.
,

SMITHSONIAN.
MISCELLANEOUS COLLECTIONS

VOL. 72

‘““EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES,
AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN ’’—SMITHSON

(PUBLICATION 2706)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1922

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BALTIMORE,

The Lord Gal

ADVERTISEMENT

The present series, entitled “ Smithsonian Miscellaneous Collec-
tions,” is intended to embrace all the octavo publications of the
Institution, except the Annual Report. Its scope is not limited,
and the volumes thus far issued relate to nearly every branch of
science. Among these various subjects zoology, bibliography, geology,
mineralogy, and anthropology have predominated.

The Institution also publishes a quarto series entitled “ Smith-
sonian Contributions to Knowledge.” It consists of memoirs based
on extended original investigations, which have resulted in important
additions to knowledge.

CHARLES» De WALCOTT,

Secretary of the Smithsonian Institution.

(iii)

CONTENTS

. Explorations and field-work of the Smithsonian Institution in

1919. May 10, 1920. 80 pp., 77 figs. (Publ. no. 2881.)

. Hotiister, N. Two new East African primates. January 22,

1920. 2 pp. (Publ. no. 2582.)

. Dixon, H. N. Reports upon two collections of mosses from

British East Africa. September I, 1920. 19 pp.,2pls. (Publ.
no. 2583.)

. Ripaway, Ropert. Diagnoses of some new genera of birds.

December 6, 1920. 4 pp. (Publ. no. 2588.)

. Maxon, Witit1am R. New selaginellas from the western United

States. December 22, 1920. 10 pp.,6 pls. (Publ. no. 2589.)

. Explorations and field-work of the Smithsonian Institution in

1920. May 12,1921. 126 pp., 138 figs. (Publ. no. 2619.)

. CLark, Austin H. Sea lilies and feather stars. April 28, 1921.

43 pp., 16 pls. (Publ. no. 2620.)

. WincE, Hertur. A review of the interrelationships of the

cetacea. July 30,1921. 97 pp. (Publ. no. 2650.)

. Britton, N. L. anp Rose, J. N. Neoabbottia, a new cactus genus

from Hispaniola. June 15, 1921. 6 pp., 4 pls. (Publ. no.
2651.)

. Foote, J. S. The circulatory system in bone. August 20, 1921.

20 pp., 6 pls. (Publ. no. 2652.)

. CyarK, Austin H. The echinoderms as aberrant arthropods.

July 20,1921. 20 pp. (Publ. no. 2653.)

. WETMORE, ALEXANDER. A study of the body temperature of

birds. December 30, 1921. 52 pp. (Publ. no. 2658.)

. AtpricH, L. B. The melikeron—an approximately black-body

pyranometer. January 25, 1922. 11 pp. (Publ. no. 2662.)

. GILMoRE, CHARLES W. A new sauropod dinosaur from the Ojo

Alamo formation of New Mexico. January 31, 1922. 9 pp.,
2pls. (Publ. no. 2663.)

. Explorations and field-work of the Smithsonian Institution in

1921. May 26, 1922. 128 pp., 132 figs. (Publ. no. 2669.)

(v)

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 72, NUMBER 1

EXPLORATIONS AND FIELD-WORK OF THE
SMITHSONIAN INSTITUTION
IN 1919

(PUBLICATION 2581)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION

1920

The Lord Baltimore Press

BALTIMORE, MD., U. 8. 2.

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CONTENTS
TLARUEETFCOKGISKCLEIONIS ey rh eo es Rte Un it sen tae Mt Ree nee Soni Wii EM) ee ea I

Geolocicalaxploration. im the Canadtam Iockiesey.s-20--0.0-+40ee oes.

Geological and Paleontological Explorations and Researches.......... 16
Expedition to South America in the Interests of the Astrophysical
OD Seiavialt Ot vais teers en mets AS ceewas Mid sts lv PIOUS Orn PCS ore toes: 24
ENS igetll aTMlsexsT © GNTTOME py. vace tes voversl vets. ste ck) shes swt, Stats) ce aun Gs atlas sano occas 28
Wine. ConlltinesGearmaer Cornero, 1 sxbhiotetacooeeasodauopousucocohouscceeess 33
JE SSPOMANCOMS -abols Seuainoy. IDYosni poker icra ais comin woe ntioa ae ean on cone c came eee an Rul
Botanical Exploration in Glacier National Park, Montana.............. 30
Explorations and Cerion Studies on the Florida Keys:..--.-.2..-......- 41
Breld Wonk on! the Misa Verde National Park, Colorado...............- 47
MiEGheOlOcGal we xCavAalOns IeATIZONaps saa .e sess yates de a. oe ae 64
A\reneoloencall Imngesineeiarorns sh (When ebael UNietzonel, 656. Gahaeencouueouse 66
Field Work on the Iroquois of New York and Canada.................... 69
Osaveminibalmiitess © allomaneyemecee ne ocee: ee SOUR HOON she same 71
_ Ethnological Studies in Oklahoma, New Mexico, and California........ in
MITEKE Oil ne Jeanne, OlbunGiae Saee od ono konk onoae aa eadecooasedd meses 7s
Maternal Culture of the Chippewa of Canada.....<:.......% 2.0.56 -. ee atom cs)

EXPLORATIONS AND FIELD-WORK OF THE SMITH-
SONTAN INST ITULION IN 1919

INTRODUCTION
An important phase of the Institution’s work, in carrying out one

‘

of its fundamental purposes, the “increase of knowledge,” is the
scientific exploration by members of the staff of regions in this
country and abroad previously imperfectly known to science. The
more important of these field researches carried on during 1919 are
here described briefly, for the most part by the explorers themselves.
Naturally the great war held in abeyance a number of proposed
expeditions, some of which it has since been possible to send out on

the resumption of a peace status.

GEOLOGICAL EXPLORATION IN THE CANADIAN ROCKIES

Geological explorations were continued in the Canadian Rocky
Mountains during the field season of 1919 with two objects in view:
(1) The discovery of an unmetamorphosed, undisturbed section of
the Upper Cambrian formations north of the Canadian Pacific
Railway ; and (2) the collection of fossils to determine the various
formations and to correlate them with the Upper Cambrian forma-
tions elsewhere.

The party going from Washington consisted of Dr. and
Mrs. Charles D. Walcott and Arthur Brown, who has accompanied
them for a number of years.

Early in July an attempt was made to reach Ghost River northeast
of Banff, Alberta, but this failed owing to the extensive outbreak of
forest fires in that region.

In August the party proceeded north from Lake Louise over Bow
Pass down the Mistaya Creek to the Saskatchewan River, and thence
up to the head-waters of the Middle Fork in the area about Glacier
Lake, where a wonderfully well preserved Upper Cambrian series
of rocks was found that had been cut across in pre-glacial time by a
deep east-and-west canyon valley, at the head of which were two
beautiful glaciers, which are illustrated by the panoramic view figure I
(Frontispiece), and more in detail by figures 5, 7, 8, 9, and ro.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 72, No. 1

‘ott, 1919

to

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

The character of the mountains about the head-waters of the
Middle Fork of the Saskatchewan is illustrated by figure 2, where a
glacial stream flows out through a deeply eroded valley with high
ridges and peaks rising in steep slopes and cliffs. The position of
the camp in Glacier Lake canyon is shown in figure 3, and the outlook
to the west toward the glacier from the camp by figure 5, and to the

Fic. 2—View looking up the Middle Fork (Howse River) of the Sas-
katchewan River to Howse Pass (5,000’) on the Continental Divide.

In the distance beyond Howse Pass the peaks of the Van Horne Range
and Mount Vaux of the Ottertail Range, and on the right and above the
Pass Mount Conway, and to the extreme right the eastern ridge of Mount
Outram.

Locality—View taken from the upper slope of Survey Peak above
Glacier Lake, about 48 miles (76.8 km.) northwest of Lake Louise station
on the Canadian Pacific Railroad, Alberta, Canada.

Photograph by Mr. and Mrs. C. D. Walcott, 1910.

east by figure 4, where the dark massive bulk of Mount Murchison
rises in cliffs above the canyon of the Mistaya River along which the
trail from Bow Pass descends.

The measured geological section begins at the foot of the ridge
at the extreme left of figure 1, and was measured in the cliffs and
slopes, and thus carried to the side of the Mons glacier shown in

NOG I SMITHSONIAN EXPLORATIONS, IQIQ 3

the central portion of figure 1, the upper limit of the section being
above the narrow vertical F-shaped snow bank directly over the
glacier. The evenly bedded rocks sloped at an angle of from 10 to
20 degrees, westward, which accounts for the great thickness mea-
sured, although the elevation above the canyon bed was not more
than 3,500 feet.

Fie. 3.—Walcott camp located in the brush beside Glacier River, 1 mile
(1.6 km.) below the foot of Southeast Lyell Gacier, which 1s shown more
distinctly in figure 5. Photograph by Mrs. C. D. Walcott, 1910.

The glaciers—The Southeast Lyell Glacier is beautifully shown
in figure 1, and also in figure 5, and more in detail by figure 6, where
it is cascading over a high cliff. Figure 1 is a profile view of Lyell
Glacier from the Continental Divide on the right to where it abuts
against the low cliff on the left. In figure 6 the foot of the glacier
is shown, along with large amounts of débris forming the terminal
moraine, also the dark mass of broken rock and débris carried on the
back of the glacier, which is shown in figures 1 and 6.

A portion of the great snow field from which both Lyell and
Mons glaciers flow is shown on the slope of Mons Peak on the left
side of figure 5. This snow field extends back of Division Mountain,

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NOZ it SMITHSONIAN EXPLORATIONS, I919Q Ti

dividing Mons and Lyell glaciers, and continues along the Conti-
nental Divide for many miles, contributing to glaciers both on the
Pacific and Eastern sides of the Rockies.

Fic. 7.—View of the ice fall of Southeast Lyell Glacier, taken from the
surface of the glacier below the fall. The locality is the same as for
figure 6. Photograph by Mr. and Mrs. C. D. Walcott, 1910.

Mons Glacier is named from Mons Peak, shown in figures 5 and
8. It flows over the high cliffs (see fig. 8), and creeping down the

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

slope, spreads out in a fan-like form toward its lower end. Seen in
profile, this portion of the glacier resembles a great animal sliding
down the sides of the broad canyon. The foot of Mons Glacier with
the stream flowing from beneath it is illustrated by figure 11.

Fic. 8—View of Mons Peak (10,114’) and the upper snow fields of
Mons Glacier, also the cascade of the glacier over the cliffs below. The
summit of the peak is more than 2 miles (3.2 km.) back of the glacier,
the intervening space being occupied by-a great snow field from which
the glacier flows. The locality is about the same as for figures 2 and 5.
Photograph by Mr. and Mrs. C. D. Walcott, 1919.

These views (figs. 5, 8, 9, 10 and 11) illustrate a glacier from its
néve at the base of Mons Peak (fig. 8) to its foot (fig. 11), where
the retreating ice is making its stand against the force of the summer
sun and wind.

Geological section.—The geological section, which is beautifully
exposed in the mountain ridge of figure I, is of such interest that it
is included in this brief account of the exploration in the vicinity of

9

EXPLORATIONS,

SMITHSONIAN

1919

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INTO i SMITHSONIAN EXPLORATIONS, I9QIQ Ed

Fic. 11.—Foot of Mons Glacier from canyon down through which the glacier
formerly extended to unite with the south end of Northeast Lyell Glacier.
The locality is about the same as for figure 6. Photograph by Mr. and Mrs.
C. D. Walcott, roto.

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MISCELLANEOUS COLLECTIONS

SMITHSONIAN

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INOS = SMITELSONIAN EXPLORATIONS, I9Q!Q I

Fic. 13.—Leaving Glacier Lake camp with the camp outfit and specimens
on the pack horses, ready for the four days’ trip to the railway. Photo-
graph by Mrs. C. D. Walcott, roto.

Fic. 14—Mount Ball (10,825’) on the Continental Divide from the

southwest slope of the Sawback Range, looking across Bow Valley.
Locality About 17 miles (27.2 km.) west of Banff, Alberta, Canada.
Photograph by Mr. and Mrs. C. D. Walcott, toro.

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.. 72

Glacier Lake. The rocks exposed in the highest cliffs of Mount
Forbes and Mons Peak belong to the great Carboniferous system
of rocks of this region. Below this series occur the Devonian rocks,
the snow-capped cliffs shown in figure 1 above Mons Glacier, form-
ing a belt 1,000 feet or more in thickness, and below these are the
strata of the Sarbach formation of the Ordovician system, and

es Se
Ae le

Fic. 15.—Mountain sheep in game sanctuary, Rocky Mountains Park,
Alberta, Canada. Photograph by Mr. and Mrs. C. D. Walcott, 1919.

beneath the Sarbach the five formations assigned to the Upper Cam-
brian series. Attention should be called here to Mount Murchison,
figure 4, where there is almost the same series as that exposed in the
cliffs on the long mountain slopes of figure 1. To the left of Mount
Murchison in figure 4 there is a low ridge formed of strata of
Middle Cambrian age which passes beneath the rocks of Mount
Murchison.

Fic. 16.—Cirkut camera in use on the south slope of the Sawback Range,
looking across Bow Valley, Alberta, Canada. Photograph by Mrs. C. D.
Walcott, 1910.

GLACIER LAKE, SECTION, ALBERTA

Thick- Thick:
Formation |No. Stratum ness || Formation |No. Stratum | ness
feet || | feet
| i}
S| 1 | Thin-bedded 700 || 1a |Compact gray | 325
“3| sarbach. gray lime- (est.) limestone. |
> stone. tt cette 0 a ot NN liar wielllolatatel siete ates store’ jeleivte [eteretet seve .
3 (1120 feet) Befivellveietepcve te ctelerelstelsis 6 Aes ayaa eet | Sullivan. 1b | Shale withinter-| 975
= (341-3 meters). | 2 Argillaceous 420 «|| | bedded layers
e) | shale. || | (1440 feet) of limestone.
| (Gara mehr) |p cme ectciew oleieisieteiele tiv'eleal|=/=ilel =i 3
| | ra | Calcareous | 235 "Z| 1d stone and
shale and lime- \|-2 J shales.
stone. VE IE
SQnH| (Goons aosodacosKde Saisieisiers sill) | |
Mons. 1b| Massive bedded) 740 | 1a | Laminated 520
i gray limestone. si limestones.
= (1480 feet) satire lteter siete Sate teeel sae geevallia! ArctOmlys: |a«cclecccceanscvcwces ue slelelatst=reis
5 (467.2 meters) | 1c | Limestone and| 320 |/D | 1b | Purple, green, | 866
&) shale. (1386 feet) and gray shale
cs Sekar | xclotsl acs Be aisjelelereiaiatetel | stoisietere’erai||nn|| (42aea meters). with layers of
o 1d | Oolitic lime- 185 | laminated
3 stone. | limestone.
a
a | BEE
2) G q
ta | Massive bedded) 1,270 ery oe serve |r |Gray an 220+
| Lyell gray lime- el Murchison. bluish-black
| . stones. Canteen) limestone,
(1700 feet) [ees eee eee ee pseezene(sieie)||| | (67 meters). | - :
1b Light gray. | 430 || | Base concealed.
RG3S-GIREBENS) 15 I thick-bedded I) | |
| 1d | limestone. |

Total thickness Cambrian strata......... Eecloraleie Terate orale ale saie! vias aivinessaiwiainie este eiee:t/seleetelals 6,226

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

On returning, a camp was made in the Bow Valley below Lake
Louise at the foot of the Sawback Range, where a brief examination
was made of the up-turned Carboniferous, Devonian, and Cambrian
formations, and fossils were collected, many of which are identical
with those found in the Glacier Lake section. One of the views
across Bow Valley is of Mount Ball (fig. 14), which is one of the
massive peaks on the Continental Divide where the Middle Cambrian
rocks have a great development.

The preliminary study of the fossils in the several formations cor-
relates them with the Upper Cambrian formations of Wisconsin and
Minnesota and the Upper Cambrian section in southern Idaho, and
to amore limited extent with that of the central belt of Pennsylvania.

GEOLOGICAL AND PALEONTOLOGICAL EXPLORATIONS AND
; RESEARCHES

Field-work in the Department of Geology has been considerably
restricted by prevailing conditions during the season of 1919.
Research work has, however, continued on the collections accumu-
lated either in years past or obtained through gift or purchase in
more recent times.

One of the most interesting acquisitions to the geological col-
lections during the year 1919, was some 15 kilograms of the meteoric
stone which fell at Cumberland Falls, Kentucky, in April of this
year. The stone, which proved to be a coarse breccia of enstatite
fragments and a dark chondritic stone, has been studied by Dr. Mer-
rill, and a paper giving his results is now in press.

Owing to the fact that the division of Mineralogy has been without
a head for two years, a large amount of work upon the collections has
been necessary and hence only a limited amount of field and research
work was possible. On his own initiative, two field trips were under-
taken by Assistant Curator Foshag, one to the mica mines about
Amelia, Virginia, where a considerable amount of study material was
collected. This included a large number of specimens of the rare
mineral microlite and an exhibition specimen of manganotantalite,
the latter species not before represented in the Museum’s exhibition
series. The old iron mine at Brewster, N. Y., was also visited and
some material for study collected. The work of the division in the
Museum’s laboratory consisted in an investigation of the hydro-
talcite minerals, resulting in the establishment of the true chemical
nature of this group. Shorter researches on miscellaneous minerals

NO. I SMITHSONIAN EXPLORATIONS, I919 7

were also carried on. Work is now under way on the study of the
mineralogy of serpentine and its relation to the chlorite group, and
also the correlation of the chemical composition and the optical
properties of rhodonite, the latter in collaboration with Dr. E. S.
Larsen of the U. S. Geological Survey.

In October various localities in Connecticut were visited by
Assistant Curator Earl V. Shannon, including the vicinity of Long
Hill in Trumbull, where tungsten has been mined in years past.
Extensive collections comprising much interesting material were

Fic. 17.—Transporting fossil specimens in Southeastern Indiana.
Photograph by Bassler.

obtained which will be studied and described. The trap quarries of
Meriden, Conn., were also visited and extensive collections of the
zeolites and other secondary minerals secured. The pegmatite locali-
ties of Collin’s Hill in Portland ; the workings of the old cobalt mine
and the old lithia mine in Chatham, and a number of feldspar quar-
ries in this region were visited and collections of the representative
minerals secured. Much of the material collected as above has been
carefully examined and descriptions will appear in forthcoming num-
bers of the Proceedings.

3

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Explorations for paleontological material were limited during
1919 to two short field trips by Dr. R. S. Bassler, Curator of Paleon-
tology, who continued the work of former years in securing certain
large showy specimens of fossils and rocks required for the exhibi-
tion series. Dr. Bassler spent a portion of June in southeastern
Indiana, first proceeding to the locality where at the end of the field
season of 1918 he had cached for safe keeping, because of inability
to secure help in getting them to a freight station, several large
exhibition slabs crowded with brachiopod shells. These slabs were

Fic. 18.—Fossiliferous strata of the Richmond formation in south-
eastern Indiana. The slab indicated is now on exhibition at the National
Museum. Photograph by Bassler.

found undisturbed, but transportation conditions proved equally bad
as in the summer before and it became necessary to employ the
primitive method shown in figure 17. By the use of burlap covering
and an abundant supply of weeds for padding, each slab was finally
slid along the rails for a considerable distance to the nearest station.

The same area in Indiana, namely, the vicinity of Weisburg where
the early Silurian rocks are well exposed, was then explored tor fur-
ther desirable exhibition specimens. Water worn slabs crowded
with animal and seaweed remains are abundant in all the creeks of

NOS 4 SMITHSONIAN EXPLORATIONS, IQIQ IQ

this richly fossiliferous region, but large rock specimens with their
surfaces unweathered are comparatively rare. Fortunately for the
present purposes, heavy freshets in the spring of 1919 had uncovered
the richly fossiliferous layers along the creek shown in figure 18, and
here several additional well-preserved exhibition slabs were secured
and transported to the freight station by the rail route mentioned be-
fore. One of these specimens, a slab several feet in length and width,
is worthy of special mention, as its surface 1s crowded with impres-
sions of the branching fossil seaweed Buthotrephis, and with excel-
lent examples of the dumbbell seaweed Arthraria. The discovery
of this specimen was most fortunate as a large slab containing an
assemblage of these ancient plant remains has long been needed for
the exhibition series of fossil plants.

In October, 1919, Dr. Bassler was detailed to proceed to Dayton,
Ohio, in order to prepare for shipment to the Museum the largest
entire American trilobite so far discovered. This unique specimen
(fig. 19) was discovered in the Richmond formation in the excava-
tions for the Huffman Conservancy Dam 6 miles east of Dayton,
which forms a part of the greatest engineering project ever under-
taken for controlling stream flow, with the exception of the Assouan
Dam along the Nile. Following the destructive floods in the Miami
valley in 1913, the Miami Conservancy District was organized to
prevent a repetition of this disaster through the control of the
Miami River and its tributaries by dams extending entirely across
their valleys. This operation involving an expense of over fifteen
million dollars has been under way for three years, and will require
three years more for its completion. The view (fig. 20) showing the
excavations in which the trilobite was found illustrates only the
beginning of one of these dams. This particular dam when com-
pleted, will extend a distance of a mile entirely across the river valley.
Normally the water will flow in its usual channel, but in flood times
it will be retained and allowed to escape gradually.

The trilobite which was found lying on its back in a hard clay bed
in the central part of the excavation was unearthed by the pick of a
workman, who believed it to be a petrified turtle. Mr. Arthur E.
Morgan, Chief Engineer of the Miami Conservancy District, recog-
nized the true nature and scientific value of the so-called turtle and
presented it to the Smithsonian Institution, where it now forms a
most instructive and unique exhibit in the hall of invertebrate
paleontology. The value of the specimen is further increased by the
fact that it has become the type of the new species /sotelus brachy-
cephalus described by Dr. August F. Foerste of Dayton, Ohio.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

During the summer of 1919, Mr. Frank Springer resumed his
field researches upon the fossil echinoderms of the Ohio Valley,
which had been interrupted by the war, with the special object of
securing additional material from the remarkable crinoidal fauna
of the Laurel formation of the Niagaran for use in his monograph
on the Silurian crinoids. The principal work was done at St. Paul,

Fic. 19.—The type specimen of /sotelus brachycephalus Foerste,
the largest known entire American trilobite. About % natural
size. Photograph by Bassler.

in Shelby County, Indiana, where Mr. Springer’s assistant, Dr.
Herrick E. Wilson, continued the systematic collecting and
detailed study of the strata begun several years ago. Collecting
in the beds at this locality is laborious and difficult, and specimens
are not abundant, but they exhibit in some respects a strong paral-
lelism with those of the Swedish and English Silurian, which renders

NO. I SMITHSONIAN EXPLORATIONS, I9QIQ 21

their comparative study one of much interest. Important new mate-
rial was obtained which is added to the collection of echinoderms
deposited in the National Museum. Mr. Springer’s monograph of
the Crinoidea Flexibilia, now being published by the Smithsonian
Institution, has been printed and only awaits binding to be ready
for distribution. It will be in two quarto volumes, one of them con-
taining 79 plates.

The Section of Vertebrate Paleontology has been unable to under-
take any field explorations during the past year, and the time of the
staff has been largely employed in preparing and mounting for
exhibition material otherwise obtained. |

Fic. 20——View of small portion of Huffman Conservancy dam near _
Dayton, Ohio, showing excavation in which the largest trilobite was found.
Photograph by courtesy of Arthur E. Morgan, chief engineer.

Through the acquisition from the veteran collector of fossils,
Mr. Charles H. Sternberg, of an excellent skeleton of the large
swimming lizard 7'ylosaurus dyspelor and a skull of the primitive
horned dinosaur Monoclonius, an interesting addition to the exhibi-
tion collection was made. Figure 21 shows the 7ylosaurus skeleton
in process of being mounted for exhibition by Mr. N. H. Boss. It
will form a panel, in half relief, and will occupy the wall space in
the northeastern part of the main exhibition hall of fossil vertebrates.
The diving pose given the skeleton was largely determined by the
position of the articulated tail as found in the ground.

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Tylosaurus was a long, slender-bodied reptile some 25 feet in
length, having the limbs modified into short swimming paddles, with
a long, powerful, compressed tail. It was predatory in habits, living
on fishes and probably also on the smaller individuals of its own kind.
One of the unique features of the skeletal structure 1s the presence
near the middle of the lower jaw of a joint which permits it to bow
outward. This feature in connection with the loose articulation at
the extremities allowed the jaws to expand and thus enabled the

Fic. 21—The swimming reptile 7ylosaurus dyspelor in process of
preparation for exhibition. Photograph by Bassler.

animal to swallow large objects. In life the body was covered by
small horny scales. A study of this specimen is being made by
Mr. C. W. Gilmore, and the results of his investigations will be pub-
lished in the Proceedings of the U. S. National Museum.

In figure 22 is shown another of the recently mounted skeletons—
that of the large extinct Rhinoceros-like mammal from the Tertiary
(Oligocene) deposits of western Nebraska, named by Osborn Bron-
totherium hatcheri. The bones of this skeleton were found em-

NOr I SMITHSONIAN EXPLORATIONS, IQT9 23

bedded in a fine sandstone. An idea of the amount of painstaking
work involved in the collection, preparation and mounting of such a
large fossil skeleton may be gained by the fact that after the bones
were collected and freed from the enclosing sandstone, 258 working
days were spent in restoring the missing parts, posing the skeleton
and making and fitting the supporting iron work.

The National Museum has now the distinction of having the larg-
est existing collection of Titanothere remains. The specimens were

Fic. 22—The Rhinoceros-like mammal Brontotherium hatcheri mounted
in the laboratory of vertebrate paleontology. Photograph by Bassler.

brought together by the late J. B. Hatcher, working under the direc-
tion of Prof. O. C. Marsh, at that time United States Paleontologist,
and it is peculiarly fitting that the only skeleton complete enough for
mounting should pertain to the species named 1n his honor.

Among other important investigations in the mammalian division
of the year is a study by Mr. Gidley of the Peccaries of the Cumber-
land Cave deposit. This includes the description of three new
species, the redefinition of the Pleistocene species hitherto described.

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS Wks W/Z

and the partial revision of the entire peccary family (Tayassuidae),
the latter involving redefinitions of the two living genera of this
group, and of the two suilline families Suidae and Tayassuidae. An
important outcome of this preliminary investigation has been to
emphasize the fact that the entire group, and especially the Pleisto-
cene species of American suillines, is in need of thorough revision.

EXPEDITION TO SOUTH AMERICA IN THE INTERESTS OF THE
ASLROPHYVSICAE OBSERV ALOR

Dr. and Mrs. C. G. Abbot sailed from New York on May 2, by
way of the Panama Canal to Antofagasta, Chile. The expedition
had three objects. First, to observe the total eclipse of the sun of
May 29 at La Paz, Bolivia; second, to enable Dr. Abbot to confer
with the officials of the Argentine Weather Bureau in relation to the
use of the observations of the solar radiation for the purpose of
forecasting weather conditions; third, in order that a visit might be
paid at the observing station of the Smithsonian Institution at
Calama, Chile, which is maintained by the Hodgkins Fund under the
direction of Mr. A. F. Moore and assisted by Mr. L. H. Abbot.

Landing at Antofagasta, the journey was continued by the English
railroad up into Bolivia over that desert which Darwin describes in
his “ Voyage of the Beagle.”’ Although the travelers had visited the
Sahara Desert in southern Algeria, and the deserts of the southwest
of the United States, there was still reserved for them a stronger
impression of a void wilderness in the Nitrate Desert of Chile.
Neither bird, beast, insect nor crawling thing, nor any vegetation
could be seen as far as the eye could reach.

Stopping a day at the observing station at Calama, in order to
repack the apparatus required for the eclipse expedition, and joined
by Mr. A. F. Moore, director of the observing station, they went on
to La Paz. The plateau of Bolivia is eminently the country of
mirage. The railroad appeared to rise out of a lake and to run inte
a lake at no distant point beyond, and all of the mountains appeared
to be islands rising out of the lake. The desolation, while not equal
to that of the Nitrate Desert of Chile, was yet very marked. Near
La Paz the country becomes cultivated with fields of grain and
vegetables, and villages of people are passed by on either side. The
mountains take on a new grandeur, especially the great mountain
Illimani, which rises to a height of 22,000 feet or more.

However one may have been impressed with the grandeur of the
mountains, he is unprepared for the view of the great canyon in

NO. I SMITHSONIAN EXPLORATIONS, I9QIQ 25

which La Paz lies. In most respects it is comparable with the Grand
Canyon in Arizona, but with several added charms. First of all, on
the farther side lies the great chain of the Andes, to which [llimani
belongs, snow-covered although so far within the tropics, while on
the nearer slope of the canyon all is green with vegetation, trickling

Fic. 23.—Total solar eclipse of May 29, 1919.

brooks running down, and the beautiful city lies upon its cluster of
hills all surrounded by the colors and broken features which might
result by combining the Grand Canyon of the Colorado with the
Garden of the Gods near Colorado Springs.

By the kind assistance of the manager of the English railroad the
eclipse station was located at El Alto, situated on the rim of the

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

canyon at about 14,000 feet elevation. There the apparatus was set
up, and on the day of the eclipse very satisfactory observations were
made, both photographs of the phenomenon and measurements of
the brightness of the sky and the nocturnal radiation. The accom-
panying photograph shows in but a feeble way how splendid was the
eclipse phenomenon on this occasion. ‘The corona extended at least
two diameters of the sun in almost all directions, with a great pro-
fusion of fine streamers. Underneath the sun, that is to say towards
the east, there hung a sickle-shaped solar prominence of hydrogen
and calcium gases, extending fully 150,000 miles outwards from the
sun and over 300,000 miles long, which cast its crimson glory over
all. The background against which this splendid phenomenon was
seen was a range of mountains, perhaps 50 miles distant, which raised
their snow-covered heads fully 20,000 feet in altitude.

From La Paz, Messrs. Abbot and Moore proceeded at once to
La Quiaca, Argentina, where they inspected the meteorological
observatory and conferred with Messrs. Wiggin and Clayton, Chief
and Chief Forecaster, respectively, of the Argentine Meteorological
Service. Under Mr. Clayton’s direction a system of forecasting by
the aid of daily telegraphic reports of the intensity of the radiation
of the sun as observed at Calama, Chile, has been worked out with
encouraging success.

Both Mr. Wiggin and Mr. Clayton were firmly convinced of the
great value of solar radiation observations for forecasting. Arrange-
ments were tentatively entered into for the transfer of the Smith-
sonian observing station at Calama to the Argentine Government,
to be located at La Quiaca thereafter. At the present time, however,
these arrangements have not yet been completed by the higher
officials of the Government of Argentina.

From La Quiaca the travelers returned to the observing station at
Calama, Chile, where Dr. and Mrs. Abbot remained about a month
before returning to the United States, during which time Dr. Abbot
worked over the results obtained with a view to discovering some
means of measuring the intensity of the solar radiation without the
long and tedious process of observing and computing which has
hitherto been necessary. In this he was so fortunate as to discover,
with the aid of Mr. Moore, a method by means of which suitable
observations taken within a period of 10 minutes may be reduced
within a period of 2 or 3 hours. Thus the intensity of the solar
radiation outside the atmosphere may be determined with greater
accuracy than by the former method which required about 3 hours
of observing and about 15 hours of computing.

SMITHSONIAN EXPLORATIONS, IQI1Q

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28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Mt. Wilson observations.—As usual, the station at Mt. Wilson was
occupied from May until October by Mr. L. B. Aldrich, who carried
on there the usual measurements of the “ solar constant of radiation,”
and some others intended to determine the intensity of the nocturnal
radiation and the intensity of skylight.

AUSTRALIAN EXPEDITION
The zoological work by Mr. H. C. Raven in Borneo and Celebes,
which has been made possible through the generosity of Dr. William
L. Abbott, came to an end in 1918, as explained in the last Explora-

Fic. 28—A pair of adult duck-
bills, the male (at left) showing
the spurs on hind feet.

tion Report (p. 35). At the close of the war Mr. Raven did not wish
to return immediately to the field. Doctor Abbott therefore arranged
to send Mr. Charles M. Hoy to Australia for the purpose of collect-
ing vertebrates, especially those which are in danger of extermina-
tion. From the point of view of the national collection of mammals
there is probably no field-work of similar scope that could approach
this in importance. The fact has long been recognized, but the
means for putting such a project into execution have hitherto been

NO. I SMITHSONIAN EXPLORATIONS, IQIQ 29

lacking. The remarkable Australian mammal fauna is now repre-
sented in the museum by only about 200 specimens. Very few of
these have been collected in accordance with present-day methods,
and no species is represented by adequate series of adults and young,
of skeletons and material in alcohol. Not only is this true of our
collection, but it is equally true as regards other museums in
America. So many of the Australian mammals are now rapidly
approaching extermination that in future there will be few oppor-
tunities for securing the material needed for a proper representation
of the fauna.

Mr. Hoy spent about two months in Washington assembling his
outfit and preparing for his trip. He left for San Francisco early in

Fic. 29.—Two young duck-bills in their nest.

May and arrived in Sydney about the end of the month. The period
from June to November has been passed at various localities in
New South Wales. While no specimens have yet been received
(December 31, 1919), it is evident that the work has been very suc-
cessful. The following passages from Mr. Hoy’s letters and reports
will give an idea of the conditions under which it has been car-
ried on:

“Travel is rather difficult just now. Each state has its own quarantine
restrictions, on account of the ‘flu’ and it is quite difficult to get from one
to the other. Then too a seaman’s strike is on, making travel by boat very
uncertain. The railroads are run by the state governments and each state has
a different gauge road and different rates.

“T have a great chance to get Platypus (duck-bill). A Mr. Burrell, whose
hobby is Platypus is going after them on the first of October. As he has

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

kindly offered to take me with him and is only interested in the-natural history,
it will be a great chance for me to get not only the adults but also the young
and possibly an egg. Mr. Burrell is after their nesting habits and is taking
men along to dig out the burrows. It is safe to say that there is not another
man who knows as much as he about the duck-bill.

“Nine weeks were spent in the Wandandian region (19 miles S. W. of
Norwra, N. S. W.) with the result of but one hundred and thirty one (131)
mammals, and one hundred and twenty-four (124) birds collected. Among
the mammals ten genera and twelve species are represented in my collection.

“The greatest agent working toward the extermination of the native animals
is the fox, next comes the cattle and sheep men who distribute poison by the

Fic. 30.—An Australian marsupial with parachute
membrane like that of the flying squirrel.

cartload in the effort to reduce the rabbits. This has also caused or helped
to cause the extermination of some of the ground inhabiting birds. Another
great agent is the bush fires which sweep over the country. These are often
lit intentionally in order to clear out the undergrowth and thus increase the
grass.

“ Owing to the great increase in the price of rabbit skins and the consequent
increase in trappers the rabbits are fast ceasing to be a pest, and in some
sections have become quite scarce. ‘The foxes, which are everywhere nu-
merous, after killing off the native mammals are now turning to the rabbits,
which also helps in their reduction. The extermination of the native mam-
mals has apparently gone much farther than is generally thought. Many
species that were plentiful only a few years ago are now almost, if not alto-

NO. I SMITHSONIAN EXPLORATIONS, IQIQ 31

gether, extinct. Diseases have also played a great part in the extermination.
The native bear died in thousands from a disease which produced a great
bony growth on their heads. A mysterious disease also spread through the
ranks of the native cat, Dasyurus viverrinus; the domestic cat also played a
great part in their extermination. Even adult specimens of Dasyurus were
often dragged in by the family cat.

“The only mammals caught in traps were Mus assimilis and Phascologale
flavipes. The others were all shot or snared. As the majority of the mam-
mals taken were nocturnal and arboreal, the headlight proved to be a valuable

Fic. 31—An Australian marsupial resembling a very
large flying squirrel.

and indispensable part of my outfit. The hunting of kangaroos and wallabies
was greatly handicapped through the lack of a rifle.

“ Perameles nasuta has been practically exterminated throughout N. S. W.,
but they are still to be found in Mosman, one of Sydney’s suburbs, so I made
a trip out there and was able to get a fine female with two young in her pouch.
This was trapped inside the Taronga Park Zoo grounds with the kind per-
mission of Mr. A. S. Le Souéf.

“The fact of Perameles nasuta being found at Mosman is probably due to
the isolation of that district from the rest of N. S. W. by the city of Sydney,
thus keeping out the introduced foxes.

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

“The country at Bulliac is a good example of what the cattle men will do
in a few years time in killing off and burning the timber and the consequent
destruction of animal life. Five years ago Bulliac was a virgin forest but
then the railroad was built and now it is devoid of living trees for miles on
each side of the track except for small and scattered patches in the more
inaccessible gullies which form the last resort of the fast disappearing mam-
mals. There are very few game laws in Australia and no one gives any
attention to the ones that are in order. The Bulliac trip has clearly shown
that the edge of the settlements, and even a short way into the wilds, is no
better than the older settled parts and in order to get results one must go
into the wilds. It is the killing and burning of the brush, by the cattle men,
that does the most to kill off the animals, and they are yearly reaching farther
and farther away from the railroads. One thing that was very noticeable was
the great abundance of the introduced rats. They seem to have driven out or
killed off practically all the native rats and I found them everywhere.

Fic. 32—Skinning a kangaroo.

“A trip was made primarily for the eggs and young of the Platypus.
Fifty miles of bank was searched and approximately one hundred burrows
were dug out only six of which contained young. These six burrows furnished
eleven young of which Mr. Harry Burrell, of Sydney, with whom the trip
was made, secured 7 while I secured 4. The intense drought which has
scarcely been broken for over twelve months (it is the worst since 1862)
has advanced the breeding of the animals somewhat so we were too late for
eggs. The Platypus although scarce is not by any means nearing extinction
and if they continue to be protected there is every hope that they
will never become extinct. Young Hydromys were also looked for but the
drought seems to have interfered with their breeding and but two embryos
were obtained. The adults themselves were very scarce and but six specimens
were obtained.

The accompanying photographs show some of the characteristic

Australian mammals obtained by Mr. Hoy.
G. S. MILter, Jr.

INOS LE SMITHSONIAN EXPLORATIONS, I9QIQ 35

THE COLLINS-GARNER CONGO EXPEDITION
The Collins-Garner Congo Expedition returned to this country in
May, after having spent more than two years in the collecting of
natural history material in French Congo. Mr. C. R. Aschemeier,

Fic. 34.

Fics. 33, 34.—Skulls of gorillas collected by Aschemeier in French Congo.
Young adult male at left; adult female at right.

who represented the Smithsonian Institution, brought back with him

the bulk of material collected. Among other things Mr. Aschemeier

collected about 2500 birds and mammals, which added invaluable

material to the museum collection.

4

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

EXPLORATIONS IN SANTO DOMINGO

Dr. W. L. Abbott continued his investigations at the eastern end
of the Island of Haiti, making two visits during the year 1919. His
first visit was to the Samana Bay region and to the mountains in the
vicinity of Sanchez and Constanza, and covered the interval from
February to June. On his second trip (July to October) he stopped
at Sosua, on the north side of the island, where he made a search
for certain birds needed to fill gaps in the material already collected :
thence to the Samana Peninsula, after which he went to the islets
Saona and Catalina, off the southeastern corner of Santo Domingo.
Before returning to Philadelphia, he spent a few days at Lake Fnri-
quillo, where he secured a few water birds of interest.

The material gathered on these two trips was varied in character,
embracing the several groups of vertebrates, as well as mollusks,
insects, and plants, with a plentiful series of archeological objects
from caves in the Samana district. Of birds alone, he obtained 27
study skins, 87 alcoholics and skeletons, and 56 eggs. Four species
of birds in this collection represented forms not hitherto possessed
by the Museum, and three or four other species were not previously
known to occur on the island. Probably the most noteworthy objects
in the bird collection were four eggs of the Dulus dominicus, known
variously as the Sigura, Palm Slave, or Palm Chat, a bird somewhat
larger than an English Sparrow, of sombre colors, but structurally
different enough from other birds to occupy a family by itself. It
““apart-
ment house” nests are sufficiently noticeable to attract the attention

is common and noisy, and lives in colonies, and its enormous

of the most casual person. The discovery of the Palm Chat was
almost coincident with that of America, for Columbus and his men
must have seen these birds and their nests when a landing was made
on this island at the end of 1492. Oviedo, the official historian of
the Indes, early made its acquaintance, and devoted a chapter to it
in his ‘‘ Hystoria natural y general de las Indias yslas,” published
in 1535. The Sigura was thus one of the earliest, and at that time
one of the best-known birds of the new continent. Various authors
of later date have described the bird, its habits and nests, but thus
far, apparently no correct description of the eggs has been given,
although a French writer, in 1851, reported them to be white and
unmarked.

The eggs sent by Dr. Abbott are authentic, and will be described
in detail elsewhere, but the illustration here given (fig. 35) will
convey a general idea of their appearance, and it may be remarked

NO. I SMITHSONIAN EXPLORATIONS, IQIQ

ios)
oni

that the eggs are rather large for the size of the bird, measuring,
roughly, about one inch by three-quarters of an inch in diameter.
The description of the nest, as furnished to Dr. Abbott by the man
who made the actual examination, indicates a radical departure from
other known types of birds’ nests, but a careful study of the interior
of the mass by a competent observer will be required before the
strange arrangement of the individual nests can be considered as

accurately known. Dr. Abbott writes: “ The communal nest was

Fic. 35—Eges of the Palm Chat (Dulus dominicus).

built in a small royal palm, about 25 feet from the ground. The nest
was about 44 feet in diameter, and about the same in height. Built
of a loose mass of sticks from I to 2 feet long, about the thickness
of bone knitting needles. There appeared to be but one entrance,
from above.. A local man (John King) climbed the tree and gave
me the description of the iternal construction of the nest. The
entrance passage was narrow and crooked and led into a large
central chamber about the size of a small bucket. The nests were
placed around this on shelves of softer materials (fine bark, etc.).

All four eggs were in one nest. The other five nests were still empty.

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

‘Tt is difficult to get into these nests, the huge smooth palm trunk
(2 feet in diameter) must be climbed, and then it is difficult to crawl
over the huge mass of sticks which lie interlaced amongst the crown
of leaf stalks.

“Most of the nests are in far larger palms than this one, and the
nests are frequently much larger in size, and probably contain much
larger colonies.”

Fic. 36.—Dense forest of giant cedar (Thuya plicata)
near Lake McDonald, Glacier National Park.

BOTANICAL EXPLORATION IN GLACIER NATIONAL PARK
MONTANA

Mr. Paul C. Standley, assistant curator in the division of plants,
spent the summer of 1919 in Glacier National Park, Montana, under
the authority of the National Park Service, for the purpose of study-

No. I SMITHSONIAN EXPLORATIONS, IQIQ Sy7.

ing the vegetation of the region. The information thus obtained is
to serve as the basis of a popular account of the plants to be published
by the National Park Service, and it is expected that a complete and
more technical account of the flora will be published by the National
Museum. All the portions of the Park usually seen by tourists were
explored, and 4,000 specimens of flowering plants and ferns were
obtained, representing over 900 species.

Aside from the scenic features of Glacier Park, one of its chief
attractions is found in the plant life. Especially striking is the-dis-
play of flowers above or near timber line, where the meadows are
solid masses of brilliant color, formed by Indian paint-brushes,

Fic. 37—Bearerass, Glacier National Park.

monkey-flowers, fleabane, asters, harebells, heather, larkspur, gen-
tians, fireweed, columbine, and a host of other plants. The most
characteristic plant, perhaps, is the beargrass (Xerophyllum tenax,
fig. 37), which occurs abundantly at middle altitudes and in alpine
meadows. It is usually about 3 feet high and bears enormous plume-
like racemes of creamy-white flowers. The narrow, tough leaves
were used by the northwestern Indians in making baskets, hence the
name of Indian basket-grass which is sometimes applied to the plant.

The Continental Divide, which traverses the Park, has a marked
influence upon plant distribution. On the east slope, whose drainage
is partly into the Missouri River and partly into Hudson Bay, the
flora is of the Rocky Mountain type, like that of Wyoming and Colo-

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 38.—Sperry Glacier, Glacier National Park. There are over
60 glaciers in the Park. Photograph by R. E. Marble.

Fic. 39.—St. Mary Lake, Glacier National Park. Photograph by
Fred H. Kiser.

NO: I SMITHSONIAN EXPLORATIONS, I9QIQ 39

rado ; while on the west slope, whose streams drain into the Colum-
bia River, the flora is more obviously related to that of the Pacific
Coast. The forests about Lake McDonald are very dense and are

Fic. 40—Gunsight Lake from Gunsight Pass, Glacier National Park.
Photograph by R. E. Marble.

composed of unusually large trees. Although not nearly so exten-
sive, they are much like those of the humid regions of Oregon and

Washington.

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 41.—Trail to Swiftcurrent Pass, Glacier National Park; Swift-
current Glacier in the distance. Photograph by Scenic America Company,
Portland, Oregon.

NO. I SMITHSONIAN EXPLORATIONS, I9QIQ 4!

In the vegetation there are represented four of the life zones
recognized by biologists. The Transition Zone is indicated on the
west slope by small areas of yellow pine timber, and east of the Park
are the prairies of the Blackfoot Indian Reservation, which extend
also within the Park boundaries along the stream valleys. The plants
here are chiefly herbs, with a few shrubs, and they belong mostly to
species which have a wide distribution over the Great Plains. by
far the largest portion of the Park is covered with the characteristic
vegetation of the Canadian Zone, which is the heavily forested area.
Above the Canadian Zone, around timber line (6,000 to 7,500 feet),
lies a narrow belt belonging to the Hudsonian Zone. The trees here
are mostly low and stunted, and their branches frequently lie pros-
trate upon the ground. Above this belt, and occupying the highest,
exposed slopes, lies the Arctic-Alpine Zone, whose vegetation is
composed chiefly of small herbaceous plants, with a few dwarfed
shrubs, mostly willows. Many of the species of this zone are
widely distributed in alpine or arctic regions of North America, and
some of them occur also in similar situations in Europe and Asia.

EXPLORATIONS AND CERION STUDIES ON THE FLORIDA KEYS

Dr. Paul Bartsch, curator of marine invertebrates, U. S. National
Museum, joined Dr. Alfred G. Mayer, the Director of the Tortugas
Marine Biological Laboratory, of the Carnegie Institution, in New
York on December 28, when they sailed south for Key West to
make an examination of the Cerion colonies discussed in previous
Smithsonian exploration pamphlets. The breeding experiments
presented an entirely new phase, in the crossing, on one of the keys,
of the native species, Cerion incanum (Binney) with one of the
introduced Bahama species, Cerion viaregis Bartsch. It was this
discovery that made it desirable to start an entirely new set of experi-
ments. Furthermore, the anatomical differences discovered in the
dissections of Cerions also made it desirable to gain material from all
the colonies now existing on the Florida Keys, in order that these
might be subjected to anatomization, to determine if Certon incanum
is really one species, or a complex, shell characters alone being
insufficient to determine this point. It was for this double reason
that a return was made to Florida on May 2, and an exploration of
the keys adjacent to Miami at once undertaken.

On the 3d Capt. Tracy and Dr. Bartsch started in the “ Darwin,”
a shallow draught launch of the Carnegie Institution, for an explora-
tion of the shores of the mainland of the lower peninsula and the

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72
outside keys. This was rendered comparatively easy, since they were
fortunate enough to find the mosquito pests practically absent.
Between May 3 and 6 they examined every sandspit on the mainland
along Key Biscayne Bay, Card Sound, Little Card Sound and Barnes
Sound. They then turned the southern portion of Barnes Sound
and came up along the outer keys, examining them in turn. They
crossed Key Largo at several places and walked long stretches in its
interior.

No Cerions were found on the mainland between Miami and the
point where the railroad enters the keys, but two colonies were dis-

Fic. 42.—Nest of swallow-tailed kite (Elanoides forficatus forficatus)
between Flamingo and Coot Bay, Cape Sable, Florida.

covered on keys a very short distance from the mainland in Barnes
Sound. One of these is situated on a sandspit covered with grass and
shrubbery on Middle Key, while the other one was found in a clear-
ing on the southeastern point of Main Key. No Cerion colonies
were discovered on the keys forming the eastern perimeter of Barnes
Sound, Little Card Sound, Card Sound and Key Biscayne Sound,
excepting Porgee Key, where a goodly quantity were obtained, some
of which have been dissected and published upon in the bulletin on
“ Experiments in the Breeding of Cerions”’ recently issued by the
Carnegie Institution. In addition to Cerions, collections of all the

other species of land mollusks available were made wherever found at

NO. I SMITHSONIAN EXPLORATIONS, I9IQ 43

all the places visited, especial stress being laid upon the securing of
series of the most beautiful of our American landshells, the tree snail
Liguus, anatomical material of all of which was preserved in order
that this group may also be subjected to a close scrutiny in an ana-
tomical way. What is said for Liguus also applies to the equally
large, though less brilliantly colored Oxystyla.

On the 7th the ** Anton Dohrn ” left Miami for Indian Key, spend-
ing the night off Rodriguez Key, and reaching Indian Key early on
the following day. By to o’clock Capt. Tracy, Mclvar, the Assistant
Engineer, and Dr. Bartsch were off in the “ Darwin” with the skiff

Fic. 43.—Characteristic vegetation where the prairie and hammock
meet, Coot Bay region, Cape Sable, Florida.

in tow for the Cape Sable region. They skirted the outside of the
Lower Matecumbe Key, passed through the viaduct and then headed
across the extensive flats of Florida Bay. The first stop was made
on Sands Key, where a burned out colony of Cerions and a well
flourishing colony of Oxystyla and Liguus were found.

The expedition next headed for Flamingo City, which was reached
at sundown. On May 9 they explored the region about Flamingo
City and the coast for several miles to the east, for Cerions, and other
mollusks. Cerions were not found, but large numbers of beautiful
Liguus and Oxystylas were noted everywhere. A trip was then
made by wagon to Coot Bay.

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Coot Bay is a fine sheet of water fringed by tall mangroves, show-
ing even at this season a sprinkling of water fowl. The evaporation
from the lake at night undoubtedly keeps the adjacent region sup-
plied with moisture, and is responsible for the large number of epi-
phytic plants which render the region a veritable hanging garden.
Orchids and tilandsias fairly draped and seemed to almost smother
the trees and shrubs. Here we saw quite a number of rare birds,
and among them the swallow-tailed kite, of which no less than six
specimens were observed in the air at one time.

On the toth an exploration was made of the region between
Flamingo and the middle of the bight between Middle Cape and

Fic. 44—Young great white heron (Ardea occidentalis) on the
lookout for his parents.

West Cape, but the search was rewarded with only a couple of dead
specimens occupied by hermit crabs which probably had been drifted
in here by the winds from the Sands Key colony. Liguus were
found in many places, and so were Oxystyla and other species.

On the evening of the 1oth the party headed for Indian Key, stop-
ping again at Sands Key, where they found a most remarkable
flight of Florida yellow throats. Every tree and every shrub seemed
to have on every branch one or more of these little fellows. On the
morning of the r1th sail was set from Indian Key for the Newfound
Harbor group, where the next two days were spent in examining the
chain of keys that extends southwest from Big Pine Key, for Cerions

NO. I SMITHSONIAN EXPLORATIONS, I919 45

and other mollusks. A very flourishing colony was found on the
northeast point of the first key southwest of Big Pine Key, where
were gathered no less than 75 specimens in a square foot of beach,
covered by grass. Another colony was found on the second key

Fic. 45.—A nest of Ward’s heron (Ardea herodias ward)
in gumbo limbo tree.

southwest of Big Pine Key. A colony was discovered likewise on
the third key southwest of Big Pine Key. The fourth key is a mere
clump of mangroves and did not contain Cerions. On the fifth key
southwest of Big Pine Key a colony had been planted in 1912, and it
is here that the crossing between the native and Bahama species has

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

taken place. On the sixth key no Cerions were found, but there
was secured the new clapper rail which has since been described
as Rallus longirostris helius, by Dr. Oberholser. Here, also, the
pictures of the Ward’s heron (Ardea herodias wardi) nest and
young were taken. This key might really be called Rail Key. In
the seventh no Cerions were found, but the great white heron
(Ardea occidentalis) was found breeding, and it was here that
we secured our photographs of this bird and the specimen, which
was shipped to the Zoological Park at Washington by parcel post

from Key West. It is believed that this is the first time this species

Fic. 46.—Young great white heron (Ardea occidentalis) in
the nest in a mangrove thicket.

has been shown in any zoological garden. The eighth and last
key is a mere clump of mangroves without Cerion possibilities.

On the evening of the 12th the expedition headed for Key West.
A large collection of Cerions was made near the Bureau of [Fisheries
station, to be used for breeding purposes at the Tortugas.

A new set of experiments was started on Loggerhead Key, Yor-
tugas. Ninety cages of bronze wire, 2’ x 3’ x 3’ high, were con-
structed in which were placed one specimen each of two species. In
addition to this nine cages 6’ x 8’ x 2’ high were built, in which 50
each of the two species were placed.

On the 19th opportunity presented itself to return to Key West on

a submarine chaser.

MO, SMITHSONIAN EXPLORATIONS, 1919 47

A summary of the results so far obtained from the breeding
experiments has recently been published as a bulletin by the Carnegie
Institution. The results are rather startling, and incline one, con-
trary to the views previously held by most people, to the belief that
Cerions are not easily influenced, if at all, by environmental factors.
The cross obtained in the Newfound Harbor colony opens up an
entirely new vista in the problem of speciation, and it was this which
was responsible for the starting of the new set of breeding
experiments.

Incidental to these investigations, which have been conducted since
1912, a list of the birds observed on the various keys has been kept
and published annually in the Year Book of the Carnegie Institution.

FIELD WORK ON THE MESA VERDE NATIONAL PARK,
COLORADO

The field-work of Dr. J. Walter Fewkes, Chief of the Bureau of
American Ethnology, at the Mesa Verde National Park, in the sum-
mer of 1919, was devoted to the excavation and repair of the pic-
turesque cliff dwelling, Square Tower House, known for many years
as Peabody House, and two low prehistoric mounds situated among
the cedars on top of the plateau. - This work was a continuation of
that of previous years and was carried on in cooperation with the
National Park Service of the Department of the Interior. As Square
Tower House has several unique structural features, the summer’s
work has added to the educational attractions of the Park. At least
two new types of hitherto unknown small-house ruins were dis-
covered, and it is believed that a new page has been added to the
history of the Mesa Verde cliff people. Dr. Fewkes was assisted in
his field-work by Mr. Ralph Linton, a temporary assistant, who con-
tributed much to the success of the work.

The main object was to gather data that may aid one to better com-
prehend the Indian civilization that arose, flourished on the Mesa
Verde, and disappeared from the plateau over four centuries ago.

Square Tower House is situated in a shallow cave at the head of
a spur of Navaho Canyon opposite Echo Cliff, about 2 miles south
of Spruce Tree Camp. It has long been considered by tourists one of
the most attractive cliff dwellings of the park, but its inaccessibility
has deterred all but the most venturesome from descending to it from
the rim of the canyon. Part of the old Indian trail (fig. 47) was indi-
cated by shallow foot holes cut in the almost perpendicular cliffs, and
previous to the past summer this was the only means of access.

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Without mutilating the vestiges of this primitive trail another was
made in the cliff near it, around which was constructed a balustrade
(fig. 48), with ladders conveniently set to aid those who wish to visit
the ruin.

Fic. 47.—The Indian trail from the canyon rim to Square Tower House.
The cliff below is almost perpendicular for over a hundred feet. <A party
of vistors aided by a rope is shown climbing along the trail. Photograph
by G. L. Beam. Courtesy of the Denver and Rio Grande R. R.

Square Tower House (fig. 49) measures 140 feet in length and
averages three stories high, with seven circular subterranean sanctu-
aries or kivas. The floor of the eastern end of the cave is composed

NOP SMITHSONIAN EXPLORATIONS, I9IQ 49

of large boulders fallen from the roof; that of the western end is
lower and comparatively level. The original entrance to the build-
ing, like that of the Cliff Palace, Far View House, and Sun Temple,

Fic. 48.—The footpath blasted in the cliff at the most
difficult part of the ancient trail where the tourists are
represented in figure 47. Photograph by Fred Jeep.

is a recess in the front wall. On the western end of the ruin there
protrude radiating walls of basal rooms, one story high, suggesting

c

2

oO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

on

a terrace. The rear wall of the cliff rises almost perpendicularly
from the floor with no recess back of the buildings. The destructive
effects of water dripping from the canyon rim are most marked
midway in the length of the building where the walls (fig. 50), now
reduced to their foundations, were formerly at least two stories high.

The walls of the ruin were in bad condition when the work began:

Fic. 49—Square Tower House before excavation and repair,
from Canyon rim.

great gaps in the masonry of the tower having rendered it in danger
of falling. The interiors of the rooms were choked with fallen
stones and the dust of ages. wo months given to excavation and
repair have put the ruin in fine condition, exhibiting a good example
of the best type of Pueblo architecture (figs. 51 and 52). The special
attractions of Square Tower House are the remains of the roofs of
two kivas and the high tower rising midway in its length.

NO. I SMITHSONIAN EXPLORATIONS, I9QI9Q 5l

The original roof beams (fig. 53) of these two kivas are almost
wholly intact. Considering how few kiva roofs on the mesa have
survived destruction in the lapse of time, especial care was exercised

Fic. 50—View of Square Tower House from the

west, before excavation and repair. Photograph by
Fred Jeep.

to preserve these and to indicate their mode of construction, and a
model (fig. 54) has been made, photographs of which, in successive
stages of construction, are given (figs. 54, 55). A good understand-

i)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

cn

ing of the construction of a typical kiva is especially important, as it
distinguishes cliff houses of the Mesa Verde from those found else-
where in the Southwest as well as in foreign lands.

Fic. 51—Square Tower House excavated and repaired as seen from
the southwest. Photograph by Fred Jeep.

Fic. 52.—View of the Tower completely repaired. Photograph
by Fred Jeep.

The kivas of Square Tower House are circular, subterranean in
position, and entered by a hatchway. [Each kiva has a fire hole F,
and near it an opening in the floor called the sipapu, L, which is very

NON 1 SMITHSONIAN EXPLORATIONS, IQIQ 53

sacred because it symbolizes the entrance to the underworld. Over
it in Hopi ceremonies is erected the altar, and through it the priests
call to their kin in the underworld. A most instructive feature in
the structure of the kiva is the means of ventilation. Between the fire
hole and the wall there is an upright slab of stone, H, a wall of
masonry, or simply upright sticks covered with clay. The function
of this object is to deflect pure air which enters the room trom a
shaft opening outside, G; the ventilator is morphologically the
survival of the doorway of the earth lodge or prototype of the kiva.

Fic. 53—View from below of one sector of original roof logs of kiva ‘A.
Photograph by Gordon Parker.

A characteristic feature of the kiva is the roof, which rests on six
mural pilasters, C; the intervals between which are called ban-
quettes, B, that (4) over the ventilator being wider and broader than
the others. The pilasters support logs, D, D’, D*, laid one above
another in the form of cribbing. Short sticks, D*, are placed at right
angles to the cribbing to prevent sagging. Upon this cribbing are
laid logs over which is spread cedar bark to support the clay covering
the roof. The hatchway, which also served for the passage of smoke,
is situated in the roof above the fire hole. In the construction of this
roof, men of the Stone Age in America were not far from the dis-
covery of the principle of the dome.

Fic. 54.—Model of a typical prehistoric kiva of the pure pueblo type.
The photograph shows the model from above (a) and from the side (0),
with first roof beams in place. Photographs by De Lancey Gill.

A, large banquette. E, pegs for ceremonial paraphernalia.
B, small banquettes. F, fire hole.

C, pilasters to support roof cribbing. G, external opening of ventilator.

D. beams of lower level of roof. H. fire screen, or pure air deflector.
D1, beams of second level of roof. I, niches for sacred meal.

D2, beams of third level of roof. Kk, floor entrance to ventilator.

D3, logs to prevent sagging of roof. L. ceremonial floor opening or stpapu.

Fic. 55.—Model of typical kiva of the pure pueblo type; a shows
construction of roof beams; b, half-covered roof and hatchway. Photo-

graphs by De Lancey Gill.

56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

The most striking feature of Square Tower House is the tower
from which it takes its name. The cave in which it 1s situated hav-
ing no recess at its back, there is consequently no refuse heap in the

Fic. 56.—Middle section of Square Tower House
from the Crow’s Nest. Photograph by Fred Jeep.

rear, such as was utilized at Spruce Tree House for mortuary pur-
poses. ‘The rear wall of the tower is formed by the perpendicular
cliff (fig. 56). As shown by windows, doorways, and remnants of

NO. I SMITHSONIAN EXPLORATIONS, I9QIQ

on
a |

O
eh

flooring, this tower is four stories high. The inner plastering
the lowest story 1s painted white with a dado colored red; its roof is
likewise well preserved.

A room near the western end, (fig. 57) of the ruin has doors and
windows closed with secondary masonry, and in the rubbish, half

filling the neighboring kivas, human bones were found, indicating

Fic. 57.—Western end of Square Tower House, excavated and
repaired. Photograph by Fewkes.

that the western end of the ruin was deserted and used for mortuary
purposes before the remainder of the ruin was abandoned.

There is no archeological evidence that the tribes to the east, north,
and west of the cliff dwellers and stone house builders of the Pueblo
area were stone masons. On the south of the area, in the valley of
the Gila, Santa Cruz, and San Pedro, looking toward Mexico, the
ancients built their houses of earth and logs, and while the prehistoric

buildings on the southern tributaries of the San Juan resemble those

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

on the Mesa Verde they are quite different from the pueblos now
inhabited. <A logical interpretation of the geographical distribution
of ruins with kivas like those of Square Tower House would not be
that the knowledge of stone masonry was introduced from Mexico,
but that the craft was acquired after the original inhabitants entered
the country, and that the pure pueblo type, or that with kivas like
those of Square Tower House, was born, cradied, and reached its
highest development in the area where it was found. But we may

Fic. 58.—Wall of Earth Lodge A, showing adobe plastering on earth; the
horizontal log is a roof beam.: Photograph by T. G. Lemmon.

take another step, and point out that the prototype of these pre-
historic kivas has a morphological likeness to “ earth lodges.”

The discovery of Earth Lodge A in this area by my assistant,
Mr. Ralph Linton, was important, considering the light it may
throw on the genesis of cliff dwellings. This ancient prototype (fig.
58) of a kiva is a semicircular isolated room with a slightly depressed
floor in which is a centrally placed firepit, the surrounding walis
being either adobe plastered on the earth or molded into clumps
shaped like rolls. In this rude sunken wall were set at an angle
posts, now charred at the free ends, all that remains of the supports

of roof and sides.

Z
2
4

SMITHSONIAN EXPLORATIONS, I9QI1Q

9

o1

Earth Lodge A was not only excavated but a shed (fig. 59) was
built over it for permanent preservation. While it is interpreted as
the prototype of a kiva, it was formerly the dwelling of a family or
other social unit dating to an epoch much older than that of the
cliff dwellers. On opposite sides of the fire hole at the periphery oi
the floor, but within the outer walls, are small square or rectangular
cists made of stone slabs set on edge. The indications are that these
were covered with sticks and clay, suggesting the so-called slab
houses. The pottery found in these cists is very crude, undecorated,
and not of the cliff house type.

Fic. 59.—Shed built over Earth Lodge A to protect it from the elements,
north end, entrance opposite. Photograph by Fred Jeep.

There are many sites resembling that of Earth Lodge A before
excavation awaiting investigation on the top of Mesa Verde. Near
it was a mound which when opened proved to be a unit-type house.
The crude masonry and rough pottery found in it indicate an advance
on the walls of an earth lodge, but the former is inferior to that of a
kiva of the highest development, suggesting that it is an intermediate
form between Earth Lodge A and Square Tower House. ‘The
spade revealed that after this room was first deserted debris had
filled the depression a few feet deep on which a new fire hole and a
grinding bin had been made of stone slabs on edge in the middle of
the depression. Later on it was again abandoned and human bones
had been thrown on the débris that formed over the grinding bin,

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

indicating that the depression had become a dump place.’ Last of all,
these were also covered by accumulated sand and soil, leaving only
the top stones of a pilaster projecting above the surface.

Fic. 60—Idol of the germ-god set by author in
cement at head of the stairway, near kiva B. Photo-
graph by Fred Jeep.

The pottery found in this crudely constructed kiva is more varied,
but still an advance on that excavated in Earth Lodge A. It may be
classified as black and white, and corrugated, but so inferior to that
typical of cliff houses that it can be readily distinguished. From this

Fic. 61—Mat made of sticks found with a skeleton in room west of
kiva D. Photograph by De Lancey Gill.

Fic. 62—Pottery rest made of agave fibre core wound with
feathered string.

62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL

ruin was taken a shard with a fine swastika, showing the antiquity of
this design so rarely found in Mesa Verde.

The general facies of the collection of artifacts from Square
Tower House is the same as in other cliff dwellings on the park,
and although a few specimens are different from those already
known, the majority corroborate, as far as age is concerned, the
testimony of the buildings. A broken fragment of the rim of a vase

Fic. 63.—Stick with carved extremity.

of the sugar bow] pattern, a type peculiar to the upper San Juan area,
was obtained from the Unit-Type House. Fragments of food
bowls corrugated on the outside, black and white on the interior,
belong to a type hitherto rare. No collector has thus far reported a
prehistoric pipe from Mesa Verde, but a stumpy straight tube of
unburnt clay, more like a “cloud blower ” than a pipe, betrayed the
fact that the cliff dwellers, like other Indians, smoked ceremonially.

NO. I SMITHSONIAN EXPLORATIONS, IQIQ 63

On their altar at the great winter solstice ceremony at Walpi, one
object of which is the increase of life by calling back the sun, the
Hopi now employ an idol representing the god of germination. This
idol is half oval in shape, the surface being painted with symbols of
corn. A similar undecorated idol (fig. 60), found at Square Tower
House, one of the best ever collected, was cemented by the author in a
conspicuous place at the head of the stairway.

An almost perfect reed mat (fig. 61), resembling those often
deposited with the dead, was found in a room of Square Tower
House. Good specimens of feathered cloth were wrapped around
skeletons of infants. A fine pottery rest (fig. 62), and a stick which

a

=< eR ew

ba

Fic. 64—Incised maze on one side of an artificially worked cubical
stone found with idol of the germ-god. The dotted line does not exist
on the specimen, but was placed there to enable the reader to trace the
meander. Photograph by T. G. Lemmon.

shows excellent carving on one end (fig. 63), occur in the collection :
there are also many bone needles, basket fragments, and other objects
similar to those elsewhere described.

A cubical stone with an incised design (fig. 64) found in the same
room as the idol of the germ god, is worthy of special mention as
the maze or labyrinth depicted upon it is unlike any pictograph yet
described from the Southwest.

Theoretically, Earth Lodge A is supposed to resemble forms of
dwellings that have survived to our day among non-pueblo tribes.

64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

It has, however, an instructive feature they do not possess, viz.,
cists made of slabs of stone set on edge. Evidences are accumulat-
ing of a culture antecedent to the pure pueblo type in which vertical
masonry predominates, but we must await more knowledge of the
construction of the houses of this epoch before speculating on the
early relations of the builders of vertical and horizontal masonry.

eh a ; Whe
we a a 3 ts ae)

Fic. 65—Square Tower House repaired, as seen from the west. Photo-
graph by T. G. Lemmon.

ARCHEOLOGICAL EXCAVATIONS IN ARIZONA
In continuation of work in Arizona for the Bureau of American
Ethnology, Dr. Walter Hough began excavation of an important
ruin in Grasshopper Valley, 14 miles west of Cibecue on the White
Mountain Apache Reservation, Arizona. The ruin consists of two

_

great mounds covered with brush and showing portions of walls.
The inhabitants, as shown by the skeletal remains, were Pueblo
Indians. Among the discoveries were a temporary camping place
of aclan while their houses were being constructed ; the use of heavy
masonry retaining walls to prevent the thrust in the earth covered
with the great structure of the pueblo; and the determination that
the house plans, sometimes called “ foundations,” and thought to be
unfinished structures, are remains of open air sheds, such as those
now in use by the Pimas. The presence of two very large debris

NORE SMITHSONIAN EXPLORATIONS, I9QITQ 65

Fic. 66.—Retaining wall of ancient pueblo at Grasshopper Valley, Arizona.

Fic. 67—Outlines of open-air sheds, ancient pueblos at Grasshopper

Valley, Arizona.

6

66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. -72

lenses, forming low mounds at 300 yards from the pueblo, is con-
sidered somewhat remarkable.

The artifacts consist of flint implements, stone axes, bone imple-
ments, and the usual variety of small objects. Of pottery, red and
gray preponderate ; a!so found are pure yellow (Hopi type) ; black
outlined with white on strong red (Chevelon type) ; obscure yellow
gray on brown with black designs (Gila type); and green glaze
design on white (Rio Grande type). Analysis of the shards from
the 20 excavations made show that the great ruin and the neighboring
members of the group may be differentiated on the basis of the

pottery fragments.

ARCHEOLOGICAL INVESTIGATIONS IN UTAH AND ARIZONA

Early in May, 1919, provision was made by the Bureau of Ameri-
can Ethnology for continuation of an archeological reconnoissance
of northwestern Arizona, inaugurated several years ago, but inter-
rupted by the recent world war. As in previous years, the work was
directly in charge of Neil M. Judd, curator of American archeology,
United States National Museum. Plans for an extensive survey of
the arid regions immediately north of the Rio Colorado were neces-
sarily abandoned because of the exceptionally dry season. Mr. Judd
then proceeded to Cottonwood Canyon near Kanab, Utah, where, in
1915, he had hastily examined several shallow caves containing pre-
historic remains.

As unforeseen conditions prevented completion of the recon-
noissance originally contemplated, so unexpected difficulties also
handicapped the Cottonwood Canyon investigations. Work con-
tinued during two weeks only, but in this short period twelve or more
caves were visited and five of them were examined with considerable
care. Most of the standing walls in these caves (fig. 68) were of the
customary stone and mortar construction, but others were found in
which adobe had been utilized almost exclusively. A third type of
architecture was that in which the walls were formed of upright posts,
bound together with horizontal willows and plastered over with adobe
mud. In such dwellings the heavy roof was ordinarily supported by
larger posts, placed as part of the wall or entirely within it. In these
ruins (fig. 69) and in the remains of other houses which had preceded
them, sandstone slabs were invariably utilized for the inner base of
the walls, the remainder being either of stone and mud or wattled
construction. Although dwellings of the three types above mentioned

NO. I SMITHSONIAN EXPLORATIONS, I9QIQ 67

Fic. 68.—Isolated rooms forming part of a small cliff village in Cotton-
wood Canyon. Several of the rooms were repaired after excavation;
the darker portions show the extent of these restorations.

«

tt eae

Fic. 69.—At the upper left are the broken walls of dwellings built
above the remains of a circular room. Fragments of wall posts and one
roof support are shown at the right; the fireplace at the left. In the
immediate foreground and in the middle of the view will be seen upright
slabs which formed part of the lower inner wall.

68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

seem to indicate different stages in constructional experience, they
could be and were found in the same cave. In such cases there was
nothing to suggest any considerable lapse of time between the periods
represented by the successive years of occupancy ; neither could it be
determined from the refuse in and about the dwellings that more than
one people had taken part in their construction.

From the minor antiquities collected it does not appear that the de-
eree of culture reached by the ancient inhabitants of Cottonwood Can-

Fic. 70—Masonry walls built above the ruins of a circular kiva, previ-
ously destroyed by fire. The banquette or bench surrounding the room
will be noted in the foreground; also, the charred fragments of several
wall posts and one roof support.

yon differed essentially from that of other primitive peoples farther
to the north.’ The pottery, generally, is of a type closely related to the
pre-Pueblo peoples south and east of the Rio Colorado, and indicates
a higher degree of experience than that noted among the ruins at
Beaver or Paragonah, for example. Wooden agricultural implements,
basketry, cotton cloth and other objects commonly found in cliff
ruins of the southwest are likewise of the well-known Pueblo type.
The results of these recent excavations tend to confirm, therefore,
the belief that in western Utah there is certain evidence of a pre-
historic people which originated some place in the northwest and

1 Smith. Misc. Coll., Vol. 66, No. 3, 1915; Vol. 66, No. 17, 1916; Vol. 68,
No. 12, 1917.

NO= SMITHSONIAN EXPLORATIONS, IQIQ 69

journeyed southward ; that during the course of their long-continued
migrations they changed rather rapidly from a semi-nomadic to a
sedentary life as they approached the Rio Colorado. Having
gained the “red rock” country and having found, for the first time,
natural caves that increased the protection afforded by their small
dwellings, they became more closely related, if not identical, in cul-
ture to those people commonly recognized as the ancestors of the
modern Pueblo Indians.

Fic. 71.—Walls of rectangular dwellings built above the remains of a
circular room. The upright slabs in the foreground formed the inner
wall base of the latter structure.

FIELD WORK ON THE IROQUOIS OF NEW YORK AND CANADA

Mr. J. N. B. Hewitt left Washington May 12, 1919, on field duty.
On the Onondaga reservation near Syracuse, N. Y., he found only
fragmentary remnants of the League rituals, laws and chants, aggre-
gating less than 2,000 native terms ; but these rituals, laws and chants
are so much broken and wasted away, and their several remaining
parts are so confused and intermixed the one with the other that
with these remains alone it would be quite impossible to obtain even
an approximate view of their original content, forms, and settings.
The texts which Mr. Hewitt has recorded among the Canadian
Iroquois aggregate more than 125,000 native terms. During the twe
weeks spent on this reservation Mr. Hewitt recorded in Onondaga

7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

text all the doctrines of the great Seneca religious reformer of the
close of the 18th century, Skanyodaiyo (the so-called ‘“ Handsome
Lake,” but which is literally “ It is a beautiful Lake ”’).

Mr. Hewitt left this Onondaga reservation May 31, 1919, going
directly to the Grand River Grant, Ontario, Canada, where the other
tribes of the six Iroquois tribes dwell. There he resumed his inten-
sive study and analysis of his recorded texts relating to the institu-
tions of the League, recording variant passages and terms when
encountered. He also obtained in detail the pattern of the wampum
strings in beads which are used in the Requickening Address of the
League. ‘Twenty-eight strings, at least, are necessary. On each
string the blue and the white beads are arranged according to a
definite pattern.

An effect of the war of the American revo‘ution on the tribal integ-
rity of the Six Nations of Iroquois was that every tribe, except the
Mohawk, was sundered into two independent bodies; and one part
of each of the divided tribes became resident on a separate reserva-
tion in the State of New York, and its public affairs became mea-
surably dissociated from those of the parts of the other tribes dwell-
ing in New York, while the complementary tribal parts removed to
Canada, where they finally settled on the Grand River Grant. So
that at first there were two Onondaga tribes, the one in New York
and the other in Canada, two Seneca tribal groups, the one in New
York and the other in Canada, two Oneida tribes, two Cavuga tribes,
and two Tuscarora tribes, similarly dispersed.

This disrupting of tribal integrity resulted in sundering the League
Federal Council into two independent units. Since the tribes in
New York State severally occupied individual reservations, often far
removed one from another, each tribe was thrown more on its own
resources than previously ; and the Federal Council composed of the
New York tribes was convened only when some matter affecting all
these tribes became urgent; and this situation naturally tended to
efface the concrete knowledge of the basic federal laws and principles
of the League from the minds of the New York tribes, so that within
50 or 60 years after this, the laws and the rituals of the original
League had become largely obsolescent, if not wholly forgotten, in
New York State.

Conversely, the tribes of the Six Nations of Iroquois who removed
to Canada and settling on the Grand River Grant elected to transact
their affairs at a semi-federal council composed of all tribal and all
federal chiefs (whose titles were not then held in New York State).

oa

NO. I SMITHSONIAN EXPLORATIONS, IQIQ FAM

This arrangement naturally tended strongly to preserve the tra-
ditional and the concrete knowledge of the distinctively federal insti-
tutions and laws and rituals. But, even here, a living and con-
structive knowledge of the institutions of the League is less definite
and is often displaced by dubious modern interpretations. So it has
become increasingly difficult, indeed, to obtain from the variant
extant versions of laws, traditions, rituals, songs and institutions,
their most probable original form.

The Onondaga tribes of New York State, dwelling 8 miles south-
ward from Syracuse, occupy the ancient seat of the Federal Council
of the League of the Iroquois. In the original structure of the
League, and still exercised by both the Canadian and the New York
federations, the Onondaga chiefs as a body exercised functions
approximating those of the presiding judge of a modern court sitting
without a jury, and had power to confirm or to refer back (but not to
veto) for constitutional reasons the decisions or votes of the Federal
Council.

OSAGE TRIBAL RITES, OKLAHOMA

In the spring of 1919 Mr. Francis LaFlesche, Ethnologist, spent
a month among the Osage Indians, gathering further information
concerning the ancient rites of that tribe and collected two rituals,
one from Wa-tsé-mo"-1", pertaining to the origin of the people
of the black bear gens, and one from Mo’-zho"-a-ki-da of the Peace
gens of the Ts1’-zhu division, as to the origin of that people.

The ritual obtained from Wa-tsé-mo?-1" contains 582 lines, divided
into 29 sections and arranged in groups according to subjects. The
first group of five sections describes the descent of the people from
the sky to the earth. The second group of four sections tells of the
appeal of the people to certain water insects who promise help. The
third group of eight sections speak of the Great Elk who brought to
light four different colored soils which he gave to the people to use
in this riteas symbols. The next group of a single section tells of the
wanderings of the people, of their meeting a man whom they learned
had descended from the stars and who gave to the people certain
sacred gentile names. Although they were warriors whose business
it was to destroy, they resolved to make the stranger their chief,
saying to one another: “ There shall be in him no anger, no violence
and he shall be a man of peace.”

The fifth group of three sections speaks of the parts of the swan
that were dedicated for use as a war standard. From this sacred
bird were taken personal names. The sixth group of four sections

72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72
recounts the finding of four different kinds of rock in their consecra-
tion for symbolic purposes. The seventh, a single section, speaks of
the finding of a female buffalo, the consecration of parts of its body

Fic. 72—Wa-tsé-mo"-i". Member of the Black Bear Gens.

and skin for ceremonial purposes. The eighth group of three sec-
tions recounts the search for a suitable kind of stone out of which
to make a ceremonial knife. On his third search the man chosen for

NO. I SMITHSONIAN EXPLORATIONS, I9QIQ 73

the duty returned with a red flint out of which was shaped a round-
handled knife and consecrated to ceremonial uses.

The ritual given by Mo?-zho®-a-ki-da contains 248 lines, divided
into 20 sections and arranged, according to subjects, into eight groups.
In the story told by Mo*-zho-a-ki-da the people of the Peace-maker
gens awakened in mid-heaven to a realization that they were a people
when a desire came upon them to descend to earth where they might
come into bodily existence. The first three sections record the down-
ward journey, of the people finding the first and second heavens
(counting downward) to be blank, but in the third heaven they met
the Man-of-mystery (the god of rain), who promised them aid in
their struggles upon the earth. The sixth section tells of the further
descent of the people, of their meeting the great Buffalo Bull who,
out of pity, gives them certain roots to use in healing their bodily
ailments. These medicinal roots are used to this day. The great
animal then gave them various kinds of corn and squash. The five
sections fo'lowing speak of the journey of the people over the earth,
of their coming to certain trees, to the waters of a river where they
met the spirit of waters, to water-plants each of which they adopt as
life-symbols. The next, a single section, tells of the coming of the
people to a dead animal which they adopt as a symbol. The signifi-
cance of the incident is not clear. In the next two sections is told of
the people coming to the center of the earth, meaning mid-summer,
the time greatest in fruitfulness (August), and to a “ beautiful
house ” which was to be their sanctuary. The next group 1s a single
section and speaks of the encounter of the people with another dead
animal (an elk), the meaning of which is not clear. The next two
sections speak of the people again coming to the center of the earth,
meaning another month (September), which, together with August,
makes mid-summer. The people at last reach another “ beautiful
house ” with many openings, wherein are to be sent all the children
of the people to be given their gentile personal names and assigned
to their proper places in the tribal and gentile order.

ETHNOLOGICAL STUDIES IN OKLAHOMA, NEW MEXICO,
AND CALIFORNIA
The year was spent by Mr. J. P. Harrington, ethnologist, in con-
tinuation of his studies of Southwest Indian languages and ethnology.
During the first five months of the calendar year he was engaged in
the study of the Ventureno and other Chumashan dialects of Cali-
fornia. [Especial attention was given to the ethnology as well as to
the language, the linguistic studies proving to be the key to the

74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

unearthing of considerable important ethnological data, notably on
subjects pertaining to material culture, sociology, and the little
habits of daily life and thought which are really fully as important
as the larger and more striking features of ethnology.

Fic. 73.—Ventureno Man.

The months of July, August and September were spent in New
Mexico in study of the Tano-Kiowan problem, and with as con-
clusive and gratifying results as could be desired. The entire struc-

NO. I SMITHSONIAN EXPLORATIONS, IQIQ 75

ture and vocabulary of Tanoan and Kiowa is one and the same, and
points to genetic unity at no exceedingly remote period in the past. A
peculiar series of phonetic shifts and changes occurring in these lan-
guages was fully investigated, and interesting studies in comparative
vocabulary were made.

Fic. 74—Hoop and pole game of the Ventureno Indians.

MUSIC OF THE PAWNEE, OKLAHOMA

In May, 1919, Miss Frances Densmore went to Pawnee, Okla-
homa, to begin a study of the music of the Pawnee Indians who live
in that vicinity.

She arrived at the time of the Buffalo ceremony which is held
every spring by the Pawnee, having for its original purpose the
securing of buffalo for food. The ceremony was held in an earth
lodge of the old type (figs. 75 and 76). Only initiates could be
present on the first day when the “painting of the buffalo skull”
took place, but, through the courtesy of the man in charge of the
ceremony, Miss Densmore attended the Buffalo dance and the Lance
dance which were held a few days later. These constituted the
second and third portions of the ceremony. During the Buffalo
dance the buffalo skull with its ceremonial decorations lay in front
of the “altar.’”’ Participants in the ceremony were seated in four
eroups, men in each group having their bodies similarly decorated
with symbolic designs. The principal singer was Wicita Blain, a

76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL 72

blind man, who at a subsequent time recorded a number of the songs
used on this occasion. These were old songs which he had received
by inheritance, and the words contain interesting examples of native
poetry. Blain also recorded ceremonial songs of the Bear dance,
which were his by right of inheritance.

One of the leading participants in the Lance dance was John
Luwak, chief of the Chaui band (fig. 77), who recorded numerous
old songs of various classes. Some unusually attractive songs were
heard during the Lance dance, but it was learned on inquiry that they

had been recently composed by the younger men of the tribe. As
old songs were desired these were not recorded. One of the old
Lance dance songs was obtained, with the words, “ Father, the band
of the dead is coming.” This was sung when the lance bearers
danced around the lodge. The study was limited to the music, no
study of the ceremonies being undertaken at this time.

Three other gatherings were attended by Miss Densmore, 1. e., a
hand game and two victory dances.

The victory dances were of unusual interest as they celebrated the
return of young men who had served in the recent war. Forty
Pawnee enlisted ; 39 returned without having suffered any casualty,
and one died of disease in France. Many of the men had been at the
front, several volunteered for a certain duty of special danger, the

No. I SMITHSONIAN EXPLORATIONS, I9Q19 ig

son of a chief served with the heavy artillery in a responsible position,
and wherever opportunity offered, they seemed to have made a
creditable record. All appeared to be in the best of health. A mem-
ber of the tribe said he “ believed this was because the people had
prayed for them, both in private and at all their public gatherings.”
The first victory dance was the occasion of the public rejoicing of
the women of the tribe, especially those whose relatives had been
in the war. This was similar to the scalp dances of the old times, and
about 40 women were in the dancing circle. War trophies were car-
ried aloft on poles, as scalps were formerly carried. In accordance
with old custom the soldiers presented their trophies to their nearest
kinswomen. One woman held a pole with a long knife fastened at
the end like the head of a spear, while below the knife was hung the
metal helmet which, with the knife, was taken from the enemy by her
son. As the women danced they lifted their trophies high in the
air, and expressed their enthusiasm in shrill cries. It was a scene of
free, native rejoicing. Miss Densmore was the only white person
present. Many war songs were sung, former deeds of valor were
related by the old warriors and the dancing continued for several
hours. The Pawnee are an emotional people, and some of the older
members of the tribe lamented with sobs and crying the diminution
of the tribe and the passing of the old ways.

The second victory dance took place several days later and was
entirely different in character. Many spectators attended, and the
building was crowded to its utmost capacity. At this dance the
returned soldiers were the principal interest, many appearing in
native costume and dancing the war dance. One such young man
told Miss Densmore that he had been at a government school prac-
tically all his life and never before had joined in a native dance. The
gathering opened with an impressive native ceremony, then speeches
were made by the chiefs of the bands, gifts were given and received
in a ceremonial manner, and the son of a chief was adopted by a
prominent member of the tribe, receiving an old and honored name.
The young man went through the simple ceremony with quiet dignity,
wearing the khaki uniform in which he had served in Europe. Once
the space around the entrance was cleared and a woman led in a
white horse, presenting it to a soldier; later a pair of white horses
were similarly presented. An interesting little drama was the “ con-
’ of the parents of the young man who died in France.

’

soling
The singing at the victory dances was characterized by songs with
words concerning the recent war, including mention of airplanes,

78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

submarines and the enemies of the allied nations. These words were
usually sung to old melodies. One woman advanced alone to a place
in front of the chiefs and, without the drum, sang such a song which
she had composed for the occasion. 'T'wo similar songs were recorded
by Luwak who related the dreams in which he heard them. He said
he “ prayed daily to Tirawahat,” saying, “ help our boys over there
so they will all come back strong and so I may live to see them again.”
One night he fell asleep after such a prayer and in his sleep “ some-
one told him that it would not be long before he would see the
Pawnee boys again.” In his dream he saw thousands of white people

Fic. 76—Pawnee ceremonial earth lodge, interior, showing location
of the “altar.” Photograph by Miss Densmore.

‘

rejoicing and heard them singing this song, “ even the oldest people

A few days later he heard of the

b

were dancing and waving flags.’
signing of the armistice, with its celebration, and accepted it as the
fulfilment of his dream. He related the dream and sang the song at a
gathering opened with an impressive native ceremony, then speeches

learned by the people.

MATERIAL CULTURE OF THE CHIPPEWA OF CANADA
In July, 1919, Miss Densmore visited the Manitou Rapids Reserve
in Ontario, Canada, through the courtesy of Mr. J. P. Wright, Indian
agent at Fort Frances, Ontario. Chippewa from other localities
in Ontario were also interviewed. The purpose of this trip was to

NO.

SMITHSONIAN EXPLORATIONS, I91Q

re;

Fic. 77—John Luwak, chief of the Chaui band, Pawnee.

8o SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

obtain data on the bead patterns, use of native dyes, medicinal herbs
and other phases of material culture among the Canadian Chippewa
for comparison with similar data already collected among the Chip-
pewa of Minnesota. Numerous old geometrical bead patterns were
noted which show an entirely different character from those in use
among the Minnesota Chippewa, and only two patterns were dupli-
cated. An interesting observation on native dyes consisted in the use
of a bright green dye made from four plants. Specimens of three
of these plants were obtained, but the fourth grew at a considerable
distance. A birchbark article decorated with roots colored with the
dye was also obtained. Green vegetable dye is not known among the
Minnesota Chippewa, so far as the present inquiry has extended.
The simpler medicinal herbs are the same as among the Chippewa
of Minnesota, but interesting variants of general customs were
observed.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 72, NUMBER 2

TWO NEW EAST AFRICAN PRIMATES

BY
N. HOLLISTER

~oeeleces

=

(PUBLICATION 2582)

GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 22, 1920

a

sis The Lord pre
BALTIMORE, &
;
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is

TWO NEW EAST AFRICAN PRIMATES
By IN. HOLLISTER

All of the East African monkeys and lemurs in the collection of
the United States National Museum have now been critically exam-
ined, and two forms have been found which require description. One
of these was collected by the Smithsonian African Expedition in
1909, and one by the Paul J. Rainey Expedition in 1911.

GALAGO SOTIKZ sp. nov.

Type from Telek River, Sotik, British East Africa. No. 184205,
U.S. Nat. Mus. Male, adult, skin and skull. (Basal suture closed.)
Collected May 22, 1911, by Edmund Heller (Paul J. Rainey African
Expedition) ; Orig. No. 1830.

Allied to Galago braccatus Elliot and Galago albipes Dollman, but
larger, with longer tail, larger hind foot, and larger ears. Darker
than braccatus and without the brightly colored limbs of that species.
General coloration much as in albipes, but legs and feet less buffy
cinnamon and toes less whitish, not sharply marked from color of
feet.

Color of type specimen.—General color of upperparts mouse gray,
washed with buffy; underfur deep mouse gray. Nose light gray;
eye rings and ears blackish. Arms and hands buffy gray, much like
color of back and sides ; legs slightly lighter than back, with suffusion
of buffy yellowish along inner side and extending over top of feet;
toes paler gray, with less intermixture of buff. Underparts gray,
heavily washed with pale yellowish buff. Tail brownish gray, more
brownish over terminal half, and slightly lighter below.

Skull and teeth—Skull averaging larger than in Galago braccatus,
with considerably larger auditory-mastoid bulla. Upper tooth row
slightly longer but last molar relatively smaller.

Measurements of type specimen.—Head and body, 200 mm. ; tail
vertebrz, 295; hind foot, 76; ear, 46. Skull: Greatest length, 49.4:
condylobasal length, 43.6; greatest breadth, 32.9; zygomatic breadth,
32.8; mastoid breadth, 26.3; interorbital breadth, 5.4; length of
mandible, 28.2 ; upper tooth row, canine to m ° inclusive, 16.8.

Three specimens from the type locality.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 72, No. 2

LS)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

LASIOPYGA PYGERYTHRA CONTIGUA subsp. nov.

Type from Changamwe, six miles inland from Mombasa, British
East Africa. No. 163327, U. S. Nat. Mus. Male, adult, skin and
skull. (Basal suture closed but not obliterated.) Collected Novem-
ber 30, 1909, by Edgar A. Mearns (Smithsonian African Expe-
dition) ; Orig. No. 7321.

Nearest to Lastopyga pygerythra tumbili Heller, but larger, with
larger skull and actually smaller teeth. Tail more grayish, less yellow,
than in tumbili, the longitudinal stripe above much less well-marked,
and the underside of tail gray, not tawny yellow.

Color of type specimen.—Face, lips, and chin black ; brow band and
cheeks buff, more or less mixed with gray; head, neck, and upper-
parts of body yellowish buff, somewhat mixed with gray ; rump and
hips more grayish, less buffy; underparts buffy. Arms and
legs outside speckled gray and buff, the inner sides clear buff;
hands and feet blackish, the fingers and toes clear black. Tail above
speckled gray and buff, beneath gray, the median line above not
sharply marked as in related forms; a bright chestnut bay spot
beneath at base, and the tip black.

Skull and teeth.—Skull larger than that of Lasiopyga pygerythra
tumbili; palate longer, extending backward considerably beyond
plane of last molars; mandible much longer and heavier. Teeth
actually smaller than in twmbili.

Measurement of type specimen.—Head and body, 570; tail, 720;
hind foot, 150. Skul! (with measurements of type of L. p. tumbili in
parentheses): Greatest length, 110 (104); condylobasal length,
&8.5 (81.2) ; zygomatic breadth, 72.6 (70.0) ; postorbital constriction,
46.8 (43.9); mastoid breadth, 59.8 (60.0) ; breadth of braincase,
57.6 (53.2); palatal length, 44.2 (41.2); length of mandible, 77.0
(70.3); upper molar-premolar row, 24.8 (26.4); middle molar,
6.0X 6.0 (6.8 6.8) ; lower molar-premolar row, 33.0 (32.5).

Three specimens from the type locality.

Compared with a series of eight examples of typical tumbili from
the Taita Hills, these three specimens from the coast region near
Mombasa are easily distinguished by the less sharply bicolored tails ;
gray, not yellowish, underside of tail; and the distinctly larger skull
with smaller teeth.

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 72, NUMBER 3

REPORTS UPON TWO COLLECTIONS OF MOSSES
FROM BRITISH EAST AFRICA

(WiTH Two PLatEs)

BY
TN. DIXON;: M.A; F. L. S.

(PUBLICATION 2583)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
SEPTEMBER 1, 1920

i >
‘ The Lord Baltimore Press 2
BALTIMORE, MD., U, S. A.
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REPORTS UPON TWO COLLECTIONS OF MOSSES FROM
BRITISH EAST AFRICA

By EN. DIXON, Wi: A. LS.

(Wits Two PLatTes)

I. THE MOSSES OF THE DUMMER-MACLENNAN EXPEDITION
TO MOUNT ELGON, 1918

Several sets of the mosses collected on this recent expedition to
Mount Elgon have been issued. I have received part of one through
Mr. T. R. Sim, of Maritzburg ; and I have also had through my hands
the sets belonging to the U. S. National Museum and to the Kew
Herbarium. The material in many cases is poor, but in others it is
good; and though the whole collection is not large, consisting of
about 50 numbers (many of which, however, contained several
species), it contains some interesting novelties, notably a species
of Holomitrium quite distinct from any of the African species, a
fine new Bryum of the Rosulata group, and an interesting new
Braunia. The types of these and of the other novelties are in the
U. S. National Museum; duplicate types are in my own herbarium.

An unusual feature of this collection is the number of species
generally rupestral, which were here epiphytic. This is notably the
case with Grimmia ovata Web. & Mohr, Amphidium cyathicarpum
(Mont.), Anomobryum robustum (the species of this genus are
normally rupestral or terrestrial), Bryum alpinum Huds., Hedwigia
albicans (Web.), and Ectropothecium lateriticolum Broth. The
suggestion may be hazarded that these mosses formed the ancient
flora of the crater, at the time when it was mostly unclothed with
vegetation, and that as the phanerogamic vegetation increased and
covered the rock surface the mosses were driven to the only sub-
stratum left for them, viz, the stems of the tree heaths, and other
wood, living or dead.

In the following list the abbreviation c. fr.=fruiting, and st.=
sterile.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 72, No. 3.

bo

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

DICRANACEAE
CERATODON PURPUREUS (L.) Brid.

Nos. 2391f, 3393b, 3414c. Mostly fruiting. The capsules are
frequently subsymmetrical, very little strumose, and only lightly
plicate, as I have found also in specimens collected by Scott-Elliot
in Central Africa; but these characters are far from constant and
may indeed be pathological.

HOLOMITRIUM MACLENNANI Dixon, sp. nov.
Cer ie 42.)

Sat robustum; habitu H. crispuli Mart. brasiliensis; laxiuscule
caespitosum, pallide olivaceo-viride. Caulis rigidiusculus, densifo-
lius, circa 3 cm. altus, parce divisus; folia e basi erecta ovata sub-
vaginante squarrosa, siccitate crispula, undulata, suberecta, superne
in acumen latiusculum actum sensim angustata, marginibus inferne
revolutis, supra dense arguteque et grosse inaequaliterque dentata;
costa sat valida, infra apicem desinens; cellulae superiores ovales,
dorso papillis nwmerosis dense scaberulae; basin versus sensim
elongatae, infimae aurantiacae, laeves, juxtacostales lineares, margi-
nem versus latiores, breviter rectangulares, pellucidae, ad alas de-
currentes perpaucae paullo dilatatae indistinctae.

Seta I-1.5 cm. alta, foltis perichaetialibus peraltis, circa tertiam
partem setae saepe attingentibus, convolutis. Theca subcylindrica,
‘circa 3.5mm. longa. Peristomium e dentibus irregularibus inaequali-
bus, in crura plura inaequalia, longa, pallida, brevissime interne
nodosa fissis instructum. Annulus multiplex.

Hab.: Heath Zone, 12,000 ft., epiphytic, rare; No. 3445.

A fine species, differing in the coarsely toothed, broadly pointed,
squarrose leaves from all the African species except H. acutum
Wright, to which it is allied; that species, however, has’ the leaves
more narrowly pointed, less sharply serrate, and with the base less
widely vaginant, and the perichaetia are much shorter. The peri-
stome teeth here are more or less equidistant, but very variable, some-
times showing a tendency to approximation in pairs ; they are very ir-
regular, but the capsules are overripe and it is difficult to know how
much of the irregularity is due to wear. Each tooth is made up of
two, three, or more vertical, filiform, red laminae, more or less
coherent, without transverse bars below, and without striolations ;
above they are pale, and closely articulate with shortly protuberant
internal nodules.

NO, 3 BRITISH EAST AFRICAN MOSSES—DIXON 5

DICRANOWEISIA AFRICANA Dixon, sp. nov.
(PI. I, fig. 1.)

Corticola; densissime caespitulosa, circa I cm. alta, flavo-viridis.
Folia conferta, madida erecta, sicca crispata, e basi anguste oblonga
sensim acuminata, carinata, acuta; marginibus inferne vel medio
leniter revolutis, supra planis, integris; costa latiuscula, variabilis,
tenuis, indistincte definita. Cellulae superiores majusculae, 6-8 »
latae, breviter rectangulares vel isodiametricae, angulis rotundatis,
laeves ; basilares omnes laxiores, lineares vel rectangulares, angulos
versus saepius dilatatae, numerosae, partem basilarem maximam
occupantes.

Autoica. Bracteae masculae internae perconcavae convolutae,
obtusae. - Perichaetium parvum, e foliis externis paullo brevioribus,
latius acuminatis, minus acutis, internis brevibus, late vaginantibus,
obtusis instructum. Seta brevis, 5-6 mm. longa, theca (vetusta)
parva, anguste elliptica, microstoma. Operculum et peristomium
non visa.

Hapb.: On tree heaths, in crater, 13,000 ft.; No. 3396.

The only two species of the genus hitherto found in Africa are the
widely distributed D. cirrata and D. crispula. The latter differs in
the plane margins of the leaves. The present species is very near
D. cirrata, but appears to be distinct in the denser tufts, smaller size
of all its parts, shorter capsule, and distinctly weaker nerve.

CAMPYLOPUS ACROCAULON (C. M.) Par.

Heath Zone and moorland, 12,000-14,000 ft.; Nos. 3412d, 239Ie,
3398b, 3407d, 3762. I determine these from the description alone,
having seen no specimens of the original plant from Kilimanjaro.
Sterile only. The color of the plant, length of stem, and evolution
of auricles vary considerably.

CAMPYLOPUS CAGNII Negri

Moorland in crater, 13,500 ft.; No. 3425, c. fr. I determine this
with some hesitation from Negri’s description of the sterile plant
from Ruwenzori. It appears to agree with that in every respect but
one: the leaves are given as 7.2 mm. in length, while here they are
only 4 to 5mm. long. The crowded, rigid, almost entire leaves, only
the comal (fructiferous) ones finely setaceous, ate rather character-
istic; the nerve is smooth at back. Capsules aggregate; calyptra
smooth at base.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE 72

POTTIACEAE
? TRICHOSTOMUM sp.

No. 3444c. A rather distinct, sterile plant, with somewhat sheath-
ing, flexuose, suberect, fragile leaves, and trichostomoid areolation,
may belong here, but its generic position is somewhat doubtful. The
description of T. usambaricum (Broth.) Broth. reads much like tt,
but I have not seen specimens.

LEPTODONTIOPSIS FRAGILIFOLIA Broth.

Heath Zone, epiphytic, 12,000 ft.; Nos. 3447g, 3772c; also No.
37506 in the Kew set. St. A very distinct plant.

TORTULA CAVALLII Negri
Nos. 3406, 3410, 3424, 3430b; mostly c. fr.

TORTULA EU-BRYUM (C. M.) Dixon

Tree heaths in crater, 13,500 ft.; No. 3426. With numerous
setae and a few old capsules, apparently small for the size of the
plant. It has not been found before in fruit. (Some of Dimmer’s
specimens issued under this number do not belong here but to
T. Cavalli.)

GRIMMIACEAE
GRIMMIA OVATA Web. & Mohr.

Nos. 3403, 3403b, 3409d, 3412c, 3418; mostly c. fr. Most or
all of these gatherings were epiphytic, on bark of trees, a very curious
condition.

ORTHOTRICHACEAE
ANOECTANGIUM sp.

Heath Zone, epiphytic, 12,000 ft.; No. 3759d. A small, delicate
species, with numerous setae and a few capsules in very old, poor
condition. It has the appearance of A. Wilmsianum (C. M.), but
shows larger, more pellucid cells, more tapering, short-acuminate
leaves, etc.

AMPHIDIUM CYATHICARPUM (Mont.) Broth.

On tree heaths; No. 3422b, c. fr.; No» 3444c¢ (p. p.),. st.

ZYGODON EROSUS Mitt.

No. 3412b, st. I have seen no specimens of Mitten’s plant, but
from the description there can be hardly a doubt that this belongs
there.

NOS, 3 BRITISH EAST AFRICAN MOSSES—DIXON 5

ORTHOTRICHUM UNDULATIFOLIUM C. M.

Tree heaths, western crater lip, 13,000 ft.; No. 3407c, c. fr. jun.
From the description there can be no doubt, I think, that this is C.
Miuiller’s plant.

ORTHOTRICHUM LEIKIPIAE C. M.

firee heathsin crater, 13,000 it.; No. 3417b (p.p.), c. fr.. A
fertile stem mixed with Braunia. The endostome segments are as
described by C. Miller, unusually broad, almost, in fact, connivent
at base. The plant is no doubt nearly allied to O. speciosum.

MACROMITRIUM HYALINUM Broth.
No. 3766, c. fr.

MACROMITRIUM ABYSSINICUM C. M.

Tree heaths, 12,000-13,000 ft.; No. 3413, c. fr.; No. 3756b, forma
laxiramea, c. fr.

After careful comparison of M. hyalinum Broth. with M. abys-
sinicum I feel very doubtful whether the two can be kept separate.
Brotherus rests his species on the smaller size and the very acute
leaves, often with hyaline tip. Comparison of Holst’s Usambara
plant, however, with W/. abyssinicum (No. 431, Schimp. Iter Abyssin.,
in Schimper’s herbarium) shows no difference, or at any rate no
constant difference, in leaf. The Central African plant appears as
a rule to run rather smaller than the Abyssinian specimens, but this
is by no means constantly the case and Dummer’s No. 3413 is as
robust as any of these. No. 3766 is smaller and agrees in dimensions
with Holst’s plant, and I have retained it provisionally under M.
hyalinum.

No. 3756b is a very peculiar plant. The stems are elongate, robust,
distantly and fairly regularly pinnate, apparently pendulous, with
none of the habit of a Macromitrium, but closely resembling some
forms of Papillaria. The leaves are very acutely long-acuminate,
often with the apex half-twisted, and when dry somewhat recurved.
I do not think, however, that the leaf form must be considered as
of any importance in comparing it with M/. abyssinicum (type), as
the leaves with which one is familiar there are the branch leaves,
while owing to the peculiar growth of this plant the bulk of the leaves
probably, and of course those of the primary stem, are of the nature
of stem leaves. A seta and capsule in my specimen are identical with
those of M. abyssinicum, while the specimen of No, 3413 in the Wash-

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.572

ington set indicates a distinct transition between this form and
ordinary MW. abyssinicum. It is, therefore, probably only a remarkable
form of that.

FUNARIACEAE
FUNARIA VOLKENSII Broth.
Moorland, 13,500-14,000 ft.; Nos. 2391b, 3414b, both c. fr.

BRYACEAE
BRACHYMENIUM FLEXIFOLIUM B. & S.

Tree trunks in crater, 13,000 ft.; No. 3423, c. fr. This agrees with
the Abyssinian specimens of Schimper’s at Kew, especially with No.
452. No. 29 has the outer teeth more strongly bordered, the endo-
stomial membrane paler and more pellucid, and the leaves with
rather smaller cells, and wider border. No. 452, however, agrees
quite well with Dummer’s plant in the peristome and cells, though
the border is somewhat wider, showing that these characters are
variable and not correlated, while here and there a leaf on Dtimmer’s
No. 3423 has the border exactly asin the Abyssinian plant. - More-
over, another specimen of Schimper’s (No. 552) has the leaves
exactly as here.

The outer teeth are densely barred, trabeculate within; the inner
membrane is rather low, the processes linear, distant.

BRACHYMENIUM STENOTHECUM Dixon, sp. nov.
(Pl. I, fig. 3.)

§ OrtTHocARPUS. Robustum, infra densissime tomentosum. Folia
confertissime interrupteque comosa, sicca erecta, haud torquescentia,
paullo flexuosa, inferne rubella, anguste oblonga, infra haud angu-
stata, supra parum dilatata non spathulata, breviter et acute acumi-
nata, marginibus inferioribus anguste recurvatis, superne dentibus
argutis tenuibus sat distincte dentatis, limbo lato flavido e cellulis
6-8-seriatis angustissimis incrassatis circumdata ; cellulae superiores
breviter hexagonae, 40-50 p longae; circa 3-41. Costa infra valida,
supra sensim angustata, in cuspidem perangustam flexuosam brevius-
culam integram excurrens.

Synoicum. Seta 3-4 cm. alta, flexuosa ; theca elongata, sat angusta,
cum collo ad 6-7 mm. longa, inclinans vel subpendula, e collo praelongo
in setam sensim attenuato anguste elliptica vel fusiformis, micro-
stoma, castanea, operculo brevi, conico, obtuso; exothecium e cellulis

INO lS BRITISH EAST AFRICAN MOSSES—DIXON a

parvis, valde irregularibus, inaequalibus, parietibus flexuosis instruc-
tum. Peristomii dentes fusco-aurantiaci, apicibus peracutis pallidis ;
opaci, plus minusve pallide marginati, dense lamellati, intus alte
trabeculati; endostomii membrana sat alta, aurantiaca, processubus
brevibus, inperfectis, irregularibus. Spori 25-30 p.

Hab.: Moorland in crater, 13,000 ft.; Nos. 3399, 3421, 3768.

The densely imbricate leaves, erect’ and little altered when dry,
little broader above and not narrowed to base, with broad border and
the long, narrow, subpendulous capsule, with very distinct collum,
distinguish this from the allied synoicous species, most of which have
the leaves twisted, or at least highly flexuose, when dry. B. abys-
sinicum C. M. has the leaves much narrowed to base and entire above,
the capsule smaller, etc.

I have figured the peristome, showing a process in the most perfect
condition, but they are mostly fragmentary and more or less adherent
to the outer teeth.

ANOMOBRYUM ROBUSTUM Dixon, sp. nov.
CE etion 74m)

E robustioribus generis. Stirps circa 2-2.5 cm. alta, pallide viridis,
subnitida. Caules julacei, foliis dense confertis ad 1.25 mm. longis,
suborbicularibus, cochleariformibus, obtusis vel obtusissime apicu-
latis, marginibus superioribus indistincte sinuolatis ; costa apud basin
60-70 p lata, rubella, medium folium versus multo tenuior, sed parum
angustata, circa tertiam quartam folii attingens. Areolatio superior
e cellulis angustissimis, vermicularibus, parietibus firmis nec incrassa-
tis instructa, inferior per tertiam partem folii laxissima.

Seta circa I.5 cm. alta; theca e collo longo, infra in setam sensim
angustato atque illic curvato, oblongo-elliptica, leniter curvata, infra
orificium paullo contracta, badia, operculo purpureo, obtuso, nitido.
Peristomium externum e dentibus aurantiacis, dense trabeculatis,
externe striolatis instructum, internum membrana praealta, hyalina,
laevi, circa 3-2 dentium altitudinem aequante; processubus brevibus,
inferne latis, superne raptim angustatis, filiformibus, subintegris, vel
angustissime rimatis; ciliis plerumque binis, swbaequilongis, articu-
latis nec nodosis, superne tenerrime papillosis. Spori parvi.

Hab.: Bamboo Zone, 9,000 ft., epiphytic; No. 3764.

The comparatively large densely crowded leaves give the stems a
robust appearance, and in this as well as in structural details it is a
markedly distinct species. A. promontorii (C. M.) Dixon has much
narrower, smaller leaves, a longer nerve, acute lid. etc.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

BRYUM ARGENTEUM L.

Nos. 3752, 3752b, c. fr. These plants afford rather more interest
than it usually falls to the lot of this ubiquitous species to present.
They show in the same gathering a form with the leaves widely
rounded at summit and obtuse, without any trace of apiculus or
hair-point, and others with long hyaline hair-points, practically identi-
cal with var. lanatum B. & S.; and all transitions between these
extreme forms occur, even (with the exception of the form with the
longest hair-points) within the limits of a single tuft.

It was gathered at an elevation of 12,000 ft., on the roofs of the
mud huts of the Ratmen. The Ratmen or Molemen—as their name
“Mese” signifies—are a small tribe of very primitive savages
inhabiting the extinct crater of this mountain.

BRYUM ALPINUM Huds.

Nos. 2391d, 3422, 3444b; No. 3422b (sparingly fruiting). These
plants were—a very unusual thing for this species—epiphytic, on tree
heaths. They differ somewhat from our northern B. alpinum in
being robust, with the nerve very stout, excurrent in a short, acute
or obtuse point, the leaves usually acute, subdenticulate at apex;
these characters however are not constant and are not correlated,
while some of the tufts are quite ordinary B. alpinum. The fruit
also agrees exactly. I have no hesitation, therefore, in placing them
under B. alpinum, which occurs also in South Africa, where, as in
the Northern Hemisphere, it shows considerable variation.

BRYUM BREVINERVE Dixon, sp. nov.
(CEN Ae snes fh)

§ Rosutata. E robustioribus generis, habitu B. spinidentis Ren. &
Card., vel B. perspinidentis Broth. Caulis validus, usque ad 10-12 cm
altus, inferne tomentosus, sat dense regulariter foliosus, vix, nist sub
tloribus, comosus; flexuosus vel hic illic geniculatus; folia erecto-
patentia, sicca flexuosa, contracta; 6-7 mm. longa, e basi angusta
decurrente, late obovata, obtusa, apiculo perbrevi plerumque reflexo ;
marginibus inferne anguste reflexis, superne sat conferte inter-
rupteque et breviter acuteque dentata; costa ad basin valida, rubra,
raptim angustata, supra perangusta, infra apicem sat longe desinens,
rarissime percurrens. Areolatio densa, e cellulis superioribus anguste
hexagonis circa 60 p longis instructa, infimis elongatis, angustis.
hexagono-rectangularibus, nec valde laxis ; marginalibus supra serie-

NO. 3 BRITISH EAST AFRICAN MOSSES—DIXON 9

bus 3-4 perangustis, incrassatis, limbum bene notatum, aliquando
rufescentem, inferne evanescentem, instruentibus. Fructus ignotus.
No. 3408b. Cliff base in thicket, western side of crater, alt. 13,500
ft.
A very fine species, quite distinct in leaf form and structure, and
especially in the short nerve, from any of its allies.

BARTRAMIACEAE
BARTRAMIA RUVENZORENSIS Broth.

Tree heaths in crater, 13,000 ft.; No. 3422c. Bamboo heath zone,
10,000 ft., No. 3444, c. fr.

BARTRAMIA STRICTULA C. M.

Moorland, 14,000 ft.; No. 2391c, st. This seems to agree with
C. Muller’s description of the above species. It is probably not dis-
tinct from the South African B. substricta Schimp.

BREUTELIA STRICTICAULIS Dixon, Smiths. Misc. Coll. 692: 21. 1918
Cliff base in thicket, 13,500 ft.; No. 3408, st.

BREUTELIA SUBGNAPHALEA (C. M.) Par.

Cliff base in thicket, 13,500 ft.; No. 3415, c. fr. This agrees per-
fectly in the vegetative characters; the seta is about I cm. long. C.
Muller says only, “seta perbrevi,’ which would seem to apply; the
seta in the Kew specimen of the original plant is perhaps slightly
shorter.

The peristome, not described by C. Miiller, is double, the outer
teeth well developed, red-brown, the inner fragmentary, pale orange-
brown.

POLY TRICHACEAE
POLYTRICHUM PILIFERUM Schreb.
Polytrichum nano-globulus C. M, Flora, 71: 408. 1888.
Nos. 2391, 3407e, 3414; all c. fr.

POLYTRICHUM HOEHNELII C. M.

Nos. 3409, 3776 (Kew set) ; both c. fr. This species, while closely
allied to P. commune, seems really distinct in the marginal toothing of
the leaves and in their position when dry, more or less spirally con-
torted with the points rigidly spreading— horride patentia.’’

ie) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL Ie 72

POLYTRICHUM KENIAE Dixon, Smiths. Misc. Coll. 697: 21. 1918

Moorland, 13,000 ft.; No. 3411, st. The present specimen agrees
well with the plant from Mt. Kenia, except that the sheathing leaf
base is not so elongated as in that; I have perhaps overrated the
importance of that character in the description.

HEDWIGIACEAE
HEDWIGIA ALBICANS (Web.) Lindb.

Nos. 3407, 3409b, 3419; all c. fr. These were growing on tree
heaths, an unusual station for what is commonly so rupestral a
species.

BRAUNIA BRACHYTHECA Dixon, subsp. nov. —
(Pl. I, fig. 5.)

Habitus, folia etc., omnino B. diaphanae, capensis, et B. secundae,
americanae. Differt solum thecae forma, Jatissime elliptica vel sub-
globosa, submicrostoma, ommnino fere sine collo, sicca vetusta sub-
plicata, ore latiore, suburceolata.

Hab.: On tree trunks in crater, 13,000-13,500 ft.; Nos. 3398,
2A 13h, 2403c; alc. fr. INOS. 3407, 34070; 2417 b-(p: pr) jest:

The vegetative characters, perichaetial leaves, and seta are so
exactly similar to B. diaphana, that I thought at first, in view of the
paucity of the capsules on my specimens, that it was possibly a case
of malformation of fruit. However, further material from the
U. S. National Museum and from Kew entirely confirmed the nor-
mality of the structure, and Mr. Sim writes to me that the capsules
on his three specimens—eight in all—are identical in the subglobose
form, scarcely tapering at neck, all except one being more or less
striate when old. In B. diaphana the capsule is narrowly elliptic, or
fusiform, being narrowed to the mouth, and with a well-defined
very gradually tapering neck (cf. pl. I, fig. 5b). Though often
somewhat wrinkled when old, moreover, it has no sign of regular
striae.

Mitten, in describing the mosses collected in Central Africa by
Bishop Hannington, refers a Kilimanjaro plant “ perfectly fruited ”
to Hedwigia (§Braunia) secunda Hook., and raises the question,
“ Are the B. sciuroides of Europe, the B. indica so luxuriant in the
Nilgiri Mts., and the Abyssinian B. Schimperi, really different, or
are they not most probably slight variations of one wide-spread

*Journ., Linn. Soc. Bot. 22: 310. 1886.

NO. 3 BRITISH EAST AFRICAN MOSSES—DIXON LEAL

species?” So far as my observation goes, B. sciuroides and B.
Schimperi stand on a somewhat different footing from the others;
but the Indian plant is most certainly identical with that which occurs
in several parts of Africa, and which has generally been known as
B. diaphana. It is éspecially frequent in South Africa, where it
fruits commonly. The fruit is described by Thériot ;* and Brotherus ’
says of it, “ Kapsel unbekannt.” I have several fruiting specimens
from South Africa, however, and the fruit occurs on several speci-
mens in the British Museum and Kew collections. Both vegetatively
and in the fruit the Indian plant is exactly identical with the African,
and as Mitten finds the Kilimanjaro plant identical with the Mexican
there can be no doubt that they must all fall under the name B.
‘secunda, As regards the Indian and African plants at least, I do
not find even the “ slight variations ’”’ which Mitten allows them. The
leaves vary in the degree of plication. They may be quite without
a hyaline point, or they may have a short hyaline tip, even occasion-
ally a quite long, flexuose hair-point, and the perichaetial leaves vary
much in length; but none of these characters shows any constancy,
nor are they correlated in any way with geographical distribution.

In the course of studying the Indian plant, however, I stumbled
upon a very unexpected thing: In the British Museum collection.
in Herb. Wilson, there are two specimens of an undetermined moss,
labelled “Indies, Winterbotham,” which are identical with my B.
brachytheca. I find no reference to it in any bryological works, and
it appears to have remained, otherwise than in Wilson’s herbarium,
quite undetected. It can scarcely be supposed that two independent
species, B. secunda and B. brachytheca, would exist side by side both
in Africa and in India, especially with the very restricted range that
appears to appertain to B. brachytheca; and I have therefore thought
it best to consider the latter as a subspecies of B. secunda.’

*Bull. Soc. Bot. Genéve IT. g: 135. 10917.

*Engl. & Prantl, Pflanzenfam. 1°: 718. 10905.

“Tt may be as well to give here the more important part of the synonymy of
B. secunda, so far as it bears on our African plant:

BRAUNIA SECUNDA (Hook.) B.S. G. Bryol. Eur, (29-30:) Braunia 3, 1846.
Hedwigia secunda Hook. Muse. Fxot. pi. 46. 1818-1820.
Neckera macropelma C. M. Syn. 2: 104. 1851.
Braunia macropelma Jaeg. Adumbr. 2: 87. 1869-1870.
Hedwigia indica Mitt. Journ. inn. Soc., Bot.3: Suppl. 123. 1859.
Braunia indica Par. Ind. 149. 1894.
Neckera diaphana C. M. Syn. 2: 105. 1851.
Braunia diaphana Jaeg. Adumbr. 2: 87. 1874-1875.

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

LEUCODONTACEAE
ANTITRICHIA KILIMANDSCHARICA Broth.

Heath Zone; Nos. 3413d, 3756, 3758b, 3772: No. 3756.acqum
good fruit, which has not been described, but I do not find any differ-
ence from that of A. curtipendula. The others are sterile.

NECKRERACEHE AE
NECKERA PLATYANTHA (C. M.) Par.

Heath Zone, principally; Nos. 3443, 3449, 3754, 3756f, 3759;
mostly c. fr. The perichaetial bracts may be three times as long as
the capsule, which, however, is not always concealed, as it may pro-
trude laterally from the perichaetium (Cf. pl. I, fig. 6).

NECKERA SUBMACROCARPA Dixon, sp. nov.
(Pl. I, fig. 7.)

Habitu foliisque N. platyanthae (C. M.) et N. macrocarpae Broth.
simillima, huic quoque speciei cauli paraphylliis numerosis praedito
similis et affinis, fructu tamen longe aliena. Perichaetium 8-10 mm.
longum, bracteis externis thecam longe superantibus. Theca im-
mersa, vel saepe e perichaetio lateraliter emergens, e seta pro more
praclonga, 2-2.5 mm. ; theca 2-3 mm., aurantiaca ; operculum conicum
breviter curvirostratum. Peristomii dentes longi, angusti, supra
dense tenereque infra grossius papillosi, non striolati, intus trabecu-
lati; endostomii membrana perbrevis, pallida, laevis; processus
anguste lineares, circa dimidiam partem dentium longitudinis
aequantes, pallidi laeves, carinati, haud rimosi, plus minusve nodosi.
Spori 25-30 wp.

Hab.: Heath Zone, 12,000 ft., epiphytic; No. 3443b.

Dummer writes on the label “ Frequent” ; but this is by confusion
with NV. platyantha, which in habit is identical or nearly so; the two
were growing intermixed, and the Washington specimen under this
number was entirely N. platyantha, which is evidently a frequent
moss on Mount Elgon, in the Heath Zone.

At first sight the fruit of the two species does not show any great
difference, but on examination it will be seen that in N. platyantha
the capsule is almost sessile and is hidden at the base of the peri-
chaetium, with the bracts two or three times its length; in N. sub-
macrocarpa the capsule itself is about the same length, but both

* Wissensch. Ergebn. Deutsch. Zentral Afrika Exped., 1907-1908, 2: 162.
IQT4.

NO. 3 BRITISH EAST AFRICAN MOSSES—DIXON 1

vaginula and seta being extremely long, together about equalling
the capsule length, it is much less immersed, often not even hidden.
The differences from N. macrocarpa Broth. may be tabulated thus:
Seta Theca Teeth Processes
N. macrocarpa Imm. 3-5mm. striolateatbase equalling teeth
N. submacrocarpa 2-2.5mm. 2-3mm. papillosetobase half length of teeth
Brotherus gives for his species certain characters derived from form
of leaf, and branching, which do not quite agree with the present
plant. I am not inclined, however, to lay much stress on these differ-
ences, as the branching and form of leaf apex appear to vary con-
siderably within the limits of the same species in this group; they
certainly do in N. platyantha. In fact the whole group of African
species, NV. Hoehneliana, N. Valentiniana, and the above mentioned
plants, are in my opinion quite inseparable from one another by
vegetative characters alone. For this reason I feel some doubt as
to the validity of N. subplatyantha Broth., which appears to be
separated from N. platyantha on vegetative characters alone. I
have not, however, seen the plant itself.

ENTODONTACEAE
LEVIERELLA FABRONIACEA ABYSSINICA (Broth.) Dixon
Fleath: Zone, epiphytic, 12,000 ft.; No; 3765, c.: fr.

FABRONIACEAE
FABRONIA sp.

Tree heaths in crater, 13,000 ft.; No. 3420. The quantity is too
- small for determination; it appears near F. Leikipiae C. M., but has
a very unusual range of denticulation, as among the leaves even of
a single plant.

HOOKERIACEAE

DALTONIA MILDREADII Broth. in Wissensch. Ergebn. Deutsch. Zentral
Afrika Exped., 1907-1908, 2: 164. 1914

No: 3423b; c. iri

HOOKERIOPSIS VERSICOLOR (Mitt.) Broth.

Without number. In quantity, and fruiting well.

= Op mcit. Lon.

T4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

THUIDIACEAE
THUIDIUM PALLIDISETUM Dixon, Smiths. Misc. Coll. 695: 8. 1918

Without number. The material agrees perfectly with Dimmer’s
original plant from Kipayo. The leaf cells are perhaps a little more
distinct and pellucid.

EYE NAC EAL
STEREODON CUPRESSIFORMIS (L.) Brid.

Nearly a dozen numbers contained this cosmopolitan species, in
- very varying forms. Three of them (Nos. 3445b, 3756g, 3756h)
represent a slender form, with narrow leaves and slender tapering
branches, which I have no doubt is the Hypnum Hoehneli: of C.
Miller; but they cannot be separated from the species, though
perhaps quite deserving of varietal rank, and I should call it Stereodon
cupressiformis (L.) var. Hoehnelii (C. M.) Dixon, comb. nov.

ECTROPOTHECIUM LATERITICOLUM Broth.

Without number. In good though somewhat old fruit. I have not
seen a specimen of the original plant, but the present specimen agrees
perfectly with the description. The only point of doubt would be
in the habitat, since the specific name of Brotherus implies a station
on stonework or brickwork; but this is not of great importance,
and it becomes still less so in view of the prevailing tendency towards
an arboreal habitat shown by the mosses of this locality.

RHAPHIDOSTEGIUM ELGONENSE Dixon, sp. nov.
(IPL 2 winter ayy)

§ AptycHus. Stirps, quoad species africanae spectantur, JK.
brachythecuformi (C. M.) et R. rivuletorum (C. M.) proxima. Sat
robustum, flavo-aureum, ramis turgidis, brevissime cuspidatis; folia
dense imbricata, saepius vix secunda, rarius paullo assurgentia, apici-
bus falcatis, 1.25-1.5 mm. longa, ovato-oblonga, supra cito angustata,
breviter acute acuminata, saepe semitorta, concava, marginibus
anguste explanatis, planis, vel angustissime recurvis, integerrimis ;
areolatio perangusta, pellucida, cellulis basilaribus aurantiacis, alari-
bus trinis magnis inflatis, supra-alaribus nonnullis majusculis pellu-
cidis.

Autoicum. Folia perichaetialia foltis caulinibus subsimilia, paullo
latiora, magis sensim acuminata, acumine latiore, swbintegro, interna

NO. 3 BRITISH EAST AFRICAN MOSSES—DIXON T5

erecta. Seta 1.25-1.5 cm. longa, laevis. Theca suberecta, cylindrica,
vix curvata, operculo curvirostro, subaequilongo.

Hab.: Heath Zone, 12,000 ft., epiphytic ; Nos. 3447, 3770.

It is rather difficult to diagnose the somewhat numerous species of
this section; but if habit, length of seta, form of capsule, and leaf
outline are taken into account, it will be found that there are no
African species very near this plant. FR. brachytheciiforme (C. M.)
is more robust, with a foliation strikingly like that of Brachythecium
albicans, and a different coloring. RR. rivuletorum is smaller, of
different habit, greener, with less crowded leaves, and horizontal or
subpendulous capsule.

BRACHYTHECIACEAE
PLEUROPUS SERICEUS (Hornsch.) Broth.
Heath Zone, 12,000 ft., epiphytic; Nos. 3759b, 3759c; st.

BRACHYTHECIUM VELLEREUM (Mitt.) Par.

Heath Zone, 12,000 ft.; No. 3756c. On tree heaths in crater,
13,000 ft.; No. 3441. Both fruiting. I have compared this with
Mitten’s plant, and there is no doubt of its identity. It is a very
striking species, but it is certainly autoicous. Mitten describes his
plant as dioicous, and I have found fruiting stems on which I have
been unable to detect male flowers, but there is no question that it
is normally autoicous.

I suspect B. gloriosum (C. M.) Par., of which the sterile plant
only was described, to be the same thing.

BRACHYTHECIUM UGANDAE Dixon, sp. nov.

Subgen. SALEBROSIUM. Robustum; luteo-aurewm, nitidum, habitu
B. salebrosi. Caules suberecti, irregulariter distanter ramosi, flexuosi,
subteretes, acutiusculi. Folia e basi subcordata late ovato-lanceolata,
in acumen anguste nec longe tenuiter acuminatum sat cito attenuata,
integerrima, profunde plicata, marginibus planis vel superne anguste
recurvis ; costa basin versus sat valida, cito multo attenuata, circa
dimidian partem folii attingens. Cellulae angustissimae, pellucidae,
alares multae, majusculae, subquadratae, inanes vel obscuriusculae,
bene notatae. Flores masculi majusculi, turgidi.

Autoicum. Perichaetia magna, foliis erectis, in acumen filiforme
flexuoso-recurvum integrum attenuata. Seta 1.5-1.75 cm. longa,
laevis; theca badia, suberecta, leniter curvata, oblonga, operculo
brevi, obtuso.

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Hab.: Tree heaths, 13,500 ft.; No. 3430. Bamboo Zone, 7,000 ft. ;
No. 3763. Heath Zone, 12,000 ft.; Nos. 3768b, 3775. All c. fr.

Very near to B. salebrosum (Hoffm.) and perhaps not specifically
distinct; but as that species has not been recorded from tropical
Africa, and as the present plant exhibits certain characters of dis-
tinction, notably the golden, glossy coloring, the terete branches,
and a suberect, only slightly curved, narrower capsule, I have pro-
visionally treated it as distinct.

BRACHYTHECIUM DUMMERI Dixon, sp. nov.
(El 2s ies, 110)

Subgen. CrrripHyLiopsis. Habitu B. stricto-patentis C. M.
capensis vel formarum gracillimarum B. implicati (Hornsch.).
Gracile, laete viride, vix nitidum, dense pinnatum, ramis circa I cm.
longis. Folia madida rigidiuscule erecto-patentia, e basi cordato vel
hastato triangularia, sensim longe acuminata, ubique denticulata ;
ramea brevius latiusque acuminata, dense et argute denticulata, con-
caviuscula, vix striata, nec plicata ; marginibus planis vel basin versus
angustissime recurvis; costa basi sat valida, superne foliis caulinis
multo, rameis paullo attenuata, circa secundam tertiam partem folu
attingens. Cellulae superiores perangustae, inferne seriebus pluribus
multo breviores, laxiores, pellucidae, alares sat numerosae, parvae,
subquadratae.

Autoicum. Perichaetia majuscula, archegoniis numerosis, foliis
erectis, latis, in acumen subfiliforme reflexum, denticulatum raptim
angustatis. Seta I-1.5 cm. longa papillosa. Theca horizontalis,
fusca, brevis, vetustate sub ore contracta, operculo conico, obtuso.

Hab.: Heath Forest Zone, 12,000 ft.; epiphytic; Nos. 3, 3425b,
B447e; 3700 3 7Omesa lic 2a.

Brachythecium stricto-patens C. M., which this resembles in having
the leaves rigidly subpatent when dry, differs in its smooth seta and
dioicous inflorescence; B. implicatum in its larger size and strongly
striate-plicate leaves. B. atrotheca Duby is more robust, with wider,
less rigid leaves and wider cells.

RHYNCHOSTEGIELLA ALGIRIANA (Brid.) Broth.

Epiphytic, on wood, Heath Zone, 12,000 ft.; No. 3447b; c. fr.
Also No. 3773 in the Kew set. This plant is exactly our European
and Northern African form, golden green. The epiphytic habit is
unusual, but is not unknown with us (the var. scabrellum, indeed, is
usually so). FR. Holstit Broth., from Usambara, ete., is a green plant

NO. 3 BRITISH EAST AFRICAN MOSSES—DIXON 17

. of a slightly different habit ; but I can find no structural differences,
and I am strongly disposed to consider it only a slight form of the
same thing.

II. A SMALL COLLECTION OF MOSSES FROM THE ABERDARE
MOUNTAINS

The mosses in the following list were collected near Mount Kenia

by Mr. A. Y. Allan in 1910, and were sent me for determination

by Rev. D. Lillie. Although the collection is small it is of unusual
interest, containing as it does the type of a new genus.

CAMPYLOPODIUM EUPHOROCLADUM (C. M.) Besch.

Nos. 395b, 398. This species has not previously been found in
Africa. It is known otherwise from Java, Tahiti, New Caledonia,
New Zealand. The second specimen above cited has very young
fruit and old setae.

[Campylopodium khasianum (Mitt.), a very closely allied species,
indeed doubtfully distinct, differs in its fruit only, so far as I have
been able to observe ; the vegetative characters described by Mitten do
not appear to hold good. There is the possibility, therefore, that the
African plant may belong there; but C. euphorocladum being a plant
of much wider distribution (and not improbably including C. khasia-
num) I have thought it best to refer this plant to the former. |

DICRANUM JOHNSTONI Mitt.
Dicranum Stuhlmannii Broth. Bot. Jahrb. Engler 20: 177. 18094.

No. 400, c. fr. I have compared this with the original of D.
Johnstoni (Kilimanjaro, H. H. Johnston 52), and it agrees perfectly.
It is also identical with D. Stuhlmannii (Stuhlmann 3290b, and
Volkens 1166, det. Brotherus). Mitten’s description of the leaves
of his species as “ linearia, sensim loriformi-angustata,” and “ unlike
any form of D. scopariwm from the narrower lower portions of the
leaf,’ is very misleading, and is no doubt the cause of Brotherus
having redescribed the plant. As a matter of fact, in Johnston’s
specimen the leaves, though occasionally (abnormally) narrow at
the base, are usually dilated there quite as in D. Stuhlmannii and
other species.

The fruit has not hitherto been described. Perichaetium about
6 mm. long, tubular, the bracts convolute with spreading points;
seta about 2 cm. long; capsule erect, symmetrical, cylindric, with very

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

gradually tapering neck and no struma, dark chestnut-brown, lightly
plicate when dry and old; lid finely subulate, erect or suberect;
deoperculate capsule about 4 mm. long.

CAMPYLOPUS JOANNIS-MEYERI (C. M.) Par.
Nan 207 ach arn:

FISSIDENS LINEARI-LIMBATUS C. M.

No. 403; st. From the description this would seem to agree exactly
with the original plant, collected in the same district.

KLEIOWEISIOPSIS Dixon, gen. nov.

Stirps habitu Astomi Hampe, sed cellulae superiores majusculae,
et folia superne distincte denticulata. Paroica; antheridia 3-4, infra
fructum, in folii perichaetialis axilla. Theca fere sessilis, in seta
brevissime sita, immersa, minima, globosa, microstoma, operculo
rostellato, annulo male evoluto persistente, calyptra parva, cucullata ;
peristomium nullum. Spori majusculi.

KLEIOWEISIOPSIS DENTICULATA Dixon, sp. nov.
(ei 2), inks 10)

Caespitosa seu dense gregaria; minuta. Stirps (vetusta) sordide
pallideque luteo-viridis ; circa 5 mm. alta, plerumque ad caulis basin
divisa. Folia erecto-patentia, sicca subcrispata, inferiora brevissima,
supra sensim longiora, superiora (fructifera) 3-4 mm. longa, e basi
concava latiore linearia, latiuscule breviterque acuminata, obtuse
acutata, carinata, marginibus planis, superne plus minusve grosse et
sat distanter denticulata, Costa infra circa 60 » lata tenuis, superne
angustata, sat pellucida, percurrens. Cellulae superiores 9-13 p, isodi-
ametricae et subquadratae vel breviter rectangulares, seriebus longi-
tudinalibus regularibus dispositae, pellucidae, perdistinctae, laeves,
basilares omnes perlaxae, rectangulares, hyalinae.

Theca profunde immersa; vaginula circa 200 p, seta 60-100 p, theca
400 » longa. Operculum subaequilongum, conico-rostellatum, curva-
tum, acutum. Calyptra parva, late cucullata. Spori 18-22 p, fusci,
conferte non alte papillosi. Exothecium e cellulis tenerrimis instruc-
tum ; infra orificium series 1-2 cellularum pellucidarum persistentium
quasi annulum imperfectum sistentes.

No. 395. With Campylopodium euphorocladum.

A remarkable little plant, the position of which is somewhat
doubtful. In areolation and denticulation the leaves are very similar

NO.. 3 BRITISH EAST AFRICAN MOSSES—-DIXON 19

to those of some species of Rhabdoweisia, and on this account it,
might be placed in the Dicranaceae. On the whole, however, it
seems to be best placed in the Pottiaceae, near Astomum and the
subgenus Kleiowetsia of Hymenostomum.

The plant is paroicous; but I suspect it may be heteroicous, as [|
have seen what seems to be a male flower below the fertile flower.

The capsule is, accurately speaking, neither cleistocarpous nor
stegocarpous. The lid is perfectly differentiated, and there is a
distinct row of subannular cells at the orifice; these may be in more
than one series. On the other hand, the lid is probably not normally
functional ; the capsule wall is of extremely delicate texture and under
pressure breaks up without the lid being detached, and this appears
to be the case also under normal conditions.

It is unfortunate that the altitude is not recorded. The association
of the plant with Campylopodium would seem to indicate a compara-
tively low level.

TORTULA ERUBESCENS (C. M.) Broth.
No. 399; st.

POLYTRICHUM COMMUNE L.
ING. 304:; St.

BRAUNIA SECUNDA (Hook.) B. S. G.

Nos. 401, 404; st. Presumably this species.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS - VOL. 72

EXPLANATION OF PLATES
PLaTE I[

Fic. 1. Dicranoweisia africana. a, Leaf, < 20; bb’, apex, X 50; c, cap-
sule, X 6.

Fic. 2. Holomitrium Maclennani. a, Plant, X 1, (left moist, right dry) ;
b, leaf, X 20; c, cells, X 200; d, peristome teeth, X 50.

Fic. 3. Brachymenium stenothecum. a, Stem, dry, X 1; a’, do., moist, X 1;
b, leaves, X 10; c, apex, X 40; d, part of peristome, XX 50.

Fic. 4. Anomobryum robustum. a, a’, leaves, X 20.

Fic. 5. Braunia. a, B. secunda, b, B. brachytheca; capsules, X 3.
Fic. 6. Neckera platyantha. a, a’, Capsule with perichaetium, X 2.
Fic. 7. Neckera submacrocarpa. a, Capsule with perichaetium, X 2.

PiateE II

Fic. 8. Bryum brevinerve. a, Stem, X 1; b, leaf, X. 10; c, cells in upper
part; X 50.

Fic. 9. Rhaphidostegium elgonense. a, Stem, X 1; 0, leaf, X 20; c, cap-
sule, X 5.

Fic. 10. Brachythecium Diimmeri. a, Stem, X 1; b, stem leaves, X 20;
c,'cells, X 200; d, perichaetium, X 5.

Fic. 11. Kletoweisiopsis denticulata. a, Plant (moist), X 1; a’, do. dry,
x 1; b, plant, X 8; c, leaf, X 20; d, leaf apex, X 40; e, upper cells, X 200;
f, basal cells, X 200; g, cells at orifice of capsule, 200.

VOL. 72; NO. 8, PL. 1

SMITHSONIAN MISCELLANEOUS COLLECTIONS

BRITISH EAST AFRICAN MOSSES

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72, NO. 3, PL. 2

Vs &

Sees Ta:

BRITISH EAST AFRICAN MOSSES

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 72 NUMBER 4

DIAGNOSES OF SOME NEW GENERA OF BIRDS

BY
ROBERT RIDGWAY
Curator, Division of Birds, U. S. National Museum

(PUBLICATION 2588)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 6, 1920

The Lord Waltimore Mress
BALTIMORE, MD., U. S. 4.
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DIAGNOSES OF SOME NEW GENERA OF BIRDS

By ROBERT RIDGWAY
CURATOR, DIVISION OF BIRDS, U. S. NATIONAL MUSEUM

OROAETUS gen. nov. (Buteonide)
Type: Falco isidori Des Murs.

Similar to Spizaétus Vieillot,” but rectrices much broader, the
width of middle pair equal to about one-fourth their length (from
point of insertion) ; feathers of legs longer, those of thighs plume-
like ; wing-tip relatively longer, the longest primary exceeding outer-
most (distal) secondary by more than one-fourth the length of wing;
tip of tail truncate; occipital plumes shorter, subcuneate; adults
with under parts striped, upper parts uniform black, and tail mottled
grayish with a broad terminal band of black ; size larger (wing more
than 500 mm.).

(“Opos, mountain; derds, eagle.)

SPECIES: Oroaétus isidori (Des Murs).

The type and only known species of this genus has been referred
by Sharpe and others to Lophotriorchis Sharpe (type, Astur kienerii
Geoffroy-St. Hilaire), but it would be difficult to find two forms of
this group (Spizaeti) more conspicuously different in structural
details or even in appearance. Lophotriorchis differs in (1) the
very broad cere, its width on top nearly if not quite equal to its
length; (2) relatively very much smaller bill; (3) densely bristled
loral region; (4) five outer primaries with inner webs deeply and
abruptly emarginated (instead of six obliquely sinuated) ; (5) very
different relative length of anterior toes, the inner (without claw)
not extending beyond the penultimate articulation of the middle toe,_
the outer very little longer, the middle toe (without claw) three-
fourths as long as tarsus instead of only about two-thirds as long;
(6) tip of tail distinctly rounded instead of truncate. In short,
practically the only character, apart from those common to the
whole group, possessed by the two genera consists in the unim-
portant ones of a similar crest and striped under parts of adults.

*With “ Falco mauduyti Daudin” =F. ornatus Daudin as type (subsequent
designation by Gray, 1840).

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VoL. 72, No. 4

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

PHAOAETUS gen. nov. (Buteonidz)
Type: Falco limnaetus Horsfield.*

Similar to Lophotriorchis in abrupt emargination of five outer
primaries and densely bristled loral region, but differing conspicu-
ously in having the cere much narrower (across top); relatively
much larger and more produced bill; in very different proportionate
length of toes, the inner (without claw) extending to decidedly
beyond penultimate articulation of middle toe; tail about three-
fourths (instead of less than two-thirds) as long as wing; wing-tip
shorter, the longest primary exceeding outermost (distal) secondary
by less than one-third the length of wing ; middle toe (without claw)
less than half (instead of three-fourths) as long as tarsus; toes and
claws much stouter, the latter relatively shorter and less acute, and
uniform dusky coloration.

(aids, brown, dusky; deros, eagle.)
Species: Pheoaétus limnaetus (Horsfield).

MORPHNARCHUS gen. nov. (Buteonide)

Type: Leucopternis princeps Sclater.

Similar to Leucopternis Kaup but differing in relatively shorter
and stouter tarsus (less than twice as long as middle toe without
claw), circular nostril, cuneate feathers of head and neck, and very
different coloration, the head, neck, chest and upper parts plain
black, the under parts (posterior to chest) white narrowly barred with
black; loral and orbital regions nearly nude.

(Mopdvos, dusky, dark; apyxés, a leader or chief.)

Species: Morphnarchus princeps (Sclater).

PERCNOHIERAX gen. nov. (Buteonide)

Type: Falco leucorrhous Quoy and Gaimard.

Somewhat like Rupornis Kaup, but relative length of anterior toes
very different, the outer toe (without claw) not longer than the
inner and extending to barely beyond penultimate articulation of
middle toe (instead of to beyond middle of penultimate phalanx) ;
middle toe relatively longer (more, instead of less, than half as
long as tarsus); tarsus less (instead of more) than one-fourth as

"This species is the type of Limnaétus Vigors, which name is antedated by
Limnetus Bowdich, the latter a synonym of Buteo.

NO. 4 DIAGNOSES OF SOME NEW GENERA OF BIRDS—RIDGWAY 3

long as wing, its upper portion more extensively feathered in front
(for one-half instead of only one-third, or less, the length of tarsus),
the naked portion of acrotarstum with much fewer (about 7 instead
of 12-13) transverse scutellae; cere much narrower across top, and
coloration radically different, the adults and young being conspicu-
ously different, the former mostly plain brownish black or blackish
brown, the latter with under parts buff or light ochraceous conspicu-
ously striped with dusky.

Also resembling somewhat the smaller species of Buteo, especially
B. brachyurus (which it resembles more in coloration than in struc-
tural details), but these differ in (1) having the wing-tip (distance
from tip of distal secondary to that of longest primary) equal to
much more than one-third the length of wing; (2) only three outer
primaries with inner webs emarginated ; (3) first (outermost) pri-
mary equal to eighth (instead of equal to tenth) ; (4) tarsus less
than one-fourth as long as wing (instead of more than one-fourth
as long), and also in possessing several of the characters distinguish-
ing Rupornis from Percnohierax.

_ (Ilepxvos, dark-colored, dusky ; ‘Tépaé, a hawk.

Species: Percnohierax leucorrhous (Quoy and Gaimard).
(Monotypic.)

HAPALOCREX gen. nov. (Rallide)

Type: Rallus flaviventris Boddaert.

Very small Rallee (wing about 66-69 mm.) with longest feather
of alula falling short of tips of longest primary coverts; bill nearly
as long as head; toes very long (the combined length of first two
phalanges of middle toe as long as tarsus, the hallux, without claw,
half as long as tarsus), and with a white superciliary stripe and
black loral stripe.

(‘A7rados, delicate; xpeé, a crake.)

Species: Hapalocrex flaviventris (Boddaert). (Monotypic.)

LIMNOCREX gen. nov. (Rallide)

Type: Porgana cinereiceps Lawrence.

Small Rallee (wing about 70-78 mm.) with longest feather of
alula extending decidedly beyond tips of longest primary coverts ;
nostril narrowly elliptical; tarsus shorter than middle toe without
claw (but longer than combined length of first two phalanges of
middle toe) ; outermost primary as long as (sometimes longer than)

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

distal secondary; bill subcuneate (tapering) in lateral profile, the
culmen elevated and more or less arched basally; and with sides,
flanks, and under tail-coverts conspicuously barred black and white.

(Aiuvyn, a pool, marsh; x«pé€, a crake.)

Species: Limnocrex cinereiceps (Lawrence) ; Limnocrex albigu-
laris (Lawrence) ; Limnocrex exilis (Temminck).

(Possibly the following species, which I have not seen, may also
be referable to this genus: Porgana enops Sclater and Salvin;
P. levraudi Sclater and Salvin, and Rallus leucopyrrhus Vieillot.)

THRYOCREX gen. nov. (Rallidz)

Type: Corethrura rubra Sclater and Salvin.

Small Ralleze (wing about 81-85 mm.) with bill not conspicuously
deeper. at base than at gonydeal angle, its width at posterior end
of nostril equal to decidedly more than half its depth at same point;
malar antia slightly anterior to the broadly rounded (convex) loral
antia ; longest feather of alula extending beyond tips of longest pri-
mary coverts ; outermost primary not projecting beyond tip of distal
secondary ; tarsus longer than middle toe without claw, and under
parts plain cinnamon-rufous.

(@pvov, a rush; xpégé, a crake.)

Species: Thryocrex rubra (Sclater and Salvin). (Monotypic.)

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 72, NUMBER 5

NEW SELAGINELLAS FROM THE:
WESTERN UNITED STATES

(With Six PLATEs) ©

BY
WILLIAM R. MAXON

(PUBLICATION 2589)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DEGEMBER 22, 1920

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NEW SELAGINELLAS FROM THE WESTERN
UNITED SEATES
By WILLIAM R. MAXON
(With Srx PLates)

Within the past twenty years a considerable number of species
have been proposed in the group of Selaginella rupestris, nearly all
of which, judged by a critical comparison of their essential though
minute characters, are undoubtedly well founded. In all, about
twenty-five species have been described from the United States.
These are so various in gross form and habit, and under a dissecting
microscope or even by use of a good hand lens show such remark-
ably diverse and constant leaf and sporophyll characters, that it is
hard to conceive of their ever having been regarded as, for the most
part, “ forms” of a single species. Extensive collecting, especially
in the Rocky Mountain region, is still necessary in order to clear up
the relationship of a few doubtful forms, and it is likely that explora-
tion in the Southwest will yield additional new species, since the
plants as a group are decidedly xerophilous or, at least, are able to
withstand long periods of drought, and so may be sought in those
arid out-of-the-way places that appeal chiefly to the natural history
collector. Specimens from any part of the southern and western
United States will, indeed, be gratefully received by the writer.

Of the six species here described the first is one of the interesting
assemblage of species growing together, often intimately associated,
in the Organ Mountains of New Mexico; the second is a plant of the
desert region of southern California, confused by Underwood with a
similar species from Zacatecas, Mexico; the third, long known to the
writer as distinct, is a related plant from Arizona; the fourth and
fifth are species of southern California, brought to light by the
_ energetic field work of a small group of enthusiastic botanists ; and
the sixth is a strongly marked plant, not uncommon in the Glacier
National Park, recently discovered during the course of intensive
botanical collecting in that region. In the lack of a monograph or a
synoptical account of the group as represented in the United States,
it has seemed especially desirable to accompany the descriptions by
illustrations. These, besides assisting in identification, will serve to

SMITHSONIAN MISGELLANEOUS COLLECTIONS, VOL. 72, No. 5

Z SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

show very well some of the more diverse forms assumed by members
of this group. All the illustrations are at natural size and represent
the type specimens in each instance, excepting only that of S. arizonica
which is of Thornber 315.

SELAGINELLA NEOMEXICANA Maxon, sp. nov.

(Pl. 1)

Plants strongly assurgent, 10 to 20 cm. long, the main stem rooting
sparingly at the extreme base, freely ramose, all the branches erect
or ascending, several times pinnate, subequal; stems (leaves ex-
cluded) mostly 0.2 to 0.5 mm. in diameter, the older ones readily
defoliate. Leaves uniform, rigidly ascending on all sides, subdistant
in attachment, imbricate but not wholly concealing the axis, 2 to
2.75 mm. long (seta included), the blades subulate-attenuate, 1.7 to
2.5 mm. long, 0.37 to 0.5 mm. broad at the base, setigerous (the seta
0.3 to 0.47 mm. long, whitish-hyaline from a greenish-lutescent base,
straight, sparingly serrulate), thin-herbaceous, spongiose at the base,
subglaucous, flat above, dorsally convex toward the narrowly obtuse
apex, sparingly pilose at the base of the deep narrow median groove,
ciliate, the cilia 12 to 20 on each side, slender, rigid, nearly straight,
mostly 0.06 to 0.125 mm. long, spreading or slightly ascending, the
apical ones reduced and more oblique. Spikes numerous, terminating
the main branches, 1 to 2 cm. long, about 1.5 mm. thick, recurved,
sharply quadrangular; sporophylls glaucous, yellowish brown with
age, readily detached, 2.4 to 2.7 mm. long (seta included), the blade
2.2 to 2.4 mm. long, 0.9 to 1.2 mm. broad, ovate, evenly long-
acuminate, setigerous (the seta 0.2 to 0.47 mm. long, greenish-
lutescent with a white tip, slightly scabrous), strongly concave, with
a deep dorsal groove throughout, freely short-ciliate, the cilia 25 to
32 on each side, close, stout, rigidly ascending, mostly 0.045 to
0.075 mm. long, the upper ones reduced. Megasporangia abortive
or wanting. Microsporangia very numerous; microspores yellow,
about 0.022 mm. in diameter.

Type in the U. S. National Herbarium, No. 591262, collected in
the Organ Mountains, Dona Ana County, New Mexico, at an altitude
of about 1,800 meters, January 9, 1909, by E. O. Wooton. There are
at hand three additional sheets of specimens collected in the same
range of mountains by Mr. Wooton on September 28, 1902, Septem-
ber 11, 1904, and March 3, 1907, the last mentioned associated with
S. rupincola Underw.

The present species has hitherto been referred doubtfully to
S. bigelovu Underw., of southern California, and this is clearly its

NO. 5 NEW SELAGINELLAS—MAXON 3

relationship. Selaginella bigelovii differs, however, in its more com-
pact habit and rigid, funiform branches, and more particularly in
details of leaf structure, the cilia being strongly oblique, very rigid,
short (mostly 0.03 to 0.04 mm. long), and pointed, often incurved ;
also, the setae are strongly scabrous throughout. Similar differences
are found in the sporophylls, those of S. bigelovu being coriaceous,
distinctly carinate, and tipped with a strongly scabrous seta.

SELAGINELLA EREMOPHILA Maxon, sp. nov.
(Pl. 2)

Plants wholly prostrate, the main stems up to Io or 12 cm. long,
coarsely radicose at intervals throughout, freely branched, forming
a close mat, the principal basal divisions subequal, divaricate, 2 or
3 times pinnate, the ultimate sterile branches very short, mostly 2 to
4 mm. long, about 2 mm. broad, involute upon drying; stems,
branches, and minor divisions all densely leafy, of pronounced dorso-
ventral aspect. Leaves crowded, in six ranks, those of the under side
the largest, about 2 mm. long, 0.5 mm. broad, exactly lanceolate,
acutish, not setigerous, ciliate (the cilia about 25 on each side, white,
spreading, mostly 0.075 to 0.125 mm. long), yellowish brown, im-
bricate, oblique-spreading, strongly secund upon drying ; leaves of the
upper side close-set, subimbricate, nearly vertical, straight or slightly
curved, deltoid-subulate, acutish, not setigerous, I to I.4 mm. long,
0.4 to 0.47 mm. broad at the base, at first bright green and sub-
glaucous, soon turning yellowish brown, flat above, broadly convex
beneath and sulcate in a median line nearly or quite to the tip, ciliate,
the cilia 6 to 12 on each side, weak, mostly spreading, about 0.1 mm.
long, similar minute hairs tufted at the base of the midrib and extend-
ing sparingly along the dorsal groove. Spikes numerous, arcuately
ascending, 6 to 10 mm. long, I mm. thick or less; sporophylls deltoid,
acute or acutish, not setigerous, mostly 1.2 to 1.4 mm. long, 0.9 to
I mm. broad, convex, subcarinate and sulcate dorsally, ciliate, the
cilia 12 to 18 on each side, spreading or weakly ascending, mostly 0.09
to 0.125 mm. long, rarely reaching the apex. Megasporangia few,
inferior, mostly basal; megaspores light yellow, 0.36 to 0.4 mm. in
greatest diameter, the commissural faces finely and deeply reticulate,
the outer face coarsely but sharply and deeply reticulate, the ridges
about 0.016 mm. broad ; commissural costae prominent, long. Micro-
sporangia numerous; microspores dull yellow, about 0.039 mm. in
diameter, long remaining associated in tetrads.

Type in the U. S. National Herbarium, No. 867484, collected in
Palm Canyon, Riverside County, California, April 4, 1917, by Ivan
M. Johnston (No. 1047) ; distributed as Selaginella parishii Underw.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Mr. Johnston’s notes accompanying the specimen read, “ Very
common in rock crevices and in their shade; Lower Sonoran Zone.
It is very hard to find a rock which hasn’t a large colony of this at its
foot. It grows with Selaginella bigelovu.”

The following additional specimens of S. eremophila, all from the
Colorado Desert region, are in the National Herbarium:

CALIFORNIA: Mountain Spring, San Diego County, alt. goo
meters, May 14, 1894, Mearns 3162. Top of Granite Mountain, seven
miles east of Julian, April 17, 1918, Bethel. Base of San Jacinto
Mountain, March, 1908, Saunders. Palm Canyon, eastern base of
San Jacinto Mountain, March, 1919, Hall. West Canyon, Riverside
County, alt. 200 meters, April 18, 1907, Parish 6111.- Tahquitz, near
Palm Springs, December 25, 1903, Dudley.

Selaginella eremophila is the plant of southern California (rare
in herbaria) which has been called S. parishii. Underwoad in describ-
ing S. parishit,, however, cited three collections, two of these from
the Colorado Desert (Parish 1200; Saunders), and the third from
Zacatecas, Mexico (Palmer 306). The California plant is specifically
distinct from the Mexican element, which, having been designated
by Underwood as the type, must bear the name S. parishti. The
dissociation of Mr. Parish’s name from so characteristic a species of
the Colorado Desert flora is especially regrettable.

Of the species hitherto described, S. eremophila is closely related

~only to S. parishu, of Zacatecas, and S. landu Greenm. & Pfeiffer,”
of Jalisco, both of which have a very similar dorso-ventral aspect.
Selaginella parishii is a more lax plant, with the ultimate branches
2 to 3 mm. broad, and larger, narrower leaves, whose characters may
be summarized as follows: Leaves of the under side 2.2 to 2.5 mm.
long, with about 18 cilia upon each side, these oblique, 0.078 to
0.125 mm. long; leaves of the upper side 1.3 to 1.6 mm. long, 0.35 to
0.43 mm. broad, with 4 to 8 cilia on each side, these oblique, 0.06 to
0.09 mm. long. The sporophylls, moreover, are broadly cordate-
deltoid, 1.5 to 1.7 mm. long, 1.3 to 1.4 mm. broad, with 25 to 30
“very oblique, close-set cilia on each side, those of the lower two-thirds
0.1 to 0.17 mm. long, forming a conspicuous fimbriate border. The
megaspores are pale yellow, about 0.42 mm. in diameter, and
delicately reticulate.

Selaginella landui is represented in the National Herbarium by a
portion of the type, Barnes & Land 2024 (San Esteban Mountains,

32 kilometers from Guadalajara, Jalisco) and by another collection

* Bull. Torrey Club 33: 202. 1906.
Ann. Mo. Bot. Gard. 5: 205. pl. 11, 12. 1918.

NO. 5 NEW SELAGINELLAS—MAXON 5

(Rose & Painter 7499) from the same locality. The essential char-
acters are as follows: Stems very firm, stiff, the branches rigid, not
intricate; leaves of lower side closely appressed-imbricate, lance-
attenuate, 2.7 to 3.2 mm. long, 0.6 to 0.7 mm. broad, with about 15 to
20 slender, mostly ascending cilia on each side in the lower half or
two-thirds (these 0.06 to 0.12 mm. long), the apical third with
pungent serratures ; leaves of the upper side crowded, rigidly vertical
or recurved, narrowly deltoid, evenly acuminate, I to 1.2 mm. long,
0.4 to 0.5 mm. broad, with about 13 to 16 cilia on each side, these
mostly oblique and incurved, 0.055 to 0.085 mm. long, the upper ones
passing into broad pungent serratures; sporophylls deltoid-ovate,
narrowly long-acuminate, 1.7 to 2 mm. long, 0.85 to I mm. broad,
variable in ciliation, sometimes with as many as 20 stiff, rigidly
ascending, mostly incurved cilia in the basal half (these 0.03 to
0.06 mm. long) and elsewhere serrate, or with short ascending teeth
along the whole margin above the extreme base (here with a few
cilia). Megaspores yellow, subglobose, about 0.33 mm. in diameter,
rugulose-reticulate, the ridges projecting sharply, less than 0.008 mm.
broad.

These three species, S. eremophila, S. parishui, and S. landi, while
readily distinguished specifically, are by no means typical members
of the group of S. rupestris, and together with the next species
(S. arizonica) forma fairly well-defined subgroup. The pronounced
dorso-ventral habit and subdimorphous leaves are doubtless to be
associated with their strongly xerophilous habitat.

SELAGINELLA ARIZCNICA Maxon, sp. nov.
(PIs)

Plants wholly prostrate, the main stems up to 20 cm. long, rooting
at intervals throughout, pinnately branched, the branches I to 1.5 cm.
apart on each side, the lower and middle ones 2.5 to 6 cm. long, twice
pinnate, the ultimate branches broadly subclavate, short, subdistant,
all the parts densely leafy, dorso-ventral, involute. Leaves crowded,
in six ranks, those of the under side the largest, appressed-imbricate,
oblique laterally, linear-lanceolate, acuminate, not setigerous, 2.5 to
2.8 mm. long, 0.5 to 0.6 mm. broad, thin-herbaceous, yellowish brown,
evenly ciliate, the cilia 18 to 22 on each side, oblique, up to 0.11 mm.
long, the apical ones reduced ; leaves of the upper side subulate, 1.8 to
2.2 mm. long, 0.3 to 0.44 mm. broad, evenly attenuate to the acutish,
whitish-marginate, short-setigerous apex (the seta dirty white, stout,
0.15 to 0.28 mm. long, serrate, often reflexed, present only in the

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

young leaves, caducous), ciliate (the cilia 4 to 8 on each side, ascend-
ing, pungent, 0.04 to 0.09 mm. long, passing into short serratures at
the apex), at first bright green, grayish with age, rigidly herbaceous
from a thick spongiose base, rigidly ascending, with age nearly
vertical in the older branches, flat above, slightly convex dorsally
and deeply sulcate nearly to the tip. Spikes ascending, terminating
the short ultimate divisions of the larger branches (or the divisions
wholly fertile), sometimes numerous, 2 to 5 mm. long; sporophylls
iarrowly ovate-deltoid, 1.5 to 1.95 mm. long, 0.75 to 0.85 mm. broad,
evenly long-acuminate, at first barely setigerous (the seta 0.15 to
0.25 mm. long, broad, pointed, serrulate, dirty white), subcarinate,
ciliate, the cilia 18 to 22 on each side, stout, rigidly ascending, mostly
0.03 to 0.06 mm. long, the upper ones greatly reduced, dentiform.
Megasporangia numerous, basal, or sometimes interspersed among
the microsporangia ; megaspores pale yellow, subglobose, 0.36 mm.
in greatest diameter, coarsely reticulate on the outer face (the ridges
sharp and narrow), finely reticulate on the commissural faces ; com-
missural costae long, delicate. Microsporangia numerous; micro-
spores orange, about 0.035 mm. in diameter.

Type in the U. S. National Herbarium, no. 694327, collected at
the foot of Soldier Trail, Santa Catalina Mountains, Arizona, altitude
about 960 meters, July 28, 1914, by Forrest Shreve. Other specimens
in the National Herbarium are as follows:

ArIzoNA: Sabino Canyon, Santa Catalina Mountains, alt. 870
meters, June 15, 1903, Thornber 315. Ventana Canyon, near Tucson,
1913, Cook. Pimo Canyon, near Tucson, February, 1913, Parish
8513. Arivipa Canyon, April, 1873, P. F. Mohr. Santa Catalina
Mountains, April 3, 1894, Toumey. Roosevelt Dam, on steep rocky
slopes, August 3, 1910, Goodding 722. Salt River Mountains, alt.
600 meters, November 9, 1913, Bailey. Ray, 1913, Bailey.

Selaginella arizonica is related to S. eremophila, and to S. land
and S. parishii, whose characters are given in detail under the last
preceding species. The disparity in size between the leaves of the
under and upper sides is far less in S. arizonica than in the others,
and it alone of the four species has the leaves and sporophylls at all
setigerous. Jt is a much larger and coarser plant than S. eremophila
and differs in nearly all technical leaf characters.

SELAGINELLA ASPRELLA Maxon, sp. nov.

(PI. 4)
Plants forming a loose mat, the main stems 3 to 6 cm. long, creep-
ing but not prostrate, rooting at intervals throughout, with a few

NO. 5 NEW SELAGINELLAS—MAXON 7

laxly ascending branches, these close, usually intricate, 1 to 2.5 cm.
long, twice pinnate, the ultimate divisions 3 to 7 mm. long, oblique,
slender, all the parts scantily leafy ; main branches (leaves excluded)
about 0.6 mm. thick, tardily defoliate. Leaves uniform, rigidly as-
cending on all sides, subdistant, decurrent, subimbricate, 2.75 to
3.2 mm. long (seta included), the blades narrowly deltoid-subulate,
1.85 to 2.3 mm. long, 0.55 to 0.7 mm. broad at the base, long-setigerous
(the seta 0.7 to 0.9 mm. long, white-hyaline, slender, subflexuous,
serrulate-ciliate throughout, the cilia up to 0.04 mm. long), charta-
ceous, inflated, concave above, broadly convex beneath, with a deep
median groove, very strongly glaucous, with a more or less well-
defined whitish-hyaline border (0.045 to 0.075 mm. broad at the
middle of the leaf), ciliate, the cilia 16 to 23 on each side, spreading,
straight or often curved, mostly 0.05 to 0.09 mm. long, the upper
ones distant, shorter, oblique. Spikes numerous, loosely aggregate
at the ends of the short apical branches, I to 2 cm. long, 1.5 to 2 mm.
broad, arcuate, sharply quadrangular; sporophylls laxly imbricate,
strongly glaucous, yellowish brown with age, 2.5 to 3 mm. long (seta
included), the blade narrowly ovate-deltoid, evenly long-acuminate,
1.9 to 2.2 mm. long, 0.85 to I mm. broad at the base, long-setigerous
(the seta stiff, straight, 0.6 to 0.8 mm. long, whitish, strongly
scabrous), carinate, with a deep dorsal groove, strongly whitish-
marginate, ciliate, the cilia 25 to 35 on each side, slightly oblique,
mostly 0.03 to 0.06 mm. long, the apical ones few and reduced.
Megasporangia few, mostly basal; megaspores pale to bright yellow,
subglobose, about 0.375 mm. in diameter, lightly reticulate on all
faces, the meshes broad, with low ridges; commissural costae
prominent. Microsporangia very numerous; microspores bright
’ orange, about 0.033 mm. in diameter.

Type in the U. S. National Herbarium, No. 867507, collected at
the west end of Ontario Peak, San Antonio Mountains, southern
California, altitude about 1,800 meters, in rocky ground, March 25,
1918, by Ivan T. Johnston (No. 1815). Other material, all from the
same range of mountains, has been examined, as follows: San
Antonio Canyon, in shelter of rock on the dry, open canyon floor,
alt. 1,725 meters, July 28, 1917, Johnston 1595; Ontario Peak, in
crevices of a sunny, exposed granite cliff, alt. 2,475 meters, December
22, 1917, Johnston 1807. The last-mentioned specimen is dwarfed,
and the leaf parts scarcely attain the size given in the description.

Selaginella asprella is a strongly marked species, without any
very close relatives. Of western United States species it is related
only to S. bigelovii Underw., S. rupincola Underw., and S. neo-

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

mexicana Maxon, all of which are much larger plants of essentially
erect growth and differ, besides, in numerous technical characters.
The very slender, rigid branches and spaced, half-appressed, strongly
setigerous leaves give the plant a scant, harsh aspect, which has sug-
gested the specific name.

SELAGINELLA LEUCOBRYOIDES Maxon, sp. nov.
(Pl. 5)

Plants very closely prostrate, the stems short-creeping, 1 to 2 cm.
long or less, closely aggregate, simply pinnate, the divisions thick,
strongly cespitose, erect, only 2 to 7 mm. long, or the terminal ones
bearing erect elongate spikes, all the parts densely leafy. Leaves
crowded, closely appressed-imbricate, mostly incurved, glaucous,
linear-subulate, uniform as to shape, variable in size, the basal ones
2.8 to 3.25 mm. long (seta included), 0.44 to 0.53 mm. broad, the
upper ones mostly 2 to 2.8 mm. long (seta included), 0.42 to 0.5 mm.
broad, all short-setigerous at the acutish whitish thickened apex (the
seta stout, white, not translucent, subflexuous, 0.125 to 0.28 mm. long,
strongly scabrous, often reflexed), ciliate (the cilia 8 to 16 on each
side, those of the basal half spreading, 0.6 to 0.13 mm. long, the
upper ones shorter, distant, ascending), thick, rigidly herbaceous,
flat or broadly concave above, convex beneath (strongly so toward
the apex), the median groove deep, broad, nearly percurrent. Spikes
relatively numerous, aggregate, 5 to 10 mm. long, about I.5 mm.
thick, erect, nearly straight ; sporophylls rigidly appressed-imbricate,
deeply concave, narrowly to broadly deltoid-ovate, evenly long-
acuminate, about 2 mm. long, 0.8 to I mm. broad, short-setigerous
(the seta white, rigid, pointed, subentire, about 0.15 mm. long or
less), ciliate-serrulate ; cilia or teeth 20 to 25 on each side, the basal
cilia not more than 0.046 mm. long, pungent, oblique, passing grad-
ually into oblique hyaline teeth toward the apex. Megasporangia
few, basal; megaspores subglobose, bright yellow, about 0.47 mm. in
diameter, the outer face obscurely reticulate, the commissural faces
manifestly so; commissural costae short, elevated. Microsporangia
numerous ; microspores orange, about 0.039 mm. in diameter.

Type in the U. S. National Herbarium, No. 982453, collected at
Bonanza Mine, Providence Mountains, southeastern California, alt.
840 meters, in crevices, rocky mountain side, March 30, 1920,
by P. A. Munz and R. D. Harwood (No. 3789). The following addi-
tional material is at hand:

NO. 5 NEW SELAGINELLAS—MAXON 9
®

CALIFORNIA: Surprise Canyon, Panamint Mountains, Inyo County,
alt. 1,400 meters, April 14, 1891, Coville & Funston 628. Vicinity
of Bonanza King Mine, east slope of Providence Mountains, Mojave
Desert, alt. g60 meters, May 11-24, 1920, Munz, Johnston & Harwood
42206.

The relationship of Selaginella leucobryoides is difficult to deter-
mine, since the plant differs not only in megaspores but in most
foliage characters from all other species of the Pacific Coast region.
The most striking characteristics are the extremely short, pure white
but opaque seta of the leaves and the condensed, rosette-like arrange-
ment of the very short branches. In habit and color there is a strong
suggestion of the tufted growth of some of the smaller species of
Leucobryum.

The Panamint Mountains plant collected by Coville and Funston
(No. 628) was mentioned as a critical form by Underwood in his
initial work upon the United States species allied to S. rupestris
It is clearly a reduced state of the present species, differing from the
type only in its lesser size. The leaves are only 1.65 to 2 mm. long
and 0.35 to 0.44 mm. broad; the seta and cilia characters are identical.
The plant collected by Munz, Johnston, and Harwood (No. 4226)
also comes from a higher elevation than the type collection and is
somewhat smaller.

SELAGINELLA STANDLEYI Maxon, sp. nov.
(Pl. 6)

Plants closely prostrate, the main stems up to 6 cm. long, finely
radicose, pinnately branched, the larger basal branches up to 2.5 cm.
long and with a few short alternate divisions, the upper branches
-simple or once dichotomous, all the divisions cespitose, rigidly
arcuate-ascending. Leaves crowded, imbricate, rigidly appressed,
those of the older stems of a characteristic bronze color, relatively
large, the blades broadly acicular, acutish, up to 2.5 mm. long and
0.6 mm. broad, with a short lutescent seta; leaves of the branches
mostly dull green, oblong-linear, 2.1 to 2.5 mm. long (seta included),
0.35 to 0.45 mm. broad, setigerous at the narrowly obtuse apex (the
seta 0.6 to 0.78 mm. long, lutescent throughout, coarsely serrulate-
scabrous), ciliate (the cilia I0 to 14 on each side, hyaline, stiff,
oblique, mostly 0.05 to 0.06 mm. long, passing into pungent serratures
toward the apex), rigidly herbaceous, flat above, convex beneath and
reddish along the deep median groove, especially toward the cymbi-
form apex. Spikes numerous, mostly geminate, erect from a curved
base, 7 to 11 mm. long, about 1.5 mm. thick; sporophylls deltoid to

ie) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. . 72

ovate-deltoid, 1.8 to 2 mm. long, 0.9 to I mm. broad, acuminate,
setigerous at the acutish tip (the seta 0.2 to,o.4 mm. long, stout, rigid,
scabrous, lutescent from a darker base), ciliate, the cilia close, 16 to
21 on each side, stiff, rigidly ascending, up to 0.08 mm. long.
Megasporangia few, basal; megaspores orange-yellow, 0.46 to 0.5
mm. in diameter, oblate-spheroidal, rugose in all aspects, the com-
missural costae short and prominent. Microsporangia very numer-
ous; microspores dull orange, about 0.032 mm. in diameter.

Type in the U. S. National Herbarium, No. 1028638, collected in
the vicinity of Sexton Glacier, Glacier National Park, Montana,
altitude 1,950 to 2,220 meters, on a moist rocky slope, August 7, 1919,
by Paul C. Standley (No. 17228). Other material studied is as
follows:

Montana (Glacier National Park): Gunsight Pass and vicinity,
alt. 1,775 to 2,100 meters, August 25, 1919, Standley 18136; August
25, 1917, Ulke. Along the trail from Many Glacier Hotel to Piegan
Pass, alt. 1,500 to 2,160 meters, August I1, 1919, Standley 17483.
Vicinity of Iceberg Lake, alt. 1,740 to 1,950 meters, July I1, 1919,
Standley 15363. Ptarmigan Lake, alt. 1,800 to 1,900 meters, August
3, 1919, Standley 16970.

ALBERTA: Tunnel Mountain, alt. 1,650 meters, June I1, 1906,
Brown 95. :

The writer takes pleasure in dedicating this excellent species to
Mr. Paul C. Standley, who, in the course of his botanical exploration
of Glacier National Park, assembled an extraordinarily rich collec-
tion of material in this group, the specimens comprising (besides
S. standleyt) S. montanensis Hieron., S. densa Rydb., and S. wallacei
Hieron., all in ample series.

Superficially S. standleyi most resembles S. watsoni Underw., of
the high mountains of Utah, Nevada, and California, in which also
the leaves have lutescent setae. It is at once distinguished from
S. watsoni, however, by the fact that the setae (which are even
darker) are not only 2 to 3 times as long but are strongly serrulate-
scabrous nearly throughout, those of S. watsont being smooth or
nearly so. The sporophylls also have longer and scabrous setae, and
the blades are much more freely ciliate, the cilia being stiff, very
oblique, and subpersistent nearly to the apex, in marked contrast to
S. watson.

VOL. 72, NO. 5, ‘Pl. 1

SMITHSONIAN MISCELLANEOUS COLLECTIONS

SELAGINELLA NEOMEXICANA Maxon

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72, NO-'5, PL.

SELAGINELLA EREMOPHILA Maxon

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72, NO. 5, PL.

SELAGINELLA ARIZONICA Maxon

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72, NO. 5, PL.

SELAGINELLA ASPRELLA Maxon

SMITHSONIAN MISCELLANEOUS COLLECTIONS

SELAGINELLA LEUCOBRYOIDES Maxon

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72, NO. 5, PL.

SELAGINELLA STANDLEYI Maxon

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 72, NUMBER 6

EXPLORATIONS AND FIELD-WORK OF THE
SMITHSONIAN INSTITUTION
IN 1920

(PUBLICATION 2619)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION

1921

The Lord Baltimore Press

BALTIMORE, MD., U. 8. A.

CONTENTS

PAGE
MGI HIs AGIAN era seyere gi tceeiny< SEKI viens «nies egoions aid Wei Me Seo hes REE eS Ss I
Geological Explorations in the Canadian Rockies. .. 2.2.02. ehe0ee. een es I
Geolocicalebield=VWorke in the! United Statesn 44-55 saeccaeceeeaer secre: 10
IFIIGEWWor eatin: WANS WE) O NGHCS) So gm Guin POO e eno puonomegouUEOOUn SoU as OUD 13
ACOORICA lesson sin AGGIES oooagocan0 b4docc0cKdobnne oon bbobooOnUauE Pye
Botanicale Collectio tmmeAhis Ca-mace ie sesee ee sene eereeie ciaine emia s ai
PNAS Urpallit Ati eRe pNE GE OM Ih se ce pcv ches si aleste tonas'atiear ot esa ses cave sich, Shane oka oem ees av doe lees aut « 39
Fiolosicalmixplorationmeine Elaine soca aco necro oe ae eceite es 43
Malacological Field-Work in California and the Hawaiian Islands....... ny)
lByoyeimicall IB Sqo lovreiniormatnl, |ekenteh\eHln waacoocaodag cones oude.e5 bn oc URS onGoeN oe 49
Botanical Easplonamon cineebritishn Giana ears oteecenieeces nae niace 54
Collections of Living Animals for the National Zoological Park......... 50
FAMiGO POLOcicalm xa editiOnmtOntdcmhatmMaSteea area ciae ae ca seacieiae cise 63
MM ee@ iby waste Viti eso beacermeyss «ta eerie ravers oe eee ache ise onn 8 ake cane ecko 75
Pield-Work onthe Mesa’ Verde National. Park. 22.05. s0046< cscs caste os 75
Picid-\onks Amons the bMiopiclndians: <<... 3... acess s.cee es vce oe ene oe 94
Archeological Investigations in Utah, Arizona, and New Mexico........ 06
aVinisiGuoiethnerbapacorand seawneeayaar hae aa ee oeecien oad oeerceecmanecn 102
OzankaCGavesrandsMoundsmne Missounlsoseeneeeenae seen one non ee: 107
mecheological Recounoissance in Hawai. «..<2..0.. sess 1s. bes we aas aes 110
Field-Work Among the Fox and Plains Cree Indians.................... III
mrcheological, explorations. im “Menmessees «havc oseccs-cu 2 se eariawdieness i)
Archeological Explorations in New Mexico «2. o:: s.c.¢ eon ac cece csccece: 120

Archeological Explorations in Eastern Texas

>, : Pee ee err ee ae

EXPLORATIONS AND FIELD-WORK OF THE- SMITH-
SONEAN INSTITUTION IN) 1920

INTRODUCTION

Investigation of the unexplored regions of the earth and extending
scientific knowledge of imperfectly known localities have from the
beginning been an important phase of the Institution’s activity in the
“increase of knowledge.” This pamphlet serves as an announcement
of the more important expeditions sent out during the calendar year
1920, and more detailed accounts of the scientific results are later
published in the Proceedings of the U. S. National Museum, Bul-
letins of the Bureau of American Ethnology, and other series of
publications issued under the direction of the Institution.

Not only has our knowledge of little known regions been increased
through the numerous expeditions sent out by the Institution during
the 75 years of its existence, but also the collections in natural his-
tory and anthropology in the Museum have been greatly enriched
thereby. The urgency of some of the field-work is illustrated by the
Australian expedition herein briefly described. The remarkable
Australian mammal life has been but meagerly represented in the
National Museum and the rapid extermination of the native animals
through various agencies renders increasingly difficult the securing
of an adequate collection. However very satisfactory shipments are
being received from the present expedition and it is hoped that a
good representation will be secured for the Museum before it is
too late.

GEOLOGICAL EXPLORATIONS: IN THE CANADIAN ROCKIES

During the field season of 1920, geological exploration was con-
tinued by Secretary Charles D. Walcott in the Canadian Rocky
Mountains, with two main points in view, (1) the determination of
the character and extent of the great interval of non-deposition of
sedimentary rock-forming material along the Front Range of the
Rockies west of Calgary, Alberta; (2) the clearing up of the rela-
tions of the summit and base of the great Glacier Lake section of
1919° to the geological formations above and below. The party
going from Washington consisted of Dr. and Mrs. Walcott and
Arthur Brown.

*Smithsonian Misc. Coll., Vol. 72, No. I, 1920, p. 15.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 72, No. 6.

Cascade Mt. (9,825)

Mt Pe (9, is

———_—— ats

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VOL. 72

OUS COLLECTIONS

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MISCELLANE

SMITHSONIAN

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SMITHSONIAN EXPLORATIONS, 1920

NO.

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VOL. 72

MISCELLANEOUS COLLECTIONS

SONIAN

SMITH

‘oz6r “ooTe@AA\ “q “DO Aq ydesSojoyg ‘AaTyea
uokues pedeys--) peosq & YSsno1y} ureyunoW oy} Jo JNO sMoy YOIYA ‘JOATY JSOYD) JO Vpls y}1OU 9y} UO puelpeay yeaI8 17 Mig |

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 5

Early in July work was begun along Ghost River northeast of
Banff and about 53 miles (85 km.) west of Calgary, Alberta. The
route taken was along the north shore of Lake Minnewanka (fig. 1,
Frontispiece) and through the Devils Gap to Ghost River, which here
runs north and south at the foot of high eastward-facing cliffs of
Cambrian limestone, capped by limestones of Devonian age. Lake
Minnewanka is a beautiful sheet of water (fig. 2) in the broad
bottom of a pre-glacial river channel, the eastward extension of which
forms the Devils Gap.

The Rocky Mountain front (fig. 3) is formed of masses of evenly
bedded limestone that have been pushed eastward over the softer
rocks of the Cretaceous plains-forming rocks. This overthrust is
many miles in extent and occurred long ago before the Devils Gap,

Fic. 5.—Devils Head (9,204 ft.), a butte rising above the cliffs of Fig. 3 on
the north side of Ghost River Gap. Photograph by C. D..Walcott, 1920.

Ghost River Gap and other openings were cut through the cliffs by
running water and rivers of ice. Great headlands (fig. 4) and high
buttes (fig. 5) have been formed by the silent forces of water and
frost, many of which stand out against the western sky as seen from
the distant foothills and plains.

It was among these cliffs that we found that the first great cliff
(figs. 3 and 4) was of lower Middle Cambrian age, and that resting
on its upper surface there were 285 feet (86 m.) of a yellowish
weathering magnesian limestone, here named the Ghost River forma-
tion, which represents the great lost interval between the Cambrian
below and the Devonian above. Sixty miles to the west, over four
miles in thickness of limestone, shales and sandstones (22,670 feet
(6,890 m.) ), occur in the lost interval of the Ghost River cliffs.

VOL. 72

MISCELLANEOUS COLLECTIONS

SMITHSONIAN

‘Oz61 JiONTeAA “q ‘D Aq
ydersojoyd “Yl JOMO] yj ur ueliquiey sradd~Q pure yyeausq syoo1 URITAOPIC, YUM 4YSI4 ay} UO a8prs ay} deo sauojsowy ueMOAaq
94 “Woes asino7y syeT FO Yj1OU Sols Tz “JOATY JoyeMAvaTD JO Aalyea 1addn JO pusd Usdjs¥a JY} 3e SID JO apis usJIsa\\—'9 “OLY

ee A

‘deo uviuo0aac ‘SHI Ueliqmuiey ssddy

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 7

es} ey ‘ b - eo 3 Ea ah ; % Se
Fic. 7.—A contented outfit on a Sunday afternoon near the head of the
Clearwater River. Photograph by C. D. Walcott, 1920.

SS aia fede ee ea

aber line (9,400 ft.) over Pipestone Pass.
Photograph by Mrs. Mary V. Walcott, 1920.

ea

Fic. 9.—Result of an hour’s fishing in Lake Minnewanka near Banff.
Photograph by Mrs. Mary V. Walcott, 1920.

Fic. 10.—The avalanche lily forces its way up through the hard snow, and
its beautiful slender green leaves and yellow flowers fairly cover the thin
outlying margins of the winter's snow. Photograph by Mrs. Mary V.
Walcott, 1920.

No. 6 SMITHSONIAN EXPLORATIONS, 1920 8)

Fic. 11.—Our pack horse “ Pinto” preferred to take a short cut at the ford
and went down in deep water and was dragged out. Getting ready to get her
up. Photograph by Mrs. Mary V. Walcott, 1920.

es ee Be oe ‘ etiam aE

Sia Ne a i: —_"

ed)

Fic. 12.—Getting “ Pinto” up on her feet in shallow water. Photograph by
Mrs. Mary V. Walcott, 1920.

iz) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Returning to the Bow Valley, the party left the Canadian Pacific
Railroad at Lake Louise and went north over Pipestone Pass to the
Siffleur River, which is tributary to the Saskatchewan. In the north-
ward-facing cliffs 25 miles (40 km.) east of the Glacier Lake section
of 1919, and 4o miles (64 km.) north of Lake Louise, a geological
section was studied that tied in the base of the Glacier Lake section
of 1919 with the Middle and Lower Cambrian formations. Return-
ing up the canyon valley of the Siffleur River to the wide upper valley
of the Clearwater River, a most perfectly exposed series of lime-
stones, shales, and sandstones of Upper Cambrian and later forma-
tions was found (fig. 6) which cleared up the relations of the upper
portion of the Glacier Lake section to the Ordovician formations
above.

The field season was marred by forest fire smoke in July and
August, and almost continuously stormy weather in September.
Some of the incidents of the trail are illustrated by figures 7-12.
The trout of Lake Minnewanka (fig. 9) increased our food supply
for days while on Ghost River, and the camp on the Clearwater was
a paradise for man and beast (fig. 7). On Pipestone Pass the
avalanch lily was found forcing its way up through the hard snow
(fig. 10), and in a treacherous ford of the Pipestone River, Arthur’s
war bag and the sugar and flour got a soaking.

The party is indebted in many ways to the officials of the Rocky
Mountains Park, and to the officials of the Canadian Pacific Railroad,
all of whom gave assistance whenever it was possible to do so.

GEOLOGICAL FIELD-WORK IN-THE UNITED STATES

Dr. R. S. Bassler, Curator of Paleontology, U. S. National
Museum, was engaged in field-work in Ohio and Illinois during the
latter half of June and the first part of July, with the result that two
large, instructive exhibits and important additions to the study series
of fossil invertebrates were obtained. Proceeding first to Northside,
Ohio, Dr. Bassler made arrangements for the shipment to Washing-
ton of a large, well-preserved fossil elephant skull which had been
purchased by the Museum through the efforts of Dr. E. O. Ulrich,
Associate in Paleontology. This specimen, discovered in glacial
gravels near Cincinnati some years ago, was long the prize exhibit of
a local saloon; in fact, it was so highly regarded for advertising
purposes that repeated offers of a considerable sum for its purchase
were invariably refused. With the coming of prohibition, its former
usefulness departed and the Museum was able to secure it for a

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 eit

nominal amount. Teeth of this species (Elephas columbi Falconer)
are contained in our collections, but such a complete skull is of rare
occurrence, there being only one or two others in North American
museums. The specimen is further valuable in giving evidence as to
the proper position of the tusks in the skull, a subject of long
controversy.

The second important exhibit secured during the trip was a slab,
measuring four by eight feet, of highly fossiliferous limestone from
the Richmond formation of Early Silurian age as exposed near

Fic. 13.—Beginning of excavation for exhibition slab of Richmond limestone
near Oxford, Ohio. Photograph by Bassler.

Oxford, Ohio. Such a specimen had long been desired for the
exhibition halls to show the advancement in life from the primitive
Cambrian forms, represented in the large Cambrian sea-beach sand-
stone exhibit, to the higher and more complex species of succeeding
geological periods, but notwithstanding the numerous occurrences of
fossiliferous limestone of Ordovician and Silurian age, it was not
until 1920 that a layer affording slabs of suitable size and sufficient
perfection of preservation was brought to the attention of the
Museum. This was discovered by Dr. W. H. Shideler, Professor

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

of Geology at Miami University, Oxford, Ohio, who most generously
assisted in quarrying out the specimen. As shown in the accompany-
ing photograph (fig. 13) representing the begining of the excavation
for the thin bedded, fossiliferous layer desired (marked -r), numerous
large blocks of stone had to be removed before the real task of
quarrying the slab was begun. The work was completed successfully
and the exhibit is now being installed in the hall of invertebrate
paleontology. The perfection of the fossil shell remains on this slab

iS i *

Fic. 14.—Surface of fossiliferous limestone slab, one twenty-fifth natural size.
Photograph by Bassler.

is evidenced in figure 14, which, however, shows only in a small degree
the distinctness of the shells upon the rock background.

Upon the completion of the quarrying operations at Oxford,
Dr. Bassler proceeded to Chicago, Illinois, where he was engaged in
the preparation of casts of type specimens of fossils contained in the
collections of the Walker Museum, University of Chicago. The
paleontological collection of the National Museum, which includes
the celebrated Walcott, Ulrich, Springer, Harris, Nettelroth, and
Rominger collections, is especially rich in type specimens of Early
Paleozoic fossils, but nevertheless the Walker Museum possesses

No. 6 SMITHSONIAN EXPLORATIONS, 1920 13

many unique types not represented at all in Washington. Permission
to prepare casts of these and thus advance our study series toward
the completeness which the National collections should attain, was
generously granted by Dr. Stuart Weller, Director of the Walker
Museum. In two weeks time Dr. Bassler was enabled to finish cast-
ing all of the Ordovician and Silurian types, leaving the remaining
Paleozoic species for a future trip. The work was done quickly by
using the modeling compound (plastocene) to make the mold from
which the cast is prepared. After dusting the fossil with talcum
powder the modeling compound is carefully pressed upon it and then
withdrawn, thus securing a clear-cut impression into which the plaster
is poured. Bubbles can be avoided by first pouring thin plaster of
Paris into the mold and distributing it uniformly with a camel’s hair
brush. The thicker plaster is then introduced as usual to fill the
cavity. When dry the modeling compound is torn away, thus leaving
the complete cast but also, unfortunately, destroying the mold.

Field-work in vertebrate paleontology was limited to a short trip
made by Mr. J. W. Gidley, Assistant Curator, in the latter part of
August to Williamsburg, Virginia, where scattered remains of a
fossil whale had been found in the Miocene strata outcropping nearby.
It was at first hoped that an entire skeleton could be secured here, but
careful search proved the bones to be so scattered and fragmentary
that no exhibition material was available although some interesting
additions to the collection of fossil vertebrates and some excellent
Miocene shells for the exhibition series were obtained.

Mr. William F. Foshag, of the Division of Mineralogy, at his own
expense made sundry trips into interesting mineral localities in Cali-
fornia and secured a considerable quantity of desirable material for
‘ the Museum’s collections, including an excellent series of borax
minerals.

FIELD-WORK IN ASTROPHYSICS

In astrophysical research the Institution was unusually active.
Early in 1920, Dr. Abbot had a long discussion and correspondence
with Professor Marvin, Chief of the United States Weather Bureau,
on the applications of solar radiation measurements to meteorology
now being officially practised in Argentina and Brazil on a basis of
daily telegraphic reports from the Smithsonian observatory near
Calama, Chile. Professor Marvin felt strongly the inadequacy of
existing solar radiation observations as a basis for studies of the
dependence of temperature on the solar variation. While the Chilean
results might be excellent, still they were for the most part not

2

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

checked by independent observations. The Mount Wilson work
yielded results on less than a third of the days, and might well be
affected by variations of atmospheric humidity incident to the site
so near the Pacific Ocean and the cities about Los Angeles. These
objections could not but be admitted by Dr. Abbot, and led him to
make a great effort to strengthen the observations of solar variation.

Mr. John A. Roebling, of New Jersey, had indicated a strong
interest in the work. In conference with Dr. Abbot in May, 1920, he
generously gave the sum of eleven thousand dollars for the purposes

oh ee ee

Be sare ea pM oR
i : he hs ay, ae :
vee eet betes, te ee
Piller SA PS
Fic. 15.—Montezuma solar observ- Fic. 16.—Montezuma solar observ-
ing station near Calama, Chile. ing station. Entrance to spectrobo-
Dwelling house, shop and garage. lometer tunnel. Also pyrheliometric
apparatus.

first, of removing the station theretofore on the plateau near Calama,
Chile, to a nearby mountain high enough above the plain to avoid
dust and smoke; second, of removing the “solar constant” outfit
from Mt. Wilson to the best mountain site available in the United
States; third, the balance for any other objects closely associated
with these investigations.

Under the zealous and able management of Director A. F. Moore,
a new observing station was selected and prepared at a place called
Montezuma, about 8 miles south of Calama on a mountain rising
about 2,000 feet above the local level, and about 9,500 feet above sea

SMITHSONIAN EXPLORATIONS, 1920 15

NO.

‘asnoy SuljamMq ‘BUINZI}UOW 3 UOI}e}S BurArasqo
“EUINZaJUOPY Je UOI}e}S BuUlALasqO AvJOS—oz7 “DIYy IeJOS dy} JOF eCwR[eD WoOIT sjersajeu Suljnep]—O6l1 ‘oy

"poyeoo] st A1OJLAJOSGO dy} YIM uO

yeod oy, “wore}s SurAtasqo 1ejOs euInza}JUOJY—'gI “OI

‘snyeiedde drajaworeyszdd pue
}BJSOJIOD) “UOI}LJS SUTATISqO ARJOS VUINZaJUO[—ZT “MJ

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

ae ie

Fic. 21.—Town of Wenden, Arizona. Mt. Harqua Hala in the background.

eA 7 eee Pao ‘ ule ee So ery

Re a fics ee SE a sane So mie.

Fic. 22.—Packing apparatus to Mt. Harqua Hala.

| me
|
\ * a a a

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 17

level. The instruments are arranged in a tunnel excavated hori-
zontally in solid rock near the summit of the mountain. Observers’
quarters, computing rooms, garage and shop are located some three
hundred feet lower in a sheltered ravine. These quarters can be
reached by the auto truck in 50 minutes drive from Calama. No road
had to be constructed, as there are no trees in the region, and a way
was found smooth enough, and of sufficiently easy grade, without
working. Within about two months of receipt of the Institution’s
telegram, Mr. Moore completed these arrangements, removed the

VE tw jill Y nw

Fig. 23.—Observatory on Mt. Harqua Hala.

outfit from its former location and recommenced observing August
5, 1920, at Montezuma with the loss of but ten days for the removal.
He regards the new site as excellent, and expresses doubt if a better
one could be found in the whole world.

In December, Mr. Moore returned to the United States, turning
over the Directorship of the Chile Station to Mr. Leonard H. Abbot,
formerly assistant, who is now assisted by Mr. Paul Greeley. Tele-
grams giving the values of solar radiation observed are sent as
heretofore to Buenos Aires and forwarded from there to Rio de

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Janeiro. Both the Argentine and Brazilian Weather: Bureaus employ
them with approval for forecasting purposes.

Dr. Abbot, with the advice and assistance of the U. S. Weather
Bureau, which carried on special observations in several localities for
the purpose, selected a site on Mount Harqua Hala, near Wenden,
Arizona, as the best place to relocate the Mount Wilson outfit. A
building, partly underground, was erected there in July and August
by local contractors. Messrs. Abbot and Aldrich occupied the Mount
Wilson Station as usual from July 1 to September 15, when the outfit

Maes ‘ o ae
its SEL ae at hall =

Fic. 24.—Coelostat and pyrheliometer, Mt. Harqua Hala.

was removed to Mount Harqua Hala, where observations were begun
on October 2, 1920. Dr. Abbot took charge and continued observing
until January 25, 1921, assisted by Mr. Fred A. Greeley. Mr. Aldrich
then relieved Dr. Abbot, and it is expected will himself be succeeded
about’ May 1, 1921, by Mr. A. F. Moore, formerly at Calama.

Thus the Institution has now in charge two first-class solar radia-
tion observatories, which are to be operated continuously hereafter
until the question of the value of the solar variation as a meteoro-
logical datum is definitely settled. Heretofore the measurements have

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 19

been secured on nearly 70 per cent of the days at Harqua Hala and on
nearly 80 per cent at Montezuma. The agreement of results on days
in common has hitherto been remarkably close and leads to the hope
of surely detecting solar variations as small as 1 per cent.

The conditions of living at Montezuma, while lonely, are not
excessively so. Frequent motor trips to the city of Calama for
supplies, and occasional visits to the copper mine at Chuquicamata,
where great kindness is experienced, help to break the monotony.
At Mount Harqua Hala, however, the isolation is excessive. There

cme RAK NN EOE SIRE nse eALERTS TRE ET . * _ ee

ee ee . ar oo eaenen

Fic. 25—Top of Mt. Harqua Hala after a snowstorm, showing fog-bank in
the background.

is a single neighbor, Mr. Ellison, a mining prospector located a mile
away, on whose three burros depends the transportation for the
observatory. It is 11 miles from Wenden to the foot of the mountain
trail, which is 5 miles more in rising about 3,000 feet. Mail 1s
received only about once in two weeks, when supplies are ordered by
heliograph signaling with Morse code to the merchants in Wenden,
at the cost of several hours hard work with the lights. Water must
be hauled from Mr. Ellison’s camp, over a mile distant and 850 feet
below, except when at rare intervals rain falls. The two observers

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

cook, wash, cut firewood from the small oak bushes and dwarf yuccas
about, and repair or alter the building or the apparatus as occasion
requires, besides carrying on the solar investigation.

During the occupation of Mount Wilson in 1920, many pieces of
research were successfully carried through by Messrs. Abbot and
Aldrich besides the measurements of solar variation. One of the
most interesting was the perfecting of the solar cooker begun several
years ago. A parabolic cylindrical mirror with polished aluminum
surface of about 100 square feet focuses the sun’s rays upon a
blackened tube filled with mineral oil communicating to an iron

Pe. “om

Wilson.

x

Fic. 26.—Solar cooker on Mt.

>

reservoir of oil in which are two baking ovens. A continuous circu-
lation of the heated oil keeps the ovens hot enough to perform all
cooking operations except frying. Excellent bread, meat dishes,
vegetables, cereals, canned fruits and vegetables and preserves were
cooked there by Mrs. Abbot, who had charge of this part of the
experiments and who was much envied for her cool kitchen and novel
appliance by the ladies of the mountain. This solar cooker was con-
structed on Dr. Abbot’s plans largely at the cost of grants from the
American Academy of Arts and Sciences in Boston, and the National
Academy of Sciences. It has proved successful, but must be regarded

No. 6 SMITHSONIAN EXPLORATIONS, I920 2.

at least for the present as rather a luxury for rural and relatively
cloudless regions, than as a generally useful appliance.

ZOOLOGICAL EXPLORATION IN AFRICA

The generosity of friends of the Smithsonian Institution made it
possible to engage Mr. H. C. Raven, well known for his previous

Fic. 27,—The Chimpanzee was one of the most inter-
esting animals in the forest. Their calling and shouting
could frequently be heard early in the mornings and on
moonlight nights. Adult female, Uganda, July, 1920.

work in Borneo and Celebes, to accompany the ‘‘ Smithsonian African
Expedition, under the direction of Edmund Heller in conjunction with
the Universal Film Manufacturing Co.,” which sailed from Brooklyn,
July 16, 1919, on the steamship City of Benares, and arrived in Cape

Town, August 13.

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

i BOR ee
ecured at Masindi, in Uganda, by jacking,
June, 1920.

ig be y
NP es 8
F)

Ps

Fic. 28.—A fine leopard s

seldom seen owing to its nocturnal habits. Budongo Forest, June, 1920.

SMITHSONIAN EXPLORATIONS, 1920

were encamped on the Kafue River about sixty miles above its confluence
with the Zambesi, in Northern Rhodesia, December, r1g1o.

Fic. 31.—The “ standard-wing night jar.” Masindi, Uganda, June, 1920.

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Le. Loe tie < Stee
Fic. 32.—Lates, a large perch of Lake Tanganyika.

wey

Fic. 33.—Kaffirs cleaning elephant skeletons at the camp of Major
Pretorius, in the Addo Bush. The South African Government has ordered
the destruction of these animals—the only herd of the kind in the world
today, living so far beyond the confines of the tropics, with the exception of
the few in the Knysna Forest. Kenkel Bosch, Cape Colony, September,

IQI0.

No. 6 SMITHSONIAN EXPLORATIONS, I920 25

Fic. 34—Wahutu men dancing at Nyanza on the shore of Lake Tan-

ganyika. Their voices, jingling bells on the ankles and stamping feet add
excitement to the scene.

Fic. 35.—Among the Wahutu who live on the coast of the lake and the
Watuzi of the mountains it is customary for a few of the best dancers to
come forward that their skill in jumping and whirling may be demonstrated
to better advantage.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

In the vicinity of Cape Town, Mr. Raven was able to collect only
insects and invertebrates, and from there he went to the Addo Bush,
where 19 days were spent in collecting small mammals and _ birds.
Going through Durban and Johannesburg, Mr. Raven spent two
weeks collecting at Ottoshoop in the Transvaal, after which he pro-
ceeded to Victoria Falls, and from there he left for the Kafue River

Fic. 36—A young chief of the Wahutu ready to
lead his men in the dance. Nyanza, Lake Tangan-
yika, February, 1920.

region, where he camped for several weeks. After spending some
weeks along the Congo, he reached Lake Tanganyika, where camp
was made for about a month. The next stop of any length was in
Uganda, where a few days over a month were spent in collecting in
the Budongo Forest. As the whole forest was in the sleeping-sick-
ness area, it was necessary to get a special permit from the district
commissioner to enter it, and the native boys had to be examined by a

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 27

ata el ad ed 20 beet ti

Fic. 37.—The Watuzi of the mountainous region northeast of Lake Tan-
ganyika in one of their very picturesque dances. Each man carries a long
lance or two, and a bow with one or more arrows.

Fic. 38.—W epairing the wall of a hut
with mud. The walls are made with wooden supports and between these are
placed rows of stalks of elephant grass partly buried in the ground and
fastened together with grass. The whole is then covered with mud and a

roof built so that it overhangs and protects the walls from the erosive effect
of rain.

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

doctor before entering the area and again on leaving it. Work here
was finished on July 14, 1920, after which Mr. Raven returned to the
United States, sailing from Cairo, September 2, and arriving in New
York, September 17.

Fic. 39.—A Dinka woman and her child at Shambe
on the upper Nile. It is a rather common sight to
see the natives cleaning their teeth with a bit of stick.
Sometimes they pound the end or split it with a knife
so that it becomes brushlike.

Though not numerically large the collections are of unusual interest
on account of the manner in which they supplement those obtained
by other expeditions in which the Smithsonian Institution has been
interested. Among the most important material may be mentioned
697 mammals (including 272 specimens from South Africa, a region
hitherto very imperfectly represented in our collections; 152 from

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 29

Fic. 40.—The Victoria Falls of the Zambesi River. Above the falls the
river is about a mile wide but drops over a cliff nearly four hundred feet
into a narrow gorge which in some places is less than one hundred and
fifty yards in width.

Fic. 41—A native village at Port Bell in Uganda, with Lake Victoria
Nyanza in the distance and to the right an acacia tree in which a colony of
weaver birds have made their nests.

3

30 SMITHSONTAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 42.—Primitive irrigating machinery, on the Blue Nile at Khartoum,
August, 1920. Oxen, donkeys or camels are used to turn such water
wheels.

*

Fic. 43.—Sheep grazing near the temple of Medina Habu in Upper Egypt.

‘No. 6 SMITHSONIAN EXPLORATIONS, 1920 31

Lake Tanganyika; the chimpanzee of Uganda), 567 birds, 206 rep-
tiles, and 193 fishes. The photographs here reproduced were all taken
by Mr. Raven, who has also prepared the legends which accompany
them.

BOTANICAL COLLECTING IN AFRICA

Dr. H. L. Shantz, Botanist, Office of Seed and Plant Introduction,
Bureau of Plant Industry, United States Department of Agriculture,
was also a member of the Smithsonian African Expedition, and his
chief objects were to secure live plants of agricultural value for intro-
duction into the United States, to study the agricultural methods of
both natives and Europeans, and to collect plants for the National
Herbarium of the United States National Museum. This work began
at Cape Town, August 13, 1919, and terminated at Port Said, Sep-
tember 2, 1920.

The vegetation of the Cape region has long been known to Euro-
pean botanists, and has received more attention than that of any other
portion of Africa. It is difficult to imagine a region which has so
many striking features. The wealth of Proteas and Ericas alone
would make this region unique. Lilies of various and beautiful types,
Iridaceae, Amaryllidaceae, and Orchidaceae, each represented by
many genera and species, the Arum lily or calla, which occurs every-
where in wet soil, and Pelargoniums, which here cover the mountain
sides, make of this Cape region a natural botanical garden, so rich and
varied that any botanist will here find plants of absorbing interest.
At Kirstanbosch there has been established the National Botanic
Gardens now under the direction of Dr. R. H. Compton. If the well-
laid plans are fully carried out, it will be possible to find here all of
the more interesting indigenous plants of South Africa, and to this
garden will come botanists, horticulturists, and agriculturists inter-
ested in South African plants. American botanists would be proud
if we had anywhere in our country a garden to compare with it.

East of Port Elizabeth in the Addo Bush, which is a low thorn
thicket of trees and vines about 15 to 20 feet high, are food plants of
especial interest. Among the succulents, none are more interesting
than the speckbroom (Portulacaria affra), a source of forage for the
wild elephant herd of the bush as well as for other large herbivora,
ostriches, cattle, sheep, or other domesticated animals. This plant
has been introduced into the United States and is doing especially
well in southern California in the lower chaparral zone and should
greatly improve the forage value of the natural range. Here are
many interesting plants such as the picturesque Acacia horrida, with

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

its large white spines, the Boer Bean, Schotia speciosa, the young
beans of which are cooked and eaten, also many interesting Aloes,
Lilies, Cotyledons, and Euphorbias. The elephants’ foot (Testu-
dinaria elephantipes) and many species of asparagus also occur here.

The Karroid plateau was seen only during the rest period, when its
vegetation, which consists of desert grasses and shrubs, is in appear-
ance identical with portions of the Great Basin in Nevada. Especially
interesting features of this desert are the great number of species of

i Rag?

Fic. 44——On the Karroid Plateau, west of De Aaar. This desert shrub,
known as Karroo bush (Pentzia), is a valuable forage plant, resembling our

bud sage of the Nevada and Utah deserts. Cattle, sheep and ostrich are the
chief grazing animals.

Mesembryanthemum, several of which are edible, and the Karroo
bush (Pentzia), a valuable forage plant, areas of which resemble in
general appearance our Bud Sage areas of Nevada and Wyoming.
Passing northwest to the region about Kimberley, there is a
scattered growth of Acacias, over an open desert grass type similar
in some ways to the vegetation of west Texas and portions of Arizona
and New Mexico. The high grasslands of the Transvaal, on the
other hand, with a grass vegetation dominated by Themeda forskalu,
reminds one of Andropogon scoparius areas in the drier portions of

No. 6 SMITHSONIAN EXPLORATIONS, 1920 33

our prairies. Here corn is the principal crop and the large ranches
with houses far apart, each with a clump of trees, make the resem-
blance to our prairies in the earlier days even more striking.

The portion of the Transvaal north of Pretoria, known as low
Veld, consists of a scattered growth of relatively small trees over a
grass cover composed of rather coarse tall grasses. This is one of
the most widely distributed types and an exceptionally interesting
one. The grasses are burned off each year and only such trees as can
resist the fires can maintain themselves under this condition. This
type becomes somewhat modified at Lourengo Marques, where the
vegetation, although quite luxuriant, still showed signs of a prolonged
drought period. Here, as in the region north of Pretoria, the Kaffir
Orange (Strychnos pungens) is abundant, and there are many species
of Acacia and Combretum. The Cashew Nut (Anacardium occi-
dentale) is everywhere a prominent tree and has the appearance of
being indigenous, although introduced from South America. It forms
a large evergreen tree and is never cut down by the natives, who prize
it both for the fruit and for the nut, from which they secure both food
and a strong alcoholic beverage. Another tree of unusual interest
is the Morula (Sclerocarya caffra), which has a valuable oil and
edible nut, with a fruit useful for making jam and an alcoholic drink.

Of the more strikingly beautiful trees of this section are the red
flowered and fernlike-leaved Delonix regia, the beautiful blue
flowered Jacaranda, and the Mahogany Bean (Pahudia quanzensis)
which produces in its large pods a large black bean with a brilliant
scarlet cup-shaped aril at the base and is one of the important timber
trees.

The vegetation at Salisbury reminds one of the low Veld above
Pretoria, a grassland with scattered trees, singly or often in clumps.
At Bulowayo the grasses are less luxuriant and the trees smaller and
more xerophytic. The vegetation about Victoria Falls, except for
the small forest irrigated by spray from the falls, is also xerophytic
in character. At Kafue the grasses seem more luxuriant but the
trees are much as at Victoria Falls. Here a number of important
fruits were secured, most of which are still undetermined. A nut
tree of unusual interest, because of its value for food and oil and the
remarkably light weight of the wood, was secured here. It has been
known as the Manketti Nut (Ricinodendron rautanenu). Here also
occurs the Beobab (Adansonia digitata), the largest tree in Africa,
useful to natives as a source of Bast fiber and as food. The acid
white pulp which fills the fruit and surrounds the seed is eaten or
dissolved in water to produce a refreshing drink.

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

The dry, open forest around Elizabethville gives way, before
Bukana is reached, to the tall grass and scattered trees which form
the great Savanna, which surrounds the tropical forest of the Congo.
The Lualaba is lined with oil palms (Elaeis guineensis) throughout
almost the whole of its course, and during much of the time swamps
of Papyrus are abundant along its course. Most interesting were the
great numbers of wild Sorghum grasses, some of which may prove

Fic. 45——Two large mango trees (Empe oribo) at Kigoma on Lake Tan-
ganyika. The trees bear two crops of fruit a year, a large crop in January and
February, and a small crop in August and September. The fruits are large,
fully five inches long, of excellent flavor and with practically no fiber around
the seed. The building in the back is the railway station.

valuable in our dry-land agriculture. The oil palm, which belongs to
the native who planted it, is probably the most useful native plant in
all Central Africa, and its oil is used by the natives as food and for
making soap with which to wash their clothes.

The dense tropical forests which cover much of the central Congo
were seen at Kindu on the Congo River, where they form a dense
canopy, but where the undergrowth is not entirely shut out. At
Kigoma and Ujiji, on Lake Tanganyika, the grassland is dotted with

NO. 6 SMITHSONIAN. EXPLORATIONS, 1920 35

large mango trees, the fruit of which is large and of excellent quality.
These trees were probably introduced by Arab slave dealers and

Fic. 46.—Watuzi chiefs dressed for the dance. They are, from
left to right, Kikovio (son of Ararawe), Mizambo (a Bagamwa
or prince of the fourth generation) and Ararawe (brother of
the head chief, Andugu).

The dress consists of an undergarment of bark cloth, made
from the bark of a fig tree, of two skins of leopard or serval
cat, one over the shoulder and one around the loins. Each carries
a long bow, decorated with banana fiber, two arrows and a long
spear.

They live largely on cattle, and inhabit the high mountain grass-
lands of Urundi. They are exceedingly tall, slender and athletic
men, are alert and pleasant and are almost untouched by white
influence.

merchants, and probably all originated as seedlings. Along the shore
of Tanganyika in Urundi many plants were collected. In the high-
land back of the lake at N’gano N’gano, the rolling hills are covered

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

with excellent grasses of high forage value, and support thousands
of cattle. The Watuzi who inhabit this region are one of the least
known but most interesting tribes in all Africa. They are tall,
slender, athletic men, with thin. lips and straight noses, and are

rw \ EE = OME SN LD See ae
Fic. 47—A grove of cocoanut palms at Dar es Salam, Tan-
ganyika Territory, with a native (Swahili) hut in the back-
ground.

The whole region about Dar es Salam and for fifteen or
twenty miles inland is occupied by an almost continuous cocoa-
nut palm grove. The notches in the trees facilitate the gather-
ing of the nuts.

probably the best athletes in the world, especially at high jumping,
which is one of their chief accomplishments. Many grasses and
many bulbous plants from this region give promise of value as plant
introductions. A great variety of bananas, beans, and cereals are
grown by these natives.

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 37

At Dar es Salam, a broad strip of coast is given over largely to the
cultivation of the coconut palm, although the streets are often lined
with wild figs, Delonix regia, and Terminalia catappa. Zanzibar is
given very largely to the cultivation of cloves, and here are also grown
many tropical fruits, and there is a small but exceedingly interesting
botanical garden. Tanga is somewhat similar to Dar es Salam in
vegetation.

Fic. 48.—Bank of the Victoria Nile at Masindi Port in Uganda. A luxuriant
growth of papyrus, water-lilies, morning glories and other water plants. Much of
Lake Kioga is covered with this type of vegetation.

es 4

While the vegetation of the north shore of Lake Victoria is rich
and varied and reminds one of that seen in the Congo and on Lake
Tanganyika, the outstanding features in Uganda are the areas of
elephant grass, Pennisetum purpureum, and the immense tracts of
bananas which extend for miles and miles, for bananas are relatively
as important to Uganda as corn is to Illinois.

There are several small tropical forests in Uganda, and at Entebbe
an unusually interesting botanical garden. The flame of the forest
(Spathodea), a prominent tree through much of tropical Africa, the

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

kiduri1 (Antiaris toxicaria), and many other trees. are prominent
here. At Entebbe one may see Para rubber, Cera rubber, Cacao,
sugar cane, yams, corn, rice, wheat, and other temperate crops har-
vested at the same time.

The principal results of the expedition were the collection of about
1,000 botanical specimens for the National Museum, and the introduc-
tion of about 1,600 plants for growth as agricultural plants in this

Fic. 49.—The Uaso Nyiro River at Archer’s Post. It flows through a_ semi-
desert country abounding in big game animals. It is lined throughout by
groves of Dum Palm.

country ; the more important being forage plants, nut plants, fruits,
and vegetables. The acquaintance was made of many men interested
in plants and agriculture, throughout the Continent, who can be of
service 1n connection with securing additional material. First-hand
observations were made of the methods of agriculture pursued by
African tribes as well as the Europeans, and a large number of photo-
graphs were taken which illustrate the natural vegetation and agri-
cultural crop methods.

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 39

AUSTRALIAN EXPEDITION

During the past year Mr. Charles M. Hoy has continued the work
of collecting material in Australia, under about the same conditions as
those mentioned in the last report on explorations. In this period
collections were made at the following localities :

“Farina, S. A.: Work in the Farina district was done at Lindhurst, 30
miles east of the town of Farina. Nineteen days were spent there, resulting
in the collection of 110 birds and 64 mammals. A few reptiles and insects
were also collected.”

“Kangaroo Island, S. A.: Twenty-six days were spent, in the field, on

*

Fic. 50.—Aboriginal grave yard, North Australia.

Kangaroo Island with the result of 85 mammals, 51 birds, and miscellaneous
reptiles, amphibians, and marine specimens collected.”

“Port Lincoln (Eyres Peninsula), S. A.: Twenty-two days were spent in
the field resulting in the collection of 86 birds and but 15 mammals. A
few miscellaneous specimens including reptiles, crustacea, etc., were also
obtained.”

“Busselton, W. A. (50 miles south): Camp was pitched 50 miles south
of the town of Busselton, on the edge of the Government Timber Reserve.
Forty days were spent in camp (May 14-June 23). The weather was the
worst that I have experienced. During the whole 40 days, there were only
three days free from rain. Over 18 inches fell in that time. It was impossible
to keep things dry and even the tent fly went green with mould. Despite
these handicaps, however, a pretty fair collection was obtained. The collec-
tion contains 94 mammals, 46 birds and a few miscellaneous alcoholic speci-
mens (reptiles and land shells).”

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 51.—A rufous rat-kangaroo from New South Wales.

Fic. 52.—Phascogale, a mouse-like marsupial from New South Wales.

No. 6 SMITHSONIAN EXPLORATIONS, 1920 4!

Ri +: s

geo a Z * *

- ef “sy - we x * . ~—> =e ” <
+S * -% oon ‘ - * Es, 4 ae, x
es . ‘ 3 ee | -

+ ? ee ae

Fic. 53—A young kangaroo. Photographed on Kangaroo Island, Australia.

Fic. 54.—An Echidna or “spiny anteater” photographed on Kangaroo
Island, Australia. The long bird-like beak projects from behind the plant
which conceals the animal’s face.

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

“Derby, W. A. (32 miles southeast): Twenty-three days were spent in
the above locality, August 7-29. The locality visited was very poor in both

mammal and bird life and a collection of only 43 mammals, 68 birds and 10
reptiles secured.”

Fic. 55.—A native of North Australia.

In the last letter received from Mr. Hoy, from Port Darwin,
Northern Territory, dated September 25, 1920, he writes, in part, as
follows :

NO. 6 SMITHSONIAN EXPLORATIONS, I9Q20 43

“T have decided to work the Northern Territory and then go overland to
Queensland via the old wagon road. I will travel in a buckboard and will go
south from here, to the Catherine River, then east along the Roper, and then
to the McArthur. On reaching the latter river I will go south along the river,
then to Anthony’s Lagoon and east across the tableland to some point in
Queensland—most likely Gloncurry. The trip will take about four months
and will have many advantages. The road is an easy one, being an old estab-
lished route with plenty of good water along the way, and the trip ought to be
productive of very good results.”

During the year two shipments were received from Mr. Hoy, the
last one arriving here in September. A total of 440 mammals, well
prepared, several of which were hitherto unrepresented in our collec-
tion, together with series of skeletal and embryological material ;
570 bird skins, with 24 additional examples in alcohol, and smaller
collections of reptiles, amphibians, insects, marine specimens, etc.,
were received.

Up to the time of Mr. Hoy’s visit to Australia the Museum had
received not over goo specimens of birds from that country; the
majority of these were old, mounted, and without precise data, some
of them dating back to the time of the Wilkes Exploring Expedition.
About 350 mounted birds, displayed at the Centennial Exposition in
Philadelphia in 1876, by the several Australian colonies, and pre-
sented to the National Museum at the close of the exposition, formed
the largest single contribution to our collection; the remainder of
the material consists of many smaller lots, obtained by gift or ex-
change at long intervals. About 140 species, including several of the
distinct forms of Kangaroo Island, are represented in the material
received from Mr. Hoy, all well prepared and with adequate data.
Some of them are quite new to the Museum collection, and others,
‘such as the lyre bird, have not previously been available in our study
series.

BIOLOGICAL EXPLORATION IN HAITI

Dr. W. L. Abbott, of Philadelphia, accompanied by Mr. E. C.
Leonard, of the National Museum, as botanical collector, made a
visit of exploration to southern Haiti from February to July, 1920.
Dr. Abbott undertook this visit chiefly that he might study the bird
life of Gonave Island, hoping to complete certain series he had col-
lected on a former visit. The island is about 30 miles long and Io
miles broad, and consists of a low mountain range bordered by a
belt of foothills that merge gradually on the north coast to a level
beach fringed by mangroves, but on the south slope descend rather
abruptly into the sea. Dense thorn thickets cover the arid mountain
sides and foothills, while the uplands, called La Table, open into

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

large grassy tracts with only occasional trees or shrubs. Due to the
scarcity of water and, in many places, to large outcrops of bare coral
rock, the island is poorly adapted to agriculture, but the grassy up-~
lands furnish excellent grazing. The forests yield a fair amount of
lignum vitae. The party spent three weeks on the north coast, work-
ing around Anse Galette and Etroite, and later a week on the south
coast in the vicinity of a small fishing village called Pikmi.

The regions visited on the mainland were St. Marc, Etang or
Manneville, Fond Parisien, La Mission, Furcy, and Pétionville, with
Port au Prince as base.

ian ex al “ot Se NBeae bu «
Fic. 56.—Royal Palm groves of Fond Parisien.

Etang is on the northwest shore of Etang Saumatre, a large lake
nearly 20 miles east of Port au Prince, bordered on the west by the
plains of the Cul de Sac, on the north by the mountains of Grand
Bois, and on the south and east by the foothills of La Selle Moun-
tains. The lake is about 50 feet below sea level and very salty.
Among the interesting natural features of the region is a series of
large springs, flowing into the lake through a belt of marshy meadows
which are covered by a short sod composed almost entirely of several
species of sedges. An excellent opportunity was offered to study
both the arid cactus forests and cat-tail marshes that occur in the
Cul de Sac.

The large royal palm groves of Fond Parisien, situated on the
southeast shore of Etang Saumatre, give it a picturesque appearance,

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 45

In old French times Fond Parisien was considered one of the most
beautiful spots in Haiti, but a flood has covered the greater part of
this once fertile district with stones and gravel, transforming it into
a desert. The large areas of black mucky swamp land to be found

1 is
yds f i
j yy
Boe j ld ; Ny ‘ . 3
Fic. 57.—“ Strangler plant” (a species of Clusia) on a
large leguminous tree near Pétionville.

here are utilized in sweet potato farming, while some cotton is grown
in the drier parts.

La Mission is a day’s journey to the south from Fond Parisien.
The most striking features of this region are the open pine forests

4

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

* _ af i Reh rd seat

Fic. 58.—Pic de Bromt (called “ Morne-cte-Weézan). A new species of grass
, was found growing on the summit of this mountain.

Fic. 59.—Weaver bird and nests in a thorn tree. Fond Parisien.

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 47

and occasional wet thickets, the latter sheltering a luxuriant growth
of ferns. The natives are, however, rapidly destroying the pine trees
both by burning the forests repeatedly and by girdling the pines for
splinters, which are sold in the markets as kindling.

Furcy and Pétionville are not far from Port au Prince. Both are
easily accessible and are among the few localities which have been
visited by naturalists.

Nearly 10,000 specimens of plants were collected, as well as a
number of land shells and insects.

The birds obtained by Dr. Abbott during this expedition numbered
201 skins, with a few alcoholic specimens and skeletons. By far the
most interesting ornithological observation made was the discovery
in some abundance of an introduced weaver bird, Hyphantornis
cucullatus (Muller), a native of West Africa. This species was
found at several points in Haiti, where it occurs in colonies and
affects much the same type of country as do related species in Africa.
Nearly completed nests, without lining, were found about the middle
of May, but no eggs had been deposited at this date. They are
strongly woven of narrow strips of palm or banana leaves, and
have an entrance at the side. An illustration of the manner of nest-
ing is shown in figure 59. Two smaller species of west African
weavers, belonging to other genera, are known to occur in Porto Rico,
where they have existed for many years, but the date and circum-
stances of their introduction, as well as those of the species dis-
covered in Haiti, are at present unknown.

MALACOLOGICAL FIELD-WORK IN CALIFORNIA AND THE
HAWAIIAN ISLANDS

On the way to the First Pan-Pacific Scientific Congress held in
Honolulu, August 2-20, 1920, Dr. Paul Bartsch, curator of mollusks,
U. S. National Museum, stopped for three days in Glacier National
Park, where some collections were made.

He also spent a day on shipworm investigation about Mare Island,
where he had placed at his disposal, by the commandant of the station,
a tug and pile extractor, and the necessary officers and men to make
every minute of his stay count, the result being a careful examina-
tion of pilings throughout the stretch of San Pablo Bay and the
adjacent shores of San Francisco Bay. This investigation resulted
in establishing the fact that the mollusk which has been doing the
damage estimated at some $25,000,000 last year is a new species of
Teredo, which Dr. Bartsch has named Teredo beachi in honor of the
commandant of Mare Island.

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE 72

Dr. Bartsch, in preparing a monograph on the land shells of the
Epiphragmophora californiensis group, was very much puzzled how
to account for the presence of two very closely related subspecies at
Pt. Pinos, California. He took this occasion to visit Pt. Pinos, and
there found that the two subspecies do not occupy the same habitat.
The smaller of the two was found in great abundance under plants
on two of the rocks lying off the Point, separated from the mainland
by a gap across which one could easily leap, while the larger sub-
species, which was rather rare, was found on the ground, buried
beneath needles and vegetable detritus. Two subspecies of the

Fic. 60—The home of Epiphragmophora californiensis Lea, a species of land
shells, off Pt. Pinos, California.

Epiphragmophora tudiculata group were found to occupy a similar
range.

Mr. Henderson and Dr. Bartsch arrived in Honolulu a few days
prior to the meeting, and this time was used for collecting land,
fresh-water, and marine shells on the island of Oahu. They also
collected mullusks during their sojourn about the wonderful crater
of Kilauea, on the occasion of the visit by the congress to the island
of Hawaii. Several stations were likewise made between Kilauea and
the Kohala coast. A large number of marine shells were secured
from the rocky shores of Honaunau Bay.

Since the first accommodations to be secured for the return were
dated September 8, the intervening time between the close of the

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 AQ

congress and this date was spent in collecting specimens. Dredgings
were made in Pearl Harbor, where the commandant placed one of
the dredges at their command. They were also rendered the neces-
sary assistance in making a search for shipworms, with the result
that a new species of Teredo was discovered, which has been named
Teredo parksi, in honor of Admiral Parks, in charge of Yards and
Docks, U. S. N. They also dredged in Maunalua Bay, on the south
side of Oahu and in Kaneohe Bay, on the east side of the same island.
Specimens were also collected on various occasions at Haleiwa Beach,
on the north end of Oahu, and the beach and shallow water adjacent
to their cottage at Waikiki were thoroughly scratched over. Trips
were also made into various parts of the mountains, where land shells
of many kinds were secured.

Another excursion carried them to the island of Maui, where
marine shells were collected wherever possible along the shore, and
land shells were secured on their ascent of the magnificent extinct
voleano of Haleakala.

One of the very interesting observations made on this trip to the
Hawaiian Islands was the finding of an existing marine flora and
fauna at a considerable elevation above the level of the sea on the
gently sloping bench at the southeast point of Hanouma Bay. This
flora and fauna consist of algae, quite a number of species of mollusks,
crustaceans, echinoderms and other marine organisms, which occupy
pools and puddles kept ever moist and supplied with fresh water by
the spray from the breaking surf, which incessantly pounds that
shore. Dr. Bartsch considers this an important observation, since the
occurrence of fossiliferous laminae bearing marine organisms be-
tween sheets of lava has been held to indicate that they were deposited
‘at or below sea level and their occurrence above this has been held as
evidence of elevation. We have here an instance which indicates
that this is not necessarily the case, for such a lamina would be pro-
duced if a new outpouring of lava were to cover up the place
mentioned.

BOTANICAL EXPLORATION IN JAMAICA

In February, 1920, Mr. William R. Maxon, Associate Curator in
the Division of Plants, United States National Museum, and
Mr. Ellsworth P. Killip, aid, were detailed to make botanical col-
lections in Jamaica. The expedition was made possible largely
through the co-operation of the New York Botanical Garden, the
Gray Herbarium of Harvard University, the Field Museum of
Natural History, the University of Illinois, the Arnold Arboretum,

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 62.—Residence at Cinchona.

NO. 6 SMITHSONIAN EXPLORATIONS, I920 51

Fic. 63—Higher peaks of the Blue Mountains as seen from the southwest;
Mossman’s Peak (unexplored) at the left, separated from Blue Mountain
Peak (2,225 meters) by Portland Gap.

Fic. 64.—In the heart of the Blue Mountains. The denuded areas are land-
slips due to erosion in areas long under cultivation in coffee.

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

and Mr. Oakes Ames in contributing to the field expenses of the
work. Two months were spent in the island, and upwards of 10,000
specimens were brought back, representing about 1,700 collection

Fic. 65.—Fern-covered bank on trail to Morce’s Gap. The
tree ferns in the center are Cyathea pubescens.

numbers. The material has been shared among the contributing

institutions, the ferns and flowering plants having already been fully
identified.

NO. 6 SMITHSONIAN EXPLORATIONS, 1920

3

on

From headquarters in Kingston field-work was carried on in several
widely separated parts of the island, the courtesy of free transporta-
tion on the Jamaica Railway having been extended by the colonial
government. Through the kindness of Mr. J. G. Kieffer, general
manager of the United Fruit Company in Jamaica, a house on the
company’s plantation at Windsor, in the rich banana region south of
Port Antonio, was placed at the disposal of the party. About 10
days was spent here, trips being made to Mooretown, Mill Bank,
Cuna Cuna Pass, and the northern foothills of the John Crow range.

> Se ae

Fic. 66.—A characteristic fern (Dicranopteris bifida), growing in vinelike
masses near Morce’s Gap.

The most productive period was one of three weeks in March,
spent in the Blue Mountain region, with headquarters at the botanical
station at Cinchona, the lease of which had been renewed by the
Smithsonian Institution in January, 1920. Located on a projecting
southern spur at an altitude of 1,500 meters, equipped with a service-
able laboratory and most comfortable living quarters, Cinchona served
as an excellent base for botanical exploration in the Blue Mountain
region, most of the peaks lying within fairly easy reach. Extensive
collections were made on John Crow Peak, at New Haven Gap and
Morces Gap, and in the vicinity of Cinchona, as well as on trips to
Thompson’s Gap, Hardward Gap, and the summit of Blue Mountain

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Peak. Over 800 numbers were collected in this region. In prepara-
tion for a projected work by Mr. Maxon on the ferns of Jamaica,
particular attention was paid to this group, nearly 200 species being
here collected.

Other regions visited include Hollymount, on the upper slopes of
Mount Diabolo, in the central part of the island, the coastal territory
of Montego Bay, and the southern border of the “ Cockpit Country,”
the last a wild, little inhabited, wooded area of innumerable limestone
sinks lying southeast of Montego Bay. Collections at Cook’s Bottom,
Mulgrove, Mocho, and Ipswich yielded a number of new or otherwise
very interesting species. Just before leaving Jamaica Mr. Maxon
made a brief visit to Pigeon and Great Goat islands, lying off Old
Harbour.

Much of the success of the trip is due to assistance extended freely
by officials of the United Fruit Company, acknowledgment of which
is gratefully rendered.

BOTANICAL EXPLORATION IN BRITISH GUIANA

Mr. A. S. Hitchcock, Custodian of Grasses, visited British Guiana,
making the trip through the co-operation of the United States De-
partment of Agriculture, the New York Botanical Garden, and the
Gray Herbarium. He left New York, October 4, 1919, and arrived
at Georgetown, October 22, stopping on the way at St. Thomas,
St. Croix, St. Kitts, Antigua, Guadeloupe, Dominica, Martinique,
St. Lucia, and Barbados. He left Georgetown, February 2, 1920,
and arrived in New York, February 16, stopping four days in Trini-
dad and one day at Grenada.

Six weeks were spent at Georgetown, and other points along the
coast; three weeks at the Penal Settlement on the Mazaruni River,
from which were visited Bartica, Kalacoon, and Kartabo; two weeks
on a trip up the Demerara, Essequibo, and Potaro rivers to Wismar,
Rockstone, and Tumatumari; and ten days on a visit to the North-
western Distict, including Morawhanna on the Barima River, Isso-
rora, the Rubber Station on the Aruka River, and the Yarikita Police
Station on the Venezuelan border.

Four sets of plants were collected including 1,134 numbers, with
extra sets of the grasses. On account of the extremely damp climate
it was necessary to use artificial heat in drying the specimens. Two
oil stoves were kept burning night and day, the specimens being
between corrugated paper. After drying, the plants were sprinkled
with naphthalene powder to prevent subsequent molding.

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 55

British Guiana has an area of about 90,000 square miles, is about
400 miles deep, and extends about 250 miles along the coast (Lat.
1°-8° N., Long. 57°-61° W.). There are three counties: Demerara,
including the drainage system of the Demerara River and to the
Abary River on the east; Essequibo, including the drainage system
of the Essequibo River and all to the west; and Berbice, including

Fic. 67.—St. Thomas. The wall is covered with coralita
(Antigonon leptopus), an ornamental vine with handsome
racemes of pink flowers. The palms are royal palms.

the drainage system of the Berbice River and east to the Courantyne
River. Georgetown, at the mouth of the Demerara River, has a
population of about 60,000. The only other city is New Amsterdam,
at the mouth of the Berbice River, with a population of about 9,000.
The entire population of the colony is about 300,000, about 4,000 of
whom are whites. The bulk of the population consists of East

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Indians (42.7 per cent) and negroes (39 per cent). The chief
product of the colony is sugar, which, together with the by-products
rum and molasses, constitutes about three-fourths of the exports.
Other products are gold, rice, balata, timber, and cattle. The country
is fairly healthy, the death rate being about 35 per 1,000 (14.8 per
1,000 among whites). Yellow fever is absent, but malaria and dysen-
tery are prevalent.

Fic. 68.—Antigua. An old sugar mill. These old stone
mills, formerly used for crushing the cane, are common on
the islands. Oxen were attached to the long beam. The
crushing or grinding is now done in modern sugar factories.

The rainfall at Georgetown is about 90 inches, with a dry season
September to November, and another short dry season in the spring.
The temperature at the same place is, in the daytime, about 88° F.
in the summer, falling four to six degrees at night, and in winter
four to six degrees lower. The temperature at night in winter rarely
falls below 75° (the minimum record for 35 years is 69°).

No. 6 SMITHSONIAN EXPLORATIONS, 1920

oF

The coastal region of the colony for 10 to 40 miles inland is a flat

plain scarcely above sea level. Back of this there is a belt with hills

7 ee
Fic. 69.—Georgetown, British Guiana.
rain tree (Samanea saman).

One of the main streets with rows of

Fic. 70—Morawhanna, British Guiana. A typical village of the interior.

as much as 200 feet high. Toward the southwest the land rises

and at the Venezuelan-Brazilian border culminates in the famous
Mt. Roraima, a table mountain rising to a height of 8,500 feet. With

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

the exception of the savannas of the southern part of the country
(Rupununi District) and a few other small areas, the whole country
is covered with dense virgin rain-forest.

Communication in the interior is almost entirely by boat along the
numerous streams, but unfortunately is hindered by falls and rapids
above the influence of the tide (30 to 60 miles).

OE > yA Me ES, eee

Sad . Wat” >. » ‘ia ee a. ‘a

Fic. 71.—A giant mora tree (Dimorphandra excelsa) in
the virgin forest of British Guiana, near Tumatumari. The
base is broad and buttressed. Attached to the trunk is a
species of Marceravia.

The scientific activities of the colony are mainly under the control
of Prof. J. B. Harrison, Director of Science and Agriculture, who
extended to Mr. Hitchcock many courtesies.

The Jenman Herbarium, an important collection of British Guiana
plants, is at the office of the director in Georgetown. There is an
excellent botanical garden with a large collection of trees and shrubs,
including a very fine series of palms.

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 9

on

COLLECTIONS OF LIVING ANIMALS FOR THE NATIONAL
ZOOLOGICAL PARK
As in former years, friends of the Smithsonian Institution, while
on expeditions abroad, collected and sent to Washington interesting
living animals for the National Zoological Park.
Mr. W. J. La Varre, who has before made collections of this kind,
visited South America and explored the upper waters of the Amazon.

Fic. 72.—Animals for the National Zoological Park awaiting shipment at
Manaos, Brazil. Photograph by La Varre.

He left the United States in June, 1919, and traveled directly to
Manaos, Brazil, the jungle metropolis about 1,000 miles up the
Amazon at the mouth of the Rio Negro. From Manaos he ascended
the Rio Negro, by small steamer and launch, into Venezuela. He
found much of interest, but no trace of the so-called “ cannibals ” of
the region, or of anything savage, either beast or man. The natives,
poor, half-breed rubber gatherers—Spanish, Portuguese, Negro, and
Indian mixtures—treated him cordially and most hospitably, sharing
their rude homes of thatch with him and giving him as much of their
food as they could possibly spare. Six months were spent with these

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

i

Fic. 73.—Girls of the Rio Negro, Brazil. Photograph by La Varre.

Fic. 74.—Native Batalao of rubber gatherers, Rio Negro, Brazil. Photograph
by La Varre.

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 61
kindly, primitive people, in geographical and zoological research, ex-
ploration, and hunting.
The general scarcity of mammal life along these famous rivers was
co) my, co)

surprising. So many rubber gatherers live here that the country has,

wad

CF eet NN cml Sg wan,

Fic. 75.—Border marker between Brazil and Venezuela, with govern-
ment officials of both countries. Photograph by La Varre.

in fact, been hunted out, and the natives themselves are often hard
pressed for food. Pacas and peccaries were plentiful, and numbers
Tapir tracks were crossed several

were shot for use in the camp.
natives.

times, and small deer were occasionally killed by the

3)

62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Curiously enough, only two wild monkeys were observed, although
examples of various species were commonly seen in the possession of
natives. Birds, on the contrary, were plentiful—among the con-
spicuous species were toucans, curassows, macaws, and _ parrots.
Mr. La Varre left the jungle in February, and arrived in the United
States near the end of March, 1920. His collection of living animals,
which he presented to the park, included a specimen of the rare black-
headed ouakari monkey (Cacajo melanocephalus), a species never
before represented in the collection. This monkey is a member of the

TBO SS GORE HAE BGT pai
Fic. 76.—White-backed Trumpeter (Psophia leucoptera) from the Rio Negro,
now in the National Zoological Park. Photograph by La Varre.

only genus of short-tailed monkeys inhabiting the New World, and is
very seldom seen in captivity.

Other animals in the La Varre collection are capuchin and squirrel
monkeys, an ocelot, two margay cats, egrets, a scarlet ibis, and a
number of parrots and paroquets. A large living specimen of the rare
and curious matamata turtle, a gift to the park from Mr. A. T. S.
Hore, of Manaos, was brought home with the lot. Another American
resident of Manaos, Mr. Edward B. Kirk, also contributed some
interesting birds, among which was a fine specimen of the very rare
white-backed trumpeter (Psophia leucoptera) from the Rio Negro.

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 63

A second collection of living Neotropical animals was brought to
the park by Dr. William M. Mann, of the Bureau of Entomology,
United States Department of Agriculture, who visited Honduras in
the spring of 1920. Dr. Mann’s duties in other lines naturally
absorbed most of his time, but he succeeded in landing in good con-
dition a number of valuable animals. His collection included pacas,
agoutis, kinkajous, squirrels, a mantled howler monkey, and some
reptiles, among which was a specimen of Rossignon’s snapping turtle,
a species rarely taken by collectors.

Other valuable animals were collected and presented to the park
by Hon. Henry D. Baker, American Consul at Trinidad, British West
Indies, and by Mr. Isaac Ellison, of Singapore, Straits Settlements.
Mr. Ellison succeeded in landing and placing in the park a thrifty
young male orang-utan, three years old. This is one of the most
interesting and valuable gifts received in many years. The animal
has now become thoroughly adapted to his new home and promises
to become .a most unusually attractive addition to the collection.

The National Zoological Park also shared in the large collection of
African animals collected and brought to America for the New York
Zoological Society by Mr. A. K. Haagner, director of the National
Zoological Gardens at Pretoria, South Africa. Included in the lot
received at Washington are a lechwe antelope and a specimen of the
Rhodesian baboon, recently discovered and described by Mr. Haagner.

ANTHROPOLOGICAL EXPEDITION TO THE FAR EAST

Under the auspices of the Smithsonian Institution and in connec-
tion with the Rockefeller Foundation and the Peking Union Medical
-College, Dr. Ales Hrdli¢ka made an extensive trip to the Far East
during the first half of 1920. The objects of this trip were continua-
tion of the studies relating to the origin of the American aborigines ;
examination of the oldest skeletal and other human remains in Japan ;
the furthering of the interests of physical and medical anthropology
in China; and a personal visit to the rapidly disappearing full-blooded
Hawaiians. The countries visited included Japan, Korea, Manchuria,
northern China, the boundary of southern Mongolia, and the islands
of Oahu and Hawaii in the Hawaiian Archipelago.

In Japan especial attention was given on one hand to the physical
characteristics of the people, and on the other to the prehistoric
anthropological collections. The latter have by now assumed con-
siderable importance. They are deposited in the universities and
medical schools of Tokio, Kyoto, Sendai, Osaka, and Kumamoto,

64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.

Se

Fic. 77—Average types of Japanese children. Photographs presented by
Mr. Tsunawo Araki.

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 65

and are being steadily added to by new explorations in prehistoric
mounds and shell heaps. They date all from the Neolithic period,
nothing earlier having thus far been discovered in eastern Asia; and
they show both an old diversity, as well as more or less relation to the
Aino and to the Japanese.

cd
iA
7
5
+

Fic. 78.—Japanese child.

In Korea special facilities were obtained for visiting the museum
at Seoul, which was found unexpectedly rich in Korean and Tur-
kestan antiquities. The people represent quite a distinct subtype of
the yellow-brown stem from that of the Japanese as well as that of
the Chinese; they are more like the western Siberian or southeastern
Russian Tatars.

One of the most interesting features in Korea are the mound burials
of the people. These mounds are all hemispherical, in contradistinc-

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 79.—Of the best in Central Japan. Photographs presented by
Mr. Tsunawo Araki.

NO. 6 SMITHSONIAN EXPLORATIONS, I920 67

tion to the conical mounds of Manchuria and northern China. They
are of various sizes; they occur singly, in clusters, and in whole
“ cemeteries "’; they are frequently assiduously cared for, and many
are most picturesquely located on the slopes of hills, where they ap-
pear to the best advantage. The Koreans are a modern race of mound
builders. The country is full of archeological remains, including some
big mounds and dolmens, and deserves much closer scientific atten-
tion than it has so far received.

Fic. 80 —A Korean Hamlet. From a few houses to large villages, they
cluster in the nooks of the hills like mushrooms.

The Manchus and the northern Chinese, particularly those of the
Chihli Province, are for the most part tall, well-built people, quite
different in bearing and even in physiognomy from the southern
Chinese, though there is no sharp delimitation. They, too, present a
fruitful field for detailed anthropological investigation.

The southern or inner Mongolians were found to be a rather
mixed lot, more so than the northern Mongolians who were visited
by Dr. Hrdlicka in t912. A series of photographs was secured here
as well as in Korea and Japan. In Japan, through the kind help of
Dr. Tsunawo Araki, there was obtained a large collection of portraits

68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLi72

of Japanese children, while the Tokyo Dental College, through the
good offices of Professor Mitsuru Okada, contributed several hun-
dreds of portraits of its students.

China, for the anthropologist and archeologist, is one vast open
museum, in which something of absorbing interest 1s met with at
almost every step. In addition the people, and particularly the students
in colleges, impress one with their native abilities. China in the
future may well be relied upon to give the world many a philosopher
and scholar of distinction.

Fic. 81.—Mounds near a Korean village. At sunrise on special days it is
not unusual to find before each mound a prostrate figure in devotion before
the grave of a father or mother.

The stay at Peking was principally devoted to assisting in the de-
velopment of the medico-anthropological work at the Union Medical
College, and the organization of the “Anatomical and Anthropological
Association of China.” The college has an excellent staff of well-
trained young workers such as Drs. Cowdry, Black, Howard, and
others, the majority of whom are seriously interested in those branches
of anthropology which are nearest the medical sciences ; and there are
bright prospects for anthropological work in other parts of China,
due to the presence there of English-speaking (mostly American)

ell te

ae mtg ii

Fic. 83.—A young matron of a good family, with baby and servant, at
Sen Sen, Korea.

70 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

medical missionaries, who are naturally interested in investigations
of that nature. ;

While at Peking, certain investigations were carried on, with the
help of Professor Davidson Black, on Chinese teeth ; and a conference
was held with the representatives of several Chinese ministries, as
well as of the Chinese Geological Survey and the Peking Union

Fic. 84.—A Korean girl.

Medical College, on the subject of the foundation at Peking of a
“Museum of Natural History of China,” the establishment of which
would mean so much for the progress of the Chinese themselves, and
for facilitating the work of foreign men of science in the Chinese
Republic. There exists already, under the direction of Dr. V. K.
Ting, a very creditable geological museum, which could serve as a
nucleus of the more comprehensive institution.

No. 6 SMITHSONIAN EXPLORATIONS, 1920 71

Fic. 85-The surgical class at Severance College, Seoul, Korea,
Dr. A. I. Ludlow operating.

PSG Sete ae

Fic. 86—Chunhuzes. A rare photograph of the North-Mongolian brigands.

72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

On the return trip a stop was made at Hawaii, and with the kind
assistance of the staff of the Bernice Pauahi Bishop Museum at
Honolulu, and particularly of Dr. Gregory and Mr. Stokes, some
studies were carried out on the native Hawaiians in several of the
most favorable localities. It was found that the Hawaiians, even

Fic. 87.—Chinese woman with artificially de-
formed feet, at Peking. This crippling deforma-
tion is still extensively practised by the Chinese.
A Chinese woman without deformed feet is not

“cc

regarded as “comme il faut.”

where free from historic admixture with whites or negroes, present
an old blend of several ethnic elements (yellow-brown, Indo-Euro-
pean, and Negro or Negrito); this blend, however, has already
reached a degree of approach to physical unity which permits the
student to deal with it as with a racial subvariety or subtype. But
this subtype is rapidly vanishing through new contacts.

No. 6 SMITHSONIAN EXPLORATIONS, 1920

NI
no

Fic. 88 Full-blood Hawaiion:

ae i.
Fic. 89.—Full-blood Hawaiian.

74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 90.—Full-blood Hawaiian.

~~. * * eo ¢@,

a” J

Fic. 91.—Full-blood Hawaiian.

No. 6 SMITHSONIAN EXPLORATIONS, 1920 75

THE OJIBWA OF MINNESOTA

During the fall of the past year, Dr. Ales Hrdli¢ka was called once
more to Minnesota to assist the Department of Justice in settling the
subject of mixed bloods and pure bloods among the Ojibwa. (See
Smithsonian Explorations for 1917.)

The whole work, now concluded, presents a good illustration of the
practical value of anthropology in certain directions. The results are
outlined in the following extracts from a letter to the Institution from
Mr. R. C. Bell, Special Assistant to the Attorney General :

The Department of Justice in 1910 instituted approximately 1500 suits in
the United States Court for the District of Minnesota involving Indian lands.
This litigation to January, 1916, has cost the government a very large sum
and little had been accomplished.

The blood status of the Indians became the determining factor and it was
ascertained that all the available evidence (principally testimony as to
genealogy) on this issue was found unreliable and always unsatisfactory.
Consequently, the Department of Justice, in the spring of 1916, procured the
services of Dr. Hrdlicka to make a physical examination of the Indians and
report his conclusions as to their status. The doctor spent more than three
months in this work and examined approximately 800 individuals.

The Department of Justice, the Department of the Interior, and counsel for
the defendants agreed to accept his findings as a basis for settlement, and
the litigation now has been terminated, resulting in the recovery of more than
$1,000,000 in land and money for the Indians; besides, the cost of the work
since 1916 has been insignificant in comparison to the cost prior to that time.

Furthermore, a commission was created by an Act of Congress to make a
roll of the allottees of the White Earth Reservation. The law required, in
addition to much other information, that the blood status of the Indians be
given. This roll, which affects the title to approximately 725,0000 acres of
land, has been completed; and the commission in preparing it followed the
findings of the doctor. Only those thoroughly familiar with the situation can
‘appreciate the far-reaching importance of this work.

FIELD-WORK ON THE MESA VERDE NATIONAL PARK

The Chief of the Bureau of American Ethnology, Dr. J. Walter
Fewkes, continued his field-work on the Mesa Verde National Park,
Colorado, during June, August, and September, 1920, in cooperation
with the National Park Service of the Department of the Interior.
Excavation and repair work was done on the Fire Temple Group and
Oak Tree House, ruins in Fewkes Canyon and at Cedar Tree Tower,
situated about a mile north of Spruce Tree House, leading to im-
portant contributions to our knowledge of the culture of cliff dwellers.

He was ably assisted in this work by Mr. J. A. Jeancon, who made
the originals of the ground plans of the Fire Temple Group and
Cedar Tree Tower here published.

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MITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

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Fic. 92—Map of Sun Temple Area in the Mesa Verde
National Park, Colorado.

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 77

The opening of Sun Point Road along the south rim of Fewkes
Canyon (fig. 92) is a most important addition to the park from the
tourist’s point of view. For about a quarter of a mile along this road
one can look down into Fire Temple (fig. 94) and Fire Temple
House, Oak Tree House (fig. 103), and the two ruins under Sun
Temple, the walls of one of which, Willow Tree House, are in the
same condition as when the cliff dwellers left the mesa; the other,

Fic. 93—Ladders into Fewkes Canyon from Sun Point Road, opposite Fire
Temple. Photograph by G. L. Beam. Courtesy of the Denver and Rio Grande
Railroad.

Mummy House, on a lower level, is notable for its fine masonry. On
the point across Fewkes Canyon, rise in full view the walls of Sun
Temple, and beyond it, nestling in the cliff, is the magnificent Cliff
Palace. The fine cliff dwelling Sunset House (‘‘ Community
House’’), a prominent ruin in Cliff Canyon, is likewise conspicuous.
The road along Fewkes Canyon from which these prehistoric build-
ings are visible has already become a very popular drive, being only
about two and a half miles from Spruce Tree Camp. The accom-

6

78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

panying map (fig. 92) shows the relative position.and approaches to
the above mentioned ruins. A fine trail, one of the most beautiful in
this area, was developed under the north rim of Fewkes Canyon, to

Fic. 94.—Fire Temple from Sun Point Road. Photograph by G. L. Beam.
Courtesy of the Denver and Rio Grande Railroad.

connect the above ruins. Three ladders (fig. 93) placed in the cliff
opposite Fire Temple enable the traveler to reach this trail, which
ends in ladders on the canyon rim opposite Cliff Palace near the point

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80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

where the discoverers stood when their eves first beheld this most
impressive cliff dwelling of the Southwest.

The most important discovery of the past season was the determina-
tion by excavation that a large cliff house was not a dwelling, but
specialized for some communal purpose. This ruin (fig. 98) was
devoted to fire worship, a cult which up to this year, although sus-
pected, had not been demonstrated as existing among cliff dwellers—
at least no specialized building for that function had been recognized.

In 1915 there was discovered on the promontory opposite Cliff
Palace a building with semicircular ground plan that was interpreted

. sei

Fic. 96.—Western end of Fire Temple Court, half excavated. Photograph
by Fewkes.

as a specialized religious structure and called Sun Temple. While
work on it was in progress a reconnoissance was made of cliff houses
situated in the canyon below, to one of which was given the name
Painted House. Like Sun Temple, it was suspected of having also
been dedicated to some religious cult of the cliff dwellers, but its true
significance was not apparent until the spade of the archeologist last
season verified this suspicion and revealed its true purpose. Painted
House, upon excavation, proved to be one of the most exceptional
cliff ruins yet recognized in the Southwest. The results of the exca-
vation have led Dr. Fewkes to designate it Fire Temple, and by

No. 6 SMITHSONIAN EXPLORATIONS, 1920 81

implication to regard it a temple of the eternal fire. Attention should
be called to the importance of the discovery that the cliff dwellers
had a New Fire Cult and possibly that rites of new fire and conserva-
tion of the same existed among prehistoric people of the Mesa Verde.

The rites of kindling the new fire among the descendants of the
cliff dwellers, as the Hopi, occur in July and November and are
known as the Lesser and Greater fire ceremonials. The act in both
is performed by means of a fire stick or drill made to rotate in a

Bae

Fic. 97.—Eastern end of Fire Temple Court. Photograph by G. L. Beam.
Courtesy of the Denver and Rio Grande Railroad.

notched board ; the same kind of fire sticks have been found in Spruce
Tree House, Square Tower House, and elsewhere.

Probably it is to the Lesser Fire ceremony at the East Mesa of the
Hopi that we should look for the nearest survival of the cliff dweller’s
rite, as in it we find the personation of a phallic being, Kokopelli,
whose picture was well preserved up to a few years ago on the wall
of the secret chamber of the Fire Temple where fire was created.
This Lesser New Fire, called Sumykoli, is celebrated by a fraternity
of fire priests, now extinct, known as the Yaya priesthood. The Yaya
priest at Hopi carries in his hand during this ceremony a rattle of

72

VOL.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

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SMITHSONIAN EXPLORATIONS,

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NO.

84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

exceptional construction. Two of these, now in the museum of the
Brooklyn Institute, were found in a cliff dwelling in the Chelly
Canyon, which seems to indicate that there formerly existed among
the cliff dwellers of that canyon a fire priesthood like the Hopi Yaya.
As the cliff dwellers of Chelly Canyon and those of the Mesa Verde
were closely related it is a fair conclusion that the latter also had a
well-developed New Fire Cult, and possibly a Yaya priesthood.

The masonry of Fire Temple points to an early epoch in the evolu-
tion of the Mesa Verde culture, possibly one contemporary with the

Fic. 101—Western end of Fire Temple Court. Photograph by G. L. Beam.
Courtesy of the Denver and Rio Grande Railroad.

first settlement in Cliff Palace cave, but anterior to the erection of the
unfinished Sun Temple, which marks a later or culminating phase of
cliff house development. Architectural features that Fire Temple
shares with Sun Temple may be interpreted by the close relationship
of fire and sun cults among the cliff dwellers. Earth Lodge A, exca-
vated in 1919, is the ancient type in the evolution of buildings on the
mesa antedating stone walls, and there are evidences of successive
stages illustrating cultural epochs from the crude Earth Lodge A,
which the earliest colonists constructed, to those of horizontal ma-

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 85

sonry (fig. 108), the highest products of the mason’s craft in pre-
historic North America. This evolution may have occurred on the
area now reserved as a national park, but was not limited to it.

After the abandonment of Fire Temple and the desertion of build-
ings of the culminating epoch that followed, people of like culture
may have still inhabited the great pueblos at Aztec and in the Chaco.
But these in time also succumbed and were deserted before the arrival
of the white man. Their descendants were amalgamated with nomadic
or non-pueblo peoples and their survivors still inhabit the modern

Fic. 102.—Bins for grinding corn in upper cave of Fire Temple
House. Photograph by J. A. Jeancon.

pueblos along the Rio Grande. Both blood and culture suffered
changes in this mixture, and architectural features remain to espe-
cially indicate the modifications. The Hopi, Zufii, and modern Rio
Grande pueblos have no specialized buildings like Sun Temple nor
Fire Temple for sun or fire cults, although they have ceremonial rooms
where they formerly kindled the new fire annually. They no longer
conserve the fire in this room, but there are legends that they did so
in former times, pointing to a remote cultural connection between the
cliff dwellers and their modern survivors, the Pueblos.

86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

The general form and arrangement of rooms at the east and west
ends of the central court of Fire Temple differ from any cliff ruin
or pueblo in the Southwest. Significant facts show that the building
(fig. 97) 1s unique, as will readily appear from the following state-
ments. Excavations in other cliff houses of the same size reveal
household utensils, as pottery, and other domiciliary objects. No
cooking places, grinding bins, or household implements were found
in the extensive work at Fire Temple. Moreover, every cliff house

of size on the Mesa Verde has one or more specialized gener-

Fic. 103 Oak Tree House from Sun Point Road. Photograph by G. L. Beam.
Courtesy of the Denver and Rio Grande Railroad.

ally circular subterranean rooms for ceremonies. There were no
kivas in Fire Temple. But the one exceptional positive feature in this
ruin that separates it widely from the cliff dwelling is stronger than
these negative evidences. In no cliff house, and indeed in no pueblo,
do we find a similar large circular fire pit filled with ashes in the center
of a rectangular court. This structure would seem to be the key to
the meaning of the whole building. That great fires were once built
in this fireplace, as the abundant ashes indicate, no one can doubt.
This fireplace is too large for an oven for culinary purposes and
although we know that the cliff people sometimes cremated the dead
the absence of calcined human bones would disprove the theory that

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 87

it was used as a crematory. A natural conclusion would be that it
was constructed for ceremonies connected with fire.

The significance of the two low-walled rectangular enclosures
one on each side of the central fire pit, is unknown. Each was par-
tially filled with soil and ashes when excavated and it is suggestive
to record that the stratum of earth above them as well as the whole
surface of the hardened floor of the court was filled or covered with
charcoal and burnt brush. Great fires must have raged over the

Fic. 104—Cedar Tree Tower before excavation. Photograph
by JeeAs seancom:

whole court in addition to that in the fireplace, after the temple was
deserted.

We find several other facts that fit in very well with the interpreta-
tion that this building was a fire temple. On the east and west ends
of the court (figs. 97, 101) there are banquettes, the former evidently
seats for those who watched the ceremonial performance in the court.
There are niches in the rear wall where possibly sacred objects may
have been placed; a wall of the cliff bears triangles and zig-zag paint-
ings, symbols of sex life ; but, most important of all, on the wall of one
of the rooms at the west end of the court there are paintings in red,

88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

one of which, now erased, represented a phallic being still associated
with New Fire ceremonies among the Hopi, while others represent
fire itself. At the New Fire ceremony among the Hopi one of the
four fraternities that celebrate it is the order of Horn Priests, who
wear on their heads imitations of the horns of mountain sheep. A
large number of paintings of mountain sheep cover the walls of the
west room of Fire Temple where fire was supposed to be kindled.
The massive walled buildings at the east and west (fig. 94) ends
of the court formerly reached to the roof of the cave, and although
two stories high no wooden beams for flooring occur in their con-

ee phe,
e's We! re ACA oo 2

Fic. 105—Cedar Tree Tower after excavation. Photograph by J. A. Jeancon.

struction. Wherever a lower story existed it was filled in with rubble
on top of which was laid an adobe floor. On the white plastering,
which is well preserved, there are numerous figures in red, mostly
triangles and symbols of lightning ; female and male symbols similar
to those in sacred rooms.

Just west of Fire Temple there is a group of rooms from which
utensils were excavated. In the floor of one of these rooms is a
vertical shaft which opens outside the house walls like a ventilator.
The former use of this structure is unknown.

Although Fire Temple was not inhabited there were undoubted
dwellings nearby. A hundred feet east of it there are two low caves,
one above the other, in which may have lived those who once made

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 89

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Fic. 106.—Below, Cedar Tree Tower and kiva; above, ground plan of
Cedar Tree Tower and kiva. Photograph by G. L. Beam. Courtesy of the
Denver and Rio Grande Railroad.

go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL: (72
use of this sanctuary. Dr. Fewkes has called the ruin (fig. 98) situ-
ated in these caves the Fire Temple House and supposes it was the
habitation of the fire priests and their familzes. The rooms in the
lower cave were fitted for habitation, and it had two, possibly three,
circular ceremonial rooms ; but the upper cave (fig. 99) is destitute of
kivas. The large rooms of the upper house (fig. 100a) may have
been granaries for storage of provisions, although possibly some of
its rooms were inhabited. In the rear of the large rooms identified
as granaries was found a small room with a well-preserved human
skeleton accompanied with mortuary pottery. One of these mortuary
offerings is a fine mug made of black and white ware beautifully dec-
orated; a specimen worthy of exhibition with the best in any mu-
seum. In the rear of the cave were three fine grinding bins (fig. 102),
with metates still in place.

The upper house 1s now approached from the lower by foot-holes
in the cliff and a ladder shown in the illustration. Evidences of
a secondary occupation of kivas in the lower house appear in double
walls and those of crude masonry without mortar, forming a rec-
tangular room built diagonally across the room. The plastering on
the rear walls of the lower house is particularly well preserved, but
there are very few rooms in addition to the kivas. One of the kivas
has in place of a deflector and ventilator shaft a small rectangular
trench enclosed by a well-made wall, as in Sun Temple.

Work was done on the large cliff ruin, Oak Tree House (fig. 103),
on the trail an eighth of a mile east of Fire Temple Group. Three
new kivas were excavated to their floors and the walls repaired, add-
ing to the four already known. No signs of these buried kivas were
visible when work began. All these kivas show fine masonry ; after
the most easterly had been used as a sanctuary for a time it was
abandoned and five well-preserved grinding bins were set in the floor
so as almost to conceal it. The upright slabs of stone and metates of
these structures were reset, showing fine examples of these prehistoric
mills. Many other novel features were brought to light in the exca-
vation of Oak Tree House, which may be regarded as one of the
most instructive ruins of the park. One of the exceptional features
of Oak Tree House is a fragment of a circular wall in the rear of the
cave, made of willow and other sticks set in mortar, like what is called
“ stick and adobe ” construction in other regions.

The ground plan of one of the kivas is semicircular and shows a
rectangular room on the straight side communicating with the cham-
ber by means of two passageways. The ventilator opens directly into
this room, whose function 1s unknown.

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 gli

In one of the grinding bins there was excavated a bundle of grass,
Koeleria cristata, of exactly the same form as the brushes with which
Hopi maidens sweep their metates after grinding meal; one more
resemblance between cliff dweller and Hopi customs.

In his classic on the “ Cliff Dwellers of the Mesa Verde,” Baron G.
Nordenskiold figured and described a nameless ruin designated a
tower (fig. 104), situated in the cedars about a mile north of Spruce
Tree House. To this ruin the author has given the name Cedar Tree

Fic. 107.—Section of floor of Cedar Tree Tower, showing
ceremonial opening. Photograph by J. A. Jeancon.

Tower, on account of an ancient cedar tree (fig. 105) hanging over
the top of the north wall. Nordenskiold closes his brief description
with the remark, ‘‘ Perhaps it should be regarded a religious building.”

The desirability of testing this surmise of the talented Swede led
the author, in August, 1920, to excavate this tower and the area about
its base, which led to the discovery that although it appeared to stand
alone there were two subterranean rooms connected with its base
situated on the west and south sides. The larger of these rooms
(fig. 106) had all the structural features of a typical kiva of a Mesa
Verde cliff dwelling. This subterranean structure, its floor excavated

Q2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

in solid rock, was circular with pedestals for supports of a roof, the
beams of which were absent, and had a central fire hole, ventilator,
and deflector. It communicated with the tower by a subterranean

Fic, 108.—Square Tower House. Photograph by G. L. Beam. Courtesy of
the Denver and Rio Grande Railroad.

passage which bifurcated, one branch opening through the tower floor,
the other into a square room situated on the southwest side, also
subterranean, partially constructed under a large rock forming a

SMITHSONIAN EXPLORATIONS, 1920

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O4. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

veritable cliff chamber. In the middle of the solid rock floor of the
tower which served as the foundation of the tower walls a circular
hole or sipapi (fig. 107), symbolic of the entrance to the underworld,
had been drilled, affording evidence that the tower was used for cere-
monials, A distant view down Soda Canyon may be had from the top
of the tower, although it is situated some distance from the rim of the
mesa and shut in by a dense growth of cedars and pinyons.

An automobile road constructed around Cedar Tree Tower was
continued through the cedars to join the Mancos road. Several
ladders were placed in position and a trail opened down the steep wall
of Soda Canyon from Cedar Tree Tower to Painted Kiva House, an
instructive cliff dwelling about a quarter of a mile away, formerly
practically inaccessible. To the west of the Mancos road about the
same distance from Spruce Tree House as Cedar Tree Tower there
is another tower of the same type, but with walls of adjacent rooms
projecting above ground. Several other similar towers have been
reported on the mesa, in the Mancos, McElmo, and Hovenweep Can-
yons, and elsewhere. The relation of a tower to kivas and other
buildings of Square Tower House is shown in figure 108.

A preliminary examination was made of the ruin at Aztec Springs
now called Yucca House National Monument, in the Montezuma
Valley, with a view to future excavation and repair of this important
site. As no satisfactory photograph of this ruin has ever been pub-
lished a view of the Lower House of this ruin taken in 1874 by
W. H. Jackson is by his permission given in figure 109. It is planned
to begin work on the Lower House of this great ruin in the spring
Or 192%

FIELD-WORK AMONG THE HOPI INDIANS

Dr. Walter Hough, curator of ethnology, U. S. National Museum,
spent the month of June among the Hopi Indians of Arizona, a tribe
with which he has been associated for 25 years. At present some
of the Hopi tribes are making rapid progress toward assimilating the
culture of the white man, while others; though becoming more and
more affected, show changes to a lesser degree. As these changes
have taken place through peaceful assimilation and were not forced
by war or other disruptive agency they present an interesting field
for ethnological research on normal modifications of social struc-
tures due to contacts. Some notes on this subject are appended.

MATERIAL WELFARE

The Hopi have prospered during the last quarter of a century.
From the period when they knew almost nothing of money the Hopi

No. 6 SMITHSONIAN EXPLORATIONS, 1920 95

have advanced to a complete familiarity with United States currency.
Purchases can be paid for with check. Some of the more advanced
have bank accounts. The Hopi were always acquisitive and frugal,
and their habits have put them in a rather good financial condition.
Their agricultural products have had a better market and the prices
are less subject to the former fixed low valuations of the local trader.

SoctaL CHANGES

The most effective cause of social changes was the enforced atten-
dance of children at government schools and the sending of children
to distant Indian schools. Causing considerable friction at first, this
has now been accepted in most cases as advantageous.

A result not prevised is the weakening of the clan discipline whose
former regulative authority was silent but powerful. The weakening
of this authority produced much laxity for a time and it was found
necessary to appoint policemen and a “ judge.” The government
found it necessary to send away girls not noticeably under parental
discipline to schools until they should become discreet. This measure
was quite opportune and had a beneficial effect.

Decay oF NATIVE CEREMONIES

In this connection there is ensuing a rapid decay of native cere-
monials. The heaviest loss to the native cult is felt by the death of
the old men of a former generation. Very shortly there will be no
one of the tribe who has not had training in school and more and more
of these will evade the call of the native religion.

Some fraternities have suspended operation and others are very
weak. Notably the woman’s harvest ceremony at Walpi has passed
out and the Snake Society there is about to discontinue.

CHRISTIANIZATION

Missions to the Hopi were carried on from time to time during a
period of several hundred years without appreciable results. When
the disintegration due to government control had progressed mis-
sionary efforts began to bear fruit.

The Baptist mission at Polacca has a stone church and a consider-
able and growing number of converts. Other branches of this mission
are doing well.

A slight cleavage is observed between the Christians and non-
Christians but no friction. The Hopi are agreed to let things work
out as they will.

96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

ARCHEOLOGICAL INVESTIGATIONS IN UTAH, ARIZONA, AND
NEW MEXICO

During May and early June, 1920, Neil M. Judd, curator of Ameri-
can archeology, United States National Museum, continued his arch-
eological reconnoissance of the arid region north of the Rio Colo-
rado, Arizona, in behalf of the Bureau of American Ethnology.
Attention was chiefly directed to the Toroweap Valley and several
caves in an extensive lava flow on the east slope of Mt. Trumbull ; to
the unwatered mesa known as Pariah Plateau ; to certain portions of
House Rock Valley not previously visited’ and to the upper two-

pe © : t - s
a a

Fic. 110.—Ruin surmounting a circular butte of white sandstone near the
eastern rim of Paria Plateau and not far from the Rio Colorado. Note the

upright slabs in the foreground—a frequent feature of prehistoric dwellings
in this region.

Be we oS ee EE a

thirds of Bright Angel Creek. In addition, a number of caves in
Cottonwood Canyon and Kanab Creek, Kane County, Utah, were
examined for evidence of ancient habitations.

In Bright Angel Creek several open ruins and three groups of cliff
houses were inspected; a complete survey was found impossible on
account of unexpected high water which prevented access to the
narrow, walled-in portion adjoining the Grand Canyon of the
Colorado.

In contrast to the conditions which were encountered in this
beautiful gorge, lack of water and forage for pack animals seriously

* Smithsonian Misc. Coll., Vol. 70, No. 2, 1918.

No. 6 SMITHSONIAN EXPLORATIONS, I9Q20 Q7

Fic. 111.—Cliff dwellings on the west side of Bright Angel Creek, above
the mouth of Beaver Creek. Through the doorway of the right-hand room
will be seen a bench, made from a cottonwood log, which forms an unusual
feature of the ruin.

Fic. 112.—A group of small storage cists near several ancient ruins at the
foot of the upper falls, Ribbon Falls Canyon, Bright Angel Creek. The in-
accessible cliffs surrounding this section of the canyon form an amphitheater

whose narrow entrance was guarded by a second fall over one hundred feet
in height.

98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

restricted investigations in the other sections above noted. The
hurried inspection of the Mt. Trumbull region was made in an auto-
mobile, it being impracticable to use horses in this parched country
during the summer months. Exposed pueblo ruins bordering the
east rim of Pariah Plateau and overlooking the Painted Desert proved
more numerous than was anticipated. In size, in arrangement of
rooms and in the quality of their masonry these ancient dwellings are
superior to those seen elsewhere in northwestern Arizona; likewise,
potsherds examined at each site exhibit greater perfection of form
and a higher decorative technique. These two factors—architectural

Fic. 113.—Part of a cliff village in a cave on the east side of Cottonwood
Canyon, near Kanab, Utah. The village includes nineteen rooms and a nearby
spring furnished excellent water for the ancient inhabitants.

and ceramic remains—alone are sufficient to connect the former in-
habitants of this region with the pre-Puebloan peoples east of the Rio
Colorado and to warrant the expectation that additional investigations
will disclose the approximate points at which the Colorado River was
crossed in ancient times.

Following his researches for the Bureau of American Ethnology,
Mr. Judd proceeded to New Mexico as director of an archeologic
reconnoissance of the Chaco Canyon National Monument. This sur-
vey was conducted under the auspices of the National Geographic
Society and had for its prime object close examination of the
aboriginal remains in the above monument with a view toward selec-

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 99

tion of a possible site for exhaustive investigation and permanent
repair.

The Chaco Canyon National Monument was created by presi-
dential proclamation March 11, 1907, and includes 18 major ruins
of very great significance in the study of ancient Pueblo life. Unlike
the vast majority of cliff houses and other ruins found elsewhere,
each of these huge buildings was constructed along preconceived plans

Fic. 114.—Grand Canyon of the Colorado from the lower
Toroweap valley, southeast of Mt. Trumbull. Evidence of
prehistoric habitations were not lacking in this region but
the dwellings were widely separated and poorly preserved.

and as a community enterprise. The studied arrangement of their
rooms and the perfection of their masonry rank them as the very
finest examples of prehistoric architectural accomplishment in the
United States. Not only did the ancient inhabitants of Chaco Canyon
excel as builders with stone, but the lesser objects, found in and about
the great communal dwellings, show that they had attained remarkable
skill as makers of pottery, ornaments and implements of various

IO0O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 115.—The north wall of Pueblo Bonito, viewed from the northeast.
This is the largest and justly the most famous of all the Chaco Canyon ruins ;
it covers nearly three acres of ground and some of its walls still stand to a
height of forty feet. When occupied Pueblo Bonito probably contained as
many as 800 rooms, sheltering over 1200 individuals.

Fic. 116.—The north-central portion of Pueblo Bonito, from the southeast,
showing its position relative to the north wall of Chaco Canyon. Sections of
fourth story walls appear in the above illustration. Like most of its neigh-
boring ruins, Pueblo Bonito was constructed in a series of terraces overlooking
a central court or plaza; the outer wall was pierced by small windows above
the first floor but had few doorways.

No. 6 SMITHSONIAN EXPLORATIONS, 1920 IOI

Fic. 117.—A portion of Kimmenioli ruin, showing the present condition of
its walls. About 135 ground floor rooms are still traceable; originally the
building was three, perhaps four, stories high. Although one of the finest
in the national monument this great communal dwelling is not so well known
as some of its neighbors in Chaco Canyon which is situated some ten miles
to the north and east.

Fic. 118.—Pueblo del Arroyo, occupying an insecure position on the very
edge of Chaco wash, 300 feet west of Pueblo Bonito. Flood waters have
exposed the remains of an older dwelling beneath the walls of the larger
structure and now threaten the latter. View from the north.

102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

sorts. The beauty and technique of such specimens warrant the belief
that these canyon dwellers had reached a plane of esthetic as well as
social development not surpassed elsewhere in the Southwest.
Although the 18 major ruins constitute the main feature of this
national monument there are, in addition, hundreds of “ small
house” remains scattered throughout the entire Chaco Canyon dis-
trict. In culture as well as in architecture these lesser dwellings were
closely allied to the larger structures. Talus pueblos and small cliff
houses are to be found along the north side of the canyon; elsewhere
the mud walls of a “ pit house” have been disclosed—a crude, semi-

Fic. 119.—Part of Pueblo Pintado, as seen from the northwest. This ruin
was visited by Lt. J. H. Simpson, August 26, 1849, on his memorable advance
into the Navaho country; the large timbers noted by Simpson and other early
explorers have since been torn from the walls, causing obvious destruction to
the latter.

subterranean shelter—older than any of the other habitations yet
observed in this region. Certain it is that comprehensive investiga-
tions in the Chaco Canyon drainage will add largely to present knowl-
edge concerning the prehistoric pueblo peoples of the southwestern
desert country.

MUSIC OF THE PAPAGO AND PAWNEE
In February, 1920, Miss Densmore went to the Papago Reserva-
tion in southwestern Arizona to continue her study of Indian music
for the Bureau of American Ethnology, residing for more than four

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 103

weeks at a government station called San Xavier Mission. The
Papago tribe was selected partly because of its desert habitat (fig.
120), the intention being to compare the phonographic records of
Papago songs with those of Arabian songs obtained from Arabs who
were temporarily in Washington, D. C. (Subsequent results proved
the value of this comparison.) According to the last census there are
7,465 Papago Indians on the reservation, but not one “ mixed-blood ”
family. It is said further that there has never been any intermarriage
between this tribe and Mexicans or Spanish. Their manner of life is
becoming modified, but many primitive customs remain and were
observed. A primitive burial place was found by Miss Densmore.

acini

Fic. 120.—Habitat of Papag

o Indians. (Photograph by Miss Densmore.)

These burial places were constructive on the side of a mountain and
consisted of low walls of rocks, roofed with timber and tightly closed
with stones. Bodies were removed after a time to make room for
other burials. A skull and a few bones remained in the tomb exam-
ined.

The subjects studied were: (1) Songs used in treating diseases
caused by spirits of dead Apaches and Papago; (2) songs connected
with the “ purification ” of returned warriors who had killed Apaches,
and (3) songs connected with dreams, games, and dances. Musical
instruments formed a subject of special investigation. A native
flageolet was obtained (fig. 12) together with the tradition concerning
its origin. The music of this instrument was phonographically
recorded and has been transcribed. The Papago beat upon an over-

104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 121.—Papago playing on native flageolet. -

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 105

turned basket instead of a drum, striking the basket with the palms
of one or both hands. “ Rasping sticks” are sometimes used with
such a basket, as shown in figure 122.

Fic. 122.—Papago and native musical instruments.

After a brief stay in Phoenix, Arizona, Miss Densmore went to
Camp McDowell (formerly Fort McDowell) and was present at a

106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

gathering of Mohave Apaches, explaining her work and securing
their consent to record songs on a subsequent visit. The adobe build-
ings of the old fort are in ruins, but a few of the smaller and more
substantial buildings remain.

From Arizona Miss Densmore went to Pawnee, Oklahoma, arriving
April 12, a few days before the Morning Star Ceremony. This is
one of the most important ceremonies of the year as it is held for the
purpose of securing good crops. At this time the “ Morning Star
Bundle” is opened and its contents displayed for several hours, while
the proper rituals are sung. Miss Densmore was allowed to enter
the lodge for a brief time and to view the sacred articles (it is said

Fic. 123.—Pawnee lodge of Morning Star Ceremony, (Photograph by
Miss Densmore.)

only one other white person has been accorded this privilege). Dur-
ing the remainder oi the ceremony, which lasted many hours, she
stayed outside the lodge (fig. 123) and make manuscript notes of the
songs. An approach to two-part music, heard at this time, had not
been previously observed. While at Pawnee a sufficient number of
songs was recorded to complete the musical study of that tribe.

In November, 1920, Miss Densmore returned to Arizona to resume
work among the Papago. The principal work was done at Vomari, a
point near the Mexican border and 80 miles from the railroad. Inter-
esting material was collected also at Sells, formerly known as Indian
Oasis. Among the subjects studied were: (1) The Papago expedi-
tions to the Gulf of California for salt and for “ medicine power,”

No. 6 SMITHSONIAN EXPLORATIONS, 1920 107

each sort of expedition having its songs; and (2) the rain-making
ceremonies, including the manufacture of tizwin. In connection with
the latter a visit was made to Santa Rosa, at the extreme north of the
reservation, where a tizwin camp and lodge were photographed. A
specimen of the cactus syrup used in making tizwin was obtained, as
well as a large basket which had been used in serving this wine.

Many sites of legendary or geographic interest were photographed,
Miss Densmore travelling more than 360 miles by auto on this trip.

The most important result of this expedition was the hearing of a
form of three-part music at a Papago dance. This was said to be a
native musical custom. So near an approach to polyphonic music has
not been hitherto observed by Miss Densmore and the subject will
receive further investigation.

OZARK CAVES AND MOUNDS IN MISSOURI

During the summer of 1919 the work of cave exploration in the
Ozark region was continued by Mr. Gerard Fowke, for the Bureau
of American Ethnology. Almost his entire time was given to a
thorough examination of two large caves in Pulaski County. The
first, known as Miller’s cave, is three miles northeast of Big Piney
postoffice. The opening is in the vertical face of a high cliff fronting
Big Piney River, with a steep talus slope beginning 30 feet below the
floor of the cave and extending to the water’s edge. The perpen-
dicular wall below, with a projecting ledge which forms the roof,
prevents a direct entrance, and the interior can be approached only
through another cave whose opening is in a ravine near by. A narrow
passage, barely large enough to admit a man in a crawling or crouch-
ing position, connects the two, and it is only through this that access
can be gained to the main cave. The inmates were absolutely safe
from molestation, as one man could defend this opening against any
number. A little stream flowing along the foot of the east side of the
cavern ensured a supply of water at all times: game was plentiful in
the neighborhood: the river abounded in fish; and fertile, level
bottom lands, easily cultivated, on either side of the stream furnished
much corn and other farm products.

A ditch and enbankment across an isthmus guarded a peninsula
on the opposite side of the river, and on both sides low house mounds
and abundant debris furnished proof of two large village sites.
Whether there was any connection between the villages and the cave
dwellers cannot be determined.

A bed of clean, pure ashes whose depth ranged from 3 to 64 feet,
according to the irregularities of the clay, was found in the cave

108 SMITHSONIAN MISCELLANEOUS. COLLECTIONS VOL. 72

reaching from wall to wall, a width varying from 45 to 70 feet. This
bed was so loose as to be almost like a snow bank; but for the most
part they were as compact as if much trampled over while wet. When
solidly packed, the mass would measure fully 800 cubic yards in
volume; but when loosened by excavation, 200 cubic yards more.
All the wood had to be carried from either the top or the bottom of
the hill, which is about 400 feet high, and passed through the small
opening from the other cave. It is safe to say no more fuel would be
used than was strictly necessary. When it is considered how little
fire is requisite for the needs of an Indian household, and that the
limited space suitable for residence would not provide sufficient room
for more than half a dozen families at a time, it is quite clear that this
amount of ashes meant a very long occupancy. [ven with continuous
habitation, several centuries would be required for such a quantity to
accumulate; and if residence was desultory and intermittent, as is
customary with roving or hunting tribes, or if it was only a winter
home for some of those living in the villages mentioned, the period
would be greatly lengthened. Yet the remains found in these ashes
were of the same character from top to bottom. The artificial objects
found numbered about 75 mortars, more than 200 pestles, hundreds of
flint knives or spear heads, numerous implements of bone, antler, and
shell; quantities of crude pottery fragments, a few tomahawks, and
two pipes. While the many mortars and pestles indicate much use of
grain, seeds, and nuts, at the same time the great amount of mammal,
bird, and fish bones showed that a large part of their sustenance was
derived from animal food. Of more than 20 skeletons found in
various stages of decay, only two were of aged individuals, most being
remains of children or young persons. The skulls were of low type.
Not an ornament of any sort was found except a few rude ones of
bone or shell. Some of the human bones, mostly those of children,
were charred and broken, and mingled with the debris of food animals
and ashes as if the flesh had been used for food, and the broken bones
thrown aside with the refuse. There was no evidence of the crema-
tion of bodies ; the condition of these bones points to the practice of
cannibalism.

The second cave explored is situated a mile south of Waynesville,
on land belonging to Dr. J. W. Sell, and was probably a temporary
camping place. Its opening is on a hillside facing Roubidoux creek,
and is easily accessible from either the top or the bottom of the hill.
A few rods back from the entrance, water stands on the floor through-
out the year: so that only the front part of the cave was used for
shelter. At the entrance is a pile of earth washed from the sloping

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 10g

surface of the hill above, over the ledge that forms the roof. This
accumulation spreads for 50 feet into the cave, but not so far on the
outside, because there it washes down the slope. From its surface to
the clay floor on which it rested the greatest depth was a few inches
over six feet. From top to bottom there was found in this cave the
ordinary debris of an Indian campfire. Buried at various depths
here and there, in the portion within the cave were 14 human skele-
tons, most of them so decayed that only a few fragments were remain-
ing. They are of persons of various ages; some of the skulls were
low, small, and flat.

A long period of occupancy is indicated by the conditions here as
at Miller’s cave. While the amount of earth heaped in front of the
cave does not seem large, yet it all has come from a space not exceed-
ing 6,000 square feet in area and most of this is bare rock with humus
of decayed vegetation existing only in the crevices or on the few flat
surfaces. All the other water from the hill runs to the slopes and does
not reach the cave. A violent storm passed over the region soon
after the work was concluded, in which 12 inches of rain fell in
three days, yet not more than a wheelbarrow load of soil was washed
down over the roof to the pile already there. It is evident that cen-
turies would be required to build up the mass, throughout which these
traces of man’s presence are scattered promiscuously.

During all the period these caves were in use no improvement took
place in the fabrication of stone implements or pottery. Specimens
found nearest the top of the ashes or dirt could not be distinguished
from those of the same class from the rock or clay floors. The inhabi-
tants remained in the same plane of culture.

The thousands of small mounds extending southward from the
upper swamp region of Missouri have long been a puzzle to arche-
ologists and until recently it has been supposed that in this state they
are confined to the southeastern portion: but in the course of
Mr. Fowke’s field-work they have been found to extend to the north
and west as well. Groups of them have been located in Oregon,
Dent, Phelps, Pulaski, Osage, and Morgan counties, the latter along
the Benton County line. Their purpose has not yet been determined,
although Thoburn’s hypothesis that they are due to the Pawnees,
whose line of migration was through the area in which they occur,
and are the remains of earth-covered houses, seems the most tenable
yet advanced. This theory implies that when this tribe passed beyond
the region in which suitable timber for supporting the weight of the
earth, and also earth adapted to such use, could be procured, they were
compelled to substitute for them small poles overlaid with grass.

8

110 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Comparison with the typical modern Pawnee earth lodge, a photo-
graph of which Miss Densmore published in her account of field-
work for 1919, corroborates Thoburn’s interpretation of these
Missouri mounds.

ARCHEOLOGICAL RECONNOISSANCE IN HAWAII

With an allotment from the Bureau of American Ethnology,
Mr. Fowke spent the entire summer of 1920 in the Hawaiian Islands,
making a careiul examination of all ancient works of whatever nature,
that could be visited with the time and opportunities at his command.

While there is abundant evidence everywhere that the old Hawaiian
people were extremely industrious, using vast quantities of stone in

Fic. 124.—Pawnee ceremonial earth lodge, exterior. Photograph by
Miss Densmore.

the construction of their temples, houses, garden enclosures, fish
ponds, and taro terraces, there was nothing discoverable among all
these remains which could be attributed to a prehistoric tribe, or to
any other race than that found in possession when the islands were
first known to the white man.

The five principal islands were visited and explored as thoroughly
as was possible in the circumstances. It was not practicable to ex-
amine every feature of interest, where there is so much demanding
attention; but at no place could there be discovered any indication
which would seem to justify excavation with the expectation of un-

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 MEA EAE

earthing beneath the top soil remains which might differ in general
character from those on the surface. There being no difference
between remains belonging to the historic period and those superficially
showing evidence of great age, it is logical to conclude that when that
branch of the Polynesian race, now known as Hawaiians, left their
home in the distant South Seas and migrated to these islands, they
found the territory without inhabitants ; and there is no reason what-
ever for supposing that any people culturally different from the his-
toric Hawauians had ever previously lived on the islands.

FIELD-WORK AMONG THE FOX AND PLAINS CREE INDIANS

Dr. Michelson, ethnologist of the Bureau of American Ethnology,
began field-work among the Fox Indians at Tama, Iowa, about the

Fic. 125——The dwelling in which the White Buffalo Dance of the Fox In-
dians is held. The building is the typical “ bark” house used by the Fox in
the summer and early fall.

middle of June. His main purpose was to restore phonetically a text
containing the autobiography of an Indian woman written in the cur-
rent syllabary which he had obtained in the summer of 1918, to correct
the translation where there was need, to elucidate some ethnological
references contained in the text, to clear up some grammatical ob-
scurities, and to work out the verbal stems so far as was feasible in
the field. All this was successfully accomplished, and Dr. Michelson
left for Saskatchewan in the latter part of July for a preliminary
investigation of the Plains Cree. The results of this investiga-
tion show that the Plains Cree are tall and have a cephalic index

LIZ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

.

Fic. 126.—An aged Plains Cree (File
Hills Agency).

£ mn si 5 =
ee

Fic. 127.—Tipi of the Plains Cree (File Hills Agency).

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 113

of about 79, and evidently are the same type as the one which
formerly occupied the Mississippi Valley, thus confirming the results
of Dr. Boas, announced in 1895. The general grammatical prin-
ciples which have been worked out for Fox apply also to Cree.
In some respects Cree is more archaic than Fox, in others less so.
Ethnologically the Plains Cree are about half way between more
typical Indians of the Plains, such as the Blackfeet, and the Central
Algonquins. An analysis of the myths and tales which cluster around
the culture hero shows that we practically have the myths and tales of
the culture heroes of the Blackfeet and Ojibwa combined. All this
is just what one would expect from the geographical position of the
Plains Cree.

ARCHEOLOGICAL EXPLORATIONS IN TENNESSEE

Mr. W. E. Myer, of Nashville, Tenn., spent September and
October, 1920, making explorations for the Bureau of American
Ethnology in the Cumberland Valley around Nashville. He dis-
covered on the H. L. Gordon farm, one mile northeast of Brentwood,
in Davidson County, the remains of an ancient Indian walled town.
These were situated in a woodland and had never been disturbed by
the plow. Their partial exploration brought to light some new and
interesting details of the life of the inhabitants. Traces of 87 house
circles and faint indications of several more could be made out. This
town covered 11.2 acres and was surrounded by an earthen embank-
ment which formerly supported a palisaded wall, equipped with
circular towers every 55 feet.

The ancient inhabitants, for some unknown reason, had deserted
.this village and the site had never afterward been occupied or dis-
turbed. The deserted structures had gradually fallen down and the
remains slowly buried under from 10 to 14 inches of earthmold. In
some of these circles portions of beautiful, smooth, hard-packed,
glossy-black floors were found. In the centers were the ancient fire-
bowls, yet filled with the ashes of the last fires kindled in these homes
before their owners left them forever. Near these fire-bowls often
could be seen the metates, mullers and other household utensils, just
as left the last time used. Underneath the floors were the stone slab
graves of the little children, one of which is shown in figures 128
and 120.

A level open space was found near the center of the town and on
the western side of this plaza was a low flat-top mound that had
originally supported some important building. Adjoining this mound

114 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

on the west was an earth circle which probably outlined the ruins of
the town house. At the center of this sacred structure, on the unique
black glossy floor, an ancient altar (fig. 130) was found. It was still
filled with the pure white ashes of what had once been the sacred fire.

Fic. 128.—Child’s grave after removal of infiltrated soil,
before disturbing mortuary vessels.
This altar was carefully preserved and is now in the Bureau of
Ethnology. The Gordon site is of much interest because here we
have an ancient Indian village just as the original inhabitants left it.

THE FEwWKES GRouP
Mr. Myer also partially explored an unnamed Indian village group
at Boiling Spring Academy in Williamson County, Tenn. At the

iD
3S

No. 6 SMITHSONIAN EXPLORATIONS, 19 115
request of many citizens of Tennessee Mr. Myer named this site the
Fewkes Group, in honor of Dr. J. Walter Fewkes, who had visited it
and recognized its possibilities a few months before.

Fic. 129.—Child’s grave after removal of body. Note
floor composed of broken pottery.

At least two different peoples have lived on this site. The earlier
people built the mounds and most of the other remains. At a later
date a small band of some other tribe located here. The earlier people
buried their dead either in hexagonal or almost circular stone slab
graves, the bodies closely flexed. The later band used rectangular
stone slab graves with the body extended full length on its back.

116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

The Fewkes Group consists of four mounds, one on each of the
four sides of a level plaza, the traces of about a dozen house circles,
and a small remnant of what was once a considerable stone slab
cemetery.

Fic. 130.—Altar.

Mound No. 2 on the map is a low oval mound situated on the
western side of the plaza. The site of this mound had been lived upon
for some time before any mound was raised. At last the mound was
commenced and raised to a height of three feet and a building for
domestic purposes erected thereon. This building was later torn down
and then the mound was raised three feet higher. The mound was
again used for domestic purposes for a period. Then a town house

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 WAVY),

or sacred ceremonial house was built on it. This sacred building also
had one of those rare, beautiful floors made of clay, smoothed, then
hardened by fire, and finally covered with a coating which is yet black
and glossy. In the center of the building, on this beautiful floor, an
altar was found. It was similar to the altar shown in figure 130.
This building had walls made of cane stalks with the leaves
attached, which had been woven in and out between the upright posts
which supported the roof. These canes may have had a coating of
earth, though no trace of it could be found, and the walls also had a

Fic. 131.—Mortuary vessel from
child’s grave.

covering of woven cane matting. In some way the building was
destroyed by fire. Earth was thrown on the remains in time to
smother its still glowing embers, which produced a large amount of
powdery charcoal containing fragments of cane stalks with the leaves
attached, and portions of the woven cane matting. After this sacred
building was burned the mound was raised one and one-half feet or
more in height. All trace of its last use has been destroyed by 85
years of cultivation.

The low mound, No. 3, on the south side of the plaza, was a burial
mound belonging to the first settlers. The mortuary vessel shown
in figure 131 came from a hexagonal grave in this mound.

118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Mound No. 1 on the north side of the plaza is 180 feet across the
base and 25 feet high. It is the most conspicuous mound in the
group. Lack of funds prevented its exploration.

House circle No. 6 was one of the group of buildings, Nos. 6, 10,
and 11, whose functions were doubtless closely interwoven. No. 6
contained in its center an altar or fire-bowl.

There was evidence that this town had either been taken by an
enemy and burned, or the ancient inhabitants, forced to flee, had
burned their homes to prevent their falling into the hands of the
invader.

Fic. 132——House circle No. 17. Cleared floor of wigwam, showing ancient
fire-bowl. Body of child was found by side of upright stone. Its head rested
within edge of fire-bowl. Top edges of upright stone slab sides of another
coffin in corner to right of women.

House circle No. 17, shown in figure 132, was a typical dwelling.
It was evidently the home of a neat housekeeper. No broken animal
bones, pottery, fragments, or other evidences of untidiness littered
the floor. The floor was of hard-packed clay and a fire-bowl was
sunk in the center of the floor. At this fire-bowl a puzzling burial
was unearthed. A child, about eight years of age, was buried by
the side of the upright stone slab, with its head resting just within
the extreme edge of the fire-bowl, whose rim had been cut away at
this point to admit the top of the child’s head. The fire-bowl was
found still filled with ashes, and although the ashes covered the top

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 I1g

of the child’s head, the head showed not the faintest trace of the
action of fire. The graves of two infants were also found in the floor
of this house.

At both the Gordon and the Fewkes groups every piece of bone
and every fragment of pottery was carefully saved and location
noted. These thousands of fragments will give a reliable record of
the food animals and practically a complete list of all the sizes, colors
and shapes of their domestic pottery.

Both these sites, when explorations were completed, were accurately
restored to their original shape for the benefit of coming generations.
The interesting altars, fire-bowls, building post-holes, and vestiges of
domestic life were carefully preserved, and again covered up so as to
allow their future study. It is strongly urged by the citizens of
Tennessee that the Fewkes Group be made a national monument.

Mr. Myer discovered a great Indian fortress on the long, narrow
point of land between the Harpeth and Cumberland rivers, at their
junction, in Cheatham County, Tennessee. This fortress consists of
a thin, double-faced bluff, about three-quarters of a mile in length,
and only from 10 to 250 feet wide along its tall and narrow summit.
It faces both rivers and has nearly perpendicular sides along its entire
length on both streams. It can be scaled with very great difficulty and
at only a few places. The Indians protected these few places of ascent
with breastworks or mounds. This was a central place of refuge for
a series of scattered Indian settlements extending about six miles up
and five miles down the Cumberland River and about five miles up
the Harpeth.

The four pipes from this region are unlike any found elsewhere
. in the valley, and probably the culture of the ancient people who used
this fort was different from any other known at present in the Cum-
berland Valley.

Ona recent visit Dr. Fewkes examined the great unexplored mound
group on Harpeth River at the mouth of Dog Creek, in Cheatham
County. This group is the remains of one of the important pre-
historic settlements east of the Mississippi.

There is a great mound, with wide earthen platforms, capping a
hill in the up-stream end of this settlement. A portion of the hill
has been artificially shaped so as to give greater prominence to the
works on top of it. Surrounding these works on the summit are
the ruins of a large edifice and other important remains. This portion
of the settlement covers about 40 acres and is said to be connected by
an embankment with the remainder of the mounds in Mound Bottom,

I20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

about two miles down the Harpeth. The Mound Bottom portion,
shown in figure 133, covering about 50 acres, contains Mound No. 2
with large platform ; Mounds Nos. 1, 4, 5, and 6; cemetery No. 7, and
other traces of a considerable population.

eee
les

Fic. 133—Mound Bottom. Other great mounds belonging to this unexplored
group.

ARCHEOLOGICAL EXPLORATIONS IN NEW MEXICO

During July, and a part of August, Mr. J. A. Jeancon, special
archeologist of the Bureau of American Ethnology, made a recon-
noissance and conducted intensive archeological work at Taos, New
Mexico. While it has been known for a long time that there was
excellent archeological as well as ethnological material to be obtained
in this valley, no archeological research has been done there before the
present work in the summer of 1920.

The great number and variety of sites seem to indicate a long
period of occupancy of the region. These sites are well-defined small
house groups, each indicating one, two, or more rooms. ‘There are
also great communal groups as that at Bagley ranch and the Arroyo
Hondo. The mounds are sometimes over 20 feet in height; the
pottery sherds scattered over these sites range from the primitive
black-and-white ware to what appears to be the so-called biscuit ware.

A small pueblo ruin at Llano was selected for excavation and, while
the yield of pottery was small, many interesting facts were obtained.
The ruin is located on the south bank of the Little Rio Grande, one
mile and a half from the plaza of Ranchos de Taos. Situated on the
edge of a high mesa it commands a magnificent view of the country
for miles around, excepting to the east, where the view is cut off by

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 121

the mountains. The fields which probably furnished the villagers
well-watered agricultural areas are situated in the valley below.

The excavation showed that there had been two occupations on the
site. In several places the remains of earlier old walls show beneath
the present walls, indicating that the first and second buildings had

Fic. 134.—Ruin at Llano, Taos Valley.

totally different ground plans. All of the walls (fig. 134) were made
of a mixture of wood ash, small stones (about the size of pebbles ordi1-
narily found in gravel), and adobe which when exposed to the air
becomes very hard. The walls of the second occupation are not as
well made as those of the first, but the floors of both resemble those
of the older villages still inhabited, where the blood of animals was
used to give them temper and polish.

[22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

The general plan of the ruin recalls those of southwestern Colo-
rado. The kiva at Llano (fig. 135) was completely surrounded with
rooms and when the roof was intact its surface was a small plaza
surrounded by the buildings of the main group. An evidence of the
two occupations is that the more recent kiva was built inside the older,
and the space between their walls had been filled in with rubbish. No
roof timbers remained in the ruin, but there were in the kiva four
upright posts upon which the former roof rested. In about the center
of the kiva there was an excellent fireplace, and in the floor directly

Fic. 135.—Kiva at Llano, Taos Valley.

adjoining the fireplace a plastered pit (fig. 135), an unusual adjunct
to the fireplace, the purpose of which is unknown. Between the fire-
place and the pit is a stone slab, one foot high and about nine inches
wide. The excavations brought to light a very small number of the
undecorated sherds of white ware; the black-and-white pottery is
unusually hard and fine, but the black or rather brownish-black ware
used for cooking and storage is very soft. The forms of and designs
on the black-and-white pottery strongly suggest the San Juan ware.
During the month of May, 1920, Mr. Jeancon made a satisfactory
reconnoissance in the country lying south and southwest of Dulce,

123

SMITHSONIAN EXPLORATIONS, 1920

NO.

Fic, 136—Ruin in La Jara Canyon.

Fic. 137.—Ruin in La Jara Canyon.

124 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

New Mexico. He found in Burns, La Jara and other canyons indi-
cations of two periods of occupation evinced by the pottery which
belongs to the old pre-Spanish black-and-white and the hard-
coiled wares. The houses of the earlier period were “earth
lodges” of a primitive type. The house remains point to the second
occupation and are in many cases built on the tops of high masses of
sandstone (fig. 136), and show the crudest workmanship. A common
site for buildings is the edge of high points (fig. 137), along the
terraced (natural) sides of the canyons. All of the later sites give

~ a

Set

Fic. 138.—Dulce Ruin.

an impression that they were more or less defensive and _ hastily
erected. Associated with these buildings are remains of metal objects,
apparently of Spanish manufacture ; many of the beams in the houses
plainly show the marks of metal tools. The pottery has a bluish-black
color, closely resembling a modern vitrified brick, and is so hard that
when struck it rings with a clear bell-like tone. The surface suggests
old Hopi ware and the decorations are similar to the older Zuni and
Acoma designs.

The pottery and associate remains probably do not antedate the
reconquest of the Southwest by the Spaniards in 1690. It is known
that some of the pueblo people fled from the Spaniards at the time of

NO. 6 SMITHSONIAN EXPLORATIONS, 1920 125

the great rebellion and took refuge in the mountains to the north.
It is probable that further investigation will show that in this area
the ruins ascribed to the second period were built and inhabited by
refugees from Zuni and Acoma.

ARCHEOLOGICAL EXPLORATIONS IN EASTERN TEXAS

Archeological field-work was carried on in 1920 by the Bureau of
American Ethnology in co-operation with the University of Texas
under direction of Professor Pearce and Professor Engerrand.
Several localities in Texas were visited and considerable intensive
work done in the eastern part of the state.

The region in eastern Texas to which most attention was given was
in the vicinity of the little city of Athens in Henderson County. Judge
A. B. Watkins of that city has long taken a keen interest in the
history and archeology of the region and was of great help to
Professor Pearce in his field-work. A number of aboriginal objects,
complete specimens of pottery, clay pipes, conch-shell cores, fine
arrowheads and human bones were obtained from an ancient burial
ground two miles northeast of Frankston on the De Rossett Farm.
One of the objects found was a “ turkey call” or whistle made from
the drumstick of a turkey. This is identical with those used not
many years ago by old white hunters on the frontier and Professor
Pearce feels sure that the white man’s use of this device was borrowed
from the Indian. Professor Pearce finds that the east Texas region
contains numerous mounds, village sites and burial places, among
which may be mentioned Nacogdoches, Panola, Bowie, Wood and
other counties.

Three interesting mounds on the Morrall Farm four miles east of
Cherokee County were investigated. The highest of these mounds
has an altitude of about 35 feet above the level valley or field in which
they are all located and is about 80 feet across at the base. This
mound is very steep and even now after long erosion its sides rise at
an angle of 45°. Mound B is 180 feet long by 75 feet wide and rises
only 15 feet above the general level. The other three mounds in the
vicinity of Athens have been ploughed into and have no regularity
in form.

On the Quate tract east of De Rossett farms there are Indian
mounds which were not regarded as very ancient. Several mounds
situated in Harrison County on the farm of Mr. Lane Mitchell, of
Marshall, were examined and remains of earth lodges with central
fire pits were reached. These mounds are probably very ancient.

126 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Numerous other sites were explored, yielding a collection of pottery,
stone implements and other objects, illustrating the life of the pre-
historic aborigines of eastern Texas.

Everything found implies that the Indians of this region lived in
relatively settled villages, had considerable agriculture, made pottery
extensively and of a high-grade, and were altogether in a considerably
higher stage of culture than were those who occupied the prairies
and plains.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 72, NUMBER 7

SEA-LILIES AND FEATHER-STARS

(WitH 16 PLATES)

BY
AUSTIN H. CLARK

(PUBLICATION 2620)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1921

The Lord Baltimore Press

BALTIMORE, MD., U. S. A.

SEA-LILIES AND FEATHER-STARS
BY Aust UNH CUART
(WitH 16 PLATES)

CONTENTS PAGE
[PRELARE Gini AA COO Oe De LS BEE TIRE ae ii ei ek ee I
Number and systematic arrangement of the recent crinoids.............. 2
she interrelationships: of the crinoid’ species..5.0.......csse-crcsensccees 2B
etapa ticestntGeunercone the enimO1dSs ses ¢) ccclssrm elo Gustewtt nc sieeve cr detetore ito sees 4
Wiviparous chinoids, and sexual differentiation... ......- 0.2: -2-.+-.-s+5.. +46 10
itiendevelopment ol the comatiulids.....4..02 00.055 cease use oc cemc secs nes 10
IR ESREMEGELIONS od Sloat chess GEOR CRD EOC OEE EEC are men a ee 12
RSA Cla Med NORA Logos Patel Sled ore AK GA is col rel se aToneis vsyekaym ceoeat s eteaeieatie Subetea tle Bah 13
MHeKcomposition om the Crinlold Skeleton. « cs. 0.0 oh dla ewiacteosles salsa s 15
SP eMCSitatlo Tito neOreth Cu ChMO1G Sirarcveis dicta sie cre eres ors oe, Ae aioreke. sels 0 #1 gains ous 15
Mieapalconrolosicale history: Of therlivine Chinoid’s...¢2 2... c. cele cnes- 16
The fossil representatives of the recent crinoid genera.................. 17
The course taken by specialization among the crinoids.................... 18
MM eROCCHEGENCeLOL ttOtal: GhINOIGS sir. see oe Sele Sakae «2 Grete s cieyee nie dl oray abs 18
sbieRrelatronnOlachinords, CO LEMPeratlre’s a. sceiccie es crecwi ns sje ciele.c eieces s wieie ets 20
LEGS Chm TE iirc eat ae er Sens Sears a LS Ue, lol ePeve esi el cpelaccrk's ets eitisiaiedele w apices dae PS
HEGCOMOPTON, Yelos cater. ssa APR Rete tae ve Acacia suas eae rerere Sh Ao RS CECE 2
(CELGIE * & cercid oe ocr ORE TIOCS BCG OC CI EL rN ICT PRIOR Se Ronee ee 24
The similarity between crinoids and plants............. GS Ei Ne aon Oat 29
Parasites; and CoMmmensalsy i... 5.555. yen Re AR a iisd ret dae, thee ios 34
eaumPUSaiarOte ties CPINOUISs,<oacles eivgeres <\heansad ays abe u.eie soa ae Bek coals oe 39
Bemucwmiaicuvedl wero tne livitle CTinlO1dS. «2/2 e ss. 0 seein occ sacked adds uh 30
Pea ema NHN MI HU Te [TLASS I cee carsy cree Pestana a ache, era loke pence et eantajeaa. 6 bath toe | ot clea eS 4O

PREFACE

Of all the animals living in the sea none have aroused more general
interest than the sea-lilies and the feather-stars, the modern repre-
sentatives of the Crinoidea. Their delicate, distinctive and beautiful
form, their rarity in collections, and the abundance of similar types
as fossils in the rocks combined to set the recent crinoids quite apart
from the other creatures of the sea and to cause them to be generally
regarded as among the greatest curiosities of the animal kingdom.
They have usually been considered as the rare, curious and decadent
remnants of an interesting animal type once important but now
trembling on the verge of extinction, and it is from this melancholy
viewpoint that they are discussed in practically all the text-books.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 72, No. 7

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

The discoveries of the last few years have shown that the living
crinoids, far from being rare or few in numbers, are abundant both
as individuals and as species, and that in all localities where the some-
what exacting conditions under which they can exist are met they
occur, sometimes in enormous numbers. The requirements necessary
for the maintenance of crinoidal hfe are unfortunately of such a
nature that, though quite generally distributed in the deeper waters
of the oceans, they become rare or very local in the littoral, which,
together with the great difficulty of preserving them in anything
approaching satisfactory form, has served to keep them enshrouded
in mystery while animals of other types engaged the attention of
investigators.

NUMBER AND SYSTEMATIC ARRANGEMENT OF THE RECENT
CRINOIDS

_ There are known from the seas of the present day 576 described
species of crinoids which are distributed in 142 genera and 28 families
and subfamilies; of these, 76 species, included in 22 genera and 6
families, are stalked, while 500 species, included in 120 genera and
22 families and subfamilies, belong to the unstalked or comatulid type.

The systematic arrangement of the families and higher groups is
as follows:

Order Inadunata.
Family Plicatocrinide (Calamocrinus, Ptilocrinus [fig. 42], Thalassocrinus,
Gephyrocrinus and Hyocrinus).
Order Articulata. :
Family Bourgueticrinide (Rhizocrinus, Bythocrinus, Democrinus, Bathy-
crinus, Ilycrinus and Monachocrinus [fig. 39]).
Family Phrynocrinide (Phrynocrinus and Naumachocrinus).
Family Apiocrinide (Proisocrinus [fig. 40] and Carpenterocrinus).
Family Pentacrinide. :
Section I: Pentacrinites (Metacrinus, Isocrinus, Endoxocrinus,
Comastrocrinus [fig. 41] and Hypalocrinus).
Section II: Comatulids (see below).
Family Holopodide (Holopus).
The comatulid section of the family Pentacrinide in the recent
seas has become so numerous, so widely spread and so diversified

that it alone has acquired all the characteristics of an order. It is thus
subdivided :

Suborder Macrophreata.
Family Atelecrinide.
Family Pentametrocrinide (figs. 49, 50).
Family Antedonide (including the subfamilies Antedonine [figs. 47, 48],
Thysanometrine, Perometrine, Heliometrine, Zenometrine, Insome-
trine and Bathymetrine).

IN@.)07 SEA-LILIES AND FEATHER-STARS—CLARK 3

Suborder Oligophreata.

Family Charitometride.

Family Thalassometride (including the subfamilies Ptilometrine [fig.
46] and Thalassometrine).

Family Calometride (fig. 45).

Family Tropiometride.

Family Colobometride.

Family Mariametride.

Family Stephanometridz.

Family Himerometride.

Family Zygometride.

Family Comasteride (including the subfamilies Capillasterinee, Comac-
tiniine [fig. 44] and Comasterinz).

THE INTERRELATIONSHIPS OF THE CRINOID SPECIES

The various crinoid species are of very different relative value. In
some comatulid genera, especially those including species with many
arms, if any one character whereby the species are commonly dif-
ferentiated be plotted on a species curve the several species will be
found to be indicated not by a series of separate triangles, but by a
succession of more or less marked nodes which are united to the mass
forming the adjacent nodes by coalesced bases in thickness equal to
from 10 to 60 or more per cent of the maximum height of the neigh-
boring nodes. Such variability and lack of absolute fixity in any one
character is as a rule reflected in all the characters, and thus there
results a species group or genus which may be compared to a small
mountain system rising out of a plain each peak of which represents
a recognized form. In such a genus every character varies between
two extremes, but there is often no correlation whatever between the
different characters. Thus every sort of combination is possible and
a very large variety is found, though the tendency is for the characters
to form more or less definitely correlated groupings and to crystallize
into certain definite types.

This type of variability is not connected with the geographical
origin of the specimens except in a very general way ; it is chiefly seen
in the multibrachiate species of the Oligophreata, and in specimens
of these species from the East Indian region, though there are one or
two good examples in the Caribbean Sea. In the Indo-Pacific many
species which occur in numerous well-marked varieties in the Malayan
region when extending their range outside of this region gradually
become more and more fixed and definite in their characters so that
individuals from, for example, Madagascar or southern Japan, are all
found to be practically uniform in their various features and to
represent the mean of the two extremes seen in a series from the
central East Indian region.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

Thus it is evident that in a number of crinoid genera, as in many
groups among the fixed and arborescent marine organisms, we have
to deal not only with the usual and well-recognized geographical
variation and the formation of geographical “ subspecies,” but also
with the formation of “ varieties” which are strictly comparable to
the accepted “ varieties” among the plants. Such difficult and intri-
cate genera as Crategus, Aster, etc., have their representatives among
the crinoids, though in the latter the number of included species is not
by any means so large.

Many plants form marked local varieties correlated with the
physical and chemical character of their immediate environment—the
type of soil, amount of sunlight, average temperature, amount of
moisture, etc-—and several of these varieties may often be found
within a relatively limited area, as in certain species of Aster. Many
crinoids do exactly the same thing as a result of local variations in
the food supply, the amount and kind of illumination, and the
temperature.

FORM AND STRUCTURE OF THE CRINOIDS

Typically a crinoid is rather abruptly divided into a slender and
more or less flexible column which supports at its summit a pen-
taradiate head or crown (figs. 39-42) the five divisions of which
each may bear anywhere from one (figs. 42, 50) to nearly forty (fig.
17, left), but most commonly two (figs. 39, 43, 46-48), flexible arms
along the ventral side of which, giving off branches to the slender
alternating lateral appendages or pinnules, runs a narrow ciliated
groove (fig. 36, left), the grooves from all the arms of each division
uniting and running in a single groove across the so-called ventral
disc to the mouth (figs. 4,5). The cilia in these grooves pick up and
convey to the mouth the minute plankton organisms, both animal and
vegetable, which serve the crinoid as food.

A single living crinoid (Holopus), lacking the column altogether,
‘is attached directly by the crown, the basal portion of which becomes
much elongated. In the Plicatocrinidz the column continues through-
out life to add new segments just beneath the crown and never, so to
speak, matures. In the large group to which all the living crinoids
except the Plicatocrinidze belong the column increases to a definite
size, the topmost columnal then enlarging and becoming permanently
attached to the crown, forming a sort of apical plate. This‘is the
general law which, like all of nature’s laws, must be liberally inter-
preted, for it is subject to curious modifications. In some types, as in
Bathycrinus, maturity comes slowly and several of these topmost

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK 5

columnals, progressively increasing in width with the growth of the
crown, are formed before a fixed attachment occurs. In the group
including the pentacrinites (fig. 41) and the feather-stars (figs. 43-
50) two widely different extremes are found. The pentacrinite
column is extremely precocious. It reaches the mature stage and
forms the definite topmost columnal when the crown is still extremely
immature.and, so to speak, as yet lacks the instinct to attach itself to
the column. The column therefore continues to grow and forms a
second topmost columnal as in Bathycrinus, though in this case the
two are separated from each other by a number of intercalated
columnals. The second, like the first, fails to become attached to the
crown and, the rapidity of stem growth increasing, a continuous series
of these topmost columnals is formed each of which immediately
after its appearance is pushed away from the crown by the formation
of another one above it, the column becoming so enormously over-
developed that the animal cannot nourish it all and so it dies away at
the distal end as rapidly as new segments are added beneath the crown.
As they are pushed backwards the numerous elements which nature
intended for topmost columnals become separated by a definite num-
ber of intercalated segments from which they are always to be dis-
tinguished by their larger size and the presence of a whorl of five long
jointed processes ending in a strong hook, the so-called cirri. In the
feather-stars the column is just as precocious as in the pentacrinites,
reaching maturity and forming the definite topmost columnal at a
very early stage (fig. 55). But in this group attachment occurs. The
column, being mature, has ceased its development ; the crown is as yet
very small and young; the topmost columnal has become attached to
. the latter and now forms an integral part of it, and it continues to
develop with it regardless of the conditions in the column of which it
was once a part. The crown becoming too large and heavy for the
column, the latter breaks away just beneath the enormously enlarged
topmost columnal, now called the centrodorsal, and the animal be-
comes free. The further development of the soft structures of the
column takes place entirely within the centrodorsal (fig. 7) from
which the numerous cirri, here crowded together instead of spaced in
whorls of five as in the pentacrinites, are extruded (fig. 10).
Feather-stars mostly attach themselves to foreign objects by means
of their usually stout and hook-like cirri which are most commonly
from one-fourth to one-third the length of the arms (figs. 33, 34),
though in a few species they are longer than the arms (figs. 32, 46).
In some the cirri are very numerous, long, slender and nearly straight,
forming collectively a sort of disc which, on the principle of a snow-

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

shoe, supports the animal on soft ooze or mud (fig. 35), and in one
family (Comasteridz) they are frequently quite absent, the centro-
dorsal being atrophied and sunken in so that its now flat surface does
not project beyond the dorsal surface of the crown (fig. 11). Penta-
crinites attach themselves to arborescent marine animals by their
cirri, like the comatulids, or lie upon the sea floor with the column
more or less coiled and the crown raised high above it. The other
crinoids, except Holopus, are either attached to solid objects by the
expanded and encrusting basal columnal or terminal stem plate
( Plicatocrinidz, Apiocrinidz, and Phrynocrinide), or send out from
the distal portion of the column a mass of root-like processes by
which, plant-like, they are able to maintain an erect position on muddy
bottoms (Bourgueticrinide). It is curious that in some of these if
the crown be lost before the stem dies a mass of roots is produced
from the upper end of the column similar to those at the lower end.

The crown of a crinoid is composed primarily of three alternating
circlets of five plates, excepting in the Plicatocrinide in which there
are only two, calling to mind the bracts, sepals and petals of a flower.
The plates of the lowest circlet (infrabasals) are small, often only
three in number, and frequently, or perhaps usually, absent in mature
animals ; the plates of the second circlet (basals) are larger and more
constant, though in the feather-stars they become transformed into
an internal septum (fig. 7, shown cut across just above the cavity
within the centrodorsal) with no hint of their original significance,
and in the Plicatocrinide there are frequently only three of them; the
plates of the third circlet (radials) are always highly developed, and
are structurally the basal plates of the arms. In a few types each of
these is doubled so that there are ten of them instead of only five
(compare figs. 49 and 50).

Following the radials there is a linear series of ossicles which
rarely remains undivided (figs. 42, 49, 50), almost invariably forking
on the second (figs. 39, 46-48) ; this gives ten arms, the commonest
number among the crinoids ; but each of these may again divide on the
second or fourth segment beyond the first division (very rarely on any
other) (figs. 40, 41) and this process may be repeated in extreme
cases as many as eight or nine times (fig. 17, left). Ordinarily all
of the five groups of arms are alike and all of the arms are of the
same length, but in the Comasteride the arms arising from the left
posterior radial, sometimes from both posterior radials, are frequently
shorter than the others, sometimes scarcely more than a third as long
(fig. 44), and they are further peculiar in lacking the ambulacral
groove (fig. 36), in bearing more numerous and more developed

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK 7

gonads, and in ending in a terminal brachial bearing a pair of pinnules
instead of in a growing tip as do normal arms.

From the second onward the arm segments bear on alternate sides
long slender jointed processes called pinnules (figs. 1, 44, 48), which,
in the comatulids, are of three types. At the base of the arms and on
the second ossicle of division series composed of four elements are
the so-called oral pinnules which lack the ambulacral groove and are
variously modified into very sensitive tactile organs (figs. 47, 48, 51)
or into stout spines (fig. 52) extending over and protecting the disc ;
following these and more or less abruptly differentiated from them
are the shorter genital pinnules (figs. 1, 51, 52), with, or, more rarely,
without (fig. 36, left) an ambulacral groove, on which the gonads are
developed ; there are usually between six and twenty of these on each
side of the arm, and distally they pass gradually into the longer and
usually very slender distal pinnules (figs. I, 51, 52) which serve
purely as food gatherers. In a few types one or more of the earlier
pinnules are lacking (fig. 43), while in the Comasteridee (figs. 15, 43)
the oral pinnules bear curious comb-like structures recalling the
pectinate antennz of certain insects. In the crinoids other than the
comatulids the pinnules are much more nearly uniform in structure
and in function; in some types (Metacrinus, Hypalocrinus, Comas-
trocrinus [fig. 41] and Proisocrinus [fig. 40]) they are rudimentary
or even quite lacking on the terminal portion of the arms. In one
comatulid (Comatulella brachiolata) many of the pinnules are modi-
fied into stout organs resembling cirri which assist the animal in
clinging to arborescent marine organisms.

The various ossicles which together form the crinoidal skeleton
are tied together by more or less closely packed bundles of fibrillze
the ends of which take the form of loops within the calcareous sub-
stance. Between most of the brachials or ossicles of the arms there
is found in addition to the ligaments a pair of ventral muscle bundles
and between the ossicles of the pinnules there are sometimes a few
muscle fibers or a small muscle bundle. Excepting in Holopus more
or fewer of the brachials are united in pairs by ligament fibrillze only ;
such unions, known as syzygies, are extremely close, at right angles
to the axis of the arm, and with the joint faces marked with radiating
ridges ; two brachials so united usually appear as a single one with a
thin dotted line across it (fig. 54). Crinoids seem to have the power
of severing the syzygies at will, and arm fracture almost invariably
takes place at these unions. As a rule syzygial pairs are regularly
distributed throughout the crinoid arm, the first being composed of
the third and fourth brachials ; their number decreases with specializa-
tion and with increase in the number of arms.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

In addition to the primary bony framework including the ossicles
of the crown—calyx plates, brachials and pinnulars—and stem and
cirri crinoids have two other skeletal systems; one, superficial, takes
the form of very numerous spicules which may increase in size to
definite plates sometimes mutually in contact protecting the soft
ventral integument; along the ambulacral grooves, especially on the
pinnules, these become more regular and better developed than else-
where (figs. 18-26) and often form large and definite side and cover-
ing plates (figs. 27-31) the former lying in the perisomic wall and the
latter, hinged to them by ligaments, in the lappets which line the
ambulacral grooves and capable of being closed down over them; the
other skeletal system is the internal, consisting of numerous spicules
and networks occurring more or less plentifully in the bands of con-
nective tissue traversing the body wall and in the walls of the
digestive canal.

In a very large 10-armed feather-star in which side and covering
plates are developed there are visible externally about 600,000 distinct
skeletal elements each of which arises from a separate center of
ossification; of these about 87,000 belong to the primary and about
513,000 to the secondary or perisomic skeletal series. In a large
comasterid with no side and covering plates visible there may be as
many as 700,000 primary skeletal elements, while in the very small
antedonids the number probably never falls below 10,000. The
greatest of the figures, however, is insignificant when compared with
the number of ossicles in the larger pentacrinites where, in the recent
species, nearly two and one-half millions are found. These figures,
large as they are, must be approximately doubled when the internal
skeleton is taken into consideration.

The internal structure of all the crinoids which have been studied
is very similar. The mouth, at the point of convergence of the
ambulacral grooves coming onto the disc from the arms (figs. 4, 5),
usually central or subcentral but often excentric or even marginal in
the Comasteride (fig. 12), leads downward through the gullet into
the digestive tube (fig. 13) which, turning to the right, makes some-
what more than one complete coil (four in some of the Comasteride),
the posterior portion bending upward and slightly forward to end at
the summit of the so-called anal cone situated on the disc between
two of the ambulacral groove trunks. Except for its canal-like pro-
longations into the arms the body cavity is filled with a mass of
mesenteries, bands and septa so that it is reduced to a minimum, and
in some cases, as in Isocrinus, has entirely disappeared.

Within the centrodorsal in the comatulids (fig. 7) and just over the
summit of the column in the other types lies the chambered organ;

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK 9

its central axis from which the partitions dividing it into five sections
arise is continued irregularly upward to the vicinity of the mouth as
the so-called axial organ, a cord of connective tissue including a
number of open spaces or lacune. Five slender prolongations, one
from each of the chambers-of the chambered organ, accompany the
axial organ for some distance, ending blindly, and each cirrus
includes a median vessel from the same organ.

Except at the point where the axial organ leaves it the chambered
organ is completely encased in a mass of nerve fibrilla from which the
large dorsal nerves of the arms and pinnules and their various
derivatives as well as the cirrus nerves arise (fig. 9) ; while over this
nervous envelope and along the nerve cords lie multitudes of wander-
ing cells which play an important part in the regeneration of lost
members. The envelope of the chambered organ and its derivatives,
including also a closely associated nerve ring about the mouth which
sends off very numerous branches including two cords running along
the ventral surface of each arm, form the chief nervous system of the
crinoids, but beneath the ambulacral grooves of the disc, arms and
pinnules there is second quite independent nervous system consisting
of a continuous thin layer of nerve fibrille.

The water vascular system of the crinoids consists of a ring canal
about the mouth and vessels radiating out from it under each of the
ambulacral grooves which they follow in their course to the pinnule
tips, sending off branches into the delicate tentacles. Attached to
the ring canal is a row of little tubes which open into the body cavity,
and communication between the body cavity and the sea water outside
the body is maintained by a number of calyx pores (5 in Rhizocrinus,
1500 in Antedon mediterranea, and still more in larger species) which
pierce the body wall.

The blood vascular system of the crinoids is very highly developed,
though the blood vessels are nothing more than intercommunicating
cavities or gaps in the connective tissue of the mesenteries, bands and
cords which in all directions traverse the body wall, the walls of the
digestive tube, the axial organ, etc.

The so-called genital cord forms an irregular pentagon about the
mouth from which five branches are given off, these running beneath
the water tubes into the arms and pinnules; but it is only in certain
of the lower pinnules in most comatulids or at the pinnule bases and
rarely in the arms in the stalked types that the sexual products are
developed. It has already been mentioned that in the short posterior
arms of certain of the Comasteridz the ambulacral grooves are absent ;
when this occurs the ambulacral nerves, water vessels and tentacles
are also absent, but the genital cord is here especially developed.

Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Along the ambulacral grooves, except in most of the Comasteridee,
and usually also internally along the digestive tube, is a usually
regular row of little round bodies, conspicuously colored in preserved
specimens though usually colorless in life, called sacculi (fig. 4.) which
remind one strongly of the glandular dots on the leaves and petals of
certain plants such as the species of Hypericacee. They are assumed
to be excretory organs.

VIVIPAROUS CRINOIDS, AND SEXUAL DIFFERENTIATION

Three antarctic comatulids are viviparous, the young developing in
special pouches or marsupia formed on the pinnules in two, but on
the arms themselves in one. In all three of these the two sexes are
easily distinguished by superficial external examination. Besides
these three comatulids only six echinoderms are known to exhibit
sexual dimorphism, four irregular echinoids, one holothurian, and
one ophiuran. Excepting for one of the echinoids (Anochanus) all
are from antarctic or subantarctic regions.

THE DEVELOPMENT OF THE COMATULIDS

The early stages of the comatulid larve are passed within the egg
membrane and the developing eggs usually hang from the bases of
the genital pinnules like little bunches of grapes. It is only after the
formation of the elements of the skeleton—the rudiments of the
terminal stem plate, a number of columnals, the infrabasals, basals
and orals—that the larva emerges as a small barrel or bean-shaped
creature with five transverse ciliated bands, an anterior tuft of long
cilia, and a deep ventral groove (fig. 56).

In the best-known species (Antedon adriatica) the length of the
free-swimming life varies very greatly, even in different individuals
of the same brood; some attach themselves after a few hours and
immediately proceed to further development, while others are to be
found still swimming about after a lapse of as much as four and a
half days. As a rule the free-swimming existence terminates after a
few hours and is rarely as long as two or three days; larve still
swimming at the end of that time are abnormal and incapable of
fixation.

After attachment the larva soon assumes the form of a curious
little stalked creature composed of a delicate column attached by a
spreading base and supporting a calyx consisting of three or five, or
often no, infrabasals which are very small, delicate, and difficult of
detection, five large united basals forming a cup in which the visceral
mass is enclosed, and five equally large orals superposed upon them

INO: 7, SEA-LILIES AND FEATHER-STARS—CLARK i

which may be opened outward or closed together over the mouth.
The plates of the radial series appear first as minute spicules in the
angles between the basals and orals, one in each of these angles, or
sometimes two in the right posterior angle. These grow very rapidly
and two additional plates appear beyond them, the outer giving rise
to two plates each of which forms the base of a rapidly growing arm.
The additional plate in the right posterior area, known as the radianal,
is always present, often appearing before any of the radials, but
sometimes, as in Antedon, not being formed until a relatively ad-
vanced stage has been reached; it grows very slowly and by the
growth of the radial just to the right of it it is shoved gradually
upward and to the left, so that when the radials have come into lateral
contact and have united into a ring it occupies a position at the edge
of the disc where it is soon resorbed. Though small and soon
disappearing in the young comatulid this is a very important plate
in many fossil types. The arms at first are very different from the
arms of the adults; they are composed of a series of exactly similar
brachials without pinnules and with all the articulations between them
the same, without muscles, and crossing the arm at right angles.
Pinnules first appear at the tip of the growing arm after from nine
to fourteen brachials have been formed, these being followed by the
pinnule on the second brachial and considerably later by the pinnules
on the intervening brachials (fig. 55). The first cirri appear on the
now enlarged topmost columnal just before, simultaneously with, or
just after, the first formation of the pinnules. Just before the
appearance of the pinnules and cirri and before the disappearance
of the radianal the larval comatulid is a remarkably perfect replica
of a fairly typical representative of the Flexibilia Impinnata, but
‘after the appearance of the pinnules and the cirri the crown under-
goes a most extraordinary transformation and rapidly assumes all
the characters of the adult comatulid, at various periods between five
or six months and two and one-half years breaking away from the
column and becoming a free-living feather-star.

The fully grown pentacrinoid young of different species show great
variation, and range between 15 mm. and 65 mm. in total length; in
some the column is very short and relatively stout, not more than one-
third of the total length, and composed of as few as 10 segments,
while in others it is very slender and much elongated, reaching four-
fifths of the total length and being composed of more than 65
segments.

In the comatulids and in the pentacrinites (possibly excepting
Metacrinus) with more than 10 arms the young always have Io arms

a

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

only until a considerable size is reached ; increase in the number of
arms is accomplished by breaking off each of the original 10 arms ©
at the articulation between the first and second (more rarely between
the third and fourth) brachials and forming on the stump an axillary
ossicle from which two arms arise, one or both of which may still
further divide.

REGENERATION

In the crinoids, especially in the feather-stars, the replacement of
lost parts by regeneration reaches an unusual extreme. It is most
highly developed in the comatulids, pentacrinites and bourgueti-
crinites, and least in the Plicatocrinide and, so far as we know, in
Holopus.

In the pentacrinites it is very common in Metacrinus, nearly as
common in Jsocrinus, less noticeable in Endoxocrinus, and relatively
rare in Comastrocrinus and in Hypalocrinus. This is not necessarily
the result of a greater inherent ability to regenerate on the part of
Metacrinus and Isocrinus, but is correlated with the fact that they
run up into the shallowest water where the wave action makes itself
felt, for the proportion of regenerated individuals decreases rapidly
with depth regardless of species.

Although inhabitants of deep water the species of Jlycrinus and
Bathycrinus lose the radials, arms and visceral mass very readily and
specimens are frequently found regenerating from the basal ring.
Similarly the species of Bythocrinus, Rhizocrinus and Democrinus
lose their arms so easily that it is very difficult to secure individuals
with the arms still attached to the basiradial cone.

The family Apiocrinide as represented in the recent seas is known
from two specimens, each representing a different genus. One of |
these (the type of Proisocrinus ruberrimus) is perfect, the other
(the type of Carpenterocrinus mollis) is only a fragment. Similarly
the family Phrynocrinidze is known only from two specimens each
representing a different genus. One of these (the type of Phryno-
crinus nudus) is perfect, though the arms were broken during cap-
ture, while the other (the type of Naumachocrinus hawatensis) lacks
the arms beyond the radials and the terminal stem plate.

Among the comatulids regeneration is very common in all littoral
species, but with increasing depth it becomes less and less frequent.
Broadly speaking it appears to occur to a much greater extent in the
Macrophreata than in the Oligophreata, though partially regenerated
cirri have been mostly recorded in the latter. It is quite possible that
this is connected with the large chambered organ of the Macrophreata ;
and it is also possible that it is the small size of the chambered organ

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK 13

which to a large degree limits the extension of the pentacrinites and
such comatulids as the Thalassometrine and the Charitometridz
toward the surface, since as a result of wave action breakage is most
common in the littoral and without a large chambered organ a crinoid,
unless unusually tough, could not repair its injuries with sufficient
rapidity to survive.

The most common mutilations are the loss of the visceral mass and
of more or less important portions of the arms, the fracture in the
latter case almost invariably occurring at a syzygy. Loss of the
visceral mass appears to cause the animal no inconvenience whatever,
and it is entirely replaced in a little more than three weeks.

ASYMMETRY

There are four types of deviation from the normal pentamerous
symmetry of the crinoids. These follow the following lines:

1. A rearrangement of the five primary groove trunks upon the
disc whereby (a) the left posterior increases in size and gives off
more branches than any of the others; (b) as a result of the anterior
migration of the mouth the two posterior become much longer and
the anterior much shorter than the others and a condition of bilateral
_symmetry is attained; (c) correlated with the anterior migration of
the mouth, all of the primary groove trunks become merged into a
horseshoe-shaped ring which skirts the lateral and anterior borders
of the disc, giving off branches to the arms, the mouth being in the
right center of the ring so that the ambulacra on the left are more
developed than those on the right, or the ambulacra leading to the left
posterior ray disappear altogether so that the ambulacra on the right
are more developed than those on the left.

. 2. A dwarfing, or an overdevelopment, of the left posterior, more
rarely of both posterior, radial with the accompanying post-radial
series.

3. The intercalation of additional radials and post-radial series
which alternate with the original five, and the associated dropping out
of one of the five radials.

4. The suppression of two of the primarily five basals.

Asymmetry is almost universal in the comatulid family Comas-
teridz, which includes the most specialized of all the recent types; in
this family the first and second types occur, though the latter is much
less common.

Asymmetry is characteristic of the genus Promachocrinus, which
is probably rightly considered as the most specialized genus in the
subfamily Heliometrinz ; here the first and third types occur.

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

Asymmetry is equally characteristic of the genus Thaumatocrinus,
the most specialized genus of the family Pentametrocrinidz ; in this
genus the third type is found.

Asymmetry exists in all of the recent genera of the Plicatocrinide,
the first, second and fourth types being represented; the third also
occurs in the fossil representatives of this family.

Asymmetry is characteristic of both of the recent genera of
Apiocrinide, which are the most highly specialized genera in the
family ; in these the second type occurs.

Asymmetry of the second type is characteristic of Holopus.

Asymmetry characterizes both of the species of Rhizgocrinus exist-
ing in the present seas, and one of the species of Monachocrinus; in
these the third type is found. ©

It appears that, no matter in what form it may manifest itself, in
the recent crinoids asymmetry is an attribute of the most special-
ized types in the groups in which it occurs. From the conditions in
the Plicatocrinide, the last remnants of the once abundant Inadunata,
it would appear that asymmetry is an attribute of phylogenetically
decadent types—types in which type senescence has so far advanced
as to inhibit the normal course of development. .

Although occurring everywhere except in the Arctic Ocean and in
the Mediterranean, Bering, Okhotsk and Japanese seas, asymmetrical
types are most frequent and most highly developed (1) in warm and
shallow water from southern Japan southward throughout the
Malayan archipelago to northern Australia and thence westward to
Ceylon, and (2) in the Antarctic and in the cold abysses.

Though present among species inhabiting the western Atlantic
from North Carolina to Brazil, and characteristic of many forms
living at intermediate depths in the western Pacific and in the Indian
Ocean, in these it is never more than slightly developed, even though
they may be very closely related to types in which it is, in other situa-
tions, carried to an extreme.

Briefly stated among the recent crinoids any wide departure from
the normal close approximation to true pentamerous symmetry indi-
cates unfavorable conditions of one or other of two main types which
are not mutually exclusive ; these two types are:

1. Internal unfavorable conditions, induced by incipient phylo-
genetical degeneration through type senescence, as in the Plicato-
crinidz which in the recent seas represent the otherwise almost ex-
clusively palaeozoic Inadunata ;. and

2. External unfavorable conditions, taking the form of

a. Phylogenetically excessive cold, and
b. Phylogenetically excessive warmth.

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK 15

THE COMPOSITION OF THE CRINOID SKELETON

The skeleton of the crinoids has the composition of a moderately
magnesian limestone, the proportion of magnesium carbonate to
calcium carbonate appearing to be a function of temperature and
rising from 7.26 per cent in the coldest water to as high as 13.74 per
cent in the tropical littoral. A trace of phosphate of lime appears
always to be present, but whether or not it is an essential constituent
is uncertain.

THE DISTRIBUTION OF THE CRINOIDS

The modern distribution of the crinoids has been chiefly the result
of the gradual differentiation in the conditions of different sections of
a once uniform sea area resulting in the evolution of more or less
distinct faunal units through the selective extirpation of different
types in different regions; combined with this there has been under
the changing conditions the evolution of a few new types and the
further specialization of others, some of which, efficient and aggres-
sive, have apparently extirpated the previous crinoidal inhabitants
from all the regions into which they have been able to extend their
range.

Changing geological and meteorological conditions affect chiefly the
shallow water of the littoral and sublittoral regions ; the intermediate
depths are but slightly affected, and the abysses not at all. Thus
faunal differentiation is most marked in shallow water, much less
marked in water of intermediate depth, and but vaguely indicated
in the abysses.

One of the results of the gradual change in conditions in a once
nearly uniform ocean has been the discontinuous distribution of many
.marine types occurring in two or more widely separated localities
which have been subjected to less modification than the intermediate
regions but which are now separated by impassable thermal or other
barriers. This is well brought out in many marine types and in the
crinoids is illustrated by the curious correspondence between the
fauna of Australia and that of the Caribbean Sea.

After the differentiation of a fauna land barriers sometimes appear
isolating a certain section from the main range. It is thus that we
account for the reappearance of Arctic types in the Okhotsk and
Japanese seas.

Geographically the littoral crinoids are divisible into a number of
more or less well-marked faunas corresponding in the main to those
indicated by other marine types. In those regions where the tem-
perature of the water is more or less uniform from the surface to

2

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

great depths, as in the Arctic, the Antarctic, the Mediterranean, etc.,
the littoral crinoids descend with it and there is then a uniformity
in the crinoid fauna from the surface almost or quite to the abysses ;
but where the temperature decreases rapidly with depth, as along the
tropical shores, the bathymetric extension of the littoral types is very
limited and the littoral fauna is underlain by an intermediate fauna
consisting for the most part of more ancient and more conservative
types which is fairly uniform the world over, though it reflects the
broader divisions of the local overlying faunas. It is from this inter-
mediate fauna that the littoral faunas on the one hand and the abyssal
faunas on the other appear to have been derived.

The distribution of the crinoids appears to be governed entirely by
temperature, pressure playing no part whatever.

There seems to be a close correlation between the bathymetric (or
thermal) range and the geographical range of the same type, a form
with a, restricted bathymetrical range having a similarly restricted
geographical range and the reverse.

THE PALEONTOLOGICAL HISTORY OF THE LIVING CRINOIDS

Of the four great divisions of the crinoids, the Camerata, the
Flexibilia, the Inadunata and the Articulata, two, the Inadunata and
the Articulata, are represented in the present seas.

The Inadunata are almost entirely confined to the Paleozoic, rang-
ing from the Ordovician to the Carboniferous, with one of the 18
gamilies (Poteriocrinide) represented in the Trias and another
(Plicatocrinidz) known only from the Upper Jurassic and from the
recent seas.

The Plicatocrinide (including the living genera Calamocrinus,
Ptilocrinus, Thalassocrinus, Gephyrocrinus and Hyocrinus and the
fossil genus Plicatocrinus) are at present entirely confined to the deep
and cold abysses ; all of the species are very rare, most of them having
only been dredged once and none of them more than twice. The
Jurassic representative of this family is also very rare, and its remains
are commonly associated with those of hexactinellid and lithistid
sponges so that it also probably lived at considerable depths. No
representatives of the Plicatocrinide are known from the Cretaceous
or from the Tertiary.

The Articulata are entirely post-paleozoic, all from the Jurassic or
later except the pentacrinites, which are also represented in the Trias.

The Phrynocrinide are not represented as fossils; the ranges of
the other families and of the genera including both recent and fossil
species are as follows:

NO. vi SEA-LILIES AND FEATHER-STARS—CLARK 7,

Bourgueticrinide: Upper Jurassic to Recent.
Rhizocrinus: Cretaceous (New Jersey); Eocene (Europe); Recent.
Democrinus: In a recent breccia at Guadeloupe which also contained a
human skeleton; Recent.
Apiocrinide: Jurassic, Cretaceous and Recent.
Pentacrinide: Trias to Recent.
Isocrinus: Triassic and Jurassic; Recent.
Zygometride: Jurassic to Recent.
Catoptometra: Jurassic to Recent.
Eudiocrinus: Jurassic to Recent.
Holopodide: Jurassic to Recent.
Holopus: Tertiary to Recent.

fie) POSSIL REPRESENTATIVES OF THE .RECENT CRINOID
GENERA

Only a very few crinoid genera include both recent and fossil
species, and these are divided into two groups, (1) those occurring
only in the western Pacific (Eudiocrinus, Catoptometra, Proisocrinus
and Carpenterocrinus, the species of the two last being apparently
congeneric with certain species of Millericrinus) and (2) those con-
fined to the western, or western and boreal, Atlantic (Jsocrinus,
Rhizocrinus and Holopus). The recent distribution of these types
is thus seen to be the same as that of the living king or horseshoe
crabs (Xiphosuridz).

Excepting only Proisocrinus ruberrimus and Carpenterocrinus
mollis, all of the species of these genera are chiefly represented in
shallow water, and these two are the only ones which do not occur
within 100 fathoms of the surface ; indeed the species of three of the
five other genera are entirely confined to water of less than 155
- fathoms depth.

The maximum representation is between the shore line and 200
fathoms, especially between 50 and 150 fathoms; as, taking the ocean
as a whole, we find at a depth of 200 fathoms a temperature of
50.1° F. and at 100 fathoms a temperature of 60.7° F. it is evident
that these genera are most abundantly represented within the opti-
mum temperature for crinoid life, which is between 50° and 65° F.
It is interesting to observe that (excepting for Proisocrinus and
Carpenterocrinus, each only known from a single dredge haul at tem-
peratures of 36° and 4o° F.) the only increase in the numbers falls
between 50° and 64°, that is, within the optimum temperature for
crinoids, and is particularly emphasized between 56° and 64°, the
emphasis within the optimum temperature range being between 60°

and 65°.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

THE COURSE: TAKEN: BY SPECIALIZATION AMONG iis
CRINOIDS

The dominant feature of the progressive specialization among the
crinoids from the earliest times to the present day has always been
a process of progressive simplification in structure, the result of a
process of progressive atrophy or suppression affecting some part or
other of the organism. Thus the more specialized types differ from
the more generalized through the atrophy or suppression of some
important structural element, while the later groups are differentiated
among themselves according to the lines which this atrophy or sup-
pression has followed.

The (recent) Articulata are distinguished from the Inadunata by
the sudden cessation of stem growth (with an apparent, though not
real, exception in the pentacrinites) after the stem has attained a
definite and fixed length, and by the extreme atrophy of the calyx
involving in most cases the complete disappearance of certain essential
elements ; the comatulids are differentiated from all the other (recent)
types by the suppression of the column excepting for the proximal
or topmost columnal which becomes permanently attached to the
calyx; Holopus is differentiated from all other (recent) genera by
the suppression of the column, the infrabasals and the basals, the
stalk being formed by the coalesced and elongated radials; the
Phrynocrinide differ from the Bourgueticrinide in the complete sup-
pression of the radicular cirri, and the Bourgueticrinide differ from
the Phrynocrinidz in the suppression of the terminal stem plate.

THE OCCURRENCE OF LITTORAL CRINOIDS

I-xcept on sandy and exposed muddy shores littoral crinoids occur
in all possible situations. Their one essential requirement is pure,
well-aerated water having a relatively high minimum salt content and
well provided with minute plankton organisms, and wherever this
condition is met within the range of the littoral species they may be
looked for in the water just below the low-tide mark or in protected
situations ; sometimes they even occur in regions left bare at low tide.

Along the shores of the Indian Ocean from southeastern Africa,
Madagascar and Mauritius to Suez, India and the Malay archipelago,
along the coasts of Australia, especially in the north, and thence north-
ward to Fokien and southern Japan, littoral comatulids of many
species are abundant—about 30 are known from Singapore alone—
particularly on reefs and rocky shores, less commonly in sheltered
situations and in eelgrass, though their occurrence is commonly more
or less local and they are frequently not to be found in apparently

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK 19

ideal places. A few species have been found by shore collectors in
New Caledonia, Lord Howe Island, Fiji, Samoa, the Marshall, Gil-
bert (Kingsmill), Pelew, Caroline, Society and Hawaiian Islands,
but throughout Oceania they appear to be relatively rare. None are
known from the Japanese coasts north of Tokyo Bay, from the Asiatic
coasts north of Fokien, from the northern or eastern shores of the
Pacific, or from New Zealand.

In the Atlantic basin littoral crinoids occur from Scandinavia and
Great Britain to the Gulf of Guinea, including the Mediterranean
area (but not the Black Sea) and the islands of the European and
African coasts, and in the region of Cape Town, and in the west from
the Bahamas and Florida to southern Brazil; but in the west they
are extraordinarily rare, there being, except in the case of Tropio-
metra, but six records, one from the Bahamas (Nemaster), one from
the Tortugas, Florida (Nemaster), one from St. Thomas (Antedon),
one from Dominica (Nemaster) and two from Brazil (Nemaster
and Antedon).

Of all the comatulids the preéminently littoral genus is Tropiometra,
and wherever this genus occurs, from South Africa to Australia,
Oceania and southern Japan, and from the southern Caribbean to
south Brazil and St. Helena, it is commonly found along the shores,
often in great abundance. In the western Atlantic, from Tobago,
Trinidad and Venezuela to southern Brazil, it is the only really
common littoral form.

A close second to Tropiometra is found in the genus Antedon,
ranging from Scandinavia and Great Britain to the Gulf of Guinea,
including the offshore islands and the entire Mediterranean basin,
and also found from St. Thomas to Brazil, all the species of which
occur along the shores, where they are often locally abundant. Only
two specimens of the American species are known, one from shore
collections at Rio de Janeiro, the other from shore collections at
St. Thomas.

The species which have actually been captured along the shore
number no less than 152, representing 38 genera and 12 families and
subfamilies, while 93 more undoubtedly occur there, making a grand
total of 245 shore-living types already known. Of these 227 are
from the Indo-Pacific region (including two from South Africa),
eight are from the region between the Bahamas and Florida and
Brazil, six are from the northeastern Atlantic north of the Gulf of
Guinea, and four are from southern Australia.

The favorite localities for shore-living comatulids are more or less
shaded situations, holes and crevices in reefs, benéath stones; in half-

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

submerged caves, on piling beneath wharves, and in the irregularities
on the outer side of breakwaters; but they are sometimes found
among gorgonians or eelgrass, on mangrove roots, and occasionally
on mud.

THE RELATION OF THE COMATULIDS TO TEMPERATURE

In the recent seas the comatulids range from the very warm water
of the tropical littoral to water with a temperature of only 28.7° F.,
considerably below the freezing point of fresh water.

The species of the genera of the Oligophreata are especially de-
veloped in the warm waters of the present seas, and they are pecu-
liarly characteristic of the warm waters of the tropical coasts. The
species which occur in this warm water are almost without exception
highly specialized, and they are especially remarkable for a great
reduplication in the number of their arms, of which they may have as
many as 150 or even more, and also for their large size.

A study of the ontogeny of the most extreme of these types shows
that the essential characters of the adults appear at an extraordinarily
early age, and also suggests that these characters do not indicate a
true phylogenetic progress which will eventually lead to the evolution
of new types, but rather a more or less pathological hyperdevelop-
ment, an abnormal exaggeration of the normal phylogenetical ten-
dencies, which will lead nowhere, but will terminate simply in the
extinction of the species in which it appears. The fundamentally
aberrant or unbalanced nature of these types is strongly indicated by
the invariable conservation of some primitive character, for example
spiny borders on the brachials, a reguiar distribution of the syzygies_
in the arms, a very primitive type of cirri or of pinnules, etc.

The species of the genera of the Macrophreata are mostly developed
in the colder waters of the recent seas, and this suborder includes all
the comatulids of the polar regions and of the abysses. The species
which are found in very cold water are almost without exception very
primitive, and they are especially remarkable for a reduplication of
the radials, a conservation of the carination of the ossicles of the
division series and of the arms, a conservation and an exaggeration
of the spines which ordinarily are found only among the young, an
abnormal shortness of the brachials and of the segments of the
pinnules and of the cirri, as well as for their very large and very
primitive pentacrinoids in which the radianal plate approaches its
original position beneath the right posterior radial; the greater part
of these species are remarkable for their very large size, and they
include among their number the largest living crinoids known; their

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK — 21

arms are never more than Io in number except in the forms with 10
radials, which may have 10 or 20.

A careful study of these types, together with a detailed comparison
between their characters and the characters shown by other species
found in water abnormally cold for their immediate phylogenetic
stock shows that the apparently primitive characters are without
doubt the result of a repression or an inhibition of the normal
phylogenetic development; furthermore, in combination with these
characters we always find other characters which indicate a condition
of very marked specialization, as for example a large number of
proximal pinnules which are provided at their tips with more or less
developed terminal combs, both of which characters are otherwise
only found in the comatulids of very warm water at the opposite end
of the temperature scale, and a great specialization of the centro-
dorsal.

The species of very cold water thus resemble the species of very
warm water in the possession of a fundamentally aberrant structure,
for they preserve and exaggerate certain very primitive characters
while at the same time they show a high degree of specialization along
other lines.

However extraordinary it may appear, in their unbalanced type of
specialization the comatulids of the coldest water agree more nearly
with the species inhabiting very warm water—nearly all of which
belong to the other suborder—than with any of the species of the
intermediate waters, and the largest species, the smallest species, and
the species with the greatest number of segments in the arms, pinnules
and cirri are found equally at both extremes of temperature.

The Oligophreata and the Macrophreata are both represented by
six families between the temperatures of 50° and 55° F., but the
Oligophreata predominate at all temperatures above this, while the
Macrophreata predominate at all temperatures below.

In the Oligophreata the greatest number of families is found be-
tween 60° and 65°, and in the Macrophreata between 50° and 66°.
For all the comatulids the greatest representation is between 55° and
65°, with the emphasis on 60°-65°.

It would therefore appear that the temperature range included
between 55° and 65° represents the temperature phylogenetically
most suitable for the recent crinoids. It is a very curious fact that
the comatulids found between 55° and 65° are all of medium size,
none very large and none very small, and that they all show well-
balanced and conservative characters.

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

So far as we can see it is with the recent species which are found
within these temperature limits that the fossil crinoids best agree,
and one might hazard the guess that it was principally, if not entirely,
within these temperatures that the crinoids of the post-palzeozoic
faunas, characterized by a very great development of the Articulata,
were developed.

FOOD

Duchassaing records that the stomach contents of a specimen of
Isocrinus decorus which he fished up in relatively very shallow water
at Guadeloupe consisted only of the remains of small crustaceans.

Bronn, summarizing previous ‘accounts, wrote that the stomach
contents of Jsocrinus were made up of the remains of small crus-
taceans, while those of the comatulids consisted of diatoms such as
Navicula, Bacillaria, Actinocyclus and Coscinodiscus, of Tethya, and
of many types of entomostraca.

W. B. Carpenter said in 1866 that in the very numerous specimens
of Antedon bifida from Arran of which he examined the contents of
the digestive cavity he never found anything other than microscopic
organisms, and the abundance of the horny rays of Peridinium tripos
made it evident that in this locality that organism is one of the
principal articles of food. But in specimens from other localities he
found a more miscellaneous assemblage of alimentary particles, the
most commonly recognizable forms being the horny casings of ento-
mostraca or of the larve of higher crustaceans.

In his account of Hyponome sarsw (the visceral mass of Zygometra
microdiscus) Lovén states that in the ambulacral grooves he found
masses consisting of minute crustaceans, larval bivalves, and other
remains of food.

In 1876 W. B. Carpenter wrote that the contents of the alimentary
canal of Antedon bifida both in the pentacrinoid stage and in the adult
consists of minute entomostraca, diatoms, spores of algz, etc., but
in his Lamlash specimens especially of Peridinium tripos, which was
usually very abundant in that locality. He also notes that the contents
of the alimentary canals of the various types of pentacrinites ex-
amined by him are of the same nature.

P. H. Carpenter says that the food of a crinoid is considerably
varied in its nature according to the character of the sea bottom on
which it lives. The horny casings of entomostraca and the larve of
larger crustacea are frequently to be found in the digestive tube
together with the frustules of diatoms, spores of alge, etc. In sec-
tions of Bathycrinus, Rhizocrinus, Isocrinus and Endoxocrinus from
deep water the silicious skeletons of radiolarians may be found in

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK 23

considerable abundance and variety. Foraminifera also form a staple
article of food for these deep-sea species, for he frequently found
Globigerina, Biloculina and other types beneath the covering plates of
the food grooves on the arms and pinnules, while the remains of their
soft parts occur in the intestines of decalcified specimens.

Seeliger believes that he recognized in the earliest food of the larve
of Antedon adriatica half-digested infusorians and different pelagic
larve. Bury found the stomach of the very young pentacrinoids of
Antedon mediterranea so filled with diatoms that the cutting of sec-
tions was rendered very difficult.

Dr. Edwin Kirk states that in the case of a number of specimens of
Comanthus japonica which he examined the contents of the intestine
were almost wholly comminuted animal matter.

At Maer Island, Torres Strait, Dr. H. L. Clark examined the
stomach contents of four comatulids (species undetermined). He
found that in each case the greater part of the food material was
green algz, chiefly unicellular though some linear forms (thread
alge) were also noted; a few diatoms were detected, and some
foraminifera. In one of the stomachs several radiolarians were seen,
in another a piece of a red alga, and in a third some fragments of
minute crustaceans. Dr. Clark also examined the stomach contents
of Tropiometra picta at Tobago which he found to consist of a
mixture of vegetable and animal food, the former predominating.
The plants were diatoms and unicellular green alge, with occasional
fragments of seaweeds ; of animals, crustaceans were most frequently
noted, but a few foraminifera were also seen; the crustaceans were
minute amphipods, copepods and crab zozas.

Dr. Th. Mortensen found that a relatively large percentage of the
pentacrinoids of Jsometra vivipara have in their stomachs the half-
digested, but still perfectly recognizable, remnants of the larve of
the same species; he even found very young pentacrinoids with the
vestibule recently ruptured and the arms not yet developed with
embryos almost as large as themselves in their mouths. He remarks
that on account of the large number of pentacrinoids found attached
in clusters to the tips of the upturned cirri—as many as 99 in one
specimen—this danger to the embryos is very real, and probably quite
a large number of them perish in that way.

LOCOMOTION
Excepting for the pentacrinites all of the stalked crinoids are
firmly attached to foreign objects or rooted in the mud and therefore
incapable of locomotion. The pentacrinites have such long and heavy

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

stems which are usually so entangled with the objects on the sea floor
and with the stems of other individuals that they are to all intents
and purposes as firmly fixed as are the other stalked types. But the
comatulids, attached by their highly mobile dorsal cirri, are able to
detach themselves and move about, though as a rule they remain
pretty constantly in one place and rarely change their position except
as a result of some unusual stimulation.

Among the comatulids locomotion is of two types, swimming and
crawling; swimming is the more usual, and apparently all littoral
comatulids can swim. In swimming the arms of each pair beat the
water alternately and at first quite rapidly, as much as I00 times a
minute, but the animal soon tires and the longest distance recorded as
covered by a swimming comatulid is less than 3 meters. The swim-
ming of the feather-stars has been likened to the flitting of small
birds in shrubbery as contrasted with sustained flight.

The young of two species, Dorometra nana and Comanthus parvi-
cirra (twice) have been captured while swimming at the surface from
ships at anchor or in plankton hauls.

Crawling is accomplished by a combined pulling and pushing, and
in the comasterids the long anterior arms are extended forward and
used for pulling while the short and stout posterior arms are used
for pushing. The rate of progress has been calculated as 85 mm. a
minute, or 5 m. an hour.

COLOR

Of all the animals in the sea there are none that exceed in beauty
and variety of coloration the shallow water crinoids. Flower-like in
form and almost flower-like in the fixity of their habit, they are also
flower-like in the variety and distribution of their pigments. But
with depth the diversity of hue diminishes so that we find the color
range of the species of the deeper water relatively restricted while
the individuals themselves, losing the almost universal spottings and
bandings of the littoral types, become comparatively plain.

Though crinoids resemble flowers in the diversity and brilliance of
their colors, their color types are quite the reverse of flower-like. In
a particolored flower the center or eye is more or less abruptly lighter
or darker than the remaining portions, or the petals are longitudinally
striped ; cross-banding of the petals is very rare. Among the crinoids
particolored specimens are usually cross-banded, regularly or irregu-'
larly, and though the tips of the pinnules may be of a different color
from that of their bases a true longitudinal striping of the arms is
very rare and a conspicuous eye is never developed.

NO. :7 SEA-LILIES AND FEATHER-STARS—CLARK 25

The published records show in the feather-stars the following
frequency for the various colors; the numbers in parentheses show
the individuals entirely of the color given, the others all the records
for that color both in unicolor and in variegated types:

LCG 194 (47) Hs lhe acre gaa o ee aes che « po ol 19)
SHO NVA ehes crac sic 3,400 <5 138 (24) Oranges coe its oe es 390 (5)
NWalnit leans Aiciccartescsuacane shee s 126 (1) Giicye Persie ae eestor Doe! (Te)
OEE ee SR Pais wea dy oe 80 (8) WiGleE rr eeweesee kee eh 15.) (6)
GRECIIgre keane eee n ees FAC 73) Stein, Pee perpen se tNaeteis oars 15 (0)
EE Dlegetiem a ckiso stones S 70 (10)

While white and yellow occur in all possible combinations, orange
does not occur with violet, black or green; red does not occur with
black; purple does not occur with black, gray or green; violet does
not occur with orange, black, gray or brown; black does not occur
with orange, red, purple, violet, gray, green, or brown; gray does not
occur with purple, violet, black, green or brown; green does not occur
with purple, black or gray ; and brown does not occur with red, violet,
black or gray.

It is of course true that this apparent incompatibility of colors is
partly due to a lack of observations and to a misinterpretation of the
colors as recorded, and the foregoing list must therefore be accepted
with considerable reservation.

In particolored comatulids the distribution of the colors on the
arms and pinnules falls into several well-marked types, which may be
arranged as follows:

I. Arms and pinnules uniform in color.

a. Arms and pinnules uniform in color, but differing in
color from the cirri.

b. Arms and pinnules uniform in color, but the ventral and
dorsal surface of different colors.

II. Pinnules of a different color from that of the arms.

a. The distal portion of the pinnules (and usually also the
arm tips) is of a different color from that of the proxi-
mal portion and the arms.

b. All of the pinnules are of a color different from that of
the arms.

III. Arms and pinnules irregularly nea mottled and blotched.
IV. Arms with more or less regular and uniform spots.

a. Spots confined to the division series and arm bases.

b. Numerous small spots generally distributed.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

V. Arms with a median dorsal stripe. :
a. A broad median stripe, lighter or darker than the color
on either side of it.

b. A narrow median stripe, always very dark.

VI. Arms conspicuously and regularly cross-banded.
a. With several more or less irregular broad bands.
b. With a broad approximately central band.
c. With broad alternating bands of equal width.
d. With narrow alternating bands of equal width.
e. With narrow well-spaced bands.

As a general rule the coloration of the pinnules follows that of
arms, but in a few types they are banded on each segment and in at
least one case this is a good specific character.

The cirri are usually unicolor, less commonly dull at the base,
becoming gradually or abruptly brighter distally, and rarely show
distinctive color types; when this occurs the color types are usually
a reflection of the color types of the division series and arm bases and
very rarely distinctive.

Particolored cirri, except those which are merely brighter distally,
fall into the following classes:

I. Each cirrus segment with a transverse band.
II. Cirri unicolor, but of a color not found in the calyx or arms.

III. Cirri blotched or spotted.

a. Cirri with irregular blotches or irregular bands.
b. Cirri with small spots.

IV. Cirri longitudinally striped.

A study of the development of the colors seems to indicate that all
colors except gray and black may arise directly from white, and that
yellow, red and brown usually arise directly from white, while violet
arises as often from white as from any other color. All colors may
develop from yellow ; gray and black are developed only from yellow,
and purple and green arise more frequently from yellow than from
any other color. Red frequently develops into purple, and occasion-
ally into yellow, violet and green. Brown develops into green and,
less frequently, into violet. Orange develops into brown...

At a depth of 55 fathoms in the sea the relative proportion of the
red rays is considerably diminished, and at 300 fathoms they have
almost completely disappeared. In a discussion of colors, color com-
binations and color types and their relation to depth, therefore, it
would seem that bathymetric divisions of 0-55 fathoms, 55-300
fathoms, 300-600 fathoms and 600 fathoms and over would be as
satisfactory as any.

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK 27,

A tabulation of the colors according to depth shows that black does
not occur below 55 fathoms; red, violet, gray and white do not occur
below 300 fathoms; purple, green and orange do not occur below
600 fathoms; yellow and brown occur at all depths.

Yellow and brown are relatively much more frequent below 300
fathoms than above; purple, green, orange, gray and white occur in
about the same relative proportions down to 300 fathoms, at that
point decreasing abruptly or disappearing altogether ; red, violet and
black decrease abruptly below 55 fathoms.

No colors are more frequent between 55 and 300 fathoms than
elsewhere. The proportion of uniformly colored species increases
markedly with depth, while the corresponding decrease in variegated
species is even more abrupt. All of the color types given occur only
above 300 fathoms. The median dorsal stripe is most common be-
tween 55 and 300 fathoms. Distinctively colored pinnules and regu-
lar crossbands occur in about the same proportion down to 300
fathoms. Distinctively colored cirri and irregular spotting or mottling
are much more common above 55 fathoms than below. Regular
spotting occurs only above 55 fathoms.

Whereas the development of color in the crinoids seems to have a
more or less definite relation to illumination, it appears to have no
relation whatever to the temperature of the water in which the
crinoids live. The various color types are almost entirely confined to
water of high or intermediate temperature; but this is undoubtedly
due to the fact that the crinoids of the colder parts of the oceans
belong to groups in which color patterns are not developed even in
their tropical representatives. This supposition is emphasized by the
‘occurrence of several well-marked and beautiful color types in the
species of the genus Antedon which are quite as well developed in the
Scandinavian species as in those inhabiting the shores of northwestern
Africa and the Mediterranean.

From the evidence at hand the following conclusions seem justified :

White, which is the original color of the pentacrinoid young and
occurs frequently in the adults, denotes the more or less complete
absence of pigment.

Yellow is the color of practically all of the more primitive forms,
and of many of the more specialized, throughout life, and with very
few exceptions (occurring in the brilliantly illuminated littoral) of
small specimens and of advanced pentacrinoids. Orange or red, in
reality an intensification of the yellow, is the color of a few primitive
forms, and of nearly all the young which are not yellow.

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

After the full size has been reached a dusky factor makes its
appearance which may be generally diffused or more or less localized,
and deepens, alters, or obscures the original colors.

Illumination of the habitat results in an intensification of the natural
colors and the very early appearance of the dusky age factor, as well
as in the appearance of a blue factor resulting in the formation of
bright greens, purples and violet, which may deepen to black.

All the comatulids living below the limit of light penetration exhibit
the basic colors, white, orange or red, only, which, though they may
become more intense toward the calyx and arm bases, are never other-
wise diversified. Each of these colors, however, may be modified by
the dusky age factor, resulting in “ dusky purple,” greenish or brown-
ish yellow, brown, orange brown, crimson, or red brown. The blue
factor is absent in these species, but in the group as a whole it gradu-
ally increases from the limit of light penetration to the surface,
causing the appearance of greens, purples and violets of increasing
intensity. |

Geographically the maximum development of color diversity ap-
pears to be in the Malayan and north Australian region, and thence
westward to Ceylon; but it is here also that the maximum develop-
ment of littoral types is found. The whole littoral and intermediate
fauna from east Africa to Oceania and southern Japan is notable for
the diversity in the coloration of the endemic forms.

On the other hand, throughout the vast extent of the east and north
Pacific we find the minimum diversity of crinoid coloration; all of
the comatulids are unicolor, most of them yellow, becoming yellow
brown, a few purplish brown or red; all of the stalked forms are
yellow.

The crinoids of the Caribbean Sea as we know them to-day are
much less highly colored than those of the Indo-Pacific region, and
this holds good for stalked as well as for unstalked types. But here
the groups which furnish the majority of the most variegated species
are absent. In the remaining portions of the Atlantic, outside of the
region of the Cape of Good Hope where the Indo-Pacific fauna in-
trudes for a short distance, we note especially the presence of the
highly colored species of Antedon, which range collectively from Rio
de Janeiro to St. Thomas and from the Gulf of Guinea to Norway,
including the Mediterranean basin; of the green or white species of
Leptometra which occur from Madeira to Scotland, including the
Mediterranean basin; and of the small green or gray species of
Hathrometra which are found from Chesapeake Bay and Portugal
northward.

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK 29

The coloring matter of crinoids is freely soluble in fresh water and
in alcohol. It is possible to keep certain species for some time in
water fresh enough to dissolve out a considerable amount of pigment
without apparent injury, while many may be partially decolorized in
a stream of fresh water while still alive.

. As a general rule comatulids preserved in alcohol, no matter what

their original colors may have been, become brown, usually a yellow-
ish, more rarely a purplish, reddish or greenish brown, later slowly
fading out to grayish white. The bands and spots often persist for
some time, though with entirely changed color values, but they
eventually disappear. On account of the wonderful diversity of the
colors in life and of the altogether extraordinary alteration of the
colors by preservation the greatest care is necessary in identifying
living specimens, especially from descriptions based upon preserved
material, for the color may or may not be a good specific index; it
usually is not.

THE SIMILARITY BETWEEN CRINOIDS AND PLANTS

Although they are animals possessing a relatively high type of
organization the crinoids are so plant-like in their outward form that
it seems worth while to explain briefly the extent of and the reasons
for this curious and striking similarity.

The roots of the stalked crinoids are of several different types
varying from a large encrusting mass with digitiform processes about
its borders to-a long slender taproot buried in the mud from which
very numerous delicate lateral roots are given off. Every type of
crinoid root can be matched among the plants, though the crinoid root
performs only one of the functions of the plant root, and that is to

‘hold the organism in place.

The stem of some of the stalked crinoids, such as Proisocrinus, is
long, smooth, slender, and enlarged toward the base, and thus
strikingly similar to the stems of many of the commonest palms, this
similarity being heightened by the numerous pinnate arms like palm
leaves at the summit. The pentacrinite stems with their whorls of
five cirri at regular intervals call to mind the stems of many plants
with narrow whorled leaves, in combination with their lily-like crowns,
especially such lilies as Lilium philadelphicum. From their resem-
blance to palms the stalked crinoids are commonly called “ sea-palms ”
in French and Spanish, while their usual appellation in English is
“ sea-lilies.”

The food of the crinoids consists of the minute plankton organisms
suspended or moving slowly about in the surrounding water. In

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

order to obtain an adequate supply of these organisms they must
intercept the maximum amount of water without, however, impeding
its flow, for it must pass by them constantly and continuously in order
to furnish them with a supply of oxygen without which they would
soon perish, as well as to deliver to them the requisite amount of
food. As the maximum area is included within a circle the crinoids
have developed a circular food-collecting apparatus consisting of
slender pinnules which, spread out in the form of a circular net,
filter the maximum amount of water while at the same time they
interrupt the flow of water to the minimum degree. In this circular
food-collecting apparatus composed of a vast number of slender
filaments we see at once the influence of the same factors which have
determined the development of the submerged filiform-dissected
leaves among the water plants ; and the similarity becomes more strik-
ing still when we call to mind such carnivorous plants as the species
of Utricularia.

The crinoid crown is almost entirely a food-collecting apparatus ;
the essential organisms of the animal are reduced to a minimum and
subordinated to the development of a structure offering a maximum
area for the interception of food particles. This is not by any means
a peculiarity only of the crinoids, for all of the other fixed and
arborescent animals, the sponges, ccelenterates, polyzoans, tunicates,
protochordates, etc., have similarly subordinated, as it were, their
whole being to the specialization of the meehanism for collecting
mobile food to such a degree that they may be differentiated often
down to genera, and sometimes even down to species, by the char-
acters found in the food-collecting apparatus alone without considera-
tion of their other structures. The polyp or polypoid individual more
or less flower-like in form or else capable of maintaining a strong
inflowing current of water is a physical necessity correlated with a
fixed existence, and the contrast between the requirements of a fixed
and an active life are nowhere better illustrated than in the echino-
derms through the comparison between the crinoids on the one hand
and the echinoids, asteroids and ophiuroids on the other.

Terrestrial plants live rooted in the earth from which and from the
surrounding atmosphere they derive all the substances necessary for
their existence. But the medium about them is so light that some
special provision must be made for the fertilization of their ova.
Thus while the crinoids, and all the other fixed marine animals, have
had to specialize, so to speak, on the development of an adequate
apparatus for food collection, the plants have had to devote their
energies to the problem of securing cross fertilization. This is largely

INOS 7. SEA-LILIES AND FEATHER-STARS—CLARK Bil

accomplished through the intermediary of insects of various types,
more rarely by small birds, which transport the pollen through the air,
and the plants have developed all sorts of artifices by which they make
their flowers attractive to these creatures. The result of this neces-
sity has been to localize in the flowers the chief differential characters
of the plants just as the same characters have in the crinoids and in
the other plant-like animals been chiefly segregated in the commonly
flower-like food-collecting mechanism.

The polyps of the plant-like animals cover the maximum area with
their arms or tentacles in order to collect the maximum amount of
food, while the flowers cover the maximum area with their petals in
order to attain maximum visibility, with the common result of a
circular expanse of symmetrically arranged parts in both cases. To
increase their efficiency by mass effect the polyps of plant-like animals
are often spiked, sometimes spirally arranged on the axis, and
occasionally grouped in umbels or in imperfect racemes like flowers,
while to counteract unusual external stresses of waves or wind both
flowers (/taoulia, etc.) and polyps (brain-corals, etc.) are sometimes
in the same way gathered together in great more or less globular and
highly resistant masses.

A very large proportion of the conspicuous flowers are pentapartite,
with five sepals and five petals alternating with them, and commonly
bracts beneath the sepals; crinoids are also pentapartite, with five
basals and five arm-bearing radials alternating with them, and com-
monly infrabasals beneath the basals. Some flowers are tetrapartite,
like the crucifers; some crinoids are the same, like Tetracrinus.
Many flowers are hexapartite, as are also some crinoids, like Hexa-
crinus. The reason for the most common occurrence of five in both
cases is probably that in the pentapartite division there lies the maxt-
mum strength.

The basals of the crinoids normally enclose the visceral mass much
as in many flowers the sepals enclose the ovary, and sometimes
(Isocrinus, etc.) they imbricate over the bases of the radials as the
sepals imbricate over the bases of the petals.

In many crinoids the arm bases are firmly united by interbrachial
plates or so closely pressed against each other that they may almost
be said to possess a gamopetalous corolla.

In a few fossil crinoids (Petalocrinus and Se) all the
arms borne by each radial are united into a single, broad, flat plate
which may be highly flexible, and the crowns of these crinoids
resemble flowers to a most astonishing degree.

3

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

At the base of the petals in the flowers are the stamens, while at
the bases of the arms in the comatulids are the elongated oral
pinnules which are usually bent inward over the sometimes central
high anal tube which in many cases looks very much like a pistil.

The essential part of the plant is the flower; roots, stem and
leaves may be dispensed with in the parasitic types, but the flower
must be developed. In the crinoids the great essential is the food-
collecting apparatus ; everything else may be reduced to a minimum,
but that must remain at an irreducible maximum. In the family
Rafflesiaceze the whole plant is reduced to nothing but a flower, which
may be very large, as much as three feet in diameter ; in the coma-
tulids the animal is little else than arms and pinnules, and the diameter
of the expanded animals in one species is about three feet.

Since the spermatozoa of the crinoids escape into the sea while the
ova remain attached to the pinnules of the female it is evident that in
this group conditions exist in a way comparable to those found in
wind-pollinated plants, and it is interesting to note that the crinoids
possess many scores of entirely separate and distinct gonads arranged
in a series along both sides of each arm on the pinnules or at the
bases of the pinnules remotely suggesting the arrangement of many
wind-pollinated flowers in catkins.

The ciliation of the crinoid larvae may be compared with the
development of the fibers on the seeds of such plants as the cotton,
and the development of the long anterior tuft of cilia with the coma
on such seeds as those of the milkweed (Asclepias) or fireweed
(Epilobium).

The color of the crinoids has already been discussed, but there are
one or two points regarding color which are of interest in this
connection.

Many of the species of Comasteridz are asymmetrical, one or two
of the arm clusters being more or less, sometimes very much, shorter
than the others, the animal developing a secondary bilateral symmetry
from an original pentamerous symmetry. Many flowers also develop
a bilateral symmetry from an original pentamerous symmetry, as is
well seen in our species of Orchidacez, Scrophulariaceze, Menthacez,
etc., and in all intermediate stages comparable to those seen in the
Comasteride, in the Campanulariaceze and Solanaceze. Because of the
coiled digestive tube the visceral mass in the center of a crinoid never
shares the pentamerous symmetry of the rest of the animal, and in
the pentapartite flowers the similarly placed ovary, excepting only
in the Crassulacez, is out of harmony with the radial symmetry of the
other structures.

NOs 7 SEA-LILIES AND FEATHER-STARS—CLARK 33

Flowers facing directly upward or directly downward are always
regular, insuring maximum visibility from all directions; irregular
flowers always are directed more or less laterally. Crinoids which
rest on muddy bottoms and therefore face directly upward (as Penta-
metrocrinus ) or which have pendent crowns (as Ptilocrinus) are also
symmetrical since this means maximum efficiency in combing the
water which may pass across them in any direction.

Among our native flowers (in eastern North America) the various
colors represented fall into three distinct groups on the basis of the
proportion of irregular to regular flowers included, as follows:

Group I: AsoutT ONE-QUARTER OF THE FLOWERS IRREGULAR

Red White Green
RemCentiol teettlar HOWERSes. st. sence scr > 7 73 72
iRenecent on imregtlar flowers... s2052s00.0 056% 22 27) 28

Group II: Nearty One-HALF oF THE FLOWERS IRREGULAR

Brown Yellow Orange
Rercentyon negtlar i MOwers.ccerccces ones ss 57 50 56
Petsceut Ol itterillar HowerS....¢s6-5-.2 6. 5+% 43 44 44

GrovuP IJ]: More THAN THREE-QUARTERS OF THE FLOWERS IRREGULAR

Blue Purple
RemAceninar Tesilar NOWErsiss lc, sede alc ce Ss sel acide ds 24 21
Pe MRED COTe MEM UIA Re HOWETS «(5.2 ci ti oc bclsis fotos ajo ss 76 70

The proportion of variegated flowers is much higher in irregular
than in regular types since in the irregular types every artifice which
will increase the visibility must be adopted. The proportion of varie-
gated crinoids is very much higher in the irregular than in the regular
species because they are all from shallow water and it is in the shallow
‘ water that the colors of crinoids are best developed. From a mini-
mum in the coldest regions the proportion of irregular flowers in-
creases to a maximum in the tropics; from a minimum in the coldest
water the irregular crinoids increase to a maximum in the tropical
littoral.

About three-quarters of all our blue, violet and purple flowers are
irregular; since blue, violet and purple seem to be the most con-
spicuous colors so far as insects are concerned a flower loses less in
visibility by being of these colors than it would by being of other
colors. Blue is only recorded from irregular crinoids, and violet
and purple are much more common in irregular than in regular types,
as a result of the occurrence of the former only in shallow water.

In the crinoids the pigment is not confined to the exterior of the
animal as in most active types, including the other echinoderms, but

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

occurs more or less generally distributed throughout the body just as
pigment is distributed throughout the interior of many plants. The
coloring matter of the crinoids is, in part at least, a lipochrome, and
other lipochromes occur in a number of flowers just as indigo, occur-
ring in many different plants, is also found in the Polyzoa and other
animals, and just as cellulose or a very closely allied substance is
found in the tunicates.

Many flowers have a sweet and attractive odor, and certain crinoids
(Tropiometra and others) also exhale a pleasant plum-like aroma,
though this is not so marked as in the case of certain polyzoans
(Flustra) ; as this pervades the whole animal it is perhaps better to
compare these types with such plants as those of the families
Menthaceze or Myricaceze, most of which are aromatic. But among
the fixed animals these plants more nearly parallel the sponges in this
respect, while the sharp principle pervading the cruciferous plants
calls to mind the very acrid secretions found in the ccelenterates.

Many plants, like nettles, have stinging hairs; in the crinoids the
secretion from the glands connected with the hair-like papilla on the
tentacles appears to possess stinging qualities. Just as cattle will not
eat nettles, so the fishes carefully avoid the crinoids.

The petals of certain flowers, as in the Hypericacee, are dotted with
so-called glands containing excretory products and often arranged in
regular rows. Along the ambulacral grooves in the ¢rinoids is a row
of minute glandular bodies also containing excretory products.

Similarity of habit and the resultant similarity or at least parallel-
ism in the problems to be met have given rise to a very close corre-
spondence in many features between the fixed and sessile animals
and the plants, though the means by which this close correspondence
has been attained differ very widely in the two classes of organisms.

PARASITES AND COMMENSALS

A very large number of organisms belonging to very diverse groups
are found more or less associated with the crinoids. The relation
between these types and the crinoid hosts runs by imperceptible
gradations all the way from true parasitism, in which the organism
feeds directly upon the body tissues or fluids of the host, to the most
casual or even accidental association.

The animals associated with the crinoids may be grouped as
follows:

I. True parasites—Animals which (1) live upon the tissues or
body fluids of the crinoids and occur either (a) internally or (b)
externally; (2) occur internally, though not feeding directly upon

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK 35

the tissues of the host; or (3) while living externally upon the sur-
face of the body and not feeding directly upon the tissues or fluids
of the host are more or less permanently fixed in position and cause
more or less extensive malformations, sometimes becoming encysted.

This class includes a few “ worms,’ a number of myzostomes, a
few crustaceans, and the parasitic gasteropods.

Il. Semiparasitic commensals—Animals which feed upon minute
organisms and have to a greater or lesser extent adopted the habit of
sucking up the food particles from the streams flowing down the
ambulacral grooves of the crinoid to the mouth, or of temporarily
entering the digestive tube and feeding upon the contained matter.

This class includes the polynoid and ophiuran parasites, most of
the crustacea, and most of the myzostomes.

III. Nonparasitic commensals—Animals which, while usually, or
commonly, found living upon or among the crinoids lead an entirely
independent existence and for the most part are found living under
similar relations with other organisms.

Here are included the foraminifera, sponges, corals, hydroids,
polyzoa, barnacles, tunicates and Rhabdopleura, as well as certain
shrimps.

IV. Casual associates—Animals which normally occur hiding
among, crawling over, or attached to other usually arborescent organ-
isms (fig. 62) from which they may or may not derive nourishment,
or which normally occur attached to any available support, and which
occasionally stray among or upon, or attach themselves to, the crin-
oids, but remain otherwise entirely independent of them.

This class includes a vast number of organisms of very diverse

types.

_ As in the case of the other arborescent marine types, and in general
among the animals that live by filtering the smaller plankton from the
sea water, the crinoids are chiefly subject to indirect parasitism, that
is to say, the creatures depending upon them for their existence
appropriate the food particles which the crinoids have collected in the
ambulacral grooves, or even which they have swallowed, instead of
consuming the tissues or body fluids directly. Of the animals which
derive a part or all of their nutriment from the body or from the
efforts of the crinoids about 10 per cent are directly parasitic, and
about 90 per cent are indirectly parasitic in varying degrees.

Of the animals which are parasitic on the crinoids nearly all may
be described as casual parasites, for they belong to genera or families
other representatives of which are nonparasitic; that is to say, they
are merely particular species which have found an easy existence in

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

preying upon the crinoids, though this mode of life has not induced
any special modification of their structure.

There is a curious and interesting correspondence between the
relations of the fixed marine organisms (including the crinoids) and
their parasites and commensals and those between parasitic and epi-
phytic flowering plants and their hosts. The barnacles, most hydroids,
polyzoans, etc., correspond very closely to the epiphytic plants, espe-
cially those of the families Orchidaceze and Bromeliacee. Rhabdo-
pleura and certain hydroids are quite vine-like in habit, ascending
crinoid stems as vines do the trunks of trees. Most parasitic plants
appropriate the unelaborated sap of the host and convert it to their
own ends; most parasites of the fixed marine organisms in the same
way appropriate the concentrated but undigested microplankton in or
approaching the stomach of the host. On land most animals are
parasitized by animals of an inferior organization; but among the
fixed marine animals the parasites for the most part belong to a
phylum with a superior organization and sometimes even to the same
phylum (ccelenterates parasitic on ccelenterates, crustaceans parasitic
on crustaceans, ophiurans parasitic on crinoids, etc.). The relations
between the fixed marine animals and their parasites are thus more
nearly the same as those between parasitic flowering plants and their
hosts. On land the various animal groups are definitely parasitic or
nonparasitic ; but many plant families, such as the Scrophulariacez,
Santalacez, etc., and even many single genera, such as. Pedicularis,
Melampyrum, Gerardia, etc., include both parasitic and nonparasitic
species, just as do many families and genera, such as Synalpheus,
Periclimenes, etc., occurring with the fixed marine animals.

The three types of parasites which are of especial interest are the
gasteropods (Stilifer, Stylina, Sabinella and Melanella), Enterog-
nathus and the myzostomes (figs. 57, 58).

The family Melanellide to which Stilifer, Stylina, Sabinella and
Melanella belong includes species showing all gradations between
free-living nonparasitic types and shell-less parasites living entirely
within the body of the host. As parasites the Melanellidz occur only
upon the echinoderms, in which group, however, they are found on
species of all the classes. Most of the parasitic forms, including all
of those occurring on the crinoids, are characterized by extraordinarily
delicate shells. Some of the species are permanently fixed in one
position on the body of the host, but others, including all those found
upon the crinoids, appear to move about and to bore into different
parts of the host. It is not a little curious that, apart from Melanella
capensis and Stylina comatulicola, all the species parasitic on the

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK a7

crinoids are always attached to the calyx plates, or to the cirrals,
brachials and pinnulars instead of to the soft ventral integument.

The curious copepod Enterognathus occurs only in crinoids, but
the family to which it belongs is well known as a parasite (or com-
mensal) of the tunicates, most of the species living in the branchial
chamber of these animals.

The myzostomes form a group of very highly specialized polychzete
annelids and are the chief parasites of the crinoids, to which animals
they are almost exclusively confined. On the crinoids they are, with
one possible exception, always ectoparasitic, though they may form
soft or calcified cysts within which they are almost completely isolated
from the outer world. An organism, possibly a myzostome, has been
reported in the ovarian cavity of Notocrinus virilis. If this really is a
myzostome, which is not unlikely as similar endoparasitic species
occur in starfishes (Asterias, Stolasterias and Ceramaster) and
astrophytons (Gorgonocephalus eucnemis and G. arcticus), we find in
the crinoids the five following groups of myzostome species:

1. Wandering species which move about freely and actively over
the body of the host, as Myzostomum cirriferum.

2. Sedentary species which rarely, if ever, leave the spot where
they have settled, as M. parasiticum.

3. Cyst-producing species which cause the formation of galls or
swellings on the arms or disc, as M. cysticolum.

4. Entoparasitic species inhabiting the digestive tract, as M.
pulvinar.

5. Entoparasitic species living in the ovaries, as Protomyzostoma
polynephris does in the astrophytons.

Thus in the crinoids we find a single group of animals which,
broadly speaking, play the part of the fleas, lice, jiggers and bots,
intestinal worms and flukes combined as we know them among the
land vertebrates.

A comparison between the myzostomes and the species of Thrips,
occurring only on flowers, is also interesting.

In the vertebrates the blood with its multitudes of red corpuscles
which when destroyed are promptly and continuously renewed is the
logical food of practically all the parasites which do not inhabit the
intestinal canal. The dilute blood of the crinoids, without structures
corresponding to the red corpuscles, has none of the features which
make the blood of the vertebrates such a reservoir of concentrated
food. But the uncountable myriads of minute organisms flowing
continuously downward along the ambulacral grooves and into the
mouth form a stream of nutrient fluid in many ways analogous to the

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

vertebrate blood stream, and it is from this source that the myzostomes
as well as most of the other parasites derive their subsistence.

The species in each group parasitic on the crinoids in those cases
in which our information is sufficient to permit us to speak with a
reasonable amount of certainty follow bathymetrically and geo-
graphically the distribution of the classes to which they belong quite
regardless of that of their hosts, and apparently, excepting possibly
in the case of Stelechopus, the most primitive of the myzostomes
parasitic on the most ancient of the living crinoids, there is not the
slightest correlation between the systematic position of the parasite
and that of the crinoid.

An undetermined internal worm, a parasitic ostracod, Laphysti-
opsis, Anilocra, Cirolana (fig. 60), Cyclotelson (fig. 61), Synalpheus,
Periclimenes, Pontoniopsis, Galathea (fig. 59), Ophiactis, Ophiomaza,
Ophioethiops, Ophiophthirius, Ophiosphera, Sabinella and Polynoé
are known as parasites on crinoids only in the Indo-Pacific region,
though Laphystiopsis, a parasitic ostracod (on fish), Sabinella, Synal-
pheus, Periclimenes, Galathea, Anilocra, Cirolana, Ophiactis and
Polynoé also occur in the Atlantic.

Collocheres, Enterognathus, Stylina and Hemispeiropsis are known
as parasites on crinoids only from the Atlantic; but all of these are
small and must be especially searched for; probably all occur in the
Indo-Pacific.

Mortensen’s parasitic worm of doubtful affinities is only known
from the Antarctic; but only Notocrinus virilis offers a suitable
habitat for it.

Thus while the myzostomes occur wherever crinoids are found the
majority of the other parasites and commensals on crinoids are con-
fined to the Indo-Pacific region, though many are very closely related
to nonparasitic Atlantic species. The chief reasons for this are
probably the absence of a richly developed littoral crinoid fauna in
the tropical Atlantic comparable to that in the Indo-Pacific region,
and the plating of the ambulacra in most of the tropical Atlantic types,
including the littoral species, which renders them unavailable as a
source of food to most of the parasitic forms.

It is interesting to note that, with the exception of the myzostomes
and the gasteropods, the great majority of the organisms which are
directly or indirectly parasitic upon the crinoids are confined to the
littoral zone. The reason for this is probably to be found in the
development of side and covering plates along the ambulacral grooves
of the pinnules, arms and disc of the crinoids from intermediate and
great depths which enables the animals to convert the ambulacral

NO. 7 SEA-LILIES AND FEATHER-STARS—CLARK 39

grooves into closed tubes and prevents the appropriation of food
particles by the ectoparasitic crustaceans, the ophiurans, and the
polynoid worms.

The larger commensals living on the crinoids are usually striped or
banded, and resemble them more or less closely in color, though in
many cases the closely related noncommensal species are quite plain.
This may or may not be the case with the myzostomes.

In regard to the parasites and commensals of the comatulids there
is one curious feature which stands out very prominently—the
majority of the records, especially of the larger and more vigorous
types, are based upon species of the family Comasteridz, probably the
most specialized of all the comatulid types.

COMMENSALISM OF THE CRINOIDS

A number of small comatulids and the young of certain others may
be considered as truly commensal, living as they do in the cavities of
large sponges and gathering the minute organisms brought to them
by the currents flowing into the afferent openings of the host. Many
others habitually cling to gorgonians or withdraw into crevices in
corals where they live symbiotically, but quite independently of the
supporting organism. ?

ECONOMIC VALUE OF THE LIVING CRINOIDS

Economically the crinoids serve no useful purpose—at least up to
now they have been put to none. They cannot be eaten, and they are
not, so far as we know, eaten by any fish or other animal that serves
as human food.

As a result of their ordinarily fixed mode of life it is possible that
they might be used to furnish an index of the density of the finer
plankton content of the water in which they live, though it is probable
that other more generally distributed animals with more or less
similar feeding habits would serve the purpose better.

Because of their beauty and delicacy of form as well as on account
of their rarity they are frequently preserved and offered for sale as
curios in Japan and China and, less frequently, in India, Oceania,
Australia and the West Indies.

In southern Japan crinoids are frequently brought up on the long
lines used for fishing in deep water in Sagami Bay. The comatulids,
because of their beauty and delicacy of form, are called “ komachi ”—
a name originally borne by an exceptionally well-favored lady of the
court upwards of a thousand years ago—while the local stalked
crinoid (Metacrinus rotundus) is known as the “ bird’s foot.” The

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

former when well preserved are sometimes sold as curios, while the
latter,always meet with a ready sale at extraordinarily high prices on
account of their rarity combined with their paleontological interest.

In China comatulids are sometimes offered for sale which have been
brought from a considerable distance.

At Barbados the local species of [socrinus, especially I. asteria, and
Holopus rangu, are occasionally to be found in the curio shops.

Among the Slavic peoples red is the color about which all their
abstract ideas of beauty, and hence of idealism, revolve. The delicate
and often gorgeously colored red Adriatic Feather-Star (Antedon
adriatica) occurs more or less abundantly along the coasts of the
largely Slavic provinces of Istria and Dalmatia where it is frequently
found in the fishermen’s nets and is sometimes brought up on their
hooks. Its beauty of form, and particularly its red color, especially
commend it to the local fishermen, who commonly take it to market
and exhibit it along with the fish offered for sale.

EXPLANATION, OF PEATES
PLATE &

Fic. 1. A Feather-Star (Antedon adriatica) with the terminology of its
parts explained.

2. Lateral view of the centrodorsal and articular faces of the radials of
of a Feather-Star (Himerometra martensi) with the terminology
of its parts explained. Fossil Feather-Stars usually consist of
this portion of the animal only.

3. The same, in ventral view.

4. The naked incised disc of Cenometra bella with the terminology of
its parts explained.

5. The plated entire disc of Neometra multicolor with the terminology
of its parts explained.

6. Lateral view of the centrodorsal and articular faces of the radials
of Pentametrocrinus japonicus.

7. Longitudinal section of the same.

8. Ventral view of the same (compare with fig. 3).

9. The dorsal nervous system of Tropiometra macrodiscus.

10. The proximal portion of Nanometra bowersi showing the differ-
ence between the large cirri about the periphery of the centro-
dorsal and the small one near its apex.

11. Dorsal view of the centrodorsal, radials and arm bases of a specimen
of Comatula rotalaria showing the centrodorsal reduced to a
stellate plate.

12. The disc of a specimen of Comatula micraster with four grooved
and six ungrooved arms.

13. The ambulacral grooves and the digestive tube of Antedon bifida
(adapted from P. H. Carpenter).

14. The arm tip of Pterometra trichopoda.

NO.

FIc. 15.

TRIG

Fic.

7

16.

17.

18.

TO.
20.
ar:
22)
23.
24.

25.
26.

27
28.
20.

30.

21.

82:

33.

34.

37.

38.

SEA-LILIES AND FEATHER-STARS—CLARK 4l

The terminal comb on the oral pinnules of Leptonemaster venustus
in external, internal and ventral view.

The tip of an outer pinnule of Capillaster multiradiata in lateral
and dorsal view, showing the long spines.

The division series and arm bases of Comanthus bennetti with, on
the right, homologous ossicles shown similarly shaded.

PLATE 2

The calcareous deposits in the perisome bordering the ambulacral
grooves and in the tentacles of the pinnules of Dorometra
parvicirra.

The same in Leptonemaster venustus.

The same in Eudiocrinus junceus.

The same in Heterometra bengalensis.

The same in Amphimetra discoidea.

The calcareous deposits in the perisome bordering the ambulacral
grooves in Psathyrometra antarctica.

The same in Pentametrocrinus varians. |

The same in Eumorphometra concinna.

The same in Sarametra triserialis.

The side and covering plates of Glyptometra tuberosa; the latter
(above) are hinged to the former and can be closed down over
the ambulacral grooves.

A side plate of Strotometra hepburniana in (upper) interior and
(lower) dorsal view.

Interior view of two side plates of Pachylometra distincta.

Ventral view of a portion of a pinnule of Pachylometra distincta
showing the side and covering plates, the latter closed down
over the ambulacral groove.

Lateral view of the side and covering plates of Pachylometra dis-
tincta, the latter partially closed down.

PLATE 3

Diagram showing the relative proportions of the arms and cirri in
Asterometra macropoda; the cirri are adapted to clinging to
very rough bottom.

Diagram showing the relative proportions of the arms and cirri in
Comactinia echinoptera; the short, strong and stout cirri are
well fitted to hold the animal securely.

Diagram showing the relative proportions of the arms and cirri in
Pentametrocrinus tuberculatus; the animal is very different in
every way from the preceding, but the cirri are of the same type.

Diagram showing the relative proportions of the arms and cirri in
Pentametrocrinus varians; the cirri collectively form a sort of
circular mat supporting the animal on soft ooze.

The grooved anterior (left) and ungrooved posterior (right) arms
of a specimen of Comatula pectinata, drawn to the same scale.

A cirrus of Capillaster multiradiata in dorsal (upper) and lateral
(lower) view, showing the dorsal spines.

A smooth cirrus in dorsal (upper) and lateral (lower) view.

Fie, 42.

43.

Fic. 44.

Fic. 45.

Fic. 46.

Fie. 48.

Fic. 49.

Fic. 50.

BIGe ss

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

PLATE 4

Crown and upper part of the column of Monachocrinus sexradiatus.

PLATE 5

Crown and upper part of the column of Proisocrinus ruberrimus.

PLATE 6

Crown and upper part of the column (above) and central portion
of the column (below) of a _ pentacrinite, Comastrocrinus
springer.

PLATE, 7

Crown and upper part of the column (a), and middle (b) and lower
(c) part of the column of Ptilocrinus pinnatus; in life the upper
part of the column is recurved so that the crown points directly
downward.

‘Comatilia iridometriformis, a species of Comasteridze with some of

the pinnules at the base of the arm lacking.

PLATE 8

A specimen of Comatula pectinata from Singapore showing long
anterior and short posterior arms (compare with fig. 36).

PEALE :

Neometra acanthaster, one of the Calometride.

BEATE -10

Asterometra macropoda, one of the Ptilometrine.
PLATE Aa

Antedon adriatica, one of the Antedonine.

IPL NIN, We

Compsometra incommoda, one of the Antedoninez.

PLATE, 13
Thaumatocrinus jungersent, a ten armed species of Pentametro-
crinide.
PLATE 14
Pentametrocrinus diomede@, a five armed species of Pentametro-
crinide.
PLATE 15

An arm of Heterometra compta, showing the difference between
the oral, genital and distal pinnules.

NO. 7

FIG. 52.
53.

54.

SEA-LILIES AND FEATHER-STARS—CLARK 43

An arm of Stephanometra echinus, showing the stout and spine-
like oral pinnules.

An arm of Stylometra spinifera, showing the mid-dorsal overlap-
ping spines.

Four syzygial pairs from the arm of Stylometra spimfera, showing
the progressive increase in the individuality of the two elements
distally.

Pentacrinoid larva of a large Feather-Star, Heliometra glacialis
(after Levinsen).

Ventral view of a larva of Antedon mediterranea early on the eighth
day (adapted from Bury).

Myzostomum costatum, dorsal view (after Boulanger).

The same, ventral view (after Boulanger).

Galathea elegans (after Potts).

Cirolana lineata (after Potts).

Cyclotelson purpureum (after Potts).

Scalpellum pentacrinarum (after Pilsbry).

PLATE: 16

A specimen of Nemaster iowensis captured in shallow water in the
Bahamas (Cat. No.. 36164 U. S. N. M.).

VOL. 72, NO. 7, PL.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

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VOL. 72, NO. 7, PL.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 72, NUMBER 8

A REVIEW OF THE INTERRELATIONSHIPS
OF THE CETACEA

BY
HERLUF WINGE
[Translated by GERRIT S. MILLER, JR.]

(PUBLICATION 2650)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1921

¥ ies The Lord Baltimore Drees

BALTIMORE, MD, U.S A.

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Pe VV Ob - Lith INTE RRELATIONSHIPS OFTHE
CETACEA

By HERLUF WINGE
[Translated by Gerrit S. Mitrer, Jr.]

In translating Doctor Winge’s “Udsigt over MHvalernes indbyrdes
Slegtskab” (Vidensk. Medd. fra Dansk naturh. Foren., vol. 70, pp. 59-142,
1918) my aim has been to give the author’s ideas as clearly and exactly as
possible rather than to make smooth English sentences. I have been much
aided by the kindness of Dr. Leonhard Stejneger, who has compared the
entire MS with the original, making himself responsible in particular for
the rendering of the adverbs ret, sikkert, vel, and vist, whose idiomatic
shades of meaning present many difficulties to one whose acquaintance with
Danish is limited to the printed language. Doctor Winge has also examined
the translation, expressed hi3 approval of it, and made some useful suggestions
for its improvement. I have added an index, a few bibliographical refer-
ences, and in some instances the generic names which are correct according
to. the International Code of Nomenclature. Brackets are used to distinguish
all additions to the original text——G. S. M., Jr.

The Cetacea * originated * from the Hyenodontide, the most primi-
tive family of the Carnivora, by way of the most typically carnivorous
members of the group such as Pterodon and Hyenodon. The oldest
known whales have such a great likeness to Hyenodon and its nearest
relatives that there can be no doubt about the relationship. Aquatic
habits have given the cetaceans their special peculiarities and have
caused their differentiation from the Hyznodonts.

As an inheritance from the highest Hyzenodonts, and as an indica-
tion ‘of relationship with exactly these animals, the most primitive
whales retained a series of special peculiarities which the Hyzenodonts
had developed in the course of their differentiation from the insectiv-
orous stage. They still had about the same dentition as the Hyzeno-
donts. All the teeth were fitted for flesh eating; the incisors and
canines were strong and hooked, the anterior cheekteeth strong,
elongated, compressed, smooth-edged; the molariform cheekteeth,
especially those of the upper jaw, had a peculiar form and all of
them were nearly alike. In the upper molars the 4th and 5th cusps
[paracone and metacone] were coalesced to form a trenchant longi-
tudinal ridge, the 1st and 2d cusps [parastyle and mesostyle] were
reduced, the 3d cusp [metastyle] was a compressed ridge, and the

*Notes are at end of paper, pages 47-93.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 72, No. 8

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

6th cusp [protocone] was reduced or absent. All of the upper molari-
form teeth including the third premolar still had the inner root,
though this was in process of reduction. All the teeth of the typical
[eutherian] dentition were present, 11 in each jaw [44 in all]. The
jaws were elongated in harmony with the long, well-developed tooth-
rows. The temporal fossa was very large, widened out by a powerful
temporal muscle. It was bounded by a high sagittal crest, by a
strong, backward-projecting occipital crest, and by an abruptly out-
standing, posteriorly heavy processus zygomaticus squame.

In addition to these peculiarities the most primitive whales had two
high characters which were perhaps inherited from the Hyzenodonts ;
at any rate they are to be found in the latter group, though less pro-
nounced: a rather large supraorbital process, and a bony palat
lengthened backward far under the posterior nares.

Radical alterations have taken place during the change from
Hyeenodont-like carnivores to true whales. In many of the mam-
malian groups there have arisen forms modified for life in the water ;
but no other aquatic mammals are modified to the same degree as the
cetaceans, nor has any other become so exclusively aquatic; only to
breathe do they raise the nose above the water in which they other-
wise are hidden.

The cetacea have used the tail as the chief implement of locomo-
tion ; the hind limbs are put wholly out of service; the fore limbs are
scarcely used for much else than steering and balancing.

The tail becomes enormous, long and thick, powerfully muscled.
It is formed in agreement with the manner in which it is wielded:
with strokes from side to side, or up and down, or with a sculling
motion. Throughout most of its extent it becomes compressed, but
at the tip 1t acquires a powerful, horizontal, caudal fin constructed of
skin folds (not present in quite young embryos of recent cetaceans).
At the front of its upper margin, in the region where the tail joins
the back, there may occur a special erect skin fold in the form of a
longitudinal crest, a dorsal fin. Most of the caudal vertebrz lose the
atrophied appearance which they have in primitive mammals; they
acquire powerful centra, heavy, flat-outspread transverse processes,
high dorsal arches with large, compressed spinous process, and
articular processes which are distinct though not mutually fitting
together. The ventral arches with the inferior spinous processes
become so large that they approach the upper arches in size. Only
the outermost caudal vertebre, which lie almost inclosed in the caudal
fin, retain the degenerate character. The tail has an influence on the

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 3

dorsal vertebre also. Its powerful muscles, which have their origin
in part on the sacral and dorsal vertebrz, and are also in connection
with the muscles of the back, widen their place along the spinal
column and stimulate the vertebrz to increase in bulk. The sacral
and lumbar vertebre come to resemble the largest, most anterior
caudal vertebrz exactly, apart from the lower arches. On a few of
the hindmost thoracic vertebre, which in the ancestral forms are
without, or as good as without, transverse processes, there grow out
powerful transverse processes on a line with and similar to those
of the lumbar vertebrz (parapophyses, apparently corresponding to
the lower section of the double, rib-bearing “ transverse process ”’ of
the anterior thoracic vertebree, which supports the rib’s capitulum ;
but in reality they most probably correspond to the upper and lower
sections combined), and on their tip they eventually bear the attach-
ment surface for the rib. Apparently this surface may be either for
the attachment of the tuberculum or of the capitulum or of the two
coalesced, but in reality it is perhaps always for the two combined
(or, more correctly, not separated). The transverse processes of the
anterior thoracic vertebre (diapophyses, the upper portion of the
double ‘“‘ transverse processes”), which in the beginning are quite
short, may eventually grow long, pushing far out to the side the
articular surface for the rib’s tuberculum which they bear at their
extremity. On all the thoracic vertebre the spinous processes finally
become high and strong.

The hind limb atrophies completely, and disappears. At length
only quite insignificant parts of its skeleton are found, hidden deep
under the skin, finally in the form of a mere little rod-shaped bone, a
‘remnant of the pelvis.” In small embryos the hind limb can, however,
still be distinguished externally.

The disappearance of the hind limb has a great influence on the
vertebral column. No longer does a pelvic bone come in contact with
any of the vertebree. In consequence the sacral vertebree completely
lose their peculiarities : their characteristic strength, their mutual firm
connection, their robust transverse processes with flattened areas for
the hip bones. They are formed exactly like the adjacent dorsal and
caudal vertebre. The movements of the spinal column become
changed in character. Bending of the column in the vertical plane,
which depends especially on the movements of the hind legs, is
reduced or abandoned, and as a result the differences in slant—
forward or backward—of the spinous processes as good as disappear,

val SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

so that all the processes alike are directed upward. The zygapophyses
become reduced and in large part lose their mutual articulations.

The fore limb, which no longer comes in contact with the earth or
bears any load, is changed into a flipper whose single function is that
of striking against the water. The entire arm becomes an oar blade.
Of the fore limb’s articulations the shoulder joint only is used; it
retains its ball-and-socket structure. All the other articulations are
held stiff. They degenerate, become flat and immovable, or are
wholly effaced. Practically the only function of the upper arm is to
support the forearm and hand. It becomes short and heavy. Its
middle portion retains its terete form, but its lower end is com-
pressed in agreement with the bones of the forearm. The radius
and ulna become very simply-formed, compressed bones, losing
muscle crests, sinew furrows and all pronounced articular sur-
faces ;,even the olecranon may wholly disappear. The mutual posi-
tion of the two bones is somewhat altered, so that they eventually
lie exactly fore and aft of each other. The hand is set somewhat
supine, fore edge downward. The carpal bones become compressed,
or more correctly flattened, pieces [like sections of a mosaic]. They
are rather indifferent as to form and number, and are immovable.
The folds of skin between the fingers are lengthened out to the finger
tips; and the hand stiffens. The claws disappear. The first and
fifth fingers are somewhat inclined to be stunted, but the other fingers,
particularly the second, tend to lengthen and to form new joints at
their tips, so that the number of phalanges may increase far beyond
the typical three. The metacarpals and phalanges are shaped almost
alike, as more or less flattened pieces of bone.” The shoulder blade
degenerates only slightly. In the most primitive whales it already
has the form which, with few exceptions, is found among the highest.
It is broadly fan-shaped, with a prominent, antrorse acromion, and a
large coracoid, but on the other hand almost without crista scapule.
Rarely it becomes narrower or lacks both acromion and coracoid.

The fact that the fore limb does not act as a support for the body
results in lessening the limb’s pressure on the chest. Another result
is that the spinous processes on the anterior thoracic vertebre lose
their special height. Still another result is that the connections
between the ribs and both the thoracic vertebree and the sternum have
a tendency to become loose or to disappear. Perhaps this tendency
is also brought out by the fact that the water pressure on the chest
during diving changes strongly. The ribs may lose the capitulum,
and the costal cartilage may practically disappear. When this happens

No. 8 INTERRELATIONSHIPS OF THE CETACEA—-WINGE 5

the sternum loses an essential stimulus and becomes reduced and
atrophied.

The head, during swimming, is held directed as firmly as possible
forward. The neck is not moved, and for this reason it becomes short
and stiff. During motion through the water the head is pressed from
the front; it is forced backward against the cervical vertebrze, which
thereby are squeezed excessively together and pressed back against
the anterior dorsal vertebree, with the ribs of which they may even
come in connection. Most of the cervical vertebrae may become
almost as thin as paper. The odontoid process of the axis becomes
short and blunt; the articular surface between the bodies of the atlas
and axis becomes almost flat. And there arises a strong tendency to
coalescence of the cervicals.’ The occipital condyles lose their pro-
jecting form and become almost flat, only quite weakly convex, pressed
in against the wall of the braincase; and the concave surface of the
atlas likewise becomes flattened out. The occipital crest in its
capacity as an attachment for the upper neck muscles is restricted ;
the points of attachment for the lower neck muscles on the basal part
of the occipital bone are effaced, and the under side of the cccipital
bone is formed more as a sheath around the gullet and windpipe.

The pressure of the water on the head when the cetacean swims
has a highly modifying effect on the skull.

From above the water presses especially during the animal’s con-
stant rising to the surface to breathe. This gives the skull.a tendency
to acquire a flat and broad upper surface, with thick bones. The size
of the horizontally outspread supraorbital process of the frontal,
which pushes itself far out over the orbit, becomes particularly notice-
able. The facial part of the cheek bone may likewise become pecu-
liarly flattened out.

From in front the water presses during forward motion, the more
strongly as the motion is faster. Its effect is to develop an unusual
strength in those bones of the face which project furthest forward,
the intermaxillary, maxillary, and vomer, as well as in the cartilaginous
nasal partition which the vomer embraces. This strengthening may
show itself in different ways: in the noticeable lengthening forward
of the bones in question, in their solid ossification, in their tendency
to coalesce. It also appears in the backward spreading of the inter-
maxillary and especially of the maxillary. The latter may extend
itself out over the facial part of the zygoma and over the frontal,
which it almost entirely covers to the hinder margin, so that the supra-
orbital foramen may pierce not only the frontal as in other mammals.

6 SMITHSONIAN “MISCELLANEOUS COLLECTIONS VOL. 72

but the maxillary as well. The cartilaginous nasal partition, the
mesethmoid, has a tendency to ossify. The incisive foramen is
narrowed and closed. In the palate the maxillary pushes itself far
backward, forcing the palatine behind it; the palatal surface of the
palatine is thus shortened. But at the same time the maxillary acts on
the palatine in such a way that it also increases in thickness. The
braincase is acted on from the front by the pressure of the water
against the forehead; from behind it is pressed by the cervical
vertebree and the neck muscles. In this manner it becomes so
squeezed that it acquires a short, broad form. Pressure is exercised
especially on the frontal and on the supraoccipital and interparietal.
These bones widen out at the expense of the parietal, whose inner-
most part is squeezed quite narrow and eventually obliterated. The
exoccipital also grows, especially noticeably downward, where it
broadens out shield-like behind the mastoid and the tympanic. The
mastoid is compressed inward between the exoccipital and the
squamosal, by both of which it is so overgrown that at last it is no
longer visible on the outer surface of the braincase.

The water pressure on the head from in front has also a great
influence on the soft parts of the face and through them on the skull.
It assists in shifting the nasal apertures. The cetacean has tried, with
the help of the nose muscles, to draw the apertures as high as possible
up on the head’s upper side, in order to be able easily to get them
raised above the water. The result has been that the nasal cartilage
has caused resorption of the anterior border of the nasal bones and
has forced them further and further back. The cartilage has also
worked itself back between the anterior median part of the frontals,
pushing the plates of the ethmoid behind it. Thus at last the nares
have acquired a position which appears to be on the forehead but
which in reality is close in front of the anterior wall of the braincase.
The moving of the nasal cartilage has been accelerated in those cases
where the facial adipose cushion which originally lay in front of the
nares and which in the first place was merely a little filling out of
fatty connective tissue has been stimulated to growth by the pressure
of the water, becoming very large, pushing the nasal cartilage back-
ward, pressing it against the front wall of the braincase and disinte-
grating the nasal bones and the plates of the ethmoid. The nasals then
become tuberformed and are pressed into the frontals. The adipose
cushion together with the nasal muscles and other neighboring
structures may exercise an enormous influence on the skull, the
anterior and upper sides of which it modifies to form its bed, the
“ facial depression.”

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 7

The water acts in a very special way where the whale lets it stream
into the mouth for the purpose of catching the small animals which
it carries in with it. In such cases it brings about huge increase in the
size of the jaws together with many other remarkable peculiarities.

For smelling there comes to be no use; this sense is not exercised,
and the nose is therefore formed in accordance with the needs of
breathing only. The ethmoid degenerates. The numerous folded
laminze of which it originally consisted disappear, while the cribriform
plate loses its nerve perforations and becomes a solid lamina of bone
on the front wall of the braincase. The nose becomes a simple
passage for air. The air, which is exposed to strongly varying
pressure and temperature, has a tendency to provide itself with
greater space by widening out the nasal passage and Eustachian tube
wherever it meets with least resistance. It may form air-sacs, partly
on the upper side of the skull over the facial bones, partly on the
under side behind the palate. Here an air-sac may spread itself
forward along the outer side of the pterygoid and palatine and back-
ward along the outer margins of the body of the sphenoid and the
basal part of the occipital, pushing itself out under the ala parva, ala
magna and the squamosal, and bounded more or less by plate-like
outgrowths from all the bones mentioned. The bony palate is
lengthened backward still more by the pushing out from the ptery-
goids of laminze which extend into the soft palate beneath the nasal
passages. This clearly takes place partly under the action of the
tongue, but doubtless still more under the influence of the larynx.
The fact that the two original outer nostrils finally coalesce into one
is an indication of the nose’s degeneration.

The lacrimal bone is reduced and eliminated, or it fuses with
the cheek bone as in many other aquatic mammals, probably because
the bone is no longer acted upon by a lacrimal duct.

The outer ear disappears from lack of use; the outer auditory
aperture is so strongly contracted that it may be difficult to find.
The bones of the inner ear acquire a peculiarity which is found
again in several other mammals that live in the sea, and which cer-
tainly in some manner or other must be dependent on aquatic life.
They are formed of unusually thick, stony-hard masses of bone; this
is especially remarkable as regards the tympanic, the inner wall of
which is thickened in a peculiar way.”

The dentition degenerates because the chewing of food is given up
as not easy to carry on satisfactorily under water. Most animals
chew with open mouth; under water the chewed food would be

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

washed away from between the teeth. The dentition is therefore
chiefly used for grasping the food and holding it fast. In the most
primitive whales the mouth did service as an implement for catching
fish. The jaws were used in exactly the same manner as in the shell-
drakes, Mergus, and they were produced forward as a long slim beak,
a kind of tweezers, influenced not only by the use to which they were
put, but also by the pressure of the water during swimming forward.
The Hyenodon-like dentition which the most primitive whales
inherited, with teeth of considerable size, diversified form, and of
typical number, at first becomes more simple. The upper molariform
teeth lose the inner cusp and the inner root, and the crown under-
goes compression. A further step in the reduction is that the crowns
of the cheekteeth, or at least of most of them, acquire a serrated
anterior and posterior margin. Next the two remaining roots,
foremost and hindmost, of the cheekteeth fuse into one, and the
serrations of the crowns are reduced and obliterated. The size
at the same time is reduced, and the form becomes simply conical so
as to resemble that of the incisors and canine, which in their turn
undergo reduction. While this is happening the number of teeth in
the long jaws is increased, no doubt because in the place of the few
quite large teeth there spring up many smaller ones; scarcely by the
actual splitting up of the few. Perhaps also in the beginning some
of the milk teeth came to take a place in the series with the permanent
ones, without, however, the entire milk dentition’s intercalation in the
permanent set. The number of teeth grows greater and greater, far
beyond the typical, while the individual teeth become smaller and
smaller. Those at the front and back of the series become especially
stunted, frequently disappearing from the intermaxillary. The
enamel covering of the teeth becomes thin or disappears entirely.
What later happens to the dentition depends on the use to which it is
put. It may happen that there comes to be no use whatever for it,
and that it consequently disappears. Or it may, wholly or in part, be
once more put to heavy use and be modified to this end; or a single
tooth may take on power while all the others atrophy.

The succession of teeth, which in the most primitive whales took
place in the ordinary way, ceases. It is not clear how this happens.
Judging from investigations of the teeth in embryos of the higher
cetaceans it might appear, at least sometimes, as if it were retained
milk teeth that are found in the adult animal’s dentition—as if the
successors to the milk teeth had cisappeared. Such, however, is
scarcely the explanation. Most probably it is really the actual perma-

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 9

nent set that is found in the adult, while those that precede and follow
(both of which have been demonstrated) disappear.’

The fact that mastication ceases and that the teeth become stunted
has a great influence on the chewing muscles and the jaws. It was
necessary for the first whales as fish catchers to be able to open the
mouth wide. The masseter muscle which has the tendency to limit
the opening of the mouth was therefore little used, and it became
restricted ; together with the muscle the region of its origin became
shrunken. This region is the anterior and median part of the
zygoma ; it is transformed into a slender bridge of bone. The tem-
poral muscle has been more used, but it also shows the tendency to
be reduced by lack of vigorous use, and it draws itself backward quite
low on the side of the braincase, losing its influence on the zygomatic
process of the squamosal. This process shrivels up like the coronoid
process of the mandible, the muscle’s point of insertion. In cases
where the under jaw becomes very large the temporal muscle may
acquire renewed strength and may spread its region of origin out
over its surroundings in an unaccustomed manner. With the atrophy
of the teeth they cease to influence the body of the mandible, which
consequently loses its original height. The alveoli become less defined
and the partition walls between them may disappear so that there
arises a common dental furrow. The articular condyle of the
mandible weakens, loses its cylindrical form, and the articular sur-
face becomes an almost flat area pointing backward at the similarly
formed glenoid fossa on the squamosal, which as good as loses its
postglenoid process and is otherwise inclined to suffer reduction. It
may happen, however, that the lower jaw becomes huge and that its
articular condyle acquires corresponding heaviness. In such cases
the condyle is curiously modified, losing the true articular surface.
This is grown over by articular ligament, and the lower jaw stimu-
lates the squamosal to grow out in prodigious size, bearing, instead
of the true articular surface, an area of attachment on a projecting
foot. The symphysis menti. long in the most primitive whales, is
restricted. The under jaw’s degeneration is also no doubt indicated
by the huge gaping posterior entrance to the mandibular canal, which
is mostly filled with loose connective tissue. It is not clear what the
reason is for this peculiarity, which was already present in the most
primitive cetacea and is found in all the later ones though sometimes
in a rather disguised form; possibly it might in some way depend on
the air-sacs of the nasal passages which lie exactly internal to this
part of the lower jaw.

10) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Whales lose their hair covering because it ceases to be of use; at
most some few degenerated vibrisse remain.

It holds good for the cetacea as for other groups of mammals that
the most primitive forms have much less brain than the later ones;
in the highest whales the brain is extremely well developed.

It likewise holds good for the cetacea as for others that the earlier
forms are smaller than the later, though dwarfs may at any time be
developed. Ordinarily whales increase noticeably in size as they
become more highly developed; the highest forms have reached
gigantic proportions.

Judging by their greater or less resemblance to the Hyznodonts
the cetacea are mutually related essentially as follows: ”

Cetacea.
I. The number of teeth is not more than typical [44]. Braincase
not telescoped, not shortened.

Archeoceti.

Zeuglodontide.
Protocetus, Progeuglodon, Zeuglodon.
II. The number of teeth is or has been more than typical. Brain-
case telescoped, shortened.

A. Nasal bones forming a roof over hinder part of nasal

cavity. Maxillary not covering frontal.
Mystacoceti.
Baleenide.
BaL2#NINI: Balena, Neobalena.
BALANOPTERINI: Lhachionectes, Plesiocetus,
Cetotherium, Balenoptera, Megaptera.

B. Nasal bones pressed into fore wall of braincase, not or
scarcely forming a roof over any part of nasal
cavity. Maxillary covering frontal.

Odontoceti.
1. Teeth not alike, the most posterior less simply
formed than the most anterior.
Squalodontidz.
Agorophius, Squalodon, Neosqualodon,
Prosqualodon.
2. Teeth now or formerly alike, simple in form.
a. Temporal fossa large, not covered over by
frontal and maxillary; zygomatic pro-
cess of the squamosal heavy, primitive
in form.

No. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE II

Platanistide.
Pontistes, Pontoporia, Lipotes, Inia,
Saurodelphis, Platanista.

b. Temporal fossa relatively small, covered
anteriorly by the widened frontal and
maxillary; zygomatic process of the
squamosal reduced, losing its primitive
form.

a. Occipital wall not especially elevated.
Delphinide.

Delphinodon, Champsodelphis,
Schizodelphis, — Heterodelphis,
Eurhinodelphis, | Argyrocetus,
Delphinapterus, Monodon,
Steno, Prodelphinus, Delphi-
nus, Tursiops, Tursio, Lageno-
rhynchus, Orca, Orcella,
“ Grampus,” Pseudorca, Globi-
ceps, Phocena, Neomeris.

8. Occipital wall highly elevated.
Physeteride. ,

XIPHINI: Argyrodelphis, Meso-
plodon, Xiphirostrum, Chonoxi-
phius, Xiphius, “ Berardius,”
Hyperoodon.

PHYSETERINI: Hoplocetus, Phy-
seterula, “ Cogia,’ Physeter.

In the form of a genealogical tree [see pp. 45-46] :
Physeteride.

Delphinide.
Platanistide.
Squalodontide.
Balende.

Zeuglodontide.

Zeuglodontide [| Basilosauride|—Of all known cetacea the
Eocene Egyptian Protocetus of the family Zeuglodontide is the most
primitive. It is known from a rather complete skull without the

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

lower jaw, and from a few vertebre and ribs. In all that is known it
stands so near to the Hy@nodontide that there would scarcely have
been any reason to separate it from them had it not been evident that
it was one of the first members of the cetacean series. The number
of teeth in the upper jaw is, as in Stypolophus and Pterodon, the
typical 11, since m* is present, while it has disappeared in Hyenodon,
the highest genus of Hyenodontide. But the form of the teeth is
most nearly as in Hyenodon, more shearing than in other members
of the family. The difference from Hyenodon is chiefly a result of
the fact that heavier use has been made of that part of the toothrow
which serves for grasping the food and that consequently the incisors
and anterior cheekteeth have increased in size. The incisors have
become about as heavy as the canine, the premolars have become
heavier and more elongated than formerly, while the molars are
weakened and m? has lost its predominance. The mouth was already
used mostly as a pair of forceps; the long, narrow, but strong, beak-
shaped jaw, in which the teeth have abundant space, bears witness
to the fact. The anterior nasal aperture is already forced consider-
ably backward; but it has, however, only reached a point scarcely
half way to the anterior margin of the orbit, and it has kept a rather
primitive form. The nasal bone is long and narrow, roofing over a
large part of the nasal cavity. The intermaxillary is strengthened
anteriorly, its body is lengthened, likewise its nasal process, though
this process does not reach to the frontal. Otherwise the inter-
maxillary does not show much deviation from the conditions ordi-
narily found in carnivores. The maxillary also is lengthened and
thickened, but is not otherwise modified to any noticeable degree.
Posteriorly it does not push itself out over the facial part of the
zygoma or over the frontal, which it merely forces slightly backward.
On the palatal surface it has not crowded the palatine bone, which
has retained its original length. The incisive foramina seem to have
disappeared. The forehead is pressed quite flat, and the supraorbital
process of the frontal has become very broad; otherwise the forehead
is unmodified. The anterior and median part of the zygoma is
already well on its way to become slender, but the zygomatic process
of the squamosal is still robust. It bears a considerable postglenoid
process, though the articular surface for the lower jaw has begun
to assume the: peculiar vertical position that it has in the higher
whales. The temporal fossa has on the whole remained primitive
in size and form. It is bounded by high crests. The braincase is
not compressed antero-posteriorly ; the frontal and supraoccipital are

No. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 13

not widened so as to encroach on the parietal. The mastoid is still
visible on the outer wall of the braincase. Occipital crest well
developed, projecting. Occipital condyles not pressed flat. On the
basal part of the occipital the impressions where the lower neck
muscles were attached are essentially unmodified in character, and the
under side of the occipital bone’s basal part is not shaped for sheath-
ing the gullet and larynx. On the other hand the exoccipital has
already acquired a noticeable widening out to the side. The hinder
part of the nasal cavity appears to be wholly undisturbed ; it must
contain a well-developed ethmoid. The bony palate is already pro-
longed backward by plate-like outgrowths from the lower margin of
both the palatine and the pterygoid. No doubt an air-sac formed by
an enlargement of the nasal passage lay on the outer side of the
pterygoid, but whether it was enclosed by outgrowths from the
adjacent bones is doubtful. The tympanic bone had already acquired
the characteristic cetacean thickening of the inner wall. The cervical
vertebre are mutually free, not strongly compressed. The odontoid
process of the axis is strong, projecting. The spinous processes of
the dorsal vertebre differ noticeably among themselves as to their
slant, some of them sloping strongly backward, others upright or
directed a little forward; those on the hindmost dorsal vertebre are
rather low. Zygapophyses apparently well developed. No project-
ing transverse processes on the hindmost thoracic vertebrae. Centra
of ordinary size. On the tip of the transverse process of a sacral
vertebra there is present a rather large area of attachment for the
ilium, although the process has otherwise already lost much of its
original character. On such ribs as are present in the fossil there is
a well-developed capitulum; the hindmost ribs lack the tuberculum
and are articulated with the corresponding vertebrz by the capitulum.

Progeuglodon (Zeuglodon osiris, Prozeuglodon atrox partim”),
also Eocene, Egyptian, has departed in dental characters not a little
from Protocetus. In the number of teeth the difference is only that
m*, small in Protocetus, is here absent. The form of the teeth has
undergone greater change: pm’ has lost the compressed form of the
crown and has become simply conical with a single root like the
incisors and canine; pm? has acquired a serrate posterior margin;
pm’, pm*, m+ and m? are strongly serrated on both the anterior and
posterior margins of the crown; in pm? and pm‘ the inner heel is
much reduced and in the two molars it has entirely disappeared. The
lower jaw is also known; it contains the typical 11 teeth. Incisors,
canine, and pm, approximately uniform, simply conical; pm,, pm,

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

and pm, with compressed crown and serrated anterior and posterior
margins; m,, m, and m, also with compressed crown, its anterior
margin smooth, its posterior margin serrate. Tooth succession
occurred in the ordinary way as it assuredly did in Protocetus also.
In all characters the skull agrees essentially with that of Protocetus.
The basal part of the occipital appears, however, to be more adapted
to the larynx and gullet. A few peculiarities of Prozeuglodon which
are not clearly demonstrated in Protocetus are: the presence of an
elongated, compressed incisive foramen on each side; the presence
of a distinct lacrimal bone; and the presence to the outside of the
pterygoid and in front of the tympanic of a considerable pit bounded
by high ridges springing from the surrounding bones, evidently the
impression of an air-sac. The under jaw already has nearly the same
peculiar form as in many highly developed cetaceans with long
symphysis menti; it has, however, a relatively large coronoid process.
But the mandibular condyle is placed low and is turned backward,
and the strange gaping hinder entrance to the mandibular canal is
present. Of the rest of the skeleton rather more is known than of
Protocetus, among other parts most of the vertebral column and the
fore limb down to the hand. There is a great similarity to Protocetus.
A difference from this genus is that no sacral vertebra is found with
the transverse process plainly acted upon by the ilium. The skeleton
of Prozeuglodon throws light on certain conditions that are not under-
stood in Protocetus. The sternum is of considerable size, with several
joints. The shoulder blade is essentially as in the higher whales.
The humerus has retained relatively much of the original form:
distinctly separated greater and lesser tubercles, a distinct deltoid
crest, and a well-developed hinge-shaped lower articular surface.
Radius and ulna have correspondingly well-developed articular sur-
faces for the humerus, are relatively only a little compressed, and
have distinct articular surfaces for the carpal bones; the ulna has a
rather large olecranon.

Zeuglodon [Basilosaurus| (Z. cetoides, Z. isis), known rather
completely as to the skeleton, occurs in Eocene strata of both the
Old and New Worlds. In most respects it resembles Prozeuglodon.
But it has acquired a highly remarkable peculiarity in the vertebral
column. While the centra in Progeuglodon are not in any direction
strikingly altered in form, in Zeuglodon the centra of most of the
hinder thoracic vertebre, of the lumbar vertebrz, sacral vertebrz and
all but the outermost of the caudal vertebrae, have become remarkably
large and especially greatly elongated, while the vertebral arches
have remained short, standing about midway on the centra, the arches,

no. 8 INTERRELATIONSHIPS OF THE CETACEA——-WINGE I5

in common with the spinous processes, widely separated from each
other. Thus in Zeuglodon the body has acquired an altogether
peculiar length, putting one in mind of the snakes. The posterior
thoracic vertebree seem to have developed considerable transverse
processes which bore the ribs on their extremities. In size also
Zeuglodon went further than its relatives. Of the hind limb there
is known a small, quite atrophied pelvic bone with articular surface
for the femur, and an even more degenerated little rod-shaped femur.”

The genera of Zeuglodonts together form the section Archzoceti,
the source from which all the higher cetaceans have originated. Pro-
tocetus has scarcely a single peculiarity, apart from its large size,
that one would not expect to find in an ancestral stock for the higher
whales. The same is true of Prozeuglodon. On the other hand
Zeuglodon, a descendant of Prozgeuglodon, has followed its own line
away from the starting point of the other whales, deviating particu-
larly in its remarkable vertebre.

The peculiarities which especially place the Zeuglodonts lower than
all other cetaceans are that the teeth are still present in the typical
number, and that the braincase is not telescoped and shortened. Of
all other whales it holds good that they, so far as they are known.
have the number of teeth raised above the typical (or that they are
descended from cetacea in which it had been raised), and that they
have the braincase more or less compressed antero-posteriorly. As
regards the form of the teeth Protocetus no doubt stands lower than
all other cetaceans; but Prozeuglodon and Zeuglodon are in this
respect scarcely more primitive than the lowest members of the higher
families. Of all the many other primitive characters that are found
in the Zeuglodonts some are, it is true, no longer to be found in the
higher families, not even among the extinct lowest forms; but for
most of them this does not hold good.

Zeuglodontide * | Basilosauride |.
I. Crowns of cheekteeth with smooth, not serrate, margins.
Protocetus.
II. Most of the cheekteeth have serrate anterior and posterior
margins to the crowns.
A. Centra of thoracic, lumbar, and caudal vertebre not
elongated.
Prozeuglodon.
B. Centra of posterior thoracic, of lumbar and caudal
vertebrz elongated.
Zeuglodon | Basilosaurus |.

ts
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Balenidze.—The group Mystacoceti with the single known family
Balenide includes whales that stand near to the Zeuglodonts ; but the
most primitive members of the group had already advanced a step
further than the Zeuglodonts. They presumably had the number of
cheekteeth raised above the typical. With that change there followed
others. The most primitive Mystacoceti must have already had the
nasal aperture pushed further back than in the Zeuglodonts. The
intermaxillary probably extended further back. The maxillary must
have been somewhat more broadened out posteriorly. The parietal
was slightly encroached upon, and the braincase was a little tele-
scoped. The spinous processes of the dorsal vertebrze presumably
slanted to a less degree in different directions. The joints at the
elbow and wrist must have almost wholly lost their primitive structure,
etc. Taken all in all, however, the most primitive Mystacoceti must
have been in general like the most primitive Zeuglodonts.

Of the many forms which the group Mystacoceti must have in-
cluded no others are known than a little circle of highly developed
genera very specially modified in their own direction; but in spite of
their remarkable development they have retained many primitive
features which are no longer found in the other, higher families.
This holds good especially in the structure of the face. Although the
narial aperture is drawn backward into proximity with the anterior
wall of the braincase the nasal bone is not wholly misshapen. It
retains part of its long, narrow form and it still roofs over the hind
part of the nasal cavity which may yet inclose very considerable
remnants of the ethmoid plates. The anterior part of the nasal cavity,
bounded by the intermaxillary, maxillary and vomer, is also relatively
primitive in structure, more open than usual, with less tendency to
closing together of the bones. And the maxillary, although it has
expanded backward, and shoved itself somewhat back both above and
beneath the frontal, has nevertheless not in any way covered the
frontal’s broad supraorbital process. A distinct lacrimal is present,
but this is not unknown among higher cetacea. The zygomatic arch
has retained more of its primitive form and strength than elsewhere.
Two outer nasal apertures are still found; they are not mutually
united. The basal part of the occipital is also to a somewhat less
degree modified than in other recent cetacea, being less specialized to
accommodate larynx and gullet.

That which more than anything else has left its impress on the
known Baleenids is their habit of not hunting after single large fish,
but of swimming with open mouth into shoals of small fish, crus-

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE V7

taceans, or other small creatures, which they allow to stream into the
mouth in multitudes along with the water. They seek to retain the
edible contents when they close the mouth and the water flows out
again between the lips. The water has thereby acquired great power
to act upon the mouth cavity from within; it distends the opening
enormously ; the jaws grow and acquire a disproportionately large
size in comparison with the braincase; the branches of the mandible
are bowed strongly outward to the sides and are widely divergent
from each other behind, while the connection between them in front
becomes quite loose. The gigantic lower jaw wears so upon the
ligaments which bind its articular head to the squamosal that the
ligaments are incited to growth. They become uncommonly strong
and spread themselves over the original gristle-covered articular
surfaces on the jaw and the squamosal, both of which surfaces they
entirely cover. They cause the squamosal to grow out as a huge
process which bears the attachment surface for the lower jaw on its
free margin. By the enlargement of the mouth cavity the squamosal
together with the articular head of the lower jaw is pushed far out
to the side and so far back that at last its free postero-external
extremity comes to lie further back than the occipital condyles. The
squamosal in its turn presses strongly on the parts which lie behind
it: on the mastoid which is squeezed inward, and on the exoccipital
which is pushed backward. In proportion to the size of the under
jaw the temporal muscle increases and pushes its region of origin
-forward over the supraorbital process. There has been no use what-
ever for the teeth; they atrophy so completely that finally they are to
be found only in the embryo as a long series of insignificant, small,

pin-shaped teeth, hidden under the skin and soon resorbed. On the
other hand the inflowing and outflowing water acted as a stimulant
on the corneous papillz of the roof of the mouth. The papille along
the margin of the upper jaw are so stimulated that they have grown
out as a close-set series of ‘‘ whale-bones ”: high, crosswise-placed,
corneous plates, the inner margin of which is frayed out into threads.
_ The entire set of whale-bones functions as an excellent instrument

for catching the solid material that flows with the water into the
open mouth. The palate is strongly acted upon by the instreaming
water, by the larynx, and by the tongue, which is pressed against it
when the water is to be expelled. The palatine bone grows and forces

itself backward, pushing back the pterygoid behind it; and the
pterygoid pushes and presses that which lies still further back ;
namely, the tympanic bulla and the region of attachment of the neck

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

muscles on the basal part of the occipital. The palatine may push
itself wholly back under the base of the occipital, and the muscle
attachment may come to lie about on a line with the posterior margin
of the occipital condyle.

In other respects also the known Baleenids have reached particu-
larly high. The supraorbital process of the frontal acquires an
unusual breadth, no doubt for the special reason that it follows the
eye, which, by the widening of the mouth cavity, is pushed out to the
side. The supraoccipital becomes very large and strongly slanting
forward under the influence of pressure by the water and by the
muscles of the neck. The transverse processes of the thoracic verte-
bree become widely projecting ; this is especially noticeable as regards
the hindmost thoracic vertebree (where the processes are para-
pophyses, while on the anterior vertebrzee they are diapophyses).
The ribs have a strong tendency to lose the capitulum and to restrict
their connection with the sternum. In most of the recent members
of the family the capitulum is absent from all the ribs, even the more
anterior, although an evident collum is present (it is, however, doubt-
ful whether it is really the capitulum that is absent from the hind-
most ribs; more likely the single articular head which appears to be
the tuberculum is in reality either the capitulum alone or the capitulum
and tuberculum undifferentiated). The sternum is so reduced that
it consists of the manubrium alone. The first finger has a tendency
to atrophy. Etc.

In the section Balzenini are found the most primitive of the family’s
known genera: Balena and Neobalena. With them the anterior
facial part of the skull has kept relatively much of the form ordi-
narily present in mammals. This is especially true of the inter-
maxillary and still more of the maxillary, which is quite slender in
front and not depressed. Body and tail are rather short, not quite
so well fitted for rapid swimming as in the others. The hand is more
primitive. Of the hind limb’s skeleton there are present, at least in
Balena, relatively quite considerable remnants, among other parts a _
stunted femur and the upper end of the tibia. The mouth is shaped
somewhat differently than in the others ; it is formed as an enormous
barrel or bag, bowed outward on all sides. Not only are the rami of
the lower jaw bent outward, but the upper jaw with the whole facial
part of the skull is also bent, arched highly upward. Both the upper
jaw and the branches of the lower jaw assume the structure of stays
in the walls of the pouch-like mouth cavity. The whale-bone plates
acquire a remarkable length. Finally the head becomes more pre-

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE I9

ponderant as regards the body than in other whales. The cervical
vertebrz are pressed together unusually strongly ; they coalesce.

Balena appears in some respects to stand on a lower level than
Neobalena. Its slender under jaw seems better to agree with the
condition primitive to the cetacea than does the strikingly massive,
strongly compressed under jaw of Neobalena in which the mandible
presumably must be especially influenced by the large under lip. Its
relatively few, ordinarily formed, slender ribs, and its correspond-
ingly rather long series of lumbar vertebre are also undoubtedly
primitive characters; in Neobalena the ribs have become unusually
numerous and the number of lumbar vertebrz is reduced to a few
bones, while the ribs, or at least most of them, have become remark-
ably broad and have to a remarkable degree lost connection with the
vertebre so that they lie loose among the muscles. Balena is no
doubt the more primitive also in the short, broad form of the hand.
The first finger is either (in B. australis) rather well developed, con-
taining two phalanges in addition to the metacarpal, or (in B. mysti-
cetus) reduced, though still retaining the metacarpal.“ The other
fingers are not much lengthened; in the median digits, however,
especially in the third, the number of phalanges may be increased to
four or five. The form of the phalanges is terete, not compressed.
In Neobalena, the hand appears to have essentially the same structure
as in Balena, but the first metacarpal is said to be absent, and the
entire hand has become narrower. The lack of a dorsal fin in Balena,
in contrast with Neobalena, is, presumably, also a primtive character ;
though the fin may have been lost. But in the adaptation of the head
as a pouch for catching small animals Balena has reached far beyond
_Neobalena. In the more primitive of the two certainly known species
of Balena, B. australis, the modification is a little less noticeable than
in the higher species, B. mysticetus; the head is slightly smaller, the
upper jaw is somewhat less bowed upward, etc. In B. mysticetus the
head becomes so huge that in full grown individuals it reaches a
third or more of the animal’s total length, the upper jaw is thrown
upward in an enormous arch, the palatine and pterygoid are forced
backward under the hindmost part of the basioccipital, etc. The
coracoid process of the scapula may be absent (in B. australis).

Neobalena must assuredly have originated from Balena, but from
one of the most primitive species of the genus, in which the head was
only a little increased in size; but since then it has gone its own way,
developing peculiarities in the form of the lower jaw, in the ribs,
vertebral column and hand.

2

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL

In the genera of the section Balenopterini the intermaxillary and
also the maxillary are rather strongly flattened anteriorly so that the
facial part of the skull has lost its primitive pointed form. Body and
tail are uncommonly long, adapted to more rapid swimming with
stronger muscles. Among the alterations produced by these muscles
are the higher spinous processes on the dorsal and caudal vertebre.
The hand is shaped more like an oar blade. The fingers are laid more
closely together and the third and fourth may have the number of
phalanges increased ; the first digit has completely disappeared. The
skeleton of the hind limb is more reduced. The mouth is modified
in its own way; its outbowing in the upward direction is slight or
absent, that to the sides and downward is conspicuous. The floor of
the mouth cavity has become to a high degree expansible, and the
intermaxillary and maxillary, like a broad, more or less flattened lid,
cover over the pouch which it forms. The Balznopterines stand
lower than the known Balznines in the condition of the cervical verte-
bree: the bones retain their freedom.

Among the known Balznopterines, Rhachionectes is one of the most
primitive. Its nasal bone is still relatively very well developed. The
breadth of the intermaxillary and maxillary in front is rather slight.
The supraorbital process is relatively weak and not strongly flattened.
The braincase is relatively only slightly telescoped so that on the
middle of its upper side there can be seen not a little of the frontal.
The supraoccipital is not especially large or forward-slanting. The
articular surface for the lower jaw on the squamosal is not pushed
out especially far downward and backward, and, when seen from
beneath, has not entirely covered the mastoid or pushed the exoccipital
very far backward. Bony palate relatively not strongly lengthened
behind. The point of attachment for the neck muscles on the basal
part of the occipital is still tubercular, and the basioccipital on the
whole is only to a slight degree shaped to accommodate the larynx
and gullet. . In contrast with its nearest recent allies Rhachionectes
stands lower in a few other respects also: an evident capitulum is
still found on some of its anterior ribs; the skin beneath its mouth
cavity is not thrown into longitudinal folds; the dorsal fin is not
present ; the hand is relatively short, and the number of phalanges is
only a little increased. It has perhaps high specializations in its
decidedly heavy under jaw, which slightly suggests Neobalena, and
in its somewhat upwardly arched facial portion of the skull.

Plesiocetus from the Tertiary of Europe, and presumably from that
of North America also, is best known from the skull. To a high

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 21

degree it resembles Rhachionectes, but appears to differ in having a
considerably more reduced nasal, like the higher Balzenopterines.

Cetotherium, also from the Tertiary of Europe and presumably of
America, which is likewise known from scarcely anything else than
the skull, is a very near relative to Plesiocetus. It is slightly more
specialized, with the articular surface for the lower jaw on the
squamosal pushed somewhat further backward, pressing more against
its surroundings, and covering the mastoid; but otherwise it scarcely
differs except in trifles.

As a pronounced contrast to Rhachionectes the genus Balenoptera,
on the other hand, stands much higher: with much smaller nasal;
with intermaxillary and maxillary more broadened in front; with
broader and flatter supraorbital process; with more telescoped brain-
case in the median upper part of which there shows itself only a little
of the frontal; with larger, more forward-slanting supraoccipital ;
with the articular surface for the lower jaw on the squamosal pushed
much further backward, wholly covering the mastoid and shoving
the exoccipital more to the rear ; with bony palate strongly lengthened
backward ; with point of attachment for the neck muscles on basal
part of occipital compressed, flattened; with the basioccipital more
shaped to the larynx and gullet ; with the skin under the floor of the
mouth cavity thrown into longitudinal folds; with a dorsal fin; with
the hand more lengthened; with frequently more phalanges in the
median fingers.

Magaptera stands yet higher than Balenoptera. Its body is
relatively not much elongated, a fact which points to its origin among
the most primitive species of Balenoptera. But in the structure of

the fore limb it has reached far beyond its relatives. On account of
some special use or other, perhaps most likely from rapid turning
about in the water, the arm has grown to an enormous length. The
forearm has become very much stretched out, and the hand is yet
more conspicuously lengthened, the number of phalanges in the third
and fourth finger increased in addition. The scapula has lost both
the coracoid process and the crest.

Balenide.™
I. Intermaxillary and maxillary narrow anteriorly, not flattened.
Balenini.
A. Mandible slender. Ribs not broad. First metacarpal
present.
Balena.

B. Mandible robust. Most of the ribs broad. First
metacarpal absent.
Neobalena.

iS)
iS)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

II. Intermaxillary and maxillary broad anteriorly, flattened.
Balenopterini.
A. Nasal relatively well developed.
Rhachionectes.
B. Nasal reduced.
1. Area of insertion of neck muscles on basal part of
occipital tubercular.

a. Articular surface for lower jaw on squamosal
not strongly pushed backward, not covering
the mastoid when seen from below.

Plesiocetus.

b. Articular surface for lower jaw on squamosal
more pushed backward, covering the
mastoid.

Cetotherium.
2. Area of insertion of neck muscles on basal part of
occipital compressed.

a. Hand not especially elongated. Shoulder
blade with crest.

Balenoptera.

b. Hand greatly elongated. Shoulder blade

without crest.
Megaptera.

Squalodontide.—The Squalodonts must have originated from
among the most primitive Baleenids which still had the teeth shaped
like those of the Zeuglodonts but increased in number, and which had
not yet begun to get the mouth refashioned into a catching-bag. Their
differences from the most primitive Balenids are due especially to
stronger pressure of the water on the facial part of the skull; most
likely the Squalodonts were from the beginning more rapid swimmers
than the Balenids. The nasal passage is pushed much further back,
not by muscle action alone, but probably especially by the influence of
the facial adipose cushion. The water both stimulates the cushion
to growth and presses it against the nasal passage. The nasal bone
is completely atrophied, almost tubercular in form, and pressed into
the frontal in the fore wall of the braincase, not or almost not cover-
ing over any part of the nasal cavity. The plates of the ethmoid are
probably pushed wholly away and the lamina cribrosa has probably
become a solid bone-plate without perforations or almost so. The
nose muscles, the pneumatic diverticulum from the nasal passage, the
adipose cushion of the snout, in short all that covers the skeleton of

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 23

the face, is pressed by the water in against the bones and has moulded
the upper side of the whole face as its bed, especially hollowed out
posteriorly. This bed, the “facial depression,” extends backward
along the sides of the nasals on the forehead. The beginning is
traceable of a remarkable peculiarity which, in the more advanced
cetaceans may become conspicuous to a high degree: an asymmetry
in the structure of the face. The head must no doubt be so held
during motion that the water comes to press not quite equally on both
sides, but more strongly on the right side than on the left. The facial
cushion therefore becomes larger on the right side than on the left,
extends its bed most on the right side, forces the nasal passage to
bend over to the left, and causes the bones of the face to develop
somewhat dissimilarly on the two sides.” The maxillary has pushed
itself posteriorly up over the frontal to such an extent that it almost
wholly covers it, also spreading out over the supraorbital process.
The zygoma appears to have been quite slender. The two nasal aper-
tures were presumably united into one. In all of these points of
difference from the Balzenids the Squalodonts agree with the higher
cetacea, of whose most primitive forms they remind one in nearly
everything, so far as they are known, except in the condition of the
teeth. .

The Tertiary North American Agorophius, which is only known
from a very incomplete skull, almost without teeth, appears to be the
most primitive member of the family. The number of teeth is not
known, but certainly, to judge by the other peculiarities of the genus,
it must have been greater than typical. Its braincase is much less
telescoped than in the other Squalodonts, also less than in any of the
known Balznids, somewhat suggesting the Zeuglodonts in being
relatively strongly constricted anteriorly between the large temporal
fossee, and in having the parietal form a considerable part of its roof.
In the other Squalodonts the braincase, so far as it is known, is so
telescoped and so broadened out to the sides that there is a wide area
between the temporal fossz although these are relatively large; also
the parietal in the middle of the roof of the braincase shows itself at
most as a narrow band. In other respects Agorophius appears to
agree well with Squalodon.

Squalodon is known rather completely from skulls from Tertiary
strata in both the Old and New Worlds. Almost nothing is known
of other parts of the skeleton. The teeth are well developed, hetero-
dont. In each jaw there are three incisors with conical crown and
single root, a canine of similar form and size, and I1, or sometimes

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

in the upper jaw 12, cheekteeth. Of these last the four or five
anterior have conical crown and single or bifid root, and the seven
posterior have compressed crown with more or less serrate anterior
and posterior margin (or only the posterior margin serrate), and
two roots, an anterior and posterior. The fact that the number of
cheekteeth is most often 11 might indicate that the increase above
the typical number, seven, was produced by the intercalation of four
milkteeth in the series with the seven permanent teeth; but there is
no decisive evidence either for or against this explanation, as the tooth
succession is not known. If the number exceeds 11 a true increase
must have taken place. As in the most primitive of the higher whales
the jaws are very elongate, narrow; the intermaxillary and maxillary
are not especially closed together, and the mesethmoid is not ossified
in front; the symphysis menti is long (the rami of the mandible may
have grown together) ; the facial depression does not extend very far
backward, the braincase is relatively only a little telescoped, the
temporal fossa is considerable, the zygomatic process of the squamosal
strong, the occipital condyle projecting.

The Tertiary European Neosqualodon is only known from pieces
of jaws. It has the number of serrate cheekteeth raised to at least
11; otherwise the characters of the dentition are not known.

The Tertiary Argentinian Prosqualodon, known from the more
essential parts of the skull, differs from Squalodon in having a much
shortened face, with the facial depression relatively strongly broad-
ened behind. The number of teeth appears to be somewhat reduced.
It is no doubt a little more primitive than Squalodon in the less
strongly telescoped form of the braincase proper.

Squalodontidz.”
I. Braincase only slightly telescoped.
Agorophius.
II. Braincase strongly telescoped.
A. Face long.
1. Number of cheekteeth relatively little increased
above the typical [44].
Squalodon.
2. Number of cheekteeth increased above the typical 44.
Neosqualodon.
B. Face shortened.
Prosqualodon.

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 25

Platanistide.—The Platanistids no doubt originated from the
most typically defined Squalodonts such as Squalodon. The most
important and perhaps in the first place the only distinction between
the Platanistids and their precursors among the Squalodonts is that
the teeth in the Platanistids are to a higher degree structurally degen-
erated. They have lost their heterodonty, have become smaller but
more numerous, all of them nearly simply conical with a single root.
On the other hand the Platanistids have retained most of the other
peculiarities in which the Squalodonts show themselves to be relatively
primitive. Especially noticeable in comparison with higher cetacea
are the following characters: facial depression rather narrow, not
much widened laterally behind, so that its outer margin covers over
the temporal fossa to a slight degree only; temporal fossa rather
large ; zygomatic process of the squamosal robust; all these peculiari-
ties are no longer or scarcely ever found among cetacea of the higher
families. The braincase appears to be rather small and not very
much compressed antero-posteriorly, this also in contrast with the
higher whales. In common with the most primitive forms of the
higher cetacean families the Platanistids have, so far as they are
known, such peculiarities: as the mutual independence and rather
considerable size of the cervical vertebrz ; as the conspicuous lack of
uniformity in the shape of the dorsal vertebre (for instance the
long, broad transverse processes of the lumbar vertebrz in contrast
with the rather short processes of most of the thoracic vertebrz ; only
on a few of the hindmost thoracic vertebree do there occur robust
transverse processes, parapophyses, which bear ribs at their tips in
the Platanistids that are known in this respect ) ; as the well-developed
heads to the anterior ribs, and probably also the coalescence, or more
correctly the non-separation, of the tuberculum and capitulum on the
hinder ribs; as the rather large, ossified costal cartilages ; as the rather
well-developed sternum ; as the presence of the first finger, the meta-
carpal at least of which is found; as the rather slight lengthening
of the middle fingers, etc.

In one single direction the known Platanistids have developed
themselves highly. They have used the jaws as a kind of delicate
forceps to seize and hold prey that did not make very strong resis-
tance. The jaws grow to an unusual length but become noticeably
slender, fine, though solid. The intermaxillary and maxillary press
close together, covering over the anterior part of the mesethmoid, and
they have a tendency to coalesce. The maxillary has pushed itself
forward anteriorly beyond the tip of the intermaxillary. In the lower

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

jaw the symphysis menti becomes very long, and the rami of the
mandible are inclined to grow together. The teeth that lie at the
front of the jaws are inclined to increase in size, probably because the
tips of the jaws come to be the most used part of the forceps. In
another respect also the known Platanistids stand high: the sides of
the facial depression are inclined to grow upward. Perhaps they also
stand high in the tendency of the pterygoid to widen itself out
unusually far backward in the outer wall of the air-sac behind the
palate, reaching back to the squamosal; a circumstance that may call
to mind both lower and higher whales, Baleenids and Physeterids.*

The genus of Platanistids which has removed itself least from
Squalodon appears to be the Tertiary South American Pontistes,
which is known from most of the skull. In relation to one or another
of its nearest allies it has the following primitive characters : the teeth,
judging from the alveoli, were small, simply formed, the anterior not
enlarged ; the toothrows stand rather distant from each other, as the
palate is relatively broad; the outer margin of the facial depression,
especially the longitudinal crest on the maxillary above the orbit, is
relatively low. A character which must be considered advanced in
comparison with the nearest relatives is the specially large number of
teeth, about 40 in each jaw if one judges rightly from the fragments
of toothrows that have been found.

Near to Pontistes but on a slightly higher level is Pontoporia
(Stenodelphis). The teeth have become smaller but more numerous,
about 55 in each jaw. Those in front have a slight tendency to be
enlarged. The toothrows are placed nearer together and the palate
is narrower. A high specialization, which also holds good for the
other recent members of the family, is the complete absence of the
olecranon.

Lipotes (known from external characters, skull, and cervical verte-
bre) and Jia are near relatives of Pontoporia. Their face is shorter,
the number of teeth is less (about 30 in each jaw in the former, about
26 in the latter), the anterior teeth show scarcely any tendency to be
enlarged. It might appear as if the two genera were, in these char-
acters, less specialized; but the explanation is presumably another.
The two genera most likely originated from forms that more nearly
resembled Pontoporia, and that had the strongly narrowed palate and
numerous small simply conical teeth, although not so many as in
Pontoporia. Lipotes and Inia appear to have used the teeth in a
special manner, most probably for the crushing of food, and the teeth
have therefore regained some of their earlier strength, have grown

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 27

and become massive, with wrinkled enamel. In compensation, how-
ever, they have become less numerous and the most posterior in the
jaws have acquired a form which is anything but primitive: the base
of the crown is more or less tubercularly widened out inward. The
narrow palate is retained. The lateral margin of the facial depression
is considerably more upturned than in Pontoporia, and the posterior
border, especially in the median region formed by the frontal, is far
more elevated.

Lipotes as compared with nia is surely the more primitive in the
greater slenderness of the teeth; on the contrary it is the less primi-
tive in having the facial depression relatively strongly widened behind.

Saurodelphis (Saurocetus, Pontoplanodes), Tertiary, Argentinian,
known from most of the skull, appears to have also originated from
Pontoporia-like animals, but it has gone in another direction than
Ima. It has retained the slender face with the narrow palate, but the
number of teeth is reduced to about 20 in each jaw. At the same time
the teeth are enlarged ; in any event they have acquired roots that are
more widened antero-posteriorly ; this is especially true of a number
of the anterior teeth in each jaw. In these teeth the root appears to
be in process of dividing in two, so that in cross-section it is almost
8-shaped, a form which, especially as regards the anterior teeth, is
quite the opposite to primitive. The lateral margin of the facial de-
pression is trenchant and highly upraised, even more than in Jnia.
The hinder margin of the depression is not only elevated in the middle
as in Ima, but is also pushed further back.

Platanista also probably traces its origin back to Pontoporia-like
creatures. It has gone further than any other member of the family
in the direction of making over the jaws into delicate forceps. The
face is so slender and the palate so narrowed that the right and left
toothrows in the upper jaw lie closely side by side; they may even,
especially at the extreme rear, where the teeth are undergoing
atrophy, be pushed into each other. Somewhat similar conditions
obtain in the lower jaw. The number of teeth is about 30 in each
jaw. Several teeth at the front of each jaw have acquired high,
pointed crown and compressed, enlarged root. The outer margin of
the facial depression has grown upward, higher than in any other
genus, especially that part of it which runs along the outer margin of
the maxillary over the orbit and front of the temporal fossa. This
part has shaped itself quite fantastically as a huge plate which rises
high upward and bends in over the posterior part of the face, which it
covers like a mask, since each plate nearly meets its fellow from the

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

opposite side. On the other hand the posterior margin of the depres-
sion is not particularly highly elevated. The eye is atrophied ; touch
more than vision probably guides in the capture of prey. The hand’s
unusually broad, rounded-off outline, with the especially short, uni-
form, well-developed and wide apart fingers, might appear more
primitive than in other members of the family; but possibly it may
be the story of a partial reversal from an earlier more flipper-like
condition.

Platanistide.”
I. Upper toothrows well separated throughout. Maxillary with
longitudinal crest not excessively large.
A. All the teeth with terete or scarcely compressed root.
1. Longitudinal crest on maxillary relatively low. Frontal
behind nasal only a little elevated.
a. Palate relatively broad. About 40 teeth in each
jaw.
Pontistes.
b. Palate relatively narrow. About 55 teeth in each
jaw.
Pontoporia | Stenodelphis |.
2. Longitudinal crest on maxillary relatively high.
Frontal behind nasal rather strongly elevated.
a. Teeth relatively slender.
Lipotes.
b. Teeth relatively robust.
Inia.
B. Teeth with compressed root; in some of the anterior teeth
the cross-section of the root is almost 8-shaped.
Saurodelphis.

II. Upper toothrows placed close together, especially behind, so
that teeth from the right and left sides may be pushed in
among each other. Maxillary with longitudinal crest
excessively large, completely covering over the face.

Platanista.

Delphinide.—The most important character—perhaps in the
beginning the only one—which has separated the Delphinids from
the most primitive Platanistids from which they sprang is the widen-
ing out of the facial depression. This broadens posteriorly to such
an extent that its floor wholly covers over the temporal fossa like a
roof formed by the frontal and maxillary together. A second

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 29

peculiarity, which in any event soon showed itself, is the reduction of
the temporal muscle through lack of use; its fossa becomes smaller
and the zygomatic process of the squamosal becomes less projecting
and less robust, losing, moreover, its primitive arched form. In the
more advanced members of the family many other modifications may
appear. The facial part of the skull, which begins by being long and
narrow, almost compressed, used as forceps, may become still longer.
Or the use as forceps may grow less and be exchanged for service as
an implement for rooting in the sea bottom; followed by alteration in
the form of the face. Or the mouth is used merely to clap together
around the prey; followed by flattening and shortening of the face.
In each instance the facial cushion contributes to the flattening of the
facial part of the skull. The extreme tip of the intermaxillary has
the tendency to be restricted, to be grown over by the maxillary and
to lose the teeth which at first were implanted in it. Its upper margin
may extend in over the anterior part of the mesethmoid and coalesce
with its fellow of the opposite side. The symphysis menti has a
tendency to be weakened and shortened. The teeth are inclined to a
further reduction, and they may disappear; but they may also be
again applied to special work and be modified in various ways. The
braincase increases in size and is more subject to pressure from in
front and from behind. The nasal passage may be pushed further
back. The nasal bone, which in the most primitive Delphinids
retained a slight trace of its earlier function as a cover for the nasal
cavity, becomes in most cases quite sunk into the frontal. The
occipital condyle, which at first is rather projecting in the usual
manner, becomes flattened out and pressed in against the wall of the
braincase. The cervical vertebrzee may coalesce. The thoracic verte-
bree acquire unusually long transverse processes which are especially
noticeable on the hindmost of the series. Most of them are dia-
pophyses except the most posterior ones; these are parapophyses.
Only the anterior ribs retain the capitulum. On the hindmost ribs
the capitulum disappears entirely, and the rib is articulated with the
tip of the long transverse process by the tuberculum only. (As in the
Platanistids the single articular head on the very hindermost ribs is
presumably formed by the capitulum or by the capitulum and tuber-
culum undivided.) The flippers may be lengthened. Etc. The ptery-
goid varies capriciously. It is true that it always spreads inward
under the posterior nares ; but it is sometimes rather widely separated
from its fellow of the opposite side, sometimes almost in contact with
it, while probably after having been in the latter condition it may

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

withdraw. The number of vertebrz also varies in a quite capricious
manner.

One of the most primitive Delphinids is no doubt the Tertiary
North American Delphinodon, the skeleton of which is known rather
completely. In comparison with its various relatives it has the
following primitive peculiarities. The teeth are present in large
numbers. They are small and nearly simply conical, some of them,
however, with wrinkles or small projections on the base of the crown,
probably mementos of the crown’s formerly serrate margins and of
its also otherwise less simple form. The facial part of the skull is
rather long and narrow. As in the related forms which are lowest
in this respect the anterior end of the intermaxillary was probably
freely projecting, tooth bearing, and not grown over by the maxillary.
The upper margin of the intermaxillary does not come in contact with
its fellow. The symphysis menti is long; nasal slightly projecting ;
cervical vertebrz distinct. It shows a peculiarity of its own in having
a longitudinal crest on the projecting lateral part of the basioccipital.

The Tertiary European Champsodelphis (judging chiefly from
Ch. ombonit, Acrodelphis) presumably stands near to Delphinodon.
It is known from scarcely anything else than scanty remains of the
skull. It shows high specialization in the modification of the rostrum
to serve as an implement for boring or rooting in the sea bottom.
The facial part of the skull has acquired an unusual length and
slenderness; the teeth have probably disappeared from the inter-
maxillary, and the upper margin of this bone was probably in contact
with its fellow through a considerable part of its extent. ,

The Tertiary European Schizodelphis (judging from S. sulcatus,
Cyrtodelphis), also known practically from the skull only, must be a
near relative of Champsodelphis with which it appears to have most
of its peculiarities in common, both the primitive characters and the
special modifications. Its most important difference appears to be
that its teeth have gone still further in the direction of simplicity ;
the only reminders of earlier, less reduced form that have remained
behind are a slight widening out of the crown’s base, which may be
found on some of the teeth, and the trenchant character of its anterior
and posterior margins.

The Tertiary European Heterodelphis, which is known from rather
considerable parts of the skeleton, undoubtedly stands close to Schizo-
delphis. Its teeth have become still more simple, with purely conical
crowns.

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 31

In the Tertiary European Eurhinodelphis, which is known from
most of the skull and from parts of the rest of the skeleton, the
transformation of the snout into a rooting implement has reached the
highest limit. The rostrum, both upper jaw and lower jaw, has
grown forward anteriorly as a long slender point, still more notice-
ably than in any of the other genera. The intermaxillary has ex-
tended itself, awl-shaped and toothless, far forward beyond the
maxillary. The tip of the lower jaw appears to be formed in a corre- -
sponding manner. The teeth are simply conical.

No doubt the Tertiary South American Argyrocetus is very nearly
related to Eurhinodelphis. It is known from a defective skull, and
appears to differ in trifles only.

The genera just mentioned of the group Eurhinodelphini form a
contrast with the group Monodontes. The latter includes the genera
Delphinapterus and Monodon, which must have originated from the
oldest Eurhinodelphines in which the tip of the snout had not been
remodeled as a rooting implement. In common with the Eurhino-
delphines (at least with Delphinodon, Heterodelphis, Eurhinodelphis,
and Argyrocetus, which are known in this respect) the Monodonts
alone among the Delphinids have the primitive character that the
cervical vertebree are mutually independent. Other indications of
low origin seem to be shown by the Monodonts in the form of the
teeth (in which one of the genera may recall Delphinodon and
others), in the decidedly short spinous and transverse processes of
the thoracic vertebrze, in the relatively short fingers, and perhaps also
in the absence of the dorsal fin. But in the flat and broad form of the
face, probably resulting from their habit of not using the jaws for
_ much else than to clap together on tender cuttlefish, the Monodonts
are more highly developed than their progenitors among the Eurhino-
delphines. The same is true of their lack of the olecranon.”

The most primitive of the known Monodonts is Delphinapterus.
It shows its primitiveness in relation to its nearest ally by its rather
ordinary dentition: the teeth are present in relatively considerable
numbers, about Io in each jaw; they are small and conical, but in the
upper jaw they are directed forward in a peculiar manner. The
teeth have disappeared from the intermaxillary.

In Monodon the teeth, with a single exception, are in process of
atrophy and disappearance; only a few of them are present in the
young. One of the foremost teeth in each maxilla has had its peculiar
destiny: it has grown forward as a “ramming-tooth,” at first no
doubt uniformly in the right and left jaw and in the male and female,

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

probably used, in a similar manner to the upper canine in the walrus
or to the tip of the snout itself in the Eurhinodelphines, for rooting
in the sea bottom. Such a function might be initiated by forward-
slanting anterior teeth like those in Delphinapterus, but later it must
have been restricted to the ramming-tooth in the left jaw of males.
The work of the males may possibly be of service to the females
because the species is gregarious. The ramming-tooth has become a
_kind of male secondary sexual character and has grown to an ex-
aggerated size as the well-known “ unicorn horn.” As a memento of
an earlier condition the ramming-tooth is still found in a reduced
form in the right upper jaw of the male, and in both upper jaws of
the female; in rare instances it may even now be found in the male
well developed on both sides | when the spiral of the two tusks is
parallel].

All the other Delphinids are contrasted with the Eurhinodelphines
and Monodonts by the partial or complete coalescence of the cervical
vertebre, the atlas and axis being always united. The genera in
question constitute a compact group, rich in forms, which traces its
origin back to low Eurhinodelphines.

Lowest of all stands the section Delphini, whose most primitive
known genus is undoubtedly Steno. This has still the primitive
Eurhinodelphines’ long, but not exaggeratedly long, narrow, scarcely
flattened fore-face, with long toothrows and long symphysis menti.
It is indeed scarcely distinguishable from the primitive Eurhinodel-
phines except by its partly ankylosed cervical vertebrae. The circum-
stances which place it low among its nearest relatives are the facts
that the teeth have fluted enamel, and that the symphysis menti is
long. The peculiarity of the enamel is presumably a slight reminis-
cence of an earlier, less-reduced condition.

Very near Steno comes Prodelphinus (probably including “ Sota-
lia’’), not differing in much else than that the enamel is smooth, not
fluted, and that the symphysis menti is shortened.

Delphinus differs from Prodelphinus in scarcely anything else than
a peculiar palate form: at the inner side of the toothrow the bony
palate is hollowed out into a long longitudinal furrow which is
especially deep behind. The intermaxillaries have a relatively strong
tendency to coalesce and to cover over the mesethmoid. A few small
teeth may be found in the intermaxillary as in Steno and Prodel-
phinus; in most of the Delphinids the teeth have wholly disappeared
therefrom.

In contrast to the Delphinines the other higher Delphinids have
the skull’s fore-face shorter and more depressed. As a beginning the

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 33

difference is only slight, but it finally increases so as to become very
noticeable. At the same time that the anterior facial part of the skull
is shortened, because the mouth is no longer used as a pair of forceps
but as a “clap-trap,” it becomes flatter and broader, while its upper
side is more pressed upon by the facial cushion. The cushion becomes
larger, especially widening itself out anteriorly and pushing into the
originally slender “beak.” The intermaxillaries, in their anterior
portion particularly, together lose their structure as an upstanding
roof-ridge, and finally become quite flat, each of the bones widened
out.

The genera of the section Legenorhynchi depart so slightly from
the more primitive members of the section Delphini, such as Pro-
delphinus, that there would scarcely be any reason to set them apart
in a special group were it not evident that they represent the begin-
ning of new series of forms.

Doubtless Tursiops occupies the lowest position. The anterior
facial part of the skull is indeed broader than in Prodelphinus, but
it has, however, not lost its form as a roof-ridge, and it has still a
considerable length.

Near Tursiops probably belongs Tursio [Lissodelphis|, which
also has the fore-face rather long, though more flattened. Another
difference is that it lacks the dorsal fin, either because it has lost it or
has never acquired it.

Lagenorhynchus (to which should probably be joined Cephalo-
rhynchus and Sagmatias, and perhaps “ Feresa”) has gone a step
further than Tursiops and Tursio in the direction of shortening and
flattening the rostrum.

Among the Delphinids in which the process of shortening and
flattening the rostrum has been more perfected the members of the
section Globicipites are contrasted with those of the section Phoceenz
by reason of their greater primitiveness. In them the crowns of the
teeth have retained their primitive conical form, while in the Pho-
ceenans the crowns have become entirely peculiar.

Orca [Orcinus] is the one among the Globicipites which has
retained most of the ordinary dolphin type in the structure of the
rostrum, particularly as regards the narrowness of the intermaxillary.
The rather short, rounded-off form of the hand might appear to be
primitive also, but various circumstances strongly indicate that it has
arisen through the shortening of an ordinary, pointed, porpoise
flipper: the number of phalanges in the second finger is rather large;
the finger is merely more strongly arcuate than usual. In the trans-
formation of the dentition to a conspicuously powerful biting imple-

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

ment Orca has gone further than any other genus of the family. It
has habituated itself to living on large prey such as seals and the
smaller cetaceans, and it even slashes into the largest. The teeth are,
it is true, relatively few, about 12 in each jaw, but in compensation
they are massive.

Orcella has reached higher than Orca in the great breadth of the
intermaxillary, but it must have originated at a level lower than that
on which Orca stands, sinte its teeth are small and rather more
numerous, while its hand is essentially like an ordinary porpoise hand.
The genus gives the impression of being a dwarf form with notice-
ably large braincase in proportion to the face.

The following genera of Globicipites must have originated from
Delphinids that were essentially like Orcella but without the dwarf-
ing. Each has gone its own way. There is, however, one peculiarity
that unites them: the hand has acquired an uncommon length and
narrowness, though in different degrees, at last with an unusual
number of phalanges in the second finger.

In “ Grampus”’ the intermaxillary has retained a breadth similar
to that in Orcella. Although the hand is long and narrow there are
only about eight phalanges in the second finger. The chief peculiarity
of the genus lies in the atrophy of the dentition: only a few and
rather small teeth remain. These are at the front of the mandible,
and with age they may entirely disappear.

In Pseudorca the hand is essentially as in Grampus. But the inter-
maxillary has acquired a very noticeable breadth anteriorly, and the
dentition is developed in a similar manner as in Orca.

The intermaxillary is conspicuously wide in Globiceps [ Globi-
cephala| also; it may be even wider than in Pseudorca. Peculiarities
of Globiceps are: that the nostril is pushed unusually far backward,
that the dentition is atrophied so that only a few, about 10, small
teeth remain, situated at the front of the jaws, and that the hand is
conspicuously long, with as many as 14 or more phalanges in the
second finger.

The section Phocene presumably originated among the most primi-
tive Globicipites or perhaps Lagenorhynchi. That which places this
group in contrast not only with the Lagenorhynchi but also with all
other Delphinids is the peculiar form of the teeth. The teeth are
present in large number and are of small size. Some of the foremost
and hindmost may have about the usual conical crown, and all of them
are single rooted. Most of the teeth, however, have the crown com-
pressed, widened out fan-wise or leaf-wise, and often with notches in

No. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 35

the margin; a form which is not only unique among the cetacea, but
the contrary to the forms found among the most primitive cetacean
genera.”

Phocena is, besides, a little broad-snouted porpoise, rather evenly
developed in all directions. A few small teeth may be present in the
intermaxillary.

Neomeris (Neophocena) is nearly related to Phocena. It differs
in having acquired a yet shorter and broader face, in having a notice-
ably spacious braincase, and in lacking the dorsal fin, which it doubt-
less has lost.

Delphinide.”
I. Atlas and axis mutually free.
A. Face long and narrow, not flattened.
EURHINODELPHINI.
1. Intermaxillary (undoubtedly) not specially elon-
gated in front of maxillary.
1. Face not noticeably elongated.
Delphinodon.
2. Face noticeably elongated.
a. Teeth with slight traces of less simple
forms.
a. Crowns of teeth partly with remains
of lateral cusps.
Champsodelphis.
B. Crowns of teeth without lateral
cusps.
Schizodelphis.
b. Teeth purely conical.
Heterodelphis.
2. Intermaxillary with tip produced far forward in
front of maxillary.
Eurhinodelphis, Argyrocetus.
B. Face relatively short, broad and flat.
MONODONTES.
1. Several teeth present in each jaw, none of them
especially enlarged.
Delphinapterus.
2. Almost toothless, a single tooth in the upper jaw of
males a gigantic ramming-tooth.
Monodon.

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

II. Atlas and axis coalesced. :
A. Anterior facial region, formed mostly of intermaxillary
and maxillary, long and narrow, not or scarcely flat,
nearly roof-shaped; intermaxillary in particular rela-
tively narrow.
DELPHINI.
1. Crowns of teeth rough; symphysis menti long.
Steno.
2. Crowns of teeth smooth; symphysis menti short.
a. Palate without grooves.
Prodelphinus.
b. Palate with a longitudinal groove on each side.
Delphinus.

B. Anterior facial region, formed mostly of intermaxillary
and maxillary, becoming relatively short, broad and
flat ; intermaxillary in particular broad.

1. Face relatively only a little shortened.
LAGENORHYNCHI.
a. Fore-face, beak, relatively long.
a. Rostrum not wholly flattened. — Dorsal
fin present.
Tursiops.
8B. Rostrum more flattened. Dorsal fin
absent.
Tursio | Lissodelphis}.
b. Fore-face relatively shorter.
Lagenorhynchus.
2. Face more strongly shortened.
a. Crowns of teeth conical, terete, pointed.
GLOBICIPITES.
a. Intermaxillary not especially broad propor-
tionally.
Orca [Orcinus |.
8. Intermaxillary more or less noticeably
broad. .
1. Flippers not or scarcely lengthened
and pointed.
Orcella.
2. Flippers lengthened, pointed.
a. Intermaxillary not conspicuously
broad anteriorly.
“ Grampus.”

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 37

8. Intermaxillary strongly broad-
ened anteriorly.

(1) Nostril not pushed espe-
cially far back. Teeth
not atrophied. Flip-
pers quite short.

Pseudorca.
(2) Nostril pushed unusually
’ far back. Teeth some-

what atrophied. Flip-
pers very long.
Globiceps| Globicephala].
b. Crowns of teeth in part compressed, widened out
leaf-wise.
PHOCEN.
a. Face relatively long and narrow. Brain-
case relatively small.
Phocena.
B. Face relatively short and broad. Brain-
case large.
Neomeris | Neophocena}.

Physeteride.—The Physeteride probably originated from the
most primitive Delphinids, from Delphinids in which the margin of
the facial depression must have been so widened that it covered the
temporal fossa, but which at the same time retained these primitive
features: zygomatic process of the squamosal relatively large and
somewhat arcuate; teeth small and conical, but still with traces of
- notching on the margin of the crown; teeth in the intermaxillary well
developed ; anterior part of mesethmoid free, not covered over by the
intermaxillaries; free though stunted lacrimal; separate cervical
vertebre ; rather short transverse processes on the thoracic vertebree ;
a well-developed capitulum on all the ribs, etc. The character that
already places even the most primitive Physeterids on a higher plane
than the Delphinids is a result of stronger action of the facial cushion.
It appears as if the Physeteride had from the very first trained them-
selves to swifter, more violent swimming than other whales, and that
the fat-pad in front of the nose had therefore been pressed in with
greater force against the facial part of the skull. The pad, together
with the nasal muscles, etc., has modified the face to an unusual
degree. Especially the posterior margin of the facial depression is
transformed, more or less conspicuously elevated. The crookedness

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL..72

in the posterior nasal passage and in the facial bones becomes more
conspicuous than in other whales. The resistance of the water has
acted upon the skull in other ways also, different in the different
groups; there is a tendency toward strengthening and coalescence of
the bones of the face, toward the appearance of projecting osseous
protuberances, etc.

The members of the family, so far as they are known in this
respect, show a peculiarity in the relation between the ribs and the
transverse processes on the posterior thoracic vertebre, in which they
form a contrast to at least the living forms of Delphinids. While in
the Delphinids the hinder ribs apparently lose the capitulum and
retain the tuberculum (the most posterior, probably having their own
history, have never had more than a single head), in the Physeterids
it is the tuberculum that disappears, while the capitulum remains.
On one or two of the hindmost ribs it may happen that the capitulum
and tuberculum can be seen at the same time, each in contact with its
“transverse process”’; but the tuberculum with its corresponding
process, a diapophysis, is in course of atrophy.”

The genera of the section Xiphiini stand lowest. In them the
occipital wall, which forms the posterior margin of the facial depres-
sion, is highly elevated in a section at the middle only, behind the nares,
and is not pushed very far back in relation to the nares. In the con-
trasted Physeterini the occipital wall is heightened through its whole
extent and more pushed backward. Likewise a primitive feature of
those Xiphiines that are known in this respect is that a more or less
distinct lacrimal bone is present, though in an atrophied condition,
spreading out especially in the roof of the orbit.

The essentially most primitive genus of the Xiphiines is no doubt
the Tertiary South American Argyrodelphis (Notocetus, Diochoti-
chus), of the group Argyrodelphini, not known from much else than
the skull. It stands lower than all other known Physeterids in having
a relatively robust and arcuate zygomatic process of the squamosal
and in the character of the dentition. There is a long row of small,
well-developed, conical teeth in both upper and lower jaw, some of
them bearing notches on the margin of the crown. On the contrary,
as compared with one or another of the other genera, it is advanced
in having the occipital wall pushed rather far back, in having a rather
large cushion-shaped outgrowth on the maxillary above the orbit, and
in having the intermaxillaries spread inward over the mesethmoid
and coming into mutual contact with age. The cervical vertebre
were free.

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 39

In all other Xiphiines the zygomatic process of the squamosal is
smaller and more atrophied, the teeth in the upper jaw disappear,
while of those in the lower jaw one or two only remain in each ramus
and these are particularly modified. A peculiarity of at least the
living forms of the section is the unusual size of the air-sac at the
outer side of the pterygoid, which is shaped to fit it. Somewhat of a
peculiarity likewise is partly the height of the spinous processes on
the dorsal and caudal vertebre (while the transverse processes are
relatively rather short), and partly the small size of the hand. The
former character indicates unusually heavy dorsal and caudal muscles,
or perhaps a widening out of the muscles in a different direction—
more upward than sideways—than in the Delphinids with specially
long transverse processes. The latter makes it appear that the hand
is somewhat disused. The cervical vertebre are inclined to coalesce
as in the Physeterines.

In the members of the group Xiphii, as contrasted with the
Hyperoodontes, the bones of the face have remained primitive to the
extent that no raised longitudinal crest is present on the maxillary in
front of and above the orbit ; at most there is found in the correspond-
ing place a weak cushion-shaped elevation. But in other ways the
bones of the face have increased in strength and have permitted them-
selves to be moulded by the pressure of the water.

In a few respects Mesoplodon is the most primitive among the
Xiphi. To be sure, the facial cushion, by pressing back against the
occipital wall, has caused the median part of the wall, formed for the
most part of projecting outgrowths of the intermaxillaries, to be
abruptly elevated. But*the cushion has not acted on the wall to such
a degree that the upper margin together with the nasal bones has been
either very strongly forced back or caused to bend forward in any
noteworthy manner; neither has it formed for itself any distinct pit
around the nares. Moreover the intermaxillaries have retained their
original relation to the mesethmoid, which they do not grow over.
On the other hand the rostrum acquires increased strength by the
ossification, as age advances, of the mesethmoid, and its coalescence
with the surrounding bones into a stony-hard mass. Of the teeth in
the upper jaw there is found in Mesoplodon at most a series of quite
small remnants, more or less hidden in the skin and scarcely leaving
any traces in the bones. In the lower jaw there is found only one
well-developed tooth. This is situated at the front of the mandible
and is peculiarly modified, having a large, compressed crown and a
big root which sometimes, presumably in the male especially, may

AO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

grow to a disproportionate size and to a high degree affect the form
of the jaw.

The Tertiary European Xiphirostrum (“ Ziphirostrum,’ “ Miozi-
phius”), which is known from parts of the skull, has in a single
respect gone further than Mesoplodon, near to which it otherwise
stands: the intermaxillaries have grown over the mesethmoid and
come in contact with each other along their upper margins. But
Xiphirostrum must have taken its origin from whales that were less
far advanced than the known species of Mesoplodon; as the mes-
ethmoid is not ossified anteriorly; the teeth in the upper jaw are
slightly less atrophied, leaving traces behind them in the maxillary ;
and at the front of the lower jaw there are two well-developed teeth
on each side.

Chonoxiphius (“ Chonesiphius’’), likewise Tertiary European and
known from parts of the skull, stands near to Xiphirostrum. It has
gone further in the modification of the face. The facial cushion has
begun to modify a special area around the nares for its bed. Here
the lateral margins of the premaxillaries are caused to grow slightly
upward, so that they together, and the bones that lie between, form
a special pit, a structure the first traits of which, more or less evident,
are found in many other toothed cetaceans. In the middle of the pit
there has arisen an erect longitudinal crest, evidently formed from
the posterior part of the mesethmoid. (The under jaw is probably
not known. )

Xiphius (“Ziphius”) appears to have originated from whales
which stood on about the same level as Mesoplodon. Its deviations
are of two principal kinds: (1) the median part of the occipital wall
is forced further back and raised higher upward, so thatthe nasal
bones, which are even more modified than in other Xiphiines and are
widened out plate-wise in front, once more come to form a forward-
bent roof over the nasal cavity, and (2) the lateral margins of the
intermaxillaries have grown upward as in Chonoxiphius, but much
more conspicuously, bounding a deep pit. As in Mesoplodon the
anterior part of the mesethmoid becomes ossified with age.

In the genera of the group Hyperoodontes, which must have
originated from the lowest Xiphii, the bony crest, a faint indication
of which is found in many toothed cetaceans running along the upper
surface of the maxillary in front of and over the orbit, becomes so
stimulated to growth by resistance of the water that it gradually
swells up to a huge hump which spreads itself over most of the face
in front of the nares. Each hump is closely appressed to its fellow

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE Al

of the opposite side, leaning inward over the intermaxillary and
mesethmoid so as to turn a broad shock-receiving surface forward.
The nares and middle part of the occipital wall are forced unusually
far backward toward the posterior boundary of the braincase.

The most primitive genus of the group is “ Berardius.’ The
longitudinal crest on the upper surface of the maxillary is well
developed and erect, but the face, however, has retained essentially
its ordinary form. At the front of the lower jaw are found two
rather well-developed teeth.

In Hyperoodon the face becomes with age wholly abnormal, since
the longitudinal crest on the maxillary elevates itself to a height
which exceeds even the highest part of the occipital wall. In the
lower jaw at the front is found only a single tooth (apart from
embryonic conditions).

The genera of the section Physeterini must have originated among
the most primitive Xiphiines, from Xiphiines in which the dentition
was still rather well developed, with teeth in the intermaxillary as
well as the maxillary; in which the intermaxillaries were free from
each other and from the surrounding bones ; in which the mesethmoid
was not ossified, etc. Their peculiarity is that the pressure of the
facial cushion on its surroundings acts differently than in the
Xiphiines, and even more strongly. The facial cushion, especially
that part of it which is formed by the adipose mass, widens out still
more, particularly outside of and behind the nasal passages. It
pushes the median part of the occipital wall far back behind the
nares, while at the same time the margins of the facial depression, at
the back and at the sides, grow high upward. The bones which form
the bottom of the facial cushion’s bed are strongly acted upon by the
stimulating mass of the cushion. They widen out. This holds good
especially of the bones in the rostrum, and chiefly of the anterior
part of the maxillary. The lower jaw on the contrary retains its
primitive narrowness.

The genera of the group Hoploceti are extinct, Tertiary, and are
only incompletely known, chiefly from fragments of skulls. It may
be concluded that the skull in essential respects is modified in the
same manner as in the highest group of Physeterids, the Physeteres,
but to a distinctly less noticeable degree. They are also less advanced
than the Physeteres in the development of the dentition, there being
a long row of well-developed teeth in both upper and lower jaws,
while the upper teeth of the Physeteres have atrophied.

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

In the European and American Hoplocetus (Balenodon, Physodon,
“ Scaldicetus,”’ etc.) the teeth are covered with enamel.

The European Physeterula has lost the enamel, it appears.

In the genera of the group Physeteres the influence of the facial
cushion on the skull has led to fantastic results. The dentition also
is modified in a strange manner. In the lower jaw the teeth continue
to be well developed; but in the upper jaw they atrophy and dis-
appear either wholly or essentially so. It is not clear what the reason
can be for this difference between the upper and lower jaws. Per-
haps the difference is connected with the great lateral broadening
out of the maxillary whereby the upper toothrows are so pushed
outward that they lose their interaction with the lower toothrows
which retain their ancestral position in the closely appressed mandi-
bular rami. There has been no hard work for the upper teeth which
might have maintained them in spite of all; the lower teeth together
with the palate must have proved sufficient, as the task is indeed
scarcely anything else than to grasp the cuttlefish which appear to be
the favorite food for this whole family as well as for various other
cetaceans, especially for those with more or less degenerate dentition.

“ Cogia”’ is in some respects the most primitive of the genera. In
the upper jaw there is still found, or may be found, a tolerably well-
developed tooth. In the skull there remains a rather considerable,
curved remnant (although very narrow and compressed) of the
heightened osseous ridge which elsewhere in the toothed whales lies
between the nares and the occipital wall; the adipose cushion has not
yet wholly destroyed it. Neither has the cushion so grown around
the outer nasal passage that it has pushed the orifice away from its
accustomed place, nor has it to any noticeable degree pressed the roof
of the braincase down. The anterior part of the face has become
broader than usual, but it is still rather short, or, more strictly
speaking, has been further shortened. On the other hand the lateral
margin of the facial depression, over the braincase, is pushed unus-
ually far out to the side and raised conspicuously high upward; it
has also acquired a unique thickness.

In Physeter nothing has remained of the upper teeth except small
vestiges hidden in the skin. The fat-cushion, which has grown
gigantically, has caused the bones of the rostrum to grow far forward
and to broaden themselves strongly at the side. The posterior margin
of the facial depression is more abruptly elevated than in any other
whale and is pushed further back. The fat-cushion has completely
overgrown and leveled off the bony wall which elsewhere lies be-

no. 8 INTERRELATIONSHIPS OF THE CETACEA—-WINGE 43

tween the nares and the occipital wall. The soft outer nasal passage is
pushed forward so that the blow-hole lies far to the front. The
adipose cushion has forced the roof of the braincase down by its
weight, and the whole braincase has sunk down below its original
level, so that the spinal marrow, or medulla oblongata, has to bend
down in an S-shaped curve to connect with the brain. The skull has
acquired a noticeably large size in proportion to the body, and most
of its bones have become conspicuously ponderous ; this is especially
noticeable as concerns the zygoma. Just as Physeter is the largest
of all the toothed whales and pushes through the water with greater
force than any other, it is the one on which the resistance of the
water has had the most powerful influence. But it is a question
whether this high development is not a menace to the creature’s life.
Irresistibly the water’s pressure has caused the facial cushion to
grow to a disproportionate degree and in its turn to call forth a skull
the size of which is without relation to brain and body. The fate of
Physeter, the most highly developed toothed whale, is much like that
of Balena, the strangest whale-bone whale; the difference is that the
pressure of the water in the one has acted most strongly on the outer
side of the head, in the other most strongly on the inner walls of the
mouth. Both animals are developed with such extravagant one-
sidedness that they appear to be in danger of certain extinction even
if their extirpation were not being worked at by man.

Physeteridz.”
I. Occipital wall highly elevated in middle only, its position close
behind the nares.
XIPHIINI.

A. Dentition primitive: a long row of rather uniform
small teeth in both upper and lower jaw.
Zyvgomatic process of the squamosal well
developed.

ARGYRODELPHINI.
Argyrodelphis.

B. Dentition atrophied: most of the teeth disappear,
leaving one or two in each lower jaw specially
modified. Zygomatic process of the squamosal
somewhat reduced. .

1. Longitudinal crest on maxillary above and in front
of orbit absent or slight.

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

XIPHII. t

a. Intermaxillaries not or scarcely forming a
cup-shaped depression around nares.

a. Intermaxillaries not covering mes-
ethmoid in front.
Mesoplodon.
8. Intermaxillaries mutually in contact,
covering mesethmoid in front.
(1) No pit formation around nares.
Xiphirostrum.
(2) Indication of pit formation
around nares.
Chonoxiphius.

b. Intermaxillaries with an elevated outer
margin forming a deep cup around
nares.

Xiphius.
2. Longitudinal crest on maxillary above and in
front of orbit well developed, swollen.

HyPEROODONTES.

a. Longitudinal crest on maxillary relatively
weak.

“ Berardius.”
8. Longitudinal crest on maxillary huge.
Hyperoodon.
II. Occipital wall spreading itself, highly elevated, across entire
braincase, and pushed far back behind nares.
PHYSETERINI.
A. Upper toothrow well developed.
HopPLoceti.
1. Teeth with enamel.
Hoplocetus.
2. Teeth without enamel.
Physeterula.
B. Upper toothrow atrophied.
PHYSETERES.

1. Distinct remains of the longitudinal crest
that originally extended from the nares
to the occipital crest.

Coo:

2. Longitudinal crest behind nares _ wholly

flattened out, obliterated. ;
Physeter.

no. 8 INTERRELATIONSHIPS OF THE CETACEA—-WINGE 4

on

The Hyznodonts, the nearest stock-forms of the cetacea among
terrestrial mammals, lived at the beginning of Tertiary times in the
northern parts of both the Old and New Worlds. They had spread
over Europe and North America and were found in northern Africa
as well. The whales must have made their appearance somewhere
within the territory occupied by the Hyzenodonts, and probably in
the oldest part of the Tertiary; in agreement with this the most
primitive cetacean that is yet known, the Hyznodont-like Protocetus
of the family Zeuglodontidz, is found in Egypt in Eocene strata.
Likewise one of the next links in the chain of cetacean development,
Prozeuglodon, was Egyptian, from the Eocene. But soon the mem-
bers of the family must have spread widely ; in any event the highest
genus, the almost fantastic, snake-like Zeuglodon |Basilosaurus},
appears to have found its way during the Eocene to all oceans.

The Zeuglodonts died out early in the Tertiary. Their highest
forms left no descendants; but from the more primitive genera of
the family sprang the new family Balznide. The oldest, tooth-
bearing forms of Balznids are as yet scarcely known. In Miocene
times, however, the family had already produced the specialized
whalebone-bearing forms, a side branch on the cetacean genealogical
tree, and they soon spread themselves to all the seas of the globe,
where they still are found. Some of the recent genera are essentially
cosmopolitan, even in the sense that individual species occur in all
seas. This holds good in part only of Balena, one of whose species,
the more primitive, B. australis, is almost cosmopolitan, while the
second, the more specialized, B. mysticetus, is confined to the northern
polar oceans. It is literally true of Balenoptera and Megaptera.
_ Two of the recent genera are confined to a smaller range: Neobalena
a relatively high genus that lives in the South Sea, where it likely
originated, and Rhachionectes a relatively low genus, in many respects
recalling extinct Miocene forms. It lives in the northern part of the
Pacific, perhaps as a kind of last remnant from an early day. The
reason why the Balzenids, in spite of their rather primitive structure,
are not wholly extinct, supplanted by the more specialized cetacea,
is probably because they have chosen a peculiar food supply: the
small creatures of the sea. Therefore they do not have very many
competitors among their kind.

From the most primitive, tooth-bearing Balznids the family
Squalodontidze branched off in Tertiary times. It had its flourish-
ing period in the Miocene, widely distributed in the oceans. The
whole family disappeared before the end of the Tertiary, chiefly,
it would appear, because it passed onward into its successors.

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL 72

The descendants of the primitive Squalodonts are the members of
the family Platanistide which appeared early in the Tertiary and
was soon widely distributed. Most of the genera have died out
again; only four, Pontoporia [Stenodelphis|, Lipotes, Inia and Pla-
tanista, have come down to the present time. These have doubtless
avoided being crowded out by higher cetaceans purely because they
have chosen a peculiar habitat, rivers and estuaries, which they have
been almost alone in utilizing.

Early in the Tertiary the family Delphinidz branched off from
primitive Platanistids. Extinct genera, especially in the Miocene,
are known from localities that were even then far apart; at present
the family is universally distributed, many of the genera and species
being nearly cosmopolitan. The family seems to be having its
flourishing period now. Only a few of the recent genera have
ranges that are somewhat restricted, as Delphinapterus and Monodon
in the Arctic Ocean, Tursio [ Lissodelphis| in the Pacific, Orcella in
the rivers of Southeast Asia and in the neighboring sea, Neomeris
| Neophocena] on the eastern and southern coast of Asia and the east
coast of Africa.

Early in the Tertiary the family Physeteridz originated from the
most primitive Delphinids. It had already reached its climax in the
Pliocene, widely distributed. Only rather few genera, but these very
highly developed, have come down to the present day. They are
widely distributed, essentially cosmopolitan.

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 47

NOTES

*(P. 1.) The present treatise on Cetacea is a continuation of the
series of memoirs on the other orders of mammals which have
appeared in E Museo Lundii, vols. 1-3, 1887-1915, and in the Vidensk.
Medd. Dansk Naturhist. Foren., vol. 68, 1917. Part of the opinions
that are here expressed have been previously published in the Vidensk.
Medd. Dansk Naturhist. Foren. for 1882, pp. 29-31, 40 and 53-55;
tbid., for 1909, pp. 5-9; Meddelelser om Gr¢gnland, pt. 21, 1902,
pp. 364-368 ; Danmarks Fauna, Pattedyr, 1908, pp. 9-10, 200-209.

*(P. 1.) On the origin of the Cetacea very different opinions
have been put forward. The idea of Brandt and others that the
Cetacea are the lowest, most reptilian mammals is now shared by
scarcely any one. Likewise the old idea of the relationship with
sea-cows was long ago laid aside. Flower’s early opinion that the
whales originated from seals, an opinion which he shared with others,
was disputed by Winge (Vidensk. Medd., 1882, pp. 53-55) and
almost abandoned by Flower himself. It was not taken up by others
except in a way by D’Arcy Thompson. D’Arcy Thompson’s opinion
(On the Systematic Position of Zeuglodon; Studies from the
Museum of Zoology in University College, Dundee, vol. 9, 1890,
pp. 1-8, with illustrations) that the Zeuglodonts, really the most
primitive whales, are not Cetacea, but near relatives of the seals, is
disproved by Lydekker (Proc. Zool. Soc. London, 1892, pp. 560-
561) and Dames (Ueber Zeuglodonten aus Aegypten, etc. ; Palaeont.
- Abhandl., herausgeg. von Dames u. Kayser, vol. 5, pt. 5, 1894, section
pp. 204-210). Flower’s ideas about the whale’s relationship to par-
ticular ungulates proper other than sea-cows have also shown them-
selves to be incorrect. The author who has most extensively ex-
amined the question in earlier times is Max Weber in his book:
Studien uber Saugethiere, ein Beitrag zur Frage nach dem Ursprung
der Cetaceen, 1886, which contains a review of earlier work on the
subject. His own conception of the history of the Cetacea was then
“dass sie einem generalisirten Saugethiertypus im mesozoischem
Zeitalter entstammen, der zwischen Carnivora und Ungulata mitten
inne steht, wohl aber nahere Beziehungen zu Carnivora hatte” (J. c.,
p. 241). In his work Die Saugethiere, 1904 (p. 581), Max Weber
sets forth the idea that “ primitive Condylarthrer ”’ were perhaps most

nearly the precursors of the whales.

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Solid ground was first reached with the discovery of Protocetus
atavus described by Fraas (Neue Zeuglodonten aus dem unteren
Mitteleocan von Mokattam bei Cairo; Geol. u. Paleont. Abhandl.,
herausgeg. von Koken, vol. 10, pt. 3, 1904. There could be no doubt
that Protocetus came from Hyznodonts and that it was itself a fore-
runner of the Zeuglodonts. Curiously, however, Fraas thought that
the origin of the Cetacea was not thereby explained. He considered
both Protocetus and with it the other Zeuglodonts as a side branch
from the carnivores which did not lead in the direction of the true
whales. There seems now, however, to be unanimity of opinion that
Protocetus, Prozeuglodon, etc., are some of the long sought pro-
genitors of the whales. In spite of all differences from the higher
Cetacea there is a multitude of resemblances to them which it would
be impossible to explain except on the basis of relationship. One has
only to think of the striking likeness in such peculiarly formed bones
as the tympanic and scapula ; their characters in the fossils are exactly
those that one would expect to find in ancestral Cetacea.

It has been said that the whalebone whales and the toothed whales
might have separate “‘ diphyletic”’ origins; Kukenthal in particular
has spoken for this view (Ueber die Anpassung von Saugethieren an
das Leben im Wasser ; Zool. Jahrbucher, Abth. fiir Systematik, etc.,
vol. 5, 1891, pp. 373-399, especially p. 384, and elsewhere). In face
of the host of agreements in numerous structural relationships which
are found in the two groups this idea is an impossibility. Just one
little bone like the tympanic, with its thickened inner wall, its mussel-
shaped outgrowth around the outer auditory aperture, its petrous
process which reaches out under the mastoid, and other details, all of
the most peculiar form, and all essentially identical in all Cetacea, is
sufficient evidence of the near relationship of all whales.

Kukenthal has put forward a “‘ Versuch, den Bau des Walkorpers

von biologischen Gesichtspunkten aus zu erklaren,” most elaborately
in Die Wale der Arktis, Fauna Arctica, vol. 1, pt. 2, 1900, section
pp. 181-203.
* (P. 3.) We owe to Abel a special treatise on the skeleton of the
hind limb in Cetacea: Die Morphologie der Hiiftbeinrudimente der
Cetaceen; Denkschr. d. math. -naturw. Klasse d. k. Akad. d:
Wissensch. Wien, vol. 81, 1907, pp. 139-195, with illustrations. A
supplement is given by Lonnberg: The Pelvic Bones of Some
Cetacea; Arkiv for Zoologi, vol. 7, No. 10, 1910, pp. 1-15, with
illustrations.

No. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 49

*(P. 4.) Much information about the structure of the hand in
the Cetacea has been collected by Kukenthal (Die Hand der Cetaceen ;
Denkschr. d. med. naturw. Ges. zu Jena, vol. 3, pt. 1, 1889, pp. 23-69,
pl. 3, and the section “ Die Brustflosse,” in Vergl. -anat. u. entwicke-
lungsgesch. Unters. an Walthieren, ibid., pt. 2, 1893, pp. 267-312,
with illustrations) and Kunze (Ueber die Brustflosse der Wale;
Zoologische Jahrbiicher, Abt. fur Anatomie, etc., vol. 32, pt. 4, 1912,
pp. 577-651, pls. 33-35), both of whom give references to earlier
works.

Kiukenthal regards it as most probable that the large number of
phalanges in the Cetacea has originated as follows: That the diaphy-
ses and epiphyses in an ordinary hand whose fingers had mostly three
phalanges have loosened themselves from each other, and have be-
come independent and uniform, all of them ossified. This explana-
tion cannot possibly be right. It is immediately contradicted by the
fact that in cetacean hands with many-jointed fingers there can be
found both diaphyses and epiphyses, ossified, in the larger of the
phalanges that are present, as Kukenthal himself has observed. If
one examines series of adult cetaceans’ hands or of embryo hands, it
is quite impossible to detect anything that could point in this direction.
It certainly should be possible to find, somewhere or other, transition
forms which would show indication that the phalanges were of unlike
origin, some of them diaphyses, others epiphyses ; but of this there is
not the slightest evidence. Neither is it probable that the forerunners
of the whales among terrestrial animals, had, even when young,
epiphyses at both ends of all the phalanges, as would be needed in
order to explain even tolerably the large number of joints in the

-Cetacea. It is true that in the Cetacea there have arisen super-
numerary ossified epiphyses, more epiphyses than in their ancestors.
But let it be noted that this has only happened in those Cetacea that
already had acquired many-jointed fingers. (The objection to the
“epiphysis-hypothesis ”’ that it could at most explain the presence of
only I2 joints in the fingers, including the metacarpal, and that it
therefore cannot hold good where the number of joints is more than
12, is met by Kitkenthal with the admission that in such instances the
number of joints is actually increased out beyond the finger tips,
bcs, 1803, p. 311.)

Another explanation which is more probable Ktkenthal himself
sets forth but regards as less happy: ‘“‘ Wurde man die Entwicke-
lungsgeschichte allein zur Losung der Frage heranziehen, so wurde
sich der Schluss ergeben, dass ausser den vier typischen Finger-

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.: 72

elementen sich immer neue anlegen, indem sie sich, in distaler Richtung
aufeinander folgend, in dem sich immer weiter vorschiebenden em-
bryonalen Bindegewebe entwickeln, so dass die letzte Phalanx die
jungste ist” (J. c., 1893, p. 311). Ktikenthal finds a reason to reject
this opinion in the fact that Leboucq and he have occasionally found
something on the outermost tip of one of the digits of a long-fingered
porpoise hand which might be interpreted as the weakest remnant of
a nail. Should it prove that remnants of a nail are found on the
extreme tip of the finger, says Kukenthal, this “ hypothesis ” con-
cerning the origin of the many phalanges cannot be correct, “ denn
dann entspricht die Spitze der Walflosse und damit die Spitze von
deren Fingern auch der Spitze der Finger der typischen Vorder-
extremitat ” (1. c., 1893, p. 312). This objection cannot hold; there
is certainly nothing to prevent that the atrophied remnant of a nail
should constantly retain its position on the finger tip as this pushes
outward further and further, whatever the method by which the
finger is elongated.

°(P. 5.) A special treatise on the cervical vertebrz of the Cetacea
is due to Reche: Ueber Form und Funktion der Halswirbelsaule der
Wale; Inaugural-Diss., 1904, with illustrations. See also De Burlet:
Beitrag zur Entwickelungsgeschichte der Wirbelsaule der Cetaceen ;
Morphol. Jahrb., vol. 50, pt. 3, 1917, pp. 373-402, with illustrations.

*(P. 7.) On the tympanic bone and its surroundings in the
Cetacea, see especially: Van Kampen, De Tympanaalstreek van den
Zoogdierschedel, 1904, pp. 299-316. Contains references: to earlier
papers on the subject. On the ear-bones themselves, see especially :
Doran, Morphology of the Mammalian Ossicula Auditus; Trans.
Linn. Soc. London, ser. 2, Zoology, vol. 1, 1878, pp. 450-464,
pls. 62, 63.

"(P. 9.) The opinion that the increase in the number of teeth in
the Cetacea above the typical eutherian number was perhaps initiated
by the intercalation of milk teeth in the permanent set was expressed
in 1882 (Vidensk. Medd. for 1882, pp. 31 and 40) at a time when no
trace of tooth succession had yet been detected in whales. The same
opinion was maintained by Max Weber (Urspr. der Cetaceen, 1886,
pp. 195 and 199), but he abandoned it (Die Saugethiere, 1904, p. 567)
after Kukenthal had demonstrated indications of the tooth succession.
Ktikenthal had found traces of germs of both forerunners and suc-
cessors to the teeth which stand in the Cetacea as the permanent set
(but which he considered as milk teeth). Perhaps the idea is wrong ;
but there is nothing in that which has thus far been discovered which

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE sul

makes its abandonment necessary, inasmuch as only a part of the
milk dentition, only four cheekteeth, were ever supposed to be inter-
calated in the permanent set (it was expressly said that an increase in
the number of cheekteeth by other means, by division or by the
formation of new elements, was necessary as an explanation when
the number exceeded 11 in each jaw) ; and the milk dentition could
perhaps be made whole again after having given up some of its
contents. In any event it is impossible to trace in detail the destiny
of the tooth-germs through their erratic course during atrophy ; there
are many possibilities.

In spite of all investigations during recent years into the develop-
ment of cetacean dentition both embryologically and phylogenetically,
no certain conception has been reached. Only this is certain, that the
cetacean dentition is derived from the typical carnivore’s, that the
teeth are increased in number while their size is decreased and their
form made more simple, and that the tooth succession has essentially
ceased, although in embryos there can still be detected traces of
several sets of teeth, as in many other mammals, faint mementos of
their forefathers among the reptiles. But of how the changes have
in detail come about we can only partly guess.

Abel is the author who has most recently reviewed the present
question. He believes that he can more nearly show how the high
number of teeth has arisen in the Cetacea; in the whalebone whales
he thinks it came about in one way, and in the toothed whales, or at
least in the Physeterine series, in another. But his arguments are not
irrefutable.

As regards the whalebone whales Abel starts from Kukenthal’s
' investigations. As has long been known from observations by
Geoffroy Saint-Hillaire and especially by Eschricht, there is found
in embryos of all recent whalebone whales, hidden in each jaw, a long
row of small atrophied teeth with conical or knob-shaped crowns,
which are resorbed without ever erupting. Frequently some of these
small teeth are seen to be mutually united; most often it is two that
come together but in rare instances as many. as four may unite. A
part of his observations on the embryonic teeth of Balenoptera
musculus, the species which he has had especially good opportunity
to investigate, Kiikenthal summarizes in the following words: “ Die
Zahl der Zahne im Oberkiefer des letzteren Embryos (that is, the
largest of those examined) ist 53; sie liegen sammtlich in gleich
weiten Abstanden von einander. Bei den kleineren Embryonen
betragt die Zahl der Oberkieferzahne, wenn wir die mit zwei resp.

4

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

drei Hockern versehenen Zahne als je einen Zahn rechnen, etwas
weniger, und zwar haben alsdann die kleinsten Embryonen die
geringste Zahnzahl. Zahlen wir jedoch in jedem Oberkiefer die
einzelnen Hocker, so erhalten wir gleichzeitig fur jeden Kiefer die
Zahl 53, dieselbe Zahl, welche wir auch bei dem grossten untersuchten
Embryo, der keine Doppelzahne besass, aufgefunden haben. Ich
habe aus dieser Thatsache bereits den Schluss gezogen, dass sich im
Laufe der individuellen Entwickelung der Bartenwale die Zahne
theilen, und dass somit aus den verhaltnismassig wenigen, aber mehr-
hockerigen Zahnen der jiingsten Stadien viele, aber einspitzige
Zahne werden. Aus den ursprtinglichen Backzahnen entwickelte
sich also durch Theilung derselben ein anscheinend homodontes
Gebiss. (Unters. an Walthieren ; Denkschr. med. naturw. Ges. Jena,
1893, p. 431). On this Abel builds further. He considers that
Patriocetus (see pp. 70-72), a Tertiary whale with rather well-
developed dentition, with unicuspid incisors and canines, and with
serrate margins to the cheekteeth, is an ancestral form of the true
whalebone whales—their immediate precursor. In passing to the
whalebone whales its teeth would be split up and the marginal cusps
would be transformed into independent teeth: ‘ Wie wir gesehen
haben, besteht das Gebiss von Patriocetus ehrlichi aus sieben zwei-
wurzligen und siebenspitzigen Backenzaihnen, von denen die drei
hinteren als Molaren und die vier vorderen als Praemolaren zu deuten
sind. Daran schliessen sich-vorne ein einspitziger Eckzahn und die
drei einspitzigen Schneidezahne an. Im ganzen stehen also I1
Zahne in jedem Kiefer.—Wenn wir die Spitzen der Zahne susammen-
zahlen, so dass wir nicht nur die Kronenspitzen der vier vorderen
Zahne, sondern auch die sieben Zacken der sieben zweiwurzligen
Backenzahne als Einzelspitzen rechnen, so ergibt sich eine Gesamt-
summe von 53 Spitzen, also genau derselben Zahl, die wir bei dem in
Einzelzahne aufgelosten Gebiss des Finwalembryos wiederfinden.—
Nach diesem Befunde kann es keinem Zweifel mehr unterliegen,
dass das Patriocetus-Gebiss mit elf Zahnindividuen und zusammen
53 Schmelzspitzen den Ausgangspunkt des Bartenwalgebisses dar-
stellt und dass die Entstehung des letzteren in der Weise erfolgt,
dass die elf Zahne sich im Verlaufe der ontogenetischen Entwick-
lung in 53 Teile spalten, so dass also schliesslich aus einem sieben-
spitzigen Backzahn sieben einzelne Spitzen durch Teilung und fort-
schreitenden Zerfall hervorgehen. (Die Vorfahren der Bartenwale ;
Denkschr. Akad. Wissensch. Wien, mathem. naturw. K1., Bd. 90,
1914, pp. 186-187)—Several objections must be raised to Abel’s
presentation of the subject.

No. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 53

Everyone who has had to do with counting the teeth of cetaceans
knows how variable the numbers may be. It is almost a miracle that
Kukenthal should have been able to find five or six embryos, or
perhaps more, of Balenoptera musculus, each of which had 53 teeth
or tooth cusps in the upper jaw (see Kutkenthal’s more special account
in Jenaische Zeitschrift fiir Naturwissenschaft, vol. 26, 1892, p. 481,
together with his paper of 1893, /. c.; in three other jaws the numbers
were less, a circumstance said to be accounted for by the fact that
some of the teeth had been resorbed). But even if it should prove
that B. musculus always had 53 teeth in each jaw of the embryo this
fact would be without bearing on the question of the original denti-
tion in the whalebone whales. It will occur to nobody to regard
B. musculus as one of the most primitive species of the genus; on the
contrary it is one of the highest, being one of the largest and most
elongated. Other species are found in the genus that stand on a
lower: level; this holds good especially of B. rostrata, and in this
species Eschricht has found the number of teeth in two embryos to

be respectively = + and 2 (Unders. over Hvaldyrene, pt. 3, 1845,

pp. 314 and 316-317). In two embryos of the same species Ktiken-
thal found 41 in the lower jaw (Jen. Zeitschr., 1892, pp. 485-486).
In two embryos of one of the highest species of the genus, B. gigas
(sibbaldii) Kukenthal found 50 in the upper jaw (J. c., p. 486). In
several embryos of Megaptera boops Eschricht has found from 46
to 51 teeth in each side of the upper jaw, and in the lower jaw rather
fewer, the least number 42 (I. c., pp. 311 and 316). Abel says, it is
true: ‘Bei jenen Bartenwalen, deren Kiefer eine geringere Zahl
_ als 53 Zahnindividuen aufweisen, handelt es sich entweder um fruhere
Embryonalstadien, wie bei dem von C. Julin beschriebenen Embryo
von Balenoptera rostrata von etwa 48 cm. Lange (41 Zahne), oder
um Reduktionserscheinungen ” (J. c., p. 188) ; but this assumption is
entirely inadmissible. Abel himself probably had an inkling of it;
he adds: “In dieser Frage miissten noch eingehendere Untersuch-
ungen auf breiterer Grundlage angestellt werden, um unsere bis-
herigen Kenntnisse in dieser Richtung zu erweitern.”

Neither can all of what Abel says about the number of teeth or
cusps in Patriocetus stand before a closer examination. It is not
certain that Patriocetus had 11 teeth in each jaw; none of the skulls
that have been found has entire jaws, the anterior part is lacking in
them all. The number 11 is therefore only a guess, and scarcely
very likely ; judging from the rest of the cranial characters one would

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

expect a departure from the typical dentition—more teeth. Of the
cheekteeth there are only known a few that are tolerably complete ;
that every one of them had seven marginal cusps is mere conjecture
and not probable; in other cetacea with serrate teeth, both Zeuglo-
donts and Squalodonts, the number of cusps varies strongly; no
cetacean is known in which the same number of cusps is found on all
the cheekteeth. Some dissociated teeth that probably are correctly
referred to Patriocetus (Squalodon ehrlichiu), figured by Suess
(Neue Reste von Squalodon aus Linz; Jahrb. d. k. k. geol. Reich-
sanstalt, vol. 18, 1868, pp. 287-290, pl. 10, figs. 1-3) and reproduced
by Brandt (Unters. foss. u. subfoss. Cetaceen Europa’s ; Mém. Acad.
Imp. Sci. St. Pétersbourg, ser. 7, vol. 20;° No. 1, 1873, plieaueeneae
T1-13), also show other numbers, one of them 9, another 10. More-
over it is doubtful whether Patriocetus can be regarded as an ancestor
of the whalebone whales; it is not precluded that with more exact
acquaintance it will show itself to stand on a higher level, nearer to
the typical Squalodonts, closely connected with Agorophius (see
note, p. 72). In short the whole calculation about the 53 teeth in the
whalebone whales and the 53-tooth cusps in Patriocetus rests on the
weakest foundation.

It is also a question whether Kitkenthal and Abel are on the whole
right in their conception of the many small teeth of the whalebone
whales as having originated by the division of fewer, larger, serrate
teeth. There is indeed scarcely any doubt that a division of the tooth
germ might be able to take place at an early stage of a tooth’s develop-
ment; but that a tooth which had already acquired serrate margins
should be able to divide is not probable; in the case of the whalebone
whales at any rate there is nothing convincing in this respect—quite
the contrary; and other cases are not known. The “ double-teeth ”
of the embryo whalebone whales are the ones that are conceived to
be serrate teeth in course of division; but they could be better ex-
plained in another manner. Their position in the toothrow is quite
erratic—sometimes far to the front, sometimes in the middle or far
back. The number of cusps is most often two, only in rare instances
as many as four. The cusps have the appearance of being of equal
rank, none can be called the chief cusp. In short, the cusps in the
double teeth appear to be small, atrophied, unicuspid teeth which
have quite casually come near each other and grown together, some-
thing which might be able to take place with special ease in the
youngest stages of the embryo when the tooth germs are crowded
together in relatively short jaws. That double teeth were produced

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 55

by the fusion of single teeth was already supposed by Eschricht
(1. c., p. 312). Should it be the case that there has taken place in
the whalebone whales a splitting up of serrate teeth, the correspond-
ing ancestral forms must probably have had teeth in which the cusps
on the fore aud hind margins of the crown had great independence
and a size very nearly the same as in the principal cusp. In that
event it would not be easy to regard Patriocetus as an ancestor ; since
in it the cusps on the fore and hind margins particularly are weak in
proportion to the main cusp, and apparently in course of atrophy.

Abel gives the following as his conception of the manner in which
the many small teeth of the Odontoceti, or at least of the Physeterids,
have arisen: “ Dieser Spezialisationsweg des Gebisses (in the whale-
bone whales) ist fundamental von jenem verschieden, den wir in der
Phylogenese des Physeteridengebisses finden. Wie ich 1905 gezeigt
habe, tritt auf dem Wege zur Entstehung der Squalodontiden
zunachst eine starke Vermehrung der mehrwurzeligen, vorn und
hinten gezackten Backenzahne ein, so dass sich das primitive Arch-
aeocetengebiss durch Vermehrung der Backenzahne im Pramolaren-
abschnitt zu dem polyodonten Squalodontidengebiss umformt. Aus
den Squalodontiden sind die Physeteriden hervorgegangen, bei
welchen das Gebiss eine Reduktion erfahrt ; dieser Spezialisationsweg
fuhrt aber zu einer Vereinfachung der Krone, Verschmelzung der
bifiden Wurzeln, Reduktion der Zackenreihen am Vorder- und Hin-
terrande der Kronen zu einer krenelierten Leiste und endlich zum
ganzlichen Verlust der Schmelzkappen ” (1. c., p. 187). Here Abel is
no doubt right in the main. It can only be objected that it cannot
exactly be said that Abel in his more special account (Die phylo-
_ genetische Entwicklung des Cetaceengebisses und die systematische
Stellung der Physeteriden ; Verhandl. Deutsch. Zool. Gesellsch., 1905,
pp. 84-96, and Les Odontocétes du Boldérien; Mém. Mus. Roy.
d’Hist. Nat. de Belgique, vol. 3, 1905) has demonstrated that it is
precisely in the premolar region that the number of teeth has been
increased in the Squalodonts; neither is it probable that the Physe-
terids originated directly from the Squalodonts. They appear to
have branched off at a higher level; probably they had their root in
common with the Delphinids.

There is no reason at the present stage for believing that the
increase in the number of teeth beyond the typical formula should
have had a different origin in the Mystacoceti and Odontoceti. In
view of the great resemblances that are everywhere found between
the two groups it is not likely that in this respect there would be a

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

difference. The method by which the increase has come about in the
Odontoceti no doubt holds good for the Mystacoceti as well ; the most
primitive forms of probably both groups had serrate teeth in aug-
mented numbers.

*(P. 10.) It has become usual to believe that the precursors of the
cetaceans were armored mammals with well-developed osseous dermal
plates. Heated support for this idea is brought forward by Kiiken-
thal (especially in the section “ Ueber Rudimente eines Hautpanzers
bei Zahnwalen,” in Vergl. -anat. u. entwickelungsgesch. Unters. an
Walthieren, part 2, Denkschr. d. med. naturw. Ges. zu Jena, vol. 3,
pt. 2, pp. 251-258, pl. 16) and by Abel (especially in the section
“T?armure dermique,”’ in Les Dauphins Longirostres du Boldérien,
Mém. Mus. Roy. d’Hist. Nat. de Belgique, vol. 1, 1901, pp. 17-32,
with illustrations). Kitkenthal has investigated recent cetaceans ;
Abel more particularly the extinct forms. (In Abel is found refer-
ence to previous literature on the subject.) Kiukenthal imagines that
the Cetacea originated from armored land-mammals with armor sug-
gesting that of the Dasypodids, and that as sea dwellers they have
jost the armor more or less completely; Abel thinks, in agreement
with Dollo, that the armature did not occur in the terrestrial pre-
cursors of the cetaceans, but that it arose in the first whales as part
of their adaptation to aquatic life along the coast, and that afterwards
it was lost in the more strictly marine members of the group.

What we have to build upon is the following :

Together with the first lot of Zeuglodon bones found in Alabama
came a few pieces of limestone containing some plate-like, very
irregular bones of various sizes. Accounts of. these bones are due
especially to Joh. Miller (Ueber die fossilen Reste der Zeuglodonten
von Nordamerica, 1849, p. 34, pl. 27, fig. 7), Carus (Das Kopfskelet
des Zeuglodon hydrarchus; Nova Acta Acad. Caes. Leop. Carol.,
vol. 22, pt. 2, 1850, pp. 382-383, pl. 39A, fig. 5), Dames und Jaekel
(section Ueber den Hautpanzer der Zeuglodonten, in Dames, Ueber
Zeuglodonten aus Aegypten, Palaeontol. Abhandl., herausgeg. v.
Dames u. Kayser, vol. 5, pt. 5, 1894, pp. 219-221, with illustration)
and Abel (1go1, l. c., pp. 24-27). From the beginning the possibility
has been thought of that the plates were dermal bones of Zeuglodon.
They have, however, most often been regarded as doubtful; perhaps
they were bones from the carapace of a sea turtle like Psephophorus
or something of the sort; usually no one has dared to say anything
positive. Abel was the first to consider it as proved that they were
dermal bones of Zeuglodon; of one of the specimens in question he

No. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 57

thinks that it can be nothing else than a piece of armor from the fore
angle of a dorsal fin, because the plates are bent toward each other
like a roof, ina manner and form that is not possible on any part of a
turtle’s carapace. As a not unessential ground for believing in the
occurrence of armor in Zeuglodon he reckons the occurrence in the
recent Delphinids Neomeris and Phocena of structures which Ktiken-
thal explains as remnants of armor.

At Radoboj in Croatia some remains have been found of a small
dolphin-like cetacean, Delphinopsis freyerii, established and de-
scribed by Joh. Miller (Bericht titber ein neu entdecktes Cetaceum
aus Radoboy, Delphinopsis Freyerii; Sitzungsber. k. Akad. Wis-
sensch. Wien, math. naturwiss. Cl., vol. 10, 1853, pp. 1-6 of separate),
and again fully discussed and figured by H. v. Meyer (Delphinopsis
Freyerii Mull. aus dem Tertiar-Gebilde von Radoboj in Croatien;
Palaeontographica, vol. 11, 1863, pp. 226-231, pl. 34) whose illustra-
tion is reproduced by Abel (J. c.), who also has personally examined
the remains. It was only imperfect remains that were found, not
much more than pieces of a flipper lying in a slab of stone; around
the bones of the hand lie numerous small disk-shaped bodies a milli-
meter or less in diameter, the underside of which is covered with
minute projecting granules arranged in parallel lines. Joh. Miiller
seems to have left undecided the question whether these bodies were
of organic or inorganic origin, although he leaned mostly to the
opinion that they were osseous scales from the skin. But H. v. Meyer
maintained that they were inorganic. His reason for this opinion
was especially that scattered among them there lie bodies of entirely
similar appearance only without markings, and these bodies are
undoubtedly inorganic. Abel on the contrary is convinced that the
small, striated disks are dermal ossicles.

In Neomeris, which lacks or as good as lacks the dorsal fin, the skin
-of the back in the place where the fin is found in its relatives, and
also somewhat further forward and backward, is divided into small,
rather regularly placed plates, each bearing a small elevation. Similar
small knobs are found, though not always, in the nearly related genus
Phocena, along the anterior surface of the dorsal fin and sometimes
also scattered in other regions. Kiikenthal, who has closely examined
these structures, thinks that they are a kind of scale, although they
have in their intimate formation only toa slight degree the characters
that are found in scales. The explanation given is that they are

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL V2

scales which are in process of atrophy—on the way to disappearing.
A concurrent reason to regard them as scales is that traces of a
scaly covering are found in some extinct whales, Zeuglodon and
_ Delphinopsis.

But to Kukenthal’s and Abel’s conception there is something to
oppose.

It has been shown that the Hyzenodonts, the most primitive car-
nivores, are the precursors of the Cetacea among terrestrial mammals.
Remains of Hyznodonts are found in great numbers in many locali-
ties; but there has never been discovered the slightest indication that
any Hyzenodont or any other carnivore has been armored. Remains
of Zeuglodonts are found in various parts of the world, but nowhere
except in the case of the specimens from Alabama have dermal
ossicles been demonstrated in connection with the skeletons. If there
had been a dermal armature it certainly would have been found some-
where or other. Besides, it cannot be said to be proved that the
plates from Alabama are not those of some kind of turtle. Anyone
who has seen the roof-shaped keel on a Psephophorus carapace, and
has seen the fragments of the carapace mixed up together, will not
allow himself to be persuaded by Abel’s word in this connection.
Finally it is improbable that Zeuglodon had a dorsal fin, since this
fin may be absent (probably not-developed rather than lost) in diverse
recent cetaceans, both Balzenids and Delphinids.

The minute plates in Delphinopsis are altogether too uncertain to
give any evidence. Their characters are, besides, so far from recall-
ing what is otherwise known of dermal bones that one is tempted
rather to regard as an error their determination as such structures.

The small callosities in the skin of Neomeris and Phocena are
scarcely the remains of a dermal armature, they are rather entirely
new structures. It is too suspicious that nothing of the sort should
be present in lower Cetacea, but that it should be in exactly some of
the very highest that it is found. The structure of the callosities,
moreover, gives no real support to the idea that they are scales.

Altogether there is no proof that the Cetacea or their ancestors
among the mammals ever have had dermal armature.

*(P. 10.) For comparison a few of the most important and most
independent synopses of the groups of Cetacea are here given.

A fundamental work in the direction of throwing light on the
mutual relationships of the Cetacea is due to Flower, who, however,
took into consideration the recent forms only. In 1866 (69), in his

No. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 59

paper on Jnia and Pontoporia (Trans. Zool. Soc. London, vol. 6,
p- 115), he gave the following synopsis:

Cetacea.
I. Mystacoceti or Balenoidea.
Balenide.
Baleninz: Balena, Eubalzna.
Balenopteride.
Megapterine: Megaptera.
Baleenopterine: Physalus, Sibbaldius, Balenoptera.
II. Odontoceti or Delphinoidea.
Physeteride.
Physeterinz: Physeter, Kogia.
Ziphiinze: Hyperoodon, Berardius, Ziphius, Dioplodon,
Micropteron.
Platanistide.
Platanistinz: Platanista.
Iniinee: Pontoporia?, Inia.
Delphinide.
Beluginee: Monodon, Beluga (=Delphinapterus).
Delphinine ?: Phoczna, Neomeris, Grampus, Orca, Pseu-
dorca, Lagenorhynchus, Delphinus, Delphinapterus
(=Tursio), Globicephalus.

His conception of the relations between the genera in the family
Delphinide Flower developed more fully in 1883 (Proc. Zool. Soc.
London). His arrangement there was as follows:

A,
a. Monodon, Delphinapterus (Beluga).
b.
a. Phoczna, Neomeris.
8. Cephalorhynchus, Orcella, Orca, Pseudorca, Globiceps,
Grampus, Feresia, Lagenorhynchus.
By
a.
a. Delphinus.
8B. Tursiops, Clymenia, Steno.
b. Sotalia.

Flower followed essentially the same arrangement as that of 1866
and ’83, but with greater clearness as to the genera, in 1891, in “ An
Introduction to the Study of Mammals Living and Extinct ” which

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

be published in association with Lydekker. The arrangement is as
follows:

Cetacea.
Mystacoceti, Balenoidea.

Balenide: Balena, Neobalena, Rhachianectes, Megaptera,
Balenoptera. Extinct Genera: Cetotherium, Herpetocetus.

Archeoceti.
Zeuglodontide: Zeuglodon.
Odontoceti, Delphinoidea.

Physeteride.

Physeterinz : Physeter, Cogia. Extinct: Physeterula, Eucetus,
Physetodon, Scaldicetus, Physodon, Hoplocetus.

Ziphiine: Hyperoodon, Ziphius, Mesoplodon, Berardius.
Extinct: Choneziphius.

Squalodontide: Squalodon.

Platanistide: Platanista, Inia, Pontoporia. Extinct: Palzopon-
toporia (Pontistes), Champsodelphis, Schizodelphis, Pris-
codelphinus, Lophocetus, Ixacanthus, Rhabdosteus,
Agabelus. .

Delphinide.

Group A. Monodon, Delphinapterus, Phoceena, Neomeris,
Cephalorhynchus, Orcella, Orca, Pseudorca,
Globicephalus, Grampus, Feresia, Lageno-
rhynchus.

Group B. Delphinus, Tursiops, Prodelphinus, Steno, Sotalia.

Max Weber (Die Saugethiere, 1904) agrees closely with the con-
ception of Flower and Lydekker. One of the greatest differences is
that a special family, Rhachianectid@ is established for Rhachionectes
and also a family Delphinapteride for Delphinapterus and Monodon.

Abel has made special researches on the fossil Cetacea, and he has
tried to determine their positions in relation to the recent forms.
Besides what he has said on the subject in his special monographs,
he has more or less completely set forth his ideas in the papers: Die .
Stammesgeschichte der Meeressaugetiere; Meereskunde, Sammlung
volksttmlicher Vortrage, 1907; Grundziige der Palaentologie der
Wirbeltiere, 1912; and Die vorzeitlichen Saugetiere, 1914. He has
not given any general synopsis except in the following genealogical
tree, published in his work Die Vorfahren der Bartenwale, 1914

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 61

(Denkschr. k. Akad. Wissensch. Wien, math. -naturw. KI., vol. go,
gee ae

Die Stamme der Wale.
I. Mystacoceti (auct.) II. Delphinoceti (nov.) III. Squaloceti (nov.)
(Bliitezeit im Pliocin.) (Bliitezeit in der Gegenwart.) (Bliitezeit im Miocan.)

Balaeno- Rhachia- Balae- Delphinidae Physete- Ziphii- Rurhino- Platani-
pteridae nectidae nidae ridae dae delphidae stidae

Acrodelphidae

Phocaenidae
Squalodont idae
Patriocetidae Agorophiidae
Zeuglodon- 4
tidae Microzeuglodon-

tidae

Creodontia.

(To the Acrodelphide are referred, among the living genera, Del-
phinapterus and Monodon, Inia and Pontoporia.)

True’s paper On the Classification of the Cetacea (Proc. Amer.
Philos. Soc., Philadelphia, vol. 47, 1908, pp. 384-391) is mostly an
account of the opinions which Abel had expressed in 1905, with some
objections and some assent.

To cetaceans both recent and fossil have been given various generic
names in addition to those which appear in the present article. These
“names are partly well known as synonyms of others; but partly the
corresponding animals are so slightly known that no certain opinion
can be had about them. References to all the names theretofore used
for cetaceans are found in Trouessart, Catalogus Mammalium tam
viventium quam fossilium, 1897-99, with supplement, 1904-5, and in
Palmer, Index Generum Mammalium, 1904; many names are also
to be found in Beddard, A Book of Whales, 1900 ; detailed references
to the North American fossil genera are due to Hay, Bibliogr. and
Catal. of the fossil Vertebrata of North America, Bull. U. S. Geol.

62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Surv., No. 179, 1902. Incorrectly formed names which cannot
be accepted as finally settled are, in the present treatise, marked
with “ ns

* (P. 13.). The backbones of the Egyptian “ Zeuglodon”’ osirts
with its short vertebre (especially Stromer, Beitr. Palaontol. u. Geol.
Oesterreich-Ungarns, etc., vol. 21, 1908, pl. 4, fig. 1), and those of
the American Z. cetoides with its long vertebre (especially Gidley,
Proc. U. S. Nat: Mus., vol. 44, 1913, p. 81) differ to. sich aydegres
that according to ordinary standards the placing of these animals in
the same genus, as has hitherto been done, is certainly out of the
question. Z. cetoides is the type of the genus Zeuglodon. By acci-
dent no special name has been proposed that can with full right be
used for the genus to which “ Zeuglodon”’ osiris belongs. But the
name Prozeuglodon seems to have become vacant and may therefore
with some propriety be used. It was proposed by Andrews (espe-
cially Tert. Vertebr. of the Faytm, Egypt, 1906) for a lot of Eocene
cetacean remains from Egypt which he united under the name
P.atrox. But according to Stromer, the type of the species, a skull,
and some of the other remains belong to the previously described
Zeuglodon isis, which is probably correctly called Zeuglodon, while
still others are referable to “ Z.” osiris. In a way therefore “ Z.”
osiris has also been called Prozgeuglodon. Possibly the name Doryo-
don (“ Dorudon”) might be used for the genus in question with
short vertebre, or, if there are several genera with short vertebre, for
one of them (see, among others, Leidy, Journ. Acad. Nat. Sci.
_ Philadelphia, ser. 2, vol. 7, 1869, pp. 428, 431, and Lucas, Proc. U. S.
Nat. Mus., vol. 23, 1900, p. 331). But Doryodon is still not suffi-
ciently known, not even after True (Bull. Mus. Comp. Zool., vol. 52,
1908, pp. 65-78, pls. 1-3) has examined the fragments on which
Gibbes founded the genus; the remains in question are altogether
too incomplete. For the American “ Zeuglodon brachyspondylus
minor” Joh. Miller and Stromer, also with short vertebrz, which
True compares with Doryodon and finds different, True (J. c.) pro-
poses to erect a new genus, Zygorhiza; but the relationship between
it and “ Zeuglodon”’ osiris is not at all clear.

“(P. 15.) The pelvis and femur of Zeuglodon cetoides are
described and figured by Lucas (Proc. U. S. Nat. Mus., vol. 23, 1900,
pp. 327-331, pls. 5-7). Both right and left innominates were found
associated with a backbone lying in the position relative to the
vertebrze in which one would expect to find them. In spite of this
circumstance Abel explained the bones in question as the coracoid of

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 63

a gigantic bird which he called “ Alabamornis”’ gigantea (Ueber den

als Beckengtrtel von Zeuglodon beschriebenen Schultergurtel eines

Vogels aus dem Eocan von Alabama, Centralblatt fur Mineralogie,

Geologie und Palaontologie, 1906, pp. 450-458, with illustrations).

Stromer (Beitr. z. Palaont. u. Geol. Oesterreich-Ungarns, etc., vol.

21, 1908, p. 146) has expressed doubts as to the correctness of Abel’s

interpretation, and Gidley (Proc. U. S. Nat. Mus., vol. 44, 1913,

pp. 649-654, with illustrations) who has re-examined the specimens,

has entirely thrown it over. It can hardly be doubted that Lucas and

Gidley are correct.

*(P. 15.) On the Zeuglodontide [Basilosauridz]| see especially:

Joh. Miiller: Ueber die fossilen Reste der Zeuglodonten von Nord-
america mit Rticksicht auf die europadischen Reste aus dieser
Familie, 1849, pp. 1-38, pls. 1-27.

Carus: Das Kopfskelet des Zeuglodon hydrarchus ; Nova Acta Acad.
Czs. Leop. Carol., vol. 22, pt. 2, 1850, pp. 373-390, pls. 39A & B.

Brandt: Untersuchungen iiber die fossilen und subfossilen Cetaceen
Europa’s ; Mém. Acad. Imp. Sci. St. Pétersbourg, ser. 7, vol. 20,
No. I, 1873, pp. 291-313, 334-340, pl. 34. Zeuglodon.

Hector: Notes on New Zealand Cetacea, recent and fossil; Trans.
and Proc. New Zealand Inst., 1880, vol. 13, 1881, pp. 434-436,
pl. 18. “ Kekenodon.”

Lydekker: On Zeuglodont and other Cetacean Remains from the
Tertiary of the Caucasus; Proc. Zool. Soc. London, 1892, pp.
558-561, pl. 36. Zeuglodon=in part Microzeuglodon.

Dames: Ueber Zeuglodonten aus Aegypten und die Beziehungen der
Archeoceten zu den ubrigen Cetaceen; Palaeontologische Ab-
handlungen, herausgeg. von Dames und Kayser, vol. 5, pt. 5,
1894, pp. 1-36, pls. 1-7.

Lucas: The Pelvic Girdle of Zeuglodon, Basilosaurus cetoides
(Owen), with notes on other portions of the skeleton; Proc.
U. S. Nat. Mus., vol. 23, 1900, pp. 327-331, pls. 5-7. |

Abel: Les Dauphins Longirostres du Boldérien des Environs
d’Anvers; Mém. Mus. Roy. d’Hist. Nat. de Belgique, vol. 1,
1901, pp. 8-9, 24-32. On the dentition and dermal armature in
Zeuglodon.

Stromer: Zeuglodon-Reste aus dem oberen Mitteleocin des Fajum;
Beitrage zur Palaontologie und Geologie Oesterreich-Ungarns
und des Orients, vol. 15, pts. 2 and 3, 1903, pp. 65-100, pls. 8-11.
Zeuglodon=in part Prozeuglodon.

64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL: 72

E. Fraas: Neue Zeuglodonten aus dem unteren Mitteleocin von
Mokattam bei Cairo ; Geologische und Palzeontologische Abhand-
lungen, herausgeg. von Koken,-vol. 10, pt. 3, 1904, pp. 199-220,
pls. 10-12. Protocetus and Mesocetus, later called Eocetus.

Abel: Les Odontocétes du Boldérien d’Anvers; Mém. Mus. Roy.
d’Hist. Nat. de Belgique, vol. 3, 1905, pp. 21-25. On the denti-
tion in the Zeuglodonts.

Andrews: A descriptive Catalogue of the Tertiary Vertebrata of the
Faytm, Egypt, 1906, pp. 235-357, pls. 20-21. Zeuglodon, Pro-
zeuglodon.

Stromer: Die Urwale (Archaeoceti) ; Anatomischer Anzeiger, vol.
33, 1908, pp. 81-88, pl. 1. A short synopsis of the most important
part of the contents of the next paper. In the explanation of
plates the name Zeuglodon (Dorudon) osiris is used ; in the suc-
ceeding paper this use of “ Dorudon” is abandoned.

Stromer: Die Archaeoceti des Aegyptischen Eozans ; Beitr. Palaontol.
u. Geol. Oesterreich-Ungarns u. des Orients, vol. 21, 1908,
pp. 106-178, pls. 4-7. Protocetus, Eocetus, Zeuglodon, Pro-
seuglodon.

True: The fossil Cetacean, Dorudon serratus Gibbes; Bull. Mus.
Comp. Zool., vol. 52, 1908, pp. 65-78, pls. 1-3. Dorudon
(=Doryodon) and Zygorhiza.

Gidley: A recently mounted Zeuglodon skeleton in the United States
National Museum; Proc. U. S. Nat. Mus., vol. 44, "913; pp. 649-
654, pls. 81, 82 al text figures.

Kekenodon was established by Hector (i.88i l. c.) on remains
from Eocene strata in New Zealand. That which has been found is
not much else than a lot of loose teeth which show strong similarity
to Zeuglodon, but it is impossible to get any certain idea of the exact
generic relationships. Hall (Proc. Roy. Soc. Victoria, n. s., vol. 23,
pt. 2, I91I, p. 262) refers it to the Squalodontidae but gives no
reasons.

Microzeuglodon was established by Stromer (Beitr. Palaont. u.
Geol. Oesterreich-Ungarns, vol. 15, 1903, p. 89) and accepted by
Abel (Odontocétes du Boldérien, 1905, p. 35). The basis of the
genus 1s Lydekker’s Zeuglodon caucasicus (Proc. Zool. Soc. London,
1892, pp. 559-561, pl. 36), based on a few remains, not certainly be-
longing together, found in Tertiary strata in the Caucasus: a small
piece of a lower jaw with four cheekteeth, only two of which are
tolerably complete, a humerus and a caudal vertebra. The teeth are
serrate on both fore and hind margins of the crown. Abel referred

No. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 65

it at first to the Squalodonts, later he put it in a separate family. It
plays an important part in Abel’s studies of cetacean descent, whether
rightly or not time will perhaps tell ; meanwhile the genus is altogether
too slightly known for anything positive to be built on it.

Eocetus described by Fraas (first called Mesocetus Fraas, not of
Van Beneden, not of Moreno) is thought to be a connecting link
between Protocetus and Zeuglodon, with long vertebrae. The remains
are still too uncertain for judgment to be passed.

*(P. 19.) Kikenthal (Vergl. -anat., etc., Unters. an Walthieren ;
Denkschr. medic. -naturw. Ges. Jena, vol. 3, pt. 2, 1893, p. 291) thinks
that the bone in the hand of Balena mysticetus, which is ordinarily
regarded as a remnant of the first finger, a first metacarpal, is not
that, but a finger before the first finger, a prepollex, in spite of the
fact that the same bone in Balena australis (as can also be seen in two
skeletons in Copenhagen) may bear two well-developed phalanges,
something that is not elsewhere seen in any “ prepollex.’ As to the
longest finger, which is usually reckoned as the third, he believes that
it is not the third but the second, and that the third is absent. The
reason for this remarkable interpretation is probably a desire to find
agreement with Balenoptera, in which he thinks he has proved that
the third finger is the one which is absent, and not the first as is gen-
erally supposed. If the first finger were present in Balena in a more
or less atrophied condition, it would be reasonable to suppose that it
was this finger which is absent in the nearly related Balenoptera,
which has only four digits; but that belief Kukenthal will not allow.
Occasionally he has found in Balenoptera musculus something re-
sembling a few atrophied phalanges lying loose in the palm between
‘the fingers that are usually called the third and fourth. These
structures Kiikenthal regards as remnants of the third finger and
thus to be proof that it is the third finger which is absent in the
tetradactylous hand. Protest against Kukenthal’s interpretation has
already been made by Braun and Kunze (see Kunze, Zool. Jahrb.,
Abth. f. Anat., etc., vol. 32, 1912, pp. 639-641). There can be no
doubt that there is here a case of malformation, a supernumerary
digit, a kind of doubling of one of the fingers. Tendencies in this
direction are indeed not rare in cetaceans, which on the whole show
great indifference as to details in the structure of their abnormal hand.

“(P. 21.) Onthe Balenidz see especially (Of the numerous works
that deal with cetaceans there are many others that might have a
claim to be mentioned. The choice that has been made here and in
the corresponding lists for other families is somewhat arbitrary.

66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL 2

Most attention has been paid to indicating papers that describe the
various forms of cetaceans, and especially to those which contain
illustrations of the fossil members of the order.) :

Cuvier: Recherches sur les Ossemens fossiles, ed. 4, vol. 8, pt. 2.
1836, pp. 250-321, pls. 226-228, with figures of skulls and other
skeletal parts of Balena, Balenoptera, Megaptera, Plesiocetus,
mostly under other names.

Eschricht: Underségelser over Hvaldyrene, 2 den Afhandl., Anato-
misk Beskr. af de ydre Fosterformer hos to nordiske Finhval-
Arter; Kgl. Danske Vidensk. Selsk. naturvy. mathem. Afhandl., |
pt. 11, 1845, pp. 203-279. 3 dje Afhandl., Om Fosterformerne i
Bardehvalernes Ernzrings- og Forplantelsesredskaber ; ibid., pp.
281-320, pls. 1-4. 5 te Afhandl., Finhvalernes Oesteologi og
Artsadskillelse ; ibid., pt. 12, 1846, pp. 225-396, pls. 4-16. Bale-
noptera, Megaptera. Figures of skulls and other skeletal parts,
of embryos and adults, of external and other characters.

Eschricht and Reinhardt: Om Nordhvalen (Balena mysticetus L.) ;

_ Kgl. Danske Vidensk. Selsk. Skrifter, ser. 5, naturv. mathem.
Afd., vol. 5, 1861, pp. 433-592, pls. 1-6. Figures of the exterior,
the entire skeleton, the skull of adult and young and other parts
of Balena, of the skull of Balenoptera and Megaptera.

Malm: Monographie illustrée du Baleinoptere trouvé le 29 Octobre
1865 sur la cote occidentale de Suede, 1867, pp. I-110, pls. 1-20,
with figures of the exterior, some skeletal parts and other features
of Balenoptera caroline=B. gigas, sibbaldit.

Eschricht: Ni Tavler til Oplysning af Hvaldyrenes Bygning, med
Forklaring af Reinhardt; Kgl. Danske Vidensk. Selsk. Skrifter,
ser. 5, naturv. mathem. Afd., vol. 9, 1, 1869. On plates 1 and 2
are found figures of the skull of embryo Balena japonica=
B. australis.

Van Beneden and Gervais: Ostéographie des Cétacés vivants et
fossiles, text and plates, 1868-80, pp. 1-634, pls. 1-67. As
regards illustrations, with respect to both recent and extinct
Cetacea, the most sumptuous work that exists. Balzenids espe-
cially pp. 29-291, pls. I-17.

Dwight: Description of the Whale (Balenoptera musculus Auct.) in
the possession of the Society, with remarks on the classification
of Fin Whales; Mem. Boston Soc. Nat. Hist., vol. 2, 1871-78,
pp. 203-230, pls. 6-7. Exterior and skeleton.

Brandt: Untersuchungen iiber die fossilen und subfossilen Cetaceen
Europa’s ; Mém. Acad. Imp. Sci. St. Pétersbourg, ser. 7, vol. 20,

No. 8 INTERRELATIONSHIPS OF THE CETACEA

WINGE 67

no. I, 1873, pp. 1-372, pls. 1-34. Contains a section, pp. 18-202,
on the then-known fossil Balenids, among them Cetotherium
and Plesiocetus. On Patriocetus, see under Squalodontide.

Brandt: Erganzungen zu den fossilen Cetaceen Europa’s; Mém.
Acad. Imp. Sci.,St»-Pétersbourg, ser..7, vol; 21, No: 6, 1874,
pp. 1-54, pls. 1-5. Contains a section on the whalebone whales,
ppe2-re,- plot.

Capellini: Della Balena di Taranto confrontata con quella della
Nuova Zelanda e con talune fossili del Belgio e della Toscana ;
Memorie dell’Accademia delle Scienze dell’Instituto di Bologna,
ser. 3, vol. 7, 1877, pp. 1-34, pls. 1-3, with illustrations of
exterior, skull, ear bones, nasal, cervical vertebrz, other skeletal
parts, etc. Balena tarentina=B. australis.

Gasco: Intorno alla Balena presa in Taranto nel Febbrajo 1877;
Atti della Reale Accademia delle Scienze Fisiche e Matematische,
Napoli, vol. 7, 1878, pp. 1-47, pls. 1-9, with figures of exterior,
skull, other skeletal parts, etc. Balena biscayensis=B. australis.

Gasco: La Baleena Macleayius del Museo di Parigi; Annali del Museo
Civico di Storia Naturale di Genova, vol. 14, 1879, pp. 509-551.
Balena australis. Description of skeleton. / |

Gasco: Il Balenotto catturato nel 1854 a San Sebastiano (Spagna),
Balaena biscayensis, Eschricht, per la prima volta descritto;
Annali del Museo Civico di Storia Naturale di Genova, vol. 14,
1879, pp. 573-608. Description of skeleton.

Van Beneden: Description des Ossements fossiles des environs
d’Anvers, 2 partie, Genres Balenula, Balena et Balenotus;
Annales du Musée Royal d’Hist. Nat. de Belgique, série pale-
ontol., vol. 4, Text, 1880, pp. 1-83, Atlas, 1878, pls. 1-39. 3 partie,
Genres Megaptera, Balenoptera, Burtinopsis et Erpetocetus,
ibid., vol. 8, 1882; Text, pp. 1-90, Atlas, pls. 1-109. 4 partie,
Genre Plesiocetus ; ibid., vol. 9, 1885, Text, pp. 1-40, Atlas, pls.
I-30. 5 partie, Genres Amphicetus, Heterocetus, Mesocetus,
Idiocetus et Isocetus ; ibid., vol. 13, 1886, Text, pp. 1-139, Atlas,
pls. 1-75.

Burmeister: Atlas de la Description Physique de la République
Argentine, sec. 2, mammif., pt. 1, Die Bartenwale der Argen-
tinischen Kiisten, 1881, pp. 3-40, pls. 1-7. Mostly on Balen-
optera. Figures of exterior, skull, vertebral column and other
skeletal parts.

Struthers: On the bones, articulations and muscles of the rudi-
mentary hind-limb of the Greenland Right-Whale, Balzena mysti-

5

68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLs 'F72

cetus; Journal of Anatomy and Physiology, vol. 15, 1881, pp.
141-176, pls. 14-17; ibid., pp. 301-321.

| Holder: The Atlantic Right Whales: A Contribution; Bull. Amer.
Mus. Nat. Hist., vol. 1, No. 4, pp. 99-137, pls. 10-13. May 1,
1883. External characters and skeleton. |

Malm: Skelettdelar af Hval insamlade under Expeditionen med Vega
1878-1880 ; Bihang till K. Svenska Vet. Akad. Handlingar, vol.
8, No. 4, 1883, section pp. 17-98, with figures of parts of skulls,
etc., of Rhachionectes and Balena.

Tullberg: Bau und Entwicklung der Barten bei Balzenoptera Sib-
baldii; Nova Acta Reg. Soc. Sci. Upsal., ser. 3, 1883, pp. 1-36,
pls. 1-7.

Delage: Histoire du Baleenoptera musculus échoué sur la plage de
Langrune; Archives de Zoologie expérimentale et générale, ser.
2, vol. 3 bis, 1885, pp. 1-152, pls. 1-21. Exterior and anatomy.

H. P. Gervais: Sur une nouvelle espece de Mégaptere (Megaptera
indica) provenant du Golfe Persique; Nouvelles Archives du
Muséum d’Hist. Nat. de Paris, ser. 2, vol. 2, 1887-88, pp. 199-
218, pls. 18-20. Figures of skeleton and skuil.

Struthers: Memoir on the anatomy of the Humpback-Whale, Me-
gaptera longimana; Reprint from the Journal of Anatomy and
Physiology, 1887-89, pp. 1-189, pls. 1-6.

Graells: Las Ballenas en las costas oceanicas de Espana; Mem. Real
Acad. Cien., Madrid; vol. 13,’ pt. 3, 1889) pp: I-11 5;plsaanas
Deals mostly with Balena biscayensis (=B. australis) from San
Sebastian. Figures of exterior, skull, skeleton.

Rios Rial: La Ballena Euskara, 1890, pp. 1-105. . Balena australis,
mostly on skeletons from San Sebastian.

Lydekker: Cetacean skulls from Patagonia; Anales del Museo de
La Plata, Paleontologia Argentina, vol. 2, 1893, pp. 2-4, pl. I.
Cetotherium.

Struthers: On the rudimentary hind-limb of a Great Fin-Whale,
Baleenoptera musculus, in comparison with those of the Hump-
back-whale and the Greenland Right-Whale ; Journal of Anatomy
and Physiology, vol. 27, 1893, pp. 291-335, pls. 17-20.

Struthers: On the carpus of the Greenland Right-Whale, Balzna
mysticetus, and of Fin-Whales; Journal of Anatomy and
Physiology, vol. 29, 1895, pp. 145-187, pls. 2-4.

Beddard: Contrib. towards a knowledge of the osteology of the
Pigmy Whale (Neobalena marginata); Transact. Zool. Soc.
London, vol. 16, pt. 2, No. 1, 1901, pp. 87-114, pls. 7-9.

No. 8 INTERRELATIONSHIPS OF THE CETACEA—-WINGE 69

Racovitza: Cétacés; Expédition Antarctique Belge; Résultats du
Voyage du S. Y. Belgica en 1897-99, Rapports Scientifiques,
Zoologie, 1903, pp. 1-142, pls. 1-4. Contains much information
on the external characters and the habits of cetaceans, especially
whalebone whales, and gives numerous references to earlier
papers on the subject.

True: The whalebone whales of the Western North Atlantic com-
pared with those occurring in European Waters with some
observations on the species of the North Pacific; Smithsonian
Contributions to Knowledge, vol. 33, 1904, pp. 1-332, pls. 1-50.
Deals with species of the genera Balena, Rhachionectes, Balen-
optera, Megaptera. Numerous illustrations, especially of skulls
and exterior. Copious references to earlier works on the subject.

Turner: The Right-Whale of the North Atlantic, Balzna biscay-
ensis : its skeleton described and compared with that of the Green-
land Right-Whale; Transact. Roy. Soc. Edinburgh, vol. 48,
1913, pp. 889-922, pls. 1-3, with text figures.

Abel: Die Vorfahren der Bartenwale; Denkschriften der k. Akad.
der Wissensch. Wien, Mathem. naturw. K1., vol. 90, 1914, pp.
155-224, pls. 1-12. Patriocetus, Agriocetus. Review of the
origin of the whalebone whales.

Roy C. Andrews: Monographs of the Pacific Cetacea, I, The Cali-
fornia Gray Whale (Rhachianectes glaucus) ; Mem. Amer. Mus.
Nat. Hist., n. s., vol. 1, pt. 5, 1914, pp. 227-287, pls. 19-28, with
figures of the exterior and of all parts of the skeleton.

Roy C. Andrews: The Sei Whale (Balznoptera borealis) ; Mem.
Amer. Mus. Nat. Hist., n. s., vol. 1, pt. 6, 1916, pp. 289-388, pls.
29-42, with figures of the exterior and of all parts of the skeleton.

G. M. Allen: The Whalebone Whales of New England; Mem.
Boston Soc. Nat. Hist., vol. 8, No. 2, 1916, pp. 105-322, pls. 8-16,
with figures mostly of the exterior. Balena, Balenoptera,
Megaptera.

Besides Plesiocetus and Cetotherium many other genera of fossil
Balenids have been described, especially in papers by Van Beneden,
Brandt and Cope, but the bases for most of them are scanty. H. Winge
(Om Plesiocetus og Sqvalodon fra Danmark, Vidensk, Medd. Natur-
hist. Foren. 1909) has attempted to estimate the value of a number of
the genera in question: Aulocetus, Mesoteras, Cetotheriopsis, Megap-
teropsis, “ Burtinopsis,” Herpetocetus, Eucetotherium, Plesiocetopsis,
Cetotheriophanes, Cetotheriomorphus, Idiocetus, Heterocetus, Amphi-
cetus, Mesocetus, Isocetus, Pachycetus, Siphonocetus, Ulias, Tre-

7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

tulias, Metopocetus, Cephalotropis, Rhegnopsis. It is apparent that
as regards most of them there is scarcely any reason for separation
from Plesiocetus or Cetotherium or from recent genera, and that the
few which appear to be more peculiar are so slightly known that they
can scarcely be classified. True (The Genera of Fossil Whalebone
Whales allied to Balenoptera, Smithsonian Misc. Coll., vol. 59, No. 6,
1912, pp. 1-8) who later went over the subject came in all essentials
to the same conclusion.

Agriocetus is most likely a whalebone whale, but it is too slightly
known to be classified. It was described by Abel (1914, l. c., pp. 188-
194, pls. 4, 5, 7) from a very imperfect and indistinct skull from
Tertiary strata at Linz, referred to Squalodon by earlier authors.
Abel regards it as a near relative of Patriocetus, a step nearer to the
true whalebone whales. Only better discoveries will show whether
he is right or not.

Perhaps. Patriocetus belongs to the family Balenid@ as it is under-
stood in the present work, but it is not sufficiently known to be
definitely placed. It was described by Abel (1914, l. c.) who has
given a full account of the history of the remnants in question. The
basis of the genus was partly some rather imperfect fragments which
previously had most often been referred to under the name Squalodon
ehrlichii, partly a quite well-preserved skull found later, all from
Tertiary strata at Linz. If Abel’s interpretations and conjectures are
right he is no doubt correct in regarding Patriocetus as a precursor of
the true whalebone whales. Abel refers it to the Archzoceti, or at
least leaves the question undecided whether it actually belongs to
this group or to the Mystacoceti (Die vorzeitl. Sauget., 1914, p. 88) ;
most probably it should be regarded as a whalebone whale, a Balzenid
with the dentition still functional. But there is reason for doubt
about certain details in Abel’s account.

Patriocetus has in the skull a remarkably strong resemblance to
Agorophius, a resemblance that was seen by Brandt (1873, l. ¢.,
p. 324) although the remains then at hand were rather insignificant ;
and Agorophius belongs incontestably to the series of toothed whales
as a near relative of Squalodon. The peculiarity which places Agor-
ophius among the Odontoceti in opposition to the Mystacoceti is that
the maxillary bone pushes itself posteriorly as a thin limina over the
supraorbital process of the frontal, but does not stop in front of it,
or push itself in under it, or content itself with also covering it with
a narrow margin anteriorly. According to the great resemblances
which are found otherwise between the skulls of Agorophius and

No. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 71
Patriocetus one would expect that the two genera would resemble
each other in this particular also; but according to Abel’s representa-
tion Patriocetus is here like the whalebone whales. According to
the photographic illustrations which accompany Abel’s paper it is
scarcely possible to see whether his exposition of the conditions is
right or not; the skull is too weathered and obscure. There is, how-
ever, a detail in his description of the boundary between the maxillary
and frontal, which probably must be wrong or at least must awaken
doubt. He says of the maxillary that it, at its postero-internal ex-
tremity, does not extend nearly so far backward as the nasal process
of the intermaxillary, which, on the contrary, like that of other whales,
extends up, far backward, alongside the outer margin of the nasal
and beyond. But there is elsewhere no cetacean, either among the
Archeoceti, Mystacoceti, or Odontoceti, in which the maxillary does
not reach postero-internally as far back as the intermaxillary or even
further, pushing itself up over the frontal. This is an inheritance
from ancestors among the carnivores or from yet more distant fore-
runners. Abel says, it is true (/. c., p. 162) that Patriocetus in this
regard resembles /thachionectes, one of the recent whalebone whales ;
but this is an error. In one of the figures of the skull of Rhachi-
onectes published by Andrews (1914, /. c., pl. 25) it can be clearly
seen that a long process from the maxillary extends along the outer
side of the intermaxillary to its hindmost end; and it is so described
by Andrews (p. 261). In the second of Andrews’ figures the process
is not visible; it is obviously broken off, as it is in the figures pub-
lished by Van Beneden (Bull. Acad. Roy. Sci. etc., de Belgique,
Seer volnas: Nore, Pebriary, 1877, pl) and» True «(1904; 1.¢.,
pl. 47, fig. 1), both of which represent the same skull (it is True’s
figure to which Abel refers). A similar injury no doubt must have
been suffered by the skull of Patriocetus; and if this process can be
broken away without leaving visible traces behind it the same might
be possible in the case of a thin plate-like process that originally
covered the supraorbital process of the frontal. How readily some-
thing of the kind can take place is shown by the type of Agorophius
(figured by Leidy, under the name Squalodon pygmeus, Journ. Acad.
Nat. Sci. Philadelphia, ser. 2, vol. 7, 1869, pl. 29, and by True,
‘Smithsonian Inst. Special Publ., No. 1694, 1907, pl.) : on the right
side of the skull large parts of the plate-like outgrowths from the
maxillary over the supraorbital process of the frontal are broken away
without having left behind any conspicuous traces on the frontal.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL 72

NI
bo

In Abel’s treatment of the dentition in Patriocetus there are also
various doubtful points. He asserts that the complement of teeth is
the one which is typical of the placentalia, ri teeth in each jaw; but
it is impossible to see how he has arrived at this conclusion. The
best skull in question lacks the anterior part of both upper jaw and
mandible, and it is impossible to say how much is lacking, or how
many teeth were implanted in the missing parts. Of teeth fixed in the
jaws there are known for the most part mere stubs—sometimes noth-
ing but roots. Besides these there are some dissociated teeth whose
position in the jaws is not certain. All that can be said is that some
of the teeth were simply conical with single root, and that most of
the cheekteeth had serrate crown and double roots. Altogether there
is not enough known to elucidate all the details of the dentition. It is
not probable that Abel should be right in his belief that the teeth
were present in the typical number. Such a cetacean as Patriocetus,
the skull of which was already highly developed in the direction of
the most advanced whales, scarcely could have had about the same
dentition as the Zeuglodonts. It is much more likely that the number
of teeth was increased above the typical as it is or has been in all the
Mystacoceti and Odontoceti which are known in this respect. In
order to believe in Abel’s representation of the facts we must see
more incontestable finds. (See also note 7, pp. 52-54.)

*(P. 23.) The asymmetry in the skull of the toothed cetaceans
has often been written about. Special treatises on the subject are
due to Pouchet (De l’asymétrie de la face chez les Cétodontes ; Nouv.
Arch. du Mus. d’Hist. Nat. Paris, 1886, pp. 1-16 of separate), Abel
(Die Ursache der Asymmetrie des Zahnwalschadels ; Sitzungsber. k.
Akad. Wissensch. Wien, Math.-naturw. Cl., vol. 111, pt. 1, 1902,
pp. 510-526, pl.), Kukenthal (Ueber die Ursache der Asymmetrie
des Walschadels; Anat. Anzeiger, vol. 33, 1908, pp. 609-618, with
illustrations) and Steinmann (Ueber die Ursache der Asymmetrie
der Wale; Anat. Anzeiger, vol. 41, 1912, pp. 45-54, with illustra-
tions) ; Pouchet and Abel refer to various earlier papers by other
writers.

As to the reason for the crookedness Pouchet says: ‘“ Nous en
ignorons l’origine.”

In 1893 Kuikenthal said very nearly the same: ‘“ Die physiologische
Ursache kennt man nicht, vielleicht ist sie in der eigenthiimlichen
Art der Locomotion vermittelst der Schwanzflosse zu suchen” (Ver-
gleich.-Anat. u. entwickelungsgesch. Untersuch. an Walthieren, pt. 2;
Denkschr. med.-naturwiss. Ges. zu Jena, vol. 3, pt. 2, p. 342), a

NO. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 73

thought that was not at the time carried further. But in 1908 Kiiken-
thal tried to give a more exact explanation. To begin with he thought
he could prove that the asymmetry in the bones of the face is found
not only in toothed cetacea, but also in the whalebone whales, though
only slightly defined. In two skulls of Balenoptera (of B. rostrata
and B. musculus) he had found certain of the facial bones just a
trifle broader on the right side than on the left. Next he discovered
that some embryos of toothed cetaceans, of Platanista, Steno, Glo-
biceps, Delphinus, Phocena, Hypercodon, had the caudal fin set
awry, not level but in such a position that the left fluke “ etwas schrag
nach aufwarts, der rechte schrag nach abwarts gerichtet war”
(p. 614). In 12 embryos of Delphinapterus the fin was, on the con-
trary, horizontal. All the embryos of whalebone whales examined
(of Balenoptera musculus and B. gigas) had the fin oblique in the
same manner as the toothed whales. How the fin is in adult cetaceans
is said to be not clearly understood; a few observations by other
investigators may, however, indicate that the obliquity is present in
the adults also. When a whale propels itself forward by means of a
sculling movement of the oblique caudal fin it is said to turn at the
same time to the left: “ Der Wal durchschneidet also bei derartiger
schrager Bewegung der Schwanzflosse das Wasser nicht genau in der
Richtung seiner Langsachse, sondern sein Weg verlauft von dieser
Geraden etwas schrag nach links zu” (p. 616). And from this is
said to result an oblique pressure of the water on the head, and con-
sequently the obliquity of the skull, since the bones on the left side
are pressed upon more than those of the right side, are made thicker,
etc.: “ Der Druck der beim Schwimmen durchschnittenen Wasser-
massen wird auf die linke Seite des Vorderkopfes starker wirken als
auf die rechte. Dieser Druck pflanzt sich durch die elastischen
Weichteile des Vorderkopfes hindurch auf die darunter liegenden
Schadelknochen fort. Die Wirkung dieses starkeren Druckes muss
sich zunachst in einer Verdickung der entsprechenden Schadel-
knochen aussern . .. .” (pp. 616-617). That the bones in the left
side of the face are not so wide as those of the right side is said to be
connected with the fact that the bones of the left side are the thickest :
“Es wird dadurch links eine kleinere Flache als rechts geschaffen,
welche den etwas starkeren Druck auszuhalten hat und damit bis zu
einem gewissen Grade einen Ausgleich gegentiber der rechten Seite
herbeiftihrt ” (p. 617).

Abel believes that the reason for the asymmetry of the skull in
toothed whales is to be found in the atrophy of the nasal bones, etc.,

74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

and in the shortening of the braincase, but cannot prove that asym-
metry would be the necessary result of these causes.

Lillie (section The asymmetry of the Odontocete skull, in Observa-
tions on the anatomy and general biology of some members of the
larger Cetacea; Proc. Zool. Soc. ‘London, 1910, vol. 2; pp. 7e"-yess
with figure) shows that the pharynx in Physeter is asymmetrical,
divided by the projecting larynx, which is strongly displaced to the
left, into a more spacious right and a narrower left section to accom-
modate the passage of food. In this circumstance should be found
the source of the asymmetry of the face ; why, is not further explained.

Steinmann starts from Kukenthal’s assertion about the asymmetry
in the candal fin. Kukenthal had said nothing as to the cause of this
crookedness, but Steinmann believes he has found it in the supposed
fact that whales originated from Ichthyosaurs and other marine
reptiles with a vertical fin, and that on its way to the horizontal posi-
tion the fin has come to rest obliquely.

Kukenthal is doubtless the one who has come nearest to the truth.
However, there are numerous objections to be raised against his
explanation. It cannot be said with any degree of correctness that
the skull in the whalebone whales has an asymmetrical face. A series
of skulls is before me, representing Balena, Balenoptera and Megap-
tera. Such asymmetry as can perhaps be shown here and there is
similar to that which is found in most mammals. I myself have
seen many cetacean embryos (in alcohol), representing both whale-
bone whales and odontocetes (of the former I have examined in this
connection embryos of 3 Balenoptera rostrata, 2 B. musculus, 5
Megaptera boops, of the latter numerous embryos representing the
genera Delphinapterus, Monodon, Prodelphinus, Delphinus, Lageno-
rhynchus, Globiceps, Phocena, Neomeris; I have been content with
examining the tails externally, | have not cross-sectioned them), but
I have not been able to convince myself of the presence of oblique-
ness in the tail which did not appear to find its explanation in artificial
pressure. I have also seen various adult newly dead cetaceans, both
whalebone whales and toothed cetacea of different kinds. It is true
I did not expressly examine them to observe obliqueness of the tail,
but I cannot recall the slightest evidence of its existence. Neither
can anything be detected in the numerous photographs of whales that
are before me. That obliqueness of the tail can actually be present
appears to be proved by Kukenthal’s figure of a cut off tail of Balen-
optera, which shows the flukes placed obliquely in relation to the
caudal vertebre ; but it must probably be an exception. Even if it

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 75

were true that the caudal fin in cetaceans was usually obliquely set

this would probably not hinder whales from swimming forward in a

straight line if they so wished. That the obliquity of the tail, if it

occurs as represented by Kukenthal, does not in any event necessarily
carry with it the asymmetry of the skull is proved by the whalebone
whales, whose tail is said to be oblique, but whose skull 1s without
asymmetry in spite of Kukenthal’s word to the contrary. And that
the asymmetry of the face is not dependent on obliquity of the tail is
proved by Delphinapterus, whose tail, also according to Ktkenthal’s
interpretation, is not oblique, but whose skull is distinguished by a
high degree of asymmetry. Should the water’s pressure work most
powerfully on the /eft side of the head it would be difficult, in spite
of Kukenthal’s attempt at an explanation, to understand why the
bones on the left side of the skull are narrow while those on the right
side are broad, or why the nasal passage is pushed over toward the
left side, a point that Kukenthal does not try to argue. Of Kuken-
thal’s explanations scarcely anything is left except the knowledge that
it is the pressure of the water which is responsible for the asymmetry
of the skull in the Odontoceti. Why the water presses obliquely is

still unknown, but the reason is not likely to be anything else than a

habit in the carriage of the head: the head presumably must be held

a little obliquely even when the animal is swimming straight forward ;

and the pressure must be strongest on the right side.

*(P. 24.) On the Squalodontide see especially :

Grateloup: Description d’un fragment de machoire fossile, d’un
genre nouveau de reptile (Saurien), de taille gigantesque, voisin
d’Iguanodon, trouvé dans le grés marin, 4 Léognan, prés Bor-
deaux, 1840, pp. 1-8, pl. Separate from Actes de l’Acad. des
sciences, belles lettres et arts de Bordeaux, vol. 2. Squalodon.

H. v. Meyer: Arionius servatus, ein Meersaugethier der Molasse ;
Paleontographica, vol. 6, 1856, pp. 31-43, pl. 6. Squalodon.

Jourdan: Descr. de restes fossiles de deux grands Mammifeéres con-
stituant le genre Rhizoprion et le genre Dinocyon; Annales des
Sciences Naturelles, sér. 4, Zoologie, vol. 16, 1861, pp. 369-372,
pl. 10. Rhizoprion=Squalodon.

Van Beneden: Recherches sur les Squalodons; Mém. Acad. Roy.
Belgique, vol. 35, 1865, pp. 1-85, pls. 1-4, with figures of the
upper jaw of S. antuerpiensis (and of the skull of S. ehrlichti=
Patriocetus).

Van Beneden: Recherches sur les Squalodons, Supplement; Mém.
Acad. Roy. Belgique, vol. 37, 1868, pp. 1-13, pl., with figure of
the under jaw of S. antuerpiensis.

76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Gervais: Du Squalodon et de sa comparaison avec le Zeuglodon;
Zoologie et Paléontologie Générales, ser. 1, 1867-69, pp. 170-182.

Fischer: Descr. d’une machoire inférieure de Squalodon Grateloupi ;
Actes de la Soc. Linnéenne de Bordeaux, vol. 27, 1869, pp. 12-22,
pli 2:

Delfortrie: Descr. d’une nouvelle machoire inférieure de Squalodon
Grateloupi dans le grés marin de Léognan, Gironde; Actes de la
Soc. Linnéenne de Bordeaux, vol. 27, 1869, pp. 133-136, pl. 5.

Leidy: Extinct Mammalian Fauna of Dakota and Nebraska; Journ.
Acad. Nat. Sci. Philadelphia, ser. 2, vol. 7, 1869, pp. 416-424,
pls. 28-30. Squalodon and Agorophius (under the name Squalo-
don pygmeus ).

Delfortrie: Un Squalodon d’espéce nouvelle dans le Miocene
superieur du Midi de la France; Actes de la Soc. Linnéenne de
Bordeaux, vol. 29, 1873, pp. 257-260, pl. 7.

Brandt: Untersuchungen uber die fossilen und subfossilen Cetaceen
Europa's; Mem. Acad: Sci. St. Pétersbourg, ser 77 voleeeo:
No. 1, 1873. Contains a section on the Squalodonts, pp. 315-332,
pl. 31-32. Squalodon, Neosqualodon (S. gastaldii) (and Patri-
ocetus, S. Ehrlichi).

Brandt: Erganzungen zu den fossilen Cetaceen Europa’s; Mém.
Acad. Imp. Sci. St.. Pétersbourg, ser. 7, vol. 21,- Nom Gime
Contains a section on Squalodon (and Patriocetus), pp. 28-47,
pls. 4, 5.

Van Beneden et Gervais: Ostéographie des Cétacés vivants et
fossiles, Text and Atlas, 1868-80, pp. 426-454, 519, pl. 28.
Squalodon.

Zittel: Ueber Squalodon Bariensis aus Niederbayern; Palzonto-
graphica, vol. 24, 1877, pp. 233-246, pl. 35.

Lortet: Note sur le Rhizoprion bariensis; Arch. Mus. d’hist. Nat.
de Lyon, vol. 4, 1887, pp. 315-319, pl. 25 bis and ter. Squalodon.

Lydekker: Cetacean skulls from Patagonia; Anales del Museo de La
Plata, Paleontologia Argentina, II, 1893, pp. 8-10, pl. 4. Pro-
squalodon.

Paquier: Etude sur quelques Cétacés du Miocéne; Mém. de la Soc.
Géol. de France, Paléontologie, vol. 4, pt. 4, Mém. No. 12, 1894,
pp. 12-17, pl. 18. Squalodon.

Lydekker: On the skull of a shark-toothed Dolphin from Patagonia ;
Proceed. Zool. Soc. London, 1899, pp. 919-922, with illustra-

tions. Prosqualodon.

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE V7.

Dal Piaz: Sopra alcuni resti di Squalodon dell’ arenaria miocenica
di Belluno; Palaeontographia Italica, vol. 6, 1900, pp. 303-314,
pls. 26-29.

Abel: Les Dauphins Longirostres du Boldérien des Environs
d’Anvers; Mém. Mus. Roy. d’Hist. Nat. de Belgique, vol. 1,
IQOI, pp. 9-10. On the dentition in Squalodon.

Dal Piaz: Neosqualodon, nuovo genere della famiglia degli Squalo-
dontidi; Abhandl. der Schweizerischen palaontologischen Gesell-
schaft, vol. 31, 1904, pp. I-21, pl.

Abel: Les Odontocétes du Boldérien d’Anvers; Mém. Mus. Roy.
d’Hist. Nat. de Belgique, vol. 3, 1905, pp. 25-38. On the
dentition.

True: Remarks on the type of the fossil Cetacean Agorophius
pygmeus (Miller) ; Smithsonian Inst. Special Publ., No. 1694,
1907, pp. 1-8, pl.

True: Descr. of a mandible and vertebre of Prosqualodon, etc. ;
Smithsonian Misc. Coll., vol. 52, pt. 4, 1910, pp. 447-456, pl. 43.

Hall: On the systematic position of the species of Squalodon and
Zeuglodon described from Australia and New Zealand; Proc.
Roy. Soc. Victoria, n. s., vol. 23, pt. 2, I9I1I, pp. 257-265, pl. 36.
Parasqualodon, Metasqualodon.

Abel: Cetaceenstudien, III. Mitteilung, Rekonstruktion des Schadels
von Prosqualodon australe (sic) Lyd. aus dem Muozan Pata-
goniens; Sitzungsber. k. Akad. Wissensch. Wien, mathem.-
Harioweeele VOl, 120, pt. 1, 1Ot2, pp.’ 57-75, pls. 1-3: . The
account contains various guesses ; Lydekker’s figures of the type
cannot be dispensed with in forming an opinion about the genus.
References to several other papers will be found in H. Winge,
Vidensk. Medd. Naturhist Foren., 1909, pp. 31-35.

Microsqualodon Abel is said to be identical with Neosqualodon Dal
Piaz. Abel (Odontocétes du Boldérien, 1905, pp. 35-36) established
the genus on the basis of Tertiary remains from Acqui which Brandt
had called Squalodon gastaldii. In a letter to Abel, however, which
Abel printed, Dal Piaz explains that these specimens, which he had
had the opportunity to see, must be referred to the earlier described
Neosqualodon.

Parasqualodon and Metasqualodon are proposed by Hall (1911,
l. c.) to include Tertiary Australian species that previously were
referred to Squalodon: S. wilkinsont M’Coy and S. harwoodu
Sanger. The genera are supposed to be nearly related to Prosqualo-
don, but they are known from loose teeth only and their status is still
uncertain.

SY
ee)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL eye

“(P. 28.) On the Platanistide see especially :

Cuvier: Recherches sur les Ossemens fossiles, ed. 4, vol. 8, pt. 2,
1836, pp. 88-90, 128-132, Atlas, pl. 223, with figures of skull of
Platanista.

Eschricht: Om Gangesdelphinen; Kgl. Danske Vidensk. Selsk.
Skrifter, 5te R., naturv. og mathem. Afd., vol. 2, 1851, pp. 345-
387, pls. 1-3. Platanista. Figures of exterior, skeleton, skull.

Burmeister: Descripcion de cuatro especies de Delfinides de la costa
Argentina en el océano Atlantico; Anales del Museo Publico de
Buenos Aires, vol. 1, 1864-69, pp. 389-445, pl. 23, 25-28. Ponto-
poria. Figures of exterior, skeleton, skull and other parts.

Flower: Descr. of the skeleton of Inia geoffrensis and of the skull of
Pontoporia blainvillii, with remarks on the systematic position of
these animals in the order Cetacea; Trans. Zool. Soc. London,
vol. 6, pt. 3, 1869, pp. 87-116, pls. 25-28.

Van Beneden et Gervais: Ostéographie des Cétacés vivants et fossiles,
Text and Atlas, 1868-80, pp. 454° -482, pls. 29-33. On almost all
the recent genera.

Anderson: Anat. and Zool. Researches, compr. an Account of the
two Exp. to Western Yunnan in 1868 and 1875; 1878; pp. 417-
550, pls. 25, 26, 28-32, 34-41. Platanista. Figures of exterior,
skeleton and soft parts.

Burmeister: Examen critico de los Mamiferos y Reptiles fosiles.
denominados por D. Augusto Bravard; Anales del Museo
Nacional de Buenos Aires, vol. 3, 1883-91, pp. 138-144, pl. 2,
figs. 12A-C. Pontistes. Figures of an imperfect skull.

Burmeister: Adiciones al examen critico, etc.; Anales del Museo
Nacional del Buenos Aires, vol. 3, 1883-91, pp. 451-460, pl. 8.
Saurodelphis. Figures of skull.

Abel: Les Dauphins Longirostres du Boldérien des Environs
d’Anvers; Mém. Mus. Roy. d’Hist. Nat. de Belgique, vol. 1,
IQOI, pp. I-95, pls. I-10. Contains a section dealing with the
Platanistide both living and extinct, and on several plates gives
figures of their skulls, mostly copied from other papers.

Abel: Cetaceenstudien, I], Der Schadel von Saurodelphis argentinus
aus dem Pliozan Argentiniens ; Sitzungsber. k. Akad. Wissensch.
Wien, mathem.-naturw. KI., vol. 118, pt. I, 1909, pp. 255-272,
pl. I, and text figures. Tries to prove that Burmeister’s figures
of the skull of Saurodelphis argentinus are to a notable degree
incorrect. Burmeister is said to have put together parts of two
different genera that should bear the names Saurodelphis and

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 79

Pontoplanodes. At present the problem cannot be said to be
solved; but there appears to be no good reason to doubt the
correctness of Burmeister’s determination. In the present paper
his presentation of the subject is followed. A piece of the beak
of Saurodelphis with the 8-shaped alveoli is in the Copenhagen
museum.
Gerrit S. Miller: A new River-Dolphin from China; Smithsonian
Misc. Coll., vol. 68, No. 9, 1918, pp. I-12, pls. 1-13. Lipotes.
Ischyrorhynchus was based by Ameghino (Caracteres diagnosticos
de cincuenta especies nuevas de Mamiferos fosiles argentinos ; Revista
Argentina de Historia Natural, vol. 1, 1891, pp. 163-165, with illus-
trations) on the anterior part of a lower jaw from the ‘
inferior del Parana.” The branches of the jaw, which are rather

* oligoceno

heavy, are grown together in a long symphysis menti; the rows of
teeth are further apart than in Saurodephis (Saurocetes), with which
Ameghino associates the genus, and the roots in cross-section are
almost what might be called ovate, not 8-shaped; the crowns are
known only imperfectly. It is not possible to decide where the genus
belongs.

Pontivaga was also based by Ameghino (J. c., pp. 165-166, with
illustration) on the anterior part of a lower jaw, from the “ oligoceno
superior ’ at Parana. The branches of the jaw are slender and are
grown together in a long symphysis. Each of them contains a long
row of small teeth, to judge by the alveoli. Ameghino places the
genus in the Platanistid@, in contrast with the Saurocetide, whether
rightly or not cannot be said.

A genus Proinia is established by True (A new genus of fossil
’Cetaceans from Santa Cruz Territory, Patagonia, etc.; Smithsonian
Mise. Coll., vol. 52, pp. 441-447, pl. 43, 1910), who regards it as a near
relative of Jnia. Perhaps True is right; but the material on which
the genus is based, a few fragments of a braincase and a cervical
vertebra all distorted by pressure, is so scanty that an error is very
possible.

Hesperocetus is established by True (A fossil toothed Cetacean
from California, etc.; Smithsonian Misc. Coll., vol. 60, No. 11, 1912,
pp. 1-7, pls. 1-2) on the strength of the anterior part of a long, nar-
row, lower jaw with long symphysis, with rows of rather strong,
conical, slightly hooked teeth with wrinkled enamel. The teeth are
placed rather wide apart and are separated by shallow depressions in
the margin of the jaw into which the tips of the upper teeth pre-

80 ; SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

sumably fitted. True refers it provisionally to the family Jniude
(=Platanistide), perhaps rightly ; there are other possibilities.

* (Pp. 26 and 31.) True (A Review of the Family Delphinide ;
Bull. U. S. Nat. Mus., No. 36, 1889, p. 10) believes he has observed a
peculiarity in the relationships of the pterygoid that should distinguish
Delphinapterus and Monodon from all other Delphinids and recall
the Platanistids: “ that in the narwhal and white whale the pterygoid
bones, instead of merely forming the walls of the posterior nares,
extend backward in the form of broad plates across the optic canal
and articulate with the squamosals.” But the case is different. We
have to do with the bones which lie in the outer wall of the air-sac
behind the palate. As may be seen in young or youngish skulls of
Delphinapterus and Monodon, the pterygoid shares in the formation
of the outer wall of the air-sac at the front only, in contrast with the
condition in Pontistes, Pontoporia and Platanista in which it, recall-
ing the Balenids and Physeterids, forms most of the outer wall (in
Inia the outer wall appears to be mostly membranaceous). As in
other Delphinids the palatine, frontal, ala magna, and squamosal all
share in bounding the outer side of the air-sac, each contributing its
section (special ossifications may also be present). In the Delphinids
under discussion the outer side of the air-sac is merely ossified more
extensively than elsewhere, a difference, however, which is one of
degree only.

“ (P. 35.) As reasons for believing that Neomeris and Phocena
among recent Odontoceti are the ones which stand nearest to Zeu-
glodon and Squalodon Abel (Dauphins Longirostres, 1901, p. 36)
mentions the following: (1) that they still have traces of “ l’ancienne
dentition hétérodonte,” (2) that teeth are still found (or more cor-
rectly may be found) in the intermaxillary, and that the intermaxillary
extends further forward than the maxillary, (3) that they still have
traces of “Varmure dermique,” (4) that the nostrils are not pushed
very far backward, and that therefore the parietal still extends up
back of the frontal. Against this view there are the following objec-
tions: (1) The form of the teeth in the two recent genera is not
primitive ; fan-like broadened crown and single root is not the form
of tooth that is found in the more primitive cetaceans of any kind.
Conical crown and a trace of double root, in most of the teeth, is the
transitional form between the tooth structure of the more primitive
and the less primitive cetaceans. Even the anterior teeth in the jaws
of Phocena may have fan-shaped crowns, where in the most primitive
whales they are unicuspid and conical. (2) Teeth in the inter-

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 81

maxillary, and a slightly projecting intermaxillary, may also be found

in Steno, Delphinus, and several other of the recent Delphinide.

(3) There is no evidence that cetaceans are descended from mammals

with bony armor ; the so-called dermal bones that were found on rare

occasions with remains of Zeuglodon and Delphinopsis are too doubt-
ful to prove anything; in the great majority of instances no dermal
armor has been found with bones of Zeuglodon and other Archeoceti.

The “ dermal armor,” that is, the small spots of more or less tubercu-

late, mosaic-like skin, in Neomeris and Phocena 1s assuredly a new

development. (See pp. 56-58.) (4) It cannot be correctly said that
the nostrils in Phocena and Neomeris lie relatively far forward. It
appears so merely because the anterior part of the face 1s somewhat
shortened and the, braincase is unusually large. In reality the nostrils
lie, with respect to the orbits and other surrounding parts, in the same
position as in most Delphinids. Neither can it be said that the
parietals are excluded from the upper side of the braincase to a less
degree than usual. As may be seen in the young skulls the parietals in

Phocena are widely separated by the large interposed interparietal

quite as usual in other Delphinids.

* (P. 35.) On the Delphinidz see especially:

Cuvier: Recherches sur les Ossemens fossiles, ed. 4, vol. 8, pt. 2.
1836, pp. 75-170, Atlas, pls. 222-224, with figures of skull and
some other skeletal parts, of most of the recent genera.

Schlegel: Beitrage zur Charakteristik der Cetaceen; Abhandlungen
aus dem Gebiete der Zoologie und vergl. Anatomie, Heft. 1, 1841,
pp. 1-44, pls. 1-6. Contains among other things a synopsis of the
Delphinids, with figures of skulls of Steno, Prodelphinus, Del-
phinus, Lagenorhynchus, all under the name Delphinus.

J. E. Gray: The Zoology of the Voyage of H. M. S. Erebus and
Terror, pts. 3-5, Mammalia, On the Cetaceous Animals, 1846,
pp. 13-53, pls. 1-30. Most of the plates give figures of skulls
of Delphinids: Delphinapterus (“ Beluga’), “ Feresa” (Orca
intermedia), Orca, Lagenorhynchus, Tursiops (under the name
Delphinus), Prodelphinus (under the name Delphinus), Delphi-
nus, Steno.

Burmeister: Descripcion de cuatro especies de Delfinides de la costa
Argentina en el océano Atlantico; Anales del Museo Publico de
Buenos Aires, vol. 1, 1864-69, pp. 367-388, pls. 21-24. Pseudorca
(under the name Globicephalus), Orca, Phocena. Figures of
exterior, skulls and other parts.

82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Owen: On some Indian Cetacea; Transact. Zool. Soc. London, vol.
6, pt. 1, 1866, pp. 17-47, pls. 3-14. Deals partly with the Del-
phinids. Skulls are figured of “ Sotalia,’ Lagenorhynchus, Del-
phinus, Prodelphinus (all called Delphinus), Orcella (called
Phocena).

J. E. Gray: Synopsis of the species of Whales and Dolphins in the
Collection of the British Museum, 1868, pp. I-10, pls. 1-30. The
plates are the same as in the previously mentioned work by Gray.

Van Beneden et Gervais: Ostéographie des Cétacés vivants et fossiles,
Text and Atlas, 1868-80, pp. 482-512, 521-605, pls. 34-60, 63, 64.
All recent genera and many fossil, among them Champsodelphis,
Schizodelphis, Eurhinodelphis.

Eschricht: Ni Tavler til Oplysning af Hvaldyrenes Bygning, med
Forklaring af Reinhardt; Kgl. Danske Vidensk. Selsk. Skrifter,
ste R., naturv. mathem. Afd., vol. 9, I, 1869. On plate 8 are
found figures of skull and teeth of Delphinapterus.

Flower: Descr. of the skeleton of the Chinese White Dolphin, Del-
phinus sinensis ; Transact. Zool. Soc. London, vol. 7, pt. 2, 1870
(72), pp. 151-160, pls. 17, 18. Prodelphinus (“Sotalia”).

Flower: On Risso’s Dolphin, Grampus griseus ; Transact. Zool. Soc.
London, vol. 8, -pt. 1, 1872, pp. 1-21, pls. 1-2. Figures of exterior
and skeleton.

Brandt: Untersuchungen uber die fossilen und subfossilen Cetaceen
Furopa’s ; Mém. Acad. Imp. Sci. St. Pétersbourg, ser. 7, vol. 20,
No. 1, 1873. Contains a section on the Delphinids, pp. 226-290,
pls. 24-30, among them Schizodelphis and Champsodelphis.

Brandt: Erganzugen zu den fossilen Cetaceen Europa’s ; Mém. Acad.
Imp: Sci. St. Pétersbourg, ser. 7, vol! 21, 1874; ppy i3=2epqme:
2-4. Champsodelphis among others.

Murie: On the organization of the Caaing Whale, Globiocephalus
melas ; Trans. Zool. Soc. London, vol. 8, pt. 4, 1873, pp. 235-301,
pls. 30-38. Ixterior and anatomy.

Van Beneden: Mémoire sur un Dauphin nouveau de la Baie de Rio
de Janeiro désigné sous le nom de Sotalia brasiliensis; Mém.
Acad. Roy. Belgique, vol. 41, 1874, pp. 1-44, pls. 1, 2, with
figures of exterior, skeleton, skull. Prodelphinus.

J. E. Gray: Feresa attenuata; Journal des Museum Godeffroy,
vol. 8, 1875, p. 1, pl. 6, with figures of skull.

Anderson: Anat. and Zool. Researches, compr. an Account of the
two Exp. to Western Yunnan in 1868 and 1875; 1878; pp. 358-
416, pls. 25, 25-A, 27-30, 33-38, 42, 43. Orcella. Figures of

exterior, skeleton and soft parts.

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 83

Van Beneden: Mémoire sur les Orques observés dans les mers
d’Europe; Mém. Acad. Roy. Belgique, vol. 43, 1879, pp. 1-33,
pls. 1-4, with figures of exterior, skeleton, skulls. Orca.

Flower: On the characters and divisions of the family Delphinide ;
Proceed. Zool. Soc. London, 1883, pp. 466-513, with figures in
the text representing the posterior part of the bony palate in
several of the genera. One of the most important papers in
elucidating the relationship among the recent Delphinids.

Lutken: Kritiske Studier over nogle Tandhvaler af Slegterne Tur-
siops, Orca og Lagenorhynchus; Kgl. Dansk Vidensk. Selsk.
Skrifter, 6te R., naturv. mathem. Afd., vols. 4, 6, 1887, pp. 337-
397, pls. I, 2, also text figures: exterior, skull, other skeletal
parts.

Lutken: Spolia Atlantica, Bidrag til Kundskab om de tre pelagiske
Tandhval-Slegter Steno, Delphinus og Prodelphinus; Kgl.
Danske Vidensk. Selsk. Skrifter, 6te R., naturv. mathem. Afd.,
vol. 5, 1, 1889, pp. 1-61, plate with a figure of the exterior and
skeleton of Steno; also figures in the text: exterior, skulls, other
skeletal parts.

True: A Review of the Family Delphinide; Bull. U. S. Nat. Mus.,
No. 36, 1889, pp. 1-191, pls. 1-47, with figures of exterior and
skulls. In the conception of genera and their mutual relation-
ships agrees closely with Flower’s conclusions.

Lydekker: Cetacean skulls from Patagonia; Anales del Museo de
La Plata, Paleontologia Argentina, vol. 2, 1893, pp. 10-12, pl. 5.
Argyrocetus.

Guldberg: On the development and structure of the Whale, pt. 1, on
the development of the Dolphin; Bergens Museums Skrifter,
vol. 5, 1894, pp. 1-70, pls. 1-7. Lagenorhynchus, Phocena, Orca.

Longhi: Sopra i resti di un cranio di Champsodelphis fossile scoperto
nella molassa miocenica del Bellunese ; Atti della Societa Veneto-
Trentina di Scienze Naturali, ser. 2, vol. 3, fasc. 2, 1898, pp. 1-60
in the separate, pls. 1-3.

Abel: Untersuchungen uber die fossilen Platanistiden des Wiener
Beckens ; Denkschr. d. k. Akad. Wissensch. Wien, math.-naturw.
Cl., vol. 68, 1899, pp. 839-874, pls. 1-4, with figures of skulls.
Cyrtodelphis and Acrodelphis=Schizodelphis and Champso-
delphis.

Abel: Les Dauphins Longirostres du Boldérien (Miocéne supérieur )
des Environs d’Anvers; Mém. Mus. Roy. d’Hist. Nat. de

6

84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Belgique, vol. 1, 1901, pp. 1-95, pls. 1-10, with figures of skulls.
Cyrtodelphis (=Schizodelphis), Eurhinodelphus.

Dal Piaz: Sugli avanzi di Cyrtodelphis sulcatus dell’, arenaria di
Belluno; Palaeontographia Italica, vol. 9, 1903, pp. 187-220,
pls. 28-31, with figures of skulls and teeth. Schizodelphis.

Abel: Eine Stammtype der Delphiniden aus dem Miocan der Hal-
binsel Taman; Jahrbuch der k. k. geol. Reichsanstalt, vol. 55,
pt. 2, 1905, pp. 375-392, with text figures. Paleophocena.

Abel: Les Odontocétes du Boldérien d’Anvers; Mém. Mus. Roy.
d’Hist. Nat. de Belgique, vol. 3, 1905, pp. 1-155, with text figures.
Includes a section on the Delphinids: Eurhinodelphis, Cyrtodel-
phis (=Schizodelphis), Acrodelphis (=Champsodelphis) , Pro-
tophocena, Pithanodelphis.

C. v. Papp: Heterodelphis leiodontus, nova forma aus den miocenen
Schichten des Comitates Sopron in Ungarn; Mitteilungen aus
dem Jahrbuche der k. ungarischen geol. Anstalt, vol. 14, pt. 2,
1905, pp. 25-61, pls, 5, 6, also text figures. Skeleton.

Abel: . Cetaceenstudien, I, Das Skelett von Eurhinodelphis coche-
teuxi aus dem Obermiozan von Antwerpen ; Sitzungsber. k. Akad.
Wissensch. Wien, mathem.-naturw. Kl., vol. 118, pt. I, 1909,
pp. 241-253, pl. 1, with a figure of the skeleton, partly conjectural.

True: Observations on living White Whales, Delphinapterus leucas,
with a note on the dentition of Delphinapterus and Stenodelphis ;
Smithsonian Misc. Coll., vol. 52, pt. 3, No. 1864, 1909, pp. 325-
330, pl. 23, with a figure of the exterior. Discusses among other
things the projections on the tooth crowns of Delphinapterus.

Lonnberg: Remarks on the dentition of Delphinapterus leucas ; Arkiv
for Zoologi, vol. 7, No. 2, 1910, pp. 1-18, with illustrations.
Taken up in part also in the paper Om Hvalarnes Harstamming ;
K. Svenska Vetenskapsakademiens Arsbok for Ar 1910, pp. 219-
259, with illustrations.

Roy C. Andrews: A new Porpoise from Japan; Bull. Amer. Mus.
Nat. Hist., vol. 30, I91I, pp. 31-51, pls. 1, 2, also numerous
figures in the text. Phocenoides. Exterior and skeleton.

Bassani e Misuri: Sopra un Delfinorinco del calcare miocenico di
Lecce (Ziphiodelphis Abeli Dal Piaz) ; Atti della R. Accademia
dei Lincei, Anno 309, 1912, ser. 5, Memorie della Classe di
Scienze Fisiche, etc., vol. 9, fasc. 2, pp. 25-38, pl. 1, with figures
of skull.

True: Descr. of a new fossil Porpoise of the genus Delphinodon
from the miocene formation of Maryland; Journ. Acad. Nat.

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 85

Sci. Philadelphia, ser. 2, vol. 15, 1912, pp. 165-194, pls. 17-26,
with figures of skull, parts of the rest of the skeleton, teeth.
Lull: Fossil Dolphin from California; American Journal of Science,
ser. 4, vol. 37, 1914, pp. 209-220, pl. 8, also figures in the text.

“ Delphinavus.”

Delphinopsis (see note 8) is placed by Abel (Jahrb. k. k. geol.
Reichanst., vol. 55, pt. 2, 1905, pp. 384; 387, in the “Subfamily
Phocenine ”’ because it has “ dermal armature.’’ The remains are so
imperfect and so uncertain that it is impossible to say where it be-
longs ; not even the family can be determined from the specimen; the
reference to Phocenine is pure guesswork.

Rhabdosteus was described in 1867 by Cope, who in 1890 (Amer.
Nat., vol. 24, p. 607) gave figures of the specimens on which the
genus was based, some remnants of a “ beak,” from a Tertiary North
American deposit. These remains Cope reconstructed in a somewhat
arbitrary manner. True (Remarks on the fossil Cetacean Rhab-
‘dosteus latiradix Cope; Proc. Acad. Nat. Sci., Philadelphia, vol. 60,
1908, pp. 24-29, pl. 6, and text figures), who has had the specimens in
question under revision, together with some others more or less
similar, says that Cope has scarcely put them together right. The
specimens may recall Eurhinodelphis and its relatives ; but the remains
are altogether too incomplete and uncertain for anything to be decided.

Lophocetus, established by Cope in 1867, best known from East-
man’s description (Types of fossil Cetaceans in the Museum of Com-
parative Zoology ; Bull. Mus. Comp. Zool., vol. 51, 1907, pp. 79-94,
pls. 1-4), Tertiary, North American, is most often placed in the
Platanistide. Brandt, however, counts it as a Delphinid (1873, 1. c.,
p. 288), most probably belonging to the “ Abtheilung der Phoczenen,”
perhaps to the genus Delphinapterus. In this determination he has
been followed by a few other authors. The most important basis of
the genus is a very imperfect skull, without teeth, with alveoli only,
so obscure that nothing positive can be said about it. According to
what can be seen of the form of the temporal fossa the genus appears
to agree best with the Delphinids. On the other hand it does not
seem possible to demonstrate anything that would especially recall
the Platanistids.

Imopsis was established by Lydekker (Proc. Zool. Soc. London,
1892, pp. 562-564, pls. 37-38) principally on an imperfect and obscure
braincase from a Tertiary deposit in the Caucasus. Lydekker places
it in the Platanistide and finds similarities with Pontistes, Stenodel-
phis (=Pontoporia), Inia, etc. It appears, however, to be of another

86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOR 72

type, very near the most usual Delphinid-type, differing from the
Platanistide especially in the roofed over temporal fossa. Its more
exact position among the Delphinids cannot yet be determined.

Cyrtodelphis is only a new name for Schizodelphis given by Abel
in 1899 (J. c.) to include a series of species which previously were
most often referred to Schizodelphis, among them the type of the
genus, S. sulcatus Gervais. Eastman (Bull. Mus. Comp. Zool., vol.
51, 1907, pp. 83-84) has already protested against this superfluous
new name as well as against the following.

Acrodelphis is likewise essentially a mere new name, a synonym of
Champsodelphis. It was given by Abel in 1899 (J. c.). At first
Acrodelphis was to include the type of Champsodelphis, Ch. macro-
genius (Laurillard) Gervais or macrognathus Brandt. Later, in 1905,
Abel excluded the type of Champsodelphis from the genus, with
doubtful right; but most of the species which he now includes in
Acrodelphis were earlier called Champsodelphis.

Paleophocena was based by Abel (1905, Jahrb. k. k. geol. Reich-
sanst., vol. 55, J. c.) on an imperfect piece of a braincase and a few
fragments of the rest of the skeleton from a Tertiary deposit on the
coast of Crimea. Abel considers it proved that this is a near relative
of Phocena. Possibly it will sometime turn out that he is right; but
for the present there is no means of deciding the question about
nearest relationship. The known piece of skull shows only such
general features that nothing exact can be said except that it comes
from a Delphinid. Only in the form of the teeth have Phocena and
its relative Neomeris a peculiarity which distinguishes them from
other quite ordinarily formed Delphinids ; but the teeth in Paleopho-
cena are not known.

Protophocena is also established by Abel (Odontocétes du Bol-
derien, 1905, pp. 139-141, with illustrations), on the anterior, very
imperfect part of a skull, without teeth, from the Tertiary deposits
at Antwerp. Abel refers it to the “ Phocenine.’ There is actually
nothing whatever, apart- from the small size, that could lead one to
think of Phocena,; on the contrary, the strong cushion-shaped swell-
ing and the widening out which the intermaxillary shows in front of
the nasal aperture suggests rather Lagenorhynchus or “ Grampus.”
For the present the question about nearest relationship cannot be
settled.

Pithanodelphis is established by Abel (Odontocétes du Boldérien,
1905, pp. 142-145, with illustrations) on the basis of Phocenopsis
cornutus du Bus from Tertiary strata at Antwerp. Abel refers it to

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 87

the “Delphinine.’ That which is known of it is an imperfect piece
of a braincase and a few other parts. The characters, so far as they
go, agree well with the ordinary dolphin type; the feature which
especially distinguishes it is that the maxillary posteriorly is bowed
inward unusually strongly behind the nasal. In this respect, how-
ever, dolphins show great variation. The more exact position of the
genus cannot be determined.

Phocenoides is established by Roy Andrews (1911, lJ. c.) to include
two recent species, one a new species, Ph. truei from Japan, the other
a species which True had called Phocena dalli, likewise from the
northern part of the Pacific Ocean. The deviations from typical
Phocena are very small; perhaps the most noticeable is that the teeth
are smaller, with the fan-like widening of the crown less pronounced.
There can scarcely be sufficient ground for generic separation.

Xiphiodelphis (“ Ziphiodelphis”’) (see especially Bassani e Misuri
and Dal Piaz, 1912, J. c.) is established on fragments of skulls from
Tertiary Italian deposits. There can be no douht that it is a near
relative of Schizodelphis, etc., but its more exact position is not yet
clear.

Delphinavus is established by Lull (1914, J. c.) on an imperfect
and compressed, indistinct skeleton from a no doubt Miocene deposit
in California. The genus is supposed to stand very near to Delphinus
in the narrow sense. The form of the palate, however, the only
character that distinguishes Delphinus from nearly related Delphinids,
does not seem to have been ascertained. One of the most important
peculiarities is that the atlas and axis are mutually free. According
to what is known it is not possible to clear up the relationship of the
genus to other Delphinids; but it ought to be especially compared
with Heterodelphis.

* (P. 38.) It is Flower who has especially emphasized the differ-
ence between Delphinids and Physeterids with regard to the relation-
ship of the hindmost ribs to the vertebree. It is likewise he, in his
paper on Jmia (Trans. Zool. Soc. London, vol. 6, 1869, pp. 98-100)
and elsewhere, who has pointed out the intermediate position of the
Platanistids. The question about the interpretation of the transverse
processes, etc., had previously been discussed, among others by Esch-
richt in his paper on Platanista (1851, pp. 369-370). Later it has
been extensively dealt with by Gerstaecker (Das Skelet des Déglings,
Hyperoodon rostratus, etc., 1887) and it is also taken up by Abel
(Sitzungsber k. Akad. Wissensch. Wien, math.-naturwiss. K1., vol.

118, pt. I, 1909, pp. 247-249).

88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

*(P. 43.) On the Physeteride see especially :

Cuvier: Recherches sur les Ossemens fossiles, ed. 4, vol. 8, pt. 2,
1836, pp. 117-247, Atlas, pls. 225, 228, with figures of skulls of
Physeter and Hyperoodon and parts of fossil skulls of Meso-
plodon, Chonoxiphius and Xiphius (all under the name
“ Ziphius”’).

Eschricht: Unders¢gelser over Hvaldyrene, 4de Afhandl., Om
Nebhvalen; Kgl. Danske Vidensk. Selsk. naturv. mathem.
Afhandl. 11te Del, 1845, pp. 321-378, pls. 5-8, with figures,
mostly of soft parts. Hyperoodon. On the dentition in the
embryo and other things.

Owen: On some Indian Cetacea; Transact. Zool. Soc. London, vol.
6, pt. 1, 1866, pp. 17-47, pls. 3-14. Contains a section on
“Cogia” under the name Physeter (Euphysetes). Exterior,
skeleton and skull are figured.

Fischer: Mémoire sur les Cétacés du genre Ziphius; Nouvelles
Archives du Muséum d’Hist. Nat. de Paris, vol. 3, 1867, pp. 41-
79, pl. 4, with figures of skulls of Xiphius cavirostris.

Van Beneden et Gervais: Ostéographie des Cétacés vivants et fossiles,
Text and Atlas, 1868-80, pp. 303-324, 514-518, pls. 18-27 Dis,
61-63. All the recent genera and most of the fossils, anione
them Xiphirostrum, Chonoxiphius, Hoplocetus.

Burmeister: Descripcion detallada del Epiodon australe (sic) ; Anales
del Museo Publico de Buenos Aires, entrega quinta, 1868, pp.
312-366, pls. 15-20. Xiphius. Figures of exterior, skeleton,
skull, and other parts.

Eschricht: Ni Tavler til Oplysning af Hvaldyrenes Bygning. med
Forklaring af Reinhardt ; Kgl. Danske Vidensk. Selsk. Skrifter,
ste R., naturv. mathem. Afd., vol. 9, 1, 1869. On plates 6 and
7 are found figures of the skull of adult Hyperoodon and of the
exterior and skeleton of the fetus.

Flower: On the osteology of the Cachalot or Sperm-whale (Physeter
macrocephalus) ; Trans. Zool. Soc. London, vol. 6, 1869, pp.
309-372, pls. 55-61.

Owen: Monogr. of the British fossil Cetacea of the Red Crag;
Paleontogr. Society, vol. for 1869, 1870, pp. 1-40, pls. 1-5, and
with figures in the text. Mostly on the skull and “beak” of
“ Ziphius” (=Xiphius, Chonoxiphius, Mesoplodon, “ Ber-
ardius”’).

Flower: On the recent Ziphioid Whales, with a descr. of the Skele-
ton of Berardius arnouxi; Trans. Zool. Soc. London, vol. 8, pt.

3, 1872, pp. 203-234, pls. 27-29.

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE &9O

Turner: On the occurrence of Ziphius cavirostris in the Shetland
Seas and a comparison of its skull with that of Sowerby’s Whale,
Mesoplodon sowerbyi; Trans. Roy. Soc. Edinburgh, vol. 26, 1872,
pp. 759-780, pls. 29, 30, with figures of skulls.

Brandt: Untersuchungen uber die fossilen und subfossilen Cetaceen
Europa’s; Mém. Acad. Imp. Sci. St. Pétersbourg, ser. 7, vol. 20,
No. 1, 1873. <A section, pp. 204-226, gives a synopsis of the
then-known fossil Physeterids.

Flower: A further contrib. to the knowledge of the existing Ziphioid
Whales, genus Mesoplodon; Trans. Zool. Soc. London, vol. 10,
pt. 9, No. 2, 1878, pp. 415-437, pls. 71-73, with figures of skulls
and skeleton.

Turner: Report on the bones of Cetacea collected during the voyage
of H. M. S. Challenger in the years 1873-1876; Report on the
Scientific Results of the voyage of H. M. S. Challenger, Zoology,
vol. 1, pt. 4, 1880, pp. 1-45, pls. 1, 2. Mostly on Mesoplodon.
Figures of skulls and other parts.

De Sanctis: Monografia zootomico-zoologica sul Capidoglio arenato
a Porto S. Giorgio; Atti della R. Accademia dei Lincei, Mem.
Cl. sc. fisiche, ser. 3, vol. 9, 1881, pp. 160-242, pls. 1-7. Physeter.
Exterior and viscera.

Capellini: Resti fossili di Dioplodon e Mesoplodon; Memorie della
R. Accademia delle Scienze dell’ Instituto di Bologna, ser. 4,
vol. 6, 1885, pp. 291-306, pl. 1, with figures of “‘ beak ” and other
parts. Mesoplodon.

Capellini: Del Ziphioide fossile (Choneziphius planirostris) scoperto
nelle sabbie plioceniche di Fangonero presso Siena; Atti della
R. Accademia dei Lincei, Mem. C1. sc. fisiche, ser. 4, vol. 1, 1885,
pp. 18-29, pl. 1, with figures of the skull. Chonoxiphius.

Malm: Om Sowerby’s hval ; Ofversigt af kgl. Svenska Vetensk.-Akad.
Forhandlingar, 1885, No. 5, pp. 121-153, pl. 9, with figures of
the skull and other parts. Mesoplodon.

Gerstaecker: Das Skelet des Doglings, Hyperoodon rostratus, ein
Beitrag zur Osteologie der Cetaceen und zur vergleichenden
Morphologie der Wirbelsaule, 1887, pp. 1-175, pl. 1, with figures
of the vertebre.

Pouchet et Beauregard: Recherches sur le Cachalot; Nouvelles
Archives du Muséum d’Hist. Nat. de Paris, ser. 3, Mémoires,
vol. 1, 1889, pp. 1-96, pls. 1-8, with figures of the exterior,
skeletal parts, teeth and their development. Physeter. Suite,
Mém., vol. 4, 1892, pp. 1-90, pls. I-12, with figures of the
exterior, viscera, etc.

go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Moreno: Lijeros apuntes sobre dos géneros de Cetaceos fosiles de la
Reptblica Argentina; Revista del Museo de La Plata, vol. 3,
1892, pp. 13-20, in the separate, pls. 10, 11. Mesocetus=Hypo-
cetus, Notocetus= Argyrodelphis.

Forbes: Observ. on the development of the rostrum in the Cetacean
genus Mesoplodon, with remarks on some of the species; Pro-
ceed. Zool. Soc. London, 1893, pp. 216-236, pls. 12-15.

Lydekker: Cetacean skulls from Patagonia; Anales del Museo de
La Plata; Paleontologia Argentina, vol. 2, 1893, pp. 4-8, 12-13,
pls. 2, 3, 6. Physodon (=Hoplocetus), Hypocetus, Argyro-
delphis. .

Moreno: Nota Sobre los restos de Hyperoodontes conservados en el
Museo de La Plata; Anales del Museo de La Plata, Seccion
Zoologica, vol. 3, 1895, pp. 1-8, pls. 1, 2. Hyperoodon. Figures
of entire skeletons and of skulls.

Benham: On the anatomy of Cogia breviceps; Proc. Zool. Soc.
London, 1901, vol. 2, pp. 107-134, pls. 8-11. See also, on the
larynx, Proc. Zool. Soc. London, 1901, vol. 1, pp. 278-300,
pls. 25-28.

Benham: Notes on the osteology of the Short-nosed Sperm-Whale ;
Proc. Zool. Soc. London, 1902, vol. I, pp. 54-62, pls. 2-4.
“ Cogiax

Grieg: Bidrag til kjendskab om Mesoplodon bidens; Bergens
Museums Aarbog, 1904, No. 3, with figures in the text. Exterior,
skull and various skeletal parts.

Abel: Die phylogenetische Entwicklung des Cetaceengebisses und
die systematische Stellung der Physeteriden; Verhandl. d.
Deutsch. Zool. Gesellschaft, 1905, pp. 84-96.

Abel: Les Odontoceétes du Boldérien (Miocene supérieur) d’Anvers ;
Mém. Mus. Roy. d’Hist. Nat. de Belgique, vol. 3, 1905, pp. 1-155,
with figures in the text. Deals largely with the Physeterids,
especially with Scaldicetus (=Hoplocetus), Thalassocetus, Phy-
seterula, Prophyseter, “ Placoziphius,’ “ Paleoziphius,’ Ceto-
rhynchus, “ Mioziphius” (=Xiphirostrum), “ Choneziphius,”
Mesoplodon.

Danois: Recherches sur l’anatomie de la tete de Kogia breviceps
Blainv. ; Archives de Zoologie expérimentale et générale, ser. 5,
vol. 6, I910, pp. 149-174, pls. 5-8, and with text figures.

Trtie: Descr. of a skull and some vertebre of the fossil Cetacean
Diochoticus Vanbenedeni from Santa Cruz, Patagonia; Bull.
Amer. Mus. Nat. Hist., vol. 28, 1910, pp. 19-32, pls. 1-5. Argyro-
delphis.

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE gti

True: An account of the Beaked Whales of the family Ziphiide in
the Collection of the United States National Museum, with re-
marks on some specimens in other American Museums; Bull.
U.S. Nat. Mus., No. 73, 1910, pp. 1-89, pls. 1-42. Mesoplodon,
“ Ziphius,’ “ Berardius,’ Hyperoodon. Figures of skulls and
skeletal parts.

Danois: Recherches sur les visceres et le squelette de Kogia brevi-
ceps Blainv. avec un résumé de Vhistoire de ce Cétacé; Arch.
Zool. expér. et génér., ser. 5, vol. 6, 1911, pp. 465-489, pls. 23, 24.

[Schulte: The skull of Kogia breviceps Blainv.; Bull. Amer. Mus.
Nat. Hist., vol. 37, pp. 361-404, pls. 35-43. June 28, 1917. ]

[Schulte and Smith: The external characters, skeletal muscles, and
peripheral nerves of Kogia breviceps (Blainville) ; Bull. Amer.
Mus. Nat. Hist., vol. 38, pp. 7-72, figs. 1-21. February 23, 1918. ]

[Kernan and Schulte: Memoranda upon the anatomy of the respira-
tory tract, foregut, and thoracic viscera of a foetal Kogia
breviceps; Bull. Amer. Mus. Nat. Hist., vol. 38, pp. 231-267,
figs. 1-16. April 18, 1918. ]

[Kernan: The skull of Ziphius cavirostris; Bull. Amer. Mus. Nat.
Hist., vol. 38, pp. 349-394, pls. 20-32. August 1, 1918. ]

Cetorhynchus was established by Gervais on remains from Tertiary

strata in southern France. The most important fragment was the
anterior portion of an under jaw (figured in Ostéographie des Cétacés,
pl. 57, fig. 12). The genus is discussed by Abel (Odontocetes du
Boldérien, 1905, pp. 94-98), who refers to it a piece of a lower jaw
from the Tertiary deposits at Antwerp. The mandible has a long
symphysis menti and a long row of close-placed alveoli for rather
large teeth. It is a peculiar fact that the alveoli are not completely
separated from each other. Only low transverse ridges separate the
teeth at their bases ; otherwise the teeth lay in a common groove. Abel
thinks that he sees in these conditions a beginning to the peculiarities
of the “ Ziphius” group. Perhaps he is right, but there are still
other possibilities.

Anoplonassa probably belongs to the group Xiphii, quite likely as

a near relative of Xiphirostrum. It was described by Cope (Proc.

Amer. Philos. Soc., vol. 11, 1871, pp. 188-190, pl. 5, fig. 5) on the

basis of the anterior part of the mandible from Tertiary deposits at

Savannah, Georgia; but it is best known from a paper by True

(Observations on the Type specimen of the fossil Cetacean Anoplo-

nassa forcipata Cope; Bull. Mus. Comp. Zool., vol. 51, 1907, pp. 97-

106, pls. 1-3). The fragment in question shows the mandibular rami

Q2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

grown together in a very long symphysis menti. They are staff-
shaped, each with a cup-like pit left by a large tooth at the very front,
a somewhat indistinct alveolus further back, and also with more or
less indistinct traces of other teeth in a degenerate dental groove.
Abel (Odontoceétes du Boldérien, 1905, p. 92) compares Anoplonassa
with Paleoziphius; but True is no doubt right in finding a greater
likeness to “ Mioziphius” (=Xiphirostrum). However, Anoplo-
nassa is still too slightly known to be exactly placed.

“ Paleoziphius” is established by Abel (Odontocetes du Boldérien,
1905, pp. 90-94, with figure) on the basis of a piece of the anterior
end of a lower jaw from the Tertiary at Antwerp. The specimen had
previously been referred by others to Champsodelphis and by Abel
himself doubtfully to Acrodelphis (=Champsodelphis). The jaw
has a long symphysis menti and a long series of alveoli left by good-
sized teeth. Abel says of it with great positiveness that it belonged to
a member of the family “ Ziphiide”’ (a group that about corresponds
to the Xiphiini of the present paper; it was, he thinks, one of the
first links in the series that leads from the oldest, many-toothed
Ziphiids to the living two-toothed forms. His reason for believing
this is that he finds the first and seventh alveoli larger than the others,
a condition that he considers a first beginning of the condition found
in the recent genera of the group. But in the photograph of the jaw
it is impossible to see this difference in the alveoli. There is the
greatest possibility of a mistake; and it cannot be asserted with any
positiveness where the genus belongs.

“ Placoziphius” is established by Van Beneden on the basis of
pieces of a skull from the Tertiary deposits at Antwerp (figured in
the Ostéographie des Cétacés, pl. 27, fig. 11). It is discussed by Abel
(Odontocetes du Boldérien, 1905, pp. 85-88), who considers it a near
relative of Physeter. In this he is no doubt right, but any final
decision is still impossible.

Hypocetus was described as a special genus by Moreno (1892,
I. c.) under the name Mesocetus Moreno (nec Van Beneden). It was
called Hypocetus by Lydekker (1893, /. c., in title and in explanation
of plates, Paracetus in text). From Ameghino (Enumération syn-
optique des especes de Mammiféres fossiles de Patagonie, 1894, p.
181) it received the name Diaphorocetus. It is based on a much-
broken skull from the Tertiary of Patagonia. Of the genus it can
be said that it no doubt belongs to the section Physeterini as a rather
near relative of Hoplocetus, but a more exact opinion is scarcely
possible.

no. 8 INTERRELATIONSHIPS OF THE CETACEA—WINGE 93

Thalassocetus is based by Abel (Odontocétes du Boldérien, 1905.
pp. 70-74, with figures) on a few pieces of forehead of skulls from
the Tertiary deposits at Antwerp. Abel is no doubt right in consider-
ing it a near relative of Hoplocetus (“ Scaldicetus’’) ; but the genus
is too slightly known to be definitely placed.

Prophyseter is based by Abel (Odontocétes du Boldérien, 1905,
pp. 82-85, with figures) on very imperfect remains from the Tertiary
deposits at Antwerp. If the interpretation of the bones is right
(whereof, according to the photographs, there seems to be some
reason for doubt) the remains represent two pieces (perhaps belong-
ing together) of the left side of a snout-tip, a piece of intermaxillary
and a piece of maxillary, both with alveoli but no teeth. Abel believes
that Prophyseter was a relative of Hoplocetus (“ Scaldicetus”), but
that it had gone a step further in the direction of Physeter, since the
upper teeth had begun to degenerate. This refers to the fact that the
alveoli in the intermaxillary appear to be in course of obliteration
after the disappearance of the teeth. With regard to this there might
be other explanations also. The specimens are too doubtful for any-
thing final to be said about the animal’s relationship.

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INDEX

Acrodelphide 61 Classifications, most important former
Acrodelphis 30, 86 58

Agabelus 60 Clymenia 59

Agorophiide 61 Cogia 42, 44

Agorophius 23, 24, 70 Creodontia 61

Agriocetus 70 Cyrtodelphis 30, 86

Alabamornis gigantea 63 Delphinapteridz 60

Amphicetus 69 Delphinapterus 31, 35

Anoplonassa 91 Delphinavus 87

Archaeoceti 10, 15 Delphinide 11, 28, 35, 46, 81

Delphini 31, 36
Delphinoceti 61
Delphinodon 30, 35
Delphinoidea 59
Delphinopsis 57, 85
Delphinus 32, 36
Dermal armor, supposed traces of 56
Diaphorocetus 92
Diochoticus 38
Dioplodon 59
Diphyletic origin 48
Doryodon 62
Dorudon 62

Eocetus 65
Eubalaena 59
Eucetotherium 69
Euphysetes 88
Eurhinodelphide 61
Eurhinodelphini 31, 35
Eurhinodelphis 31, 35
Feresa 33
[Globicephala] 34, 37
Globicephalus 59
Globiceps 34, 37
Globicipites 33, 36
Grampus 34, 30
Hand, structure of 49
Herpetocetus 69
Hesperocetus 79
Heterocetus 69
-Heterodelphis 30, 35
Hoploceti 41, 44

Argyrocetus 31, 35
Argyrodelphini 38, 43
Argyrodelphis 38, 43
Asymmetry of skull 72
Aulocetus 69
Balena 18, 21
Balenidz 10, 16, 21, 45, 65
Balznini 18, 21
Balznodon 42
Baleznoidea 59
Balenoptera 21, 22
Balenopteride 61
Balzenopterini 20, 22
[Basilosauride] 11, 15, 63
[Basilosaurus] 14, 15
Beluga 59
Beluginae 59
Berardius 41, 44
Burtinopsis 69
Cephalorhynchus 33
Cephalotropis 70
Cetacea, classification of 58
origin of 47
Cetorhynchus 91
Cetotheriomorphus 69
Cetotheriophanes 69
Cetotheriopsis 69
Cetotherium 21, 22
Champsodelphis 30, 35
ombonii 30
Choneziphius 40
Chonoxiphius 40, 44

95

96 INDEX

Hoplocetus 42, 44 Paleopontoporia 60
Hyznodon 1, 12 Paleoziphius 92
- Hyznodontide 1, 12 Paracetus 92
Hyperoodon 41, 44 Parasqualodon 77
Hyperoodontes 40, 44 Patriocetidz 61
Hypocetus 92 f Patriocetus 70
Idiocetus 69 Phalanges, increased number of 49
Inia 26, 28 Phoczena 35, 37
Iniidz 80 Phocenz 34, 37
Iniinae 59 Phocenidze 61
Iniopsis 85 Phocznoides 87
Ischyrorhynchus 79 Phocenopsis 86
Isocetus 69 Physalus 59
Ixacanthus 60 Physeter 42, 44
Kekenodon 64 Physeteres 42, 44
Kogia 59 Physeteridz 11, 37, 43, 46
Lagenorhynchi 33, 36 Physeterini 41, 44
Lagenorhynchus 33, 36 Physeterula 42, 44
Lipotes 26, 28 Physetodon 115
[Lissodelphis] 33, 36 Physodon 42
Lophocetus 85 Pithanodelphis 86
Megaptera 21, 22 Placoziphius 92
Megapteropsis 69 Platanista 27, 28
Mesocetus 65, 69, 92 Platanistide 11, 25, 28, 46
Mesoplodon 30, 44 Platanistine 59
Mesoteras 69 Plesiocetopsis 69
Metasqualodon 77 Plesiocetus 20, 22
Metopocetus 69 Pontistes 26, 28
Micropteron 59 Pontivaga 79
Microsqualodon 77 Pontoporia 26, 28
Microzeuglodon 64 Pontoplanodes 27
Microzeuglodontidz 61 Preepollex 65
Mioziphius 40 Priscodelphinus 60
Monodon 31, 35 Prodelphinus 32, 36
Monodontes 31, 35 Proinia 79
Mystacoceti 10, 16 Prophyseter 93
Neobalzna 18, 21 Prosqualodon 24
Neomeris 35, 37 Protocetus 11, 15
Neophocena 35, 37 atavus 48
Neosqualodon 24 Protophocena 86
Notocetus 38 Prozeuglodon 13, 15, 62
Odontoceti 10 atrox 13
Orca 33, 36 Pseudorca 34, 37
Orcella 34, 36 Pterodon 1, 12
[Orcinus] 33, 36 : Pterygoid bone 80
Origin of the Cetacea 47 Rhabdosteus 85
Pachycetus 60 Rhachianectidz 60

Paleophoczena 86 Rhachionectes 20, 22

Rhegnopsis 70
Rhizoprion 75
Sagmatias 33
Saurocetide 709
Saurocetus 27
Saurodelphis 27, 28
Scaldicetus 42
Schizodelphis 30, 35
sulcatus 30
Sibbaldius 59
Siphonocetus 69
Sotalia 32
Squaloceti 61
Squalodon 23, 24
Squalodontidz 10, 22, 24, 45, 75
Steno 32, 36
Stenodelphis 26, 28
Stypolophus 12
Teeth, increased number of 50
Thalassocetus 93
Tretulias 69-70

INDEX

Tursio 33, 360
Tursiops 33, 36
Ulias 69
Xiphii 39, 44
Xiphiini 38, 43
Xiphirostrum 40, 44
Xiphius 40, 44
Xiphodelphis 87
Zeuglodon 14, 15
brachyspondylus minor 62
caucasicus 64
cetoides 14
isis 14
osiris 13, 62
pelvis of 62
Zeuglodontide 10, II, 15, 45, 63
Ziphiide 61
Ziphirostrum 40
Ziphius 40
Ziphodelphis 87
| Zygorhiza 62

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 72, NUMBER 9

NEOABBOTTIA, A NEW CACTUS GENUS
FROM HISPANIOLA

WitH Four PLates

BY
N. L. BRITTON AND J. N. ROSE

(PUBLICATION 2651)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JUNE 15, 1921

at

The Lord Baltimore Press

BALTIMORE, MD., U. 8. A.

<

NEOABBOTTIA, A NEW CACTUS GENUS FROM
HISPANIOLA

By N. L. BRITTON anp J. N. ROSE
(With Four Pirates)

Among the earliest cacti described were those obtained by Plumier,
more than two hundred years ago, mostly from the island of His-
paniola, better known as Santo Domingo. These were characterized
briefly by him in 1704 and his illustrations of them were published
in 1755 by Burmann; Lamarck gave most of them binary names in
1783 under the genus Cactus, and in 1828 De Candolle referred all
the cereoid forms to the genus Cereus; the other species described
by Plumier have been referred by various authors to Mammuillaria,
Cephalocereus, Pilocereus, Rhipsalis, Melocactus, Pereskia, Opuntia
and Nopalea. We have experienced great difficulty in definitely
identifying the plants from the illustrations of Plumier, since these
are largely diagrammatic. As the type locality is generally given,
however, the identification of all of them may eventually be made
fairly definite.

In 1920 when Dr. W. L. Abbott and Mr. E. C. Leonard were start-
ing for Haiti, we asked them to collect both living and herbarium
specimens of all the cacti met with. As good fortune directed, they
spent a considerable time on the Cul-de-sac, where Plumier collected,
and so probably obtained several of the species which he described.
About 20 species of cacti were observed by them there. One of
these, which has proved to be an undescribed genus, is the subject
of this article. The Cul-de-sac is the bottom of an old salt lagoon,
which now has an altitude of 20 feet or more above the sea. It is
a coral formation and an ideal habitat for many kinds of cacti. Here
they appear in thickets or literally as forests, forming the dominant
feature of the landscape. This region lies north and east of Port-
au-Prince, extending from the bay of Port-au-Prince to Lake
Saumatre.

After the return of Dr. Abbott and Mr. Leonard in September
of the same year with these valuable specimens we wrote, at the
suggestion of Mr. Leonard, to Mr. H. M. Pilkington, an American
business man then stationed at Port-au-Prince, asking him to pro-

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 72, No. 9

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

cure additional material. In the latter part of December, 1920, Mr.
Pilkington returned to New York and brought with him two large
boxes of plants, containing two sections of the trunk, several living
plants, and fruits of the new genus, as well as specimens of four
other species, with field notes and photographs.

NEOABBOTTIA Britt. and Rose, gen. nov.

A treelike cactus with a smooth upright terete trunk and a much
branched top, the branches strongly winged or ribbed, normally from
the distal end of the preceding branch, but sometimes from below the
tip and usually in the same plane; ribs thin and high, very spiny;

FLIKGQ) Ihe FIG: 2:

Fics. 1 and 2.—Flower and fruit of Neoabbottia. Natural size.

flowers nocturnal, small, tubular with a narrow limb, borne several
together at the distal end of a terminal branch from a small felted
cephalium ; perianth persisting on the ovary ; perianth-tube and ovary
bearing small scales with short wool and an occasional bristle in their
axils; perianth-segments very small; throat of flower a little broad-
ened at the top, bearing many stamens; style slender; fruit oblong,
turgid, nearly naked, deeply umbilicate; seeds minute, black,
muricate.

A monotypic genus of Hispaniola, dedicated to Dr. W. L. Abbott.
a patron of natural history.

Type species, Cactus pamculatus Lam.

NO.Q NEOABBOTTIA, A NEW CACTUS GENUS—BRITTON AND ROSE 5

NEOABBOTTIA PANICULATA (Lam.) Britt. and Rose

Cactus paniculatus Lam. Encyl. 1: 540. 1783.
Cereus paniculatus DC. Prodr. 3: 466. 1828.

Six to ten meters high or even higher; trunk woody, 30 cm. in
diameter, the wood close-grained, yellowish white ; bark of the trunk
1.5 cm. thick, brown, not spiny in age, smooth; branches 4 to 6 cm.
broad, strongly 4-ribbed, rarely 5-ribbed, occasionally 6-ribbed or
winged; ribs thin, 1.5 to 2.5 cm. high, their margins somewhat
crenate, the areoles borne at the base of the sinuses, 1.5 to 2 cm.
apart ; spines 12 to 20, acicular, brownish to gray, 2 cm. long or less ;
cephalium 1 to 1.5 cm. in diameter, becoming elongated and angled ;
flowers straight, 5 cm. long, with a limb about 3 cm. broad; tube 6
to 7 mm. long, about 18 mm. in diameter, with walls 5 to 6 mm.
thick; inner perianth segments greenish white, short-oblong, about
I cm. long, obtuse; throat 18 mm. long, covered with numerous
filaments, these with a knee near the base and pressing against the
style; stamens and style included; ovary and flower tube tubercled,
the former with short tubercles, the latter with oblong ones (some-
times 1.5 cm. long), each ending in a depressed areole subtended by
a minute scale; areoles bearing a tuft of brown felt and an occa-
sional brown bristle; fruit oblong in outline, 6 to 7 cm. long, 4 to 5.5
cm. in diameter, turgid, nearly naked; rind green, thick, hard; seeds
rounded above, cuneate at base, with a large lateral depressed hilum.

Collected near Port-au-Prince, Haiti, on the Cul-de-sac by Dr.
W. L. Abbott and Mr. E. C. Leonard, April, 1920 (no. 3500) ; also
at the same locality by Mr. H. M. Pilkington, December, 1920: also
a single branch by Dr. Paul Bartsch at Thomazeau in 1917 (no.
221). Here doubtless belongs W. Buch’s specimen, described in a
note under Cereus paniculatus by Dr. 1. Urban in his Flora Domin-
gensis."

This plant was described by Plumier* as follows: Melocactus
arborescens, tetragonus, flore ex albido. This description was re-
peated by Tournefort,’ with the addition of a single word, in 17109.
Plumier’s drawing of this plant was published long after his death
by Burmann as plate 192 of the Plantarum Americanum and upon
this plate Lamarck based his Cactus’ paniculatus, which De Candolle

‘Symbol Antillane seu Fundamenta Flore Indiz Occidentalis, 8: 462. 1920.
*Catalogus Plantarum Americanum, 19. 1703.
* Histoire des Plantes, 1: 653. 17109.

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

a little later took up as Cereus paniculatus. Ever since, the plant
has usually passed under the latter name, with an occasional reversal
to the earlier one.

Until recently, the species has been known only from this old
illustration and these brief descriptions. The Abbott and Leonard
material consists of wood sections and herbarium specimens of
branches, flowers, fruit, and seeds, supplemented by living specimens
and by fruit and flowers in formalin, together with several habit
photographs. These convince us that the plant belongs to neither
Cactus nor Cereus, but to an undescribed genus. In habit it resem-
bles Dendrocereus, its branches resemble Acanthocereus, and the
small limb of the flower resembles Leptocereus; but the plant differs
from all of these in bearing several flowers at the ends of terminal
branches and in developing a kind of cephalium. In the last respect
it approaches Neoraimondia, near which we would place it in our
present classification. ]

Although at first Neoabbottia is weak and only 4-angled, suggest-
ing Acanthocereus, it forms ultimately a thick woody trunk. The
full-grown plant in habit and branches much resembles Dendrocereus
nudiflorus of eastern Cuba, but it has much smaller and different
flowers and seeds.

The following remarks are from the careful field notes of Mr.
Pilkington, made in December, 1920.

“ Grows to a height of 50 feet, in light sandy arid soil of recent
ocean bottom. Known to natives as ‘ Gadasse.’ No use is made of the
plant except burning the dead branches for torches. The wood so
used is called ‘ Bois Chandelle’ or ‘ Bois Flamboyant ’—‘ Candle-
Wood’ and ‘ Flaming-Wood,’ from the bright smokeless light. Fruit
falls when ripe; rind soon decays, leaving seed in a mass retained
in shape by a mucilaginous pulp. The young plant develops a bul-
bous root with a simple upright stem made up of several joints and
later giving off lateral branches which come off from the upper end
of other branches; the main stem is 4 to 6-winged, but as it grows
older becomes square, pentagonal, or hexagonal, according to the
original number of ribs on each joint, and in age terete or nearly
so with the ribs showing as mere lines, bearing the scars of the old
spines; the branches are more numerous on one side of the main
branch and these always lie in the same plane, the ribs when of the
same number being opposite those of the main joint. This dispo-
sition of the joints causes the main stem to bend or curve and the
whole has a striking resemblance to the flat antlers of moose and

NO.Q NEOABBOTTIA, A NEW CACTUS GENUS—-BRITTON AND ROSE 5

elk. This arrangement is shown in the mature tree, although the
intermingling of the several branches gives the general effect of an
ordinary tree-top.

“The natural pruning of the tree comes about through the death
of branches caused by epiphytic plants, the breaking off of branches
by the weight of a clambering cactus, and the attack of insects which
live in the fleshy joints. These insects are much sought after by a
red-headed woodpecker. ;

“Flowers are borne at the extreme tip of the terminal joints and
never from the sides, the fruit appearing therefore always at the
tips. A single fruit always grows directly in line with a rib, but when
several fruits grow from the same terminal bud they are compelled
to radiate at right angles to the axis of the joint. Four fruits from
one joint is the highest number observed, two only usually appearing
to be normal. As the ovary develops the flower shrinks, dries, and
appears finally as a brown protuberance attached to the apex of the
mature fruit. The old flowers at length fall off the mature fruit,
leaving a well-defined umbilicus. The fruit measures 6% to 7 inches
in circumference. When ripe it is waxy, smooth, yellow with faint
streaks of pink radiating from the base; flesh same color as rind,
glutinous, firm, slightly acid to taste, hardly edible.

“Seeds are embedded in a secretion which in water produces a
remarkable bulk of mucilaginous jelly, which is mildly acid and not
unpleasant to taste. Fruit does not seem to be attacked by birds and
is never eaten by natives. Successive crops of fruit appear from
this same bud cluster at the top of the terminal joint, each crop absorb-
ing some of the substance of the joint ; the joint shrinks and solidifies,
the ribs become furrows, the center enlarges, and finally all becomes
a woody mass of varying dimensions, as long as 3 inches, thus
forming what you have called a ‘cushion,’ but which is really an
atrophied joint after several years of fruit-bearing.”

The nature of the cephalium is not well understood, but it seems
to be an abortive joint. It first appears like a large felted areole
from which several flowers are produced; it slowly elongates and
finally becomes 7 cm. long or more, still producing the flowers at the
tip. When very old most of the felt wears off, leaving a stubby 4
or 5-angled joint ; the areoles, however, are not borne on the angles
as in normal branches, but in the depressions or furrows between the
ridges. In these furrows the areoles form a continuous band of felt
from the base to the top of the joint. One of these flower-bearing
joints which Mr. Pilkington has sent is 5 cm. long and we have esti-

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 7/2

mated that it has borne 20 sets of flowers and fruits and may possibly
be 20 years old. While all the flowers we have seen are terminal,
it is possible that they may sometimes occur from other places on
the terminal joint. In one specimen examined we have found an
enlarged areole near the base and one on the side of the terminal joint,
which suggests that they had been flower-bearing. Plumier’s illus-
tration, which is not accurate, shows numerous lateral flowers. The
stubby tlowering joints, while usually solitary, appear sometimes in
pairs.

EXPLANATION: OF PLATES

PLATE I

Plumier’s original illustration of Neoabbottia paniculata, reduced; repro-
duced from plate 92 of Burmann’s Plantarum Americanum.

PLATE 2
Neoabbottia paniculata (Lam.) Britt. & Rose <A. B. Two types of growth.

PATE

Neoabbottia paniculata (Lam.) Britt. & Rose. A, a plant growing in the
open; B, a plant growing in a thicket.

PLATE 4

A. Neoabbottia paniculata (Lam.) Britt. & Rose. Upper part of a plant.

B. Neoabbottia and Cephalocereus. a, An elongated branch of Neoabbottia;
e, a terminal branch of Neoabbottia, fruiting for the first time; c and f, stubby
branches of the same, which have produced fruit for many years; b and d,
small plants of Cephalocereus polygonus, growing epiphytically on Neoab-
bottia.

VOL. 72, NO. 9. PL. 1

SMITHSONIAN MISCELLANEOUS COLLECTIONS

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SMITHSONIAN

MISCELLANEOUS COLLECTIONS

VOL. 72, NO. 9, PL. 2

NEOABBOTTIA PANICULATA (Lam.) BRITT. AND Rose

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MISCELLANEOUS COLLECTIONS

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Sgt igh x

VOL.

72, NO. 9, PL. 3

NEOABBOTTIA PANICULATA (LAM.) BRITT. AND ROSE

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72, NO. 9, PL. 4

A. NEOABBOTTIA PANICULATA (LAm.) Britt. AND Rose

a

B. NEOABBOTTIA AND CEPHALOCEREUS

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 72, NUMBER 10

THE CIRCULATORY SYSTEM IN BONE

WitH Six PLATES

BY
US ROORE, MD:

(PuBLICATION 2652)

GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
AUGUST 20, 1921

The Lord Baltimore (Press
BALTIMORE, MD., U. 8. A. iy,

“

Thi CIRCULATORY “SYSTEM IN BONE

By Jy Se EOOTE, M.D.

Professor of Pathology, Medical Department of Creighton University,
Omaha, Nebraska

(WitH 6 PLaTEs)

INTRODUCTION

This article on the circulatory system in bone is the result of a
continued study of the comparative histology of bone published in a
monograph entitled “ A Contribution to the Comparative Histology
of the Femur,” Smithsonian Contributions to Knowledge, Vol. 35,
No. 3, 1916.

In that monograph were described the structural bone types and
the type combinations as they were observed in cross sections of the
femora of various animals from amphibians to and including man.”
The circulation within the bone substance was not seen at that time,
as cross sections do not show it properly.

In 1919 a casual preparation of a tangential section of the femur
of a domestic turkey disclosed a very remarkable circulation in the
bone substance and it was this disclosure that led to further examina-
tion of tangential sections of the bones of different animals, the draw-.
ings and descriptions of which are here presented.

Later (1919-20) the structure of and circulation in the bone of the
_ fish, as seen in the Mascalonge, Esox, were studied, compared with

the bone types of later vertebrate animals and added to the list of
bones examined.

THE CIRCULATORY SYSTEM IN BONE

The circulatory system in bone, as usually described, is limited to
the blood vessels of periosteal membranes, medullary canals and
diploe of long and cranial bones respectively, little being known about
the circulation within the bone substance itself.

The circulation described in this article is situated in the bone
substance of the mandible and cranial bones of the fish, Mascalonge,
_Esox, in the walls of the long bones of the amphibian, reptile, bird,

1Number of sections of femora described and drawn in the monograph, 440.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 72, No. 10

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

bat, other mammals and man, in the cranial bones of man, in the inner
wall of the oth rib and in the infra- and supra-spinous fossae of the
scapula of man. The blood supply of the lower jaw and cranial bones
of the fish is derived from the dermal vessels which enter the bone
at various points and assume parallel positions in the bone substance,
while the blood supply of the long bones of the higher vertebrates
comes from two sources, viz. the periosteal vessels which send off
many small branches into the bone through entering canals (canals
of Volkmann and other canals), and the medullary arteries which
pass obliquely through the walls of the shafts into the medullary
canals where they divide into ascending and descending branches
from which small vessels are sent off into the walls of the bones and
here become continuous with the vessels from the periosteum.

The blood supply of the flat bones, such as the cranial, is derived
from the vessels of the pericranial and endocranial membranes which
send off branches into the outer and inner tables of these bones com-
municating by way of the central medullary diploe.

From a study of a large number of bone sections * there are found
to be three structural types and various type combinations * which
enter into the formation of bone: these are the first, composed of
lamellae ; the second, composed of laminae; and the third, composed
of Haversian systems. These three types are combined in various
proportions in the bones of different animals. The circulations
which are found to be present in the different types of bone also
present variations which are sufficiently distinctive in character to
form two circulatory types, viz., the branching, and the plexiform.
The branching type, composed of tree-like branches, is found in the
first type bones, the plexiform, composed of small, large and irregu-
larly shaped meshes enclosed by small vessels, is found in the second
and third type bones, while combinations of the branching and plexi-
form circulations are present in structural type combinations.

In the demonstration of the circulations some difficulty arises in
the preparation of the bone slides. In small animals like the frogs
or other animals of the same size it is practicaily impossible since
the long bones are almost always round and tangential sections of
them are necessarily flat. In large animals the long bones have flat
areas of sufficient extent to make satisfactory slides.

In the preparation of bone sections for the purpose of showing the
circulation, a flat surface of bone is selected and as large a piece as

* Number of sections of bones examined up to the present, 1000.
* A Contribution to the Comparative Histology of the Femur, by J. S. Foote,
M. D., Smithsonian Contr. to Knowl., Vol. 35, No. 3, 1916.

NO. IO THE CIRCULATORY SYSTEM IN BONE—FOOTE 3

possible is sawed out, tangential to the diameter of the bone, and
ground down to a suitable thinness which is determined by the ap-
pearance of the circulation.

It ‘is not the thinnest possible section, measured in microns, that
is most desirable, but one of sufficient clearness to show the circula-
tion. During the grinding process it is necessary to examine the
section at short intervals in order to ascertain just when to terminate
the process.

The following sections taken from the bone of fish, amphibian,
reptile, bird, mammal, and man have been studied, described, and
drawn for the purpose of showing the structure and the circulation
which belongs to it.

CROSS AND TANGENTIAL SECTIONS OF THE LOWER JAW AND
TANGENTIAL SECTION OF THE CRANIAL BONE OF THE MAS-
CALONGE, ESOX, A FISH OF WISCONSIN WATERS

CROSS SECTION OF A PORTION OF THE LOWER JAW
Pie Gen eleA\

The section is composed of parallel disks of bone substance situated
between vascular channels or clefts. The bone substance does not
show the presence of lacunae with their canaliculi. Very minute
parallel canaliculi extend across the disks from one channel or cleft
to another. A wave effect is given to the section by the undulating
forms of the clefts. This arrangement of channels in the bone sub-
stance produces a very fine channel circulation. At short intervals
within the clefts, as at C, figure 1, plate 1, may be seen small objects
from which radiate minute canaliculi presenting the appearance of
the canaliculi radiating from their lacunae, but these are in the vas-
cular clefts and not in the bone substance.

TANGENTIAL SECTION OF A CRANIAL BONE OF THE MASCALONGE
IPAb, any IRE, ae

In this section are seen parallel rows of objects of various shapes
situated in the bone substance. Some of them are circular with dark
crescents on one side, while others are very irregular in shape. No
canaliculi extending outward from them can be seen. For this
reason they cannot be lacunae in the bone substance, as osteoblasts
occupying such lacunae would be without a blood supply. On ac-
count of their general circular character these objects appear to be
different sections of the vascular clefts of the bone and if any bone-
producing cells are present they must be within the vascular clefts. .

4. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

CROSS SECTION OF ANOTHER PORTION OF THE LOWER JAW OF THE MAS-
CALONGE SHOWING THE EARLY DIFFERENTIATIONS OF
HAVERSIAN SYSTEMS
IPs iy LOWY

These are small vascular canals surrounded by clear areas—in
some instances lamellated, in others, homogeneous. Between the
vascular canals is the bone substance with very fine channels. The
vascular canals with surrounding clear areas become Haversian sys-,
tems as vascular differentiation progresses. This early form of
Haversian differentiation has a wide distribution since it is found in
bone sections from fish to and including man. It is much more prom-
inent in the lower orders of vertebrates than in the higher. Thus
two early phases of the circulation are found in the bone of the
mascalonge, viz., the channeled and early Haversian differentiation.

TANGENTIAL SECTION OF THE INNER RIDGE OF THE LOWER JAW OF
THE MASCALONGE SHOWING THE BLOOD VESSELS
IBity ity MC)

The blood vessels vary in size. They are parallel with each other.
Some are branching. Around the outside of their delicate connective
tissue walls are seen fine plexuses of nerves. Between the vessels
is seen a fine channeled bone substance without lacunae. —

CROSS AND LONGITUDINAL SECTIONS OF THE FEMUR OF A
MEDIUM SIZED BULLFROG, RANA CATESBEIANA

(The large, medium sized and small bullfrogs have different degrees of bone
differentiation.) *

CROSS SECTION SHOWING THE STRUCTURE
Tein iis JENKeS at

The type of bone is an early first. The inner wall is composed of
bone substance in which are radiating, vascular, bush-like channels
extending from the external to the internal circumferential lamellae
and between which are round or oval lacunae with short branching
canaliculi embedded in the bone substance and communicating with
the bush-like radiations by means of their canaliculi. The bush-like
vascular radiations appear to be segmented in some instances due
to their oblique positions and the plane of the cross section. In the
outer wall these radiating channels are absent and the bone is com-
posed of lamellae with oval lacunae and radiating canaliculi.

*Tdem.

NO. IO THE CIRCULATORY SYSTEM IN BONE—FOOTE 5

LONGITUDINAL SECTION OF THE SAME FEMUR SHOWING THE
CIRCULATION
Pu. 1, Fie. 5

The oblique entering canals in the shafts of the femora of bull
frogs, for the passage of the medullary arteries, are found in the bone
of the large and medium sized but not in the small animals. As the
femur of the frog is small and round the circulation in a tangential
section could not be seen. The general plan, however, is shown in a
longitudinal section of the inner wall as represented in figure 5,
plate 1. The letter C is placed at the center of the shaft. In this wall
are two sets of six or seven oblique vascular canals entering the shaft
from the periosteal surface and united in such a manner as to form
slanting m-shaped converging loops. These loops, beginning near
the extremities of the shaft, converge toward the lineal center and
medullary surface. From these loops and their stems minute can-
aliculi are sent off into the bone substance where they communicate
with the lacunae. Between the two sets of converging loops are short
segments of loops extending across the wall transversely. In the
outer wall the loops are not seen in longitudinal section nor are the
radiating vascular channels seen in the cross section of this wall.

CROSS AND TANGENTIAL SECTIONS OF THE FEMUR OF THE
REPTILE, ALLIGATOR MISSISSIPPIENSIS

CROSS SECTION SHOWING THE STRUCTURE
Pei. bite. 6

This section is composed of three wide, concentric rings of incom-
pletely differentiated Haversian systems alternating with two narrow,
concentric laminae. The vascular canals surrounded by clear areas
similar to those seen in the bone of the fish are early forms of
Haversian systems. The laminae are more advanced in differentia-
tion than the Haversian systems. The bone, as a whole, shows an
early second and third type structure.

TANGENTIAL SECTION OF THE SAME FEMUR, SHOWING THE
CIRCULATION

124b aie EWC G

The circulation is in the form of a vascular plexus. The section
is situated below the periosteal surface near the posterior ridge.
Several entering canals, without surrounding lamellae, are seen in

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

cross section in the bone substance through which periosteal vessels
pass into the bone. The vascular plexus is very extensive and has
a general longitudinal direction. The meshes are irregular in size
and shape. In some portions of the section vascular expansions are
found. The blood vessels are round and occupy similarly shaped
channels in the bone substance. They are composed of very thin
connective tissue walls, without smooth muscle, and are striated
spirally and longitudinally. The exact purpose of the vascular ex-
pansions is not clearly understood. They are found too frequently
to be accidental, and, as will be noted later, are more prominent in
the branching than in the plexiform types of circulation. They may
have an important physical value.

CROSS AND TANGENTIAL SECTIONS OF A SECOND TYPE BONE
'. OF BIRDS AS SEEN IN THE FEMUR.OF A DOMESTIC
TURKEY, MELEAGRIS GALLOPAVO

CROSS SECTION SHOWING THE STRUCTURE
~ Pie Tene

The section is composed of concentric laminae separated and
crossed by vascular canals. The wall of the bone is divided into
nearly equal segments by large radiating canals extending from the
medullary canal to the periosteal surface. From these canals are
sent off lateral, small, parallel canals which divide the wall of the
bone into laminae. The laminae are interrupted in the anterior wal!
and posterior ridge by incompletely differentiated Haversian systems.
It was the femur of the turkey which first called attention to the
variations in structure and circulation.

TANGENTIAL SECTION OF A FEMUR OF THE TURKEY, SHOWING
CIRCULATION

Pia, iGo

This section consists of a very rich, small-meshed plexus of vessels
situated between the laminae and having a general longitudinal direc-
tion. In the central portion of the section two plexuses can be seen,
one above the other, with a lamina of bone between them. ‘This is
one of the most extensive circulations observed in long bones and
with such a blood supply as is here shown, this becomes a vascular
organ of great importance.

NOs 10 THE CIRCULATORY SYSTEM IN BONE—FOOTE 7

RECONSTRUCTION OF A SECOND TYPE BONE WITH ITS INTER-
LAMINAR CIRCULATION

Piezehice 10

This figure shows a reconstruction of a second type bone with its
circulation. The drawing represents three concentric laminae with
two interlaminar plexuses of blood vessels. The vessels are uniform
in size, composed of thin connective tissue walls and the vascular
expansions are not at all prominent. It is difficult to understand why
such a rich blood supply should be required in bone for its nutrition.

CROSS AND TANGENTIAL SECTIONS OF AN EARLY DIFFERENTI-
ATION OF A THIRD TYPE BONE IN BIRDS AS SEEN IN THE
FEMUR OF A TURKEY BUZZARD, CATHARTES
AURA SEPTENTRIONALIS

CROSS SECTION SHOWING THE STRUCTURE
Bae eatGsrT

This bone is more advanced in differentiation than that of the
domestic turkey since it is composed of third type structural bone
units instead of the second. These units are enclosed between the
external and internal circumferential lamellae and form nearly the
whole bone structure.

TANGENTIAL SECTION OF THE FEMUR OF THE SAME ANIMAL SHOWING
THE CIRCULATION

Pie 2 hice TZ

The meshes of the circulatory plexus are longer and enclose larger
bone areas than those seen in the femur of the turkey. The vascular
expansions are not as prominent as they are in some of the other

bones, but may be seen here and there. A general plexiform plan of
circulation is evident, but an elongation of the meshes in the central
portion shows a slight variation in the circulatory distribution.

TANGENTIAL SECTION OF THE FEMUR OF A DOMESTIC
CHICKEN, GALLUS

Eig, EME ai:

The type of structure is a first and early third. Several entering
canals without surrounding lamellae are present. The circulation
appears as a rich plexus of small blood vessels coming off from a
central vessel which extends lengthwise of the section. The meshes
of the plexus are small, round, oval or irregular in shape and the
lacunae of the bone substance are round or oval with short, branching
canaliculi.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

TANGENTIAL SECTION OF THE FEMUR OF A PRAIRIE CHICKEN,
TYMPANUCHUS AMERICANUS, SHOWING THE
CIRCULATION

12, ZAI, TA

The type of bone is first. A few entering vascular canals, without
surrounding lamellae, are seen in the bone substance. The circula-
tion is a rich plexus of blood vessels with small, round and irregularly
shaped meshes. It is situated nearer the periosteal than the medullary
surface. The lacunae of the bone substance are round or oval and
have short bushy canaliculi.

TANGENTIAL SECTION OF THE FEMUR OF A DOMESTIC DUCK,
ANAS DOMESTICA, SHOWING THE CIRCULATION

Bie Ze Ie, iis

The type of bone is an early second. The section has two rich
plexuses of blood vessels, one above the other, with bone substance
between them. The meshes are quite regular in form and somewhat
larger than those found in the other birds examined. Here and there
are seen entering vascular canals without surrounding lamellae. The
lacunae of the bone substance are round or oval with short bushy
canaliculi.

CROSS AND TANGENTIAL SECTIONS OF THE FEMUR OF A
FRUIT BAT, PTEROPUS (CELEBES)

CROSS SECTION SHOWING THE STRUCTURE
lee, 2 Leey, ie)

The section is composed of a wide central ring of lamellae per-
forated in the inner wall by a few vascular canals extending length-
wise of the bone. Internal circumferential lamellae surround the
medullary canal and poorly differentiated external lamellae surround
the section. The lacunae are oval with straight canaliculi.

TANGENTIAL SECTION OF THE SAME FEMUR SHOWING THE
CIRCULATION

Pie we EiGerty,

In this section are seen the vascular canals of the inner wall extend-
ing from above downward and inward. The canals are parallel with
each other and some of them are branched. They were absent in the
outer wall.

NO. IO THE CIRCULATORY SYSTEM IN BONE—FOOTE 9

CROSS AND TANGENTIAL SECTIONS OF A SECOND TYPE BONE
IN MAMMALS AS SEEN IN THE FEMUR OF A LAMB, OVIS

CROSS SECTION SHOWING THE STRUCTURE
IPL, 2 Ree Tits}

The section is composed of concentric laminae which are more com-
pletely differentiated than they are in birds. Here and there the
laminae are interrupted by Haversian systems of a late differentiation
and the concentric canals between the laminae are widened at inter-
vals and around them are incompletely differentiated lamellae form-
ing an early aberrant type of Haversian system. The posterior ridge
is composed of Haversian systems of a late differentiation.

TANGENTIAL SECTION OF THE FEMUR OF THE SAME ANIMAL SHOWING
THE CIRCULATION
Teab, 24, IRuCh a0)

The circulatory plan of arrangement in this bone is not precisely
like that observed in the second type bone in birds. While it is plexi-
form in character there is about it more or less irregularity in distri-
bution approaching the branching type of circulation. Near the
sides of the drawing the plexus has a long mesh; while in the central

portion it has more of a branching character. The vascular expan-

sion seen in the center of the drawing gives the impression of a
distributing point in the circulation. A few entering vascular canals
are seen.

CROSS AND: TANGENTIAL SECTIONS OF ANOTHER SECOND
TYPE BONE IN MAMMALS AS SEEN IN THE FEMUR OF
THE MEXICAN BURRO

CROSS SECTION SHOWING STRUCTURE
Pia RiGw 20
The section is composed of concentric laminae interrupted by
Haversian systems of different degrees of differentiation. The struc-

ture is similar to that seen in the femur of the lamb and a large
number of other mammals which have the same type bone.

TANGENTIAL SECTION OF THE FEMUR OF THE SAME ANIMAL,
SHOWING THE CIRCULATION

Pie EiGs 2
The type is plexiform and the meshes of the plexus are much more
regular in shape than those seen in the femur of the lamb. There is
no evidence of a branching character. The vascular expansions are
not as prominent as they were in the femur of the lamb.

ie) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

TANGENTIAL SECTION OF THE FEMUR OF AN ELK, ALCES
AMERICANUS, SHOWING THE CIRCULATION

Pi. 3, Hies22

The type of bone is second. Several perforations in the bone sub-
stance for the passage of blood vessels are present. Some of them
are surrounded by enclosing lamellae and some are not ( Volkmann’s
canals). The circulation is an extensive plexus of blood vessels with
round and irregularly shaped meshes. It has a general direction
lengthwise of the bone.

TANGENTIAL SECTION OF THE FEMUR OF A BELGIAN HARE,
LEPUS, SHOWING THE CIRCULATION

lets 2h, IEE, 3}

The type of bone is an early second and third. Entering vascular
canals through which blood vessels are seen to pass and ramify in
the bone substance are present in the central portion of the section.
The canals are not surrounded by lamellae. The circulation is a com-
bination of a branching and plexiform distribution. The vascular
expansions are prominent. One is seen in a vessel shortly after
entering the bone. It is a matter of observation in bone circulations
that the type of circulation varies with the type of structure.

TANGENTIAL SECTION OF THE FEMUR OF A BULLDOG,
SHOWING THE CIRCULATION

Press EiGa2A

The type of bone is second and third. Entering vascular canals,
without enclosing lamellae, are seen in the bone substance through
which blood vessels are passing. The circulation is a dense branching
and plexiform combination. The vessels are small in diameter, fre-
quently branching, the union of their branches producing a plexiform
emecn

CROSS AND TANGENTIAL SECTIONS OF THE FEMUR OF A
MONKEY, MACACA RHESUS

CROSS SECTION SHOWING THE STRUCTURE
[Pits sh, lainey 2s

The type of bone is first and third. The section is composed of
lamellae interrupted by Haversian systems of early and late differ-
entiations. Crescents of late Haversian differentiation are found
bordering upon the medullary canal in the anterior inner and pos-
terior outer wall. The lacunae of the bone are long with straight
canalicult. The principal structure is lamellar and the type is much
more first than third.

NO: TO THE CIRCULATORY SYSTEM IN BONE—FOOTE ie

TANGENTIAL SECTION OF THE SAME FEMUR, SHOWING THE
CIRCULATION

BraserhiGs 26

A few entering vascular canals without surrounding lamellae are
present. The circulation is a dense branching type which, here and
there, presents a slight appearance of the plexiform distribution. The
blood vessels are large and trunk-like in places and a very few vas-
cular expansions are present.

CROSS AND TANGENTIAL SECTIONS OF THE FEMUR OF A
9 MONTHS’ HUMAN WHITE FETUS

CROSS SECTION SHOWING THE STRUCTURE
Pie a eG 277

The section is composed of short laminae with central canals widen-
ing and shortening until they are transformed into Haversian sys-
tems.. External circumferential lamellae surround the section (pos-
terior wall excepted) and directly underneath the lamellae is a row
of advanced differentiations of Haversian systems. The medullary
canal is small, irregular in shape and situated in the anterior half
of the section. The posterior wall is composed of Haversian systems
of advanced differentiations and in the mid line where the walls of
the bene unite is a narrow radiating layer of bone substance which
disappears as complete union takes place.

TANGENTIAL SECTION OF THE SAME FEMUR SHOWING THE
CIRCULATION

Pr, 3, Fie. 28
The circulation is plexiform in type, the general direction of which
is longitudinal and slightly oblique. The vessels forming the plexus
have short branches which seem to disappear in the bone substance
within the meshes. Vascular expansions are present, one being
shown in the left central portion of the figure.

CROSS AND TANGENTIAL SECTIONS OF THE FEMUR OF A
WHITE: CHILD 9, YEARS. OLD

CROSS SECTION SHOWING THE STRUCTURE
Pie 34 BGO

The type of bone is first and third. Around the periphery of the
section is a wide crescent shaped band of lamellae enclosing numerous
vascular canals of the earliest Haversian differentiation, such as were

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

observed in the bone of the fish, plate 1, figure 2. Around the medul-
lary canal are the internal circumferential lamellae forming an enclos-
ing ring of irregular widths. Within this ring are also found
vascular canals of early Haversian differentiation extending longi-
tudinally. Between the external and internal lamellae above
described, is a ring of Haversian systems of late differentiation,
deficient in the anterior wall and increasing in width in the outer
lateral wall as it reaches the posterior ridge. The lacunae are long
and narrow with straight canaliculi.

TANGENTIAL SECTION OF THE SAME FEMUR SHOWING THE
CIRCULATION

1Ett, 3, JME, AO

Entering canals with and without enclosing lamellae are of fre-
quent occurrence. The circulation is a combination of the branching
and plexiform types. The central portion is branching and the
lateral, plexiform, vascular expansions are not as prominent as they
are in many other sections. The section shows the circulatory com-
bination of type conforming to the structural type combination.

CROSS AND TANGENTIAL SECTIONS OF THE FEMUR OF ADULT
MAN, A WHITE MALE

CROSS SECTION SHOWING THE STRUCTURE
IPG, ah IEE, Bit

The bone is third type in differentiation with the exception of a
portion of the anterior wall where the remains of first type bone are
found. The section is composed of completely differentiated Haver-
sian systems, some of which are senile. The external circumferential
lamellae are fragmentary, while the internal are complete.

TANGENTIAL SECTION OF THE SAME FEMUR, SHOWING THE
CIRCULATION

Pra hic..32

The type of circulation is plexiform with wide, irregularly shaped
meshes. Entering vascular canals surrounded by lamellae are seen
here and there in the section. Vascular expansions are prominent.
This section is taken from the bone represented in plate 4, figure 31,
near the periosteal surface of the left side of the drawing and near
the posterior ridge. It is difficult to think of the circulation as shown
in plate 4, figure 32, as belonging to that locality.

NO. IO THE CIRCULATORY SYSTEM IN BONE—FOOTE 13

TANGENTIAL SECTION OF THE FEMUR OF MAN, A WHITE MALE,
SHOWING CIRCULATION

PreAmiitGs 338

The type of bone is first and third. Numerous entering vascular
canals with and without enclosing lamellae are present. The circula-
tion is a dense branching and plexiform distribution of blood vessels
with small meshes of various shapes. Vascular expansions are
numerous and large. The lacunae of the bone substance are round,
oval, and long and narrow with the canaliculi which belongs to each
degree of differentiation. The density of the circulation varies in
different bones of the same structural type and also in bones of
different type combinations.

TANGENTIAL SECTION OF THE FEMUR OF MAN, A WHITE MALE,
SHOWING THE CIRCULATION

Pi 4s Bie 34

The type of bone is first and third. The type of circulation is
branching, changing to plexiform. Entering canals, for the most
part without enclosing lamellae, are few in number in this section.
The circulation shown in the drawing is situated in the external
lamellae which encloses the internal third type differentiation. It is
a large branch dividing into many small ones which, by uniting, form
a plexiform distribution.

TANGENTIAL SECTION OF THE FEMUR OF MAN, A WHITE MALE,
AGE 50, WHO DIED OF PULMONARY TUBERCULOSIS

Prn4 ere: 35

The type of bone is third. The type of circulation is essentially
branching although a coarsely plexiform arrangement can be distin-
guished. Entering canals with and without enclosing lamellae are
seen in the bone substance. The blood vessels are irregular in size
and shape and show varicose enlargements very frequently. Whether
‘or not this varicose condition and irregularity in the circulation in
bone are indices of similar changes in the general circulation cannot
be told; but they may be thought of in connection with the pathologi-
cal condition present in this individual.

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

THE CIRCULATION IN FLAT BONES
CROSS, LONGITUDINAL, AND TANGENTIAL SECTIONS OF A
HUMAN FRONTAL BONE
CROSS SECTION SHOWING THE STRUCTURE
Pr. 4; ere 36

The bone is composed of outer and inner tables united by a central
cancellous diploe. The two tables are composed of first type bone
enclosing a few Haversian systems in cross section and a few short
segments of blood vessels. The diploe is a coarse, cancellous bone
with large, small, and irregularly shaped cavities enclosed by first
type bone walls. The walls are composed of lamellae in which
Haversian systems are seen in cross section. This section was taken
from the vertical portion of the frontal bone.

LONGITUDINAL SECTION OF THE SAME FRONTAL BONE, SHOWING THE
STRUCTURE

TPAD, 2, EIR, Sys

This section has practically the same structure as the cross section,
as may be seen by comparing the drawings. This section is cut
at right angles to that seen in plate 4, figure 36. It is taken from the
same region, and Haversian systems, in cross section, are found in
both situations. The Haversian systems therefore run at right angles
to each other, which can hardly be accounted for on mechanical

grounds.

TANGENTIAL SECTION OF THE OUTER TABLE OF THE SAME FRONTAL
BONE TAKEN FROM THE SAME REGION AS FIGURE 36, PLATE 4,
SHOWING THE CIRCULATION :

Pin 4s RIG. 35

The section is situated nearer the external surface of the bone than
the dipl6e. Numerous entering canals with and without enclosing
lamellae are found in the bone substance. The circulation is branch-
ing in type. The vascular expansions are large and numerous and
appear to form physical centers of distribution. The blood vessels
are relatively large and frequently branch.

ENTERING VASCULAR CANALS OF THE OUTER TABLE OF THE
FRONTAL BONE

[ee iy IRC, S{0)
There are two forms, the one at the left in the drawing without

enclosing lamellae (Volkmann’s canals), and the one at the right
with enclosing lamellae. The Volkmann’s canals are smaller than

NO. IO THE CIRCULATORY SYSTEM IN BONE—FOOTE 15

the others. As soon as the blood vessels pass through the external
surface of the bone they send off branches into the planes of their
divisions and form the branching distributions there.

TANGENTIAL SECTION OF THE INNER TABLE OF THE SAME FRONTAL
BONE, SHOWING THE CIRCULATION

Pia 5 .h1G. 40

In this section the bone is perforated by numerous entering canals
with and without enclosing lamellae. The blood vessels from the
cerebral surface enter the bone by these canals and find their way to
the diploe. The circulation within the bone is branching in type and
situated nearer the cerebral surface than the diploe. Many vascular
expansions are present with their incoming and outgoing vessels.

TANGENTIAL SECTION OF THE OUTER TABLE OF A HUMAN
PARIETAL BONE, SHOWING THE CIRCULATION

PETS biG 4t
The section is taken from the central portion of the bone. Thé
type of bone is first. Entering vascular canals surrounded by lamellae
are numerous. The circulation is branching in type. Vascular ex-
pansions are large and numerous. Those in the center give the im-
pression of distributing points in the circulation.

TANGENTIAL SECTION OF THE INNER TABLE OF THE SAME PARIETAL
BONE SHOWING THE CIRCULATION

Iie, Sy, JEN, A

The section is situated nearer the cerebral surface than the diploe.
Entering vascular canals with enclosing lamellae are numerous. The
circulation is branching in type and not as dense as that of the outer
table. The vascular expansions are many and large. The type of
bone is first.

TANGENTIAL SECTION OF THE SQUAMOUS PORTION OF THE
TEMPORAL BONE OF MAN, SHOWING THE CIRCULATION

Pi 5) 1G) 43

The type of bone is first. Entering canals with and without en-
closing lamellae are present. The circulation is branching in type
and the vascular expansions are large and very numerous. The ex-
pansions vary in size and frequency of occurrence in different sec-
tions. In some instances they are very small and few in number, in
others, large and numerous, and they are much more prominent in the

16 SMITHSONIAN. MISCELLANEOUS COLLECTIONS VOL. 72

branching than in the plexiform type of circulation. They are
especially prominent in this section.

TANGENTIAL SECTION OF THE OUTER TABLE OF A HUMAN
OCCIPITAL BONE, SHOWING THE CIRCULATION

Pr. 5, Fic. 44

The section is taken a short distance above the foramen magnum.
The circulation is a dense branching type situated near the external
surface of the bone. There are many vascular expansions from
which branching vessels take their departure. Entering canals with
and without surrounding lamellae are present in the first type bone
substance.

TANGENTIAL SECTION OF THE INNER TABLE OF THE SAME OCCIPITAL
BONE, SHOWING THE CIRCULATION

PL: 5, Fic. 45

The type of bone is first. Entering canals with and without sur-
rounding lamellae are numerous. The section is situated near the
cerebral surface. The circulation is branching in type and very
dense in character. Large branches extend in different directions
and from these, small branches are sent off into the bone substance.
The vascular expansions are not as numerous as those in the outer
table, figure 44.

TWO VASCULAR EXPANSIONS AS THEY WERE SEEN IN THE OUTER
; TABLE OF THE OCCIPITAL BONE

PL. 5, Fic. 46

The vascular expansions are observed in all branching and in
many sections of the plexiform type of circulation. They are nearly
always filled with fat globules with dark, wide contours which are
so closely faceted together that they completely occupy the expansion
cavities. This would, of course, block the circulation if it existed
in the living bone. The condition was misleading in significance
until it was noticed that the fat globules were found in ground sec-
tions and not in thin unground bone plates ; so that they are probably
produced by the melting heat of friction during the grinding process
and fill the expansions. The expansions probably have a physical
value as distributing centers in the circulation or they may be for
the purpose of establishing and maintaining a circulatory equilibrium
in bone.

NO. 10 THE CIRCULATORY SYSTEM IN BONE—FOOTE 17

TANGENTIAL SECTION OF A.HUMAN INFERIOR MAXILLA,
SHOWING THE CIRCULATION

12, Gi, [Res Ay

The section is taken from the bone just in front of and below the
left central incisor tooth. The type of bone is first. Entering canals
with and without lamellae are present. The circulation is a branch-
ing plexiform type having a slanting direction from above downward
and forward. The meshes are long and irregular in shape. Vascular
expansions are numerous.

TANGENTIAL SECTION OF THE INFRA-SPINOUS FOSSA OF A
HUMAN SCAPULA, SHOWING THE CIRCULATION

IPI canes, IRC ts!

The type of bone is first with here and there an Haversian system.
A few entering canals with enclosing lamellae are seen in the section.
The circulation is a branching and plexiform type, the former pre-
dominating. Vascular expansions are prominent. The one in the
center of the drawing suggests a distributing center.

TANGENTIAL SECTION ‘OF THE SUPRA-SPINOUS FOSSA OF THE SAME
SCAPULA, SHOWING THE CIRCULATION

Pi. 6, Fic. 49

The type of bone is first. A few entering canals with and without
surrounding lamellae are present. The circulation is branching in
type with a few vascular expansions. It takes an oblique direction
from above downwards and toward the vertebral border.

CROSS AND TANGENTIAL SECTIONS OF THE 9TH RIB OF ADULT
MAN, MALE WHITE

CROSS SECTION SHOWING THE STRUCTURE
PEVG RP IGa50

This is a third type bone composed of Haversian systems of a late
degree of differentiation. The systems extend lengthwise of the rib
and enclose a cancellous center.

TANGENTIAL SECTION OF THE INNER WALL OF THE SAME RIB,
SHOWING THE CIRCULATION

Pr. 6, Fic. 51

The section is taken from the posterior third of the bone. The
circulation is an elongated plexus. In the center of the section is a
vascular expansion which suggests a center of distribution for the
blood vessels.

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

DEVELOPMENTAL ADVANCES IN THE CIRCULATION
DETERMINE THE STRUCTURAL TYPES OF BONE

It is generally supposed that bone is preeminently mechanical in
function on account of its position in the body and its rigid character.
It forms the skeleton of the animal body, supports its weight, gives
attachment to muscles by means of which locomotion is possible,
serves as a framework upon which the viscera are hung, affords pro-
tection and gives efficiency to the laboratory of chemical activities
constantly in operation during life. It is chiefly for these reasons
that we attribute a mechanical function to bone. This thought is
further strengthened by the microscopic structure so familiar to us
either from personal observation or from text books of histology,
in both of which bone is represented, usually, as composed of Haver-
sian systems.

There are however, as shown before, three structural types of
bone, the first, second, and third.

The first type is composed of bone substance or lamellae, the
second of laminae which are produced by a vascular separation of
lamellae into parallel divisions, and the third, of Haversian systems
which are produced by arranging lamellae around small central vas-
cular canals called Haversian canals. The Haversian system, there-
fore, is the most complex and highly organized bone unit. The types
of structure follow the advancing changes in the circulation, since
the second type bone is not recognizable until the first has been separ-
ated into laminae by parallel vascular canals. There are two general
types of circulation, the branching and the plexiform, each one giving
an individual character to the type of bone it produces. Bone de-
rived from connective tissue membranes retains the branching circu-
lation of those membranes, while bone derived from cartilage has
the plexiform circulation.

In going from a branching to a plexiform type of circulation, the
type of bone advances from the first to the second or third. The
blood supply in a plexiform circulation is greater in volume than it
is ina branching circulation, for the reason that there are more blood
vessels in a given area in the former than in the latter. This in-
creased blood supply adds an increased physiological value to the
bone units of structure and is the foundation of advancement in
tissue values: so that blood supply is determined by the extent, plan
of distribution, or type of circulation and the type of tissue, phys-
iologically considered, by the blood supply, or in this particular
instance, the type of bone is determined by the type of circulation.

NO. 10 THE CIRCULATORY SYSTEM IN BONE—FOOTE 19

THE STRUCTURAL UNITS OF BONE—FIRST, SECOND, AND
THIRD, OR LAMELLAE, LAMINAE, AND HAVERSIAN
SYSTEMS—ARE NOT ESSENTIALLY MECHANICAL

If all bones were Haversian system bones, or, if there was any one
bone which always has Haversian systems as the predominating
structural units, no matter in what individual it was found, we might
be convinced that their constant presence was sufficient evidence of
a mechanical function. But when we know that there are three bone
units of structure and that no one of them is constantly present in
bones of different animals as the only unit, and not even in bones of
individuals of the same birth,’ the idea of a purely mechanical func-
tion of bone is severely shaken if not abandoned. ‘The three units,
the lamella, lamina, and Haversian system, are found in bones of
various animals in such great proportional and locational confusion
that their mechanical purposes disappear and we are obliged to
look further for an explanation of their strange and unexpected
occurrences.

Again, if we examine these units from a mechanical viewpoint
and then observe their locations in the bones of different animals
we will find that they are not always found where the same mechan-
ical conditions would require them and are very often found where
they could serve no mechanical function of any importance. Lamellae
are layers of bone substance, laminae are strata composed of lamellae,
and Haversian systems are hollow cylinders composed of a variable
number of lamellae enclosing a central canal. Of the three units,
therefore, the Haversian systems offer the most mechanical service
by construction and are the best adapted to support weight, with-
stand muscular stress, and serve the general requirements of a
skeleton. We would, then, naturally expect to find them constantly
present in such long bones as the femur, tibia and fibula of quad-
rupeds and bipeds and generally absent from the flat bones of the
head and face. But in these respects we are disappointed. They
are found, as a pure type, in the long bones of only a few mammals
and are absent as predominating units in the long bones of a large
number of mammals including the three races of man, black, yellow-
brown and white.* Haversian systems in their later degrees of dif-
ferentiation are not found at all in amphibians, reptiles, birds, bats,
monotremes, marsupials and many of the edentates, although the
same mechanical functions are demanded by their vocational habits.
In the femur, tibia, fibula, humerus, radius, ulna, clavicle, metacarpal
and metatarsal bones of man, they are present as predominating

5 Idem.
5 Idem.

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

units in some and occur very infrequently in other individuals. In
the flat bones of the cranium and in the irregular bones like those
of the pelvis a few are found where it is difficult, if not impossible,
to understand how they could perform any mechanical function of
any value.

In the human frontal, parietal and occipital bones, a few Haver-
sian systems may be found at right angles to each other; in the
squamous portion of the temporal bone they extend vertically; in
the hard palate, antero-posteriorly ; in the superior and inferior
maxillae, horizontally ; in the spine of the scapula, horizontally; in
the crest of the ilium horizontally ; and in the ribs horizontally. In
these different situations the mechanical functions of the Haversian
systems do not seem to answer any requirement which is common
to them all. Furthermore, they are not invariably found in these
situations and even may not be present at all. Their presence or
absence or situation within the bone is more satisfactorily explained
if we assume that developmental advances in the circulation deter-
mine the types of bone.

CONCLUSIONS

A study of the circulations as they were observed in the foregoing
slides leads one to the following conclusions :

1. That a cross section of bone gives no idea of the plan of
circulation.

2. That there are two types of circulation, the branching and the
plexiform, and these are seen only in tangential sections.

3. That the plan of circulation in bone derived from connective |
tissue is branching; derived from cartilage, plexiform.

4. That the circulation in a first type bone is branching, in the
second and third, plexiform.

5. That the plexiform is a more advanced type of circulation than
the branching and represents a later plan of vascular distribution
and a more advanced degree of bone differentiation.

6. That bone is a very vascular organ.

7. That blood vessels are composed of extremely thin and finely
striated connective tissue walls, without smooth muscle, occupying
canals in the bone substance.

8. That vascular expansions ‘occur in bone circulations and, by
their positions, suggest equalizing blood pressures and uniformity
in the circulation.

g. That developmental advances in the circulation determine types
of bone.

SMITHSONIAN MIS” ELLANEOUS COLLECTIONS VOL. 72, NO. 10, PL. 1

Cross section of the bone substance of the lower jaw of a Mas-
calonge, Esox, showing vascular channels.

Tangential section of a cranial bone of the Mascalonge, showing
parallel rows of small objects in the bone substance.

Cross section of the lower jaw of the Mascalonge, showing the
early differentiation of Haversian systems.

Tangential section of the inner ridge of the lower jaw of Masca-
longe, showing the circulation.

Cross section of the femur of a medium sized bullfrog, Rana
catesbeiana, showing the structure.

Longitudinal section of the femur of a medium sized bullfrog,
Rana catesbeiana, showing the looped vascular canals of the
inner wall. C, center of the shaft.

Cross section of the femur of an alligator, Alligator mississippi-
ensis, showing the structure.

Tangential section of the same, showing the circulation.

Cross section of the femur of a domestic turkey, Meleagris gallo-
pavo, showing the structure.

Tangential section of the same, showing the circulation.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL 72, NO. 10, PL. 2

19

Reconstruction of a second type bone with its circulation.

Cross section of the femur of a turkey buzzard, Cathartes aura
septentrionalis, showing the structure.

Tangential section of the same, showing the circulation.

Tangential section of the femur of a domestic chicken, Gallus, show-
ing the circulation.

Tangential section of the femur of a prairie chicken, Tympanuchus
americanus, showing the circulation.

Tangential section of the femur of a domestic duck, Anas domestica,
showing the circulation.

Cross section of the femur of a fruit bat, Pteropus (Celebes), showing
the structure.

Tangential section of the same, showing the circulation.

Cross section of the femur of a lamb, Ovis, showing the structure.

Tangential section of the same, showing the circulation.

Cross section of the femur of a Mexican burro, showing the structure.

Tangential section of the same, showing the circulation.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72, NO. 10, PL. 3

Tangential section of the femur of an elk, Alces americanus, showing
the circulation.

Tangential section of the femur of a Belgian hare, Lepus, showing
the circulation.

Tangential section of the femur of a bulldog, showing the circulation.

Cross section of the femur of a monkey, Macaca rhesus, showing the
structure.

Tangential section of the same, showing the circulation.

Cross section of the femur of a 9 months fetus (human), showing the
structure.

Tangential section of the same, showing the circulation.

Cross section of the femur of a white child 9 years old, showing the
structure.

Tangential section of the same, showing the circulation.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72, NO. 10, PL. 4

mbes Tae ay
tran a} Lp:
Steele 3

ht

i iB i

Cross section of the femur of an adult white male, showing the
structure. ;

Tangential section of the same, showing the circulation.

Tangential section of the femur of an adult white male, showing the
circulation.

Tangential section of the femur of an adult white male, showing the
circulation.

Tangential section of the femur of an adult white male, age 50, who
died of pulmonary tuberculosis, showing the circulation.

Cross section of a human frontal bone, showing the structure.

Longitudinal section of the same, showing the structure.

Tangential section of the outer table of the same bone, showing the
circulation.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72, NO. 10, PL. 5

Sea
: :

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Fes ASP ot eee a
1) Pay (, v
, E PALY Te

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Fic. 39. Entering vascular canals of outer table of a human frontal bone.

4o. Tangential section of the inner table of the same bone, showing the
circulation.

41. Tangential section of the outer table of a human parietal bone, show-
ing the circulation.

42. Tangential section of the inner table of the same bone, showing the
circulation.

43. Tangential section of the squamous portion of a human temporal
bone, showing the circulation.

44. Tangential section of the outer table of a human occipital bone, show-
ing the circulation.

45. Tangential section of the inner table of the same bone, showing the
circulation.

46. Vascular expansions filled with fat globules as seen in outer table of
occipital bones.

47. Tangential section of human inferior maxilla in front of and just
below left central incisor tooth, showing the circulation.

12, NO. 10, PL. 6

VOL.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 72, NUMBER 11

THE ECHINODERMS AS ABERRANT ARTHROPODS

BY
AUSTIN H. CLARK

E- INC,

LMT WIG
$e GE'A
hinges

(PUBLICATION 2653)

GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JULY 20, 1921

»*

The Lord Galtimore Press

BALTIMORE, MD., U. S. A.

THE ECHINODERMS AS ABERRANT ARTHROPODS
By AUSTIN H. CLARK

CONTENTS

PAGE
A eeesaaee ee ce See Paaothe Ceee a e Sa cc oe acta ke hie ccces bide wk due I
ihe dominant ‘characteristics of the echinoderms 3... ....00. 0. ce eee ees 2
Migewleevets Gt CGR OC MOM enis vc. De Une Le ciujaldodars Gels eins saalca ch(haveceaces 2
The change from mid-somatic to inter-somatic development in the crinoids. 2
Poe ueE Mons system Of Ee ECHINOGERMS toes. oo. arose ca cis spec va ceceecews 3
SOME CHIT O Ce Gimin Cel Omi ey ye rd a fie eh eee cl oa sale OURS aver, Gist oreo dial we eclo bs 5
SP MULE GE VAGEI AY GSU GhGI Marae Se voc celeste co atts deeleddessvieclashogoseans 6
Pee cEMMOUenM VASCIAG SYSTEM A. odd aciecasac ccs seneSecaesiccevancs oT)
Semen OCehine SIKCLETOM, < o hais eise eel sievecre.\ gs ce csc ads aieqess sees cece. 8
EA SSESECAT VS hia NOS" CRON (6 oR A le Ae ce rr 9
Pepe OES GO pEINE CHIMOLGAS sh 2i-' acs o ae cle saa is ole aed e.s'a eerdaleeedieleds cass 9
Characters of a hypothetical crinoid with mid-somatic development only. 10
Sa MIMS aC. He ®DATMACICS) © cies <a oe «c/o atacra os 6,0 sca‘ec os eielocug'e'ewe 0/4 10
hexcrinordssand) thesabertant) DATMACIES! A... 0 cles cic sislecic-cve'ds Guess ccs secs II
inl met aMAG Schl TE MGu Stat SHES. c/s/acct: s-ice diss 2.0 sys sth, ole's abe s lave dae icles leas 14
enersed WEGIINS ATIC ther DOIttle-StATS).v)00 ashes ced ce tcceie'ce es dees wbaeeae 17
The feeding habits of the echinoderms and of the crustaceans.......... sf
Professor Patten’s interpretation of the affinities of the echinoderms.... 18

PREFACE

Of all the larger animal groups there is none which has been the
subject of such diversified opinion regarding its affinities and sys-
tematic position as the Echinodermata: Originally assumed to be
related to the ccelenterates on account of their radial symmetry, the
echinoderms were later placed near the flat-worms, the annelids and
the chordates. At present they are regarded as representing a very
distinct and isolated group, some considering them as allied to the
chordates, while others emphasize their points of similarity with the
annelids; most zoologists, however, are reluctant to commit them-
selves regarding their probable affinities.

For many years I have been convinced that the echinoderms are
by no means such anomalous creatures as they appear to be, and that,
in spite of their many and extraordinary peculiarities, they are un-
doubtedly closely allied to the crustaceans, and especially to the bar-
nacles. In the following pages the reasons for this belief are given.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 72, No. 11

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

THE DOMINANT CHARACTERISTICS OF THE. ECHINODERMS

The echinoderms are very anomalous forms, and their relationships
to other animals are masked not only by a highly perfected radial
symmetry, but also by a unique development of all the organs of the
body. Their outstanding features are the presence of a vascular, a
respiratory, and a superficial skeletal system, the last composed of
articulated (calcareous) elements, the absence of gill clefts, and the
sharp division of the body externally into (five radial) segments.
In these features they agree only with the arthropods.

THE LARV#: OF THE ECHINODERMS

The unique larve of the echinoderms, which are of very varied
types, vermiform, bean-shaped with five ciliated rings and a long an-
terior tuft of cilia, auriculariz, bipinnariz, brachiolarie, plutei, etc.,
and differ more or less widely among themselves in the details of their
development, are always at first bilaterally symmetrical, which may
be accepted as an indication that the echinoderms are derived from
bilaterally symmetrical ancestors.

The larvze do not grow directly into the adults, but the latter for
the most part arise from new growth within the larval body, the
structures peculiar to the larvae being absorbed; in a few cases the
development is direct.

There is little in the structure or in the development of the echino-
derm larve which is comparable to the structure or to the develop-
ment of the larve of any other animals, and it is evident that the
extraordinary features exhibited by the adult echinoderms have been
projected so far forward in the ontogeny as quite to destroy the
value of the larve as phylogenetic indices.

THE CHANGE FROM MID-SOMATIC TO INTER-SOMATIC DE-
VELOPMENT IN THE CRINOIDS

Perhaps the most interesting feature connected with the mor-
phology of the crinoids, and one which it is necessary especially to
emphasize in order to understand the relationships between them and
the other echinoderms, is the abrupt change in the regions of bodily
growth and extension which takes place beginning with the formation
of the arms. It is this sudden change from interradial to radial, or
from mid-somatic to inter-somatic, development which occurs at
the commencement of arm formation that has always proved the
chief stumbling block in the way of a correct interpretation of these
animals.

NO. EI ECHINODERMS AS ABERRANT ARTHROPODS—CLARK &

In the echinoderms the primary appendages, the teeth, the water
pores, and the genital openings, and in the crinoids the primary
nerve cords as well, are all interradial in position. There can be no
doubt, therefore, that the dividing lines between the five half somites
fall in the middle of the so-called radial areas, and in the echinoids
and asteroids a sharp line of division is always maintained down the
middle of the radial series of plates throughout life, while no such
dividing line is found in the interradial regions.

In the young crinoids each of the somatic regions is completely
walled in by two large superposed plates, a dorsal basal and a “ ven-
tral’ oral; beneath the basals there are usually 3-5 small infrabasals
alternating with them and corresponding with the oculars of the
echinoids which, since they are entirely absent in large groups, and
are usually more closely associated with the column than with the
calyx, are probably to be interpreted as a dissociated columnal.

The young crinoid therefore has its body protected by ten large
somatic shields, five dorsal and five ventral, the latter with the primi-
tive appendage under the median line.

The arms first appear as evaginations in the intersomatic lines at
the plane of separation between the dorsal and ventral plates. The
evidence is that the skeleton of the arms is double, half being derived
from the somite on either side; but whatever may be the ultimate
genesis of their skeleton, the arms arise as linear and almost im-
mediately biramous appendages taking their origin from the inter-
somatic planes.

From this point onward the development of the animal is entirely
intersomatic ; the peristome and its underlying nerve ring, the water
tube ring about the mouth, the blood vascular ring, the genital cord,
and the ccelomic cavities all send off radial branches which increase
in length as the arm grows, while along either side of the peristomial
extensions (ambulacral grooves) there is formed progressively a con-
tinuous series of reduplications (ambulacral lappets) of the little
flaps with their associated tentacles, the latter in communication with
the radial water tube, which surrounded the mouth in the “ pre-
brachial” stage.

THE NERVOUS SYSTEM OF THE ECHINODERMS

In the chordates the central nervous system never becomes sep-
arated by mesodermal tissues from the tract of ectoderm from which
it originated in the embryo. Sedgwick remarks that this is a feature
of all echinoderms in so far as the ventral nervous system is con-

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

cerned, and when this nervous system is removed from-the surface
the removal is effected by invagination. |

But the only nervous system found in the echinoderms which in
its details is at all comparable to the central nervous system of the
other higher invertebrates is the so-called apical nervous system of
the crinoids, which first forms relatively late in life, and which ap-
pears to arise in connection with the coelomic epithelium.

It seems to me that this nervous system of the crinoids, which is
possibly (though not by any means probably) represented by the so-
called mesodermal nerve plexus in the starfishes, but which is quite
unrepresented in the other echinoderms, affords the best indication
of the probable affinities of these animals, and at the same time its
high state of development suggests that the crinoids have departed
less widely from the ancestral type than have the other classes.

In the developing crinoid the ectoderm of the surface of the body
becomes more or less disintegrated, and its cells to a greater or lesser
extent pass inward and intermingle with the cells of the underlying
mesoderm so completely that they can in no way be distinguished
from them, the body wall being formed of an ectoderm-mesoderm
complex in which the cells of the two types cannot be differentiated. ©

On the inside of this body wall, apparently in connection with the
coelomic epithelium, the apical nervous system arises; but in view
of the mixed nature of the component cells of this wall it seems not
illogical to assume that this apical nervous system is in reality
formed from cells which, originally ectodermal, have infiltrated
through the underlying mesoderm and now appear as if they belong
to the ccelomic epithelium. If this hypothesis can be accepted it is
obvious that the apical nervous system of the crinoids is in no way
comparable to the nervous system of the chordates, and as this is the
only echinodermal nervous system comparable with the nervous
system of other animals it naturally follows that no affinity with the
chordates can be inferred from the nervous system of the other
echinoderms. ;

In the crinoids the larvze become attached by the ventral side of the
anterior end, and the column is a development of the preoral lobe, aris-
ing therefore from the place where in the larve the anterior nerve
mass, just in front of the mouth, is situated. The mouth moves from
the ventral surface onto the left side and then migrates upward,
away from the point of attachment, until it comes to lie at the pole
opposite the latter, that is, at the posterior end near the anus.

NO. II ECHINODERMS AS ABERRANT ARTHROPODS—CLARK 5

The larval nerves disappear ; but some time after the torsion of the
animal is completed a new nerve mass forms in exactly the place
that the preceding nerve mass occupied, now become the point just
above the top of the column. It is thus natural to assume that the
central mass of the apical nervous system in the adult crinoid, ap-
pearing in the region previously occupied by the anterior nerve mass
of the larval crinoid, corresponds to it, and therefore that it repre-
sents the anterior nerve mass in other invertebrates. But as a result
of the torsion which.the animal has undergone the right ccelomic sac
has become extended anteriorly and, following the enteric wall, has
reached over onto the left side; from its anterior end it gives off five
anteriorly directed diverticula which at a later stage become cut off
and give rise to the chambered organ about which the central nerve
mass forms a close and almost complete investment.

The central nerve mass gives off five stout cords which immediately
branch, the two branches from each being connected by one or two
commissures, and then joining the similar branches from the nerve
trunks on either side to form the radial nerves of the division series
and arms. In each segment of the arms and pinnules, the nerve cord
gives off from a ganglionic swelling four branches, two dorsolateral
and two ventrolateral. In addition to these nerve cords, which may
be either single or double, and are in some types represented by two
widely separated parallel cords, each cirrus in its central canal con-
tains a prolongation from the chambered organ ensheathed in nerve
fibers continuous with those of the central nervous mass, and five
similar prolongations from the chambered organ and the central
nerve mass extend downward into the stem.

THE ECHINODERM CQELOME

Much has been made of the fact that in the echinoderms the
cceelome is enteroccelic in origin, as in the Brachiopoda, Chetognatha,
Chordata, and probably the Phoronida, while in the developing mol-
luscs, annelids and arthropods the ccelome is not enteroccelic in origin.

Sedgwick notes that in all the chordates except the tunicates the
coelome in its first state in the embryo shows more or less marked
traces of three divisions, the anterior or proboscis ccelome, which in
the Vertebrata and Enteropneusta is single and in Amphio.vrus double ;
the collar or middle ccelome, which is always double, and the trunk
ccelome, which is double and which in the Vertebrata and in Am-
phioxus becomes metamerically segmented. In the echinoderms
there seem to be indications, at least, of a similar tripartite division ;

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

there is the anterior ccelome, which is sometimes single (Asterina)
and sometimes double (Echinus), the hydroccele, which is probably
fundamentally double, though in some cases-(holothurians and cri-
noids) only one hydroccele sac is formed, and the posterior ccelome,
which is always paired. But in those chordates in which the entero-
coelic origin of the ccelome is clearly presented these three divisions
of it always come off from the enteron separately, while in the
echinoderms the enteron at most gives off only one pair of ccelomic
sacs; and whereas in the chordates the middle (collar) ccelome is
never more closely associated with the anterior than with the posterior,
in the echinoderms it is always closely associated with the anterior
ccelome, being developed from it and remaining connected with it by
the stone canal throughout life.

Whatever its origin the ccelome is clearly homologous in all these
types so that the manner of its development may be considered as due
to special mechanical or other limitations imposed by conditions in
the early stages—size, form, amount of yolk, etc—and not to phy-
logenetic causes.

THE WATER VASCULAR SYSDEM

The most extraordinary structure of the echinoderm body is the
water vascular system. This arises as a narrow dorsolateral out-
growth from a portion of the ccelome which unites with an ecto-
dermic infolding on the anterior aboral surface. From this develop
the stone canal and the madreporites. The ectodermic opening places
the hydroccele in communication with the exterior, so that the organ
has often been compared, in whole or in part, to an annelid excretory
organ or nephridium.

But Professor Patten has pointed out that it is much more like
one of the typical excretory organs of the arthropods (shell gland,
green gland, coxal gland) which consist of thin walled coelomic sacs
with a thick walled tubular outgrowth of varying length united to
a short duct infolded from the ectoderm.

The five primary tentacles or tube feet of the echinoderm larva
according to Professor Patten represent five modified thoracic ap-
pendages; an outgrowth of the underlying somite grows into each
appendage in typical arthropod fashion, but instead of breaking up
into separate muscles for the appendage it remains permanently in
the form of a membranous diverticulum of the hydroccele and becomes
the distal end of a radiating water vascular canal. Only the distal
end of the original appendage separates from the body as the primary

NO. II ECHINODERMS AS ABERRANT ARTHROPODS—CLARK 7

tentacle ; the remainder of the appendage, however long it may event-
ually become, may be regarded as lying in the surface ectoderm, de-
veloping on either side as it increases in length paired cirri which
become the double row of tube feet for each arm into each of which
a prolongation of the water vascular canal extends.

Primarily, then, each of the five primitive thoracic appendages, one
from each of the five half metameres of which the echinoderm body
is composed, contains a tube of nephridial intent leading into a canal
opening to the exterior by a pore. The anterior and posterior ends
of the series of half metameres join, and the excretory canal becomes
a ring canal from which grow out five long radial canals giving off
branches to the tentacles or tube feet as these are formed.

The excretory function of the water vascular system of the echino-
derms is reduced to a minimum if, indeed, it can be said to exist at all.
Its action is chiefly that of an hydraulic system whereby power origin-
ating in a great number of weak and dissociated muscle fibers scat-
tered along and within the water tubes and their branches is unified
and transmitted to the hollow tube feet, tentacles, and other appen-
dages, while at the same time the constant inflow and outflow of
water through the madreporic openings, especially when these com-
municate with the body cavity as they do in the gill-less crinoids,
undoubtedly serve to a greater or lesser extent the purpose of
respiration.

THE ECHINODERM VASCULAR SYSTEM

The vascular system, which is especially well developed in the
holothurians and echinoids, is formed of a peculiarly modified con-
nective tissue in which the fibers are sparse and which contains inter-
communicating spaces without an epithelial lining. The fluid in these
spaces does not appear to undergo any definite movement. Typically
there is a circumoral tract with radial prolongations which lie be-
tween the radial water vessel and the radial nerve cord, an annular
aboral tract in which the generative rachis is embedded and which
sends off extensions to the genital organs, and in holothurians and
echinoids a considerable development in the mesentery and on the
gut wall.

In the barnacles no heart is ever present, and the lacunar channels
in which the blood circulates are for the most part ill defined. ,

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

THE ECHINODERM SKELETON

MacBride and others have remarked that the presence of calci-
fied skeletal tissue in the mesoderm of the body wall is a character
found in the echinoderms and vertebrates only among ccelomate
animals. It does not seem to me that this can be considered as any
indication of affinity between these groups.

Many vertebrates have uncalcified mesodermal skeletons, and
others have only partially calcified skeletons. The uncalcified skeleton
of the notochord resembles in structure the parenchyma of the solid
tentacles of certain ccelenterates, and is quite different from anything
found in the echinoderms. Calcareous deposits of greater or lesser
extent occur in the mesoderm of barnacles, brachiopods, rotifers and
cestodes, as well as in the mesoglcea of sponges, and the calcareous
skeletal structures of some ccelenterates are mesodermal in origin.

In certain of the early cystideans the skeleton appears to have
been wholly or chiefly chitinous, and their surface exactly resembles
that of the phyllopod and other crustaceans preserved in the same.
rocks. Itis not improbable, therefore, that the calcareous exoskeleton
of the echinoderms of the present seas developed from a chitinous
body covering through an exoskeleton composed of chitin with an
increasingly greater amount of inorganic matter such as we see today
in most of the larger crustaceans.

In the developing crinoid the ectoderm of the surface of the body
more or less completely disintegrates and its component cells largely
pass inward and intermingle with the cells of the underlying meso-
derm, so that in the crinoid the outermost layer of the body is almost
as much mesodermal as it is ectodermal. This. being the case, no
matter what its phylogenetic relationships and tendencies are, the
formation of an ectodermal skeleton has now become impossible as
there is no continuous ectoderm from which to form it. A cal-
careous mesodermal skeleton appears, the first rudiments of which
are formed in the deeper layers but soon move to a more superficial
position enclosing the body in a calcareous investment formed of large
and definite plates. Just before the appearance of the arms there are,
in addition to the columnals, 13-15 thin cribriform films lying just
below the surface and fitted edge to edge, including 3-5 infrabasals,
5 basals alternating with them, and 5 orals superposed upon the
latter. ;

Now although this skeleton is mesodermal and calcareous, the re-
lations between it and the enclosed body of the animal are entirely
different from the relations‘ between the vertebrate skeleton and the

NO. II ECHINODERMS AS ABERRANT ARTHROPODS—CLARK 9

body organs, being on the contrary quite like the relations between
the chitinous or more or less calcified skeleton of the arthropods
and their enclosed body. The crinoid skeleton is a superficial
(though not external) skeleton enclosing the body and giving off
articulated appendages; it therefore resembles the skeleton of the
arthropods more closely than it does that of any other animals. The
fact that it is mesodermal and calcareous seems to me to be, in view
of its development in every way like an ectodermal chitinous exo-
skeleton, and especially in view of the fact that it lies outside of the
ventral nervous system which runs along or just within its inner
surface, of purely secondary significance.

Crinoids are undoubtedly descended from animals with an ar-
ticulated exoskeleton, and their articulated superficial skeleton is
calcareous instead of chitinous as would be expected solely because
of the disintegration of their ectoderm in the young stages.

In the arthropods we find in the sessile barnacles the beginnings
of a transition from a chitinous exoskeleton to a calcareous mesoder-
mal superficial skeleton, and from the conditions in these animals it
is not difficult to supply the connection between the crustacean and
the crinoid skeleton.

AUTOTOMY IN THE CRINOIDS

The crinoids with more than ten arms increase the number of their
arms by breaking off the larval arms at the base, the stump forming
an axillary from which two or more arms arise. This is primarily
due to the inability of the brachial skeleton, a rigid calcareous invest-
ment of the dorsal and dorsolateral portions of the arm, to keep pace
with the other brachial structures in development, and is therefore
distantly comparable to the moulting so characteristic of the
arthropods.

THE APPENDAGES OF THE CRINOIDS

The appendages in the crinoids are of two kinds. From the base,
and always in connection with the chambered organ and the central
nerve mass of the apical nervous system, arise uniserial jointed ap-
pendages ending in a strong hook. These are especially developed
in crinoids unprovided with a stem, and serve both as tactile organs
and for attachment. In their position as anterior organs and in
their function as tactile and grasping organs, as well as in their uni-
serial structure, they recall the antennz of the barnacles.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

About the ventral surface there are five long arms typically forking
at the base, thus representing biramous ventrolateral appendages and
calling to mind the biramous thoracic appendages of the barnacles.

There can be no question of any direct homology between the cirri
of crinoids and the antenne of barnacles, or between the thoracic
appendages of barnacles and the arms of crinoids, but both sets of
organs have the same functions and the same location in each group,
and are constructed on a similar plan, so that it is not impossible to
regard them as parallel manifestations of the same ancestral ap-
pendicular plan, a plan not occurring in the animal kingdom outside
of the arthropods and crinoids.

CHARACTERS OF A HYPOTHETICAL. CRINOID WITH MiB
SOMATIC DEVELOPMENT ONLY

Let us imagine a crinoid with entirely mid-somatic development
and with entire instead of half somites, that is, with bilateral instead
of radial symmetry. We would have a body composed of five broad
somites each covered with a broad arched plate (tergum) to the edge
of which is articulated a flap (oral, corresponding to a pleuron) ;
within this would be the radial (epimeron) at the inner edge of
which arises a biramous appendage. A body with five terga and five
pairs of biramous appendages with their bases covered by pleura
from which they are separated by epimera would certainly be con-
sidered as crustacean in character, and if it were attached by the head
end, with the mouth upward, it would unhesitatingly be pronounced
a barnacle, its deficiencies and anomalies of organization being as-
cribed to. degeneration.

THE CRINOIDS:AND THE BARNACLES

The crinoid develops from a highly anomalous larva, with a so-
called vestibule suggesting a partial development of a bivalved cover-
ing, which attaches itself by the anterior end like the cypris larva of
a barnacle and turns a half somersault bringing its mouth upward
and opposite the point of attachment, also like a barnacle; so far its
development equally well suggests that of a polyzoan from a cypho-
nautes larva; but in its further growth it develops a superficial
skeleton as does an arthropod of a sort already seen in rudimentary
form in the barnacles, with the chief nerve cords, which are highly
developed and arthropod-like, running over or just within its internal
surface as in the arthropods, and uniramous anterior (oriented from
the central nerve mass) and biramous ventrolateral appendages, °

NO. II ECHINODERMS AS ABERRANT ARTHROPODS—CLARK TEL

both sets of which have functions similar to those of the correspond-
ing appendages in the barnacles.

Thus in spite of the utter dissimilarity in the details the broader
features of the structure of the crinoids and the barnacles as well as
of their development are seen to be similar, or at least comparable,
or perhaps it should be said that there is less divergence between
crinoids and barnacles than there is between crinoids and any other
organisms.

THE CRINOIDS. AND THE ABERRANT BARNACLES

The crinoids represent a derivative from a branch of the same
arthropod stock that gave rise to the barnacles, but they have gone
much further; half of each of the five segments of which the body
is composed fails to develop so that the body is composed of five
half segments joined in a circle with the central organ of the ventral
nervous system at one pole and the mouth, which has moved poster-
iorly and come to lie near the anus, at the other; the development
of the body structures after early youth suddenly becomes entirely
inter-somatic instead of mid-somatic ; the peristomal region becomes
enormously enlarged and extended, resulting in the formation of a
sort of lophophore ; and the appendages have been suppressed, or at
least appear in a very modified form late in life. Certain hereditary
tendencies show themselves after the animal has, so to speak, re-
covered from the profound ontogenetic shock resulting from the
loss of half its body, in the appearance of articulated uniserial tactile
and grasping organs at the neural pole (the original anterior end)
and of articulated biramous appendages used for food gathering or
for locomotion along the ventrolateral border.

Crinoids, like barnacles, are sessile, pedunculated, attached by
hook-like processes, or unattached.

The strong probability that the arthropod stock which by profound
modifications gave rise to the barnacles also gave rise to the crinoids
is indicated not only by the asymmetry in the Verrucide, in which
the operculum consists of the scutum and tergum of one side only,
those of the other side being fused to form one half of the wall which
is completed, on the side of the movable opercular plates, by the
greatly developed and displaced rostrum and carina, but also by the
anomalous parasitic forms which have developed among the former
in which the aberrant features are so fundamental that they have
been thrust forward into the ontogeny so far as to modify profoundly
the form and structure of the nauplius. These forms also show that

12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

the occurrence of complicated ramifying. roots, so highly developed
in some crinoids, is an inherent possibility in the barnacles.

A combination of the asymmetry of the Verrucide (inherent also
in very many other crustaceans, and especially noticeable in the .
Paguride and Bopyride) carried to its logical conclusion in the com-
plete atrophy of one side, with the modifications of the body seen in
Spherothylacus or Sarcotaces in a less extreme form, the roots of
the Rhizocephala, and a skeleton formed after the manner of the
plates in the shell of the Operculata, furnishes all the elements
needed for recombination to form the crinoid. It may be well to
call attention to the fact that outside the Cirripedia there is no group
in which all the morphological peculiarities exhibited in the crinoids
coéxist—indeed they are not to be found in all the rest of the animal
kingdom together.

In this connection it may be worth while to review the salient
features of the more important of the aberrant barnacles.

The Rhizocephala are exclusively parasitic barnacles, the most
degenerate of all parasites; in the adult stage they are distinguished
from normal barnacles by the entire absence of all traces of segmen-
tation and of appendages, and at all stages they lack an alimentary
canal; every trace of arthropod organization has disappeared. Nearly
all of them occur on decapod crustaceans.

The body has the form of a simple sac, or may be divided into
numerous similar sacs, attached by a short peduncle from which
root-like processes ramify throughout the body of the host; these
absorptive roots appear to be absent in the aberrant genus Duplorbis.
The body proper is completely enveloped by the mantle which usually
has a narrow aperture capable of being closed by a sphincter muscle ;
in Sylon the opening is double, and in Thompsonia, Clistosaccus and
Duplorbis the mantle cavity is completely closed.

The mantle commonly is attached to the visceral mass by a narrow
mesentery near which on either side are the paired (more rarely
unpaired) openings of the male and female genital organs. In the
different genera the external form varies considerably, and with it
the position of the mesentery and of the genital apertures.

Thompsonia, the most aberrant and highly specialized of all the
parasitic barnacles, consists of nothing but a diffuse system of branch-
ing and sometimes anastomozing mycelium-like roots continuous
throughout the body of the host and all arising from a single original
larva; the peripheral division of the root system passes out into the
walking legs, abdominal swimmerets and tail fans and there gives
rise to numerous (up to more than 500) small sacs consisting of a

NO. II ECHINODERMS AS ABERRANT ARTHROPODS—CLARK 13

thin mantle without muscles and containing only an ovary, without
generative ducts, testis or nerve ganglion. During development the
visceral mass disintegrates so that at the time of hatching the mantle
contains a great number of cypris larve ready to emerge. The
escape of the larve is contemporaneous with, or soon followed by,
a moult of the host. The empty shells of the external sacs are carried
away with the cast skin, and the terminal swellings of the root system
emerge as a new crop of external sacs.

In Peltogaster the body has an elongated sausage shape, with the
mantle opening at one end, and is attached by the peduncle about the
middle of its length. The mesentery is longitudinal on the proximal
side (next the peduncle).

In Sacculina the whole body is flattened in the plane of the mesen-
tery and has assumed a secondary and superficial bilateral symmetry
about a plane at right angles to this and coinciding with the median
plane of the host. In other genera, such as Lern@odiscus and Tri-
angulus, the symmetry becomes still more complicated.

In Clistosaccus and in Sylon the genital organs are unpaired.

The peduncle perforates the integument of the host and gives off
on the inside the absorption roots which, in the case of Sacculina,
penetrate into all the organs of the host with the exception of the
gills and heart, and extend to the terminal segments of the legs and
into the antennules and eye stalks. In Duplorbis, in which the root
system appears to be absent, the peduncle is hollow, its cavity com-
municating with the closed mantle cavity and opening at the other
end into the body cavity (hemoccele) of the host.

Apart from a single nervous ganglion (absent in Thompsonia)
which lies close to the mesentery near the female genital openings,
the only organs present are those of the generative system.

Spherothylacus is parasitic on a simple ascidian (Polycarpa), liv-
ing attached by ramifying roots to the inner wall of the branchial
sac. The globular body is enclosed in a mantle which has a small
opening. There are no appendages, but there is a complete alimen-
tary canal with mouth and anus, the latter near the mantle opening.

The two known species of Sarcotaces live embedded in the muscles
of fish; an alimentary canal is said to be present, and there are no
roots.

In the Ascothoracica, all of which are parasitic in Zoantharia or
in echinoderms, the mantle may have a bivalved form (Synagoga
and Petrarca), or it may form a capacious sac (Laura) much larger
than the body with which it is connected by a narrow neck and havy-
ing only a small opening to the exterior. In Dendrogaster the mantle

14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

is still more developed and is produced into branched lobes. In
Laura the mantle is covered with stellate papilla penetrating the
tissues of the host and presumably absorptive. In all cases the mantle
contains ramifications of the enteric diverticula and portions of the
gonads. In Laura the body is divided into six “ thoracic ” and three
limbless “abdominal” somites, and ends in a caudal furca. In
Petrarca and in Dendrogaster the body is unsegmented.

In these three genera a pair of preoral appendages is present and,
except in Laura, are armed with hooked spines suggesting that they
are organs of fixation. They are inserted, at least in Laura, at the
sides of the buccal region, and are more or less enveloped by the
mantle.

The cement glands appear to be absent, and the mouth parts are
more or less reduced, but appear to be adapted for piercing.

The thoracic appendages are biramous and articulated only in
Synagoga. In Laura they are uniramous and indistinctly segmented,
and the first pair are long and slender. In Petrarca they are still
further reduced, and in Dendrogaster they are represented only by
some indistinct papillz.

In all three genera the gut ends blindly, and the hepatic diverticula,
which are large, extend into the mantle. The nervous system is
reduced. An eye is said to be present in Synagoga. In Laura the
oviducts open at the base of the first pair of cirri.

The larva of Laura is a nauplius lacking the frontolateral horns ;
in Dendrogaster the larva hatches as a peculiar cypris with only five
pairs of biramous thoracic limbs.

In the Apoda the curious Proteolepas bivincta is elongated and
maggot-like, with no trace of a mantle nor of appendages other than.
a pair of adhering antennules. The mouth parts, borne on the first
“segment,” seem to be adapted for piercing and sucking. The ali-
mentary canal is greatly reduced; according to Darwin only the
cesophagus is present, and there is no trace of stomach, rectum or
anus.

THE CRINOIDS AND THE STARFISHES

In the crinoids during their development the mouth moves from
the ventral surface onto the left side, indenting the left hydroccele
and the left posterior coelome, and continues its migration until it
comes to lie at the posterior end beside the anus. When it has
reached this point the hydroccele ring closes. As a result of this
movement the right posterior ccelome has also shifted and come to

NO. II ECHINODERMS AS ABERRANT ARTHROPODS—CLARK 15

lie on the aboral (originally anterior) side of the gut and the preoral
lobe of the larva becomes enclosed by the rows of skeletal elements
(apical plates) which are developed outside the right posterior
coelome in all echinoderms except holothurians ; these plates are first
laid down in a horse-shoe shaped ring which later closes, as does
the hydroccele, to form a complete ring.

In the asteroids the closure of this curved row of plates is effected
far from the point of origin of the preoral lobe on the right or right
dorsal side of the larval body; in the crinoids it is effected at the
anterior end of the larval (posterior pole of the adult) body and
encloses the preoral lobe just as the hydroccele does in asteroids.

The larve of both crinoids and asteroids attach themselves by
the preoral lobe ; but whereas in the crinoids the preoral lobe is quite
free of the circumoral vessel and arises from the apical or aboral
surface of the adult, in the asteroids the preoral lobe is encircled
by the water vascular ring and its withered vestige springs from the
oral surface of the adult disc.

In the asteroids the mouth has shifted from the ventral surface onto
the left side of the body, but has gone no further. Since the preoral
lobe disappears as an appendage from the oral surface within the
hydrocceele ring it is evident that the apical nervous system of the.
crinoids, which appears to be intimately connected with the preoral
lobe, must be represented by a ventral nervous system in the
asteroids.

In the crinoids the circumoral nerve ring and its extensions be-
neath the ciliated ambulacral grooves of the disc, arms and pinnules
is associated only with the latter and with the ventral surface of the
tentacles. In the asteroids the similarly situated ectodermal ner-
vous tracts are in connection with a diffuse ectoneural plexus found
throughout the ectoderm and at the mouth with an endoneural plexus
which is the central portion of the so-called endodermal nervous
system. The deep oral nervous system, consisting of a double cord
in each radius just within the radial nerve thickening of the ectoneu-
ral system and centering in a more or less complete ring about the
mouth, said to be exclusively motor in function, possibly corresponds
to the deep oral system of the crinoids which consists of paired cords,
one on either side of the water tube ; but the latter is connected with
the apical nervous system and the former is not. The apical nervous
system of the asteroids, motor in function, consists of a cord in the
mid-radial line of each arm and appears to develop from the dorsal
peritoneum with which it remains in continuity. It differs from
that of the crinoids in being radial (intersomatic) instead of inter-

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

radial (mid-somatic) in position, in consisting of single cords in-
stead of paired cords with commissures, and in lacking a well de-
veloped central organ. From its lack of any relation to the preoral
lobe and its general indefiniteness of structure it is difficult to see how’
it can be in any way homologous with the apical nervous system
of the crinoids. It is probably a special feature peculiar to asteroids.

The apical nervous system of the crinoids, from which the whole
animal with the exception of the peristomal region and its extensions
is innervated, is absent in the asteroids, or rather it has become merged
into the circumoral structures and their derivatives.

It has been shown that the crinoids before the development of the
arms are encased in plates developed over the somatic divisions, but
that beginning with the appearance of the arms the development be-
comes wholly intersomatic or radial. In the asteroids the develop-
ment is radial from the first appearance of the plates, the interradial
(mid-somatic) body covering seen in the young crinoids not appear-
ing at all.

In the crinoids there are at first no plates belonging to the ventral
surface, the basals being dorsal and the orals ventrolateral, but the
latter become ventral plates after the formation of the arms, which
themselves are composed of a series of dorsal ossicles carrying exten-
sions from the ventral structures on their ventral surface.. In the
asteroids the plates at their first appearance represent those of the
crinoids after the formation of the arms as far as their bifurcation,
minus the orals. There is a central plate, corresponding to the infra-
basals in the crinoids ; about this are five basals, corresponding to the
five basals of the crinoids; beyond and alternating with these are five
terminals, corresponding to the radials of the crinoids, but always
single and showing no indications of a primarily paired condition as the
crinoid radials do. On the opposite (ventral) surface are five pairs of
plates, one pair in each radial division, corresponding to the first two
post-radial plates in the crinoids, but side by side instead of tandem.

Thus whereas the extensions of the crinoid body—the arms—are
just on the border between the dorsal and ventral surfaces and are
composed dorsally of dorsal ossicles and ventrally of extensions of
ventral structures, in the asteroids the dorsoventral edge of the body
has moved dorsally so that the dividing line between the dorsal and
ventral surfaces falls between the radials and the succeeding plates,
and the whole ventral surface is encased in plates which are repre-
sented in the crinoids on the dorsal surface only.

The asteroids therefore differ from the crinoids in the temporary
attachment of the larvee; in the relatively slight alteration in the posi-

NOs. Tal ECHINODERMS AS ABERRANT ARTHROPODS—CLARK 17

tion of the mouth; in lacking the apical nervous system, which has
either completely disappeared or has become transformed into a
ventral nervous system centering in a ring or rings about the mouth;
in omitting the early mid-somatic development, the development of the
body being inter-somatic from the first; and in having moved the
border between the ventral and dorsal surfaces dorsally so that all
the post-radial plates now lie on the ventral surface, the arms being
formed by an extension of the body in the plane dividing the crinoid
radials from the plates succeeding.

THE SEA-URCHINS AND THE BRITTLE-STARS

The echinoids and ophiurans differ still more widely from the
crinoid type. Their larvze, except in special cases, are extraordinarily
developed plutei which have no attached stage at all, and are char-
acterized by the small size of the preoral lobe, by the great development
of the post-anal portion of the body, and by the possession of a special
larval skeleton supporting the arms which is later resorbed.

Whereas the asteroids differ from the crinoids in transferring the
post-radial plates from the dorsal to the ventral surface and thereby
forming a ventral skeleton of primarily dorsal elements, the echinoids
have gone further and have eliminated the dorsal surface altogether
except for a ring of plates about the periproctal region and the small
area within it, the globular body being composed of plates representing
the ventral plates of the asteroids. The perfection of an entirely new
type of compact radially symmetrical body from the crinoid through
the asteroid, simulating the compact radially symmetrical ccelenterate
body, has furnished a starting point for new development, and bilateral
symmetry, superposed upon the perfected radial symmetry, has reap-
peared and in some cases (as in Pourtalesia) has been carried to an
extreme.

The ophiurans are phylogenetically parallel to the echinoids, but
their line of specialization is entirely different. In them the relation
of the dorsal to the veritral surface has remained as in the asteroids,
but the radial (intersomatic) extensions of the body have become
narrowed and consolidated into highly efficient jointed appendages |
from which all non-essential structures have been eliminated.

THE FEEDING HABITS OF THE ECHINODERMS AND THE
CRUSTACEANS
Corresponding with the progressive specialization in their structure
it is interesting to note a progressive specialization in the feeding
habits of the echinoderms. The crinoids are plankton feeders, like the

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

barnacles. The starfishes are largely carnivorous, feeding especially
upon molluscs, but some swallow mud out of which they digest the
organic matter. The echinoids feed upon vegetable matter or upon
organic detritus, and many are mud or sand swallowers. The
ophiurans feed largely upon detritus or swallow mud, but many are
ectoparasitic upon coelenterates and crinoids from which they steal
the food in or on its way to the stomach.

Thus along with the widening of the gap between the structure of
the true barnacles and that of the echinoderms there is a similar
divergence in their feeding habits ; from plankton feeders they become
simply scavengers and parasites.

The recent crustaceans as a whole show exactly the same line of
specialization from the phyllopod, which feeds on minute organisms,
to the decapod, which feeds largely on carrion or detritus, or is
ectoparasitic on ccelenterates or on crinoids, and this development of
the feeding habit, with the emphasis on the scavenging activities, is
characteristic of these two groups alone.

PROFESSOR PATTEN’S INTERPRETATION OF THE AFFINITIES
OF THE ECHINODERMS

Although I arrived at the conclusion that the echinoderms and the
arthropods are in reality closely related more than ten years ago and
in a short paper published in April, 1910, that conclusion is readily
to be inferred, Professor William Patten was the first to attempt to
explain the relationships of the echinoderms to the arthropods in
detail. His reasoning is so entirely different from mine that it is
worth while to repeat his arguments here.

He says: “ The echinoderms are notable for their contrasts and con-
tradictions. Their outward appearance and their pronounced radial
structure distinguish them from all other animals, and at first
sight suggest a very primitive organization similar to that of the
coelenterates. On the other hand they display a high degree of his-
tological and anatomical specialization that is in marked contrast with
their low grade of organic efficiency. They begin their early em-
bryonic development with a bilaterally symmetrical body and with clear
indications of metamerism, only to change it in the later stages for one
that is radially symmetrical and in which all outward traces of
metamerism have disappeared. After a short free-swimming larval
existence they attach themselves, neural side down, by means of larval
appendages and a cephalic outgrowth; they then turn neural side up
and remain so attached for life; or in some cases they give up their

NO. II ECHINODERMS AS ABERRANT ARTHROPODS—CLARK 19

sessile existence and again become free, moving slowly about, neural
side down. There are, therefore, three chief characteristics of the
echinoderms that demand our first consideration: (1) The early
bilateral symmetry and metamerism; (2) the sessile life and mode
of attachment by cephalic outgrowths; and (3) the asymmetry.
There appears to be but one explanation for these remarkable condi-
tions, which is as follows: The early development of bilateral sym-
metry and metamerism in the echinoderms, and the presence of a
telocele and telopore in place of the more primitive gastrula and
blastopore, clearly indicate that they had their origin in bilaterally
symmetrical animals of the acraniate type that had already acquired
a considerable degree of complexity. These ancestral forms prob-
ably belonged to the cirriped group, for before the latent asymmetry
becomes effective the young echinoderm larva resembles a cirriped in
its form, mode of attachment, and subsequent metamorphosis more
than it does any other animal. The radiate structure of the later
stages was due to a persistent local defect, or to the absence of a
definite part of the embryonic formative material, which in turn
created a condition of unstable equilibrium, the result of which is that
the whole side, following the path of least resistence, bends toward
the defective area, forming an arch that increases in curvature until an
approximate equilibrium is again attained by the union of the two
ends to form a circle. The original half metameres and segmental
organs are then arranged in radiating lines, thus creating a new
radiate type and a new set of internal conditions that dominate the
future growth of the organism. If we assume that a strongly marked
asymmetry, such as that which occurs so frequently as an abnormality
in Xitphosura, or even as a normal character in the Bopyride and
Paguride, was a fixed feature of the hypothetical ancestral cirripeds
and was capable of a successful organic adjustment, we shall have a
perfectly simple and natural explanation of the origin and structure
of the echinoderms.

“The young asteroid larva is said to attach itself voluntarily at
first, and for a short time only ; later it becomes permanently attached,
head first and neural side down, in the same remarkable manner as a
young cirriped, both the cephalic appendages (which are thick walled
and muscular, with a long basal portion and a short terminal knob
studded with small adhesive papille, greatly resembling the minute
adhesive antenne of the cirripeds and parasitic crustaceans) and the
adhesive disc taking part in the process. The young crinoid larva
attaches itself wholly by means of the cephalic disc, as the adhesive

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

appendages appear to be absent. Its first position is. with the neural
or oral surface down, as in the cypris stage of the cirriped. The disc
then elongates, forming a slender cephalic stalk or peduncle, and the
larva turns a somersault, bringing its neural side uppermost. Mean-
while the vestibule, or peribranchial chamber, which at first is small
and temporarily closed, enlarges, then ruptures, and the five ap-
pendages project from the cup-like head in typical cirriped fashion.
In certain of the representatives of the recent echinoderms, such as
the asteroids, the fixed stage is temporary, while in certain others,
such as the echinoids and holothurians, it appears to be omitted alto-
gether and the young echinoderm, after its metamorphosis, again
acquires a limited power of locomotion. But in most primitive echino-
derms, such as the stalked crinoids, blastoids and cystideans, a perma-
nent attachment by an elongated cephalic stalk, in typical cirriped
fashion, was the almost invariable rule, and no doubt represented the
primitive condition for the whole class. When an echinoderm does
become free it acquires only a very limited power of locomotion and of
coordinated movement. Its characteristic lack of efficiency in this
respect is due not so much to its simple structure as to the fact that
its freedom was gained at a late period in the phylogeny of a very
ancient group in which sessile inaction was the prevailing condition.
It is often assumed that a sessile or parasitic mode of life is the initial
cause of degeneration. The various anatomical peculiarities common
to the copepods, cirripeds and acraniates do not bear out this con-
clusion. The fact that in these diverse subphyla we see the same shift-
ing of cephalic appendages to the hzemal side, the same cephalic out-
growths, and the same degeneration of the neuro-muscular organs,
indicates that there are certain initial defects or peculiarities of
germinal material common to the whole group, and that these are the
underlying cause of defective organization, the defective organization
being in every case of such a nature that a sessile or parasitic or
vegetative mode of life is the only one possible.”

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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 72, NUMBER 12

A STUDY OF THE BODY TEMPERATURE
OF BIRDS

BY
ALEXANDER WETMORE

(PUBLICATION 2658)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 30, 1921

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Pes huyOr THE BODY:TEMPERATURE OF BIRDS
By ALEXANDER WETMORE,

BIOLOGICAL SURVEY, U. S. DEPARTMENT OF AGRICULTURE

| CONTENTS ates
nr ineaie Redvers ana t Pae eo ere roe Wee RR ee 2 Sie cen adr a Ba'e dere = iors hele I
Methoduoheseccunine: avian temiperaiiiheSia tis cie 6. -vlss.< aici wic/ere gene ae) ose ceeres 3
Diurnal rhythm in temperature.......... Ge Renee daa at ee Pt ee 5
Table 1. Time of maximum and minimum temperatures in birds..... 5
Weaniahton ni gemperattinesihi relation £0) SEX... casi «ms. = ciel treleisine «cles 2 lela a 9
Exrepnaleremperattine: inn relation. to. bodily heat..s.2.2sc..0s- sce oe ee deans 12
Diverse miscellaneous factors in their relation to body temperature....... 16
1) SoH EsPELEWCAE (aR Non C1 ANS EO 5 OP a reer 18
Table 2. Temperatures of young or nestling birds................... 19
Methodroretempenatiure: control imebirds.. sac. s)sc-.)sis esis. aeeie's vis 0 ove wvegoi os 21
SHoCMAcHMECEOl tempehatliter COMO a. vce a4.» dew fe slo ed tuelelees ches wieyewlere ya « 28
Discussion of differences in average temperatures...........0.ec eee eee nee 30
ECAC CUO Mente LEGS meas Me saree etc aie eros) a cis oy aire che Rr ayinre Sorevauaersrtoyeline Gh ele eye as 22
Maples. sSoummeatys on temperature necOrdS...0..--.2..:5+ccceseses- 36

Table 4. Average temperature of families of birds, summarized from
WANDA, 6 5 AWS cro’ oul rok eon td ois CIEL MELE RECESS ie rE ene ee renee 47

Table 5. Temperatures of species of birds not included in Table 3,
LAKCHEnGOMMiaviell Ale MIELE: ateNa.1,ctorste-che @#/eleve alee eieyelnve cree cls 48
LED OSEARE NOURI. “ey GA que Ciel Be a Ocoee co CNTR OREN ee 51

INTRODUCTION

The subject of the body temperatures of the many and varied
species that compose the great vertebrate class of birds is one that
in the past has been rather slightly treated by those interested in
avian physiology. Statements to the effect that the bodily tempera-
tures of birds are higher than in others in the group of classes com-
posing the Vertebrata, are current in many zoologies and text books,
but on the whole, literature gives few definite statements of fact on
the subject, and observations have been restricted to a comparatively
small number of species. In the course of other work many hundreds
of birds have been handled in the flesh by the present writer, and
after some thought methods were devised for the taking and record-
ing of body temperatures in as accurate a manner as possible. This
was a little known field of endeavor and at first no guides as to method

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 72, No. 12

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

were available, so that mistakes were made, and early records had to
be abandoned. Continued experimentation led to definite methods
that promised accuracy, and it is upon these that the following account
is based. The results of studies covering the period from January,
1912, to October, 1919, are included herewith, giving a total of 1,558
records covering observations on 327 species of birds distributed
among 50 families. If the number of observations seems small
compared to the length of time involved it must be considered that
the taking of temperatures was not always feasible as the difficulties
attending the task were manifold and varied.

The results of this work are embodied in a series of tables which
form the foundation upon which this report is based. In addition
to the 327 species of birds on which observations regarding body
temperature have been made by the writer personally, a supplementary
compilation has been given (in table 5) embodying published records
on this subject for other species, in order to render this account of
the body temperatures of birds as complete as practicable. In this
final table records are given for 89 forms, so that in the following
pages may be found definite statements regarding the temperatures
of 416 species of birds. It is hoped that the work of securing such
records may be continued until a much greater amount of informa-
tion covering many more species is available. :

It was the original intention to incorporate in this paper a table
giving in detail the individual records upon which the work has
been based. Such a course was desirable as it would have furnished
a mass of data far greater than any hitherto available for those who
may be interested in using such information in lines of research other
than that in which it has been utilized here. In addition it would have
given opportunity to check up more carefully the deductions that I
have made. Cost of printing of tabular matter has proved so high
at the time of publication that it has been necessary to omit such a
detailed statement and to supply the records only in a condensed form.
The table itself is, however, deposited in the files of the Smithsonian
Institution and may be consulted by those interested in using it.

From his studies in the subject of avian body temperatures the
writer has ventured to come to certain conclusions and deductions,
some in accordance and some at variance with modern ideas as previ-
ously accepted. If these seem sufficiently substantiated to meet the
approval of others, then may the time and labor required in the
compilation of these data be deemed justified. The following pages
are respectfully submitted for attention, with the hope that part if

NO. 12 BODY TEMPERATURE OF BIRDS—WETMORE 3

not all of this labor may gain the stamp of approval from those who
may be interested in such lines of research.

In carrying on these investigations the writer has been indebted to
the Smithsonian Institution for a grant of seventy-five dollars from
the Hodgkins Fund, for the purchase of specially constructed ther-
mometers required, and for certain other minor expenses incurred.

METHOD OF SECURING AVIAN BODY TEMPERATURES

In recording the body temperatures of birds it was necessary to
work with thermometers that permitted a considerable range in regis-
tration as it was found that the degree of bodily heat was high or
low according to the species in hand. All records were made in the
Fahrenheit scale by means of clinical thermometers similar in form
to those used by physicians. For the present investigation it was
necessary to have these instruments specially made to give accurate
registration ranging from 95°+ to 115°+. Ordinary clinical ther-
mometers do not register above 110°+ so that they could not be
utilized, for certain passeriform species frequently have temperatures
ranging above that figure.

While engaged in field work these special thermometers were
carried constantly in suitable carrying cases. When a bird was shot
a temperature reading was taken when the specimen could be reached
immediately. If there was delay in retrieving for any reason, an
accurate temperature record could not be made, so that it was possible
to secure records in less than one-half of the birds that were collected.
In taking temperatures of specimens of small or medium size the
thermometer was placed in the opened mouth of the bird, and worked
down until the lower end was entirely within the cavity of the thorax,
usually until it reached the proventriculus. With birds of larger
size the reading was taken through the anus, with the thermometer
thrust in through the cloaca well into the canal of the large intestine.
In either case it was necessary to penetrate the body cavity to secure
a correct reading. The peripheral circulation of birds is poor, so
that often there may be a difference of two to five degrees between the
temperature of the upper part of the oesophagus and that of the body
cavity proper. A similar variation was noted between the tempera-
ture of the cloaca and the large intestine above it.

The thermometers used were self-registering, and were so con-
structed that they reacted immediately to any increase in heat. The
highest point of a reading was reached in a very few seconds, but
to insure accuracy the instrument was held in position for a period

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

ranging from half a minute to a minute. After each reading the
thermometer was shaken down to be ready for instant use when
needed again.

Temperatures were read to tenths of a degree, and were tran-
scribed at once in a note book opposite the name of the bird, together
with the date and other needed information. In each case the time
of day was noted to the nearest quarter hour. Thus a temperature
taken at five minutes past ten in the morning was recorded as 10.00
a.m. and one at ten minutes past ten as 10.15 a.m. Further refine-
ment in recording the time of day was considered useless, as save on
certain meridians there is always slight variation between the actual
time as shown by the height of the sun, and the time adopted for
universal local use as a matter of convenience. The time of observa-
tions recorded during the continuance of the so-called “ daylight
saving’ regulations of 1918 and 1919 has been corrected in each
‘instance to normal.

In order to be prepared at all times to secure temperature records
of birds it was necessary to have thermometers constantly at hand
while in the field. The physical labor involved in field work at times
is arduous, so that though carrying cases of an improved type were
used many instruments were destroyed in spite of every precaution.
Some were broken through accident, others were crushed by birds
while records were being secured, and a few were lost. In all a
considerable number were used in securing the records given.

The records of temperatures of birds secured have been taken
throughout the year, but the localities where this work has been
carried on have all been in the limits of the United States in temperate
regions where extreme cold in winter has not been encountered. As —
a matter of fact very few of the observations have been made at a
time when the temperature of the air was below +25° Fahr. For
this reason it has not been necessary to adopt means for warming the
thermometers, or for keeping them warm, immediately previous to
their use. During the early course of the observations recorded
herein experiments were made in heating thermometers by holding
them in the mouth of the observer when it seemed probably that they
were to be used in a short time. In this way the temperature of the
bulb and the glass for a short distance above was warmed to +98°,
more or less (depending upon the condition of the mouth). It was
soon found however that the results gained with warm and cold
thermometers were so nearly identical that it was impossible to dis-
tinguish between them. Where observations on the body tempera-
tures of birds are to be made at times when the air 1s at zero Fahren-

NO. 12 BODY TEMPERATURE OF BIRDS—WETMORE 5
heit, or even lower, it would seem probable that it would be necessary
to take some steps to use thermometers that had been previously
heated at least to a slight degree. This fact may be readily determined
by observation. It is probable that save in the most extreme cold
weather error from using unwarmed instruments would not amount
to more than a few tenths of a degree.

The observations recorded in the tables at the close of this paper
are to be regarded at best as approximations to the actual tempera-
ture of the birds handled. In treating data of this nature taken
by force from the lower animals we can only assume that we are
correct in our observations. Through experience we may establish
what we consider as the normal limit of variation, but in many cases it
is difficult to be assured that we are absolutely correct. Through long
experience the writer believes that the records presented are a portrayal
of the conditions as nearly correct as may be with the instruments used.

DIURNAL RHYTHM IN TEMPERATURE

The question of the diurnal rise and fall of temperature in a few
species of birds and animals has been carefully investigated by Simp-
son and Galbraith.’ These authors, on two different occasions, secured
records of the body temperature of living gulls, starlings, sparrow-

TABLE I.—Time of maximum and minimum temperatures in birds
(Taken from Simpson and Galbraith)

Time of Time of Time of | Time of
Species Facet ceicies Species Hi ohan qleepaiaee
temp. temp. temp. temp.

Domestic owilich. «3.00 pim.|)3-00/a.mi.||| “ Plawle” ....0. 4.00 p.m.} IT.00a.m.
Domestic fowl ¢..] 3.00p.m.| 3.00a.m.]] “Hawk” ........ | 4.00p.m.| T.00a.m.
Bantam fowl¢ ...| 6.00 p.m./12.00a.m.|| Thrush ........./12.00p.m.|12.00a.m.
Batirammnowleo es 4) "O.00 pin. 4!00asm.||| Dhriuish .. 0.4...) 1.00 p.m.) -1.00'a.m,
Domestic duck ¢. .| 9.00a.m.| 3.00a.m.|| Starling.........| 3.00 p.m.|I2.00a.m.
Domestic duck 9. .|12.00 p.m.|} 3.00a.m.|| Starling.........| 6.00p.m.| 3.00a.m.
Domestic pigeon ¢| 6.00 p.m.|12.00a.m.|| Starling......... | 3.00 p.m.| 3.00a.m.
Domestic pigeon ?/12.00 p.m.|12.00 a.m.|| Owl.............) 3.00a.m.| 9.00a.m.
Sea ciili?, Aees oa 3100\p-tnl-|12:00.a.1.|| Owls... ss. oc. | 4.00a.m./10.00 a.m.
“Seagull. ec. 12.00 p.m.| 3.00a.m.|) Owl .....| 4.00a.m.| 7.00 p.m.
jackdawid acs. « 1.00 p.m.} 1.00a.m.)| Owl -++.+) 4,00a.m.) 1.00 p.m.
Jackdaw @ .......| 1.00p.m.| I.00a.m. |

hawks, a kestrel, thrushes, several owls, and domestic fowls, ducks and
pigeons, at three-hour intervals for period of a week. The results
obtained when tabulated gave curves that agree essentially with simi-
lar curves taken for man and other mammals. (Variation in the time

* Journ. of Phys., Vol. XXXIII, 1905, pp: 225-238.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

of high and low temperature in individuals belonging to the same
species, as shown in the table on page 5, are probably to be explained
on the grounds of individual temperament in the birds under experi-
ment. Thus a bird that soon became accustomed to handling would
give a slightly different series of readings from one that remained
wild and that struggled violently whenever approached.)

In the case of those species normally active during the hours of day-
light a constant diurnal rhythm was indicated, with a gradual rise
until late in the afternoon and then a corresponding decrease until
early in the morning. In owls, species of nocturnal habit, the tem-
perature curves were reversed, the highest point being observed late
at night and the lowest during the day. It was found on the whole
that the temperature curves of daylight-loving species were similar
to those of diurnal mammals save that the point of highest tempera-
ture in the birds came earlier in the afternoon and that of low tem-
perature earlier in the morning.

Hildén and Stenback * record a series of experiments in which birds
were confined in a dark room and their activities regulated by means
of artificial light. Light was turned on from 6 p.m. to 6 a. m., with
a more brilliant illumination from 9 p.m. to 3 a.m. to correspond to
the brighter portion of the middle of the normal day. After the
second day the diurnal birds studied in general adapted themselves
to this change in condition in such a way that the temperature rhythm
was reversed, the highest point in the record for each twenty-four
hours coming after midnight instead of afternoon. When the experi-
ments were terminated and the birds again led a normal life in relation
to daylight the temperature curves at once adjusted to the normal
rhythm.

In studies made by the present writer the range of diurnal tempera-
ture is well shown by records for certain of the owls. Thus a male
barn owl (Aluco pratincola) killed at 3.00 p.m., as it flew from a
perch in a cottonwood tree had a temperature of 1o1.9°. The day
was bright and clear and the bird had in all probability been at rest
since early morning. Another male, shot at 8.30 p.m., while quarter-
ing back and forth across a level flat near the Gila River in Arizona,
showed a body heat of 105.0°. This bird was seen coursing about for
several minutes and had evidently been hunting for food for some
time. The variation in the screech owls (Otus asio) also is instruc-
tive in connection with the same points. A male and a female taken
by hand at 3.30 p.m., from a low pine at the edge of a swamp in

NO. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 7

western Florida, gave temperatures of 101.8° and 102.7° respectively.
A second male shot an hour later registered 100.7°. These birds
had been resting quietly for the entire day. In direct contrast is the
condition found in a male killed by moonlight at 8.15 p.m. in the
Chiricahua Mountains in southeastern Arizona. On this evening
screech owls had been actively calling for over an hour before this one
was secured. The bird taken had a temperature of 105.4°. A female
secured in the Dragoon Mountains, Arizona, at 8.45 a.m., not long
after it had retired for the day, registered a body heat of 105.3°.
In the case of those species active during the hours of daylight the
difference is less marked, due in large part to the fact that the majority
of birds examined had been killed during the period of their normal
activity when little or no temperature range was evident that could
be correlated directly with time. However, it is possible to cite a few
cases showing this regular variation.

The series of mourning doves (Zenaidura macroura) obtained may
be discussed as pertinent in the matter of diurnal variation in birds
of diurnal habit. In four males secured between 5.00 a.m. and
5.45 a.m., the range in temperature is from 108.0° to 109.8°, the
four records giving a mean of 108.9°. Five other birds shot between
5.00 p.m. and 6.00 p.m. under similar conditions give a variation of
from 109.3° to 110.4° with a mean of 109.6°. An average difference
of .7° is thus shown. All of these birds were feeding but in the first
set killed early in morning the body heat was still comparatively low.

As stated above, most of the records were secured at hours when
the temperature was naturally high so that little time variation is
shown. However, the following may be quoted in addition to the
above. An ash-throated flycatcher (Myiarchus cinerascens) taken at
7.30 a.m. had a temperature of 108.6°. Another taken at 10.00 a. m.
registered 110.0° and a third shot at 11.00 a. m. showed a body tem-
perature of 111.8°. All of these birds were males. A male Canadian
warbler (Wilsonia canadensis) shot at 7.30 a.m. registered 106.7°
and another taken at 9.45 a. m. gave a temperature of 107.6°. Females
of the same species taken at 7.30 and 9.00 on the same morning had
a bodily heat of 107.7° and 107.0° while a third bird of the latter
sex killed at 12.00 p.m. showed 108.3°.

The daily increase in bodily temperature in our smaller birds may
be exemplified also by the following: On September 12, 1919, at
Plummers Island, Maryland, the writer spent the day in observing
small birds, many of which were in their southward migration. There
had been a heavy rain the previous evening, and later high wind had
come up with a considerable fall in temperature. The early morning

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

was damp and cold so that insectivorous birds were more or less
inactive until the sun was well up in the sky. As the air became
warmer the birds, correspondingly, became more sprightly. A female
magnolia warbler (Dendroica magnolia) secured at 8.30 a. m., hada
bodily temperature of 107.0°. At this time birds of similar habit
were just commencing to move about and feed. A male of the same
species shot at 9.30 a.m. had a temperature of 108.2° and a second
male taken at 10.15 a.m. had a temperature of 108.3°. Other war-
blers taken between 9.00 a.m. and 10.30 a.m. gave records similar
to the last. Thus a female bay-breasted warbler (Dendroica castanea}
shot at 9.15 a.m. registered 108.8° and a male killed at 10.00 a.m.
registered 108.7°. A male black and white warbler (Mniotilta varia)
secured at 9.15 registered 108.5°. In the observations of this fore-
noon it was noted that the red-eyed vireo (Vireosylva olivacea) was
astir much earlier in the day than the warblers. This activity appar-
ently was reflected in the body temperature as a female taken at 9.15
gave a record of 109.7°, a reading distinctly higher than that of the
warblers taken at the same time, although average temperatures for
the two groups on the whole are about the same. .

In perusal of the data used in preparing this paper many cases are
found of range in temperature not correlated with time. These,
however, cannot be taken as destroying the value of those instances
where early and late readings were available that show a distinct
rhythm or increase from early to late. As stated above, the bulk
of the records are made during the period of high activity when the
bodily temperature approaches a maximum. Range in temperature
then is to be attributed to other causes. Had records made early in
the morning been available in all of these cases there can be no ques-
tion but that an increase in body heat correlated with time would have
been shown.

The daily variation in temperature is much more in birds of small
size than in those of greater bulk. Thus Simpson and Galbraith’
found the daily range in the “ thrush” (Turdus merula?) to be from
6.8° to 7.5°, in starlings (Sturnus vulgaris) from 5.2° to 7.2°, while
in the domestic fowl it amounted to 1.9° and in the domesticated duck
from 1.6° to 1.8°. The marked difference in these cases is noticeable.

The diurnal variation in temperature that has been noted may be
attributed directly to the metabolism of the individual as reflected
by its activities. Wuth movement and the digestion of food heat is
generated. Although a part of this is dissipated through the usual

4 Journ. of Phys., Vol. XX XIII, 1905, p. 237.

NO. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 9

channels there is a gradual accumulation that warms the tissues and
thus is held. This accumulation reaches its height near the close of
the day’s activities. At night the body is inactive so that at once the
production of heat is lessened. The stored up heat energy of the
day therefore is slowly dissipated and is replaced only in part, so
that there is a steady lessening of the body temperature until the
beginning of a new period of activity. With this there is a sudden
jump again in the production of heat and a corresponding increase
in body temperature. In the case of our small birds that migrate by
night the increased activity induced must of necessity make a decided
break in the daily temperature rhythm. No data on this point is avail-
able, but it must be supposed that the prolonged flights that are made
tend to bring the temperature above the normal for individuals at rest.
A part of the accumulated heat secured during the day may be attrib-
uted to the continual ingestion and digestion of food, as the nutriment
thus secured would be provocative of renewed energy. As digestion
is rapid the stomach is soon empty when no food is being taken in
so that the temperature of night-flying migrants may be held down to
some extent by a decrease without constant renewal in the stored
nutriment in the tissues.

Although in birds there seems at present no indication of a marked
seasonal variation in degree of temperature yet we may suppose that
the total amount of heat produced by the body may be slightly more in
summer than in winter. As the period of daily activity is one of high
temperature (considered on the basis of diurnal species) it will be
readily seen that that period of activity is much longer, in species
living north or south of the tropics, during a day in June than during
a corresponding period in December. There is therefore a contrast
in many birds in the total amount of heat produced in the seasons of
summer and winter even though no birds are known to hibernate.
Although it may be supposed that the increased activity in summer
among our small birds may be offset in part by brief siestas taken in
the heat of the day, still it would not seem that the difference between
the two seasons would be anywhere near compensated. The quantity
of body heat produced during the summer period may therefore be
considered greater than for the winter.

VARIATION IN TEMPERATURE IN RELATION TO SEX

Variation in body temperature correlated with sex has been well
established in mammals, where the temperature of the female is stated
to average slightly higher than in the male. Thus Roger’ records a

*Richet’s Dict. de Phys., Vol. III, 1892, p. 96.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

slightly higher temperature in girls than in boys, although the subjects
of experiment were too young to exhibit striking sexual differences.
Martins * notes a higher temperature in females than in males of the
domestic duck, and Simpson and Galbraith* make similar observa-
tions with regard to other birds in those cases in which the sex of
the individuals in hand was known at the time of experiment or was
determined later by dissection. A few observations made by Hildén
and Stenback* do not bear this out, as they cite higher temperatures
in males than in females. Apparently these authors misunderstood
Simpson and Galbraith as they state that these observers also record
higher temperatures in males than in females although the reverse
is true.

In my-own observations there is found in some species a convinc-
ing agreement with the findings of Simpson and Galbraith, Martins
and Roger in this matter, as where a sufficient amount of data is avail-
able the average temperature of the female is usually slightly higher
than that in the male. Certain exceptions to this rule will be noted
later. The difference in favor of the female where present is rather
slight, being usually only a part of a degree. Thus in the green-
winged teal (Nettion carolinense) the temperature of the male (19
records) averaged 106.1°, of the female (8 records) 106.6°, a differ-
ence of .5° in favor of the female. In Traill’s flycatcher (Empidonax
trailli) males (6 records) averaged 108.0° and females (4 records)
108.6° a difference of .6°. Numerous other instances will be noted
in table 3 but need not be cited here as the two given will serve to
exemplify the statement made above.

The present work, however, has emphasized the fact that in cer-
tain groups the temperature of the male on the average is distinctly
higher*than in the female. Apparently this is true in the herons
(Ardeidae) as in three species in that group we have the following
averages: Great blue heron (Ardea herodias) male (2) 104.8°,
female (2) 103.7°; snowy heron (Egretta candidissima) male (5)
104.8°, female (8) 104.0° ; and black-crowned night heron (Nycticorax
naevius) male (3) 103.5° and female (2) 102.6°. A similar differ-
ence is best shown perhaps in certain of the shore-birds, as the phal-
aropes, where it is indeed striking. In the northern phalarope
(Lobipes lobatus) males (9 records) averaged 107.6° and females
(17) only 106.6°. Males (10) of the Wilson’s phalarope (Stegano-
pus tricolor) gave 106.3°, and females (18) only 105.7°. The same

* Journ. de Phys., 1858, p. ro.
* Journ. of Phys., 1905, p. 237.
* Skandinavisches Arch. ftir Phys., Bd. 34, 1916, pp. 382-413.

NG: 52 BODY TEMPERATURE OF BIRDS—-WETMORE ~ Wu

holds true of the avocet (Recurvirostra americana) where 14 males
averaged 106.6° and 12 females only 104.9°. <A good series of read-
ings for the black-necked stilt (Himantopus mexicanus) showed the
same average (105.8°) for both males and females. Through the
family Scolopacidae the general average on the whole showed a
balance in favor of the males save in a few instances. Among the
Charadriidae two species, the black-bellied plover (Squatarola squata-
rola) and the killdeer (Owyechus vociferus) show a balance in favor
of the female, while in the snowy plover (A£gialitis nivosa) and the
mountain plover (Podasocys montanus) the reverse is true. In some
passerines the two sexes average about the same in degree of bodily
heat. Thus in the yellow-headed blackbird (Xanthocephalus xantho-
cephalus) males (29 records) averaged 108.3° and females (18
records) 108.2°. Similarly in the house finch (Corpodacus mexicanus )
males (20 records) averaged 108.9° and females (7 records) 108.8°.
In these species we have, therefore practically an agreement in both
sexes as the difference noted, amounting to only one-tenth of one
degree may well result from accident in securing the records.

It appears therefore that in many species of birds temperatures of
females are higher than those of males. In a considerable number,
however, the two sexes average about the same, though with more
information it may be found that there is a slight difference in favor
of the females. In a few cases there is found a higher temperature
in males than in females. This last is true in the Ardeidae, the Pha-
laropodidae, the Scolopacidae and in part in the Charadriidae.

In the average run of species of homoiothermal animals the differ-
ence in temperature between male and female may be ascribed to the
needs of sexual activity and reproduction. At least this higher tem-
perature seems correlated with certain phases of reproduction. In
many birds the care of the young devolves upon the female and she
has the higher body temperature. In the Phalaropodidae, on the con-
trary, it has long been known that the duties of incubation and the
rearing of the young fall to the lot of the male, so that in connection
with this it would appear that he has developed a higher body tem-
perature than the female. In the case of the Scolopacidae the writer
is prepared to state from his own observations that incubation and
brooding of the young may fall largely upon the male in the willet
(Catoptrophorus inornatus) and it is suspected (though not yet
proven) that this may be true of the majority of the species in this
family. Males of this group examined in breeding season often
exhibit areas upon the lower surface of the body bare of feathers
where the skin is thickened and vascular as in birds that have been

12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

incubating. This is true also in the Recurvirostridae. In the case
of the herons definite knowledge is lacking so that I venture no state-
ment in regard to their sexual aberration in body temperature.
Admitting that the incubation of eggs and the brooding of young
necessitate a higher temperature in the parent, it appears that in those
species where this duty falls upon the female she has a higher average
temperature than the male. Where the male performs these duties
the reverse is true. As in many cases, in particular among the pas-
serines, these cares are shared about equally by both parents, we may
expect in such species a close approximation in body temperature
in the two sexes, a supposition that is well borne out by the data given
in table 3. On the basis of this reasoning it may be permissible to
theorize further with regard to the shore birds. In many species
here the cares of the family are undoubtedly shared by both parents
though as has been said, in a good many forms this duty falls on the
males alone. However, in those that have been investigated, the
greater part show a higher temperature in males than in females.
In this group then, where the male is the home drudge taking over
all the family cares and leaving the female in freedom after the deposi-
tion of the eggs the condition may be assumed to be a primitive one.
Males of other species have become emancipated in part from this
domestic yoke so that the task of rearing offspring is shared in part
by their spouses, though this has occurred so recently that adjustment
is not complete and the body of the male still develops a higher
average temperature. Such statements however must be taken with
reserve and cannot be considered as applying to other groups of birds.

EXTERNAL TEMPERATURE IN RELATION TO BODILY HEAT

In the cold-blooded vertebrates heat generation within the body is
slow, while the processes that act in controlling radiation are imper-
fectly active. In consequence the animal chills or is warmed in close
harmony with fluctuations in heat of its surrounding element. Such
creatures of necessity are sluggish when they encounter low tempera-
tures and become more active when well warmed. When cooled below
a certain point they become torpid and dormant. It would seem that
animals of such habit have the means utilized in equalizing or resist-
ing high temperatures better developed than those that might assist
them in overcoming cold. Otherwise turtles, frogs, or lizards would
be killed when basking in the intense heat of a midsummer sun. This
equalization of heat must be accomplished largely by the lungs in
Reptilia, as skin glands that might serve this purpose are absent. The

NO. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 13

amphibians, with their poorly developed lungs, gain the same end
by evaporation and radiation through their moist glandular skins.
There is a definite limit, however, to the degree of heat that these cold-
blooded animals may endure. In Arizona I have seen a Gila monster
(Heloderma suspectum) fresh from the desert perish when the sack
in which it was confined was inadvertently left exposed in the noon-
day sun for a period of fifteen or twenty minutes.

Although in these cold-blooded animals there is a direct reaction
to external cold, with birds the case is entirely different. Some investi-
gators in making studies of avian body temperatures have been careful
to record the temperature of the atmosphere and to cite this data in
connection with their other records. After due consideration I have
not done this as I do not consider that there is any constant relation
between the normal temperature of the surrounding medium and that
of the body cavity in birds. After making careful records of avian
body temperatures at all seasons of the year, I am, in light of the
records available at present, unable to recognize any constant differ-
ence between body temperatures made in the same species at seasons
of marked heat or marked cold. Where the individual is in normal
health and is sufficiently supplied with food, the agencies of tempera-
ture control will tend to maintain an even body heat. Any variation
that may occur, other than that incident to the daily rhythmic rise
and fall of body heat, may be attributed to some other condition that
under normal conditions would disappear within a comparatively
short period through a readjustment of the bodily functions. Any
bird may, through inclement exposure, become thoroughly chilled and
so have a greatly reduced temperature but such a condition cannot
be considered normal. Thus an immature white-faced glossy ibis
(Plegadis guarauna) exposed for half an hour to a severe rain and
hail storm became so chilled that it could scarcely stand and shook
violently with cold. When warmed with hot towels and dried out
once more it was restored to its normal condition and soon was run-
ning about on the floor of the laboratory so far recovered that it
mischievously began to torment other smaller birds confined with it.

Birds, however, may be divided roughly into two classes with regard
to their ability to adjust to external temperature. The first category
includes those able to withstand any reasonable degree of cold, while
in the second are included those species that migrate to regions where

_cold in any degree is not encountered at the approach of inclement
weather. Broadly speaking, the question of difference between these
two groups is not so much one of change in external temperature as it
is of food supply. Thus species that feed on flying or crawling insects,

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

or on fresh fruits, must leave before a supply of- this food fails.
Others that search out insects in hibernation, dried berries, or live
on seeds, pay little attention to the approach of chilling weather. The
question seems on the whole one of adequate food supply, that the
organism may receive its life-giving elements constantly. However,
forms that habitually experience cold weather during a part of the
year must have a greater development of reaction for temperature
regulation than do those of the other groups. Some species from the
category of those accustomed only to a hot climate may experience
severe cold without harm if supplied with proper food. Others suc-
cumb under these unusual conditions. Thus, Mr. N. Hollister, Super-
intendent of the National Zoological Park, informs me that red, blue
and yellow macaws (Ara macao) confined in large flight cages, remain
outdoors at Washington, D. C., during the winter in perfect health.
An allied species however, the blue and yellow macaw (Ara ararauna),
was unable to withstand the cold and perished, though it throve during
warmer weather. From information available it seems that both
of these macaws in their normal range inhabit the tropical zone, and
are subjected to the same general conditions of life.

There is a marked decrease in body temperature where food is not
obtained 1n suitable amount while the bird is subjected to cold. This
may be seen readily among our smaller insectivorous birds where they
are caught by a sudden return of cold weather during their northward
migration in spring. Decrease in bodily temperature from this cause
may be illustrated by the following:

During the latter part of May, 1916, I was stationed at a small
field laboratory in Utah near the point where Bear River enters Great
Salt Lake. For several days preceding the evening of May 23 the
weather had been mild, and small migrant birds that nested in the
mountains had left the middle of the valley for the uplands. On the
night in question a cold wind with a driving rain came on and con-
tinued until ten the following morning, and there was little rise in
temperature of the air until late afternoon. A few Audubon’s, pileo-
lated and yellow warblers and an occasional small flycatcher appeared
in the willows, and until noon there was a steady flight of swallows
down the river toward some haven on the flats below. To escape the
driving wind, the latter flew low over the river or beat along behind
shelter of the willows that fringed the stream. Hundreds passed,
travelling in little flocks so that for a time there seemed to be no end of
the constant procession of passing birds. These small birds were not
obtaining food as no insects were to be had and in consequence many
were suffering from a lowered vitality. This was reflected in the body

NO. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 15

temperatures of individuals that I collected. Thus, two male violet-
green swallows (Tachycineta thalassina lepida) registered 103.8° and
106.8°, respectively, a female black-throated gray warbler (Dendroica
nigrescens) 105.4°, a female Audubon’s warbler (Dendroica audu-
bont) 105.6° and two purple martins (Progne subis) 104.8° and
105.0°. Such abnormal records, due manifestly to lack of sufficient
food, were not included in the register giving the normal average
and range of temperature. That they were abnormal may be ascer-
tained by referring to other records given for these species.

Birds that remain in regions where they are exposed to cold, be-
come more heavily feathered before the winter season so that there is
less radiation of heat externally. Correspondingly, in summer the
feathered covering is thinner, and the feathers themselves often
become more worn so they are less burdensome. With increased cold
there is apparently some readjustment to maintain the bodily heat
at its normal point. Were this not so the individual would become
affected so unfavorably that with prolonged exposure it would perish.
We must suppose a more rapid metabolism and a conservation of the
resultant energy in order to overcome this. Such a condition is not
difficult to imagine in hawks, crows, and other birds of large size, but
is wonderful when such feathered mites as the kinglets, creepers and
chickadees are considered.

The part that the feet and tarsi of birds play in equalizing the
body temperature is difficult to state. In the majority of birds the
space from the lower end of the tibio-tarsus to the tips of the toes
is covered with skin in which are developed more or less perfect horny
scutes (the whole forming one of the most evident reptilian features
visible in the living bird.) The blood supply is of fair quantity clear
to the tips of the toes, as blood trickles from slight wounds in the
foot. There is no evidence of a forced circulation yet in many cases
warm blood must be conveyed constantly to these parts to avoid frost-
bite. Many ducks, grebes and other aquatic birds remain during
winter just south of the line of ice. Swans, phalaropes and loons
appear in the Arctic regions with the first breaking up of ice. I have
seen auklets, puffins, and murrelets swimming and diving for hours
in Bering Sea with no apparent discomfort at a time when the tem-
perature of the water registered +39° Fahrenheit. Mallards and
other wild ducks frequently clamber out of the water and stand about
on ice for considerable periods without visible hardship. How
these birds overcome or avoid the effect of cold upon their feet is a
mystery as yet. Although supplied with a certain amount of blood,
as has been stated, the feet and tarsi of birds are more often cold or

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

cool to the touch than otherwise in spite of the high body heat. In
fact they are seldom warm save in the hottest weather. Frozen feet
and toes are not uncommon among domestic fowls but are seldom
encountered in wild birds. The skin of the feet and tarsi is smooth
and oily, so that in the case of aquatic species water does not adhere
when the extremities are exposed to the air, thus preventing danger
through the formation of ice. Aquatic birds at rest frequently draw
up the feet one at a time beneath the long feathers covering the flanks,
and ducks often rest on frozen ground with both feet drawn up in this
manner, so that relief is available when needed.

In some groups of birds the tarsi and at times the toes are well
protected by a covering of feathers that prevent the radiation of
heat. Such a development is found in ptarmigan and certain other
grouse, in our owls of northern habitat, and in the sand grouse and
rough-legged hawks. A similar covering, though less dense and
heavy, is found, however, in other birds that never encounter severe
cold. Thus the tarsus is feathered in whole or in part in some trogons,
in whippoorwills, in certain species of edible-nest swiftlets (Collo-
calia) and many others. The entire tarsi and upper surfaces of the
toes are feathered in the Old World martin (Delichon urbica). A\l-
though this covering is present in Delichon, in the bank swallow
(Riparia riparia) a species that also breeds regularly far north in
Arctic regions, the feathering is restricted to a small tuft on the
posterior face of the tarsus near its lower end. Where this covering
of the legs and feet is found in species that at present do not seem
to require it for protection it may be supposed that it has persisted
after an ancient need causing the growth has disappeared, or that
it has developed as a correlated structure, perhaps ornamental in
nature. Thus in tropical owls the feathering of tarsi and toes is
greatly reduced, although in northern species it is very heavy.

On the whole it would appear that radiation of heat through the
lower extremities is comparatively slight.

DIVERSE MISCELLANEOUS FACTORS IN THEIR RELATION TO
BODY TEMPERATURE

Previous sections have covered various phases of variations in
temperature due to sex, daily temperature rhythm and other condi-
tions. It remains to consider a few miscellaneous factors that affect
this matter. Some of these are normal and some abnormal.

The ingestion of large masses of food will frequently cause a
sudden decrease in body temperature in a bird of small size. The
matter swallowed if cold will absorb warmth until it has acquired a

NO. I2 BODY TEMPERATURE OF BIRDS—-WETMORE 17

degree of heat equivalent to that of the tissues inclosing it. In this
way a distinct lowering of internal heat may be occasioned. On
Sept. 12, 1919, while watching two yellow-throated viroes (Lanivireo
flavirons) that were feeding in company I saw one after much effort
swallow a very large caterpillar. Both of these birds were collected
and were found to be immature females. The bird that had eaten the
caterpillar five minutes before it was killed registered a body tem-
perature of 107.2°, while the other gave a reading of 108.1°. This
difference of .9° was attributed to heat absorbed by the large mass
of recently ingested food. This may be considered a normal variation.

In small birds bathing may also occasion a slight decrease in body
temperature where the plumage becomes thoroughly wet. The heat
taken up during evaporation incident to drying the feathers may
occasion an appreciable drop in body heat. As an example of this,
a verdin (Auriparus flaviceps) that had just bathed, taken June 16,
1919, near Arlington, Arizona, gave a temperature of only 106.0°,
while others of this species ranged from 106.5° to 107.6°. Variation
from this cause is slight, however, and would not be appreciable save
in species of very small size.

Many persons with whom the writer has discussed the question of
the taking and recording of the body temperatures of birds have
expressed the belief that the shock produced in, the bird when it is
shot is sufficient to increase the bodily temperature to a marked degree.
Such statements have come in particular from physicians and others
of similar training. Experiment and observation have shown, how-
ever, that this is not true.. On various occasions by accident or inten-
tion birds have been killed in such a way that they were instantly
riddled by shot, so that all functions of the body, nervous as well as
circulatory must have ceased instantly when the bird was struck,
and this on occasions when the individual in question had no reason
to suspect danger. Temperatures of such specimens show no varia-
tion from those of birds taken in a more normal way. As a matter
of fact it has transpired that the shock of wound in birds serves rapidly
to reduce their body heat after a period of from thirty to sixty
seconds. Thus a wing-tipped bird, with an injury that is compara-
tively slight, will be found usually to have a temperature below
normal after a period of two minutes has elapsed from the time that
it was injured. With more serious injuries the fall in body heat may
be so great that a record made on a living bird four or five minutes
after it was shot is so low that it must often be considered as abnormal.
As an example of this I may cite the case of a cinnamon teal ( Quer-
quedula cyanoptera) that was struck in such a way that the sight of

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

both eyes was destroyed though other injury was not present. This
bird rested quietly on the water, while I went for a boat in order to
retrieve it. On reaching the spot fifteen minutes later the body tem-
perature of the duck was found to register only 102.0°, a reading
considerably below normal. Many other examples of a similar sort
in the case of birds bearing only slight wounds have come under
observation. The rapid reduction in body heat is due perhaps to an
abnormal exchange and radiation through the air-sacs.

TEMPERATURE OF YOUNG

During the course of this investigation occasional opportunities
were presented for securing temperatures on nestling birds or on
young of species that leave the nest as soon as hatched. These have
not been used in securing the average temperatures for each species
given in table 3 or for the family records in table 4, as they showed
some variations from readings for adults. The results obtained are,
however, of considerable interest and are presented in tabular form
herewith in order that they may be discussed briefly. In this table
the few species included are grouped under family headings. In
the second column is given the temperature and under remarks is
included a statement of the approximate age of the individual and the
manner in which the temperature was taken. In certain passerine
species, where axillar temperatures were taken, the end of the ther-
mometer was held closely in the hollow between the folded wing and
the body. Birds utilized from the same nest or brood are grouped in
brackets.

Of the species listed those belonging to the families Colymbidae,
Laridae, Anatidae, Rallidae and Recurvirostridae are precocial while
those listed under Columbidae, Tyrannidae, Hirundinidae, Mniotilti-
dae, Mimidae, Paridae, and Turdidae are altricial. A difference in the
temperature records in the two groups is readily apparent upon
examining the table. In the group of precocial birds there is on the
whole less variation and the temperatures given closely approxi-
mate those of adults of the same species. The only wide divergence
in this group is in the case of the three-day old young of the black-
necked stilt (Himantopus mexicanus). It so happened, however,
that the temperatures of these three birds were taken upon a cold
raw day, when the young were very evidently affected by the external
cold, perhaps through lack of sufficient food.

In the case of the altricial species considerable variation is present
and these birds show a much lower average temperature than adults.

NO. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 19

TABLE 2.—T able of temperatures of young or nestling birds

Tempera-

Species fare Remarks
CoLYMBIDAE .
Podilymbus podiceps.....| 104. Newly hatched. (interthoracic).
y
“fe oe
Spee alah TOO
‘a So eae) LOL) i Lron one brood:
ss Seen Sete A LOO
LARIDAE
Sterna forsieris ........-|) 107-3: | Len days old (rectal).
e S Seiad ah] yy HOO ss Soe aka “
oy Deets |) O52) a1 Sevyen, > * s
oy 3 mratene io ietenate s I0l.5 ier ES e
ns 2 BOSD OC oor 104.5 Six ° s 42
NUZTA..... : even
Hydrochelidon nigr 105.4 |S Cente x
ANATIDAE
Anas platyrhynchos......| 105.8 | One-third grown “
Chaulelasmus streperus..| 104-5 | One day old Py
‘ce ‘ 106. if ‘6 ‘é ““ “
“e ia 107.3 “e ee ce “ee
“ee 6c 107.8 “ec 6c “cc ‘cc
“ce 77 . 107.8 a3 ce «e ‘c
Querquedula cyanoptera. 107.7 Suen NEE a z
“ ‘6 int 100.5 ‘ce “c “ ‘
“ce (73 105.6 6 ‘é 3 ‘6
“e ‘“c . 100.3 ‘ce ‘6 ‘ ‘sé
Spatula clypeata......... 102.7 | One-half grown “
“ce ce ‘7 104.0 oe “ec “ce ee
Marila americana........| 102.9 | Twohours old “
ee “cc An 102.7 a3 “cc “ce “ec
“ re Pe etesa: Whivedays ald”.
“c “cc 100.8 “cc “cc “ “<e
ce GLU ae aE oem TOo6.0 “ec 6“ ‘ce “
ve ce iS deh. Io = Cc oe ee ce oe
“ce ce get eee “cc é se “ec
is ee ee Ot Ot Pemy in‘) Pst
a os UieE ele T0020" "Two weeks" *
Erismatura jamaicensis..| 104-2 | Tendays “ ss
RALLIDAE
Fulica americana ........| 108.6 | One-third grown “
a Oo | tat ie een 109.0 | One-half Se heen
“ 6c 106.6 6c “cc 6c (73
“ “e Ciseae ee 104.5 oe e (a3 “e
RECURVIROSTRIDAE
Recurvirostra americana.| 103.9 One week old ps
“ce ce 104.3 oe oe “ce ce
ce ia ; 104 5 oe f ce ins
“ee “cc i 103.6 “6 “cc ‘ec “
. es .| 104.4 | Newly hatched.
Himantopus mexicanus .. 95.8 | One day old 9
““ 6c be 95.3 oe “ ‘ec a3
“ee “cc ~ ce “ce “ce “ec
0s 97.
- es ..| 105.7. | One-half grown “
CoLUMBIDAE
Zenaidura macroura.....| 106.5 | Two weeks old ‘*
TyYRANNIDAE ;
Mvytarchus crinitus.......| 103.3 Nestling, 12 days old (axillar).
ce oe 103 8 “ce ae ae “ce “e .
ce “oe a ae 103.2 ce ae ae a3
oe ce te 103.8 ae ce ee ae “
“e ee =e ; ce oe ae ce * =
wee 103.7 (interthoracic)

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

TABLE 2.—Continued

Species Temper Remarks
HIRUNDINIDAE : ¢ ;
Petrochelidon lunifrons..| 103.4 \| Nestling, 2 weeks old (interthoracic).
“ce ia3 om 102.8 “ec “ce “ce ee ce
MNIOTILTIDAE ; ;
Geothlypis trichas ....... 100.2 | Nestling, 7 days old (axillar).
ce et ek IOI 8 “e “e ce “ce “
“c Ce ax» Satd ak a 102.5 ce ce of “ce “
ce OO bi Tyan te 102.8 ce ce “cc ce be
MIMIDAE
Dumetella carolinensis... 105-2} Neetiine (asia:
¥ 3 105.7 One brood.
f 2¢-%| | Nestling, 8 days old (axillar)
éc go? | One brood.
97.9
PARIDAE :
Penthestes gambeli ...... 101.4 | Nestling nearly grown (interthoracic).
7 SF A Wet feo 97.0 One brood.
TURDIDAE
Sialia currucoides........ 97.6) | Nestling, week old (interthoracic).
x SENEGAL ri oes 96.7 J | One brood.

These helpless young are evidently as dependent upon brooding by
a parent to maintain their bodily heat as are eggs before hatching.
Apparently the body temperature may be considerably reduced, how-
ever, without permanent injury so that the body heat may sink as low
as 97° without death resulting. Even where nestling birds have devel-
oped contour feathers the temperature still remains considerably below
the average for the adult. When the bird leaves the nest at once there
is agreement between the degree of bodily heat that it develops and
that present in the adult. ,

The single observation recorded for a young mourning dove is
apparently anomalous as it averages higher than those given for other
altricial birds. This is of interest as the doves have distinct affinities
with groups having precocial young but in the Columbidae the imma-
ture birds, though covered with down at birth, are confined to the
nest until able to fly. Further study of young doves and of other down-
covered young that do not leave the nest when first hatched, as young
hawks, owls, turacos and others, will be of interest.

From this discussion it may be stated with apparent certainty that
in birds with precocial young the mechanism of temperature control
is well organized at birth, while in species with altricial offspring this
power is so feebly developed that these birds are largely dependent
upon the parents for heat. The ability of perfect temperature con-

NO. 12 BODY TEMPERATURE OF BIRDS—WETMORE 21

trol is not fully matured until the young leave the nest. The early
development of this faculty in precocial young is in line with their
advanced stage as regards securing food, general activity, and ability
to care for themselves. According to Pembrey, similar statements
regarding young birds have been made by Edwards.’

METHOD OF TEMPERATURE CONTROL IN BIRDS

Bodily heat in all animals is caused by tissue changes during active
work performed by various organs or parts. Mills * states that bodily
heat, though arising in great part from actual oxidations that take
place in the system, is in its entire amount best defined as the out-
come of all chemical processes that take place in the organism. In
so-called cold-blooded vertebrates the combined energy or rate of these
chemical changes is slow, so that heat is given off by the body almost
as rapidly as it is generated. In the groups that we class as warm-
blooded, Aves and Mammalia, these changes are more rapid and
intense so that heat generation may be in excess of heat radiation. In
the warm-blooded group there is also a more or less perfect control of
heat radiation when the body is normal in health. In homoiothermal
animals there is, therefore, an approximation to the maintenance of
a fairly uniform internal temperature, and the animal remains inde-
pendent of the ordinary rise and fall of the degree of heat of its
external medium.

Bayliss * considers control of heat production (probably in muscles)
as the primitive method of temperature control. Among homoio-
thermal animals, the Monotremes (both Echidna and Ornithorhyn-
chus) have the lowest body temperatures, as the average for these
species is only 85.6° Fahr. In the case of Eciidna all regulation of
temperature appears to be through change in heat production as this
animal possesses no sudoriferous glands and shows no apparent change
in respiration at high temperatures. In cold weather it hibernates and
maintains a temperature only slightly above that of the air. The
duck-bill (Ornithorhynchus) has the power of regulating both heat
loss and heat production so that its temperature is maintained at a
more even level. The marsupials are intermediate in this respect
between monotremes and higher mammals.

* Schafer, E. A., Text-book of Phys., London, Vol. I, 1898, p. 804.

2“ De l’influence des agens physiques sur la vie, Paris, 1824. (Not seen by
the present writer.)

* Animal Physiology, p. 461.

“Principles of General Physiology, 1915, pp. 458-459.

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

It has been estimated by Helmholtz * that in the human body heat
lost through transpired air amounts to 5.2%, through the water of
respiration to 14.7% and through the skin to 77.5%. The remaining
amount disseminated is given off in egesta or is consumed in warming
ingesta. The part played by the skin glands in regulating temperature
in the mammal is readily seen. The distribution of skin glands varies
in different groups, though such glands are known in all save
Cetaceans, elephants, Echidna, and some others. It is claimed, for
example, that in the dog skin glands are present only in the legs or feet
but in this case the open mouth and protruding tongue act as organs for
reducing excess body heat. The presence of skin glands in the horse
is readily observed in an animal that is hard driven in warm weather.

Amphibians agree with mammals in the presence of many integu-
mentary glands, though in this group the use of these structures is
in some ways different in purpose. In reptiles and in birds, the two
classes joined in the supergroup Sauropsida, skin-glands are practi-
cally wanting and no case is known in which glands similar to those
in mammals are found. In birds the development of feathers with
their filamentous barbs and barbules, as a body covering, would not
have been possible had sudoriferous glands been present in the skin.
Excretion of fluid through such glands inevitably would have soiled
such delicate structures as feathers and ultimately have destroyed
them: The diffusion of heat through the skin in birds is confined to
the amount, notably small in quantity, that is, given off by direct
radiation. It is a fact easy of verification that the skin in birds is defi-
cient in blood supply when compared with mammals. Only a compara-
tively small amount of blood, therefore, can be cooled to any extensive
degree through the agency of the skin.

As a matter of fact the feathers that form a loose covering over
the bodies of birds are not adapted to the radiation of heat but on the
contrary tend to conserve it and hold it within. Though the contour
feathers lie smoothly one upon the other yet they are permeated and
separated by innumerable air-spaces varying in size from the tiny
interstices between barbs, barbules, and barbicels in individual
feathers, to the broader areas separating one feather from another.
These all go to make up series of more or less closed air cells that act
efficiently as non-conductors and serve to retain the bodily heat
within. The use of so-called “dead” air-spaces between walls as
a protection against conduction of heat and cold is too well known to
make further explanation of this factor necessary.

* Smith, R. M., Physiology of Domestic Animals, 1880, p. 696.

NO. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 23

This lack of heat regulation by means of the skin would throw the
vital work of temperature control directly upon the respiratory system.
In this fact then we have a ready explanation for the presence of the
great series of pulmonary air-sacs that are developed throughout the
avian class as a whole. Birds in order to maintain a high rate of
metabolism, necessary to continued activity without reference to shifts
and changes in the temperature of their surrounding media, have been
forced to develop &n auxiliary to the small amount of heat that may
be thrown off through the lungs. This has led to the evolution of the
air-sacs that, while connected by ostia directly with the lungs, radiate
throughout the coelom and penetrate the bones to serve as an agency
of temperature control. In other words, safety to the organism
demanded that if activity be great and continued, there be some safe
release for the excess heat developed during rapid muscular move-
ment.

The proper function of the air-sacs has been a moot point for many
years and has given rise to considerable discussion. Some have con-
sidered that these sacs acted as reservoirs to replenish air in the lungs,
as containers that, balloon like, raised the weight of the bird in flight,
or that the presence of these open spaces reduced the relative specific
gravity of the body. While the idea of the true use of the air-sacs
in birds as organs of temperature control was arrived at independently
by the present writer, subsequently an admirable exposition of the
same fact has been found in an account by J.-M. Soum.* This author
in turn believed that the discovery of this fact originated in an hypoth-
esis first advanced by De Vescovi.’ W. P. Pycraft* also has adopted
this view as he states that “the air stored in these reservoirs serves
not only for respiratory purposes, but also as regulators of tempera-
ture, thereby compensating for the lack of sweat glands.” With this
comment, however, he goes no further, as he gives no details to support
this statement." M. Soum, however, made an admirable exposition
of his hypothesis. He pointed out that all birds possess air-sacs,
have a covering of feathers, and lack skin glands, and all have a
high temperature. To correlate these facts he believed it necessary
to consider the air-sacs as a means of temperature control. The addi-
tional facts that I am able to bring forth leave no doubt as to the cor-
rectness of this belief.

*Soc. Linn. de Lyon, Vol. XLII, 1805, pp. 153-157.

* Res Zoologicae, Ann. 1, No. 1, Rome. (This publication I have not seen.)

* History of Birds, London, 1910, p. 17.

* With regard to statements by other authors consult also Headley, Structure
and Life of Birds, pp. 100-103.

24 SMITHSONIAN MISCELLANEOUS. COLLECTIONS VOL. 72 .

To continue, as the statement given in the preceding paragraphs
becomes more clear, it seems evident that the bird owes its high
development, when compared with the reptile, to the growth of these
air-sacs as well as to a complete double circulation of the blood. The
truth of this statement is apparent when it is considered that the
Crocodilia among reptiles possess a double circulation so nearly per-
fect that only a comparatively small amount of venous blood finds its
way into the purified stream of the trunk arteries. Yet these crea-
tures are “cold-blooded” and become dormant when subjected to
cold. In other words, their body processes function so slowly that
when they encounter an outside temperature below a certain point
heat is given off by the body more rapidly than it can be produced.
It follows then that the bodily activities ebb lower and lower until
finally they are practically at a standstill.

With animals as active as are birds means of relief from overheating
must be well organized ; the extension of the air-sacs through the body
cavity is excellent for this purpose. The walls of the sacs are very
poorly supplied with blood so that heat is not radiated directly by
means of special circulatory vessels. The thin walls of the sacs, how-
ever, are brought into intimate contact with the trunks bearing the
blood stream and their principal branches while in addition the sacs
closely invest the glands and organs that generate heat. It is claimed.
that the liver produces more heat than any other organ so that the
blood from the hepatic drainage is warmer than any other in the body.
The liver itself is partly enclosed by air-sacs, while the venous
trunks coming from it adjoin sacs that give excellent opportunity for
the casting off of excess heat. Ramifications of air-sacs in the bones of
the body are not uniform in distribution and appear to follow no set
plan. Some species have the osseous system highly pneumatic
throughout while in others this pneumaticity is greatly reduced.
When air-sacs are present in bones invested by considerable muscle
masses they may be considered as developments that tend to further
the proper radiation of excess heat. Thus air cells in the keel of the
sternum and the coracoids would aid in controlling heat generated in
the pectoral muscles and supplement the work of those divisions
of the sacs that underlie the body of the sternum and penetrate from
the thorax into the cervical region.

Evolution of the air-sacs beyond their normal development of five
main pairs that fill the body cavity and the cervical region apparently
has been partly beyond control. The presence of numerous cells
between the skin and muscles in brown pelicans may be supposed to
break the impact of the water as the birds dive for food. The pres-

NO. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 25

ence of these same air-pads in the white pelican and, possibly, in the
man-o’-war bird may be explained by considering that they were
developed as a protection while diving from a height and that they
have persisted now that these birds have altered their mode of securing
food. Similar air-pads in the screamers, however, cannot be ex-
plained by the same argument. Similarly there seems no adequate
explanation on the basis of use for pneumaticity in the pedal phalanges
of the Bucerotidae or in the pygostyle of the Picidae.

Many physiologists have supposed that air-sacs have been developed
by birds to impart lightness to the body, especially to the bones.
Anatomists, however, have pointed out that while the main air-sacs
are more or less uniform in growth, in many cases the bones are highly
pneumatic in species not especially noted for strong or prolonged
flight. The hornbills, already cited as having the osseous system more
extensively permeated with ramifications of air cells than any other
group, are not known to be especially active on the wing.

In the most recently published extensive account of avian air-sacs
Bruno Miller * considers that air-sacs serve no special physiological
function but that they give bulk to the bird body without adding
to its weight. This author continues with the statement that the
connection of the air-sacs with the lungs comes from their manner of
development, and that this connection serves merely to “ assist in
renewing the air in the trachea.” Reflection and study of the facts
of avian anatomy show, however, that this line of reasoning is unten-
able. Bats among mammals fly with the utmost ease and yet possess
no such system of air cells as permeates the body in birds. Some of
our most ancient birds from the standpoint of phylogeny are flightless,
have been in this condition for millions of years, and yet have as
perfect a system of air-sacs as are found in forms noted for their
powers of flight. Fossil remains of an ostrich have been found in
the Pliocene deposits in the Siwalik Hills in India, an indication of
the ancient ancestry of our present-day struthious birds. To those
who would adhere to the theory of Muller as propounded above the
highly emphysematous condition prevailing in the screamers (Anhima
and Chauna) and others may seem of importance but the condition,
as has been said, may be ascribed more to an exaggerated develop-
ment, unchecked because it had no particular significance to the
organism as a whole. Otherwise we must expect a similar condition
in other species, as the Old World vultures and American buzzards

* Air-sacs of the Pigeon, Smithsonian Misc. Coll. (Quart. Iss.) Vol. 50,
pt. 3, Jan. 16, 1908, pp. 403-404.

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

(Cathartidae) that like the screamers spend hours in soaring high
in air.

The action of the air-sacs in controlling the body temperature may
be demonstrated clearly by the following. A house sparrow (Passer
domesticus) was caught across the neck in a spring mouse trap of the
“Out o’ Sight” type. The bird struggled and fluttered violently
for approximately 60 seconds while at the same time ingress or egress
of air to or from the lungs was prevented by compression of the
trachea against the edge of the trap platform by the spring of the
trap. I reached this bird as it became quiet and found that the body
temperature (interthoracic) registered 114° Fahr. The violent, sus-
tained muscular exertion had produced a considerable amount of heat
that could not be given off as the accustomed outlet was blocked. The
temperature, therefore, rose several degrees above the usual maximum
for this.species. These same factors operate occasionally when the
trachea of a wounded bird is clogged with blood that prevents the
. passage of air. The air current must be cut off quickly, however, as
the temperature falls rapidly in a wounded bird even when it is
struggling.

During hot weather it is common to see birds breathing rapidly
with the mouth held open. This facilitates the rapid inspiration and
expiration of air from the lungs. Cooling of the mucous membranes
of the posterior portion of the mouth may also be of slight aid in
reducing the excess internal heat. In the case of some young birds,
as, for example, young herons, there is in connection with this habit
of breathing with open mouth another development to aid in regulating
the internal temperature of the body. When overheated these birds
open the mouth widely so as to expand the capacious mouth cavity
and pharynx while at the same time the skin on the sides of the upper
throat is vibrated with great rapidity. The inner walls of the pharynx
and upper throat in the birds in question are highly vascular so that
the currents of air set in motion aid in cooling the blood exposed in
the radiating blood vessels found near the surface in the moist mucous
lining. Conversely it may well transpire that the checking of the
rapidity of interchange of air between the branches of the bronchi
and the air-sacs during extremely cold weather may bring about a
storage or an increase in internal heat. In other words the heat of the
body cavity may be held at a higher level by the cessation of inhala-
tion of constant supplies of cold air into the air-sacs. We may imagine
a delicate adjustment here that will vary expiration of heated air
at need. With the air-sacs acting thus as heat reservoirs the ability

NO. I2 BODY TEMPERATURE OF BIRDS—-WETMORE 27

of some species of birds to withstand bitter cold winter weather may
be better understood.

Temperature control among birds is less perfect in juvenile than
in adult individuals so that the action described in the case of young
herons in the preceding paragraph is of great aid to control of heat
through the air-sacs. As temperature is poorly regulated in young
individuals parent birds often find it highly necessary to shelter their
offspring when these are reared in exposed nests. On hot cloudless
days, therefore, one bird of each pair remains constantly at the nest
during the warmer part of the day, and intervenes its body and partly
spread wings between the young in the nest and the burning rays of the
sun. I have seen many young herons and ibises perish when the
adults were driven from their nests on hot days, and during field work
in rookeries of these birds have made it a point to visit them during
the cooler portions of morning or afternoon, or to come on days
- when the sky was overcast by clouds in order to prevent such mortality.
As another evidence of poorer temperature control in young birds I
may add that in several cases I have seen immature coots (Fulica
americana) die, apparently of sunstroke, when unduly excited while
exposed to the burning rays of a western sun. Adult birds seem able
to react against these circumstances in such a way that they do not
succumb but often exhibit evident signs of severe suffering. It is
probable that the more perfect development of the feathered covering
in adult birds is of as great advantage in this as the increased efficiency
of the heat regulatory organization in the body.

On a few occasions I have observed a further development of the
function of temperature control by air-sacs in certain forms of birds
while in the fledgling state. Those who have had occasion to work
in summer in marshes densely grown with rushes will agree that at
times the heat encountered is almost overpowering. Ina few instances
in such situations I have observed young yellow-headed blackbirds
recently from the nest, resting quietly with the cervical air-sacs
immensely swollen so that the lower part of the neck was greatly
enlarged. The whole gave the appearance of some unwholesome
tumorous growth and at first I was under the impression that the
birds were diseased. On handling them, however, the sacs rapidly
subsided and the birds seemed normal in every respect. The same
phenomenon has been observed in young savanna sparrows and in
young red-winged blackbirds. In these cases I was forced to conclude
that the distended air-sacs form an insulation or protection against
heat from without. In other words, that the enlarged cavity of
the sac acted as a dead air space protecting the blood stream in the

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

larger vessels from becoming overheated. The importance of the
enlarged cervical sacs in preventing excessive heating of the carotid
artery carrying blood to the head may readily be seen.

SIGNIFICANCE OF TEMPERATURE CONTROL

In common parlance animals are divided in two groups distin-
guished as those with “ warm blood” or “cold blood,” according to
their condition as regards body heat. Though two classes may be
recognized without difficulty, the criterion implied in these two terms
is not exact as a “cold-blooded” animal temporarily may have its
body temperature raised toa high degree. The distinction between the
two, in fact, is not one of actual degree of heat, but rather one of main-
tenance of a more or less uniform temperature in the group defined
as possessing ‘‘ warm blood,” and of fluctuation in bodily temperature
in those distinguished as “ cold-blooded.” To express this idea with
exactness, the first group of animals is said to be homoiothermal, and
the second poikilothermal, terms proposed by Bergman* in 1847.

It will be admitted without question that the possession and main-
tenance of warm blood is of advantage to any animal. We may
suppose, therefore, with what amounts to some certainty that this
faculty when once gained, would not be lost. On the basis of this
assumption it may be concluded further that the first vertebrates were
cold-blooded, an hypothesis in line with facts of evolution as they
are understood and accepted at the present time. Whether these types
developed in regions of equable temperatures or in areas with moder-
ate seasonable changes is a matter of no moment in the present discus-
sion. In either case these early vertebrates as they extended their
ranges, encountered barriers erected by cold during a part of the year.
Groups successful in coping with this condition developed an ability
to undergo certain periods, longer or shorter in length, in the state
of suspended animation that we term hibernation, and then to revive
and carry on their activities as before with return of a period of
increased warmth. In meeting these conditions of cold it was of
advantage to develop increased resistance to the torpor induced. In
other words, it was an advantage to the organism to maintain its
activity at lower and lower temperatures. In order to accomplish
this it was necessary to evolve a mechanism for temperature control
in the body, and for regulation of the rate of production of heat from
ingested food elements. When once begun, such control would prove

*Gottinger Studien, Vol. I, 1847, p. 593.

*

—-

NO. 12 BODY TEMPERATURE OF BIRDS—WETMORE 29

of value not only in overcoming cold but in enabling the organism to
withstand excessive heat.

It seems probable that in our living fishes there is little actual tem-
perature control. In Amphibia, this regulation is developed to some
extent, and it has progressed somewhat farther in modern reptiles.
In the bird, however, the regulation of body temperature has reached
its highest point, though birds stand second to mammals from an
evolutionary standpoint. Proof of this is found in the fact that birds
have the highest body temperatures known, and that none of them
hibernate (in spite of ancient beliefs to the contrary). Where con-
ditions become too unfavorable, birds, through their power of flight,
pass readily to regions where the environment is more clement. They
are enabled, therefore, to foster their powers of temperature control
and keep them at the highest pitch. Small mammals, on the contrary,
are more or less sedentary and in many cases must still undergo
hibernation in order to maintain themselves in regions with cold
winters. As they must always hibernate in order to survive there is,
in their case, less incentive to develop temperature and temperature
control beyond a certain point.

Thermogenic centers or areas in the central nervous system devel-
oped for temperature control have been studied in mammals and have
been fixed tentatively by some in or near the corpus striatum. Others
would recognize a cortical heat center. Seemingly this matter has
received little attention in birds and it would be unwise in the absence
of definite data to decide that this function is vested in the same areas
in this group when the wide separation between birds and mammals
is considered. It may be assumed as certain that heat production and
heat control are under nervous direction and that these two functions
are directly concerned in whatever mechanism has developed for
temperature control. ;

The origin of the warm-blooded animal may be attributed to natural
selection in which certain individuals showed a slight reaction against
temperature conditions producing hibernation in their fellows. In
other words, these favored ones were able to remain active in a tem-
perature a few degrees colder than others of their kind. With this
tendency as a basis and with strains developing in which this tendency
was perpetuated it followed that there were evolved groups of species
with a more independent metabolism in regard to the degree or the
lack of heat of the surrounding medium. “ Warm blood ”’ therefore
arose in a struggle against enforced hibernation. During evolution
of the vertebrates it may be that among living groups of today, warm

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

blood arose first among the birds." We may suppose, however, that
primitive pro-avian creatures (on the borderline between small-
brained reptile and large-brained bird) were cold-blooded and that
they were subject to hibernation as is any reptile today. However,
it seems probable that Archaeopteryx, the most primitive of known
birds, was warm-blooded, as impressions of feathers are shown dis-
tinctly in the slabs of stone containing the remains of these creatures.
These marks made by feathers indicate the development of a body
covering designed to retain heat, a circumstance unknown in any
cold-blooded vertebrate. How far back we may safely trace this sup-
posed warm-blooded ancestral bird creature is problematical but in
this connection attention may be called to the supposition that there
are grounds for believing that Pterosaurs, among ancient reptiles,
possessed warm blood. As warm blood permitted greater mental and
physical activity it was natural that the mammal should also develop
this faculty, though it seems probable that this function arose inde-
pendently in the Reptilian-Avian and Mammalian groups.

DISCUSSION OF DIFFERENCES IN AVERAGE TEMPERATURES

Attention has been called to the general statement that the body
temperatures of birds vary as a rule from low to high as the species
change from those considered low in the scale of development to those
farther advanced. Agreement with this theory is shown in part in the
data summarized in table 4. Thus grebes, the tctipalmate groups
(Anhingidae, Phalacrocoracidae and Pelecanidae) and herons are in
general low in average body temperature, while gulls, shorebirds,
pigeons and cuckoos are high. Many apparent discrepancies to this
broad statement may be noted. ‘These must be left for the present
without attempt at explanation save to note that knowledge of the
actual evolution of groups in birds is slight, while new facts constantly
demand a revision of the status of many forms. Whether the varia-
tions in body temperature here noted may have significance time alone
can tell. It is probable that temperature level is of value as a criterion
only between the most primitive and the most highly developed groups
and that the great mass of intermediate orders and families may in
some cases in themselves develop high or low temperature according
to their actual needs.

* According to Osborn (Origin and Evolution of Life, 1918, p. 236) primitive
mammals arose during the Jurassic period. The earliest known birds are
found in deposits of this same age but are so highly specialized that it is
evident that they were preceded by a long line of pro-aves of more ancient
origin than the early mammals.

NO. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 31

When ranged by families the highest temperatures noted are to be
found in pigeons, cuckoos, woodpeckers and in the great passerine
order beginning (in table 4) with the Tyrannidae and ending with the
Turdidae. It must be noted too that the range of body temperature
among ducks is in general comparatively high. Gulls and shorebirds
show a general agreement, compatible with their close relationship
as now commonly accepted. The quails (Odontophoridae) seem to
have a temperature high for birds that have been considered compara-
tively low in development. The observations recorded for the owls
probably do not represent a true average as many of the readings upon
which the mean is based were taken during the day when temperature
in these birds normally is at low ebb. The low average given for the
kingfishers is based upon a small number of observations and may be
incorrect. Humming-birds, with their tiny bodies seem to have a con-
siderable range in temperature, but as a whole fall low in body warmth.
This apparent lack of heat may be due in part to the small bulk of
their bodies in comparison with the size of the thermometers used.
Part of their heat may have been absorbed and dissipated by the glass
of the inserted instrument.

Observations upon the greater part of the species of woodpeckers
found in the United States reveal an almost uniform high level of
body temperature. The general range and the limits of variation
from high to low are similar to those of passerine families. As in
other families individuals large in body show a general lower tem-
perature and a smaller limit of variation than do some others of
smaller bulk. The records on the whole are-so uniform that further
comment regarding them is superfluous.

An examination of the species and families of passerine birds
reveals much of interest. Of the twenty-two families for which
records are given eleven or exactly one-half, have a mean temperature
averaging below 108°. It will be noticed that most of these families
are those having only a small number of species represented in the
records. In several instances observations were available on one
species alone and only in the crows, swallows, vireos, wrens and
nuthatches is the number of species available comparatively large.
The Hirundinidae (seven species) with an average temperature of
106.7° is the only family in the order falling below 107°. Seven
families, the Tyrannidae, Alaudidae, Fringillidae, Tangaridae, Bomby-
cillidae, Mimidae and Turdidae, have mean temperatures higher than
108.5°. These seven families include 86 of the 203 species of passeri-
form birds represented, or approximately 42 per cent. The Alau-
didae show a mean temperature of 109.4°, which is higher than for any

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

other group, but is not comparable, as this family is represented by only
one species, the horned lark (Otocoris alpestris). A number of
species included in other families have a higher average temperature
than this so that this must be discarded. Among the others the
Bombycillidae, Mimidae, and Turdidae each show an average of 108.9°
while the three families remaining in the category under discussion
vary only two- or three-tenths of a degree below this. From 108.6° to
108.9° would seem therefore to be the maximum for families of
perching birds where records are available for a number of species.

A comparative examination of some of the species of perching birds
brings out still other facts of interest. It has already been stated that
the swallows as a group possess the lowest average body temperature.
In the seven species examined in this family only one, the rough-
winged swallow (Stelgidopteryx serripennis) , showed an average body
heat above 107.5°. Turning to the higher temperatures it is found
that in the Tyrannidae there are five species in which the mean tem-
perature for the male or female is 110° or more. The Fringillidae
include three species in this category and the Corvidae, Icteridae,
Mniotiltidae and Turdidae each possess one. In other words there
‘are records for twelve species in all in which this is true. The highest
average temperature for both sexes is that of the western wood pewee
(Myiochanes richardsoni) with a mean of 110.2°. The highest single
reading believed to be valid was found in this species in an indi-
vidual killed in the Graham Mountains, Arizona, at two o’clock in the
afternoon on June 25, 1919. This bird, shot dead as it rested quietly
in the shade of a cottonwood fell to the ground without a struggle.
When the temperature was taken the extraordinary reading of 112.7°
was secured. From the data at hand it is indicated that the highest
average body temperatures for a number of related species may be
found in the Tyrannidae. This statement is made only tentatively as
further observation may show that other groups are equal in this
respect. It is not unusual for individual birds in several other groups
of perching birds to register 110° or more as shown by the column
of maximum temperatures in table 3, and accident of association of
such high records might give a high average. Only by recording many
extended observations can error from this cause be reduced to a mini-
mum.

EXPLANATION OF TABLES

The data secured during this investigation into the body tempera-
ture of birds are summarized in two tables that are given in the
pages that follow. The table giving in detail the individual records,

NO. 12 BODY TEMPERATURE OF BIRDS—WETMORE 33

which it was found necessary to omit from this paper owing to the
excessive cost of publishing tabular matter, is deposited in the files
of the Smithsonian Institution and may be consulted by those who
wish to use the data contained in it. ;

The order of arrangement and the nomenclature, followed is that
of the third edition of the American Ornithologist’s Union Check-list
of Birds, published in 1910." By referring to this check-list physiolo-
gists and others interested in these tables, who may not have made
detailed studies of birds, will have no difficulty in ascertaining the
application of the names that are used, and the relationship of the
various forms that are treated. At one time the writer intended, in
publishing this information, to use names of birds in accordance with
the most modern findings in nomenclature, and to arrange them in
a sequence of families that would express his own ideas in classifica-
tion. The latter idea was commendable as it tended to place the
species into what may be considered a somewhat more natural sequence
that showed a tendency (not universal, however) for a gradual
increase in degree of bodily temperature from forms low in the scale
to those conceded to be higher in development. With regard to the
names to be employed it was soon seen that changes were so rapid that
they tended to bewilder even those more or less adept in such matters,
while to workers in other fields, they would be wholly unintelligible
without great expenditure of time in looking up and verifying the
various authorities. As the present contribution is not one of research
into systematic ornithology but rather a treatise designed to throw
light upon the physiological and more general aspects of our science,
this scheme of using such a classification was abandoned and another
plan was adopted.

In table 3 is given a synopsis of the information of all of the records
secured with the average, minimum and maximum temperatures sum-
marized for males and females of the species treated so far as this
data is available. In this table attempt is made to arrange the matter
in order of convenience for reference. The name and sex of the bird
are followed by the mean temperature. After this are given the mini-
mum and maximum range, the number of records available, and a
symbol that indicates the manner in which they were taken, the abbre-
viation R. meaning rectal and I. interthoracic. In this table subspecific
names are ignored entirely and all information is grouped under

+ Check-list of North American Birds, prepared by a Committee of the
American Ornithologists’ Union, Third Edition (Revised), New York, 1910,
Pp. I-430, 2 maps.

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

specific names. The mean temperature for each sex is given where
information for both males and females is available.

In grouping this information a departure has been made from a
method that has been utilized by many authors that gives the minimum
temperature, then the maximum, and then the mean. In the present
connection the mean temperature for each species is considered the
most important fact and is therefore placed first, nearest the name
of the bird concerned. Records showing the minimum and maximum
range follow immediately where they are readily available in case this
information is desired.

In table 4 is given another summary in which mean temperatures
for each of the families of birds represented is tabulated. The name
of the family is followed by the number of species represented in the
present studies. Following this are mean, minimum and maximum
temperatures with the mean as the most important fact given first.
The data in these three columns are taken from the column of mean
temperatures in table 3. In other words, this is a summary based
upon the mean temperatures alone of the various species.

The laborious work of securing the averages in these various tables
was performed with the aid of a computing machine. This not only
greatly lessened the labor and expedited the work in hand but also
made the results less liable to error than would have been the case
had it been necessary to perform so many computations mentally.

In a final table (table 5), is given a compilation of temperature
records for species of birds that I have not been able to examine
personally in the flesh. This table has been taken from available
literature and includes only those records for which it has been possible
to assign specific names with certainty. Where a record is listed
simply as “ gull,” “hawk,” etc., it has been discarded. No attempt
has been made to cite all records available for each species but simply
to give enough to indicate the body temperature in relation to other
forms. Many published notes have been discarded for lack of certain
identification, while in utilizing other records I have simply quoted
what I have found with no assumption as to accuracy of statement.
Records are given for &9 species of birds in addition to those found
in table 3. The table has been made as complete as practicable but no
claim is made that it includes all records that have been published.

The system of nomenclature to be used in recording the data in
table 5 has given considerable trouble. The records cited cover
birds from all parts of the world. This material is listed according

NO. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 35

to the arrangement found in Sharpe’s Handlist * and the names used
are given in accordance with this list in most cases. Ina few instances
relating to North American birds, to avoid confusion with the system
of names found in the previous tables, the names given are those of
the A. O. U. Checklist for tg10 as in tables 3 and 4. Such deviations
from the general rule are indicated by reference to a suitable footnote.
This has eliminated confusion in generic names that might otherwise
arise, as for example in the case of the two closely related scaup ducks,
where the lesser scaup is given in table 3 as Marila affinis, while fol-
lowing Sharpe in table 5 the greater scaup would appear as Fuligula
maria. By using the name in the A. O. U. Checklist this is changed
to Marila marila thus dispelling any uncertainty as to the relation-
ship of the two birds in the minds of those not familiar with the
changes that have occurred in the application of generic names to
these birds.

*Sharpe, R. B., Handlist of the Genera and Species of Birds, 5 vols.,
1899-1909.

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

TABLE 3.—Summary of records of body temperature in birds

Temperature
: S as No. of
eae ; Mean | Mini- | Maxi- | 7ecords
mum mum
CoLy MBIDAE
AEchmophorus occidentalis .... | M. | 10t.3 I R.
Colymbus aurtius ~~. ti 02 se we M. | 103.7 I ce
. iL a anes ee F. TOAEO a eae mace I Le
Colymbus nigricollis oc. 025.00 M. | 104.2 | 103.0 | 104.6 4 ee
“ Sih lade cae Eas eed hele 104.9 | 104.7 | 105.2 2 a
Podilymbus podiceps ..........| F. O23 7.8 | upevets Dred I a
LARIDAE
LLGIUS CONPOTHICUS Ti oti tctmorars x6 M. | 106.7 | 106.2 | 107.2 3 R.
is ay |, Aueneetict Teter ee evorce F. TOO MS a waste: Cote I iS
Larus delawarensis ......%.... [Fest Mi TOOn |S I .
OKUSMOUTICLLIIGR ee ae eee | IME Pp i@s6G) | 4 fe Bhan I -
LL OAD Gopoe dose ats 2 | SMS) SOS sie LOAR Sele tOOFo 3 s
ae ele Laser heey ea eat ) IRE 105.9 | 105.0 | 106.6 4 i
Larus philadelplia ............ VEST) HOGS S| roa) eae I -
SiLEU0 COSPIO@s ae Seieee- ee nee Ni O53 LOZe 7a erOOnS 2 a
. We aE Mer neh chien br ek F LOZROd|een acne ae I
Sterna maxima ........02.0+:. M 105.6 I a
f Se ean ewer tae hasten 1B 106.8 I
Sterna sandvichensis .......... IME SW) HOOLO)') oaoe ie I Me
SHEVMG” FOFSUCTH | Sawa uee ete hens M.S 10623, |) 105.6") "10720 4 s
e Se Se: AA ee ea eee Tg F LOOMS G || LOSA yal Oss 4 a
Hydrochelidon nigra .......... M 106.9 | 106.0 | 107.3 4 uy
% i? ih sae, So F 107-0) |) T0085) | eLa7eO 6 a
RYNCHOPIDAE
VW GHOPS MUGKOs sites aioe ae M. | 105.0 | 104.5 | 105.9 2 R.
ANHINGIDAE
Anhinga anhinga ............. IW OS I Os, 67 I 1.
PHALACROCORACIDAE
Phalacrocorax auritus ........ M, 4) 1062)" 10h. 4-\167.0 4
‘< NAR eae ty ae oie F WOO UTI |) Wold .3 2 HY
PELECANIDAE
Pelecanus erythrorhynchos ... M 103.5 | 103.0 | 104.0 2 R.
ty Bosal Pn ale MOBS | Aas e Bae I et
Pelecanus occidentalis ........| M. | 104.6 | 104.2 | 105.0 2 i
ss ee Sp Ny ants F 104.5 | 103.4 | 106.3 4 in
ANATIDAE
MOR Quis “SErrGtor tase sceeeie te MEE sl BOy 5a 0 ocete ee wu RS
e Bah oh aise irate al aoc F. 107.5 | 106.3 | 108.6 2 %
Anas platyrhynchos ........... M. | 106.4 | 105.1 | 108.0 4
op ete Ciiinccran sans IR 106.I | 105.4 | 109.0 Gj oF
Chaulelasmus streperus ....... Wl} abe) Ps So Rone I N
Mareca americana ............ WL A ORES le Got eae it <
Nettion carolinense ........... Mie |) 166. 1 ||) 104.21) 10870 19 -
i PCAN OSS ous arn g ne F. 106.6 | 104.4 | 109.8 8 .
Querquedula discors’.......... IM: 5) 108:6i-| 107.7) |" 100.4 2 ¢
ui Ue aaa ean Ee F. TOZ AG cll Ges ete nas I ee
Ouerquedula cyanoptera ...... M. | 106.5 | 105.0 | 108.2 Ff Ke
ad (ook pear ge ws F. 108.0 | 105.8 | 109.1 5 ke
SHO? ANIGHO, a55Sebesedcace M. | 105.8 | 104.6 | 107.7 3 &
eC es ats a Ah Pe 107.3 | 106.3 | 100.0 B i
1D riled CHEMO 655 ooN seo ood obc Mel to. t «| st04e4 a |etOsso 9 <
. DR AELE snake Lara ee Re eae F. 107.5 | 106.9 | 108.0 2 a
Marila americana ............. Mi" | 106.3: | Ted.) To8s1 4 “
vs Pie F. 109.9 ; I 4

NQ. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 37
TABLE 3.—Continued
Temperature
Species Sex No. of
Mean | Mini- | Maxj- | records
mum mum
ANATIDAE—C ontinued
DVEQNLAS GRIMES, Ve Siac Pile ceraesc 3 hol M. | 106.5 | 105.8 | 107.0 3 Re
io POT iii AA Sete ei aepa tats M F, 106.4 | 104.4 | 108.0 7 ‘¢
Erismatura jamaicensis M. | 106.2 | 103.9 | 108.2 2 :
o We Micah eee eae (a. 108.0 | 105.4 | 109.7 3 :
Chen hyperboreus ..........66. F. TOZ RFE Meee chan I “
Brania canadensis ............ NI ATOOWON |e eiyen eae I Ry
s HES Dreier Ga Ae gape Fe: 105.8 | 104.6 | 107.0 2 -
IBIDIDAE
CVAD AUN he os okoe Se Oe oe F. LOSS Ontneeree: B Sear I IRe
Plegadis guarauna ........+.4. ES 105.1 | 104.4 | 105.6 5 se
ARDEIDAE
Botaurus lentiginosus ......... MESS eLOAC ON nce... spiel I R.
PAO O ENO OLUS. a his sa esteres eele si « M. | 104.8 | 104.5 | 105.0 2
$ op Soe Heke neue F. LO: 71 103.4) | TO3kG Be x
Egretta candidissima .......... M. | 104.8 104.0 | 105.2 5 s
a Meg aerate wesfecks Bx |) 104.0!) 102.2"| ‘16652 8 e
Hydranassa tricolor .........%. He OelOse5 |-10525. 1 105.5 z s
Butorides virescens ........... F. TOSRO uae cere ee I i
Nycticorax nycticorax ........ Mer | 0385 | 102. 4.|' Tose 3 <
= Tee Ds Oe vectra bas F, LO2,G | TOE <G)|" 1087 2 E
ARAMIDAE |
Aramus vociferus ........0005. M. | 104.5 | 104.3 | 104.6 2 R.
RALLIDAE
Railus virginianus ............ M. | 105.5 I ile
a oh gM A ec eR eRe 18, TOS OM eee Mae I is
Fulica americana ..........05. 2 |1°106.7| 105.9 | 100.0 12 R.
PHALAROPODIDAE
IE} OSMAN Brae COCR AOL Mr (107.6. |-305.6 | 109.0 9 iB
& Re Pe a en ae Re re F. | 106.6 | 103.2 | 108.7 17
Steganopus tricolor ........... M. | 106.3 | 104.8 | 108.6 10 o
uy HS Ste 3 pee Ss Be estO5274|\ 45030) 1074 18 3
RECURVIROSTRIDAE |
Recurvirostra americana ...... M. | 106.6 | 104.7 | 108.9 14 R.
Mi Dt ak eee ae ie 104.9 | 104.I | 106.4 12 i
Himantopus mexicanus ........ M. | 105.8! 104.6 | 106.9 10 -
- date (a aS F. 105.8 | 103.8 | 108.4 9 a
ScOLOPACIDAE
Gallinago delicata ............ M. | 106.3 , I R.
eee Gia Mase stink? IRE LOSE OU sera pee I Z
Macrorhamphus griseus ....... Me. 106. F< |, 103-4" |108. 0 9 =
i: Cae te iS) | 1e85.2:.| 102. a | 16824 10 =
Pisobia maculata ...........-. By «| 10740'\|. 107.0%) 107-0 2 1b
IPAS OUSENOGATOU Toate we cece bale ee M. | 107.9 | 106.5 | 108.6 4 s
g en hale (nk aR eae a BeeaqevOve2? tee eens I os
Pisobia minutilla ....:........- M. | 106.1 | 106.1 | 106.2 2 “
. Se eee gee wietes 6. 2 F. 106.6 | 105.9 | 107.8 6 ie
Peltdnavalpimm: <).c..20s.3.2 66. F. TOOLS! los eae Pitas I fe
EV CACICTESNINQUNL | ole wr ces cise oe M. | 107.4 | 106.4 | 108.2 4 sf
9g he Sh bot ee F, | 107.3 | 106.0 | 108.4 21 =
Calidris leucophaea ........... Miele 107223 /seirny. Fah te I
ee CN Sa a ee 12. 107.1 | 106.3 | 108.8 Gi i
TAPABSGTEGOW he. cc tk ea Caw s M. | 105.9 | 104.6 | 106.5 4 R.
Ee Up) os A Eee ae ee F. TOS 0) | t02.. 3) LO7n Ss 27, vi

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS

TABLE 3.—C ontinued

Temperature

: ax in No. of
trai P Mean | Mini- | Maxi- | Tecords
/ mum mum
ScoLopAcIDAE—C ontinued |
Totanus melanoleucus ......... M. | 107.0 | 106.5 | 107.6 4 Re
x ety oy ALN paket F. 106.4 | 105.4 | 108.0 6 uy
TOTAMUSMH OI PES ee eee eG M. 106.2 | 105.8 | 106.6 2 ‘f
Se itt Vie an F. | 106.9 | 105.4 | 108.2 4 :
Helodromas solitarius ........ Wie) PROGR bee Tolane I e
Catoptrophorus semipalmatus..| M. | 106.7 | 105.3 | 108.9 7 e
| F. | 106.6 |'104.6 | 108.3 | fo ae
Actitis macularia ...... sitet sae VE SOS a4 oars Aes I i
Numenius americanus ......... Mi) 205265) T04s:3 aoe. a 4 Ik,
CHARADRIIDAE |
Squatarola squatarola ......... M:;* | e671 | 205.3") “16770 4
¥ a S oneete ne tide halts KOO trai eae I
Oxyechus vociferus Epahatte acetate | M.. | 107.0 | 106.3 | 108.3 5 Ms
PASM ah eee Be) 107.) OGsT | Tob 36 5 2
AEgialitis semipalmata ........ F, NOOVOM| acre sian I ce
AEgialitis nivosa Abe ae eale M. | 106.8 | 106.4 | 107.3 2 es
me eeeutCRc PNT IG F...| 105.1] 104.2 | 106.6 3 8
Ochthodromus wilsonius ...... EAB TOS 200) tan eae bis I r
Podasocys montanus .......... IM || OS eaa i elO4 ese OSes 2
= Se Wr Pate AS F. | 106.7 | 106.6 | 106.8 2e sf
ODONTOPHORIDAE |
Colinus virgimianus ¢......5 0.6 M. | 106.8 I R.
SS) RoR ahve NO ae ae F 107.4 ue I -s
Onc Orie Picla ho. muss eee cok fot M 107.4 | 106.8 | 108.2 3 K
ars Kr re Pee F 107.8 = I S
Lophortyx californica ......... M. | 108.0 |. 107.4 | 108.7 4 *
Daa Pr ae ee eee Es | O70) | LOZ. ON |aOsey a Ms
CoLUMBIDAE | |
Columba fasctaia? ts acne... M | 108.0 a I R.
Zenaidura macroura .......... ME). MOORO) || sO7-38 eure II ie
§ pA ise diets eed ae F, 107.7 | 107.4 | 109.7 2 ie
Melopelia astatica ............. My 4 T0825), mo7.3)| sii. Ss 177 he
See Seite ee ete F. 108.9 | 107.4 | 109.7 8 ate
Chaemepelia passerina ........ Mi) 107525) TOOr6 aimnorne 2 Ip
CATHARTIDAE
Cathartes auras st Raia wien tee : M. | 103.8 I R.
BuUTEONIDAE
Circus: hudsonis :............ IR 105.5 i iD R.
ACCIPILErYUCIOR .bakae. foe leek 109.0 | 108.0 | II0.0 2 f
Bureouvorecusencs aan ene M. 105.0 | 104.2 | 105.8 2
Buteo, swaiwisont ©. 3.5 Ji ecs.02.. | Ma) 105.1 | 104.6 | 108.3 2 ne
- ROM Ota re 8 F. 105.5 Ay. I 3
FALCONIDAE
Falco PENS OTINUS ornate hee 18. 105.2 | 104.2 | 107.3 4 R.
Falco sparverws jc. 5: se cuns Mi 5) 106.81 hex I S
ALUCONIDAE
Aluco pratincola: ...-.542. 002: M. | 103.5] 101.9 | 105.0 2 R.
STRIGIDAE |
Asio wilsonianus .........-00.%- |i 10d4e2 I R.
AL SUONTLCIUNIVE SU ee ee WU | OA Soop AD I -
Oinscastor eee Lk M. “| 102.6 | 100.7 | 105.4 3 le
mo Saas ABP MSU ea MEE LS A 4 F. 104.0 | 102.7 | 105.3 2 5
Olas Hammeolus, oo vposnene M 102.5 I W

i

NO. I2 BODY TEMPERATURE OF BIRDS—WETMORE 39
TABLE 3.—C ontinued
Tepapendvare
: . TA; No. of
pees oa Ree ei | eat PPeCOr ES
mum mum
+ age |
StTRIGIDAE—C ontinued
Bubo virginianus <...0....5.66 F, 103.8 | 102.0 | 105.8 3
Speotyto cunicularia .......... 1% 106.6 . s I as
CUCULIDAE
Coccyzus americanus .......... iB. 108.1 | 106.9 109.3 3 ig
Coccysus erythropthalmus ..... IM) edo Gy |< ss/e. Lae I *
ss mG Air wide F. 108.3 | 108.2 | 108.5 2 H
ALCEDINIDAE |
GER VEL MUCV ON We temecvot niet cre soyeiar- Mi |s164.6 | 102.6). | 195.3 Be Hie
PIcIDAE
Dryobates villosus ..........4. M. | 108.2 | 108.0 | 108.4 2 iis
. a Deak ana A F. | 108.7 | 108.3 | 109.0 2 XG
Dryobates pubescens .......... Meme 20752) | 9100.27) 108.2 7 i
| Sea ath es ten cto c She i 108.3 | 107.9 | 108.7 2 a
Dryobates borealis ............ M. | 108.2 | oars I ss
S PS Waki. 108.5 | 108.2 | 108.8 2 ‘
Dryovates Sc@laris: ..2 0s a0. 2 eNO Ze sh MLOOnG. | e107 40 4 S
2 ail el ieee tea a eS if, 109.8 BAA oh byt I a
Dryobates nuttalli ............ M. | 108.0 | 108.0 | 108.0 3 is
is FE Me Ws cei a ole Ime 108.4 | 108.1 | 108.7 2 rf
Dryobates arigonae ........... M. | 108.2 | 108.0 | 108.4 2 “
PAGOIGES GFCUCUS \.)2..\5 ee ee Ys M. | 108.1 a I :
Picoides americanus:..........| M. | 107.0 | 106.5 | 107.4 2 &
* Pine nk ep ere ea ire, F. 107.4 oe I .
Sphyrapicus varius ........... M. | 109.4 tA! I <
. Liadeed Wises cai ole ct F, | 107.8 | 106.8 | 109.0 3 e
Sphyrapicus ruber ........606 M. | 107.0 : I
e ad ad ths Spc 8 ts eS F. 108.5 I %
Sphyrapicus thyroideus ........ Fee | pat Oech || cae Nee I
< mae eT ets 2 F. 106.3 | 105.2 | 107.3 2 ex
Phloeotomus pileatus ......... NES | Srozeo! | 06%Or |) 10722 3 R.
Melanerpes erythrocephalus ...| M. | 108.0} ... as I i
os nook dae 108.6 ae I <
Melanerpes formicivorus ......| M. | 108.6 | 108.0 | 109.2 3 £
es Fea enoSe 2a, lO 75 OSs 2 os
Asyndesmus lewist .........0+- Wile? dprutetsielonll see 4 I ‘f
% eld Wi hemirata sole a Bye) |pt07e3 ares I ie
Centurus carolinus ............ M. | 109.4 | 109.4 | 100.4 3 §
Centurus uropygialis .......... M. | 108.7 Lalas I 3
- eee WE Gs cone ss Ho) hae7 20 sh ints yar i
Golaptes SGUraius © ia oak...) 38 M. | 109.1 | 108.3 | 110.0 2 a
ONE PEES (EGREM tanto. lam tae o 8 Mle) kaye a =] I af
kg Fo eS. Ba ed se bates F. 108.0 u <
Colaptes ‘chrysotdes. 2200+. 5.. M. | 105.8 | I f
e POR pie ee a Bue} 1086 I cL
CAPRIM ULGIDAE
Antrostomus vociferus ........ M. | 108.4 I il
Phalaenoptilus nuttalli ........| M. | 107.2 I s
Chordeiles virginianus ........ VERS | mOOR2m hoe te Stes I S
se Mtns A504 2 S56 F. 105.7 | 105.0 | 106.4 2 §
Chordeiles acutipennis ........ M.* pao7s8 Wo rezs 7 |} 107.6 2 2
<i ae pee F. LO7 14: ere Som ane I et
MiIcropopIDAE
Ghactura pelagica, ....5. 2.45. F, 107.2 Saath I Vs
Aéronautes melanoleucus ...... M. 106.0 | 105.7 | 106.3 | 4
«“ « ely huessa) |) 105.2 tos a fe =

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.
TABLE 3.—C ontinued
Temperature
Species Sex ne ; No. of
Riot, Mini- Maxi- records
‘TROCHILIDAE
Archilochus colubris .......... NVI ee OGsOt eres ee I
e red Dechert ek F. 101.4 | 100.8 | 102.0 2
Archilochus alexandri ......... M. LO32 0) | § 10247110326 3
= Sh, ate, Me PRR NS F, LO237A |e LOT Aa enone 3
Selasphorus platycercus ....... F, Oly Si are Nees I
S@lasphorus rufus .06': Jas%.. NE) 802525) "TORTS 102-6 eS
Cynanthus laturostris ........-- M. | 105.9 | 105.1 | 106.5 3
TYRANNIDAE
Tyrannus tyrannus ........... er HOPI Wag a I
Lyrannus verticalis ..0-..+..0s M. | 108.5 | 108.3 | 108.6 2
Lyrannus vociferans .........-. M. LO ARCO) Ion greece ee I
i hit Le eee F. NOS20 le peer ieee I
Mvarchus cinerascens ......... Mena) TOOkS || TOS sOnennnss 4
a PM ls? kone ase ge F. TO ||) 100. 2h eriteo 2
Myiarchus lawrencei .......... M. | 109.5 | 109.2] 109.8 2
e ia aoe shies F. L107 : eee I
sayornis pPloebe«. a... sie bees Whe leat@ays0).'|] S56 é Pees I
Sayorms nigricans ...........- VE | OOH Te LOS: Ol) e1OO87, 3
or ee pam iats chet roca F, IO ROT Gro seat I
Nuttallornis borealis .......... M. 109.0 | 108.0 | I10.0 2
y aL 8 Soe Be ake Soe ie LOD Atal, cise hee xigs
Mytochanes pertinax .......... F. TIO RSy| LOG. 7a elite o 5
Mytochanes virens .........4.. IMIS | atsje@) | aise ook I
Myiochanes richardsoni ....... INE, |) BPTo).(e) i) sWoys}.(on|| wt Fe 4
3 St a ak F. CUOLA Herre te pcre I
Empidonax flaviventris ........ M. | 108.0 | 107.4 | 109.0 6
vi regret mukty a Boe Wee. F. 108.2 | 107.6 | 108.7 4
Empidonax diffictlis ........... Ni |S t0822) |" 1070s 100-8 3
Empidonax virescens .......... M. | 108.8 | 108.4 | 109.2 2
3% APR Pee yarn eS F, EOS AOm Wires eh I
Empidonasx tratllt ............. M. | 108.0 | 107.5 | 108.7 6
3 eieieteenea mater ate F. 108.6 | 107.2 | 109.6 4
Empidonax minimus .......... IES ROY IS NP es one Lee I
mi Warne Mettces eae 15, 108.3 | 107.6 | 109.0 2
Empidonax hammondi ........ WES aoe ee ae I
3 Ae gta toy 12, 107.9 | 107.3 | 108.7 B
Empidonax wrighti ........... MER eTOSi3 |= 107 .7an snOGr 3
Pyrocephalus rubinus ......... M. | 108.6 | 108.2 | 109.0 2
ALAUDIDAE
Otocoris: alpestas: si, Wake ceateon M. | 109.4 | 108.6] 110.4 B
4 al, yelcathniid skeen 1B 109.4 | 108.6 | 110.3 6
CorvIDAE
PIC NPICG Sete eee «ioe he ee M. | 107.3 | 106.4 | 108.6 9
cs Boe Stee ats enna ees F. 107.1 | 106.2 | 108.8 9
PAGO MAL QUD Ge retire ie ae he WES? | aecapssea 5 Se sae I
*: Riad Pon ee ear OM Utena ID. LOZa7 |) LOO OAS 2
CVanocwia (Cristatd ere icc toees M. | 108.4 | 108.0 | 108.8 2
i near... Ser F. 109.3 cp I
Cyanocitta stellerit ............ ME. RISEOZS AAP take rs es I
a a eet ah Eh IR 107-9) | LOZ ss Lose 2
Aphelocoma cyanea ........... Mi 8) 108.2) |) L07ez We rOses 5
eS Sr PS 2 a bin F, 107.9 | 107.1 | 108.6 2

Wi

|
|
|

NO. 12 BODY TEMPERATURE OF BIRDS—WETMORE Al
TABLE 3.—Continued
Temperature
; * No. of
Ra ae Rneeae i Meum | Mai. 9 | Pece rae
mum mum
CorvIDAE—C ontinued :
Aphelocoma woodhousei ...... M. | 108.6 | 108.4 | 108.7 2 ie
ie Spay ne pak: cea F. TONG rele ok ate 3 I fe
Aphelocoma californica ....... Vie 107-5) |) 107.3) 107.0 2 pe
: ee tas tes Coes F, HOSS TA |e ise She I i
Aphelocoma sieberi ........... M. | 108.5 | 108.2 | 108.7 2 +
a +; 1B, 108.4 | 108.2 | 108.6 2 3
Pertsoreus obscurus ........... M. | 108.2 | 107.8 | 108.6 2
Me MAREE aus che char cere iets F. TUCO Faecal [a ga, : I e
(CORUS. COREA Se SAO ODD EAD OT ING NO 7ie4 iene aie I [Pe
e Cae ths ct SNS EERIE F. 108.6 | 107.6 | 109.6 2 »
Corvus cryptoleucus .........- INE LO77. 02) 107.09) 10727 2 S
a Gr liseece eee or Te TOOUG. |eanre seaoie I &
Corvus brachyrhynchos ....... M. | 107.9 | 107.6 | 108.4 4
Nucifraga columbiana ........ VES a0. Gall nis... SFist I 2
hs Oe radar an F. 107.1 | 106.8 | 107.4 2 :
Cyanocephalus cyanocephalus ..|. M. | 106.6| .... eran I 1s
se : ieee Ne Tore pace I Lee
IcTERIDAE
Dolichonyx orysivorus ........ IME eat O72 OF sree ae I Tie
WHOV GENIUS] GLE A jercg see. 4s Nipes posal aloes) tose 4 eS
z Nah ove cek ehel a aleraiet 12, 108.2 | 108.0 | 108.3 2 s
Xanthocephalus xanthocephalus| M. | 108.3 | 107.0 | 110.0 29 o
= BS 108.2 | 106.6 | 110.3 18 ye
Agelaius phoeniceus ........... MS 168: '3)°| 106.5" || Tro. 3 10 rs
3 DP eae Eee F. 108.1 | 106.7 | 108.7 6 <
PU GAUUS THICOLOL eo ois con sos awe os | FOSS eI), se aor I e
a ine) AVA eer artes iB. 108.6 | 108.0 | 109.0 4 4
Sturnella neglecta ............ VERO. OF rier RiGee I a
ss on iE ti eee 1, 108.4 | 107.8 | 109.0 2 a
Tckerus PATISOTUM «oo cee cc ce ces M. | 107.9 | 107.2] 108.4 4 if
cs 2g a enc ree es F. 108.1 | 107.8 | 108.3 2 .
Icterus cucullatus .........0..- Mee | S10O. 38 | 1OS.3el Bros: 2 5
e Mf 3 te ec ee ees F. 107.6 at athe I %
VGCtETWS SPUTIUS Dee ives 8 INE. || rietclehip ile, apt A oe I a
WESERUS NG GIONS 62.0 ca haha Bais M. | 1090.1 | -tog22: | ‘09.2 2 my
Ugiemes UN GCK oo. eke. o's F, 108.0 | 107.8 | 108.2 3 S
Euphagus carolinus ........... AY, ees am oa ls ch oO GLO O) 2 :
i HEN i A ae et F. 109.7 | 109.6 | 110.0 3 ¥
Euphagus cyanocephalus ...... M. | 108.7 | 108.6 | 108.8 2 «s
. STARR Ts ean eae F. 1OZs De eLOGe Te | NOGE 2 <
Ouiscalus quiscula ............ F. LOGO i cee: A I :
Megaquiscalus major ......... M. | 109.4 | 109.3 | 109.6 2 y
Ue et are F. TOSCOR| meet ae I R.
FRINGILLIDAE
Hesperiphona vespertina ...... M. | 100.5 I T.
bi Phy eee Say chen G F. EOOIO0|) kA Se aay I -
Carpodacus purpureus ........ M. | 108.3 | 106.7 | 109.5 6 “s
: WAIN pb sayite seh. F. TOOKS | Mavs ye ht I
Carpodacus mexicanus ........ M. | 108.9 | 108.0 | I10.0 20 3
: Cp eS Ree iB, 108.8 | 108.6 | 109.2 7 :
Lowia curvirosira .......0000.. M. | 108.2 | 107.9 | 108.6 3 oe
TFOTADAVCUCOPECKG, 22.2 feos revels F. 109.4 | 108.9 | 109.9 2 *¢
Passer domesticus ............| M. | 107.6 | 106.8 | 108.6 7
MOE Sais cs Yen F. 107.9 | 107.0 | 108.8 2 be

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

TABLE 3.—Continued

Temperature
Species Sex ia ean my
Mean | Minit | Maxi
FRINGILLIDAE—C ontinued
Astragalinus tristis 06.0... e M. | 107.7 | 106.0 | 109.9 4 fr
rs Br Wa Aaeete fabs hos F. 108.1 | 106.6 | 109.7 2 a
SPIMUSSPINUS Pune hs Caw kate M. | 108.3 | 107.4 | 108.8 3 s
i BSE Cet ta Ware Sateen Saya F. 107.3 | 106.7 | 107.6 3
Calcarius lapponicus ..2.::....| F. | 107.0} «... sree I :
Passerculus sandwichensis ..... M. | 108.3 |} 106.6 | 110.0 5 *
Pe en ee eet 1m, 109.2 | 108.4 | 110.0 3 %
Ammodramus savannarum ....| M. | 108.6] .... “sae I s
Passerherbulus henslowi ...... IV pale LO 2 ees: nee I o
Chondestes grammacus .......| F. 1Og,.2 |) LOSS3|/nr0.2 2 i
Zonotrichia querula’.........- Mt me822:-) 107", 5, |) noes 3 ne
a Men ch nceeratiacee ote IR iWotsste) o 65 o seat I a
Zonotrichia leucophrys ........ M. | 109.0 | 106.8 | 110.2 4 se
re Fe een Sec 18 108.9 | 108.8 | 109.1 2 5
Zonotrichia coronata .........- VISGs tO Siac aeceeae ee I s
M ehaeein ta Ari cacy ache F. 109.5 | 109.4 | 109.6 2 HR
Zonotrichia albicollis ......:.. Mj) E10-2+)| 100s 4e) TIO. 5 2
Spizella monticola ...........- IME 8] S808 50. | 28 os I if
u ok LAME rae eee ee 1, 108.6 | 107.3 | 109.5 8 Pe
Spizella passervind .........208% MEN) MOO. Gan ee ayer I ‘*
7 SEE i osc toutes tee F. NOLO 0) || Stes ie I 24
Spizellan pallida. ascmewaeeeese M. | 108.0 | 107.5 | 108.4 3 “
Spigella brewert .0.....2..20% Wil, * |) alse | nal. || tits. & 2 Ite
re Sper artes yates baer 18. 108.7 | 108.0 | 109.4 2 ‘i
Spiele pusilla d (Le Meee ce nee MEMS t07—e4 letoGe2 spose 5 >
4 Vi ante a Ata tel Apt Esa POR A251 eo. Fis I
Junco hyemalis’...... ses ees os M. | 108.6 | 106.8 | 110-0 8
' Le anak eee ne ALL F 108.8 | 107.1 | 110.3 3 :
UPA) OALGUAIS bobeddnsosens 6c M 109.2 | 108.5 | 110.6 y i
Re. ees peat ees 5a F 108.5 | 108.6 | 109.0 2 hs
Junco phaeonotus ......2...00% M 108.3 | 108.0,| 108.9 4 .
‘a 5 her Wed Habba eectaoie. sete te F 108.8 | 107.9 | 109.7 6
Amphispiza bilineata M 108.0 | 107.9 | 108.0 3 rs
i SAP whist one F. TMO)GO || oe oe iebels I =
Amphispiza nevadensis ........ 12 109.4 | 108.6 | 110.0 3 .
Pencacaaesitualis sna eee te Wale wall ioyestial Gao BY es I SS
Aimophila ruficeps ............ Mie 1098) |) LOO. 4a entO.2 2 a
Melospiza melodia ............ M. | 109.1 | 108.0 | 109.8 II ie
e POR ee hcgees hy cae tee F. LOO D | LOdeas |e LLone 4 x
Melospiza lincolmi .............| M. | 107.8 | 107.2 | 108.5 2 ee
Melospiza georgiana .......... M. | 108.8 | 107.9:| 109.8 3 a
* Rol are eta oe F TOG Geil) fetens ahs aie I of
Passerella AUGcd) sore seene ee M 109.3 | 108.5 | 110.0 3 ‘s
e oF Dia nee ee F. 109.3 | 108.6 | 109.9 2 i
Pipilo erythrophthalmus ....... M. | 110.0 | 109.6 | 110.3 3 e
Papilommnacwlarisn nae aiecteiene F. 109.I | 108.4 | 110.3 3 ye
PApilo tii scus (te ocpes sore cee M. | 107.9 | 107.6 | 108.0 3 eS
Pipl CPASSalis ss Wnt aie cpt aks M. | 108.5 | 108.0 | 109.6 3 as
ii Ri Aang ehet Ousiecciben rae ade ape B 107.8 a I
Pipilovaveriag. sate sees eed en OO n| I ‘
f UVES Stucten Gm tess F. THO Aaluueeraa sae I e
Cardinalis cardinalis .......... Mis) 10083) |e 1o7. One ntose 7 s
¥ SUM ND x ccg Seok Se F. 109.3 | 108.6 | T10.0 2 “§

NO. I2 BODY TEMPERATURE OF BIRDS—WETMORE 43
TABLE 3.—Continued
Temperature
: No. of
aes eS Mean | Mini- | Maxi- | Tecords
mum mum
| |
FRINGILLIDAE—C ontinued | |
Zamelodia ludoviciana ........ M. | 107.8 | 107.3 | 108.2 eae: it
s daw ONES Fe” ate F 116 eer ae ee tia | I s
Zamelodia melanocephala ..... M 108.5 108.2 | 109.0 5 «
as Seine ee he ate F. 108.5 | 108.0] 109.5 | 4 ss
Gutraca caerulea... sic. cede + Mz 4 108.0: |-107.0: | 108:6' |= 3 ‘e
Passerind cyane@ .......300.0% Mrs 0207.6: |. 107-5. | 210756 2 5
- CF erate ey Se HREOC) fetes oc litre se ¢ I 5
Passerina amoena ............ Vie OS sOnile meres < aot I i
% Pech ech ke ee ee F. 108.2 Be edt: © rea I
Sipie@ QMeMCANG.. 2s se. vee Mee 108-3 | 107:0"| Teele. 8% ‘6
Calamospiza melanocorys ...... M. | 108.3 | 108.2 | 108.4 | 2 -
TANGARIDAE |
Piranga ludoviciana .......... M.. | 108.4 |108:1 | 108:7.| 4 IL
ms cakes gtd oh her Sate Bea ie TO3 A rence: Pees || I s
Piranga erythromelas ......... NE 107.8 "|. 106,85 | T6925.\) 24 i.
es a tia ie ea Co ert ere 1B 107.3 ee eae I f
Paring Wepatca> os. 00d oe M. | 108.6 | 107.2 | 109.4 5 “
: 4 eae Se Se ee F. 110.2 | 110.0 LEOPAN ee as
TAR ONTG TUDO ae ee hes oss ke hace Vip TOO RO ley rahe pte a I ie
ez is Ue Baaeen Pet Re eee a an F 109.3 | 1090.1 | 109.5 | 3 e
HIRUNDINIDAE
IETANOTO ISVIGIS Mec AEE E Me | 107.4 | 10720. | 107-7 4 2
‘ EES ey teeth ot F. MOO Gel awe ste mies I
Petrochelidon lunifrons ....... Mae | -106.:3" | 10570: | To7 x4 4 es
Hirundo erythrogastra ........ Veet eTOORS arose wa tal I He
Iymidiprocne bicolor... 00.0.6... M. | 106.8 | 106.8 | 106.8 | -2 s
Fe 5) ea Se oe F 107 .0''|'"£06,0-|'107.9)|" 2 co
Tachycineta thalassina ........ M 105.7 | 104.6 | 106.8 2 4
+ Meer F. TO5e5 noeee aBaS I ¢
Riparia riparia .........+.+.6. | M. | 105.7 | 105.4 | 106.2 4
Stelgidopteryx serripennis ..... M. | 108.8 | 108.7 | 109.0 2 ie
BoMBYCILLIDAE | |
Bombycilla cedrorum ......... M. | 108.2 | 107.2 | 109.2 3 I;
ss So Os Wa F 109.7 | 109.0 | II0.7 | 3 s
PTILOGONATIDAE
Pha'nopepla mitens ........... F., 107.4 | 107.4 | 107.4 2 Te
LANTIDAE |
Lanius ludovicianus ........... NS) 1075) |e ae I if
+ OU oe F 108.0 | 107.3 | 109.3 3 s
VIREONIDAE |
Vireosylva olivacea ...........- Min) 10825" |107-95|| 109.27) 92 ie
‘ ii a ae F. TOOT eee pete I Ue
Vireosylva philadelphica ...... M. | 107.3 | 106.8 | 107.8 2 “
PP OSMIUE CHUN 5 «is on ea ole we M. | 107.5 | 107.0 | 107.9 3 ss
s See ME, Sere, F 107-6) || 1076) | O77, 2
Lamvireo flavifrons ........... M iofsis(0) ll “aes Perec I ef
as Sen Var us F 107.7 | 107.2 | 108.1 2 a
Lanivireo solitarius ........... M 108.0 | 107.8 | 108.3 2 eS
ie 4 beet Pcie fs F. 107.3 | 107.3 | 107.4 2 He
VSCOM GVISEUS: io. sje snd ns aes F. TOA) || parce oe Facieh I 4
VALCOMBULIONY Ciiscs dnvodiee iscses Ala ae TOZ_O! eee one I -
RECON aa ree: cole! soe ss M 107.5 | 106.5 | 108.3 3 =
BE St ee ao ee F TOE Se Laas Foe I ie

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS . VOL 7z

TABLE 3.—C ontinued

Temperature
P ic coast —— | No. of
ey ice Mean (| Mini- | Maxi 4) "eceade
mum mum
MNIOTILTIDAE

Mniotilia varia ............ S| Me P108:00| 107-0: | -re0r6 5 Te
ae Bay cas lhe, Scions acalaseatt Es TOSs On| meet -taeye I ES
Helmitherus vermivorus ...... M. | 107.4 | 107.2 | 107.5 2 ‘.
if pee erent ra 1D TOS 200 ste aye eae I Ss

Ver mivora. PINUS ..i2-.000.-0s- M. | 106.9 I &
Vermivora chrysoptera ........ M. | 109.4 I =
“ apo moh A OE ce te ES TOSS2 uieierioe Siac I 5
Vermavora lucige (....6-.5+. 26s M. | 108.7 | 108.6 | 108.9 2 és
; Sg ko tayk Rena ae 195 108.8 | 108.0 | 109.5 2 a
Vermivora virginiae .......... M. | 108.0 | 107.8 | 108.3 2 -
7 eS aes atic 1B 108.2 | 108.0 | 108.5 2
Viermaviora Colt. <i: was mre ciate M7) 107.37. 10653") 109.1 2 iy
in Cotes att are Miapaxecsn ee F, LO7MS lO7e 2 ae LOpea 2 -
Vermivora peregrina .......... M. | 107.9 | 107.5 | 108.0 4 2
~ Een arte reese F, 10) s ke Al gra ane I 7
Compsothlypis americana ...... M. 4) 107.2%) 106.7 | 160728 3 ©
: Meet erie Crane FE; LOS.A foo. oe Gee? I g
Peucedramus olivaceus ........ MES LOZAS Ll) perce: BS Ba I if
Dendroica tigrina .......--..0- M. | 108.8 | 107.6 | 109.9 2 f
Dendroica aestiva ...........+. M. | 108.6 | 107.9 | 109.8 3 .
Dendroica caerulescens ........ M. | 108.0 | 107.5 | 108.5 6 ie
Hd a gs Ream ae ee F. 108.6 | 106.8 | 108.4 2 s
Dendroica coronata ... 3.2 c.6 F. 108.5 | 107.6 | 109.0 4 x
Dendroica auduboni .......... M. | 107-7 | 106.9 | 108.4 3 Ee
* - cco Cet een le Et LOS OF 2s eae eer I sf
Dendroica magnolia ........... M. | 108.0 | 106.6 |- 109.3 5
= iia he ok Sagas einer Ee wO7eG lOO manosee 4 a
Dendroica cerulea ............ Wee Ai oT Ve0) | Gee a Pea I ss
Dendroica pennsylvanica ......| M. | 108.6 | 107.2 | 109.4 6 ce
i Se RW eis d cicte F. 108.4 | 108.0 | 108.7 2 =
Dendroica castanea ..........: MS St07200| 107.22! TOSa7 7 2
i Boe BAR ance F. 108.8 | 108.7 | 108.9 | 3 s
Dendroica striata .............| M. | 107.8 | 107.2 | 108.5 3
« i rm ee beatin F. 107.4 | 107.0 | 107.8 2 cs
Dendrovea sfuse@ Wea eee cere M. | 107.8] 107.8 | 107.8 3 y
ca Teams teete a aot Ads Lelsyhinte Be 108.2 | 107.8 | 108. 2 €
Dendroica dominica ..........- M. | 108.3 | 108.0 | 108.6 2 A
Dendroica graciae .........-+: M. | 108.0 | 107.2 | 109.5 4 Ai
Dendroica nigrescens ......... F. TOS Zrii mewetere Lgetis I S
Dendroica virens ...........-- M. | 108.0 | 107.6 | 108.5 9 i
Dendroica vigorst ..........+.- M. | 108.3 | 108.0 | 108.6 2 S
a SANT ue atin bok a emerge Ie F. NOOS2/ | eer shiek I .
Dendroica palmarum .......... M. | 108.9 | 108.0] ro9.5 3 is
" Sein RR Re sink eee 19. 108.3 | 107.8 | 1090.2 4 -
Dendroica discolor is. ccs.. 0+ - M. | 108.0 | 107.6 | 108.4 2 ©
Seiurus aurocapillus .......... Me | 107.4. |\ 07st). 105.0 3 <
Seiurus noveboracensis ........ M. | 108.5 | 106.2 | 109.7 4 i“
te aT oi tak secoes ate che F. TOO IO) th yeecsas Agit I e
Seiurus motacilla ............. M. | 100.4 I
Oporornis formosus ........+4 MA) TOF A ies AS I ie
Oporornis philadelphia ........ M. | 107.7 | 107.3 | 108.0 3 s
Oporornis tolmiei ............ M. | 108.2 | 107.0 | 109.4 2 “
ve aon ney See Yel Ps F TO? ol) sees eke I is

NO. 12

BODY TEMPERATURE OF

TABLE 3.—C ontinued

BIRDS—W ETMORE

45

Species

Sex

MNIOTILTIDAE—C ontinued
Geothlypis. trichas .o.0555 560s
“ a3
Icteria virens
“ce “cc

ee

ey

Wilsonia citrina
Wilsonia pusilla
“ “c“

wie 10,0) .9) wie 0 e106 en 6
acre (siis 0) is) (e616 «) ws ee
|
ee]
o fel, *: e:16,"6, ee 8 0) ete.
ee Ce ee CE a)
Cr ec ey

ee er)

MoTAaAciLLIDAE
Anthus rubescens
‘
Anthus spraguei
CINCLIDAE
Cinclus mexicanus
MIMIDAE
Oreoscoptes montanus

O40) a6 10)/8).0 (0)'6).0)6 «60
CCN gt Se Cir uC Be Sar |

Mimus polyglottos ............
“c “
Dumetella carolinensis ........
“oe “
Toxostoma curvirostre ........
“ “
Toxostoma crissale ..........+.
“ “cc
TROGLODYTIDAE
Heleodytes brunneicapillus ....
Catherpes mexicanus

Thryothorus ludovicianus
“cc ce

Thryomanes bewicki
“ it3

Troglodytes aédon ;
Nannus hiemalis

?
Cy
a ee ey

CERTHIIDAE
Certhta familiaris .............
“ “c

ee oe |

SITTIDAE
Sitta carolinensis ...0...c.c000e
“ oe
Sitta canadensis .........0000
“ce ia3

silells) isis)» |<) 6i0 6.4 6

OPEC PMN ted Ciosiaes wd eee es
Sitta pygmaea

SisiSin ela s]is = «6 6 66 ve

by

2

Temperature

Mean | Mini- | Maxi-

| mum mum
TOGHOM |Wreves's oar
108.1 | 106.6 109.0
109.2 | 108.0 | 110.4
O/C) oe BON sealers
TOOROM pais. c ee
107.7 | 106.8 | 108.6
OORO: Raa Sire
LOZ. 2) (P1057) || 107-0
107.6 | 107.0 | 108.3
TOOLS Neracrs trees
TOOFOM eae sears
108.5 | 108.0 | 109.0
BEG On| ae See:
107.9 | 107.6 | 108.1
109.1 | 108.4 | 109.8
108.6 | 108.6 | 108.6
107.3
109.0
NOORZ) eck Sea
109.2 | 108.1 | 110.4
109.0 | 108.4 | 109.6
TOSHON eae SANE
108.7 | 108.6 | 108.8
TOSBON |G aieee hettes
109.7
108.5
109.5
108.5
TO7ROy | eee Sates
108.9 | 108.4 | 100.7
108.4 | 108.2 | 108.7
107.9 | 107.4 | 108.2
LOZ 50 e107 5 1107.0
TOO 2 alent. Sie
107.3 | 100.6 | 107.9
107.4 | 106.2 | 109.2
105.7 | 105.2 | 106.6
107.7 | 107.1 | 108.5
TOF A, Nos ar ee a
107.7 | 106.8 | 108.3
107.7 | 106.0 | 108.8
107.9 | 107.5 | 108.2
107.9 | 107.8 | 107.9
TOOe2 4 ecto ot Rice
107.8 | 107.8 | 107.9

No. of
records

HHH HD HY RH HR i] tb b&b & MWHHWNHR OK SH NW HE

eh WNT NWPRW HOH

NHWULWwW

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

TABLE 3.—C ontinued

Temperature
; se No. of
we 7 Mean (| Mitui- |) Max) Seeeeee
mum mum
PARIDAE

Baeolophus bicolor a). iceia.» cia. M. | 108.8 | 108.6 | 109.0 3 ils
= BEM a bib yan cutie scrcee te F. 109.4 | 109.2 | 109.6 2 a
Baeolophus inornatus ......... Ni 10875210727 4s TnONo 4 «
i Pads a hate tle arcte se FB; 108.6 be I Ge
Baeolophus wollweberi ........ MiP 10756 acts iF z
E peter Pal Mesiee rs annie F. 108.5 | 108.3 | 108.7 | 2 =
Penthestes atricapillus ........ M. | 107.9 ee. oe I as
. en te eens | F. 108.7 | 108.2 | 109.6 3 ee
Penthestes carolinensis ........ M. | 108.0 | 106.6 | 109.4 4 ze
es ips on EAD Spree to F. 108.3 | 107.5 | 109.3 8 S
Penthestes sclaterd oi... seed «. M.. "00723" |) "10653"|a168.4 2 93
oe OASIS MA ney city ke 229 F. 107.8 ae I u
Penthestes gambeli .........-.. M. | 109.0 ote I ¢
f SON Ey Bah Ate. te cae F. 108.3 | 107.9 | 108.6 2 “6
Psaltriparus minimus M. | 106.0 | 106.0 | 106.0 2 =
ss daa Mat ath NN? 3 F. 105.3 | : I x
Psaltriparus plumbeus ........ M. | 108.2 | es. oa «“
me CD a ante arc, F. 107.4 ise I a
Auriparus flaviceps ........... M. | 106.2 | 106.0 | 106.5 2. S
y UMW seks rece eh aes F, 107.5 | 107.5 | 107.6 2 xf

CHAMAEIDAE |
Chamaea fasciata ............. M. | 106.8 7 Lo ee qs
FY (Rae rocmton, F. 108.0 | 108.0 | 108.1

SYLVIIDAE |
Regus SGUrap Gs coi iscsi sis oes NEN LOOLON I 106.35 20723 4 ifs
- Se pa Re Beker AR a F. 107.7 ze I “
Regulus calendula 3..4%.-. 5. M. | 106.0 | 106.0 | 106.1 2
es papi Mg A eg F. 107-7 + |) 207.3 OSes ss a
Poltoptila caerulea ............ Mee | 10727 || 106.0 1210647, 5 #
s Fora ge Rad A ae eae F, 107.6 = a I Wy
Polioptila plumbea ............ M. | 107.8 | 107.5 | 108.0 2 se

TURDIDAE |
Myadestes townsendi ......... F, 109.3 | 108.6 | 110.0 2 1
Hylocichla mustelina .........- M. | 109.0 | ce I ay
Hylocichla fuscescens ......... M. | 109.0 | 108.6 | 109.4 2 i:
Hylocichla altciae ............. M. | 100.6 eee Ghee: I e
Lat ear Ra a ae 108.3 | 108.0 | 108.6 z Ss
Hylocichla ustulata ......:.... M..-}108.3°| 107:2 | 109.5 2 .
: an hah ois pate F. LTO. AS eT TOR mOsS 2 z
Hylocichla- gutiaia ..6...-..45- M. | 109.3 | 108.4 | 109.8 6 se
ss Pas onan deka, wk 18 107.6 ae om I 3
Planesticus migratorius ....... Ma) 10978" 107.0 | a0 rieo 8 "s
de Se MR aes F. 109.7 Se I x
Sali *stalis ne scdiccm ice ceh eect M. | 108.1 | to8.0 | 108.2 2 ie
- SDR Sa a at lee aan ee F, 108.0 I -
Siga MeXICANE Taiee. ores os M. | 100.6 I be
Sialia currucoides ..........%. M. -| 108.0 | I =

NO. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 47

TABLE 4.—Average temperatures of families of birds, summarized from

Table 3
Temperature
Famil No. of : No. of
y species records
Mean Minimum | Maximum
Keaimenidael iss aia es 4 103.6 101.3 4.9 10
Weer laces a ka ser rae 10 100.3 105.1 LO7eOMen 38
Rynchopidae ......... I HFEVSE (0) Rnd gue ep nee ee ae 2
Anhingidae 2244.4... I TO Siem tel be eye aaa I
Phalacrocoracidae .... I 100.2 106.1 106.3 F
Pelccamidae 12... 2 103.6 103.5 103.8 9
PNTAATIGAE eons ieicioie < 14 107.0 ° 105.8 109.9 98
Pbadidae = 200°. siiew a's, <3 2 106.5 105.1 108.0 6
PMR GeICAG serena fostive.e 6 104.3 102.6 105.8 26
IAGAINIIG AC weisves eee. c I LOR GR wiles has aN WiNM ec tocdey 2
Ralilidaene.ty mcm conte: 2 105.9 105.5 106.7 14
Phalaropodidae ...... 2 106.6 105.7 107.6 54
Recurvirostridae ..... 2 105.7 104.9 106.6 45
Scolopacidae™.:..:.... 15 100.7 105.1 109.1 140
Charadridaé .: 2.8... 6 106.7 105.1 109.0 26
Odontophoridae ...... 3 107.6 | 106.8 108.0 13
Columbidaey =. sce 4 1098.2 |) 17.2 109.0 30
Gathartidae, ase: case. I HOSE Osea mane Aras I
Bieonidae’ of... 22. 4 106.0 | 5.0 109.0 8
Fie MINU AL... eee orcs 2 106.0 | 5.2 106.8 | g
PINE OUNGGE Lh. cela. ce oe i TOGO de Riceas ant Mk Meera | 2
Strietdaen sh oases. oe 6 TOSL GE ero? Pe sirteata Ghia 12
(Gtrctindaew es. eu re 2 108.7 108.1 109.6 | 6
Aleedinidae’ <2... .. I TOARON Hip ees eee 2
PiGidaek 220. e sols ota ce a 20 108.0 | 105.8 109.8 | 64
Gaprimimloidae ss3. 2. - 4 107.1 105.7 108.4 | 8
Micropodidae ........ 2 106.2 105.3 107.2 7
Mrochilidac 3.2.2... . 5 103.2 101.4 106.0 15
iyrannidae S+..!0 vse. 19 TOSe7in le, LOO. 2 110.4 71
Nam GIdaGa, wasn ae eeieer I TOS ARO Me tact tage <td See. 9
GOTrvAdae’y = sas sec at 14 107.9 | 100.6 TIO). 3 62
Nictenidder cet sass 15 108.4 107.0 110.3 106
Brinewlidaes 2s... «.2 46 108.6 106.2 110.4 236
dlianmarnidaen's a. ec <6: 4 108.6 107.3 110.2 21
lnfiribhate hints es ea geno nine 7 106.7 | 105.5 108.8 23
Bombycillidae ........ I NOCROD | NeLOG=2 109.7 6
Ptilogonatidae ....... I OWL vale Oeantest shh Ra ote 2
Ibayaihiceye.) Winans Gaon ee I 107.8 107.5 108.0 4
Wineontdae: -jacne sss e's 8 107.7 106.3 109.7 23
Wintotlttd alesse ncrcee AI 108.2 106.6 III.0 163
Motacillidae’ <s5..¢5 0. 2 108.5 107.9 109.1 | 6
Ginchidaes sansa... I 107.3 Pee te ‘lec: I
WN Uitsani cl ave veetae ese sietcrs as 5 108.9 108.0 100.7 15
Troglodytidae ....,... 7 107.6 105.7 108.9 30
Genthiaae: wines eet I 107.5 107.4 107.7 5
Sithicaemeresmheir reas =. 4 107.5 106.2 107.9 17
TEAK EC ENS” Brio AO cco ORGS 10 107.9 105.3 109.4 44
Whatnaerdacwecdccns = I 108.0 108.0 108.1 2
SyilvdiGadelee pron. ces, 4 107.3 106.0 107.8 18
PT OIG AS othe aeons 10 108.9 107.6 110.4 33

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

TABLE 5.—Temperatures of species of birds not included in Table 3, taken
from available Iiterature

Temperature © -
oT
Species Ree Ne F S Reference
& Mean cae ara 5 é
STRUTHIONIDAE |
Struthio camelus ..... M.|104.0).....51....-|)% |Bergtold) 1917, p52
ee 5 ....| F.|101-0/100.0,102.0| 2 |Bergtold, 1917, p. 52.
DROMAEIDAE | |
Dromaeus novae- heat TOD a2 ts moe eee ? |Sutherland, 1899, p. 780.
hollandiae. | |
CASUARIIDAE |
Casuarius intensus ...| ? |tor.8).....|.....| ? |Sutherland, 1890, p. 78o.
Casuarius beccarti ...|) |102.5)..... eee ? Sutherland, 1899, p. 780.
APTERYGIDAE | | |
Apteryx mantelli ....|M.100.9 99.3 100.7) 2 |Sutherland, 1809, p. 780.
Apteryx haasti ...... M.|100.s|.....]...-.| I |Sutherland, 1890, p. 780.
TINAMIDAE
Rhynchotus rufescens. eT ORM ele tele. peters ? Sutherland, 1899, p. 780.
Nothura maculosa ... ? 104.9,102.5 108.3 3 Sutherland, 1899, p. 789.
CRACIDAE | |
Crax globicera ....... M.|106.4|.-...|-----| 3 |Bergtold, 1917, p. 53.
a ie MY ee ek are By LOOcaltroe areca 4 |Bergtold, 1917, p. 53.
PHASIANIDAE |
Francolinus natalensts.| ? |107.9|.....|---+-| ? |Bergtold, 1917, p. 53.
Phasianus torquatus..|M.107.5).....|...--| 2 |Bergtold, 1917, p. 53.
: ys «| F. 1106.0! allie 1 |Bergtold, 1917, p. 53.
Gallus gallus ..2.2..; M. 106.5 105.6 107.552 Simpson and ' Galbraith,
| | 1905, D. 237.
i sh fe Sa F. 106.7,105.5 107.452 |Simpson and Galbraith,
| | 1905, P p. 237.
; “ (Bantam). M. 106.4 105.2 107.352 Simpson and Galbraith,
| | “SEOOS; paze7.
ee: FS G3 F. 106.9 105.6 107.852 Simpson and Galbraith,
| 1905, Pp. 237.
Pavo cristatus ......- ? |107.1/104.9 109.4) 2 | Milne- eta. H., 1863,
|e
Numida meleagris ...| ? |110.0|.....|.---.| ? |Pembery, 1898, p. 791.
Meleagris gallopavo.. }? TOOK Ole welll > |Pembery, 18098, p. 791.
CoLUMBIDAE |
Columba livia fe 105.6 103.9 105.651 Simpson ee Galbraith,
(domestic). ee | | [° 4) HgOs pa 287.
Columba livia | F. |106.4/105.1/107.5|52 ‘Simpson. re Galbraith,
(domestic). | | | | 1905, Pp. 237.
Columba phaeonota...| ? |110.0|.....|-----| I pe IQI7, p. 55.
Geotrygon montana...| ? \110.0|.....|..-..| 1 |Bergtold, 1917, p. 55.
RALLIDAE |
Rallus crepitans*..... P. 1TO4:L2| nese @ ac ~ lesa] WWEDEE, SOTO: apa ame
PMC: GHG: dicen! ued PES ROALO| as Swale are 1 Milne-Edwards, H., 1863,
SPHENISCIDAE | pe 18:

Eudyptula minor ....| ? |102.1.100.0 104 2 ? |White, 1916, p. 56.
PROCELLARIIDAE |
Procellaria pelagica...| ? |103.6).....|.....| 1 |Simpson, 1912a, p. 31.
PUFFINIDAE |
Puffinus tenuirostris*. ? |T01.6|/100. 0|103.2
.lyg.{100.2| 99. 4/I01.0
Fulmarus glacialis.... ? |101.8|100.5/103.3|
Daption capensis ....| ? ‘103. 6 102. 8 104.3

White, 1916, p. 46.

White, 1916, p. 46.

Martins, 1858, p. 32.
Brown-Sequard, 1858, p. 44.

& ot sw-u

1A, O. U. Checklist, 1910.

NO. 12 BODY TEMPERATURE OF BIRDS—-WETMORE 49

Wee 5.—Continued

Somateria mollissima.

Temperature ls =|
; oy
Species Pilea c S| Reference
3 Mean mum) mum 2 +
DIOMEDEIDAE | |
Diomedea exulans ...| ? |105.2)103.2107.4. 7 Brown-Sequard, 1858, p. 43.
Thalassogeron chloro-| ? \105.81c5.2 106.3 2 Brown-Sequard, 1858, p. 43.
rhynchus | lagi
ALCIDAE | ers
WA etOFE 2.23. . <a ss IMETOR Ble alas. ate _1 Simpson, 1912a, p. 32.
‘i ES be a ae 1 ELA htt ea Pog ea) Ll Simpsons, Lola) (p.: 32:
Ur irodle sn... - M.|104.3 102.3 106.1 2 Simpson, 1912a, pp. 27-32.
Meera ee F, |104 5,103.6 105.4 3 Simpson, 1912a, pp. 27-32.
Uria lomvia ......... ? |104.9|103.5 106.7; 8 |Martins, 1858, p. 32.
Cepphus grylle ...... M. 105. 1/104, 1105.8) 6 Simpson, 1912a, pp. 27-32.
CET CRE ak oa F. |105.6|104.3 106.7) 7 |Simpson, 1912a, pp. 27-32.
Fratercula arctica ....| ? |105.3\105.2105.5{ 2 |Martins, 1858, p. 32.
LARIDAE tis
Larus ridibundus .. FRO S| cick aisle see 5) Martins, Taso, pf. 32)
Larus argentatus ....| ? |108.2 106.9 109.4/10 Martins, 1858, p. 32.
EOGUS TUSCUS! 5c... . < ? |107.0/106 3/107.7, 3 |Simpson, 1912a, p. 31.
TATUS CONUS 2 ek oss s ? |107.1105.9,107.7| 8 |Simpson, 1912a, p. 31.
Larus glaucus ....... ? /105.3|/103.6/107.0/12 |Martins, 1858, p. 33.
Pagophila eburnea ...| ? |104.8'103.9 106.2) 3 Martins, 1858, pp. 32-33.
Rissa tridactyla ...... e 106.6 103.8 108.3 16 Simpson, 1912a, p. 31.
Megalestris catar- ? |104.2/103.2 105.4, © Brown-Sequard, 1858, p. 44.
rhactes.
STERCORARIIDAE | |
Stercorarius para- TOO. Bye k. s|2 wae =| I White, 1916, p. 46.
siticus. | | |
CHIONIDIDAE | |
Chionis minor ....... ? |104.0 | ..| 2 Eydoux and Souleyet, 1838,
-EURYPYGIDAE | p. 458.
Eurypyga helias ..... PM IEOZ Alin 2 Si aaec I Bergtold, 1917, p. 54.
ANATIDAE |
Cygnus olor .........|M.105.9/105.7/106.3|) 3 Martins, 1858, p. 33.
is The, Wa ahase res oe F, |105.9/105.6 106.3) 4 |Martins, 1858, p. 33.
Cairina moschata ....| ? |107.7\105.5|108.2|16 |Martins, 1858, p. 37.
AER, SPOWSG. ssa clue « PR VEOT Ol cag beac En SIMPSON) «lotsa, DP. 31
Anser (domestic ? |106.4/104.4 107.696 Martins, 1858, pp. 34-36.
goose).
Anser albifrons ......| ? |109.1|..... | apa: I Martins, 1858, p. 36.
Cygnops's cygnoides..| ? |109.1|108.3 109.9] 4 |Martins, 1858, p. 33.
Branta bernicla ...... CelTCOVOlr. oe |. .<.ce| F-lMartins, «1888, “p, 36:
Tadorna tadorna . ? |108.8'108.3'109.2| 3 |Martins, 1858, p. 36. :
Anas platyrhynchos* |M.\106.4'105.6 107.2.42 |\Simpson ge Galbraith,
(domestic). | 1905, p. 23
Anas platyrhynchos* | F.|107.0 105.9 107.741 |Simpson et "Galbraith,
(domestic). | TOOS De 2a0e
Anas rubripes? ...... ? |105.8105.2106.7, 3 |Simpson, 1912a, p. 31.
Mareca penelope ..... ? |108.5'106.6 109.5/18 |Martins, 1858, p. 40.
Marila marila* ...... ? |108.8 107.9 1109.9 7 |Martins, 1858, p. 36.
Clangula clangula ? 104.7 Ber Pare oee| 1 |Simpson, IgI2a, p. 31.
Oidemia nigra ....... E 106.3/105.6,107.0 Simpson, I912a, p. 31.

108. 4/104.1 109.8 9 Martins, 1858, p. 306.

1A. O. U. Checklist, rgr1o0.

50

TABLE 5.—Continued

SMITHSONIAN MISCELLANEOUS

COLLECTIONS VOL. 72

1A, Gs Cheatin: Fons

Temperature © oh
alse)
Species B S Reference
Bataan Mini- Maxi- 5 9
oF | mum | mum iy, ta
| |
PHALACROCORACIDAE | |
Phalacrocorax carbo. ? \103.6102.0104.5)12 Simpson, 1912a, p. 31.
Phalacrocorax M.|104.7|/102.9 105.6] 7 |Simpson, I9I2a, pp. 27-32.
graculus. | | | |
Phalacrocorax | F’. }104.8/103.2)106.1|/12 |Simpson, 1912a, pp. 27-32. -
graculus. | | | |
SULIDAE | |
Sula bassane 7.7. 2.4. ? |107.0/104.5/108.1| 7 |Simpson, 1912a, p. 31.
FALCONIDAE | |
Astur palumbarius ... ? \107.2 106.8 107.4, ? Hildén and Stenbick, 1916,
. | | pp. 382-413.
Gypaétus barbatus....| ? \105.8).....|..... ? |Milne-Edwards, H., 1863,
| |_ p. 17.
Archibuteo lagopus ..|M.|105.8|.....)..... 1 Bergtold, 1917, p. 53.
Falco mexicanus ..... M.|106.6. | 1 |Bergtold, 1917, p. 53.
BUBONIDAE |
Strix varia* ......... ? /102.6/102.6 102.6) ? |Weber, 1918, p. 31.
MomotTipAE | |
Momotus paraensis ? |104.1) 1 |Bergtold, 1917, p. 55.
MIcROPODIDAE * | |
Micropus apus ....... Dyes 3 ? Pembery, 1808, p. 7o1.
MUSOPHAGIDAE
Turacus corythaix P |104.2).....).. | t |Bergtold, 1917, p. 55.
CUCULIDAE | |
Geococcyx cali- M.|107.4). | 1 \Bergtold, 1917, p. 55.
fornianus.
MIMIDAE ‘ | |
Toxostoma rufum ? |100).O% 3.23. lhacetttesty t Weber, 1918, p. 20.
TURDIDAE |
. DLurdus pilarts 52... Pi GEOVO|S 2220 e We ? |Pembery, 1808, p. 791.
Hylocichla iliaca ..... Pa BOQE ONS ee iesllccsterece ? |Pembery, 1808, p. 701.
Hylocichla musica ....| ? [105.6 101.2,108.7. 29 Simpson and Galbraith,
| 1905, p. 237.
ce «“ .. 2? 106.0 101.5,108.3 21 Simpson and Galbraith,
BOMBYCILLIDAE * 1905, p. 237.
Bombycilla garrula*..|M.|108.0|.....|..... 1 |Bergtold, 1917, p. 56.
bd 4) || F.| 107) 1|106.2|107.8) 3:|Bergtold,; ‘1917, ps:
PARIDAE |
Parus major ........- P |III.2! | ? |Pembery, 1808, p. 7901.
MNI0TILTIDAE |
Oporornis agilis ..... LOO" O tanec have kare 1 Weber, 1918, p. 20.
FRINGILLIDAE (eet
Ligurinus chloris ....| ? 106.8 106.1/107.9 ? Hildén and Stenback, 1916,
liane | pp. 382-413.
Pyrrhula pyrrhula ...| ? 107.9 | 1 |Milne-Edwards, H., 1863,
Lope] p. 17.
Emberiza citrinella ..| ? |109.8|.....|.....| ? |Pembery, 1808, p. 791.
Plectrophenax nivalis.| ? [709 .6 109.2|110.1) 2 |Milne-Edwards, H., 1863,
| Oe ee ce
Calcarius ornatus ....|M.|109.6].....|....-| t |Bergtold, 1917, p. 57.
Passerherbulus POO VAG ac 1 |Weber, 1918. p. 30.
caudacutus’*. : ;
Peucaea cassini* ..... WWE UGtS} OWE natool laiciorrs 1 |Bergtold, 1917, p. 57.

NO. 12 BODY TEMPERATURE OF BIRDS—WETMORE 51

TABLE 5.—Continued

Temperature ‘°C =
= _|53
Species he a eae a8 Reference
E [Mean | mum | mum [3 ®
| ig *
STURNIDAE | |
Sturnus vulgaris ..... ? |106.7 101.9 109.117 |Simpson and Galbraith,
A F 1905, Pp. 237. :
ate ? |107.8/104.3 110.255 |Simpson and Galbraith,
; | | 1905, p. 237.
3 Le A ? |106.5 103.5 108.755 |Simpson and Galbraith,
| 1905, P- 237. :
ete Oe rn cts _? |106.6 103.3 109.256 |Simpson and Galbraith,
| | 1905, Pp. 237.
PARADISEIDAE
Paradisea apoda ..... PPR IMOO Win gmlacss i) L Meerstold, 1927, 1.; 55.
CoRVIDAE | |
Coloeus monedula ...|M.\107.0)105.2 108.456 |Simpson and Galbraith,
; | | 1905, Dp. 237.
Rr re ...| F.|107.7|106.2 108.857 |Simpson and Galbraith,
se si ; | 1905, Pp. 237.
Psilorhinus morio ...\ ? \|1I0.0|.....|..... | 1 |Bergtold, 1917, p. 57.
Pyrrhocorax alpinus.. ? |107.7|. ....|-----| I |Milne-Edwards, H., 1863,
| | | D. 17.
BIBLIOGRAPHY

AMERICAN ORNITHOLOGISTS’ Union. Check-List of North American Birds.
Third edition (revised), New York, 1910, pp. 1-430, 2 maps.

Bayiiss, W. M. Principles of General Physiology, London, 1915, pp. 455-459.

Bepparp, F. E. The Structure and Classification of Birds. London, 1898,
PP. i-xx, 1-548, 252 text figs.

Berctotp, W. H. A Study of the Incubation Periods of Birds. Denver, 1917,
pp. I-I09.

British OrNITHOLOGISTs’ Union. A List of British Birds. London, 1915,
pp. i-xxii, I-430.

Brown-Sequarp, E. Note sur la basse Température de Quelques Palmipédes
Longipennes. Journ. de Phys., 1858, pp. 42-46.

Cameron, A. J., and Browntez, T. I. The Upper Limit of Temperature Com-
patible with Life in the Frog. Trans. Roy. Soc. Canada, Ser. III, Vol. IX,
1915, Sec. IV, pp. 67-84.

Depretz, M. C. Recherches expérimentales sur les causes de la chaleur
animale. Ann. de Chimie et de Physique, Vol. 26, 1824, pp. 337-364.

Heaptey, F. W. The Structure and Life of Birds. London, 1895, pp. i-xx,
1-412, 77 text figs.

Hivpén, A., and Stenpdcx, K. S. Zur Kenntniss der Tagesschwankungen
der Korpertemperatur bei den Vogeln. Skandinavisches Arch. fiir Phys.,
Bd. 34, 1916, pp. 382-413.

Kinc. Temperatures of Quadrupeds, Birds, Fishes, Plants, Trees, and Earth
as ascertained at different times and places in Arctic America during
Captain Back’s Expedition. Edinburgh Phil. Journ., New Series, Vol.
XXI, 1836, pp. 150-151.

Knowtton, F. H. Birds of the World. New York, 1909, p. 3.

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Litiiz, F. R. The Development of the Chick. New York, 1908, pp. 326, 330
and 331.

Martins, C. Mémoire sur la Témperature des Oiseaux Palmipédes du Nord
de l’Europe. Journ. de Phys., 1858, pp. 10-41.

Mitts, W. A Short Chapter in Comparative Physiology and Psychology.
Trans. Roy. Soc. Canada, Series II, Vol. XII, Pt. I, Sec. IV, pp. 291-300.

A Text-book of Animal Physiology. New York, 1889, pp. 1-700.

Mitne-Epwarps, H. Lecons sur la Physiologie et l’Anatomie Comparée de
Homme et des Animaux. Paris, 1863, Vol. VIII, pp. 1-02. i

MircHeLt, P. C. The Childhood of Animals. New York (1912), p. 184.

Morcan, A. M. Further Observations on the Cormorants and Bird Tempera-
tures. South Australian Orn., Vol. II, July 1, 1916, pp. 178-183.

Notes on the Food and Temperatures of Cormorants. South
Australian Orn., Vol. III, July 1, 1917, pp. 75-78.

Muuter, B. The Air-sacs of the Pigeon. Smiths. Misc. Coll. (Quart. Iss.),
Vol. 50, No. 1724, 1908, pp. 365-414, 12 text figs.

Newton, A. A Dictionary of Birds. London, 1896, pp. 3-6.

Pempery, M. S. Animal Heat. In Text-book of Physiology, edited by E. A.
Schafer, London, Vol. I, 1898, pp. 785-867.

Pycrart, W. P. A History of Birds. London, 1910, pp. i-xxx, 1-458, 37 plates,
50 text figs.

Suarpe, R. B. A Hand-List of the Genera and Species of Birds. London,
Vols. I-V, 1899-1900.

Smpson, S. Observations on the Body Temperature of Some Diving and
Swimming Birds. Proc. Roy. Soc. Edinburgh, Vol. XXXII, 1912, pp.
19-35.

An Investigation into the Effects of Seasonal Changes in Body Tem-
perature. Proc. Roy. Soc. Edinburgh, Vol. XXXII, 1912, pp. 110-135.
Stmpson, S., and GALpraitTH, J. J. An Investigation into the Diurnal Varia-
tion of the Body Temperature of Nocturnal and Other Birds, and a few

Mammals. Journ. of Phys., Vol. X XXIII, 1905, pp. 225-238.

SmitH, R. M. The Physiology of the Domestic Animals. London, 1889, pp.
693-608, 416 text figs.

Soum, J.-M. Deuxieme Note sur les sacs Aériens des Oiseaux. Soc. Linn. de
Lyon, Vol. XLII, 1805, pp. 149-161.

SuTHERLAND, A. The Temperature of Reptiles, Monotremes and Marsupials.
Proc. Roy. Soc. Victoria, N. S., Vol. 9, 1897, pp. 57-67, 1 plate.

On the Temperature of the Ratite Birds. Proc. Zool. Soc. London,
1899, pp. 787-790.

Waroiaw, H. S. H. Note on the Temperature of Echidna aculeata. Proc.
Linn. Soc. New South Wales, Vol. XLIII, Pt. 4 (for 1918), March 26,
1919, pp. 844-840, 2 text figs.

Weeser, J. A. Bird Temperatures. Abstract of Proceedings, Linn. Soc. of
New York, No. 30, 1918, pp. 28-31. .

WiepersHEIM, R. Comparative Anatomy of Vertebrates. Third (English)
Edition, 1907, pp. 33-35.

Wuirte, S. A. An Ornithological Cruise among the Islands of St. Vincent and
Spencer Gulfs, S. A. Emu, Vol. XVI, 10916, pp. 1-15.

The Cruise of the Avocet in Search of Skuas and other Things.
Reprinted from The Register, Adelaide (1916), pp. 46, 56.

Wortz, A. Production de la chaleur dans les étres Organisés. Paris, 1848,
pp. 1-38.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 72, NUMBER 13

THE MELIKERON—AN APPROXIMATELY
BLACK-BODY PYKANOMETER

BY
L. B. ALDRICH

(PUBLICATION 2662)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 25, 1922

ay

-

The Lord Bat more Press

BALTIMORE, MD., U. SAL

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THE MELIKERON—AN APPROXIMATELY BLACK-BODY
PYRANOMETER

By L. B. ALDRICH

INTRODUCTION

The instrument about to be described is the outgrowth of the
experience of Dr. Abbot and myself in the use of the pyranometer,
and was planned in many discussions between us, as we walked
together to and from the office. The pyranometer (described in
Smithsonian Miscellaneous Collections, Vol. 66, Nos. 7 and 11,
and Vol. 69, No. 9) has proved of great value in a wide range of
radiation measurements. We have long felt the desirability and need,
however, of a radiation measuring instrument of equal sensitiveness
which would be perfectly absorbing and radiating for all wave-
lengths by virtue of its form. Existing types of instruments, such
as the pyranometer, bolometer, Angstrom’s pyrgeometer, compensa-
tion pyrheliometer, etc., all use a blackened flat surface upon which
the radiation falls and is mostly absorbed. -For the usual range
of wave-lengths, for which the percentage absorption of the blackened
surface is well known, these instruments are highly satisfactory.
But in measuring radiations from bodies at comparatively low tem-
peratures, grave doubt arises [with these instruments] because of the
uncertainty of the absorptive power of a blackened flat surface for
rays of long wave-length.

In the new instrument we have tried to produce one embodying an
approximately “ black-body ” absorber, and still to retain as far as
possible the advantages of the simple pyranometer. The melikeron
is not as sensitive nor as quick-acting as the pyranometer, yet we
have been very well pleased with its behavior. A detailed description
of the instrument follows.

DESCRIPTION OF THE MELIKERON

The name of the instrument, first suggested by Dr. Abbot, is the
Greek word peduxnpoy, honeycomb. That portion of the instrument
which absorbs the radiation to be measured is somewhat like a honey-
comb in shape.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 72, No. 13

- 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Sheet’ therlo,” an alloy having a low temperature coefficient of
resistance, was rolled out into a strip about a meter long and as thin
as possible. (In the first instrument made, this strip was about
0.05 mm. in thickness. This was about the limit of thinness obtain-
able by rolling between cold rollers. For the second instrument, a
strip of one-half this thickness was produced by rolling between hot
rollers. This was done by the mechanician of the University of
Wisconsin Physics Department, through the kindness of Dr. C. E.
Mendenhall.) With a straight edge, the strip was cut to one-half
inch in width, and then pressed out in a specially prepared die, to
assume the alternately flat and zigzag shape shown in figure 3. When
this long strip was held together in a square frame, there were formed
200 small triangular tubes with walls in common, each tube one-half
inch in depth and about 2.5 mm. on a side. The open end of this
honeycomb of triangular tubes forms the absorbing area of the
instrument.

The advantage gained by the large number of cells is that the outer
ones protect the inner ones from loss of heat, so that notwithstanding
the very large area of the walls of the cells compared to their open
ends, the central cells, losing only at front and rear, change tempera-
ture about as much as flat strips presenting equal areas would do for
the same intensity of radiation. We invoke, in other words, the
guard-ring principle.

Before the long, crinkled strip was pressed into this square shape,
each apex was coated with thin shellac, the whole baked in an oven for
some hours, and this process repeated several times. Thus the whole
strip, when formed into its final shape, was insulated, each part from
every other that could come in contact with it, and a current of elec-
tricity could be sent through its whole length.

On the walls of the central cluster of tubes formed by the bending
of the strip were fastened four thermo-electric elements, of fine copper
and nickel wire. The junctions were symmetrically placed 2.5, 5.0,
7.5 and 10.0 mm. respectively along the length of the tube and insu-
lated from it by thin tissue paper. These wires were brought out on
the lower end of the tubes and connected in series. The constant
temperature junctions were buried in wax on the under side of the
glass plates f, f (fig. 2) and the outer leads were soldered to the bind-
ing posts a, a, (fig. 1). The two ends of the therlo strip were con-
nected by copper wires to binding posts a’, a’ (fig. 1).

* Obtained from the Driver-Harris Wire Co.

NO. 13 THE MELIKERON—ALDRICH 3

Referring to figures 1 and 2, we may see how the honeycomb is
mounted. Figure 1 is a view looking vertically down upon the instru-
ment with shutter (¢) wide open. Figure 2 is a vertical cross-section.
Two nickeled copper plates (i, 1), each with square holes 3 cm. on
a side are placed one about 6 mm. above the other. The plates are held

@

2 ----2] OO
He, yy
TULUM BLE
WV AP
nuanaaaaaay
ie

Fie. 3.

together by the three posts (b) and the space between the plates
around the square hole enclosed with’a copper box (m) attached to
the upper plate. The wires leading to the four binding posts (a, a,
a’, a’) pass through holes in this copper box. Four nickeled copper
clips (c) are screwed to the top of the upper plate, and four more
to the bottom of the lower plate. Each of these eight clips holds in

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

place a silvered glass plate (f), each of which glass plates is beveled
along its inner edge. These beveled glass edges serve to support the
therlo strip in its square form, four around the upper edge of the
honeycomb and four around the lower edge. The upper four also
serve to determine the area which absorbs radiation. The silvered
glass mirror (g), below the honeycomb, is placed, as shown, at a
small angle to the face of the honeycomb and serves both to protect
from the wind and to reflect radiation escaping from the lower face
of the honeycomb back upon the sides of the tubes. The rod (k)
screws into the plate (1) and affords a means of mounting the instru-
ment in any desired position. The hemispherical shutter, (¢), nickeled
on the outside and blackened inside, operates from the handle (7),
just as in the pyranometer. The optically figured ultra-violet crown
glass hemisphere (d) serves the same purpose as in the pyranometer
and may be used or not, according to whether or not it is desired to’
cut off the exchange of long waves between the instrument and the
object to which it is exposed.

The melikeron is similar to the pyranometer in principle. In place
of a small flat absorbing surface we substitute a large absorbing area
consisting of the above described honeycomb of triangular tubes.
Radiation falling normally passes through and is reflected back upon
the walls by the rear mirror. Radiation not falling normally strikes
the walls of the tubes and after one or more reflections is absorbed.
For the purpose of somewhat increasing the blackness of the honey-
comb, only the lower two-thirds of each tube is painted with lamp-
black, the upper one-third remaining a metallic reflector. Thus the
number of regular reflections before final absorption is increased and
the loss by diffuse reflection near the upper end reduced, because the
diffusely reflecting and radiating lampblack lies so far below the
aperture that the latter subtends only a small angular area as viewed
from the blackened surface.

METHOD OF USE

For nocturnal radiation, or for the measurement of radiation
exchange between the instrument and an object at lower temperature,
the melikeron is used like the pyranometer. That is, an electric
current is passed into the therlo strip producing heat sufficient to
exactly compensate for the loss of heat by radiation. Knowing the
current used, the resistance of the strip, and the other constants of the
instrument, the amount of heat radiated is computed as with the
pyranometer.

NO. 13 THE MELIKERON—ALDRICH 5

For measurements on the sun, daylight sky, or any radiation from
bodies at higher temperature than the instrument, the simple “ first-
swing” pyranometer method is not applicable, since the slow-acting
melikeron prevents a definite first-swing of the galvanometer and
requires several minutes to complete the galvanometer deflection. An
almost equally simple and satisfactory method applicable to constant
sources of radiation was suggested by Dr. Abbot, however, namely:
To open the shutter and expose to the radiation to be measured until
the galvanometer deflection is constant, then close the shutter and
instantly introduce sufficient current to keep the galvanometer at the
same reading.

CONSTANTS OF INSTRUMENTS AND TESTS MADE

As mentioned above, two copies of the melikeron have been made.
The second instrument embodies several improvements, notably a
thinner therlo strip, and the tipping of the rear mirror at a slight
angle to the honeycomb face. The constant of each instrument and
tests made with each are given below.

Melikeron No. 1—The constant of the instrument may be apeined
in two ways:

(1) By computation from the dimensions and properties of the
instrument ;

(2) By direct comparisons on the sun with a silver disk pyrhelio-
meter or other standardized instrument.

Only the first of these methods was used for the constant of Meli-
keron No. 1. As compared with the second method, this method is
difficult and inaccurate, because of the uncertainty of such corrections
as the amount reflected from the end surfaces of the thin metal com-
posing the tubes, the amount lost by reflection and radiation from the
upper portion of the tubes, the decrease in total aperture due to the
unavoidable indentations around the edge, etc. A rough determination
was made of the computed constant of Melikeron No. 1, as follows:

Area of aperture formed by beveled glass edges = 5.83 cm.”
Estimated decrease in area of unused portions= .30 “

6c

Corrected area = 5.53

Resistance of therlo strip = 0.945 ohms.

Then 945 X 60 = 2.45= constant Melikeron No. 1, appli-

4.183 X 5.53 cable to reduce C* readings to calories per (ar }.

This is the constant without the glass hemisphere. With glass hemi-
sphere, allowing for the reflection loss at two glass surfaces, the con-

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

stant becomes 2.66 for the rays of short wave-length for which glass

is highly transmissible.
Test experiments of three kinds were made with Melikeron No. 1:
(1) Using an incandescent lamp source, comparisons were made

by interchanging Melikeron No. 1 and Pyranometer A. P. O. No. 5.

TABLE I
; P Calories Ratio
Calories .
Date, 1920 Conditions (by melik- (Oy Bree meWkeron:
eron No. 1) eter | pyranom-
No. 5) eter
Feb. 11..|Carbon lamp, 60 cm directly above..| .0538 .0648 . 830
Feb. 12..|Same, except angle 25° from zenith.| .0522 .0537 .972
Feb. 13..;/Mazda “Daylight” lamp, 30 cm.| .0558 .0562 -995
above, 13° from zenith.
Feb. 13../Same source, 8 cm. aboveand 25cm.| .0245 .0237 1.033
east, 72° from zenith.
Feb. 14. .)Same. oye 30cm. above, 74° from) .0502 .0517 -972
zenith.
Feb. 14..|Same source, 30 cm. above, 43° from| .0435 .O512 .850
zenith.

Both instruments were leveled, and the source placed at varying angles
and distances. To insure constancy of the source, storage batteries
were used. ‘The results are summarized in table I.

As would be expected, since in Melikeron No. 1 the rear mirror is
perpendicular to the honeycomb tubes, the instrument does not mea-
sure the full amount of radiation falling normally or nearly so, for
this passes through the tubes to the rear mirror and is reflected by
it directly back without being absorbed. For incidence greater than

Taste II
(February 18, 1920)

(Current)? by | (Current)? by Calori : Constant of

meykeron | pyrgcometer'| “elikeron | P¥¥geometer
Withee Splatter snmtae -01334 .00304 | .0327 10.85
No 7. ef hatches .01756 .00350 || .0430 12.28
Wath | atid tent sida . 01284 . 00286 .0314 10.97
With ‘“ INA are yea .01246 .00282 -0305 10.81
No Ls Pry Sateen RS 01483 .00313 . 0364 11.61
No as Bea. Peta as .01573 .00314 .0385 12.26
With es Oe Pack . 01096 .00259 .0269 10.40
fe) ‘s fe ats Ab mcrae - 01300 .00256 .0321 12.55

Mean constant No. 22 with R. S. =1
Mean constant No. 22 no R.S. =12.18.

NO. 13 THE MELIKERON—ALDRICH Vf

5 degrees from normal, however, good agreement is shown between
the pyranometer and the melikeron. 4
(2) Comparisons were made between Melikeron No. 1 and Ang-
strom Pyrgeometer No. 22, with and without the interposition of a
rock salt plate. A flat copper vessel (fig. 4) 90 x 86 cm., blackened
on the front surface by painting with lampblack-alcohol-shellac paint
and filled with ice-cooled water, formed the source, the instruments
being at room temperature. A double shutter (s), sliding horizontally
close to the copper vessel, exposed or screened the source. The instru-
ments could be quickly exchanged, each mounted with absorbing

Fic. 4.

Fic. 5.

surface vertical, facing the copper vessel, and 65 cm. from it. Alter-
nate comparisons were made with and without a 1 cm. rock salt plate
(r), figure 4, interposed directly in front of the instrument aperture.
Using the above computed constant of the melikeron, values of the
constant of Pyrgeometer No. 22 were determined (table II).

The absolute value of these results is of little weight, but the
markedly lower value of the constant of Pyrgeometer No. 22 for the
case where waves longer than 20 are excluded, seems to indicate the
greater “blackness”? of the melikeron as compared with the other
instrument for rays of very great wave-length.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.. 72

(3) A rough determination of the constant o of Stefan’s formula
was made. A wooden case and water jacket were fitted around Meli-
keron No. 1 to protect it from temperature fluctuations of the sur-
roundings. This jacket extended over the face of the instrument,
leaving an aperture 3.64 cm. in diameter at 7.03 cm. from the honey-
comb face. Two hollow-chamber black bodies were made of double
walled galvanized iron vessels (a and 0, fig. 5) filled between the walls
with stirred water, one at room temperature, the other containing a
mixture of ice and water. The melikeron and surrounding jacket
just filled the aperture of either of these black bodies, and could be
quickly moved from one to the other aperture. The results are sum-
marized. Details and necessary corrections which have been intro-
duced in these results here will be found in the forthcoming Vol. IV,
Annals of the Astrophysical Observatory.

Temperature of 2
== (Current)? Orem eal o (calories per cm.? per min.)
B. B. Dee) 1845 Be ce b”
19.63 0.44 .003807. -00054 8.45X10-— (Mean of 7)
28.30 0.40 -005852 .01434 8.53107? (Mean of 7)

Value usually accepted (Smith. Phys. Tables, 7, p. 247) 8.26107".

These values are not given as new determinations of sigma. They
have little weight for this purpose. They are given to show that not
only does the melikeron agree with the standardized pyranometer for '
short-wave radiation observations but it also agrees well with the best
work for long-wave rays.
Melikeron No. 2.—The constant of this instrument was determined
with more care, and by both methods above mentioned.
(1) Computed constant.
Area of aperture formed by beveled glass edges=2.42x
2 AR => 1e0 (CMl,-
Therlo strip=80 cm. long and .003 thick, making an end
cross-sectional area of 0.24 cm.?
Assume 40 per cent loss by reflection from this edge and
this correction becomes .096 cm.?
Area of incomplete triangles along edge of aperture=
0.36 cm.?
Radiation entering 4 this area is lost=o.18 cm.?

NOS 13 THE MELIKERON—ALDRICH 9

Assume direct absorption for solar rays’ of therlo strip=
70 per cent, then of the other 18 cm.? 30 per cent is
lost=.054 cm.? Hence, the total correction for in-
complete triangles is .054+.18=.234 cm.”

Area of irregular indentations that can lose energy by
direct radiation=.06 cm.?

Of this, 4 is lost=.03 cm.? (because radiation from one
side of perpendicular is lost, other side is absorbed).

Total losses of all kinds=.096+ .234+.03 =.36 cm?

Corrected area=5.86—.36=5.50 cm.?

Resistance of strip=1.555 ohms.

Computed constant (without glass hemisphere) =

1.555 X60
4.183 X 5.50

= 4.05

(2) Observed Constant.

The constant determined by comparison with pyrheliometers may
be given more weight. The melikeron was mounted equatorially and
a hood placed around it, similar to that supplied with the pyranometer,
exposing the instrument only to the sun and a small area of sky
around it. A double, ventilated shutter, blackened below, served to
cut off the radiation at intervals. The first comparison was made on
Mount Wilson, California, August 29 and 30, 1920, using secondary
pyrheliometer No. IV. All the following values are without glass
hemisphere.

First determination—Melikeron mounted so that sun rays fall nor-
mally on the instrument.

Calories by Pyr. No. IV (per ae

(Current)? of Melikeron—= .348 (mean of 3).

1.468

BABS

Second determination-—Melikeron mounted so that sun rays strike
the instrument at an angle of 8°.5 (cos.=.989).

's 1.468 (mean of 3).

4.22=constant Melikeron No. 2.

Calories by Pyr. No. IV=1.437 (mean of 3).
(Current)? of Melikeron= .360 (mean of 3).
1.437 X .089
360
From the ratio of these two results it appears that 6.8 per cent
of the normal beam is absorbed and scattered, probably largely by

= 3.95 = constant of Melikeron No. 2.

*¥For rays of great wave-length the absorption is much less, so that this part
of the loss would be increased. The difference cannot be serious, however,
because this correction is after all very small.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

the silvered surface of the rear mirror. Thus 3.95 is regarded as
more nearly correct for ordinary work with beams which (unlike
direct run-rays) subtend large angles.

The second comparison was made at Mount Harqua Hala, Arizona,
by Dr. Abbot, November 10, 1920, using secondary pyrheliometers
Sz Li Now32and ASPAOTNora:

First determination—Melikeron normal to sun’s rays.

Calories by Pyr. No. 32 and No. 9 =1.531 (mean value).
(Current)* of Melikeron—= .347 (mean value).

1.531

347

Second determination—Melikeron at 7° angle to sun’s rays (cos.
7° =.992).

Calories by Pyr. No. 32 and No. 9 = 1.538 (mean value).
(Current)? of Melikeron= .378

1.538 X .992
378

This result shows for normally incident rays a considerably greater
absorption and scattering by the rear mirror than was the case in
the comparison of August 30. From the deteriorated appearance of
the silvered mirror on November Io this was quite to be expected for
sun rays, but the deterioration was probably quite negligible for
earth rays. The best constant, then, of Melikeron No. 2 without glass
hemisphere is the mean of 3.95 and 4.04, or 4.00 which is now the
adopted value, applicable for all rays not at strictly normal incidence.

At Mount Wilson and Mount Harqua Hala numerous comparisons
on the night sky were made between Melikeron No. 2 and Pyrgeo-
meter No. 22. The two instruments, leveled, were mounted at the
same height and within less than 6 inches of each other. Exactly
similar bright tin-box shutters were used on both instruments. Using
the above adopted constant of Melikeron No. 2, a value of the con-
stant of Pyrgeometer No. 22 was obtained each time. The results are
summarized in table III.

The mean of all, under these varying conditions of air temperature
and water-vapor content is 9.72. There is perhaps some evidence in
these values that the constant of Pyrgeometer No. 22 is a function of
both air temperature and water-vapor content. But further compari-
sons under a wider range of air conditions are needed to confirm it.

To illustrate this indication, values are given in the table computed
by the formula:

Constant = 11.50—3.12p—1.47 (t—60°)*%.

= 4.41 = constant of Melikeron No. 2.

= 4.04 = constant of Melikeron No. 2.

NWO. 13s THE MELIKERON—ALDRICH 13!

They fit the observations much closer than the mean. This would
be expected. To increase either the humidity or the temperature is to
diminish the proportion of the extreme long-wave rays. The insertion
of the salt plate in the above reported experiments with Melikeron
No. 1 had a similar tendency. Hence, in view of the earlier observa-

Taste III

Date, oo | Wet and dry() | Saucons |pyrgeometer £0 pated | oc

Mt. Wilson Wet Dry mm. ete:
August 25.. eee iB aay 9.36 |—0.36| 9.59\—0.23
ee ees | eee eases
August 28 ...... he eed oe 9.78 |+0.06] 9.90—0. 12
INTISTISH One. es: | pe ae ao 10.33 |+0.61! 10.19+0.14
Mt. Harqua Hala
September 29... ae Fone ae 9.40 |—0.32) 9.23/-+0.17
September 30... a eee ae 9.70 |-0.02| 9.990.290
@ctober t..2..... ee ae a 9.36 |—0.36) 9.42—0.06
@ctober.2: .. 2.4. fee oe re 10.02 |+0.30) 10.06—0.04
Octoberi3.:..... | ae poe ce, 9.95 |+0.23) 9.65+0.30

tions, from the analogy, we should expect by increasing humidity or
temperature to reduce the observed pyrgeometer constant. The
observations are in harmony with this view.

It is hoped many further experiments with the melikeron may soon
be made with a view to a better knowledge of the behavior of long
wave-length radiation in our atmosphere and as emitted by bodies at
low temperatures.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 72, NUMBER 14

A NEW SAUROPOD DINOSAUR FROM
THE OJO ALAMO FORMATION
OF NEW MEXICO

(WitH Two PLATES)

BY
CHARLES W. GILMORE

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

(PUBLICATION 2663)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 31, 1922

The Lord Baltimore Press

aber gly BALTIMORE, MD., U. S. A. :

rc
+
,
7 a

a NEW SAUROPOD DINOSAUR FROM THE OJO
ALAMO FORMATION OF NEW MEXICO

By CHaries W. GILMorE,
ASSOCIATE CURATOR, DIVISION OF PALEONTOLOGY, UNITED STATES NATIONAL MUSEUM

(Witu Two Piates)

INTRODUCTION

In a brief note* I have recently announced the discovery by Mr.
John B. Reeside, Jr., geologist of the United States Geological Survey,
of Sauropodous dinosaur remains in the Upper Cretaceous of New
Mexico, and it is now proposed to give a more detailed account of the
specimens and of their exact geological occurrence than was possible
in the preliminary notice.

The remains so far recovered consist of a left scapula and a right
ischium, both in a good state of preservation. The great importance
of these particular bones lies in the fact that the remains of Sauro-
podous dinosaurs have not previously been known to occur above the
Lower Cretaceous in North America, so that the extension of their
geological range into the Upper Cretaceous is of the greatest paleon-
tologic and geologic interest.

Tt is particularly fortunate that this discovery should have been
made by a trained geologist of Mr. Reeside’s attainments, and in a
section so well established as to preclude the possibility of question as
to their late position in the geological column.

That these bones pertain to a member of the Sauropoda is indicated
by their immense size and also by their close general resemblance to
homologous elements of the typical Sauropoda from the Morrison
formation. Marked differences in details, however, more especially
in the ischium, in conjunction with the very late geological occurrence,
makes it necessary to establish a new genus and species for their
reception, for which the name Alamosaurus sanjuanensis is proposed.

DESCRIPTION
ALAMOSAURUS, new genus
The characters of this genus are included in the following descrip-
tion of the type species:

1Science (N. S.), vol. LIV, 1921, p. 274.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72, No. 14

iS)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

ALAMOSAURUS SANJUANENSIS, new species
Plates I, 2

Type.—Cat. No. 10,486, U. S. N. M., consists of the nearly com-
plete left scapula.

Paratype.—Cat. No. 10,487, U. S. N. M., consists of the nearly
complete right ischium.

Type locality—Barrel Spring Arroyo, one mile south of Ojo
Alamo, San Juan County, New Mexico.

Horizon.—Ojo Alamo formation, Upper Cretaceous.

Collector —J]. B. Reeside, Jr., June, 1921.

The scapula and ischium designated as the type and paratype,
respectively, were found in the same geological horizon, but some
200 feet distant from one another. While it is quite possible that
both may pertain to the same individual, it is thought best to regard
them as distinct until their closer relationship can be more positively
established.

Scapula.—The scapula is in a good state of preservation except for
the loss of a portion of the proximal or articular end (see pl. 1)
where it projected above the ground and was weathered away. The
suprascapular end is also incomplete, though apparently only the
border is missing. In size this bone rivals the largest of the Camara-
saurus scapulae described by Messrs. Osborn and Mook,’ since as
preserved its greatest length is 155 cm. (60 inches), and it is con-
servatively estimated that the total length of the complete bone would
have been at least 170 cm. (68 inches).

In outline, as shown in plate 1, the blade of the scapula differs from
any described form in that there is a gradual widening of the shaft -
from below upward to the superior end, there being no especial expan-
sion of the anterior border as found in Camarasaurus, nor rapid
superior expansion of both borders as found in Diplodocus and
Haplocanthosaurus. While this portion of the bone is heavy it is not
so massive as in Camarasaurus, being much thinner. The superior
end is flattened out, though the external surface becomes convex trans-
versely as the middle of the bone is approached. From end to end
the bone is curved as in other members of the Sauropoda. Both
anterior and posterior borders thin out to sharp edges, this condition
continuing downward half its total length. Immediately above the
point where the anterior border begins to turn upward to form the
prescapular expansion of the lower end the border becomes thick-

*Memoirs Amer. Mus. Nat. Hist., new ser., vol. 3, pt. 3, 1921, p. 341, fig. 74

NO. I4 A NEW SAUROPOD DINOSAUR—GILMORE 3

ened and rounded. The posterior border, however, continues down-
ward as a fairly sharp edge to the downward swing of this border to
form the glenoid socket where the bone rapidly thickens transversely.

The spine or ridge on the lower external surface extends from the
base of the shaft’ in an anterior direction to the anterior-superior
border, and at right angles to the longitudinal axis of the bone. This
ridge is not greatly elevated except that on the side toward the cora-
coidal border the bone is rapidly and deeply excavated, forming a
muscle fossa of great extent. On the upper side of this ridge the
surface of the bone slopes off gradually to the border, there being
no excavation or superior fossa such as is found in so many Sauropod
scapulae.

MEASUREMENTS Connie
Createstileneth of scapula, (as) preserved!) i2..2..).0...5-.- 155
# rhe ee M: Gestimatedi im. teidae cra rieeis rs Sete 170
Greatest breadth oh superior ends. secs cee cas «ciate ae en's cle erehs 45
Weastebreadthimo ie SHatb tes me Sirs ase aia te meus Sine qpesieete scents ere 20
Greatest breadth. inferior Coblique) o. . 0.6.00 2.c.e lee ten 82
pelitckiressrorsshatt atrCentensyee sls. crere, sets oo) ste aiete te levalesievetsre = 29

Ischium.—A large bone found in the same horizon but some 200
feet distant from the scapula described above is identified as the right
ischium of a Sauropod dinosaur. This bone differs so from other
Sauropod ischia that its true nature was determined with difficulty.
That it pertains to a member of the Sauropoda is indicated by its
large size and also by its general resemblance, though differing
markedly from any described form. It is characterized by its extreme
shortness, and especially by the lack of the long, slender posterior
extension so characteristic of other Sauropod ischia.

The proximal portion is nearly complete, lacking only a small por-
tion of the sharp inner edge of the acetabular border. The distal end,
though not perfect, apparently lacks but little of being complete.
Likewise the thin inner border below the articulation for the pubis
is slightly imperfect. Except for the missing portions mentioned,
the bone is in a remarkably fine state of preservation.

The expanded proximal end is unusual, not so much because of
its great antero-posterior extent, but on account of the great dorso-
ventral diameter, and especially the great length of the pubic articula-
tion which extends distalward more than one-half the total length of
the bone. Below the pubic articulation the inner border presents a
thin, sharp edge, and the flattened distal portion gradually diminishes
in width to the distal end. This end is apparently without distal

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

expansion, though the incomplete surfaces makes this point slightly
uncertain. Neither can it be definitely determined whether the ischia
met on the median line, though I am inclined to think they did. The
rounded and somewhat thickened posterior border is deeply concave
from end to end. The sweep downward from the iliac articulation is
especially pronounced. On the posterior external surface at the mid-
length of the bone is a raised ridge with roughened surface marking
the point of insertion for a strong muscle.

The form and principal features of this bone are well shown in

plate 2:

MEASUREMENTS Cettinenees

Greatest length aces pile eee See hoe eee Otel ee ae 81
Gireatestawic tho ta pro xt e tC rs yeyeeseery- ce eee ee teen 44.5
Greatest width at lower end of pubic articulation............ 2
Greatest lensthiion pubic artetlarisun faces srr ieee B75
Greatest transverse diameter of articular end for ilium...... 10.5

RELATIONSHIPS

The scapula cannot be closely correlated with any of those of
described genera, and the ischium differs so much in its details from
those with which it has been compared as to indicate an animal with
a considerably different pelvic structure than any of the Sauropoda
with which we are acquainted to-day. That both of these bones
pertain to the same individual cannot be proven, but that both are
Sauropod in aspect seems certain.

In size the scapula approaches Camarasaurus, but it differs by the
non-expansion of the upper anterior border and the very much thinner
blade; from Diplodocus it is to be distinguished by its larger size and
the direction of the spine in relation to the longitudinal axis. In the
present specimen this angle is approximately 90°, whereas in Diplodo-
cus and Amphicoelus it is acute. The scapulae of Apatosaurus,
Amphicoelus, and Brachiosaurus are more slender and with a much
more constricted shaft at their narrowest width. Haplocanthosaurus
is very much smaller and has a very different outline.

GEOLOGICAL OCCURRENCE

At my request Mr. Reeside prepared the following note on the
stratigraphy :

NOTE ON THE STRATIGRAPHY OF SAN JUAN COUNTY, NEW MEXICO,
WITH ESPECIAL REFERENCE TO THE OCCURRENCE OF DINOSAURS
The oldest rocks exposed in San Juan County, New Mexico, have been

assigned to the McElmo formation of Jurassic or Lower Cretaceous age. The
overlying rocks, of Upper Cretaceous and Tertiary age, have been divided into

NO. 14 A NEW SAUROPOD DINOSAUR—GILMORE 5

a number of units named in ascending order as follows: Dakota sandstone,
Mancos shale, Point Lookout sandstone, Menefee formation, Cliff House
standstone, Lewis shale, Pictured Cliffs sandstone, Fruitland formation, Kirt-
land shale with included Farmington sandstone member, Ojo Alamo sandstone,
Puerco formation, Torrejon formation, and Wasatch formation. The Point
Lookout sandstone, Menefee formation, and Cliff House sandstone comprise
the Mesaverde formation of the older literature and the Pictured Cliffs sand-
stone, Fruitland formation, and Kirtland shale, the Laramie formation. The
Dakota sandstone contains coal beds and other plant remains and grades into
the overlying Mancos shale. The formations from the Mancos shale to the
Pictured Cliffs sandstone, inclusive, are marine except parts of the Menefee
formation which are brackish and fresh water deposits with coal beds. The
lower part of the Fruitland formation contains a transition series of brackish
water beds and the upper part and all of the overlying formations are fluviatile
deposits. The Mancos shale represents in large part the Benton shale and
Niobrara formation of the region east of the Rocky Mountains. Its extreme
upper part, however, is the equivalent of the basal part of the Pierre shale.
The Mesaverde group, Lewis shale, and Pictured Cliffs sandstone contain
invertebrates of Montana age, and the Fruitland and Kirtland formations,
plants, invertebrates, and reptiles of Montana age. These beds definitely
assignable to the Upper Cretaceous, 7. ¢., from Dakota sandstone to Kirtland
shale, inclusive, are a comformable series 5,500 feet thick, of which about
4,000 feet are of Montana age. The age of the Ojo Alamo sandstone is in dis-
pute. It has been assigned by some writers on the basis of its dinosaur fauna
to the Montana group and correlated with the Judith River beds. It is
separated from the Kirtland shale by a widespread unconformity and has been
correlated on that ground by other writers with the Denver and Raton forma-
tions of post-Montana age. The Puerco and Torrejon formations contain
large mammalian faunas and are usually placed in the Tertiary, though some
writers would place them in the Cretaceous. The Wasatch formation is uni-
versally accepted as Tertiary.

Dinosaur remains have been found in the Fruitland formation, throughout
the Kirtland shale, and in the Ojo Alamo sandstone. The sauropod bones
found in June, 1921, came from the lower part of the Ojo Alamo sandstone
on Barrel Spring Arroyo, one mile south of Ojo Alamo. A detailed section
at this locality is as follows:*

Ojo Alamo sandstone: Feet
Sandstone weonotomenaticn, tops EGLOCEG suc. cs ctis.cese clei eacle oss <)s05 ae 15+
Shales dankveneenish) oray7. aq: o<.- clo oe 2 oe ore sie Se oe See A to 7
Sandstone, soft, nearly white, crossbedded; contains gray argilla-

CCOUS Stheaksr and abTO will. COMCEETIONSE. .hysc seus chas ome stasis» oo sle 21
Shale, wine red, with local gray sandstone lenses.................. 5
Sandstone, soft, white, crossbedded; contains brown concretions in

have: HONG ag DH glance ay Rie Rae eee ae ee Broa aera oS SR a ee a 10
SAM StOme MULO Wile. Plaky: Sf eLEUCIMOUS. sche cnt)iccietani seo es sie ete else
Shalewdanc<apliisheray, tO purple, Samdyeecscs cso: coves cc ccle cs ve 4

*See Bauer, C. M., Stratigraphy of a part of the Chaco River Valley. U. S.
Geol. Survey Prof. Paper 908, pl. 69 and pl. 70, 1916. This locality is shown
as locality 67 on plate 69 and the stratigraphic section as section R on plate 7o.

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Ojo Alamo sandstone :—Continued. Hes
Sandstone, soft white, conglomeratic; contains brown concretions;
horizon’ of the sauropod and other bomes.:.---)...+-..--.neees 6
Sandstone, yellow to brown, conglomeratic; contains an abundance
of siliceous pebbles as large as 3 inches in diameter............ 5
Unconformity.

Kirtland shale:
Shale, gray to drab, with several wine red layers; scattered dinosaur

ID OLLES, ants (sale cee ie to BO Rae Sue terereeue ellehe as RC ey ire ae eee 30
Sandstone with lenses of grit, fine conglomerate, and many clay

PEM SES 5 PP etree Nee ee otal ret bres occa ee ete gna 10
Shale: toraiy. ore aes fect nels ee oo re ae oie ae ae ea ne er 20+
Farmington sandstone member: brown indurated sandstone and

SHAY WMAlS 2 etl aaasi huis Pics se eesheks ee eueiel wiehotars} state ede ogc hey eee 80
Shale, gray to drab, and sandstone, soft, gray-white.............. 1000+

Fruitland formation:
Sandstone, shale, and coal.

Directly associated with the bones of Alamosaurus are many other
fragmentary and undeterminable dinosaur bones, teeth of carnivor-
ous and Ceratopsian dinosaurs, dermal plates of an armored form,’
turtle fragments, and crocodile bones. At nearly the same horizon
in adjacent localities on Barrel Spring Arroyo there were obtained
part of the frill of an undetermined Ceratopsian' different from
known forms, dermal plates of an armored dinosaur, incomplete
vertebrae of a carnivorous dinosaur as large as Tyrannosaurus,’
fragments of a Ceratopsian frill marked with radiating vasicular
grooves like those of Triceratops, but indeterminable.”. This horizon
is also the source of the maxillary and fragments of a skull collected
by Sinclair and: Granger and identified by Brown as Kritosaurus
navajovius.”

From the uppermost part of the Kirtland shale near this locality
have been collected specimens that are closely related to species known
to be of Montana age: Kritosaurus navajovius Brown, skull and

* Gilmore, C. W., Reptilian faunas of the Torrejon, Puerco, and underlying
Upper Cretaceous formations of San Juan County, New Mexico: U. S. Geol.
Survey Prof. Paper 119, p..65, 1910.

* Idem, p. 65, pl. 26, fig. 2.

*Tdem, p. 67.

“Gilmore, Vertebrate faunas of the Ojo Alamo, Kirtland, and Fruitland
formations: U.S. Geol. Survey Prof. Paper 98, p. 287, 1916.

* Sinclair, W. J., and Granger, Walter, Paleocene deposits of the San Juan
Basin, New Mexico: Am. Mus. Nat. Hist. Bull., vol. 33, p. 303, 1014.

NO. 14 A NEW SAUROPOD DINOSAUR—GILMORE H

other bones*; Monoclonius sp., horn core and fragments of frill’;
armored dinosaur suggesting a Belly River genus,’ humerus; a car-
nivorous form suggesting Dryptosaurus or Dynamosaurus, dentary *;
undeterminable fragments of other trachodont, ceratopsian, and car-
nivorous dinosaurs; turtle and crocodile bones.

In 1910 the late Dr. S. W. Williston® reported the discovery of
a Sauropod coracoid in the Trinity Sand of Oklahoma, Lower Creta-
ceous in age, which, in so far as western North America is concerned,
represented the latest occurrence of Sauropod dinosaurs, up to the
time of the present discovery. In the eastern United States, Sauro-
pod dinosaurs (Astrodon, Pleurocoelus) have been known as occur-
ring in the Arundel (Potomac) formation since Marsh first described
them in 1888, but for a long time the Arundel was correlated with
the Morrison formation (Atlantosaurus beds) of the west, but more
recently, largely on paleobotanical evidence, it has been referred to
the Lower Cretaceous. A recent restudy of the Arundel vertebrates °
appears to indicate a higher position in the Lower Cretaceous than
has previously been given them. It is also of interest that the Mary-
land Sauropoda are found associated with the remains of other dino-
saurs having undoubted Upper Cretaceous affinities, as is the case with
the bones now under discussion.

It thus appears that these specimens, found under conditions which
allow no question of doubt to be raised, furnish the first indisputable
evidence of the occurrence of Sauropodous dinosaurs in the Upper
Cretaceous of North America.

REPORTED DISCOVERIES OF SAUROPOD REMAINS IN UPPER
CRETACEOUS DEPOSITS

There have been a considerable number of reported occurrences of

Sauropod dinosaur remains in Upper Cretaceous deposits in various

parts of the world. Those recorded are from India, southern France,

South America, Madagascar, German East Africa, and Egypt. These

1Brown, Barnum, The Cretaceous Ojo Alamo beds of New Mexico, with
description of the new dinosaur genus Kritosaurus: Am. Mus. Nat. Hist.
Bull., vol. 28, p. 260, 1910.

Gilmore, C. W., Vertebrate faunas of the Ojo Alamo, Kirtland, and Fruit-
land formations, U. S. Geol. uae Prof. Paper No. 98, pp. 283-284, fig. 28;
p. 285, 1916.

? Brown, Barnum, idem, p. 27

* Gilmore, C. W., idem, p. 287.

* Gilmore, C. W., idem, p. 288, pl. 73, fig. 1.

* See Larkin, Pierce, Journ. Geol., vol. 18, No. 1, 1910, p. 93.

* Gilmore, C. W., Proc. U. S. Nat. Mus., vol. 59, 1921, pp. 581-594, pls. 110-114.

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

finds, however, have been called into question for one cause or another,
so that as the evidence stands to-day, their exact status is very unsatis-
factory. :

Below I shall briefly review these various discoveries, although no
attempt will be made to critically re-examine the evidence, however
desirable that may be. I feel that in this problem I should defer to
those having a wider knowledge of geological structures, and especi-
ally to those who have available extensive collections of Sauropodous
dinosaur materials with which to make the necessary comparisons.

Taken in chronological order these reported finds are as follows:

1. Apparently the first Sauropod remains to be described from the
Middle Cretaceous or above were those found in the Lamenta beds,
probably Cenomanian, near Jabalpur, India. These specimens were
described by Falconer * in 1862, without name, and it was 1877 before
their Sauropod nature was recognized by Lydekker,’ who redescribed
them under the name Titanosaurus indicus, the type being a post-
median caudal vertebra.

2. In 1893, Lydekker* described various dinosaur bones from
the Guaranitic beds of Patagonia, referring them to the Sauropod
genera Titanosaurus and Argyrosaurus. These identifications were
based upon numerous vertebral centra, limb and foot bones, frag-
mentary parts of the sacrum, pelvis, etc. The limb bones appear to
have been in a splendid state of preservation and are certainly sauro-
pod in aspect, though this assignment is seriously questioned by
Nopesa as I shall show later. Hatcher * observes that the Guaranitic
beds “are referred to the Upper Cretaceous both upon stratigraphic
and paleontologic evidences. Just where they should be placed in
that series cannot be determined until we know more of the Dinosaurs
contained in them. At present it seems not improbable that they will
prove to be the equivalent of the Laramie of North America, as they
have long been considered by Dr. Ameghino and others.”

3. In 1899 Depéret® recognized the Sauropod genus Titanosaurus as
occurring in the Danian of the Montague Noire of southern France.
This determination was based upon a femur, an identification to
which, as in the former case, Nopcsa makes serious objection as to
its validity. He says:° “I wish briefly to draw attention to the
fact that the Upper Cretaceous Titanosaurus, as known from the

* Paleontological Memoirs, vol. 1, 1868, p. 418, pl. 34, figs. 3, 4, 5.
* Rec. Geol. Surv. India, vol. 10, 1877, pp. 38-41.

* Anales Museo de La Plata, vol. 2, pt. 1, 1893, pp. I-12, pls. 1-5.
*Amer. Journ. Sci., vol. 9, 4th ser., 1900, pp. 94, 95.

* Bull. Soc. Geol. France, 3d ser., vol. 27, 1890, p. 692.

* Geol. Mag., n. s., Dec. 5, 1910, p. 261.

INKOR dls A NEW SAUROPOD DINOSAUR—GILMORE 9

Montague Noire in France and from the Cretaceous formation of
Argentina, and perhaps also from East Africa, has nothing to do with
the Sauropoda, but belongs to the Trachodontid Orthopoda, as proved
by the abundant Transylvanian material at my disposal.’ He then
goes on to point out that these dinosaurs referred to Titanosaurus
are generically identical with Telmatosaurus, a heavily built Tracho-
dontid animal from Transylvania, of which the structure is largely
known from undescribed material, the name Titanosaurus being
applicable only to the English Wealden Sauropod described in 1887.'

Feces, 1907, Thevenin~* described certain fossils from Madagascar
that were referred to the Sauropod genera Bothriospondylus and
Titanosaurus. Vhese are Cenomanian in age.

5. In 1907, Dr. E. Frass made the interesting discovery of Sauropod
dinosaurs in southern German East Africa. These were at first
thought to be of Upper Cretaceous origin, but since it now appears to
be universally recognized that these animals are from the Lower
Cretaceous and Upper Jurassic, they may be dismissed from further
consideration in the present connection.’

6. Dr. W. D. Matthew, in a recent letter, informs me that Stromer
has some fine Sauropod material as yet undescribed, “‘ with that
extraordinary Spinosaurus* from the Upper Cretaceous of Baharich
Oasis, Egypt.” According to Stromer these are Cenomanian-Albian
in age.

From this brief review of the reported finds of Sauropod remains
of Cenomanian age or above, it will be seen that much doubt exists
either as to their proper identification or to their exact geological
position. In the light of this more recent discovery, under conditions
that permit hardly a question of doubt to be raised as to either identity
or stratigraphic position, it would appear very probable that a critical
re-examination of the evidence would show the original determination
of some of these finds to be valid in all respects. That Sauropod
dinosaurs continued to exist until after the Cenomanian, and even
into the Danian, there is every reason to believe.

_*In this Nopcesa is mistaken for the name would certainly be applicable to
the genotype which is the specimen from India described by Lydekker in 1877
and again reviewed in the Quarterly Journal of the Geological Society of
London, p. 156, in connection with the Wealden Sauropod cited by Nopcsa.
It is also of interest to note that Seeley, in the same article, p. 160, regarded
the specimens from India as being insufficient for purposes of identification,
or to enable the relations of the animal to be determined.

* Annales de Paleontologie, 1907.

* See Schuchert, Bull. Geol. Soc. America, vol. 29, No. 2, 1918, p. 264, for
citations of articles and discussion of the age of the Tendagura series.

* Abh. Bayer. Akad. Wiss., Nov., I9I5.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72, NO 14, PL. 1

LEFT SCAPULA OF ALAMOSAURUS SANJUANENSIS

Type. About one-tenth natural size

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72, NO 14, Pl. 2

NB

RIGHT ISCHIUM OF ALAMOSAURUS SANJUANENSIS

Paratype. About one-tenth natural size. 1. External view. 2. Internal view

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 72, NUMBER 15

EXPLORATIONS AND FIELD-WORK OF THE
SMITHSONIAN INSTITUTION
IN 1921

Car,
ae THSOS O ry,
Cae

(PUBLICATION 2669)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1922

The Lord Baltimore (ress

BALTIMORE, MN,, U. S. a.

Li

PAGE
LURE OVC III CIE VG (1 NS RAEN i cp ait Oe re RO cc Re = ER OD I
Geological Explorations in the Canadian Rockies................00.e000% I
Paleontological Field-Work in the United States. ..........0:..61se.0. 2% 22
Astrophysical: Field-Work in Arizona and im Chile. 2.0.0. 0.6.e00.50056% 30
Botamcabebaspedition: to the Onientt..ic woul vee caccs eee bi euns Gab celawen 33
Biological Exploration in the Dominican Republic....................... 44
PeSpetitenes ih WeLCLEMiby o> 2.41 keoaes phe wat Ree Ree ne i ereoe ale isa og ik eee 47
Bntomolocicalebexneditionmtoy Alaska erin sae ities e seine sicisenee caeaee ne 52
Archeological Field-Work on the Mesa Verde National Park............ 64
Archeological Collecting in the Dominican Republic................:.... 83

Archeological Reconnaissance of the Cahokia and Related Mound Groups. 92

Archeological Investigations at Pueblo Bonito, New Mexico............. 106
Archeological Field-Work in South Dakota and Missouri................ L7
Field-Work on the Kiowa, Pueblo, and California Indians................ m2

Archeological Field-Work on the Susquehanna River, Pennsylvania...... 127

EXPLORATIONS AND FIELD-WORK OF THE
SMITHSONIAN INSTITUTION IN 1921

INTRODUCTION

The exploration and field-work conducted by the Smithsonian In-
stitution is one of the means employed for the “ increase and diffusion
of knowledge,” the purpose of the Institution as stipulated in the
will of James Smithson, its founder. Attention is directed whenever
possible to regions which have previously been imperfectly explored
from a scientific point of view, and during the seventy-five years of
its existence, the Institution’s field parties have been able to make
notable additions to existing knowledge as well as to provide vast col-
lections of biological, zoological, and anthropological material for the
exhibition and study series of the United States National Museum, a
branch of the Institution.

During the past year, the effectiveness of the Institution’s limited
funds for this work has been so reduced by the prevailing high costs
that it was not possible to take part in as many expeditions as is
customary. The more important of those which did take the field are
briefly described in the present pamphlet, which serves as an announce-
ment of the results obtained, many of the expeditions being later
treated more fully in the various series of publications under the direc-
tion of the Institution. The photographs here reproduced were for the
most part taken by the field-workers themselves.

GEOLOGICAL EXPLORATIONS IN THE CANADIAN ROCKIES

The geological work by Secretary Charles D. Walcott in the
Canadian Rockies was in continuation of that of the field seasons of
1919, 1920, for the purpose of securing data on the pre-Devonian
strata of the Sawback range in Ranger Brook Canyon, and a recon-
naissance of the pre-Devonian formations to the northwest as far as
the headwaters of the North Fork of the Saskatchewan River, Alberta.

The season was an unusually cold and stormy one. The party
started with a pack train from Banif, June 30, and returned September
30. During this period there were 35 stormy days, 28 cloudy and cold
days (20° to 45°) and more or less snow fell on 20 days in August and

SMITHSONIAN MISCELLANEOUS COLLECTIONS. VOL. 72, No. 15

i ae

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| i anoramice vieyv K 1ewan r loo ¥ s ) r r N ¢ yunts Outram 0,07¢ and F¢ es (12,102 ; Glac La aly , and
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NV Alver
oking west up the iver toward Moun u ( C ke cany - eN pias ;

Locality: The view is fro i
4 ma point about 47 § (75.2 k ve f Lake ise Stati : i if i |
I out 47 miles (75.2 km.) northwest of Lake Louise Station on the Canadian Pacific Railway, Alberta, Canada. (C.D. Walcott, 1921.)

e

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MISCELLANEOUS COLLECTIONS

SMITHSONIAN

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9 hic 3.—Upturned Devonian, Ordovician, and Cambrian strata southwest of Badger Pass, at head of Cascade Creek and northeast of canyon of Johnson Creck
Locality: Southeast side of canyon leading up from Johnson Creek to Badger Pass in Sawback Range. Position of camera about ten miles in air line east of Lake Louise Station
on the Canadian Pacific Railway, Alberta, Canada (( D. Walcott, 1021.)

Fic 4. Thompson Pass on Continental Divide \fountain on s¢
Thompson Pass about 63 miles (101.3 km.) northwest of Lake Louise Station on the ( n Pacific Rai , Alberta

(‘1z61 ‘WORM ‘G ‘O SIN pue A) “epeued eyeq vy ‘KEMIICY WjIOVg ueIpeuey 94} UO UOT}eIS asInoT
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1921

XPLORATIONS,

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VOL, 72

COLLECTIONS

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MISCI

SMITHSONIAN

SMITHSONIAN EXPLORATIONS, 1921

15

NO.

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6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

September. While on the trail 30 camps were made, but owing to
weather conditions and to the fact that the snow remained on the slopes
and cliffs above timber line, a relatively small amount of productive
work was accomplished.

The section studied near the head of Ranger Brook Canyon of the
Sawback Range about 12 miles (19.3 km.) northwest of Banff, was
from the base of the Devonian limestones down through the post-
Cambrian (Ozarkian) Mons formation and the subjacent Lyell and
Sullivan * formations of the Upper Cambrian.

Fic. 8—Camp on the lower eastern slope of Fossil Mountain looking north
toward the head of Red Deer River.

The character of the formations is indicated by figures 2 and 3,
which show the southwesterly slope of the highly inclined beds (45°
to 70°) and the saw-tooth-like effect caused by the unequal rate of
erosion of the massive bands of limestone and the softer, more friable
sandy and clay shales. Towards the northwest end of the Sawback
range at the Red Deer River the Black and White Douglass mountains
stand high above the surrounding ridges. (Fig. 5.) Oyster Mountain

*See Exploration pamphlet for 1919, Smithsonian Mise. Coll., Vol. 72,
No. I, 1920, p. 15.

ue 0} surjurod st JJOITVAA “SAT

1921

XPLORATIONS,

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SMITHSONIAN

NO. 15

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8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

(fig. 6) has been cut out by erosion from the limestones between
Douglass and Fossil mountains, and figure 6 illustrates the crumpling
of shaly limestones by thrusting of a series of massive limestone strata
against them during the period of displacement of the great series of
formations of this part of the Cordilleran ranges.

Fossil Mountain, named from the presence of Devonian corals, is
about 9 miles (14.4 km.) northeast of Lake Louise Station and faces
Baker Creek Pass on the east. It has a good section of Devonian and

Fic. 10.—Wild flower camp on northwest side of Johnson Creek Pass.
(Mrs. Mary V. Walcott, z921.)

pre-Devonian rocks on its eastern slope. There is a fine outlook from
camp at the east foot of the mountain.

The broad U-shaped valley (fig. 9) between Fossil and Oyster
mountains has been eroded in the shale and thin bedded limestones
that pass beneath Fossil Mountain; this formation is one of those in
the Sawback Range that is readily worn away, with the result that the
agencies of erosion followed by the glaciers have made a valley al-
together disproportionate to the present erosion agencies, water, frost
and snow.

At a camp in the heart of the Sawback Range on a tributary of
Baker Creek leading up to Johnson Pass there was a wonderful

NO. 15 SMITHSONIAN EXPLORATIONS, I921 9

exhibit of wild flowers in bloom. Mrs. Walcott counted 82 species
within a short distance of the tents. A spring-fed pond supplied camp
water; dead pines and spruce, firewood; and a grass covered snow-
slide slope, abundant feed for the horses.

The moss pink (fig. 11) and the beautiful Dryas octopetala were
very abundant, but heavy frosts in August killed nearly all the plants
and few of the flowers went to seed.

On our way north we crossed over Pipestone Pass and down the
Siffleur River. Clearwater River heads in glacial gravels on the east
side of the Siffleur about two miles north of Pipestone Pass. Figure
13 is a view looking west through the Clearwater Pass and across to
the high cliffs on the western side of Siffleur Canyon.

Twenty-five miles further to the northwest at the point where the
south branch (Mistaya Creek), the middle branch (Howse River),
and the north branch unite to form the Saskatchewan River, there
are some beautiful and instructive views of the surrounding mountains.
Figure I (frontispiece) is a fine view of the head of the river, with
Howse River in the left background and the North Fork beyond the
island on the right. The Mount Forbes massif on the left is a superb
mountain mass and in the distant center is Division Mountain at the
head of Glacier Lake Canyon which we visited in 1919; on the right
Survey Peak and beyond two unnamed points. The Glacier Lake
section of the pre-Devonian and Upper Cambrian formations was
studied on the northern slopes of the Mount Forbes massif as illus-
trated by figure 1 (frontispiece) of the Smithsonian exploration
pamphlet for 1919, and the rugged cliffs and peak of Mount Forbes
are shown by text figure 14 of the present number.

Twelve miles northeast of Mount Forbes the cliffs of Mount
Murchison (fig. 15) rise high above the dark forested slopes and
present a view of the Devonian and pre-Devonian formations that is
unequalled in all this region of peaks, cliffs and broad canyon valleys.

Opposite Mount Murchison on the north side of the Saskatchewan,
Mount Wilson (fig. 16) presents another section of the pre-Devonian
formations, the upper end of which is a massive white quartzite formed
of the sands of the beaches over which the Devonian Sea deposited
thick layers of calcareous sediments abounding in the remains of
corals and various invertebrates of the time. On the west, Mount
Wilson rises directly above the North Fork of the Saskatchewan which
here flows through a narrow picturesque inner canyon (fig. 17).

1Smithsonian Misc. Coll., Vol. 72, No. 1.

IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

FG, 12.—Dryas octopetala below Johnson Creek Pass.

II

EXPLORATIONS, I921

SMITHSONIAN

NO. 15

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NO. 15 SMITHSONIAN EXPLORATIONS, 1921 7

The trail up the North Fork follows the bed of the river most of the
way to its head beneath Wilcox Pass. The same is true of the trail
up the west branch called Alexandra River, and its northwest exten-
sion named Castleguard River, by the Interprovincial survey of the
boundary between Alberta and British Columbia. Near the union of
Castleguard and Alexandra Rivers there is a fine view of the peaks
along the Continental Divide and Alexandra glacier. On one of the

Fic. 19.—Mount Wilson and glacier from the southeast, with the eastern
section of the broad syncline, of which Mount Wilson is the western section, on
the right.

Locality: View taken from south shore of Saskatchewan River about two
miles (3.2 km.) east of Mistaya Creek and 47 miles (75.2 km.) northwest from
Lake Louise Station on the Canadian Pacific Railway, Alberta, Canada. (Mrs.
Mary V. Walcott, 1921.)

misting days of early September a photograph of Alexandra glacier,
Queens Peak, and Mount Alexandra was taken from the river bed
and is reproduced as figure 20.

Castleguard River heads in a deep, rather broad canyon at the foot
of the Castleguard glacier. Thompson Pass is on the southwest and
high barrier ridges on the northeast. On the summit of the latter
great terraced buttes occur with narrow side facing the line of drainage
(fig. 21). These outlying buttes are formed of the alternating hard

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TITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 22.—Waiting for the pack horses to be brought up to receive camera boxes,
dunnage bags, blanket rolls, tent, and small impedimenta.

Fic. 23.—Horses foraging through the snow.

NO: 25 SMITHSONIAN EXPLORATIONS, 1921 FA

and soft bands of limestone and shale of the Sullivan * formation, and
they form a somewhat unique topographic feature, and are the top of
the world at this point.

Thompson Pass is one of the scenic features of the Continental
Divide when viewed from the high Alpine valley on the northeast side
of Castleguard Canyon. The Pass is low (6,511’ or 1,984 m.), and
bold high ridges lead up to mountain summits on either side (fig. 4).
A view taken on a misting day shows Watchman Lake (6,050’ or

Fig. 24.

A snowy morning on upper Pipestone River.

1,844 m.) and above it Cinema Lake (6,400’ or 1,950 m.) on the north-
east slope of the Pass. On the south Watchman Peak (8,674' or
2,634 m.) which lies in front of Mount Rice (10,745’ or 3,275 m.)
and on the right Mount Bryce (11,000’ or 3,352 m.) and Bryce glacier,
which is at the head of the middle branch of Castleguard River. The
Castleguard glaciers flow down from Mount Castleguard (10,090’ or
3,075 m.), which is a fine peak a few miles northeast of Mount Bryce.
Figure 4 is a fine illustration of a misting day along the Continental
Divide. We were camped for a week on the south side of the Alpine

*Smithsonian Misc. Coll., Vol. 72, No. 1, 1920, p. 15.

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

upland in the foreground, and on each day numerous squalls of fine
snow or frozen mist would sweep over from Thompson Pass or
Bryce glacier.

I do not know the origin of the names of Rice and Bryce, but it
is probable that the mountains were named in honor of Sir Cecil
Arthur Spring-Rice and Lord James Bryce.

As the result of unfavorable weather not more than one-third of
the work planned was completed when the late September snow drove
us back to the railroad. The morning we broke camp to go to Lake
Louise Station the horses were pawing away the snow to get at the
grass beneath (fig. 23), and the snow was very beautiful on the trees
and along the stream below camp (fig. 24). The trail was obscured
by it and to make matters more complicated, snow driven by a strong
east wind beat into our faces during the seven hours march. The
next day the sun came out and the storms were forgotten except for
the wonderful snow scenes along the trail down the Pipestone River.

The Commissioner of the Canadian National Parks, Hon. J. B.
Harkin, and the members of the Parks Service in the field, from
Superintendent to Park Warden, were most helpful, and the same is
true of the officials and employees of the Canadian Pacific Railway.

PALEONTOLOGICAL FIELD-WORK IN THE UNITED STATES

Field-work by the Department of Geology of the U. S. National
Museum during 1921 was carried on by three members of the Division
of Invertebrate and Vertebrate Paleontology.

Dr. R. S. Bassler, Curator of the Division of Paleontology, in
cooperation with the Geological Survey of Tennessee spent the
month of July in field-work in the Central Basin of that State, where
he was occupied in collecting geologic material and in mapping and
studying the economic resources of the Franklin quadrangle in
Williamson County, south of Nashville. This area of about 250 square
miles is of economic interest, on account of phosphate and oil shale
possibilities. It is also classic ground for the paleontologist because
of the numerous outcrops of Ordovician and later Paleozoic forma-
tions which afford a wealth of fossils. During the course of the
mapping, Dr. Bassler was able to collect a considerable number of
these fossils needed in the museum study series and was also fortu-
nate in securing several large exhibits illustrating various geological
phenomena. Among the latter is a large mass of limestone composed
entirely of the dismembered calices and columns of a large species of
crinoid or sea lily in which the individual fragments are perfectly

INO: 15 SMITHSONIAN EXPLORATIONS, I92I 23

ca

Fic. 25.—Contorted and cross bedded phosphate rock, Franklin, Tenn. (Photo-
graph by Bassler.)

Fic. 26—Massive limestone with an intercalated coral reef, near Franklin,
Tenn. (Photograph by Bassler.)

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

preserved and admirably illustrate the formation of a limestone
through the accumulation of this type of animal remains. Material
was also secured, both for the exhibition and study series, illustrating
the origin of the phosphate beds of the locality through the removal
from a phosphatic limestone of the easily soluble calcium carbonate
by the leaching power of surface waters. Such material is represented
in figure 25 showing a rock outcrop where a porous limestone is over-
laid by the contorted and crossbedded rock which upon such leaching
gives rise to the phosphate.

Among the interesting stratigraphic results secured was one show-
ing the efficacy of coral reefs of the Ordovician in rock formation.
The massive limestone about fifteen feet thick shown in figure 26
represents a middle Ordovician formation here containing but a single
reef but within a distance of ten miles the number of intercalated coral
reefs has so increased that the formation attains a thickness of over
250 feet.

An ancient Indian village near Brentwood, Tennessee, was visited
during this trip in the interest of the Bureau of Ethnology. The
object of the visit, namely the determination of the length of time
since the village was deserted, proved to be, however, outside of the
domain of geology.

Upon the completion of this work Dr. Bassler proceeded to Spring-
field, Illinois, where with the permission of Dr. A. R. Crook, Chief
of the Museum, he prepared casts of the type specimens of invertebrate
fossils contained in the [linois State Museum collections. The aim
in this work is to make the national collections of invertebrate fossils
as complete as possible in its representation of type specimens, a
work which was further advanced in the early part of January by a
visit to the Walker Museum of the University of Chicago, where
the casting of all the Paleozoic species which had remained unfinished
on the occasion of a former trip was completed.

Through the courtesy of Mr. E. J. Armstrong, of Erie, Pennsyl-
vania, Dr. Bassler was enabled to visit all the classical Silurian and
Devonian localities in northwestern Pennsylvania and western New
York during the latter half of September. The object of this trip
was to obtain a field knowledge of the detailed geology and to collect
carefully selected sets of fossils illustrating the numerous formations
of this region. This work was successful and the many large collec-
tions of Devonian fossils in the museum hitherto lacking exact strati-
graphic data can now be determined and arranged in the detail neces-
sary to-day.

INOS eS SMITHSONIAN EXPLORATIONS, I9Q21 25

In April, Mr. C. W. Gilmore, the Associate Curator of Vertebrate
Paleontology was authorized to undertake a trip into New Mexico,
“for the purpose of making collections of geological material for
the National Museum and determining the advisability of preserving
certain lands in northern New Mexico for national monumental pur-
poses.” Mr. Gilmore was obliged to report that:

9

Since the many square miles of “bad lands” surrounding the reserved area
are equally fossiliferous and in places present much more favorable territory
for the recovery of fossil remains than any observed within the boundaries
of the monument, and also since the greater part of these surrounding areas
lie within Pueblo Grants over which federal control has been relinquished,
there would be no advantage in retaining governmental control of so small a
part of the area as is represented in the proposed monument.

Mr. Gilmore did, however, find a contiguous fossiliferous area in
the Santa Clara Pueblo Grant and secured for the museum a well-
preserved skull and other bones of a small rhinoceros, and in an ad-
joining Pojoaque Pueblo area remains of an extinct camel. The
most promising area for collecting would appear to lie within land
grants over which the government has at present no control.

In January, this same year, Mr. J. W. Gidley, Assistant Curator
in this Division, was authorized in cooperation with the United States
Geological Survey to conduct field explorations in the San Pedro and
Sulphur Springs Valleys of southern Arizona and on the completion
of this work to visit the La Brea asphalt deposits of southern Cali-
fornia and from there go to Agate, in Nebraska, for the purpose
of securing other exhibition material. The work in Arizona was
eminently successful, Mr. Gidley shipping some 24 boxes having
an aggregate weight of 5,000 pounds. The bulk of this collection, he
reports represents “a practically new Pliocene fauna containing about
60 vertebrate species, most of which are mammalian.”

In detail Mr. Gidley reports essentially as follows:

“ The geological structure of the San Pedro Valley will be published
in detail by Doctor Bryan of the United States Geological Survey.
It, however, may be noted here that this beautiful desert valley, now
drained by the Rio San Pedro (which, rising near the Mexican border,
runs nearly north-northwest, emptying into the Gila River, more than
a hundred miles away), narrows and deepens as it runs northward
from Benson leaving relatively small and scattered areas of sedi-
mentary deposits which may contain fossil vertebrate remains. Most
of our work, therefore, was confined to the upper valley, which forms
a rather wide basin bounded on the east by the Dragoon mountains,
on the west by the Whetstone Range, and on the south by the Tomb-

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

stone mountains, and extends northward a few miles below the town
of Benson.

‘ Erosional exposures in this general region are quite extensive, but
time and funds being limited the work done on this expedition was
confined entirely to two promising localities of relatively small area,
previously located by Doctor Bryan. One of these is situated on
the west side of the valley, about two or three miles due south of

Fic. 27—General view of fossil bearing exposure at Curtis Ranch locality,
looking across the San Pedro Valley. Partly excavated bones of Glyptodon
in foreground. (Photograph by Gidley.)

Benson, the other on the east side, at the head of a large ‘ wash’
three miles east of the Curtis ranch which is situated on the state
road about 14 miles south-southeast of Benson and an equal distance
northwest of Tombstone. The latter locality occupied the greater
part of my time and yielded by far the greater amount of material,
although the number of species later collected in the Benson locality,
slightly exceeded those found here.

“ Among the larger, and, from the museum standpoint, more im-
portant specimens secured at this locality are included parts of two
skeletons of a new species of mastodon, and parts of three skeletons
of a large armored edentate, Glyptotherium, which when restored
should make a striking exhibition piece.

NO. 15 SMITHSONIAN EXPLORATIONS, I92I 27

“Other material obtained here consists of remains representing a
wide variety of species which include a large and a smaller species
of camel, the latter apparently closely related to the South American
guanaco; two or three species of horses, a species of deer; a small
extinct antelope of the Merycodus type; a carnivore related to the
dog-wolf group but more primitive in some respects than any of the
living forms; several new species of the rodent group, but all belong-

Fic. 28.—Portion of the carapace or bony skin covering of a Glyptodon, parti-
ally excavated. Curtis Ranch locality. (Photograph by Gidley.)

ing to modern genera; two species of land turtles, and a species of
bird not yet determined.

“ At the close of this work, which had nearly exhausted the original
allotment for field expenses, an additional sum was granted, whereby
it was possible to proceed with a desired investigation planned for
earlier, in the Sulphur Springs Valley near Willcox.

“T arrived at Willcox on the 15th of March. As found ona previous
visit the conditions were not such as would inspire enthusiasm over
the prospects of a good collecting-field. The surrounding country
stretched away for miles in every direction almost as level as a floor,
with no erosional exposures ; and had not recent fossil remains already

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

been discovered through the digging of a shallow well in the vicinity
no one would have suspected their presence here. Several years
earlier fossil bones had also been found at nearly the same depth
(about 9 feet) in another well, now filled in, which had been dug
at a distance of about 250 feet from the present open one. It was
thus assumed that the fossil-bearing gravel deposit was of rather wide
extent, and that by making a long stripping with plow and scrapers, a
considerable area of “ pay gravel” might be uncovered and worked
at comparatively small expense. The spot chosen as being most
promising was naturally that between the two wells.

“At Willcox, the services were procured of a reliable man with
teams, plow and scrapers and this work was put into execution.
Thanks are here due Mr. Harris, a local real estate agent, who lent
valuable aid in this connection. I was also indebted to this gentleman
for permission to put through the project, for the locality worked
was on deeded land which he had in charge.

“As the stripping progressed, it became evident that the strata, or
layers, of deposits passed through did not conform to the section
exposed in the abandoned well. Hence, on reaching the 7-foot level
three prospect holes, about 15 feet apart, were put down to a depth
of about 6 feet, or 4 feet lower than the top of the gravel deposit in
the well. In none of these holes was there any sand or gravel en-
countered thus proving that the gravel exposed in the well was part
of an ancient stream channel of limited lateral extent. This discovery
of course caused a complete abandonment of the trench excavation
work, and the remainder of our time was spent in ‘ mining’ the
gravel from the sides of the well as far as was considered safe to do
so. In this way several good fossil horse teeth were procured.

“From Willcox, I went by way of Tucson to Feldman, arriving there
about noon of the 29th of March, where I was joined by Dr. Bryan.
Feldman is a ranch and post office in the lower valley of the San
Pedro, about 90 miles north-northwest of Benson and about 10 miles
above the junction of the San Pedro with the Gila. The valley here
is very much narrowed and deepened, the river bed being nearly
2,000 feet lower than at the Curtis ranch. The gradient of the
streams and ‘ washes’ emptying into the San Pedro in this vicinity
is very steep and benches and divides rise quite abruptly on either
side. Erosional exposures one might expect to find here under
these conditions are very much reduced by a heavy covering of gravel
of relatively recent age. But paleontological evidence for confirming
the age of this part of the valley was so much desired, a special effort

INOS U5 SMITHSONIAN EXPLORATIONS, 1921 29

Fic. 29.—Base of skull of mastodon with tusks in position, partially excavated.
Curtis Ranch locality. (Photograph by Gidley.)

Fic. 30.—Searching for small mammal jaws in excavation made in collect-
ing one of the mastodon skeletons. Curtis Ranch locality. (Photograph
by Gidley.)

3

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL iz

to procure it was considered worth while. However, the few days
spent here met with little success, and owing to the great inconvenience
of continuing it further without more complete field equipment, the
project was abandoned. On the morning of April 2, we left Feldman,
returning to Benson via Tucson, and the next morning began a
systematic search for fossils at a locality about two to three miles
south of the town. During our earlier stay at the Curtis ranch we had
made one short visit to this locality, the material obtained then sug-
gesting a slight difference in age, or phase, between these deposits
and those of the Curtis locality. The material obtained at this place
is fragmentary and abounds mostly in remains of mammals of small
size, intermixed with which were bones of birds of several species
sufficiently well preserved for their determination, and a new species
of box turtle. Here remains of thirty-four species of vertebrates were
recovered. This collection, together with the material obtained at
the Curtis ranch locality, in which 26 species are represented, makes
up a very considerable fauna which should not only do much toward
definitely determining the age of the beds of the San Pedro Valley,
but will also throw valuable added light on the at present very little-
known animal life of the upper Pliocene of America.”

From Arizona, Mr. Gidley proceeded to Los Angeles; California,
where he passed a week studying the museum of the southern branch
of the University of California and in examining the well-known
asphalt bone deposits of the Rancho la Brea. From Los Angeles, he
proceeded on the 16th of April to Agate, Nebraska, prepared to carry
out a second detail of field-work mentioned above. He was unfortu-
nate here in encountering bad weather, but succeeded in securing for
the museum a block of the bone-bearing sandstone some 34 by 54 feet
and 14 inches in thickness. This was shipped to the museum and
preparation for exhibition is now under way.

ASTROPHYSICAL FIELD-WORK IN ARIZONA AND IN CHILE

As stated in last year’s Exploration pamphlet,’ the solar radiation
work of the Smithsonian Astrophysical Observatory was removed
from Mount Wilson, California, to Mount Harqua Hala, Arizona, in
September, 1920, in order to observe under better sky conditions,
and in a more favorable place for continuing the observing the whole
year round. Under the charge of Dr. C. G. Abbot the work was

* Smithsonian Misc. Coll., Vol. 72, No. 6.

NO. 15 SMITHSONIAN EXPLORATIONS, I92I 31

established and continued at Mount Harqua Hala until January 20,
1921, when it was taken in charge by Mr. L. B. Aldrich. He remained
until May 20, 1921, when he was relieved by Mr. A. F. Moore,
formerly director of the Smithsonian private observing station at
Calama and Montezuma, Chile. Under Mr. Moore’s charge, the
work has been continued steadily at Mount Harqua Hala, with the
assistance of Mr. F. A. Greeley.

The Smithsonian Institution maintains from the income of the
Hodgkins fund a similar station at Mount Montezuma, near Calama,
Chile, under the direction of Mr. L. H. Abbot assisted by Mr. P. E.
Greeley. From this Chilean station daily telegrams are forwarded
to Buenos Aires, Argentina, giving the observed value of the solar
constant of radiation for the day. These data are employed regularly
by the Argentine Weather Bureau for weather-forecasting purposes.

While the Smithsonian Institution is not yet in position to champion
the use of statistics of solar variation for meteorological forecasts, the
great interest which its studies of solar variability have aroused here
and abroad seems clearly to warrant the continued maintenance of
the Arizona and Chile solar stations under the best possible observ-
ing conditions for several years, until a satisfactory basis for a test
of the solar variability as a weather-forecasting element has been laid.

The present year has unluckily proved unfortunate at both stations.
At Mount Harqua Hala the spring months were very hazy, the
summer and autumn months unusually cloudy, with almost unpre-
cedentedly heavy rainfall. At Montezuma the cloudiness of the
earlier months was quite unprecedented. During August and Sep-
tember a disarrangement of the apparatus caused apparently by earth-
quake, combined with illness of the director, led to the loss of many
observing days.

In October, Dr. Abbot began an inspection trip to Montezuma,
arriving at the station on November 15, and remaining until December
14. During this interval of 30 days, the observers fortunately were
able to determine the solar radiation on 26 days, and generally with
three or four closely agreeing determinations per day. All of the
apparatus was readjusted and improved to the most perfect state of
fitness. Many of the results in these conditions proved of a higher
grade than ever before observed. In fact it would be hard to con-
ceive of anything which could add now to the excellence of the
Montezuma station and outfit.

The accompanying illustrations show the desolate, rainless char-
acter of the region ; figure 31 shows the mountain top with the observ-

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

Fic. 31—Summit of Mount Montezuma. Observing cave near the top.

Fic. 32.—Garage, Shop and Dwelling, Mount Montezuma,

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 33

ing cave; figure 32, the group of buildings comprising the observer’s
quarters, the shop and the garage; and figure 33 the entrance to the
observing cave with such observing apparatus as is employed outside
during observations of the solar constant.

It is possible to drive the automobile on high gear clear to the
observer’s quarters which are situated at the head of a canon sheltered
on the west by a rise of several hundred feet from the strong west
winds of afternoon. There is almost invariably practically complete

Fic. 33.—Pyranometer, coelostat, pyrheliometers and theodolite with L. H.
Abbot, Director at Mount Montezuma.

absence of wind for several hours after sunrise, a thing highly favor-
able to morning work.

The observing cave near the top of the mountain is less than
10 minutes walk from the observer’s quarters. It is only necessary to
go up twice a day, once to observe, and again at 8.30 P. M. to signal
the observed value to Calama, whence it is telegraphed to Buenos
Aires.

BOTANICAL EXPEDITION TO THE ORIENT

During the summer and fall, 1921, Dr. A. S. Hitchcock, system-
atic agrostologist of the Department of Agriculture and custodian of
the section of grasses of the Division of Plants in the U. S. National

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

Museum, visited the Orient for the purpose of collecting and studying
the grasses, especially the bamboos. He left Washington April 25
and returned December 23, visiting the Philippines, Japan, China,
and Indo China. Six days were spent at Honolulu on the way over.
Collections were made at the following places: Philippines, Manila,
Los Banos, Baguio; Japan, Keelung (Formosa), Yokohama, Tokio,
Nikko, Lake Hakone, Mount Fuji, Kyoto, Nagasaki; China, Shang-

Fic. 34.—A peasant’s hut near Gotemba, Japan. The roofs of the building are
thatched with coarse grass. The bundles are for firewood.

hai, Nanking, Kuling, Hongkong, Canton, Wampoa, Yingtak, Shiu-
chow, Lohfau Mountain, Macao, Island of Hainan, Pakhoi; /ndo-
China, Haiphong, Hanoi, Vinh, Hue, Tourane.

The countries were visited in the order named so that collections
might be made at the most favorable season for grasses.

Collecting in the vicinity of Manila is not very satisfactory as
the native flora has been largely replaced by introduced species. From
Los Banos, the seat of the Agricultural College, a trip was made to
the summit of Mount Makeling about 3,500 feet high. This moun-
tain is of especial interest to botanists as it is the most accessible
region for the virgin forest, most of which has disappeared from the
vicinity of Manila. On this mountain was met one of the worst pests
of the eastern tropics, the leeches. At upper altitudes in the rain

NO. 15 SMITHSONIAN EXPLORATIONS, I92I 35

forest these vile worms are found in countless numbers. They attach
themselves to the skin and suck the blood with great avidity and con-
stant vigilance is necessary to prevent serious damage.

Japan is not very favorable for the collecting of grasses as it is
mostly a forested region and there is comparatively little open country.
The bamboos were of interest as there are many species. In the
Lake Hakone region the hills were covered for miles with a single

: wae. Coats is

Fic. 35.—Hills near Lake Hakone, Japan. The vegetation on the distant
slopes is almost exclusively a single species of bamboo (Arundinaria chino),
4 to 8 feet high.

species of bamboo (Arundinaria chino), 4 to 8 feet high, often to the
exclusion of everything else.

China on the other hand was very rich in grasses. One of the
surprises of the trip was to find so much open grass land in a country
that is said to be very thickly populated. The cities of China are
very much crowded and the valley lands are intensively cultivated,
but the hills are unoccupied and almost unused. This is in striking
contrast to our own western regions where, except in National Forests
and other protected areas, the grass lands are extensively grazed.
The basic reason for this condition in China appears to be the risk
from bandits. The valley lands can be protected but the hills are
open to the attack of robbers.

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

China was entered at Shanghai, a large comparatively modern city,
much under the influence of foreigners. Here is the only American
post office outside of the United States or its possessions. Mail can
be sent from here under frank or with United States postage stamps.
The two other places visited in central China were Nanking and
Kuling. At the former city is the University of Nanking, a flourish-
ing missionary institution, which extended many courtesies to Doctor

Fic. 36.—A street scene in Shanghai.

Hitchcock. Nanking is a thoroughly Chinese city showing little
foreign influence. Like most Chinese cities it is surrounded by a
high wall, this one being 32 miles in length and 30 to 50 feet high.
Kuling is a resort on a mountain south of the treaty port Kiu Kiang,
where the missionaries and other foreigners of central China con-
gregate during the summer.

During the visit of Doctor Hitchcock the Yangtse River was in
flood and the rice fields of the valley were covered with water. The
unfortunate peasants were in the water up to their waists or even
to their shoulders cutting the rice and placing it in small circular

INO: D5 SMITHSONIAN EXPLORATIONS, 1921 B97

Fic. 37.—A typical valley at Nanking, China, showing intensive cultivation.
There is a fish pond in the left foreground. The hills on each side of the
valley are covered with grass, much of which will be cut and used for fuel.

Fic. 38.—A ricksha party just after passing out through one of the main gates
of Nanking China. The city wall is about 50 feet high.

ios)
CO

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 39—The Yangtse Valley above Nanking in flood. View from a river
steamer.

Fic. 40.—Slender pieces of split bamboo drying in the sun. From these joss
sticks are to be made.

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 39

boats. The bundles were supported on the ends of crossed poles on
the dikes to hold them out of the water to dry.

Fic. 41 —A clump of bamboo, Canton, China. A common ornamental plant.

The gateway to south China is Hongkong, a very mountainous
island owned by the British, the peak being 1,800 feet high. There is
here a botanic garden and a herbarium. Canton lies up the river
west of Hongkong about 80 miles. Opposite Canton on the island of
Honam is the Canton Christian College, where Doctor Hitchcock

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

made his headquarters. xcursions were made to Yingtak and
Shiuchow on the North River north of Canton, to Lohfau Moun-
tain east of Canton and north of Sheklung, to Wampoa 10 miles east
of Canton, where the Wilkes Expedition made collections, and to
Macao, a Portuguese possession 40 miles from Hongkong and the
oldest foreign settlement in this region.

A more extended trip was made in company with Mr. McClure
of the Canton Christian College, to Indo-China and the Island of

et tes
igs

t

3
P35
wee

Fic, 42.—A street scene in Yingtak, on the North River, about 80 miles north of
Canton. The bundles of stalks are to be used for firewood.

Hainan. Going from Hongkong to Haiphong, a stop was made at
Pakhoi on the southern coast of Kwantung Province. Here forty-
six species of grasses were obtained in a few hours on the sandy
areas and rocky hills. Haiphong is the port of Tongking. Indo-
China is a French Colony (officially French Indo-China), consisting
of five divisions, Tongking, Annam, Cambodia, Cochin-China, and
Laos. The objective in Indo-China was Hue, the capital of Annam.
Loureiro, a Portuguese botanist, resided here and published in 1790 a
flora of Cochin-China and it was to determine the identity of many of

NOZ 15 SMITHSONIAN EXPLORATIONS, I92I 41

his grasses that this interesting city was visited. To reach Hue one
goes by rail to Hanoi and then south to Vinh, the present terminus of
the railroad that is to be built to Hue and ultimately to Saigon.
Beyond Vinh one goes by auto-bus over good roads about 175 miles.
A trip was made to Tourane on the coast, connected with Hue by
railroad.

On the return trip from Haiphong to Honkkong, a stop was made
in Hainan, landing at Hoihow on the north coast. Hainan is a
seldom-visited island about 180 miles long, belonging to China.

Fic. 43.—A wayside shrine at Yingtak, China. These shrines are common but,
like the present one, often suffer from neglect.

Through the kindness of Doctor McCandliss, a missionary in charge
of a hospital at Hoihow, we were able to penetrate to the interior of
the island as far as Kachek where there is a branch missionary station.
The journey was made by boat on the river the first day and on foot
the second and third days. From Kachek a trip was made up the
river into the foothills of the Five-finger Mountains. Traveling in
Hainan as in many other parts of China is chiefly by chair carried
by two coolies.

Traveling in China is mostly by rather primitive methods. Modern
steamers ply along the coast and on the larger rivers and there are a
few railroads. The sampan, a small partly covered boat propelled by

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

oars, is common in the harbors. In the cities where the roads are
wide enough the ricksha (jinrikisha) is used. This is a two-wheeled
cart, mostly now with pneumatic tires, drawn by a coolie, and holding

Fic. 44—A sampan at Shiuchow. This is the common type of small boat
used on the rivers of south China. The bamboo pole is used to push the boat
in shallow water. Oars are used in deeper water.

one person. In the narrow streets of the cities where there is not
room for rickshas, and on the country trails or paths, chairs are
commonly used. These are covered seats supported by two poles
and carried by two coolies. Long journeys in them are far from com-

NOs 15 SMITHSONIAN EXPLORATIONS, 1921 43

fortable. In the part of China visited animals are little used for
transportation of any kind. Freight is carried on land by man-
power, one man with a pole supporting two weights, two men with

Fic. 45——A specimen of the traveler’s tree, growing in the botanical garden
at Hue, the capital of Annam, French Indo-China. The plant is a native of
Madagascar.

a pole supporting one weight, heavy loads on rude wheelbarrows, in
the cities heavy loads, as much as a ton, on carts pulled and pushed by
several men.

One of the curious sights to one visiting China for the first time
is the enormous number of graves distributed at random over the

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

country. Thousands of little mounds are to be seen on every hand,
some hemispherical and grass-covered, some more elaborate, with
stones or masonry.

The agriculture of China is intensive and in some ways much in
advance of ours. The rice fields show usually a perfect even stand,
and the amount per acre is the maximum. It represents a large amount
of labor as every stalk is set out and harvested by hand.

The botanical results of the trip were very satisfactory, a large and
valuable collection of grasses having been made.

BIOLOGICAL EXPLORATION IN THE DOMINICAN REPUBLIC

In November, 1920, Dr. W. L. Abbott revisited the Dominican
Republic, working in both the Samana Peninsula and the region lying
between Sanchez (at the head of Samana Bay) and Puerto Plata,
on the north coast. Already familiar with much of this territory, he
was able to investigate a number of new and very interesting localities.
Two weeks was spent at Sanchez; three weeks in the vicinity of
Samana, a town on the south coast of the Samana Peninsula about
20 miles east of Sanchez, and on the mountain known as Pilon
d’Aztcar ; seven weeks at several stations along the railroad connect-
ing Sanchez and Puerto Plata, among which were Villa Riva, Pi-
mentel, Cotuy, Mao, and Navarrete; two weeks in the easternmost
portion of the peninsula, in visiting Las Cacaos, Rojo Cabo, and Cape
Samana; one week on the south coast of Samana Bay in the vicinity
of San Gabriel; and one week in the region of Old Heart River, in
the north-central part of the peninsula.

Contrasting with the remaining part of Hispaniola, the population
of the Samana Peninsula is chiefly English-speaking, due to the fact
that Samana was settled by a colony of Philadelphia negroes under
President Boyer of Haiti in 1820-22. The region is well watered
and has a luxuriant vegetation, and provisions are plentiful and rela-
tively cheap. The hills extending north to the coast from Pil6n
d’Azucar are covered with unbroken forests.

The Yuna River forms a vast swamp, which occupies the entire
region at the head of Samana Bay and extends along the railroad
for a distance of 12 miles. West of this swamp region, in the vicinity
of Villa Riva, Pimentel, and Cotuy, are vast stretches of grassy
savannah. The soil is fertile, and the inhabitants are industrious and
prosperous. Beyond this region the land, except along the streams,
becomes arid and the towns, such as Guaybin, Navarrete, and Mao,
are small, poorly provisioned, and lacking in enterprise.

NO. 15 SMITHSONIAN EXPLORATIONS, I92I1 4

ont

q

Fic. 46—View down Rio Mao from schist outcrop near Bulla; cliffs of con-
glomerate in the distance.

Fic. 47,—View along the Rio Mao, near Cercado de Mao.

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Numerous caves provide an interesting feature on the south coast
of Samana Bay, one of them comprising nearly the whole interior of
San Gabriel Isle. A cave at the mouth of Naranjita River contained
a quantity of Indian bones and pottery.

A collection of about 4,000 plants was procured, representing 1,460
numbers. Of these about 20 per cent are ferns, one being an interest-
ing new species of Anemia.

The birds obtained by Doctor Abbott on this visit totaled thirty-one
skins, with a few skeletons and eggs, chiefly representing species not
previously collected by him. Of particular interest is a whip-poor-will

Fic. 48.—Harbor of Puerto Plata, looking north from Monte Isabel de Torres.

(dntrostomus), closely related to a species found in Cuba, but not
hitherto recorded from Santo Domingo. On the natural grassy plains
on the north side of the island he secured several specimens each of
the local form of the grasshopper sparrow (Ammodramus savannarum
intricatus), and of the stone-plover or thick-knee (Ocedicnemus
dominicensis ), both new to the museum collections. The thick-knee
belongs to a family of birds resembling overgrown plovers, and is
related to them. It occurs in the West Indies only in Santo Domingo,
but allied forms are found in suitable localities in Central and South
America. The family is chiefly an Old World one, and for the most
part tropical in distribution. The Santo Domingo species is well-
known to the natives, under the name “‘ boukera,” and tame individuals

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 47

are often kept about the houses for the purpose of ridding the
premises of insects and spiders.

In addition many land shells and a considerable quantity of
ethnological material were secured. Doctor Abbott left New York
about the middle of December, 1921, on another expedition to the
island, but thus far no information or material has been received
from him.

EXPERIMENTS IN HEREDITY

Progress in the experiments in heredity conducted under the joint
auspices of the Smithsonian and Carnegie Institutions by the writer,
Dr. Paul Bartsch of the U. S. National Museum, have from time to
time been published in this pamphlet and in the Year Book of the
Carnegie Institution. A summary of the results attained up to 1920
was published as “ Experiments in the Breeding of Cerions”’ in 1920,
volume 14 of the Department of Marine Biology of the Carnegie
Institution, pp. 3-55, pls. 1-59.

The reported loss of the Cerion colonies introduced into the
Tortugas which were said to have been wiped out by the hurricane of
September, 1919, made it necessary to revisit the Bahamas to secure
additional breeding material for the heredity experiments. Accord-
ingly, passage was secured at Miami on the power schooner “ Tecoma ”
for Nassau, New Providence, on May 18, and there the services of the
power boat “ Standard J” were secured for a trip to Andros.

The desired adolescent specimens.of Cerion viaregis were obtained
along King’s Road, Bastian Point, South Bight, Andros, with con-
siderable difficulty because the agricultural efforts on the part of the
local population have shifted to the ground that was occupied by the
Cerion colonies during our 1912 visit.

The colony of Cerion casablancae has met with even greater mis-
fortune, for sheep and pigs have been introduced into the region
occupied by this species, and the larger vegetation has been cut down
in order to furnish more opportune habitat for grass culture. These
new environmental conditions promise well to exterminate this colony.
The necessary material for the experiments was secured with great
difficulty.

A trip was next made through South Bight to the western end of
Andros and then back to the eastern shore through Middle Bight. On
this journey many stops were made and Cerions were gathered in
large numbers. The localities from which they were taken were care-
fully listed so that it will be possible to go back to the same spot in

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

the future and gather material for comparison with that now resting
in the National Museum.

There were several points of interest as far as the physical features
of the locality visited were concerned. In 1912 the waters of the
western end of South Bight were of a creamy consistency and the
land areas adjacent low flats, mud cracked, with flakes of oolitic
rock. On the present visit South Bight was found to be a perfectly
clear stretch of water with well-packed bottom with an abundant
growth of aquatic plants, while the land adjacent gave the impres-
sion of moss covered flats. The green element, however, was due to
blue-green algae, which appear to serve as a binding factor.

The trip was enlivened by an iguana hunt, which resulted in the
securing of several of these large lizards which are now in the collec-
tion of our Zoological Park.

Returning to Nassau, five days were spent exploring the cays off
the northwestern shore of New Providence and the adjacent main-
land. Here large collections of Cerions were made, the location of
each colony being carefully noted, so that these likewise may serve
as a check series for comparison with future generations produced in
place.

On June 3 Dr. Bartsch returned to Miami and on the following day
set sail for the Tortugas, stopping to examine the various plantings
along the Florida keys.

It was a pleasure to find that the hybrid colony on Newfound
Harbor Key, around which the greatest interest centers just now, had
escaped being wiped out by the hurricane. Evidently the rain preced-
ing the hurricane had caused the Cerions to take to the ground, as
they are wont to do for foraging purposes under such circumstances,
and the dense mats of grass here had kept them from being swept
away by the floods that had passed over them, a most fortunate state
of affairs. A large number of dead specimens were nevertheless
found, which have been placed in the National Museum for record.

Incidentally, it may be stated that another almost fledged young
great white heron was discovered on White Heron Key, the island that
furnished the specimen that was shipped to the Zoological Park two
years ago. The present specimen, which is probably a younger brother
or sister of the former sending, was also transmitted by parcel post
to the Zoo, where it arrived in good condition.

In “ Experiments in the Breeding of Cerions,” there are given on
page 46 detailed measurements of 100 specimens representing the
check series of Cerion crassilabris from Balena Point, near Guanico

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 49

Bay, Porto Rico, which were planted on Loggerhead Key in 1915.
These were figured on plates 48 to 50. On page 47 measurements
were given and on plate 51 figures of 36 adult shells of the first Florida
grown generation which were gathered in January, 1919. This year a
much larger series of first generation material was found, and 200 of
such specimens were measured.

The summaries of these measurements show that no appreciable
changes in measurements have taken place in the first generation of
Florida grown Cerion crassilabris. The measurements in size all fall
within the limits of variation, as denoted in the check series, excepting
one, 1. @., a single specimen which was found among the 200 of the first
Florida grown generation that had a diameter 0.2 mm. less than any
in the check series. There is no doubt that one could find an indi-
vidual giving such a measurement among the specimens on the native
heath of this species, for the check series was not a selected one, but a
hundred specimens taken at random.

COMPARISON OF MEASUREMENTS OF FIRST FLORIDA GROWN CERION
CRASSILABRIS WITH THE CHECK SERIES

Greatest

No. whorls Altitude diameter

Kee ate Giteckesentesin..4.1s-no 9.55 22.18 12.41
: SS ean Pirst. generation ..... 9.13 22.36 11.89
Gresicse diameter. ....:. Check SRICG me ciao scer 10.5 27.5 13.9
First generation ..... 10.4 25.7 13.2
Tee a raebae. sx oe Wilecktsemiesy ear)... 8.5 19.0 10.6
First generation ..... 8.6 19.6 10.4

It is interesting, therefore, to note that so far as the first genera-
tion of this Porto Rican Cerion is concerned, it is in complete agree-
ment with the facts adduced from the two Bahaman species.

The hurricane of 1919 destroyed the cages in which had been placed
a specimen of each of two species, in order to determine their ability
to hybridize, and to note the results of such crosses as might be
observed from such selected individuals.

A new set of cages was therefore prepared. Eleven groups of
these cages consist of four compartments, each a cubic yard in size.
The septa between compartments are double wire walls to prevent
possible mating through the meshes of the fine Monel metal wire
screen. In each of these cages there were placed a Hymenocallis
plant, some grass and dead wood rubbish, in other words, habitat
conditions which were found to be favored by Cerions at the Tortugas.
Then two half-grown specimens, one of Cerion viaregis and one of
Cerion incanum from Key West, were placed in each of the forty-four

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

compartments. These cages are securely anchored, and every pre-
caution has been taken to make sure that the mollusks will be confined
within them, and that no extraneous individuals can find entrance.
The cages are arranged as shown in the following diagram, and a
better idea of them may be formed from the photograph (fig. 50).

Tic. 49.—Diagram showing arrangement of cages.

Cages No. 45 and No. 46 are of the same size as those last men-
tioned. In cage 45 were placed 183 young of Cerion incanum from
Key West, in order to determine what percentage of these will reach
maturity. In cage 46 was placed an abnormal specimen of Cerion
viaregis. This had a spiral keel, which may be the result of an injury,
although Doctor Bartsch was unable to discover any sign of it.
With it was also placed a normal specimen of Cerion viaregis in order
to determine if this character might be transmitted to offspring.

In addition to these, five groups of cages were made which have the
same size as the four unit cages, but they have only one partition in

NO: 15 SMITHSONIAN EXPLORATIONS, 1921 51

the middle, thus making them 3 by 6 feet, and 3 feet high. In these
there were placed the following combinations :

No. 47, 25 each of Cerion incanum and Cerion viaregis.

No. 48, 25 each of Cerion incanum and Cerion casablancae.

No. 49, 25 each of Cerion incanum and Cerion uva.

No. 50, 25 each of Cerion incanum and Cerion crassilabris.

Fic. 50—A portion of the monel metal wire cages used in Cerion breeding
experiments.

No. 51, 25 each of Cerion viaregis and Cerion uva.

No. 52, 25 each of Cerion viaregis and Cerion crassilabris.

No. 53, 25 each of Cerion casablancae and Cerion uva.

No. 54, 25 each of Cerion casablancae and Cerion crassilabris.

No. 55, 25 each of Cerion uva and Cerion crassilabris.

In cage 56 there were placed 203 young of various sizes of the huge
new form collected in Middle Bight, Andros, which Doctor Bartsch
has called Cerion mayort.

Two additional species were introduced this year on Loggerhead
Key, one Cerion mayori, as above stated, and the second, Cerion
incanum, as also stated above, but of this species a large colony was

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

as

also placed about the water tower at the northern end of the island,
in order to have additional material if it should be needed for breeding
purposes in the future,

While at the Tortugas a careful bird census was made, as usual.
By the use of a blind, a series of photographs of the beautiful roseate
tern, nesting here abundantly, was secured. The accompanying illus-
tration shows one of these birds together with an unhatched egg and
a babe.

Fic. 51.—Roseate tern, young, and egg. Bush Key, Tortugas, Fla.

ENTOMOLOGICAL EXPEDITION TO ALASKA

In May, 1921, Dr. J. M. Aldrich, Associate Curator of Insects, U.S
National Museum, was detailed to collect insects in Alaska, especially
in the interior, The museum had very little material from Alaska,
except from the coast region. The government railroad, extending
from the southern coast north to Fairbanks, was nearing completion,
and offered opportunity for travel not heretofore existing. It ap-
peared also that the completion of the railroad would probably lead
to an increase of population which would create greater interest in
the insects of the region.

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 53

Doctor Aldrich left Seattle May 30. The steamship made some
stops for unloading freight, enabling him to collect one day at Skag-
way and one day at Valdez. The coast region is fairly familiar to
tourists, with its innumerable islands, steep shore-line, snow-capped
mountains and numerous glaciers (figs. 52-56). Seward was reached
on Juneg. The government railroad begins at this point and close con-
nections were made with a waiting train. The railroad passes over
rugged mountains in the Kenai peninsula close to several large
glaciers ; it then descends to sea-level at Turnagain Arm, keeping near
the shore line to Anchorage. This was the first collecting point which
might be considered to represent the fauna of the interior. Although

Fic. 52—Cannery near Juneau, Alaska.

it is on tide-water it is behind the coast range and has the dry climate
characteristic of the interior. The town is on a level glacial plain,
several miles wide, covered with a light forest and having a thin soil
upon quite recently deposited gravel. The forest is composed of
spruce, aspen, birch, alder and willow. After several days collecting
here the journey northward was resumed. Steel had been laid as far
as Hurricane, 285 miles from Seward. On arriving here Doctor
Aldrich was furnished a horse by the Alaskan Engineering Commis-
sion and rode along the right-of-way for 85 miles across Broad Pass
and down the Nenana River to Healy, which was at the time the
terminus of the rails laid southward from Nenana on the Tanana
River. Only casual collecting was done until Healy was reached, but
here it was necessary to wait several days for baggage to be brought
from Hurricane by wagon. This proved to be a very good collecting

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.. 72

point as it 1s at the mouth of the canyon on the edge of the Yukon
Valley, thus combining to some extent the mountain and plain fauna.
After five days here, Doctor Aldrich went north on the railroad to
Nenana, collected there for only part of a day and continued the fol-
lowing day on the narrow gauge line, recently acquired by the govern-
ment, to Fairbanks, his destination. It had been intended to spend

-

Fic. 53 —North side Lynn Canal near Skagway, Alaska.

Fic. 54.—Glacier on Lynn Canal, Alaska.

most of the collecting time in the vicinity of Fairbanks, but the trip
had taken much longer than expected, so he stayed only a week at
this point.

The Tanana Valley at Fairbanks is typical of the Yukon Valley in
general, as far as the species of insects are concerned. Although it
is within about 100 miles of the Arctic Circle, it has a fairly hot
summer on account of the extremely long period of sunshine in the
day. Some farms are developed and the government experiment sta-
tion has been demonstrating for many years that the usual garden

NOT LS SMITHSONIAN EXPLORATIONS, 1921 5

on

vegetables of the northern states as well as some cereals can be grown.
The aspect of the light forest is much like parts of northern Minnesota
and the regions about Lake Superior generally: the insects collected

resid’ 2S ova

Fic. 55.—Port Althorp, Alaska (merely a cannery).

Fic. 56.—Looking north from Tannel Station, Alaska. Valley filled with glacial
gravel in part very recent.

were mostly species occurring in the region named and eastward to the
Adirondacks and New England.

A return trip was made along the same route, with stops at Healy
and at some of the construction camps on the unfinished part of the

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE 72

Fic. 57.—A good insect collecting ground on Ship Creek, near Anchorage,
Alaska.

Fic. 58.—Homesteader’s cabin near Anchorage, Alaska.

un
N

INO eS SMITHSONIAN EXPLORATIONS, I921

Fic. 59.—Outskirts of Anchorage. Log houses make up almost the whole
town, and are the usual thing in Alaska.

+
i ee

Fic. 60.—Outskirts of Anchorage, Alaska. Half-cleared land.

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

road. The bad condition of the wagon road along the right-of-way
south of Healy had reduced wagon travel to a very low stage. The
only wagons using the road were those of the Alaskan Engineering

Fic. 61—Spruce forest on Chulitna River near Mount McKinley.

Fic. 62.—Hurricane, a construction camp on the government railroad 285 miles
north from Seward, Alaska.

Commission, carrying supplies to the camps. As each wagon turned
back on unloading, and only a few were in use at the time, considerable

delay was encountered in getting baggage moved from Healy to

NO. I5 SMITHSONIAN EXPLORATIONS, IQ2I 59

Hurricane. This delay could have been used to good advantage for
collecting except for the fact that the weather became cloudy and
windy and very unfavorable. Doctor Aldrich, after several days
delay, went on to Anchorage and spent a few more days collecting
there while awaiting his baggage. Here the weather was again favor-
able so that the result was very good. Resuming his journey Doctor
Aldrich went to Seward with the intention of spending at least ten
days in getting a collection of the insects of the humid coast region.
The weather, however, gradually became more rainy, greatly limiting
the result and finally making it expedient to take the boat from Seward
about a week after arrival.

Fic. 63.—Middle fork of Chulitna, a little south of Broad Pass. Corduroy
bridge of the Alaskan Engineering Commission.

The expedition resulted in the accession of about 10,000 specimens
of Alaska insects, nearly all from the interior region. As far as they
have been studied up to the present time they indicate three somewhat
distinct faunal regions in the territory covered.

First, the maritime fauna consisting of the insects living upon the
seashore and depending upon the ocean for necessary conditions of
existence. Insects of this group extend down the coast, in many cases
as far as the State of Washington and some even so far as San
Francisco; while it is presumed that they would also be found more
or less in the Asiatic side of Bering Sea.

The second element is that of the humid mountain region along the
coast; a considerable part of this fauna extends to Puget Sound,

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.

ee.

eh

Fic. 64.—Contractors’ cabins on the line of the government railroad near
Broad Pass, Alaska.

Ftc, 65.—Looking northward down the Nenana, Alaska. Unfinished govern-
ment railroad in foreground and down left side.

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 61

Mount Rainier, and in less degree, to other mountains of the Pacific
northwest. The relation of this element to the Asiatic fauna is very
little known.

The third element of the Alaska fauna, as far as observed, is that
of the dry interior and especially of the Yukon Valley, which has
many elements in common with Northern Minnesota, Wisconsin and
Michigan, Ontario, the Adirondack Mountains of New York and the
White Mountains of New Hampshire. Many of the insects of this
group also occur in the Rocky Mountains of Colorado and no doubt
further exploration will show that they occur in other mountains
of the western United States. Those which represent a more northern

Fic. 66.—Town of Healy in the lignite belt on the Nenana River, Alaska.

range also reappear in Labrador collections, and presumably extend
across North America although we have no collections from inter-
mediate points. This element contains many species known from
Finland and the Scandinavian Peninsula in Europe, presumably ex-
tending in their distribution across Russia and Siberia.

In most orders of insects Alaska has a comparatively large fauna.
There are very numerous species of the two-winged flies, or Diptera ;
and from Doctor Aldrich’s long experience with this group he natur-
ally paid especial attention to collecting in this order. Bumble bees
and wasps are conspicuous insects everywhere on flowers; and in the
absence of darkness bumble bees were observed to work as late as
10.30 at night in Fairbanks. Grasshoppers were strikingly scarce,
only two species being found and in all but half a dozen specimens.

5

62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Mosquitoes in the interior are exceedingly abundant, as is well known.
Especial attention was given to them in collecting, and two species
previously undescribed were among the material brought back. It
appears, however, that the most troublesome species are the same ones
which occur in somewhat less numbers in the Pacific northwest in
occasional favorable localities. Horse flies are very numerous in the
region at Fairbanks where they are commonly called moose flies since
the moose is more common than the horse.

ni Pet
*, <r ee “Y :
ERSTE PS

Fic. 67——Construction camp at Nenana Bridge, north of Healy, Alaska.

The common house fly was not found at any point in Alaska. Con-
tinuous attention was given to this matter, and collections were made
at the garbage dumps in Anchorage and Seward; while at Ketchican,
the southernmost town in Alaska, grocery stores, restaurants and a
cannery were carefully examined early in August without finding any
of the flies. Other garbage-feeding flies were studied at every pos-
sible point and one new species of blow-fly was collected. The absence
of several scavenger flies which are common in the United States was
noted.

The exploration of Alaska, especially the interior, from an entomo-
logical point of view is important in itself and also forms a link in the
study of a much broader problem—that of the entire Holarctic fauna
which extends almost continuously around the globe in the vicinity of

NORE SMITHSONIAN EXPLORATIONS, 1921 63

the Arctic Circle. It isa matter of great scientific interest to determine
how much of this northern fauna is the same in the new world as in
the old, and also to determine how much of the fauna further south,

Fic. 68.—Fairbanks, Alaska, and adjacent country from top of a building.

Fic. 69.—Looking up the Cheva River eastward from Fairbanks, Alaska. Some
farms cleared and cultivated on the slopes of the distant hill.

as for instance in the United States, has been derived from this
northern region. It is hoped that opportunity will arise to carry this
exploration much farther not only in Alaska, where as yet merely a
beginning has been made, but also in other northern regions as for
instance Labrador, Greenland and Siberia.

64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

ARCHEOLOGICAL FIELD-WORK ON THE MESA VERDE
NATIONAL PARK

During May and June, 1921, Dr. J. Walter Fewkes, Chief of the
Bureau of American Ethnology, continued his archeological work of
former years on the Mesa Verde National Park, Colorado, the brief
season’s field-work being financed with a small allotment from the
Bureau of American Ethnology.

The site chosen for field operations was the Mummy Lake cluster
of mounds, a typical prehistoric southwestern village situated 44 miles
north of Spruce-tree Camp. One of the mounds in this village,
excavated in 1916, is now known as Far View House. The surface
contours of the remaining mounds differ somewhat, indicating that the
buildings hidden in them have different forms, but excavations are
necessary to determine the use of these buildings. It has long been
known that some of the prehistoric pueblos of our southwest had
rooms called kivas for religious purposes, but only within the last
year has it been recognized that there was sometimes added to these
kivas a complex of rooms, also for ceremonial purposes. Several of
these specialized religious structures have already been described, but
there remain many other mysterious mounds beckoning the archeol-
ogist for excavation and accurate identification. How many different
types of buildings designed solely for ceremonials there are in our
southwest, time will reveal.

The word house (#7, Hopi) is applied in prehistoric cliff-dwellers’
nomenclature to a compact collection of inhabited rooms, secular and
religious (fig. 70). A pueblo is such a communal dwelling; but a
group of uninhabited rooms, each and all constructed for ceremonial
purposes, should bear another name. The discovery of Sun Temple
introduced archeologists to a type of southwestern buildings not in-
tended for habitations, but for a specific communal purpose supposed
to be religious. Fire Temple, on the Mesa Verde, is also regarded as
such a specialized building and is likewise believed to have had a
religious use. Similarly, Cedar-tree Tower and Far View Tower
were not habitations but communal buildings with a religious function.
The “Lower House” at Yucca House National Monument, the
“ Great Kiva” at Aztec, and similar great kivas situated in the Chaco
Canyon and elsewhere on tributaries of the San Juan River morpho-
logically belong to this type. All these may be called temples. There
are many large buildings never inhabited but now in ruins scattered
over the southwest, the use of which is doubtful. Among these are

65

1921

y

XPLORATION

E

SMITHSONIAN

NO. 15

“T ‘oay Aq ydessojoyq )

(‘peosiey spuesc oly pue Jaauaq Asajinoy ‘weag
‘yieq [PUONRN eps9A esayy ‘JUIOg UNS WoO] :a0R[eG YD pue

gduia | ung—oZ ‘o1T

66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

the so-called fire houses or “ houses ” of the Hopi fire people and the
twin mounds conspicuous on the mesa top above Sikyatki, which may
on excavation be found to have been devoted solely to religious
purposes.

rae
tS aed

Fic. 71.—Far View Tower and Kiva, partially exca-
vated. Mesa Verde National Park. (Photograph by
Fewkes. )

This specialization in the San Juan Valley of buildings showing
functional differentiation in structure is indicative of a high cultural
development. It is instructive to find that it is confined to prehistoric
stages of development and is most abundant in areas where sedentary
inhabitants had disappeared before the advent of Europeans.

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 67

The plan of the work of the Bureau on the Mesa Verde National
Park in 1921 was to investigate a conspicuous and centrally placed
mound not far from Far View House. The indications are that this
was an ancient necropolis of the Mummy Lake Village, combining
subterranean rooms or kivas with a large cemetery situated on the
southern side of a high tower. Unfortunately, this cemetery had been
rifled several years ago by vandals; but the many fragments of pottery
found on the surface betray features important in cultural comparisons.

Far View Tower was relatively an ancient building ; its architectural
form is characteristic and its pottery decidedly archaic as compared
with that of the golden epoch of geometric decoration from Cliff
Palace or Spruce-tree House. We may never know in what century
this tower. was built, but its construction can be referred to an older
epoch than the great cliff dwellings of the park, which were probably
inhabited as late as 1300 A. D. The refuse heaps of cliff houses
have so little depth that a stratification or superposition of pottery
shards is too small to afford satisfactory evidence of long occupancy.
In historic refuse heaps of pueblos now inhabited they are thicker and
the stratification method has proved advantageous; but nothing that
was not already known has been added to our knowledge of the
sequence of prehistoric pottery of cliff houses by this method of study.
No Mesa Verde refuse mound has yet shown any difference in the
character of pottery found on its surface and at its base. The pottery
fragments of mounds containing relics of earth lodges are as a rule
cruder than others. The pottery from the cemetery or necropolis of
Far View Tower is rudely decorated ware, while that from Far View
House is finer, but not as well made as that from Spruce-tree House.
It is probably older than the pottery from Far View House, but both
are more ancient than the pottery from Spruce-tree House.

Far View Tower (fig. 71), like Cedar-tree Tower, has one and pos-
sibly more subterranean rooms or kivas on the south side, but the
latter lacks the large cemetery. The use to which Far View Tower
was put and the significance of the relation of the accompanying kivas
to it were probably not very different from those at Cedar-tree
Tower, discovered last year (1920). Evidently the complex was
devoted to some archaic cult, like fire worship.

In addition to the work above mentioned, Doctor Fewkes also
excavated Painted Kiva House, a small prehistoric cliff dwelling
situated on the Mesa Verde a short distance north of Cedar-tree
Tower, under the rim of the west side of Soda Canyon. This ruin
was excavated and described by Baron Nordenskiold, who called it
Ruin g. It contains remains of two well-made kivas of the regular

68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

circular Mesa Verde type and of several granaries and living rooms.
The approaches to it from the mesa rim are very precipitous-and it
was necessary to construct four ladders and otherwise improve the
trail to enable visitors to see it.

On the walls of one of its two kivas there survives a very good
example of decorated plastering. As shown in the accompanying
illustration (fig. 72) there is a dado or lower part of the kiva wall
which is painted red, and on its upper edge there are arranged at
intervals clusters of triangular symbols (three in number) around
which extends a row of dots. The Hopi identify these triangles as

Fic. 72.—Interior of kiva, showing mural decoration, niches, and_ pilaster.
Painted Kiva House, Mesa Verde National Park. (Photograph by Fewkes. )

symbols of butterflies. They are of common occurrence on the walls
of several kivas and survive in certain secular rooms of the cliff
dwellers. These triangles with surrounding dots occur constantly on
the oldest cliff-dweller pottery, as shown in the accompanying figures.
The ventilator shaft is represented in the painted kiva by a tortuous
passage, extending under walls and opening some distance from the
room. It is spacious enough to serve as an entrance into the cere-
monial chamber. Although Baron Nordenskidld made extensive ex-
cavations in Painted Kiva House and devoted several pages of his
memoir to a description of it and the specimens he found there, many
objects (fig. 73) remained in rear chambers which were found in
192i,

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 69

eal

Fic. 73—Snowshoe frame. Painted Kiva House, Mesa Verde National Park.
Size: 14% inches by 9% inches. (Drawn by Mrs. George Mullett.)

L sal

Fic. 74—Rim basket. Painted Kiva House, Mesa Verde National Park.
Size: 15% inches. (Drawn by Mrs. George Mullett.)

7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Among the instructive specimens collected in Painted Kiva House
should be mentioned a rim basket (fig. 74) and a woven headstrap of
yucca fiber. The unique object shown in figure 75 reminds one

Fic. 75.—Unidentified object. Painted Kiva House,
Mesa Verde National Park. Size: 534 inches long.
(Photograph by De Lancey Gill.)

of Navaho “ bugaboos,” sometimes found farther down the San Juan
but not yet recorded from Mesa Verde. In a rear room which gave
every evidence of having been a granary or bin for storage there were
found numerous ears of corn with kernels entire, beans, and squash
seeds. The belief is widespread that cliff-dweller seed corn when

NO. 15 SMITHSONIAN EXPLORATIONS, I9Q2I 71

planted will germinate, but all experiments in that direction have
failed. There is no hope that any greater success will reward experi-
ments made with corn from this granary. In the centuries that have
elapsed since the mesa was deserted, corn seed left behind has lost
its vitality.

The walls of a ruin called Mummy House, situated almost directly
under Sun Temple, are among the most carefully constructed on the
park. This ruin has one kiva which was cleaned out but not repaired.
A mummy (now in the Mesa Verde Park Museum) was found in this
ruin several years ago. Above it is Willow-tree House, practically
inaccessible. Ladders were put in place connecting the trail up the
canyon with Mummy House. A typical form of cliff house called
Oak-tree House, before and after repair, is shown in figures 76 and 77.

One of the important ruins on the Mesa Verde, called Step House
by Nordenskiold, is situated in a cave 5 miles west of Spruce-tree
Camp. It presents to the archeologist one of the most instructive prob-
lems on the Mesa, and should be put in shape for visitors. In the floor
of this cave, which has been considerably dug over by Nordenskidld
and others, there was material bearing on a most interesting chrono-
logical problem, viz., the age of the cliff houses; for the artifacts in
this place represent two different epochs in the cultural history of the
pure pueblo-cliff-dwelling type. Out of the floor of the cave there ~
projects the edges of upright slabs of stone showing the existence of
cists like those in Earth Lodge A. These suggest the slab-house cul-
ture; but at the other end is a building in the highest form of hori-
zontal masonry. The probability is that the former is the older con-
struction or that it was built by the most ancient people of the park,
who lived and were buried in that end of the cave, designated by
Nordenskiold a cemetery. Here we have evidences, both architectural
and ceramic, of former earth lodges or fragile walled buildings of the
prepuebloan or archaic culture. The original dwelling built by
people when they moved into Step House Cave was an earth lodge,
and the dwelling with horizontal masonry and kivas, at the other
end of the cavern, was a later development. The pottery of the
former is more archaic than that of the latter. Figure 78 illustrates
the most highly developed Mesa Verde pottery. We have, in other
words, indications of two distinct stages of development in Step
House Cave—one the earth lodge and the other the pure pueblo or
kiva style; the former or earth construction situated at one end of the
cave, the latter stage at the other. This evidence of two stages of

(‘peorrey spurs‘
Oly pue JaAUIqd Asojinoy ‘“wresag “TJ ‘09*) Aq ydeisojoyd ) ‘redat a10joq ‘asnoFy 9913-yeO—9Z ‘DLA

VOL. 72

NS

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SMITHSONIAN

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1921

XPLORATIONS,

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SMITHSONIAN

NO. 15

I9AUAC{

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yeO JO woos spppryy—ZZ ‘oy

74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

development in the same cave is derived from both ceramic and archi-
tectural studies. The indications are that after the earth-lodge con-
dition was outgrown the floor of the cave where the evidence occurs
was used as a cemetery, and the survivors constructed their new
homes at the other end of the cave in the form of cliff houses,
Although no satisfactory scheme of the chronological sequence of
different types of Mesa Verde pottery has been worked out, it is
most important to pay some attention to its bearing on the age of the
above-mentioned buildings.

The mortuary pottery (fig. 79) from the Far View Tower cemetery
belongs to a primitive type quite unlike any yet recorded from Mesa
Verde cliff dwellings. The most exceptional features are the numer-
ous varieties of coiled, corrugated, undecorated ware. Figure 8o,
restored froma fragment, and figure 81 show one of these exceptional
bowls. 'A-similar bowl with a blackened inner surface occurs else-
where in the southwest, as on the Little Colorado, but has never been
described from the Mesa Verde. A comparison of ceramic objects
from the cemetery of Far View Tower (fig. 82) indicates it belongs
to an ancient type related to Earth Lodge A, described in the explora-
tions pamphlet for 1919.. Attempts have been made to show an
architectural evolution from an earth lodge with roof and walls of
logs and mud into buildings constructed of well-laid horizontal stone
masonry. There is a chronological development in technique, form
and decoration of pottery from the simple to the complex, but those
who have studied cliff-house pottery have not yet succeeded in arrang-
ing the different kinds in chronological sequence.

Each ceramic area in our southwest has its distinct facies. Mesa
Verde pottery excels all others in its geometrical decoration. Con-
ventionalized designs and life figures on it are few in number and
crude in execution, but linear designs are abundant and varied. In
the prehistoric Hopi pottery, where there are few life figures and the
majority of designs are geometric or highly conventionalized, there is
nothing showing successive steps in the development of designs. In
those ruins where geometric figures (fig. 83) predominate there is
little to show their evolution. The pottery from the Mimbres Valley,
New Mexico, decorated with both fine geometric and realistic figures,
gives us no clue to evolution of different typical naturalistic designs.
Apparently the three types, geometric, conventional, and realistic, are
distinct from their very origin and it is difficult to prove that one type

1Smithsonian Misc. Coll., Vol. 72, No. 1.

NO. 15 SMITHSONIAN EXPLORATIONS, I921 75

Fic. 78.—Mug ; black on white ware. Fire Temple House, Mesa Verde National
Park. Size: 4 by 4inches. (Photograph by De Lancey Gill.)

L

Fic. 79.—Archaic black on white ware; coarse decoration, Far View House
Village. Necropolis, Mesa Verde National Park. Size: 5% inches.

76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 80.—Bowl; indented corrugated ware with black interior, Far View
House Village. Necropolis, Mesa Verde National Park. Size: 4% by 2%
inches. (Repaired by W. H. Egberts.)

Fic. 81.—Detail of indented corrugated bowl, figure 80, Far View House
Village. Necropolis, Mesa Verde National Park. (Drawn by Mrs. George
Mullett.)

NO. 15 SMITHSONIAN EXPLORATIONS, I921 77.

preceded another in evolution. For the present, then, our knowledge
of sequence of types of pottery is largely derived from descriptions
and not generalizations. But our archeological method permits us to
determine the main features of a stage of culture among the Indians
of which little is historically known. For instance, previous to the
year 1915 we were ignorant of the manners and customs of the people

Fic. 82.—Archaic black and white ware, coarse decoration,
Far View House Village. Necropolis, Mesa Verde National
Park. Size: 3% by 3 inches. (Photograph by De Lancey
Gill.)

who inhabited the Mimbres Valley, New Mexico. Documentary his-
tory is silent about them. Through archeological studies data are
being brought to light year by year by which our knowledge of these
Indians is greatly advanced. Pictures on ancient pottery often impart
more information than written descriptions and are most important in
the study of lost races. During the last few years Mr. E. D. Osborn,
of Deming, New Mexico, has from time to time sent to the bureau
many unique photographs of mortuary bowls (figs. 84-86), some of

6

78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. yi

Fic. 83.—Decorated pottery from Mesa Verde National Park. a, trian-
gular geometric design; b, hatched and terraced line; c, black triangles in
concentric series; d, central triangle with curved lines at angles; e, unknown
geometric design; f, S-shaped design. (Drawn from photograph, by Mrs
George Mullett.)

NO. 15 SMITHSONIAN EXPLORATIONS, 1921

Fic. 84.—Decorated pottery from Mimbres Valley, New Mexico, Osborn
Collection. a, bird trap; b, gambling game; c, emergence of man from
lower world; d, white outline on black ground; e, two fishes; f, two negative
pictures of fishes. (Drawn from photograph, by Mrs. George Mullett.)

80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

WSS!

b\ Saeed

eee
i

-- --"

~

Fic. 85.—Decorated pottery from Mimbres Valley, New Mexico, Osborn
Collection. a, unknown bird, from back, with outstretched wings; b, feath-
ers used in geometric decoration; c, three-headed turkey; d, parrot, head
often repeated as club-shaped design on Casas Grandes pottery; ¢, humming
birds and flowers; f, unidentified flowers. (Drawn from photograph, by
Mrs. George Mullett.)

NO! 15 SMITHSONIAN EXPLORATIONS, I92I SI

Fic. 86.—Decorated pottery from Mimbres Valley, New Mexico, Osborn
Collection. a, unknown fish with feathered horn; b, animal heads like
swastika: c, sun with four tail feathers; d, geometric ornaments; e, geo-
metric ornaments; f, geometric ornaments. (Drawn from photograph, by
Mrs. George Mullett.)

82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

which are decorated with well-made pictures showing hitherto un-
known features of prehistoric life in that valley. Similar pictures
have been reproduced in former reports, but several specimens lately
discovered are the most instructive yet found. References to a few of
these close this account.

The food bowl (fig. 84a) apparently represents a hunter snaring
birds. He carries three nooses in his hand and in three of the snares
that are set are birds, while a fourth is empty. On the opposite side
of the bowl there are two other birds that possibly have been captured
earlier.

Figure 84b represents a prehistoric game of “ stick dice.’ In this
design three of the “canes” or dice are represented in a rectangular
enclosure around which are seated the players. The stakes are arrows
shown in a receptacle deposited above the picture.

Two fishes shown in figure 84e call to mind the unusual method of
representing certain life figures, men, birds, and other animals, on
other pieces of pottery. The background of the two fishes of figure 84f
is black, the bodies white; a negative picture common on ware from
Casas Grandes, Mexico, and peculiar to the inland basin in which the
Mimbres lies. The upper beak and eye of the head of the well-drawn
parrot is shown in figure 85d. This conventionalized head often
occurs without the body of a bird or any realistic likeness to a parrot
in the decoration of pottery from Casas Grandes and it is interesting to
note in this connection that Mr. Osborn claims to have found a mound
a few miles from Deming, New Mexico, in which the pottery is
practically the same as the well-known Casas Grandes ware.

The body of the animal represented in figure 86a is serpentine, but
the shape of the head and the possession of fins suggest a water
monster. The horn with a cluster of feathers occurs in a similar
painting without fins, and may be a representation of the Horned or
Plumed Serpent.

As is true of decorations on prehistoric Hopi ware, the feather
is sometimes used as a decorative element. The identification of the
use of this motive was made by a comparison of the undoubted bird
with outstretched wings and well-marked symbolic wing feathers
shown in figure 85a, and the existence of four clusters of a like design
in figure 85>. A study of over a hundred decorations, realistic, con-
ventional and geometrical, taken from Mimbres pottery indicates that
this lost people of southern New Mexico had reached a very high
stage of ceramic decoration. There is evidence that this art was
somewhat influenced from outside but mainly developed where it was

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 83

found. It is one of several localized culture areas related to but not
necessarily belonging to the pueblo with which it has affinity. It is
most closely affiliated with that of Casas Grandes and the southern
part of the plateau in which it lies. The environment of this plateau
is Mexican, climatically speaking, and the culture will probably be
found to correspond. While superior to the Casas Grandes and all
other prehistoric Indian pottery in variety and the accuracy with
which human and animal figures are drawn, it shares enough with
it to hold a place in the same group.

ARCHEOLOGICAL COLLECTING IN THE DOMINICAN
REPUBEIC

While engaged in a biological exploration of this republic in 1921
and previous years, Dr. W. L. Abbott of Philadelphia incidentally
made a collection of aboriginal Indian antiquities on the north coast,
especially around Samana Bay and the region between it and Puerto
Plata, as well as in other parts of the island. No systematic excava-
tions were attempted ; the majority of the specimens were either pur-
chased or otherwise obtained. The localities where individual speci-
mens were said to have been found are mentioned in the legends under
the illustrations. This accession contains many specimens, one or two
of which merit special notice, even if it anticipates a final report.

There is in this collection an exceptionally good water jar of unique
form upon the neck of which are incised rude figures of animal or
human heads. The body of this jar (fig. 87), instead of being round
is roughly four-sided, its base flat, neck constructed bottle shaped.
Another bowl (fig. 88), spherical in form, is also unique and the
incised figure covers much of the upper surface.

In the collections of every West Indian archeologist there are speci-
mens of burnt clay heads called “zemis” (idols) by the natives.
These objects are not idols but broken handles of bowls, portions of
which sometimes adhere to them. As broken specimens they teach
very little, but if the jar from which they were broken be restored
they become instructive. The results of Mr. Egbert’s clever recon-
struction of the bowls to which three of the handles belong are shown
in figure 80, a, b, and c.

The decoration of Santo Domingo pottery, like that from prehistoric
Porto Rico, as a rule is limited to handles or lugs of bowls and vases.
These heads are attached to the rims of jars or bowls and give us a
means of classification. They fall naturally into three distinct types:
First, and most common (fig. 89a), those where the handles are oppo-
site each other, the handle represented as looking into the bowl;

84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

second, a less common type, those with handle faces looking outward;
and third (fig. 89c), rarest of all, those with human or animal heads
attached to the rim by the back of the head or lying along the rim

Fic. 87.—Unique vase. Cueva de Roma, Dominican Republic. Size: 834
inches. U. S. Nat. Mus. No. 316445.

of the bowl with their axis parallel to it. Santo Domingo pottery as a
rule is a coarse biscuit ware, its surface waterworn but smooth, ap-
parently sometimes formerly covered with a red slip, showing, how-
ever, no evidence of a glaze. Although in bolder relief than that made
by the prehistoric potters who preceded the Carib in the Lesser

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 85

Antilles, the ceramics of the aborigines of the Greater Antilles are
more closely related to the work of the Huaxtecs of Mexico than to
that of the aborigines of South America.

There are in the Abbott collection representatives of all types of
those Antillean idols characterized as three-pointed stones: one (figs.
go and 91) with head on the anterior point; another (fig. 92), a
second type characterized by a head on the side of the cone; a third

k

cna

Bees

Fic. 88—Globular bowl of thin ware. Locality, Yaqui del Norte, Dominican
Republic. Size: 534 inches. U.S. Nat. Mus. No. 293016.

type has the cone modified into a head; and lastly one smooth, undec-
orated specimen, referred to a fourth type. The specimen represented
in figure 93 belongs to the first type and has on each side of the base
of the cone two shallow circular pits; each of these pits represents a
joint of the fore and hind limbs, both of which are cut in relief on the
side. Although similar pairs of pits are known on several specimens
and accompanying forelegs or arms sometimes appear in relief, no
specimen with two pits both having relief representations of limbs
has been recorded.

86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES 72

GC.

Fic. 89.—Restored pottery from shards collected in the Dominican Republic,
by Dr. W. L. Abbott. Restoration by W. H. Egberts. a, food bowl with effigies
on rim, facing inward; b, effigy bowl with handles in form of heads, facing
upward; c, food bowl, handles in form of heads transversely placed on rim.
Size: a, 9% inches; b, 6% inches; c, 1634 inches. U.S. Nat. Mus. No. 316454.

NOX 15 SMITHSONIAN EXPLORATIONS, IQ21 87

<bean laa nati

‘

Fic. 90.—Three-pointed stone of first type, from side. Constanza, Dominican
Republic. Size, 534 inches x 2% inches. U.S. Nat. Mus. No. 309536.

Fic. 92.—Three-pointed stone of second type, from side. Constanza, Dominican
Republic. Size: 2% inches x 1% inches. U.S. Nat. Mus. No. 309537.

VOLA72

MISCELLANEOUS COLLECTIONS

SMITHSONIAN

88

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a

yuo1y— £6 “91g

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 89

There are only seven known specimens of three-pointed stones of
the second group, and the U. S. National Museum now has five of
these, one of which we owe to Doctor Abbott.

He has also added to the museum collection the three especially fine
Antillean amulets shown in fgure 94. The form of one—that figured
in the middle—is unique. These objects are supposed to have been

ow 7

a. v. Ge

Fic. 94.—Three marble amulets. Locality, Guayubin, Yaqui River, Domin-
ican Republic. a, 2% inches, U. S. Nat. Mus. No. 316448; b, 4% inches, No.
316446; c, 2x6 inches, No. 316447.

used as fetishes and to have been tied to the foreheads of warriors
when they went into battle, as described by Gomara and other early
writers.

The cylindrical object of clay with incised figure shown in figure 95
belongs to a type concerning the use of which there has been con-
siderable discussion. These specimens have been identified as rollers
for stamping pottery with the design incised on their surfaces ; but if

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VOL. 72

SMITHSONIAN MISCELLANEOUS COLLECTIONS

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we judge from the similar objects of aborigines of Venezuela they
were more probably used for stamping fabrics or even for printing
certain totemistic or other designs on the face or body.

There is in the Abbott collection an artificially worked stone (fig.
96), about a foot in length, which appears to have been used as a

Fic. 98.—Stone Cassava grinder. Yaqui del
Norte, Jarabacoa. Size: 1234 inches x 19% inches.
U. S. Nat. Mus. No. 292908.
baton, possibly a badge of office. One end bears incised designs
representing eyes and mouth suggesting a human head.

Figure 97 resembles outwardly a pestle, but a closer examination
shows that it is made of clay, a material impossible for an effective
grinding implement. It has many pits on the under surface (shown
in the figure) which suggests that it was functionally like the cylinder
above mentioned used for imprinting paint patterns on the human
body or woven fabrics.

Q2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

One of several flat stone objects collected by Doctor Abbott having
extensions, two “ handles” on the rim, is shown in figure 98. In
shape and especially in the form and position of the handles these stone
implements resemble graters—generally of wood—specimens of which
are still in use in Haiti. Stone graters are novelties and those col-
lected by Doctor Abbott are the first of this material added to the
museum. It is probable that the surface of this stone was formerly
covered with some kind of matrix in which were set sharp stones
arranged in an ornamental design that has now completely disap-
peared, leaving no trace of its former presence.

All the above-mentioned specimens are referred to the Tainan or
most advanced neolithic culture of the West Indies, that originated
and flourished in the Haiti-Santo Domingo and Porto Rico areas in
prehistoric times. The three-pointed idols, stone collars, elbow stones,
and characteristic pottery separate the Porto Rico Tainan from that
of Jamaica, eastern Cuba, and the Bahamas, which belong to another
closely related culture that may be called Cuban Tainan.

The pottery of the aborigines of the Lesser Antilles belongs to an
allied prehistoric Tainan culture that was submerged by the Caribs,
who inhabited these islands when discovered by Europeans, at the
close of the 15th century. The fine addition that Doctor Abbott has
made to our West Indian collection all belongs to the true Tainan cul-
ture which reached its highest development in Espanola and Porto
Rico.

The archeological specimens from the West Indies presented to the
museum by Doctor Abbott are very valuable and as time goes on will
be more and more appreciated by students of the history of man in
the Antilles.

ARCHEOLOGICAL RECONNAISSANCE OF THE CAHOKIA AND
RELATED MOUND GROUPS

David I. Bushnell, Jr., collaborator of the Bureau of American
Ethnology, conducted during 1921 a reconnaissance of the remarkable
mound groups in the vicinity of the great Cahokia Mound. The in-
formation secured at this time, added to notes made during frequent
visits in the past, has been used in preparing the following sketch of
the interesting region.

It is quite evident that long before Pére Marquette discovered
and passed the mouth of the Missouri, during his journey down
the Mississippi early in the summer. of 1673, the region immedi-
ately below the confluence of the two great streams had been an
important center, a gathering place, of the native inhabitants of the

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 93
valley. Mound groups, village sites, and burial places remain to

indicate the presence of a numerous people before the coming of
Europeans, and the innumerable objects of native origin encountered

WORTH GROUP-

5 b ie
5 Lie . es ‘SCALE IN MILES-

Fic. 99.—Map showing location of mound groups.

in the region bear evidence of their skill in working the available
materials.

Immediately below the mouth of the Missouri, on the left or
Illinois bank of the Mississippi, the river bluffs become more distant
from the stream and consequently the lowlands are in some places
6 or 8 miles in width from east to west. Shallow lakes covered much

7

904 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

of the surface and some parts were heavily timbered. As indicated
on the accompanying map, figure 99, four mound groups stood in the
lowlands east of the Mississippi and a fifth was on the opposite bank,
land now covered by the city of St. Louis. And as is shown on the
map the five groups were placed with a certain degree of order to one
another, with the great mound, Cahokia, rising near the center of the
area.

But who were the builders of the mounds, the most important
groups in the Mississippi Valley? The question may never be
definitely answered although it is more than probable they should
be attributed to a tribe or tribes known in historic times but who may
have become greatly reduced in numbers and relative importance
before the coming of the French. Evidently the historic Algonquian
tribes did not reach the eastern bank of the Mississippi until about
the beginning of the seventeenth century, and it is doubtful if others
of this linguistic family had preceded them. Siouan tribes when moy-
ing from the eastward may have traversed the region, but there is no
reason whatsoever to attribute the great mound groups which form
the subject of this sketch to either the Algonquian or Siouan tribes.
The works were probably raised by a southern tribe, a southern people
who at some time before the arrival of the Algonquian tribes, or
the migration of the Siouan tribes from the eastward, occupied the
region, later to move elsewhere, possibly to return southward. These
may have been the ancient Natchez, the Chickasaw, or some other
Muskhogean tribe of whom we possess no historic record ; however,
a careful examination of the mode of construction and the contents
of one or more of the mounds may enable us to arrive at some con-
clusions regarding their origin.

The great Cahokia Mound which rises from the level alluvial plain
near the center of the area, is somewhat less than 6 miles east of the
Mississippi and 10 miles east of south of the mouth of the Missouri.
It is a truncated pyramid, of rectangular form, with a broad terrace
extending from the south side which continues in a graded way or
approach. The sides of the work face the cardinal points, as do those
of the lesser rectangular mounds of the group. Its maximum eleva-
tion is about 100 feet. Its extreme length including the approach is
1,080 feet, and its width from east to west is 710 feet. The base
covers an area of approximately 16 acres. Viewed from the east, as
in figure 100, it appears quite regular in outline and is clearly defined
from base to summit. A small conical mound formerly stood on the

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96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

upper plateau near the southeastern corner but it was removed many
years ago. The northwestern portion of the great mound is deeply
gullied and very irregular in contour ; it 1s a question whether this part
of the structure was ever completed.

Cahokia is the largest earthwork in the United States and one of
the most remarkable monuments left by the native tribes. Fortunately
it remains in its original condition, practically untouched since the
coming of Europeans, and in this condition it should be preserved.
With each succeeding generation, as the lesser mounds and other
earthworks disappear by reason of the cultivation of the soil or the
requirement of the land for other purposes, this great terraced work
is destined to become of greater popular interest and immediate steps
should be taken to make certain its preservation.

The several groups, as indicated on the map, may now be described
in detail.

NORTH GROUP

Eleven mounds constitute this group which stands on the north
side of Long Lake, near the station of Mitchell. They are about
three and one-half miles east of the Mississippi, nearly midway across
the lowlands and some seven miles west of north of Cahokia. When
the group was surveyed March 13, 1900, it was not possible to deter-
mine the original shape of several of the mounds. The land had been
cultivated for many years and this, with the constant washing and
wearing away of the surface, had caused the works to assume an
entirely different appearance from their original condition.

The largest mound of this group stood apart and to the west of the
main cluster. It was practically destroyed years ago at the time of
the construction of two railroads which pass through it, but parts of
the work may now be traced between and on either side of the tracks.
Many remarkable objects of stone and copper were recovered during
the destruction of the structure.

As is shown on the map the large mound stood to the west. The
mound nearest it on the east, as determined by the survey of r1go0,
was 1,200 feet distant and at that time had a maximum elevation of
9.3 feet above the plain, and was of circular form with a diameter of
approximately 237 feet. Eastward from this mound are other units
of the group. The highest mound of the group at that time measured
10.4 feet, but undoubtedly the large work to the west was originally
much higher than any now standing.

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 Q7

South of the lake, away from the main group, is a single, isolated
mound. Others may have stood within the area, all traces of which
have disappeared.

CENTRAL OR CAHOKIA GROUP

Surrounding the great Cahokia Mound, which has already been
briefly described, were many lesser works, about seventy in number,
some of which were more than 4o feet in height. Some were rec-
tangular, others were circular and although at first glance they appear
to have been placed without definite order, nevertheless it is quite
evident that in several instances they were so arranged as to create
inclosed areas, thus conforming with the position of the mounds of
the three lesser groups to the north, west, and south of the central
cluster.

Unfortunately the large majority of the mounds east and west of the
great central structure have been much reduced and modified by the
plow, while several have been practically destroyed and a slight rise 1s
all that remains to indicate their position. The inclosure formed of
the smaller mounds on the east is clearly defined and gives the impres-
sion of having been intentionally planned and arranged, but for what
purpose may never be determined. And although many of the lesser
mounds have thus lost their original form and appearance, Cahokia
remains the most important and impressive native work in the Valley
of the Mississippi. As the great mound now stands it should be pre-
served: to permit its destruction would be a calamity, an irrepar-
able loss to future generations.

The rectangular work immediately southwest of Cahokia was
occupied from 1810 until 1813 by a small body of Trappist monks,
during which time their garden was on the southern terrace of the
great mound. According to the survey of 1875-1876 from which all
measurements now given are derived, this lesser mound was 25 feet
in height, its base line from north to south was 180 feet and from
east to west 200 feet. Just south of this is a small circular work. A
short distance east of south of the latter stands a conical mound which
rises 44 feet above the plain, having a diameter at base of 150 feet.
Immediately east of this is a rather irregular mound 46 feet in height,
and possibly other units of this remarkable cluster were even higher
and more extensive than these. A rectangular mound southeast of
the preceding was, according to the survey mentioned, 40 feet in
height, with its base extending 300 feet from north to south and 250
feet from east to west. This reference to several of the lesser works

98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

4

Fic. 101.—Airplane photograph showing Cahokia in upper right center.
Mounds to the south and southwest are also defined, likewise the country
northward. Camera pointed west of north.

Pile? Se Lae
Fra, 102—Airplane photograph showing Cahokia in the unner left corner.

The rectangular mound in the center of the picture, just south of Cahokia,

rises 46 feet above the original surface. A light snow covers the ground.

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 99

eit eT

Fic. 103.—Airplane photograph showing mound north of Cahokia, partly
removed. Camera pointed west.

tes) ee

Fic. 104—Airplane photograph showing mound about 1% miles west of
Cahokia. One of the most perfect of the group, and probably quite similar in
appearance to the large mound of the St. Louis group which was removed in
1869. Camera pointed northeast.

100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

will serve to convey an idea of the magnitude of the group as a
whole ; the most important prehistoric site in the entire valley.

It is of interest to be able to reproduce at this time four aerial
pictures of units of the Cahokia group, and these are believed to be
the first photographs of American mounds or earthworks to be taken
from the air. The negatives, with others, were made during the winter
of 1921 and 1922 by Lieut. Harold R. Wells and Lieut. Ashley C.
McKinley, stationed at Scott Field, Belleville, Illinois, under instruc-
tions of Major Frank M. Kennedy.

Unfortunately, weather conditions during the winter were not
favorable for aerial photography, and although many attempts were
made ground haze and smoke interfered greatly with the work. As
Major Kennedy wrote in part February 6, 1922, after mentioning the
mines and factories in the vicinity of the mounds: “‘ These activities
produce a large amount of smoke which seems to settle near the
ground and form a blanket two or three hundred feet thick.” Never-
theless the four pictures are shown to record the first attempt to
photograph mounds from an airplane.

On the summit of the bluffs northeast of Cahokia, as indicated on
the map, are two mounds of great interest which command a wide
view of the lowlands extending to the Mississippi, and beyond. Both
are of conical form and rise 30 feet or more above the original surface.
One, as it appears from the foot of the bluff, is shown in figure 105.

A view of the bluffs, with the beginning of the lowlands which
slope westward to the bank of the Mississippi, is reproduced in
figure 106. This is looking northward from a point southeast of
Cahokia.

Extending from the main group which surrounded the great mound,
in a direction south of west and following a slight ridge, is a chain of
works which terminated in an irregular group of smaller mounds near
the bank of the Mississippi. It is to be regretted that all units of
this group have now disappeared.

WEST OR ST. LOUIS GROUP

There formerly stood on the right, or west bank of the Mississippi,
on the summit of the high bluff within the limits of the present city
of St. Louis, an interesting group of mounds, twenty-seven or more in
number. <All have now disappeared but fortunately their positions
were indicated on early maps of the city.

One of the earliest as well as most detailed descriptions of the
mounds was that prepared by members of the Long Expedition,
more than a century ago. At that time they stood north of the settled

NO. 15 SMITHSONIAN EXPLORATIONS, I92I IOI

Fic. 105 —Conical mound on summit of the bluff northeast of Cahokia.

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Fic. 106.—Looking northward from near the road leading to Belleville,
showing the eastern border of the lowlands which extend westward to the
Mississippi.

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VOL.

MISCELLANEOUS COLLECTIONS

SMITHSONIAN

102

NOT LS SMITHSONIAN EXPLORATIONS, 1921 103

portion of the town and were in their primitive condition, but soon
the settlement was to extend northward and the mounds were destined
to be leveled. A view of St. Louis from the east, taken from the
Illinois bank of the Mississippi during the year 1840, is reproduced in
figure 107. Far to the north of the principal structures of the town,
on the extreme right of the picture, stands the large detached mound.
The main group was below, probably near the middle of the picture.
The large isolated work was more than 1,400 feet north of the main
cluster which formed an inclosure, thus conforming with the arrange-
ment of the mounds on the opposite side of the river. This most
important work was of oval form, with the maximum diameter of its
base, from north to south, 319 feet, and from east to west 158 feet.
The dimensions of the summit plateau were 139 feet and 11 feet.
Height 34 feet. On the eastern side, facing the river, was a terrace
resembling that on the south side of Cahokia, which was 79 feet from
east to west and probably extended the entire length of the structure.
At the time of the destruction of the great mound in the year 1869
a most remarkable cavity was discovered within it. This was a burial
chamber which could be traced for a distance of 70 feet and part had
previously been removed. It had probably been constructed of logs
over which the mass of earth had been deposited and shaped. Within
were encountered human remains in the last stages of decay, and
associated with these were vast quantities of shell beads and other
objects. This was truly a remarkable structure and one which should
have been preserved, but unfortunately it shared the fate of the lesser
mounds of the group, all traces of which have now disappeared.

SOUTH GROUP

The southern part of the American Bottom—a name long applied
to the lowlands occupied by the ancient works mentioned in this
sketch—across the Mississippi from Jefferson Barracks, becomes
quite narrow, the bluffs approach the river and are, in some places, a
scant mile from the low marshy ground which was formerly covered
with water the greater part of the year. But the land extending along
the foot of the bluffs at this point was evidently at one time occupied
by a village of some importance which stood in the midst of a group
of mounds. This may be designated the south group and in some
respects resembles the north or Mitchell group, already described.

The site was visited by the writer during the latter part of October,
1921, at which time a plan of the group was made, this now being
included on the general map. As is indicated there are now seven

TO4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

mounds standing on the lowland and one, a large conical structure,
on the bluff to the east. It is said that until a few years ago, at the
time of the construction of several railroad embankments, five mounds
extended in a row southward from the one now remaining nearest
the bluffs, consequently these, together with the five now remaining,
formed an inclosure quite similar to the north group. Northward
from the main cluster or inclosure, are two detached mounds, both
large and prominent. The group as a whole and as it originally stood,
must have been as interesting and imposing as either the north or

Fic. 108.— Village site and mounds at Bixby, with bluffs beyond.

west groups as already described, and all were probably of equal
importance to their builders.

Unfortunately, the majority of the remaining units of the group
have been greatly reduced and modified by the plow and consequently
it is not possible to determine their original size or form. However, it
is evident the second mound from the south, on the west near the
Mississippi, was rectangular and quite large. It appears to have been
oriented with its sides facing the cardinal points, as were the units of
the other groups, including the great mound. At the present time
it is worn down by long-continued cultivation and now measures about
12 feet in height, with a diameter of 200 feet. A photograph looking
eastward from the summit of this work is reproduced in figure 108.

INO ans SMITHSONIAN EXPLORATIONS, I921 105

This is a view over the plain once occupied by a native village and
shows the bluffs in the distance. Fragmentary pottery and objects of
stone are now found scattered over the intervening ground.

As has been mentioned, and as is shown on the map, a conical mound
stands on the bluff just east of the main group. It is not on the
highest point, not on the summit, but on a commanding spot visible
from miles away, north and south, and from far westward across the
Mississippi. It is on the bluff in the exact middle of figure 108, and a
closer view, taken from the south, is shown in figure 109. This
resembles the two mounds on the bluffs northeast of Cahokia and
is of equal interest.

Fic. t09.—Conical mound on bluff east of Bixby.

No other area of equal size in the entire valley of the Mississippi
appears to have been of so great importance to the native tribes as that
mentioned in this sketch. Here they reared their greatest monument,
Cahokia, and surrounded it with many lesser works. The several
distinct clusters should be considered units of a greater group, in
which the massive terraced work stood as the central structure. This
was the gathering place of a numerous people, but when or whence
they came can never be known. Now, two and one-half centuries
after the region was first entered by the French, at which time Illinois
tribes were occupying small villages near the banks of the Mississippi,
the majority of the ancient mounds have disappeared, but Cahokia
remains and it should ever stand. It must be saved as have the
pyramids of Egypt; a monument of another race whose origin is
shrouded in mystery.

100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 72

ARCHEOLOGICAL INVESTIGATIONS AT PUEBLO BONITO,
NEW MEXICO

Mr. Neil M. Judd, curator of American archeology, U. S. National
Museum, began work during the year on a five-year archeological
project undertaken by the National Geographic Society, mentioned
in the Smithsonian Exploration Pamphlet for 1920,’ centering about
Pueblo Bonito, one of the largest and most important prehistoric
ruins in the United States. Mr. Judd left Washington for New
Mexico on May 1 and shortly thereafter began operations in the

a

x

Fic. 110.—Pueblo Bonito, from the northwest, showing the vast accumula-
tions of fallen wall material and wind-blown sand which cover the ruin. The
present height of the north wall is indicated by the three figures in the left
cn (Photograph by O. C. Havens. Courtesy of the National Geographic
Society. )

a

great ruin; his staff consisted of seven assistants with Navaho and
Zui Indians employed for the actual work of excavation.

The first few weeks were largely devoted to development of a water
supply sufficient for the expedition camp, to transporting equipment
and provisions from the railroad, 62 miles distant, and to removal of
several hundred tons of fallen wall material and wind-blown sand
which had accumulated in that section of the ruin selected for the
season’s explorations. Following these preliminaries attention was

*Smithsonian Misc. Coll., Vol. 72, No. 6.

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 107

directed, respectively, to the central and southeastern portions of the
pueblo. The central wing was considered of prime importance since
it included the Great Kiva, the civil and religious heart of Pueblo
Bonito; the southeastern quarter was chosen because its masonry,
apparently the most recent of all in the village, suggested that antiq-
uities found in this area would illustrate the very apex of cultural
advancement by the ancient Bonitians, thus forming an index for sub-
sequent discoveries.

Fic. 111.—Zufi workmen pointing out features of the masonry in Pueblo
Bonito, which is far superior to that in their own village. The skill exhibited
by the ancient artisans was a source of constant admiration to these modern
Pueblos. (Photograph by Neil M. Judd. Courtesy of the National Geo-
graphic Society.)

Altogether, fifty secular rooms and five kivas were excavated during
the summer. In addition, a number of dwellings previously opened
were cleared of their individual accumulations of wind-deposited sand
and other débris. An outstanding result of this work was identifica-
tion of three distinct types of masonry, each illustrating the dominant
construction method at a given period during occupancy of the village.
It is, of course, still too early to designate the factors which brought
about these various styles in building, just as any present effort to
trace the ground area formerly occupied by each of the three types

108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 112.—Central portion of Pueblo Bonito, from the south, showing the
north cliffs of Chaco Canyon towering above the ruin. Some of the rooms
were so large that the initial work of excavation could be done directly with
teams. (Photograph by O. C. Havens. Courtesy of the National Geographic
Society. )

Fic. 113.—An excavated kiva in Pueblo Bonito, showing the low encircling
bench and, above this, the roofing timbers which overlap above the pilasters.
At the left will be seen the decayed fragments of upright hewn planks which
stood between the dome-shaped roof and the circular wall of the chamber.
(Photograph by O. C. Havens. Courtesy of the National Geographic Society.)

NOZ 15 SMITHSONIAN EXPLORATIONS, 1921 10g

= ‘ eon e

Fic. 114.—Excavating the Great Kiva. The block of masonry in the middle
is the fireplace; that in the lower right, an inter-pillar compartment. The piles
of stone at the top consist of blocks retained for repair of the ancient walls.
(Photograph by O. C. Havens. Courtesy of the National Geographic Society.)

Fic. 115——The Great Kiva and its surrounding rooms, as seen from the
cliffs north of Pueblo Bonito. This remarkable structure is 52 feet in diam-
eter; it was the largest and most important ceremonial room in the village. A
trench for stratigraphical examination of the west refuse mound will be noted
at the upper left center. (Photograph by O. C. Havens. Courtesy of the
National Geographic Society. )

8

I1O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

would be premature. Dwellings were razed and replaced by other
structures as Pueblo Bonito grew in size and population.

Those walls which appear to have formed the nucleus of the village
are crude and irregular; the rooms they inclose are relatively small
and low of ceiling. In contrast to these, walls of the second type
exhibit an infinite amount of patience and attention to detail. They
consist of rather large uniform blocks of friable sandstone, dressed
on the face only, laid in adobe mud and chinked with innumerable
small, thin chips. Equally marked in its variation from that in the
oldest houses is the masonry of the third type mentioned. In this,
uniformly thin tablets of laminate sandstone were utilized with a
minimum of adobe and little or no chinking. Larger blocks were
frequently laid in bands both for the decorative effect produced and
as bonds to hold the masonry veneer to the earthy core of the wall.
Beneath the floors of a large number of the rooms excavated during
1921 were found the razed walls of older structures in which a differ-
ent style of construction prevailed.

These principal variations in masonry may represent merely local
developments—the will of ascendant influences in Pueblo Bonito—but
it seems more reasonable to believe that each came in upon a wave
of immigration from other regions. Among the collections made
during the summer are specimens of pottery characteristic of the Mesa
Verde cliff-dwellings in Colorado, of the prehistoric ruins in the
Kayenta and Gila River districts of Arizona and of the Rio San
Francisco, New Mexico. The very number of these objects would
indicate not that they had been introduced through intertribal com-
merce but rather that their makers had come to dwell at Pueblo Bonito,
bringing with them their own distinctive arts and industries. On the
other hand, it is manifest that the prehistoric Bonitians maintained
an active trade with other primitive folk at a great distance from their
terraced village in Chaco Canyon. The quantity of Pacific coast
shell—used for beads, pendants and other ornaments—copper bells
from central Mexico and especially skeletons of the great macaw
(Ara macao), furnish abundant proof that adventurers from Pueblo
Bonito or friendly traders from distant valleys braved the rigors of
open desert travel long before the Spanish conquistadores introduced
the horse and other beasts of burden.

The circular kivas in Pueblo Bonito, as elsewhere, were both
council chambers where clan representatives met for consultation and
religious sanctuaries in which secret ceremonies were enacted and prep-
arations made for public rituals to be held in the open courts of the

NO. 15 SMITHSONIAN EXPLORATIONS, I92I IP ItAt

Fic. 116.—Repairing third-story walls in Pueblo Bonito. Some of these high
walls had been so weakened by vandalism and the elements that their repair
was necessary before excavation could safely be undertaken beneath. The
work will serve, also, to preserve the present height of the walls for many
years to come. (Photograph by O. C. Havens. Courtesy of the National
Geographic Society. )

Br2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

lc. 117.—Repaired walls of the third type of masonry, showing occasional
bands of thicker blocks inserted for strength and decorative effect. Corner
doorways are not uncommon in Pueblo Bonito; they provided a direct means
of communication between neighboring dwellings occupied by members of the

same family or clan. (Photograph by Neil M. Judd. Courtesy of the National
Geographic Society. )

SMITHSONIAN EXPLORATIONS, 1921

Fic. 118.—A Zuni Indian in an ancient Bonitian doorway. The excellence of
the masonry and the trueness of the corners are well illustrated in this picture ;
a typical lintel of pine poles will be noted at the top. (Photograph by Neil M.
Judd. Courtesy of the National Geographic Society.)

Ii4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 119.—A trench 20 feet deep was cut in the west refuse mound in order
to obtain chronological data. Potsherds deposited during the early occupancy
of Pueblo Bonito were quite different from those found near the surface of the
mound. (Photograph by O. C. Havens. Courtesy of the National Geographic
Society.)

NO. 15 SMITHSONIAN EXPLORATIONS, I92I It5

village. These important structures were constructed both below
the level of the plazas and among the living rooms, in which latter
case the surrounding walls were so arranged as to simulate the required
subterranean position. In certain features of construction and equip-
ment, however, Bonitian kivas—judging from the five already ex-
cavated—differ from those heretofore examined in other sections of

, that

The ruin
Its dwellings
Courtesy of the National Geo-

; its outer wall was unbroken except for

Excavations thus far have disclosed but one entrance,

Fic. 120.—Pueblo Bonito as seen from the north wall of Chaco Canyon.
(Photograph by Charles Martin.

is semicircular in shape and covers more than three acres of ground.

were terraced upward from two inner courts

small, elevated ventilators.

from the south.
graphic Society. )

the Southwest. The ventilator shaft is connected with a manhole in
the room through a hidden tunnel ; sub-floor chambers are sometimes,
but not always, present; the primary roof supports or pilasters have
been so specialized as to lose their original stability and to take on a
new function, that of depositories for ceremonial offerings.

The Great Kiva possesses several:noteworthy features not found in
the lesser structures of its kind. It is a room of exceptional size,

116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

being 52 feet (15.85 m.) in diameter with a ceiling formerly It feet
(3-35 m.) high. The central portion of its flat roof was supported by
four masonry pillars each of which had a separate foundation of low
grade, soft coal. On the east and west sides of the chamber, between
the pillars, were built-in receptacles, probably for containing cere-
monial paraphernalia. A fire box with protective screen stood at the

Fic. 121.—A naive example of Bonitian engineering. In an obvious attempt
to hold up a huge section of cliff which threatened to topple upon their village,
the ancients placed pine props under the weathered section and covered these
with a great terraced mass of masonry. The north wall of Pueblo Bonito
stands at the left. (Photograph by Charles Martin. Courtesy of the National
Geographic Society. )

south side and, opposite this, a flight of narrow steps led to an elevated
room in which a central block of masonry represented the “ altar.”
Excavation of the kivas and secular rooms in Pueblo Bonito is
contributing in large measure to our knowledge of the prehistoric
sedentary peoples of the Southwest. Chronological data from the
vast accumulations which comprise the adjacent refuse mounds is
expected to illustrate not only the character and extent of local cul-
tural development but to serve also as a medium of correlation between
the ancient Bonitians and other aboriginal peoples of the south-

NO. 15 SMITHSONIAN EXPLORATIONS, I92I M7

western United States. Through such data it is hoped ultimately to
arrive at the approximate age of this famous center of pre-Columbian
civilization.

The National Geographic Society proposes, as an essential feature
of its Pueblo Bonito Expedition, to conduct dependent researches
which will seek to determine the ancient source of water supply ; the
agricultural possibilities of Chaco Canyon in prehistoric times; the
rapidity of subsequent sedimentation; the age and probable source
of the large timbers used in roofing the dwellings of Pueblo Bonito, and
the geophysical changes, if any, brought about since abandonment
of the great ruin. These are lines of investigation which may result
in information of far-reaching significance and yet they have been
generally neglected, heretofore, in connection with archeological
explorations.

ARCHEOLOGICAL FIELD-WORK IN SOUTH DAKOTA AND
MISSOURI

In the fall of 1921 Mr. W. E. Myer, a voluntary collaborator of
the Bureau of American Ethnology, investigated sites in South Dakota
and western Missouri, known to have been occupied by the Omahas
and Osages in early historic times, after they had come in contact with
the whites but before they had been changed thereby to any con-
siderable extent.

Especial attention was paid to any resemblance to the ancient cul-
tures found in the valleys of the Ohio, Cumberland, and Tennessee
rivers. This line of research was suggested by certain traditions of
both the Omahas and Osages, as well as some of the other branches of
the great Siouan linguistic family, that they had at one time lived
east of the Mississippi River, on the Ohio, and elsewhere, and after
many wanderings, stopping here and there for years, finally reached
their present sites in South Dakota and western Missouri.

THE OMAHA SITES

Mr. Francis La Flesche reported that the traditions of his people,
the Omahas, stated that they had occupied two important villages on
what the Omahas call ‘‘ The Big Bend of the Xe,” at some time in the
seventeenth or eighteenth century. These traditions also told of
many important events while the Omahas dwelt on these two sites.

Aided by these traditions, Mr. Myer was enabled to locate these
two ancient villages. He found one of these on the Big Sioux River,
at its junction with Split Rock River, designated Split Rock site in
this report.

118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

He found the other site where the Rock Island Railroad now
crosses the Big Sioux River, about 10 miles southeast of Sioux Falls.
It is designated here the Rock Island site.

ROCK ISLAND SITE

Sometime in the seventeenth century the Omaha and Poncas re-
moved from the Pipestone regions in Minnesota and finally, after
some further wanderings, built a fortified town on the Big Sioux
River at the Rock Island site. While living in this fortified Rock
Island site they were attacked and defeated by an enemy, most prob-
ably the Dakotas, and finally forced to leave the region. Before
leaving, they buried their dead from this fight in a mound on this
site. This burial tradition was confirmed by excavations made by
Mr. A. G, Risty and Mr. F. W. Pettigrew, who report finding a
considerable amount of human bones in one of the mounds. Some
glass beads and small copper bells of white man’s make were also
found in one of the mounds on this site. There is evidence that this
site was occupied somewhere between 1700 and 1725.

SPLIT ROCK SITE

After leaving the Rock Island site, the Omahas and Poncas roved
without long permanent settlements for several years, but finally
returned to their beloved Xe and built a permanent village at Split
Rock site on “ The Big Bend” at the junction of the Big Sioux and
Split Rock rivers.

The month of October, 1921, was spent exploring this Split Rock
site. Many interesting relics of the Omahas were here unearthed,
which throw new light on the life of these people before they had
been very much changed by contact with the whites.

There is a group of 30 mounds on the ridge between the two rivers
marking the site of that portion of the old town occupied by the
Omahas. Ona hill one-half mile to the east was a group of ten more
mounds, occupied by the Poncas before they split away from the
Omahas at this old town.

On the tall ridge 14 miles to the west, by following the clues furn-
ished by the traditions, three low mounds were discovered. These
were said by the traditions to have been on the site of the lookouts for
the main village. These lookout mounds command a view, ranging
from 6 to 15 miles, on all sides. The mounds on this Split Rock site
appear to have nearly all been used for burial.

NO. 15 SMITHSONIAN EXPLORATIONS, I92I 11g

The exploration of mound No. 1 showed that the Indians had
selected for its site the summit of a beautiful little knoll on the edge
of the steep bluff-like bank of Split Rock River. In the soil of this
summit they dug a shallow pit, about 12 feet by 6 feet, and 2 feet deep.
In this shallow pit bones belonging to five bodies had been placed.
Several of these bodies appeared to have been buried after decay of the
flesh. One body appeared to have been buried in the flesh, closely
flexed, and this human bundle placed in the pit. The position of the

Fic. 122.—A portion of the layer of human bones on floor of charnel pit.

skeleton of a horse with a crushed frontal bone showed that when
this body-bundle had been placed in the pit, a large horse, about seven
years of age, had been led to the knoll and there killed, on the edge
of the pit, by the side of this body-bundle. Then, over all these, a low,
round-topped mound, 60 feet across base and 54 feet in height, had
been raised.

Mound No. 2, the largest mound of the group, was near the center
of the village. It was round-topped, 110 feet across base, and Io feet
high. This mound proved to be of considerable importance. In
beginning its construction, a rectangular charnel pit, 12 feet by 14
feet, and 2 feet deep, had been dug in the surface of the soil near the
center of the town. This empty pit was then thoroughly coated with
a white layer, about $ inch in thickness. This white coating was
made from calcined bones.

120 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

The bottom and sides of this white pit were then probably covered
with soft furs. This is indicated by a thin black layer of animal
matter next to the white coating.

On the floor of this fur-lined pit, bones representing about 50
human beings had been laid. These bones had been brought from
elsewhere after the decay of flesh. The bones presented the appear-
ance of belonging to bodies which had either been left unburied, as
on some battle field, or of belonging to scaffold burials. This solid
layer of compressed broken and decayed human bones entirely covered
the floor of the charnel pit to a depth of from 2 inches to 6 inches.

Fic. 123.—Bone flesher.

Portions of this layer of human bones, before it had been disturbed,
are shown in figure 122.

On top of this solid mass of human bones traces of the thin fur layer
were also found. Over this soft, warm fur covering a layer of bark
was laid, and over this bark earth had been spread to a depth of from
3 to 6 inches. This layer of earth was then smoothed and pressed
down, and on this surface a white coating, similar to that on the
bottom and sides, had been spread. Thus, these human bones, en-
closed in their layer of warm furs, were completely incased by this
white layer, very much as the filling of a pie is enclosed by the crust.
Only one small, cylindrical copper bead was found with all this
mass of bones.

On the exterior of this communal charnel pit, on all four sides, the
separate burials of several adults and two small children were found.
With these outer burials were found several objects. Amongst these
was the bone flesher shown in figure 123. With a compact bundle of

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 TI

bones belonging to two adults was a small pile of 30 circular orna-
ments of shell like those shown in figure 124. These ornaments had
probably been attached to some garment in the original temporary
burial and removed from the decayed garment when placed with the
bones in this new burial.

Fic. 124.—Shell ornaments.

No object of white man’s manufacture was found on this site.
There is evidence that this site was occupied by the Omahas some-
where between 1725 and 1775.

The Omahas and their kindred, the Poncas, lived together at this
Split Rock site. It was here that some of the most important events
in the history of the Omahas and Poncas took place. While living
here the long hostilities between the still united Omahas and Poncas
and their old enemies, the Cheyennes and Arikaras, were ended by

122 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

a peace which was concluded with great ceremony at this Omaha-
Ponca town. At the urgent request of the Arikara the sacred chant
and dance of the calumet was used to cement this great peace pact. In
this manner the Omahas and Poneas for the first time came into
contact with this the most profoundly binding and sacred ceremony
known to savage man.

At this site the age-long association between the kindred Omahas
and Poncas was broken. The tradition does not give the cause ot
their separation; but for some reason the Poncas, after having lived
with the Omahas through their long slow wanderings in the regions
east of the Mississippi and through the lower and middle reaches of
the Missouri Valley, left their kindred and formed a separate tribe.

It was at this site that the Omahas first came to possess the white
man’s horse, which was to play such an important part in the later
Omaha life. The tradition tells that neither the Poncas nor the
Omahas had possessed horses until after their separation at this site.
The finding of the skeleton of a horse in a mound on this site is
one of the many evidences which confirm this tradition that the
Omahas remained at this site after the Poncas split away, and shows
the Omahas were still living here when they first obtained horses.

OSAGE SITES

In Vernon and Bates counties, western Missouri, near the junction
of the Osage and Marmiton rivers, Mr. Myer found several sites
known to have been occupied by the Osage Indians in early historic
times, shortly after they had come in contact with the whites.

Two of these early historic Osage sites, the village of the Grand
Osage and the Little Osage village, were probably located. These
were visited by Zebulon Pike in his journey of exploration in 1806.

The site of the village of the Grand Osage was at the junction
of the Marmiton and Little Osage rivers, in Vernon County.

The probable site of the Little Osage village of Pike was at the
Perry and MacMahan coal mine, about 2 miles northwest of the
village of the Grand Osage. Old settlers stated that decayed lodge
poles were still standing and many other signs of Indian occupancy
were to be seen at this Little Osage site as late as 1840. The present
appearance of this site is shown in figure 125.

A considerable collection of surface finds from this site shows no
objects of white man’s manufacture; but local tradition says frag-
ments of brass kettles, old gun barrels, early bullets, and other objects
of white man’s manufacture have been found here.

NO. 15 SMITHSONIAN EXPLORATIONS, 1921 123

Fic. 125.—Site of Pike’s Little Osage Village.

Fic. 126.—Halley’s Bluff.

124 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

Fic. 127.—Site of cache pits at base of Halley’s Bluff.

NO. 15 SMITHSONIAN EXPLORATIONS, I92I 125

Two of these surface finds throw light on the extent of aboriginal
barter. One of these is a broken obsidian implement. The nearest
source of this material is probably in the Rocky Mountains, some
1,000 miles to the west. Another is a shard of Mesa Verde pottery,
the nearest source of which is in the Mesa Verde culture region around
the southwestern corner of Colorado, about 800 miles to the west.

The largest Osage village in Vernon County is at what is still known
as Old Town, on Old Town creek, about 33 miles south of Pike’s
village of the Grand Osage. This site covers about 40 acres and is
the best known of any of the Osage sites. It has yielded a large
amount of iron axes, gun barrels, gun locks, fragments of brass
kettles, glass beads, and other articles of early white manufacture.
Along with these large quantities of shell beads, flint arrow heads,
broken pipes, and other objects of purely aboriginal origin were
found. Old Town culture furnishes an excellent example of Indian
culture in the days of early contact with the whites.

The most picturesque Indian site in this Osage region is Halley’s
Bluff on the Osage River, about 1} miles down stream from where
the Marmiton and Marais des Cygnes unite to form the Osage River.
A photograph of a portion of this bluff is shown in figure 126. There
is evidence showing occupancy of this bluff by Indians long before
the coming of the white man and probably before the coming of the
Osages.

The long summit of the bluff shows many small, low heaps of stones
and other Indian signs. The sheltered spaces at the foot of the over-
hanging cliffs were out of reach of the highest waters and were
sheltered in large degree from the winds and rains. Here, in these
dry, sheltered spaces, these ancient people lived and worked. They
dug about 20 cache pits at present about 5 feet in depth, in the mod-
erately soft red sandstone.

FIELD-WORK.ON THE KIOWA, PUEBLO, AND CALIFORNIA
INDIANS

At the end of July Mr. J. P. Harrington, ethnologist, proceeded to
California to continue his studies of the Indians of the Chumashan
area of that state. Place-names, material culture, and sociology, all
these branches being closely related to language, were especially in-
vestigated and all obtainable data recorded. By rare good fortune
several dozen old ceremonial songs were obtained on the phonograph,
with full notes and translation where possible, these songs having
not been in use since the middle of the past century. The songs were

126 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 72

accompanied by the beating of the split-stick, and the rendition, while
not what might be desired, will doubtless be adequate for transcription.
They belong to several distinct cycles. Interesting comparisons were
drawn between the California Indian culture and that of the south-
west. The sweathouse is certainly the same as the kiva. The Cali-

Aged Mission Indian informant. (Photograph
by Harrington.)

Fic. 128.

fornia phratries correspond to the dual division of the Pueblos. The
dancers who represent demons are the Pueblo katcinas. These re-
semblances also extend to many minor features.

Nor was the linguistic side of the work neglected, ethnology and
linguistics, and in fact archeology, of necessity going hand in hand
in this difficult field. This linguistic work is of the greatest im-
portance since it furnishes material for comparison with all the
related languages.

NO. I5 SMITHSONIAN EXPLORATIONS, I92I 127

Mr. Harrington’s field studies reveal the fact that the language
of the Kiowa, who are now settled in Oklahoma but formerly had
eastern Montana as their habitat, is closely and genetically related to
that of the Taos and other Tanoan tribes of New Mexico, which have
typical Pueblo culture. Thus, the interesting fact is established that
the Taos speak a dialect of Kiowa just as the Hopi, farther west, speak
a divergent Shoshonean. These studies also make it clear that Keres
and Zufi are related to each other genetically, and furthermore to
Tano-Kiowan and Shoshonean, the languages all having a common
origin.

ARCHEOLOGICAL FIELD-WORK ON THE SUSQUEHANNA
RIVER, PENNSYLVANIA

ina july, 1921, Mr. John L. Baer, Acting Curator of American
Archeology of the United States National Museum, examined for

Fic. 129.—Petroglyphs, Bald Friars, Fic. 130.—Petroglyphs on Miles Is-
Md. land, Susquehanna River, Pa., near
Mason-Dixon Line.

the Bureau of American Ethnology a number of instructive picto-
graphs at Bald Friars and Miles Island in the Susquehanna River.

These occur about one-fourth the distance between Bald Friars
Station and Conowingo Station, on the Columbia and Port Deposit
Railroad.

All the rocks upon which petroglyphs are found seem to have been
polished before the petroglyphs were cut in them. The top surfaces
of most of the rocks bearing petroglyphs were marked with cups and
circular grooves, some of which were concentric. Some of the rocks
were fractured destroying the continuity of pictures that originally
existed. Upon one large rock broken from its original position pos-
sibly by ice are carved two slender fishes headed up-stream. The rock
upon which they were found suggests a good stand for shad fishing
with a net.

128 SMITHSONIAN. MISCELLANEOUS COLLECTIONS VOL. 72

On a group of low rocks to the northwest of Miles Island is a
peculiar arrangement of figures. On one side of a tectiform rock
are two concentric circles with radiating spokes, a cup, and two semi-
circular concentric grooves, while on the ridge and extending down on
the opposite side of the roof-like rock is a figure that might represent
an animal.

During the same trip, Mr. Baer spent several days on Mount
Johnson Island, Susquehanna River, and on the near-by flats below

Fic. 131.—Petroglyphs on Miles Is- Fic. 132.—Petroglyphs, Bald Friars,
land, Susquehanna River, Pa., near Md.
Mason-Dixon Line.

Peach Bottom, Lancaster Co., Pa., seeking further evidences of the
bannerstone workshop in which he has been interested for a number
of years. He brought back with him a number of broken and unfin-
ished slate bannerstones, flint pecking stones, polishing stones and
other utensils showing evidences of a considerable sized workshop on
the island. A synoptic series from this workshop showing the differ-
ent stages in the manufacture of the bannerstone has been placed on
exhibit in the Pennsylvania case in the American Archeological col-
lection of the National Museum.

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