtc). Even in the
Brachyura (fig. 51 D, E) the protocephalon is readily identified as
such, though dorsally (D) it is much reduced, and is concealed in a
pocket beneath the anterior margin of the carapace; ventrally (E)
it carries a large epistomal plate and a small labrum. In the Stoma-
topoda, on the other hand, the protocephalon is highly developed (fig.
51 F, G), and its integumental sclerotization is broken up into several
distinct plates (d, e, f, g), which, however, can in no sense be re-
garded as representing a “‘segmentation” of the head. The mandibles
of the more generalized type found in the Malacostraca (B) are
identical in their structure and musculature with the mandibles of
Entomostraca (fig. 50 E, H).

The Amphipoda and the Isopoda (including Apseudidae), with
regard to the structure of the head, do not appear to be properly
classed with the rest of the Malacostraca, since the head (fig. 50 I, J)
is an intimate combination of the gnathal somites (J7+J/J+I1V )
with the protocephalon (Prtc), and thus resembles in its composition

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS I2I

Fic. 51.—Cephalic structures of malacostracan Crustacea in which the defin-
itive head is the protocephalon, as in Anostraca (fig. 50 A).

A, Anaspides tasmaniae Thomson (Syncarida), protocephalon and appen-
dages, anterior view. B, same, mandibles and muscles, posterior view. C,
Spirontocaris polaris (Decapoda-Macrura), showing protocephalon (Prtc)
detached from carapace. D, Callinectes sapidus Rathbun (Decapoda-Brachy-
ura) protocephalon and appendages, dorsal view. E, same, protocephalon,
anterior view. F, Chloridella panamensis Bigelow (Stomatopoda), protocephalon
and appendages, ventral view. G, same, protocephalon detached from carapace
(Cp), dorsal view.

a, primary (dorsal) articulation of mandible; rAnt, first antenna (antennule) ;
2Ant, second antenna; Bud, basendite; c, secondary (anterior) articulation of
mandible; Cp, carapace; Cxrpd, coxopodite; d, anterior (ocular) division of
protocephalon; E, compound eye; e, ocular plate of protocephalon; Epst, epi-
stoma; Expd, exopodite; f, postocular dorsal plate of protocephalon; g, pos-
terior (antennular) division of protocephalon; J, J, dorsal promotor and re-
motor muscles of mandibles; J/-VJII, second to eighth somites; k, adductor
ligament of mandibles; KL, adductor muscles of mandibles; Lm, labrum; Prtc,
protocephalon; r, rostrum of mandibular somite; T/pd, telopodite (palpus).

122 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

the head of Nebalia and of such entomostracan forms as Apus,
Daphnia, and others, though in form it often has a striking resem-
blance to the head of a hexapod mandibulate. However, in both the
amphipods and the isopods the head usually includes also the first
maxilliped somite and its appendages (fig. 50 J, rMxp), and may in
addition bear the second maxillipeds. The mandible acquires a secon-
dary anterior articulation with the cranium (I, J, K, c), by which
its action is limited to a hinge movement on a horizontal axis between
its two articular points (K, a, c). The same mandibular mechanism
has been independently developed in the decapod Crustacea and in the
pterygote Hexapoda. While the head structure of the Amphipoda
and Isopoda sets these groups apart from other Malacostraca, it does
not necessarily relate them to any other group.

The final type of head developed in the Arthropoda is that char-
acteristic of the myriapods and hexapods, and must have evolved in
the common ancestors of these groups represented in the post-
crustacean, protomyriapodan section of the main arthropod stem
(fig. 54). The head of all the myriapod and hexapod groups is a
highly standardized structure, composed of the protocephalon and
the three gnathal somites, so closely united that little evidence of the
original segmentation remains, except in the presence of the appen-
dages (fig. 53 A), and even here the evidence is obscured by the loss
of the second antennae. In early ontogenetic stages, however, the
gnathal somites are entirely distinct from a large anterior cephalic
lobe that usually includes the second antennal somite, which may bear
vestiges of its former appendages. The Protomyriapoda must have
had compound eyes, since eyes of the compound type recur finally
in the Hexapoda; they likewise must have carried paragnathal lobes
of the head from the Crustacea to the Hexapoda, though these organs
have disappeared in the modern myriapodous forms. The maxillary
appendages probably were no more specialized in the protomyriapods
than in modern Chilopoda (fig. 53 A, C). The mandibles lost the
telopodites, but they developed a special feature of which no sug-
gestion is to be found in the Crustacea, namely, a mobile gnathal lobe,
the lacinia, movable by muscles arising in the mandibular base and
on the walls of the cranium. The mandibular lacinia is retained as
a movable lobe in modern Symphyla (fig. 52 E, Lc) and Diplopoda;
in the Chilopoda it is not separated from the stipital region of the
mandible (fig. 53 E, F), though it is provided with strong stipital
and cranial muscles (F, 13, 10); in the Pauropoda and Hexapoda
(fig. 52 F) apparently it has united with the stipes (St), producing
a solid jaw of the crustacean type, and its muscles have disappeared.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 123

Fic. 52.—Symphyla, Diplopoda, and Thysanura.

A, Scutigerella immaculata Newport (Symphyla). 3B, same, maxilla. C,
same, labium. D, labium of Machilis sp. (Thysanura). E, mandible of
Scutigerella. KF, mandible of Machilis. G, gnathochilarium of Fontaria vir-
giniana (Drury) (Diplopoda). H, thirteenth body segment of Scutigerella,
ventral view. I, seventh abdominal segment of Machilis, ventral view. J. thir-
teenth body segment of Scutigerella, lateral view. K, last leg of Scutigerella,
posterior view. L, terminal segments of Scutigerella, lateral view.

a’, a”, a’, cranial articulations of mandible, maxilla, and labium; Cd, cardo;
Cer, cercus; Cx, coxa; dac, dactyl (clawlike remnant of dactylopodite) ; ficc,
cranial flexor muscle of lacinia; Fm, femur; Ga, galea; H, head; Lc, lacinia;
Plp, palpus; Ptar, pretarsus; St, stipes; Stn, sternum; Sty, stylus; Tar, tarsus;
Tb, tibia; Tel, telson; 1Tr, first trochanter ; 2Tr, second trochanter (prefemur) ;
un, lateral claw (unguis) of pretarsus; Vs, eversible vesicle; 1-16, body segments.

124 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

The maxillary appendages in Symphyla and Hexapoda have acquired
two endite lobes of the stipes (lacinia and galea), but the palpi have
been lost in Symphyla (B).

The last important event in the evolution of arthropod head appen-
dages was the union of the bases of the second maxillae to form a
single median organ, the so-called labium. The labium took its origin
in the common ancestors of the Symphyla, Diplopoda, Pauropoda,
and Hexapoda, which constituted the third and most prolific branch
of the arthropod stock (fig. 54). The primitive structure of the
labium is best preserved in the more generalized hexapods (fig.
52 D); in the Symphyla (C), Pauropoda, and Diplopoda (G) it has
lost the telopodites, or palpi, and in the diplopods it forms at least a
part of the complex gnathochilarium (G).

Crampton’s (1928) phylogenetic conclusions drawn from com-
parative studies of the arthropod head differ radically in some respects
from the concept of arthropod relationships here deduced from the
same source. Crampton believes that the first arthropods derived
from annelid precursors were probably prototrilobites, and that from
the latter were evolved in one direction the Trilobita and Chelicerata,
in another the Protocrustacea, which last in turn produced the higher
Crustacea, while finally, from the malacostracan Crustacea were
evolved the Myriapoda and Hexapoda.

To the writer it would seem that if the Protarthropoda are con-
ceded to have been derived from wormlike ancestors, whether anne-
lidan or protonychophoran, by a sclerotization of the integument and
a jointing of the appendages, they must have taken on at once a
centipedelike form. According to the theory here proposed, therefore,
a long, unbroken line of slender polypodous arthropods has persisted
from the ancient protonychophoran progenitors to the modern chilo-
pods. Along this line have been carried the features common to all
the arthropods, while new characters evolved in the main line itself
have been distributed to subsequent lateral branches, where in some
cases they have persisted in their original state, in others they have
still further evolved, and in still others they have been lost.

The first lateral branch from the arthropod stem was that of the
Prototrilobita (fig. 54), which produced the Trilobita and the Chelic-
erata. In this branch cephalization united the first four somites with
the acron to form the trilobite “head,” and continued in the Chelicer-
ata until the “prosoma” contained six and eight somites. Meanwhile,
in the main protarthropod stem, cephalization produced a more simple
head (protocephalon) consisting of the acron and only the first somite,
but the appendages of the second somite were converted into a pair

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 125

of jaws. The protarthropods thus developed into Protomandibulata.
At this point arose the crustacean branch, in which the simple proto-
cephalon is still the definitive head in a large number of forms, though

“uy ef

Fic. 53.—Head and mouth parts of Chilopoda.

A, head of Scutigera forceps Raf. B, poison claws (first legs) and second
body segment of Lithobius sp., ventral view. C, first and second maxillae of
Lithobius, ventral view. D, head of Lithobius with maxillae removed, ventral
view. EE, right mandible and associated head structures of Lithobius, ventral
view. F, right mandible of Lithobius, dorsal view. G, hypopharynx, hypo-
pharyngeal suspensoria, and preoral mouth cavity of Lithobius, ventral view
(labrum and clypeus removed).

a’, primary posterior articulation of mandible; Ant, antenna; c, secondary
anterior articulation of mandible; Clp, clypeus; Cx, coxa; E, eye; For, foramen
magnum; h, ventral inflection of cranium; HAp, hypopharyngeal apodeme;
Hphy, hypopharynx (metastoma); HS, hypopharyngeal suspensorium (ful-
tura) ; L, leg; Lm, labrum; Md, mandible; mr, mandibular rod; 1Mx, 2M, first
and second maxillae; ProC, preoral mouth cavity; Scr, subcoxa; Sin, sternum ;
T, tergum; 2, 2, frontal and clypeal muscles of labrum; 3, 4, frontal and clypeal
muscles of hypopharyngeal suspensorium; 5, 6, 7, cranial muscles of same; 8,
ventral dilator muscles of pharynx; 9, cranial muscle of mandibular stipes; Jo,
cranial flexor of mandibular lacinia (origin lateral on cranium) ; 17, adductor
muscle of mandible; 12, protractor muscle of mandible; 13, stipital flexor of
mandibular lacinia.

in several groups a more extensive head has been evolved by adding
to the protocephalon the following three, four, or five somites. Cepha-
lization, however, continued also in the main protomandibulate line,
and produced here a composite head of standardized structure in

126 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

which the three gnathal somites were intimately combined with the
protocephalon, while the appendages of the protocephalic somite
(second antennae) were suppressed. The Protomandibulata now be-
came Protomyriapoda. A composite head has thus been produced
along three separate lines of arthropod evolution, but in each case
with characteristic differences.

The Protomyriapoda had all the characters common to the several
groups of arthropods finally derived from them, and also older
characters earlier transmitted to the Crustacea, which later appear in
one or more descendent groups, and are lost in others. From the
protomyriapods there arose the final persistent arthropod branch, ‘the
Protosymphyla, while the main stem continued into the relatively
generalized modern Chilopoda (fig. 54). The Protosymphyla de-
veloped a labium by the union of the bases of the second maxillary
appendages, and so characteristic is this feature of all their descen-
dents, including the modern Symphyla, Diplopoda, Pauropoda, and
Hexapoda, that this group as a whole might be distinguished as the
Arthropoda labiata. The chilopods have developed few special fea-
tures other than the conversion of the first legs into a pair of poison
claws, but they have lost certain features of the protomyriapods.

COELOMIC ORGANS OF ADULT ARTHROPODS

In no modern adult arthropod is there retained a complete series
of coelomic sacs, but remnants of the coelom are preserved as the
lumina of the gonads and genital ducts, of various glands having an
excretory or accessory genital function, and perhaps of other glandu-
lar structures. The embryonic development of the coelomic sacs of
the Onychophora very probably recapitulates fairly closely the phylo-
genetic history of the coelomic sacs in both the Onychophora and
the Arthropoda. The primitive coelom of these animals undoubtedly
consisted of a full series of paired segmental cavities, each opening
to the exterior through a ventral diverticulum of the coelomic wall
connected with the ectoderm mesad of the base of the corresponding
appendage. The cavities must have served for the accumulation of
excretory products, and for the retention of the developing germ
cells, and the outlets gave vent to both the excreta and the
gametes (fig. 34 A). The more primitive annelids do not have per-
manent coelomic openings, and it seems doubtful that the simple
coelomoducts of the Onychophora had a common origin with the
metanephridia of the higher Annelida, since the metanephridia are
cutgrowths of the posterior walls of the coelomic sacs and each opens
through the segment following.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 127

Early in the evolution of the common ancestors of the Onychophora
and Arthropoda, judging from the embryonic development of modern
Onychophora, the coelomic cavities were differentiated into dorsal
compartments (fig. 34 B, a) containing the proliferation centers of
the germ cells in their walls, and into ventral compartments (b) open-
ing through the coelomoducts (c, d). With the complete separation
of the two series of compartments (C), the dorsal compartments
became gonadial sacs (G) and the ventral compartments (b) became
nephridial sacs. The gonadial sacs, being deprived of outlets, united
with one another on each side and formed a pair of longitudinal
gonadial tubes (E, G), which retained exit passages through one
pair of coelomic sacs that maintained their integrity and served as
genital ducts. The ventral sacs and their respective coelomoducts
were transformed into specific segmental excretory organs, or ne-
phridia. It is thus clear that the genital ducts are not “modified
nephridia,” as they are often said to be, but that the genital ducts and
the nephridia are separate products of the primitive open coelomic
sacs, and hence, when once individually established, cannot be inter-
changeable in function. However, because of the variable position of
the genital ducts in the Arthropoda, it is evident that a different pair
of coelomic sacs has been retained in different groups to serve as
genital outlets.

Excretory organs of coelomic origin in the Arthropoda are repre-
sented by the coxal glands of Chelicerata and the nephridial head
glands of Crustacea, and perhaps also by certain head glands of °
Diplopoda, Chilopoda, and apterygote Hexapoda. The coxal glands
of the Chelicerata, with one exception, consist of a single pair of
excretory organs situated in the prosoma and opening at the bases
of the appendages. Each gland in its fullest development is a com-
posite structure composed of several lobes or saccules derived from
coelomic sacs and united upon a common tubular base, the so-called
“stolon,” or “labyrinth,” composed of glandular cells and tubules, and
is connected with the exterior by one or two segmental ducts. The
organ is, therefore, variable in features that might be supposed to
vary in a composite structure of such a nature, as in the number of
coelomic sacs involved, the number of segmental openings, and the
position of the openings. The excretory head glands of Crustacea
include a pair of antennal glands (“green glands”) and a pair of
maxillary glands (‘‘shell glands’). The first are present in the adult
stage only in the Malacostraca ; the second occur in the Entomostraca
and in some Malacostraca ; both pairs are present in Nebalia (Manton,

128 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

1934). The maxillary glands are usually simple tubes or sacs, but
the antennal glands may take on a highly complex structure.

The coxal glands of Limulus are a pair of large brick-red organs
lying in the sides of the prosoma. Each organ consists of four suc-
cessive glandular lobes arising from a common longitudinal stolon
composed of numerous connecting tubules, and of a long coiled duct
that proceeds from an end-sac in the fourth lobe and opens behind
the base of the fifth appendage (third leg). According to Patten and
Hazen (1900) the nephridial lobes are developed from masses of
mesodermal cells derived apparently from the somatic walls of the
coelomic sacs of the second, third, fourth, and fifth somites. Similar
masses of cells in the first and sixth somites degenerate and disappear.
The duct arises as a tubular diverticulum of the fifth coelomic sac,
which latter becomes the fourth nephridial lobe. A short terminal
part of the definitive duct is formed as an ectodermal invagination at
the external orifice of the mesodermal duct.

The coxal glands of Arachnida are best known from the work of
Buxton (1913, 1917, see also Petrunkewitch, 1933, and Chickering,
1937). A relatively primitive condition is found in the araneid groups
Liphistiomorphae and Mygalomorphae, in which each gland has two
saccules, one in the third, the other in the fifth segment, both con-
nected with a long convoluted tubular labyrinth, from which two
outlet ducts proceed to the exterior, one opening behind the third
appendage, the other behind the fifth. Such an organ would appear
. to be a composite structure formed by the union of three consecutive
segmental glands. In certain genera of the Amblypygi group of the
Pedipalpida the gland of the fifth segment is shown by Buxton (1917)
to be an independent organ opening separately on the fifth segment.
In the Uropygi each gland has two saccules but only a single opening,
which is on the third segment. All other Arachnida have but a single
saccule for each lateral gland and a single outlet, but the opening is
at the base of the second appendage (pedipalp) in Solpugida and
Palpigradida, at the base of the third appendage (first leg) in Ara-
neida, excepting the two groups above mentioned, and at the base of
the fifth appendage (third leg) in Scorpionida and Phalangida, as
in Limulus. Buxton calls attention to the correspondence of the
coxal glands of Solpugida and Palpigradida with the salivary glands
of Onychophora, the organs in each case having their opening on
the second postoral body somite.

Studies on the development of the arachnid coxal gland appear to
leave no doubt that the organs are derivatives of coelomic sacs with
coelomoducts formed as direct diverticula from the sacs as are the

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 129

coelomic ducts of Onychophora. Brauer (1895) has shown that in
the embryonic development of the scorpion there are formed five pairs
of diverticula from the coelomic sacs of somites J//J, JV, V, VI, and
VIII, respectively, of which those of the fifth and eighth somites
acquire openings to the exterior. The coelomic sacs and their diver-
ticula in the fifth somite develop into the definitive coxal glands, the
coelomic diverticula of the eighth somite become the genital ducts,
and the sacs and diverticula of the other segments disappear. Accord-
ing to Kishinouye (1894) the development of the coxal glands in
the araneid genera Lycosa and Agelena shows that each organ is a
composite structure formed of small parts of the coelomic sacs of
somites J/J, IV, and V, but only the first acquires an opening to
the exterior.

The nephridial glands of the Crustacea, being individual organs,
resemble the simple nephridia of the Onychophora rather than the
composite coxal glands of the Chelicerata. Each organ consists of a
mesodermal end-sac, a mesodermal canal, which may become highly
complex in form, and a short exit duct of ectodermal origin (see
Cannon and Manton, 1927, and Manton, 1930). The embryogeny of
the crustacean excretory glands, however, is in some cases compli-
cated by an indirect course of development.

The antennal gland of Hemimysis lamornae is said by Manton
(1928) to be formed from two masses of cells derived from the
antennal mesoderm, one of which produces the end-sac, the other the
canal. The canal becomes attached distally to the ectoderm, and a
small ingrowth from the latter forms a short ectodermal exit duct.
Where the canal touches the wall of the sac, a compact group of 7
or 8 cells bulges into the lumen of the canal, and at this point the sac
and the canal become united, but the only visible communication
between them, Manton says, is by fine rather vague channels passing
through the nephrostome cells. According to Vogt (1935) the an-
tennal mesoderm of Mysis relicta produces only the canal and a sheet
of connective tissue membrane in the base of the antenna, to which
the canal becomes attached. A group of 8 cells in this membrane
then forms the nephrostome. The true end-sac, Vogt claims, is con-
structed from adventitious connective tissue cells that wander into
the antenna from the thoracic segments and form the end-sac beneath
the nephrostome membrane. Vogt contends that the development of
the antennal gland of Mysis relicta so closely resembles the develop-
ment of an annelid nephridium that the two organs must be homolo-
gous structures, the nephrostome membrane of Mysis representing
a dissepiment in the annelid. To the writer a parallelism in the two

9

130 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

cases is far from evident, and the development of the mysid antennal
gland seems better explained as a secondary modification of the
developmental processes that give rise to the coxal glands of Arachnida
and the nephridia of Onychophora.

Most of the tracheate Mandibulata have a series of head glands
pertaining to the gnathal somites, the openings of which lie mesad of
the appendage bases, or are displaced anteriorly or posteriorly when
the bases of the two appendages of a pair are united. Some of these
glands have been shown to have an apparent excretory function,
because of their property of eliminating from the blood particles of
carmine injected into the body of the animal, and such glands also
have a complex structure, usually described as consisting of a saccule,
a labyrinth, and a duct. Hence, various writers have claimed that
glands of this type represent nephridial organs corresponding with
the excretory head glands of Crustacea, though little evidence as to
their embryonic origin has been produced.

The gnathochilarial glands of the Diplopoda have been shown by
Bruntz (1903) to collect injected carmine from the blood, and they
are said by Heathcote (1886) to be derived from the mesoderm in
embryonic development. Likewise, according to Bruntz (1908) and
Philiptschenko (1928), a pair of labial glands of apterygote insects
have an excretory function and a nephridialike structure. These
glands open either separately (Campodea, Japyx) between the hypo-
pharynx and the labium, or (Machilis, Lepisma) their ducts unite in
a common median duct, and are joined by the ducts of a pair of
“posterior salivary glands.” The labial glands of the apterygote in-
sects, particularly those of Thysanura, would so evidently seem to be
the same as the labial glands of pterygote insects, which are commonly
found to be ectodermal organs, that it is difficult to believe they are
not homologous structures, regardless of their function. In the
Chilopoda, according to Fahlander (1938), there are present generally
three pairs of head glands associated with the bases of the gnathal
appendages, but in addition there is another pair having a complex
structure suggesting an excretory function. These glands have each
two openings, one mesad of the base of the first maxilla, the other
behind the base of the second maxilla. Fahlander contends, there-
fore, that each gland has been formed by the union of two nephridial
organs pertaining to the maxillary somites. The morphological status
of all such glands must yet be determined by a study of the embryonic
development.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA

SNODGRASS I31

THE GENITAL DUCTS

The student of arthropod phylogeny is confronted at every turn
with the vexing problem that arises from the different position of
the genital outlets in the various arthropod groups, and in recent
years much discussion has been given to the question as to how the
heterogoneate condition came about (see Tillyard, 1930, 1932, 1935,
Snodgrass, 1933, 1936, Reynolds, 1935, Imms, 1936). Two phases
of the problem have been somewhat confused, namely, that pertaining
to the position of the openings of primary lateral ducts, and that
pertaining to the position of secondary median ducts. The opening
of a median duct is subject to migration, usually in a posterior
direction; the openings of lateral ducts are closely associated with
particular segments, since the lateral ducts themselves represent
specific pairs of segmental coelomic sacs.

The possible migration of lateral genital ducts is narrowly restricted
because of the limitations imposed by the transverse segmental nerve
trunks ; a secondary median duct formed by invagination of the ventral
wall of the body, however, lies beneath the ventral nerve cords, and
may, therefore, become lengthened from one segment to another by
an extension of its connection with the body wall. There is no evi-
dence to support Tillyard’s (1930) contention that segmental gonads
were once connected by a common duct, which has retained a single
definitive opening on different segments in different arthropods, be-
cause when the germaria were segmentally arranged they were con-
tained in the dorsal parts of segmental coelomic sacs with individual
openings to the exterior, and the serial union of the dorsal parts of
the coelomic sacs has produced the definitive tubular gonads opening
through a single pair of coelomic sacs, while the ventral parts of the
other sacs discharging through the coelomoducts became nephridial
sacs. Likewise, Tillyard’s (1935) second proposal that a heterogo-
neate condition has arisen by a variation in the number of somites
formed before or behind the primary genital somite cannot be accepted
for the reason that somite formation in the genital region is primitively
teloblastic.

Inasmuch as the primary lateral genital ducts represent specific
coelomic sacs that have been retained to serve as genital outlets, a
segmental difference in the position of the genital openings is to be
explained only as the result of mutations that have been effective in
the organizer of the zone of teloblastic growth, which determines
what particular pair of coelomic sacs shall be utilized as genital exits.
A branching of the embryonic lateral ducts has been observed by

132 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Heymons and by Wheeler in Dermaptera, and, according to Heymons
(1901), the definitive ducts of Scolopendra are formed from two
united pairs of coelomic sacs. In such cases we have, perhaps,
examples of the supplanting of one pair of exit sacs by another pair.

The heterogoneate condition of modern arthropods, therefore, must
be the result of mutations that occurred among ancestral forms. The
faculty of mutation affecting the position of the genital ducts was
carried over into the entomostracan branch of the Crustacea, and was
not entirely extinct in the early forms of the Hexapoda. Moreover,
in the Chilopoda, as in the Onychophora, there still exists a variability
as to the segment of the genital ducts, for, though the genital outlet
is always on the subterminal segment in Chilopoda and on the ante-
penultimate segment in Onychophora, the genital segment is not
morphologically the same somite in all cases, since the number of
somites preceding it may be quite different in different genera. In
the Geophilomorpha, furthermore, the number of pregenital somites
is said to vary even among individuals of the same species.

VII. PHYLOGENETIC CONCLUSIONS

1.—A planulalike creature with an open posterior blastopore was
probably the ancestor of the Metazoa. A creeping form adapted to
feeding on a subsurface by the forward elongation of the blastopore
on the under side of the body might readily have evolved into a
worm by the partial closure of the blastopore, producing thus an
alimentary canal with a ventral subapical mouth and a terminal anus.
The subapical position of the mouth differentiated the animal into an
acronal sensory region, or prostomium, and a postoral visceral and
motor region, the body, or soma in a restricted sense.

2—The unsegmented progenitors of the annelids were probably
small, creeping, wormlike creatures having a simple alimentary canal,
a mouth on the anterior part of the under surface of the body, and a
terminal anus. Locomotion on solid surfaces was effected by a ventral
clothing of cilia, and body movements were produced by a system of
muscle fibers on the inner surface of the body wall, derived from
the ectoderm. The body cavity was a blastocoelic haemocoele, and
was largely occupied by lateral bands of a mesoblastic parenchyma
proliferated in the gastrula stage from endodermal or ectodermal
teloblastomeres. The nervous system consisted of longitudinal and
circular nerve tracts centering in ganglionic cell groups of the pro-
stomium, which latter eventually united to form a “brain.” Sensory
organs may have included tactile tentacles and photoreceptive “eye
spots” located on the prostomium.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 133

3.—The annelidan progenitors acquired a more effective body
movement by the attachment of the longitudinal somatic muscle fibers
at several successive rings on the body wall, and by the accompanying
formation of transverse muscular septa at the resulting integumental
grooves. The body region of the wormlike animal in this way became
differentiated into a small number of consecutive motor units, the
primary somites. To regulate the new muscular mechanism of
metameric movement, there was developed from the body surface
of contact with the substratum a new somatic nervous system in the
form of ventral nerve cords with ganglia corresponding with the
myotomes. The primary and secondary nervous systems were then
unified by a connection of the ventral nerve cords with the brain,
and the somatic elements of the primary system disappeared. The
ingrowth of the septal muscles cut the parenchymatous mesoblast
bands into segmental blocks, and the latter became excavated by
cleavage spaces (primitive coelomic cavities) for the accumulation
of body fluid containing waste products. Excretory organs, if present
at this stage, were simple nephridial tubules extending from the ecto-
derm into the haemocoele, where they were associated with the meso-
blast cavities. The inner parenchymal cells lining the cavities formed
epithelial coelomic sacs, but the outer cells, being still an undifferen-
tiated tissue, were converted into muscle fibers and connective tissue.
The secondary muscles thus formed reinforced the primary somatic
muscles already present, and eventually became the major part of
the muscular system. The germ cells remained in a mass of undiffer-
entiated tissue near the posterior end of the body, and the gametes
were liberated probably through a pore or temporary rupture of the
body wall. The primitive segmented worms evolved in this manner
from unsegmented progenitors were the ancestors of the annelids.

4.—To increase the reproductive function, the subterminal genital
region of the primitive oligomerous annelids was enlarged by the
successive generation of new somites from its undifferentiated tissue.
A series of secondary teloblastic somites duplicating the structure
of the primary somites was thus interpolated between the primitive
body of the worm and a small postgenital terminal cone containing
the anus. The multiplying germ cells spread into the haemocoele of
the new somites, and groups of them became lodged in the walls of
the coelomic sacs. The ripening germ cells were now discharged
into the coelomic cavities, which latter thus became gonadial as well
as nephric in function. Since the coelomic sacs as yet probably had
no permanent openings, the gametes must have been liberated through
temporary pores of the body wall, through secondary genital openings

134 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

into the nephridia, or by the autotomous separation of the genital
somites. At this stage the generalized annelids had acquired the
fundamental characters common to the higher Annelida, the Ony-
chophora, and the Arthropoda. ns

5.—The increase in the size of the body by the addition of the telo-
blastic genital somites created a demand for a still greater efficiency
of locomotion, and, according to the nature of the response to this
demand, two divergent groups of worms were evolved from the
generalized annelids. The members of one group acquired segmental
clusters of eversible and retractile chaetae serving as adjuncts to the
somatic muscular system by maintaining a hold on surfaces of con-
tact; the members of the other group developed segmental pairs of
lobelike outgrowths of the body wall containing extensions of the
somatic muscles, which served as primitive legs. The chaetae-bearing
forms gave rise to the Chaetopoda; the lobopod forms were the
ancestors of the walking Onychophora and Arthropoda.

6.—From the primitive chaetopods were evolved the several groups
of modern annelids. By the extension of open tubes from the pos-
terior walls of the coelomic cavities to the exterior, a more efficient
type of excretory organ (metanephridium) was developed, which
could serve also for the liberation of the gametes. The Polychaeta
are distinguished particularly by the elaboration of external structures
of various kinds, while the Oligochaeta and Hirudinea have achieved
a higher development of internal organs and functions. The loco-
motor powers of the Polychaeta were increased by the development of
lateral lobes of the body wall supporting the segmental groups of
chaetae, and in most forms each lateral pair of chaetigerous lobes
eventually combined to produce a single locomotor organ, the para-
podium. The parapodia served for progression on solid surfaces,
and became also effective adjuncts to the undulatory motion of the
body during swimming.

7.—The lobopod annelids became further differentiated from their
chaetopod relatives by a chitinization of the entire cuticula, and by
the suppression of all the cephalic tentacles except one pair probably
corresponding with the palpi of the Polychaeta. They also acquired
outlets from the coelomic sacs to the exterior, but the exit ducts were
formed as diverticula from the ventral walls of the sacs and opened
each on the segment of its sac mesad of the base of the corresponding
leg. The germ cells were located in the walls of the dorsal parts of
the coelomic sacs, and the primitive coelomoducts discharged both
excretory matter and the gametes. The coelomic sacs, however, soon
became divided into dorsal gonadial compartments and ventral nephric

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 135

compartments. The gonadial sacs of each lateral series united with
each other, forming thus a pair of tubular gonads, which opened to
the exterior through one pair of undivided coelomic sacs and their
outlet ducts. The ventral nephric sacs now became exclusively excre-
tory reservoirs, and, with the coelomoducts, formed a series of
nephridial organs along each side of the body. As a result of the
conversion of the original coelomic sacs into gonadial sacs and
nephridial sacs, the haemocoele was restored as the definitive body
cavity.

At this stage of their evolution, the lobopod annelids assumed the
status of Protonychophora. Some of the protonychophorons retained
the flexible integument of the worms; others developed a sclerotiza-
tion in the cuticula, and thus acquired an external skeleton of cuticular
plates. The soft-skinned forms, preserving some of the general
aspects of their annelidan ancestors, evolved into the modern Ony-
chophora; the armored forms gave rise to the Protarthropoda. Since
the members of both groups were well adapted by their leglike appen-
dages to a walking mode of progression, many of their descendants
found an advantageous habitat on land.

8.—The Onychophora retained the cylindrical wormlike form, but
they lost the segmented structure in the integument and musculature.
The lobiform appendages became more efficient locomotor organs
through the development of an incipient segmentation, and the acqui-
sition of terminal claws, but the first postoral appendages were con-
verted into a pair of jaws. The single pair of prostomial tentacles
took an apical position by migrating forward on the dorsal surface
of the head, but their nerve tracts were united by a commissure in the
posterior part of the brain. The eyes retained the annelid type of
structure. The somatic nerve cords, which presumably must have
been ganglionated in the segmented generalized annelids, became
simplified by a redistribution of the neurocytes, and took widely
separated positions along the sides of the body. The ganglia of the
jaw somite, however, united with the cerebral ganglion of the pro-
stomium and became posterior lobes of the brain. The coelomic sacs
of the penultimate somite, regardless of the total number of somites
in the body, were retained intact to serve as genital outlets; the
persisting remnants of most of the other coelomic sacs became small
end-vesicles of the coelomoducts, which formed nephridial excretory
organs.

9.—The Protarthropoda, because of the hardening of the integu-
mental cuticula, lost the flexibility and contractility of their annelidan ©
ancestors and onychophoran relatives, and, to compensate, developed

136 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

a mechanism of telescopic movement between successive body seg-
ments by the simple device of retaining nonsclerotized areas in the
posterior parts of the primary segments, thus establishing a secondary
segmentation in which the longitudinal muscles became interseg-
mental instead of intrasegmental in action. The sclerotized appen-
dages necessarily became segmented into individually movable parts,
and their movements became more specifically controlled by body
muscles inserted on their bases. The protarthropods retained the
annelid structure of the nervous system, and the independence of
the first postoral ganglia of the ventral nerve cords. The prostomial
appendages (antennules) assumed an anterior position by a forward
migration below the eyes, with the result that in the arthropod brain
the antennal lobes lie beneath the optic lobes, and the brain takes a
vertical position by contrast with the horizontal position of the ony-
chophoran brain. Lateral eyes of the compound type were first
developed in the Protarthropoda. Because of the origin of the Protar-
thropoda from Protonychophora, the protarthropods were equipped
with a series of nephridial organs like those of the Onychophora, and
their internal reproductive organs were of the onychophoran type.
The segmental relations of the genital ducts, however, were subject
to mutation, and the position of the gonopores was, therefore, dif-
ferent in different forms, as shown by the highly variable position
of the genital outlets in modern arthropods.

The Protarthropoda, having an annelid ancestry, and being directly
derived from wormlike protonychophorons by a sclerotization of the
integument and a jointing of the appendages, could scarcely take on
other than a centipedelike form and structure, though they did not,
of course, have the composite head and other specialized features of
present-day myriapods. The number of body segments was variable,
and potentially large, since the production of new somites in the
zone of growth was not limited. The cephalic appendages (anten-
nules) were filamentous, the lateral eyes primitively compound. The
body appendages were probably all ambulatory legs with little differ-
entiation among them, each composed of seven segments. The
dactylopodites were provided with extensor and flexor muscles aris-
ing in the propodites. Aquatic forms probably had branchial epipo-
dites on the coxopodites. Perhaps the majority of the protarthropods
lived in shallow water near the ocean shore, where they inhabited the
bottom or aquatic plants, but probably also they occurred abundantly
in débris along the beach, and very likely some the them were to be
found in damp places on the land. The genital openings being on
specific body segments, propagation took place by sex mating, though

No. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 137

fertilization of the eggs was probably external. Postembryonic de-
velopment was anamorphic. The first major diversification of the
Protarthropoda gave rise to the ancestors of the Trilobita and the
ancestors of the Mandibulata (fig. 54).

1o.—The Trilobita preserved the uniform, generalized structure
and segmentation of the protarthropod appendages, but otherwise they
became highly specialized by a lateral extension of the margins of
the body segments, taking on thus a broad, flattened form except for
a median elevation giving passage to the alimentary canal. Further-
more, the first four postoral segments became intimately united with
one another and with the prostomial acron to form a solid anterior
body section, or prosoma, the so-called “head,” bearing the labrum,
the eyes, the antennules, and four pairs of postoral ambulatory
appendages. Basal endites of the anterior appendages may have
served as feeding adjuncts, but the trilobites, so far as known, devel-
oped no specific jaws. The Trilobita were entirely marine animals,
but they lived at the bottom of the water, and their legs show few
deviations from the ambulatory type of structure, except for the high
development of branchial lobes from the lateral surfaces of the
coxopodites. The extended tergal margins covering the gills probably
formed respiratory chambers. The position of the genital openings
in the trilobites has not been discovered, but, because of the close
relation between the Trilobita and the Chelicerata, the genital aper-
tures may be expected to be found on the fourth postcephalic segment.
The Trilobita became extinct by the end of the Paleozoic period of
geological history, but from a branch of the primitive pre-Cambrian
prototrilobites were evolved the Chelicerata.

r1.—The Chelicerata are distinguished from the Trilobita by the
union of several additional somites with the head to form a more
extensive prosoma, by the loss of the acronal appendages (anten-
nules), by a greater differentiation among the somatic appendages,
and by the forcipate structure of the reduced first appendages. Very
commonly, also, there is an extra podomere in at least some of the
legs, the patella, interpolated between the femur and the tibia. In
modern forms the nephridial organs are suppressed in most of the
somites, but some of them are retained as coxal glands, and (except
in Pycnogonida and some Acarinida) the genital openings occur
always on the eighth postoral somite. The Chelicerata have become
the most sepecialized of all the arthropods, there being little in their
body form and general organization suggestive of the ancestral centi-
pede type of structure, which is so evident throughout the mandibulate

138 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

branch. The Chelicerata include the Xiphosurida, the Eurypterida,
the Arachnida, the Acarinida, and very probably the Pycnogonida.

12.—The Xiphosurida are undoubtedly the closest living repre-
sentatives of the Trilobita. The xiphosurid prosoma has the same
structure as the trilobite head, and the same composition except for
the addition of three extra somites and a part of the eighth somite.
Likewise, the opisthosoma corresponds with the trilobite pygidium
extended forward to include all the somites behind the prosoma, so
that in the Xiphosurida there is no intermediate “thoracic” region
of free segments. Such fossil forms as Belinurus and Prestwichia
would appear to be intermediate between modern Xiphosurida and
Trilobita, and the Middle Cambrian Naraoia (see Walcott, 1931,
. fig. 1) must be related to the xiphosurid line somewhere close to the
trilobites. The first six prosomatic appendages retain the leg type
of structure, except for the reduction and chelicerate form of the
first pair. The seventh appendages are reduced to a pair of small
lobes, the chilaria, and the following six have the form of broad
plates formed chiefly by epipodite lobes, those of the last five bearing
lamellate gills. The genital openings in both sexes are on a median
ventral fold of the eighth segment united with the bases of the
opercular appendages of this segment.

13.—The Pycnogonida, judging from some of their structural
features, such as the union of the anterior body segments, the pos-
terior position of the dorsal eyes between the bases of the third pair
of appendages, the presence of a patellar segment in the legs, and
the chelicerate structure of the first appendages, are to be classed
with the Chelicerata; but because of their many unique characters,
including the occurrence of multiple genital openings, it is impossible
to connect them closely with any other of the chelicerate groups. It
may be noted, however, that species with eight pairs of legs have
presumably the same number of somites in the prosoma as have the
Xiphosurida.

14.—The Eurypterida and the Arachnida differ from the xipho-
surids in having only six segments in the prosoma, and this character
together with various other features of their organization shows that
these two groups are more closely related to each other than is either
group to the Xiphosurida. On the other hand, the Eurypterida have
certain characters of the xiphosurids that leave little doubt of their
common ancestry with the latter, and their descent from trilobite
stock. The general resemblance of the eurypterids to scorpions sug-
gests a relationship between the two, but the theory of Versluys and
Demoll (1920, 1923) that the Eurypterida and Xiphosurida are

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 139

derived from primitive aquatic scorpions cannot be maintained against
the evidence of close relationship between the Xiphosurida and the
Trilobita. The Arachnida, as invaders of the land, had to evolve
organs for aerial respiration, and the lamellate gills of their aquatic
progenitors borne on the abdominal appendages were structures
readily convertible into “lung books” by invagination into pockets of
the integument (see Lankester, 1885). In addition, however, tracheal
ingrowths of the body wall were developed in the Arachnida, as they
have been in nearly all the other terrestrial arthropods.

15.—The Protomandibulata preserved the slender, polypodous,
centipedelike form of the primitive protarthropods, but they acquired
as a distinctive character a pair of jawlike feeding organs, the
mandibles, developed from the bases of the second postoral appen-
dages. Probably long before the evolution of the mandibles, the first
somite had been united with the prostomial acron to form a primitive
composite head, or protocephalon, bearing the acronal sensory organs,
the mouth, and the first pair of postoral appendages, which last
became a second pair of antennae. The two pairs of appendages
following the mandibles were reduced and modified to serve as acces-
sory feeding organs. The other appendages were probably all leglike
in form, as in modern centipedes, and were 7-segmented, since a
patella does not occur in the mandibulate branch of the arthropods.
The circulatory system still retained the basic structure of that of
the generalized annelids; respiration probably was branchial, the gills
being carried on epipodite lobes of the coxopodites, as in the Trilobita ;
the nephridial organs were perhaps suppressed in most of the body
segments, but those that remained were of the onychophoran type of
structure. The reproductive organs were closed gonadial tubes open-
ing in each sex through a single pair of ducts formed from a pair
of coelomic sacs, but the segmental position of the genital openings
varied in different forms according to what particular pair of coelomic
sacs served as gonadial outlets.

The primitive Protomandibulata probably inhabited both the water
and the land, since from them were early evolved the aquatic Crus-
tacea, while the main branch developed into the terrestrial Proto-
myriapoda, from which have descended the modern myriapods and
the Hexapoda.

16.—That the Crustacea are derived from crawling, centipedelike
protomandibulate ancestors is attested by the retention in all the higher
forms of ambulatory appendages having the same structure as the
limbs of terrestrial arthropods. Many forms, however, have become
adapted in part or entirely to swimming by a modification of the

I40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

appendages, and the special development of an exite lobe of the basi-
podites has given rise to a characteristic biramous structure of the
limbs. The primitive protocephalon is retained as the definitive head
in the Anostraca and in most of the Malacostraca, but in the majority
of the Entomostraca and in the Leptostraca, Amphipoda, and Isopoda
from three to five gnathal somites have been united with the proto-
cephalon to form a more extensive cephalic structure. A carapace is
variously developed in many groups, either from the cephalognathal
region, or from the gnathothoracic region, but there is no true cephalo-
thorax formed by an intimate union of cephalic and thoracic somites
as in the Chelicerata. The mandibles have no movable lobes such as
those of the myriapods ; in most forms the jaws preserve the primitive
monocondylic articulation with the head, but in the higher Mala-
costraca they are secondarily dicondylic. The genital openings are
variable in position in the Entomostraca, but are fixed with respect
to a specific segment in the Malacostraca. The hatching of the young
at an early embryonic stage has resulted in the development of
‘ specialized swimming larval forms representing more primitive an-
cestral stages in their general structure than the immediate protar-
thropod ancestors of the crustaceans. The great antiquity of the
Crustacea is shown by the occurrence of highly evolved forms in the
Cambrian period contemporaneous with the oldest known trilobites.
17.—The Protomyriapoda, being the direct descendants of the
protarthropods, perpetuated the generalized arthropod form after the
trilobites, the chelicerates, and the crustaceans had branched off as
side issues and taken on variously specialized forms. During their
evolution the protomyriapods acquired the structures characteristic
of their descendants, which include the modern Symphyla, Diplopoda,
Hexapoda, and Chilopoda. The three gnathal somites became inti-
mately united with one another and with the protocephalon, forming
the standardized head of the above-mentioned groups, composed of
the acron and four postoral segments. The compound eyes and the
first antennae of the Protomandibulata were retained, but the second
antennae became reduced and eventually were lost, though their
ganglia were preserved as tritocerebral lobes of the brain. The
mandibles lost the telopodites, but each had a strong gnathal lobe
(lacinia) movable by a muscle arising within the coxopodite and by
another arising on the cranial wall. The two postmandibular maxillary
appendages were modified by a reduction of the telopodites and by
other adaptations to serve as accessory feeding organs. Since the
Symphyla and some of the more generalized Hexapoda have lateral
hypopharyngeal lobes (superlinguae) resembling the paragnatha of

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS I4I

Crustacea, it is possible that these structures were transmitted from
the Crustacea to the symphylids and hexapods through the Proto-
myriapoda, though they have been lost in modern Diplopoda and
Chilopoda. The legs of the protomyriapods were all alike and retained
the generalized 7-segmented structure, but the extensor muscle of
the pretarsus was lost, leaving only the flexor muscle, which, for
more effective action, shifted its origin from the tarsus into more
proximal segments of the leg. This last feature is a distinctive char-
acter of all the descendants of the Protomyriapoda. Nephridial excre-
tory organs were supplemented or replaced functionally by Malpighian
tubules of the proctodaeum. The position of the genital openings
was probably in general posterior, but variable. Postembryonic de-
velopment was anamorphic, the young being hatched with a small
number of segments, and the full number acquired by teloblastic
generation in the subterminal zone of growth.

The Protomyriapoda undoubtedly were terrestrial, and the larger
forms may have developed tracheal invaginations on various parts of
the body for respiration, but there was no definitely established
tracheal system transmitted alike to all the descendent groups of
terrestrial mandibulates. The probable characters of the Proto-
myriapoda are summarized as follows by Imms (1936) :

(1) The head bore a single pair of antennae and two pairs of jaws, viz.
mandibles and maxillae: the second maxillae were probably a subsequent acqui-
sition. (2) The trunk was composed of a variable and indefinite number of
sub-equal segments, each bearing a pair of legs. It is probable that anamorphosis
was universal and was continued throughout the life of the animal. (3) The
gonads opened to the exterior by paired apertures, and the segmental disposition
of the orifices probably varied in different families and depended upon that of
the coelomoducts involved..... (4) The alimentary canal was probably a
simple straight tube, while the excretory organs were little more than procto-
daeal outgrowths or pockets; an accessory excretory function was probably
performed by the fat-body. (5) Respiration was probably cutaneous in many
forms and partially tracheate in others. The tracheae were presumably in the
form of groups of unbranched tubuli devoid of taenidia and bearing a general
resemblance to those of Diplopoda. Each group of tracheae opened laterally
by means of simple cryptlike, segmentally arranged spiracles: in some forms a
pair of spiracles was probably located also on the head.

From the Protomyriapoda there emerged a specialized lateral
branch, the Protosymphyla, from which have been evolved in one
direction the progoneate modern Symphyla, Pauropoda, and Dip-
lopoda, in another the opisthogoneate Hexapoda, while the general-
ized myriapodan stock has more directly continued into the modern
Chilopoda.

142 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

18.—Since modern Symphyla combine features of the progoneate
Diplopoda and Pauropoda on the one hand, and of the opisthogoneate
Hexapoda on the other, there can be little question that they are direct
descendants of common ancestors of these two groups. Modern
Symphyla, however, are linked more closely with the progoneate
forms by the anterior position of the gonopore, the segmentation and
structure of the legs, and the retention of the movable laciniae of the
mandibles. The Protosymphyla, therefore, gave rise to an opistho-
goneate branch that became the Protohexapoda.

In general appearance the Protosymphyla probably resembled their
modern representatives, but retained certain features of the Proto-
myriapoda that have been transmitted to the hexapod line, though lost
in the progoneate descendants. The legs were all alike and had the
7-segmented protomyriapod type of structure, but the coxopodites
bore each, mesad of the telopodite base, a small stylus and an eversible
vesicle, as in modern Symphyla (fig. 52 H), which structures are pre-
served also on the abdomen of some of the apterygote insects (I). The
appendages of the last body somite became reduced to styliform cerci.
The head appendages included a pair of antennae, a pair of mandibles,
and two pairs of maxillae. The lateral eyes must have been compound,
because compound eyes have been transmitted along the arthropod
line from the Trilobita to the Xiphosurida, the Crustacea, and through
the Protosymphyla to the Hexapoda. The protosymphylan mandibles
had the protomyriapodan structure, movable laciniae being well de-—
veloped, and palpi absent. The first and second maxillae retained
the palpi and each acquired two basal lobes (lacinia and galea), fea-
tures transmitted to the hexapods, though the palpi have been lost in
the progoneate branch. The bases of the second maxillae, however,
became united to form a single appendage, the labium, an organ so
characteristic of all the descendants of the Protosymphyla that the
group as a whole, including Symphyla, Pauropoda, Diplopoda, and
Hexapoda, might well be designated the “Labiata” (fig. 54).

19.—The direct descendants of the progoneate branch of the proto-
symphylids are the modern Symphyla, but at an early period there
were evolved from the symphylid line the common ancestors of the
Diplopoda and Pauropoda. The Symphyla retain the generalized
structure of the body and appendages (fig. 52 A), but of the 16 body
segments evident in the dorsum of most forms, 3 are without appen-
dages. The legs (K) show the diplopod type of structure in the rela-
tively large size of the second trochanter (277) and the smallness of
the femur (/7), but the coxae do not appear as typical leg segments,
since each pair apparently is confluent in a large posterior division of

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 143

the venter of the body segment (H, Cx), carrying mesad of the base
of each telopodite a small stylus (Sty) and an eversible vesicle (Vs).
The end segments of the legs are reduced to small dactyls (K, dac),
but each has an accessory claw (umn) arising from its base. The first
legs are usually reduced in size and lack tibiae. The last body segment
bears a pair of cerci (L, Cer), which presumably are homologues of
the legs or possibly of the styli of the preceding segments. Com-
pound eyes are absent. The mandibles preserve the movable laciniae
(E, Lc); the maxillae have both laciniae and galeae (B), but the
palpi are small or vestigial; the labium (C) is a simple flap without
palpi. Lateral lobes of the hypopharynx (superlinguae) are present
at least in Scutigerella, as shown by Hansen (1930), and a pair of
slender apodemal arms extend into the head from the hypopharyngeal
base. The single median genital aperture is situated on the anterior
part of the venter of the fourth body segment, but since the paired
gonopores of Pauropoda and Diplopoda are on the third body seg-
ment, the median genital outlet of the symphylids might be supposed
to have migrated secondarily into the fourth segment.

20.—The Diplopoda are a specialized branch of the early Symphyla,
in which the somites back of the fourth postcephalic somite are united
in pairs to form double segments. The mandibles are well developed
and have strong movable lacinial lobes, but there is only one post-
mandibular appendage of the head, the gnathochilarium (fig. 52 G),
the morphology of which is uncertain, though the organ is probably
either a combination of the maxillae with the labium, or the labium
alone. The legs of the first body segment are absent, and there are
no cerci on the last somite. Body segments are numerous in most
forms, and all but the first few are generated teloblastically in pairs
during postembryonic development. The paired gonopores are on
the third postcephalic somite at the bases of the second pair of legs.
The Pauropoda are probably an early branch of the Diplopoda, in
which a union of the somites in pairs had already taken place, and
the first legs had been much reduced but not yet obliterated. Special
characters of the pauropods are the lack of movable laciniae on the
mandibles, a weak development of the gnathochilarium, and a branch-
ing of the antennae beyond the fourth segments.

21.—The Hexapoda resemble more closely the Symphyla than any
other of the modern arthropods, a fact recognized by several of the
earlier writers, and Packard (1898) first formulated a definite theory
of the origin of insects from symphylid ancestors. Recently the
evidence in favor of this theory has been more thoroughly reviewed
in the light of present-day knowledge of the apterygote hexapods by

144 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Imms (1936), who shows that the most plausible concept of the
ancestry of insects is that of symphylid derivation. The important
difference between modern Hexapoda and Symphyla is in the position
of the genital openings, the symphylids being progoneate, the hexa-
pods opisthogoneate. It is necessary to assume, therefore, that the
Protohexapoda were evolved from an opisthogoneate branch of the
Protosymphyla.

The Protohexapoda became differentiated as a hexapod group
through the concentration of the locomotor function in the first three
postcephalic segments, with the consequent division of the body into
a motor thorax and a visceral abdomen. The abdominal appendages
were reduced, modified for purposes other than locomotion, or sup-
pressed, but in most cases the abdominal coxal remnants united with
the sternal plates of the segments and preserved the styli and eversible
vesicles inherited from the Protosymphyla, though on the thorax these
structures were lost. The number of body segments was limited to
14 somites and a simple terminal lobe (telson) containing the anus.
The persistent appendicular organs of the last somite were styluslike
cerci, as in Symphyla. It is probable that the true telopodites of all
the abdominal segments were absent. The mandibles became solid
jaws by a complete fusion of the lacinial lobes with the coxopodites,
and thus came to resemble the mandibles of Crustacea, but the
maxillae and labium retained the generalized protosymphylan struc-
ture. The hypopharynx consisted of a median lobe and two lateral
lobes, as in Symphyla, and had a pair of basal apodemes giving attach-
ment to muscles of the gnathal appendages. The eyes were compound.
The protohexapods were opisthogoneate insofar as the paired genital
apertures were located on the posterior part of the abdomen, but the
exact position of the ducts and their outlets was still subject to
mutation, as shown in the variable position of the genital outlets in
modern forms.

The discrepancy in the position of the genital openings as between
Symphyla and Hexapoda raises the chief difficulty in relating the
hexapods directly to the symphylids. The opisthogoneatism of the
Hexapoda, however, is more truly a heterogoneate condition, which
in a broad sense applies to the entire group of labiate mandibulates,
for the primary genital ducts open on the third postcephalic somite
in Diplopoda and Pauropoda, on the eighth in Collembola, on the
tenth in female Pterygota, on the thirteenth (primitively) in male
Pterygota, and on the fourteenth in Protura. Since the primary
gonopores of the hexapods are always fixed with specific segments,
as in Symphyla, Pauropoda, and Diplopoda, the opisthogoneate con-

3

NO.6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 145
dition in the Hexapoda is not comparable with that in the Chilopoda,
in which the genital outlet, though always subterminal, may be on a
quite different somite in different forms because of the variable
number of somites that may precede it. There is reason for believing,
therefore, that the opisthogoneate condition of the Hexapoda has been
acquired secondarily, and that it is a derivative from the progoneatism
of Symphyla and Diplopoda, rather than from the opisthogoneatism
of Protomyriapoda represented in modern Chilopoda. The establish-
ment of the genital openings on the posterior part of the body in the
Hexapoda was very probably an adaptation correlated with the con-
centration of the locomotor function in the thorax.

22.—An early specialization among the Protohexapoda gave rise to
the modern entognathous Diplura, Protura, and Collembola, small
hexapods characterized by a retraction of the mandibles and maxillae
into pouches of the head wall closed ventrally by the labium. The
identity in the structure of the mouth parts would alone suggest a
phylogenetic unity among the above-mentioned groups, but the latter
show also a peculiarity in the development of the hypopharyngeal
apodemes, which structures, instead of projecting as free arms into
the head, as in myriapods and Machilidae, take the form of long
internal ridges that, in Diplura and Collembola, diverge posteriorly
from the base of the hypopharynx as sclerotic linear inflections of
the membranous integument along the folds between the gnathal
pouches and the inner surface of the labium. In Protura the two rods
are united for a part of their length. These superficial apodemes
give attachment to the same muscles as do the internal apodemes of
other forms, and in Collembola they support an elaborate “tentorial”’
superstructure. In many other respects the entognathous hexapods
are widely different from one another, and their inter-relationships
are by no means clear. Except for the common characters above
mentioned, they might be supposed to have had quite separate origins
from protosymphylan or protohexapod ancestors (see Imms, 1936,
fig. 11). They represent abortive lines of evolution that have not
led to higher forms.

The Diplura depart least from the gees branch that has given
rise to the winged insects, since they retain the abdominal styli and
cerci, and have the usual hexapod position of the genital openings.
The Protura preserve a remnant of the primitive anamorphism of
the hexapod ancestors, inasmuch as the last two somites are formed
during postembryonic development, but they lack antennae, styli, and
cerci; the small appendicular organs on the first three abdominal
segments may be coxal remnants of limbs, with eversible vesicles in

10

146 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

one family. The paired genital ducts in both sexes open on the
eleventh abdominal segment. The Collembola are the most aberrant
of all the hexapods, and in some ways the most primitive. They
have only nine body segments, and the single genital opening is on
the fifth abdominal segment. There can be no question that the Col-
lembola are derived from more generalized ancestors having a greater
number of segments, but since, in their phylogenetic history, segment
formation in the zone of growth has ceased after the establishment
of the genital ducts in the eighth somite, it is fruitless to look for
evidence of the ancestral segmentation in the embryogeny of present-
day Collembola. The three pairs of appendicular organs on the
collembolan abdomen are unique in structure, and give little suggestion
of homology with the abdominal appendages of Symphyla, Diplura,
and Thysanura, though it may be supposed that the collophore is a
pair of united eversible vesicles, and that the two paired appendages
are highly developed styli. (For a fuller discussion of the special
features of the Collembola, see _Imms, 1936.)

23.—The main evolutionary line of the early hexapods led from
the opisthogoneate branch of the Protosymphyla directly into the
Machilidae, since in this family are best preserved the coxal accessory
structures of the symphylids (fig. 521) along with the normal ecto-
gnathous mouth parts. Moreover, it was in the ancestors of the
Machilidae that the characteristic ovipositor of the hexapods had its
inception, and, therefore, from the machilid line have been evolved
the Lepismatidae and the Pterygota. The common ancestors of these
last two groups developed two special features in the head structure.
One was the acquisition of a secondary anterior articulation of the
mandible on the cranium, giving the jaw a hinge movement on a longi-
tudinal axis, which brought about a reorganization of the mandibular
musculature, giving the principal function of abduction and adduction
to the dorsal muscles, and reducing the ventral muscles to a condition
of such little importance that they have completely disappeared in
the higher Pterygota. The other feature was the development of the
endocranial framework known as the tentorium, characteristically
present in Lepismatidae and Pterygota, but foretokened in Machilidae.
The tentorium is evidently a product of the hypopharyngeal apodemes
and of a transverse bar developed in the back of the head from lateral
invaginations. Both structures are present in Machilidae, but are
not united. In Lepismatidae the anterior apodemes are reflected
directly from the cranial margins and are united posteriorly with the
transverse bar, producing a typical tentorium. In the Pterygota the
roots of the anterior arms take a submarginal position on the cranium,
and in higher forms they have migrated to the facial aspect of the head.

NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—SNODGRASS 147

The hexapod structure, with the locomotor function centered in
the thorax, apparently gave little if any advantage over the polypod
structure for ordinary terrestrial life, but it furnished a condition
particularly fitted for the development of wings. Hence, with the
appearance of alar lobes on the thorax, the evolution of these lobes
into organs of flight was readily accomplished, and the pterygote
insects quickly achieved a great superiority over the other arthropods.
While there is much to suggest that the winged insects are most
closely related to the apterygote thysanurans, their direct origin from
the latter is questionable. It is difficult to explain, for example, how
it comes about that the pterygote Ephemeroptera and Dermaptera
have paired genital openings while secondary median ducts are already
established in the Thysanura, with openings on the same segments as
in the higher Pterygota.

24.—The Chilopoda are the conservatives among the arthropods;
they are the least-modified descendants of the Protomyriapoda, and
in certain phases of their embryogeny they still follow the course of
development in the Onychophora. The gnathal appendages are prob-
ably more generalized than in any other of the Mandibulata; though
the bases of the mandibles are deeply sunken into pouches of the head
wall, they have strongly musculated lacinial lobes (fig. 53 E, F), and
the two maxillary appendages (C) are but little modified except by
reduction of the telopodites and a partial union of the coxopodites.
The suspensorial sclerites of the hypopharynx maintain connections
with the cranial margins, and bear the apodemes on which the ventral
muscles of the gnathal appendages are attached. The characteristic
specialization of the chilopods is the conversion of the first legs into
a pair of poison claws (B). Most of the other body appendages
retain the structure of simple 7-segmented legs, though at the base
of each is an extensive subcoxal sclerotization suggestive of that in
the insect thorax. The last two pairs of legs are reduced and modified
to serve as genital accessories, and consequently there are no terminal
cerci. Styli and eversible vesicles are absent. The genital opening is
always on the last somite before the telson, but since the total number
of somites is variable, the genital segment may be a quite different
somite in different chilopod groups. Anamorphic postembryonic
development persists in some forms, while in others segmentation
is complete at hatching.

25.—Evolution may be accepted as a fact, but the true history of
phylogeny can never be demonstrated. Though the main branches of
the genealogic tree of any major group of animals are fairly evident,
an endeavor to follow in detail the phylogenetic connections between
more closely related forms invariably leads into a maze of difficulties,

VOL. 97

SMITHSONIAN MISCELLANEOUS COLLECTIONS

148

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NO. 6 ANNELIDA, ONYCHOPHORA, AND ARTHROPODA—-SNODGRASS 149

for it is seldom found that all characters will fit into a scheme of
relationship that attempts to relate every feature in one form with a
similar feature in another. It must be recognized that various struc-
tural adaptations have been often independently developed in approxi-
mately the same way. A successful adaptation will be equally valuable
in many groups, and it is, therefore, not surprising that an adaptive
structure should independently recur either in distantly related or
in closely related groups. To distinguish between such structures
and those that have had an identical origin, however, is one of the
most uncertain tasks of the phylogeneticist, but the very condition of
uncertainty injects into the study of phylogeny the element of per-
sonal opinion which gives to phylogeny that controversial status by
which it never lacks in interest. Every biologist must have a working
creed of phylogeny, but he should not too implicitly believe its tenets.

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"SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER q

"PAWN EE ARCHEOLOGY

(Wirn Six PLATEs)

BY

~ WALDO R. WEDEL

Assistant Curator of Archeology
U. S. National Museum

(PUBLICATION 3484)

) GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OGTOBER 19, 1938

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER 7

THE DIRECT-HISTORICAL APPROACH IN
PAWNEE ARCHEOLOGY

(Wir Srx Prates)

BY
WALDO R. WEDEL

Assistant Curator of Archeology
U. S. National Museum

(PUBLICATION 3484)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 19,{1938

The Lord Baltimore Press

BALTIMORD, MD., U. 8. As

foe DIRECT-HISTORICAL APPROACH IN PAWNEE
ARCHEOLOGY

By WALDO R. WEDEL
Assistant Curator of Archeology, U. S. National Musewn

(WitH S1x PLaTEs)

When the University of Nebraska Archeological Survey was
established in 1929, its then director, Dr. W. D. Strong, envisaged
two primary objectives. The first was a preliminary survey of the
State, including both surface reconnaissance and sampling excava-
tions, designed to give a general bird’s-eye view of the area as a
whole. With this was combined a second aim, namely, an effort to
locate and work such sites as could be definitely identified with villages
visited and recorded by the early white explorers in eastern Nebraska.
It was believed that by isolating and clearly defining the archeological
characteristics of the historic peoples a whole series of sites could
soon be removed from the category of unknowns; and furthermore,
that a comparison of materials so identified with earlier remains in
the region might open lines of attack which would permit the
establishing of a time sequence extending “from the known historic
into the unknown prehistoric.” Toward this second objective a
serious beginning had already been made by A. T. Hill, of Hastings,
Nebr., who since 1922 had accumulated a considerable quantity of
archeological materials from sites identified as Pawnee through criti-
cal study of early nineteenth century maps and narratives. This
collection, as well as numerous valuable historical leads, was
promptly made available to Dr. Strong and his coworkers, and it
became the starting point for the study of Pawnee archeology. In
this paper it is proposed to review very briefly the methods and some
results of this approach to prehistory in the Pawnee area.

It was not chance alone that prompted selection of the Pawnee for
the first systematic attempt at isolating a historic archeological com-
plex in Nebraska. Aside from Hill’s pioneer labors, consideration
was given to the fact that this tribe was one of the largest, best known,
and most powerful in the entire Plains area. Among the semi-
sedentary so-called village tribes of the Missouri valley, including
both Caddoan and Siouan groups, probably none shows evidence

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97, No. 7.

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

for a longer occupancy of its historic locale than the Pawnee.
Furthermore, of all the Nebraska peoples, the Pawnee appear to have
offered the most effective and prolonged resistance to the host of
alien practices introduced by the whites and to have retained longest
their own customs. As to documentation, allusions to the Pawnee
may be found from almost the very beginnings of recorded European
penetration into the interior United States, although it is true that
many of the seventeenth and early eighteenth century sources of
information leave much to be desired. Prior to about 1800, hazy
geographical concepts, occasional tribal shiftings, and the often hear-
say origin of the explorer’s observations made impossible the record-
ing of village locations with the exactness necessary to permit their
individual identification today. After that date, thanks to the lucid
narratives and excellent maps of such men as Dulac, Pike, Lewis
and Clark, Long, and others, the historical record has enabled us to
correlate with reasonable certainty the native towns with known
archeological sites. Excavations in sites so identified have revealed
the distinguishing characteristics of historic Pawnee culture, insofar
as these include nonperishable material traits. As the term is now
used in Nebraska prehistory and in this paper, historic Pawnee
archeology refers to the antiquities from documented village sites
where the Pawnee are known to have been living in or after circa
1800." Needless to say, throughout this period the archeological
picture can be greatly enriched through the ethnographic observa-
tions of many of the white travelers.

During the nineteenth century, the Pawnee villages with but two
or three apparent exceptions were centered about the confluence of
the Loup with the Platte River. Both of these streams flow in a
general easterly direction through broad flat-floored valleys inclosed
on either side by lofty bluffs. Above the mouth of Shell Creek the
native towns stood on terraces or second bottoms well out of reach
of floods; below this point suitable terraces are mostly lacking and
the sites are situated on the bluffs with the river sweeping past their
bases. The tree-fringed watercourses are in marked contrast to the
dry rolling, formerly grass-covered, uplands which lie beyond the
valley margins. To the natives the latter were suited only for hunting
and it was the fertile river bottoms, with an abundance of wood,
water, arable ground, and shelter, that determined the location of
their villages.

* For a discussion of historic Pawnee archeological remains see Wedel, 1936,
and Strong, 1935, pp. 55-61.

NO. 7 PAWNEE ARCHEOLOGY—WEDEL 3

The extreme limits of the known Pawnee settlements were, to the
west, near St. Paul on the Loup and Central City on the Platte; to
the east, downriver, they ran to Leshara or Yutan on the Platte
(see fig. 1 for location of all sites discussed herein). Within this
120-mile stretch of river valley they shifted back and forth as fancy
or circumstance dictated, leaving it only for their seasonal hunting
excursions. The exceptions, it may be noted, included two sites on
the Republican near the Kansas-Nebraska line and one on the Blue
near Blue Springs, Nebr. That this nineteenth century restriction
of habitat was in effect long before will become apparent presently
when certain additional historical and ethnographic facts are con-
sidered. Here it is desired to add only the observation that all of
these village sites, in addition to a somewhat decadent aboriginal
material culture, yield also many articles of iron, copper, brass, and
glassware.

Within this same area, but of even more limited distribution, are
found other sites whereon the native remains are far more abundant,
of superior quality, and associated with much smaller quantities of
white contact material. These sites extend along the Platte-Loup
riverway from Schuyler on the east to the vicinity of Genoa on the
west, a distance of approximately 50 miles; they are mostly on the
north bank, but one is also known on the south side. Generally, the
sites are large (from 15 to I00 acres or more) and compactly
arranged ; not infrequently they seem to have been located on bluffs
or hilltops with an eye to defensibility and in a few instances they
were further protected by earth walls and ditches. To date about a
dozen have been placed on record. The sites are particularly abun-
dant from Monroe westward, where for more than 8 miles remains
occur almost continuously along the Loup and on the lower portion
of Beaver Creek. In the aggregate these antiquities cover many
hundreds of acres, and prior to introduction of modern farming
operations, innumerable house circles, middens, and artifacts were
to be found. Because of their occurrence in the very heart of the
historic Pawnee habitat and since they yielded smaller amounts of
contact material than the identified nineteenth century Pawnee sites
while exhibiting many similarities to the latter, it was thought that
they might prove to be an earlier, if still post-EKuropean, phase of
Pawnee culture. Consequently, in 1931, as a sequel to the study of
the historic Pawnee, two of these protohistoric’ sites were partially

* Protohistoric sites -yield limited amounts of glass and metal trade wares,
indicating their occupancy, at least in part, since the arrival of Europeans. They

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

examined by parties from the University of Nebraska. About 8 weeks
were devoted to excavation of houses and middens at the Burkett
site near Genoa and at the Gray-Wolfe site north of Schuyler. All
but one week of this field-work was in direct charge of the present
writer, under the supervision of Dr. Strong and with much active
assistance in the field from Mr. Hill. A detailed description of the
findings has been published recently by the University, and the
remains have been assigned to the “Lower Loup Focus of an
unnamed aspect of the Upper Mississippi Phase.” * A wealth of
additional information has since been gathered by Mr. Hill for the
Nebraska Historical Society at three other protohistoric sites near
Genoa. This latest work, completed in 1936 and as yet unpublished,
included the opening of 10 houses, a number of large and _ prolific
caches, and the collecting of several thousand artifacts, all at sites
lying within 4 or 5 miles of the Burkett site. Pending future analysis
and detailed comparison, it must suffice to say that preliminary exami-
nations indicate a close similarity between this material and that
already described in print from the Burkett and Gray-Wolfe sites.
In passing it may be noted also that extensive surface collections
from most of the other protohistoric sites in the immediate locality
diverge in no significant respect. In short, a fairly uniform and
consistent cultural complex seems to be manifested at the sites
designated on the map as belonging to the Lower Loup Focus.
Historic archeology in Nebraska received added stimulus in the
summer of 1935, when Hill explored the large protohistoric Leary
site on the Nemaha River in the extreme southeastern corner of the
State. This has been elsewhere described and identified as Oneota.
Midwestern archeologists are inclined to view the Oneota culture
in Iowa and adjacent States as possibly early Siouan.* There are
indications that the Leary site was inhabited contemporaneously with
or possibly slightly earlier than the known sites of the Lower Loup

differ from historic sites in that the written records are too general to permit
their individual identification with villages actually visited by white men. In
time they antedate 1800.

*Dunlevy, 1936, pp. 147-248 (quot. p. 216). A discussion of the placing of
the Lower Loup Focus in the McKern taxonomic system is beyond the scope
of this paper. However, it may be pointed out that at least four of the nine
Upper Mississippi Phase determinants listed by Deuel (F. C. Cole and T. Deuel,
Rediscovering Illinois, table 2, p. 214, 1936) are unreported from the Lower
Loup Focus and incidentally from the historic Pawnee as well. The present
writer regards as debatable the assignment of either complex, or of a hypo-
thetical aspect which might include both, to the Upper Mississippi Phase.

* Hill and Wedel, 1936; Griffin, 1937.

NO; 7 PAWNEE ARCHEOLOGY—WEDEL 5

Focus but no documentary record exists as to the tribe which
inhabited it. It definitely antedates the historic Pawnee sites of the
nineteenth century. This is of some interest because there are Pawnee
traditions pointing to early residence of the tribe somewhere in this
section of southeastern Nebraska, suggesting the possibility of a
generic connection with the Oneota.

As regards the relation of these three postcontact archeological
complexes to one another, dissimilar conclusions have been reached
by different field and laboratory workers. Strong expressed the
belief that the sites now labeled collectively as the Lower Loup Focus
probably represented a very early historic horizon directly ancestral
to the somewhat simpler and decadent Pawnee culture of the nine-
teenth century. His use of the term “protohistoric Pawnee” in speak-
ing of these remains reflects a view with which the present writer
has elsewhere indicated his general agreement.’ .Dunlevy, on the
other hand, dissenting after her detailed analysis of material from
‘two of these sites, was persuaded that the Lower Loup Focus is
more closely related to the Oneota than to the historic Pawnee.’
Since these differences of viewpoint occur among individuals dealing
with substantially the same materials, it seems worthwhile to re-
examine the data on which they rest.

In the accompanying table the presence or apparent absence of
traits has been indicated for each of the three cultural complexes
above mentioned. The traits, totaling 120, have been grouped in
seven categories which, with exception of ceramics and miscellaneous
items, are based upon function rather than on form or substance.
Traits for the historic Pawnee and the Lower Loup Focus have been
compiled largely but not exclusively from published sources. In the
absence of complete analyses for the recently worked sites, the data
therefrom have been incorporated in and added to a check list based
on the published studies. Actually, this somewhat superficial treat-
ment involved no changes in the list other than its slight expansion
to include a larger number of traits. Data on the Oneota Aspect,
including three Wisconsin variants or foci, have been drawn from a
list furnished by W. C. McKern, of the Milwaukee Public Museum,
which has been supplemented by the published report on the Leary
site in Nebraska. No attempt has been made to weight the various
elements or to determine the degree to which a particular trait may
be present in one or another of the groups. It has not always been

° Strong, op. cit., pp. 68, 207; Wedel, op. cit., pp. 38-42, 74.
°Dunlevy, op. cit., p. 216.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

possible to refine the traits as fully as desired, owing to differences
in terminology in the sources used and to inability to examine all the
material at first hand. It is believed, however, that the data are
sufficiently extensive and representative to be strongly indicative of
trends, at least.

TABLE 1.—Presence or Absence of Traits in the Historic Pawnee, the Lower
Loup Focus, and the Oneota Aspect

A B Cc
Lower
Historic Loup Oneota
Pawnee Focus Aspect

I. ARCHITECTURE AND VILLAGE COMPLEX

Villages
1; Warge, intensively occupied /sites-actr sacien «irae - x x x
2. Walledvor defensively located’. sei s).j0 0 oestriol se x
3 ENumeroussoutside Caches. «ang sain eeces + ese x
Houses .
4. Shallow semisubterranean circular earth-covered... x x
5. Vestibule entrance in east or south............:.-; x Se
Gy Onlimedkcentrall stinepitecrcmes cemeteries x ex
7. Bison-skull shrine opposite door............++.++8- Bx x
82) Hour, saan Central yPOStS. eee sc anette oleae onl eise mses x x
One Moreithan oun, centhalepOstSererriietn tty rretelterl- BS x
10. One or two rows of widely spaced outer posts...... oe x
TTS Lniside scache sets woroetac cnc meister chaise ere eke ers a x
12. Numerous small, closely set, slanting wall posts..... se x
II. Ceramic COMPLEX
Temper
Lgee MGM bean a ceeie alot rer choot we topelee eet taste carne loynnettete take ol x x
PAS hell Meiers seyes easie Sas rele shea etal sO ieye os akelerecnen ans x x
Texture
15. Fine to medium coarse.......secescesseeccsecvcens x Ex: 5k
Structure
WO Tiley ofan stearte te chaser ace tefoyste Mies eia cheeks ovat aots ole =linsoraleke Gai x x x
7s GAM AE pete eon tase sets et elena eke ra fave ote evade estate teuaiot Xs
Hardness
18. 1-4, softer predominating. ...........eeeeeeeeeeoees ot
19. 3-6, 4-5 predominating. ..........2..csseceseeenees Ba x
Surface finish
20) irresularlys smoothed! i).)5 ochre sins ee = ane ais ee se x x x
ar. Polished (mpertecthy ) o..(c.. tae imi = eevee el =imlels © = ote von 5% x x
22. Marked by grooved paddle..............-+e+eeeeee 5g x
Color
23. Light to dark gray and buff, dull terra cotta........ x x x
Thickness
QAg fe -Ne- INCH CANDE LO. ce see toate ioelt hemi oa ice ee eryae x x x
Lip form
BS, SWanen andes, cals eens Pesisio en reel olode ae tietercie es eel steals Bk x esd

by. HERG Eh Sealers BH ere OIRO cr racteeretet ete estrone kena rete fatelrs x x x

NO. 7 PAWNEE ARCHEOLOGY—W EDEL Fj,

TABLE 1.—Presence or Absence of Traits in the Historic Pawnee, the Lower
Loup Focus, and the Oneota Aspect (continued )

A B Cc
Lower
Historic Loup Oneota
Pawnee Focus Aspect

Rim form
27 eee oh direct, flaring.) .Ss-ince ccd sciseteee ote e x x BS
2am Collar orsloraced!s.j0's. selene erect arene cine Bi BG
Ay, Clarsianeel’ 5 agcocad. nla tohacsest ty Taya io ontohavete ketotee eve elateetere te x
Neck form
30. Line of juncture between rim and body............ x BX x
Sit, Milos jnaemopincer! Goagaasotooocssssossoocudoudctoe x Ex 5
Orifice
£3 DAMS 1 Oa CR BAM ys etc shat do A\citerelaveueta oe Stars Sa eno eee x x x
RENO UETIGLIM eyes 2) 05 safe aia sm loa sapvie Gree ae eR x ex x
5 Aiea O) i cl ieee rou fouraro Fo les eiccfol So sence ko he ate seuSpavenet ay meeene arevereiey toate x Exe
Shoulder form
3 SamaELNOTITL CMM pes pec eve. sree Ocelot ore teial che esto soe ake ee RRO BK: x xe
Basal form
3 Orem INO UNG Ora are spas i er aveholins ide eats eas cle S cases ORR ere aR x se x
Byam SUD CONTCAl se Mirae Wise retets vice ee oe ern e eo ere BG x
Handles
Bowe NaGhOWsLONDLOaG mihlatesthapiuices palredae emcee ex: esa
3 OS etlLE OO Py dave Giloieo eval odie rere eies aid oral Sootsthuelic elas. sie eecelespatevers Ba ox
AOmeN tennatescollarmtabssotinm handlestemaes eerie 5
AlTeammma VALTTLi EDL pas treer sieree arate revert evs oie. store sso orlel caete a eoyatie caters 5 x
Decoration
AS MIE TINR AB ucts lels Bracalaxctos Sata ( dis)a.0s wisi 90 oo Sarees Ae hae x x BS:
Ase Shomiaermarea to lip, neck (plait. «<a> serderretele x x x
44, yineised rectilinear parallel line-motifs:..,.2...... 5 sis x oe Xx
Ace Opposeduscriesson marallelplinestrciimcriae eeticicemicce x cg x
AO Herring poneand. Chevron On fim occ ees ee one ce x Bs
47. Concentric pendent chevrons inside rim............ Xs
AG. aConcentric circle motif and/or CrosS.:.... 0 sec ss cx:
Ao: Geometric series on lines and dots. ..... =. +. --se= x
EOmdnalledtotmiltuted decoratiOnassasemitemaeriscie eee crn x
Miscellaneous
EST We SITLL WM EIOSULS sor 5.0 oie ove ictwle: voile telePoret home ege «| oretaterete « x x
BZ oimallidecorateds dishtall- 1eutinese-reiceeciees eee x x
POM SevOreLeduwashiOL pSCUdO-Skipmcietentrelttelettctetaere tte x os
BAe eerfonateds pottery, Gisksr jaro isye\ser tee atatels sielefelers Xu x
Hoe otelrdsawathe handles! /asc seiereieve ae eieracietnestore erates x
One CitsnerdsrandspisectedsvesselSenme-ieesereeteets x
III. HorticutturE AND Foop-GATHERING
57. Intensive horticulture, with maize, beans, etc........ Be x x
Re, Jaloessanevale Oni lovteral WeemulEVS) Gon omo boca qdondoodde x Ba BG
EG MV VOOGEIe MOAT ecjscictesceslecorn cicvoteratavn eyater cietaleretsleceronete Ex

60. Stone mortar: irregular, shaped, flattened surface... x x x

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

TABLE 1.—Presence or Absence of Traits in the Historic Pawnee, the Lower
Loup Focus, and the Oneota Aspect (continued)

A B Cc
Lower
Historic Loup Oneota
Pawnee Focus Aspect
IV. Mirirary AND HUNTING CoMPLEX

61. Arrowpoints, small triangular unnotched.......... px

xs x
62. Knives: diamond-shaped, beveled................-- x x
Ogee knives ovalland/Ofetlalccasm eer ace tee eee px D¢ x
64. Scrapers: small to’medium planoconvex............ ae x x
65. Scrapers: large elliptical quartzite or sandstone.... x Be
GO. Dall s ica tik secoteclt eo ee ie cee a er ene eX x x
67. Abraders: paired longitudinally grooved sandstone... x EX: x
68. Abraders: amorphous pumice lumps...............- ee
Gor Mauls oroovedia nce cat tacit ence aoe nee x x By
ME: PARKES MOTOOVEG (idee Sea oe OR eo ? x
7x, Celts: polished diorite or hematite. 2.50... 30 0e en ce x x
TO mEtAmiierStOlessipittedamarrr ements cies otieee iors EK: 5x2
730 eANdz-shaped elkhorn hideyscrapersmees eee a ee x
74. Deerhorn “cylinders” or tapping tools.............. x
75. MOCEEMOG tip HAKELS afore ClerNcrst sehen avers cit ors sto aero Be EX
76. Deerhorn projectile points, conical, socketed........ x
77. Bone projectile points, socketed, square or conical... x:
gion JBOMe confacinils foros, GwemiMEls ocongbacensoosuses Ex
7Ow Dundes mor cane (aLhOwshaitsis) mercer ameter x
80. Perforated ribs (arrowshaft straighteners)......... BG x x
oi... Notched fleshing tools or erainerSsc-c.2- 0522+ ae ese x
82) Shouldemablades scrapers nee seam meee eine xe
835uCelthke: antlermsctaperssanneee eee x
84. Mctapodial sbeamenrsmeeeoe cee nate eee eee Xs
85. sBone -fishhoolss hia. Seiaatecs ee ce eee oe eee x

V. Dress, TEXTILES, AND ADORNMENT
86; B1son-haig clothvand/ or cordages..- eerie ee x Be
872 SA wis yun 5 od stats ass «ai syed = aia ater eet telece eieee eae eae Peer etavE x x Xe
88. -Eveleted: needles.) SER LN ae ene eicees pieiarteroeiate Xe
86... Plume: holder #2 \.2)2 aoe Soe hae see eaten: 5
0)... Roachvspreadenss smnccrme tse permet stores we ote cine exe
OT. “Combs i.0)s se signed ooeaeee sch ek eee see eaten x
Oza Braceletsipand ome coOngetsnieerieicicmccieia etmek oe bs x a
Oey lettbolerlyemtas, (dob )onesunooceoBoovoenGoodc se BS x
04. uPolished@honertubess. teh occ cc tere een Ba x x
O52 vRush ma ttins Misrescielete os cinis operetta easton x
96: (Flat polished-bone:mat needles: o.. sei strc c= ol x
O72) Lwinedbagstotweretall materialist oe BG
98. Shell ornaments, variously shaped......-..-.-..-.-- x x x

VI. CEREMONIAL COMPLEX
60,. Primary extended: burials. soc ictar seis steps cer x
160)" Primary flexed burtals)s.<c0< sc evarets nesin< lois Muatnlars See ola xe Ba x

Tor: Secondary bundle burials ©. cin sce o em cee x

a8 Rare, probably atypical.

NO. 7 PAWNEE ARCHEOLOGY—WEDEL

TABLE 1.—Presence or Absence of Traits in the Historic Pawnee, the Lower
Loup Focus, and the Oneota Aspect (continued)
A B c

Lower

a

Historic Loup Oneota

VI. CEREMONIAL CoMPLEX—C ontinued
EO Zao Gray Cx MALMUEUTE 1420.5) nai g sists Se ete oe ee eee x

exe
Fog eB ucial tm dis pits or cachesy « .)52.- sis, <are aisiera ale Be xs x
TOAD Uctaleinvon winder mounds. ee aeeieeeimaeie cetiae BX:
105. Gaming stones ?; bun-shaped, flat pitted face....... EX:
TOON Gy pstiime ChyStals wOnkeds . en. cs s aan recites esos ae
107. Shaped balls of crystalline stone (grave finds)...... x
Boa Wihetstenes: ‘(srave: finds))is sued. saeco edits x
HOO Es pPesTOr PoOliShediStOMe as se sera seicieclae ete ete el > oe Xs
HOMES PESTOLM Gla yates crane ret fro tieyet sionals Dieters sre wae er otetebelats x
111. Pipes: elbow-shaped or equal-armed............... se 5 x
112. Pipes: “Siouan” type, stem projects beyond bowl.... x
BS Meri pe sir CISke DO Willers ercracineca ae os soteero are levees to dhepaltes x x
SIANMPEA NES) mee CMAGY ocsose snake etek laaiesisee ees x
fine Ornamented: animal, Skulls cies... oswsase lanes x x

VII. MisceLLANEOUS
WTOSMPlTTEISeC SON CAD eLSivsrety losis sie alee tye tre cierele ors .cls steers x x x
Mpg mes ISGHIE GEIL S PODS 15 afsis)ace)clsn etayeraie Sis saa esaie ea 5
Lia banned mussel shell Spoonse + 2.2 cic aes oes: x
TANG) Meme) MMA aeO GIGS wrserchel cy chensbave tenses: 2) codyeve.vs cVSie ie Sis -Spaneya matron BS x a
POP sally WnOnes (SCONEM: £1DS)) <c wcteuien sie orsie esa d sees ae x sm

ANALYSIS OF TABLE: SUMMARY

Total number of traits—120
Historic Pawnee has 80, or 66.6 percent of total
Lower Loup Focus has 82, or 68.3 percent of total
Oneota Aspect has 74, or 61.6 percent of total

“Universal” traits—30, or 32.5 percent of total

Pawnee Focus Aspect

39 universals in 80 historic Pawnee elements.......-......--. 48.8 percent
39 universals in 82 Lower Loup Focus elements............. 47.6 percent
39 universals in 74 Oneota Aspect elements................. 52.7 percent

Out of total of 120 traits—

26 occur only in historic Pawnee and Lower Loup Focus..... 21.7 percent
9 occur only in Lower Loup Focus and Oneota Aspect...... 7.5 percent
3 occur only in historic Pawnee and Oneota Aspect......... 2.5 percent

On basis of 81 nonuniversal traits these percentages become
respectively 32, 11, and 3.7.

Traits occurring in only one complex—

SCOT IGWE AWC ee fae eis eer eee Ee PN aerSLcne Pee Whol oie = eitsbcte sieve fot erasayieks 12

Mery eyeet WM ST AIOE CLI sca 2— cab rnis footed chegedane eller ae elevates n ntons cov tec one Sear nb) Sayaka 9 'en's

Beata WAS BECE G Aisiirsied = aiclers wiciciace seine aias lintels # te ets)arasaiet win) e's) 6.0.0! nloisierave me 23

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Analysis of the table shows first that out of the total of 120
different elements the historic Pawnee and the Lower Loup Focus
have, respectively, 80 and 82 (66.6 and 68.3 percent), and the Oneota
Aspect has 74 (or 61.6 percent). Of the 120 traits, furthermore,
39 are common to all three culture complexes. Since this represents,
respectively, 48.8, 47.6, and 52.7 percent of those found in each
complex, it is evident that there is a strong underlying relationship.
These “universals” include elements in practically all of the categories,
but occur least commonly under the “Architecture and Villages”
heading.’ As regards specific relationships between any two of the
three complexes, we find that 26 traits, or 21.7 percent, occur only in
historic Pawnee and the Lower Loup Focus ;* 9, or approximately 7.5
percent, only in the Lower Loup Focus and the Oneota;° and 3, or
2.5 percent, only in historic Pawnee and Oneota. Since it is these
relationships within the defined universe of three which are the
principal concern here, we may reduce our totals and sharpen the
above differentiations by omitting the “universal” traits. Thus, using
the 81 nonuniversals as our basis, the percentages become, respectively,
32, 11, and 3.7. Whichever set of figures is taken, it is apparent that
the table indicates very nearly three times as many traits in common
between the historic Pawnee and the Lower Loup Focus (and in no
other) as in the Lower Loup Focus and the Oneota.” Evidently the
suggested connection between the first two complexes, considered on
purely archeological grounds alone, is considerably closer than that
between the second pair. This is the more striking in view. of the
previously indicated fact that the Lower Loup Focus flourished at
the very beginning of European contact and approximately con-

™The single rectangular earthlodge floor found at the Leary site has not
been included in the present table since there seems to be general agreement
among field workers that this type of structure is not characteristic of the
Oneota. I am inclined to agree with McKern’s suggestion that the occurrence
of earthlodges in the western Oneota sites “may be due to the taking on of foreign
traits after leaving the area of earlier occupation.” (Letter of Oct. 28, 1937.)

®Including among others nine in architecture, besides such elements as de-
cided predominance of grit tempering, use of grooved paddle in surfacing pottery,
small decorated “fishtail” figurines of clay, large elliptical quartzite hide scrapers,
bone paint “brushes,” notched fleshers, ornamented animal skulls (rare), etc. In
the trait list these are Nos. 2, 4-13, 19, 22, 28, 31, 37, 41, 46, 51-53, 65, 72, 81,
93, I15.

°Including five in ceramics, besides diamond-shaped beveled knives, platform
disk pipes, scored ribs (tallies?), and antler tip flakers, Nos. 14, 34, 38, 39, 54,
O275 e103, 020:

” Cf. Dunlevy, op. cit., p. 216.

NO. 7 PAWNEE ARCHEOLOGY—WEDEL ie

temporaneously with the Oneota, whereas the Pawnee traits are based
on sites inhabited one or more centuries later toward the close of the
tribe’s residence in Nebraska. The conclusion seems inescapable
that the Lower Loup Focus stands in very much closer and more
direct relationship genetically to the later historic Pawnee than to the
contemporaneous Oneota peoples.”

With the Oneota culture and its probable Siouan connections we
shall not further concern ourselves here. Its role in the development
of later native civilization west of the Missouri is not yet clear,
although it probably introduced into the Pawnee area various innova-
tions in ceramics, pipe-making, stone-working, and certain other
fields of activity. At the moment, there is no reason to regard it as
in any sense basic to historic Pawnee culture, since its contributions
seem to have been rather in matters of detail.

Bearing directly on the question of the nineteenth century Pawnee
and their postulated descent from the Lower Loup Focus are certain
noteworthy nonarcheological considerations. These seem to have been
generally overlooked by those who challenge such a correlation on
grounds (1) that the Pawnee have no legends concerning the sites,
and (2) that the recent occupancy of the region by that tribe proves
nothing as to its connection with the older remains. Both points
can be met squarely with recorded data. Thus, to take up the first,

4 The kinds of traits comprising similarities and dissimilarities in the respective
pairings is perhaps of as much significance as the absolute numbers. For ex-
ample, while many of the hunting and skin-dressing practices were similar
throughout, important differences are probably implied in the presence of fish-
hooks and metapodial (split leg bone type) beamers in the Oneota. Both the
latter- items are widespread throughout the eastern United States, incidentally
occurring also in prehistoric cultures in the Plains. The Pawnee and Lower
Loup peoples apparently did not fish, and the outstanding feature of their skin-
working industry was its distinctly Plains character; e. g., large elliptical
quartzite scrapers, the notched fiesher, bone paint “brushes,” and probably the
adzlike elkhorn hide scraper. At least a part of the subsistence economy of
the Oneota, as well as the supposed bark or thatch house type, mound burials,
extended use of woven mats, and a number of other items which this group alone
of the three possesses, all tend to link them with eastern peoples and stamp them
as comparatively recent arrivals west of the Missouri. The Pawnee and Lower
Loup Focus peoples, on the other hand, resemble each other closely in virtually
every fundamental respect and such common elements among them as the earth-
lodge, pottery, horticulture, and other less distinctive items clearly have con-
siderable historic depth in the eastern Plains. Onto this horticultural base they
had grafted a hunting complex of western type, differing considerably but evi-
dently well attuned to the peculiarities of the former. The successful integration
of the two modes of life, both involving local ingredients, would in itself suggest
a considerable period of adjustment in loco.

12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

on at least two occasions, Pawnee Indians have claimed certain of
the protohistoric sites as the former dwelling places of their tribe.
In 1867 Hayden collected a number of potsherds from “a Pawnee
village site on Beaver Creek, Nebraska . . . . ,” some of which were
subsequently figured by Holmes.” Hayden nowhere records the
exact location of his finds, but Hill has since shown that two very
large and almost contiguous protohistoric sites occur on the right
bank of Beaver Creek a short distance above its mouth, while 2 or
3 miles to the southwest is the Burkett site (fig. 1, nos. 16-18). The
ceramic and other remains from the three are very similar, and they
were undoubtedly inhabited by the same people and at about the
same time. In all probability Hayden’s specimens which are of Lower
Loup Focus type were picked up on one of these locations. It is,
therefore, noteworthy that he says:
No Pawnee Indian now living knows of the time when this village was in-
habited. Thirty years ago [i. e., about 1837] an old chief told a missionary that
his tribe dwelt there before his birth, but he knew nothing of the use of stone
arrowheads, though, he said, his people used them before the production of iron.
When the “production of iron” here began is not known, but the
old chief’s story tends to imply habitation of the site in question prior
to the middle of the eighteenth century. The claim gains support from
another tradition recorded by Bruce in his account of the North
brothers and their Pawnee scouts.” This is much more explicit and
telling. It alludes to a battle which took place long ago between the
Pawnees and the Poncas, when 500 of the latter made a treacherous
but unsuccessful attack on a Skidi Pawnee village on Shell Creek:
north of Schuyler. The time of this alleged raid is wholly unknown,
but it could not have taken place recently because there is no historic
record to indicate that the Skidi, or for that matter any other Pawnee
band, dwelt on Shell Creek as late as 1775 or after. Interestingly
enough, at the precise locality where the old Skidi village is said
to have stood, is the Gray-Wolfe site, one of the first of the Lower
Loup Focus to be intensively studied and also one of the two on
which the complex as defined is based. (See fig. 1, nos. 24 and 25.)
Finally, in a myth explaining the formation of the Skidi federation,
Murie locates by streams two of the ancient villages. One of these
was on the Elkhorn River, the other on Looking Glass Creek.” This,
if far less definitive, is still suggestive, since the lower course of the
latter is sprinkled with not one but several related protohistoric sites.

“ Holmes, 1903, pp. 200-201 and pl. 177; Hayden, 1872, pp. 411-412.
* Bruce, 1932, pp. 42-43.
““ Murie, 1914, p. 554.

PAW NEE ARCHEOLOGY——-WEDEL 13

NO.

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kerry) “SZ WSIM “St auIeIQoOW “VI yooig asioH 9 says
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14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Insofar as they are any clue, legends are thus seen to point toward a
Pawnee authorship for at least some of the sites.

It is unnecessary to stress the fact that mere areal concurrence of a
nineteenth century tribe and a certain archeological complex is, per se,
no proof of direct relationship. In the case of the Pawnee this par-
ticular argument has never been used except as a possible corroborative
circumstance. However, a careful study of the documentary history
of the tribe tends to strengthen rather than weaken its force. Here
it is possible to pass in review only a few of the more significant
points; for further details the reader is referred to recent publica-
tions on the Pawnee and citations therein. Prior to the last quarter
of the seventeenth century the sources are inconclusive as to the
location of the tribe. Coronado, in 1541, places the province of
Harahey, tentatively identified as Pawnee territory, north of Quivira.
Later Spanish documents locate Quivira somewhere in central
Kansas and its people are believed to have been the Wichita. If these
identifications are correct, they suggest the presence of the Pawnee
in southern or central Nebraska at this early date. A century and a
quarter after, in 1666, Perrot mentions the Panys but without defining
their habitat. Bandelier notes their presence as captives in New
Mexico in the seventeenth century observing that they were not
uncommonly ransomed from the Yutes and Apaches.” By 1673,
however, they had become sufficiently well known to be shown on
Marquette’s map, as also on that of Hennepin in 1678. Before 1680
the Spanish in New Mexico heard rumors of Frenchmen among the
Pawnees, and, wherever the location is given, subsequent narratives
consistently place the Pawnee on the Rio Jesus Maria, north of
Quivira. This stream is identified by historians with the Platte.”
For the eighteenth century there are many more records, as well as
numerous maps showing ethnic distributions in the Missouri drain-
age. Curiously enough, with all the unrest and tribal movements
manifested therein from time to time, the Pawnee are almost always
shown as a relatively stable group localized west of the Missouri
on streams identifiable with the Loup, Platte, and possibly Republican
Rivers. Particularly interesting in this connection is the 1718 Delisle
map of Louisiana and the Mississippi River,” because it depicts
with remarkable accuracy the geographical details of the present
Nebraska region (fig. 2). It shows the Pani (Pawnee) in 12 villages

* Wisconsin Hist. Soc., Coll., vol. 16, pp. 15, 27, 1902.

** Bandelier, 1890, p. 185, n. 4.

™ Thomas, 1035, pp. 12, 37-

8 Delisle, G., Carte de la Louisiane et du Cours du Mississippi. Paris, 1718.

NO. 7 PAWNEE ARCHEOLOGY—WEDEL I5

on the “Riv des Panis,” unquestionably the Platte, about the mouth
of a large unnamed tributary entering from the north. Comparison
with modern maps leaves little room for doubt that this tributary
denotes the Loup, on whose banks the Panimaha (Skidi Pawnee)
are represented, also with 12 villages. This is the first really con-

vincing cartographic evidence that the Pawnee were established in

ae grande ;

pravies *

ler Octotata
5

Padoucas
a.

Onager

Fic. 2—Portion of the Delisle map (1718) showing the
Pawnee towns on the Loup and Platte Rivers in east-central
- Nebraska.

the Loup-Platte region in considerable numbers in the first quarter
of the eighteenth century. Taken in conjunction with the data
gleaned from earlier narratives, it adds strength to the view that this
tribe has occupied its historic nineteenth century locale since the
very beginning of white explorations.

Of much concern to the archeologist using the so-called direct
historical approach is the question of when European manufactures

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

first began to reach his area. The discovery of such materials may
offer an opportunity to determine approximately the time of occu-
pancy of the sites or levels in which they occurred. Sometimes it is
possible to identify beads or other trinkets with types known to have
been made at certain stated periods in Europe. There are, of course,
limitations to the method, and it must be-used with due caution. Such
objects as glass beads, copper bells or ornaments, and other small
trinkets may have, and probably very often did, spread from village to
village and from tribe to tribe, wholly independent of the trader
after their original acquisition by the natives. They might thus
precede the white man by several years. Also it is possible that the
earliest traders left no written records, or that such as they may have
left were lost or for other reasons remain unknown today. Still,
where trade goods occur in small but consistent amounts in several
related and neighboring sites, it seems reasonable to believe that a
steady and direct, if perhaps limited, traffic had been established, and
that historical records may offer valid clues as to the approximate time
involved. It is theoretically possible that stray pieces reached the
central Plains indirectly from New Mexico through the expeditions
of Coronado (1541), Bonilla and Humana (1594), Ofiate (1601), and
others, or as a result of raids against the Spanish settlements or their
Apache and puebloan proteges. -These, however, must have been
of minor consequence. As a matter of fact, the Spaniards credit the
rival French from Canada with introducing firearms, metal kettles,
axes, and the like to the Pawnee,” but it is not certain just how early
this trade began. The first Frenchman to penetrate the region west
and south of the Great Lakes is generally believed to have been
Nicolet, who in 1634 visited the Winnebago and Illinois in what is
now southern Wisconsin and northern Illinois.” Owing to the
hostility of the Iroquois and for other reasons,-this voyage of explora-
tion was not immediately followed up. It seems extremely doubtful
that there was any appreciable commerce with tribes west of the
Missouri prior to about 1650. By 1680 the Spanish had reports of
French trade goods among the Pawnee on the Platte and in 1706
their Apache allies killed a French couple somewhere in what is now
northeastern Colorado. All this leads to the inference that regular
trade was established in the central Plains region sometime between
1650 and 1700. It is worth noting that from the first the Spanish
records relating to French activities in this area uniformly link with

* Thomas, 1935, pp. 12ff.
* Butterfield, 1881.

NO. 7 PAWNEE ARCHEOLOGY—WEDEL 17

them the Pawnee who seem to have been in firm possession of the
Platte valley.

Archeological findings leave no room for doubt that some at least
of the sites belonging to the Lower Loup Focus were inhabited
during a period when commercial intercourse was still comparatively
limited in volume. Moreover, the European beads and other ma-
terials so far studied from these sites, insofar as they can be dated,
appear to be of types used in the Indian trade not prior to the latter
seventeenth or eighteenth centuries. Finally, no early contact sites
have been found in the region, other than those belonging to this
complex, which could possibly be connected with the Pawnee or
which can be viewed as the residence of settled Indians in contact
with early traders. .

The historical background as here reviewed sheds significant light
on the contention that the Lower Loup Focus may represent some
group other than the Pawnee, not necessarily ancestral or even
related to them. In the latest published work on this complex, it is
suggested that “possible migration could account for the settling of
different peoples in the same locality.” Early in this discussion it
was pointed out that the village sites of the Lower Loup Focus,
although of comparatively restricted distribution, are both numerous
and very large. Moreover, since all those so far excavated have
consistently yielded limited quantities of copper, glass beads, and
(rarely) iron, it follows that they must have flourished for a time
after white influences had penetrated into their locality. Even
granting that all were not inhabited simultaneously, they undoubtedly
indicate the presence here in protohistoric times of a populous, firmly
established, and presumably potent ethnic group. Let us assume for
the moment that this group was not ancestral nor even related to the
Pawnee. We then have the somewhat difficult situation of a numerous
and powerful tribe, resident for many years (witness the innumerable
middens, earthlodge sites, etc.) in the very heart of the’ Pawnee
territory, clinging to it until after trade contacts had been established
with Europeans (circa 1650 or later), and then emigrating so un-
obtrusively and so completely that the Pawnee, who must have
followed closely on their heels so as to be firmly settled in the region
by Delisle’s time (1718), retained no tradition of their existence.
This would not only do violence to Pawnee traditions linking that
group with the protohistoric Lower Loup Focus, but would also
require an explanation for the apparent absence of any legends of

* Dunlevy, op. cit., p. 215.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

an earlier tribe, unrelated but with very similar culture, whom the
Pawnee could reasonably be thought to have displaced since estab-
lishment of European contacts. Such a theory, furthermore, would
presumably postulate a comparatively late incursion for the Pawnee,
which is at variance with the ethnographic indications. Pawnee ma-
terial culture of the nineteenth century, as has been stated, is pretty
clearly a composite based essentially on two distinct and funda-
mentally divergent economies—one horticultural and sedentary, the
other hunting and nomadic. The significant constituents of the former,
irrespective of their ultimate origin, are now known to have been well
established west of the Missouri in prehistoric times. Those of the
latter, in part rooted in the very remote past, were shared with
numerous other historic tribes of the Plains and particularly with
the western bison hunters. The Pawnee seem to have combined
the two in harmonious fashion, and so far as adjustment to environ-
mental and ethnic conditions goes, give no evidence whatever of
having been recent arrivals in the Nebraska region.

There are other clues. Dunbar has shown how the placement of
villages relative to one another has modified certain linguistic usages
in accord with local geography.” During the later years of their
residence in Nebraska there were seldom more than three or four
villages—in other words, usually one for each of the four bands.
At times two or more bands might occupy a single town, but the Skidi
seem always to have remained more or less aloof. Both Murie and
Grinnell present evidence supporting the view that subgroups within
each of the main bands formerly constituted separate villages.”
Murie credits the Skidi with 13 of these originally. This interesting
observation may partially explain the general tendency of the early
explorers to assign, usually from hearsay, as many as a score or
more towns to the Pawnee nation. Incidentally, too, it may have
archeological implications since the Pawnee locality abounds with
small and widely scattered precontact earthlodge villages which appear
to have a number of features in common with the later ones. The
sudden disappearance of the many small prehistoric villages and the
presence of a few very large fortified towns in protohistoric times
is an archeological puzzle which still awaits solution. Finally, the
mythology of the Pawnee is replete with local Nebraska place names
such as the Platte, the Loup, the Republican, Nemaha, and others.”
There are migration legends, to be sure, but none which afford any

* Dunbar, 1880, p. 251.
* Murie, 1914, pp. 549-556; Grinnell, 1893, pp. 231-230.
** Dorsey, 1906.

NO. 7 PAWNEE ARCHEOLOGY—WEDEL 19

proof of recent arrival. Three of the five “sacred places” of the tribe
were on the Loup and Platte within 50 miles of their junction; the
other two were in southern Nebraska and northern Kansas”; and
a number of their myths and tales relate directly to this neighborhood.

It must be apparent by this time that there exists little else than
academic grounds for questioning the presence of the Pawnee as
a firmly ensconced tribe in the Platte-Loup region since at least the
coming of the whites. The data of tradition, history, ethnography,
and mythology all support this inference. Moreover, the numerous
archeological similarities between the historic Pawnee and the earlier
Lower Loup Focus reflect essentially the same dual mode of life.
Viewed in the light of history, the differences in materials from the
two complexes are not so great as to strain the probability of a
common authorship. They involve details rather than fundamentals.
The greater richness, abundance, and variety of remains on the proto-
historic sites indicate a general level of cultural achievement far
above that of the historic Pawnee. If, as is very probable, this
superiority extends to the nonmaterial side of life as well, then the
protohistoric period may be regarded as the climax of social, cere-
monial, and political development in the Pawnee area. The culmina-
tion must have been reached before 1750. Thereafter came a steady
decline which left the nineteenth century peoples in possession of a
much simpler and clearly decadent cultural heritage, though the
recorded myths as well as many political and ceremonial survivals
hark back to the older and. better days. Such a regression is perfectly
in keeping with the contemporary history of the area: increased pres-
sure from hostile tribes, growing commercial intercourse and terri-
torial quarrels with the whites, new diseases, and a generally more
desperate struggle for sheer existence, all of which left scant
leisure for cultural advancement.

The leads for future research on this problem are very clear.
It is imperative first of all that thorough analyses be made of all
available archeological materials from sites of the Lower Loup Focus.
These should be carefully compared with similarly detailed studies of
collections and data from documented sites of the nineteenth century.
Needless to say, identities are not to be expected in all details, since
individual, village, and probably band preferences were undoubtedly
active factors. The element of time, too, must ever be borne in mind,
for over a period of two or three centuries considerable changes are
expectable. Another line of attack which has so far been totally

* Grinnell, op. cit., pp. 358-359.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

neglected in this connection is physical anthropology. Skeletal re-
mains either supposedly or certainly attributable to the Pawnee are
by no means plentiful, as the early cemeteries remain undiscovered,
and the later ones have suffered woefully at the hands of vandals.
There is a disturbing possibility that scaffold burial and subsequent
dismemberment may have been practised in the early period. Still,
careful examination of the material thus far recovered might further
illuminate the issue. For obvious reasons, it will probably never be
possible to prove empirically that the inhabitants of any one of the
Lower Loup Focus sites spoke a Pawnee dialect, since the individual
sites cannot be linked with recorded towns. Thus the identification
made on other grounds must remain a probability—a very high one,
it is true, but still a probability. To maintain from this that the sites
are not Pawnee, however, seems a captious argument, particularly
in face of the very strong circumstantial evidence in every other
respect. On the whole, it may be soundest and perhaps least confusing
to retain a nonlinguistic designation for these protohistoric remains,
‘at any rate for the present. For this purpose the term suggested by
Dunlevy and used in this paper is as appropriate as any.

SUMMARY AND CONCLUSIONS

In the foregoing pages the relationships between one historic and
two protohistoric archeological complexes in Nebraska have been
briefly discussed. These are respectively the Pawnee of the nineteenth
century, the Lower Loup Focus, and the Oneota Aspect. From the
evidence of archeology, history, tradition, mythology, and ethnography,
as outlined herein, the following major facts emerge:

(1) Village sites assignable to the Lower Loup Focus, 10 or more
in number, occur only in the very heart of the historic Pawnee region
about the confluence of the Loup and Platte Rivers.

(2) These sites nearly all yield limited amounts of historical ma-
terials, indicating their occupancy at least into very early contact
times.

(3) Historic maps and documents show that the Pawnee villages
since virtually the earliest contact times were localized in and about
this region.

(4) On the basis of available archeological evidence alone, sites
of the Lower Loup Focus show a much closer relationship to the
later historic Pawnee culture than they do to the contemporaneous
Oneota sites.

(5) Pawnee traditions link that tribe directly with several of the
protohistoric Lower Loup Focus sites.

NO. 7 PAWNEE ARCHEOLOGY—WEDEL 21

(6) Neither history, ethnography, nor recorded traditions offer
any proof that another sedentary horticultural tribe inhabited this
locality since the arrival of Europeans.

These six points sufficiently refute the objections so far raised
against identification of the Lower Loup Focus with the Pawnee
tribes. There is, therefore, no reason whatsoever for abandoning the
hypothesis outlined by Strong wherein the Lower Loup Focus is
considered a protohistoric phase of Pawnee culture.

LITERATURE CITED
BANDELIER, A. F.
1890. Contributions to the history of the southwestern portion of the United
States. Cambridge.
Bruce, RoBerr.
1932. The fighting Norths and Pawnee scouts.
BUTTERFIELD, C. W.
1881. History of the discovery of the Northwest by Jean Nicolet. Cincinnati.
Dorsey, G. A.
1906. The Pawnee Mythology, pt. 1. Carnegie Inst. of Washington.
Dunsar, J. B.
1880. The Pawnee Indians: their history and ethnology. Mag. Amer. Hist.,
vol. 4, no. 4, pp. 241-281.
Dun tevy, M. L.
1936. A comparison of the cultural manifestations of the Burkett (Nance
County) and the Gray-Wolfe (Colfax County) sites. Chapters in
Nebraska Archeology, vol. 1, no. 2, pp. 149-247. Univ. Nebraska.
GriFFIN, J. B.
1937. The archeological remains of the Chiwere Sioux. Amer. Antiquity,
vol. 2, no. 3, pp.. 180-181.
GRINNELL, G. B.
1893. Pawnee hero stories and folk-tales. New York.

Havyopen, F. V.
1868. Notes on Indian history, etc. Ann. Rep. Smithsonian Inst. 1867, pp.
AII-412.

Hirt, A. T., and WenveEt, W. R.
1936. Excavations at the Leary Indian village and burial site, Richardson
County, Nebraska. Nebraska Hist. Mag., vol. 17, no. 1.
Hotmes, W. H.
1903. Aboriginal pottery of the eastern United States. 2oth Ann. Rep.,
Bur. Amer. Ethnol., pp. 200-201 and pl. 177.

Murik, JAMES.
1914. Pawnee Indian Societies. Anthrop. Pap., Amer. Mus. Nat. Hist.,
vol. II, pt. 7.
Strone, W. D.

1935. An introduction to Nebraska archeology. Smithsonian Misc. Coll.
vol. 93, no. Io.
Tuomas, A. B. ;
1935. After Coronado. Univ. Oklahoma Press.
WEeEDEL, W. R.
1936. An introduction to Pawnee archeology. Bur. Amer. Ethnol., Bull. 112. -

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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLS 97a NOe ieee |

2

Scenes in the Pawnee village on the Loup River near Genoa, Nebr., in 1871.
This was the last northern settlement of the tribe prior to its final removal
to the Indian Territory circa 1875. (Photographs by W. H. Jackson.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOR 97 siNO revi cane

1. The Wright site near Genoa, Nebr., showing type of bluff top village location
preferred by the Pawnee in protohistoric times; Beaver Creek valley at
right. (Courtesy of the Nebraska State Historical Society. )

2. Excavated floor of protohistoric Pawnee earthlodge showing circular outline,
central firepit, postholes, and short vestibule doorway ; Wright site. (Cour-
tesy of the Nebraska State Historical Society. )

SMITHSONIAN MISCELLANEOUS COLLECTIONS Vere ON ore NO) erage a tenmiscs

1. Excavated floor of protohistoric Pawnee earthlodge at Larsen site, on Looking-
glass Creek; showing central firepit, surrounded by four primary and three
circles of secondary post molds. Note the peculiar arrangement of postholes
at the rear of the floor, opposite the entrance, where the family shrine was
traditionally placed. (Courtesy of the Nebraska State Historical Society.)

2. Excavated floor of late historical Pawnee earthlodge near Leshara, occupied
probably after 1850. This lodge had eight central roof supports, a raised
altar platform at the rear directly opposite the doorway, and a sill of baked
clay across the inner end of the entrance passage. Another house floor may
be seen in the background. (Courtesy of the Nebraska State Historical
Society. )

SMITHSONIAN MISCELLANEOUS COLLECTIONS WOT ES ity IN@e tha Lbs 2

1. Restored pot of late Pawnee type from Archer,
Nebr.; height 9 inches. (Courtesy of the
Nebraska State Historical Society. )

2. Restored vessel of protohistoric Pawnee type from the Wolfe
site near Schuyler; height 4% inches. (Courtesy of the
Nebraska State Historical Society. )

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nate Bek at; VOLUME 97, NUMBER 8

®

THE FAR NORTHWEST,
1649-1851
- oi (Win 18 PLATES)
“ BY
, DAVID I. BUSHNELL, JR.
(PUBLICATION 3485)

GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 30, 1938

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE oi NOsmoieieed

Pit on lei = 2G Ce:
in | age

Bll vi MOOR BEET ON

Uae ae aS

Senses TISH-RAWA VILLAGE, AND THE KLAMATH, BELOW THE ENTRANCE
OF THE SALMON’?

Drawn by Capt. Seth Eastman, from original sketch by George Gibbs,
October 1851.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 97, NUMBER 8

DRAWINGS BY GEORGE GIBBS IN
THE PAR NORTHWEST,
1549-185]

(WiTH 18 PLATES)

BY
DAVID I. BUSHNELL, JR.

(PUBLICATION 3485)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 30, 1938

The Lord Baltimore Preas

BALTIMORE, MD., U. % A.

iN)

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10.
Te:

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13.
14.
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IFEUSTRATIONS

PLATES PAGE

“The Tish-rawa village, and the Klamath, below the entrance of the
Salmon.” Drawn by Capt. Seth Eastman from original sketch by
George Gibbs. October 1851. (Frontispiece )

Shoshonee Falls of Snake River, August 15, 1849. 1. The Canon below

theytalllss, ¢2:ocbhe fall'sr:. Wchras bya toe ete ocean Eee ae 6
Ravine in mountains of Burnt River, Baker County, Oregon. Sep-

Rem De ee PTOA Om ceses onic oa aioe ciera Siere Pe Unt ORE ECE ETE TS ete 6
On the trail in Oregon. Ascending hill from Deschutes River. Octo-

WG Tape RT SAO ye sec ceed eave h ones oe Stay eae ay STA ATC LcL eR ePLs 6
Burial canoe on bank of the Columbia at mouth of La Camas. Creek.

@CLOWEIA SONOS Ops aock. et tote Ae ee EI Meee ee 6
Canoe on the Columbia River near Oak Point. October 1850............ 6

Chiefs of tribes in the valley of the Willamette. 1. Slacum. Chief of
tribe at the falls of the Willamette, probably the Clowwewalla. May
1851. 2. Joe or Alquema. Chief of the Santiam band of the Calapooya. 6

Valley of the Willamette. 1. Champoeg and French Prairie. April 1851.

2, Woe Willers ere Chennneces, Wileiy i8Sitcaesscoocacacccposccocc 10
At Oregon City, 1851. 1. Oregon City and the Falls of the Willamette.

2. Indians taking salmon at the falls of the Willamette. June 1851.. 10
Hudson’s Bay Company’s post at Fort Vancouver. July 1851.......... 10

At the Hudson’s Bay Company’s Post. 1. Fort Vancouver, July 2,
1851. 2. Catholic Chapel at Fort Vancouver, July 1, 1851. 3. Officers

Ouazterss ColambiaeBarracks a) iilye2 05 le eee eee eeneeeeee 10
Sononm, Caio, /NWGUSE Tig, WIS, sAons co ocoecsaconoeccedn saddens 14
Wpper end of Clear Lake, California, August 10, 1851...........-... 14
Redwood Tree, 52 feet in circumference. September 6, 1851........... 16

Sketches in the valley of the Klamath. 1. Woman and child, at junction
of Klamath and Trinity Rivers. October 6, 1851. 2. Young married
woman, at junction of Klamath and Trinity Rivers. October 6, 1851.
So Wows esi Selbiovorn Iesnveie, INiohneiaaloese 14, MIS cocoon encceneeucc 16
Views in the valley of the Klamath. 1. The Klamath above the fish
dam. October 9, 1851. 2. The Klamath. Signal tree of the Indians.
OCOD ETAT TSS iy) eee oo tae sisrsuntolaerey otek: icin Or aed ee 20
stherShastessutie ande/alleyas October 27. 1oSleneaenee see eee eee nee 20
Young Chief of the Weit-spek tribe, probably Mec-ug-gra. Drawn by
Capt. Seth Eastman from original sketch by George Gibbs. October
TS TSM ee es oP oneal wee Eee ES ae ie cite oe Se eee 20

TEXT FIGURES

Carvings in wood and bone from lower Columbia River............... 8
Wroodensbowlltromesohoalwater (Sayeman sameeren eee tena 9
Bow, hats, and headband from Klamath River, California............. 16
Basketseriomue ameathmavetan Calitot ita eereer ree erent ane 18

Spoons andetrays) trom Klamath Raver) Calionnias. see oseee ose eee 19

a a > are sere -Predt
sf ‘et ane 7
aa eae ae en Bya ae
AS oo a pete: 1 3s Oa + Sees,
Bao T Cet Cas ae" eee

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2

DRAWINGS BY GEORGE, GIBBS IN THE FAR
NORTHWEST, 1849-1851

By DAVID I. BUSHNELL, Jr:

(WitH 18 Prates)
INTRODUCTION

George Gibbs, whose drawings form the subject of this article,
was born July 17, 1815, at Sunswick, Long Island, New York, near
the present Astoria. He died in 1873. At the age of 17, failing to
receive the desired appointment to West Point, he accompanied an
aunt to Europe, where he devoted 2 years to travel and study. Return-
ing to New York, he soon entered Harvard, where he was graduated
in law in 1838, and subsequently entered the law office of Preston
Hall. However, the profession did not appeal to him, and during the
next few years he wrote several works on historical subjects which
were highly acclaimed.

The Far West was now becoming of interest, and the mystery of
the wilderness appealed to many. Gibbs was among those who were
thus attracted, and in 1849 he accompanied the Mounted Rifle Regi-
ment to Oregon, where he arrived early in October. That same
autumn he became deputy collector of customs at Astoria and was
later attached to the Indian Commission in Oregon. In 1851 he was
a member of the McKee party and visited the northwestern part of
California. During the journey he learned much concerning the
various native tribes with whom he came in contact, especially those
who were encountered in the valley of the Klamath. Later he settled
near Fort Steilacoom, Washington, where he devoted much time
to the study of the languages of the different tribes and prepared
extensive vocabularies which, together with brief lists of words, are
now in the Smithsonian Institution. Although interested primarily
in linguistic studies, Gibbs collected ethnological material, much of
which is preserved in the collections of the National Museum, being
among the earliest specimens gathered in the’ country beyond the
Rocky Mountains.

Gibbs was preparing to return to New York and so wrote to his
friend, Prof. S. F. Baird of the Smithsonian Institution: “North
West Boundary Survey, Fort Walla-Walla, Nov. 16, 1860. I arrived

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97, No. 8

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

bo

here yesterday on my way to Washington.” He next wrote to Prof.
Baird trom “261 ‘Greene Street, New York, Heb; 5.) 186m, ssoon
after he reached his home.

The correspondence between Gibbs and Professor Baird continued
through many years, both while Gibbs was in the west and after his
return to New York, and later when he lived in Washington. The
letters are most interesting, and many refer, in addition to the work
in which both were engaged, to places and persons now known only
in history.

WITH THE MOUNTED RIFLE REGIMENT TO OREGON, 1849

As previously mentioned, Gibbs accompanied the Mounted Rifle
Regiment to Oregon in 1849, being one of many civilians who
reached the valley of the Columbia at that time.” The regiment was
under command of Brevet-Col. W. W. Loring, and the expedition
started “with about 600 men, 31 commissioned officers, several women
and children, the usual train agents, guides, and teamsters, 160
wagons, 1,200 mules, 700 horses, and subsistence for the march to
the Pacific.”

An interesting account of the trip has been preserved; * it was
presented as (p. 126): “‘A report, in the form of a journal, to the
Quartermaster General, of the march of the regiment of mounted
riflemen to Oregon, from May io to October 5, 1849, by Major
O. Cross, quartermaster United States army.”

Excerpts from the journal will shed light on the dangers and
difficulties with which all were confronted:

Major Cross left St. Louis May 10, 1849, and ascended the
Missouri to Fort Leavenworth where he arrived g days later. “On
inquiring at the fort I learned that the troops were ten days in
advance of me, which was a very long start, as my mode of travelling
was the same as that of the regiment.” The next day he left for
Fort Kearny. ‘‘My outfit was as indifferent a one as ever left for
any station, much less the Rocky mountains.”

It is not known with which of the groups Gibbs was then traveling,
but he was probably with the troops that had left Fort Leavenworth
about the time Cross was departing from St. Louis.

‘Bancroft, Hubert Howe, The works of ... vol. 30, History of Oregon,
vol. 2, 1848-1888, p. 81. San Francisco, 1888.

*The report was made by Maj. Osborne Cross to Maj. Gen. T. S. Jesup,
Quartermaster General, and was incorporated in the report of the latter to
C. M. Conrad, Secretary of War. 31st Congr., 2d Sess., Senate Ex. Doc.
No. 1, pt. 2, Washington, 1850.

No. 8 DRAWINGS BY GEORGE GIBBS

BUSHNELL 3

Again the journal (p. 143): “June 5.—Large trains could be seen
this morning wending their way along on both sides of the Platte.
The river here is nearly three miles wide, interspersed with islands,
some of which are thinly covered with very small cottonwood and
willow.” That day the wagons, 160 in number, were overhauled and
many were repaired.

June 7— ‘To-day buffalo were seen for the first time, which
created no little excitement.”

June 19.—*“I visited Chimney Rock this morning, as the command
wended its way along the river.”

June 22.—Arrived at Fort Laramie. “Fort Laramie is situated
on Laramie’s creek, a rapid stream, about 60 yards wide, with a firm,
pebbly bottom. This stream rises among the Black Hills to the west,
and falls into the North Platte, about half a mile below the fort.

“This fort is built in the form of a quadrangular figure, and of
unbaked clay, or adobes; the wall is about twenty feet high, with
a small palisading on a part of it. There are two block-houses at
the corners, diagonally from each other . . . Over the main entrance,
which faces the river, there is also another small block-house. The
buildings are made inside, the wall forming a part of them.” There
were no trees about the fort. Game was formerly plentiful, but
“has greatly diminished since emigrants have made it the great
thoroughfare to Oregon and California.” Fort Laramie is 639 miles
from Fort Leavenworth.

August 1—‘It was at the side of the river, and at this place,
that I saw the celebrated spring generally known as the Steamboat
spring ... This place is immediately at the point where the two
trails turn off for California and Oregon, and within a short distance
Gmane Salt lake. 6...”

August 4—‘We descended a long hill, which brought us into a
sandy plain, which extends to Fort Hall, and on the banks of the
Ort NEL 2 4) ae”

August to—‘“Our encampment last .evening seemed to be the
terminus of Snake River valley, as the appearance of the river
entirely changed after a march of about five miles, which brought
us to the American falls . . . The scene was truly magnificent . . .”
Many rapids in the river, islands and masses of rock in the stream.

August 11—“We crossed Ogden’s river about 12 o'clock. The
road turns off to the south for California, which was taken by the
Californians who were still along i

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

August 15—A day of much interest for Gibbs. Left camp at 4
in the morning and, as the journal continues (p. 196) :

We travelled, however, rapidly for about eight miles ... until we arrived
at the creek again. At this place we waited for our wagons, which soon came
up; and, having assisted them out of the cafion, which was no easy work, we
continued on until the middle of the day, when we again came to the banks of
the river, which were at least two or three hundred feet in height. I attempted
to descend into the valley through which the river ran, for the purpose of
procuring water, but it was so fatiguing, both for myself and horse, that I
returned without being able to accomplish it.

It was at this place we could easily hear the sound of a waterfall, which,
from the noise, we at first supposed might have been the Little Falls of Snake
river; but, as we were still twenty miles from that point, we were soon
satisfied that it did not proceed from there, or the small cascade on the opposite
bank, which is mentioned by Colonel Fremont as the Subterranean river; and
we were much surprised to learn, the next day, that within ten miles of this
place there is a cascade, which, in height, is not surpassed by the Niagara
Falls. The guide, who was with the command, having travelled this route very
often, was shown the place by an Indian, and took Mr. Gibbs, of New York,
and Lieutenant Lindsay to the place, who pronounced it one of nature’s great
wonders. The river here becomes a little contracted, and passes through a
chasm of solid rock; it commences to fall about a quarter of a mile above the
last pitch, and, after forcing itself among loose rocks which lay in its way,
takes a perpendicular pitch of at least 160 feet, and it is even thought to be a
greater height. They descended to the foot of the falls, and after much difficulty
and some length of time, where they were better able to judge more accurately
of its great height; and there seems to be but one opinion, that it equalled in
grandeur, in proportion to the column of water, the Niagara Falls. Having
been the first who had ever taken the trouble to examine them carefully, and
wishing to change the name said to have been given by a priest many years
since, they decided on that of the Great Shoshonie falls, instead of Canadian,
as being the most appropriate.

The road does not pass there, and probably its nearest point is not less than
eight or ten miles, which is probably the reason why it is so little known, for
I have never seen it mentioned by those who have trapped in this country for
eas mene

We continued our journey until sundown, when we came to the foot of the
little falls on Snake river, commonly called the Little Salmon Falls, and
encamped for the night immediately on the banks of the river.

The drawings of the falls made by Gibbs that day are reproduced
in plate 2. This was the first mention of Gibbs by Major Cross.
The expedition continued through the mountains until (p. 210):

September 4—Mountains were to be seen all around, and it appeared a
mystery how we had extricated ourselves from those left behind us with so
little difficulty, or how we were to pass those ahead of us. This brought us
again on Burntwood creek, where we encamped for the night...

The ravine through which the Burntwood passes is too narrow to be culti-
vated, but the soil is rich and ought to yield well. The evening was spent in

NO. 8 DRAWINGS BY GEORGE GIBBS—-BUSH NELL 5

reaching the tops of some of the highest mountain hills, where the view of
the adjacent country well rewarded us for our trouble; a few scattering hem-
locks were seen in the ravine where we made our encampment, and the distant
hills and ravines beyond were interspersed with several groves of cedar and
pine. Our encampment lay in a fork formed by Burntwood creek and a little
DyOo kev atcha seimtoMmite es elles]

The party was now moving in several groups, and it is evident
that Gibbs was not always with Major Cross; this explains the
difference in the dates that often appear on the sketches made by
Gibbs from those of the entries in the journal.

On September 22, Gibbs made a sketch of the Columbia from the
mouth of Deschutes River. From this point the wagon train, with
which Gibbs must have been traveling, moved southward up the right
bank of the Deschutes River. During the morning of October 2,
the train ascended the steep cliff near the river. A sketch made
at that time reveals the wagons, each drawn by eight mules, form-
ing a long line extending from the camp at the foot of the cliffs to
the summit. It is an interesting drawing of a subject seldom recorded
(pl. 4).

Leaving the Deschutes River, the expedition passed through the
Cascade Range, and, on October 5, Gibbs made several sketches of
the forest scenery, to which he attached the legends: ‘‘Burnt forest
in Cascade Mts.,” and “Cascade Mts. Cedar & firs,’ and again on
October 9, “Forests of the Cascade Mts. Cedar & fir.”

The expedition had now arrived at its destination. Gibbs continued
on to Astoria where he became Deputy-collector of Customs, soon
to become attached to the Indian Commission.

ON THE COLUMBIA RIVER, AUTUMN OF 1850

Two drawings of exceptional interest, made by Gibbs on the banks
of the Columbia during the month of October 1850, are reproduced
in plates 5 and 6.

The first of these shows the
Columbia river, at mouth of Chamus Creek,” and is a beautiful
example of Gibbs’ work. Chamus Creek is believed to have been the

oe

Prow of dead Canoe on Bank of

stream now known as La Camas Creek, which flows into the Columbia
River near the southeast corner of Clarke County, Washington,
about 15 miles above Vancouver. This was within the Chinookan
country. Whether this canoe was placed on a scaffold or rested on

“Burnt River flows eastward and joins Snake River in the southern part of
Baker County, Oregon.

) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

the ground is not known, but as so little has been recorded concerning
the burial customs of the people of the region this sketch is of
special interest.

A brief reference to the strange form of burial was made by
Lieutenant Broughton, of the Vancouver Expedition,’ who explored
the lower Columbia during the autumn of 1792. He was near Cape
Disappointment, on the Washington side of the mouth of the
Columbia, and wrote (vol. 2, p. 54): “At this place was found
the remains of a deserted Indian village, and near it three large
canoes supported from the ground, each containing dead human
bodies. These canoe coffins were decorated at the head and stern
with rude carved work, and from their decayed state seemed to have
been thus appropriated for a great length of time.”

Soon the Lewis and Clark party reached the valley of the Columbia.
They encountered the same peculiar burials and left a more detailed
account of the manner in which the canoes were placed, and of
the various objects deposited in them.’ They stated (p. 429) :

The Chinnooks, Clatsops, and most of the adjoining nations, dispose of the
dead in canoes. For this purpose a scaffold is erected, by fixing perpendicularly
in the ground four long pieces of split timber. These are placed two by two,
just wide enough apart to admit the canoe, and sufficiently long to support its
two extremities. The boards are connected by a bar of wood run through them
at the height of six feet, on which is placed a small canoe, containing the body
of the deceased, carefully wrapped in a robe of dressed skins, with a paddle,
and some articles belonging to the deceased, by his side. Over this canoe is
placed one of a larger size, reversed, with its gunwale resting on the crossbars,
so as to cover the body completely. One or more large mats of rushes or flags
are then rolled round the canoes, and the whole secured by cords usually made
of the bark of the white cedar. On these crossbars are hung different articles
of clothing, or culinary utensils. The method practised by the Killamucks
differs somewhat from this; the body being deposited in an oblong box, of
plank, which, with the paddle, and other articles, is placed in a canoe, resting
on the ground.

Later accounts of the curious form of burial are to be found,
but the earlier descriptions are usually the more interesting. How-
ever, as remarked in the Lewis and Clark journal (p. 429), “Those
who first visit the ground, can only be expected to furnish sketches
rude and imperfect.”

* Vancouver, Captain George, Voyage of Discovery . . . 3 vols. London, 1798.
° Lewis and Clark, Travels to the source of the Missouri river and across
the American continent to the Pacific ocean... in the years 1804, 1805, and

1806. London, 1814.

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no. 8 DRAWINGS BY GEORGE GIBBS—-BUSHNELL 7

As previously mentioned, it is not known whether the canoe, the
prow of which was sketched by Gibbs, was placed on a scaffold
when in use or had always rested on the ground as shown in the
drawing. The drawing suggests that the prow was rather massive
and heavy, but there is no way to judge its size.

The Chinookan tribes who occupied both banks of the lower
Columbia excelled in carving wood and bone. On January 20, 1806,
when near the mouth of the Columbia on the south side, the Lewis
and Clark party were among the Clatsops with whom they maintained
a friendly intercourse. The narrative of the expedition refers to the
skill of the natives in making many articles used in and about their
houses, described as “large wooden buildings, varying in length from
twenty to sixty feet, and from fourteen to twenty in width.” The
narrative continues (p. 432):

They are... very dexterous in making a variety of domestic utensils,
among which are bowls, spoons, skewers, spits, and baskets. The bowl or
trough is of different shapes, sometimes round, semicircular, in the form of a
canoe, or cubic, and generally dug out of a single piece of wood, the larger
vessels having holes in the sides by way of handles, and all executed with great
neatness. In these vessels they boil their food, by throwing hot stones into
the water, and extract oil from different animals in the same way. Spoons
are not very abundant, nor is there any thing remarkable in their shape, except
that they are large and the bowl broad... The usual plate is a small mat of
rushes or flags, on which every thing is served.

Later, when the expedition was at the Cathlamah village, also on
the Columbia and not far from the Clatsops, certain customs of the
people were recorded in the narrative (p. 493):

This village we have already described, as situated opposite to the seal
islands: on one of these the Indians have placed their dead in canoes, raised on
scaffolds, above the reach of the tide. These people seem more fond of carving
in wood than their neighbours, and have various specimens of their taste about
the houses. The broad pieces supporting the roof and the board through which
doors are cut, are the objects on which they chiefly display their ingenuity,
and are ornamented with curious figures, sometimes representing persons in
a sitting posture supporting a burden.

Beautiful examples of the work of the people near the mouth of
the Columbia are shown in figure 1. Three of the carvings are in
wood and one in bone. The latter, a knife handle, has on the end a
remarkable representation of a raccoon, Procyon lotor, with the eyes
indicated by copper inlays. The club is made of cedar and is rather
light for the purpose indicated. All were collected by George Gibbs
probably in 1850 or 1851. Another bowl obtained by him in the
vicinity of Shoalwater Bay, on the coast a short distance north of

dS SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

the mouth of the Columbia, 1s reproduced in figure 2. Similar pieces
were undoubtedly seen by Lewis and Clark a generation earlier.
The second sketch made during the autumn of 1850, plate 6, bears
the legend “‘Columbia River near Oak Point, Oct. 1850.” The point
is on the right bank of the Columbia about midway between the mouth
and Vancouver, and was so named by Lieutenant Broughton in 1792.°
When going up the river they arrived at a spot “where, for the first

Fic. 1—Specimens collected by George Gibbs on the lower Columbia.

a, bone knife handle, length 84 inches, U.S.N.M. no. 708; b, club for killing
fish, wood, length 18 inches, U.S.N.M. no. 651; c, spoon, wood, length of figure
on handle 3% inches, no number; d, bowl, wood, diameters 6 and 8 inches,
U.S.N.M. no. 601.

time in this river, some oak-trees were seen, one of which measured
thirteen feet in girth; this obtained the name of Oak Point.”

The canoe is the most interesting feature of the sketch. To quote
again from Lewis and Clark’ (pp. 433-434):

The industry of the Indians is not confined to household utensils: the great proof
of their skill is the construction of their canoes. In a country, indeed. where so

* Vancouver, op. cit. vol. 2, p. 61.
“Op: (cit:

NO. 8 DRAWINGS BY GEORGE GIBBS—BUSH NELL 9

much of the intercourse between different tribes is carried on by water, the in-
genuity of the people would naturally direct itself to the improvement of canoes,
which would gradually become, from a mere safe conveyance, an elegant ornament.
We have accordingly seen, on the Columbia, canoes of many forms, beginning
with the simple boats near the mountains, to those more highly decorated,
because more useful nearer the mouth of the Columbia. Below the grand
cataract there are four forms of canoes: the first and smallest is about fifteen
feet long, and calculated for one or two persons: it is, indeed, by no means
remarkable in its structure, and is chiefly employed by the Cathlamahs and
Wahkiacums among the marshy islands. The second is from twenty to thirty-
five feet long, about two and a half or three feet in the beam, and two feet in
the hold. It is chiefly remarkable in having the bowsprit, which rises to some
height above the bow, formed by tapering gradually from the sides into a
sharp point. Canoes of this shape are common to all the nations below the
grand rapids.

Fic. 2—Wooden bowl collected by George Gibbs at Shoalwater Bay.
Diameters 84 and 103 inches, U.S.N.M. no. 602.

The other types of canoes, larger than those just described, need
not be mentioned. Evidently the canoe sketched by Gibbs belonged
to the second group mentioned by Lewis and Clark, those which were
“common to all the nations below the grand rapids.’ There is no
allusion in the early narratives to the use of sails and masts in the
native craft. The mast and sail shown in the sketch had been adopted
after contact with Europeans.

OREGON, 1851

A letter from the Acting Commissioner of Indian Affairs, dated
October 25, 1850, addressed to J. P. Gaines, A. H. Skinner, and
Beverly S. Allen, stated* (p. 114):

*In Annual Report of the Commissioner of Indian Affairs... 1850. Wash-
ington, 1850.

10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

“Gentlemen: I have been officially notified of your appointment as
‘Commissioners to negotiate treaties with the several Indian tribes
in the Territory of Oregon, for the extinguishment of their claims
to lands lying west of the Cascade Mountains, under the act of 5th
June last’; and am directed by the Hon. Secretary of the Interior to
prepare appropriate instructions for your observance in the discharge
of the duties of your office.” The region was briefly described, the
tribes were mentioned in a vague manner, and the letter then con-
tinued: “It will probably be best for you to treat first with the
Indians in the white settlements, particularly in the Waullammette
Valley—and to treat separately with each tribe .

Evidently the three commissioners were active during the ensuing

9

months. In a joint communication to the Commissioner of Indian
Affairs, dated Champoly, April 19, 1851,’ they transmitted (p. 205) :
“a treaty concluded, on the 16th instant, with the Santian band of
the Callapooya tribe of Indians, by which they cede to the United
States a portion of the Willamette valley, about eighty miles in
length and about twenty in width. And also a treaty, concluded this
day, with the Twallalty band of the same tribe, including a country
about fifty miles in length and about thirty in width... Their
numbers are, of the Santian band, 155, and of the Twallaltys, 65.”

Gibbs was associated with the Commissioners when the treaties
were made.

Among the Gibbs material in the Bureau of American Ethnology,
Smithsonian Institution, is a manuscript designating where and when
he prepared many of the vocabularies. One record is of interest at
this time as it refers to events at Champoeg in April 1851: ”

Kalapuya.—My own vocabulary of this language was obtained April 4, 1851,
while the Commission was engaged in a treaty with them at Champoeg. It
is of the Si-yam-il, or as generally called Yamhill band, living on the river of
that name, which empties into the Willamet from the coast range. The Twallatys
(Twalati), and the Luckamukes (Luk-a-mai-yuk) speak the same dialect. The
Santiam band, on the east side of the Wilamet, a rather different one. It was
given by Thomas and Antoine, Chiefs.

Molele—Obtained at the same place. This was received from an Indian of
the band inhabiting the upper waters of the Santiam.

Many drawings were made at this time, four of which are now
reproduced. Others show different parts of the valley as it appeared
during the spring of 1851.

*In Annual Report of the Commissioner of Indian Affairs ... 1851. Wash-

ington, 1851.
2 Bur. Amer. Ethnol. Manuscript Catalog No. 742.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES Sit, NO 8, beans

tantédg sf

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1. Champoeg and French Prairie, April 1851.

2. The Willamette River at Champoeg, May 1851.
VALLEY OF THE WILLAMETTE

SMITHSONIAN MISCELLANEOUS COLLECTIONS vol Siig NOS yn len

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Oregon City and the Falls of the Willamette.

Sees eS, JS

2. Indians taking salmon at the Falls of the Willamette, June 1851.
AT OREGON CITY, 1851

WAGIE 7/q INOS fly TAL io)

SMITHSONIAN MISCELLANEOUS COLLECTIONS

’S POST AT FORT VANCOUVER, JULY 1851

.

HUDSON’S BAY COMPANY

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO oii NOm om rls tid

Hidrrms bogbo Hevk, Sirs eeuortt
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1. Fort Vancouver, July 2, 1851.

Catherine Cherfol at #} bansrned.
pity th LOS).

2. Catholic Chapel at Fort Vancouver, July 1, 185r.

3. Officers’ quarters, Columbia Barracks, July 2, 1851.
AT THE HUDSON’S BAY COMPANY’S POST

NO. 8 DRAWINGS BY GEORGE GIBBS—-BUSHNELL en

Portrait sketches of two chiefs are given in plate 7, one being
that of a chief of the Santiam band of the Callapooyas, but not one
who had contributed to the vocabulary. This is well drawn, and the
sloping forehead reveals the effect of artificial flattening. It was
probably a good likeness.

The second portrait is that of “Slacum, Chief of tribe at Falls of
Willamette (Upper Chinooks),’ and was drawn a fev days after
the treaty was made with the Callapooya. Slacum may have been a
chief of the Clowwewalla, belonging to the Chinookan family, a
tribe that occupied the region bordering the falls of the Willamette
River, the site of Oregon City. The name Slacum was probably
derived from that of an American naval officer who visited the
region in 1836 “to obtain information in relation to the settlements
on the Oregon river.” " He prepared an interesting, although brief,
account of the native tribes then living on the Willamette, part of

11

which follows. When ascending the river:

The first tribe of Indians are the Kallamooks, on the left bank, on a small
stream of the same name, 30 miles from its mouth: 2d are Keowewallahs, alias
Tummewatas or Willhametts. This tribe, now nearly extinct, was formerly
very numerous, and live at the falls of the river, 32 miles from its mouth, on
the right bank. They claim the right of fishing at the falls, and exact a
tribute from other tribes who come hither in the salmon season (from May
till October). Principal chiefs deceased. This river at the present day takes
its name from this tribe. 3d. ‘“Kallapooyahs” occupy lodges on both sides of
the river. 4th. “Fallatrahs” on a small stream of same name, right or west bank.
sth. Champoicho—west bank. 6th. Yamstills—west bank. 7th. Leelahs—both sides.
8th. Hanchoicks. All these 5 tribes speak Kallapooyah dialects, and are doubt-
less of that tribe, but at present are divided as designated, and governed by
chiefs as named. All these tribes do not exceed 1,200.

The Willamette was a beautiful stream, as Slacum wrote, “even
in midwinter, you find both sides clothed in evergreen, presenting
a more beautiful prospect than the Ohio in June . . . On the right
the land rises gradually from the water’s edge, covered with firs,
cedar, laurel, and pine. The oak and ash is at this season covered
with long moss, of a pale sage green, contrasting finely with the
deeper tints of the evergreens.”

Gibbs appreciated the beauty of the region and revealed it in two
sketches reproduced in plate 8.

The first is a view of “Champoeg and the prairies beyond,’
looking over the Willamette; the second shows the banks of the
stream with the variety of trees and shrubs, with spring foliage.

" Slacum, William A., Memorial of ... praying compensation for his ser-
vices . . . 25th Congress, 2d Session, Senate Doc. 24. Washington, 1838.

12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Champoeg was on the right bank of the Willamette, at the northern
end of French Prairie, the origin of which name was explained by
Bancroft,” who wrote (pp. 70-73) :

As their terms of contract expired, the Hudson’s Bay Company began to
retire its servants, giving them choice lands not too far removed from its
benign rule. This was the origin of the French Canadian settlements in the
beautiful Valley Willamette ... French Prairie, the tract where the servants
of the fur company began their planting in the Willamette Valley, extended
from the great westward bend of that river south to Lac La Biche about
twenty-five miles ... The landing at the crossing of the Willamette on the
east side was known as Campement du Sable, being a sandy bluff and an
encampment at the point of arrival or departure for French Prairie. Two
miles above this point was Champoeg, the first settlement.

The falls of the Willamette, when surrounded by the primeval
forest and in its natural condition, was a place of great beauty. And
as it was here that many Indians from the scattered villages would
come to get their supply of salmon, it was likewise a place of great
importance to the native inhabitants of the valley. But about the
year 1840 a settlement was begun at the falls, Oregon City, and soon
all was changed, although the few remaining Indians continued to
take salmon at the falls, as others had done through generations.

Two sketches by Gibbs, reproduced in plate 9, show the falls, and
Oregon City as it appeared in June 1851. An Indian is portrayed
spearing fish from a canoe, another is seen standing on a fishing
stage, in the right center of the sketch, using a net at the foot of the
falls. Fish are also shown leaping from the water. The upper draw-
ing is a view of Oregon City with the falls just beyond.

Oregon City was visited by Major Cross on October 5, 1849,
after the completion of the trip from Fort Leavenworth” and was
described in these words (pp. 227-228):

The city of Oregon is not a very prepossessing place in its appearance, for,
like all new places in the western country, the stumps and half-burnt trees lie
about in every direction. It is immediately at the Willamette Falls, hemmed

in by the river in front, and a ledge of rocks immediately in rear and very
close to the city.

Leaving Oregon City, Gibbs evidently continued down the Wil-
lamette and next visited Fort Vancouver, which had been erected
during the years 1824-1825. Sketches of the fort, and one of
Columbia Barracks a short distance away, are shown in plates 10, IT.

“Bancroft, Hubert Howe, The works of ... vol. 29, History of Oregon,
vol. 1. San Francisco, 1886.

“Op. cit. Jn 31st Congr., 2d Sess., Senate Ex. Doc. No. 1, pt. 2, Washington,
1850.

No. 8 DRAWINGS BY GEORGE GIBBS—-BUSHNELL rs

A concise description of Fort Vancouver, printed in 1840,"
explains many of the details of the drawings. To quote (pp. 19-20) :

On the north side of the Columbia, and a quarter of a mile from it, stands
Fort Vancouver, the principal establishment of the Hudson’s Bay Company
west of the Rocky Mountains. It consists of a number of wooden buildings
within a stockade, serving as dwelling-houses, stores, magazines, and work-
shops; and near it are other small buildings inhabited by the laborers, together
with a saw-mill and grist-mill. The whole number of residents at the place
is about eight hundred, of whom a large proportion are Indians or half-breeds.
Several hundred acres of land near the fort are under cultivation, producing
wheat, barley, oats, pease, potatoes, &c., in abundance; and the stock of cattle
is also considerable.

It was a place of great activity, surrounded by many tribes who
spoke different languages and had strange manners and ways of life.

Maj. Osborne Cross mentioned Fort Vancouver in his journal: ”

Fort Vancouver, which is the headquarters of the Hudson’s Bay Company,
is on the right bank of the river. It is situated on a beautiful plain, about five
miles long, and probably is three quarters of a mile wide. The country gradually
rises, and runs back for ten or fifteen miles, passing through several plains,
some of which are cultivated. On one of these plains there is an excellent
seminary, where the children from the fort and the neighborhood are educated.

Immediately in rear of the fort, and on the rising ground, the company of
artillery under Brevet Major Hatheway have put up temporary quarters, and
have made themselves very comfortable.

The latter became Columbia Barracks, and the temporary quarters
were soon replaced by others of a more permanent nature. The
Officers’ Quarters at the barracks, as they appeared 2 years later,
were sketched by Gibbs July 2, 1851 (pl. 11, fig. 3). At that time
they formed an attractive group of buildings facing Fort Vancouver,
with the Columbia beyond, while a short distance in the rear was
the edge of the forest which extended off to the north.

Gibbs did not remain many days in the vicinity of the fort, but
turned southward to California where he joined the McKee party
and soon set out to explore the northwestern part of the State and
to visit the many native tribes some of whom may never before
have come in contact with the white man.

IN NORTHWESTERN CALIFORNIA, 1851

The journal of the expedition into northwestern California, pre-
pared by Gibbs and later mentioned by McKee in his letter of March

™ Greenhow, Robert, Memoir, historical and political, on the Northwest coast
of North America ... 26th Congr., Ist Sess., Senate Doc. 174. Washington,
1840.

oe Opacity pa 220:

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

13, 1852, was published by Schoolcraft the following year.” It is
a valuable account of a journey through a part of the country never
before carefully studied, and describes briefly the native inhabitants
of the rough, mountainous region who occupied secluded valleys in
the vicinity of the rivers, often difficult to discover.

Pencil sketches made by Gibbs of scenes along the route reveal
much of interest and beauty encountered in the wilderness and are
now reproduced for the first time. Statements in the journal which
tend to describe or explain the drawings will be quoted, although
much of equal value, but not referring to the sketches, must neces-
sarily be omitted.

This will be followed by excerpts from McKee’s account of the
expedition.

JOURNAL OF THE EXPEDITION OF COLONEL RepicK M’KeErE, UNITED STATES
InpIAN AGENT, THROUGH NortTH-WESTERN CALIFORNIA. PERFORMED
DurING THE SUMMER AND I*ALL or 1851. By GrEorRGE GIBBS.

Monday, Aug. tr——Colonel M’Kee and party, escorted. by Major Wessells,
and a detachment of thirty-five mounted riflemen, left Sonoma this morning,
and moved over to Santa Rosa, encamping a little beyond Carillo’s ranch...
The general route proposed to be followed by the expedition, was up Russian
river to its sources, down Eel river to Humboldt bay, and thence over to the
Klamath, ascending that to the neighborhood of Shasté Valley, should the season
Meron, (Jel, 14]

Continuing northward, the party soon reached Clear Lake where
they remained several days. Large groups of Indians assembled,
and a treaty was entered into. “In personal appearance, many of
the Clear Lake Indians are of a very degraded caste; their fore-
heads naturally being often as low as the compressed skulls of the
Chinooks ... A vocabulary of this language was obtained from
the Indian who accompanied us, and who spoke Spanish sufficiently
to be enabled to interpret with his people.” On the next day,
August 19, the proposed treaty was explained to the assembled
Indians. A region of great natural beauty (p. 109):

Surrounded on every side by mountains, this valley is completely isolated
from the adjoining country, there being no access except by difficult trails...
The principal valley upon the lake is that upon which we encamped, lying on
the western side, and extending from mount M’Kee towards the head. The

‘extent of this may be stated at ten miles in length, by an average width of
four. A more beautiful one can hardly be pictured. Covered with abundant

** Schoolcraft, Henry R., Information respecting the history, condition and
prospects of the Indian tribes of the United States, pt. 3, pp. 99-177. Phila-
delphia, 1853.

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SNOILO3S11090 SNOSNVITS0SIN NVINOSH_LIWS

No. 8 DRAWINGS BY GEORGE GIBBS—-BUSHNELL 15

grass, and interspersed with groves of superb oaks of the most varied and
graceful forms, with the lake and its green margin of tulé in front, and the
distance bounded everywhere by precipitous ranges, it combines features of
surpassing grandeur and loveliness. Flowers of great variety and elegance
abound, the woods are filled with game, and in the season innumerable flocks
of water-fowl enliven the shores. [Pl. 13.]

Wednesday, Aug. 20-—The council was again assembled, and the treaty
explained to them as engrossed . . . As regards the suitableness of the reserva-
tion for its purpose, there can hardly be a doubt. The spot is isolated to a
degree unusual even on the Pacific; abounds in all that is necessary for a large
number of people in their savage state, and is capable of being made in the
highest degree productive by cultivation.

Saturday, Aug. 30—This valley, called by the Indians Ba-tem-da-kai, we
supposed to be on the head of the south fork of Eel river, and so were informed
by our guide and other mountaineers; but a belief exists, as we afterwards
found ... that it is, on the contrary, the head of the river before spoken of
as entering the coast to the westward ... A few Indians visited us, and were
directed to call in the adjacent tribes.

The entire party remained in camp the following day.

Sunday, Aug. 31—Quite a number of Indians were assembled and presents
distributed, but no treaty attempted; for our Clear Lake interpreter, although
able to comprehend them, could not explain freely in turn. Their language,
however, is clearly of the same family as that of the tribes at the head of
Russian river, and those last encountered. The total number in the vicinity, as
near as could be ascertained, was about six hundred souls . . . They pluck their
beards, and some of them tattoo. Many had their hair cut short, but others
wore it turned up in a bunch in front, or occasionally on the back of the head
... The average height of these men was not over five feet four or five
inches ... We saw no women...

I took the opportunity of to-day’s halt, to ascend the hills on the eastern
side of the valley. The view from this point was beautiful, the stream winding
in serpentine form along the margin of the plain, fringed with oaks and firs,
and the long slopes beyond diversified with forest and prairie. To the east
rose heavy ranges of mountains, between which and the yet more distant
Sacramento chain, a wide and deep gap indicated another valley, supposed to
be the source of the main fork of Eel river.

The next day the trail led through a mountainous section, “crossing
deep arroyas and then ascending a broken ridge between the waters
of the south and middle forks [of Eel river].”” Some Indians were
encountered who “had robes of deer skin, dressed with the hair on,
over their shoulders. They belonged to a wild mountain tribe, the
terror of the valley Indians ... Of their language and affinities,
nothing is known.”

September 5.—The trail crossed the river and passed a grove of
redwoods. During the day a few Indians were encountered, and
Cp aek23))0:
two or three of them were of larger stature than usual, and one was really
a fine-looking young fellow. They wore the deer-skin robe over the right

3

10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

shoulder, and carried the common short bow, backed with sinew la, fig. 3],
and arrows pointed with stone, both tolerably well made. With all these
Indians, the arrow-points are fastened into a short piece of wood, which in
turn is fixed, though but loosely, into the shaft. The quiver, of dressed deer-
skin, holds both bow and arrows. They had also, suspended round the neck,
small nets, neatly made after the fashion of the common game-bag; the twine,
which was very even, being of course their own work.
The last part of our march led us into a thick redwood forest.

Frc. 3—Specimens collected by-George Gibbs on the Klamath River, California.

a, sinew-backed bow, length 34 inches, U.S.N.M. no. 649; b, basketry hat,
diameter 7 inches, depth 4 inches, U.S.N.M. no. 7556; c, basketry hat, diame-
ter 73 inches, depth 4 inches, U.S.N.M. no. 7558; d, headband, U.S.N.M.
no. 7520; ¢, two sections of d, X approximately 2.5.

Saturday, September 6.—Indians visited the camp but they were
of little interest, and (p. 124):

I endeavored in vain to get from them the names of articles at hand, parts
of the body, &c., as they either could not or would not understand the object
of the inquiry; nor was our Clear Lake Indian more successful after his
method...

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SNOILO31100 SNOSNVITISOSIN NVINOSHLIWS

no. 8 DRAWINGS BY GEORGE GIBBS—-BUSHNELL 7

Our camp was a very pretty one, the little prairie being level and rich,
and encircled by a magnificent redwood forest. One tree near the tents I
measured, and found it to be fifty-two feet in circumference, at four or five feet
from the ground, and this although the bark and a portion of the wood were
burned away ... [Pl. 14.]

From September 29 until the morning of October 9, the party
occupied a camp established at the junction of the Klamath and
Trinity Rivers. Gibbs did not make a separate entry for each day
spent at the camp, but between the days mentioned devoted much
time to the study of the Indians with whom he was in contact. Many
tribes were represented at the gathering, possessing similar manners
and ways of life. To quote briefly (p. 139):

With regard to their form of government, at least that of the Klamath and
Trinity tribes, the mow-ce-ma, or head of each family, is master of his own
house, and there is a sci-as-lau, or chief, in every village . .. The lodges of
these Indians are generally very well built; being made of boards riven from
redwood or fir, and of considerable size, often reaching twenty feet square. The
roofs are pitched over a ridge-pole, and sloping each way ; the ground being usually
excavated to the depth of three or four feet, and a pavement of smooth stones
laid in front. The cellars of the better class are also floored and walled with
stone. The door always consists of a round hole in a heavy plank, just
sufficient to admit the body; and is formed with a view to exclude the bears,
who in winter make occasional and very unwelcome visits.

The people were described as being superior to any previously met,

and with countenances denoting greater force and energy of character, as well

as intelligence . . . The superiority, however, was especially manifested in the
women, many of whom were exceedingly pretty; having large almond-shaped
eyes, sometimes of a hazel color .. . their only dress the fringed petticoat, or

at most, a deer-skin robe thrown back over the shoulders, in addition. The
petticoat with the wealthier, or perhaps more industrious, was an affair on
which great taste and labor were expended. It was of dressed deer-skin; the
upper edge turned over and embroidered with colored grasses, the lower cut
into a deep fringe, reaching nearly to the knee, and ornamented with bits of
sea-shell, beads, and buttons ... The same round basket-cap noticed before,
is worn by the Klamath women [b, c, fig. 3], figures of different colors and
patterns being worked into it. They tattoo the underlip and chin in the manner
remarked at Eel river; the young girls in faint lines, which are deepened and
widened as they become older, and in the married women are extended up
above the corners of the mouth... The children are carried in baskets
suspended from the head, after the manner shown in the sketch.

The original sketch to which this refers is reproduced in plate 15,
figure 1. A picture of a “Young married Woman,” also made at
the forks of the Klamath and Trinity Rivers, October 6, 1851, is
reproduced in plate 15, figure 2.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

Continuing (p. 141):

The dress of the men consists, generally, of a pair of deer-skins with the
hair on, stitched together. Sometimes, however, a noted hunter wears a couple
of cougar skins, the long tails trailing behind him; and other again, on state
occasions, display a breech-clout of several small skins, sewed into a belt or
waistband .. . They are not as skilful in the preparation of dressed skins as
the Oregon Indians, and the use of those dressed on both sides is mostly
confined to the women.

Fic. 4—Specimens collected by George Gibbs on the Klamath River, California.

a, food bowl of twined basketry, diameter 104 inches, depth 23 inches,
U.S.N.M. no. 7568; b, food bowl of twined basketry, diameter 114 inches,
depth 3 inches, U.S.N.M. no. 7563; c, cooking basket, diameter 94 inches,
depth 4 inches, U.S.N.M. no. 7567; d, cooking basket, diameter 9+ inches,
depth 45 inches, U.S.N.M. no. 7553.

On the morning of October 9 the expedition left the mouth of
the Trinity and continued up the valley of the Klamath. About
2 miles above the forks they arrived at—
the Hai-am-mu village, and visiting one of the lodges, found the inhabitants

engaged in cooking and eating. The meal consisted of fish and acorn porridge,
made by mixing the flour in a basket [c, d, fig. 4], in which the water is kept

no. 8 DRAWINGS BY GEORGE GIBBS—-BUSHNELL 19

boiling by means of hot stones. Of the acorn flour they likewise make a
sort of bread, which they bake in the ashes. They had several spoons, very
neatly made of bone or horn [a, b, fig. 5]. At this village there was a large
fish-dam; a work exhibiting an extraordinary degree both of enterprise and
skill . . . We camped opposite the high point which forms a land-mark from
the Bald Hills, and which gives the name Bluff creek to a stream entering
from the northwest, called by the Indians Otche-poh. Upon the other side of
the river was an Indian village, the Sehe-perrh; the first belonging to the

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Fic. 5.—Specimens collected by George Gibbs on the Klamath River, California.

spoon made of elk antler, length 54 inches, U.S.N.M. no. 7510; b, spoon
made of elk antler, length 6 inches, U.S.N.M. no. 7511; c, shallow food tray,
diameter 7 inches, depth 2 inches, U.S.N.M. no. 7561; d, shallow food tray,
diameter 103 inches, depth 145 inches, U.S.N.M. no. 7562; e, tray for draining
liquid from food, diameter 12 inches, depth 245 inches, U.S.N.M. no. 7562.

tribe occupying the middle section of the river, and of which the Quoratem
or Salmon river Indians may be considered as the type. [PIl. 16, fig. 1.]

The party encamped on the bank of the Klamath about 1 mile
above the mouth of Salmon River and there remained from the after-
noon of October 11 until the morning of the 13th. This was a rugged,
mountainous region through which it was difficult to pass.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

P. 150.—Salmon river, or as it is called by the Indians, the “Quoratem,” is
the largest of the affluents of the Klamath, with the exception of the Trinity .. .
Upon the Klamath, both above and below the junction, are Indian villages of
some size, prettily situated on high platforms of rock projecting over the
water, and shaded by groves of oaks and bay trees; while below, the river,
compressed in its channel, rushes boiling over rapids. The accompanying
sketches were taken, one from near our camp, representing the Tish-rawa
village, and the Klamath, below the entrance of the Salmon [pl. 1]; the other
from a mile higher up, showing the course of the Klamath through the
mountains above the forks [pl. 16, fig. 2]. The tree on the right hand of the
latter represents one of the signal or “telegraph” trees of the Klamath Indians.
These, which are among the most conspicuous features of the scenery upon
the river, occur near every village. They are always selected upon the edge
of some hill, visible to a considerable distance in either direction. Two trees,
one trimmed in the form of a cross, the other with merely a tuft on the top,
represent each lodge; and in time of danger or of death, a fire kindled beneath
them, informs the neighboring tribes of the necessity or misfortune of its
occupants.

Scott’s Valley, with “the snowy peak of Shasté lying to the south-
east, towering above all,” was reached on the afternoon of October 21.
During the following days Gibbs, with other members of the party,
made trips to the surrounding country to examine the valley and to
contact the native tribes. On Sunday, October 26, he rode to Shasté
Butte City, some 25 miles from the camp, and the next morning
(p. 165):
rode to the top of a range of hills about four miles distant, for the purpose
of obtaining a view of the country. The prospect here was very extensive,
commanding the northern and eastern portions of the plain, and extending
south-easterly, to Mount Shaste, which was distant about thirty miles . .
From the same point of view we could see Mount Pitt, or more properly Piit
mountain, so called from the traps formerly dug near it, by the Indians; and
the noted land-marks of the Oregon trail, the “Pilot Knob,” on the Siskire
range to the north, and the “Black or Little Butte,’ to the south. [PI. 17.]

Members of the party, including Gibbs, started from Scott's
valley camp about noon on November 6 to return to San [rancisco.
They passed over the same trail to their old camp on the Klamath,
near the mouth of the Salmon, where they arrived on the 12th.
Pheres((p.5174)<
several of our old acquaintances among the Indians visited us; and I succeeded
in persuading a pretty girl, the chief's daughter, to sit for her portrait. The
likeness was sufficiently good to be recognised, though it certainly did not
flatter the very gentle and pleasing expression of her face ... [pl. 15, fig. 3].
We found the Indians of the village which had been burnt down, rebuilding
their houses for the winter. The style was very substantial, the huge poles
requiring five or six men to lift. These lodges, it may be mentioned, are
usually dismantled in summer, when the inhabitants live in temporary bush huts.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES 9i7,, (NOs 8 BE) 16

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2. The Klamath. Signal tree of the Indians on right. October 13, 1851.
VIEWS IN THE VALLEY OF THE KLAMATH

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SNOILO31100 SNOANVIISOSIN NVINOSHLIWS

No. 8 DRAWINGS BY GEORGE GIBBS—-BUSHNELL 21

After encountering many difficulties and delays, Gibbs arrived
at San Francisco December 28, 1851, “having been absent on the
Expedition nearly five months.”

The specimens collected by Gibbs on the Klamath River in
California, which are now in the United States National Museum
and many of which are shown in figures 3, 4, and 5, are believed
to have been obtained by him during the autumn of 1851. Several
of the pieces represent forms of objects mentioned specifically in
his journal as being used by the Indians who were met during the
journey up the valley of the Klamath, and these may have been
collected at that time. All are excellent examples and were evidently
chosen with care.

The bow, made of yew, is strengthened on the back by sinew
which, although originally glued to the wood, has become detached.
The cord, as shown in the photograph, 1s 34 inches in length.

The headband, d, figure 3, is described in the old list of specimens
as “a ribbon for the hair,’ and is an unusual piece to have been
preserved. If extended it would be about 20 feet in length, but it
is arranged in 15 coils, held together at one place by an end being
passed around the coils several times and fastened. The average
width is 4 inch, the average thickness about 3/g inch. As shown
in the enlarged photograph, e¢, figure 3, it is formed of two twisted
cords of: vegetal fiber, each cord being made of two strands which
are loosely twisted. The two cords of fiber are held together, parallel,
by a light-colored grass, braided so as to allow the cords to remain
separated. At irregular intervals the braided grass is omitted, thus
allowing the two twisted cords to be exposed. There is a small loop
at one end of the cords.

All baskets, basket hats, and trays are beautiful pieces and may
be the oldest existing examples from that part of California.

The shallow tray, no. 7561, d, figure 5, is reenforced on the
bottom by the attachment of pliable twigs or shoots, probably of the
willow, which are bent to form circular bands, each band or ring
being composed of two coils. Two such bands were used, one
rather small and the other near the outer edge.

The two small spoons, a, b, figure 5, are typical examples and
show the effect of much use.

tbo
to

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

MINUTES KEPT BY JOHN McKEE IN NORTHWESTERN
CALIFORNIA, 1851
On October. 15, 1850, a letter was addressed to Redick McKee,
Geo. W. Barbour, and O. M. Wozencraft, by the Acting Commis-
sioner of Indian Affairs, Washington, which read in part “ (p. 121):

Gentlemen: I have the honor to enclose herewith a copy of a letter from
the Secretary of the Interior, by which you will find that your functions and
salaries as Indian agents are suspended; and that you are appointed, with the
sanction of the President, commissioners “to hold treaties with various Indian
tribes in the State of California,” as provided in the Act of Congress, approved
Sept. 30, 1850. Your commissions are also enclosed.

The three newly appointed commissioners met and entered into
treaties with various tribes. They had worked together, but on
May 15, 1851, wrote from Camp Barlow, San Joaquin River, to the
Commissioner * (p. 224) :

We have now concluded, in view of the almost interminable extent of
country to be traversed in carrying out our instructions, to cease as a board,
and address ourselves to the work individually. We have made a temporary
division of the State into three districts, for the purpose of negotiating treaties
with the various tribes, upon the general plan submitted in our joint letter of
1toth March. For our respective districts, lots were drawn to-day, and the
northern fell to the writer (R. MckKee,) the middle to O. M. Wozencraft, and
the southern to George W. Barbour.

With the coming of immigrants, prospectors, and settlers, unrest
and uncertainty developed among the Indians of northern California,
and a general uprising was feared. As a result of the existing con-
ditions it was deemed necessary for a commission to visit the various
tribes in the endeavor to create friendship, to enter into treaties, and
to set apart reservations where all could live in peace and security.
Many of the scattered tribes, or groups, occupied secluded valleys,
reached by narrow trails over mountains or through rocky ravines
and often difficult to discover; nevertheless, the venture proved of
much importance.

Redick- McKee led the Expedition, and on July 29, 1851, wrote
from San Francisco to Luke Lea, Commissioner of Indian Affairs,
Washington, saying in part” (p. 128):

“In Annual Report of the Commissioner of Indian Affairs . . . 1850. Wash-
ington, 1850.
“In Annual Report of the Commissioner of Indian Affairs . . . 1851. Wash-

ington, 1851.
“The journal or Minutes kept by John McKee, secretary, on the expedition
from Sonoma, through northern California, and many letters pertaining to the

NO. 8 DRAWINGS BY GEORGE GIBBS—-BUSHNELL 23

At present I can form no satisfactory estimate of the amount of appropriations
our treaties will require ... The largest estimate will fall below the cost of
a Califorma Indian war, if one should unhappily become general... I am
anxious to get off, so as, if possible, to make the journey before the rainy
season sets in. I propose taking the Clear Lake Indians on my route, who are
said to number three or four thousand, and will endeavor to make pacific
arrangements with them before passing over the coast range. On the Klamath
and Trinity rivers, from all the accounts I have received, I shall find large
bodies of the largest, wildest, most intelligent, and warlike Indians in
California.

The commission, accompanied by an escort of 36 dragoons under
command of Maj. W. W. Wessells, United States Army, left Sonoma
August II, 1851. As stated in the journal (p. 134): “Agent McKee
moved with his escort from Sonoma, at 8 o’clock a. m., and the
command is now encamped 19 7/10 miles up the Sonoma valley, on
the Russian river valley trail. Mr. George Gibbs has been employed
ass@hinook antetpreter . . 27°

Two days later, August 13, the party left the Sonoma valley and
followed a trail on the west side of Russian River, and that night
encamped “five or six miles below the first cafion, or defile, through
which the stream flows.” The next day the commission was joined
by Gen. J. M. Estelle and staff, of the 2d division of the California
militia, who had been sent by the Governor to assist in effecting
treaties with the Indians residing near Clear Lake and Russian River.

On the night of August 16 the entire party camped on the bank
of Russian River, then moved on to (p. 136):

Camp Lupiyuma, near Clear lake, August 17, 1851—R. McKee and party,
composed of secretary, and Gibbs as interpreter, with a sufficient number of
pack-mules to transport provisions and such presents as are designed for the

Indians; also ten head of cattle, with a detachment of ten dragoons in charge
of Major Wessells as an escort, all under the guidance of two Indian guides,

undertaking, were included in the Report of the Secretary of the Interior...
March 17, 1853. Special Session, Senate Ex. Doc. No. 4. The following
quotations, unless otherwise stated, are froin that document.

* McKee sent a lengthy letter to the Commissioner of Indian Affairs dated
“Camp at Big Bend of Eel River, September 12, 1851,’ in which he said
(p. 181): “At Sonoma I was fortunate in securing the services of George
Gibbs, formerly of New York, and recently attached to the Indian commission
in Oregon. He is acquainted with the Tchinook (Chinook) language, and the
jargon spoken by all the tribes on the borders of Oregon and California. He is,
moreover, a practical topographical engineer; has kept a journal of our entire
route, and will furnish me, I hope, in time for my final report, a correct map
and reconnaissance of the trail from Sonoma, showing the exact position of
all the important rivers, lakes, mountains and valleys, together with a synopsis
of the various dialects of the tribes we have met.”

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

left the main camp at an early hour this morning, and commenced ascending
the mountains dividing the Russian river and Clear Lake valleys, following a
narrow, precipitous trail leading in many places through a dense forest, with
oak and chemisall undergrowth ... We are encamped upon the table-lands
immediately adjoining the lake. Several Indians have visited camp this evening,
and we expect to have several chiefs in council to-morrow.

On the morning of the 18th “according to agreement a number
of the chiefs and braves of the Clear Lake Indians” met the com-
mission in council. Much of interest and importance transpired
during the succeeding hours. The Council again convened on
Tuesday afternoon, August 19, when McKee explained to the
Indians the nature of the proposed treaty, and stated that he would
give them, in the name of the President, “all of the Clear Lake
valley proper, upon condition they would all live in it peaceably, and
agree that all other tribes the President may send among them to
live should .be received as brothers, &c.’”’ To this the Indians agreed.

The party returned to the main encampment on Russian River
and soon continued northward. During August 25, they crossed the
mountains between the Russian and Eel Rivers and that night
camped in a valley near an old Indian village. The Indians were
“entirely nude and very wild.” Many Indians appeared at the camp
the following morning, and the commissioner explained the nature
of the treaties made with the Indians at Clear Lake and Russian
River, after which there was a mutual understanding for peace.

The expedition continued through the mountains, but advanced
slowly over unknown trails. They left camp early on August 29 but
were soon forced to stop in a canon, a place where sufficient water
could be obtained for the animals. During the day 15 or 16 Indian
men, all entirely naked, visited the camp. From them it was learned
that many Indians were living in “a long valley on the headwaters
of the middle fork of Eel river.” The party had been unable to
discover the valley, although they had spent several days in hunting
for it. The Indians called the valley Ba-tim-da-kia.

During September 5 and 6, the expedition rested at Camp Red-
wood, on the South Fork of Eel River.

On September Io, the camp was at “Big Bend of Eel river,’ where
the party remained several days. Gibbs had, as always, been active
in obtaining information concerning the Indians, and as mentioned
in the journal (p. 151): “Some words, relating to sensible objects,
have been obtained by Mr. Gibbs. The names of tribes could not be
ascertained, nor their numbers. But he has learned that all the Indians
around Humboldt bay, and as far up Eel river as Van Dusen’s fork,
say fourteen miles, speak the same language. Above the forks a

No. 8 DRAWINGS BY GEORGE GIBBS—-BUSHNELL 2

on

ditferent dialect is spoken, but so as to be understood by the different
tribes.” And at the same camp two days later “‘about a dozen naked
Indians hanging around camp were supplied with food and some
clothing.”

The following morning, September 13, Gibbs, with two other
members of the party, left camp to “proceed in a canoe down Eel
river to its mouth.” This was to enable him to explore the country
south of the river, and to meet the Indian inhabitants of the region
in the endeavor to have them visit the camp. Gibbs returned to the
main camp on the evening of September 14, after having reached
the mouth of the river, and stopping at 10 or 12 Indian rancherias
on the banks of the stream where they distributed presents. That
same evening “the express despatched to Port Trinidad returned .. .
with despatches from the Indian department at Washington.”

“Camp of Humboldt City, September 15, 1851.—Finding it impos-
sible to collect the Indians, or to hold proper communication with
them, R. McKee moved camp with the escort this morning, and
encamped at this place, after a march of twelve miles...” On
the following day they crossed Eel River at low tide and moved a
few miles to Bucksport. An entry in the journal that day states
(p. 154): “The bands of Indians living upon Eel river have no per-
manent place of residence, but move from river to mountain and
from mountain to river, as the season for fishing and gathering nuts
and berries arrives; and among the different bands, though not at
actual war, no friendly intercourse exists.”

Soon they resumed their journey northward, encountering many
Indians as well as white settlers, and so approached one of the most
important centers. They arrived at (p. 156):

Bloody Camp, September 27, 1851, three miles from the junction of the
Klamath rivers—Reached this camp, upon the top of a mountain, after a very
tedious march of twelve miles. Several Indians were seen upon the trail to-day,
but fled to the woods when approached. Fine grass and water on this mountain.
The agent has gone forward this morning to examine the pasture, &c., near the
junction, with a view to the removal of our camp to a suitable place for a
treaty ground, if one can be found. The country around the junction is a wild
mountainous region, entirely unfit for cultivation, and indeed can scarcely be
travelled by pack-mules. The Indians are said to be numerous, and subsist
chiefly on the salmon and salmon trout, which the rivers afford in great
abundance, and on the berries, nuts, &c., obtained on the mountain sides. Deer,

elk, bear, &c., are quite plenty, but the Indians kill but few, as their only arm
is the bow and arrow.

Two days later, September 29, McKee accompanied by part of
his guard, established a camp at the junction of Klamath and Trinity

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Rivers where they remained until October 9. Many Indians gathered
near the camp (p. 157):

The Indians here are a very fine-looking race, low in stature, with smooth,
regular features. The men are nearly nude, and never seen without the bow
and quiver of arrows, exhibiting considerable skill in their construction. The
women wear petticoats of deer-skin, dressed and ornamented with tassels, beads,
shells, &c. Some of them are very handsomely made. Strings of beads and
shells are also worn about the neck, and ornaments of every description are
highly prized.”

On October 6, a treaty was concluded with the many groups of
Indians who had gathered for that purpose at the junction of the
Klamath and Trinity Rivers, and that night (p. 162): “Presents of
blankets, shirts, pants, beads, shawls, handkerchiefs, &c., &c., were
distributed by the light of large fires; after which the Indians all
separated for the night, well pleased.” Among the bands present
that day was the “Wetch-peck band” of which Mec-ug-gra was
chief, . (Pl. 18:)

On October 9, the party moved from the vicinity of the junction
of Klamath and Trinity Rivers, crossed the Klamath at Durkee’s
ferry, and established camp near Bluff Creek. During the day they
traversed a rough mountain trail. A sketch made at that time shows
a small Indian encampment in the bend of the river.

The Indians near Salmon River speak a different dialect from those below
them.

Camp Cor-a-tem, near mouth of Salmon river, October 12, 1851—R. McKee
remained at this camp to meet the Indians of this neighborhood as agreed. At
10 o'clock about 150 men and women were assembled. Each band arranged

* Quantities of trade beads were distributed among the Indians of California.
They were received from several sources, and the following letter will tell
where some were obtained (p. 360): Letter from P. B. Reading to Luke Lea,

Indian Commissioner— eS :
Washington, September 10, 1852.

“Str: Please notice below a memorandum of articles which I would recom-
mend to be purchased for presents to be distributed among the Indians in
California. It will be well to make the purchases in New York, as it is quite
uncertain if Mr. Beale, the superintendent, could find the articles in California:

Small whites porcelain beads 4..c.-v-seyntesesaioricle easyer teers $4,500
Small blacks porcelamimbeadsar eee erenaeie- eeececieceerieet 2,000
SmallpmedmpercelainybeadSteere reese ten ncee cre eer eeer 3,000
ikargenclassebeadss assontedemne scenic tiie 2,000
Miike yer red) ssp Tle Srays.a, vote eacy'ssie ese eee eevee eeeeicbr nero 2,500
Gay=colored tshawilsms-coey aus coc ocinern ed oa eon see 1,500

15,500

“It will be necessary that the packages be made up to the weight of about
200 Ibs., and well secured in oilcloth coverings, in order to prevent damage to
contents, as the present is the rainy season.”

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO Si NO ose to)

i ;

& iene Wert- shek Chu
Ney Fem -

a

YOUNG CHIEF OF THE WEIT-SPEK TRIBE, PROBABLY MEC-UG-GRA

Drawn by Capt. Seth Eastman, from original sketch by George Gibbs,
October 1851.

No. 8 DRAWINGS BY GEORGE GIBBS—BUSHNELL 27

separately, communicating with them through Mr. C. W. Durkee, and he
through the Wetchpeck Indian from the junction of Klamath and Trinity .. .

The number of Indians living near this camp, around the mouth of Salmon
river, is about 225 souls, all told. They compare favorably in size and appear-
ance, and intelligence, with the Indians below; speak a different dialect, though
they intermarry with them. Their houses are built of slabs split out from
redwood timber, in which a family of ten or fifteen will reside, relying principally
upon fish for a subsistence.

The following day, October 13, the party was in “Camp on
Klamath river, 12 miles above mouth of Salmon river.” The trail
over which they passed that day was difficult and dangerous, and
“several of our animals fell from the trail and rolled down the
mountain, but were recovered and brought into camp.” Gibbs made
another sketch of the rugged mountain scenery.

“Camp in Scoti’s valley, October 21, 1851 . . . Our route to-day
led up the Klamath river to the north of Scott’s river; thence up
Scott’s river to Scott’s bar, where a large number of miners have
been and are at work; thence crossing Scott’s river, and over a
high, steep mountain into this valley ...’’ The party remained in
the valley several days, during which time the surrounding country
was examined. On the 24th “Messrs. Gibbs, Kelsey, and Woods
have been sent out to explore and examine this valley, and adjacent
hills and mountains, relative to its adaptation for an Indian reserva-
tion.” Two days later, October .26, “Mr. George Gibbs was des-
patched to Shasta plains to examine that part of the country.”

“Camp in Scoti’s valley, October, 27, 1851—This is the day
appointed for the Indians in Shasta and Scott’s valleys, and on
Scott’s river, to assemble at this camp .. . The citizens of Shasta
Butte city assembled.” ~

Remained in Scott’s valley until November 6, when they “com-
menced the march for the coast by the same trail we came over
from Durkee’s ferry.” Continued on through the mountains and on
November 11 “crossed to the east side of the. Klamath at an Indian
rancheria, swimming our animals—no accidents.” ;

“November 12, 1851.—Detained some time this morning getting
our mules from the mountain, and crossing our goods over in canoes:
swimming the mules over, they became alarmed, and two were
drowned. This to us is a serious loss . . . Started at 10 o’clock, and
reached our old camp ‘Coratem,’ near the mouth of Salmon river, at 4.”
And the next morning “Mr. Gibbs and myself took a canoe and
three Indians this morning and descended the river, passing many

*On July 30, 1852, McKee wrote to Luke Lea, Commissioner of Indian
Affairs, Washington, and referred to happenings “in the neighborhood of
Yraka (late Shasta Butte city).” (P. 353.)

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

dangerous ripples or rapids, and at 4 o'clock reached Durkee’s ferry,
mouth of Trinity river.”

The expedition was disbanded and Gibbs went to San Francisco.

Quotations from two letters written by Redick McKee, to Luke
lea, Commissioner of Indian Affairs, Washington, will shed light
on subsequent events (pp. 294-297) :

San Francisco, March 1, 1852.
... Since I wrote I have received from Mr. George Gibbs his report, or
journal, of the expedition to northern California, accompanied by a very beauti-
ful map of the country traversed, and sundry vocabularies of the languages
spoken by the tribes we visited. These I design sending to you by the mail
which takes this; but our friend, General S. D. King, of the land survey
department, is making a copy of the map, and the Senate Committee on Indian
Affairs desires me to afford them a reading of Mr. Gibb’s views as to the
reservations made for the Indians, at a meeting appointed for the 4th instant .. .

P. S. Mr. Gibbs having forwarded some sketches to Mr. Schoolcraft by the

last mail, I will, with this, send the vocabularies.
The second letter reads in part:
San Francisco, March 13, 1852.

Sir: My last despatch was dated Ist instant, and accompanied a_ sealed
package of vocabularies, prepared by Mr. George Gibbs. I have deposited in
the post office, to go with this letter, Mr. Gibb’s map of my route through
northern California, and his manuscript journal of the expedition. This journal,
the map, and the sketches forwarded by last steamer to Mr. Schoolcraft, will,
I hope, be neatly and carefully published. They will throw some additional
light upon a part of this State, not previously explored. On this subject, I
enclose letters from Mr. Gibbs to the honorable Senators Hamilton Fish and
Truman Smith, and to H. R. Schoolcraft, esq., which you will please read,
and then deliver.

It is now possible, after the lapse of many years, to present the
sketches “neatly and carefully published,” together with others which
were made by Gibbs before he joined McKee on the journey into
northwestern California.

During subsequent years, until his departure from the Pacific coast
late in 1860, Gibbs’ interest in the Indians continued. He made
vocabularies among the native tribes scattered over a wide region,
and gathered ethnographical material in California, Oregon, and
Washington, on Puget Sound and far up the Columbia. As such
material was at that time so plentiful, it is evident he selected choice
specimens to be carried, or sent, to his home in New York. If all
the material thus collected could be brought together, it would prove
of special interest as representing the work of tribes then living in
their primitive state, maintaining manners and customs that had been
followed and practiced for generations, but which were soon to be
lost or changed through contact with those who came to claim and
occupy the country.

rk OCTOBER 10, 1938

din¥ ar

VOLUME 97, NUMBER 9

Apa

_ SMITHSONIAN MISCELLANEOUS COLLECTIONS

“BY.

Bally ger H. G. DEIGNAN

Division of Birds, U. S. National Museum

(PUBLICATION 3486)

: GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 97, NUMBER 9

A NEW NUTHATCH FROM YUNNAN

BY
H. G. DEIGNAN

Division of Birds, U. S. National Museum

°, yt AF : |
A gaiTHsONS™ I

CA ITV Tes
Seen GTO ooo

(PUBLICATION 3486)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 10, 1938

The Lord Waltimore ress

BALTIMORE, MD., U. 8S. A.

A NEW NUTHATCH FROM YUNNAN

By H. G. DEIGNAN
Division of Birds, U. S. National Museum

A series of 26 specimens of the giant nuthatch, representing the
material in the United States National Museum, the American Mu-
seum of Natural History, and the Academy of Natural Sciences,
Philadelphia, shows that the bird of northwestern Yunnan is quite
distinct from magna, based on specimens from Karen-ni. For the
new form I propose the name

SITTA MAGNA LIGEA, n. subsp.

Type.—Adult male, U.S.N.M. no. 297271; collected at Likiang,
northwestern Yunnan, 8,200 feet, August 1923, by Dr. Joseph F.
Rock. :

Diagnosis —Readily distinguished from Sitta magna magna by its
markedly shorter and laterally slenderer bill, although not separable
by color characters or other measurements. The culmens of nine males
from northwestern Siam and the Shan States measure (from the
base of the rhamphotheca) 30.3 to 32.5 mm (average: 31.4 mm) ;
of eight females from the same localities, 29.5 to 32.5 mm (average:
31 mm). On the other hand, the culmens of four males from the
mountains of Likiang measure 26 to 28.5 mm (average: 27.1 mm).
The difference in the “heaviness” of the bill is scarcely to be shown
in figures, but is very obvious to the eye.

Range.—Northwestern Yunnan: the Likiang mountains, 8,200 to
10,000 feet ; Yung-pei.

Remarks.—I have not seen topotypical magna, but feel justified in
considering Siamese specimens, taken about 50 miles east of Karen-ni,
to belong to that form, especially since they agree perfectly with
Shan States examples which British authors have held to be identical
with Karen-ni birds.

Two Shan birds, whose measurements are not included above, are
not fully mature, as is indicated by the texture of the plumage. In
these specimens, the bill is as short as in the Likiang examples, but
at the same time as thick laterally as in fully adult Shan specimens.
a combination which gives it a peculiarly blunt appearance for a
nuthatch.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97, No. 9

bo

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

The sexes of this species are easily separable in series. The male
has the dark markings of the head wholly glossy blue-black; in the
female they are dull black or black washed with gray, with a tendency
to gloss only on the shoulders. In the male the mantle is slaty-blue;
in the female paler slaty and much less suffused with blue. The female
has the underparts pale gray, more or less suffused throughout with
buff or pale chestnut ; the male has the same parts wholly a soft blue-
gray, with buffy suffusion confined to the lower abdomen. The
chin and throat are white in either sex, but in the female they are
faintly washed with buff.

The “Fauna of British India” describes the upper plumage simply
as “‘slaty-blue,” and fails to mention one of the most striking char-
acters of the species. Actually, the whole portion of the head, neck,
and upper back included between the broad black stripes is quite
different from the mantle, the front and crown being a soft blue-
gray, changing gradually to a still paler gray upon the nape and
upper back, which is sharply defined from the color of the remaining
upperparts. In the female the light portion is very faintly suffused
with buff, and, because the adjacent colors are less contrasting, is
slightly less conspicuous than in the male.

The front and crown in either sex may have more or less black
streaking in the gray. In the series before me, these parts vary in-
dividually from immaculate gray to almost solid black.

My thanks are due to John T. Zimmer, who has kindly sent me the
Burmese material from the Rothschild collection, and to R. M. de
Schauensee, who has lent me the Yung-pei specimen of ligea, as well
as his valuable series of magna from Siam.

meant "SMITHSONIAN MISCELLANEOUS COLLECTIONS ©
Silene 4 VOLUME 97, NUMBER 10.

FOURTH CONTRIBUTION

|, 10 NOMENCLATURE OF CAMBRIAN

i POSSIES

BY
GHARLES ELMER RESSER

Curator, Division of Invertebrate Paleontology,
U. S. National Museum

(PUBLICATION 3487)

GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 17, 1938

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER 10

FOURTH CONTRIBUTION
TO NOMENCLATURE OF CAMBRIAN
FOSSISe

BY
CHARLES ELMER RESSER

Curator, Division of Invertebrate Paleontology,
U. S. National Museum

(PUBLICATION 3487)

GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 17, 1938

The Lord Battimore Drees

BALTIMORE, MD., U. 8. A.

FOURTH CONTRIBUTION TO) NOMENCLATURE OF
CAMBRIAN FOSSILS

By CHARLES ELMER RESSER

Curator, Division of Invertebrate Paleontology,
U. S. National Musewmn

This is the fourth paper in a series dealing with nomenclatural
changes necessary for Cambrian fossil species.’ In this paper non-
trilobitic and non-American species are included. It was hoped all
necessary changes in sight could go in this fourth contribution, but
that proved to be impossible. Several descriptive papers now in
press or nearing completion involve previously described species,
‘and thus make additional nomenclatural changes. Moreover, it is
the practice to withhold references of species to new genera where
undescribed species are in hand which will make better genotypes.
Needed changes are also withheld in instances where someone is
known to be at work on the faunas which contain improperly named
species.

The nontrilobitic species are discussed first, grouped according to
classes. The trilobites are again placed in alphabetical order accord-
ing to genera.

BRACHIOPODA
PATERINIDAE Schuchert
MICROMITRA Meek, 1873

Micromitra minutissima (Hall and Whitfield)
Kutorgina minutissima Hatt and WuitrFieEtp, U. S. Geol. Expl. 4oth Par.,
WO, 4h he By Ws Boye wh wy, wes, ie, Wes ans yy
Kutorgina sculptilis Watcott (part), U. S. Geol. Surv. Mon. 8, p. 20, pl. 1,
fig. 7b; pl. 9, fig. 7, 1884.
Micromitra sculptilis WaAvcotr (part), idem, 51, p. 341, pl. 3, figs. 5b, c, 1912.

The Nevada form is distinct from M. sculptilis of Montana in
having coarser ribs. For this reason the old species name is restored.

1 Resser, Charles Elmer, Nomenclature of some Cambrian trilobites, Smith-
sonian Misc. Coll., vol. 93, no. 5, Feb. 14, 1935; Second contribution to nomen-
clature of Cambrian trilobites, idem, vol. 95, no. 4, Apr. 1, 1936; Third
contribution to nomenclature of Cambrian trilobites, idem, vol. 95, no. 22, Apr.
5, 1937.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97, No. 10.

bo

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

Middle Cambrian, Eldorado; (loc. 58) east side New York and
Secret Canyons, Eureka District, Nevada.
Cotypes—U.S.N.M. no. 245514, b.

Micromitra burgessensis, n. sp.
Micromitra (Iphidella) pannula Watcott, Smithsonian Misc. Coll., vol. 67,
no. 9, p. 481, pl. 106, fig. 16, 1924.

This species conforms in every respect to the characteristics of
Micromitra even though only one valve has been found. The growth
lines are pronounced and are wavy and close together. Ribbing is
well developed, and the ribs have the usual irregularities. Because
of the crinkly growth lines and particularly because of the preserva-
tion of long slender spines which extend far beyond the margin, this
shell was regarded as the same as J. pulchra. However, a glance at
the illustrations shows that the two forms are quite different and
that pulchra is a true [phidella.

Middle Cambrian, Burgess; (loc. 35k) Burgess Pass, near Field,
British Columbia.

Holotype-—U.S.N.M. no. 69646.

PATERINA Beecher, 1891

Paterina troyensis, n. sp.

Scenella retusa Watcotr (part), U. S. Geol. Surv. Bull. 30, p. 126, pl. 12,

fig. 3a, 1886 (not fig. 3=Stenothecoides).

The specimen doubtfully referred to Scenella retusa by Walcott
is a brachiopod belonging to Paterina. It is a small form only 1.5 mm
long and 1 mm high. About halfway back a depression follows the
position of an ordinary growth line, but this is regarded as an acci-
dental feature. The shell appears to be nearly smooth, but this may
be due to imperfect preservation.

Lower Cambrian, Schodack; (loc. 27) Troy, New York.

Holotype —U.S.N.M. no. 15369.

Paterina zenobia (Walcott)
Micromitra (Paterina) stissingensis? Watcorr (part), U. S. Geol. Surv.
Mon: 51, p. 353, pl. 3, fig. Ie, 1912.
Micromitra (Paterina) stissingensis ora Wa.cortt, idem, p. 354, 1912.
Micromitra genobia Watcort, idem, p. 342, text fig. 23, 1912; Smithsonian
Misc. Coll., vol. 67, no. 9, p. 481, pl. 106, figs. 1-7, 1924.

Middle Cambrian, Burgess; (loc. 35k) Burgess Pass, (locs. 61],
35k’) West slope Mount Field; (loc. 14s) Mount Stephen near
Field, British Columbia.

Holotype —U.S.N.M. no. 58311; plesiotypes, nos. 56907, 51483,
6963 1-7.

INOS -LO CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 3

IPHIDELLA Walcott, 1905

Iphidella fieldensis, n. sp.

Micromitra (Iphidella) pannula Watcorr (part), Canadian Alpine Journ.,
vol. I, no. 2, pl. 1, fig. 1, 1908; U. S. Geol. Surv. Mon. 51, p. 361, pl. 14,
Liou ie TOL:

This species was confused with J. pannula but there is little re-
semblance. J. fieldensis is characterized by a fairly well developed
diamond pattern on the older part of the shell, which is more and
more obscured toward the margins by increase in prominence of
the wavy growth lines.

Middle Cambrian, Stephen; (loc. 14s) Mount Stephen; (loc. 35k)
Burgess Pass, near Field, British Columbia.

Holotype-—Walker Mus., Toronto.

Iphidella pulchra, n. sp.
Micromitra (Iphidella) pannula Wa.cortt, Research in China, vol. 3, Carnegie
Inst. Publ. 54, pl. 1, fig. 13, 1913; Smithsonian Misc. Coll., vol. 67, no. 9,
p. 482, pl. 106, fig. 17, 1924.

This is a very beautiful brachiopod. Well-preserved specimens
show “spines” extending far beyond the margins of the shells. 7. pul-
chra carries the diamond pattern to its extreme development. Very
large specimens show the pattern interrupted somewhat in the outer
part of the shell by rather heavy growth lines.

Middle Cambrian, Burgess; (loc. 35k) Burgess Pass; (loc. 14s)
Mount Stephen, near Field, British Columbia.

Holotype —U.S.N.M. no. 59801.

OBOLIDAE King
LINGULELLA Salter, 1866

The family Obolidae includes many brachiopod genera, chief of
which in Cambrian strata are Obolus and Lingulella. Each contains
many species, and to both genera are assigned a number of sub-
genera. Discrimination of oboloid brachiopods is an inexact pro-
cedure at present because no one has yet determined what the charac-
teristics essential for classification may be. Relative size and shape,
and sometimes shell ornamentation, are the only usable criteria.

Proper generic names are not available for many Cambrian species.
It is evident that most, if not all, of the subgenera in Obolus and
Lingulella are valid genera, but these two names themselves are in
question. One thing is certain, namely, that Obolus must be confined
to post-Cambrian species. Lingulella is founded on L. davisit McCoy,

4. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

also a post-Cambrian brachiopod. Unfortunately the specimens of
L. davisi are poorly preserved; consequently it is difficult to deter-
mine whether Cambrian forms are congeneric.

Distinctions of generic rank between the Cambrian species referred
to Obolus and those placed in Lingulella seem to be wanting. In
fact the assignment of species to these genera has been on the basis
of shape alone, the wide forms being called Obolus. Pending revi-
sion of these brachiopods, Cambrian species which formerly would
have been placed partly in Obolus and partly in Lingulella are prefer-
ably referred to Lingulella.

Lingulella bridgei, n. sp.
Lingulepis acutangula Brivce (part), U. S. Geol. Surv. Prof. Paper 186-M,
p. 244, pl. 68, figs. 4, 5, 1037.

The stratigraphic position of the sandstone containing this species
has not yet been determined with certainty.

This is a large form for which reason Bridge confused it with
Lingulepis acutangula. However, it lacks the necessary restriction
of the ventral valve at the apical end, tapering too slowly and evenly
to be referred to Lingulepis.

The ventral valve figured by Bridge is broken away on the sides
toward the apex, thus causing it to approach the form of Lingulepis.
A smaller, more perfect ventral valve shows the true proportions of
the shell, and while the valve comes to a rather sharp point, the mar-
gins approach the apex as slightly convex lines. The dorsal valve
figured by Bridge is fairly complete but evidently somewhat flattened.

L. bridgei has the usual shell composition. A ventral valve about
15 mm long is 10 mm wide.

Upper Cambrian, Lion Mountain ?; { mile east of Sandy, Blanco
County, Texas.

Cotypes.—U.S.N.M. no. 93009.

Lingulella burnetensis, n. sp.
Obolus matinalis Waucort (part), U. S. Geol. Surv. Mon. 51, p. 400, pl. 8,
figs. 1a, b, 1912; Brice (part), in Sellards, Adkins, and Plummer,
Univ. Texas Bull. 3232, pl. 2, fig. 14, 1932.

This is a rather wide form, which is perhaps the reason for its
reference to O. matinalis in spite of its larger size. There is some
question as to the specific identity of the specimens from the two
localities. Since the holotype is in sandstone it may come from the
Lion Mountain sandstone member. The other specimens (loc. 67)
are clearly in the Aphelaspis zone.

NO. IO CAMBRIAN FOSSILS, 4TH CONTRIBUTION—-RESSER 5

This species is characterized by its broad shape. The ventral valve
has a sharp apex but widens rapidly, so that the holotype is about
14 mm long and 12 mm wide.

Upper Cambrian, Cap Mountain; (locs. 67, 67c) near the top
of Potatotop, 7 miles northwest of Burnet, Texas.

Holotype —U.S.N.M. no. 52420; paratype, no. 524109.

Lingulella alia, n. sp.
Obolus matinalis Wautcotr (part), U. S. Geol. Surv. Mon. 51, p..400, pl. 8,
fig. 1k, 1912; BripcE (part), in Sellards, Adkins, and Plummer, Univ.
diexas Bullls232%pl. 2) fig, 15, 1032!

This is a smaller brachiopod than the Cap Mountain species,
L. burnetensis. Both valves are very wide, the holotype dorsal valve
being about 9 mm long and g mm wide. Even though the ventral
valve appears to be less wide, measurements show but little difference
between length and width.

Upper Cambrian, Wilberns; (loc. 71) Cold Creek Canyon, San
Saba County, and (loc. 14c) Baldwin’s Ranch, Cold Creek, Llano
County, Texas.

Holotype—U.S.N.M. no.. 51566.

Lingulella monticola, n. sp.
Lingulella perattenuata Wavcotr (part), U. S. Geol. Surv. Mon. 51, p. 523,
Dl Diy wey Uy UCT,

This small, neat form was referred by Walcott to L. perattenuata.
The ventral valve is about 6 mm long and 3.5 mm wide, and the
rate of taper is nearly rapid enough to warrant reference to Lingulepis.
Otherwise the species conforms to the norm for the genus. This
form is more slender than the older species, L. longula.

Upper Cambrian, Cap Mountain (Lion Mountain member ?) ; (loc.
67c) Potatotop, 7 miles northwest of Burnet, Texas.

Holotype —U.S.N.M. no. 27424.

Lingulelila longula, n. sp.
Lingulclla perattenuata Watcotrt (part), U. S. Geol. Surv. Mon. 51, p. 523,
Peet htess. 1, 2, LOL: :

L. perattenuata is thus far confined to the Black Hills and the
name must be restricted to Whitfield’s specimens. Unfortunately
they do not warrant the precise drawings Walcott presents. The
form from the Middle Cambrian of the Grand Canyon is an unde-
scribed species. The specimens in sandstone from locality 70 also
represent a new species, but since most of the material from that

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

locality belongs to the Wilberns formation, the horizon of the sand-
stone is in question.

L. longula is associated with Obolus sinoe, from which its elongate
shape readily distinguishes it. Compared to L. perattenuata, L.
longula averages much smaller in size and is more slender. Only
dorsal valves are figured by Walcott but ventral valves are present
in the collection. The shell has the usual growth lines well marked.
The smaller type specimen is about 5 mm long and 3 mm wide.

Upper Cambrian, Hickory; (loc. 68y) Packsaddle Mountain, 11
miles southeast of Llano, Texas.

Cotypes—U.S.N.M. no. 2742

to

Lingulella hilli, n. sp.
Obolus (Lingulella) acutangulus Watcott, Proc. U. S. Nat. Mus., vol. 21,
Ps 3025 Dla 279 nie Ome pleeSe fessele 2TSOG:
Lingulella acutangula Watcott (part), U. S. Geol. Surv. Mon. 51, p. 474,
Seto Sae lame Geman lneesy mae
Lingulella arguta BripcE, U. S. Geol. Surv. Prof. Paper 186—M, p. 245, 1937.

Bridge recently pointed out that Walcott’s L. acutangula is not
related to Lingulepis acutangula of Roemer with which it was identi-
fied, and further suggested that this brachiopod is L. arguta. How-
ever, Bridge failed to note that several species from different hori-
zons were included by Walcott in L. arguta.

Walcott’s original identification of Roemer’s species was based on
the Hickory sandstone specimens described here as L. Jill. Sub-
sequently (1912) Walcott added Wilberns specimens which belong
to a much larger species. The matrix of L. hilli is a black rock,
consisting mainly of hematite in which there are scattered poorly
rounded grains of clear quartz. The brachiopods are not compressed
and many specimens retain a limy shell. This material was collected
by R. T. Hill about 1885 from an undetermined locality in Llano
County. When the shell is dissolved, many specimens show the inter-
nal features, which explains Walcott’s choice of them to illustrate the
species even though the locality is unknown.

L. hilli varies in size, averaging about 7 mm long and 4.5 mm
wide for the ventral valve, with the dorsal relatively shorter. The
other features are those usually characterizing Lingulella_ species.

L. hilli is larger and wider than L. arguta from Nevada.

Upper Cambrian, Hickory; Llano County, Texas.

Cotyvpes—U.S.N.M. no. 35240.

NO. 10 CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 7,

Lingulella texana Walcott
Obolus sinoe Watcotr (part), U. S. Geol. Surv. Mon. 51, p. 415, pl. 26,
figs. 2-2d, I9I2.
Lingulella texana Waucott, Smithsonian Misc. Coll., vol. 53, no. 3, p. 71,
pl. 8, fig. 5, 1908; U. S. Geol. Surv. Mon. 51, p. 535, pl. 40, figs. 3, 3a.

Walcott’s description stresses the radiating striae on the “outer
surface” of the shell as the diagnostic feature. When it is remem-
bered that the ribbing of a Lingulella shell is exposed only by exfolia-
tion of one or more layers, this species loses its distinctive feature.
This explains why specimens which retained the outer shell layers
were referred to Obolus sinoe.

Upper Cambrian, Wilberns; (loc. 70) Baldy Mountain, 8 miles
northwest of Burnet, and (loc. 69) Honey Creek, 8 miles southeast
of Llano, Texas.

Holotype.—U.S.N.M. no. 51806; paratype, no. 51805; plesiotypes,
no. 51627.

Lingulella lochmanae, n. sp.
Obolus (Lingulella) acutangulus Wa.cott (part), U. S. Geol. Surv. Mon.
Finepaa 7A. plei7, Tests ds eed) k m=o; TOL:
Lingulella arguta LocHMAN, Journ. Pal., vol. 12, no. 1, p. 85, pl. 18, figs.

17-19, 1938.
Lingulella acutangula Brice (part), in Sellards, Adkins, and Plummer, Univ.
Texas Bull. 3232, pl. 2, fig. 8, 1932.

This is a medium-sized species which falls between such species
as L. alia, L. burnetensis and L. bridgei, on the one hand, and the
small species like L. texana, L. longula, and L. hilli on the other.
There is naturally variation in size. A large ventral valve is about
13 mm long and 8 mm wide.

The specimens from locality 68 are doubtfully referred to the
species. In fact, it is not certain that all the others belong to one
species.

Upper Cambrian, Cap Mountain; South Fork Morgan Creek, 4
miles northwest of Highway 29; Lion Mountain sandstone?; (loc.
68) near top of Packsaddle Mountain, 12 miles southeast of Llano;
Wilberns; (loc. 69) Packsaddle Mountain, and (loc. 69) Honey
Creek, 8 miles southeast of Llano, all in Texas.

Holotype—vU.S.N.M. no. 95017; paratypes, nos. 27410, 27412,
51644. we
Lingulella nina (Walcott)

Obolus tetonensis ninus WaAucott (part), U. S. Geol. Surv. Mon. 51, p. 418,
pl. 11, figs. I-Ig, I912.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

This species characterizes Honey Creek limestone of Oklahoma.
The Texas form, from the slightly older Lion Mountain sandstone,
formerly referred to L. nina, is a larger brachiopod.

Upper Cambrian, Honey Creek; (loc. 9r) 15 miles northwest of
Fort Sill, and other localities in Oklahoma.

Holotype and paratypes —U.S.N.M. no. 51643.

Lingulella turneri, n. sp.
Lingulella (Lingulepis) acuminata Watcotr (part), U. S. Geol. Surv. Mon.
51, p. 545, pl. 34, figs. 3a-d, 1912.

This form was apparently placed in Lingulepis acuminata he-
cause L. nevadensis was included with it.

L. turneri is characterized by an average size for the ventral valve,
about 9 mm long and 6 mm wide. There is a tendency toward flatten-
ing of the front marginal outline.

Upper Cambrian, Emigrant; (locs. 7x, 7z) 2$ miles southeast of
Benders Pass, Silver Peak Range, Nevada.

Cotypes.—U.S.N.M. nos. 56967, 51884.

LINGULEPIS Hall, 1863

Lingulepis burnetensis, n. sp.
Lingulella (Lingulepis) acuminata Wa.cotr (part), U. S. Geol. Sury. Mon.
51, p. 545, pl. 42, figs. 1k-n, 1912.

This species is smaller than either Lingulella bridgei or Lingulepis
llanoensis. Compared with the latter L. burnetensis has about the
same width, but the shell is shorter and the ventral valve tapers less
evenly. One specimen g mm long is 6 mm wide.

Upper Cambrian, Wilberns; (loc. 70) Baldy Mountain, Morgans
Creek, 8 miles northwest of Burnet, Texas.

Cotypes—U.S.N.M. no. 51891.

Lingulepis llanoensis, n. sp.
Lingulella (Lingulepis) acuminata Watvcotr (part), U. S. Geol. Surv. Mon.
51, D. 545, pl. 42, fig. I, 0, 1912.

Walcott figured only the interior of the dorsal valve, but a very
fine example of the ventral valve lies against it. This species is
slender, a ventral valve 12 mm long being only about 6 mm wide. The
ventral valve tapers at a nearly even rate to the apex.

Upper Cambrian, Wilberns; (loc. 14b) Cold Creek, 2 miles south
of the San Saba county line, Llano County, Texas.

Holotype and paratypes—U.S.N.M. no. 51892.

NO. IO CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 9

Lingulepis nevadensis, n. sp.
Lingulella (Lingulepis) acuminata Waucotr (part), U. S. Geol. Surv. Mon.
51, p- 545, pl. 34, figs. 3, 3e, 1912.

This brachiopod is associated with Lingulella turneri and evidently
formed the basis on which Walcott referred both species to
L. acuminata. ;

L. nevadensis is characterized by its relatively small size. Both
figures of Walcott represent ventral valves, which show the usual
degree of variation. The shell is not restricted toward the apex to
any great extent. The growth lines are wavy.

Upper Cambrian, Emigrant; (loc. 7z) about 24 miles southeast of
Benders Pass, Silver Peak Range, Nevada.

Cotypes—U.S.N.M. no. 51884.

ACROTRETIDAE Schuchert
ACROTHELE Linnarsson, 1876

Acrothele walcotti, n. sp.
Acrothele colleni Watcotr (part), U. S. Geol. Surv. Mon. 51, p. 640, text
fig. 55, 1912; Smithsonian Misc. Coll., vol. 67, no. 2, p. 25, pl. 4, figs. 5-5c,
1QI7.

This species is Characterized by the coarseness of both the ribs
and the growth lines. The outline of both valves is more transverse
than in A. colleni, from which A. walcotti further differs in the
coarseness of the surface markings.

Middle Cambrian, Ross Lake shale; (loc. 35c) 1 mile east of
Hector, Mount Bosworth; and (loc. 63j}) Popes Peak, 14 miles south-
west of Stephen, British Columbia.

Holotype and paratypes——U.S.N.M. no. 51410.

ACROTRETA Kutorga, 1848

Acrotreta aurumensis, n. sp.
Acrotreta microscopica Waucotr (part), U. S. Geol. Surv. Mon. 51, p. 693,
pl. 67, figs. 2-2d, 1912.

Walcott recognized the distinctness of the Nevada species from
A. microscopica but argued that the ditferent shape was due to pres-
sure. Neither the Texas nor the Nevada specimens are compressed
or distorted. A. aurumensis has a more elevated ventral valve and is
larger than A. microscopica. There is not as much variation in shape
as is shown in the illustrations. In fact figure 2a is erroneously
drawn, for that shell has the same outline as fig. 2.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

This species is characterized by the erectness of the ventral valve
so that the false area is about at right angles to the base of the shell.

Upper Cambrian; (loc. 8 0) 2 miles north of Aurum, Schell Creek
Range, Nevada.

Cotypes.—U.S.N.M. no. 52119.

BILLINGSELLIDAE Schuchert
WIMANELLA Walcott, 1908

Wimanella walcotti, n. sp.
Wimanella simplex Wavcott (part), U. S. Geol. Surv. Mon. 51, p. ror, text
fig. 64, 1912; Smithsonian Misc. Coll., vol. 67, no. 2, p. 26, pl. 4, figs. 7-7¢,

1917.

This species is represented by many specimens in fine-grained,
hard shale. Most of the ribbing and growth lines have been smoothed
out by pressure. Fine ribs show in cross light and the growth lines
must have been rather stronger than usual. The hinge line is straight
and the undistorted valve forms a little more than a semicircle.

Middle Cambrian, Ross Lake; (loc. 63j) Popes Peak, 14 miles
south of Stephen; and 1 mile east of Hector, Mount Bosworth,
British Columbia.

Cotypes—U.S.N.M. nos. 63713-5, 51407.

Wimanella rossensis, n. sp.
Wimanella simplex Watcotr (part), Smithsonian Misc. Coll., vol. 67, no. 2,

p. 26, pl. 4, figs. 8-8c, 1917.

Limestone nodules in the Ross Lake shale contain an abundant
fauna among which is a species of JV’imanella different from JI’. wal-
cotti in the enclosing shale. Preservation in granular limestone in
contrast to partial flattening in shale, accounts for some of the differ-
ences at first apparent. However, not all can thus be accounted for.
W’. rossensis is much higher in the ventral valve, thus creating a
larger area. Growth lines are heavy, but vary more in size than in
IV. walcotti.

Middle Cambrian, Ross Lake; (loc. 63j) Popes Peak, 14 miles
south of Stephen; and 1 mile east of ‘Hector, Mount Bosworth,
British Columbia.

Cotypes.—U.S.N.M. nos. 64716-9.

NO. LO CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER Il

GASTROPODA
PALAEACMAEIDAE Grabau and Shimer
HELCIONELLA Grabau and Shimer, 1909

Helcionella GRaBAU and SuHiIMEr, N. A. Index Foss., vol. 1, p. 607, 1900;
SMETANA, Roz Ceske Akad., vol. 27, no. 8, p. 3, 1918.

Grabau and Shimer erected Helcionella on Stenotheca rugosa
without recognizing the fact that four species were included in Wal-
cott’s figures copied by them. Many of the numerous errors of gen-
eric reference and specific determination are not included in the
present brief study. Confusion possibly still exists between Helcio-
nella and Stenotheca. In fact, this and related, or similar forms,
should undergo detailed monographic study in the near future.

Authors compare Helcionella with Palaeacmaea, distinguishing the
former on the basis of the submarginal position and incurved shape
of the apex. It may be that when the interior structure of Helcionella
is obtained, the genus will be found to be synonymous with Parmor-
phorella, which it resembles outwardly.

Genotype: Metoptoma? rugosa Hall.

Published species of Helcionella are listed together with their syno-
nyms. New names and new species are then discussed.

Atlantic Province
Helcionella annulata Smetana
o (Calloconus) ava Smetana =H. ava
. avus excentrica Smetana = H. excentrica
ss cingulata Cobbold
comleyensis Cobbold
emarginata Cobbold = H. emarginata
lata Smetana
ri lata glabra Cobbold
maxima Cobbold
= media Cobbold
4 oblonga Cobbold
oy pompeckji Cobbold
cf. rugosa Kiaer = H. kiaeri
i rugosa acuticosta Kiaer = H. norvegica
tjerovicensis Smetana
tenuis Smetana

smu)

North America
Helcionella cingulata ? Poulsen = Indeterminable fragment
= Stenothecoides elongata
4 elongata Walcott A g 5.
= labradorica

a pricet Resser and Howell

TZ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

China
Flelcionella ? clurius (Walcott) (Poorly preserved)
c rugosa chinensis (Walcott) = H. chinensis
Helcionella rugosa orientalis (Walcott) = {77 POETS ;
H. shantungensis
ce ? simplex Walcott

acuticosta pacifica Saito=H. pacifica

Helcionella rugosa (Hall)

Metoptoma ? rugosa Hatt, Pal. New York, vol. 1, p. 306, pl. 83, fig. 6, 1847.

Stenotheca rugosa Watcorr (part), U. S. Geol. Surv. Bull. 30, p. 128, pl. 12,
figs. 1b, c, 1886 [not fig. 1 = H. curticei; 1a= H. fordi; 1d, e= Stenothe-
coides troyensis]; idem, toth Ann. Rep., p. 617, pl. 74, figs. 1b, c, 1891
[not fig. 1=H. curticet; 1a =H. fordi; 1d =H. sp.; 1e= H. walcotti;
if, g =H. halli; th, i= Stenothecoides troyensis].

Helcionella rugosa Grabau and Shimer (part), N. A. Index Foss., vol. 1
p. 607, fig. 81ob, 10909.

,

Great confusion was introduced into this species by illustration
of the New York forms in 1886 and 1891. After that other workers
assumed these animals to be highly variable and referred many
diverse forms to //. rugosa.

Available specimens were segregated according to kind and locality.
Thereupon, several species appeared as clearly defined units and are
here defined. H. rugosa occurs only in the Hudson valley; at all
other localities throughout the world the name was misapplied.

After careful comparison the name rugosa was restricted to the
form which agrees with Hall’s original description and illustration.
Possibly if the original type could be studied this choice might have
to be altered.

Lower Cambrian, Schodack; (loc. 27) Troy, and other localities,
New York.

Holotype —A.M.N.H. no. 212; plesiotypes, U.S.N.M. no. 15365.

Helcionella halli, n. sp.
Stenotheca rugosa Watcotr (part), roth Ann. Rep. U. S. Geol. Surv., p. 617,
pl. 74, fig. 1g, 18901 [see H. rugosa].

This species has a more circular outline than any other from New
York. It is also high and has its apex overturned parallel to the base
of the shell.

Lower Cambrian, Schodack; (loc. 35) 14 miles north of Bald
Mountain, northwest of Greenwich, New York.

Holotype.—U.S.N.M. no. 96472.

NO. 10 CAMBRIAN FOSSILS, 4TH CONTRIBUTION——-RESSER 13

Helcionella walcotti, n. sp.
Stenotheca rugosa Watcott (part), toth Ann. Rep. U. S. Geol. Surv., p. 617,
pl. 74, fig. Ie, 1891 [see H. rugosa].

This is the most common species in the Hudson valley. It has the
coarsest ribs, and differs further in that the ribs are angular while
in other species they are rounded in contour.

Lower Cambrian, Schodack; (loc. 33) North Greenwich, and
many other localities in New York.

Holotype-—vU.S.N.M. no. 17456.

Helcionella curticei, n. sp.
Stenotheca rugosa Watcotr (part), U. S. Geol. Surv. Bull. 30, p. 128, pl. 12,
fig. 1, 1886; idem, toth Ann. Rep., p. 617, pl. 74, fig. 1, 1891 [see H.
rugosa).

The holotype lies in the matrix, besides a good example of
H. rugosa, so that it is easy to compare the two. H. curticei, which
is named for the collector, is twice as high as //. rugosa, has a more
rounded elliptical outline and coarser ribbing.

Lower Cambrian, Schodack; (loc. 27) Troy, New York.

Holotype-—U.S.N.M. no. 96473.

Helcionella fordi, n. sp.
Stenotheca rugosa Wawcotr (part), U. S. Geol. Surv. Bull. 30, p. 128, pl. 12,
fig. Ia, 1886; idem, toth Ann. Rep., p. 617, pl. 74, fig. 1a, 1891 [see H.
rugosa).

H. fordi has a much rounder elliptical outline than H. rugosa.
The rugosity is stronger than in H. curticet, from which H. fordi
further differs in greater relative height.

Occurrence same as preceding.

Holotype.—U.S.N.M. no. 96474.

Helcionella tatei, n. sp. :
Stenotheca rugosa Tate, Trans. Roy. Soc. South Australia, vol. 15, p. 183,
pl. 2, fig. 4, 1892.

This species is characterized by its wide base. The sides slope
evenly and the apex is overturned, but only near its tip. As shown
in the figure the ribs are angular, but they are not pronounced. There
are also vertical striations.

Lower Cambrian; Androssan, South Australia.

Holotype.—Australia; plastotypes, U.S.N.M. no. 96475.

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Helcionella sp.
Stenotheca rugosa WaAxcott (part), 1oth Ann. Rep. U. S. Geol. Surv., p. 617,
pl. 74, fig. Id, 1891 (see H. rugosa).

This Atlantic Province form referred to H. rugosa cannot be found
in the collections as it evidently never was properly marked as a type.
Until the specimen is located, its description cannot be written.

Lower Cambrian, Etcheminian; Conception Bay, Newfoundland.

Helcionella kiaeri, n. sp.
Helcionella cf. rugosa Krtarr, Skft. Vid. Kristiania, 1916, Mat.-Natur. K1.,
WO As ih uO, wl, 2, ane, 4 Woz

This is a rather stout, coarse-ribbed form but is not as large as
some of the New York species. Its outline is evidently nearly
circular.

Lower Cambrian, Strenuella limestone; Témten, Ringsaker,
Norway.

Holotype.—Pal. Mus. Oslo. no. 61b.

Helcionella norvegica, n. sp.
Helcionella rugosa acuticosta Krarr, Skit. Vid. Kristiania, 1916, Mat.-Natur.
IK, WO A, os Ae, wll Ay wes A, wel, won,

This is a small form characterized by strongly curved outlines front
and back, with the apex turned beyond go°. The ribs are sharply
angled.

Occurrence same as preceding.

Holotype-—Pal. Mus. Oslo no. 15.

Helcionella erecta (Walcott)
Stenotheca rugosa erecta WaA.cott, 10th Ann. Rep. U. S. Geol. Surv., p. 617,
pl. 74, fig. 4, 1801.

Lower Cambrian, Etcheminian; (loc. 41) Manuels Brook, Con-
ception Bay, Newfoundland.
Holotype-—U.S.N.M. no. 18311.

Helcionella acutacosta (Walcott)
Stenotheca ? rugosa acutacosta Wat.cortt, toth Ann. Rep. U. S. Geol. Surv.,
p. 617, pl. 74, figs. 2-2b, 1801.

Occurrence same as preceding.
Lectotype (fig. 2) and paratype —U.S.N.M. no. 18310.

Helcionella curvirostra (Shaler and Foerste)
Stenotheca curvirostra SHALER and Forrstr, Mus. Comp. Zool., Bull. 16,
p. 30, pl. 1, fig. 8, 1888; WaAtcott, 1oth Ann. Rep. U. S. Geol. Surv.,
De OLS. plaza ioe TOWTSOm.

NO. IO CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 15

Lower Cambrian, Hoppin; North Attleboro, Massachusetts.
Holotype —U.S.N.M. no. 96476.

Helcionella cobboldi, n. sp.
Stenotheca abrupta ? Copporp, Geol. Mag., dec. 6, vol. 4, p. 156, pl. 4, figs.
28, 29, I9I9.

This species is similar to H. abrupta, but differs in being more
slender and the ribs do not project so far beyond the marginal outline
of the shell.

Lower Cambrian, Hartshill; Nuneaton, Warwickshire, England.

Cotypes.—Sedgwick Mus.

Helcionella abrupta (Shaler and Foerste)
Stenotheca abrupta SHALER and Foerster, Bull. Mus. Comp. Zool., vol. 16,
p. 29, pl. 1, figs. 9a, b, 1888; Watcort, roth Ann. Rep. U. S. Geol. Surv.,
p. 617, pl. 74, figs. 6, 6a, I8or.

Only the smaller of Foerste’s specimens is to be found in the col-
lections. These types were never marked and the material is rather
fragile. The species must be restricted to the form first described.

Lower Cambrian, Hoppin; North Attleboro, Massachusetts.

Cotypes—U.S.N.M. no. 96477.

Helcionella alia, n. sp.
Stenotheca curvirostra GRABAU, Occ. Papers Boston Soc. Nat. Hist., vol. 4,
p. 638, pl. 31, fig. 13, 1900; GrRABAU and SHIMER (not Shaler and Foerste),
N. A. Index Foss., vol. 2, p. 373, fig. 1674d, 1910.

This species is stouter and longer than H/. curvirostra and has more
numerous ribs.

Lower Cambrian, Weymouth; Sandy Cove, Cohasset, Massa-
chusetts.

Holotype.—Boston Soc. Nat. Hist. no. 11964.

Helcionella grabaui, n. sp.
Stenotheca abrupta GRABAU (not Shaler and Foerste), Occ. Papers Boston
Soc. Nat. Hist., vol. 4, p. 637, pl. 31, figs. 12a-c, 1900; GRABAU and
Sumer, N. A. Index Foss., vol. 2, p. 373, figs. 1674a-c, 1910.

This is an erect species, but even then is more curved than 1.
abrupta. Grabau’s original figures do not agree one with the other,
but angle of view and the difference between mold of interior and
exterior would account for this. This species is characterized by the
large number of annulations, 10 being shown in the figure.

Lower Cambrian, Weymouth; Nahant, Massachusetts.

Cotypes.—Boston Soc. Nat. Hist. no. 11962.

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Helcionella pauper (Billings)
Stenotheca pauper Bittines, Canadian Nat., new ser., vol. 6, p. 479, 1872.
Stenotheca rugosa paupera Watcott, 10th Ann. Rep. U. S. Geol. Surv.,
DOLE ple 7A whe. 7 Leon.

This species must be restricted to the form from Newfoundland.
Billing’s type was not figured, but Walcott states that he compared
the type with the specimen he figured in 1891.

Lower Cambrian, Etcheminian; (loc. 41) Manuels Brook, Con-
ception Bay, Newfoundland.

Flolotype—Nat. Mus. Canada; plesiotype, U.S.N.M. no. 18312.

Helcionella foerstei, n. sp.
Stenotheca rugosa pauper SHALER and Foerster, Bull. Mus. Comp. Zodl.,
VO eLOm paecOn ples Teno aeeoo:

Compared with the specimen Walcott figured as H. pauper, the

Massachusetts species is much more curved and has stronger ribs.
Lower Cambrian, Hoppin; North Attleboro, Massachusetts.
Holotype —U.S.N.M. no. 96478.

Helcionella terranovica, n. sp.
Stenotheca ? rugosa levis Watcott (part), 1oth Ann Rep. U. S. Geol. Surv.,
p. 617, pl. 74, fig. 5, 1891 (see H. levis).

This is a much taller and stouter form than the other specimen
Walcott referred to his variety levis. This species is stouter and more
erect than H. levis and has fewer, much larger, annulations.

Lower Cambrian, Etcheminian; (loc. 41) Manuels, Conception
Bay, Newfoundland.

Holotype —U.S.N.M. no. 96479.

Helcionella striata, n. sp.
Stenotheca levis GRABAU (not Walcott), Occ. Paper Boston Soc. Nat. Hist.,
WOlly 4h joy hey joll, Gin, sties, 1G}, WoL

This form is shaped somewhat like H. terranovica, but the illustra-
tion shows pronounced longitudinal striations.

Lower Cambrian, Weymouth; Pleasant Beach, Cohasset, Massa-
chusetts.

Holotype.—Boston Soc. Nat. Hist. no. 11961.

Helcionella recurva, n. sp.
Stenotheca pauper GRaBAu (not Billings), Occ. Papers Boston Soc. Nat.
Hist., vol. 4, p. 639, pl. 31, fig. 14, 1900; Grabau and Shimer, N. A. Index
‘-Foss., vol. 2, p. 374, fig. 1674e, I9Q1o.

NO. IO CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 17

This species is much more curved than H. foerstei and besides
has coarser ribs.

Lower Cambrian, Weymouth; Sandy Cove, Cohasset, Massa-
chusetts.

Holotype.—Boston Soc. Nat. Hist. no. 11963.

Helcionella levis (Walcott)
Stenotheca ? rugosa levis WaA.cotr (part), 10th Ann. Rep. U. S. Geol. Suryv.,
PAOLA ple 74s nes) Sa) 1oOr.

Walcott included two forms in his variety levis. The smoother
one is chosen to represent the species and the other described as
FH. terranovica.

Lower Cambrian, Etcheminian; (loc. 41) Manuels, Conception
Bay, Newfoundland.

Holotype—vU.S.N.M. no. 18313.

Helcionella elevata (Cobbold)
Scenella elevata CoppBoLtp, Quart. Journ. Geol. Soc. London, vol. 76, pt. 4,
p. 364, pl. 24, fig. 36, 1921.

Cobbold pointed out that this species did not conform to the re-
quirements of Scenella but because it lacked pronounced concentric
rugosities reference to Helcionella was more difficult. Now that wide
variation in this respect is known to exist in Helcionella, the shape
of H. elevata may be relied on and the species referred to Helcionella.

Lower Cambrian, Olenellus limestone; Comley, Shropshire,
England.

Holotype.—Geol. Surv. no. RR1211.

Helcionella wheeleri (Walcott)
Stenotheca wheeleri Watcott, Canadian Alpine Journ., vol. 1, no. 2, pl. 1,
fig. 7, 1908. :

Middle Cambrian, Stephen; (loc. 14s) Mount Stephen, near Field,
British Columbia.
Holotype —U.S.N.M. no. 96480.

Helcionella belliana (Walcott)
Platyceras (?) bellianus Wavcorr, Canadian Alpine Journ., vol. 1, no. 2,
pl. 1, fig. 13, 1908.

Occurrence same as preceding.

Holotype —U.S.N.M. no. 96481.

Helcionella romingeri (Walcott)
Platyceras romingeri Watcort, Proc. U. S. Nat. Mus., vol. 11, p. 442, 1888;
Canadian Alpine Journ., vol. 1, no. 2, pl. 1, fig. 14, 1908.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Middle Cambrian, Stephen; (loc. 14s) Mount Stephen, near Field,
British Columbia.
Holotype —U.S.N.M. no. 96482.

Helcionella orientalis (Walcott)
Stenotheca rugosa orientalis Waucott, Proc. U. S. Nat. Mus., vol. 29, p. 16,
1905.
Helcionella rugosa orientalis Watcotr (part), Research in China, vol. 3,
Carnegie Inst. Publ. 54, p. 91, pl. 5, fig. 15a, 1913 (not fig. 15=H.
shantungensis ).

The species is restricted to the form at the first locality, which ac-
cording to Dr. Walcott’s practice is the type locality.

Middle Cambrian, Changhia; (loc. C21) Changhia, Shantung,
China.

Holotype —U.S.N.M. no. 57776.

Helcionella shantungensis, n. sp.
Helcionella rugosa orientalis Wa.cotrt (part), Research in China, vol. 3,
Carnegie Inst. Publ. 54, p. 91, pl. 5, fig. 15, 1913 (see H. orientalis).

This species is wider at the base than H. orientalis and the rugosi-
ties are not so coarse. The striations are also finer and weaker.

Middle Cambrian, Changhia; (loc. C18) east of Changhia, Shan-
tung, China.

Holotype—U.S.N.M. no. 57775.

Helcionella triangularis (Matthew)
Stenotheca triangularis MATTHEW, Trans. Roy. Soc. Canada, vol. 3, sec. 4,
p. 58, pl. 6, figs. 15, 15a, 1886.
Middle Cambrian, St. John; Hanford Brook. New Brunswick.
Holotype and paratype-—Walker Mus. (?).

Helcionella manuelensis (Matthew)
Plumulites manuelensis MATTHEW, Trans. New York Acad. Sci., vol. 15,
p. 145, fig. 1, 1896.
Middie Cambrian, Manuels; Manuels Brook, Conception Bay,
Newfoundland.
Holotype—Walker Mus. (?°).

SCENELLA Billings, 1872

Diverse forms are referred to Scenella, particularly post-Cambrian
species. A list of Cambrian species referred to the genus is given
with cross references.

NO. I0 CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER Ig

Atlantic Province
Scenella antiqua Kiaer
= depressa Kiaer
? discinoides Schmidt (possibly two species )
elevata Cobbold = Helcionella elevata
reticulata Billings
? tuberculata Schmidt
North America
Scenella conula Walcott
: retusa Ford = (part = Paterina troyensis)
varians Walcott = (part S. amit)
amu Matthew

“cc
“
“ce

“ce

“cc

“cc

China
Scenella clotho Walcott = Scenellopsis clotho
‘a Salto" aesaeton
dilatatus Walcott = S. dilatata

“

Scenella amii (Matthew)

Metoptoma amit MattHeEw, Trans. Roy. Soc. Canada, 2nd ser., vol. 8, sec. 4,
(D> Wily, FN Ty ve, Uz Io)

Scenella varians Watcort, Canadian Alpine Journ., vol. 1, no. 2, pl. 1, fig. 6,
1908.

[Not Scenella varians Watcott, U. S. Geol. Surv. Bull. 30, p. 127, pl. 12,
figs. 2, 2a, 1886.]

Middle Cambrian, Stephen; (loc. 14s) Mount Stephen, near Field,
British Columbia.

Holotype-—Walker Mus. (?) ; plesiotype, U.S.N.M. no. 96483.

Scenella columbiana (Walcott)
Crania ? columbiana Watcortt, Proc. U. S. Nat. Mus., vol. 11, p. 441, 1889.
Philhedra ? columbiana voN HUuENE, Verhandl. Russ.-kais. Min. Gesell. St.
Petersburg, 2nd ser., vol. 36, pt. 2, pp. 216, 208, 1890.
Philhedra columbiana Watcortt, U. S. Geol. Surv. Mon. 51, p. 724, pl. 81,
fig. 10, IQ12.

Walcott reports the finding of two additional imperfect specimens
in 1907. Examination of these specimens leads one to doubt their
identity with the type. It is possible that they are merely the apical
portions of Nisusia.

The type is a minute form not very well preserved. It certainly
is not a brachiopod, so the best reference is to Scenella. This speci-
men may be merely a poorly or rather unusually preserved example
of Scenella ami. However, since no other such example has been
noted among the hundreds of Scenella ami specimens in the collec-
tion, this form may retain its specific rank.

Middle Cambrian; (loc. 14s) Mount Stephen, above Field, British
Columbia.

Holotype-—U.S.N.M. no. 58307.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

SCENELLOPSIS, n. gen.

A peculiar form from China, referred with reservation to Scenella,
seems to be a gastropod.

Scenellopsis is a small limpetlike shell with somewhat eccentric
apex. Both growth and radiating lines are present but are preserved
in two different specimens. Muscular impression, evidently branch-
ing, produces folds on the outer surface.

Genotype.—Scenella clotho Walcott.

Scenellopsis dilatata (Walcott)
Scenella ? dilatatus Watcott, Proc. U. S. Nat. Mus., vol. 30, p. 570, 1906;
Research in China, vol. 3, Carnegie Inst. Publ. 54, p. 87, pl. 5, figs. 2, 2a,
1913. '
Middle Cambrian, Changhia; (loc. C70) southeast of Tungyu,
Shansi, China.
Cotypes—U.S.N.M. nos. 57767, 8.

Scenellopsis clotho (Walcott)
Scenella clotho WaAucortt, Proc. U. S: Nat. Mus., vol. 20, p. 12, 1905; Research
in China, vol. 3, Carnegie Inst. Publ. 54, p. 86, pl. 5, figs. 3, 3a, 1913.

Middle Cambrian, Changhia; (loc. C18) East of Changhia, Shan-
tung, China.
Holotype —U.S.N.M. no. 57769.

Scenellopsis saitoi, n. sp.
Scenella clotho Sarro (not Walcott), Journ. Faculty Sci. Imp. Univ. Tokyo,
See, 2, -vol.4, pe 3: p-°350; pl. 3, hes. 18; 10;-1036:

This species resembles S. clotho, but, as pointed out by Saito, the
apex is not as high and the apertural margin is less upturned. Saito’s
figures show that the marginal outline is also much more flattened at
the broad end.

Lower Cambrian, Misaki; near Chungwa, northern Chosen.

Holotype—Geol. Inst. Imp. Univ. Tokyo.

HYOLITHES Eichwald, 1840

Eichwald established Hyolithes on H. acutus from Ordovician strata
of Estonia. The genus seems to be valid for other species extend-
ing from the Lower Cambrian into the Devonian. In the earlier
years Hyolithes was confused with various cephalopod genera, par-
ticularly Orthoceras but this confusion was cleared up long ago. Sev-
eral authors attempted classification of the family but no one has
completed the job.

NO. 10 CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER ai

Matthew erected the genus Camarotheca for the hyolithid forms
which show internal septa and a structure analogous to the cephalopod
siphuncle. Examination of many species shows that cameration de-
pends on preservation and may be found in any species. The other
genera excepting Camarotheca are apparently all valid, or at any rate
express definite and distinctive structures.

Orthotheca Novak is valid only for Ordovician organisms, which
are unrelated to the rounded tubes ofttimes placed in Hyolithes.
Therefore, the practice, followed by the author and others, of as-
signing tapered tubes with circular cross-section to Orthotheca, is
improper.

Revision of Cambrian Hyolithidae is greatly needed. Not only
must a series of genera be established but the species themselves have
been so carelessly handled that most are of little value. Recent strati-
graphic studies occasioned review, in part, of H. primordialis and
H. billingst. Several of the resulting nomenclatural changes are
recorded below.-

Hyolithes gregarius (Meek and Hayden)
Theca (Pugiunculus) gregaria MEEK and HAypeN, and Proc. Acad. Nat.
Sci. Philadelphia, p. 436, 1861; Amer. Journ. Sci., 2d ser., vol. 33, p. 73,
text fig. 3, 1862.
Theca gregaria MrEK and HAynEN, Pal. Upper Missouri, Smithsonian Contr.
Knowl., vol. 7, p. 5, text figs. a-d, 1865.
This species averages much smaller than H. primordialis to which
it was referred and, therefore, is again recognized.
Upper Cambrian, Depass; Powder River, Big Horn Mountains,
Wyoming.
Cotypes.—U.S.N.M. no. 1181.

Hyolithes gallatinensis, n. sp.
Hyolithes primordialis Wawcott (part), U. S. Geol. Surv. Mon. 32, pt. 2,
p. 454, pl. 63, fig. 2, 1800.

This wide species is characterized by a very slightly convex pos-
terior surface and an evenly, semicircular anterior side. The surface
is smooth in the specimens available but they may be interiors only.
The lingual extension is broken away.

Upper Cambrian, Dry Creek; (loc. 151c) Crowfoot Ridge, Galla-
tin Range, Yellowstone National Park, Wyoming.

Holotype —U.S.N.M. no. 35218.

Hyolithes whitei, n. sp.
Hyolithes primordialis ? Wuite, Geogr. Geol. Expl. Surv. West tooth Merid.,
Prelim: Rep. Invert. Foss: p- 6; 1874, Idem, vol. 4) pt. 1, py 37,-plh. 1,
figs. 5a-e, 1875.

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Hyolithes billingsi Waxcotr (part), U. S. Geol. Surv. Bull. 30, p. 134, pl. 13,
figs. 1, ta-d, 1886; idem, 1oth Ann. Rep., p. 620, pl. 75, figs. 1, 1a-d,
1891; Lestry (part), Geol. Surv. Pennsylvania, Rep. P4, p. 294, figs.
1889; GraBAu and Suimer, N. A. Index Foss., vol. 2, p. 3, figs. 1211e-g,
1Q10.

HI, whitei is a rather small species with a thick shell. The pos-
terior side is nearly flat; the anterior is angulated giving the tube a
nearly equilateral triangular outline. The lateral corners are rounded.

Lower Cambrian, Pioche; (loc. 31a) southeast of Pioche, High-
land Range, Nevada.

Cotypes——U.S.N.M. nos. 8579, 15377.

Hyolithes poulseni, n. sp.
Hyolithes billingst Poutsen, Meddels. Grgnland, vol. 70, p. 254, pl. 15, figs.
A, Fil, MOA7,
Hyolithus (Hyolithus) billingst Poutsen, idem, vol. 87, no. 6, p. 22, pl. 3,
fig. 6, 1932.

This species has a flat posterior side and a rounded anterior face.
It expands much more rapidly and is larger than H. billingsi. The
anterior surface is striated.

Lower Cambrian, Cape Kent; Cape Kent, North Greenland.
Bastion ; Hyolithus Creek, East Greenland.

Cotypes and plesiotypes——Min. Mus. Copenhagen; plastotypes,
US. NIM ne; 70055:

CRUSTACEX
STENOTHECA Salter, 1872

Stenotheca Hicks (Salter MSS.), Quart. Journ. Geol. Soc. London, vol. 28,
Dp. 180; 1872; Watcort, U.*S. Geol. Surv. Bully) 30, ps 128; 1886:
MattTHew, Trans. Roy. Soc. Canada, vol. 8, p. 132, 1891; GRABAU and
Suimer, N. A. Index Foss., vol. 2, p. 373, 1910; Copsotp, Geol. Mag.,
vol. 71, p. 463, 1934.

Watsonella Graspau, Occ. Pap. Boston Soc. Nat. Hist., vol. 4, p. 631, 1900.

Stenotheca, a generic name attached to museum specimens, was
published without description in 1872. In 1886 Walcott described
the genus but quite clearly based his remarks on Helcionella rugosa
and its allies. Consequently his description is that of the gastropod
genus Helcionella. More recently Cobbold restudied Salter’s types
and presented a generic diagnosis.

In 1891 Matthew recognized the crustacean characters of Steno-
theca, in which he was followed by Grabau and Shimer. The latter,
however, assigned only gastropod species to the genus. Cobbold
assigns the genus to the Notostraca.

NO. 10 CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 23

It will be observed that Cobbold’s generic description of Stenotheca
agrees in every detail with that of Watsonella Grabau. The illustra-
tions also agree ; therefore Watsonella must be regarded as a synonym
of Stenotheca.

All known species of Stenotheca are confined to the Atlantic
province.

Genotype.—S. cornucopia Salter.

The following are regarded as valid species of Stenotheca:

. angusta Cobbold

. concentrica Matthew
. cornu Wiman

. cornucopia Salter

. crosbyit (Grabau)

. hicksiana Matthew
. lata Cobbold

. nasuta Matthew

. radiata Matthew

HHHHY
HAHY

Species formerly referred to Stenotheca are given, together with
their synonyms.

. abrupta ? Cobbold = Helcionella cobboldi
. abrupta Grabau and Shimer = H. grabaui
. abrupta Shaler and Foerste = H. abrupta
. (Parmorphorella) acadica Matthew = Parmorphorella acadica
. clurius Walcott = Helcionella ? clurius
. curvirostra Grabau = H. alia
curvirostra Shaler and Foerste= H. curvirostra
Stenothecoides elongata
S. labradorica
. levis Grabau = Helcionella striata
. pauper Billings =H. pauper
. pauper Grabau =H. recurva
. rugosa Billings = H. rugosa
H. curticet
H. fordi
. rugosa Walcott =< H. walcotti
Hi. halli
Stenothecoides troyensis
. rugosa Sears = Helcionella abrupta
. rugosa Tate=H. tatei
? rugosa acutacosta Walcott = H. acutacosta
. rugosa aspera Noetling = Pseudotheca waagen
rugosa erecta Walcott = Helcionella erecta
rugosa chinensis Walcott = H. chinensis
? rugosa levis Walcott = H. levis
rugosa orientalis Walcott = H. orientalis
rugosa pauper Shaler and Foerste = H. foerstei
. rugosa paupera Walcott = H. pauper
. simplex Walcott =H. ? simplex
. triangularis Matthew = H. triangularis
. wheeleri Walcott=H. wheeleri

HH

. elongata Waleott ={

H AHHH H HHHHY

HAUHHHHHHNHHHHY

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Stenotheca crosbyi (Grabau)
Watsonella crosbyi GRABAU, Occ. Pap. Boston Soc. Nat. Hist., vol. 4, p. 632,
pl. 31, figs. 9a-f, 1900.
Lower Cambrian, Weymouth; Pleasant Beach and Sandy Cove,
‘Cohasset, Massachusetts.
Cotypes.—Boston Soc. Nat. Hist. nos. 11951-11954.

STENOTHECOIDES, n. gen.

Stenothecoides is related to Stenotheca, but is much narrower and
longer, with, a relatively wider aperture. The genus is characterized
by the long narrow carapace which is curved longitudinally. This ~
curvature is usually much greater in cross-section, amounting to a
fold along the apex. The apertural outline increases from the narrow
anterior end almost evenly to a broadly rounded rear margin. The
marginal outline of some species is not bilaterally symmetrical, curv-
ing out more on one side than the other. The surface is marked with
growth lines surrounding the anterior end, each extending itself rear-
ward at a greater rate, so that while the anterior ends remain close
together, the lines become longer and longer, reaching to the posterior
portion of the test.

Genotype.—Stenotheca elongata Walcott.

Stenothecoides elongata (Walcott)

Stenotheca elongata Watcort, U. S. Geol. Surv. Mon. 8, p. 23, pl. 9, figs. 2,
2a, 1884; idem (part), Bull. 30, p. 120, pl. 12, figs. 4a, b, 1886 (not fig.
4== S. labradorica).

Helcionella elongata Watcort, Smithsonian Misc. Coll., vol. 67, no. 3, p. 63,
1017.

Middle Cambrian, Eldorado ; (ee. 55b) West slope Prospect Peak,

Eureka District, Nevada.
Holotype —U.S.N.M. no. 15364.

Stenothecoides labradorica, n. sp.
Stenotheca elongata Watcotr (part), U. S. Geol. Sury. Bull. 30, p. 129,
pl. 12, fig. 4, 1886.

This species was identified with that from the Eureka District,
Nevada, but the drawing shows that it is clearly distinct. S. labra-
dorica is shorter and wider and the apertural margin evidently is not
as symmetrical as S. elongata. The growth lines are also somewhat
weaker.

Lower Cambrian, Forteau; L’Anse au Loup, Labrador.

Holotype —U.S.N.M. no. 14883.

NO. 10 CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER

LS)
wn

Stenothecoides troyensis, n. sp.
Stenotheca rugosa Watcort (part), U. S. Geol. Surv. Bull. 30, p. 128, pl. 12,
figs. Id, e, 1886 (see Helcionella rugosa) ; idem, 1oth Ann. Rep., p. 617,
ple 74,85. Th, 1, tka.

This a very small species, and is much wider than any of the others.
Longitudinal curvature is accentuated at the anterior end so that a
slightly recurved apex is developed.

Lower Cambrian, Schodack; (loc. 27) Troy, and other localities
in New York.

Holotype —U.S.N.M. no. 96484.

Stenothecoides poulseni, n. sp.
Undetermined lamellibranch Poutsen, Meddels Grgnland., vol. 87, no. 6, p. 29,
pl. 7, figs. 1-4, 1932.

It is possible that more than one species is represented by the speci-
mens illustrated. Growth lines are coarse and irregular. This species
is so constricted laterally toward the anterior end as to form a distinct
apex.

Lower Cambrian, Ella Island; south coast Ella Island, east
Greenland.

Cotypes——Min. Mus. Copenhagen.

' TUZOIA Walcott, 1912

Tuzoia argenta (Walcott)
Leperditia? argenta Watcott, U. S. Geol. Surv. Bull. 30, p. 146, pl. 8,
fig. 5, 1886.

Middle Cambrian, Ophir; (loc. 30a) 1 mile below Argenta, Big
Cottonwood Canyon, Wasatch Mountains; and (loc. 54a) Blacksmith
Fork, Bear River Range, Utah.

Holotype —U.S.N.M. no. 15401.

UNCERTAIN POSITION
UROTHECA Matthew, 1899

Urotheca parasitum, n. sp.
Hyolithellus flagellum Watcott (part), Smithsonian Misc. Coll., vol. 67,
MO, By jo, Ad, la Se ankeeeel, woz:

This species consists of narrow, long flexible tubes which had one
or more longitudinal grooves. It is a much smaller tube than U.
flagellum. The type and other specimens grew on shells of Wimanella,
which raises the question whether all species of Urotheca had such
growth habit.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Middle Cambrian, Ptarmigan; (loc. 63}) Popes Peak, 14 miles
southwest of Stephen, and (loc. 35c) Mount Bosworth, British
Columbia.

Cotypes—U.S.N.M. no. 63723.

COLEOLOIDES Walcott, 1890
Coleoloides hectori (Walcott)
Hyolithellus hectori Watcott, Smithsonian Misc. Coll., vol. 67, no. 2, p. 27,
Pls, ose Oz.

This species is referred to Coleoloides on the basis of its shape and
the presence of striations. It differs from C. typicalis in that the
striations are coarser, and the tube itself is of greater diameter. Too
little of the tube preserves the striated outer surface to show whether
the striations in C. hectori have a spiral course, but they seem to be
slightly off parallel position in the short portion preserved.

Middle Cambrian, Ross Lake; (loc. 35c) 1 mile east of Hector,
Mount Bosworth, British Columbia.

Holotype —U.S.N.M. no. 63721.

AGNOSTIDA
AGNOSTUS Brongniart, 1822
Agnostus yellowstonensis, n. sp.
Agnostus bidens Watcortt, U. S. Geol. Sury. Mon. 32, pt. 2, p. 455, pl. 63,
figs. 4, 4a, 1890.

This species is not the same as the Middle Cambrian 4. bidens
Meek. It is characterized by strong dorsal furrows in both shields,
the anterior glabellar lobe being sharply separated. The published
illustration of the pygidium is misleading because the artist failed
to eliminate perspective when drawing the convex shield and there-
fore extended the axis too far rearward. In fact the space between
the rhachis and rear margin is half the width of the pleural lobes.

Upper Cambrian, Dry Creek; (loc. 151¢) Crowfoot Ridge, Gallatin
Range, Yellowstone National Park, Wyoming.

Cotypes.—U.S.N.M. no. 35222.

TRIEOBITA
BAILIELLA Matthew, 1885
Bailiella Resser, Smithsonian Misc. Coll., vol. 95, no. 4, p. 15, 1936.

Several species must be added to the list published in 1936. Some
of them were overlooked, one correction was expected to appear else-
where and others have since been described under Conocoryphe.

NO. IO CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 27

Bailiella artagena (Howell)
Conocoryphe? artagena Howe tt, Bull. Geol. Soc. Amer., vol. 48, no. 8, p. 1169,
pl. 3, fig. 7, 1937.

Middle Cambrian, St. Albans; St. Albans, Vermont.
Holotype.—Princeton Univ. no. 9925.

Bailiella frangtengensis (Reed)

Conocoryphe frangtengensis REED, Mem. Geol. Surv. India, Pal. Ind., new ser.,
vol. 21, mem. 2, p. 7, pl. 2, figs. 9-12, 1934.

Middle Cambrian; Frangteng Hill, Hundwara, Kashmir, India.
Cotypes—Geol. Surv. India nos. 15593-6.

Bailiella lantenoisi (Mansuy)
Conocoryphe lantenoist MANsuy, Mem. Serv. Geol. I’Indo-Chine, vol. 5, fasc.
I, p. 30, pl. 4, figs. 6, 7; pl. 5, fig. 3, 1916; KoBayAsuI, Journ. Faculty
Sci. Imp. Univ. Tokyo, sec. 2, vol. 4, pt. 2, p. 218, pl. 23, figs. 13, 14, 1935.

Middle Cambrian ; Tien-fong, Tonkin.

Bailiella sejuncta (Reed)
Conocoryphe sejuncta Reep, Mem. Geol. Surv. India, Pal. Ind., new ser.,
vol. 21, mem. 2, p. 8, pl. 2, figs. 13, 14, 1934.

Middle Cambrian; Wadapur, Hundwara, Kashmir, India.
Cotype—Geol. Surv. India nos. 15597-8.

Bailiella ulrichi (Resser and Endo)
Conocoryphe ulrichi Resser and Enpo, in Endo, Iwanami Lecture Ser., Geol.
Pal., p. 54, figs. 5-8, December 1031.

Middle Cambrian, Tangshih ; Tang-shih-ling, 2 miles southeast Yen-
tai Colliery, Manchoukuo.

BLOUNTIA Walcott, 1916

Blountia polita, n. sp.
Arionellus sp. Watcott, U. S. Geol. Surv. Mon. 32, pt. 2, p. 463, pl. 65, fig. 2,
1899.

Walcott illustrated this pygidium as a cranidium. The artist drew
“glabellar” furrows, but when his pencil marks were washed off the
specimen, no trace of axial furrows remain.

B. polita is characterized by its flat lateral profile, and the rather
elongate shape due to the lack of narrowing as the pleural lobes
unite behind the axis. The dorsal furrow is shallow, but the axis is
marked rather clearly by a change in the profile.

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Upper Cambrian, Pilgrim; (loc. 151b) between Pebble and Soda
3utte Creeks, Yellowstone National Park, Wyoming.
Holotype —U.S.N.M. no. 96488.

BRISCOIA Walcott, 1924

Briscoia texana (Walcott)
Dikelocephalus texanus Watcott, Smithsonian Misc. Coll., vol. 57, no. 13,
Pp. 372, pl. 65, fig. 4, Ior4.
Upper Cambrian, Wilberns; (loc. 70a) Baldy Mountain, near
Morgans Creek, 8 miles northwest of Burnet, Texas.
Holotype —U.S.N.M. no. 58621.

CHARIOCEPHALUS Hall, 1863

Chariocephalus Hatt, 16th Ann. Rep. New York State Cab. Nat. Hist., p. 175,
1863.
Dartonaspis MILLER, Journ. Pal., vol. 10, no. 1, p. 29, 1936.

Miller failed to observe that the criteria on which he based Dar-
tonaspis are exactly those which led to the separation of Jrvingella
from Chariocephalus. These relationships are discussed under
Irvingella.

Chariocephalus knighti (Miller)
Dartonaspis knighti Mixer, Journ. Pal., vol. 10, no. 1, p. 20, pl. 8, figs. 34, 35,
1936.

Upper Cambrian, Boysen; Bull Lake Creek, Wind River Range,
Wyoming.
Holotype-—Columbia Univ. no. 12626.

CLEVELANDELLA Resser, 1938

Clevelandella volux (Walcott)
Saratogia volux Watcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 108
pl. 35, figs. 2, 2a, 1916.

’

Upper Cambrian, Eau Claire; (loc. 78a) Eau Claire, Wisconsin.
Holotype -—U.S.N.M. no. 61714.

CHUANGIA Walcott, 1911

Chuangia suni, n. sp.
Chuangia batia Sun, Pal. Sinica, ser. b, vol. 1, fasc. 4, p. 58, pl. 4, figs. 4a-e,
1924.
This species is narrower than C. batia. The eyes are situated
somewhat more forward and are perhaps a little larger. The species

NO. IO CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 29

also has a wider preglabellar area. Glabellar furrows are evidently
very faint.

Upper Cambrian, Kaolishanian; Pagoda Hill, Taianfu, Shan-
tung, China.

Cotypes——Geol. Surv. China nos. 579-583.

DEADWOODIA, n. gen.

This trilobite genus is represented by a single cranidium. How-
ever, it appears among undescribed material of Ironton age, from
several localities and consequently description is warranted.

The cranidium is characterized by a very large, nearly quadrate
glabella, highly arched in both directions and standing high above the
fixigenes. Glabellar furrows essentially absent. Occipital furrow
deep, but narrow. Brim is wide, concave as a whole. Preglabellar —
area slightly convex, dipping steeply down from the dorsal furrow.
Anterior furrow shallow, with the essentially flat rim turned nearly
into a horizontal position. Fixigenes narrow, as well as the postero-
lateral limbs. Eyes rather large, strongly bowed, and palpebral lobes
separated by deep furrows.

Genotype.—Ptychoparia (Liostracus) panope Walcott.

Deadwoodia panope (Walcott)
Ptychoparia (Liostracus) panope Watcott, Proc. U. S. Nat. Mus., vol. 13,
p. 275, pl. 21, fig. 13, 1890.

Upper Cambrian, Deadwood (Ironton equivalent) ; Spring Creek
Canyon, about 7 miles southwest of Rapid City, Black Hills, South
Dakota. |

Holotype —U.S.N.M. no. 23856.

EHMANIA Resser, 1935

Ehmania ? agatho (Walcott)
Asaphiscus ? agatho Watcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 391,
pl. 63, figs. 9, 9a, 1916.

Upper Cambrian, Nolichucky; (loc. 123a) Big Creek, 4 miles
northeast of Rogersville, Tennessee.

Cotypes—U.S.N.M. nos. 62819-20.

ELVINIA Walcott, 1924

Elvinia roemeri (Shumard)
Dikelocephalus roemert SHUMARD, Amer. Journ. Sci., 2d ser., vol. 32, p. 220,
1861.

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Elvinia roemert Wa.cotr (part), Smithsonian Misc. Coll., vol. 75, no. 2,
p. 56, 1924; idem, no. 3, p. 88, 1925; BrinGe (part), U. S. Geol. Surv.
Prof. Pap., 186-M, p. 251, pl. 67, figs. 2a, b, 3a, b; pl. 69, figs. 1-8, 10, 15,
1937.

E. roemeri is not a widespread species, but many other forms have
been mistakenly identified with it. Without Shumard’s specimens it
is impossible to know which Texas species received the name E.
roemeri. Bridge studied Roemer’s original material and commented
on the other species of the genus, but he failed to select a type for
FE. roemerit. Inasmuch as Shumard’s types are lost and Shumard
specifically states that figure 2a of Roemer is the species he was de-
scribing, it is logical to choose that specimen as the lectotype. Bridge
went so far as to say that this specimen “is the nearest thing to a
type that exists.”

Upper Cambrian, Wilberns; $ mile east of Camp San Saba and
(loc. 141) 1 mile west of Cherokee, in San Saba County; (loc. 70)
Morgans Creek, 8 miles northwest of Burnet, Texas.

Lectotype and paratypes ——Univ. Bonn; casts, U.S.N.M. no. 95485.

Elvinia texana, n. sp.
Elvinia roemeri BrivcE (part), in Sellards, Adkins, and Plummer, Texas
Univ. Bull; 3232; p: 323, pls 2) figs: 17, 18; 31933; BRIDGE, (part). U.S
Geol. Surv. Prof. Pap. 186-M, p. 251, pl. 69, fig. 15, 1937.

This form is about the same size as E. roemeri but is relatively
longer. The width across the fixigenes immediately in front of the
eyes is less than in /£. roemeri, and the profile is less highly arched
in both directions. The glabella of E. texvana is rather sharply trun-
cate in front, and the anterior facial sutures extend almost straight
forward.

No pygidium has been definitely assigned to this cranidium. Sev-
eral are present in the collection, but only one is good enough to as-
sign to a species. By its size and other features this pygidium seems
to be more logically referred to a larger species.

Upper Cambrian, Wilberns; (loc. 68) Packsaddle Mountain, 12
miles southeast of Llano, Texas.

Holotype-—U.S.N.M. no. 93013.

Elvinia shumardi, n. sp.

Elvinia roemert Watcotr (part), Smithsonian Misc. Coll., vol. 75, no. 2,
p56, pk rr, fig: 3, 1924; adem, no. 35 p. 88, pl: a7, figs.) Os13- e10258
BripcGeE (part), in Sellards, Adkins, and Plummer, Texas Univ. Bull.
3232, p. 323, pl. 2, fig. 19, 1933; BripcE (part), U. S. Geol. Surv. Prof.
Pap. 186-M, p. 251, pl. 60, figs. 9, 10, 1937.

NO. 10 CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER or

This is a large species, attaining an average size about twice that
of E. roemeri. The anterior suture diverges more than E. texana,
although the relative width immediately in front of the eyes is about
the same. FE. shumardi is decidedly flat in lateral profile. Longitudi-
nally the species has a highly arched profile, but without being greatly
curved. The neck furrow is discontinuous, a distinctive feature of
the species.

It should be added that Walcott selected this species to represent
Shumard’s E. roemeri. This selection cannot stand because Roemer’s
figure 2a was specifically mentioned by Shumard.

Upper Cambrian, Wilberns; (loc. 70) Morgans Creek, about 8
miles northwest of Burnet; and (loc. 68) Packsaddle Mountain, 12
miles southeast of Llano, Texas.

Holotype —U.S.N.M. no. 70259; paratypes, no. 70260, I.

Elvinia bridgei, n. sp.
Elvima roemeri Bripce (part), U. S. Geol. Surv. Prof. Pap. 186-M, p. 251,
pl. 60, figs. 19-21, 1937.

Unfortunately only an incomplete cranidium of this species was
illustrated. Although none of the numerous cranidia in the collection
are complete, many are more so than the specimen illustrated. How-
ever, this cranidium shows sufficient of the width of the head and
the depth of the furrows to characterize the species. The occipital
furrow is deeper and wider than shown. The eyes are in a position
about parallel to the course of the dorsal furrow. The preglabellar
area is bulged so that the rather straight anterior furrow is deep,
and is marked by irregular longitudinal ridges. The rim is rather
wide and swollen, separated by a deep anterior furrow. This species
is much more highly arched longitudinally than laterally.

The associated pygidium is of normal type, and has the rim turned
up rather sharply.

There are several other species of Elvinia in the Oklahoma collec-
tions, but, since they require illustration, are not included in this
paper.

Upper Cambrian, Honey Creek; (loc. 89v) NE. 4 Sec. 9, T. 1 S.,
R. 1 W., 4 miles southeast of Hennepin, Arbuckle Mountains,
Oklahoma.

Cotypes.—U.S.N.M. no. 93025.

Elvinia missouriensis, n. sp.
Elvinia roemeri Briwce (part), U. S. Geol. Surv. Prof. Pap. 186-M, p. 2511,

PleeOO, fies. 12, 13) 1037.

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Bridge figured one of the two species present at the locality. The
holotype is a well-preserved cranidium.

This species is characterized by its relative narrowness immedi-
ately in front of the eyes. The eyes are not parallel to the dorsal fur-
row, but make a wide angle with the axis. Eyelines are rather heavy.
The cranidium is arched considerably laterally, this being accentuated
by the pronounced downward depression of the anterior angles of
the brim. Longitudinally the head is also highly arched, the greatest
curvature being in the anterior half. The preglabellar area is bulged
a bit and the thickened rim is sharply upturned.

Upper Cambrian, Davis; (loc. 11k) near shaft of Federal Lead
Mine No. 4, Flat River, Missouri.

Holotype —U.S.N.M. no. 93011.

Elvinia dakotensis, n. sp.
Elvinia roemeri Bripcr (part), U. S. Geol. Surv. Prof. Pap. 186-M, p. 251,
pl. 60, figs. 14, 16, 1937.

This species is nearest like E. missouriensis, differing in the round-
ing of the anterior portion of the glabella and the slightly narrower
width immediately in front of the eyes. E. dakotensis also has less
rugged relief in the various portions of the brim. In FE. dakotensis
the glabella does not rise so much as the fixigenes, and the longi-
tudinal curvature, which is considerable, forms a rather even curve.

The libragenes are large and wide, with a strongly curved outer
margin, particularly toward the genal angle. The genal spine is short
and small. The associated pygidium has an axis which stands well
above the flat pleural lobes.

Upper Cambrian, Deadwood; (loc. 88a) northern part of Dead-
wood, Black Hills, South Dakota.

Cotypes—U.S.N.M. no. 93024.

Elvinia utahensis, n. sp.
Elvinia roemert Brivce (part), U. S. Geol. Surv. Prof. Pap. 186-M, p. 251,
Dk Co) inky, 2A, 10S ye

This species is characterized by rugged relief due to the depth of
all furrows, the arching of the preglabellar area and of the fixigenes
and the depression of the anterior angles of the brim. The eyes are
large and situated rather far forward.

Upper Cambrian, Orr; (loc. 32t) Fandango Spring Canyon, east
side Dugway Range, Utah.

Holotype —U.S.N.M., no. 93026.

NO. 16 CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 33

Elvinia matheri (Walcott)
Ptychoparia matheri Wa.cott, Smithsonian Misc. Coll., vol. 57, no. 9, p. 268,
pl. 44, figs. 15-17, 1912; Bripce (part), U. S. Geol. Surv. Prof. Pap.
186-M, p. 254, pl. 60, fig. 11, 1937.

Upper Cambrian, Potsdam; (loc. 110) east side of Whitehall,
New York.
Cotypes—U.S.N.M. nos. 58585-7.

GENEVIEVELLA Lochman, 1936

Genevievella LocHMAN, Journ. Pal., vol. 10, no. 1, p. 40, 1936.
Llanoaspis LocHMAN, idem, vol. 12, no. 1, p. 80, 1938.

When this genus was established, only the genotype was known.
Two species were added from Texas, and the descriptions of eleven
from the Appalachians are in press; consequently this is a well-
represented trilobite genus. Thus far the genus is confined to the
Crepicephalus zone.

Llanoaspis modesta and L. undulata Lochman become Gene-
vievella.

IRVINGELLA Ulrich and Resser, 1924

Irvingella was applied to species which differ from Chariocephalus
in a limited degree. Chariocephalus has large eyes situated rather
far forward. The facial suture joins the dorsal furrow anterior to
the eye, by which arrangement the anterior fixigene is eliminated
and the brim is caused to be a single bar not attached to the fixigenes
at the anterior angles. On the other hand Jrvingella has even larger
eyes than Chariocephalus, the extra length being attained by the rear
portion of the palpebral lobes extending farther back. At the same
time the eyes are slanted outward from the axis of the head. Above
all, Jrvingella differs from Chariocephalus by having anterior fixigenes
which unite with the brim.

Irvingella tumifrons (Hall and Whitfield)
Chariocephalus tumifrons Hatt and WuitrFiELp, U. S. Geol. Expl. goth Par.,
vol. 4, p. 224, pl. 2, figs. 38, 39, 1877.

Upper Cambrian, Secret Canyon; Pogonip Mountain, White Pine
District, Nevada.
Holotype —U.S.N.M. no. 24561.

KOCHINA Resser, 1935

Kochina? lux (Walcott)
Ptychoparia lux Watcott, Smithsonian Misc. Coll., vol. 67, no. 3, p. 90,

Diet2ntie. 53) 1OT7:

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Lower Cambrian, Mount Whyte; (loc. 61d) southwest slope Mount
Shaffer, British Columbia.
Holotype—U.S.N.M. no. 64387.

LONCHOCEPHALUS Owen, 1852

Lonchocephalus verrucosus (Whitfield)
Conocephalites verrucosus WHITFIELD, Amer. Mus. Nat. Hist. Bull. 1, p. 146,
pl. 14, figs. 9-12, 1884.

Upper Cambrian, Potsdam; near “Post office,’ Ausable Chasm,
New York.

Cotypes—A.M.N.H. no. 280.

MENOMONIDAE Walcott
MENOMONIA Walcott, 1916

Menomoma Wa.cort, Smithsonian Misc. Coll., vol. 64, no. 3, p. 161, 1916.
Millardia Watcort, idem, p. 163, 1916.

Walcott referred three genera to the Proparian family Menomo-
nidae, but he failed to recognize that two of them were identical.
Evidence for a primitive aspect about Menomonia and Dresbachia
may be questioned. Evidently Walcott based his idea of this on the
numerous thoracic segments of M. calymenoides. Examination of
available specimens fails to prove that this extraordinarily long thorax
belongs to Menomoma, although association in three instances sug-
gests the possibility. Even if the genotype has 42 thoracic segments,
one can hardly consider this alone as evidence of primitive structure.

It seems rather that the Menomonidae are a highly specialized
group descended from species now placed in Alokistocare.

Menomonia is a characteristic trilobite of the early Upper Cam-
brian Cedaria zone; thus far being confined to that zone. Whitfeld’s
original specimens of the type species are on small pieces of rock
with the types of Cedaria woosteri.

Millardia was distinguished from Menomomia by differences of
brim and in having fewer thoracic segments. There is a difference
in the brim of M. calymenoides and of M. semele but it involves no
altered structure and the gap between the two species is bridged by
intermediate forms. M. calymenoides has a much swollen brim,
whereas other species obtain the same rigidity by arching the short
brim. Many specimens referred to Muillardia retain the libragenes,
giving them a different aspect, which caused their reference to a
separate genus.

Dresbachia is a distinct, but closely related genus. Compared with
Menomonia, the glabella is essentially the same but the fixigenes are

NO. 10 CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 35

much larger. The large libragenes—the part of this trilobite most
commonly preserved—are attached to a very short brim, which is
reduced to little more than a line, terminating in a swollen knob like
the rim Menomonia. In other words Dresbachia is characterized by
the lateral shortening of the preglabellar area to a mere line for the
attachment of the libragenes which lies wholly in front of the glabella.

Besides the genotype and the species formerly referred to Millardia,
several new species from the Appalachian Nolichucky formation are
in press.

Menomonia avitas (Walcott)
Millardia avitas Watcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 165,
pl. 28, figs. 5-5e, 1916.

Upper Cambrian, Warrior; (loc. 107k) 2 miles north of Benore,
Center County, Pennsylvania.
Holotype —U.S.N.M. no. 61629.

Menomonia magnagranulata (Lochman)
Millardia magnagranulata LocHMAN, Journ. Pal., vol. 12, no. 1, p. 84, pl. 18,
fig. 3, 1938.

Upper Cambrian, Cap Mountain; southwest side of Lion Moun-
tain, Highway 29, 9 miles northwest of Burnet, Texas.
Holotype —U.S.N.M. no. 95015.

Menomonia optata (Hall)
Conocefhalites optatus HA, 16th Ann. Rep. New York State Cab. Nat.
Hist., p. 222, pl. 5A, fig. 7, 1863. Trans. Albany Inst., vol. 5, p. 195, 1867.
Ptychoparia optata MILLER, North Amer. Geol. Pal., p. 539, 1880.
Millardia optata Watcort, Smithsonian Misc. Coll., vol. 64, no. 3, p. 165,
pl. 28, figs. 4, 4a-f, 1916.

One cannot be certain that Hall’s original type and the specimens
figured by Walcott represent the same species. At any rate Hall’s
original is a very small specimen.

Upper Cambrian, Eau Claire; Willow River Falls, Trempealeau,
and other localities in Wisconsin and Minnesota.

Holotype.—A.M.N.H. no. 322; plesiotypes, U.S.N.M. nos. 61626-8.

Menomonia semele (Walcott)
Millardia semele Wau.cott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 166,
pl. 28, figs. 3-3c, 1916.

Upper Cambrian, Weeks ; (loc. 30 n) Weeks Canyon, House Range,
Utah.
Cotypes.—U.S.N.M. nos. 61622-5.

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Menomonia texana, n. sp.
Millardia avitas LocHMAN (not Walcott), Journ. Pal., vol. 12, no. 1, p. 84,
pl. 18, figs. 4-5, 1938.

This species evidently was referred to M. avitas because it has
smaller granules than 1. magnagranulata. The reference to the Penn-
sylvania species cannot stand. M. avitas has a few scattered large
granules on the glabella and fixigenes; the remainder of the cephalon
is smooth to finely granulose or ornamented with lines. The eyes of
M. avitas are elevated on rather long stalks. This may aiso have
been the case in MM. texana as the eyes are broken off. M. texana is
further characterized by a long slender glabella and the eyes are
rather far forward.

Upper Cambrian, Cap Mountain; southwest side Lion Mountain,
Highway 29, 9 miles northwest of Burnet, Texas.

Holotype-—Mount Holyoke Mus. no. 655; paratype, no. 656.

Menomonia lochmanae (Lochman)
Millardia avitas LocHMAN, Journ. Pal., vol. 12, no. 5, p. 460, pl. 56, figs. 27,
28, 1938. :

This species differs from J/. avitas in the more incurved anterior
margin and considerably larger fixigenes.

Upper Cambrian, Petit Jardin; Cape St. George, Newfoundland.

Holotype-—vYale Peabody Mus. no. 15822.

NORWOODIA Walcott, 1916

Norwoodia Wa.cortt, Smithsonian Misc. Coll., vol. 64, no. 3, p. 168, 1916.
Whitfieldina RESSER, idem, vol. 95, no. 22, p. 27, 1937.

Just after the 1937 paper appeared, the genus Norwoodia was re-
considered in conjunction with studies in the southern Appalachians.
As soon as the extraneous species then in Norwoodia were removed,
it became clear that Whitfieldina is not a valid genus, because its type
species is a typical Norwoodia, contrary to the misleading published
figures.

Norwoodia quadrata (Whitfield)
Conocephalites quadratus WuttFIELD, Ann. Rep. for 1879, Wisconsin Geol.
Surv., p. 47, 1880; Geol. Wisconsin, vol. 4, p. 180, pl. 1, figs. 15, 16, 1882.
Whitfieldina quadrata REssER, Smithsonian Misc. Coll., vol. 95, no. 22, p. 27,
1927.

Upper Cambrian, Eau Claire; Eau Claire and other localities in
Wisconsin.
Cotypes——Univ. Wisconsin.

NO. I0 CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER a7

OLENOIDES Meek, 1877

Olenoides expansus (Walcott)
Dicellocephalus ? expansus Wa.cott, U. S. Geol. Surv. Mon. 8, p. 45, pl. 9,
fig. 19, 1884.
Dolichometopus ? expansus Watcort, Smithsonian Misc. Coll., vol. 64, no. 5,
p. 368, pl. 53, figs. 5, 5a, 1916.

Middle Cambrian, Eldorado; (loc. 55b) east slope Prospect Moun-
tain, Eureka District, Nevada. .
Holotype-—U.S.N.M. no. 15450.

ORYCTOCEPHALUS Walcott, 1886

Species of this genus have not been carefully discriminated, and
specific names were very carelessly used. A brief revision clears the
way for erection of the necessary new species.

Oryctocephaius, as now constituted, contains two groups of species.
The valid species previously recognized are listed under the two groups.

. walcotti Resser
. saltert Reed

. mdicus (Reed)
. kobayashi Saito
. orientalis Saito
. Saitoi, n. sp.

Group of O. primus (genotype)
O. primus Walcott
O. burgessensis, n. sp.
O. walkeri Matthew
Group of O. reynoldsi
O. reynoldsi Reed

SSIS ISTS ES

In addition, one undescribed species of the O. primus group and
several of the O. reynoldsi group are in hand.

Oryctocephalus burgessensis, n. sp.
Oryctocephalus primus KopayAsuHt, Journ. Faculty Sci. Imp. Univ. Tokyo,
SECuOmVOlLe4 pts 2p lA7- plats. osama NOs5e

Kobayashi picked up a photograph in the National Museum collec-
tions and published it as O. primus, but this illustration was pre-
pared by Walcott because he recognized that it was not O. primus.

O. burgessensis is the most abundant species of the genus in the
Burgess shale. It is characterized by a normal cranidium, seven tho-
racic segments, and slender pygidial spines of even length. The rear
segments of the pygidium are turned back practically parallel to the
axis. The pygidial spines of even length cause this species to look
much like O. walkeri, but the rear pygidial segments are turned back
more sharply.

Middle Cambrian, Burgess; (loc. 35k) Burgess Pass, near Field,
British Columbia.

Holotype-—U.S.N.M. no. 96487.

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Oryctocephalus indicus (Reed)
Zacanthoides indicus REED, Mem. Geol. Surv. India, Pal. Ind., ser. 15, vol. 7,
p. 9, pl. 1, fig. 15, 1910.
Oryctocephalus cf. reynoldsi REEp, idem, p. 12, pl. 1, figs. 22-24, 1910.
Reed recognized the generic position of the cranidium but referred
the pygidium to Zacanthoides. Since both come from the same bed,
at the same place, they can be regarded as representing one species.
O. indicus is chiefly distinguished from O. salteri, which occurs
in beds at least 400 feet higher in the section, by the longer pygidial
axis with proportionate reduction of the pleural lobes.
Middle Cambrian (horizon 2); Parahio Valley, Spiti, India.
Cotypes.—Geol. Surv. India plastotypes, U.S.N.M. no. 964809.

Oryctocephalus saitoi, n. sp.
Oryctocephalus cf. reynoldsi Satro, Japanese Journ. Geol. Geogr., vol. 11,
no. 3, p. 232, pl. 27, figs. 16-20, 1934.

Saito recognized the distinctness of this species from O. reynoldsi
and O. orientalis. He pointed out that it has one more pair of glabel-
lar furrows than in O. reynoldsi.

Middle Cambrian, Ptychoparia beds ; near Hwangju, Hwanghaido,
Chosen.

Cotypes.—Geol. Inst. Imp. Univ. Tokyo nos. 513, 516, 574.

Oryctocephalus walcotti Resser
Oryctocephalus walcottt RESSER (part), Smithsonian Misc. Coll., vol. 97,

OR Sh fon Oy I Tate, Ash Tate

In the description of this species a cranidium of Oryctocare getkiet
was mistakenly assigned to it. The description was written before
the genus was studied ; consequently, the fact that much better material
is available from the Spence shale was overlooked.

Middle Cambrian, Lakeview; (loc. 37n) Lakeview, Pend Oreille
Lake, and Spence; (loc. 55c) 5 miles southwest of Liberty, Idaho.

Holotype-—U.S.N.M. no. 95038.

PARABRISCOIA Kobayashi, 1935

Parabriscoia flabellifera (Hall and Whitfield)
Dikellocephalus flabellifer HAtyt and Wuitrietp, U. S. Geol. Expl. 4oth
Par., vol. 4, p. 227, pl. 2, figs. 29-30, 1877.

Upper Cambrian, Secret Canyon?; west side Pogonip Mountain,
White Pine District, Nevada.
Holotype —U.S.N.M. no. 24569.

NO. 10 CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 39

PTARMINGIA Raymond, 1928

Ptarmingia longula, n. sp.
Bathyuriscus cf. rossensis Wa.cott, Smithsonian Misc. Coll., vol. 67, no. 2,
p. 40, pl. 5, figs. 6, 6a, 1917.

This species has a longer head than P. rossensis. The glabella is
long, expanding only a little; eyes larger and prominent. A few
widely scattered granules occur on the elevated portions of the test
but appear to be much more numerous on exfoliated specimens.

The associated pygidium is very highly arched transversely and
has a rather even border, with a large spine at the anterior angles.

Middle Cambrian, Ptarmigan; (loc. 63m’) 1 mile east of Hector,
south slope Mount Bosworth, British Columbia.

Holotype —U.S.N.M. no. 63734; paratype, no. 63735.

PTEROCEPHALIA Roemer, 1849

Pterocephalia multicincta (Hall and Whitfield)
Dicellocephalus multicinctus Hatt and Wuuitrietp, U. S. Geol. Expl. goth
Rare vols 45 p: 226) pl. 2) fis. 37, 1877.
Pterocephalia sanctisabae BripcE (part), U. S. Geol. Surv. Prof. Pap. 186-M,
p. 240, pl. 68, figs. 37, 38, 1037.

Upper Cambrian, Secret Canyon; Eureka District, Nevada.
Holotype —U.S.N.M. no. 24640; plesiotype, no. 93021.

Pterocephalia laticeps (Hall and Whitfield)
Conocephalites (Pterocephalus) laticeps Hatt and Wuitrietp, U. S. Geol.
Expl. goth Par., vol. 4, p. 221, pl. 2, figs. 4-7, 1877.
Pterocephalia sanctisabae BrivcE (part), U. S. Geol. Surv. Prof. Pap. 186-M,
p. 248, pl. 68, figs. 40-43, 1937.

Upper Cambrian, Secret Canyon; west side of Pogonip Mountain,
White Pine District, Nevada.
Cotypes—U.S.N.M. nos. 24562, 24561, 24579.

Pterocephalia dakotensis, n. sp.
Pterocephalia sanctisabae Brince (part), U. S. Geol. Surv. Prof. Pap. 186-M,
p. 246, pl. 68, figs. 27-30, 32-36, 39, 1937.

This species averages about the same size as the largest specimens
of P. sanctisabae. It is characterized by rather large eye lobes, set
nearly parallel with the axis. The pygidium is ovate, partly fattened
in the shaly matrix.

P. dakotensis has a less flaring course of the anterior facial suture
than P. sanctisabae, the eyes are more nearly parallel to the axial line

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

of the head, and are larger. The course of the pygidial furrows is
also more evenly curved and the border is wider behind the axis.
Upper Cambrian, Deadwood; Whitewood Canyon, Deadwood,
Black Hills, South Dakota.
Cotypes—Univ. Iowa; U.S.N.M. no. 93020.

Pterocephalia bridgei, n. sp.
Pterocephalia sanctisabae BripcE (part), U. S. Geol. Surv. Prof. Pap. 186-M,
p. 246, pl. 68, figs. 25, 26, 1937.

This species is characterized by a wide brim, a relatively long
glabella, and the curved course of the anterior faciai suture. The
pygidium assigned to the species has a narrower border than P.
sanctisabae, and its rear margin is notched. The surface of the crani-
dium behind the eyelines and on the glabella, is nearly smooth. An-
terior to these parts the irregular longitudinal folds are rather heavy,
and the surface is marked by the usual horizontal, anastomosing lines.

Upper Cambrian, Davis; (loc. 11k) near shaft Federal Lead mine
no. 4, Flat River, Missouri.

Holotype and paratype —U.S.N.M. no. 930109.

Pterocephalia oriens, n. sp.
Pterocephalha sanctisabae Brivce (part), U. S. Geol. Surv. Prof. Pap. 186-M,

p. 246, pl. 68, fig. 24, 1937.

Associated with P. bridget is another, possibly smaller, species
characterized particularly by its highly elevated palpebral lobes. The
eyes are large, and the eyelobes are sharply separated by a deep fur-
row. The surface is ornamented similarly to P. bridget except that the
anastomosing lines are heavier.

Upper Cambrian, Davis; (loc. 11k) near shaft Federal Lead mine
no. 4, Flat River, Missouri.

Cotypes—U.S.N.M. no. 96485.

Pterocephalia potosiensis, n. sp.
Pterocephalia sanctisabae Brince (part), U. S. Geol. Surv. Prof. Pap. 186-M,
p. 246, pl. 68, fig. 23, 1937.

Only a few specimens were obtained in a rather large collection, so
that this species seems to be rare. The presence of a large libragene
may be interpreted as indicating that P. potosiensis grew to a much
larger size than the holotype indicates, or that there is more than
one species present.

P. potosiensis has a wider and longer brim than other Missouri
species, but it is shorter than that of P. bridget. The new species is

NO. 10 CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER Al

characterized particularly by well-developed anastomosing lines on the
glabella and libragenes.
Upper Cambrian, Davis; (loc. 11e) southwest of Potosi, Missouri.
‘Holotype —U.S.N.M. no. 93018.

Pterocephalia ulrichi, n. sp.
Pterocephalia sanctisabae BriwcE (part), U. S. Geol. Surv. Prof. Pap. 186-M,
p- 246, pl. 68, figs. 19, 21, 1037.

This is a large, wide species with a wide flaring brim. In fact,
this is the widest species thus far found. The glabella is wide and
rounded in front. Likewise the anterior margin of the cranidium is
more rounded than in most species. The dorsal furrow is both wide
and deep, so that the glabella and cheeks next to it have considerable
relief.

The pygidium is rather wide, the border of moderate width and
the rear margin evidently not notched.

Upper Cambrian, Honey Creek; (loc. 9q) 15 miles northwest of
Fort Sill, and (loc. 91L*) northeast of Big Baldy, Wichita Moun-
tains, Oklahoma.

Holotype—U.S.N.M. no. 93016; paratype, no. 93014.

Pterocephalia silvestris, n. sp.
Pterocephalia sanctisabae Bripcr (part), U. S. Geol. Surv. Prof. Pap. 186-M,
p. 246, pl. 68, fig. 20, 1937.

This is also a wide form and has a very large brim. Compared
with P. ulrichi, the brim is longer and the dorsal and glabellar furrows
shallower.

Upper Cambrian, Honey Creek; (loc. 91a) 4 miles southeast of
Hennepin, West Timbered Hills, Arbuckle Mountains, Oklahoma.

Holotype—U.S.N.M. no. 93015.

Pterocephalia deckeri, n. sp.
Pterocephalia sanctisabae Brince (part), U. S. Geol. Surv. Prof. Pap. 186-M,
p. 246, pl. 68, figs. 22, 31, 1037.

This species is particularly wide opposite the front end of the gla-
.bella. The eyelines are heavy and the eyes prominent by reason of
their upturned position, but the palpebral lobes slope up only moder-
ately. The surface is highly ornamented in the usual fashion, but the
anastomosing lines on the glabella and fixigenes have such a fine mesh
that the surface appears to be granulated.
The pygidium is rather narrow and high.

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

Upper Cambrian, Honey Creek; (loc. 89v) 4 miles south of Hen-
nepin, west Timbered Hills, Arbuckle Mountains, Oklahoma.
Holotype-—U.S.N.M. no. 93017.

PTEROCEPHALINA, n. gen.

For many years the genotype has been referred from one genus to
another, which, besides those listed below, include Anomocare, Niobe,
Saukia, and Platycolpus. The pygidium resembles that of both
Pterocephaha and the Saukinae but does not fit into either. Careful
search of the collections has revealed a partial cranidium which also
has affinities with Pterocephalia but cannot be placed in that genus.

Pterocephalina is characterized by a broad bordered pygidium in
which the wide doublure is almost severed by the deep notch in the
rear margin of the tail. The axis, which is subcylindrical, extends
almost to this notch, being connected with it by a sharp postaxial ridge.
Unexfoliated specimens show that pleural fusion is far advanced,
but the furrows are traceable on the upper surface nearly to the border.
The holotype shows the underside and hence the extent of the doublure,
but not the pleural furrows.

The cranidium assigned to the species is incomplete. It consists of
a large prominent, highly arched glabella, on which the rear pair of
furrows are represented by slight, rearward directed indentations.
Eyelines present. Eyes evidently of moderate size. Fixigenes less
than half glabellar width. Brim very wide, apparently with a flat,
sharply upturned rim. Wide preglabellar area is striated by coarse
irregular lines among which are interspersed scattered lumps. The
entire cranidium is highly arched in both directions.

Genotype-—Dikellocephalus bilobatus Hall and Whitfield.

Pterocephalina bilobata (Hall and Whitfield)
Dikellocephalus |Pterocephalus| bilobatus Haru and Wuitrietp, U. S. Geol.
Expl. 40th Par., vol. 4, p. 226, pl. 2, fig. 36, 1877.
Dicellocephalus bilobatus Watcott, U. S. Geol. Surv. Mon. 8, p. 40, 1884.
Platycolpus bilobatus Wa.cott, Smithsonian Misc. Coll., vol. 57, no. 13, p. 349,
IQI4.

Upper Cambrian, Secret Canyon; (loc. 65) east side Sierra Canyon,
opposite Pinnacle Peak, and (loc. 61) south of Hamburg Mine, Eureka
District, Nevada.

Holotype-—U.S.N.M. no. 24568.

NO. IO CAMBRIAN FOSSILS, 4TH CONTRIBUTION—RESSER 43

SAUKIELLA Ulrich and Resser, 1933

Saukiella junia (Walcott)
Saukia junia Watcotr, Smithsonian Misc. Coll., vol. 57, no. 13, p. 378, text
fig. 17, 1914.

Upper Cambrian, Signal Mountain; (loc. 12j) 2 miles southwest
of Signal Mountain, Wichita Mountains, Oklahoma.
Holotype —U.S.N.M. no. 60677.

SINOSAUKIA Sun, 1935

Sinosaukia bella (Walcott)
Ptychaspis bella Watcott, Proc. U. S. Nat. Mus., vol. 30, p. 585, 1906;
Research in China, vol. 3, Carnegie Inst. Publ. 54, p. 180, pl. 17, fig. 9,
1913.

Upper Cambrian, Fengshan; (loc. C 74) east of Fang-lan-chon,
Shansi, China.
Holotype —U.S.N.M. no. 58124.

TELLERINA Ulrich and Resser, 1933

Tellerina rustica (Walcott)
Saukia rustica Watcott, Smithsonian Misc. Coll., vol. 57, no. 13, p. 383, text
figs. 18-19a, IQT4.

Upper Cambrian, Signal Mountain; (loc. 12j) 2 miles southwest of
Signal Mountain, Wichita Mountains, Oklahoma.
Holotype —U.S.N.M. no. 60678; paratypes, nos. 60679-8o.

TONKINELLA Mansuy, 1916

Tonkinella kobayashi, new name.
Tonkinella breviceps KopAyAsHti, Journ. Faculty Sci. Imp. Univ. Tokyo,
Secu EVOle Ay pty 2. p: 150, pl. 05, hes. On G, Gy 1935:

This name was previously used for a species in Kashmir.
Middle Cambrian, Taiki; Neietsu, south Chosen.
Cotypes.—Geol. Inst. Imp. Univ. Tokyo.

"SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER 11

E “AN ASSAY METHOD FOR GROWTH-PROMOTING
|| SUBSTANCES UTILIZING STRAIGHT GROWTH
OF THE AVENA COLEOPTILE

(WitH ONE Beinn)

BY
ROBERT L. WEINTRAUB

Division of Radiation and Organisms,
Smithsonian Institution

(PUBLICATION 3488)

GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DEGEMBER 31, 1938

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER 11

AN ASSAY METHOD FOR GROWTH-PROMOTING
SUBSTANCES UTILIZING STRAIGHT GROWTH
OF THE AVENA COLEOPTILE

(WitH ONE PLATE)

BY
ROBERT L. WEINTRAUB

Division of Radiation and Organisms,
Smithsonian Institution
6

(PUBLICATION 3488)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 31, 1938

Tbe Lord Waltimore Press

BALTIMORE, MD., U. S. A.

y

AN ASSAY METHOD FOR GROWTH-PROMOTING
SUBSTANCES UTILIZING STRAIGHT GROWTH
OF. THE AVENA COLEORTILE

By ROBERT L. WEINTRAUB

Division of Radiation and Organisms, Simithsonian Institution
(WitH ONE PLATE)

The most delicate available methods for the determination of plant
growth-promoting substances involve the direct measurement of the
effects of these substances on the growth of suitable plant test objects.
The most commonly employed indicator is the decapitated Avena
coleoptile. Two general techniques are available ; one makes use of the
elongation (straight growth) resulting from the application of the
substance in question symmetrically with respect to the long axis
of the coleoptile, the other utilizes the curvature produced by uni-
lateral application of the growth-promoting substance. The latter
method, which has been more widely used, has been described repeat-
edly (see e.g., Boysen-Jensen, 1936; Went and Thimann, 1937;
Avery, Burkholder, and Creighton, 1937) and need not be detailed
here. As Went and Thimann (1937, p. 51) point out, “The con-
venience of curvature methods rests upon two facts: (1) the residual
growth, after decapitation, is the same on both sides of the plant and
thus is automatically eliminated from the measurement—no controls
are necessary ; and (2) only one measurement need be made; there is
no zero reading.” It should be noted, however, that the first condi-
tion, namely, the uniformity of the residual growth, is true only dur-
ing the first 2 hours following decapitation (cf. fig. 20, Went and
Thimann, 1937), and this limits the length of the test period.

Thus the curvature test measures not the maximum amount of
curvature (growth) which can be induced by the applied substance,
but rather the mean rate of curvature during a given period. During
this period the rate is not constant and may even change in sign (cf.
Schneider and Went, 1938). The factors which cause a reduction in
the curvature rate and therefore in the amount of curvature at the
end of the test period are: (1) gravity, which causes a geotropic curva-
ture in the opposite direction ; (2) the effect of “physiological regen-
eration” of the tip, and (3) the lateral transport of the applied growth-

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97 No. 11

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

promoting substance across the coleoptile, producing a growth ac-
celeration on the far side of the plant. The influence of the last-named
factor is very marked in the case of a number of substances which
show relatively little or no activity by the curvature method but have
considerable effect on straight growth (cf. table XII, pp. 137-139,
Went and Thimann, 1937). This lateral transport is greater the
higher the concentration of growth-promoting substance applied; the
net result is a decrease in the sensitivity of the test.

Methods employing straight growth have been employed occasion-
ally but have not come into routine usage, largely because of the
inconvenience in measuring the growth. The present report describes
an assay procedure utilizing straight growth of the coleoptile of Avena
sativa, in which the sensitivity of the response and the ease and ac-
curacy of measurement are at least as great as in the commonly used
curvature test. In addition, the method offers a number of other tech-
nical advantages.

The procedure will be outlined briefly first, and then each step will
be discussed in greater detail.

SUMMARY OF METHOD

Seeds are planted on agar slants in small test tubes at a determined
distance below the rim of the tube. The seedlings are germinated and
grown under controlled conditions. When the coleoptiles have at-
tained a given length they are decapitated level with the rim of the
tube and the leaf is withdrawn completely. Blocks of agar, containing
the growth-promoting substance to be tested, are placed terminally
upon the entire cut surface of the stump. After some time a shadow-
graph is made in the usual manner. The length of the coleoptile which
extends above the rim of the tube represents the growth increment
during the test period. It can be measured very easily with a dissect-
ing microscope equipped with an ocular micrometer.

TEST TUBES, AND RACKS

Soft glass or Pyrex tubes having an inner diameter of about 15
mm and a length of 7 cm have been found satisfactory. Trials with
larger tubes indicate that within reasonable ‘limits the size is imma-
terial. A simple rack for the tubes may be constructed by boring a row
of holes in a wood block. The holes should be of such depth and

1 Avena sativa var. Markton has been used exclusively. The seeds were ob-
tained through the courtesy of Mr. T. Ray Stanton, of the U. S. Department
of Agriculture.

NO. II ASSAY METHOD FOR GROWTH SUBSTANCES—WEINTRAUB 3

diameter that the tubes slip in easily and stand upright; 2.5 cm
between centers allows sufficient room to manipulate the tubes. The
length of the block, of course, determines the number of tubes and
will depend upon the size of the photographic paper, incubator, etc.,
which one uses. For ready identification of the racks and the shadow-
graphs some suitable design (e. g., a letter or number) may be punched
or drilled in a strip of sheet metal which is fastened against the back
of the rack (see pl. 1). Machine-made test tubes usually have suff-
ciently uniform rims; if the tubes-are made by hand from glass tub-
ing it 1s necessary to grind the rim at right angles to the long axis of
the tube. A mark should be made on the tube at a given distance
below the rim; if it is desired to change the depth of planting in dif-
ferent experiments a glass-marking pencil is convenient ; otherwise, a
scratch made with a file or carborundum wheel furnishes a permanent
mark.

THE AGAR SLANT

The same purified agar which is used for the test blocks is suitable.
The concentration of agar should not be less than 0.8 percent. Greater
concentrations, up to 2 percent, give equally good results; 0.9 to
I percent agar has been routinely used. The agar may be made up in
nutrient solution if desired but tap or distilled water have given uni-
formly satisfactory results. The growth rate of the coleoptile will be
found to depend upon the composition of the agar. The simple device
used in bacteriological laboratories is very convenient for filling the
tubes with the melted agar. This consists of a funnel connected by a
short length of rubber tubing to a glass tip and provided with a pinch
clamp. The tubes are placed in the rack, filled up to the mark and the
whole rack is tilted backward through about 60° so that the agar
solidifies in a slant. The angle of the slope (and of the planted seed)
should be such that the coleoptile grows erect without being required
to curve. One hundred tubes can be charged with agar in about 8
minutes.

PLANTING AND GERMINATING SEEDS

The husked seed is pressed gently against the surface of the agar
slant with the groove side down and the embryo at the level of the
mark on the tube. The seeds may be soaked in water before plant-
ing or planted without previous soaking. Soaking does not affect
the growth rate or the sensitivity of the plants. Furthermore, husked
seeds planted dry on the agar absorb water nearly as rapidly as

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

if they are immersed in water, so that there is no advantage in pre-
liminary soaking. Planting of dry seeds obviates a second handling
of the seedlings. Under the conditions routinely employed in this
laboratory (growth continuously from the time of planting in red
light (Wratten Safelight, series 0) at 25° C. and about go percent
relative humidity) the coleoptiles attain a length of 25 mm at about
65 hours after planting. The growth rate at this time, and for the
next 24 hours, is approximately 0.9 mm per hour. Plants grown on
agar slants in small tubes as described have shown less individual
variability than those handled in any other way, as on filter paper, or
sand, in the usual glass Avena holders, or on porous stone wicks.
One hundred seeds can be planted in the tubes in about Io minutes.

LENGTH OF COLEOPTILE

In connection with the size of the coleoptile used for the test, three
factors have been studied. These are the total length of coleoptile,
the length of the tip decapitated, and the length of the stump used.

The growth rate of the basal portion of the coleoptile decreases as
the total length of the coleoptile increases, and if 20 to 25 mm of the
coleoptile tip are removed, the stump makes practically no growth
when a plain agar block is applied. If coleoptiles are used under these
conditions no controls are necessary. However, the sensitivity (used
here as the amount of growth in excess of the control which is pro-
duced by application of a given amount of growth substance) of the
basal portion also decreases rather rapidly as the total length of the
coleoptile increases. The 20-mm stumps of 40-mm coleoptiles have
practically no residual growth under the conditions of the test, but
do have a rather high sensitivity. It is possible, therefore, to use
coleoptiles of this length without controls. More commonly, however,
13-mm stumps of 24- to 27-mm coleoptiles have been used because the
plants are ready for the test nearly a day earlier. A control set must,
of course, be included.

It is not intended to suggest that it is necessary to employ plants
of just this length or even that the described conditions are optimal,
but merely to indicate the technique which has given satisfactory
results. As a matter of fact, since the test is essentially comparative,
a few millimeters variation in the length of the test plants is of no
consequence provided the plants are randomized throughout the
different sets.

NO. If ASSAY METHOD FOR GROWTH SUBSTANCES—WEINTRAUB 5

DECAPITATION

It is essential that the cut surface of the coleoptile be exactly at the
level of the test tube rim and that the cut be clean and horizontal (at
right angles to the long axis of the coleoptile). Otherwise the plants
may bend and measurement will be difficult. Decapitation can be per-
formed quite rapidly by making a small cut partially through one (or
two opposite ) sides of the coleoptile with a thin safety razor blade held
flat against the rim of the tube and bending the coleoptile toward the
cut with the fingers or forceps until it breaks. The leaf is pulled out
completely. It is often possible to break off the coleoptile and withdraw
the leaf ina single motion. One hundred coleoptiles can be decapitated
in 30 minutes or less.

AGAR LESt BEOECKS

In the development of the method, weighed amounts of dehydrated
agar were mixed with aqueous solutions of known concentrations
of indole-3-acetic acid, or of auxin-a.” Similar results have been
obtained with both of these growth-promoting substances. The test
blocks were prepared with an apparatus similar to that described by
DuBuy (1938).

Thimann and Schneider (1938) have reported that the concentra-
tion of agar in the test blocks is of considerable importance in the
Avena curvature test. In general they found that a given concentra-
tion of indole-3-acetic acid produced larger curvatures the lower the
agar concentration. Similar, although less marked, differences have
been found in the present study of straight growth. The use of 1.5
percent agar has been adopted as a general procedure.

SIZE OF, TEST BLOCKS

Went (1928) concluded that with 0.9 mm* blocks the curvatures
are proportional to the absolute amount of growth substance in the
blocks. Van der Weij (1931) and Thimann and Bonner (1932) con-
cluded that the curvatures are proportional to the concentration of
growth substance in the block. The data of Thimann and Bonner indi-
cate that the rate at which the growth substance passes from the
block to the plant is proportional to its concentration in the block at
any moment. The change in concentration of growth substance in the
block during any given period will be less the greater the volume of

2A solution of pure crystalline auxin-a was very generously supplied by
Prof. F. Kogl, of the University of Utrecht.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

the block. Therefore the larger the block the longer the time during
which the induced growth will be proportional to the original con-
centration of applied growth substance. Much larger blocks can be
applied terminally than can be applied unilaterally.

With blocks as large as 26 mm* it has been found that the straight
growth rate of the decapitated coleoptiles remains constant for at least
6 hours. As blocks of this size can be manipulated very conveniently
they have been adopted.

The test blocks are applied so as to cover the entire cut surface of
the coleoptile stump. A small drop of water or gelatin solution may be
previously applied to the cut surface in order to insure good contact
with the block. About 20 minutes are required for the application of
100 blocks.

Schneider and Went (1938) have shown that the length of time
between decapitation and application of the blocks is of considerable
importance in the curvature test. This has been confirmed by Thimann
and Schneider (1938). In the straight growth method, on the other
hand, the response to applied growth substance has not been found to
be significantly influenced by the interval between decapitation and
application of the blocks, at least within the limits of 5 to 120 minutes.

LENGTH OF TEST PERIOD

The greatest sensitivity is obtained with the longest test period
during which the growth in excess of the control is proportional to
the concentration of applied growth-promoting substance. That is,
under such conditions the absolute useful amount of growth is the
greatest, and consequently the measurement can be made with great-
est accuracy. Actually, it has been found that a test period of 3 to 4
hours is quite adequate. In the present study a 4-hour period has
been generally used. In practice it is not essential to use a test period
of any exactly predetermined length. Hence, if it is inconvenient to
terminate the test at precisely 4 hours, there is no objection to making
the test period several minutes shorter or longer. In any comparable
series, of course, the test periods for all the sets should be the same.

ENVIRONMENTAL CONDITIONS OF TEST

Thimann and Schneider (1938) have reported that the growth
response of coleoptile sections to indole-3-acetic acid depends upon
the conditions of illumination of the seedlings during the previous
development ; maximal response was found when the plants received
red light during the first several hours of germination and were kept

NO. II ASSAY METHOD FOR GROWTH SUBSTANCES—WEINTRAUB v

in darkness thereafter. It has not been determined whether this is
true also of the coleoptile stumps attached to the seeds as employed in
the present technique. As has been mentioned previously, constant
illumination with red light has been used, since this permits absolute
reproducibility in successive lots of plants and is much more con-
venient when successive lots are grown concurrently in a single dark
room. It has been found, however, that the sensitivity is not appreci-
ably different whether the plants are kept in darkness or given red
light during the test period itself.

With an adequate water supply to the roots of the plants, consid-
erable differences in atmospheric humidity do not influence the sensi-
tivity. No significant difference in growth rate was found between
plants at 100 percent and at 75 percent relative humidity even though
the test blocks shrink very considerably at the lower humidity.

MEASUREMENT OF GROWTH

For measurement of the shadowgraphs a dissecting microscope
equipped with a 14 X ocular anda 2 X objective has been used. The
ocular is provided with a I-cm scale subdivided into 100 divisions.
One mm on the scale (10 divisions) corresponds to 0.5 mm on the
shadowgraph so that the length can be read directly to 0.05 mm.
The uncertainty in measuring is of the order of one scale division.
Since the growth of the control plants in four hours is about 0.7 mm,
this corresponds to an error of about 7 percent ; in the test plants, which
make more growth, the error of measurement is correspondingly re-
duced. Furthermore, the error tends to be minimized when the aver-
age of a number of plants is taken.

RESULTS

The usefulness of the Avena coleoptile as a test object rests upon
the fact that the induced growth is proportional, within certain limits,
to the concentration of applied growth-promoting substance. That
such a proportionality does exist was demonstrated by Thimann and
Bonner (1933), and has been confirmed repeatedly in the present
study. Figure I represents the relationship between growth and
growth substance concentration which has been found with the pro-
cedure here employed. It will be seen that the curve is a typical Black-
man curve with a very short transition region, very similar to that
obtained originally by Went (1928) for the curvature test. The work
of Thimann and Schneider (1938) indicates that, in the curvature test

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

at least, the form of the curve may vary greatly according to the
technique used. No evidence of a similar situation in the straight
growth response has been obtained as yet. ;

DISCUSSION

In addition to the theoretical preferability of utilizing straight
growth rather than curvature, the present method offers several tech-
nical advantages. The usual glass holders, which are time-consuming
to make, difficult to clean and easily broken, are eliminated. The solid

(MM)

INCREASE OVER CONTROL

| sae) | L |

0.05 0.10 0.15 0.20 0.25
CONCENTRATION INDOLE-3-ACETIC ACID (MG/LITER)

Fic. 1—Relation between concentration of applied growth substance
and straight growth.

root medium provides firm anchorage for the seedlings which facili-
tates the operations of decapitation, removal of the leaf, and applica-
tion of the block. The use of test tubes greatly expedites the removal
and rearrangement of the plants in the racks in preparing uniform
sets. The large test blocks are more easily applied and make better
contact when applied terminally. No handling of the seedlings is
necessary between the time of initial planting and of testing.

These practical advantages, as well as some others, apply also to the
use of agar in test tubes for the growth of seedlings to be used in the
curvature test. The seeds may be planted very close to the top of the
tubes so that practically none of the coleoptile is obscured in photo-
graphing. The tubes allow the plant to be revolved about its long axis

NO. II ASSAY METHOD FOR GROWTH SUBSTANCES—WEINTRAUB 9

so that the plane of curvature can be placed parallel to that of the photo-
graphic paper. Deseeding may be very easily accomplished with a small
section lifter or with forceps.

Some disadvantages of the method should be mentioned also. The
size of the plants used is a relatively critical factor in comparative
studies. It has been found that with the uniform conditions employed
the time at which the seedlings will be ready for use can be predicted,
at the time of planting, to within 2 or 3 hours. It is essential that this
be considered in planning the various operations. The individual vari-
ability of the seeds used is such that only about 75 percent of the plants
are ready at one time. In a limited series, therefore, there will be con-
siderable waste. In an extended series of tests, involving a few hun-
dred plants, if the larger plants are used first the smaller ones will
attain a suitable size by the time they are needed so that more than
go percent of the planted seeds can be utilized.

SUMMARY

An assay method for growth-promoting substances, which utilizes
straight growth of the Avena coleoptile, is described. The method
appears to possess a number of theoretical and practical advantages
over the widely used curvature test.

LITERATURE CrrED

Avery, G. S., Jr., BuRKHOLDER, P. R., and CrercHton, H. B.
1937. Avena coleoptile curvature in relation to different concentrations of
certain synthetic substances. Amer. Journ. Bot., vol. 24, no. 4,
pp. 226-232.
BoysEN-JENSEN, P.
1936. Growth hormones in plants. English translation by G. S. Avery, Jr.,
and P. R. Burkholder. McGraw-Hill Book Co.
DuBuy, H. G.
1938. A method for extracting growth substances from pigmented tissues.
Journ. Agr. Res., vol. 56, no. 2, pp. 155-158.
SCHNEIDER, C. L., and WENT, F. W.
1938. A photokymograph for the analysis of the Avena test. Bot. Gaz.,
vol. 99, no. 3, pp. 470-496.
THIMANN, K. V., and Bonner, J.
1932. Studies on the growth hormone of plants. II. The entry of growth
substance into the plant. Proc. Nat. Acad. Sci., vol. 18, pp. 692-701.
1933. The mechanism of the action of the growth substance of plants.
Proc. Roy. Soc., ser. B, vol. 113, pp. 126-149.
Tuimann, K. V., and SCHNEIDER, C. L.
1938. The role of salts, hydrogen-ion concentration, and agar in the re-
sponse of the Avena coleoptile to auxins. Amer. Journ. Bot., vol.
25, no. 4, pp. 270-280.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. ‘97
VAN DER WEI, H. G.

1931. Die quantitative Arbeitsmethode mit Wuchsstoff. Proc. Kon. Akad.

Wetensch. Amsterdam, vol. 34, pp. 875-8092.
WEntT, F. W.

1928. Wuchsstoff und Wachstum. Rec. Trav. Bot. Neerl., vol. 25, pp. 1-116.
Went, F. W., and Turmann, K. V.

1937. Phytohormones. The MacMillan Co.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 97, NUMBER 12
(End of Volume)

THE SPENCE SHALE AND ITS FAUNA

(Witu Srx Priates)

BY
GHARLES ELMER RESSER

Curator, Division of Stratigraphic Paleontology,
U. S. National Museum

(PUBLICATION 3490)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 20, 1939

The Lord Baltimore Press

BALTIMORE, MD., U. 8 Ae

THE SPENCE SHALE AND ITS FAUNA

By CHARLES ELMER RESSER
Curator, Division of Stratigraphic Paleontology,
U. S. National Museum

(WitH Six PrateEs)
INTRODUCTION

The Spence shale occurs in the northern Wasatch Mountains, in
northeastern Utah and adjacent portions of Idaho. Thus far neither
this shale nor its fauna has been found south of Brigham, but it is
present in both the Wasatch proper and its eastern prong, the Bear
River Range. Fossils are everywhere present, sometimes in great
abundance and variety.

In 1896 R. S. Spence sent some very interesting and well-preserved
Middle Cambrian fossils to Dr. Charles D. Walcott. Correspondence
shows that additional material was sent during the next Io years.
Late in the summer of 1906 Dr. Walcott moved camp from Black-
smith Fork to the eastern slopes of the Bear River Range in the
vicinity of the locality discovered by Mr. Spence. A brief description
of the sections studied in 1906, together with formational names,
was published by Walcott in April 1908 (1g08a). In December of
the same year (1908b) the sections were republished in more detail,
including preliminary faunal lists. Unfortunately, the several sec-
tions measured in both divisions of the Wasatch were combined into
a composite section, thereby obscuring essential stratigraphic facts.

The Spence shale was defined (Walcott 1g08a) as “argillaceous
shales” about 30 feet thick with “an extremely abundant and varied
lower Middle Cambrian fauna,” and in the fuller description (Wal-
cott 1908b) the words “and sandy shale’? were added. This thin
stratigraphic unit was regarded as a member at the base of the Ute
formation. It is not a mappable unit and therefore deserves recog-
nition only for its abundant and striking fauna.

The type locality for the Spence shale is in Spence Gulch, which
is situated on the eastern slopes of the Bear River Range, about
5 miles southwest of Liberty and 15 miles west of Montpelier, Idaho.
Similar beds, but with few fossils, are recognized to the south near
Garden City. On the western slope of the Bear River Range, the

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 97, No. 12.

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Spence shale fauna occurs in the base of the Ute formation in Black-
smith Fork. Deiss (1938) remeasured this section, which forms the
standard for the northern Wasatch region, and concluded that the
Spence was not present in Blacksmith Fork. But these studies indi-
cate that Walcott’s original assignment was probably correct. In the
Wasatch proper Spence shale fossils were obtained from Two Mile
Canyon, near the northern terminus of the range, and from a belt
of unknown extent several miles north of Brigham.

At the latter locality the fauna is found in more calcareous shales
than elsewhere. Usually, the Spence shale is a rather soft argillaceous
shale, but the fossils are never completely flattened. It seems that
fossiliferous lime nodules occur at most localities; at some places
these are small, very hard pyritiferous concretions, and at others they
consist of crystalline limestone. Irregular oolitic limestone layers
are also evidently developed locally.

Shortly after the large collections were made from Spence Gulch
in 1906, illustrations of the more conspicuous elements of the fauna
were prepared. The 1912 monograph includes the brachiopods, and
from time to time trilobite species were described incidentally in
other papers. Even the present paper does not describe every species,
but it gives a fairly accurate concept of the fauna, only a few obscure
forms remaining. Neither are the large number of embryonic speci-
mens considered.

The strata in the Pend Oreille Lake region (Resser, 1938) are
the only precise equivalents of the Spence shale. Close affinities exist
southward in the Wasatch and southwestward from and including
the Ophir shale of the Oquirrh Range. Exact correlations are not
attempted until several other faunas have been studied.

In order to save many repetitions of locality descriptions, they are
placed here in full, and reference is made to them in the text by
number only.

Locality 55c——Middle Cambrian, Spence shale; about 5 miles
southwest of Liberty, 15 miles west of Montpelier, Bear River Range,
Idaho.

Locality 54L.—Middle Cambrian, Spence shale; Blacksmith Fork,
about 10 miles east of Hyrum, Bear River Range, Utah.

Locality 55e.—Middle Cambrian, Spence shale; mouth of first
small canyon south of Wasatch Canyon, east of Lakeview Ranch,
44 miles north of Brigham, Wasatch Mountains, Utah.

Locality 20x—Middle Cambrian, “Langston,” Spence, Ute; near
top of gulch, about 2 miles north of Brigham City, Wasatch Moun-
tains, Utah.

NO. 12 SPENCE SHALE AND ITS FAUNA—RESSER ; 3

Locality 5g—Middle Cambrian, Spence shale; Two Mile Canyon,
2 miles southeast of Malad, Wasatch Mountains, Idaho.

Locality 32x%—Middle Cambrian, Ute, Spence, Bloomington;
Wasatch Canyon, 5 miles north of Brigham, Wasatch Mountains,
Utah.

It will be observed that part of the collections from all but lo-
calities 55c and 541 consists of other than Spence shale fossils.
Consequently, assignment of species to the Spence shale is certain
only for locality 55c where all fossils came from a thin bed with
no chance for admixture, and locality 5g on the western side of the
Wasatch Mountains. Species believed to represent the Spence fauna
have been chosen from the collections of localities 55e, 20x, and 32x.
This choice is based on lithology and generic assemblages on hand
pieces of rock.

Westonia ella (Hall and Whitfield) was described originally from
locality 55e. At several places this species presumably appears in
the Spence shale fauna, but evidently is present also in the higher
beds of the Ute formation. However, since the species was not found
at Spence Gulch, it has been omitted from the illustrations.

Zacanthoides is present also in localities 55e and 20x, but the species
are not identifiable owing to the fragmentary nature of the material.

The Archaeocyathinae are possibly represented (pl. 1, fig. 39).

DESCRIP TION OM SPECIES
ALGAE

MORANIA Walcott, 1919
MORANIA, sp. undet.
Plate 1, fig. 40
A few Spence shale surfaces show flat algae of the Morania type.
Morania is found in highly fossiliferous argillaceous shales almost
everywhere in Lower and Middle Cambrian strata.

Locality 55c.
Figured specimen.—U.S.N.M. no. 96491.

ECHINODERMATA
EOCRINUS Jaekel, 1918
EOCRINUS LONGIDACTYLUS (Walcott)
Plate 1, figs. 41, 42

Eocystites?? longidactylus Watcott, U. S. Geol. Surv. Bull. 30, p. 94, pl. 5,
fig. 3; pl. 6, fig. 1, 1886.
Eocrinus longidactylus JAEKEL, Pal. Zeitschr., vol. 3, p. 24, 1918.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97
e
The identification of this species is not altogether certain, but
as far as the state of preservations allows, careful.comparison seems
to indicate that it should be so identified.
Localities 55c and 55e.
Cotypes—U.S.N.M. no. 15315; eee no. 96492.

WORMS

SELKIRKIA Walcott, 1911
SELKIRKIA SPENCEI, n. sp.
Plate 1, figs. 34, 35

This species from the Spence. shale averages smaller than S.
major. The rate of taper is also less, so that the margins of S. spencet
are almost parallel.

Localities 55c, 55e, and 54L.

.

Cotypes —U.S.N.M. no. 96493.

BRACEMOPODA

MICROMITRA Meek, 1873
MICROMITRA LEPIDA, n. sp.

Plate 1, figs. 11-13

This species is characterized by its large size, many of the shells
measuring 10 mm long and 7 mm wide. The ventral valve is rather
highly elevated. The dorsal valve preserves internal markings which
are the first to be found in any species of the genus. The surface of
both valves is ornamented with the usual strong growth lines, and
both valves show fine ribbing.

Locality 55c.

Cotypes—U.S.N.M. no. 51458.

IPHIDELLA Walcott, 1905
IPHIDELLA GRATA, n. sp.
Plate 1, figs. 14-18

This brachiopod was referred by Walcott to /. pannula, but its large
size alone distinguishes it from the numerous species referred to
I. pannula, except certain ones in the Grand Canyon and from lo-
calities northward of Montana. The characteristic ornamentation is
beautifully developed and covers practically the entire shell with
equal intensity. The hinge line is rather long and on the dorsal valve

NO. 12 SPENCE SHALE AND ITS FAUNA—RESSER 5

is nearly straight. The crenulated ornamentation gives way some-
what at the outer margin of old shells to strong growth lines.
Localities 55c¢ and 54L.
Cotypes—U.S.N.M. nos. 51448, 27444.

LINGULELLA Salter, 1866
LINGULELLA EUCHARIS, n. sp.

Plate 1, figs. 1-3

The Lingulella species in the Spence shale was referred to the
Upper Cambrian species, L. desiderata. L. eucharis is of medium
size averaging about 3 mm in length. The illustrations show that it
averages broader than any of the numerous species referred to L.
desiderata. Growth lines are normally defined, and the usual faint
ribbing shows where the outer surface of the shell is exfoliated.

Localities 55¢ and 55e.

Cotypes—U.S.N.M. nos. 51704, 51826.

ACROTHELE Linnarsson, 1876
ACROTHELE AFFINIS, n. sp.

Plate 1, figs. 19-22

The Spence shale species of Acrothele was referred to A. subsidua,
but it averages larger and differs further in the more central position
of the apex. The ventral valve had considerable elevation. The ex-
terior of both valves is marked by strong growth lines. Well-pre-
served outer surfaces show characteristic crenulations. The average
size of the shells is about 8 mm.

Localities 55c, 55e, and 20x.

Cotypes—U.S.N.M. no. 52015.

ACROTRETA Kutorga, 1848
ACROTRETA DEFINITA Walcott
Plate 1, figs. 7-10

Acrotreta definita Watcort, Proc. U. S. Nat. Mus., vol. 25, p. 584, 1902; U. S.
Geol. Surv. Mon. 51, p. 683, pl. 64, figs. 2, 3, 1912.

Locality 55c.
Holotype and paratypes —U.S.N.M. no. 35270.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 7.

ACROTRETA LEVATA, n. sp.
Plate 1, figs. 4-6

Another species with a very high ventral valve occurs with A.
definita. Besides the height of the ventral valves, this species is char-
acterized by its strong growth lines and the narrowness of the false
pedicle groove.

Locality 55c.

Cotypes—U.S.N.M. no. 52108.

WIMANELLA Walcott, 1908

Walcott proposed II’imanella for smooth, nonplicate brachiopods.
He failed to observe that his specimens of the type species were not
well preserved and that the ribs were effaced. Because of this error
many species of I]’imanella have been referred to Nisusia. In fact,
most Middle Cambrian forms referred to Nisusia are WVimanella, but
thus far the genus does not extend above the Middle Cambrian.

WIMANELLA SPENCEI (Walcott)
Plate 1, figs. 27, 28

Nisusia (Jamesella) spencei Watcort, U. S. Geol. Surv. Mon. 51, p. 737, text
fic, '62) pl. (93; figs. -7,°7a, 1912.

Localities 55c, 20x, and 32x.
Cotypes—U.S.N.M. no. 52435; plesiotypes, no. 96408.

WIMANELLA RARA (Walcott)
Plate 1, fig. 29

Nisusia rara Watcott, (part), Smithsonian Misc. Coll., vol. 53, no. 3, p. 97,
pl. 9, fig. 13a, 1908; U. S. Geol. Surv. Mon. 51, p. 729, text fig. 60, 1912.
It is possible that this single shell is not a good species but merely
a peculiarly preserved specimen of W. spencet.
Locality 55¢c.
Holotype—U.S.N.M. no. 52295.

WIMANELLA NAUTES (Walcott)
Plate 1, figs. 23-26

Nisusia (Jamesella) nautes Watcortt, U. S. Geol. Surv. Mon. 51, p. 734, pl. 93,
figs. 6-6b, 1912.

Locality 55c.
Cotypes.—U.S.N.M. no. 52432; plesiotypes, no. 96497.

NO. I2 SPENCE SHALE AND ITS FAUNA—RESSER

“SI

GASTROPODA
HYOLITHES Eichwald, 1840
HYOLITHES CECROPS Walcott
Plate 1, figs. 36-38

Hyolithes cecrops Watcott, Smithsonian Misc. Coll. vol. 67, no. 2, p. 27,
pl. 5, figs. 3-3c, 1917.

Hyolithes idahoensis REssER, idem, vol. 97, no. 3, p. 5, pl. 1, figs. 57, 58, 1938.

This is an abundant species, but owing to its large size and shale
matrix, few complete specimens are available. Numerous operculae
are preserved. Assembly of many specimens shows that the Spence
shale form, together with that in the Rennie shale, is identical with the
Ross Lake shale species.

Ross Lake; (loc. 63j) Popes Peak, 14 miles south of Stephen, and
other localities, British Columbia.

Rennie; (loc. 37m) North Gold Creek, Pend Oreille Lake.

Spence; localities 55c, 55e, and 54L.

Cotypes—U.S.N.M. no. 63724; plesiotypes, nos. 95021, 96496.

HYOLITHES ORNATELLUS, n. sp.

Plate 1, figs. 30-32

This is the most highly ornamented species of Hyolithes known.
The anterior side is flat and depressed below the rounded margins,
while the posterior surface is arched. Strong striations run at right
angles to the axis on the anterior surface, but on the posterior are
parallel with the front margin.

‘Locality 55c.

Cotypes—U.S.N.M. no. 96494.

“QORTHOTHECA” SOLA, n. sp.
Plate 1, fig. 33

A single conical tube was found among the thousands of Spence
shale fossils. This shell has nearly a circular cross-section and is
about 8 mm long and 4 mm wide at the aperture.

Locality 55c.

Holotype-—U.S.N.M. no. 96495.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

AGNOSTIDA
AGNOSTUS Brongniart, 1822
AGNOSTUS BONNERENSIS Resser
Plate 2, figs. -24-26

Agnostus bonnerensis RESSER, Smithsonian Misc. Coll., vol. 97, no. 3, p. 6,
ply Ta higs: 10; 17,) 1038:
This species is abundant in the Spence shale.
Middle Cambrian, Lakeview; (loc. 37n) Lakeview, Pend Oreille
Lake, Idaho.
Locality 55c.
Plesiotypes.

U.S.N.M: no. 96499.

AGNOSTUS BRIGHAMENSIS, n. sp.

Plate 2, figs. 27-29

Photographs of this species are mounted to show the associated
fauna, Oryctocephalus, Pagetia, Clavaspidella, and Lingulella.

Compared with A. bonnerensis, this species differs first in its more
circular outline for both shields. Little further difference is noticeable
in the head. A sharp difference is brought about in the pygidium
by the deeper transverse rhachial furrows but more particularly by
the failure of the rear rhachis to penetrate the pleural lobe.

Locality 20x.

Holotype and paratypes —U.S.N.M. no. 96500.

TRICOBIMaE
PAGETIDAE Kobayashi
PAGETIA Walcott, 1916

PAGETIA CLYTIA Walcott

Plate 2, figs. 30-32

Pagetia clytia Watcorr, Smithsonian Misc. Coll., vol. 64, no. 5, p. 408, pl. 67,
figs. 2-2e, 1916.
In spite of the great abundance of this trilobite it is difficult to find
really good specimens.
Localities 55c and 20x.
Cotypes——U.S.N.M. nos. 62862-7.

NO. 12 SPENCE SHALE AND ITS FAUNA—RESSER OF

ZACANTHOIDEA Swinnerton
ZACANTHOIDES Walcott, 1888
ZACANTHOIDES IDAHOENSIS Walcott
Plate 3, figs. 18-20

Zacanthoides idahoensis Wa.cotrt (part), Smithsonian Misc. Coll., vol. 53,
no. 2, p. 26, pl. 3, figs. 1, 6, 1908. [Not figs. 2-5, 1o= Z. holopygus; figs.
7=Z. disjunctus; figs. 8, 9, 11 = Z. gradatus.]

This species is now restricted by elimination of three other species
included among the original illustrations. Walcott mentioned the
different forms present but did not carry his observations to their
logical conclusion.

Locality 55c.

Lectotype —U.S.N.M. no. 53434; paratypes, nos. 53432, 53438.

ZACANTHOIDES ADJUNCTUS, n. sp.
Plate 3, figs. 13, 14

Zacanthoides idahoensis \WALcoTT (part), Smithsonian Misc. Coll., vol. 53, no. 2,
p. 26, pl. 3, fig. 7, 1908. (See Z. idahoensis.)

Walcott figured a pygidium of this species. The pygidium of Z.
adjunctus is characterized by a wide axis and extensive fusion of the
pleura, so that the tail makes a far more solid plate than in Z.
idahoensis. The outer spine is long but is free only as far as the end
of the axis. The second pair of spines is fairly long, but slender,
and the remaining three pairs of spines are short and sharp-pointed.

Locality 55c.

Holotype and paratypes ——U.S.N.M. nos. 53429, 53427.

ZACANTHOIDES GRADATUS, n. sp.
Plate 3, figs. 15-17

Zacanthoides idahoensis Watcorr (part), Smithsonian Mise. Coll., vol. 53, no. 2,
p. 26, pl. 3, figs. 8, 9, 11, 1908. (See Z. idahoensis.)

This species is characterized by a rather broad pygidial axis. The
marginal spines decrease in size slightly for the first three pairs ; then
abruptly for the remainder. It is thus that the species is distinguished
chiefly from Z. idahoensis in which the pygidial spines continue to
decrease at a fairly even rate from the outer to the inner pair.

The cranidium is referred to the species on the basis of its wide
glabella.

Locality 55c.

Holotype —U.S.N.M. no. 96521; paratypes, nos. 53430-I, 3.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLO?

ZACANTHOIDES ABBREVIATUS, n. sp.
Plate 2, figs. 7-9

Z. abbreviatus is represented by fewer specimens than most other
species in the Spence shale. It is characterized by a rather wide axis
and a considerable degree of fusion in the pygidium. Like Z. ida-
hoensis the pygidial spines decrease in size at a regular rate from the
outer to the inner pair. But these spines are shorter, the axis stouter
and the degree of fusion of the pygidium greater than in 7. zdahoensis.

Locality 55¢c.

Holotype and paratypes.—U.S.N.M. no. 96501.

ZACANTHOIDES SERRATUS, n. sp.
Plate 2, figs. 4-6

This small species is relatively uncommon. Walcott had illustra-
tions of this distinctive pygidium prepared, but recognizing it as a
separate species, did not publish the figures. Aside from the slender
axis throughout, the rather long thoracic pleura and the pygidial struc-
ture characterize the species. In it the spines are fused but remain
distinet, terminating in rather broad short spines. These spines termi-
nate at nearly a straight line, hence give the rear of the pygidium
its serrate margin.

Locality 55c.

Holotype and paratypes—U.S.N.M. no. 96502.

ZACANTHOIDES HOLOPYGUS, n. sp.
Plate 2, figs. 10-12

Zacanthoides idahoensis Watcott (part), Smithsonian Misc. Coll., vol. 53, no. 2,
p. 26, pl. 3, figs. 2-5, 10, 1908. (See Z. idahoensis.)

At first it was thought that this small species was merely a young
stage of one of the larger forms, but careful sorting shows that such
is not the case. Z. holopygus varies in length from less than one-
eighth of an inch to more than 2 inches. As a whole this trilobite
has a more even oval shape than most species of Zacanthoides, which
is due to the fact that the thoracic terminations are relatively broader.
It is the most common Spence shale species.

Z. holopygus has rather large eyes, and the anterior facial suture
diverges sharply, leaving rather long anterior angles. The pygidium
is fused into a solid shield, including all marginal spines except the

NO. 12 SPENCE SHALE AND ITS FAUNA—RESSER oe

outer pair, the other spines being reduced to a serrated border. The
long thoracic spine is not on the fifth but the last segment.

Locality 55¢c.

Holotype—U.S.N.M. no. 96522; paratypes, nos. 53435-6-7, 53440.

DOLICHOMETOPINAE Walcott
BATHYURISCUS Meek, 1873
BATHYURISCUS ATOSSA Walcott
Plate 5, fig. 15

Bathyuriscus atossa Watcotr, Smithsonian Misc. Coll., vol. 64, no. 5, p. 336,
pl. 48, figs. 2, 2a-b, 1916. ReEsser, Smithsonian Misc. Coll., vol. 93, no. 5,

p. 15, 1035.

Locality 55c.

Lectotype -—U.S.N.M. no. 62642; paratypes, nos. 62643-4.

BATHYURISCUS BRIGHAMENSIS, n. sp.
Plate 5, figs. 3, 4

This species is fully typical of the genus. It is characterized by a
rather wide glabella, nine thoracic segments, and a rather wide
pygidium. A broad indentation notches the rear margin rather deeply,
and the anterior marginal pygidial spines are small.

The holotype evidently is a pathological specimen, for the anterior
pleural segments on the right side have coalesced and in healing an
injury have produced an extraordinarily long spine.

Locality 20x.

Holotype and paratype —U.S.N.M. no. 96524.

POLIELLA Walcott, 1916
POLIELLA CARANUS (Walcott)
Plate 2, fig. 21

Bathyuriscus (Poliella) caranus Watcorr, Smithsonian Misc. Coll., vol. 64,

no. 5, p. 350, pl. 46, fig. 5, 1910.
Poliella caranus REsseER, Smithsonian Misc. Coll., vol. 93, no. 5, p. 44, 1935.

Locality 55c.
Holotype —U.S.N.M. no. 62628.

E2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

POLIELLA ANTEROS Walcott
Plate 2, figs. 19, 20
Bathyuriscus (Poliella) anteros Watcort, Smithsonian Misc. Coll., vol. 64,
no. 5, p. 349, pl. 46, fig. 5, 1916.

Locality 55c.
Holotype.—U.S.N.M. no. 62622.

GLOSSOPLEURA Poulsen, 1927
GLOSSOPLEURA BION (Walcott)
Plate 5, figs. 1, 2
Dolichometopus bion Watcort, Smithsonian Misc. Coll., vol. 64, no. 5, p. 363,
pl. 52, figs. 2-2c, 1916.

Glossopleura bion ReESSER, Smithsonian Misc. Coll., vol. 93, no. 5, p. 32, 1935.

Locality 55¢c.

Cotypes—U.S.N.M. nos. 62709-12.

GLOSSOPLEURA SIMILARIS, n. sp.
Plate 5, figs. 9-11

This species is similar to G. producta. The eyes, however, are
shorter and the palpebral lobes smaller, and in the pygidium fusion is
less complete. The species is characterized by a rather narrow pygidial
border, a narrow doublure, and segmentation in both axis and pleural
lobes. The palpebral lobes are strongly bowed. Fine irregular lines
cover the surface.

Localities 55¢ and 54L.

Holotype—U.S.N.M. no. 96525; paratypes, no. 96520.

GLOSSOPLEURA UTAHEWNSIS Resser
Plate 5, fig. 16

Bathyuriscus productus Watcorr (part), U. S. Geol. Surv. Bull. 30, p. 217,
pl. 30, figs. 1, 1b, 1886.

Bathyuriscus anax Waxcotr (part), Smithsonian Misc. Coll., vol. 64, no. 5,
p. 335, pl. 48, fig. 1b, 1916.

Glossoplewra utahensis RESSER, Smithsonian Misc. Coll., vol. 93, no. 5, p. 33, 1935.
The pygidium, identified with Clavaspidella anax, is refigured.
Locality 55e. (Holotype locality 30a, Big Cottonwood Canyon,

Wasatch Mountains. )

Holotype and paratypes.

U.S.N.M. nos. 62641, 62639.

NO. 12 SPENCE SHALE AND ITS FAUNA—RESSER T3

GLOSSOPLEURA GIGANTEA, n. sp.
Plate 5, fig. 17

This is the largest species known in the genus, and is moreover
one of the largest Middle Cambrian trilobites known except, of
course, the enormous species of Paradoxides. The holotype is over
'5 inches long, and other fragments show that the average size is
nearly as great.

This species is characterized by a large glabella, somewhat swollen
in front. There appear to be 8 thoracic segments. Compared with
other Middle Cambrian species, the pygidium of G. gigantea has a
distinctly circular outline.

Locality 20x.

Holotype —U.S.N.M. no. 96527.

CLAVASPIDELLA Poulsen, 1927
CLAVASPIDELLA BITHUS (Walcott)
Nee Fy ner, 12
Bathyuriscus? bithus Waucort, Smithsonian Misc. Coll., vol. 64, no. 5, p. 340,

pl. 47, figs. 4, 4a, 1916.
Clavaspidella bithus Resser, Smithsonian Misc. Collvolkio3s nes 5) py Zom935

Locality 55c.
Lectotype —U.S.N.M. no. 62635; paratype, no. 62636.

CLAVASPIDELLA ANAX (Walcott)
Plate 5; igs. 5, 6

Bathyuriscus anax Wat.corr (part), Smithsonian Misc. Coll., vol. 64, no. 5,
sss .D 4S, figs. t, 1a, c,d, 1o16.
Clavaspidella anax RressErR, Smithsonian Misc. Coll., vol. 93, no. 5, p. 21, 1935.

Localities 55e and 32x.
Lectotype —U.S.N.M. no. 62637; paratypes, nos. 62638-40.

ORYCTOCEPHALIDAE Raymond
ORYCTOCEPHALUS Walcott, 1886
ORYCTOCEPHALUS WALCOTTI Resser
Plate 2, figs. 15-18

Oryctocephalus walcotti Resser (part), Smithsonian Mise. Coll., vol. 97, no. 3,
p. 9, pl. 1, fig. 23, 1938.

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE-97,

This species is common in the Spence shale.

Middle Cambrian, Lakeview; near Lakeview, Pend Oreille Lake,
Idaho.

Localities 55c and 5¢.

Plesiotypes—U.S.N.M. nos. 96503-4.

ORYCTOCARE Walcott, 1908
ORYCTOCARE GEIKEI Walcott
Plate 2, figs. 22, 23
Oryctocare gcikei Watcott, Smithsonian Misc. Coll., vol. 53, no. 2, p. 23, pl. 1,
figs. 9, 10, 1908.

No additional specimens of this rare trilobite were found when
the thousands of specimens from the locality were again examined.
The mistaken reference of this species to Oryctocephalus and Utia
in the Lakeview limestone of Pend Oreille Lake, Idaho, has been
adjusted elsewhere.

Locality 55c. Lakeview ; near Lakeview, Pend Oreille Lake, Idaho.

Lectotype and paratypes.—U.S.N.M. nos. 53426-28.

LEIOSTEGIDAE Bradley
OLENOIDES Meek, 1877
OLENOIDES WAHSATCHENSIS (Hall and Whitfield)
RiaterA nese ie

Dikellocephalus wahsatchensis Hatt and Wuitrietp, U. 5 Geol. Expl. 4oth
Pat: vol..4; ps 243,)pl.. 0, mie. 35.) lore

Dikellocephalus ? gothicus Ean and WuHuitTFIELD, U. S. Geol. Expl. 4oth Par.,
vol. 4, p. 242, pl. 1, fig. 36, 1877

Olenoides wahsatchensis Waucott, U. S. Geol. Surv. Bull. 30, p. 189, pl. 20,
figs. 2, 2a, 1886.

Locality 20x.
Cotypes—U.S.N.M. no. 15447.

OLENOIDES BRIGHAMENSIS, n. sp.
Plate 3, figs. 8, 9

This is a small species associated with Ehmaniella and differs from
the other two Wasatch species in several respects. The cranidium
does not depart from the norm, and is rather highly arched in both
directions. The pygidium has six pairs of marginal spines which are
rather short.

Locality 20x.

Holotype and paratype —U.S.N.M. no. 96513.

SNOy 12 SPENCE SHALE AND ITS FAUNA—RESSER 15

OLENOIDES EVANSI, n. sp.
Plateray figsseaiea

This species is characterized by a quadrate glabella similar to species
of Kootema, but the pygidium has the true Olenoides feature of sepa-
rate pleura. It has seven pairs of spines. The pygidium of O. evansi
resembles that of O. elongatus due to its triangular shape and elonga-
tion to accommodate seven spines. The occipital ring and thoracic seg-
ments each bear a small median spine or tubercle, which are lacking
from the axial rings of the pygidium.

A crushed and poorly preserved hypostoma occurs with portions
of a few pygidia and is thought to belong to the species.

Locality 55e.

Holotype and paratype.—U.S.N.M. no. 96514.

KOOTENIA Walcott, 1888
KOOTENIA IDAHOENSIS, n. sp.
Plate 3, figs. 1-3

This species belongs to the serrata group of the genus and is a
six-spined form. The cranidium has the usual quadrate outline and
proportions, and the surface is ornamented with lines and granules.
The neck spine is slender and not very long.

The thorax has seven segments terminating in rather blunt spines,
which have the usual elongate, scaly granulations.

Six blunt spines margin the pygidium. All spines are coarsely
granulated.

Locality 55c.

Holotype and paratypes——U.S.N.M. no. 96505.

KOOTENIA SPENCEI, n. sp.
Plates sigs) 4505

Only two incomplete pygidia have been found of this seven-spined
species. Unfortunately, neither pygidium is well preserved. The
pleural grooves remain on exfoliated specimens. Six pairs of the
marginal spines are long and slender, possibly attaining a length
greater than the pygidium. Each spine had a medial furrow for a
considerable distance from the border. The seventh pair of spines is
short.

Locality 55c.

Holotype and paratypes —U.S.N.M. no, 96500.

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

KOOTENIA MATHEWSI, n. sp.
Platess nese Ow

This species belongs to the seven-spined group. It is characterized
by long, round spines, the longest likely exceeding the length of the
pygidium. The seventh pair is short and slender.

This species is very similar to K. spencei, being distinguished by
the smaller spread and curvature of the spines and their lack of
median grooves.

Locality 20x.

Holotype and paratype-—U.S.N.M. no. 96507.

KOOTENIA GRACILIS, n. sp.

Plates ness nies 12

This beautiful trilobite is associated with Ehmaniclla and Olenoides
brighamensis in the thin limestone layers intercalated in the shales.

K. gracilis is one of the few six-spined species in the genus. Evi-
dently the seventh pair, which is usually greatly reduced, has become
altogether obsolescent. The glabella is long, but otherwise the cra-
nidium is normal in all respects. Fusion is normal in the pygidium
and the long, slender spines are straight and hence well spread out.

Locality 20x.

Holotype and paratype-—U.S.N.M. no. 96508.

PTYCHOPARIDAE Matthew
ALOKISTOCARE Lorenz, 1906
ALOKISTOCARE IDAHOENSE, n. sp.
Plate 4, figs. 8, 9
This species has 23 or 24 thoracic segments and is finely granulated
on the outer surface. Exfoliated surfaces show lines and punctation.
The genal spines extend back to about the fifth thoracic segment.
The pygidium is completely fused.

Locality 55¢c.
Holotype and paratype -—U.S.N.M. no. 96507.

ALOKISTOCARE SPENCENSE, n. sp.

Plate 4, figs. 10, II

This is also an abundant species associated with A. idahoense.
Compared to that species, A. spencense is characterized by a narrower

NO. I2 SPENCE SHALE AND ITS FAUNA—RESSER 17

cranidium. In keeping with that narrowness, the trilobite as a whole
is more slender.

Localities 55c, 55e, and 20x.

Holotype and paratypes—U.S.N.M. no. 965106.

ALOKISTOCARE LATICAUDUM, n. sp.
Plate 4, figs. 15-19

This species is characterized by a wide brim and wide, long, genal
spines, a wide pygidium and 17 thoracic segments. The genal spines
extend back to about the 14th segment. Fusion has not obliterated
pleural furrows in the pygidium.

Localities 55¢ and 55e.

Holotype and paratype —U.S.N.M. nos. 96517, 8.

ALOKISTOCARE SEPTUM, n. sp.
Plate 4, figs. 5-7

This is a wide-tailed form like A. laticaudum. It ditfers in having
a narrower glabella. But the distinctive feature is the great genal
spines which extend almost to the pygidium.

Locality 55c.

Holotype and paratype.—U.S.N.M. no. 96520.

ALOKISTOCARE PUNCTATUM, n. sp.
Plate 4; figs. 20, 21

Reduction of brim brings this species closer to Chancia, than most’
species referred to Alokistocare. But since a rim is not differentiated
by thickening, this form is called Alokistocare.

The cranidium is rather wide and the anterior margin not so much
curved. On exfoliated specimens the eye lines are heavy and the
surface is strongly punctate. One specimen retains 15 thoracic seg-
ments, indicating a thorax of possibly 20 segments.

Locality 55e.

Holotype and paratypes —U.S.N.M. no. 96519.

CHANCIA Walcott, 1924
CHANCIA EBDOME Walcott
Plate 4, figs. 12-14
Chancia ebdome Watcortt, Smithsonian Misc. Coll., vol. 75, no. 2, p. 55, pl. 10,
fig. 4, 1924. Idem, no. 3, p. 80, pl. 17, fig. 26, 1925.

Locality 55c.
Holotype.—U.S.N.M. no. 70274.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

CHANCIA EVAX Walcott
Plate 5, figs. 18, 19

Chancia evax Wat.cotr, Smithsonian Misc. Coll., vol. 75, no. 3, p. 81, pl. 17,
fig. 27, 1925.
Localities 55¢ and 55e.
Holotype —U.S.N.M. no. 70275.

CHANCIA ANGUSTA, n. sp.
Plate 5) figs. 13) 14

This species differs from C. ebdome chiefly in that the cranidium
is narrower, and that the surface of the head is more finely granulose.
C. angusta is characterized by a glabella which tapers at the usual
rate and attains slightly more than half the cranidial length. Three
pairs of furrows are faintly defined, the rear pair being directed
sharply backward. Eyelines are well developed on the under side of
test. Eyes moderately elevated.

Exfoliated specimens have all furrows accentuated, and the doublure
impression modifies the appearance of the unfurrowed brim.

Locality 55¢c.

Holotype and paratypes—U.S.N.M. no. 96523.

ALOKISTOCARELLA Resser, 1938
ALOKISTOCARELLA SPENCEI, n. sp.
Plate 3, fig. 10

This species is represented by several cranidia, all of which are, »
unfortunately, almost entirely exfoliated. This, of course, accentuates
all furrows and ridges. A. spencei is characterized by a truncate
glabella of normal size and an upturned narrow rim. The width
of the fixigene at the eye is about equal to that of the glabella at the
same point, which makes the cranidium wide.

Locality 55¢c.

Holotype.-—U.S.N.M. no. 96509.

ELRATHIA Walcott, 1924
ELRATHIA SPENCEI, n. sp.
Plate 6, figs. 15-17
This species has 17 thoracic segments. The cranidium is wide, the
glabella occupying about three-fourths its length. The small pygidium
is characteristic of the genus, and it has an indented rear margin.

Locality 55¢c.
Holotype and paratypes —U.S.N.M. no. 96540.

NO. I2 ; SPENCE SHALE AND ITS FAUNA—RESSER IQ

ELRATHIA RARA, n. sp.
Plate 6, fig. 18

This species is based on a single individual, consisting of the
cranidium and 17 segments. Most of the cranidium is exfoliated, and
shows strong striations on the preglabellar area. A narrow, straight
rim is demarcated.

Locality 55c.

Holotype —U.S.N.M. no. 96541.

ELRATHINA Resser, 1937
ELRATHINA OFFULA, n. sp.
Plate 2, figs. 13, 14

This genus and species is represented by only a few cranidia. The
illustrations show the relative proportion of the several parts, the
characteristic constriction of brim width, and the slightly concave
brim, with the rim only weakly defined. Eyes are rather small.

Locality 55c.

Holotype and paratype —U.S.N.M. no. 96510.

EHMANIELLA Resser, 1937
EHMANIELLA QUADRANS (Hall and Whitfield)
Plate 6, figs. 28-32

Crepicephalus? (Loganellus) quadrans Hatt and Wuirrietp, U. S. Geol. Expl.
40th Par., vol. 4, p. 238, pl. 2, figs. 11-13, 1877. ;
Ptychoparia quadrans Waucott, U. S. Geol. Surv. Bull. 30, p. 199, pl. 20,
figs. 4, 4a, b, 1886; Smithsonian Misc. Coll., vol. 64, p. 204, 1916.
Ehmaniella quadrans REssER, Smithsonian Misc. Coll., vol. 95, no. 22, p. 10, 1937.
Localities 20x, 55e, and 54L.
Holotype and paratypes—U.S.N.M. no. 15432; plesiotype, no.
96539.

CLAPPASPIS Deiss, 1939

The genotype, C. typica Deiss, comes from the Pentagon shale of
Lewis and Clark Range, Montana. Average size and shape, normal
development of eye lines and furrows, and eyes situated slightly be-
hind the middle of the head, characterize the genus. All species are
evidently ornamented by some sort of granulated surface. Deiss did
not have any entire individuals and so could not illustrate the thorax.
Several Spence shale individuals have 14 thoracic segments.

Five species in the Spence shale are referred to Clappaspis.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Clappaspis appears to be related to Ehmamniella or at least most
closely resembles that genus. The only differences of consequence are
the slightly narrower preglabellar area and the granulated surface, for
the pygidia are clearly constructed on the same plan.

CLAPPASPIS SPENCEI, n. sp.
Plate 6, figs. 5, 6

This is the widest of the five species recognized in the Spence shale.
It is further distinguished by two sets of granules rather evenly dis-
tributed, the smaller ones being more numerous. Exfoliated speci-
mens have coarse irregular striations on the preglabellar area.

Locality 55c.

Holotype and paratypes —U.S.N.M. no. 96530.

CLAPPASPIS IDAHOENSIS, n. sp.
Plate 6, figs. 26, 27

Three complete individuals represent this species. The cranidium
is narrower than C. spencei and has a shorter, more conical glabella.
C. idahoensis is characterized by scattered granules of uneven size.
The thorax has 14 segments.

Locality 55c.

Holotype and paratypes —U.S.N.M. no. 96531.

CLAPPASPIS LANATA, n. sp.

Plate 6, fig. 7

A single cranidium represents this species, which has a narrow
cranidium, long glabella, and the surface is closely crowded with small
granules. Rather heavy striations occur on the preglabellar area.

Locality 55c.

Holotype —U.S.N.M. no. 96532.

CLAPPASPIS CORIACEA, n. sp.
Plate 6, figs. 11, 12

This species is rather wide. Coarse granules are widely scattered
over the surface, which is covered with a small set of closely crowded
granulations. Exfoliated specimens appear pitted.

Locality 55c.

Holotype and paratypes —U.S.N.M. no. 96533.

NO. I2 SPENCE SHALE AND ITS FAUNA——RESSER 21

CLAPPASPIS DOTIS, n. sp.
~ Plate On igss 135,04

This species is represented by the largest number of specimens.
It is characterized by a short glabella, and closely crowded granula-
tions which are of medium size.

Locality 55c.

Holotype and paratypes—vU.S.N.M. no. 96534.

FAMILY UNDESIGNATED
VISTOIA Walcott, 1925
VISTOIA? MINUTA, n. sp.

Plates2y igs cree

One of the illustrated specimens of this species is mounted as a
cranidium, but this may be only an accidental resemblance. Certainly
most of the examples available are pygidia. Whenever such trilobites
as these retain their full relief, difficulty is experienced in distinguish-
ing heads and tails. Consequently, the question is not brought to a
final conclusion.

Even though this trilobite is very small, it is referred to Vistoia
chiefly because it fits no other Middle Cambrian genus in any re-
spect. It may well be that entire specimens will show that the cra-
nidium disagrees with that of Vistoia prisca and then a new genus
Gan) be erected.

Aside from its small size, V. ? minuta is characterized by sim-
plicity. The pygidium has a simple, highly arched semicircular out-
line. An axis about a third the width of the pygidium is faintly out-
lined by shallow dorsal furrows, which do not unite in the rear except
by the faintest trace. One specimen retains several simple thoracic
segments.

Localities 55¢ and 5¢.

Holotype and paratypes——U.S.N.M. no. 96511.

UTIA Walcott, 1925
UTIA CURIO Walcott
Plate 2, figure 3
Utia curio Watcort, Smithsonian Misc. Coll., vol. 75, no. 2, p. 60, pl. 14, fig. 4,

1924. Idem, no. 3, p. 119, pl. 15, figs. 11-14, 1925. RESSER (part), idem,
vol. 97, no. 3, p. 9, pl. I, figs. 19, 20, 1938.

Several hundred specimens of this unique trilobite have now been
segregated. One of the cotype cranidia has about nine thoracic seg-
ments attached, but no pygidium has been assigned to the species.

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

Locality 55c. Lakeview; (loc. 37n) Lakeview, Pend Oreille Lake,
Idaho.
Cotypes.—U.S.N.M. nos. 70235-7; plesiotypes, no. 95041.

BYTHICHEILUS, n. gen.

Trilobites of less than average size. Cranidium rather large.
Glabella well defined except in front where the dorsal furrow
coalesces with the depression in the brim. Fixigenes about as wide
as the glabella. Eyelines formed by abrupt depression of anterior
fixigenes. Eyes rather large, situated back of the cranidial midpoint.
Brim less than one-third cranidial length. Narrow, striated upturned
rim clearly defined. Preglabellar area depressed in a peculiar man-
ner, which suggests generic name. Libragenes narrow with moderate
genal spines. Striated rim sharply upturned almost to the tip of the
genal spines.

Thorax with 14 segments, rather straight, parallel to each other,
divided evenly by pleural furrows and bent down rather abruptly at
geniculation.

Pygidium small, completely fused, with dorsal furrow shallow and
not clearly defined around the rear.

Genotype.—B. typicum, new species.

BYTHICHEILUS TYPICUM, n. sp.
Plate 6, figs. I-4

The generic description and illustrations present the specific fea-
tures adequately.

Locality 55c.

Holotype and paratypes ——U.S.N.M. no. 96537.

BYTHICHEILUS ALVEATUM, n. sp.
Plate 6, figs. 8-10

This species differs from B. typicwm in having a longer depression
in the preglabellar area, a less upturned rim, less indentation of the
front of the glabella by the median depression, and stronger eye-
ridges due to the more abrupt slope of the brim.

B. alveatum is characterized by a curved anterior margin, fixigenes
about three-fourths as wide as the glabella, and by a depressed pre-
glabellar area. Exfoliated specimens show pronounced glabellar fur-
rows and strong eyelines.

Locality 55c.

Holotype and paratypes —U.S.N.M. no. 96538.

NO. I2 SPENCE SHALE AND ITS FAUNA—RESSER 23

SPENCIA, n. gen.

Small trilobites characterized by a rather large, slightly tapered
glabella which extends to the anterior furrow. Brim reduced to
thickened rim, wider in the middle than at the ends. This causes the
anterior furrow to join the dorsal furrow in front of the glabella.
Fixigenes convex, anteriorly nearly as wide as the glabella. Eyelines
fairly prominent. Eyes small, situated slightly behind the middle of
the head.

Libragenes small, with short but sharp-pointed genal spines.

Thorax has 16 segments. The pygidium is small, trilobate, and
well fused.

Surface of cranidium and ridges of the thoracic segments marked
by scattered granules.

Genotype.—S. typicalis, new species.

SPENCIA TYPICALIS, n. sp.
Plate 6, figs. 22-25

This species is characterized by scattered granules and a rather
wide space where the anterior and dorsal furrows join.

Locality 55c.

Holotype and paratypes—U.S.N.M. no. 96535.

SPENCIA PLENA, n. sp.
Plate 6, figs. 19-21

This species is characterized chiefly by the swollen rim and the
narrowing of the preglabellar area or rather the anterior and dorsal
furrows. Scattered granules occur on the more elevated portions of
the cranidium.

Locality 55c.

Holotype and paratypes—vU.S.N.M. no. 96536.

STAUROHOLCUS, n. gen.

Small trilobites ; only cranidia known. Glabella about three-fourths
the length of cephalon, without furrows, truncate with rounded
anterior corners, tapering only slightly. Fixigenes fully as wide as
glabella. Eyelines faint, but clearly defined. Eyes small, situated
about the mid-point. Occipital spine present. Brim rounded at an-
terior corners. Rim thickened in center, upturned. Preglabellar area

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

depressed in center, giving rise to the appearance of the anterior and
dorsal furrows crossing each other in the center.

Naime.—oravpos = cross ; oAxos = furrow.

Genotype.—S. typicalis, new species.

STAUROHOLCUS TYPICALIS, n. sp.
Plate 5, figs. 7, 8

The illustrations and generic description portray the specific char-
acters.

Locality 55c.

Holotype and paratypes —U.S.N.M. no. 96528.

REFERENCES

DEIss, CHARLES.
1938. Cambrian formations and sections in part of Cordilleran trough.
Bull. Geol. Soc. Amer., vol. 49, pp. 1067-1168.
1939. Cambrian stratigraphy and trilobites of northwestern Montana. Geol.
Soc. Amer., Spec. Paper 18.
IRESSERS | Co9hien ge
1938. Middle Cambrian fossils from Pend Oreille Lake, Idaho. Smithsonian
Misc. Coll., vol. 97, no. 3.
Watcort, C. D.
1908a. Nomenclature of some Cambrian Cordilleran formations. Smith-
sonian Misc. Coll., vol. 53, no. 1, April.
1908b. Cambrian sections of the Cordilleran area, Smithsonian Misc. Coll.,
vol. 53, no. 5, December.

EXPLANATION OF PLATES

All figures are natural size unless otherwise designated. Species without
locality references are all from locality 55c.

PLATE I

LEG HOM GHELTOES. SUSE BOT ESbc 55 lose pb oo pono unos ne bosecoumod péouo 5
Pica Wentralavalver(<<3))).
Fics. 2, 3. Two dorsal valves (xX 3.5).

Acratretaleudta ew SPECS... Gunes ne Vleet are eee 6
Fic. 4. Narrow false pedicle groove ( 6).
Fic. 5. Interior of ventral valve (x 6).
Fic. 6. Interior of dorsal valve (x 3).

Acrolreta sae pia alcorts ns ncctance Cae eee ace eae eae 5
Fic. 7. Exterior ventral valve ( 4).
Fics. 8, 9. Exterior and interior of dorsal valves (x 4).
Fic. 10. Interior ventral valve ( 4).

Micromiiva lepidas newispecieSs.%34,. 22eian in ciontee aes ee ee ee 4
Fic. 11. Somewhat crushed ventral valve (> 1.5).
Fics. 12, 13. Exterior and interior of dorsal valve (xX 1.5).

NO! £2 SPENCE SHALE AND ITS FAUNA—RESSER 25
PAGE
Ie ILOMO LALO PTIEW) ‘SPECIES «3,553 aa ne ream intra eee Eee eee 4

Fics. 14, 18. Ventral valves (xX 3).
Fics. 15, 17. Molds of the exterior surface of dorsal valves (X 2).
Fic. 16. Dorsal valve (X 3).
FEROLRCLOMIUINIS§. TIEW, “SPECIES '« Acc 5,015 ona Oe OR e E Ere en. 5
Fic. 19. Ventral valve (X 2).
Fic. 20. Dorsal valve (X 1.5).
Mies. 21, 22. Interiors -(>X< 2:5).
Hamanellmenautes (Walcott) tax. ces eect coe ae eae at ok eae 6
Fig. 23. Ventral valve (xX 2). ;
Fic. 24. Dorsal valve (X 2).
Fic. 25. Dorsal valve (X 3).
Fic. 26. Ventral valve (X 1.5).

ignamelloms ences: (WalGOtt )'.0<:¢..cc:c5 ee esa eee ae oe eee ee 6
Fics. 27, 28. Ventral valve and mold of exterior.

LCR Od CNVAL COLE) =. scc, hess 8c eo sais ee Per 6
Fic. 29. Only specimen known.

MOMMIES MORNOLEIIUS ITIEW SPECIES = 215 2 sists oe cen Giesinre Oe pee he eee 7
Fics. 30, 32. Posterior surface (X 2).
Fic. 31. Anterior surface (x 2).

MONE Aline S010, SICW SPECIES. bi cjeia idivsios see ee SURE Dea ate ee emer yi

Fic. 33. Round conical tube (X 2).

De IRt A SPencer: “NEW "SPECIES. 5 << ccgdecs coe cs Been Abe e. be os oohen Seek 4
Fics. 34, 35. Several flattened tubes.

RES CEEROPIC NV ALCO 25. o\ciaisaias clva'ss ose edad os smclele alee ee eee aeons
Fic. 36. Posterior surface.
Fic. 37. Specimen showing cameration.
Fic. 38. Operculum.

PA GGHILEO GU CUIUSI ROR o eraih sca line)a's G!siahe eisvahe'gea, 218 £ e/ivsta wes wapeleracomme nae ear aaee ete eibe 3
Fic. 39. Sides of tubes shown in lower and “ends” turned toward

observer in upper portion ( 6).

N

VOOR GMEES PY SeaLITIO Cle a caia)2) 3, Sones -hace inva ies oc Se deinen mele ee Rune ORE oes EE 3
Fic. 40. Irregular ovate pieces of the alga.
Baeemusvonguactyius (Walcott) .< . 6 c.ccossoces ounces seh tnness oesee es 3

Fic. 41. Plate with the most common style of ornamentation (x 6).
Fic. 42. Poorly preserved calyx with several attached arms (X 2.5).

PLATE 2

LASTOLARININUL: TICW SPECIES i.0106-6 Dx Ao ariiafers sede cleriern e atelier bhieye 21
Fic. 1. Supposed cranidium (X 6).
Fic. 2. The holotype pygidium (xX 6).

TOG VOM NVAlCOtt secs wold goon 31d Oe Se raid PER e coe Gib sateoehe rcteusinreeiae 21
Fic. 3. Dorsal view of cranidium (X 2).
Aaconinoldesservatrus, Tew. SPECIES. «cc... 5 sorte sae eee ee oer sieteteieys = 10

Fic. 4. Cranidium retaining left libragene (x 2).
Fic. 5. Pygidium, with exfoliated spines (xX 2).
Fic. 6. Holotype with pygidium and thorax (X 2).

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 97

ZLacanthoides aborevidius. Mew SpeGleSs.. oe ein aaa eie ea ee eee 10
Fics. 7 (X 2), 8. Small pygidia showing the stout axis.
Fic. 9. Nearly complete holotype.

Zactanthotdes polopygus; Mew SPpeCieS... 20 sen anes oa at ew Dye Woe crema ro Sere 10
Fic. 10. Holotype.
Fic. 11. Several individuals.
Fic. 12. Pygidium (xX 2).

Birathina offiula, mew SpCGles sso e asics sic cieirairekeuteleeie eee Seater eect ere 19
Fics. 13 (X 2), 14 (X 1.5). Paratype and holotype cranidia.
Orvctocephalus cdl GovtimReSSeri raster tencvetetecicle 1+ aioe te cesar ie ate 13
Fic. 15. Pygidium, thorax and doublure with attached hypostoma
(X15).

HrGSe on @Oxales) salen Grandia.
Fic. 17. Pygidium (XX 2).
Poliella anteros (Walcott) )x).64 .¢) a. 27- as: eh ae Aho eins ciety bare a ere 12
Fic. 19. Holotype.
Fic. 20. Underside with hypostoma.

Roltellaccananus {(WialCott))s ences erica ehe creed ere eek lero re en ra aeier II
Fic, 21. Holotype (Xx 2).
Orvetacare’ getkes. NValcott® -2..035. cc sao anne oe eae Rete: ee eae ae mae 14

Fic. 22. Holotype (X 5).
Pres 225 seycidiminn (Cxe2)-
ACGHOSEUS BOMMENEMSIS, JIRESSEL. fra clacietcicie eter ce reasietrrec ie ate cree erent 8
Fic. 24. A good example (X 6).
Fics. 25, 26. Separate shields (x 6).
AGROStUSLDHIGHOMeNsIS, Mew SPECIES ere eee eer akan ener 8
Fic. 27. Holotype pygidium ( 6).
Fic. 28. Cranidium, Pagetia and Lingulella (x 6).
Fic. 29. Pygidium, Oryctocephalus and Clavaspidella (xX 3).
Locality 20x.
Pagetiacls tia Wal cout ne paises tee oe ae ele ahaa /< antes eee eee 8
Fic. 30. Several entire individuals (X 6).
Fic. 31. Cranidium ( 6).
Fic. 32. Pygidium (xX 6).

IMoOoteMiandanocnsis mew SDCClesnme cet eerie cme oe ee ere eeecio 15
Fic. 1. Holotype.
Fic. 2. Partially exfoliated pygidium.
Fic. 3. Cranidium with occipital spine.
Riootema-spencet. Mew; SPECIES: we < sinks os feels ones vie Seattle tea eae as 15
Fic. 4. Holotype pygidium (xX 1.5).
Fic. 5. Portion of pygidium showing pleural development (X 1.5).
Kootenia mathewsi, new species.......... rE IN tS BO AUN oe oe ee ee 160
Fic. 6. Holotype pygidium.
Fic. 7. Squeeze of an imperfect cranidium.
Locality 20x.

NO. I2 SPENCE SHALE AND ITS FAUNA—RESSER

ienmaes brighamensis, New SpeCieS......<. Gan voce dvs Vass d oclen ae Ho dade we
Fic. 8. Cranidium.
Fic. 9. Squeeze of pygidium.
Locality 20x.

PAELOGAL CLIC. SPCNCEL, TIEW SPECLES® ./<a) gies dalomen,oou a ve eeen asset o's ee ale
Fic. 10. Holotype cranidium.

PE aaeGrCilis? TENG ‘SPECIES: « sc. silsiee «ohn onde erin ane chy ani
Fic. 11. Squeeze of cranidium.
Fic. 12. Holotype pygidium.
Locality 20x.

AnniMrnordes ad punctus,. NEW SPECIES)... jos 2s 4. e.aca cen acess aa a
Fic. 13. Holotype.
Fic. 14. Well-preserved pygidium.

PANCMINOUOCSMOROGGIUS: TIEW ‘SPECIESe... .. gcc a0 hunted ete aeie cis soa sels eee
Fic. 15. Well-preserved pygidium.
Fic. 16. Cranidium with right libragene.
Fic. 17. Holotype.

PAIGUNIMOLAES AAAHOEMSIS; NEW SPCCICSs oscsgieten aces ons ecsescnoems aaa elses
Fic. 18. Cranidium with occipital spine.
Fics. 19, 20. Nearly complete individuals.

PLATE 4

Olenoides wahsatchensis (Hall and Whitfield) ................0.2 ec ceeeee
Fic. 1. Squeeze of cranidium.
Fic. 2. Pygidium.
Locality 20x.

DICH OTMESREUGIVSH, TIEW » SPECIES eco.5 5 coca Soe» 6 «9's oo ora re Pctevnatefaapal state ieeeanerete
Fic. 3. Cranidium
Fic. 4. Holotype.
Locality 55e.

lakistocare sepium, New SPeCi€S... ii. .sceeecseccss see oeseseeccuabenen
Fic. 5. Individual showing brim well.
Fic. 6. Small cranidium.
Fic. 7. Holotype.

Ellapistocare wdanoense: tlew SPCCIES. . 26.00. cenee e+ se ees ote eee
Fic. 8. Libragene (X 1.5).
Fic. 9. Holotype.

Alokistocare spencense, new species.............- ace A SEN oR renee
Fic. 10. Holotype.
Fic. 11. Another individual showing pygidium and changed aspect

due to broken brim.

(QUOTE RCD COMET NVC Ob cis, sarin och cielocacslo cholo Siete s Ie cake ose ee ie gi eer eeerate
Fic. 12. Cranidium.
Fic. 13. Cranidium and displaced libragene.
Fic. 14. A well-preserved specimen.

18

16

14

15

17

16

16

17

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Q7

PAGE
Alokistocare latecoudunn news SpeCleSecac ai oe een eee eee eee eee 17
Fics. 15, 16. Cranidia in calcareous shale.
Locality 55e.
Fic. 17. Incomplete individual.
Fic. 18. Holotype.
Fic. 19. Specimen lacking the libragenes.

ALORASLOCON EM PUNGLOLUNY. NEW, {SDECIES-e a rie ace eit ta eee ieee 17
Fic. 20. Cranidium and portion of thorax.
Fig. 21. Cranidium.
Locality 55e.

Glossopleuraabion “(NValcott)< basen: cence eens citar e ee aye 12
Fic. 1. Cranidium and impression of pygidium.
Fic. 2. Large pygidium.

Bathyuriscus brighamensts, mew SpeCies.....-2-.-s.e..5-+2--4reeess- eekle es II
Fic. 3. Holotype.
Fic. 4. Impression of pygidium.
Locality 20x.

Glavasiidellaxanax (CW alcott)c: cass v ride sede acces eee = cent 13
Fic. 5. Cranidium.
Fic. 6. Lectotype.
Locality 55e.

SraunoMwolewsary picalismmewa SPEClEStie qermens acid oe ciaenteeiancine ir aeiere erate 24
Fic. 7. Exfoliated cranidium.
Fic. 8. Holotype cranidium (x 2).

Glossopleuxa sumilaris new SpeCleSenne ease eerie erie eeiaer 12
Fic. 9. Cranidium and libragene with Hyolithes cecrops.
Fic. 10. Pygidium.
Locality 56 L.
Fic. 11. Holotype pygidium.

Glawspidellasbithws (\WVialcott) acre eceeeere ee eee eee 13
Fic. 12. Lectotype pygidium.

Ghancia-anguistal, Mew SPpeCles's 27s cies sieve cre. ores a aera ne tener Pee tector 18
Fic. 13. Paratype.
Fic. 14. Holotype.

BiG Vusns CUSTauoSSa: NVAlCOttae ner nic eine atic ee note Lee ner Il
Fic. 15. Holotype.

Glossopleuma WignensisMRESSELe ae iene ate a ree bie ae or nore et ket he eee oer 12
Fic. 16. Paratype pygidium.
Locality 55e.

Glossopleuyvasguganiea mew, SpECles. aaa oricine cae cee erie eee 13
Fic. 17. Holotype (<5).
Locality 20x.

Chancia tevar Niall Cotten aaseotcon eRe Meee LR COM TOE Or are ore 18
Fic. 18. Plesiotype cranidium.
Fic. 19. Holotype.

NO. 12 SPENCE SHALE AND ITS FAUNA—RESSER 29

PAGE

Fic. 1. Complete individual and cranidium ( 1.5).

Fic. 2. Individual lacking libragenes (X 2.5).

Fic. 3. Holotype (X 2).

Fic. 4. Small example (xX 3).

inp paspiss spencel. NEW. SPECIES... + «sos a5/aen one ieee a ae ie erie 20
Fic. 5. Exfoliated cranidium.
Fic. 6. Partially exfoliated holotype (x 2).

Penn a EM IAMITI ThE W, - SPECIES... a. a4 5 sig aes OR ee Ste Sele ee eres srse sieeree 20
Fic. 7. Holotype (x 1.5).
SEICNEU US UVUCHLUM. IEW SPECIESA ta. ose 4 ces mos Soe elmeteisinis sui leer aeiererne 22

Fic. 8. Exfoliated cranidia (X 2.5).
Fic. 9. Exfoliated conventionally lighted cranidium (X 2).
Fic. 10. Holotype (X 2.5).
GlappaspesGORiaced, NEW-SPECIES = << s.5¢< <.siscer c's sas cperom ice oie ein’ s oe sieloveiers aheiere 20
Fic. 11. Cranidium.
Fic. 12. Holotype cranidium (x 2).
LSP USMAG LS ell ewWhe SPECIES scx soa oy «2 cts eo alow aise a SS ae eee 21
Fic. 13. Cranidium, with Spencia plena (xX 2).
Fic. 14. Holotype (X 2).
ia ERESHENGCI. NEW SPECIES. . os 2. dels les does sce steaks Gece oceans 18
Fig. 15. Holotype (xX 2).
Fic. 16. Smaller individual retaining libragenes (x 2).
Fic. 17. Exfoliated specimen (xX 2).

PERE IEL ee TIC SD CCLES ore 55-4 is) vo ahs ticle odes et oa eee de Clee meas 19
Fic. 18. Holotype (xX 2).
PE PEa EIEN EME SPECIES. «5% o.oo to einea die eo cen ss neem ee es ane 23

Fic. 19. Partially exfoliated holotype cranidium (x 3).
Fic. 20. Small cranidium (> 3).
Fic. 21. Cranidia with Bythicheilus alveatum ( X 2).
De MPETAEU CRIN TIEW, SPECIES’. o.0)5.2 5 6.5, oie 0.66015 sole a0 cine erates nsere ome reiateratel see 23
Fic. 22. Cranidium showing granules (X 1.5).
Fic. 23. Small cranidium (x 1.5).
Fic. 24. Holotype (X 2).
Bre. 255 Cranidium (< 1-5)-
Ola uasric faGeENSIS, NEW SPECIES: - +... 2 sce sens 22 enbile se beatae 20
Fic. 26. Holotype.
Fic. 27. Two smaller individuals.
Ehmaniella quadrans (Hall and Whitfield) ............--..--.es sees eens 19
Fic. 28. Original type of cranidium (X 2.5).
Fic. 29. A less distorted cranidium (X 3).
Fic. 30. Associated libragene (X 2.5).
Fic. 31. Pygidium (xX 3).
Locality 20x.
Fic. 32. Limestone nodule with cranidia and fragments of pygidia.
Locality 55¢e.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES 97, NOS 425 PEs

BRACHIOPODS, HYOLITHES, AND OTHER FOSSILS
(See explanation of plates at end of text.)

VOL. 97, NO. 12, PL. 2

SMITHSONIAN MISCELLANEOUS COLLECTIONS

AGNOSTIDA AND TRILOBITES

(See explanation of plates at end of text.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. 97;, NOD12;>PEI%3

KOOTENIA, OLENOIDES, AND ZACANTHOIDES
(See explanation of plates at end of text.)

12. PEs

VOL. 97, NO.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

Oe

bi

rr

Ui

, AND CHANCIA

ALOKISTOCARE

OLENOIDES,

(See explanation of plates at end of text.)

VOEW97,, NOD 12; PES

SMITHSONIAN MISCELLANEOUS COLLECTIONS

SPENCE SHALE TRILOBITES

(See explanation of plates at end of text.)

12, PL. 6

VOL. 97, NO.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

SPENCE SHALE TRILOBITES
(See explanation of plates at end of text.)

i?