1

VOL. I

JANUARY, 1947

No. 1

505- °|
P 1 17

PACIFIC

SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

IN THIS ISSUE: St. John — Sandalwood on Oahu, Hawaiian Islands
• Macdonald, Shepard, and Cox — Tsunami of April 1, 1946, in
the Hawaiian Islands • Sherman, Kanehiro, and Fujimoto —
Dolomitization in Semi- Arid Hawaiian Soils • Fisher and
Baldwin — The Red-Billed Leiothrix in Hawaii ® NOTES

Published by
THE UNIVERSITY OF HAWAII
HONOLULU, HAWAII

BOARD OF EDITORS

A. Grove Day, Editor-in-Chief, Department of English, University of Hawaii
Ervin H. Bramhall, Department of Physics, University of Hawaii
Vernon E. Brock, Director, Division of Fish and Game, Territorial Board of
Agriculture and Forestry

Harry F. Clements, Plant Physiologist, University of Hawaii Agricultural
Experiment Station

Robert B. Dean, Department of Chemistry, University of Hawaii
Charles H. Edmondson, Zoologist, Bishop Museum, Honolulu, T. H.
Harvey I. Fisher, Department of Zoology, University of Hawaii
Frederick G. Holdaway, Head, Department of Entomology, University of
Hawaii Agricultural Experiment Station
Maurice B. Linford, Head, Department of Plant Pathology, Pineapple Research
Institute, Honolulu, T. H.

A. J. Mangelsdorf, Geneticist, Experiment Station, Hawaiian Sugar Planters’
Association, Honolulu, T. H.

Harold St. John, Chairman, Department of Botany, University of Hawaii
Chester K. Wentworth, Geologist, Honolulu Board of Water Supply

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ly Continued on inside back cover ]

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

VOL. I JANUARY, 1947 No. 1

Issue printed December 10, 1946

CONTENTS

PAGE

The History, Present Distribution, and Abundance of Sandalwood on Oahu,

Hawaiian Islands. Harold St. John . 5

The Tsunami of April 1, 1946, in the Hawaiian Islands. G. A. Macdonald,

F. P. Shepard, and D. C. Cox . 21

Dolomitization in Semi-arid Hawaiian Soils. G. Donald Sherman, Yoshinori

Kanehiro, and Charles K. Fujimoto . 38

Notes on the Red-billed Leiothrix in Hawaii. Harvey I. Fisher and Paul H.

Baldwin . 45

Notes:

Recommendations of Pacific Science Conference, National Research Council . 52

Micronesian Expedition of University of Hawaii, Summer of 1946 ... 60

Survey of Micronesia by U.S. Commercial Company, 1946 . 62

Cover drawing by A. S. MacLeod

Pacific Science, a quarterly publication of the University of Hawaii, appears in January,
April, July, and October. Subscription price is three dollars a year; single copies are one
dollar. Check or money order payable to University of Hawaii should be sent to Pacific
Science, Office of Publications, University of Hawaii, Honolulu 10, Hawaii.

505.^

.*? \\7

The History, Present Distribution, and Abundance of
Sandalwood on Oahu, Hawaiian Islands:
Hawaiian Plant Studies 14 1

Harold St. John2

INTRODUCTION

Today it is a common belief of the resi¬
dents of the Hawaiian Islands that the san¬
dalwood tree was exterminated during the
sandalwood trade in the early part of the
nineteenth century and that it is now ex¬
tinct on the islands. To correct this impres¬
sion, the following notes are presented.

There is a popular as well as a scientific
interest in the sandalwood tree or iliahi of
the Hawaiians, the fragrant wood of which
was the first important article of commerce
exported from the Hawaiian Islands.

For centuries the sandalwood, with its
pleasantly fragrant dried heartwood, was
much sought for. In the Orient, particularly
in China, Burma, and India, the wood was
used for the making of idols and sacred
utensils for shrines, choice boxes and carv¬
ings, fuel for funeral pyres, and joss sticks
to be burnt in temples. The distilled oil was
used in numerous medicines, perfumes, and
cosmetics, and as a body rub. The thick oil
pressed from the seed was used as illuminat¬
ing oil.

Though long believed to be native to
India, the " white sandalwood,” Sant alum

1This is the fourteenth of a series of papers
designed to present descriptions, revisions, and
records of Hawaiian plants. The following papers
have been published as Bernice P. Bishop Museum
Occasional Papers: 10 (4), 1933; 10 (12), 1934;
11 (14), 1935; 12 (8), 1936; 14 (8), 1938; 15
(1), 1939; 15 (2), 1939; 15 (22), 1940; 15 (28),
1940; 17 (12), 1943; and nos. 11, 12, and 13 are
in press.

2 Chairman, Department of Botany, University
of Hawaii.

album L., the species first commercialized,
is now considered to have been introduced
into India many centuries ago and cultivated
there for its economic and sentimental val¬
ues. Only in more recent times has it at¬
tained wide distribution and great abun¬
dance in that country. It is certainly indige¬
nous in Timor and apparently so all along
the southern chain of the East Indies to east¬
ern Java, including the islands of Roti, We-
tar, Sawoe, Soemba, Bali, and Madoera. The
earliest voyagers found it on those islands
and it was early an article of export, reach¬
ing the markets of China and India ( Skotts-
berg, 1930: 436; Fischer, 1938). The in¬
sufficient and diminishing supply of white
sandalwood gave it a very high and increas¬
ing value. Hence, trade in the wood was
profitable even when a long haul was in¬
volved.

the genus Santalum

The genus Santalum contains several spe¬
cies and most, if not all, of these are accept¬
able alternatives or substitutes for S. album.
Hence, each newly discovered stand of any
species of sandalwood attained great eco¬
nomic value. There are now 19 accepted
species of Santalum occurring naturally from
Java to Juan Fernandez, Hawaii, and the
Bonin Islands. These species are found as
follows :

Java to Timor — S. album L.

New Guinea — S. Macgregorii F. Muell., S. papu-

anum Summerh.

Australia — S. lanceolatum R. Br., S. obtusifolium

R. Br., S. ovatum R. Br.

5

6

PACIFIC SCIENCE, January, 1947

New Caledonia, Loyalty Islands, New Hebrides — *
S. austrocaledonicum Vieillard
Fiji — S. Yasi Seem.

Tahiti, Raiatea, Austral Islands, Rapa, Marquesas
— S. insulare Bertero

Henderson Island — S. hendersonense F. B. H. Br.
Juan Fernandez — 5*. fernandezianum Phil, (now
extinct)

Hawaiian Islands — S. ellipticum Gaud., S. Frey-
cineiianum Gaud., S. haleakalae Hbd., S. lanai-
ense Rock, S. paniculatum H. & A., S. Pilgeri
Rock, S. pyrularium Gray
Bonin Islands — S. boninense (Nakai) Tuyama

THE SANDALWOOD ERA IN HAWAII
The Hawaiian people were familiar with
the pleasant aroma of the iliahi. They called
its wood laau ala (or laau aala), meaning
"fragrant wood,” and they sprinkled finely
powdered heartwood on their kapa or bark
cloth to perfume it.

Just how the Hawaiian sandalwood trade
began is not known. Perhaps the Hawaiians
showed the pleasantly scented wood to the
white voyager or "haole.” Perhaps, as has
been suggested, logs of iliahi were included
in firewood furnished to sailing vessels in an
island port and some sea captain, veteran of
the China trade, recognized the sweet per¬
fume of sandalwood oil coming from the
burning wood in the cook stove. The actual
beginning is not certainly known, but the
first record is as follows (Bradley, 1942:
26-27; see Ingraham, 1791: 15, 19, 20, 23,
24) :

The earliest specific mention of sandalwood at the
Islands is attributed to Isaac Ridler, a deserter
from the "Columbia.” Ridler was at Kealakekua
Bay in March 1790, during the visit of the "Elea-
nora”; a year later, he told Captain Joseph Ingra¬
ham that while at Kealakekua Captain Metcalfe
had engaged in "taking in sandalwood.” Metcalfe
may have purchased the sandalwood for use as
firewood; it is also possible, if we may credit
Ridler’s testimony, that he was the first to carry
sandalwood from the Hawaiian Islands to China
for commercial purposes.

In 1790, Captain John Kendrick of the
"Lady Washington” left two men on Kauai
to collect sandalwood for his next return
from Boston. Before May, 1791, Captain

William Douglas of the "Grace” (Ingra¬
ham, 1791: 15, 19) put two men ashore on
Kauai to collect sandalwood, "which wood
he had discovered or a wood similar to it.”

There is doubt about the accuracy of these
early records, for Amasa Delano (1817:
399; see Bradley, 1942: 27) observed in
1801 that the first sandalwood shipped from
the islands to China was not the real sandal¬
wood and was of such inferior quality that
the Chinese merchants refused to purchase
it.

From 1790 to 1810 sandalwood may have
been exported, but if so, in very small quan¬
tity, for little record is found. Then, in
1809, two brothers, the American ship cap¬
tains Jonathan Winship of the "Albatross”
and Nathan Winship of the "O’Cain,”
started on a voyage that established the san¬
dalwood trade. After trading for furs on
the coast of Oregon, they sailed in October,
1811, for Honolulu, where they and Cap¬
tain William Heath Davis of the "Isabella”
took on cargoes of sandalwood. The ships
sailed to Canton, where the fragrant wood
was sold at a large profit. Returning to
Honolulu, the three captains persuaded King
Kamehameha I to grant them a monopoly of
the sandalwood and cotton trade for 10 years.
Loading five ships, the three captains sailed
to Canton and thus established a highly re¬
munerative traffic. While they were in
China, the startling news came that America
and Great Britain were at war, and when
the captains returned to Honolulu they
found Kamehameha unfriendly. He can¬
celed their trade monopoly and refused to
renew it, his changed attitude being due,
they reported, to British influence (Davis,
1816: 52-53).

Thereafter, though no longer a trader’s
monopoly, the sandalwood trade developed
rapidly and throve from 1815 to 1826. The
successive Hawaiian kings at first followed
the example of the shrewd Kamehameha I
and kept sandalwood as a royal monopoly,

Sandalwood in Hawaii — St. John

7

but later they shared it with the higher
chiefs. Taxes, to be paid in sandalwood,
were levied on the district chiefs, who in
turn levied them on their retainers, the peo¬
ple. Wanting some of the white man’s sail¬
ing ships, the king bought several, paying
for them, it is said, with an equal or a double
tonnage of sandalwood. For measuring the
amount, a rectangular pit the size of the
greatest length, breadth, and depth of the
hull was dug, then filled with sandalwood
logs (Rock, 1916: 10). These pits are still
reported in various parts of the islands, and
there is one, identified by the late Albert F.
Judd, at about 800 feet elevation on the
Kapalama-Nuuanu ridge, on the grounds of
the Kamehameha Schools, Honolulu.

After the death of Kamehameha I in
1819, his successors ruled, but without the
sagacity and shrewdness of the great king.
Their sandalwood monopoly still seemed to
them an inexhaustible source of wealth.
The wood was marketed in China by the
picul (133 V3 pounds) and its value fluctu¬
ated from $3.00 to $18.00; during the pros¬
perous years 1825 to 1827 it sold at from
$10.50 to $14.00 a picul. In 1822-23 a
total of 20,000 piculs was sold in Canton.
The trade was a great resource for Kame¬
hameha I, but for the next two kings, Liho-
liho and Kauikeaouli, and their womenfolk,
it was a bonanza. They paid $77,000 for a
brig and its cargo. They bought frame
houses shipped ready-cut, silks, woolens,
liquors, cut glass, and luxuries of many other
kinds. They paid $800 for a looking glass
and $10,000 for a brass cannon. With the
consent of the eager, competing traders,
these purchases were accepted on credit for
sandalwood to be delivered later. These
reckless purchases piled up royal debts in
1821 of 22,000 piculs — or $220,000 calcu¬
lated at $10 a picul, the quotation for 1821-
23 (see Mathison, 1825: 5)— and by 1824
of about $300,000.

Not only were the logs of sandalwood the
prime article of commerce, but for some
years they served as a medium of exchange
in the absence of a local coinage. This fact
is recorded by James Hunnewell (1895:
16), who as a ship’s officer made several
voyages to Hawaii. When, on one of the
voyages, the cargo was only partly disposed
of by sale, Hunnewell was left in Honolulu
with the remaining part of the cargo to sell.
He wrote, concerning the years 1817 and
1818:

Business seems to have been conducted, to a very
considerable extent, by barter. Sandal wood was
the chief article, indeed it might almost have been
called the standard coin, although Spanish silver
more nearly reached that definition. There is con¬
stant mention of sticks or piculs of the wood, but
none of money. May 14th [1818] is a note, "Sold
40 looking glasses for 4 piculs wood”; next day,
"Sold the remainder of the muslin, 2 pieces, for
31 piculs wood received.” These examples are
enough to show the nature of trade.

The king and the chiefs, pressed for pay¬
ment, put pressure on the people, whose
labor made the trade possible and who re¬
ceived little or none of the profit. There are
numerous contemporary accounts of the
forced gathering of sandalwood. All the in¬
habitants able to go were ordered into the
hills in search of the precious wood. The
trees were cut down and chopped into logs
6 to 8 feet long; then with adzes the bark
and sapwood were chipped off. Men and
women tied the logs to their backs with the
fibrous leaves of the ti ( Cordyline termi¬
nal/ s) and trudged to the measuring pit or
to the shore. As many as a dozen small ships
were used to haul the logs to Honolulu.
Ellis (1826: 275) at Olaa, Hawaii, encoun¬
tered a chief with about 400 people bearing
sandalwood from the mountains. There are
also accounts of the gathering of logs by
two to three thousand people near Waipio,
Hawaii. The following account by a British
sailor ("Old Quarter Master,” 1839: 220-
221 ) does not- seem to have been noticed be¬
fore, and it is here quoted because of its

8

PACIFIC SCIENCE, January, 1947

charm and because it gives a picture of gath¬
ering the wood at night:

January, 1827, weighed and made sail, and after
a fine passage anchored within the reefs, at Wha-
hoo [Oahu], where we found an American ship.
. . . Sometime before our arrival the Government
had purchased a vessel from an American Mer¬
chant called the Chinchili; this vessel was to be
paid for in sandal wood, which was as usual levied
in certain quantities from each of the Chiefs. . . .

The time of cutting it is appointed by the King,
and that is invariably at night, but for what cause
I could never find out; this time was appointed
while we lay here, and the Captain was invited to
accompany the young King to view the scene; our
cutter was ordered to take the party, and it took
us nearly all the forenoon to get to the spot. When
the party took horses and left me in charge of the
boat, I asked and obtained leave to ascend the hill
in the evening; the ascent was painful and fatigu¬
ing, but it fully repaid me by the pleasing sight
that met my eyes; there stood a vast number of
men assembled, each with a torch made from
sandal wood, which burns bright and clear, at a
certain signal they dispersed; each taking his own
way to cut his load, accompanying his labour with
a song, to which the whole band within hearing
join in chorus; the song we understood not, but in
the calm of a beautiful night it was calculated to
inspire delight. After the labour of two or three
hours the wood is collected together, each Chief
inspecting his own lot, judging of the quality by
the colour and weight; it is then taken to the
water’s edge where it is piled end on ready for
boats to take away; the people then returned to
their homes, and we to the young King’s country
house, after which I went to the boat.

As the king bought greater and greater
quantities of imported goods, his demands
for sandalwood in taxes became greater and
more frequent. Tax records were kept, some
of which are still extant and may be con¬
sulted at the Public Archives, Honolulu. The
writer studied them carefully in hope of
finding exact records, including the locali¬
ties where sandalwood was cut. The records
are few, fragmentary, and difficult to inter¬
pret. One of the best is a sheet kept by the
Spaniard Francisco de Paula Marin, who was
a councilor or business manager of Kame-
hameha I. The year date is lacking. Each
center entry is the name of a district on
Oahu or of the chief ruling that district.

Then, in three columns appear the day and
month, the number of piculs, and the num¬
ber of pieces, all written in Spanish. In
brackets is here suggested the modern spell¬
ing or translation, as follows:

Sandalwood Tax Record for Oahu by
Don Francisco Marin

Guaguigui [Waikiki]
dia 24 de Septiembre 240 piculs
25 50

18 de Septiembre

Taguipu [Kahuku?]

168

16

Cayrua [Kailua]

1200

16

Guaymanamano

[Waimano]

1200 10 pedazos [pieces]

Caneoje [Kaneohe]

16

800 5

17

Teylla [Kealia]

1200

17

Cajanu [Kahana?]

120

23

Camejameja [Kamehame-
ha, the governor of part
of Oahu]

4400

27

1080

29

230

28

Tonuaunau [unidentified]
580

23

Guallanae [Waianae]

415

Marin’s journal, in manuscript translation
in the Public Archives, also gives some data
on the gathering of sandalwood taxes.

Marin’s Journal No. 2

27 Sept. 1811. Marin cutting Wood for King.

26 Dec. 1811. Marin goes to cut Wood with the
Minister.

4 Aug. 1812. This day the King made a Con¬
tract with Captain Guynan [Winship] and Capt.
Debes [Davis] not to sell fragrant wood to any¬
one but to them.

25-26 Oct. 1814. Kealakekua, Hawaii. Went
on shore to see King. Spoke to him about per¬
fumed wood. He gave no answer.

18 Feb. 1818. This day weighing wood — 116
piculs.

Journal No. 4

1 March 1819. Quinopu & Quehomaquea left
for Waimea to fetch sandalwood [they were prob¬
ably tax collectors, now unidentified].

Sandalwood in Hawaii — St. John

9

Journal No. 5

2 Dec. 1819. Poqui [Boki] buys boat of Capt.
Luis for twice its full of Sanlegue [sandalwood].

19 March 1821. This day Craymoca [Kalani-
moku] started to collect sandalwood and Poqui
[Boki] for Quallanae [Waianae].

22 Oct. 1821. Pitt [Billy Pitt or Kalanimoku,
the king’s prime minister and treasurer] bought
schooner "Asta” for 1280 Piculs of Sanlegud [san¬
dalwood].

Journal No. 6

15 Jan. 1822. Sandalwood for caulking.

8 Aug. 1822. Embarked sandalwood on ship
America.

It will readily be seen that these historic
documents are incomplete and lack details
as to the exact localities where the sandal¬
wood trees grew.

The tax levies increased, becoming more
and more exacting, and as all chroniclers
agree, they became an intolerable burden on
the people. As the easily accessible sandal¬
wood stands had been felled, the people had
to climb farther and farther into the wet,
cold mountain forests and the quests were
no longer like idyllic song fests. The people
were driven to the task, and many died of
exposure in the mountains. While they
were away in the interior, crops and taro
patches were neglected, so that famine came
to the islands and took its toll of the king’s
subjects.

Eventually the shipments decreased and
the wood gathered was smaller and of poor
quality. In 1828-29, 13,000 piculs of san¬
dalwood were shipped. In 1830-31 the
shipments, wholly of small and crooked
sticks, brought only $1.50 a picul. This
poor yield finished the sandalwood trade in
the Hawaiian Islands. Says Mesick (1934:
140):

A few years later [than 1825], seeing that the sup¬
ply was rapidly diminishing, the government took
steps to conserve it; but despite this tardy measure
it is only occasionally that sandalwood trees are
found in the Hawaiian forests.

A similar summary (Kuykendall and
Gregory, 1926: 116-117) reads:

The reckless way in which the trees were cut de¬
stroyed the forests. Very little effort was made to
preserve the young trees or to replace those which
were cut down. In a few years sandalwood almost
disappeared from the islands. Even to-day, a hun¬
dred years after the trade was at its height, only
a few small groves are to be found.

These statements emphasize the depletion
of the stands of sandalwood, but exaggerate
the present scarcity.

bennett’s locality for sandalwood

Since locality records given by the earlier
botanists and explorers are few and are often
lacking in detail, in general they are not
quoted here. However, there is one of real
interest. Dr. George Bennett (or Bennet)
was a British physician who accompanied
the Rev. Daniel Tyerman on a world tour.
A detailed and readable report on the Chris¬
tian missions and also an account of their
general observations was edited by James
Montgomery (1831). The pair visited
many island groups, some very remote. In
the decade after their return to England,
Bennett published several scientific or botan¬
ical accounts; then, after settling in Aus¬
tralia, he continued work in natural history.
While in England he published "An account
of the sandal wood tree” (1832). A speci¬
men of Santalum which he had collected on
Oahu was deposited with his other collec¬
tions in the Botanical Museum at Berlin-
Dahlem; no other herbarium seems to have
any duplicates of his collections.

Professor J. F. Rock ( 1916: 19-21) stud¬
ied this specimen in Berlin and stated in his
notes that it agreed perfectly with an authen¬
tic Gaudichaud specimen of S. Freycine-
tianum Gaud. The data are: "Santalum,
Sandalwood Tree, Native name Iliahi or
Lauhala. Wouhala, Oahu, Sandwich Islands,
December, 1830. The tree is of slow growth
and inhabits elevated and rocky situations.

10

PACIFIC SCIENCE, January, 1947

G. Bennett.” Rock’s paper is a detailed
monograph of the Hawaiian species of San-
talum. For S. Freycinetianum he cites only
four collections from three localities :
"Wahou” [Oahu], Gaudichaud, in 1827;
Palolo, Rock nos. 10,063 and 12,512; and
Bennett’s from "Wouhala.” It is obvious
that the Hawaiian common name lauhala
does not apply to Santalum. It means
"leaves of the hala tree” ( Pandanus ) and
was and is in very common usage in refer¬
ence to mats, baskets, etc., plaited from the
leaves of the Pandanus; thus Bennett’s use
of it for Santalum can be rejected as an error.
Rock gave no explanation of the locality
"Wouhala.” The subsequent standard gazet¬
teer of the Territory of Hawaii (Coulter,
1935) includes not only the place names on
the current official topographic maps, but
also the obsolete names or spellings from all
available older maps. It does not include the
place name Wouhala or anything like it.

The lengthy journal of Tyerman and Ben¬
nett has been scanned for data on this point.
They arrived on Oahu in December, 1821,
and spent more than 3 months on the island.
Bennett gives an account of his ascent on
April 26, 1822, of Erihi [Diamond Head],
of climbing "the great mountain Punch¬
bowl,” and of climbing on April 28 the
Koolau divide, not far from the Nuuanu
Pali, "one of the highest accessible points
in this island.” He and his companion made
a tour around the island of Oahu, but he
does not in this account mention his find¬
ing of the sandalwood at "Wouhala.” Ben¬
nett later mentioned the sandalwood and
this locality, and gave a helpful description
of the tree in his book (I860: 420-421),
but this is a condensed and paraphrased ac¬
count from the fuller one (1832: 260-261)
which is here quoted:

On the 10th of December, 1829 {1821], I visited
the district of Wouhala (Island of Oahu); on
ascending a high hill, the plains on the summit
were found covered with dry grass, and various
plants and shrubs, and at some parts deep wooded
glens formed most picturesque and beautiful scen¬

ery. Among the specimens of plants, &c., I col¬
lected were the following: —

A species of Cyathodes, called pokeawi by the
natives, bearing small red berries; the same native
name is given to red beads, from their resemblance
to the berries of this shrub. A species of Phyto¬
lacca, called poporo-tumai by the natives: the
berries (which grow erect in long bunches) yield
a reddish brown juice, used for dyeing the native
cloth; the berries externally are of a purplish red
colour; the leaves of the shrub are cooked and
eaten.

On the plains was found a species of Dian ella,
named uki by the natives, bearing small berries of
a mazarine blue, which are used by the natives in
making a permanent blue dye. The Pyrus ^nthyl-
lidifolia of Smith (in Rees’s Cyclopaedia ), and
more recently the Osteomelis ^nthyllidifolia of
Lindley (in the Linnean Transactions') , called ure
by the natives, was very abundant; it is a small
shrub, bearing berries of a white colour, contain¬
ing a reddish juice of sweet and astringent taste;
the flowers are white and fragrant.

The mamati or cloth plant, also named oreyna,
the U rtica argentea; the bark is used in the manu¬
facture of the native cloth, and also produces a
flax which might form a useful article of com¬
merce. A species of Scaevola, named nouputa by
the natives, was also abundant on the hills, bear¬
ing yellow flowers.

A shrub, attaining the elevation of 9 or 10 ft,
called karia or taria by the natives, was abundant,
but the only specimens gathered had abortive
flowers.

A small tree, called lumma by the natives, had
the leaves when young of a beautiful red colour,
and the foliage has a peculiar appearance, appar¬
ently from minute glands situated on the upper
and under surfaces. There is also a shrub (prob¬
ably a Bass ia), called ohava, the seeds of which
yield a red dye, used by the natives to stain their
cheeks and fingers. A species of Gnaphalium,
called poina by the natives, was also abundant.

Of the uwara, or sweet potatoes (Convolvulus
Batatas et var.); which are much cultivated at
the Sandwich Islands, there are seventeen varieties.

On the declivities of the hills, and in the ravines,
the tui tui, or candle nut tree (Aleurites triloba)
is seen abundant; the whiteness of its foliage ren¬
dering it a conspicuous object. The whiteness is
occasioned by a fine white powder on the upper
surface of the leaf, which is readily removed by
the finger. Under it the leaf is found of a dark
green colour. The young foliage is thickly covered
with this white powder; the older leaves have
little, or are entirely destitute of it. The foliage
of this tree varies much in form, depending on
the age of the tree or leaves. The flowers grow
in erect clusters, are small, white, and possessed
of very little fragrance; the fruit is of small size.

Sandalwood in Hawaii — St. John

globular, rough externally, and contains oily nuts,
which, when baked and strung on a reed, are used
by the natives of most of the Polynesian Islands
as a substitute for candies or lamps, and burn
with a clear and brilliant flame. The tree is
branchy, attains an elevation of 30 ft. in height,
and a circumference of 3 or 4 ft., the timber being
of soft quality is useless, except as firewood. A
gum is yielded by this tree, both spontaneously,
and on incisions being made in the trunk. It is of
a yellowish colour, inodorous and tasteless; the
natives chew it, but the suspicious family [Eu¬
phorbia^*?] to which the tree belongs would ren¬
der caution requisite in its use. I tried it, however,
as mucilage for the suspension of some balsams,
without any ill effects arising from it.

The turmeric plant ( Curcuma longa), called
oreina by the natives, is abundant wild; the root,
as well as that of the noni (Morinda dtrifolia),
is used for dyeing their native cloth of a bright
yellow colour.

The foregoing account is quoted exactly
as published, including the curious render¬
ing of the scientific names, some unaccented,
some with grave, some with acute accents,
some in roman type, some in italics or partly
so or even with only one letter in italics.3
Probably Bennett’s underlining of the scien¬
tific names to be set in italics in his manu¬
script was hasty, and the underlinings did
not equal the names, though he intended
them to do so. The printer did not interpret
them thus; rather, he followed the copy
exactly.

The plants mentioned in Bennett’s ac¬
count are now known as pukeawe ( Styphelia
T am eiameiae) ; popolo (Phytolacca sandwi-
censis) ; uki (Dianella sandwicensis) ; uulei
( Osteomeles anthyllidifolia) ; m a m a k e
(Pipturus albidus) ; naupaka kuahiwi (Scae-
vola Gaudichaudiana) ; kalia (Elaeocarpus
bifid us) ; lama (Diospyros Hillebrandii) ;
kealia or alaea laau (Bixa Orellana) ; enaena
( Gnaphalium sandiuicensium ) ; uala (1 po¬
rno ea Batatas) ; kukui ( Aleurites moluc-
cana) ; olena ( Curcuma longa) ; and noni
(Morinda citri folia) . Explanatory remarks
are here appended for several of these
plants.

3 A few vowel accents unavailable to the printer
of Pacific Science have been omitted. [Editor.]

11

The naupaka kuahiwi does not have yel¬
low flowers. There is one native species
which does, Scaevola glabra, but this grows
only on the very crest of the mountains in
the Cloud Zone. All indications are that
Bennett did not on this day climb to the high
peaks, but rather was describing collections
made at the lower edge of the forest, in
grassland or dry open forest. On Oahu, one
of the commonest bushes and one of the first
to be encountered on approaching the forest
is S. Gaudichaudiana, which has white flow¬
ers with delicate magenta lines on the veins
near the throat, but the flowers, when with¬
ering and drying on the bush, turn yellow¬
ish. It seems certain that this was the
"nouputa” he noted. In confirmation of this,
there was a specimen of S. Gaudichaudiana
from Oahu, collected by Bennett, in the Ber¬
lin Herbarium ( Skottsberg, 1927: 26) .

Bennett’s bush the "karia” is certainly
kalia (Elaeocarpus bifid us) , though truly a
tree. The native name is a clear indication
and the abortive flowers settle the question,
for even today nearly eve£y inflorescence
through insect injury develops into a large,
bright red, abnormal growth almost like a
witches’-broom.

The "lumma” is evidently the lama
(Diospyros Hillebrandii) , which is conspic¬
uous in the lower forest in spring because of
its flush, or luxuriant young growth with
abundant new crimson leaves, which is even
more showy than the similar red flush on the
cultivated mango tree. The leaves of the
lama are not glandular, but both surfaces,
and particularly the upper, are strongly
salient rugose reticulate, with thin tissue be¬
tween the meshes of the heavy close net¬
work. When looking through a leaf towards
the light, Bennett may have mistaken the
light intervals for glands.

Bennett’s record of the name "ohava” is
difficult to identify. There is no such name
in the Hawaiian dictionaries or botanical
works. The only similar name is hoawa

12

PACIFIC SCIENCE, January, 1947

( Pittosporum ), which native shrub or tree
does not fill the bill. It seems fairly certain
that instead, his tree is the introduced shrub
kealia or alaea laau (Blxa Orellana). This
is an American tropical shrub, of early intro¬
duction into the Hawaiian Islands, but just
how early cannot now be definitely stated.
Dr. W. T. Brigham (1911: 158), in his ac¬
count of the dye plants used for coloring
kapa (bark cloth), wrote:

This shrub was formerly cultivated here for the
red dye obtained by macerating the seed pulp, and
has become naturalized in places. ... I found it
growing apparently wild in 1864 in Nuuanu and
on the barren plains east of Kawaiahao church.
... I believe that the old Hawaiians used the plant
as a useful dye at least a century ago.

It is still commonly cultivated and occa¬
sionally naturalized in the islands. Brigham’s
estimate of its introduction would put the
date at 1811. If this is correct, by 1831 there
would have been plenty of time for it to have
become dispersed among the Hawaiian vil¬
lages. The Hawaiians have always been great
plant lovers and skillful gardeners. Each
attractive or useful plant has been spread
among them with great rapidity.

The kealia is not a native plant, but
neither is the uala ( Ipomoea Batatas) nor the
olena ( Curcuma longa ), which latter Ben¬
nett thought was growing wild. As Bennett
was more zoologist than botanist, perhaps
he should be excused for thinking the olena
a wild plant. Many other explorers and
botanists have been fooled by the gardens
of the Polynesian peoples. Some of their
crops, which need such culture, are planted
and tended in well-cultivated fields. Others
which grow well without care are planted at
the base of the trees in a forest or on a grassy
hillside where they grow as if wild, until
the planter comes and harvests his crop.
Many a botanist visiting a strange tropical
island has recorded such plants as wild or
native, when the situation is exactly the re¬
verse, the plants being introduced economic
species planted and owned by a tiller of the

soil. The noni ( Morinda citrijolia) is also
a cultivated plant, but it seeds and tends to
spread in the lowlands.

Bennett (1832: 257) gave more details
concerning the sandalwood ( Santalum Frey-
cinetianum) :

At the Sandwich Islands, the tree is named iliahi
or lauhala, signifying sweet wood ( lau , wood,
hala, sweet) [an error, the meaning being "leaf
of the hala tree”; Bennett should have written
laau, wood, aala, sweet]; and, when young, it is
of very elegant growth. At Wouhala (Island of
Oahu), I observed numbers of the young trees,
some of which were covered by a profusion of
beautiful flowers of a dark red colour; the flowers,
however, are often observed to differ in colour on
the same tree, and even on the same stalk; they
grow in clusters, some having the corolla exter¬
nally of a dark red colour, and internally of a
dull yellow; others having it entirely of a dark
red, and others again have the corolla partly red
and white externally; the young leaves are of a
dark red colour, and give an elegant appearance
to the tree. This was not observed in the species
found at the Island of Erromanga; indeed, the
species found at the Sandwich Islands had a more
handsome appearance in its growth than that at
Erromanga. At the Sandwich Islands, two varie¬
ties of the wood are observed by the natives, de¬
pending, however, only on the age of the tree; the
young or white wood is called lau, keo keo {lau,
wood, keo keo, white);, and the red wood, lau,
hula hula {lau, wood, hula hula, red). [These are
now written laau, wood; ulaula, red.] As before
stated, the wood, when taken from a young tree,
is white, containing but a small quantity of oil;
as the tree increases in growth, the wood becomes
of a yellowish colour, and the oldest and best is
of a brownish red colour.* The different varieties
of the wood depend, therefore, on the age of the
tree; and are of three kinds, white, yellow, and
red; of which the yellow and red, from containing
the largest quantity of oil, are most esteemed in
the Chinese market, where the wood is principally
used, the expressed oil being mixed with pastiles,
and burned before their idols in the temples. The
Chinese are said to procure the oil by rasping the
wood, and then expressing it through strong can¬
vass bags.

* The wood is frequently buried, and the sap
[sapwood] allowed to rot off: and this is consid¬
ered to improve its quality. [Author’s note.]

By assembling all the data from the plant
associations and from the habitat, it has
proved possible to spot Bennett’s locality,
’'Wouhala,” for the sandalwood. The asso-

Sandalwood in Hawaii — St. John

13

dated plants are characteristic of the lower
edge of the dry forests. This habitat, up a
high hill with dry, grass-covered plains on
top, dissected by deep, wooded glens, is well
described by him. Together these data apply
to only one section of Oahu — the Leilehua
plains or the broad pass between the Koolau
and the Waianae Mountains. To avoid the
deep, steep-walled gulches cut by the larger
permanent streams from the Koolau Range,
the old foot and horse trail over the pass
kept close to the Waianae Mountains, swing¬
ing from Pearl Harbor toward the old ham¬
let of Lihue, keeping on the right (or east)
the largest stream, Waikele, and its tribu¬
taries Kipapa and Waikakalaua Streams, all
of which have cut deep gulches across the
plateau or its slopes. On the upper reaches

of Waikele Stream is Pouhala, about 1 mile
south of Schofield Barracks and 1 mile north
of Robinson Camp 1 (see Oahu topographic
map, 1938 edition). The Wouhala of Ben¬
nett is surely Pouhala of today, because the
description applies, the place is close to pres¬
ent localities for sandalwood, and Bennett is
known to have journeyed through this area.

PRESENT DISTRIBUTION ON OAHU
Accompanying this discussion is a map
showing the present occurrence of Santalum
Freycinetianum Gaud., a species restricted
to the island of Oahu, and the only arbores¬
cent species on that island. This tree was
doubtless the sole source of laau aala for the
inhabitants of Oahu. On the map, the solid
black dots mark localities for specimens with

14

PACIFIC SCIENCE, January, 1947

exact locality data, such as part of valley, alti¬
tude, etc.; half-black dots mark the approxi¬
mate place for specimens with indefinite
data, such as name of valley or ahupuaa
(land division) but nothing more; circles
mark the localities without substantiating
specimens but recorded in the field notes of
William Meinecke, E. Y. Hosaka, or the
writer. All these collections and records are
subsequent to 1907 and so give a good index
of the present distribution, which is wide,
extending nearly from end to end of both
mountain ranges. The tree is found on both
sides of the Waianae Mountains from 500
to 2,400 feet altitude, but there is a break
at Kolekole Pass, which is now treeless, de¬
nuded, and much eroded. Probably the tree
occurred originally all across this stretch.
Its absence now may be due to denudation
by lumbering, forest fires, or cattle grazing.
On the other hand, this low pass was a
standard route of travel and was easily acces¬
sible to people coming either from the west¬
ern villages in the Waianae region or from
the broad Leilehua or Schofield saddle on
the east. Its absence may also be due to
despoliation by people in the sandalwood
trade or by those in search of timber or fuel.

In the Koolau Range the sandalwood now
occurs from 550 to 1,800 feet altitude, from
the Niu-Wailupe Ridge to Kaunala and
swinging around the northern end to Laie.
The sloping north end of the Koolau Range
is lower, and therefore does not receive as
heavy a rainfall as the more elevated parts.
It is noteworthy that on the windward or
northeast side of the mountains and the
windward shore, southeastward from Laie,
there are no stations for the sandalwood. The
whole shore and mountain slope is windier
and rainier, so that the typical dry lower
forest is not well developed here. Though
koa ( Acacia Koa Gray) is present, it does
not form a definite zone, rather occurring
scattered among other trees characteristic of
wet forests. It will be recalled that in the old

sandalwood tax collection record by Marin
there were entries of 1,200 piculs from
Cayrua, 800 from Caneoje, and 120 from
Cajanu. If these names are correctly trans¬
literated as Kailua, Kaneohe, and Kahana,
they are districts where no tree sandalwood
grows today, and where it is improbable that
any ever did grow, the region being too wet.
Yet these districts paid a tax in sandalwood.
K. P. Emory has suggested to the writer that
the men of these districts which lacked san¬
dalwood may have been assigned to cut wood
in crown lands in another district and thus
to work off their taxes. The remaining dis¬
tricts all have sandalwood in the mountain¬
ous (mauka) inland sections. "Camejameja”
was probably not the king, but Kamehameha,
governor of part of Oahu. The collections
entered under his name were made on several
dates and are much larger than those from
any single district.

Almost down to the present there has
been a conspicous gap in the distribution of
sandalwood on the lee (or southwestern)
side of the Koolau Range, from Manoa to
Moanalua. There is no old record, no pub¬
lished record, no herbarium specimen to
represent this region, despite the fact that it
is the most thoroughly botanized section in
the Hawaiian Islands. All the early explor¬
ers came to Honolulu Harbor; later, the resi¬
dent botanists mostly lived in Honolulu, and
all of them repeatedly explored these ridges
and valleys just behind Honolulu. As the
habitat is suitable, this 8 -mile gap is curious.
Sandalwood is attractive, easily recognized;
it thus has both historic and sentimental
interest, and most botanists collect it every
time they see it. Consequently the gap is
probably not due to lack of collections. An¬
drew Bloxam (1925: 38) made a record of
interest in his connection. On May 13,
1825, he and the botanist James Macrae
made a trip up Nuuanu Valley toward the
Nuuanu Pali, traversing the 4 or 5 miles

Sandalwood in Hawaii — St. John

15

from the lower region, with its huts and taro
patches, to the thick woods. He stated:

We found a great variety of ferns and other plants
among which the ginger plant was very prominent.
We saw several of that beautiful tree the Eugenia
malaccensis, or Malacca apple, in full bloom with
its bright scarlet flowers, the dooe dooe, or oil nut
[now called kukui. Aleurites moluccana ] was very
common. We could not find one sandalwood tree,
all had probably been cut down about here for
the purpose of barter.

This passage testifies to the early despolia¬
tion of the stands of sandalwood.

Still in search of records in the region
behind Honolulu, the writer made inquiry
of malacologists. The terrestrial and arbo¬
real snails on the Hawaiian Islands are num¬
erous, large, and of beautiful form and col¬
oring. For more than a century they have
been sought both by amateur nature lover
and professional naturalist. As collections
have accumulated it has become evident that
many of the numerous species and subspecies
have very restricted ranges, such as a single
valley, or part of a valley, or even a single
clump of trees. The land shell collectors be¬
come well acquainted with these localities
and the species of trees favored by the snails.
Thus, many of the collectors have an exten¬
sive and accurate knowledge of the native
flora. Dr. C. M. Cooke, Jr., of the Bishop
Museum, Honolulu, remembers seeing from
1890 to 1892 several sandalwood trees on
the Kapalama-Waiolani ridge, just south¬
west of Napuumaia (in Nuuanu) at about
1,600 feet altitude. This was one of his best
collecting localities, and he clearly remem¬
bers several sandalwood trees with their
bright red, young leaves. George S. Water-
house likewise remembers seeing sandal¬
wood trees in the same locality between 1903
and 1906. He cut and carried down a good-
sized log of the wood, which he kept for
many years. He visited the locality fre¬
quently, for his brother, Fred Waterhouse,
had a mountain cabin on the ridge at Na¬
puumaia. This site is at 1,870 feet elevation

on the west ridge of Nuuanu (see Kaneohe
Quadrangle, Advance Sheet, surveyed in
1928). These two reports are considered
fully trustworthy, and indicate that S. Frey-
cinetianum persisted in Nuuanu as late as
the beginning of the present century.

Finally, the writer was delighted on Jan¬
uary 9, 1944, to find a single tree, 27 feet
high and 10 inches in diameter, on the
Kahauiki-Kalihi Ridge at 1,400 feet alti¬
tude. It was a vigorous, healthy tree, near
the crest of the ridge in the Koa Zone. This
collection, filed as H. St. John no. 20,444,
fills in the gap and allows the conclusion
that the sandalwood originally extended the
length of the mountains and occurred on all
the ridges behind Honolulu. Its scarcity or
absence there now seems due to extermina¬
tion. To supply the sandalwood trade, the
large population of Honolulu would natur¬
ally seek the trees on the nearest accessible
ridges, which were those behind the city.
The resulting destruction of the trees by
cutting as well as by heavy and destructive
grazing by cattle nearly exterminated the
sandalwood in these areas.

The former territorial superintendent of
forestry, Charles S. Judd (1927: 43), in
giving a detailed account of the sandalwood,
stated :

It is not likely, however, that sandalwood will
soon, if ever, become much of a factor as a natural
resource in the Territory because of the fact that
it is of very slow growth, and before it can be
artificially propagated with success, much more
must be learned of its alleged parasitic habit of
growth.

The sandalwood is widespread and occa¬
sional in the Waianae Mountains. In the
Koolau Range (except behind Honolulu) it
occurs on the lee side on nearly every sec¬
ondary ridge, and is actually common at the
lower edge of the forest in the Koa Zone
and occasional up into the forest. Young
trees are numerous, but so, too, are old trees.

No data have been published as to the
rate of growth of this tree. At Kalauao

16

PACIFIC SCIENCE, January, 1947

there are trees 20 feet tall, 6 inches in diam¬
eter; at South Opaeula Gulch trees 20 feet
tall, 1 foot in diameter; at Laie-Malaekahana
ridge a grove with trees up to 20 feet tall, 1
foot in diameter; and at Puu Peahinaia a
tree 30 feet tall, 18 inches in diameter.
Charles S. Judd (1939: 36) reported and
printed a photograph of the largest tree he
found, in Waola Valley, Kawailoa, 30 feet

Fig. 2. Largest known sandalwood tree ( Santa -
lum Freycinetianum ) on Oahu, 28 inches in diam¬
eter, Kawailoa, September 28, 1943. The epiphytes
are Nephrolepis exaltata. The figure is Harold St.
John.

high, 21.3 inches in diameter. Territorial
forest ranger Tom McGuire and the writer
found a still larger one on September 28,
1943 (Fig. 2). It is near the north crest of
a northern side ridge descending from the
Anahulu Trail about one-fourth mile inside
the Forest Reserve boundary at about 1,300
feet altitude. The tree had many branches
and a wide spreading crown. As its trunk
forked 4 feet from the ground, it was meas¬
ured at 3 feet from the ground. It was 25
feet tall, 78^ inches in circumference, 28
inches in diameter.

RATE OF GROWTH

It is natural to inquire as to the rate of
growth of S. Freycinetianum. A wood sec¬
tion was taken from a branch iy^ inches in
diameter, collected in 1932 by the writer at
Paumalu on the Pupukea-Kahuku Trail at
the north end of the Koolau Range. The
wood was so hard that it was necessary to
boil it for 3 days before it could be cut with
a microtome knife. To the naked eye, faint,
fairly regular rings were visible in a cut stub
or in the transverse, stained microscopic sec¬
tion. Under the low-power lens of the com¬
pound microscope, these rings were also
faintly visible. The wood was dense, of
closely massed tracheids, small in diameter,
thick walled, and uniform in all parts of the
rings. The vessels were in radial rows, but
irregularly spaced. The ray cells were uni¬
form. The distinct banding which distin¬
guished the rings was due to a grouping of
cells with dark contents, probably tannin.
The 15 or so rings might be annual rings,
but this is not certain. Conditions are favor¬
able for growth at all times of year on Oahu,
as there are never freezing temperatures.
There are, nevertheless, definite seasons to
which all plants respond, flowering mostly
in the spring and summer. There is a belief
that these seasons are determined by a rainy
and a dry season, but this does not seem to
be substantiated. Weather records made

Sandalwood in Hawaii — St. John

17

available by the Hawaiian Pineapple Com¬
pany give data for three stations in the gen¬
eral region from which the wood sample was
taken: at Opaeula, Paumalu, and Waimea
No. 2. The last, in the upper pineapple
fields by the lower edge of the forest above
Waimea Camp at about 900 feet elevation,
was the closest. Its yearly rainfall was very
unequal in successive years. There were, of
course, periods of greater rainfall, but at
times two or three such wet periods occurred
during a single year. These came often in
December and August, but not constantly.
Considerable differences between the rain-

Fig. 3. Sandalwood ( Santalum Freycinetianum )
tree 16 inches in diameter, Anahulu Trail, Kawai-
loa, 1,500 feet altitude, September 28, 1943, in
lower, open forest, with dense fern undergrowth
of Cibolium Chamissoi and Dicranopteris linearis.
The figure is Tom McGuire.

fall even at adjacent stations indicate that the
rain comes mostly from local showers. Sun¬
shine and temperature records were not kept,
but these would doubtless show a correlation
with the rainfall. The rainfall records fail
to show any regular annual maxima which
could be correlated with growth rings. The
rings noted in the sandalwood specimen are
probably not annual rings, but rather wet
season or sunny season rings, of which there
are one or several each year. Hence, the
rings do not clearly indicate the age of the
stem. Long-term, precise observations are
needed on trees grown beside a weather sta¬
tion before one can interpret rings such as
those noted in the sandalwood.

Most of the trees of S. Freycinetianum
known today are less than 18 inches in diam¬
eter (Fig. 3). It seems probable that all or
nearly all of these have grown since the end
of the Sandalwood Era in 1830. Larger trees
such as the one on the Anahulu Trail, 28
inches in diameter, may well have been vig¬
orous young trees that escaped destruction
at that time.

LOCATION OF ORIGINAL SANDALWOOD
FORESTS

The present stands of S. Freycinetianum
on Oahu are at the lower edge of the dry
forest, and the tree is common, beginning at
1,000 or 1,200 feet altitude. Below .this are
outlying stations at much lower altitude: at
Kawailoa, by the rim of the gulch north of
the road running straight inland from Ash¬
ley, are healthy young trees at elevations as
low as 600 feet (specimens collected Sep¬
tember 28, 1943, St. John no. 20,364).
This is 500 feet lower than, and 2 or 3 miles
distant from, the lower edge of the forest.

Another record in the Koolau Range is
of a single tree 20 feet tall, 18 inches in
diameter, at 550 feet elevation on the north
slope of Kipapa Gulch, 1%. miles above the
old territorial highway, and 2 1/> miles below
the lowest forest stand in the gulch at about

18

PACIFIC SCIENCE, January, 1947

750 feet altitude. This record was an ob¬
servation made by E. Y. Hosaka while study¬
ing the flora and vegetation of Kipapa
Gulch.

A third low station is in the Waianae
Mountains, where a single 20-foot tree oc¬
curs in a dry lowland gulch below Puu Kuua
at 500 feet elevation (specimens collected
November 22, 1931, St. John no. 11,166).
This is 2,000 feet below, and 3 miles dis¬
tant from, the lower forest line near Palehua
and Puu Manawahua, where the tree is
again found. From the persistence of these
isolated trees in the lowlands, even good-
sized trees, there are indications that the
sandalwood was originally much more com¬
mon in the lowlands. There are also his¬
toric and other records which indicate the
same early distribution.

Commander Charles Wilkes (1845, vol.
4: 78-79), reporting on a trip of the nat¬
uralists W. P. Rich and W. D. Brackenridge
in 1840 from Waialua southward, says:

The next day they proceeded on their way to Ho¬
nolulu, across the plain between the two ranges
of mountains. This plain, in the rainy season,
affords abundance of food for cattle in three or
four kinds of grasses, and is, as I have before
remarked, susceptible of extensive cultivation by
irrigation from the several streams that traverse it.
The largest of the streams is the Ewa [Waikele].
Scraggy bushes of sandalwood and other shrubs
are now scattered over a soil fit for the cultivation
of sugar-cane and indigo.

Even in the time of Kamehameha I the
sandalwood had been much depleted, so
that this monarch put a kapu (ban) on the
cutting of young trees. A contrary action,
but arising from the same scarcity, was the
burning of grassland or forest areas by the
natives. This was done on the central plain
of Oahu in order to detect standing or fallen
logs of sandalwood by the sweet odor of
their smoke. The flames, of course, killed
many young sprouts and seedlings and pre¬
vented the recovery of the depleted stands of
sandalwood or other trees.

B. Seemann (1853, vol. 2: 83), who vis¬
ited Oahu in May, 1849, wrote:

The Oahu Sandal-wood ( Santalum paniculatum,
Hook, et Arn.) [this Oahu species is now known
to be S. Freycinetianum Gaud.], the Iliahi, or
Laau ala (fragrant wood) of the Hawaiians, is
now to be found in only one place, called Kuaohe,
where it grows on the slopes of hills, close to the
sea. Of the splendid groves, with the produce of
which formerly so many ships were laden, but a
few isolated bushes, which do not exceed three feet
in height and an inch in diameter, remain, and
these would probably disappear had they not been
protected by the law, and thus escaped being con¬
verted into fuel.

There is also an exact German translation
of this passage (Seemann, "Die Flora von
Oahu," 1853: 31). The geographic name
"Kuaohe" is unidentified, though Thomas
G. Thrum (1904: 72) suggested doubtfully
that it might be Kaneohe. This is very un¬
likely because the two names are dissimilar
and the tree does not occur in that humid,
wet region. As no recorded geographic name
for any land division of whatever size is
known that coincides with "Kuaohe," it re¬
mains unidentified. The name means "bam¬
boo ridge."

The father of Edward Y. Hosaka is
Yahei Hosaka, a farmer who settled in 1908
in Kipapa Gulch, Waipio, on the valley
floor 31/2 mdes upstream from the bridge on
the old paved territorial highway. On both
sides of the gulch above his home the steep
slopes had a forest cover with many sandal¬
wood trees. The settlers cut the trees,
destroying the forest to clear the land for
the cultivation of pineapples. The koa
{Acacia Koa) and ohia lehua {Metrosideros
collina ssp. polymorpha ) were used for fire¬
wood. As the sandalwood did not make
good charcoal, they cut and burned it for
mosquito punk.

According to a statement of Dr. H. L.
Lyon in 1942, cord wood was formerly cut
and hauled to Honolulu, where it was used
for cooking and heating. In 1910-11 he
examined piles of this cord wood in Wa-

Sandalwood in Hawaii — St. John

19

hiawa, cut in the near-by gulches and ridges
and destined for the kitchen stoves of Hono¬
lulu. A considerable proportion of the logs
were sandalwood.

Numerous writers have recorded the deci¬
mation of the native forests on the Hawaiian
Islands by many agents such as sandalwood
cutters, grazing cattle, and fire, and by lum¬
bering for firewood, timber, or charcoal.
Hence, there is no doubt that the forests,
until the coming of the white man, covered
a large portion of the islands. On Oahu the
forest did not stop at the 1,000- or 1,500-
foot line, but came down in places right to
the sea. What trees made up this forest can¬
not be completely known, but the trees now
found at the lower, drier edge of the forest
probably combined to form a forest over
part of this area. They are the koa ( Acacia
Koa ), olopua ( Osmanthus sandwicensis) ,
iliahi ( Santalum Freycinetianum) , aalii
(Dodonaea viscosa ), ohia ha ( Eugenia
sandwicensis ), ohia lehua ( Metrosideros
collina ssp. polymorpha vars.), pukeawe
[Styphelia T ameiameiae) , kopiko [Straus sia
Mariniana ), and naupaka kuahiwi [Scaevola
Gaudichaudiana ) . The Leilehua or Scho¬
field plains between the two mountain for¬
ests were densely and continuously forested.
On the lowest, driest slopes, an open forest
or savanna doubtless spread, largely of wili-
wili [Erythrina sandwicensis ) but also con¬
taining the ohe ( Reynoldsia sandwicensis) .

Charles S. Judd (1927: 43) expressed the
opinion that the "sandalwood evidently
never occurred in pure stands, but was found
in small groups or as scattered individuals";
but Seemann (1853, vol. 2: 83) wrote of the
splendid groves which formerly existed. To
the writer, Judd’s conclusion seems a better
description of the present status than a re¬
construction of the past. Professor J. F.
Rock (1916: 13-14) had the same view as
Seemann :

Sandalwood must certainly have formed a large
percentage of the tree growth in the drier regions

or mixed forests, in the early days, before the
value of the wood became known to the natives of
these Islands. It must have existed in pure stands,
or as forests, or it would have been next to impos¬
sible to export as much as $400,000 worth per
annum.

S. Freycinetianum has recovered so far to¬
day as to be common and widespread, and at
numerous parts of Kawailoa, Waimea, and
Kaunala, it now forms the lower forest at
1,000 to 1,200 feet altitude. This is at the
lower border of the present native forest,
but not the natural lower limit of the sandal¬
wood in former times. As E. Y. Hosaka
observed during his 92 trips from 1931 to
1934 to study the vegetation of Kipapa
Gulch, the sandalwood grew well there at
the present lower edge of the forest at about
1,200 feet altitude. Lower down on the
sides of the gulch at 700 and 800 feet alti¬
tude there were other sandalwood trees,
definitely healthier and more vigorous than
those above. This was true despite the fact
that the surroundings were nearly denuded
of native plants and trees, and were more
exposed, drier, and more eroded.

The heaviest stands of former times were
doubtless at lower elevations, below the Koa
Forest Zone of that time. The existing out¬
lying stations of old trees as low as 500 feet
elevation, and the greater luxuriance at
lower elevations, give an indication of where
the old sandalwood stands were. As is true
today, there were small stands on ridges or
dry habitats far up into the forested moun¬
tains up to 2,000 or 2,400 feet altitude.
Doubtless the large stands grew at even
lower elevations than those of today, ap¬
proaching or even reaching the shore in
regions of moderate rainfall such as Kawai¬
loa and Waimea, at the northern end of the
Koolau Range. To judge by the many thou¬
sands of piculs of heartwood gathered, san¬
dalwood must have been abundant. Hence,
the writer deduces that there were heavy
stands of sandalwood either abundant in, or
dominant in, a forest zone from about 300

20

PACIFIC SCIENCE, January, 1947

to about 1,000 feet altitude, below the Koa
Zone and above the Wiliwili Zone. Prob¬
ably the tree occurred on the lower, dry
slopes of nearly every secondary ridge lead¬
ing from the Waianae Mountains and on
those leading from the leeward side and the
north end of the Koolau Range. The north¬
ern and southern slopes of the Schofield sad¬
dle apparently had much more extensive
stands of sandalwood, from Waimano to
Honouliuli, and from Pupukea to Makaleha.
This is a deduction from various types of
evidence here assembled, which, for the first
time, give an indication of the location of
the principal stands of sandalwood on Oahu.

Though decimated by the sandalwood
trade, the tree persisted on Oahu, survived
the overgrazing and forest recession, and is
now common and widespread at its former
upper limit, now the lower forest line on the
lee side of the Koolau Range and on both
sides of the Waianae Mountains. Most of
its present stations are protected from
further destructive exploitation by their sit¬
uation within the Territorial Forest Reserves.

REFERENCES

Bennett, George. An account of the sandal
wood tree (Santalum), with observations on
some of the botanical productions of the Sand¬
wich Islands. Mag. Nat. Hist. ( Loudon's ) 5:
255-261, 1832.

- Gatherings of a naturalist in Australasia.

xii+456 p., 8 pi., 24 fig. John Van Vorst, Lon¬
don, I860.

- , 1831. See Montgomery.

Bloxam, Andrew. Diary of Andrew Bloxam,
naturalist of the "Blonde.” 96 p., 5 fig., 9 pi.
Bernice P. Bishop Mus. Spec. Pub. 10. Hono¬
lulu, 1925.

Bradley, Harold W. The American frontier in
Haivaii: The pioneers, 1789-1843 . xi-f-488 p.
Stanford Univ. Press, 1942.

Brigham, William T. Ka hana kapa: The mak¬
ing of h ark-cloth in Hawaii, iv-f-273 p., 131 fig.,
48 pi., 27 col. pi. Bernice P. Bishop Mus. Mem.
3. Honolulu, 1911.

Coulter, John Wesley. A gazetteer of the Ter¬
ritory of Hawaii. 241 p., 13 fig. Hawaii Univ.
Res. Pub. 11. Honolulu, 1935.

Davis, C. Sandwich Islands. North Amer. Rev.
3: 52-53, 1816.

Delano, Amasa. A narrative of voyages and
travels. 598 p., 3 pi. E. G. House, Boston, 1817.

Ellis, William. Narrative of a tour through Ha¬
waii. 442 p. H. Fisher & P. Jackson, London,
1826.

Fischer, C. E. C. Where did the sandalwood tree
(Santalum album Linn.) evolve? Bombay Nat.
Hist. Soc. Jour. 40(3) : 458-466, 1938.

Hunnewell, James. Voyage in the brig " Bor¬
deaux Racket,” Boston to Honolulu, 1817, and
residence in Honolulu, 1817-1818. 19 p. Ha¬
waii. Hist. Soc. Papers 8. Honolulu, 1895.

Ingraham, Joseph. Log of the brig rfHope,” 1791.
36 p., 2 pi. Hawaii. Hist. Soc. Reprints 3. Ho¬
nolulu, 1918.

Judd, Charles S. The natural resources of the
Hawaiian forest regions and their conservation.
Hawaii. Forester and Agr. 24(1): 40-47, 2 pi.,
1927.

- Report of Territorial Forester. Hawaii

Bd. Commrs. Agr. and Forestry Bien. Rpt. 1938:
32-58, 4 pi., 1939.

Kuykendall, Ralph S., and Herbert E. Greg¬
ory. A history of Hawaii, x-j-375 p., illus.
Macmillan, New York, 1926.

Mathison, Gilbert Farquhar. Narrative of a
visit to Brazil, Chile, Peru, and the Sandwich
Islands during-the years 1821 and 1822. xii-f 478
p. C. Knight, London, 1825.

Mesick, Lilian Shrewsbury. The kingdom of
Hawaii. 400 p. Porter, Honolulu, 1934.

Montgomery, James (ed.). Journal of voyages
and travels by the Rev. Daniel Tyerman and
George Bennet, Esq. 2 vol. F. Westley & A. H.
Davis, London, 1831.

"Old Quarter Master.” Thirty-six years of a
seafaring life. 336 p. W. Woodward, Portsea
(England), 1839.

Rock, Joseph F. The sandalwoods of Hawaii; a
revision of the Hawaiian species of the genus
Santalum. Hawaii Bd. Commrs. Agr. and For¬
estry, Div. Forestry Bot. Bui. 3: 1-43, 13 pi.,
1916.

Seemann, Berthold. Narrative of voyage of -
H.M.S. " Her aid” during the years 1843-31. 2
vol. Reeve, London, 1853.

- Die Flora von Oahu. Bonplandia 1:

30-32, 1853.

Skottsberg, C. The geographical distribution of
the sandalwoods and its significance. Fourth
Pacific Sci. Cong. Proc. (Java), 3: 435-440,
map, 17 fig., 1930.

Thrum, Thomas G. The sandalwood trade of
early Hawaii. Thrum’s Hawaiian Almanac and
Annual for 1903, 43-74. Honolulu, 1904.

Wilkes, Charles. Narrative of United States ex¬
ploring expedition, 1838-1842. 5 vol. Lea &
Blanchard, Philadelphia, 1845.

The Tsunami of April 1, 1946, in the Hawaiian Islands

G. A. Macdonald, F. P, Shepard, and D. C. Cox1

INTRODUCTION

The tsunami which struck the shores of
the Hawaiian Islands on the morning of
April 1, 1946, was the most destructive, and
one of the most violent, in the history of the
islands. More than 150 persons were killed,
principally by drowning, and at least 161
others were injured. Property damage
reached about $25,000,000.

The wave attack on Hawaiian shores was
far from uniform. The height and violence
of the waves at adjacent points varied greatly,
and not always in the manner which would
have been expected from superficial inspec¬
tion and a study of the existing literature on
tsunamis. Therefore, a detailed study of the
effects of the tsunami has been made, in an
effort to understand the observed variations,
and in the hope that the principles estab¬
lished may help lessen the loss of life and
property in future tsunamis. Space is not
available in the present short paper to discuss
findings in detail, or even to present all the
evidence for all the conclusions. These mat¬
ters will be treated in detail in a longer paper
( Shepard, Cox, and Macdonald, in prepara¬
tion) .

Acknowledgments: We wish to thank the
many persons who furnished information
during the course of the field study. We are
also especially grateful to M. H. Carson, H.
S. Leak, H. W. Beardin, and W. K. Sproat,
who supplied measurements of the high-

1 U. S. Geological Survey, Scripps Institution of
Oceanography, and Hawaiian Sugar Planters’ As¬
sociation Experiment Station, respectively. This
paper is published by permission of the Director,
Geological Survey, U. S. Department of the In¬
terior.

water level in areas not visited by us; H. W.
Iversen and J. D. Isaacs, who supplied addi¬
tional measurements on Oahu; A. F. Robin¬
son and Dexter Fraser, who furnished de¬
scriptions of the wave effects on Niihau and
Lanai, respectively; the Hawaii County Engi¬
neer’s Office, which supplied a map showing
the extent of flooding in Hilo; and the U. S.
Coast and Geodetic Survey, which supplied
data on the earthquake and the record of the
Honolulu tide gage, and permitted the use,
in advance of publication, of C. K. Green’s
manuscript on the tsunami along the shores
of North and South America. Howard A.
Powers, Seismologist of the Volcano Obser¬
vatory at Hawaii National Park, aided greatly
in the investigation on the island of Hawaii.
Miss Maude Jones, Archivist of the Territory
of Hawaii, and Miss Margaret Titcomb,
Librarian of the Bishop Museum, aided in
locating records of past waves. C. K. Went¬
worth and H. S. Palmer aided greatly in dis¬
cussions. Wentworth and Walter Munk
read and criticized the manuscript. J. Y.
Nitta prepared the illustrations.

DEFINITION OF "TSUNAMI”

The name 'Tsunami”2 is applied to a long-
period gravity wave in the ocean caused by a
sudden large displacement of the sea bottom
or shores. A tsunami is accompanied by a
severe earthquake, but the earthquake does
not cause the tsunami. Rather, both are
caused by the same sudden crustal displace¬
ment. The waves of a tsunami have a period

2 Also spelled "tunami,” the Japanese equivalent
of the letter / being pronounced ts in English. It
appears preferable, however, to use the phonetic
spelling in English, avoiding thereby much incor¬
rect pronunciation.

21

22

PACIFIC SCIENCE, January, 1947

of several minutes to an hour as contrasted
with several seconds for ordinary storm waves
caused by wind, a wave length of scores of
miles as contrasted with less than 500 feet
for wind waves, and a speed of hundreds of
miles an hour as contrasted with less than 60
miles an hour for wind waves. Tsunamis
are also sometimes termed "seismic sea
waves,” and are popularly known as "tidal
waves.” The latter term is patently undesir¬
able, as the waves have no connection what¬
ever with the tides. "Tsunami” is used herein
in preference to "seismic sea wave” because
of its greater brevity, and because the etymo¬
logical correctness of the term "seismic sea
wave” appears open to question.3

HISTORY OF TSUNAMIS IN HAWAII

Tsunamis probably reach Hawaiian shores
on an average of more than one a year. Most
of these are small, however, and generally
escape notice except when their record is
recognized on tide gages. Earlier tsunamis
in Hawaii have been discussed by Jaggar
(1931: 1-3) and Powers (1946: 3). The
accompanying table lists all the tsunamis
noticed on Hawaiian shores, in the period of
written history, of which record could be
found, together with their sources if known.
A total of 27 are listed, or an average of one
every 4.7 years since 1819. Most of them,
however, did little damage. During the same
interval there are listed five severe tsunamis
which caused extensive damage, an average
of one every 25.6 years.

Other violent waves have been termed
"tidal waves” in the newspapers, but were
more probably storm waves. Such were the
wave which hit Maliko, Maui, on January
28, 1895, and those which struck Kauma-

3 The adjective "seismic” is derived from the
Greek root seismos, meaning earthquake, and is
defined as pertaining to, produced by, or charac¬
teristic of an earthquake. The waves in question
are not, however, characteristic of most earth¬
quakes, even those of submarine origin, and are
not produced by earthquakes.

lapau on Lanai, and Nawiliwili on Kauai, on
May 30, 1924.

It will be noted that only two of the 27
tsunamis listed in the table were of local
origin. With the exception of the numerous
volcanic earthquakes on the island of Ha¬
waii, which seldom cause tsunamis, the
Hawaiian region is only moderately active
seismically (Gutenberg and Richter, 1941:
84-85 ) . The great majority of the tsunamis
reaching Hawaii originate in the highly
seismic border zone of the Pacific. Of the
22 tsunamis from known sources listed in
the table, five came from near South Amer¬
ica, one from near Central America, one
from near California, three from near Alaska
and the Aleutian Islands, five from near
Kamchatka, three from the Japanese area,
and one from near the Solomon Islands. Of
the five severe tsunamis, three originated

TABLE 1

Hawaiian Tsunamis

DATE

SOURCE

DAMAGE

IN HAWAII

AVERAGE

SPEED OF

WAVES

1819 Apr. 12

nearest coast

Unknown

Unknown

mi. per hr.

1837 Nov. 7

South America

Severe

—

1841 May 17

Kamchatka

Small

—

1868 Apr. 2

Hawaii

Severe

—

1868 Aug. 13

South America

Severe

—

1869 July 25

South Amer¬

Moderate

—

1872 Aug. 23

ica (?)

Hawaii

Small

1877 May 10

South America

Severe

—

1883 Aug. 26

East Indies

Small

—

1896 June 15

Japan

None

478

1901 Aug. 9

Japan (?)

None

—

1906 Jan. 31

Unknown

None

— —

1906 Aug. 16

South America

Small

—

1918 Sept. 7

Kamchatka

Small

456

1919 Apr. 30

Unknown

None

—

1922 Nov. 11

(distant)
South America

None

450

1923 Feb. 3

Kamchatka

Moderate

432

1923 Apr. 13

Kamchatka

None

438

1927 Nov. 4

California

None

462

1927 Dec. 28

Kamchatka

None

438

1928 June 16

Mexico

None

462

1929 Mar. 6

Aleutian Is.

None

492

1931 Oct. 3

Solomon Is.

None

447

1933 Mar. 2

Japan

Small

477

1938 Nov. 10

Alaska

None

496

1944 Dec. 7

Japan

None

425

1946 Apr. 1

Aleutian Is.

Severe |

490

23

In Hawaii, it was recorded on the instrument
of the U. S. Coast and Geodetic Survey lo¬
cated on the campus of the University of
Hawaii in Honolulu, and on those of the
Hawaiian Volcano Observatory at Kilauea
on Hawaii. The epicenter of the earthquake
has been located by the Coast and Geodetic
Survey at latitude 53.5° N. and longitude
163° W., and the time established as lh
59m a.m. Hawaiian time ( 12h29m Greenwich
time) (Bodle, 1946: 464). It may be as¬
sumed that the tsunami originated at the
same place and time as the earthquake. The
place of origin was thus 2,241 miles N. 8.5°
W. of Honolulu, and 2,375 miles N. 12° W.
of Hilo (Fig. 1).

Fig. 1. Map of the Pacific basin, showing the position of the Hawaiian Islands, the place of origin
of the tsunami of April 1, 1946, and the distribution of seismically active belts around the Pacific in
which tsunamis are likely to originate.

Tsunami of April 1, 1946 — Macdonald et al.

near the coast of South America and one in
the Aleutian area, and one was of local
origin. One tsunami of moderate intensity
came from near Kamchatka, and another
probably from South America.

GENERAL FEATURES OF THE APRIL, 1946,
TSUNAMI: ORIGIN AND NATURE
OF THE WAVES

The tsunami of April 1, 1946, was caused
by a movement of the sea bottom on the
northern slope of the Aleutian Deep, south
of Unimak Island. The same crustal move¬
ment gave rise to a violent earthquake, re¬
corded on seismographs all over the world.

24

PACIFIC SCIENCE, January, 1947

The time of arrival of the waves in the
Hawaiian Islands is known with certainty
only at Honolulu. The record of the Hono¬
lulu tide gage (Fig. 2) shows that the first
rise started at about 6:33 a.m. (Green, C. K.,
1946: 491), though the exact time can¬
not be stated closer than 2 or 3 minutes.
The drum of the water-stage recorder at the
Waimea River, on Kauai, revolves too slowly
to give an accurate indication of time, but the
first rise appears to have started there at
about 5:55. At Hilo, electric clocks were
stopped at 7:0 6, and a brief power failure
occurred at 7:18. These have been inter¬
preted by Powers (1946: 2), probably cor¬
rectly, as the time of arrival of two wave
crests at Hilo. From other considerations,
discussed briefly elsewhere (Shepard, Mac¬
donald, and Cox, in preparation), it appears
probable, however, that the crest at 7:06

was the second wave at Hilo, not the first.
If so, allowing for the observed 15-minute
interval between later waves, the first rise
at Hilo probably started at about 6:45. Com¬
puted from these times of arrival, the ap¬
proximate average speed of the tsunami from
its origin to Honolulu and Hilo was, respec¬
tively, 490 and 498 miles an hour. On en¬
tering shallow water the waves decreased
greatly in speed. The waves moving up
Kawela Bay, on Oahu, were estimated by
Shepard to be moving only about 15 miles
an hour. Similar low speeds near shore were
reported by other observers, and are com¬
parable to the speed of 20 miles an hour
recorded in San Francisco Bay (Green,
C. K., 1946: 492).

The interval between the first and third
wave crests, as recorded on the Honolulu tide
gage (Fig. 2), was about 25 minutes, indi-

HOURS

Fig. 2. Record produced on the tide gage in Honolulu Harbor by the tsunami of April 1, 1946.

Tsunami of April 1, 1946 — Macdonald et al.

25

eating an average interval between early wave
crests of approximately 12.5 minutes. The
interval between the first wave crest and the
succeeding trough was 7.5 minutes, however,
indicating a wave period of 15 minutes at
the beginning of the disturbance. This cor¬
responds with the mean wave period of 15.6
minutes found by Green ( 1946: 499) at
Honolulu and eight other stations on the
coasts of North and South America. At the
mouth of Nuuanu Stream in Honolulu, C. K.
Wentworth observed an interval of approxi¬
mately 15 minutes between successive bores
ascending the stream, and a wave period of
about 15 minutes was observed by J. B. Cox
and D. C. Cox at Waikiki at about 7 :45 a.m.
Observations elsewhere were poor, but in
general indicated an interval not far from
15 minutes between the early waves of the
series. The interval between later waves at
Honolulu (Fig. 2) and elsewhere was
shorter and less regular, probably because of
the arrival of chains of waves traveling by
somewhat different routes, refracted around
different sides of islands, and reflected at
various points, as well as traveling by the
most direct route. Probably contributing to
the irregularity of later waves were wind
waves and also the free-period oscillations,
in harbors and channels, known as "seiches.”
If the period of the waves is assumed to be
15 minutes, and the average speed to be
about 489 miles an hour, the average wave
length from crest to crest was about 122
miles.

Direct observations on the height of the
waves in the open sea are lacking, but theo¬
retical considerations indicate that the height
probably did not exceed 2 feet from crest
to trough.4 If so, the small height combined
with the very great wave length should have
made the waves imperceptible to ships at sea.
That such was indeed the case is indicated

4 Based on the assumption of a 10-foot wave in
10 feet of water, and the variation of the wave
height inversely as the fourth root of the depth.

by the fact that the master of a ship lying
offshore near Hilo could feel no unusual
waves, although he could see the great waves
breaking onshore. Crews of fishing boats in
the Hawaiian area also reported no unusual
conditions at the time of the tsunami, al¬
though heavy storm waves were running.
The few reports of violent waves of great
height from ships at sea were probably occa¬
sioned by storm waves, together with the
knowledge that a tsunami was taking place.

The nature of the waves sweeping up on
to Hawaiian shores varied greatly from place
to place. At some places the water rose
gently, flooding over the coastal lands with¬
out the development of any steep wave front.
At such places most of the damage resulted
from the violent run-back of the water to the
sea. At some localities, although the gen¬
eral water surface rose gently, ordinary storm
waves moved in over the top of the broad
swells of the tsunami, and there at least part
of the damage was caused by the storm
waves. At most places, however, the waves
of the tsunami swept toward shore with
steep fronts and great turbulence, causing a
loud roaring and hissing noise. Locally, the
wave closely resembled a tidal bore, the steep
front rolling in over comparatively quiet
water in front of it. Behind the steep front,
the wave crest was broad and nearly flat,
with smaller storm waves superimposed upon
it. Such bores were best developed in bays
and estuaries, but waves of closely similar
form were observed crossing shallowly sub¬
merged reefs upon otherwise open coasts.

At many places the violence of the waves
moving shoreward was sufficiently great to
tear loose heads of coral and algae, up to 4
feet across, and toss them onto the beach
as much as 15 feet above sea level. Locally,
blocks of reef rock weighing several tons
were quarried at the outer edge of the reef
and thrown onto the reef surface.

Between crests, the water withdrew from
shore, exposing reefs, coastal mud flats, and

26

PACIFIC SCIENCE, January, 1947

harbor bottoms for distances up to 500 feet
or more from the normal strand line. The
outflow of the water was rapid and turbu¬
lent, making a loud hissing, roaring, and
rattling noise. At several places houses were
carried out to sea, and in some areas even
large rocks and blocks of concrete were car¬
ried out onto the reefs. Sand beaches were
strongly eroded by the outgoing water.
People and their belongings were swept to
sea, some being rescued hours later by boats
and life rafts dropped from planes.

At a few places, generally but not exclu¬
sively on the sides of the islands away from
the wave origin, the first wave was reported
to have been the highest. At those places,
the rise was generally of the quiet sort.
There are, however, no instrumental records
showing the first wave to have been the high¬
est, and it is possible that at places reporting
the first wave as the highest, earlier waves
may have been overlooked. Much more gen¬
erally the third or fourth wave was reported
to have been the highest and most violent.
The third crest was the largest at the Hono¬
lulu tide gage (Fig. 2). At other localities
the sixth, seventh, or eighth waves were said
to have been the highest. At Waimea River,
Kauai, the sixth crest was higher than any
other, both in absolute level and in its height
above the preceding and succeeding troughs.

In general, if not everywhere, the size and
violence of the waves increased to a maxi¬
mum with the third to eighth waves. The
oscillations then gradually decreased in am¬
plitude over a period of at least 2 days, but
with occasional waves which were larger
than those just before and after them. Such
temporary increases in wave height prob¬
ably resulted from mutual reinforcement by
the essentially simultaneous arrival, in phase,
of waves which had traveled different paths,
or from the coincidence of tsunami waves
with storm waves or seiche oscillations.

Measures of the height of the waves ap¬
proaching shore in shallow water, but before

they dashed up on shore, are poor. At
Kawela Bay, Oahu, Shepard estimated the
height of the waves advancing across the reef
to have been as much as 18 feet, and observ¬
ers estimated the height of the waves cross¬
ing the reef off Lanikai, on Oahu, to have
been about 7 feet. Photographs taken at Hilo
show the top of the breakers to have been 2 5
feet above the normal bay surface where they
struck Cocoanut Island, but the waves may
have increased considerably in height in
crossing the breakwater, and the effect of
dashing up on the shore was probably already
present, further exaggerating the height.
Photographs of some of the late waves at the
mouth of the Wailuku River, in Hilo, show
them to have been 6 to 8 feet high (Plate
8), and early waves undoubtedly were
higher. In general, these heights correspond
fairly closely with the measured heights to
which the water dashed on the shore at those
localities. At any rate it appears clear that
the waves not only slowed down, but in¬
creased in height on entering shallow water.
George Green (1838: 457-462) states that
the wave height varies inversely as the fourth
root of the depth of the water.

Most observers reported the first move¬
ment on Hawaiian shores to have been a
withdrawal of the water. However, the only
available instrumental records, at Honolulu
and Waimea, both indicate the first move¬
ment to have been a rise. The instrumental
records are probably more reliable than the
reports of untrained observers. The initial
rise at Honolulu was small (Fig. 2), and a
similar small rise at other localities may eas¬
ily have been overlooked. Certainly it would
have been less impressive than the large
withdrawal of the water from shore as the
succeeding trough approached. It is inter¬
esting to note that the records of tide gages
along the coasts of North and South America
obtained by C. K. Green (1946: 497) all
show the initial movement to have been a

Tsunami of April 1, 1946 — Macdonald et al.

rise, with amplitude of about one third that
of the ensuing trough.

HEIGHTS REACHED BY THE WAVES
ON HAWAIIAN SHORES

Measurements of high-water marks have
been made around the shores of all five
major islands of the Hawaiian group. The
measured heights are shown on Fig. 3 to 7.
All heights are stated in feet above lower
low water. At each point sea level was esti¬
mated, the height of the high-water mark
above that level was measured by means of
hand level or steel tape, and the measure¬

27

ment reduced by means of tide tables to
height above lower low water. Some inac¬
curacy undoubtedly has entered in the esti¬
mation of mean sea level, but it is believed
that the heights are probably accurate to
within 1 foot. The levels measured include:
points indicated by eyewitnesses as the upper
limit of the water, lines of flotsam or swash
marks, the upper limits of soil and vegeta¬
tion scouring, levels of consistent scratching
and barking on trees, and the upper level of
staining on the walls of buildings.

The measured heights of high-water
marks range from 53 feet at Pololu Valley

Fig. 3. Map of the island of Kauai, showing heights reached by the water during the tsunami of
April 1, 1946. Heights are in feet above lower low water.

28

PACIFIC SCIENCE, January, 1947

on Hawaii, 54 feet at Waikolu Valley on
Molokai, and 45 feet at Haena and Kilauea
Point on Kauai, to 2 feet at Kaunakakai on
Molokai, 2 feet at Milolii and Hoopuloa on
Hawaii, and less than 2 feet at the head of
Kaneohe Bay on Oahu. Causes of the varia¬
tions in height will be discussed in a later
section.

Most of the heights measured are, of
course, not the heights of the actual waves,
but rather the heights to which the water
was driven on shore. On a vertical cliff di¬
rectly across the path of the wave, this height
may theoretically amount to twice the height
of the actual wave. On slopes less than ver¬
tical, or on cliffs at an angle to the direction
of wave advance, it should be somewhat less
than twice the wave height. This measure
represents the height of dash of solid water,
but very abundant spray may be thrown
much higher. Moreover, storm waves riding
on the crest of the broader swells of the
tsunami undoubtedly added in places to the
height to which water dashed on shore.
There are places where normal trade-wind
waves are flung to a height nearly as great
as that reached by the tsunami, and many
places, particularly on shores facing away
from the origin of the tsunami, where waves
of heavy storms reached appreciably higher
than did the waves of the tsunami.

It is not possible to make reliable esti¬
mates of the magnitudes of these complicat¬
ing factors, as there are too many unknown
elements involved. However, it is probable
that most of the water heights recorded for
the tsunami on the northern and eastern
sides of the islands were appreciably in¬
creased by these factors.

FACTORS INFLUENCING THE HEIGHTS
AND INTENSITIES OF THE WAVES

It may be assumed that the size and speed
of the waves approaching the islands from
the open ocean to the north were essentially
the same throughout the length of the Ha¬

waiian Archipelago. Differences in height
reached by the water and in violence of wave
attack along Hawaiian shores must be at¬
tributed to local influences modifying the
size and behavior of the waves.

The factors found to have affected the
height and intensity of the waves during the
tsunami of April 1, 1946, are:

1. Orientation of the coast line with respect
to the point of origin of the tsunami.

2. Shape of the island.

3. Exposure to storm waves.

4. Submarine topography.

5. Presence or absence of reefs.

6. Configuration of the coast line.

7. Merging of waves from different direc¬
tions, or of different types.

Orientation of the coast line with respect to
the point of origin of the tsunami. — In gen¬
eral, the heights reached by the water were
greatest on the sides of the islands facing
the origin of the waves, and lowest on the
sides away from the wave origin. This is
evident from even a cursory inspection of
the maps (Fig. 3 to 7). Heights average
consistently greater on the northern than on
the southern sides of the islands. All the
extreme heights were measured on the north¬
ern or northeastern sides. Conversely, most
of the lowest figures were found on the
southern and southwestern sides. It appears
almost self-evident that this should be so.
Waves striking northern shores retain their
full force, whereas the refracted waves strik¬
ing southern shores suffer a diminution in
force and height. This effect is discussed for
wind waves in Breakers and surf (U. S. Navy
Hydrographic Office, 1944: 12-13). No
wave can be refracted or reflected without
losing some of its force.

Shape of the island. — Waves were refracted
around circular or nearly circular islands
much more effectively than around angular
or elongate islands. This fact had a marked
effect on the height and violence of waves on

Tsunami of April 1, 1946 — Macdonald et at.

29

the southern shores. Thus the water reached
considerably greater heights along the south¬
ern coast of the nearly round island of
Kauai ( Fig. 3 ) than along the southern coast
of the angular and elongate island of Molo¬
kai (Fig. 5), even though the heights along
the northern coast of Molokai were on the
average perhaps a little greater than those
on the northern coast of Kauai. The contrast
between the very high average height on the
northern coast of Molokai and the very low
average height on the southern coast is
greater than that between the two sides of
any other island, although the difference
between the extreme highs and lows is al¬
most exactly the same as on the island of
Hawaii (Fig. 7).

Exposure to storm ivaves. — At the time of
the tsunami, large storm waves had been run¬
ning for several days. As already pointed
out, these storm waves riding in on the backs
of the broad swells of the tsunami in places
undoubtedly increased the height to which
the water dashed on shore. Moreover, in
other places, where the rise in water level
due to the tsunami was gentle, storm waves
on top of the tsunami were responsible for
much of the damage. The generally greater
violence of the waves on the windward
(northern and northeastern) coasts as com¬
pared to that on the leeward coasts may have
been in considerable part the result of the
large storm waves which were driving in on
the windward coasts. Places on the wind-

Fig. 4. Map of the island of Oahu, showing the heights reached by the water during the tsunami
of April 1, 1946. Heights are in feet above lower low water.

30

PACIFIC SCIENCE, January, 1947

ward coasts which were sheltered from the
storm waves also experienced less violent
waves. Thus at Kalaupapa, on the sheltered
side of the peninsula on the windward side
of Molokai, both photographs and the testi¬
mony of observers indicate that the rise of 2 5
feet caused by the tsunami was not violent.
On the windward coasts, much of the rapid
variation in intensity of wave attack may
have resulted from the caprice of storm
waves.

Submarine topography. — Owing to their
great wave length, the waves were somewhat
affected by the ocean bottom throughout
their course. However, the effect of the bot¬
tom increased greatly as the waves moved
into shallow water, and caused a slowing of
the wave, an increase in its height, and a
steepening of its front. A direct evidence of
the increase in height of the waves in shal¬
low water was afforded by the lesser heights
reached by the water at the ends of certain
peninsulas projecting into deep water and
not prolonged seaward by pronounced
ridges, as compared with the heights on ad¬
jacent shores rising from shoal water. Thus
at the end of Kalaupapa Peninsula, on the
northern coast of Molokai (Fig. 5), the
water dashed only 7 feet above normal sea
level, distinctly less than do the waves of

ordinary storms; whereas on the coasts ris¬
ing from shoal water both east and west of
the peninsula, the water swept up to heights
of 30 to 54 feet. At the end of Keanae Pen¬
insula, on the northern coast of Maui (Fig.
6), the tsunami reached heights only a little
greater than large trade-wind waves.

Submarine ridges and valleys, particularly
those pointing toward the wave source, were
of great importance in their effect on the
strength of the waves. The best examples of
the effect of ridges are found on the north¬
ern coast of Kauai. A long ridge extends
in a direction slightly west of north from
Haena, to a depth of about 8,000 feet
(Plate 2). Another extends northeastward
from Kilauea Point, to even greater depths.
The greatest heights (45 feet) reached by
the water on the shores of Kauai were at
the heads of these two ridges (Fig. 3). An¬
other ridge extending northwestward from
the western coast of Kauai is probably re¬
sponsible for heights of 35 to 38 feet at its
head. Long ridges projecting from Kaena
and Kahuku Points on Oahu similarly caused
an increase in wave heights there as com¬
pared to the heights on both sides (Fig. 4) .
The ridges projecting eastward north of Hilo
Bay and at Cape Kumukahi on Hawaii had,
on the other hand, no such pronounced

Fig. 5. Map of the island of Molokai, showing heights (in feet above lower low water) reached
by the water during the tsunami of April 1, 1946.

Tsunami of April l, 1946 — Macdonald et al.

effect on the heights at their heads; but it
should be noted that they extend across the
general direction of wave advance, not
toward it.

The greater heights reached by the water
at the heads of submarine ridges are not
difficult to explain. The ridge has a greater
effect in limiting the movement of water
particles in the advancing wave than does
the deeper water alongside it. Consequently
the portion of the wave over the ridge is
retarded more than that away from the ridge,
and the wave front becomes bent, with its
concavity directed toward the ridge head.
The result is a focusing of wave force on the
shore at the head of the ridge (U. S. Navy
Hydrographic Office, 1944: 13).

Similarly, in moving toward shore along
the axis of a submarine valley, the part of

31

the wave in the deep water along the valley
axis moves faster than that in shallower
water on the two sides. In consequence the
wave front becomes bent, with its convexity
toward the valley head. In the vicinity of
the valley head the force lines ( orthogonals )
of the wave are diffused or spread apart, and
over any unit area the force of the waves
striking shore is greatly decreased.

An example of the effect of a submarine
valley in lessening the force of the waves
at its head is found at Kahana Bay, on Oahu
(Fig. 4). There the waves dashed to
heights of 11 to 17 feet on the coasts north
and south of the bay, but reached heights of
only 4 to 7 feet in the bay itself. A small
submarine valley extends 2 miles northeast¬
ward from the bay, to a depth of 150 feet.
An example on a much larger scale is af-

Fig. 6. Map of the island of Maui, showing heights (in feet above lower low water) reached by
the water during the tsunami of April 1, 1946.

32

PACIFIC SCIENCE, January, 1947

forded by the zone of small heights along
the northwestern shore of Kauai (Fig. 3), at
the head of a broad swale extending outward
to oceanic depths. The broad valley-like de¬
pression off the eastern coast of Hawaii south
of Hilo Bay probably also was somewhat
effective in reducing the heights reached by
the water along that coast. Although fairly
great, ranging from 16 to 19 feet, the
heights there are not much greater than those
reached by ordinary storm waves.

Presence or absence of reefs.— The presence
of a well-developed fringing reef appears
to have had a decided effect in reducing the
intensity of wave onslaught. Along the reef-
protected northern coast of Oahu the heights
reached on shore by the waves were on the
average decidedly less than on the unpro¬
tected northern coasts of Molokai and Ha¬
waii, or on the less protected northern coast
of Kauai. The best-developed coral reef in
the Hawaiian Islands fills Kaneohe Bay on
Oahu, where it has a width of about 3 miles.
Despite the fact that the broad mouth of
Kaneohe Bay is open to the north and north¬
east, the tsunami produced a rise in water
level at the bay head which was so small as
to be hardly perceptible to observers, and, so
far as could be determined, nowhere ex¬
ceeded 2 feet. Along the shore north of the
bay the heights ranged from 4 to 10 feet,
and on the end of Mokapu Peninsula south¬
east of the bay the heights reached more than
20 feet (Fig. 4) .

The lesser heights along the southern
shore of Molokai were probably partly due
to the wide protecting reef. The effect of the
reef in reducing wave violence along that
shore is well shown at places where channels
cross the reef. There the waves striking the
shore at the heads of the channels were dis¬
tinctly larger than those reaching shore on
each side of the channel. Thus at the- head
of a small channel which crosses the reef
just west of the mouth of Kainalu Stream the
water rose 1 1 feet, damaging houses, where¬

as just east and west of this channel the
water rose only 7 to 8 feet.

Configuration of the coast line. — It is gen¬
erally considered that the effects of tsunamis
should be intensified near the heads of V-
shaped embayments. Such embayments great¬
ly increase tidal fluctuations, as in the Bay
of Fundy, and might be expected to act like¬
wise on the similarly long waves of a
tsunami. Imamura (1937: 125-127) states
that as such a wave rolls up a V-shaped em-
bayment its height increases in inverse ratio
to the width and depth of the bay, and cites
examples of such increases in height of the
waves toward the bay head during Japanese
tsunamis. Consequently, special search was
made for this phenomenon in funnel-shaped
bays on Hawaiian shores. No good examples
could be found. Hilo Bay would appear to
be an almost ideal site for such funneling,
but measurements around its shores show no
systematic increase in heights toward its head
(Fig. 7 and Plate 1). Similarly there was a
lack of increase in heights toward the head
of the broad V-shaped embayment on the
northern coast of Maui. Possibly the ex¬
treme height of 54 feet at Waikolu Valley,
on the northern shore of Molokai, may have
been partly the result of funneling between
Kalaupapa Peninsula and the point and small
islands just east of the mouth of the valley.
At both Pololu Valley on Hawaii and Pele-
kunu Valley on Molokai, the water level was
higher at the bay head than on the walls of
the bay part way out. However, at Pololu
Valley, and probably also at Pelekunu, this
level was the result of a local upsurge where
the waves crossed the beach. Conversely,
several bays were found in which the heights
reached by the water were less at the bay
head than near its mouth.

Several small steep valleys, debouching
into small bays, were found in which the
water rose to appreciably greater heights
along the valley axis than on the sides near
the bay mouth or opposite the beach. Thus,

Onekahakoba

Plate 1. Map of the Hilo area on the island of Hawaii, showing the heights reached by the water, the area of flooding, and the portion of
the breakwater destroyed (shaded portions) during the tsunami of April 1, 1946. Heights are in feet above lower low water.

Plate 2. Map of the Hawaiian Islands, showing submarine topography (after H. T. Stearns).

Plate 3A. Wreckage left by the tsunami along Kamehameha Avenue, Hilo. Buildings on the left-
hand (seaward) side of the street have been pushed into the street, some more or less intact, others
as heaps of debris. Photo by Francis Lyman.

Plate 3B. House in Keaukaha, east of Hilo, carried inland about 100 feet by the waves. The
house in the background was above the reach of the water. Photo by G. A. Macdonald.

Plate 4A. Mouth of the Wailuku River at Hilo, showing the advance of one of the later waves
into the river mouth. Photo taken near the trough between two waves, showing very low water, and
the waves starting up the river as the next crest approaches. The steel span from the distant railroad
bridge is visible in the middle distance. Photo by Francis Lyman.

Plate 4B. A minute or so later, the waves are sweeping turbulently up the river. Photo by Francis
Lyman.

Plate 5 A. The very high stage of the water, in Wailuku River at Hilo, reached 3 or 4 minutes
later than the stage shown in Plate 4B. Photo by Warren Flagg.

Plate 5B. Scarp 5 feet high cut by the tsunami at the head of the beach at Moloaa, Kauai. The
roots were exposed by removal of the enclosing soil. Photo by G. A. Macdonald.

Plate 6B. Coral heads thrown up on the beach at Kaaawa, Oahu, by the tsunami. Photo by G. A.
Macdonald.

Plate 7 A. Grove of pandanus trees pushed over, and blocks of coral thrown up on the shore plat¬
form by the tsunami near Haena, Kauai. Photo by F. P. Shepard.

Plate 7B. Small boat washed inland and left stranded by the tsunami near Pier 1, Hilo. Photo
by G. A. Macdonald.

Plate 8. Bore advancing past the railroad bridge at the mouth of the Wailuku River, Hilo. Note the steep front, the turbulence of the water
behind it, and the placidity of the water in front of it. Photo by Shigeru Ushijima.

Tsunami of April l, 1946 — Macdonald et al.

33

Fig. 7. Map of the island of Hawaii, showing heights (in feet above lower low water) reached by
the water during the tsunami of April 1, 1946.

34

PACIFIC SCIENCE, January, 1947

in the small bay just south of Hanamaulu
Bay, on the eastern shore of Kauai, the water
rose only 25 feet on the bay sides, but swept
up the small valley at its head to a height
of 40 feet. At Moloaa, on Kauai, the water
reached an altitude of 40 feet in the axis of
the valley, but only 30 to 35 feet on the bay
walls. Again, at Honouliwai, on Molokai,
the water reached a height of 27 feet oppo¬
site the beach, but went 6 feet higher up the
valley. These are merely specialized exam¬
ples of effect, upon the rush of water up on
shore, of a topography above sea level which
served to concentrate the inrushing water.

Merging of waves from different directions.
— Wave crests traveling by different routes
may arrive at a given locality simultaneously,
giving rise to a wave of greater size than
either. Likewise, the simultaneous arrival by
different routes of a wave crest and a wave
trough may effectually cancel out both. Thus,
variations in the size and intensity of waves,
particularly on the sides of the islands away
from the wave origin, may result from the
arrival, either in or out of phase, of two
wave trains. During the tsunami of 1946
several examples of the formation of a large
wave by the juncture of two smaller ones
were observed. Thus, in the Keaukaha area
east of Hilo, witnesses described the arrival
of a wave from the north simultaneously
with one from the northeast, which built up
a very high crest at the place of juncture.
At the head of Maunalua Bay, on the south¬
eastern shore of Oahu, two waves were seen
to advance up channels across the wide reef,
move toward each other parallel with the
shore, and meet, throwing water upward like
the spray from a geyser. The water dashed
up on shore to a height of only 3 feet except
at the place of juncture, where it swept over
the top of a sandspit 5 feet above sea level.

Progressively southward around the shores
of Kauai, the average height of the high-
water marks gradually decreases, and along
much of the southern shore it is 6 to 12 feet

above sea level. However, in a zone 3 or 4
miles wide it ranges from 15 to 18 feet. This
zone is almost directly across the island from
the direction of wave origin, and probably
represents the area in which the waves re¬
fracted around opposite sides of the island
met and reinforced each other.

DAMAGE BY THE TSUNAMI

Damage by the tsunami can be divided
into structural damage, damage by erosion
and deposition, and damage by flooding. The
total property damage has been estimated
by the office of the Governor, Territory of
Hawaii, at about $25,000,000. Space per¬
mits only a brief review of the types of dam¬
age. The numbers of dwellings destroyed
and damaged by the tsunami on the major
islands are listed in Table 2 on page 36.

Structural damage includes damage to
buildings, roads, railroads, bridges, piers,
breakwaters, fishpond walls, and ships.
Frame buildings at low altitudes along
Hawaiian shores suffered extensive dam¬
age. Some were knocked over, by the force
of the waves, by cutting away of the sand on
which they stood, or by destruction of the
foundations. Others were bodily washed
away from their foundations. Some had
walls pushed in by the force of the water,
and in a few residences the water went on
through the house and took out the opposite
wall. As with earthquakes, there was a tend¬
ency to reduce the few two-story buildings
to a single story, by destruction of the lower
story. It is noteworthy that houses which
were well built and tied together internally
could be moved for considerable distances
without suffering severe damage. Even more
striking was the fact that houses elevated on
stilts a foot to several feet above the ground
survived the waves much more effectively
than did those built directly on the ground.
Apparently the water was able to pass under
such houses without greatly disturbing them,
unless it was deep enough actually to float

Tsunami of April 1, 1946 — Macdonald et al

the house off the stilts. The few reinforced
concrete structures in devastated areas suf¬
fered little or no damage except that caused
by flooding.

The railroads along the northern coast of
Oahu and in Hilo were wrecked, partly
through destruction of the roadbed, but
largely because the tracks were shifted off the
roadbed, either inland or shoreward. Locally
rails were torn loose, but more generally the
track was moved en masse, a motion probably
aided by the buoyancy of the ties. Coastal
highways also were partly destroyed, largely
by undercutting as the water returned sea¬
ward, but partly by the direct force of the
waves. Several highway and railway bridges
were destroyed. Most appear to have been
partly or entirely lifted from their founda¬
tions by the rising of the water under them.
The head of the pier at Waianae, Oahu, was
damaged in the same manner. At the Wai-
luku River, in Hilo, an entire span of the
steel railroad bridge was torn loose and
carried 750 feet upstream, passing under but
not damaging a highway bridge. At Kole-
kole Stream, 11 miles farther north, an entire
leg of the high steel railroad trestle was re¬
moved and carried upstream about 500 feet.

Part of the end and much of the shed of
Pier 1 in Hilo was wrecked by the force of
the wave. Most of the damage on Pier 2,
however, resulted when heavy pontoons,
which had been moored near by, were washed
across the pier. The wharves at Kahului on
Maui were flooded, but sustained little struc¬
tural damage.

The upper part of the breakwater at Hilo
was about 61 per cent destroyed (Plate 1).
Blocks of rock weighing more than 8 tons
were lifted off the breakwater and dropped
both inside and outside it. Destruction was
limited, however, to the part above water or
that only slightly submerged. The average
depth of water over the destroyed sections
after the wave was only about 3 feet. The
breakwater at Kahului, Maui, also suffered

35

minor damage. At both Hilo and Kahului
the breakwaters appear to have reduced mate¬
rially the height and violence of the waves
in the enclosed portions of the harbors.

Many small boats were washed ashore and
damaged. Railroad cars were overturned on
Oahu, Maui, and Hawaii. Many automobiles
were wrecked. The loose stone walls of fish-
ponds along the southern coast of Molokai
were partly thrown down. The mill of the
Hakalau Sugar Company, situated only about
10 feet above sea level at the mouth of Haka¬
lau Gulch on the island of Hawaii, suffered
severe damage.

Erosion by the tsunami resulted in the
partial removal of some sand beaches, in
some places causing a retreat of the shore
line for several tens of feet, cutting of small
scarps, and forming of large beach cusps at
the heads of beaches; locally, erosion caused
stripping away of a small amount of soil. The
erosion was largely concentrated high on the
beach, severaLfeet above sea level. Some of
the sand from the beaches was carried inland
and redeposited. At Haena, Kauai, the high¬
way was buried under 4 feet of sand, and
thinner layers of sand covered roads on
Oahu.

Flooding caused much water damage to
house furnishings and personal property.

LOSS OF LIFE AND PERSONAL INJURY

The following table summarizes, by islands,
the number of persons killed, injured, or
missing as a result of the tsunami. The fig¬
ures were supplied by the American Red
Cross. Most of the deaths were by drowning.
By far the heaviest toll was at Hilo, with 83
known dead and 1 3 missing. Those listed as
missing have been missing for more than 2
months, and must be presumed dead, bring¬
ing the total number of probable dead to
159. Great as it was, this loss of life was
moderate compared to that in some other
tsunamis, such as that of 1896 in the Sanriku

36

PACIFIC SCIENCE, January, 1947

district in Japan, which took more than
27,000 lives (Byerly, 1942: 72).

TABLE 2

List of Casualties during the Tsunami of
April 1, 1946

ISLAND

KNOWN

DEAD

MISS¬

ING

IN¬

JURED*

HOMES

DEMOL-

ISHEDf

HOMES

DAM-

AGEDf

Hawaii

87

34

153

283

313

Maui

9

5

2

65

144

Oahu

9

0

0

67

335

Molokai

0

0

0

13

14

Kauai

10

5

8

60

130

Total

115

44

163

488

936

159

* Injury sufficiently serious to require hospitalization,
f Homes only; other buildings not included. Data from
Lewers and Cooke, Ltd.

MITIGATION OF DISASTERS RESULTING
FROM FUTURE TSUNAMIS

There is no Hawaiian shore which is ex¬
empt from tsunamis. The most likely sources
of devastating tsunamis are the North Pacific
and South America. The areas heavily hit by
the 1946 tsunami are probably those most
likely to be hit hard again by tsunamis from
the North Pacific. Violent tsunamis from
Central or South America might, however,
cause much more damage than did the 1946
tsunami along eastern and southern coasts.
There is also possibility of serious damage
on western shores by a tsunami from Japan,
particularly if the tsunami occurred during
a heavy southwesterly storm. Tsunamis of
local origin might do heavy damage on any
shore.

It is obviously impractical to consider the
removal of all dwellings from Hawaiian
shores because of the danger from tsunamis.
It might, however, be advisable to prevent or
restrict building in certain areas of greatest
danger, particularly in centers of heavy pop¬
ulation, such as the waterfront at Hilo. Con¬
struction of suitable sea walls might also be
advisable in places. Sea walls cannot, how¬
ever, be built high and strong enough to hold

the water back completely, and an open zone
should be left back of the wall in which the
water pouring over the wall can use up its
energy in turbulence. Any construction per¬
mitted in such areas should be of a wave-
resistant type, such as reinforced concrete.
These wave-resistant buildings would have
the added virtue of serving as a line of de¬
fense for frailer structures behind them.
Frame structures in rural areas should be
built up off the ground, and far enough back
from the edge of the beach to reduce the
danger of undercutting. They should also be
properly reinforced and tied together.

It appears inevitable that future tsunamis
will cause loss of property on Hawaiian
shores, but loss of life from all except tsu¬
namis of local origin could be largely or en¬
tirely avoided. A system of stations could be
established around the shores of the Pacific
and on mid-Pacific islands, which would ob¬
serve either visually or instrumentally the
arrival of large long waves of the periods
characterizing tsunamis. The arrival of these
waves should be reported immediately to a
central station, whose duty it would be to
correlate the reports and issue warnings to
places in the path of the waves. It should be
possible in this way to give the people of the
Hawaiian Islands enough warning of the ap¬
proach of a tsunami to permit them to reach
places of safety. The effectiveness of the
warning, however, would depend on educa¬
tion of the public on the necessity for leaving
areas of danger, and on the efficiency of the
local organization in spreading the warning
and evacuating the threatened areas. Event¬
ually it should also be possible to state, at the
same time, which areas are likely to suffer the
most damage. Before that can be done, how¬
ever, we need more knowledge of the be¬
havior of tsunamis on Hawaiian shores, par¬
ticularly tsunamis from sources in the east¬
ern and western Pacific, and a more complete
picture of the submarine topography around
the Hawaiian Islands.

Tsunami of April 1, 1946 — Macdonald et al.

37

SUMMARY

The tsunami which reached the shores of
the Hawaiian Islands on April 1, 1946, was
the most destructive in the history of the
islands. Generated by a sudden shifting of
the sea bottom on the northern slope of the
Aleutian trough, the waves traveled south¬
ward to Hawaii with an average speed of 490
miles an hour, an average wave length of
about 122 miles, and a height over the deep
ocean of about 2 feet. Effects on Hawaiian
shores varied greatly. Locally the water
dashed more than 50 feet above sea level and
swept as much as half a mile inland. Else¬
where the rise in water level was very small,
and waves were gentle. Property damage was
heavy but loss of life was moderate.

The heights and intensities of the waves at
different points were influenced by position
on the island toward or away from the source
of the waves, offshore submarine topography,
presence or absence of coral reefs, shore-line
configuration, mutual reinforcement or inter¬
ference by waves traveling different paths,
and the presence or absence of storm waves.
Loss of property during future tsunamis can
be reduced by proper construction, by erec¬
tion of sea walls, and by restricting or pro¬

hibiting construction in certain especially
dangerous areas. Loss of life can be nearly
or entirely eliminated by the establishment
of a suitable system for warning of the ap¬
proach of waves.

REFERENCES

Bodle, R. R. Note on the earthquake and seismic
sea wave of April 1, 1946. Amer. Geophys.
Union Trans. 27: 464-465, 1946.

Byerly, Perry. Seismology, x+256 p., 58 fig.

Prentice-Hall, New York, 1942.

Green, C. K. Seismic sea wave of April 1, 1946,
as recorded on tide gages. Amer. Geophys.
Union Trans. 27: 490-500, 1946.

Green, George. On the motion of waves in a
variable canal of small depth and width. Cam¬
bridge Phil. Soc. Trans. 6: 457-462, 1838.
Gutenberg, Beno, and Charles F. Richter.
Seismicity of the earth. 131 p., 17 fig. Geol.
Soc. Amer. Spec. Paper 34. New York, 1941.
Imamura, Akitune. Theoretical and applied
seismology. 358 p. Tokyo, 1937.

Jaggar, Thomas A., Jr. Hawaiian damage from
tidal waves. Volcano Letter 321: 1-3, 1931.
Powers, Howard A. The tidal wave of April 1,
1946. Volcano Letter 491: 1-3, 1946.

Shepard, Francis P., Gordon A. Macdonald,
and Doak C. Cox. The tsunami of April 1,
1946. [In preparation.]

United States Navy Hydrographic Office.
Breakers and surf. 52 p., 19 fig., 4 pi. U. S.
Navy Hydrog. Off. Pub. 234. Washington,
D. C., 1944.

Dolomitization in Semi-arid Hawaiian Soils1

G. Donald Sherman, Yoshinori Kanehiro, and
Charles K. Fujimoto2

INTRODUCTION

Soils developed under a limited rainfall
usually show the influence of salinization by
calcium or sodium salts. In the presence of
appreciable quantities of sodium salts, soil
carbonates are converted to sodium carbonate
and give rise to a condition known as black
alkali. Recently workers have found soils in
which the salinization had been caused by
magnesium salts. Ellis and Caldwell (1935)
found a magnesium clay "solonetz” in cer¬
tain Manitoba soils in which 80 to 90 per
cent of the adsorbed cations were calcium
and magnesium. In addition, the calcium-to-
magnesium ratio in these soils varied from
1.6 to 1.0. Alicante ( 1933) reported a
Philippine soil which contained twice as
much magnesium as calcium in a soluble
form. Kudrin and Rozanov (1938), work¬
ing with "sierozem” soils, found a high con¬
tent of adsorbed magnesium especially in the
"solonchaks.” Rost and Chang (1941) di¬
vided the solonchaks of the Red River Val¬
ley of Minnesota into two groups : those
having 20 to 50 per cent of their exchange¬
able cations as magnesium and those having
more than 85 per cent of their exchangeable
cations as calcium. In addition, they found
a direct relation between exchangeable mag¬
nesium and gypsum in the soil. Thus, there
is evidence that magnesium salts may play
an important role in the salinization pro¬
cesses in soil.

Salinization of a soil with magnesium salts

1 Published by permission of the Director of
the University of Hawaii Agricultural Experiment
Station as Technical Paper 137.

2 University of Hawaii Agricultural Experiment
Station, Honolulu, Hawaii.

would be expected to give rise to the forma¬
tion of magnesium carbonate, a very insolu¬
ble compound of magnesium. However, if
a magnesium sulfate salinization occurred in
a soil rich in calcium carbonate, then dolo¬
mitization might occur in the soil with the
formation of two relatively insoluble com¬
pounds, dolomite ( double carbonate of cal¬
cium and magnesium) and gypsum. The re¬
sults of some recent studies suggest that this
process does occur in soils. Alway and Zet-
terberg ( 1935) found that the molecular
ratio of calcium carbonate to magnesium car¬
bonate varied from 1.51 to 4.80 in calcareous
Minnesota soils. Later work on Minnesota
soils ( Sherman, 1937) showed the occur¬
rence of dolomitization in soils which have
a high water table of waters rich in magne¬
sium sulfate. In some of these soils the
molecular ratio of calcium carbonate to mag¬
nesium carbonate approached 1.0. The car¬
bonates in the dolomitized area did not vis¬
ibly effervesce when treated with cold dilute
hydrochloric acid even though the soils con¬
tained as much as 40 per cent carbonates.
When pure dolomite is treated with cold
dilute hydrochloric acid, the ensuing effer¬
vescence is barely perceptible. This relative
inaction of a carbonate to cold hydrochloric
acid is characteristic of dolomite. The car¬
bonates from the area rich in magnesium
were identified as dolomite and distinguished
from calcite and other carbonates by a stain
technique ( Sherman and Thiel, 1939) . Pel¬
lets containing gypsum and long needle-like
crystals of gypsum were found imbedded in
the clay below the dolomitized layer of soil.
The nature of the occurrence of gypsum in.
these forms suggested their secondary origin.

38

Dolomitization in Hawaiian Soils — Sherman et al.

39

In Hawaii, Lyman and Dean ( 1938) have
reported soils containing twice as much mag¬
nesium as calcium. They found these soils
in low-lying areas which had been under salt
water in the past. The authors have found
other Hawaiian soils rich in magnesium in
areas which had never been under sea water.
These soils in general occur only in the dry
areas of the Hawaiian Islands.

The dark gray soils of Lualualei Valley,
Waianae Valley, and Makaha Valley on the
west slope of the island of Oahu have a low
calcium-to-magnesium ratio, as shown in the
present study. Under a very limited annual
rainfall of 16 to 20 inches per year, these
soils were developed on alluvial materials
brought down from the Waianae Range.
These soils are intermingled with numerous
outcrops of coral rock (95 per cent calcium
carbonate) and a few outcrops of basaltic
rocks. The soils developed on these mate¬
rials (coral and basaltic rock) can be readily
distinguished from the alluvial soils by color.
The native vegetation in the area is that
characteristic of semi-arid regions, the domi¬
nant vegetation being algarroba ( mesquite ) .
Because of the highly dispersed condition of
the soil and the consequent retarded penetra¬
tion of water, the "A” and "B” horizons of
the soil profile have been very poorly de¬
veloped. Crystals of gypsum were found in
some of the subsoils. In soil above the layer
containing gypsum, the carbonates gave a
very weak effervescence when treated with
cold dilute hydrochloric acid and a violent
effervescence with hot acid. This test would
indicate the presence of dolomite in this soil.

The object of this study is to determine
whether dolomitization is taking place, and
if so, to what extent it is occurring in these
soils and its possible relationship to soil¬
forming processes.

EXPERIMENTAL METHODS

Soil samples from several areas of soils
having a high percentage of their exchange¬

able cations as magnesium were collected,
on the basis of profiles which showed evi¬
dence of dolomitization. These samples were
analyzed for water-soluble salts, for ex¬
changeable cations, and for the composition
of the carbonates.

1. Water-soluble salts were leached from the soil
by shaking 25 grams of soil with a liter of dis¬
tilled water and allowing the mixture to stand
overnight. It was then filtered on a Buchner

. funnel and the soil was washed with another
liter of distilled water or until no test was given
for sulfates in the leachate. The filtrate was
evaporated to a small volume and the organic
matter was destroyed by acid oxidations with a
few drops of hydrogen peroxide. Calcium was
determined by the standard volumetric method,
using potassium permanganate as the oxidizing
agent. Magnesium and sulfate were determined
gravimetrically as magnesium pyrophosphate
and barium sulfate, respectively.

2. Exchangeable cations were extracted by neutral
normal ammonium acetate solution containing
70 per cent ethyl alcohol to reduce the solubil¬
ity of the calcium carbonate. Calcium and mag¬
nesium were determined by the same methods
as were used for the water-soluble ions. Potas¬
sium was determined volumetrically by the
cobaltinitrite method (Volk and Truog, 1934),
and sodium was determined gravimetrically as
sodium zinc uranyl acetate (Barber, 1928).

3. The carbonate analysis was made by a method
described for the analyses of dolomitic carbo¬
nates (Sherman, 1937). This method involved
the determination of carbonate carbon dioxide
by decomposing the carbonates with normal
hydrochloric acid solution, followed by absorp¬
tion of the carbon dioxide in a 0.5 N sodium
hydroxide solution in an absorption tower. Be¬
fore entering the tower, the carbon dioxide was
passed through a silver sulfate-sulfuric acid
solution to remove any hydrochloric acid fumes
from the digestion flask. The digestion flasks
were heated to boiling in order to insure the
complete decomposition of the dolomite. After
the decomposition of the carbonates, the so¬
dium hydroxide solution in the absorption
tower was drained into a volumetric flask. The
tower was washed with five portions of hot
carbon-dioxide-free distilled water to remove
all the sodium hydroxide. The carbonates were
precipitated in the volumetric flask by the addi¬
tion of barium chloride and the solution was
then made up to volume. The precipitate was
allowed to settle and an aliquot was taken from
the clear portion of the liquid. The excess so¬
dium hydroxide was determined in the aliquot

40

PACIFIC SCIENCE, January, 1947

by titration with a standard acid. A blank
determination was made by the same method,
using only the chemical reagents.

To determine the composition of the carbo¬
nates, another sample of soil was extracted
with the amount of half-normal hydrochloric
acid calculated as necessary to decompose the
determined quantity of carbonates in the soil,
this amount being based on carbonate-carbon
dioxide determinations. To insure a slight ex¬
cess of the acid, 0.1 ml. was added. Previous
work has shown that only the carbonates are
attacked by this quantity of acid (Sherman,
1937). The mixture w7as diluted with distilled
water and then heated to boiling to insure the
complete digestion of the carbonates. The mix¬
ture was filtered and the soil washed free of
chloride with hot distilled water. Calcium and
magnesium were determined in the filtrate by
using the usual standard methods for these
elements.

EXPERIMENTAL RESULTS

Samples of surface soils were collected
from various parts of Lualualei Valley, Wai-
anae Valley, and Makaha Valley on the island
of Oahu, and in Table 1 is given the content
of exchangeable calcium and magnesium in
these soils. In every soil, magnesium consti¬
tuted a very appreciable part of the cations,
accounting for 20 to 45 per cent of the total
exchangeable cations present in the soil.

Samples from a soil profile representing
a typical area which showed indications of
dolomitization were analyzed for composi¬
tion of the water-soluble salts, quantity of
each exchangeable cation, and composition

TABLE 1

The Content of Exchangeable Calcium and Magnesium in Surface Soils from Makaha Valley,
Waianae Valley, and Lualualei Valley on the Island of Oahu

LOCATION OF SOIL

pH

EXCHANGEABLE

Ca

EXCHANGEABLE

Mg

RATIO

Ca:Mg

m.e./100 gm.

m.e./lOO gm.

Makaha Valley .

7.3

22.12

16.06

1.38

Waianae Valley .

7.1

24.15

19.69

1.23

7.4

39.36

13.62

2.90

6.3

27.40

14.21

1.93

6.9

27.38

19.05

1.44

7.3

36.80

14.69

2.50

7.3

40.00

14.79

2.70

Lualualei Valley ......

7.1

56.80

16.15

3.49

6.4

29.40

18.62

1.58

6.9

31.98

19.16

1.67

7.0

26.10

17.77

1.47

7.8

33.89

9.89

3.43

7.6

37.42

9.64

3.88

TABLE 2

The Composition of the Water-soluble Salts of a Lualualei Valley Soil Which Contains

Dolomitic Carbonates

pH

WATER-SOLUBLE SALTS M.E. PER 100 GM. SOIL

RATIO

DEPTH OF SOIL SAMPLES

Ca

Mg

SO*

Ca:Mg

inches

0 6 .

7.8

0.84

0.43

0.52

1.95

6-12 .

7.7

0.76

0.43

0.36

1.77

12 18 .

7.5

0.72

0.48

0.40

1.50

18-24 .

7.5

0.80

0.41

0.41

1.95

24-30 .

7.2

8.48

2.18

10.10

3.89

30-36 .

7.3

26.60

3.63

30.12

7.33

36-42 .

7.3

26.78

4.36

31.08

6.14

Dolomitization in Hawaiian Soils — Sherman et al.

41

In Table 3 are given the data obtained for
the quantities of exchangeable cations in the
soil from the different parts of the soil pro¬
file. The quantity of exchangeable calcium
was found to be markedly greater in the soils
from the part of the profile which showed
the accumulation of gypsum. In this part of
the profile, calcium amounted to 80 to 90
per cent of the total exchangeable cations in
the soil, whereas in the soil above the zone
of gypsum accumulation it amounted to 72
per cent of the total exchangeable cations.
Exchangeable magnesium showed an oppo¬
site relationship, since it amounted to 25 per
cent of the total exchangeable bases in the
soils of the profile above the zone of gypsum
accumulation and in the gypsum zone it ac¬
counted for 9 per cent of the cations. The
ratio of exchangeable calcium to magnesium
was the lowest in the soils above the zone of
gypsum accumulation. The quantity of ex¬
changeable potassium and sodium found in
the soils of this profile was small, amounting
to less than 4 per cent of total exchangeable
cations.

The results of the analysis of the carbonate
fraction are given in Table 4. The amount
of total carbonates found in any soil horizon
of this profile was not great. Analysis of
carbonates in the soils of this area has failed
to show a zone of carbonate accumulation in

TABLE 3

The Quantity of Exchangeable Cations and the Percentage of Each Cation of the Total
Exchangeable Cations in a Lualualei Valley Soil Which Contains Dolomitic Carbonates

DEPTH

OF SOIL

EXCHANGEABLE CATIONS IN M.E.

PER 100 GM.

PERCENTAGE OF TOTAL

EXCHANGEABLE CATIONS

Ex¬

RATIO

SAMPLES

change

capacity

Ca

Mg

K

Na

Ca:Mg

Ca

Mg

K

Na

inches

0-6 . .

65.8

48.0

15.8

1.4

0.6

3.04

72.9

24.0

2.1

1.0

6-12 . .

57.2

40.8

14.5

1.0

0.9

2.81

71.3

25.3

1.7

1.6

12-18 . .

54.9

39.3

14.2

0.5

0.9

2.78

71.6

25.9

0.9

1.6

18-24 . .

48.0

34.5

12.1

0.3

1.1

2.85

71.8

25.2

0.6

2.3

24-30 . .

52.2

42.5

8.2

0.3

1.2

5.18

81.4

15.7

0.6

2.3

30-36 . .

74.4

66.4

6.5

0.2

1.3

10.21

89.2

8.7

0.3

2.0

36-42 . .

76.1

67.4

7.2

0.2

1.3

9.36

88.6

9.4

0.3

1.7

of the carbonates. The soil at a depth of
24 inches and deeper contained grains and
needle-like crystals of gypsum. The carbo¬
nates in the soil above the zone of gypsum
accumulation effervesced weakly when treated
with cold dilute hydrochloric acid; this fact
suggested the possibility that a high percent¬
age of the total carbonates was in the form
of dolomite. The data obtained in the analy¬
sis of this profile were similar to those ob¬
tained from similar soil profiles.

The data in Table 2 were obtained in the
analysis of the water-soluble salts extracted
from the soil samples of the selected profile.
The greatest quantity of water-soluble salts
was found in the part of the profile which
contained the crystalline gypsum. The cal¬
cium and sulfate amounted to approximately
27 and 31 milliequivalents per 100 grams in
this horizon. The water-soluble magnesium
increased with the increase in sulfate but the
increase was less than that shown by the cal¬
cium. The milliequivalents of water-soluble
calcium and magnesium were equal to the
milliequivalents of sulfate in soils rich in sul¬
fate. This would suggest that both calcium
and magnesium were combined with the sul¬
fate in this part of the profile. The lowest
calcium-to-magnesium ratio was found in the
zone overlying the zone of gypsum accumu¬
lation.

42

PACIFIC SCIENCE, January, 1947

any soil profile. In this profile, magnesium
carbonate was found in the greatest quan¬
tities in the soils above the zone of gypsum
accumulation. The portion of carbonates in
the form of dolomite was found to be great¬
est in the 12- to 18-inch layer of soil, which
corresponded to the zone of weakest effer¬
vescence with cold dilute hydrochloric acid.
The molecular ratio of calcium carbonate to
magnesium carbonate was 1.12 for this layer.
The carbonates were considered to be in the
form of dolomite, since if they were a mix¬
ture of calcium and magnesium carbonates
the quantity of calcium carbonate present
would have been sufficient to give a very vio¬
lent effervescence with cold dilute acid. Sher¬
man and Thiel (1939) found that when car¬
bonates of the soil were 87 per cent or more
dolomite they gave a characteristic efferves¬
cence with cold dilute acid which was similar
to that given by pure dolomite. Approxi¬
mately 94 per cent of the carbonates in this
soil layer are in the form of dolomite. In the
soil from the zone of gypsum accumulation,
calcium carbonate constituted approximately
80 per cent of the total carbonates.

DISCUSSION

This study has presented an instance of
the dolomitization of a calcareous soil by the
action of magnesium salts. The occurrence
of dolomitization in soil, a process by which
the double carbonate of calcium and magne¬
sium is formed, has been established by
chemical analysis of the soil carbonates, by
weakness of effervescence of the carbonates
when treated with cold dilute hydrochloric
acid, and by the presence of one of the two
end products of the process in the soil, gyp¬
sum. The formation of dolomite in these
soils is associated with the soil-forming pro¬
cesses, since the materials which make up the
alluvial parent material do not contain dolo¬
mite as a mineral. The soil is rich in mag¬
nesium silicate minerals and thus provides a
source of magnesium as soil weathering pro¬
gresses.

The mechanism by which the dolomite is
formed may need some explanation, since
calcium exceeds magnesium in both the
water-soluble salts and exchangeable cations.
The soils of this area receive most of their
precipitation during very short intervals of

TABLE 4

The Composition of the Soil Carbonates of a Lualualei Valley Soil Which Contains

Dolomitic Carbonates

DEPTH OF

SOIL

SAMPLES

SO*

PERCENT¬

AGE

SATURA¬

TION

WITH

Ca-fMg

CARBONATE ANALYSIS

Efferves¬
cence in
cold dilute
HC1

Total

carbonates

MgCOs

Carbo¬
nates as
dolomite

Ratio

Mols

CaCOs

to

Mols

MgCOs

inches

m.e./lOO gin.

per cent

gm. /100 gm.

gm. /100 gm.

per cent of

soil

soil

total

0-6 . .

0.52

96.9

strong

6.06

1.50

54.1

2.56

6-12 . .

0.36

96.6

weak

4.62

1.35

63.9

2.03

12-18 . .

0.40

97.5

very weak

3.44

1.47

93.9

1.12

18-24 . .

0.41

97.0

weak

0.87

0.21

51.7

2.64

24-30 . .

10.10

97.1

strong

0.84

0.09

23.8

6.82

30-36 . .

30.12

97.9

very strong

2.55

0.23

19.6

8.60

36-42 . .

31.08

98.0

very strong

3.67

0.28

16.6

10.27

Dolomitization in Hawaiian Soils — Sherman et al.

43

the year. Each year these soils become very
dry, often to the extent that they show wide
and deep cracks. During these periods of
drought the salts are brought toward the sur¬
face by capillary rise of water. In the process
of drying, much of the calcium salt is precipi¬
tated at lower levels, while the magnesium
salt, owing to its higher solubility, remains
in solution. At such periods the soluble mag¬
nesium may exceed the soluble calcium and
thus create conditions favorable to the for¬
mation of dolomite by the action of magne¬
sium sulfate on calcium carbonate. When the
soils become wet the salts are leached to a
lower level in the soil, from which the more
soluble salts (magnesium) can rise again.
This would be a possible explanation of the
gypsum accumulation in the subsoil of these
soils and of the formation of dolomite di¬
rectly above the gypsum layer.

The formation of dolomite and gypsum
as end products of this process should be
expected under arid or semi-arid conditions,
since these two compounds represent the
most insoluble compounds of calcium and
magnesium which could be formed in the
presence of calcium carbonate. According to
Leather (1913), the presence of magnesite
together with calcite in soil is extremely
doubtful and if magnesite did exist it would
only be present in traces. It has been found
that when the dolomitic carbonates are
treated with acid they give up calcium and
magnesium carbonate in approximately a 1 : 1
ratio. This fact and the effervescence which
is characteristic of dolomite establish the
identity of the type of carbonate formed. The
formation of gypsum will result from the
action of magnesium sulfate on the calcium
carbonate and its precipitation from solution
during wetting and drying of the soil. This
process would lead to the formation of gyp¬
sum in the crystalline forms found in these
soils.

Dolomitization in soils can be expected to
occur under soil conditions similar to those
described in this report. It is likely that
dolomitization has occurred to some extent in
many calcareous soils.

CONCLUSIONS

As a result of this study of the composi¬
tion of the soil carbonates in a representative
soil profile from Lualualei Valley, Makaha
Valley, and Waianae Valley, the following
conclusions may be drawn:

1 . Dolomitization of the carbonates has been
established by the chemical composition
of the soil carbonates, by the effervescence
in cold dilute hydrochloric acid charac¬
teristic of dolomite, and by the presence
of one of the two relatively insoluble end
products, gypsum, which will be produced
when this process occurs in the presence
of sulfates and calcium carbonate.

2. In the soil horizons where dolomitization
has occurred, 50 to 94 per cent of the car¬
bonates are in the form of dolomite.

3. In order that the process of dolomitiza¬
tion may occur, the presence of soluble
magnesium salts is required. The soils of
this area were found to be rich in mag¬
nesium, since this cation accounted for
20 to 45 per cent of the exchangeable
cations in the soil. The drying of the soil
during dry weather concentrates and in¬
creases the soluble magnesium until it ex¬
ceeds the calcium in solution due to the
precipitation of calcium as gypsum, and
thus leads to conditions favorable to the
formation of dolomite. Periodic rainy and
dry seasons are essential to the dolomiti¬
zation process.

4. The soil below the zone where the dolo¬
mitic carbonates exist contained numerous
pellets filled with crystalline gypsum and
crystals of gypsum. In this zone of gyp¬
sum accumulation the calcium-to-magne-
sium ratio is very high.

44

PACIFIC SCIENCE, January, 1947

REFERENCES

Alicante, M. M. A survey of the soils in the
district of the Philippine Milling Company,
Mindoro. Philippine Sugar Assoc. Res. Bur.,
Ann. Rpt. Dir., 1933: 69-72, 1933.

Alway, F. J., and Jean M. Zetterberg. Relative
amounts of calcium carbonate and magnesium
carbonate in some Minnesota subsoils. Soil Sci.
39: 9-14, 1935.

Barber, H. H., and I. M. Kolthoff. A specific
reagent for the rapid determination of sodium.
Amer. Chem. Soc. Jour. 50: 1625-1631, 1928.

Ellis, J. H., and O. G. Caldwell. Magnesium
clay "solonetz.” Third Internatl. Cong. Soil Sci.
Trans. 1: 348-350, 1935.

Kudrin, S. A., and A. N. Rozanov. The char¬
acterization of serozems having a high content
of adsorbed magnesium. Pedology (8): 83 6-
855, 1938.

Leather, J. W. Soil problems. Pusa Agr. Res.
Inst, and Col, Sci. Rpts., 1913-14: 16-17, 1913.

[Abstract, U. S. Off. Expt. Stas., Expt. Sta. Rec.
33: 513, 1915.]

Lyman, Clarence, and L. A. Dean. The cal¬
cium-magnesium ratio of some Flawaiian soils.
Hawaii Univ. Agr. Expt. Sta. Rpt., 1938: 49,
1938.

Rost, C. 0.: and P. C. Chang. Exchangeable
bases of solonchak of the Red River Valley.
Soil Sci. Soc. Amer , Proc. 6: 354-359, 1941.
Sherman, G. Donald. The occurrence of dolo¬
mite in the subsoils of the Red River Valley.
[Master’s thesis on file at University of Minne¬
sota, 1937.]

- and G. A. Thiel. Dolomitization of

glacio-lucustrine silts of Lake Agassiz. Geol.
Soc. Amer. Bui. 50: 1535-1552, 1939.

Volk, N. J., and E. Truog. A rapid chemical
method for determining the readily available
potash of soils. Amer. Soc. Agron. Jour. 26:
537-546, 1934.

Notes on the Red-billed Leiothrix in Hawaii

Harvey I. Fisher and Paul H. Baldwin1

The name Japanese Hill Robin often
used in Hawaii for Leiothrix lutea (Scopoli)
is an unfortunate misnomer, as is Pekin
Nightingale, because the bird does not occur
in the wild in Japan nor does it occur as far
north as Shanghai, much less Pekin or Pei-
ping.

Henshaw (1911: 6 3) recommended the
introduction of this species into Hawaii as
early as 1911 and noted that it lives to some
extent on insects and small fruits. Caum
(1933: 38-39) states that the species was
first introduced into the Territory of Hawaii
in 1918 by Mrs. Dora Isenberg, who liber¬
ated several birds on the island of Kauai.
However, the records kept by the Territorial
Board of Agriculture and Forestry indicate

1 Department of Zoology and Entomology, Uni¬
versity of Hawaii, and Bernice P. Bishop Museum,
Honolulu, respectively.

several small importations previous to this
time (Table 1). These probably were made
for cage purposes, but some of the long-time
residents believe Leiothrix was established in
the Hawaiian Islands before 1918 by escapees
from cages.

In Table 1 are summarized all importa¬
tions of the species shown by official records.
It is worthy of note that the table does not
include the importations of 1918 and 1928
from San Francisco mentioned by Caum
{loc. cit.). There may have been other un¬
recorded introductions. The table does show
a total of 41 importations, including some
1,037 individuals. These individuals have
been released on all the larger islands of the
southeastern end of the Hawaiian Chain; the
islands are Kauai, Oahu, Molokai, Maui, and
Hawaii.

From official records the only indication

TABLE 1

Ports of Embarkation and Summary of Dates and Numbers of Leiothrix Imported into Hawaii

YEAR

PORT OF EMBARKATION

TOTAL

Hongkong

Sydney

Shanghai

Kobe

Yokohama

1911 . .

1

1

1913 . .

2,6

3

11

1915 . .

4

4

1916 . .

5

2

• . . . .

7

1917 . .

l, 4, 6, *3

14

1918 . .

....

2

2

1919 . .

2, 12, 12

26

1922 . .

1

1

1923 . .

3

3

1924 . .

2

2

1928 . .

*3* 50*

40, 30, 19

142

1929 . .

6, 184

190

1932 . .

• • • •

1

1

1933 . .

1

7

8

1934 . .

75

75

1935 . .

....

....

28

28

1936 . .

1, 4, 1, 80

200

....

100, 37, 36

459

1937 . .

1

2

....

60

63

Totals '.

181

2

208

354

292

1,037

45

46

PACIFIC SCIENCE, January, 1947

we have of the original home of these intro¬
duced birds is the name of the port from
which they were shipped, and this tells
little. It was hoped that the subspecific deter¬
mination of the Hawaiian birds might aid
in revealing the native home of the intro¬
duced stock. However, H. G. Deignan, of
the United States National Museum, has
graciously examined 15 of our specimens
from the Hawaiian Islands and has identified
them as Lelothrix lutea lutea. The type local¬
ity of this race is the mountains of Anhwei
Province south of the Yangtze. Since lutea
ranges over all of China south of the Yang¬
tze and east of the Kansu frontier, the origi¬
nal home of the breeding stock now in the
Hawaiian Islands cannot be further specified.

DISTRIBUTION IN THE HAWAIIAN ISLANDS

Caum (1933: 39) noted that large flocks
were present on Kauai and rightly assumed
that the birds were breeding successfully on
that island. However, he was uncertain about
the status of the species on other islands,
simply saying that the birds were "reported”
to be breeding on Molokai, Maui, and Ha¬
waii and "believed” to be breeding on Oahu.
Subsequent unpublished observations have
indicated that the birds are reproducing and
are increasing the extent of their ranges on
Hawaii, Oahu, Molokai, Kauai, and Maui.
The bird is abundant, widespread, and nest¬
ing successfully on Hawaii. It is numerous
and nesting in the Koolau and Waianae
mountain ranges on Oahu. In early June,
1945, adults were observed feeding nearly
fledged young in the mountain valleys at
Mapulehu, Molokai, at an elevation of 500
feet. On Maui it is generally present in the
moderately wet to wet forests of Haleakala
and probably West Maui.

The vertical distribution on individual
islands is unrecorded except on Oahu, Molo¬
kai, Maui, and Hawaii, where we have ob¬
served the species. On Oahu the species
ranges from 400 to 3,000 feet in the Koolau

Range and probably goes to about 4,000 feet
in the Waianae Mountains. In other words,
its vertical distribution includes all elevations
available except those below 400 feet, and it
seems likely that the species might be found
below this level if all suitable cover condi¬
tions in deep valleys were to be investigated.
Although the altitudinal range is consider¬
able, the birds are most abundant above 600
to 800 feet; this fact may be due to the lack
of sufficient cover at lower elevations.

On Hawaii the bird has been found chiefly
at elevations ranging from near sea level
(Puna district and windward Hawaii) to
7,500 feet on Mauna Loa. We may assume
that the upward limit of their livable range
on Hawaii is between 8,000 and 9,000 feet.
However, there is no definite upper limit to
the distribution in fall and early winter,
when the birds band together and wander all
over the island. Flocks have frequently been
seen at more than 13,500 feet on Mauna Loa,
but these flocks are soon reduced by deaths
caused by exposure and starvation.

Temperature may be a factor in limiting
upward distribution; this is indicated by in¬
creased mortality rates above 9,000 feet on
Hawaii, where the mean temperature is 40°
to 50° F. with but little seasonal variation.
However, it is not likely that temperature is
the important factor below this altitude, and
even above 9,000 feet, food, wind, and cover
conditions are probably the more important
limiting factors.

Throughout its range (vertical and hori¬
zontal) the species is limited to the more
moist, forested areas having a dense under¬
story of plant growth. On Oahu this means
it is restricted to the mountain valleys; sel¬
dom is the bird observed on the higher ridges
except during its movements from one valley
to another. Despite the bird’s proclivity for
moist valleys, rainfall apparently is not the
determining factor except possibly in areas
having less than 40 inches a year, for yearly

Red-billed Leiothrix in Hawaii — Fisher and Baldwin

47

averages of precipitation in its observed
range on Oahu vary from 30 to 200 inches.
On Hawaii the species is rare or absent in
areas having less than 20 inches a year;
this is true in the leeward districts. In the
Kau district, where rainfall is 20 inches or
less in much of the lowland, Leiothrix is rare
below 3,000 feet and absent below 2,500
feet. On Maui the annual rainfall in parts
of the range of the species is 350 inches.

Presence or absence of surface water may
be a significant factor. The species has often
been observed bathing in water up to 2 inches
in depth. Further, the birds are most fre¬
quently observed within a few hundred yards
of such water. We have on several occasions
noted birds drinking rain water caught in
basins formed by the large fallen leaves of
the ti plant ( Cordyline terminalis ) and the
kamani tree ( Terminalia Catappa) .

HABITAT IN THE HAWAIIAN ISLANDS

A cover of dense vegetation near the
ground is the major characteristic of the habi¬
tat of Leiothrix. Without exception the birds
are found in areas having such a growth. It
seems immaterial whether or not a canopy is
formed above this undergrowth, as long as
there is a thicket 10 to 15 feet in depth. We
have never found this species feeding, nest¬
ing, or even remaining long in the large
plantings of imported ironwood ( Casuarina )
or exotic Eucalyptus; there is rarely any
ground growth in such places. The birds are
frequently found in guava ( Psidium Gua-
java ) thickets not having a high canopy.

On Oahu the species is most abundant in
the dense understory of forested and partly
forested areas on the floors and steep slopes
of the valleys. Usually there is in this habitat
a stream, a small pond, or some temporary
catch basins (fallen leaves, concavities in
rocks) to hold rain water. On the extremely
steep slopes of Manoa Valley, Oahu, at 1,100
feet where the rainfall is between 100 and
150 inches annually is a "typical” habitat.

Here are scattered ohia lehua ( Metrosideros
collina var. polymorpha) trees up to 12 and
15 feet in height, interspersed with koa
( Acacia Koa ) trees up to 30 feet high and
with kukui nut ( Aleurites moluccana) trees.
These three kinds of trees form in many
places a loose, open canopy. In certain areas
the kukui trees form a dense canopy. The
understory consists of ti plants ( Cordyline
terminalis) up to 8 feet in height, a few
Gardenia Remyi trees, and an occasional
white hibiscus [Hibiscus Arnottianus) . The
lowest-growing vegetation is made up chiefly
of palm foxtail [Set aria palmi folia) and oi
shrubs [Stachytarpheta cayennensis) . This
growth is in many places so dense that it is
difficult to penetrate. A further factor in
creating an almost impenetrable thicket is
the presence near the ground of many fallen
branches. Also, in a few places the inter¬
twining branches of the hau tree [Hibiscus
t iliac e us) add to the maze (Fig. 1).

Fig. 1. Thicket formed by branches of the hau
tree ( Hibiscus tiliaceus ) in Nuuanu Valley, Oahu,
at 1,000 feet elevation.

On Hawaii this babbler is most abundant
in forests from 1,000 to 5,000 feet elevation
and with 40 or more inches of rain each year.
Forests with a variety of habitats, such as koa
forest partly modified by cattle grazing and
with thimbleberries [Rub us rosaefolius) and
other fruiting plants, harbor large numbers
of the species. The density reaches 80 to

48

PACIFIC SCIENCE, January, 1947

100 birds an acre at times at Kipuka Puaulu
(4,000 feet). Guava thickets at elevations
of 1,000 to 2,000 feet are also tenanted in
good numbers. However, virgin tree fern
forests favorable to the native Hawaiian
birds are in some areas without Leiothrix. A
forest of this type at the Twin Craters ( 3,800
feet) in Hawaii National Park usually had
not more than one individual per acre. Simi¬
lar forests of the Upper Olaa Forest Reserve
near a farming district exhibited a somewhat
greater density.

On Maui in the Kipahulu Valley the vir¬
gin, undisturbed ohia lehua forests (eleva¬
tion 4,000 to 6,000 feet and rainfall 250 to
350 inches a year) had very few birds of this
species. In koa forests in this same valley,
but between 2,000 and 4,000 feet elevation
and with a rainfall of 150 to 250 inches a
year, the population density of this species
was slightly greater, an indication that the
bird was generally present, but uncommon.

The birds are seldom seen more than 15
feet above the ground and then only when
the ground cover is very thick, or when the
birds are in heavy tree foliage. Only occa¬
sionally can they be observed in the higher
strata of the ohia and koa trees. Most fre¬
quently the birds are heard in the low thickets
and not seen. If one is quiet it is possible to
observe the bird flitting from shrub to shrub
next to the ground. These movements com¬
bine short fluttering flights and hops of a few
inches.

FLOCKING

We have seldom observed a flight of more
than approximately 50 feet, but others report
that flights of 200 feet are not uncommon in
the more open parts of the range of the
species. Longer flights are more frequently
found during the winter, when the birds are
banded together into traveling groups of 20
or even 100 birds. Some of the flocks in
favorable places seem to confine their move¬
ments to a limited area, as at Kipuka Puaulu

on Hawaii. However, some observers think
the flocks represent groups migrating into
the lowlands for the winter. There is a
movement at this time, but it is more of a
general diffusion into peripheral areas not
heavily occupied during the breeding season,
and it is by no means limited to downward
movements, for, as mentioned previously, in
winter the birds appear conspicuously in the
ohia lehua forest of Mauna Loa at high ele¬
vations.

By May at Kipuka Puaulu the bands have
broken up, and most of the birds are segre¬
gated by pairs. By late summer (July and
August) small bands of 4 to 12 birds are
common. Consequently, it appears that flocks
are re-formed almost immediately after pa¬
rental duties at the nest are completed. How¬
ever, some of the birds remain in flocks
throughout most of the breeding season
( March through June ) .

NESTING

Perhaps nesting occurs at all elevations
found in the spring and summer range of
Leiothrix, but on Oahu nests have been
found only between 500 and 2,500 feet ele¬
vation. On Hawaii the highest elevation at
which a nest was found was 6,100 feet at
Kipuka Kulalio. Without exception on Oahu
the nests have been less than 10 feet from
the ground and located in the densest parts
of the understory; one nest was only 18
inches above the ground. On Hawaii nests
have been found from 3 to 7 feet from the
ground. Choice of the nest site seems to
depend not so much on the particular kind
of vegetation as on its density, for nests on
Oahu have been found in staghorn fern
(Dicranopteris linearis ), oi, palm foxtail,
and guava, on the finer branches of low-
growing hau trees, and attached to the stems
of ti leaves. On Hawaii, sites selected for
nests included dense shrubs (aalii, Dodonaea
viscosa; pukeawe, Styphelia Tameiameiae) ,
the branches of trees (mamani, Sophora

Red-billed Leiothrix in Hawaii — Fisher and Baldwin

49

chrysophylla ), and fronds of tree ferns
( Cibotium Chamissoi) .

Occupied nests have been found between
March 3 and May 7 on Oahu, and from early
March through June on Hawaii. It is likely,
however, that further investigation on both
islands would reveal a longer breeding
period.

On March 3, 1945, a nest with four eggs
was found at 1,200 feet on the west side of
Manoa Valley, Oahu. A complete descrip¬
tion of this nest follows, because we have
been unable to find any published account of
the nest of this species. The nest was located
in a well-shaded spot beneath a koa tree and
in a dense growth of the exotic palm foxtail.
It was well concealed by the rank growth of
the foxtail and would have passed unnoticed
had not the bird flushed when the observer
stopped some 18 inches from the nest, which
was 36 inches from the ground.

Support for the nest was a fork near the
terminal twigs of an oi shrub ( Stachytarpheta
cayennensis) ; the heaviest branch of the fork
was only 2 mm. in diameter. The nest was
attached to the fork by only a few fibers from
the leaves of the foxtail, which were wound
around the twigs and woven into the rim of
the nest. While not swinging as free, for
example, as the nest of an oriole, the nest
was definitely pendulous.

The measurements of the outside of the
top of the ovoid nest were 9 cm. by 11 cm.;
corresponding dimensions of the inside were

4.5 cm. by 6 cm. Over-all depth of the nest
was 8.5 cm., but the depth of the cavity was

5.5 cm. From these figures we may conclude
that the walls of the nest vary between 2 and
3 cm. in thickness.

All materials used in constructing the nest
came from one plant, the palm foxtail,
which was available within a few inches of
the nest site. On the outside, entire leaves
(some 18 inches long and iy4 inches wide)
were loosely wrapped around the nest and

tied into the nest by small fibers (fibrovascu-
lar bundles) from the leaves of the foxtail.
However, the bulk of the nest was woven of
longitudinally split leaves from which the
epidermis and mesophyll had disappeared,
leaving only 8 to 15 parallel fibers. The
lining of the nest consisted of single fibers
from the same plant; they were loosely laid
in to form a springy inside surface for the
nest. The only part of the nest not coming
from the leaf of this grass was three rootlets
(the largest was 25 cm. long and 1 mm. in
diameter) woven into the rim of the nest.
The rootlets gave additional rigidity to the
free edge.

Because of certain differences in construc¬
tion and in materials used it seems desirable
to describe another nest, found May 18,
1940, at 6,100 feet at Kipuka Kulalio,
Mauna Loa, Hawaii. It was suspended in an
aalii tree from the crotch of a forked twig
at a height of 6]/2 feet from the ground and
was secured by grass stems bent over the
arms of the crotch and incorporated into the
wall of the nest. The outermost part of the
nest was a sling of grass woven vertically
from the rim on one side, under the nest, to
the opposite rim. This was the sole support,
as there was no base of twigs. Inside this
sling, grass was woven in various directions
to make up the wall, which was about 3 cm.
thick. Grass was worked in horizontally
along the rim. There was a lining of loosely
woven grass, strands of which ran diagonally
up the sides to bend down at the rim at
angles of about 80° and run again across the
floor of the nest. The whole structure was
firm, but not compact. Light could be seen
through the sides. A few leaves had been
placed near the bottom and in the bottom.
The materials in the nest were 85 per cent
grass (60 per cent stems and 25 per cent
leaves), 15 per cent leaves of aalii, a trace
of rootlets in the rim, and a few feral goat
hairs in the lining. Measurements of this
nest were:

50

PACIFIC SCIENCE, January, 1947

GREATEST

LEAST

DIAMETER

DIAMETER

DEPTH

cm.

cm.

cm.

Outside

. . 11.5

9.0

9.0

Inside .

. . 5.5

4.5

5.5

EGGS

The egg is shiny, with an extremely pale-
blue ground color and a ring of red-brown or
purple splotches around the larger end. A
few pale-brown splotches are found on the
blunt end and the sides of the egg, less often
at the apex. Small brown specks and dark
scrawlings may overlie the brown and purple
splotches, especially at the blunt end.

Weights and measurements of eggs from
two clutches are given below. The weights
were taken 3 days before the eggs hatched.

WEIGHT LENGTH

DIAMETER

gm.

mm.

mm.

Clutch 1

eggl • . . .

2.4

20.9

16.5

egg 2 • ■ • -

2.4

20.3

16.4

egg 3 • • • •

3.0

21.3

16.5

Clutch 2

eggl . . . .

20.4

15.4

egg 2 . . . .

20.3

15.5

egg 3 • • • •

20.0

15.4

egg 4 . . . .

20.9

15.6

The number of

eggs

observed in

a clutch

varied between two and four, with an average
of about three (two nests with two eggs, two
with three eggs, and three with four eggs).

HATCHING

The earliest hatching observed on Hawaii
was on March 14, and the latest was on June
1 6. On Oahu these dates were March 10 and
May 15.

Cracks were found in all three eggs of a
clutch at 3:00 p.m. on May 22. The cracks
appeared to have resulted from activity in¬
side the egg. In one egg the cracks were just
distal to the largest diameter. A second egg
had cracks in various places around its great¬
est circumference, while the third had only
one small hole immediately distal to the
greatest diameter. By 5:00 p.m. two of the
eggs had hatched. The shells had been re¬

moved from the nest and could not be found
on the ground; apparently they were carried
away by the parent. By 5:35 p.m. the third
egg had broken open. The shell was broken
in a ring just proximal to the largest diame¬
ter. The crown of the embryo was pressed
against the cap of the shell. The hatching
bird flexed its neck and then extended it,
shoving the shell cap forward and freeing
its head. After resting a few moments it
worked its legs back and forth twice, simul¬
taneously kicking its body forward and the
shell backward.

DESCRIPTION AND DEVELOPMENT
OF THE YOUNG

When the nestlings were 1 hour old they
appeared as follows: skin, rich reddish apri¬
cot, except where feather follicles caused gray
color as in humeral tract, dorsal surface of
neck, upper part of wing, and mid-dorsal
region, and over eye; bill, reddish apricot,
except upper mandible (gray between nares
and tip); egg tooth, whitish yellow; rictus,
light yellowish apricot; legs and feet, apricot;
claws, yellow. Down feathers were present
above eye, along occiput and mid-dorsal line
from interscapular region to posterior edge
of oil gland.

When the young were 24 hours old the
skin color was less reddish, and the gray
feather tracts were more pronounced and ex¬
tensive; plumules, absent in ventral tract;
eyes, still closed and gray; legs, yellow-apri¬
cot; skin, loose in tibiofibular region and on
neck, tight over belly; belly, more prominent
and bulging than at hatching.

When the birds were 7 days of age the
first primary feather was 15.5 mm. long and
still within its sheath; bill, orange at edges,
gray on ridge and side; rictus, whitish; legs,
salmon.

When they were 11 days old the first
primary was 33.5 mm. long and two thirds
out of its sheath; central rectrices extended
3 mm. and 1 mm. out of their sheaths; plu-

Red-billed Leiothrix in Hawaii — Fisher and Baldwin

51

mules, gone from back but present on crown
and femoral tracts, sparse on crural tract;
feathers forming fluffy covering over entire
body. The primaries were lined with golden-
yellow; breast, yellow-green; back, dark yel¬
low-green; ventral tracts, whitish yellow-
green; undercoverts of tail, yellow (still in
sheaths ) ; secondaries, black and dark green.
The bill was dark salmon, but superficially
gray; rictus, whitish along edge; legs, yellow¬
ish-tan; and claws, yellow-orange at tips.

The weights of the young are summarized :

AGE SIBLING A SIBLING B SIBLING C

gm. gm. gm.

1 day . . . 3.1 3.0 3.2

7 days . . . 13.0 13.9 13.5

11 days. . . 15.0 _ 15.2

When the birds were 1 hour old, their
movements were sluggish; the head was
lifted to receive food, and swallowing was
accomplished. When 24 hours old the nest¬
lings could squirm around; they rested on
their abdomens. They threw back their heads
and opened their mouths frequently. Beg¬
ging behavior and ejection of feces could be
brought about by touching the nestling.
When not active they lay curled up on their
sides with legs and toes flexed. At 7 days of
age they still were unable to crawl. At 11
days one sibling left the nest when the ob¬
server tipped it to look inside. It escaped
into the bushes by moving along the ground
in short flights (3 feet) . The nestlings could
perch well, and they shifted their footholds
to compensate for movements of the nest.
Low-pitched alarm notes were uttered when
they were trying to escape capture.

FOOD

Animal matter found in the stomachs of
13 birds from Hawaii National Park came
from various species of Hymenoptera, Dip-
tera, Lepidoptera, and Mollusca. Plant ob¬
jects included fruits of the thimbleberry
( Rubus rosaefolius ) and other fruits and
stems. The proportion of animal to plant

matter varied from all animal to all plant
food, but the proportions usually were be¬
tween 40:60 and 60:40 by volume. Grit
was typically present in the gizzard.

Observational data on Oahu indicate that
the birds also feed on the fruits of strawberry
guava ( Psidium Cattleianum ) , overripe pa¬
payas, petals of small flowers, and small new
buds of various plants. Leiothrix has also
been seen to capture insects which flew near
its position, but in feeding on insects it does
not fly out from a perch in typical flycatcher
fashion.

BIRD MALARIA

Blood smears from 11 specimens of Leio¬
thrix from Hawaii were examined by the
U. S. Fish and Wildlife Service. Plasmodium
vaughani was reported from one smear made
from a bird taken near Kipuka Puaulu in
Hawaii National Park.

INTERACTION WITH OTHER ANIMALS
Although at times Leiothrix seems to be a
noisy, aggressive bird, we found no antago¬
nism toward other avian species except on
one occasion. An observer was at a nest of
the species, and under the stress of excite¬
ment one of the adults drove a foraging
Olive-green Creeper (Paroreomyza hairdi
mana) from the nest tree.

Except for the Hawaiian Hawk on the
island of Hawaii, there is no evidence to
indicate that other avian species molest Leio¬
thrix in any way. Elsewhere in the Hawaiian
Islands, the only possible predators on this
species are mammalian — the rat, the mon¬
goose, wild dogs and cats, and feral hogs —
and these would exert pressure on the popu¬
lation by destroying eggs and young.

REFERENCES

Caum, Edward L. The exotic birds of Hawaii.
Bernice P. Bishop Mus. Occas. Papers 10 (9),
55 p., 1933.

Henshaw, Henry W. Report of the Committee
on the Introduction of Birds into the Hawaiian
Islands. Hawaii. Forester and Agr. 8: 61-64,
1911.

NOTES

A number of policies for the co-ordinated devel¬
opment of scientific research in the Pacific were
adopted by the Pacific Science Conference of the
National Research Council, which met at the Na¬
tional Academy of Sciences, Washington, D. C.,
on June 6-8, 1946. More than one hundred mem¬
bers of the conference from various divisions met
with a number of liaison members from the State
Department, War Department, Army Air Forces,
Navy Department, U. S. Coast Guard, Depart¬
ment of the Interior, Department of Agriculture,
Department of Commerce, U.S. Commercial Com¬
pany, U. S. Office of Education, U. S. Public
Health Service, Smithsonian Institution, American
Council of Learned Societies, American Council on
Education, and Social Science Research Council.
Special guests were invited from a number of
foundations interested in Pacific research.

The general and specific recommendations
adopted by the Council, given below, should be of
interest to all persons and organizations concerned
with the systematic and methodical advancement
of scientific studies in this vast area.

GENERAL RECOMMENDATIONS

1. Declassification

That this Conference strongly recommend
that, insofar as practicable, all Pacific Island
materials and information now the property
of government agencies and organizations be
declassified and made available to recognized
scientific organizations in accordance with
recommendations from the proposed Pacific
Science Survey.

2. Conservation

That throughout the Pacific area every effort
be made:

a. To protect and preserve areas, objects and
living species of flora and fauna having
scientific, historic, or aesthetic significance,
through appropriate conservation legisla¬
tion, including the establishment of na¬
tional parks, nature monuments, and re¬
serves.

b. To take necessary measures to insure the
preservation of flora and fauna in their
native environment.

To set aside certain wilderness areas that
are to be maintained inviolate except for
essential scientific studies.

d. To determine which species are in danger
of extinction and to take special measures
for their protection and preservation.

e. To avoid the deliberate introduction of
exotics wherever indigenous fauna or flora
will be endangered, and to keep records of
the intentional and accidental introduction
and spread of exotic forms of animal life.

/. To minimize accidental introductions, by
more effective quarantine efforts.
g. To apply caution in the use of insecticides
(such as DDT), rodenticides (such as
1080), herbicides, and other chemical con¬
trols of organisms, and to carry out thor¬
ough researches on the effects of such
chemicals on all forms of life, including
independent investigations before, during,
and after the applications.

That conservation regulations and the im¬
portance of protecting vanishing species from
extinction be brought to the immediate atten¬
tion of the establishments of the Armed
Forces to prevent indiscriminate shooting or
other practices that might cause the extinction
of vanishing species of flora and fauna.

3. Fellowships

That the continuing organization arrange
for research fellowships at varying financial
grades for competent graduate students, and
for grants-in-aid to established scholars, in¬
cluding local inhabitants, in the several fields
of science involved, as a part of the me-,
chanics of staffing research.

That funds be made available to foster the
interchange of information on the physiology,
biochemistry, and biophysics of plants of im¬
portance in the Pacific area, to allow personal
contact in this field between workers of vari¬
ous nationalities, and to make possible the
translation and publication of research results
obtained during the war in former enemy and
enemy-occupied territories.

4. The Pacific Foundation War Memorial

That the Pacific Science Conference approve
the concept of the Pacific Foundation: "Es-

Recommendations of Pacific Science Conference.
National Research Council

c.

52

NOTES

53

tablished as a memorial to all those who
served with the Armed Forces of the United
States in the Pacific area.” The purpose of
this Foundation is to create a living war
memorial devoted to the advancement of
knowledge through research and conservation.

5. Field Stations

That scientific research base stations be es¬
tablished in Hawaii and Guam to work in
close co-operation with existing research or¬
ganizations and institutions in these areas.

That subsidiary stations be established in the
following categories:

a. Floating stations consisting of vessels
equipped for specific fields of research.

' b. Advance base stations for both marine and
terrestrial research on various types of
islands and at the extremes of environ¬
mental conditions available.
c. Liaison stations to promote, in co-opera¬
tion with allied agencies, research in the
following areas: the Solomon Islands,
Australian New Guinea, French Oceania,
Indonesia, the Philippines, and the Gala¬
pagos.

That steps be taken toward the establish¬
ment of a base research station for various
types of scientific investigation in the
Galapagos Islands, making use of existing
installations. This base should be of a
permanent nature because of the impor¬
tance of maintaining continuous oceano¬
graphic, biological, and meteorological
records from this island outpost of South
America. By way of specific illustrations
of projects for this station, it may be
pointed out that various elements of the
land fauna are little known; that the extra¬
ordinary humid zone of the south face of
the larger islands offers the opportunity
for a unique ecological mountain transect,
especially from Academy Bay or Indefa¬
tigable Island; that the more barren coasts
and islands afford simplified ecological
conditions, comparable to those of Arctic
islands, and provide a veritable field labo¬
ratory in themselves; and that the biologi¬
cal interest of these islands is so great that
conservation measures, under the control
of such a research station, are urgently
required.

6. Science Appraisal

That a survey be made of the state of our
knowledge in the various fields of science in
the Pacific. The appraisal would have for its
main object the compilation of a guide to
what has been done in these fields, including

a bibliography of the basic contributions. As
a result of the survey, an investigator would
know where further research in any field is
most needed. The publication resulting from
the appraisal would be a guide for investi¬
gators and administrators. In addition, the
guide would help in two specific kinds of
important undertaking:

a. Conservation measures to be agreed upon
by international action.

b. Commercial policies to be evolved in pro¬
gressive steps as international agreements
are made necessary by problems of the use
and distribution of natural resources.

7. Documentation Centers

That documentation centers be set up at
Washington and at Honolulu, some of their
functions to be:

a. Distribution of bibliographies on special
fields.

b. Maintenance of a clearing house on cur¬
rent researches and projects.

c. Publication of a list of American and for¬
eign scientists (with addresses) who have
active interest in the Pacific. This list
should be cross-referenced as to:

(1) The islands or ocean areas with which
each scientist has had first-hand expe¬
rience.

(2) The specific fields in which each scien¬
tist is most qualified to furnish co¬
ordinating information.

(3) The fields in which each scientist may
have become an authority even though
he may not have visited or worked in
the area.

d. Establishment of an archive of publica¬
tions, translations, and manuscripts deal¬
ing with Pacific researches.

8. Internships

That internships on Navy Survey vessels be
provided for one or more scientists, under the
supervision of an experienced scientist, to
gather scientific information and collections
from all areas in which these vessels may
operate.

9. Pacific Congress

That the projected Pacific Science Survey
encourage and assist in the organization of
the Seventh Pacific Science Congress as soon
as possible, in order to further co-ordination
of research already in progress or being
planned and to perfect arrangement for co¬
operation among countries of the Pacific
Basin.

10. Check Lists of Flora and Fauna

That distribution check lists of the different

54

PACIFIC SCIENCE, January, 1947

animal and plant groups in Oceania be pre¬
pared and published.

11. Descriptive Geography of Micronesia
That a Descriptive Geography of Micronesia
be compiled, covering land forms, floral and
faunal ecology, and human geography.
Wherever possible, fullest use should be
made of visual presentation of data, includ¬
ing aerial photographs that have already been
taken by Army and Navy Air Forces.

12. Scientific Appraisal at Bikini

That means be provided for continuation of
the appraisal of biological consequences of
the atomic bomb tests at Bikini, over a period
sufficiently long to cover the repopulation of
the waters and lands with plant and animal
life.

13. Specific Scientific Recommendations

That every assistance be given to the imple¬
mentation of the specific recommendations
for scientific research formulated at the con¬
ference in the fields of the anthropological
sciences, the earth sciences, oceanography and
meteorology, the plant sciences, public health
and medicine, and the zoological sciences.

RECOMMENDATIONS PERTAINING TO
INTERNATIONAL CO-OPERATION

1. That the Pacific Science Conference recog
nizes the urgency of international co-oper*
tion in scientific research and to that emJ
recommends to the Congress of the Unite '
States that the Act of August 9, 1939, whiU*
authorizes the United States to co-opera ^
with the American Republics in scientific
undertakings of mutual interest, be amende^
to authorize such co-operation with all it
eign countries.

2. That the governments of the countries
the Western Pacific be invited to consider th'-
establishment of visitors’ facilities at the prffi •
cipal centers where considerable research
facilities for- resident scientists already exi*'J

3. That a committee be appointed to investigate
and recommend avenues of collaboration with
the United Nations and other international
organizations.

4. That the Pacific Science Conference urge the
establishment, in co-operation with allied
agencies, of liaison stations to promote re¬
search in the following areas: New Zealand,
the Solomon Islands, Australian New Guinea,
French Oceania, Indonesia, the Philippines,
and the Galapagos.

5. That the proposed Pacific Science Survey:

a. Collaborate with interested institutions and
individuals, American and foreign, in the
preparation of a series of regional floras.
It is suggested that a beginning be made
by drawing up plans for the publication
of (1) a Flora of Micronesia and (2) a
Flora of the Philippines, with a judicious
amount of preliminary field work in both
areas.

h. Encourage field work looking toward pub¬
lication of other regional floras in the Pa¬
cific and Oriental areas.

That, to forward these objectives, requests
from countries interested in securing
American co-operation be welcomed in
order to further the preparation of re¬
gional floras, which, in preliminary edi¬
tions, may extend only to the genera.

That field work be encouraged in those
areas for which the data in hand are ob¬
viously inadequate, as, for example, the
New Hebrides and the Solomon Islands.

6. That the Pacific Science Conference is highly
gratified to learn of the establishment of the
"Institut Francais d’Oceanie” and appreciates
the opportunity afforded by the invitation of
the French Government to American scientists
to make use of the institute’s facilities and to
co-operate with French scientists in furthering
knowledge of the Pacific area through a wide
range of scientific research.

7. That encouragement be given' {a) to the
establishment by the government of the
Netherlands Indies of a scientific research
station at Hollandia, Dutch New Guinea,
and (A) to the use of former Army installa¬
tions for this purpose. The location of a
scientific field station at Hollandia would
greatly facilitate the scientific exploration of
many parts of New Guinea. It is recom¬
mended that, if established, the station be
provided with a small vessel suitable for
coastwise operations and, if possible, plane
facilities. American scientists would welcome
an invitation to utilize the facilities of such a
station and to co-operate with scientists work¬
ing there.

’ 8. That the survey of the algae and algal re¬
sources of Philippine and Indonesian waters,
begun before the war by collaboration with
Philippine and Dutch scientists and institu¬
tions, be continued and be extended to cover
the Pacific by drawing into collaboration all
agencies and persons willing to contribute
toward this end. The food and fertilizer
values of algae, and also their value as raw

NOTES

55

materials in the preparation of commercial
products, should be appraised.

9. That international correlation and standardi¬
zation of nomenclature and of methods of
measurement be established by international
committees representing all nations working
in the general Pacific area. (This relates par¬
ticularly to land forms, rock and soil types,
geological timetables, etc.)

10. That the Pacific Science Conference go on
record as expressing its recognition of the
urgent need for support in the rehabilitation
of scientific libraries and scientific collections
destroyed during the war.

11. That in the Philippines a scientific center
supported by private funds be established as
an aid to scientific work and studies. Such a
center should co-ordinate its activities with
existing government bureaus and institutions.

SPECIFIC RECOMMENDATIONS

A co-ordinated program of scientific research
for the Pacific Islands (under American or for¬
eign administration) has been formulated. Recom¬
mendations are as follows:

I. DIVISION OF ANTHROPOLOGICAL
SCIENCES.

General preamble. It is recommended that this
Conference, in collaboration with local inhabit¬
ants, encourage investigation of problems concern¬
ing the welfare of the people of the Pacific Islands.

A. General studies that should be stressed in¬
clude:

1. A comprehensive anthropological survey,
covering each of the major divisions of
the subject, priority to be given to an
ethnographic study of Micronesia.

2. Human geography.

3. A survey of Micronesian linguistics and
the establishment of standards of pho¬
netic transcription, with the publication
of textbooks of native languages.

4. A survey of the social, economic, and
political structure of the present-day cul¬
tures of Micronesia.

5. Intensive study of the effects upon Mi¬
cronesian societies of non-indigenous con¬
tact, such as Spanish, German, Japanese,
and American culture, as well as of alien
civil administration.

6. Survey of Micronesian nutrition, diet,
dietary therapeutics, food habits, and
production and preparation of food.

B. Specific studies that should be initiated as

soon as possible include:

1. A physical anthropological survey of the
Micronesians.

2. Problems arising from increase and de¬
crease of population.

3. Child growth and development.

4. Race mixture.

5. Land utilization in Micronesia, to be
studied with a view to determining those
areas best suited for indigenous food
crops and those best suited for commer¬
cial crops.

6. Systems of land tenure, fishing rights,
and property concepts in Micronesian
cultures.

7. Cultural conditioning (including the ef¬
fects of the school system) of the child
from infancy to maturity, to be studied
in significant Pacific areas by anthropol¬
ogists, psychologists, and educators.

8. Native trade, and methods of developing
natural resources.

II. DIVISION OF EARTH SCIENCES.

A. Scope. The Earth Sciences as herein defined
include Physical Geography, Geology, and
Geophysics.

B. Regions of interest. It is recommended that
this Conference encourage a research pro¬
gram in the Earth Sciences for the entire
Pacific Basin, with particular emphasis on
Micronesia.

C. Recommended Investigations.

1. Geology. Systematic geological surveys
and areal geological mapping of selected
key islands and, later, of island groups
in the Pacific as a basis for scientific re¬
searches in paleontology, stratigraphy,
petrology, and physiography; terrain
studies and engineering-geologic inter¬
pretations of water supplies, construction
materials, foundation conditions, min¬
eral resources, and soil types.

2. Soil Science. Reconnaissance soil surveys
of the whole Pacific area and detailed
soil surveys of regions of agricultural
importance, including research through
field stations and field working parties
for each of the principal soil types as to:

a. Genetic formation in relation to en¬
vironment.

b. Mechanical properties in relation to
engineering needs.

c. Crop adaptability and response to ir¬
rigation and fertilization in relation
to production, nutrition, conservation,
flood control, and land use.

3. Agricultural Committee. Appointment
of a Committee or Division on Agricul¬
ture under the Pacific Science Survey, to
assist administrators of islands in the co¬
ordination of action programs designed
(a) to increase food production and im-

56

PACIFIC SCIENCE, January, 1947

prove efficiency in growing all agricul¬
tural and forest products, whether in¬
tended as food or as industrial raw ma¬
terials for local industries and for com¬
merce, and (b) to conserve soils, waters,
and forests.

4. Gravity Investigations. Gravity observa¬
tions at several atolls and at selected
points about the shores of the lands and
islands of the Pacific. It is recommended
that a U. S. Navy submarine, using
Vening-Meinesz pendulum apparatus, be
employed for one year to make gravity
observations at sea, particularly in the
vicinity of the Aleutian trench and other
trenches. These observations are intended
to increase knowledge of geological
structure, of seismological conditions,
and of deflections of the vertical.

5. 'Precise Position Determinations. Employ¬
ment of all possible means to improve
the quality of basic astronomic position
determinations and extend first-order tri¬
angulation when feasible, in order to
obtain better basic geographic positions
for airports and for Loran and other
navigational installations and for general
aerial mapping.

6. Seismology. Establishment and perma¬
nent maintenance of four additional seis¬
mological stations containing modern
seismographs at selected points in the
western Pacific, to improve the science
of locating and studying Pacific area
earthquakes, to increase knowledge of
the geologic structure of the area, to
study the relations between earthquakes
and seismic sea waves, and to evaluate
earthquake and seismic hazards.

7. Microseisms. Additional study of extra-
tropical meteorological disturbances in
the north Pacific and Alaska area,
through extending to the Aleutian Island
area the observation and study of micro¬
seisms and their relation to meteorology.

8. Seismic Prospecting. Promotion of seis¬
mic prospecting work about the subma¬
rine trench areas and at selected points
about the shores and in the depths of the
Pacific, for determination of sediment
depths and of underlying rock structure.

9. Structure of Atolls. Investigation of the
structure of typical atolls, including sub¬
merged, atoll-like structures, by seismic
prospecting and core drilling, magneto¬
meter prospecting, gravitational pros¬
pecting, and detailed bathymetric sur¬
veys of their flanks and approaches.

10. Volcanology. Descriptive geological and
geophysical observations of various types
at a number of active volcanoes, par¬

ticularly any volcano showing unusual
activity. Modern seismic, gravimetric,
electric, and magnetometric techniques
should be applied to the determination
of underground structure in volcanic re¬
gions.

11. Physics of the Ionosphere and Tropo¬
sphere. Investigation of atmospheric fac¬
tors influencing the propagation of elec¬
tromagnetic radiation at all frequencies,
in relation to radio communication and
geomagnetism and to telemetering and
tracking at extreme altitudes.

12. Geomagnetism. Establishment and main¬
tenance of permanent magnetic observa¬
tories in the Aleutians, in the Philip¬
pines, at Samoa, and at Christmas Island
or Jarvis Island, for continuous record¬
ing of magnetic variations; employment
of two field magnetometer parties for
one year to investigate magnetic condi¬
tions at islands and along the shores of
the Pacific, and complete repetition of
such investigation after an interval of
five years.

Suitable airborne magnetic instruments
should be utilized and further improved
as necessary for investigations generally
over the Pacific Ocean area, and two
planes so equipped should be employed
for one year in general magnetic work.
These studies are designed to improve
nautical and aeronautical charts, to in¬
vestigate magnetic anomalies as related
to geologic structure and volcanism, and
to achieve other scientific results.

13. Hydrology. Investigation of infiltration
of rainfall, stream flow and run-off, and
flood hazards, and preparation of isohy-
etal maps; investigation of storage and
diversion for irrigation and hydroelectric
power development, and of ground water
resources; and research in dynamics of
erosion.

14. Mineral Industries. Initiation of pros¬
pecting operations to evaluate the im¬
portance of mineral deposits in the local
economy, as the information on these
deposits becomes available from geologi¬
cal studies.

15. Mapping Activities. It is recommended
that hydrographic, topographic, geophys¬
ical, and submarine contour mapping be
supported and correlated.

16. Geologic Timetable. Gradual establish¬
ment of a standard stratigraphic se¬
quence of geologic formations in the
Pacific. In order to understand the suc¬
cession of geologic events in the area, a
geologic timetable correlating the rock
formations of the islands with those of
the surrounding mainlands is essential.

NOTES

57

III. DIVISION OF OCEANOGRAPHY AND
METEOROLOGY.

The Division recommends:

A. That the United States immediately initiate
preparations for a comprehensive program
of investigations in the marine sciences in
the Pacific area and that the following topics
be a part of the general program of investi¬
gation.

1. Currents.

a. The vertical, horizontal, and seasonal
distribution of tidal and non-tidal
currents.

2. Interrelations of Sea and Atmosphere.

a. Data on waves: height, length, and
period, including frequency and direc¬
tion of waves of different magnitudes
by months.

h. Heat exchange with atmosphere.

c. Water exchange with atmosphere.

3. Distribution of Physical Properties.

a. Complete survey, by areas and sea¬
sons, of temperature and salinity in
the ocean, the variability of these
characteristics, and the factors affect¬
ing this variability.

b. Transparency of sea water and how
it is affected by seasonal variations in
coastal waters.

4. Distribution of Chemical Properties.

5. Characteristics of Sea Bottom.

a. Composition, firmness, color, and geo¬
logical character of bottom sediments,
in offshore areas, channels, and har¬
bor mouths.

b. Beaches and wave zone, including
trafficability characteristics of beaches
and shallow water (hardness, cohe¬
siveness, mechanical composition, and
bearing capacity).

c. Harbor and coastal silting and erosion
resulting from waves and currents.

B. That the co-operative tidal program now in
operation by the Coast and Geodetic Survey
be continued and expanded in the western
Pacific until observations are obtained over
a sufficient period of time for a satisfactory
analysis of the Pacific tides.

C. That support be given to the maintenance
and expansion of the network of meteoro¬
logical surface and upper-air stations in the
Pacific area, including weather reconnais¬
sance squadrons and special stations set up
for the purpose of observing sferics, micro¬
seisms, sea swell, and radar as these relate
to weather. The program would include:

1. Study of medium and high altitude mete¬
orology (above 5,000 feet), particularly

of the northern and western areas.

2. Basic theoretical work on the cause and

maintenance of fog and a synoptic and
geographic study of the distribution of
fog in the North Pacific.

3. Study of the thermal, moisture, and wind
micro-structure of the lower layer of at¬
mosphere (less than 5,000 feet).

4. Research in formation, structure, and
motion of typhoons.

5. Study of meteorological conditions caus¬
ing anomalous propagation of ultra-high-
frequency radio and radar in the west¬
ern Pacific area.

D. That support be given to the establishment
of an office to supply information regard¬
ing locations of meteorological stations and
available meteorological observations, and
to furnish meteorological advice to agencies
or individuals planning scientific research
projects in the Pacific area.

E. That, if expeditions are organized in the
Pacific area, provision be made for taking
detailed sea-surface, meteorological, and
radiation observations that are necessary to
investigate the energy exchange between
ocean and atmosphere and to study other
problems.

F. That, since the already available data from
the Pacific area must be thoroughly studied,
analyzed, and evaluated before a compre¬
hensive and well-integrated program in
oceanography (marine sciences) can be de¬
cided upon:

1. An office be established under the Na¬
tional Research Council to study, analyze,
and evaluate the data and make the re¬
sults available to the planning group.

2. This office be adequately staffed by a
head scientist and necessary clerical and
technical assistants.

G. That, because of the different requirements
in the various areas and in the several
fields of science, four vessels be procured,
equipped, and operated for general investi¬
gations on the high seas and that eight ves¬
sels be procured, equipped, and operated
for regional investigations. One of the four
large vessels and the eight smaller vessels
should be used primarily for research in the
natural resources of the sea and, in addition,
should be equipped to participate in general
oceanographic studies conducted by means
of the other three large ships.

IV. DIVISION OF PLANT SCIENCES.

A. Whereas there has been a lack of correla¬
tion among the botanical endeavors of scien¬
tists of different countries, and

Whereas the most rapid progress and scien¬
tific co-operation among these countries in
the study of botanical problems of the Pa-

58

PACIFIC SCIENCE, January, 1947

cifk will be brought about by personal con¬
tacts.

Be it resolved that botanical missions be sent
to Japan, China (especially Formosa), and
Korea ( 1 ) to obtain specimens, photographs
of type and other important specimens, pub¬
lications, translations of publications, and
manuscripts, (2) to ascertain the needs of
Pacific and Oriental botanists for correspond¬
ing scientific materials from America, and
(3) to arrange for co-operation among the
scientists of the participating countries.

B. Whereas ethnobotanical investigations are
frequently neglected by both botanists and
anthropologists, and

Whereas the field of ethnobotany is one in
which much useful research may be done by
workers in other fields.

Be it resolved that the National Research
Council recommend methods of securing
comparable linguistic terminology on eco¬
nomic botany and primitive agriculture from
the boundaries of India through Polynesia.

C. Whereas the experiment station of the
South Seas Bureau of the Japanese Govern¬
ment at Kolonia, Ponape, has a large collec¬
tion of economic plants, two large modern
buildings, and many acres of tillable land,
and

Whereas both wet land and well-drained
land have been under active cultivation, and
the area is well suited for experiment sta¬
tion work,

Be it resolved that agricultural experimenta¬
tion be continued at this station.

D. Whereas the food production capacity of
many Pacific Islands might.be increased by
judicious enrichment of the flora through
plant introduction, and

Whereas many other products of utility
might be made available to isolated peoples.

Be it resolved that measures be taken to
introduce widely throughout the Pacific new,
useful, disease-free plants (including forest
trees), favoring those that may have two or
more utilities, such as use for lumber, for
food, for fiber, for tannin.

E. Whereas modern modes of transportation
and increased contacts of the island groups
of the Pacific with each other and with the
continents are greatly increasing the danger
of spread of plant diseases.

Be it resolved that, whenever possible, a sur¬
vey of parasitic fungi and other disease-pro¬
ducing organisms be made a part of plant
science projects sponsored in the Pacific area,
and

Be it further resolved that every precaution
be observed to prevent the spread of dis¬
eases and pests (including weeds) . Increased
attention should be given to quarantine reg¬
ulations and renewed studies should be
made of the efficacy of quarantine measures.

F. Whereas certain problems of marine biology
are of practical significance in the operation
of ships,

Be it resolved that the following studies be
undertaken :

1. Biological study of the distribution and
habits of marine fouling organisms.

2. Investigation of the distribution and
causes of phosphorescent waters.

3. Survey of fouling in certain areas of im¬
portance to shipping in order 'to deter¬
mine maximum depth of significant ma¬
rine growth.

4. Survey in representative areas of the
main marine growth forms, their seasons
of attachment and rates of growth.

5. Distribution of kelp and other large
algae.

V. DIVISION OF PUBLIC HEALTH AND
MEDICINE.

The Division recommends: That co-operative
studies be conducted by qualified workers in the
field of medicine, together with workers in the
allied sciences, as follows:

1. Problems of the Pacific that have been
demonstrated to be of particular impor¬
tance to "non-immune whites.”
a. Diseases particularly prevalent in the
Pacific and of major importance:
dysenteries, malaria, hepatitis, dengue
complex, and tropical dermatitis; also
diseases of lesser but peculiar impor¬
tance: scrub typhus, schistosomiasis,
filariasis, and Japanese B encephalitis.
h. Dissemination and implantation of
disease, with particular reference to
quarantine procedures.

c. Possible deleterious effects of the
widespread use of DDT.

d. Utilization of special opportunities
that may arise in the study of respira¬
tory diseases.

e. Determination of factors that have
led to the absence of arteriosclerosis
and hypertension, which are pre¬
sumed to be largely or entirely absent
in certain native population groups of
the Pacific.

/. Nutritional problems that arise from
residence in the tropics.

NOTES

2. Problems that have been demonstrated
to be of particular importance to native
populations.

a. Diseases of special importance to na¬
tives, such as tuberculosis, yaws, ma¬
laria, leprosy, and helminthiases.

b. Medical education for native doctors
and nurses of the area.

c. Feasibility of instituting modern pub¬
lic health procedures in certain native
groups.

d. Native nutrition.

VI. DIVISION OF ZOOLOGICAL SCIENCES.

General preamble. It is proposed that this con¬
ference aid and stimulate a broad basic program
of zoological collecting in, and a study of, the land
and water areas that fall within its scope. To this
end it is recommended:

1. That a "clearing house” be established
to assemble data on unstudied scientific
materials from the Pacific; that such
materials and specimens be located and
listed as to scope, place, time of collec¬
tion, and collector; that such informa¬
tion be made available to all interested
scientists; that staff and funds be secured
for the organization, study, and identifi¬
cation of existing Pacific collections and
for the preparation of reports, such staff
and funds to be used to strengthen the
appropriate departments of existing in¬
stitutions.

2. That special attention be given to the
survey of the fish life of the Pacific, in¬
cluding:

a. Preparation of a bibliography of the
entire Indo-Pacific fauna.

b. Collection of specimens in all parts of
Oceania.

c. Revisions of the fauna, group by
group, at several institutions.

d. Preparation and publication of a
check list and successive revisions.

3. That the governments of the Philip¬
pines, of Canada, of the United States
(especially in the Territory of Hawaii
and the States of Washington, Oregon,
and California), and other countries con¬
cerned, be encouraged to:

a. Undertake a thorough, closely inte¬
grated investigation of the tuna-like
fishes of the tropical Pacific.

b. Co-ordinate their research with the
needs of the industry.

4. That researches be conducted on the
reef and lagoon fisheries of the Micro-
nesian Islands and that the products of

, these fisheries be allocated for native use.

59

5. That a vessel and crew be furnished to
facilitate a survey of the fishes and mol-
lusks of the Carolines.

6. That advanced studies of evolution be
supported by the establishment of ade¬
quate laboratory facilities in the Hawai¬
ian Islands, where advantage can be
taken of the unique development of
Drosophila and of other animals and
plants in that area, which may yield data
of utmost importance bearing on the
evolution of living organisms.

7. That the zoological survey of the Pa¬
cific area be implemented by the provi¬
sion of funds for permanent staff and
visiting fellows to utilize the facilities of
the biological stations proposed.

8. That the zoological work of shore and
floating biological stations relating to the
marine fauna be co-ordinated with bio¬
logical oceanographic work undertaken
as part of the total survey of the Pacific
envisaged.

9. That investigations in animal husbandry
in the Pacific region be undertaken to
cover:

a. Determination of existing livestock
populations, including adaptabilities.

b. Possible improvement of livestock pro¬
duction by the use of better adapted
breeds and improved species, such as
Indian cattle.

c. Study of existing livestock diseases
and parasites on the islands.

10. That early and continued attention be
given to the following biological prob¬
lems:

a d Comprehensive investigations of the
zooplankton.

b. 1 Determination of causes and seasonal

variation of phosphorescent waters.

c. 1 Analysis of the character and preval¬

ence of background noises of animal
origin. This will involve studies of
(1) sound production, and (2) geo¬
graphical and seasonal distribution of
sound-producing animals and their
ecological relationships.

d. Regional studies on the biology of
fouling and boring organisms.

e. Researches on the biology of reef¬
building corals, with particular refer¬
ence to composition of populations
and to growth rates in different areas.

/. Ecological studies of poisonous and
otherwise dangerous animals.

g. Ecological studies of termites.

h. Survey of the populations and major
breeding grounds of the larger marine
birds.

1 This should be correlated with other researches and pro¬
grams in physical oceanography and fisheries biology.

Micronesian Expedition of University of Hawaii,
Summer of 1946

Intensive field research in several branches of
natural science was carried on in the islands of
Micronesia during the summer of 1946 by groups
of University of Hawaii faculty members. Based
on a reconnaissance made by a team of four
professors in this region in December, 1945, the
summer program initiated research in this oceanic
area, which has long been closed to most American
scientists. The University of Hawaii surveys were
planned and carried out through its Pacific Islands
Research Committee, headed by Dean Paul S.
Bachman. Transportation, housing, and other
facilities were furnished by the thoroughgoing
co-operation of United States Navy officials.

Most of the scientific investigations in Micro¬
nesia have been made by the Germans and Japa¬
nese, under which nations the islands have been
administered for the past half century; and these
studies, some of which still have value, were pub¬
lished in the German or Japanese languages.
Much investigation, however, still remains to be
carried on in fields that have not been touched or
have been treated inadequately. With the likeli¬
hood that these islands will continue for some
time under the control of the United States, it
appears obvious that neW and full information,
reported in English, will be urgently demanded.
The University of Hawaii, the American univer¬
sity closest to Micronesia, is in an advantageous
position geographically, and from it other studies
of the islands will be launched and continued in
the future. Its faculty includes men who by train¬
ing, experience, and interest are well fitted to carry
on these studies; many of these men have already
done field work on other Pacific islands. The
administration, moreover, has recently filled posi¬
tions with other men who have particular qualifi¬
cations for conducting studies in this area.

A scientific study of Micronesia has important
practical values. The published findings will con¬
tribute to a better acquaintance with these terri¬
tories newly under American protection and to
an understanding of the clash of cultures in these
scattered islands — traditionally steppingstones in
the oceanic travel lanes between Asia and the
middle Pacific; and these findings will provide a
storehouse of scientific knowledge to be drawn
upon by workers in many fields. Information is
needed at once if the United States is to carry on
a policy of developing the government, education,
and economy of the Caroline, Marianas, and Mar¬
shall groups.

A summary of the field work sponsored by the
University in the several branches of natural
science during the summer of 1946 here follows.

BOTANY

A party consisting of Dr. Harold St. John,
chairmali of the Department of Botany, Dr. Don¬
ald P. Rogers, assistant professor of botany, and
Richard S. Cowan, graduate assistant in botany,
left Pearl Harbor on August 7 on LSM 382, a
vessel which served as their base for most of the
trip. In this Navy ship they surveyed the islands
of Kwajalein, Likiep, Ailuk, Utirik, Mejit, and
Wotje; thereafter they visited by seaplane the
islands of Namu, Jaluit, Ailinglapalap, and Ebon.
(Part of the route was planned to avoid duplicat¬
ing efforts of scientists visiting Bikini on Opera¬
tion Crossroads.) They returned on September 12
by air from Kwajalein, headquarters of the mili¬
tary government of the Marshall group.

One purpose of the trip was to make a general
botanical exploration of as many of the Marshall
atolls as was possible in the available time.
Though these islands do not possess a large flora,
they are little known because of their remoteness.
Special attention was given by the University team
to the ethnobotany of the crops of the natives;
this subject, often neglected by the agriculturist
and the botanical seeker of new species, might well
reveal facts on the origins of certain crops, and
studied in collaboration with qualified anthropol¬
ogists might give new evidence on the migration
routes of Micronesian and Polynesian native
groups.

One to four days were spent on each of the
islands studied, a time sufficient for a satisfactory
initial exploration. In recent years, it was found,
most of the islands have been turned into copra
plantations, but all species of the higher plants
reported by Chamisso in 1817 were found except
one; these native plants were found fringing the
seashore or sprouting in the coconut plantations.
One of the most prominent of native trees is the
pandanus — called "bop” by the Marshallese, who
distinguish by name at least 20 varieties. A num¬
ber of observations were made of driftwood logs
found on island shores; several of these were
apparently from trees native to northwestern
United States. Study of these drift logs should
give data on ocean currents and on the spread of
certain plants in the Pacific.

60

NOTES

61

An obvious gap in botanical knowledge of the
Pacific is found in the lack of collections of fungi,
algae, lichens, and bryophytes. Except for certain
parasitic families which have been studied in Ha¬
waii, the fungi of the main Pacific area are quite
unknown. Identification of the forms collected
should be a welcome contribution to mycology and
should eventually add to present knowledge of
distribution patterns of Pacific fungi and perhaps
raise new problems in comparative morphology.
For algae the region has been almost equally terra
incognita, and the bryophytes and lichens, except
those of Hawaii, have not previously been col¬
lected. The results of the summer expedition
should eventually lead to the publication of sev¬
eral studies on the fungi and other cryptogams of
the Marshall Islands.

Some 625 collections of higher plants and 500
of lower plants were brought back for further
study at the University. These specimens will
eventually be placed in the Bishop Museum for
permanent record. Many photographs from the
islands, both black and white pictures and color,
were taken, as well as a number of 8 -mm. motion
picture films.

ZOOLOGY AND BACTERIOLOGY

A party consisting of Dr. Robert Hiatt, chair¬
man of the Department of Zoology and Entomol¬
ogy, Dr. Harvey I. Fisher, assistant professor of
zoology. Dr. Floyd W. Hartmann, acting chair¬
man of the Department of Bacteriology, and two
assistants, Leo Fortess and Eveni Levi, left by air
from Honolulu on July 17, and after a 5 -day stay
at Guam, went to the island of Yap by LCI. After
a stay of almost a month, they left Yap on August
22 by air and returned, via Saipan and Guam, to
Honolulu on August 28. Dr. Fisher collected ver¬
tebrate specimens and Dr. Hiatt collected inver¬
tebrate specimens.

The ornithology, mammalogy, and herpetology
of the Micronesian chain have been virtually un¬
known to English-speaking students. Study of the
vertebrate fauna in this area is of interest from the
viewpoints of taxonomy, distribution, and natural
history, because detailed studies are available for
certain surrounding regions. One purpose of the
survey was to note the movement of bird species
from the Asiatic continent southward and also
eastward into the Palaus and the Marianas chain,
as well as movement northward within this chain;
in other words, to compare the avifauna of Yap
with that of Peleliu and Guam. A complete zoolo¬
gical exploration of the Yap islands was made.
About 150 skins of birds and mammals and some
50 alcoholic and skeletal specimens were brought
back for further study. In addition, a number of
reptiles were collected. It is hoped that present
investigations may demonstrate means of saving
from extinction some of the particular endemic

vertebrate species now isolated on small groups
of islands or on individual islands; certainly the
work will provide much new material for studies
in morphology, natural history, ecology, and dis¬
tribution.

No comprehensive survey of the invertebrate
fauna of the Pacific islands formerly mandated to
Japan has previously been made; indeed, the dis¬
tribution of species west of Hawaii is virtually
unknown. Studies of the specimens collected will
provide information concerning the taxonomy,
ecology, and distribution of these invertebrates.
Thousands of marine invertebrates were collected
and placed in preservatives for shipment to the
University for further examination. Subtidal areas
were explored with the aid of "skin” diving and
"self-contained” diving techniques. The aim in
exploring Pacific invertebrate life was not merely
to identify and map the distribution of inverte¬
brates in the Yap group, but also to compare these
results with the dispersal of species previously
known to exist in the Philippine and Indo-China
area to the west and with that of species from the
Great Barrier Reef of Australia to the south. Such
studies might reveal the main routes of dispersal
of species from the Asiatic mainland east into the
Pacific. It is expected that future explorations will
concentrate on island groups lying between Yap
and the Hawaiian Islands.

Dr. Hartmann, in addition to collecting a num¬
ber of autopsy specimens from rats in co-operation
with Dr. Alicata on a study of the incidence of
leptospirosis in the Carolines (see next section),
carried on research on dental caries among the
school children of Yap. Duplicate saliva speci¬
mens for several hundred children whose physical
examinations and dental records were made avail¬
able by Navy personnel were submitted to bac¬
teriological examination. The Yap leper colony
was visited, and observations were there made on
administration, sanitation problems, and progress
in rebuilding dwellings. A number of important
bacteriological problems present themselves in the
Caroline Islands, and these problems will have to
be faced by the new American administration.

A large number of black and white photo¬
graphs, as well as 200 colored 35-mm. slides and
2,000 feet of 16-mm. color motion pictures, were
brought back as records of the 6-week visit to
Micronesia.

PARASITOLOGY

Dr. Joseph E. Alicata, head of the Department
of Parasitology, University of Hawaii Agricul¬
tural Experiment Station, left by air on July 18
on a visit to Truk and Ponape in the eastern
Carolines, and returned August 31.

Parasitic diseases are of paramount concern in
the Pacific area and are responsible for retardation
in the development of island areas and in the

62

PACIFIC SCIENCE, January, 1947

expansion of the livestock industry. Although
scattered reports are available on the occurrence
of parasitic diseases of man in Micronesia, very
little is known about their actual prevalence; nor
is much known about parasites which affect do¬
mestic animals. Information acquired through the
present study and similar explorations planned for
the future should assist in the adoption of more
adequate health measures for man and, through
parasite control, improved quality and quantity
of animal products. The geography of parasitic
diseases of the Pacific area and the factors respon¬
sible for the spread of such diseases need much
investigation in order to formulate intelligent con¬
trol measures.

The major purpose of the trip was to collect
ectoparasites and endoparasites of economically
important animals such as cattle, swine, and poul¬

Survey of Micronesia by U. S

An economic survey of Micronesia by 25 spe¬
cialists attached to the U. S. Commercial Company
was completed in the autumn of 1946. It was
carried out primarily to discover how fully these
formerly Japanese-mandated islands could be de¬
veloped to promote the welfare of the native
populations.

A comprehensive report is in preparation for the
benefit of government agencies and others con¬
cerned. It is expected that this will contain factual
data of value in determining future policies with
reference to the islands, particularly as regards the
fostering of a self-sustaining native economy and
the avoiding of commercial exploitation.

Teams of scientists were assigned to remain in
the various areas for a minimum of 3 months of
observation. In addition to six area economists,
specialists on the study included botanists, hor-

try, and to investigate the possible occurrence of
murine leptospirosis or Weil’s disease in Micro¬
nesia. Data collected in the Hawaiian Islands in¬
dicated that this disease might be widespread in
the Pacific area where rainfall is usually high and
rodents (the agency which usually transmits the
disease) are abundant. A number of rodents were
trapped in the Caroline Islands. Examination of
kidney samples from these rodents for leptospirae
will provide a basis for better judgment on the
recognition and control of this disease in man.
Dr. Alicata also carried on a study of the inci¬
dence of helminthic infection among the natives ;
of Ponape and Truk. He experimented as well
with the use of copper sulfate preparations for
control of the giant land snails which destroy
much island vegetation.

Commercial Company, 1946

ticulturists, agronomists, animal husbandry spe¬
cialists, entomologists, soils experts, nutritionists,
mineral geologists, water geologists, fish special¬
ists, and specialists in handicrafts.

An LCI, made available by the Navy, carried the
several specialists from island to island throughout
Micronesia, and in port served them as a floating
laboratory and base of operations. This vessel
returned to Honolulu at the end of the survey,
carrying specimens, records, and samples of native-
made products. Many of these latter will find
their way into the collection of the Bishop Mu¬
seum.

The study was carried on originally under the
direction of Dr. Douglas L. Oliver, who retired in
September as head of the agency for the Middle
Pacific. At present the work is headed by Richard
B. Black, former expedition director.

APRIL, 1947

No. 2

E4CIFIC

SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

IN THIS ISSUE: Alicata — Parasites of Domestic Animals
in Hawaii • Palmer — Fault at Waimea, Oahu • Rogers —
Fungi of the Marshall Islands , Central Pacific Ocean • Dean
and Hawley — Chloride and Oxygen Analysis Kit • St. John —
New Species of Car ex (Cyperaceae) from Fiji • NOTES

THE UNIVERSITY OF HAWAII
HONOLULU, HAWAII

BOARD OF EDITORS

A. Grove Day, Editor-in-Chief, Department of English, University of Hawaii

Ervin H. Bramhall, Department of Physics, University of Hawaii

Vernon E. Brock, Director, Division of Fish and Game, Territorial Board of
Agriculture and Forestry

Harry F. Clements, Plant Physiologist, University of Hawaii Agricultural
Experiment Station

Charles H. Edmondson, Zoologist, Bishop Museum, Honolulu, T. H.

Harvey I. Fisher, Department of Zoology, University of Hawaii

Frederick G. Holdaway, Head, Department of Entomology, University of
Hawaii Agricultural Experiment Station

Maurice B. Linford, Head, Department of Plant Pathology, Pineapple Research
Institute, Honolulu, T. H.

A. J. Mangelsdorf, Geneticist, Experiment Station, Hawaiian Sugar Planters’
Association, Honolulu, T. H.

G. F. Papenfuss, Department of Botany, University of California, Berkeley 4,
California

Harold St. John, Chairman, Department of Botany, University of Hawaii

Chester K. Wentworth, Geologist, Honolulu Board of Water Supply

SUGGESTIONS TO AUTHORS

Contributions to Pacific biological and physical
science will be welcomed from authors in all parts
of the world. Manuscripts should be addressed to
Dr. A. Grove Day, Editor, Pacific Science, Uni¬
versity of Hawaii, Honolulu 10, Hawaii. Use of
air mail for sending correspondence and brief
manuscripts from distant points is recommended.

Manuscripts will be read promptly by members of
the Board of Editors and by other competent
critics. .

Manuscripts may run from 1 Fo 30 pages in
length. Authors should not overlook the need for
good brief papers presenting results of studies,
notes and queries, communications to the editor,
or other commentary.

Preparation of Manuscript

Although no manuscript will be rejected merely
because it does not conform to the style of Pacific
Science, it is suggested that authors follow the
style recommended below and exemplified in the
journal.

Title. Titles should be descriptive but brief. If a
title runs to more than 40 characters, the author
should also supply a "short title” for use as a
running head.

Manuscript form. Manuscripts should be typed
on one side of standard-size, white bond paper
and double-spaced throughout. Pages should be

consecutively numbered in upper right-hand cor¬
ner. Sheets should not be fastened together in any
way, and should be mailed flat. Inserts should be
either typed on separate sheets or pasted on proper
page, and point of insertion should be clearly indi¬
cated.

Original copy and one carbon copy of manuscript
should be submitted. The author should retain a
carbon copy. Although due care will be taken, the
editors cannot be responsible for loss of manu¬
scripts.

Introduction and summary. It is desirable to state
the purpose and scope of the paper in an intro¬
ductory paragraph and to give a summary of
results at the end of the paper.

Dictionary style. It is recommended that authors
follow capitalization, spelling, compoundings ab¬
breviations, etc., given in Webster’s New Inter¬
national Dictionary (unabridged), second edition;
or, if desired, the Oxford Dictionary. Abbrevia¬
tions of titles of publications should, if possible,
follow those given in U. S. Department of Agri¬
culture Miscellaneous Publication 337.

Footnotes. Footnotes should be used sparingly and
never for citing references (see later). Often,
footnotes may better be incorporated into the text
or omitted. When used, footnotes should be con¬
secutively numbered by superior figures through-
[ Continued on inside back cover ]

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

Vol. I APRIL, 1947 No. 2

.

Previous issue published January 5, 1947

CONTENTS

PAGE

Parasites and Parasitic Diseases of Domestic Animals in the Hawaiian Islands.

Joseph E. Alicata . 69

Fault at Waimea, Oahu. Harold S. Palmer . 85

Fungi of the Marshall Islands, Central Pacific Ocean. Donald P. Rogers ... 92

A Chloride and Oxygen Analysis Kit for Pond Waters. R. B. Dean and R. L.

Hawley . 108

A New Species of Carex ( Cyperaceae) from Fiji. Harold St. John . . . . 116

Notes:

Facilities for Research in the Natural Sciences in the Hawaiian Islands . . 119

Cover drawing by A. S. MacLeod

Pacific Science, a quarterly publication of the University of Hawaii, appears in January,
April, July, and October. Subscription price is three dollars a year; single copies are one
dollar. Check or money order payable to University of Hawaii should be sent to Pacific
Science, Office of Publications, University of Hawaii, Honolulu 10, Hawaii.

Parasites and Parasitic Diseases of Domestic Animals in the

Hawaiian Islands1

Joseph E. Alicata2

Parasites of animals have gained entrance
to the Hawaiian Islands for a century or
more largely with the importation of infected
animals from various parts of the world.
Because of the mild climatic environment
and other favorable factors, these parasites
have become established and now constitute
an agricultural problem of considerable eco¬
nomic importance. To what extent the peo¬
ple of these islands will be successful in
keeping other animal parasites and vectors
from entering, especially with the expansion
of air and sea transportation, remains to be
determined. Much is being done, however,
through quarantine, inspection, and other
Territorial and Federal regulatory measures
to prevent the introduction of additional
disease-producing organisms and vectors of
disease.

The parasites now present in domestic
animals in the Hawaiian Islands are to a large
extent the same as are found in continental
United States. This is true because most of
the animals found in the Islands have come
from that area. There are a few parasitic
forms, however, which have undoubtedly
been introduced from the Orient. These
include, at least, Fasciola gigantica Cobbold,
the common liver fluke of cattle, and Hy-
menolepis exigua Yoshida, a tapeworm fre¬
quently found in chickens. In spite of the
many parasitic diseases which have been in¬

1 Published with the approval of the Director
of the University of Hawaii Agricultural Experi¬
ment Station as Technical Paper 150. Manuscript
received November 25, 1946.

2 Parasitologist, University of Hawaii Agricul¬
tural Experiment Station, Honolulu, Hawaii.

troduced from continental United States,
some of the important ones affecting the
blood, such as anaplasmosis and piroplas-
mosis of cattle and dourine of horses, either
have not been introduced or have failed to
become established.

The present paper represents a resume of
internal and external parasites, and of their
intermediate hosts, if any, which have been
reported up to the present time from domes¬
tic animals in the Hawaiian Islands. Special
reference is given to certain species which are
of economic importance. Whereas consider¬
able data are available on parasites of chick¬
ens, cattle, horses, and swine, information on
those of other animals is up to the present,
time inadequate or entirely lacking. The
chief sources of information on the external
parasites reported in this paper have been
the scattered reports published by various
entomologists in the Islands. Data dealing
with internal parasites (protozoa, round-
worms, tapeworms, and flukes) and any of
their intermediate hosts have been secured
largely, except as indicated, from the various
reports and observations made by the writer
during the past several years.

Parasites of Poultry
protozoa

Coccidial organisms, Eimeria tenella Rail-
liet and Lucet, are the most important pro¬
tozoa affecting chickens. Infection with these
parasites is as troublesome in Hawaii as it is
anywhere else.

Pigeons in Hawaii are commonly infected

69

70

with the malarial organism Haemoproteus
columbae Celli and Sanfelice (Alicata,
1939c) . This protozoan lives in the red blood
cells and may be responsible for the produc¬
tion of anemia and low vitality. It is spread
among pigeons through the bite of the pigeon
fly, Pseudolynchia canariensis (Macquart),
which is generally distributed (Bryan, 1934) .
Histomonas meleagridis (Smith), the causa¬
tive organism of "blackhead” in turkeys, is
responsible for sporadic outbreaks of this
disease in various parts of the Islands.

ROUNDWORMS

Up to a few years ago gizzard-worms,
Cheilospirura hamulosa (Diesing), were
widespread among chickens and turkeys in
the Territory and were responsible for
anemia, emaciation, and deaths, especially
among chickens. The infection was checked
after the discovery and control of the follow¬
ing arthropods, which were found serving as
intermediate hosts (Alicata, 1936; 1938 b\
1939c): (coleoptera) Carpophilus dimi-
diatus Fabricius, Dactylosternum abdominale
(Fabricius), Dermestes vulpinus Fabricius,
Epitragus diremptus Karsch, Euxestus sp.,
Gonocephalus seriatum (Boisd.), Litargus
balteatus Lee., Oxydema fusiforme Woll.,
Palorus ratzeburgi (Wissm.), Sitophilus
oryzae (Linn.), Tenebroides nana Melsh.,
Tribolium castaneum (Herbst) , and Typhaea
ster corea Linn.; (orthoptera) Atracto-
morpha ambigua Bolivar, Conocephalus sal-
tator (Sauss.), and Oxya chinensis (Thun.) ;
(amphipoda) Orchestia platensis Kroyer.

The eyeworm, Oxyspirura mansoni (Cob-
bold), which utilizes the burrowing roach,
Pycnoscelus surinamensis (Linn.), is com¬
mon in chickens in the Islands (Alicata,
1936). This eyeworm has also been found
in the English sparrow ( Passer domesticus
[Linn.]) (Illingworth, 1931), the mynah
bird ( Acridotheris tristis [Linn.]), and the
Chinese dove ( Streptopelia chinensis [Sco-
poli] ) . On this account, these wild birds are

PACIFIC SCIENCE, Vol. 1, April, 1947

believed to assist in spreading the infection
in nature. The parasites are located in the
inner corner of the eyeball and in the nictat¬
ing membrane. In heavy infestations there
is a puffiness around the eye, and inflamma¬
tion frequently results in blindness. Infested
birds often wink their eyes continually, and
the irritation causes the bird to scratch the
eye with the claws for relief. The process of
scratching frequently causes mechanical in¬
jury to the eyeball and development of
secondary bacterial infection. Eyeworm in¬
fection is most prevalent in dry areas with
loose sandy soil in which roaches thrive. As
a means of controlling this disease, the writer
has advocated the maintenance of giant toads,
Bufo marinus (Linn.), in poultry yards.
These toads are insectivorous and devour
roaches readily.

The poultry ascarid, As car id ia gal It
(Schrank), and two species of cecal worms,
Heterakis gallihae (Gmelin) and Subulura
brumpti (Lopez Neyra), are common among
chickens. S. brumpti is the most prevalent,
and unlike H. gallinae requires an interme¬
diate host, which may be any one of the
following (Alicata, 1939^; Cuckler and Ali¬
cata, 1944): (coleoptera) Alphitobius
diaperinus (Panz.), Ammophorus insularis
Boh., Dermestes vulpinus Fabricius, Gono¬
cephalus seriatum (Boisd.), and Tribolium
castaneum (Herbst); (orthoptera) Cono¬
cephalus saltator (Sauss.), Oxya chinensis
(Thun.); (dermaptera) Euborellia annu¬
li pes (Lucas). The intestinal roundworm,
Ornithostrongylus quadriradiatus (Steven¬
son), has been found common in pigeons in
the Islands and is believed responsible for
unthriftiness and losses among pigeons (Ali¬
cata, 1939c).

Other roundworms of chickens which re¬
quire an intermediate host include the crop
worm, Gongylonema ingluvicula Ransom,
and the proventricular worms, Tetrameres
americana Cram and Dis pharynx spiralis
(Molin) . In continental United States G. in¬
gluvicula has been found to utilize the beetle

Parasites of Domestic Animals in Hawaii — Alicata

71

Copris minutus Drury as an intermediate
host. In Hawaii, the related beetle C. incer-
tus (Say) may be found to serve as a suitable
host. T. am eric ana is known to utilize any
of the following as intermediate hosts in the
Islands (Alicata, 1938c): (coleoptera)
Dendrophilus sp. (probably D. punctatus
Herbst), Dermestes vulpinus Fabricius, E pi-
tragus diremptus Karsch, and Gonocephalus
seriatum (Boisd.); (orthoptera) Blattella
germanica (Linn.); and Gonocephalus salta-
tor (Sauss.); (dermaptera) Euhorellia an-
nulipes (Lucas); (amphipoda) Orchestia
platensis Kroyer. The sow bug, (isopoda)
Porcellio laevis Latr., serves as intermediate
hosts for D. spiralis (Alicata, 1938c), which
often produces deep ulcerations of the pro-
ventricular wall.

FLUKES

The cecal fluke, Postharmostomum galli-
num (Witenberg), commonly infects adult
chickens raised on the ground. Extensive
cecal hemorrhages have been found asso¬
ciated with infection by this parasite. Recent
studies have shown that two common land
snails, Eulota similaris (Fer.) and Subulina
octona (Brug.), serve as intermediate hosts
(Alicata, 1940).

TAPEWORMS

Tapeworms are of common occurrence in
chickens. Those known in Hawaii include
the following: Choanotaenia injundibulum
(Bloch), Hymenolepis carioca (Magalhaes),
Hymenolepis exigua Yoshida, Raillietina ces-
ticillus (Molin), and R. tetragona (Molin)
(Alicata, 1938c).

Various arthropods in Hawaii are known
to serve as intermediate hosts for the above-
mentioned tapeworms, as follows (Alicata,
1938c; Hall, 1929) : C. injundibulum:
(coleoptera) Dermestes vulpinus Fabri¬
cius, Epitragus diremptus Karsch, Gono¬
cephalus seriatum (Boisd.), and (diptera)
Musca domestica Linn.; H. exigua: (amphi¬

poda) Orchestia platensis Kroyer; R. cesti-
cillus: (coleoptera) Dermestes vulpinus
and Gonocephalus seriatum; R. tetragona:
probably various species of ants, especially
those of the genera Pheidole and Tetra-
morium. Members of this group of ants
( P . vinelandica and T. caespitum) are known
intermediate hosts of R. tetragona in conti¬
nental United States (Jones and Horsefall,
1935).

ARTHROPODS

Various species of lice are known to in¬
fest poultry in the Islands. These include
the following (Illingworth, 1928^): chick¬
en body louse, Eomenacanthus stramineus
(Nitzsch); chicken head louse, Lipeurus
heterographus Nitzsch; common hen louse,
Menopon gallinae (Linn.), also found on
turkeys and guinea hens; fluff louse of
chickens and turkeys, Goniocotes hologaster
Nitzsch; large chicken louse, Goniocotes
gigas Taschenberg; large turkey louse, Goni-
odes stylijer Nitzsch; peafowl and guinea hen
louse, Menopon phaeostomum (Nitzsch);
turkey louse, Lipeurus gallipavonis Geoffroy;
and the chicken wing louse, Lipeurus caponis
(Linn.) .

Mites found on chickens include the red
mite, Dermanyssus gallinae (De Geer); the
wing mite, Pterolichus obtusus Robin; and
the body mite, Megninia cubitalis (Megnis)
(Alicata et al., 1946). Included likewise is
the tropical fowl mite, Lyponyssus bursa
(Berlese); this mite has also been reported
common in nests of English sparrows and
mynah birds. It is known to invade houses,
where it bites human beings and causes skin
irritation (Zimmerman, 1944).

Other arthropods of poultry include
the sticktight flea, Echidnophaga gallinacea
(Westwood) (Illingworth, 1916); the
pigeon fly, Pseudolynchia canariensis (Mac-
quart) (= Lynch ia maura Bigot), generally
widespread among pigeons (Bryan, 1934);
and the biting louse of pigeons, Columbicola
columbae (Linn.) (Zimmerman, 1944).

72

PACIFIC SCIENCE, Vol. 1, April, 1947

Parasites of Cattle
protozoa

Four species of coccidia, Eimeria cylin-
drica Wilson, E. bovis Zublin, E. zurnii
Rivolta, and E. bukidnonensis Tubangui,
have been recovered from the feces of young
calves (Cuckler and Alicata, 1943). Al¬
though no severe cases of bovine coccidiosis
have been recorded in the Islands, reports
elsewhere indicate that infection may pro¬
duce bloody diarrhea and emaciation.

ROUNDWORMS

In a recent survey involving the examina¬
tion of about 375 cattle raised on various
islands and slaughtered in Honolulu, the
following percentages of roundworm infec¬
tions were found (Cuckler and Alicata,
1943): gullet worms, Gongylonema pul-
chrum Molin, 54.3 per cent; stomach worms,
Haemonchus contortus (Rudolphi), 0.9 per
cent; intestinal roundworms, Bunostomum
phlebotomum (Railliet), 6.7 percent; Coop -
eria punctata (v. Linstow), 4.0 per cent;
C. pectinata Ransom, 0.3 per cent; and the
skin filarid, Stephanofilaria stilesi Chitwood,
89. 8 per cent. Eggs of Trichuris ovis ( Abild-
gaard) and Strongyloides sp. (probably
S. papillosus) have at times been found in
the feces of cattle in Hawaii.

The intestinal roundworm, Oesophago-
stomum radiatum (Rudolphi), and the lung-
worm, Dhtyocaulus viviparus (Bloch), have
also been noted by the writer. Lungworm
infection is believed to be of considerable
importance, especially among calves, in some
sections of the Islands, and deaths resulting
from this parasite have been recorded (Wil-
lers, 1945).

Of the above roundworms, Stephanofilaria
stilesi and Gongylonema pulchrum require
an intermediate host in their development.
The intermediate host for S. stilesi is un¬
known. G. pulchrum is known to utilize one
of various coprophagous beetles and roaches
as intermediate host in continental United

States (Alicata, 1935). Of insects reported
as hosts, Aphodius lividus (Oliv.), Dermes-
tes vulpinus Fabricius, and Blattella german-
ica (Linn.) occur in Hawaii.

FLUKES

Two species of flukes have been recorded
from cattle in the Islands. One species con¬
sists of an unidentified rumen fluke reported
by Hall (1936), and the other species is the
liver fluke, Fasciola gigantica Cobbold.

Liver-fluke infection is the most important
parasitic disease of beef and dairy cattle. In¬
fection with this parasite was first reported
by Dr. A. Lutz (1892) as being common on
four of the larger islands. Although at that
time the parasites were reported as Fasciola
hepatica Linn., more recent study has shown
them to be F. gigantica (Alicata and Swan¬
son, 1937). The importation of this fluke
into Hawaii is not clearly understood, but it
is believed to have come from the Orient
with the introduction of water buffaloes. It
is of interest to note that the limnaeid snail,
Fossaria ollula (Gould), which serves as the
intermediate host, has Japan and China as its
geographic range (Alicata, 1938^). This
snail is widely distributed in Hawaii and is
common in streams and swampy lowlands.

The maintenance of fluke infection in Ha¬
waii, as elsewhere, is dependent on various
factors of which topography, climatic condi¬
tions, and agricultural practices are very im¬
portant. The Hawaiian Islands represent the
summits of a 2,000-mile range of volcanic
mountains which vary from coastal to centric
or eccentric in position. The mountains de¬
scend to the ocean abruptly, in steep walls
or by gradual transition over relatively flat
land with very little drainage. These poorly
drained lowlands and valleys, especially on
the windward side, often present rather ex¬
tensive swamps. Rainfall is most prevalent
in winter months, but showers during other
seasons of the year are sufficient to maintain
swampy conditions. These wet areas and the
mild Hawaiian climate encourage snail prop-

Parasites of Domestic Animals in Hawaii — Alicata

73

agation the year round as well as the de¬
velopment and hatching of fluke and other
parasite eggs. Moreover, agricultural prac¬
tices in the Islands have encouraged rather
than hindered the maintenance of fluke in¬
fection. With ample supplies of vegetation,
cattle have been allowed to graze continu¬
ously. Many dairymen have long been in
the habit of feeding cut forage from wet or
swampy areas to cattle. These practices have
been largely responsible for the widespread
fluke infection. This disease is gradually be¬
ing brought under control largely through

(1) use of copper sulfate for the control
of the snail vector in swamps or streams,

(2) use of forage grass cut from dry areas,
and (3) treatment of infected animals with
hexachloroethane. This synthetic compound,
although first used in fluke control in Europe
in 1926 (Thienel, 1926), was first utilized
on a large scale in the United States by the
University of Hawaii Agricultural Experi¬
ment Station (Alicata, 1941 <2).

TAPEWORMS

The infective larval stage "bladderworm”
of Taenia saginata Goeze has occasionally
been found present in the musculature of
cattle in the Islands, according to a personal
communication received from Dr. A. H.
Julien, Federal meat inspector. The larvae
reach maturity in the intestine of man, fol¬
lowing ingestion of improperly cooked beef.
Cattle acquire the infection as a result of
eating vegetation contaminated with human
feces containing eggs of this parasite. It is
generally believed that most cases of human
infection occur among immigrants from the
Orient, especially the Philippines.

ARTHROPODS

Several dipterous larvae are knov/n to be
parasitic on cattle. In a recent examination
of 303 animals (Cuckler and Alicata, 1943),
26.1 per cent showed evidence of the cattle
grub, Hypoderma lineata (De Villers), a fly

first reported in the Islands in 1906 (Bryan,
1934). This parasite is recognized as very
injurious to cattle, causing loss of flesh and
decreasing the value of the skin for leather.
Some years ago, Mr. O. A. Pickerill of the
Hawaii Meat Company in a personal com¬
munication reported that during the year
1934, of 15,099 hides examined, 4,252, or
28.16 per cent, were grubby.

In recent years a report was made of at¬
tacks or "fly strike” of blowflies on young
calves on the island of Kauai (Holdaway,
1943; 1945). Observations indicated that
three species of flies were involved, Chry-
somia megacephala (Fabricius), C. rufifacies
(Macquart) , and Lucilia sericata (Meigen) .3
These flies ordinarily breed in carcasses and
other animal matter. However, they may de¬
posit eggs in a number of different places on
recently born calves. The eggs hatch and the
larvae or maggots feed on the surface layer
and cause an inflamed, malodorous wound.
Infested calves become spiritless and, unless
suitably treated, die in a few days.

Auricular myiasis of cattle caused by the
larvae of C. megacephala, C. rufifacies, and
Fannia sp. have been reported by Zimmer¬
man (1944). Species of adult flies which
are pestiferous on cattle in the Islands in¬
clude the horn fly, Siphona irritans (Linn.)
(— Lyperosia irritans [Linn.]), and the
stable fly, Stomoxys calcitrans (Linn.) (Bry¬
an, 1934).

Lice, Haematopinus eurysternus (Nitzsch)
(Cuckler and Alicata, 1943) and Bovicola
hovis (Linn.) (Zimmerman, 1944), have
occasionally been found on cattle. General
emaciation or unthriftiness is usually asso¬
ciated with infestation.

The spinose ear tick, Otobius megnini
(Duges), which was first noted in recent
years (Alicata, 1941b; Cuckler and Alicata,
1943; Zimmerman, 1944), is widespread on
beef cattle. Of 357 cattle examined from

3 According to a personal communication from
F. G. Holdaway, the identification of Lucilia is
tentative.

74

PACIFIC SCIENCE, Vol. 1, April, 1947

Hawaii, Oahu, and Maui, 160, or 44.8 per
cent, showed infestation (Cuckler and Ali-
cata, 1943). In several instances the ticks
were found in large numbers filling the en¬
tire ear canal. These ectoparasites are known
to puncture the tender skin of the ear and
suck blood. The wounds thus caused often
ulcerate and a condition known as ear canker
results.

Parasites of Swine
protozoa

There are two types of protozoa infecting
swine in the Islands. They are frequently the
cause of dysentery, especially among young
animals. Included are the coccidia, Eimeria
debliecki Douwes, E. scabra Henry, and
E. s pinos a Henry, and the ciliate, Balanti¬
dium coli (Malmsten). Various forms of
unidentified amoebae and flagellates of un¬
known pathogenicity are also frequently
noted in the feces of swine.

ROUNDWORMS

In 1938 an examination of the feces of
103 grown pigs from the islands of Oahu
and Kauai (Alicata, 1939 b) revealed the
following incidence of parasite eggs: Ascaris
suum Goeze, 21 per cent; Oesophagostomum
dentatum (Rudolphi), 32 per cent; Strongy-
loides sp., 43 per cent; Trichuris suum
(Schrank), 7 per cent.

Adult roundworms which have been re¬
covered at necropsy from swine include
the following (Alicata, 1938 d)\ stomach
worms, As car ops strongylina (Rudolphi)
and Hyostrongylus rubidus (Hassall and
Stiles); kidney worms, Stephanurus denta-
tus Diesing; lungworms, Choerostrongylus
pudendotectus Vostokov and Metastrongylus
elongatus (Dujardin). Larvae of Trichinella
spiralis (Owen) have also been found en¬
cysted in the musculature of a domestic pig.

Kidney worms and lungworms are most
frequently found among hogs raised in open

hog lots. According to a personal commu¬
nication received from Dr. R. N. Beddow,
Veterinarian, Territorial Board of Health, of
25,234 hogs slaughtered in Honolulu dur¬
ing 1945 and 1946, 2.8 per cent showed
adult kidney worms in the kidney fat. This
undoubtedly represents a partial incidence of
infection in swine, since no observation was
apparently made on the presence of young
migrating worms, which are frequently
found in the liver and other parts of the
body.

Of the above roundworms, lungworms
are known to require earthworms as an in¬
termediate host. At least two species of un¬
identified earthworms recovered from hog
lots around Honolulu have been found by
the writer to harbor infective lungworm
larvae. It is reported that in Hawaii there
are about a dozen species of earthworms of
the genus Pheretima (Williams, 1931) . The
stomach worm, A. strongylina, utilizes one
of various coprophagous beetles as interme¬
diate host in continental United States (Ali¬
cata, 1935) ; in Hawaii, beetles of the genus
Aphodius possibly serve in this capacity.

Because of the occurrence of the first
laboratory-proved case of human trichinosis
in Hawaii in 1936 (Alicata, 1938^), the
writer, under the auspices of the Territorial
Board of Health, conducted a survey to de¬
termine the source and prevalence of trichina
infection in nature. This survey revealed the
following information: of 61 domestic and
41 wild hogs examined from the island of
Hawaii, 1 and 6, respectively, were found
infected; of 2,130 rats and 70 mongooses
examined, 57 and 17, respectively, showed
infection. . No trichinae were found in 92,
130, and 30 domestic hogs examined from
the islands of Maui, Oahu, and Kauai, re¬
spectively. Of 1,904 rats and 22 mongooses
examined on Maui, 1 and 2, respectively,
were found infected. Of 352 and 601 rats
examined from Oahu and Kauai, respec¬
tively, none showed infection.

It is of interest to point out that from

Parasites of Domestic Animals in Hawaii — Alicata

75

1936 to 1945, 58 cases of human trichinosis
have been reported in the Islands by the
Territorial Board of Health. Most of the in¬
fected persons had eaten, or were suspected
of having eaten, improperly cooked wild
pork or products made from wild pork ( Ali¬
cata, 1938*?). According to records of the
Territorial Board of Agriculture and Fores¬
try, during the 8-year period from 1933
through 1940 inclusive (Tinker, 1941),
32,724 wild hogs, or an average of 4,090 a
year, were killed on five of the larger islands.
Because of the moderately high incidence of
trichinosis in wild hogs, meat from these
animals is believed to constitute a health
menace unless proper precautions in cook¬
ing, preserving, and refrigerating are taken.
Mention may also be made that of 133
human diaphragms examined at random at
autopsy in Honolulu, 7.4 per cent harbored
trichinae larvae (Alicata, 1942).

FLUKES

Mo flukes are known to be present in
domestic hogs in Hawaii. However, the liver
fluke of cattle, presumably Fasciola gigan-
tica, has been reported from wild pigs (Ship-
ley, 1913). Wild hogs are descended from
domestic forms which have escaped and now
roam wild in the mountains, swamps, and
waste lands of the Islands.

TAPEWORMS

No adult tapeworms are found in swine,
but the infective larval stage "bladder worm"
of Taenia hydatigena Pallas has been found
attached to the liver and omentum of swine
(Alicata, 1938 d). These larvae are known
to reach maturity irf the intestinal tract of
dogs.

ARTHROPODS

The hog mange mite, Sarcoptes scabiei
suis Linn., is prevalent on swine in the
Islands. The louse, Haematopinus adventi-
cius (Neum.) (= H. suis [Linn.]), is also

present (Illingworth, 192 8b). Infestation
with these ectoparasites is very often asso¬
ciated with malnutrition and unhygienic sur¬
roundings.

Parasites of Horses
protozoa

No reports are available on protozoan
parasites of horses in the Islands.

ROUNDWORMS

According to a recent survey (Foster and
Alicata, 1939), horses in Hawaii harbor at
least 25 species of roundworms, as follows:
Strongylus equinus Mueller, S. edentatus
(Looss), S. vulgaris (Looss), Triodon-
tophorus serratus (Looss), T. brevicauda
(Boulenger), Gyalocephalus capitatus Looss,
Poteriostomum imparidentatum Quiel, Cya-
thostomum coronatum (Looss), Cylicocercus
catinatus (Looss), C. goldi (Boulenger),
C. pater at us (Yorke and Macfie), Cylicoste-
phanus calicatus (Looss), C. longibursatus
(Yorke and Macfie), C. minutus (Yorke
and Macfie), C. asymetricus (G. Theiler),
Cylicocyclus nassatus (Looss), C. lepto-
stomus ( Kotlan ) , Cylicodonto phorus bicoro-
natus (Looss), C. euproctus (Boulenger),
T rich 0 strongylus axei (Cobbold), Parascaris
e quorum (Goeze), Oxyuris equi (Schrank),
Probstmayria vivipara (Probstmayr), Ha-
bronema muscae (Carter), and H. micro¬
stoma (Schneider). Of the above parasites,
S. vulgaris has been found to be somewhat
common, a fact suggesting that verminous
arthritis and aneurism, caused by the larval
stage of this roundworm, are not infrequent
among horses in the Islands.

The roundworms of the genus Habronema
listed above are known to utilize elsewhere
various species of flies as intermediate hosts.
(Hall, 1929). In Hawaii, the house fly,
Musca domestica Linn., may transmit H. mus¬
cae and H. microstoma, and the stable fly,
Stomoxys calcitrans Geoffroy, may transmit
IF. microstoma.

76

PACIFIC SCIENCE, Vol. 1, April, 1947

FLUKES

According to a report by Hall (1936),
liver flukes collected in 1894 from a horse in
Honolulu were sent to the U. S. Bureau of
Animal Industry. These flukes were orig¬
inally diagnosed as Fasciola hepatica Linn.,
but a recent re-examination by Mr. A. Mc¬
Intosh of that Bureau revealed that they are
F. gigantic a (Cobbold). Moreover, veteri¬
narians on the island of Kauai have verbally
reported to the writer the finding of fas-
ciolid flukes in livers of horses. Thus far the
writer has not confirmed these observations.
In recent years the examination of the livers
of five horses pastured in known fluke-
infested areas failed to reveal liver fluke in¬
fection. In addition, a horse and a mule fed
experimentally 650 and 2,300 infective liver
fluke cysts, respectively, failed to show evi¬
dence of flukes or fluke lesions when autop-
sied a few months later (Alicata and Swan¬
son, 1938). It appears that equines only
rarely become infected with liver flukes.

TAPEWORMS

Two species of tapeworms, Anoplocephala
perjoliata (Goeze) (Foster and Alicata,
1939) and A. magna (Abildgaard) (Swan¬
son, 1939), have been reported from horses
in the Islands. The intermediate host for
each of these parasites is unknown.

ARTHROPODS

The larvae of the 'Tot flies,” Gastro-
philus intestinalis (De Geer) and G. nasalis
(Linn.) (Foster and Alicata, 1939), are
commonly found attached to the stomach
wall of horses in the Islands. Adult flies,
Stomoxys calcitrans (Linn.), are also pesti¬
ferous on horses.

Parasites of Sheep and Goats
protozoa

No reports are available on the protozoa
of sheep and goats in the Islands.

ROUNDWORMS

A recent examination (Cuckler, 1943) of
a group of sheep from the island of Kahoo-
lawe revealed the following incidence of
roundworms: stomach worms, Haemonchus
contortus (Rudolphi), in 6 of 15 examined,
and Trichostrongylus instabilis Railliet in 3
of 10 examined; intestinal worms, Cooperia
punctata (V. Linstow), in 3 of 10 examined,
and Nematodirus spathiger (Railliet) in 1
of 10 examined.

FLUKES

Specimens of liver flukes collected from
sheep in Honolulu were submitted to the
U. S. Bureau of Animal Industry in 1892
(Hall, 1936). These specimens, which were
originally identified as Fasciola hepatica
Linn., are doubtlessly F. gigantica, since the
former is not known to have occurred in the
Islands.

tapeworms

Unidentified larval tapeworms or "bladder
worms,” probably those of Taenia hydati-
gena Pallas, attached to the liver and peri¬
toneum of sheep have been noted by Dr. A.
H. Julien, Federal meat inspector (personal
communication). The larvae of T. hydati-
gena are known to reach maturity in the
intestinal tract of dogs.

ARTHROPODS

In the examination of 60 sheep from the
island of Kahoolawe (Cuckler, 1943), 43
harbored the spinose eartick, Otobius meg-
nini (Duges). Reports also indicate the
occurrence on sheep of the "sheep tick,”
Melophagus ovinus (Linn.) (Bryan, 1933;
Muir, 1928); the "head maggot,” Oestrus
ovis Linn. (Bryan, 1933) ; and the "Oriental
blowfly,” Chrysomyia megacephala (Fabri-
cius) (Bryan, 1934). The sucking louse,
Linognathus africanus Kellog and Paine, and
the biting louse, Bovicola caprae (Gurlt),

Parasites of Domestic Animals in Hawaii — Alicata

77

have been reported from goats (Zimmerman,
1944).

Parasites of Dogs
protozoa

Canine coccidiosis (species unknown) is
known to be present in dogs in Hawaii.

ROUNDWORMS

The roundworms known from dogs in
Hawaii include the following: intestinal
roundworms, Toxocara cams (Werner),
Toxascaris leonina (V. Linstow), Ancylos-
toma caninum Ercolani, Trie hurls vulpis
(Frohlich), and the heartworm, Dir o filar la
immltls (Leidy). Heartworms are believed
to be common in the Islands. Of the three
species of mosquitoes in Hawaii, Culex quin-
quefasclatus Say, Aedes aegypti (Linn.), and
A. albopictus (Skuse), the first two have
already been incriminated as intermediate
hosts for heartworms (Hall, 1929). In a
check list of parasites of dogs and cats, Dik-
mans (1945) lists the lungworm Filariodes
osleri (Cobbold) from Hawaii. The life
cycle of this parasite is unknown.

TAPEWORMS

Dipylidium caninum (Linn.) is the only
tapeworm noted in dogs in Hawaii. This
tapeworm is known to utilize fleas and lice as
intermediate hosts (Hall, 1929). Ctenoce-
p halides felis '( Bouche ) ( Pemberton, 1926)
and Trichodectes latus Nitzsch (Swezey,
1931 ) , which could serve as hosts, are found
on dogs in Hawaii. Infective larvae "bladder
worms” of Taenia hydatigena Pallas have
been found attached to the liver and omen¬
tum of swine (and sheep ?) in the Islands;
from this finding it may be inferred that the
adult stage of this parasite is found in dogs.

ARTHROPODS

Arthropods present on dogs in Hawaii
include the following: fleas, Ctenocepha-

lides felis (Bouche) (Pemberton, 1926)
and Echidnophaga gallinacea (Westwood);
lice, Trichodectes latus Nitzsch (Swezey,
1931); a species of kangaroo lice, Hetero-
doxus longitarsus Piaget, collected from a
dog in Honolulu (Pemberton, 1934); ticks,
Rhipicephalus sanguineus (Latreille) (Van
Zwaluwenburg, 1934); and undetermined
species of mange mites.

Parasites of Cats
protozoa

No reports are available on the protozoan
parasites of cats in the Islands.

ROUNDWORMS AND TAPEWORMS

Little information is available on round-
worms and tapeworms of cats in the Islands.
The following were collected by the writer
from a stray cat in Honolulu: (round-
worms ) stomach worms, Physalo ptera
praeputialis Von Linstow; hookworms, Am
cylo stoma caninum Ercolani; lungworms, Ae-
lurostrongylus abstrusus (Railliet); (tape¬
worms) Taenia taeniae for mis (Batsch) and
Dipylidium caninum (Linn.). Immature
acanthocephalids, determined by Dr. H. J.
Van Cleave as Arhythmorhynchus sp., have
been collected by the writer from the small
intestine of a cat. Dr. Van Cleave believes
that the cat is not the natural host. Acantho¬
cephalids of this genus are predominantly
parasites of water birds.

Among the above parasites, the life cycle
of the stomach worm is unknown. The lung-
worms are known to require snails or slugs
as intermediate hosts (Hobmaier and Hob-
maier, 1935). The land snail Subulina
octona (Brug.) was reported by Van Volken-
berg (1937) as serving as intermediate host
in Puerto Rico; in Hawaii the writer has
found that the land snails S. octona and
Eulota similaris (Fer.) may serve for that
purpose. Cats may also acquire lungworms
from eating infected mice, the latter acquir-

78

in g the parasite as a result of eating infected
snails. In mice, the larvae of lungworms
migrate to the musculature, where they be¬
come encysted (Van Volkenberg, 1937).
The tapeworm, T. taeniaeformis, is known
to utilize rats or mice as intermediate hosts;
the infective larval forms, "bladder worms,"
are commonly found in the liver of these
rodents in Hawaii. The tapeworm, D. cani¬
num, utilizes fleas or lice as an intermediate
host ( see parasites of dogs ) .

ARTHROPODS

The cat flea, Ctenoce p halides felis
(Bouche), and the sticktight flea, Echidno-
phaga gall mace a (Westwood), are common
on cats in Hawaii; the former is also com¬
monly found under houses frequented by
cats. The biting louse, F el i cola suhro strata
(Nitzsch), has been collected from cats
(Zimmerman, 1944).

PACIFIC SCIENCE, Vol. 1, April, 1947

Parasites of Rabbits

Very little is known about parasites of
rabbits in the Islands. Liver coccidia, Eimeria
stiedae Lindemann, have been noted by the
writer on several occasions. The scab mite,
Psoroptes communis Furstenberg (Pember¬
ton, 1946), which is commonly found in
the ears, has been reported from rabbits.

ACKNOWLEDGMENTS

The writer wishes to acknowledge the
assistance of Drs. E. W. Baker, H. S. Barber,
E. A. Chapin, H. E. Ewing, W. S. Fisher,
J. O. Maloney, C. R. Shoemaker, E. W. Staf¬
ford, and F. X. Williams, who from time to
time have assisted in identifying various local
arthropods reported in this paper. Acknowl¬
edgment is also made to Dr. F. G. Holdaway
and Mr. C. E. Pemberton for helpful sugges¬
tions made in connection with some of the
entomological aspects of this paper.

Summary of Host List of Parasites and Intermediate Hosts Recorded in Hawaii

a

NAME OF PARASITE LOCATION IN HOST INTERMEDIATE HOST*

Roundworms:

cat ( Felis domestica)

Aelurostrongylus abstrusus

Lungs

Gastropoda: Subulina octona? Eulota
similaris 2

Rodentia: Mus mus cuius 3

Ancylostoma caninum

Small intestine

Physaloptera praeputialis

Stomach

(Unknown)

Arhythmorhynchus sp.

Small intestine

(Unknown)

Tapeworms:

Dipylidium caninum

Small intestine

Siphonaptera and Anoplura (see parasites
of dog)

Taenia taeniaeformis

Small intestine

Rodentia: Mus mus cuius? Rattus rattus
alexandrinus,1 Rattus rattus norvegicus ?
Rattus rattus rattus 1

Arthropods:

Ctenocephalides felis

External

Echidnophaga gallinacea

Attached to skin

Felicola substrata

External

CATTLE (Bos taurus)

Protozoa:

Eimeria bo vis

Small intestine

Eimeria bukidnonensis

Small intestine

* Legend: (1)= infection found in nature; (2) = determined experimentally; (3) = reported elsewhere
for animals similar to those occurring in Hawaii.

Parasites of Domestic Animals in Hawaii — Alicata

79

Eimeria cylindrica
Eimeria zurnii
Roundworms:

Bunostomum phlebotomum
Cooper id pectin at a
Cooperia punctata
Dictyocaulus viviparus
Gongylonema pulchrum

Small intestine
Small intestine

Small intestine
Small intestine
Small intestine
Lungs
Esophagus

Haemonchus contortus

Oesophagostomum radiatum
Stephanofilaria stile si
Strongyloides sp. ( papillosus ?)
Trichuris ovis
Flukes:

Fasciola gigantica
(Rumen fluke)

Tapeworms:

Taenia saginata (cysticercus)

Fourth stomach
or abomasum
Cecum and colon
Skin

Small intestine
Cecum

Liver

Rumen

Muscles

Arthropods:

Bovicola bovis
Chrysomyia megacephala
(larva)

Chrysomyia rufijacies (larva)

Fannia sp. (larva)
Haematopinus eurysternus
Hypoderma lineatum
Lucilia sericata ? (see text)
(larvae)

Otobius megnini

External
In wounds and
external
In wounds and
external
External
External
Under skin
External

Inside ears

Coleoptera: Aphodius lividus? Dermestes
vulpinus2

Orthoptera: Blattella germanica2

(Unknown)

Gastropoda: Foss aria ollula1
(Unknown)

Artiodactyla: Bos taurus 1 (cattle are inter¬
mediate hosts; man is the final host)

Protozoa:

Eimeria tenella
Roundworms:

Ascaridia galli
Cheilospirura hamulosa

Dispharynx spiralis
Gongylonema ingluvicula
Heterakis gallinae
Oxyspirura mansoni
Subulura brumpti

chicken ( Gallus gallus )

Ceca

Small intestine
Gizzard

Proventriculus

Crop

Ceca

Conjunctival sac
Ceca

Coleoptera: Carpophilus dimidiatus ,2
Dactylosternum abdominale ,2 Dermestes
vulpinus,2 Epi tragus diremptus / Euxestus
sp.,2 Gonocephalus seriatum ,* Litargus
balteatus,2 Oxydema fusiforme 2 Palorus
ratzeburgi,2 Sitophilus oryzae2 Tene-
broides nana? Tribolium castaneum2
Typhaea stercorea 2

Orthoptera: Atractomorpha ambigua2 Co-
nocephalus saltator2 Oxya chinensis2
Amphipoda: Orchestia platensis1
Isopoda: Porcellio laevis 1
(Unknown; probably coprophagous beetles)

Orthoptera: Pycnoscelus surinamensis 1
Coleoptera: Alphitobius diaperinus* Am-
mophirus insular is? Dermestes vulpinus,1

* Legend: (x) = infection found in nature; (2) = determined experimentally; (3)= reported elsewhere
for animals similar to those occurring in Hawaii.

80

PACIFIC SCIENCE, Vol. 1, April, 1947

Tetrameres am eric ana

Flukes:

Proventriculus

Postharmostomum gallinum

Ceca

Tapeworms:

Choanotaenia infundibulum

Small intestine

Hymenolepis exigua

Small intestine

Raillietina cesticillus

Small intestine

Raillietina tetragon a

Small intestine

Arthropods:

Dermanyssus gallinae

External

Echidnophaga gallinae e a

External

Eomenacanthus stramineus

External

Goniocotes gigas

External

Goniocotes hologaster

External

Goniodes stylifer

External

Lipeurus caponis

External

Lipeurus heterographus

External

Lyponyssus bursa

External

Megninia cubitalis

External

Menopon gallinae

External

Pterolichus obtusus

External

dog ( Canis ;

Protozoa:

(Coccidia of undetermined

species)

Intestine

Roundworms:

Ancylostoma caninum

Small intestine

Dirofilaria immitis

Heart and pulrr

nary artery

Eilariodes osleri (see text)

Lungs

Toxascaris leonina

Small intestine

T oxo car a canis

Small intestine

Trichuris vulpis

Cecum

Tapeworms:

Dipylidium caninum

Small intestine

Taenia hydatigena ?

Small intestine

Arthropods:

Ctenocephalides felis

External

Gonocephalus seriatum ? Tribolium
castaneum 2

Orthoptera: Conocephalus saltator ? Oxya
chinensis2

Dermaptera: Euborellia annulipes 1
Coleoptera: Dendrophilus sp.,1 Dermestes
vulpinus ? Epitragus dir empties? Gono¬
cephalus seriatum?

Orthoptera: Blattella germanica ? Cono¬
cephalus saltator2
Dermaptera: Euborellia annulipes1
Amphipoda: Orchestia platensisx

Gastropoda: Eulota similaris? Subulina
octona?

Coleoptera: Dermestes vulpinus? Epitragus
diremptus? Gonocephalus seriatum 1
Diptera: Muse a dome Stic a2
Amphipoda: Orchestia platensis 1
Coleoptera: Dermestes vulpinus? Gono¬
cephalus seriatum?

(Probably ants of the genera Pheidole and
Tetramorium )3

Diptera: Aedes aegypti? Culex quinquefas-
ciatus 3
(Unknown)

Siphonaptera: Ctenocephalides felis? Pulex
irritans 3

Anoplura: Trichodectes latuss
Artiodactyla (see parasites of swine)

* Legend: C) = infection found in nature; (2) = determined experimentally; (3) — reported elsewhere
for animals similar to those occurring in Hawaii.

Parasites of Domestic Animals in Hawaii — Alicata

81

Echidnophaga gallinacea External

Heterodoxus longitarsus External

Rhipicephalus sanguineus External

Trichodectes latus External

(Mites of undetermined species) External

goat ( Capra hire us )

Arthropods:

Bovicola caprae External

Linognathus africanus External

guinea fowl ( Numida meleagris )

Arthropods:

Menopon gallinae External

Menopon phaeostomum External

Roundworms:

Cyathostomum coronatum
Cylicocercus catinatus
Cylicocercus goldi
Cylicocercus pateratus
Cylicocyclus leptostomus
Cylicocyclus nassatus
Cylicodontophorus bicoronatus
Cylicodontophorus euproctus
Cylicostephanus calicatus
Cylicostephanus longibursatus
Cylicostephanus minutus
Cylicosternus asymetricus
Gyalocephalus capitatus
Habronema microstoma

Habronema muscae
Oxyuris equi
Parascaris equorum
Poteriostomum imparidentatum
Probstmayria vivipara
Strongylus edentatus
Strongylus equinus
Strongylus vulgaris
Trichostrongylus axei
Triodontophorus brevicauda
Triodontophorus serratus
Flukes:

Fasciola gigantica
Tapeworms:

Anoplocephala magna
Anoplocephala perjoliata
Arthropods:

Gastrophilus intestinalis
(larvae)

Gastrophilus nasalis (larvae)

HORSE (Equus

caballus )

Large intestine
Large intestine

«

Large intestine
Large intestine
Large intestine
Large intestine
Large intestine
Large intestine

Large intestine
Large intestine
Large intestine
Large intestine
Large intestine
Stomach

Diptera: Musca domesticap Stomoxys

Stomach

calcitransz

Diptera: Musca domestical

Colon

Small intestine
Large intestine
Colon

Large intestine
Large intestine
Large intestine
Stomach

Large intestine
Large intestine

Liver

Gastropoda: Fossaria ollula 1

Small intestine

(Unknown)

Cecum

(Unknown)

Stomach

Stomach

* Legend: (3)= infection found in nature; (2) = determined experimentally; (3)= reported elsewhere
for animals similar to those occurring in Hawaii.

82

PACIFIC SCIENCE, Vol. 1, April, 1947

Arthropods:

Menopon phaeostomum

peafowl (Pavo cristatus)
External

Protozoa:

Haemoproteus columbae
Roundworms:

Ornithostrongylus

quadriradiatus

Arthropods:

Columbicola columbae
Pseudolynchia canariensis

pigeon ( Columba livia domestica )

Blood Diptera: Pseudolynchia canariensis 1

Small intestine

External

External

rabbit ( Oryctolagus canicular is)

Protozoa:

Eimeria stiedae Liver

Arthropods :

Psoroptes communis External

Roundworms:

Cooperia punctata
Haemonchus contortus
Nematodirus spathiger
Flukes:

Fasciola sp. ( gigantica ?)
Tapeworms:

Taenia hydatigena ?
(cysticercus)

Arthropods:

Chrysomyia megacephala
(larvae)

Melophagus ovinus
Oestrus ovis (larvae)

Otobius megnini

sheep (Ovis aries )

Small intestine

Fourth stomach

Small intestine

Liver Gastropoda: Fossaria ollulcd

Attached to liver, Artiodactyla (see parasites of swine)
mesentery, and
omentum

In wounds and
external

External

Nasal cavities and
sinuses of head

Inside ears

Protozoa:

Balantidium coli
Eimeria debliecki
Eimeria scabra
Eimeria s pinos a
Roundworms:
Ascaris suum

swine (Sus scrofa domestica)

Large intestine
Large intestine
Large intestine
Large intestine

Small intestine

* Legend: 0) = infection found in nature; (2) = determined experimentally; (8)= reported elsewhere
for animals similar to those occurring in Hawaii.

Parasites of Domestic Animals in Hawaii — Alicata

83

As car ops strongylina Stomach

Choerostrongylus pudendotectus Lungs

Hyostrongylus rubidus
Metastrongylus elongatus

Stephanurus dentatus

Strongyloides sp.

Trichin ell a spiralis

Trichuris suum
Flukes :

Fasciola sp. ( gigantica ?)
Tapeworms:

Taenia hydatigena
(cysticercus)

Arthropods:

Haematopinus adventicius
Sar copies scabiei suis

Stomach

Lungs

Adults in kidneys
and kidney fat;
immature forms
in liver and
other internal
organs

Adults in small
intestine, larvae
in muscles

Cecum

Liver

Attached to liver,
mesentary, and
omentum

External

External

'(Unknown; probably coprophagous beetles
of genus Aphodius )

Macrodrili (earthworms, probably of the
genus Pheretima )

Macrodrili (earthworms, probably of the
genus Pheretima')

Gastropoda: Foss aria olluld1

Artiodactyla: Sus scrofa domesticap Ovis
aries;1 Canis familiaris3 is the final host

Protozoa:

Histomonas meleagridis
Roundworms:

Cheilospirura hamulosa
Arthropods:

Goniocotes hologaster
Goniodes stylffer
Lip ear us gallipavonis
Menopon gallinae

turkey ( Mel e agr is gallopavo )

Intestine, liver

Gizzard (See parasites of the chicken)

External

External

External

External

* Legend: (x) = infection found in nature; (2) = determined experimentally; (3) = reported elsewhere
for animals similar to those occurring in Hawaii.

REFERENCES

Alicata, J. E. Early developmental stages of
nematodes occurring in swine. U. S. Dept. Agr.
Tech. Bui. 489. 96 p. 1935.

• - Poultry parasites. Hawaii Agr. Expt. Sta.

Rpt. 1936: 79-82, 1936.

- Observations on the life history of Fasciola

gigantica, the common liver fluke of cattle in
Hawaii, and the intermediate host, Fossaria
ollula. Hawaii Agr. Expt. Sta. Bui. 80. 22 p.
Honolulu, 1938 {a).

- The life history of the gizzard worm

( Cheilospirura hamulosa) and its mode of
transmission to chickens, with special reference
to Hawaiian conditions. Livro Jubilar Prof.
Travassos, Rio de Janeiro, Brazil. 3: 11-19,
1938 (b).

- Studies on poultry parasites. Hawaii Agr.

Expt. Sta. Rpt. 1937: 93-96, 1938 (c).

- Studies on parasites of swine. Hawaii

Agr. Expt. Sta. Rpt. 1937: 96, 1938 (d).

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PACIFIC SCIENCE, Vol. 1, April, 1947

- A study of Trichinella spiralis in the Ha¬
waiian Islands. Pub. Health Rpt. 53: 384-393,
1938 0).

- Preliminary note on the life history of

Subulura brumpti, a common cecal nematode of
poultry in Hawaii. Jour. Parasitol. 25: 179-
180, 1939 (a).

- Parasites of swine. Hawaii Agr. Expt. St a.

Rpt. 1938 : 78-79, 1939 ( b ).

- Poultry parasites. Hawaii Agr. Expt. Sta.

Rpt. 1938: 79-82, 1939 0).

- The life cycle of Postharmostomum galli-

num, the cecal fluke of poultry. Jour. Parasitol.
26: 135-143, 1940.

- Studies on control of the liver fluke of

cattle in the Hawaiian Islands. Amer. Jour. Vet.
Res. 2: 152-164, 1941 ( a ).

- Spinose ear tick is found on cattle in the

Territory. Hawaii Farm and Home, Oct. 1941:
15, 30, 1941 ( b ).

- Preliminary report of studies on typhus,

leptospirosis and trichinosis in Honolulu. Rpt.
of Activity, Parasitol. Project, Pub. Health
Com., Chamber Com. Honolulu, 1941. 29 p.
Honolulu, 1942 (mimeographed) .

- , F. G. Holdaway, J. H. Quisenberry,

and D. D. Jensen. Observations on the com¬
parative efficacy of certain old and new insecti¬
cides in the control of lice and mites of chickens.
Poultry Sci. 15: 376-380, 1946.

- , and L. E. Swanson. Fasciola gigantica,

a liver fluke of cattle in Hawaii, and the snail,
Fossaria ollula, its important intermediate host.
Jour. Parasitol. 23: 106-107, 1937.

- , and L. E. Swanson. Experimental feed¬
ing of liver fluke cysts to a horse and a mule.
Hawaii Agr. Expt. Sta. Rpt. 1937: 89, 1938.

Bryan, E. H. A review of the Hawaiian diptera
and description of new species. Hawaii. Ent.
Soc. Proc. 8: 399-468, 1934.

Cuckler, A. C. Sheep parasites. Hawaii Agr.
Expt. Sta. Rpt. 1941-1942: 49, 1943.

- , and J. E. Alicata. Parasite survey.

Hawaii Agr. Expt. Sta. Rpt. 1941-1942: 46-48,
1943.

- , and J. E. Alicata. The life history of

Subulura brumpti, a cecal nematode of poultry
in Hawaii. Amer. Micros. Soc. Trans. 63: 345 —
347, 1944.

Dikmans, G. Check list of the internal and ex¬
ternal animal parasites of domestic animals in
North America. Amer. Jour. Vet. Res. 6: 211-
241, 1945.

Foster, A. O., and J. E. Alicata. Notes on para¬
sites of horses in Hawaii. Helminthol. Soc.
Wash. Proc. 6: 4-8, 1939.

Hall, M. C. Arthropods as intermediate hosts of
helminths. Smithsn. Inst. Misc. Collect. 81. 79
p. Washington, D. C., 1929.

- Problems of parasitism in Hawaii. Rev.

de Parasitol., Clin, y Lab. 2: 367-383, 1936.

Hobmaier, M., and A. Hobmaier. Mammalian
phase of the lungworm Aelurostrongylus ab-
strusus in the cat. Amer. Vet. Med. Assoc. Jour.
40: 191-198, 1935.

Holdaway, F. G. Blowfly attack on young calves.
Hawaii Agr. Expt. Sta. Rpt. 1941-1942: 127-
129, 1943.

- Blowfly strike of young calves in Hawaii.

Hawaii. Acad. Sci. Proc., 1940-1943: 17, 1945.
Illingworth, J. F. Notes on the hen flea (" Echid -
nophaga gallinacea” Westw.). Hawaii. Ent.
Soc. Proc. 3: 252-254, 1916.

- Lice affecting poultry in Hawaii. Hawaii.

Ent. Soc. Proc. 7: 41-42, 1928 ( a ).

- Insects collected in the pineapple growing

section on the Island of Lanai, August, 1927.
Hawaii. Ent. Soc. Proc. 7: 42-46, 1928 (b).

- Manson’s eye worm distributed by English

sparrow. Hawaii. Ent. Soc. Proc. 7: 461, 1931.
Jones, M. F., and M. W. Horsefall. Ants as in¬
termediate hosts for two species of Raillietina
parasitic in chickens. Jour. Parasitol. 21: 442-
443, 1935.

Lutz, A. Zur Lebensgeschichte des Distoma he-
paticum. Centbl. f. Bakt. u. Parasitol. 11: 783-
796, 1892.

Muir, F. Report. Hawaii. Ent. Soc. Proc. 7: 4,
1928.

Pemberton, C. E. Report. Hawaii. Ent. Soc.
Proc. 6: 221, 1926.

- Heterodoxus longibursatus Piaget. Haivaii.

Ent. Soc. Proc. 8: 394, 1934.

- Report. Hawaii. Ent. Soc. Proc. 12: 463,

1946.

Shipley, A. E. Entozoa. Fauna Hawaiiensis 2:
427-441, 1913.

Swanson, L. E. A note on the parasite fauna of
the Hawaiian Islands. Helminthol. Soc. Wash.
Proc. 6: 29-30, 1939.

Swezey, O. H. Report. Hawaii. Ent. Soc. Proc.
7: 361-362, 1931.

Thienel, M. Neue Arbeiten zur Medikamentosen
Bekampfung der Lebergelseuche. Munchen.
Tierdrztl. Wchnschr. 71: 771-772, 1926.
Tinker, S. W. Animals of Hawaii. 190 p. Tongg
Pub. Co., Honolulu, 1941.

Van Volkenberg, H. L. Lungworm in cats is
widespread. Puerto Rico Agr. Expt. Sta. Rpt.
1936: 76, 1937.

Van Zwaluwenburg, R. H. Report. Hawaii.

Ent. Soc. Proc. 8: 360-361, 1934.

Willers, E. H. Report of the Territorial Veteri¬
narian. Hawaii Bd. Commrs. Agr. and Forestry
[ Bien.H Rpt. 1944: 62-85, 1945.

Williams, F. X. The insects and other inverte¬
brates of Hawaii sugar cane fields. 400 p. Ad¬
vertiser Pub. Co., Honolulu, 1931.

Zimmerman, E. C. A case of bovine auricular
myiasis and some ectoparasites new to Hawaii.
Hawaii. Ent. Soc. Proc. 12: 199-200, 1944.

I

Fault at Waimea, Oahu

Harold S. Palmer1

INTRODUCTION

This paper is planned to show that a fault
cuts the shore line of the island of Oahu at
Waimea Bay, 7 miles southwest of the north
point of the island. It is thought that this
is the first time that a fault of appreciable
offset has been unequivocally detected in the
island of Oahu.

Stearns (1935: 82, 173, 414) makes little
mention of faults in his comprehensive re¬
port on the geology of Oahu. He thinks that
a certain cliff in the Waianae Range, buried
by later lava flows, may have been caused by
faulting, although he says that the cliff may
well have some other origin. He also de¬
scribes several minor faults.

In regions underlain by extensive, uniform
beds of rock, faulting is often manifested by
an offsetting of readily identifiable layers
along the surface of fracture. In many places
the offsets may be seen on valley sides or in
artificial excavations. This type of evidence
of faulting is not applicable in Hawaii be¬
cause the individual lava flows vary so much
in texture and in thickness that they are not
readily identifiable. Continuous tracing of
an individual flow is of little use because
most flows are rather narrow, and not ex¬
tensive.

In the uneroded, younger parts of the
Hawaiian Islands, bold cliffs have been made
by faulting. The block of rock on one side
of the fault has been dropped, so that a bold
cliff leads up from the edge of the dropped
block to the edge of the stationary or rela¬
tively raised block. The infacing cliffs at

1 Department of Geology, University of Hawaii.
Manuscript received October 22, 1946.

Kilauea and the cliffs that cut the shore line
of the island of Hawaii at South Point and
at Kealakekua Bay are almost certainly such
fault cliffs.

In the strongly weathered and eroded older
parts of the Hawaiian Islands, any fault
cliffs that may have existed formerly have
become much subdued and are no longer
conspicuous features of the landscape. The
existence of such faults can be demonstrated
only by less direct evidence.

In the present paper the hypothesis is
adopted that a fault extends inland from
Waimea Bay in a direction a little south of
east, and that the block on the north side has
been raised relative to the block on the south
side. Several consequences of this hypothesis
are deduced, and the consequences are found
to agree with features actually existing in this
part of Oahu.

I am greatly indebted to Dr. Chester K.
Wentworth and to Dr. A. Grove Day for
carefully reading the manuscript and giving
me valuable suggestions for its improvement.

OFFSETTING OF THE SHORE LINE

Where a coastal region has a seaward slop¬
ing surface, and is cut by a fault at a large
angle to the shore line, the block that is rela¬
tively raised will have its shore line shifted
seaward. This kind of offsetting is diagram-
matically illustrated in Figure 1, in which
the upper sketch represents a dome-shaped
island, such as the Hawaiian volcanoes build.
The lower sketch shows a similar dome cut
by a fault perpendicular to the shore line.
The block on the left is the up thrown side
and has its shore line displaced seaward from

85

86

PACIFIC SCIENCE, Vol. 1, April, 1947

the shore line on the right, or downthrown,
side. The accompanying maps of the Wai-
mea, Oahu, region (Fig. 2 and 5) show
that the shore line has a general trend from
northeast to southwest, but that the part
northeast of the bay, if continued southwest-
ward, would lie about two thirds of a mile
seaward of the southwestern part of the
shore line. A fault at Waimea would result
in this sort of offsetting of the shore line.

OFFSETTING OF RESTORED
CONTOUR LINES

The volcanic domes sketched in Figure 1
have topographic contour lines drawn on
them, and it will be noted that on the faulted

Fig. 1. Diagram to explain offsetting of a shore
line by a fault.

dome the lower contour lines are offset like
the shore line, but to a lesser distance. The
shore line, of course, may be thought of as
the contour line of zero altitude. If a fault
cut the Waimea Bay region, we should be
able to find traces of offsetting of some of
the contour lines.

Figure 2 was prepared by tracing as light
lines the 500-, 1,000-, 1,500-, and 2,000-foot
contour lines from a topographic map of
Oahu (U. S. Geol. Survey, 1938) as the first
step. The contour lines of the original map
were slightly smoothed and generalized in
tracing. Next, smoothly sweeping, curved,
heavy lines were drawn so as to be tangent

Fig. 2. Generalized contour map of the Wai¬
mea, Oahu, region. The light lines are the present-
day contour lines; the heavy, smoothly curving
lines are restored contour lines; the heavy, broken
line shows a reasonable course of the fault, with
spurs extending toward the downthrown side; and
the straight, heavy line is the line of the profile
of Figure 6.

to the seaward salients of the actual contour
lines, and thus to form "envelopes” around
the present-day contour lines. These smooth,
restored contour lines show approximately
the shape of this part of the Koolau volcanic
dome as it was before dissection by stream
erosion had roughened it. The restored con¬
tour lines at 500, 1,000, and 1,500 feet have
their northeastern parts offset seaward with
respect to their southwestern parts in the
same way that the shore line is offset, and,
presumably, for the same reason.

The precise course of the fault is obscured,
but the heavy, broken line of Figure 2 shows
a reasonable approximate position. The short
lines perpendicular to the several segments
indicate the downthrown side of the fault.

HEIGHTS OF WAVE-CUT CLIFFS

Volcanic domes are subject to erosion by
streams and by waves. If we neglect the
work of streams for the moment, and con¬
sider only the work of waves, we realize that
the sloping shores would become cliffed, as
is suggested in the upper sketch of Figure 3.
The sketch cannot show it, but the cliff would

Fault at Waimea, Oahu — Palmer

87

continue below sea level a short distance,
and a wave-cut bench would extend seaward
from the foot of the cliff. If after the cut¬
ting of the cliff the dome-shaped island were
faulted, the cliff along the raised part would
of course have its crest higher than the cliff
crest on the lowered part, as is suggested in
the lower sketch of Figure 3. If there were
in addition a lowering of sea level or a rais¬
ing of the whole island, the wave-cut bench
might be exposed and would be higher on
the upthrown block than on the downthrown
block.

Fig. 3. Diagram to explain how a fault may
change the height of sea cliffs.

At some time in the past, waves cut con¬
spicuous cliffs along this part of the shore
line of Oahu, just as waves have cut the
present striking cliffs along the Hamakua
Coast of the island of Hawaii. Later a lower¬
ing of sea level exposed the wave-cut bench,
on which reefs had been built in the mean¬
time by corals and associated organisms.
Streams carrying detritus from inland have
built alluvial slopes on the emerged plat¬
form. As one drives along Kamehameha
Highway, which is on the platform and in
general fairly close to the shore, one can see
that the wave-cut cliffs are decidedly higher
northeast of Waimea Bay than southwest of
the bay. This difference in heights of cliffs
agrees with the hypothesis of a fault at Wai¬
mea Bay.

CONCENTRATION OF PERENNIAL
DRAINAGE

Streams on a symmetrical volcanic dome
tend to take courses that radiate from the
central summit, somewhat like the spokes
of a wheel, as is suggested by Figure 4, a

Fig. 4. Diagrammatic map to explain the con¬
centration of drainage by a fault.

diagrammatic map of the stream courses on
half of a dome-shaped island. But if the
island is faulted, some of the stream courses
may be diverted, as is shown at the left in
Figure 4. The fault is shown by a straight
line with short perpendicular lines on the
downthrown side. The fault is seen to divert
the headwaters of four streams into one
course, which is thereby given an unusually
large volume of water and an increased
ability to erode. This appears to be what has
happened at Waimea Canyon, Kauai, where
the stream has cut that remarkably deep
canyon. The lowering of the stream bed,
partly by downfaulting and partly by its own
erosion, has brought it closer to the water
table, so that the stream is more generously
fed by underground water at all times, and
receives some ground water even in the drier
seasons.

At the right in Figure 4 another fault is
shown, but one about perpendicular to the
shore line. Since the course of this fault is
radial and essentially parallel to the radial

88

PACIFIC SCIENCE, Vol. 1, April, 1947

stream courses, it does not concentrate stream
flow to an appreciable extent. The situation
at Waimea, Oahu, would seem to be midway
between these two types, for our hypothesis
supposes a fault only moderately oblique to
the shore line.

When Figure 5 was drawn, all the stream

Fig. 5. Drainage map of the Waimea, Oahu,
region, showing the concentration of perennial
reaches of the streams near the fault.

courses of the region were traced off from
the topographic map. Some generalization
resulted during the process of tracing. The
intermittent reaches of the streams are shown
as broken lines and the perennial reaches are
shown as heavier, continuous lines. Except
in the southeast corner it would be hard to
find a point more than a third of a mile away
from some stream course, which amounts to
saying that the region has been rather in¬
tricately cut up into valleys and ridges.

The perennial reaches of the streams, how¬

ever, show a decided concentration inasmuch
as the three that are longest lie rather close to
one another and rather close to the position
of the suggested fault. The concentration of
perennial courses is another item that agrees
with the present hypothesis.

PROFILE ACROSS WAIMEA

The heavy, straight lines extending from
northeast to southwest in Figures 2 and 5
show the line along which the profile of
Figure 6 was constructed. The profile was
prepared from the preliminary sheets of the
most recent topographic map of Oahu (U. S.
Geol. Survey, n.d.), with a horizontal scale
of a little over 3 inches to the mile. The
vertical scale of Figure 6 is a little over four
times as large as the horizontal scale. It will
be seen that the high parts of the profile to
the left (northeast) of Elehaha Stream are
at altitudes around 800 feet, whereas those to
the right (southwest) of Kaiwikoele Stream
are about 200 feet lower. This difference
would result from the vertical offsetting of
the two blocks by the suggested fault.

TWO-CYCLE TOPOGRAPHY ON THE
UPTHROWN BLOCK

Where the earth’s crust is stable, a stream
will at first cut a rather narrow and deep
valley, but as time goes on the valley becomes
wider and more flaring because the rocks of
the valley sides weather, become weak, and
gradually slump or creep toward the stream.
Thus a wider valley, with more gently slop¬
ing sides, will be made.

Fig. 6. Topographic profile of the Waimea, Oahu, region, parallel to the shore line.

Fault at Waimea, Oahu — Palmer

89

If the region is later faised, or tilted, so
that the stream is given a steeper gradient
and a great velocity, the stream will revert
to a predominance of deepening over widen¬
ing, and will cut a new, inner, narrow valley
along the trough of the older, outer, broader
valley. This process results in what is known
as "two-cycle topography,” or "valley-in¬
valley topography,” characterized by the
break in the transverse profile where the
upper and lower valley sides meet. The upper
valley was cut during an earlier cycle of ero¬
sion and the lower valley during a later cycle.

The hypothesis that there is a fault at
Waimea supposes the block on the northeast
to have been raised. If this is true, the rais¬
ing should have rejuvenated the streams and
should have produced two-cycle topography.
Such topography is found between Pupukea
and Kaunala, in an area extending for about
3 miles parallel to the shore line, and lying
inland from the crest of the wave-cut cliff.
Unfortunately the contour intervals of the
maps do not permit constructing a profile
that will show the inner and outer valleys. In

Figure 7, the inner valley runs from right to
left. The photographer stood on a part of
the older, upper valley slopes which extend
to the middle distance. The pineapple fields
on the far side, and the pasture to the right
of the pineapple fields, are also on the
older, upper valley slopes. The sides of the
younger, inner valley are too steep to be
tilled, and are partly in brush, partly in grass,
and partly bare of vegetation.

THE FAULT SCARP

Where faulting has recently raised one
crust block with respect to the adjacent block,
a cliff or fault scarp separates the two blocks,
and at first is likely to be a conspicuous
feature of the landscape. As time goes on,
however, the scarp becomes weathered and
eroded back, and changes into a more Or less
subdued and inconspicuous land form. Such
is the scarp of the Waimea fault, now that
it is rather old. The best place known to the
writer from which to view this subdued
scarp is the bridge across the small Lauhulu

Fig. 7. View of two-cycle or valley-in-valley topography on the upthrown fault block.

90

PACIFIC SCIENCE, Vol. 1, April, 1947

Stream, 2.1 miles from Anahulu Bridge
at Haleiwa, along Kamehameha Highway,
toward Waimea Bay. Figure 8 was photo¬
graphed from this point. To the right is the
old, wave-cut cliff. The cliff leads up to
fields of sugar cane, which show up in a
rather light shade. The cane fields are on the
top of the downthrown block. Beyond the

Fig. 8. Distant view of the eroded and subdued
scarp of the Waimea, Oahu, fault, from Lauhulu
Bridge, looking northeastward.

cane fields (and, in the picture, above the
cane fields ) is the eroded and subdued scarp
of the upthrown block, marked by the darker
shade of the trees that grow on it. The pic¬
ture, of course, cannot show Waimea Valley,
but it lies between the cane fields and the
wooded slope. Valleys that are due solely to
erosion will normally have ridges of the same
height on both sides, but since the northeast
side of Waimea Valley rises some 200 feet
higher than the southwest side, the valley is
not solely the product of stream erosion.

DIRECTION OF MOVEMENT

A fault scarp results from the vertical
movement of the block of rock on one side
of the fault relative to the block on the other
side. If one block is actually stationary, the
other may move either up or down; or it may
be that both blocks move but that one moves
farther than the other; or the two blocks may
move in opposite directions.

At most of the faults in the Hawaiian
Islands the higher block seems to have re¬

mained relatively 'stationary while the lower
block moved downward, probably as a result
of the removal of support from below. The
infacing fault scarps at Kilauea, for example,
are thought to have resulted from subsidence
of the central block at times when the magma
column has receded. The last event of this
sort was in May, 1924, when the diameter
of Halemaumau, the inner pit, was increased
from about one third of a mile to about two
thirds of a mile, by the engulfment of the
central part, leaving a scarp of sheer cliffs
several hundred feet high. A month earlier,
subsidence along a fault at Kapoho near the
east point of the island of Hawaii made a
scarp 8 to 12 feet high, and the sea flooded
a part of the block that dropped downward.
Numerous other examples of presumed
downward movement of fault blocks in
Hawaii could be cited.

It is the writer’s opinion, however, that at
the Waimea, Oahu, fault the lower block re¬
mained relatively stationary while the higher
block was actually raised, because this as¬
sumption appears to be called for by the two-
cycle topography on the higher block.

Two-cycle topography results when a re¬
gion that is fairly well advanced in the ero¬
sion cycle has its streams rejuvenated by
having their gradients increased. This in¬
crease of gradient might be due to either
( 1 ) a lowering of sea level or ( 2 ) a raising
of the land area. If the rejuvenation in ques¬
tion were due to a lowering of sea level, then
similar rejuvenation and similar two-cycle
topography should be found in many of the
older parts of the Hawaiian Islands. In con¬
trast, if the rejuvenation were due to unique
uplift of this particular bit of Oahu, other
parts of the Hawaiian Islands would not be
rejuvenated and would not develop two-cycle
topography. Inasmuch as the writer knows
of no other places with similar two-cycle
topography, he concludes that uplift of the
higher, or northeastern, block took place.

The writer freely admits that he cannot
show the mechanism by which the northeast-

Fault at Waimea, Oahu — Palmer

91

ern block was raised. A tentative suggestion
is that a considerable body of molten matter
was intruded under this block, perhaps at the
time of the eruptions at Diamond Head and
at numerous other young vents in the south¬
eastern part of the Koolau Range.

CONCLUSION

It is not to be expected that one can de¬
tect faults in Hawaii by the offsetting of lava
flows, because the individual flows are of
small lateral extent and because there is great
variation in texture and thickness from place
to place within each lava flow.

It is believed that faulting occurred at
Waimea, Oahu, because this region shows a
number of features that would have resulted
from faulting, and for which no other ex¬
planation comes to mind. These features
include :

1 . Offsetting of the shore line.

2 . Offsetting of the restored contour lines.

3. Higher wave-cut cliffs on the up-
thrown side.

4. Concentration of perennial stream
courses near the fault.

5. A step in the profile across the fault.

6. Two-cycle topography on the up-
thrown block.

7. A subdued scarp such as prolonged
erosion would make of a fault cliff.

The cumulative evidence of these features
seems to indicate faulting, although actual
offsetting of the rock layers has not been
found.

REFERENCES

Stearns, Harold T., and Knute N. Vaksvik.
Geology and ground water resources of the
island of Oahu, Hawaii. 479 p., 34 fig., 33 pi.
Ter. of Hawaii, Div. Hydrog., Bui. 1, 1935.

U. S. Geol. Survey. Advance sheets of Haleiwa,
Kaipapau and Laie quadrangles of the topo¬
graphic map of Oahu. Surveyed in 1929 and
1930. Scale 1:20,000; contour intervals 10 and
50 feet, [n.d.]

U. S. Geol. Survey. Topographic map of the
island of Oahu, City and County of Honolulu,
Hawaii. Scale 1:62,500; contour interval 100
feet, 50-foot contour added. 1938.

Fungi of the Marshall Islands, Central Pacific Ocean

Donald P. Rogers1

INTRODUCTION

During the late summer of 1946 a party
from the Department of Botany of the Uni¬
versity of Hawaii visited a number of islands
of the Marshall group with the purpose of
collecting the species which occur there. The
party consisted of Dr. Harold St. John, pro¬
fessor and chairman of the department,
Mr. Richard S. Cowan, graduate assistant,
and the writer. Transportation and living
and working quarters were provided by the
United States Navy; the journey from Hono¬
lulu to Kwajalein, and thence to the more
northern islands, was made on a naval ves¬
sel, and to the islands south of Kwajalein
by seaplane.

Although only 10 of the 34 atolls and
single islands making up the archipelago
were visited, these lie in both the northern
and southern parts of the group, and in both
the Radak (eastern) and Ralik (western)
chains of which it is composed; and earlier
visits by Dr. St. John to other atolls indicate
that the 10 visited show nearly all the diver¬
sity of climate, topography, and vegetation
exhibited by the larger number. It is true
that all the islands on which the party landed
were inhabited, and it is possible that studies
of some of the uninhabited islands would
have given a somewhat different impression
of the vegetation. This limitation, however,
probably is of importance only for the col¬
lector of phanerogams; since stands of un¬
disturbed native vegetation occurred on the
inhabited islands also, it is unlikely that the

1 Department of Botany, University of Hawaii.
Manuscript received December 2, 1946.

others would have proved much more favor¬
able for mycological study.

Fungi were collected on eight atolls and
a single island (Mejit); the localities are
here listed, in order from north to south,
with the dates on which they were visited.

Atoll

(or single island)

Utirik
Mejit I.

Ailuk

Likiep

Wotje

Namu

Ailinglapalap

Jaluit

Ebon

Island

Utirik

Nankarai

Marab

Marme

Ailuk

Eniarmij

Biebi

Likiep

Ormed

Riri

Namu

Leuen

Ailinglapalap

Imrodj

Ebon

Date (1946)

September 1-2
September 3
August 31
August 3 1
August 31
August 30
August 29
August 29
August 28
September 4-5
September 5
August 16
August 16
August 20
August 20
September 9-12

Most of these names are those given on U. S.
Hydrographic Office charts 5413 and 5414.
The other localities are shown, but not always
named, on advance proofs of H. O. charts
6007, 6018, 6020, and 6022. In Ailuk
Atoll the ninth island north of Ailuk Island
is called by the inhabitants of the atoll
"Marab”; that name is used in this report,
although the proofs of chart 6022 desig¬
nate it by the apparently nipponized name
"Marappu.” The island immediately south
of it (and eighth to the north of Ailuk),
unnamed on the charts, is called "Marme”
by the Marshallese. The thirteenth island of
the series running north from Ailuk has the
native name "Nankarai.” In Likiep Atoll,
Biebi is shown on H. O. chart 6020. The

92

Fungi of the Marshall Islands — Rogers

island next to Biebi on the northeast, and
connected to it by a stretch of reef exposed
at low tide, is unnamed on the charts; ac¬
cording to Marshallese informants it is called
"Eniarmij.” Riri in Wotje Atoll lies next to
Ormed on the east; it is named on chart
6018. Imrodj in Jaluit Atoll is shown im¬
mediately above "N. E. Pass” on chart 5414,
and named on 6007.

August and September fall within the
rainy season for the Marshalls, and there had
been recent rainfall, enough not only to
moisten the soil but even to soak the fallen
logs, on all the islands except Ebon; often
there were showers or heavy rains during
the collecting trips. On Ebon there had been
no rain for some time, and the paddies de¬
voted to the cultivation of Cyrtosperma
contained mud but no standing water; never¬
theless, soil and vegetable debris were damp
and supported an abundant growth of fungi.
On the evidence of the vegetation, as well
as of published accounts, there is a consider¬
able increase in rainfall in the archipelago as
one goes from north to south; the vegetation
of the northern islands is somewhat more
open and poorer in species, and lichens and
other epiphytes are distinctly less abundant,
than in the islands south of Kwajalein.
Mejit, in the northern part of the group and
also considerably east of its nearest neigh¬
bors (Utirik, Ailuk, and Wotje), appears
much wetter than they. Ebon, the southern¬
most atoll of the archipelago, alone of those
visited supports the dense tangle of under¬
growth and the abundant epiphytes that
usually enter into the description of a jungle.
Any further attempt to characterize the vege¬
tation is beyond the scope of this report, and
must be left for Dr. St. John’s account of
the vascular plants.

I have been able to discover few dis¬
cussions of the fungi of the Marshalls.
Ehrenberg (1820) described Scaphophorum
* Agaricoides, Boletus Katui, B. sanguineus,
Sphaeria jur, and S. profuga from collec¬
tions by Chamisso in the Radak chain; these

93

were reduced to synonymy ( S . Agaric oides
= Schizophyllum commune, B. Katui =
Polyporus xanthopus, B. sanguineus = P.
sanguineus ) or redescribed, and validly pub¬
lished, by Fries (1821: 371, 504, 505;
1823: 431, 488) ; the two species of Sphaeria
were transferred to Metasphaeria by Saccardo
(1883: 182). Hennings (1897) listed or
described from Jaluit 13 species, from col¬
lections bySchwabe and byFinsch: Auricula-
ria auricula- judae, Polystictus sanguineus,
Schizophyllum Alneum, Pomes amhoinensis,
Polyporus Kamphoeveneri, Marasmius c al¬
io pus var. jaluitensis P. Henn., M. Pandani-
cola P. Henn., Psathyrella disseminata,
Psathyra Schwaheana P. Henn., Hypholomd
jaluitensis P. Henn., Galera sp. (confertae
aff.), Pleurotus Schwaheanus P. Henn., and
Lachnea jaluitensis P. Henn. Schumann and
Lauterbach (1901: 37-63) combined Ehren-
berg’s and Hennings’s lists, treating Psathyra
Schwaheana under Pratella and Pleurotus
Schwaheanus under Agaricus, and, for some
incomprehensible reason, reporting Meta¬
sphaeria jur as (( Metasphaeria Jus.” Volkens
(1903: 84) listed only species already re¬
ported. No other reports have come to my
attention. Thus, unless there are studies pub¬
lished during the Japanese occupation which
have been overlooked, the recorded myco-
biota of the Marshall Islands embraces 16
species. The University of Hawaii party
brought back approximately 425 specimens
of fungi, of which 126 specimens, repre¬
senting 34 species (2 of them new) , are here
listed. The study of this material is con¬
tinuing, and other reports will be published.

In the interests of brevity, in listing speci¬
mens only the atoll is given where but one
island of the cluster furnished collections of
fungi, and the date is omitted unless collec¬
tions were made on an island on more than
one day. Except where otherwise noted, the
collector is D. P. Rogers and the numbers are
”D. P. R. . . .” Portions of all Marshallese
specimens have been deposited in the Bishop
Museum (occasionally indicated by the sym-

94

bol BISH); portions of all collections of
Myxomycetes except three (duplicated by
other specimens from the Marshalls) , and of
all species of Heterobasidiomycetes, are in
the herbarium of the University of Iowa
( SUI ) ; certain specimens cited are in the
Farlow Herbarium (FH) ; portions of nearly
all collections are in the writer’s herbarium.
Where material is sufficiently abundant,
wider distribution has been or will be made.
The synonymy given is not intended to be
complete;, for most species only a few of the
more satisfactory and more readily available
descriptions are cited.

I am indebted to the University of Ha¬
waii, which through Dean Paul S. Bachman
and its Pacific Islands Research Committee
assumed the financial burden of the expedi¬
tion; to Captain John C. Hammock, Com¬
modore Benjamin F. Wyatt, Lieutenant H.
B. Wessinger, and those other officers of
the United States Navy who facilitated our
journey to the Marshalls; to Professor G. W.
Martin of the University of Iowa, who named
a number of the Myxomycetes and made
available extensive manuscript notes on Auri-
cularia; to Professor J. N. Couch of the Uni¬
versity of North Carolina, for notes on
material of Septobasidium; to Miss Hilda
Harris of the Farlow Library, who verified
citations of works not available in Hawaii;
and to Dr. St. John and Mr. Cowan, whose
assistance and companionship made the
journey an agreeable as well as a productive
one.

Myxomycetes

1. Arcyria denudata (L.) Wettst. —
Utirik, 1657; Mejit, 1492 , 1642, 1660;
Likiep: Likiep I., 1651; Jaluit, 1649; Ebon,
1631.

2. Arcyria nutans (Bull.) Grev. —
Likiep: Biebi L, 1652.

3. Ceratiomyxa fruticulosa
(Muell.) Macbr. — Mejit, 1628; Wotje:

PACIFIC SCIENCE, Vol. 1, April, 1947

Ormed I., 1661, Riri I., 1629; Ebon, 1634.
4. COMATRICHA TYPHOIDES2 (Bull.)

Rost. — Mejit, 1643; Jaluit, 1647; Ebon,
1636.

3. Cribraria tenella Schrad. — Jaluit,
1650, 1662.

6. Dictydium cancellatum (Batsch)
Macbr. — Ailuk: Ailuk I., 1653.

7. Fuligo septica (L.) Wiggers —
Ebon, 1627, 1638.

8. Hemitrichia serpula (Scop.) Rost.
— Namu: Leuen I., 1648; Ebon, 1635.

9. Hemitrichia stipitata (Mass.)
Macbr. — Ailuk: Ailuk I., 1654, 1706;
Wotje: Ormed I., 1640; Ebon, 1632.

10. Hemitrichia vesparium (Batsch)
Macbr. — Ailuk: Ailuk I .,1710; Ebon, 1637.

11. Perichaena depressa Lib. — Ebon,
1630.

12. Physarella oblonga (Berk, and
Curt.) Morg. — Ailuk: Ailuk I., 1708; Jaluit,
1646.

13. Physarum tenerum Rex — Mejit,
1641; Ailuk: Ailuk I., 1655.

14. Physarum viride (Bull.) Pers. —
Utirik, 1659.

15. Physarum Wingatense Macbr. —
Ailuk: Nankarai I., 1712; Wotje, Ormed I.,
1645, Riri I., 1626, 1639.

16. Stemonitis fusca Roth — Utirik,
1658; Ailuk: Ailuk I., 1656; Likiep: Likiep
I., 1625.

17. Stemonitis splendens Rost. —
Likiep: Biebi I., 1644.

None of the Myxomycetes collected is
sufficiently uncommon or unusual to be

2 As in all other matters, I have attempted to
follow the existing provisions of the International
Rules with respect to orthography. Cf. Rec. XLIII,
and the addition voted at Amsterdam.

Fungi of the Marshall Islands — Rogers

worthy of comment. The circumscription and
nomenclature are those of Macbride and
Martin (1934).

Phycomycetes

18. Glaziella aurantiaca (Berk, and
Curt.) Cooke, Grevillea 11: 83, 1883; Lloyd,
Mycol. Writings 7: 1203-1204, pi. 248,
fig. 2484, 1923; Boedijn, Buitenzorg Jard.
Bot. Jour. 3 ser. 11: 57-66, fig. 1-7, 1930.
Xylaria aurantiaca Berk, and Curt., Linn.
Soc. [London] Jour. Bot. 10: 382, 1868.
Glaziella vesiculosa Berk., Naturhist. For.
Kjobenhavn Vidensk. Meddel., 1879-80:
31, 1879; Thaxter, Amer. Acad. Arts and
Sci. Proc. 72: 334, pi. 4, fig. 88-94, 1922.

Fructifications tuberiform, bladderlike, approxi¬
mately isodiametric or flattened, plicate, scarlet,
growing paler toward the base, tough-membran¬
ous, hollow, opening by an irregular fissure either
at the base or above, about 1.5-5 cm. in greatest
diameter, early partly enveloped in a white bys-
soid veil, later free, the surface slightly roughened;
hyphae of the veil 4.5-5 in diameter, their walls
asperulate, about 0.5 n thick, the lumen inter¬
rupted at intervals of 15-60 n by thin septa; wall
of fructification composed of compact outer layers
of hyphae with inflated cells up to 10 ^ in diam¬
eter; of medullary hyphae loosely interwoven,
septate, 2-3 /* in diameter; and of embedded
chlamydospores; chlamydospores ellipsoid, about
300 X 200 /*, with a rigid wall 17-19 thick and
an outer apparently gelatinized membrane 5-7
thick, enclosing fluid material with numerous oil
globules.

On the surface, or under loose fragments,
of well-decayed vegetation, chiefly leaves of
Cocos nucifera. Ebon, R. S. Cowan and
D. P. R. (D. P. R. 1343).

The fungus is characterized adequately for
recognition merely as having hollow, red,
membranous fructifications; the few micro¬
scopic details given are only for possible
microscopic comparison with other material,

for it is less than certain that the Ebon col-

•

lection is conspecific with those reported
from the American tropics. Thaxter writes
of the "conspicuous orange yellow color” of
G. aurantiaca; and the specific epithet seems

95

(except by Boedijn) to have been taken
without challenge at its face value. The
carrying of a copy of Ridgway to the humid
tropics seemed too hazardous, and no color
comparison could be made in the field; but
the fructifications certainly were not orange.
My recollection is that the deepest-colored
would come close to Scarlet-Red,3 or at least
Scarlet, and the paler basal portions were not
yellowish, but a clear rosy pink. The col¬
lectors of Boedijn’s specimens gave the color
as "dunkel steinrot” and "orangerot.” Pre¬
sumably neither Thaxter nor Berkeley nor
Curtis ever saw the fungus in living condi¬
tion; and in 2 months the Marshalls collec¬
tion has faded to various orange-yellow
shades, and in both color and consistency
could very well be taken for slightly de¬
fective dried peaches or apricots. Without
knowledge of fresh American material it
would be rash and useless, therefore, to
attempt to distinguish the fungus from
G. aurantiaca by its color. Boedijn, whose
admirable account gives an excellent idea of
the Ebon material (except that the living
specimens here reported were not in the least
gelatinous), found no obstacle to the treat¬
ment of his specimens under the early name.
Mr. Cowan found the greater number of
fructifications (11 were collected) in deep
shade under a tangle of bushes in a dense
grove of coco palms. The species is reported
from half a dozen islands of the West Indies,
from Brazil and Mexico, and from two
islands in the Netherlands Indies. Lloyd
(loc. cit.) described a second species from
Nicobar Island, which in Thaxter’ s opinion
(Stevenson and Cash, 1936: 3) may be iden¬
tical. There are other synonyms.

Basidiomycetes

19. Tulasnella allantospora Wakef .
and Pears., Brit. Mycol. Soc. Trans. 8: 220,
fig. 7, 1923; Rogers, Ann. Mycol. 31: 190,

3 Color-names capitalized are used in the sense
of Ridgway (1912).

96

pi. 6, fig. 5, 1933; Martin, Iowa Univ.
Studies in Nat. Hist. 18 (3): 18, pi. 1,
fig. 4, 1944.

On rotten, friable wood of Ar to car pus
incisus. Ebon, IX. 10.46, 1366.

Like a number of European and American
specimens, that from Ebon was a barely
perceptible bloom when living, and dried is,
except under the binocular, wholly invisible.
It can readily be recognized in microscopic
preparations by the four inflated epibasidia
and the thick-allantoid spores. The latter in
the collection cited are mostly 7-7.5 X 3.5-4
P , relatively stout, like the type, but very little
tapered toward the ends, more like some
Iowa material. Previously reported from
western Europe and from the northeastern
quarter of the United States, the southern¬
most station being in Missouri. Martin
(1939: 242) has reported two other species
of Tulasnella from the tropics: T. violea,
a common species of western Europe, the
northeastern states, and southeastern Canada,
from Colombia; and T. sphaerospora, un¬
known elsewhere, from Panama. Gloeotu-
lasnella calospora occurs in Hawaii.

20. Ceratobasidium cornigerum
(Bourd.) Rogers, Iowa Univ. Studies in
Nat. Hist. 17: 5, pi. 1, fig. 2, 1935; Martin,
ibid. 18 (3): 14, pi. 1, fig. 1, 1944.

On decorticate wood of Art o car pus in¬
cisus. Likiep: Likiep I., 1301.

A grayish-pruinose growth showing under
the microscope repent mycelium; obovate,
aseptate hypobasidia bearing (usually) four
tubular epibasidia like those of a tremella;
and oblong-ellipsoid spores. The shape of
the spores reaches the extreme of elongation
known in this species, quite like Figure 2(g)
of the Iowa paper. Hitherto reported from
the Austrian Tyrol and central France in
Europe, and in the Western Hemisphere from
southern Canada and the northern states. In
comparing the Likiep specimen with main¬
land material, I observed, in D. P. R. 144

PACIFIC SCIENCE, Vol. 1, April, 1947

from Iowa, basidia with five well-developed
epibasidia. More than four epibasidia or
spores apparently are developed elsewhere in
the Tremellales only in Gloeotulasnella.

21. Sebacina cinerea Bres., Fung.Trid.
2: 99, ph 210, fig. 2, 1892; Rogers, Iowa
Univ. Studies in Nat. Hist. 15 (3) : 12, pi. 1,
fig. 4-6, 1933; McGuire, Lloydia 4: 37,
pi. 5, fig. 91-94, 1944; Martin, Iowa Univ.
Studies in Nat. Hist. 18 (3): 39, 1944.
Exidiopsis cerina Moll., Protobas. 85, 167,
1895. Thelephora cinerea (Bres.) Sacc. and
Syd., Syll. Fung. 16: 183, 1902; nec Pers.
ex Fr., Syst. Mycol. 1: 453, 1821.

On rotten, friable wood of Artocarpus
incisus. Jaluit, 1380.

A thin cinereous-buff crust, under the
microscope showing irregularly linear gloeo-
cystidia with often yellow-granulose content,
ellipsoid cruciate-septate basidia, and obtuse
cylindric spores. The fructification cited is
young, and thinner than is usual for the
species. Its spores, which have the form
characteristic of S. cinerea, are smaller than
in most collections (7. 5-8. 5 (-10) X 4-4.5
(“5) /x), and near the lower limit of size
published for the species. The form of the
basidia and the degree of compactness of the
hymenium, both significant characters in the
subgenus Bourdotia to which the species be¬
longs, are typical.

The two synonyms given are an addition
to those cited in previous discussions. The
comparison of Moller’s descriptions with
good material of S. cinerea leaves little need
for hesitation in equating his Exidiopsis to
the present species. Acceptance of that
synonymy extends the range of S. cinerea to
southern Brazil; its known range already
stretches from Ontario to Panama, from
Massachusetts to Oregon, and in Europe
from Finland to Italy.

22. Sebacina dubia (Bourd. and Galz.)
Bourd.; Rogers, Iowa Univ. Studies in Nat.
Hist. 15 (3): 11, ph 1, fig. 1-3, 1933;

Fungi of the Marshall Islands — Rogers

97

McGuire, Lloydia 4: 31, pi. 4, fig. 73-74,
1941; Martin, Iowa Univ. Studies in Nat.
Hist. 18 (3): 37, 1944. Heterochaetella
dubia (Bourd. and Galz.) Bourd. and Galz.,
Hym. Fr. 51, fig. 30 [1928].

On dead wood of Cocos nucifera. Ebon,
IX.9.46, 1391.

The fructification is distinctly cretaceous
and, unlike that in most specimens of the
species, clearly visible to the naked eye — and
about as conspicuous as a thin dab of white¬
wash. Under the lens it is seen to consist of
numerous more or less separated whitish
conical spicules; under the microscope each
of these may be seen to be composed of a
number of parallel cystidia, surrounded and
conglutinate by a film of indistinct hyphae
which ascend for fully two thirds of the
height of the spicule and bear on their sur¬
face cruciate-septate basidia with four very
short epibasidia continued in long, subulate
sterigmata. The cystidia of the Ebon speci¬
men differ somewhat from those developed
in other collections; but as noted elsewhere,
the same may be said of almost any speci¬
men: they are a variable lot. The cystidia,
rather than being straight, thick-walled and
with a narrow lumen at the base, and thin-
walled and with a dilated lumen at the
summit, as often in Peniophora sect. Tubuli-
jerae, are contorted, often several times sep¬
tate, constricted at some septa or for the
entire length of some of the segments, and
with the wall only slightly thickened. They
show considerable resemblance to those of
Peniophora pallidula. Some, however, ap¬
proach the regularity of more typical Seba-
cina dubia. Known from widely separated
localities in temperate Europe and North
America; reported by Martin [loc. cit .) from
Brazil.

23. Sebacina farinacea sp. nov. Fig. 1.

Fructificatio viva alba, farinaceo-ceracea, mar-
gine attenuate), zona peripherica sub lente e gran-
ulis albis distinctis (cystidiis) composita, exoleta
pallide griseo-pruinosa, sicca cretacea vel leviter
sordida; hyphae nodoso-septatae, 1.5-3 A* diam.;

paraphyses sparsi, tenues, ramosi; cystidia e cor-
poribus cylindraceis 3-4 a* diam., valde granulis
calcareis incrustatis, composita, stipitata, subfusi-
formia, 20-33 X 7.5-10 a*; gloeocystidia linearia,
irregularia, matura suco luteo granuloso suffulta,
17-50 X 5-6 /^; basidia cruciato-septata, ellipsoi-
dea vel oblonga, 14-16.5 (-18) X 8.5-11.5 a*, epi¬
basidia quattuor 2-2.5 A* diam., long, ad 16 a q
gerentia; spo'rae oblongo-ellipsoideae vel obovatae,
8-9.5 X 6-7 A*, per repetitionem germinantes.

Fructification when fresh pure white, becoming
slightly creamy (a little lighter than Cartridge
Buff), adnate, separable in small bits, farinaceous-
waxy, under the binocular with minutely farina¬
ceous surface, thinning out at the margin to a peri¬
pheral zone composed of minute white granules
(cystidia) arising out of a thin transparent waxy
film; old specimens grayish-pruinose; when dry
chalky white to barely cream (paler than Ivory
Yellow) or in older fructifications nearly as dark
as Cartridge Buff; hyphae distinct, with clamps
throughout, 1.5-3 A1; paraphyses scattered, reach¬
ing the surface, mostly inconspicuous, the stalk
linear, 2-3 A* in diameter, with peg-like or con¬
torted branches 1 a* in diameter; cystidia arising
at the margin and later engulfed by the hymenium
and lying at its base, composed of a subcylindric,
thin-walled cell 3-4 a* in diameter and heavy coarse
mineral incrustation, at maturity subfusiform, ob¬
tuse, stalked, 20-33 X 7.5-10 a*; gloeocystidia aris¬
ing near the base of the fructification, somewhat
sinuous, irregularly inflated or constricted, thin-
walled, the content early hyaline and homogene¬
ous, later yellow, granular, refractive, resinoid,
17-50 X 5-6 a*; basidia subtended by a prolifer¬
ative clamp, ellipsoid, becoming oblong and trun¬
cate, 14-16.5 (-18) X 8.5-11.5 A*, longitudinally
cruciate-septate, bearing 4 epibasidia 2-2.5 A* in
diameter and up to 16 a* or more in length; spores
oblong-ellipsoid to obovate, 8-9.5 X 6-7 A*, ger¬
minating by repetition.

On dead fibrous leaf sheaths, leaf bases,
and husks of Cocos nucifera.

Hawaii: Oahu: University of Hawaii
campus, Manoa, 11.12.46, I. A. Abbott and
D. P. R. (D. P. R. 633); 11.17.46, 1132;
III.20.46, 1. A. Abbott and D. P. R. (D. P. R.
1173), and 1176; III. 3 1.46, 1243; VI.2 1.46,
1327, 1328; X.29.46, 1884, type (in herb.
D. P. R., BISH, SUI) ; X.30.46, 1888;
XII.4.46, 1931.

marshall islands: Ebon, IX. 10.46,

1388.

To the naked eye Sebacina farinacea pre¬
sents the appearance of one of the innumer¬
able small white Thelephoraceae — perhaps

98

PACIFIC SCIENCE, Vol. 1, April, 1947

Fig. 1. Sebacina far mace a: A-I, basidia; J-M, spores; N-Q, cystidia; R-S, gloeocystidia; T, para-
physis. C, F, I, K, M, N, O, Q, S = D.P.R. 1888; A, B, D, E, G, H, J, L, P, R, T = D.P.R. 1931.

Fig. 2. Sebacina petiolata: A-H, basidia; I-M, spores; N, paraphysis; O, gloeocystidium. All D.P.R.
1473.

All drawings were made with camera lucida under oil-immersion objective and 10X ocular at 1675 X.
and reduced in reproduction to approximately 1000 X- Fig. 1 B and I and 2 C, D, and F-H are incom¬
plete (optical section) and do not represent two-celled basidia.

Fungi of the Marshall Islands — Rogers

a thin growth of Corticium suecicum or
Peniophora Samhuci. In structure it is most
nearly allied to those species of Sebacina
subgenus Bourdotia with well -developed
paraphyses, such as S. Galzinii; its gloeo-
cystidia are similar, as are the arrangement
of the basidia and their frequent elongate
form. It is, however, much less gelatinous,
and more resistant to separation of the
hymenial elements under the cover-glass. On
the other hand, its possession of paraphyses
and commonly elongate basidia separate it
from the group of more arid species of which
S. caesio-cinerea, for example, is a member,
and it does not show the sheath of empty
basidia surrounding the fertile hypha that is
so characteristic of those species. The heavily
encrusted cystidia, resembling those of Penio-
phora pubera, are apparently unique in Seba¬
cina. In fact, their place of formation — in a
peripheral zone, where they mature before
any other hymenial elements are developed
— seems to be unique in the whole known
range of the Basidiomycetes.

The subgenus Heterochaetella is defined
by the possession of cystidia. Sebacina jari-
nacea shows no indication of affinity to the
species of Heterochaetella other than its
cystidia; and even that exists only because
the term "cystidium” is employed for a
number of very different structures. The
affinities of S. jarinacea being rather with
S. Galzinii, it is assigned to the subgenus
Bourdotia.

Since the available keys to Sebacina all
make use of the dichotomy "gloeocystidia
present : cystidia present” in one form or
another (Bourdot and Galzin, 1928: 17;
Rogers, 1935: 37; McGuire, 1941: 11; Mar¬
tin, 1944: 35), S. jarinacea can be included
most readily by the insertion of an additional
choice :

"Both gloeocystidia and encrusted cysti¬
dia present. . S. jarinacea

Sebacina jarinacea can be found after
almost any prolonged rainy period on the
dead sheaths of leaves attached to young coco

99

palms. Probably it could be collected more
abundantly if the bases of the leaves of
mature trees were less inaccessible. The Ebon
specimen is the only one found elsewhere —
on the rotting husk of a coconut. It is quite
old and poor, but recognizable.

24. Sebacina petiolata sp. nov. Fig. 2.

Fructificatio viva gelatinosa, opalea vel hyalina,
luteo-, roseo-, vel cyaneo-tincta, 0.5-3 mm. crassit.,
sicca hyalino- vel ochraceo-vernicosa; hyphae ple-
rumque distinctae, nodoso-septatae, 2-2.5 At diam.,
sub basidiis ad 5 At expansae; paraphyses fili-
formes, apicem versus furcatae vel fruticulosae,
raro nodoso-septatae, 1-3 At in diam.; gloeocystidia
primo hyalina, homogenea, serius luteo-granulosa,
irregulariter subcylindracea vel subfusiformia, ob-
tiisa, 39-150 X 4-7. a*; probasidia primo clavata,
hypobasidia matura obpyriformio-clavata, cruci-
atim septata, 21-32 X 9-12.5 At, ad apicem lobata,
epibasidia 4 crassa, 10-50 X 3—5.5 At gerentia,
basidiorum stipitibus ab hypobasidio separatis a
septis eisdem verticalibus parietem versus curvatis;
sporae hyalinae, ellipsoideo-oblongae vel ellipsoi-
deo-subglobosae, 7—1 1 X 6-8 At, per repetitionem
germinantes.

Fructification gelatinous, hyaline or yellowish-,
pinkish-, bluish-, or plumbeous-opalescent, 0.5-3
mm. thick, the margin attenuate or abrupt and cili-
ate, when dry hyaline- or ochraceous-vernicose;
hyphae mostly distinct, with clamps throughout,
2-2.5 At in diameter, broadened up to 5 M just be¬
low the basidia; paraphyses filiform, near the tips
forked to bushy, occasionally nodose-septate, the
basal portion 1.5-3 At in diameter, the tips con¬
torted, nodulose, unencrusted, about 1 At in diam¬
eter, finally obscure; gloeocystidia with hyaline
homogeneous content, becoming coarsely yellow-
granular, irregularly subfiliform, subcylindric to
fusiform or clavate, obtuse, 39-150 X 4-7 a*;
basidia arising as clavate bodies, becoming obpyri-
form-clavate, the hypobasidia cruciate-septate,
lobed at the summit, the inflated terminal portion
16-24 X 9-12.5 At, the stalk 20-27 X 3-4.5 a*.
separated from the fertile summit by the longitu¬
dinal walls, which curve away from their line of
intersection to meet the outer wall, the epibasidia
very thick near their bases, 10-50 X 3-5.5 A*;
spores hyaline, evenly oblong to ellipsoid-oblong,
9-11 X 6-7.5 A*, or ellipsoid-subglobose, 7-9 X
6-8 A*, germinating by repetition.

Growing over bark of a dead branch
(unidentified), over an old fungus, and on
wood of Acacia Koa, Aleurites moluccana,
Artocarpus incisus, Hibiscus Arnottianus,
Messerschmidia argentea, Psidium Guayava,
Samanea Saman (= Pithecolobium Saman ),

100

P and anus sp., and bark and wood of Cocos
nucifera.

Cuba: Blanco’s Woods,. Soledad, Cien-
fuegos, VII. 1.41, IF. L. White 596; San
Bias, Santa Clara Prov., VI. 20-2 1.41, IF. L.
White 407, 427 (all in FH).

Hawaii: Oahu: east slope Manoa Valley
(500-900 ft.), XI. 18.45, 1218; Tantalus
Trail, Pauoa, VII. 2 1.46, 1331; S. branch of
N. fork of Ekehanui Gulch (1,800-2,400
ft.), Honouliuli, III. 17.46, 1169 ; Kupehau
Gulch above pipe-line trail (2,000 ft.),
Honouliuli, X. 13.46, R. S. Cowan 179 .

marshall islands: Utirik, IX.1.46,
1525, 1548; Mejit, 1436, 1441, 1450, 1513,
1514, 1545; Ailuk: Marab I., 1571, Ailuk
I., 1665, 1701; Likiep: Likiep I., on log of
Cocos, VIII.28.46, 1475, type (in herb.
D. P. R., BISH, SUI, FH); Wotje: Ormed
I., IX.4.46, 1385, Riri I., 1500; Namu:
Leuen I., 1403, 1431; Jaluit, 1617; Ebon,
IX. 9.46, 1393.

A continuous, usually thick gelatinous
layer, composed (under the microscope)
of branched paraphyses, gloeocystidia, and
cruciate-septate basidia whose fertile distal
part is separated from the stalk-like basal
part by the intersecting vertical walls, which
turn out to meet the outer wall of the hypo-
basidium. The apparent color varies con¬
siderably, according to the thickness, the
color of the substratum which shows through
the clear or opalescent basidiocarp, and
probably other factors. The collection from
Kupehau when fresh was pinkish — Pallid
Mouse Gray to Pale Vinaceous-Fawn, where¬
as the one from Ekehanui, not far away, was
in part Pale Olive-Gray and in part colorless
and merely pruinose; others were noted
in the field as being yellowish-opalescent,
bluish-opalescent, blue-white, iavender-giliy,
light neutral gray, and deep blue-gray. In
texture and presence of gloeocystidia and
paraphyses S. petiolata is related to S. Gal-
zinii and S. umhrina, from both of which it
differs in the manner of branching of the

PACIFIC SCIENCE, Vol. 1, April, 1947

paraphyses, in clavate basidia, and in spores.
The presence of the stout stalk separated by
the vertical septa from the fertile portion is
the clearest recognition-character. The new
species may be accommodated in Martin’s,
McGuire’s, or Rogers’s keys (earlier cited
under Sebacina jarinacea ) by the addition of
a third choice to the dichotomy separating
S. umhrina and S. Galzinii (or S. Pulula-
huana) :

"Gloeocystidia similar to those of S. Gal¬
zinii ( Pululahuana)\ spores oblong, ellip¬
soid-oblong, or ellipsoid-subglobose, 7-11
X 6-8 /x; basidia subclavate, the stalk sepa¬
rated by the vertical walls from the fertile
summit . . . S. petiolata”

25. Stypella minor Moll., Protobas.
77, 166, pi. 4, fig. 7, 1893; Martin, Iowa
Acad. Sci. Proc. 36: 128 [1930]; Iowa
Univ. Studies in Nat. Hist. 16 (2): 147,
fig. 2, pi. 6, 1934; ibid. 18 (3): 34, 1944.

On dead rhachis and charred wood of
Cocos nucifera, bark and wood of Pandanus
sp., dead wood of Art o car pus incisus, bark
and wood of Carica Papaya, and decorticate
log of unidentified species. Ailuk: Marab I.,
1572, Ailuk L, 1699; Mejit, 1454; Wotje:
Riri I., 1760; Jaluit, 1585, 1587, 1606,
1610, 1614; Ebon, IX.10.46, 1799.

When young, and about the margins when
mature, formed of distinct minute gelatinous
pustules, which later are confluent into a
reticular or apparently continuous, uneven¬
surfaced layer. This inconspicuous but char¬
acteristic fungus, originally described from
southern Brazil, has since been reported as
occurring more generally in tropical Amer¬
ica and in the eastern and central states at
least as far north as Wisconsin and Massa¬
chusetts. It certainly occurs and has been
reported (under other names) in western
Europe, and has several times been collected
in Hawaii.

Four of the specimens here listed are in
all respects quite typical, and are not dis¬
tinguishable from Iowa material. The five

Fungi of the Marshall Islands' — Rogers

collections made on Jaluit, however, show in
varying degrees a more luxuriant growth;
the separate lenticular or tuberculiform ba-
sidiocarps are confluent to form not merely
a reticulate aggregation, or the thin film
which seems to be the most homogeneous
fructification developed in temperate lands,
but a tough, separable membrane, held to¬
gether by a dense subicular layer, almost
cartilaginous in consistency, of conglutinate
hyphae. In some of the specimens this layer
on drying has remained intact, forming
a hyaline, translucent pellicle, glistening
(under the binocular) from light reflected
from the sides of the minute polygonal
areoles which represent the originally sepa¬
rate basidiocarps, and from their very short
narrowed stipe-like bases, which remain
separate even in the older parts of the fruc¬
tification, and which can be made out through
the continuous surface. The spores and ba-
sidia of the Jaluit material also average
somewhat larger than those of most other
specimens. There is, however, considerable
overlapping in these as in other respects;
the best developed of the Jaluit specimens is
well matched by one from Hawaii, while the
more delicate specimens or areas are not
distinguishable from some of the Hawaiian
and North American ones. It is certain that
conditions of temperature and humidity on
Jaluit at the time of collection were ex¬
tremely favorable for the development of
tremellaceous fungi, and to the weather may
be ascribed the extreme forms here noted.

26. Guepinia Spathularia ( Schw. )
Fries; Coker, Elisha Mitchell Sci. Soc. Jour.
25: 177, pi. 23, fig. 14; 51; 64, fig. 5, 6, 9,
1920; Brasfield, Amer. Midland Nat. 20:
221, pi. 3, fig. 64-66, 1938; Lloydia 1:
155, 1938; Martin, Iowa Univ. Studies in
Nat. Hist. 18 (3): 30, 1944. Merulius
Spathularia Schw., Naturf. Gesell. zu Leip¬
zig Schr. 1: 92, pi. 2, fig. 1-3, 1822.

On bare wood, or growing out through
fissures in the bark, of P and anus sp. Utirik,

101

1554; Ailuk: Marme I., 1750; Mejit, 1440,
1455, 1508.

Guepinia Spathularia as here treated
includes all those fungi with fructifica¬
tions cartilaginous-gelatinous in consistency,
orange or brownish-orange in color, flabelli-
form or incised-flabelliform, tomentose on
the compressed stipe and one surface of the
expanded summit, with forked basidia and
one-septate spores. All of the Marshallese
collections have the basidiocarps less ob¬
tuse at the distal margin, more translucent,
brighter (and redder) in color, and less
tomentose on the sterile surface, than any
material at hand from temperate regions,
and than any shown in the illustrations cited.
Microscopically, although the spores may be
slightly more slender, no significant differ¬
ences appear to exist. A considerable num¬
ber of species have been described from the
tropics, most of them differing little from
G. Spathularia. Several recent authors have
indicated doubt of the autonomy of certain
or all of these species; and it seems neither
necessary nor prudent, in the absence of any
considerable amount of tropical material for
comparison, to attempt at present a decision
on the distinctness of these segregates or of
the Marshall Islands material.

27. Auricularia ampla Pers. in Gaudi-
chaud, Bot. Freycinet Voyage autour du
Monde . . . Uranie et . . . Physicienne 177,
1827. Exidia ampla (Pers.) Lev., Ann. Sci.
Nat. Bot. 3 ser. 5: 159, 1846. Hirneola
ampla (Pers.) Fries, K. Vetensk. Akad.
Hand!., 1848: 146, 1849.

Fructifications when fresh thin, flexible, leathery-
gelatinous, the dorsal surface uniformly pilose,
cinereous-gray, -pink, or -buff, or in old specimens
fuscous, glabrescent, the hymenial surface more or
less pruinose, dull flesh-color, purplish-rosy, dull
purplish, or in older specimens blackish; solitary
or gregarious to densely imbricate-caespitose,
eccentrically peltate or sessile by a narrowed pos¬
terior margin or produced into a short stipe-like
sterile base, up to 5 (rarely -20) cm. in breadth,
convex and even or slightly plicate above, cupulate
and even or in larger specimens shallowly venulose
below; when dry thin, flexible, quite fragile, trans-

102

PACIFIC SCIENCE, Vol. 1, April, 1947

lucent, pallid-cinereous to buffy-cinereous above or
in old specimens sometimes dark and nearly gla¬
brous, below purplish-black to slaty-black, con¬
tracted and usually considerably wrinkled; in sec¬
tion composed of a hymenial layer about 70 a*
thick, a medullary layer about 700 a* thick, and a
compact dorsal layer about 20 thick underlying
the tomentum; hymenial layer consisting of
densely compacted basidia, of paraphyses 1.5-2 v*
in diameter with brownish branching indistinct
tips exceeding the basidia and forming a dense
continuous superficial layer, and of slender con¬
torted subhymenial hyphae; medullary layer of
rather loosely arranged hyphae with irregular
often open clamps and confluent, highly gelatin¬
ized walls, apparently 2-3 M in diameter but much
thicker if the limits of the walls could be observed,
mostly parallel to the surface in the middle and
perpendicular in the upper and lower quarter;
dorsal layer consisting of an inner colorless half
of heavily stainable hyphae 1 At in diameter,
densely compacted, perpendicular to the surface,
and with little gelatinized wall material between,
and an outer pigmented layer abundantly pene¬
trated by the bases of the surface hairs; hairs up
to 225 X 4.5-7 /*, thick- walled and aggregated in
conical tufts; basidia linear or slightly clavate,
40-50 X 4.5-5 A*, 3-septate; spores allantoid,
15.5-17 X 4.5-5. 5 A*.

In the Marshalls usually on wood of Cocos
nucifera, with one collection on Artocarpus
menus. Wotje: Ormed I., 1353, 1354,
1379, 1383; Namu: Namu I., 1404, 1414,
1434; Ebon, 1338.

A thin, leathery, often ear-like basidio-
carp, pilose and ashy to nearly glabrous and
brownish-black above, and rosy to purplish-
black below. The most abundant growth
was encountered on Ormed, where large
clusters had developed on standing dead
trunks of coco palms, many of which had
been topped by the blockaded Japanese gar¬
rison for their edible terminal buds, while
others had been cut by artillery fire or bomb-
blast. The Marshallese name, "lajiling kiji-
lik” (rat’s ear), is applied without much
discrimination to many bracket- or ear-like
or even stipitate Basidiomycetes — polypores,
Pleurotus, and others — but there was some
suggestion that this species is the true rat’s
ear, and on Ormed the name was given only
to this Auricularia. Strangely enough, it is
not eaten in the Marshalls, whereas in

Hawaii the same species, under the name
"pepeiaoakua” (ghost ear) is regarded, not
mistakenly, as good food.

Persoon’s description [loc. cit.) reads as
follows:

Afuricularia] magna resupinata substipitata,
inferne pubescens lutescente-grisea, superne laevis
nigricans. Pers.

In insulis Mariannis inque Moluccis.

Pour la configuration et la couleur, cette espece
ressemble beaucoup a V auricularia sambuci [i.e.,
A. auricularis (S. F. Gray) Martin, Amer. Mid¬
land Nat. 30: 81, 1943]; mais elle s’en distingue
par sa partie fertile, qui n’est presque pas veineuse,
et parce qu’elle pousse en-dessous une sorte de
stipes ou appendice long de quelques lignes. A ces
caracteres, il faut ajouter V habitat, qui pourtant
ne doit pas avoir indue sur la forme et la dimen¬
sion? P.

In this there is, first of all, an obvious error:
the description has the basidiocarp inverted;
it is not pubescent below, but above. (It
may be noted that Fries and succeeding gen¬
erations of mycologists made the same mis¬
take for all species which, like this, were
placed in the. genus Hirneola. See Fries
\loc. cit.'], p. 144; and 1825: 93.) The
presence of a stipe, furthermore, is here no
more than fortuitous; it is not a discrete
structure, but an indistinct prolongation of
the basidiocarp, and probably no more than
an occasional response to the position of the
latter, or to such an external condition as
development through a fissure in the bark.
The same development occurs in northern
specimens of A. auricularis. Neither is the
configuration of the hymenium a constant
character; among the numerous collections
at hand some are quite smooth, and others as
venulose, and as ear-like in their convolu¬
tions, as any specimen of A. auricularis. The
word "resupinata” need not be misleading
if it be noted that Persoon (1822: 97) uses
the same adjective in describing the familiar
northern A. auricularis. What is left, then,
of Persoon’s account is a description of a
large species of Auricularia, yellowish-gray-
pubescent above, blackish below, greatly re¬
sembling A. auricularis, and occurring from

Fungi of the Marshall Islands — Rogers

103

the Marianas to the Moluccas. The size of
the Marshalls specimens is not remarkable,
nor does the largest specimen at hand, a
Hawaiian basidiocarp 20 X 12 cm., signifi¬
cantly exceed the maximum given by Mar¬
tin (1944: 65) for A. auricularis — which,
as it happens, is also described by Persoon as
"sat magna”; but the typical development of
the tropical fungus seems to be a larger one.
The yellow-ashy dorsal surface and blackish
hymenium are characteristic of mature speci¬
mens, grown in strong light, of the form
under discussion, which is probably the one
Auricularia of the Pacific islands resembling
A. auricularis. Whether the two are dis¬
tinct has been doubted; probably reports of
A. auricula-judae from Oceania, such as.
Hennings’s, cited earlier, and Patouillard’s,
refer to the fungus here called A. ampla. By
its more strongly pilose basidiocarps, con¬
spicuously rosy young hymenia, and some¬
what larger spores it is at least readily dis¬
tinguishable, and one inclined to question
the distinctness of the Pacific form should
note that the differences are much greater in
living than in preserved material. Unsatis¬
factory as it is, Persoon s account gives as
sound a basis for the identification of his
fungus as do those of later authors whose
names have been more commonly adopted.
Briefly, that basis is an abbreviated descrip¬
tion which, allowance being made for ob¬
vious error, is quite applicable, as far as it
goes, to the fungus at hand, and with that
description a geographical limitation which
seems to confine the application to the one
species. Reference to the original valid de¬
scriptions of accepted species of Auricularia
will show that they are no better founded. It
is at least highly probable that the form
under consideration is A. ampla, and that
name should be superseded only if reason
appears to doubt the correctness of its appli¬
cation, or if an earlier name, likewise prob¬
ably applicable, exists.4

Dried mature specimens from the Mar¬
shalls mostly show on the dorsal surface a

color that cannot be exactly matched in
Ridgway — more tawny, and less greenish,
than Olive-Buff or Deep Olive-Buff; the
grayer specimens are Pale Olive-Gray to
Mouse Gray; the hymenium is Light Quaker
Drab to Blackish Plumbeous. A specimen
soaked for spore-printing (and still alive)
showed the hirsute surface Pallid Mouse
Gray and Light Mouse Gray to a tawny
color like that of the dried material, and the
hymenium Light Heliotrope-Gray to Dark
Purple -Drab, Dark Vinaceous-Gray, and
Heliotrope-Slate; certainly young specimens
were redder when first collected. Freshly
collected Hawaiian specimens have the dor¬
sal surface Tilleul Buff and Vinaceous-Buff
to Sorghum Brown, Drab-Gray, and Drab,
and the hymenium Russet- V inaceous to
Sorghum Brown. The colors of the very
brightest young hymenia have not been
recorded; according to my recollection they

4 Through the great kindness of Dr. A. J. Lam
and R. A. Maas Geesteranus of the Rijksherba-
rium, Leiden, I have recently (11.18.47) been
permitted to study adequate fragments of the
type of A. ampla and a photograph of the entire
type collection. The two fructifications there pre¬
sent measure respectively 4.5 X 3.5 and 8X7 cm.;
the dorsal surface is evenly pilose, and Cinnamon
Brown to Prout’s Brown (Mr. Maas Geesteranus
writes that the smaller specimen is glabrescent) ;
the ventral surface is nearly smooth (only a few
low folds appearing in the photograph), not
pruinose, and Plumbeous Black or Black. The
layers shown in section are the same as those here
described in the Marshalls material except that for
about 75 ^ above the basidia the medullary layer
includes a continuous mass of colorless, amor¬
phous, refractive material not seen in any Mar¬
shalls specimen sectioned. Persoon’s type and the
Hawaiian and Marshall Islands specimens agree
very closely, and their identity may be considered
to be established. The name A. ampla which
heads this note cannot, however, be retained; it is
antedated by Exidia cornea (Ehrenb.) ex Fries,
Syst. Mycol. 2: 222, 1822 {Auricularia cornea
Ehrenb., Hor. Phys. Berol. 91, pi. 19, fig. 9, 1820),
a name applied to the same fungus collected on
Oahu by Chamisso. The valid name is then Auri¬
cularia cornea ([Ehrenb.} ex Fries) Ehrenb. ex — .
I have been unable to find that this binomial has
been validly published; it seems unlikely that a
still earlier name will be discovered.

The type of A. ornata is now lacking at Leiden.

104

PACIFIC SCIENCE, Vol. 1, April, 1947

would be at least as red as Purplish Vina-
ceous.

28. Auricularia ornata Pers. in
Gaudichaud, Bot. Freycinet Voyage autour
du Monde . . . Uranie et . . . Physicienne
177, pi. 2, fig. 4, 1827. A. adnata Lyon in
Pvock, Hawaii Coll. Publ. Bui. 4: 33, 1916.

Fructification when fresh tough fleshy- to car¬
tilaginous-gelatinous, adnate over much of its area
or centrally peltate-rooting, disciform, lobed-
disciform, or by confluence elongate, 0.5-7. 5 cm.
in diameter, the margins at first closely appressed,
later often reflexed to 0.5 cm., the lower (hyme-
nial) surface pruinose, soft brown (Pale Brownish
Drab to Natal Brown), in young specimens
smooth to slightly rugose, in older with strong,
irregularly forking, vein-like thickenings more or
less radially arranged, the upper surface where
free coarsely pilose, zonate, the zones alternately
of long and of short hairs, mostly dark brown
(Snuff Brown to Bister), but interspersed with
light bands (about Pinkish Buff), in young speci¬
mens with appressed margins the pilosity shown
only as a narrow buffy margin; when dry hard, in
consistency like dried cartilage, the hymenium
strongly pruinose, plumbeous (Cinereous to Dark
Plumbeous), with younger portions lighter and
browner (Drab-Gray); in section composed of a
hymenial layer about 85 A* thick, a medullary layer
about 1000 At thick, and an abhymenial layer about
30 A1 thick; the hymenial layer consisting of a pig¬
mented superficial portion about 15 v thick com¬
posed of the indistinct, branching ends of para-
physes 1-1.5 ^ in diameter, and a basidial portion;
the medullary layer consisting of hyphae 3-9 A* in
diameter with irregular, often open clamp connec¬
tions and greatly gelatinized walls, densely com¬
pacted and perpendicular to the surface in the
lower and upper 100 At, looser and more or less
parallel to the surface in the middle; the abhy¬
menial layer brown, nearly opaque, compact and
(in the free parts of the basidiocarp) supporting
a pilose layer about 70-300 /t thick (according to
the surface zonation) of free brown hyphae;
basidia linear or often narrowly claviform or
basally ventricose, 3-septate, 52-70 X 5.5-7 (rarely
-8.5) /t; spores thick-allantoid, with stout truncate-
apiculus, 12.5-15.5 (-17) X 5.5— 6.5 A*.

On bark and decorticate wood of Messer -
schmidia argentea.

line islands (Pacific): Palmyra I.,
VIII. 20. 24, H. F. Bergman [A], [£], det.
by Lyon as A. adnata.

marshall islands: Utirik, 1549; Ailuk:
Marab I., 1568, Ailuk I., 1518 ; Wotje:
Riri I., 1372; Namu: Leuen I., 1408 .

A leathery growth, brown above and
below when moist and active, apparently
limited to the single substratum Messer-
schmidia (= Tournejortia ) argentea, and
to hard, elevated, little decayed branches of
that species. The earliest of the Marshalls
collections was made in a soaking rain, and
the fungus was at first taken to be a some¬
what thicker and softer relative of Stereum
hirsutum. The highly characteristic brown
color disappears from the hymenium on
drying, and is replaced by a pruinose slate
not too different from the appearance of
other species of Auricularia. Since A. ornata
is one of the far too numerous lost species,
the original description is here reproduced:

A[uricularia} pileo dimidiato reflexo tomen-
toso, zonis fuscis pallidisque variegato, inferne
venoso nigricante. Pers.

In insulis Mariannis.

Cette plante est Tune des plus jolies especes du
genre. Elle a par sa partie superieure quelque
similitude avec V auricularia mesenterica, Pers.
Mycol. Europ. 1, p. 97. Mais sa surface inferieure
est garnie de veines comme dans V auricularia sam-
buci ( tremella auricula, Linn.). P.

The illustration shows a dimidiate basidio¬
carp with prominent zonation on the dorsal
surface, more like a sketch of Coriolus versi¬
color than like any Auricularia, and certainly
not remotely like the almost resupinate fun¬
gus on Messerschmidia. But having soaked
a dried specimen to secure a spore-print, I
happened to pick away a few marginal bits
of the bark to which it was attached, in order
to discover how much of the dorsal surface
was free from the substratum. The surface
revealed was zonate not merely because
of the varying lengths of the hairs which
covered it, but "variegated by fuscous and
pallid bands,” exactly as described and illus¬
trated by Persoon. It seems likely that the
material from which the description was
drawn up either had been removed from its
substratum by the collector or had been pried
away, as was mine, to discover the character
of the dorsal surface. The resemblance noted
to Auricularia mesenterica may possibly ac-

Fungi of the Marshall Islands — Rogers

count for some of the reports of that species
from Pacific lands. Fries (1838: 555) lists
A. ornata as a possible synonym of A. mesen¬
teric a, with the note, "sec. Montagn. non
differt” (Montagne having, it would appear,
studied Persoon’s specimens). Material at
hand of A. mesenterica shows a much thicker
and more deeply pilose basidiocarp, with the
hyphae of the medullary layer much more
highly gelatinized and more widely spaced,
and with an ochraceous, rather than chiefly
brown, dorsal surface. The two species are
undoubtedly closely related; on the basis of
available specimens, I should consider them
amply distinct. There seem to be no zonate
species described in the genus other than
A. mesenterica and A. ornata; the latter is as
well characterized as any member of Auricu-
laria, and there can be little doubt that the
collections here cited belong to Persoon’s
species.

The fungus is quite as certainly Lyon’s
Auricularia adnata. The type of that species,
collected "on Tournejortia trees" on Pal¬
myra Island, 1,100 miles south of Oahu, is
not at hand; but Dr. H. L. Lyon has recently
confirmed Bergman’s identification of the
two later Palmyra collections (from the same
substratum), indicating that the specimen
now marked "A" is the more typical. These
are indistinguishable from the Marshall
Islands material. Other adnate species have
been described in Auricularia; of these only
A. peltata Lloyd seems well enough char¬
acterized to be recognizable without special
study. Material at hand of this species shows
uniformly smaller and more strongly venu-
lose basidiocarps attached over nearly all
the dorsal surface by the uniformly pallid
tomentum; it is quite a different fungus from
A. ornata. It is at least odd, if not significant
of relationship, that Lloyd’s type, as well as
one of the specimens more adequately de¬
scribed by Ahmad (1945: 242) grew on
Cordia "myxa” — like Messerschmidia, an
arborescent member of the Boraginaceae.

105

29. Septobasidium sp. (bogoriensi
affin. ) ; cf. Boedijn and Steinmann, Buiten-
zorg Jard. Bot. Jour. 3 ser. 11: 205, fig. 25,
26; pi. 18 (b), (c), 1931; Couch, Genus
Septobasidium 213, pi. 61, fig. 3-10; 103,
fig. 4-7; 104, fig. 10-12, 1938.

On living prop roots, and more rarely on
bark of living trunks, of Pandanus pulp os us
and Pandanus spp. Namu: Namu I., 1421,
Leuen I., 1417; Ailinglapalap, 1480; Jaluit,
1474, 1577, 1578; Ebon, IX.9.46, 1335,
1336, 1389 .

This fungus formed conspicuous lichen¬
like encrusting patches, light gray or lila-
ceous-gray, on a considerable part of the
Pandanus trees on the islands where it oc¬
curred; the quantity of material collected was
limited only by the time that could be profit¬
ably spent cutting it off the trunks, and the
space available for drying. Although super¬
ficially resembling the grayish lichens which
occurred abundantly in the same places, the
Septobasidium could readily be recognized
as a member of that genus by the irregular
discontinuous development of its margin, by
the evident elevation of its surface on short
brownish pillars (whereas the lichens clung
closely to the bark), and by the brown cen¬
ters of the older fructifications. It occurred
on all the islands visited south of Kwajalein;
whether it is found there also is not known,
since the two islets of Kwajalein Atoll on
which the party landed were almost com¬
pletely denuded of native vegetation. Al¬
though a careful search for the Septobasidium
was everywhere made, none was discovered
north of Kwajalein. On the drier atolls,
such as Ailuk and Wotje, where the lichen
incrustation was also sparse or wanting, this
lack was to be expected; but although on
Mej it the vegetation and lichens indicated a
climate fully as favorable for the growth of
fungi as that of Namu, the Septobasidium,
it seems safe to say, does not occur there.
Some reported hosts of S. bogoriense, such as
Citrus and Hibiscus, grow on islands where

106

Septobasidium was abundant on Pandanus,
but the fungus was found on none but the
latter plant. Parts of all collections have
been sent to Dr. J. N. Couch, who will dis¬
cuss the fungus in a future publication of
his.

30. Pellicularia isabellina (Fries)
Rogers, Farlowia 1: 99, 1943. Hypochnus
isabellinus (Fries) Fries; Burt, Mo. Bot.
Gard. Ann. 3: 222, fig. 12, 1916. Tomen-
tella isabellina (Fries) Hohn. and Litsch.;
Bourd. and Galz., Hym. Fr. 482, fig. 121
[1928]. Botryobasidium isabellinum (Fries)
Rogers, Iowa Univ. Studies in Nat. Hist. 17:
11, pi. 2, fig. 5, 1935.

On wood, husks, and dead leaf -sheaths of
Cocos nucijera, and on old polypore. Ailuk:
Ailuk I., 1738; Mejit, 1443, 1312; Likiep:
Eniarmij I., 1690; Namu: Leuen I., 1419;
Jaluit, 1604.

A delicate ochraceous tomentose growth,
under the lens minutely tufted, under the
microscope composed of stout, short-celled
hyphae branching at right angles, of rough-
walled subglobose spores, and of obpyriform
basidia. The mature spores are thin-walled
when abstricted; but in the Namu specimen,
and to a slighter extent in others also, the
wall soon becomes considerably thickened.
I have seen one other similar tropical col¬
lection.

31. Pellicularia lembospora Rogers,
Farlowia 1: 109, fig. 8, 1943.

On bark of rotten log of Ochrosia parvi-
flora. Likiep: Biebi I., 1672.

A cream-colored, very slight and loose,
hypochnoid growth; distinguishable under
the microscope by its small navicular spores
mostly 7-8 X 3 /x. The collection consists of
a few minute pallid patches of this, the per¬
fect stage, scattered among the much more
robust, tawny, imperfect fructification of
the same fungus. See under Oidium tomen-
tosum, below. Previously reported from
British Guiana and Cuba, and present also

PACIFIC SCIENCE, Vol. 1, April, 1947

in Hawaii; a tropical relative of the more
nearly ubiquitous P. vaga.

32. Pellicularia vaga (Berk, and
Curt.) Rogers, Farlowia 1: 110, fig. 9, 1943.
Corticium botryosum Bres.; Bourd. and
Galz., Hym. Fr., 241 [1928].

On bark, wood, and husks of Cocos nuci¬
jera, and wood of Pandanus sp., Artocarpus
incisus, and an undetermined dicot. Utirik,
1439, 1801; Ailuk: Ailuk I., 1703, 1746,
1737; Mejit, 1447, 1448, 1310, 1313, 1320;
Likiep: Likiep L, 1477; Wotje: Ormed I.,
1370, 1343; Jaluit, 1613.

A pallid or buff hypochnoid growth with
the characteristic very wide mycelium of the
genus and naviculiform spores (fusoid, de¬
pressed on one side) mostly 10-12 X 4.5 /x.
The commonest member of the genus in the
Marshalls, as on the northeastern and north¬
western coasts of the American mainland
and, apparently, in western Europe.

Fungi imperfecti

33. Helicomyces roseus Link; Linder,
Mo. Bot. Gard. Ann. 16: 271, pi. 12, fig.
5-7, 1929.

On husks of Cocos nucijera. Ebon,
IX. 10.46, 1803 (with 1338).

A grayish bloom, imperceptible except
under considerable magnification, composed
of hyaline linear conidia coiled like a watch-
spring, and very short hyaline conidiophores.
The fungus was discovered only when the
specimen earlier cited of Sebacina jar inace a
was being examined under the binocular. It
is less conspicuous macroscopically than the
quite inconspicuous Iowa specimens, deter¬
mined by Linder, with which it was com¬
pared, but agrees in all other respects.

34. Oidium tomentosum (Berk, and
Curt.) Linder, Lloydia 5: 204, pi. 5, fig. E,
1942.

On bark of rotten log of Ochrosia parvi-
jlora and unidentified dicot wood. Likiep:
Biebi I., 1672, 1674.

Fungi of the Marshall Islands — Rogers

107

Pilose, loose, reddish-tawny; composed,
as are all species of Pellicularia, of very
thick, short-celled hyphae branching at right
angles; fruiting by globose conidia borne on
short peg-like lateral teeth. As in Hawaiian
collections, this imperfect stage is much
better developed than the perfect stage,
Pellicularia lembospora. Reported from the
Bahamas and Cuba.

REFERENCES

Ahmad, Sultan. Higher fungi of the Panjab
plains: IV. Lloydia 8: 238-244, fig. 1-3, 1945.
Bourdot, Hubert, and Amedee Galzin. Hy-
menomycetes de France Heterobasidies-Homo-
basidies Gymnocarpes. iv + 761 p., 185 fig. Bry,
Sceaux [1928].

Ehrenberg, Christian Gottfried. Fungos a viro
clarissimo Adalberto de Chamisso, sub auspiciis
Romanzoffianis in itinere circa terrarum globum
collectos, enumeravit novosque descripsit et
pinxit. In Nees ab Esenbeck, C. G., Horae
physicae berolinenses, 77-104, pi. 17-20. Sump-
tibus Adolphi Marcus, Bonnae, 1820.

Fries, Elias Magnus. Systema mycologicum. vol.
1. lvii + [i] + 520 p. Officina Berlingiana,
Lundae, 1821. vol. 2. p. 1-274 [1822]; 275-620
+ [1]. Officina Berlingiana, Lundae, 1823.

- Systema orbis vegetabilis. Pars I. Plantae

homonemeae. viii + 374 p. Typographia Acad-
emica, Lundae, 1825.

- Epicrisis Systematis Mycologicae seu syn¬
opsis Hymenomycetum. xii + [ii] + 610 p.
Typographia Academica, Upsaliae, 1838.

Hennings, P. Einige Pilzarten von dem Mar-
shallinseln. {Berlin ] Bot. Gart. u. Mus. Notizbl.
1: 226-229, 1897.

Macbride, Thomas H., and G. W. Martin. The
Myxomycetes. viii -f 2 + 339 p., 21 pi. Mac¬
millan, New York, 1934.

McGuire, J. M. The species of Sebacina (Tre-
mellales) of temperate North America. Lloydia
4: 1-43, pi. 1-5, 1941.

Martin, G. W. New or noteworthy fungi from
Panama and Colombia: III. Mycologia 31:
239-249, 18 fig., 1939.

- The Tremellales of north central United

States and adjacent Canada. Iowa IJniv. Studies
in Nat. Hist. 18 (3) : 1-88, pi. 1-5, 1944.

Persoon, C. H. Mycologia Europaea. Sectio
prima. [2] + 356 p., 12 pi. Impensibus . . .
Palmii, Erlangae, 1822.

Ridgway, Robert. Color standards and color no¬
menclature. iv + 44 p., 53 pi. Published by the
author, Washington, D. C., 1912.

Rogers, Donald P. Notes on the lower Basidio-
mycetes. Iowa Univ. Studies in Nat. Hist. 17:
1-43, pi. 1-3, 1935.

Saccardo, Pier Andrea. Sylloge fungorum om¬
nium hucusque cognitorum. vol. 2. 815 +ii-
lxix + 77 p. Sumptibus auctoris, Patavii, 1883.

Schumann, Karl, and Karl Lauterbach. Die
Flora der deutschen Schutzgebiete in der Siidsee.
xvi + 613 p., 1 map, 22 pi. Gebr. Borntraeger,
Leipzig, 1901.

Stevenson, John A., and Edith K. Cash. The
new fungus names proposed by C. G. Lloyd.
Lloyd Libr. Mus. Bot., P harm, and Materia Med.
Bui. 35. 209 + [1] p. 1936.

Volkens, G. Die Flora der Marshallinseln. {Ber¬
lin'] Bot. Gart. u. Mus. Notizbl. 4: 83-91, 1903.

A Chloride and Oxygen Analysis Kit for Pond Waters1

R. B. Dean and R. L. Hawley2

INTRODUCTION

Fishponds in the Hawaiian Islands are
shallow brackish ponds which are frequently
fed by underwater springs as well as surface
flows of fresh water and sea water. The
chloride content of the water is an indication
of the quantity of sea water present and is
one factor which affects the growth and sur¬
vival of many of the plants and animals
utilized as food by the fish. The presence of
dissolved oxygen is absolutely essential for
animal life in the ponds. Low oxygen con¬
centrations in some regions indicate stagnant
waters and bottom putrefaction which make
these areas unfit for fish. In the course of
fisheries studies in the Hawaiian Islands it
was therefore necessary . to make a large
number of salinity and oxygen determina¬
tions on fishpond waters.

Since both chloride and oxygen may vary
severalfold in a single pond, and since most
living organisms are insensitive to concentra¬
tion changes which are less than 1 per cent
of the normal value, it is not necessary
to make extremely accurate determinations.
Methods were desired for the rapid analysis
of chloride and oxygen at the side of the
pond so that any unusual observations could
be checked before one left the pond.

The apparatus described in this paper was
designed to provide sufficient equipment and
reagents for 25 chloride, and 50 dissolved

1 Research Paper 5, Cooperative Fisheries Re¬
search Staff of the Territorial Board of Agriculture
and Forestry and the University of Hawaii.

2 Department of Chemistry, University of Ha¬
waii. Dr. Dean is now at the University of Ore¬
gon; Mr. Hawley is a student at George Washing¬
ton University Medical School. Manuscript re¬
ceived December 2, 1946.

oxygen determinations. Two auxiliary half¬
liter bottles contain enough reagents for
an additional 50 chloride determinations.
Chloride up to 25 gm. per liter (sea water
is about 20 gm. per liter) can be determined
with an accuracy of 0.06 gm. of Cl per liter.
Dissolved oxygen up to 20 cc. per liter can
be determined with an accuracy of 0.1 cc.
per liter on a sample of 10 cc. of water. The
apparatus weighs about 25 lb. and can be
used in a rowboat, if necessary. A single
chloride determination requires about 3
minutes; an oxygen determination requires
about 8 minutes.

Although there are several well-known
titrimetric methods for chlorides, none of
them is suitable for use on micro-samples
out-of-doors where it is difficult to observe
a colorimetric end point. An electrometric
method of detecting the end point which
substituted a galvanometer needle for the
color change appeared to be more suitable
for our purposes. None of the published
electrometric methods was quite satisfactory
and an entirely new method was developed
which will be dealt with in another publi¬
cation (Dean and Hawley: unpublished).
This method makes use of two wire elec¬
trodes, a galvanometer, a few radio re¬
sistances, and a dry cell. A relatively low
resistance circuit is formed which is not
sensitive to high humidities, although it
should not be soaked in water.

Krogh (1935: 131-133) has described a
modification of the Winkler method for dis¬
solved oxygen which was well suited to our
purpose, with slight modifications. Krogh
used the conventional starch indicator to de¬
tect the end point. We have substituted

108

Chloride and Oxygen Analysis Kit — Dean and Hawley

109

Foulke’s dead-stop electrometric end point
(Foulke and Bawden, 1926: 2045 ff.), be¬
cause the starch iodine indicator is unreliable
above 25° C. and our field temperatures fre¬
quently exceed 30° C. Essentially the same
electrical circuit is used for both the oxygen
and the chloride end points: only the elec¬
trodes are different.

The unknown solutions are titrated with
standard solutions from a micrometer syringe
burette (see Trevor, 1925: 1111; Dean and
Fetcher, 1942: 237; and Dean: unpub¬
lished). This burette has many advantages
over conventional gravity- feed burettes. It is
more compact, there is no drop error and no
parallax error, and better than 0.1 per cent
accuracy is possible on a total volume of
only 1 cc. of reagent. Micro methods are
obviously necessary to conserve reagents if a
small apparatus is to carry enough to permit
50 or more determinations. Two syringes
are used interchangeably with the same
micrometer head in this apparatus, so that it
is not necessary to clean and refill the burette
when changing from chlorine to oxygen
analyses.

DESCRIPTION OF THE APPARATUS

All of the equipment is contained in a
wooden box which measures 9X9X12
inches (see Fig. 1). The electrical wiring is
enclosed in a mahogany case which is firmly
attached to the right-hand side of the box.
The galvanometer is a needle type of instru¬
ment which has a sensitivity of 0.25 micro¬
amperes per scale division and a critical
damping resistance of 1,800 ohms.

The electrical circuit is shown schemati¬
cally in Figure 2. The two switches Sx and S2
are combined in a telephone type of toggle
switch. In the central position both switches
are open and no current flows. When the
switch is moved to either side, current flows
from the battery through the series of re¬
sistors in the upper line. Suitable taps take
off about 10,200 and 1,018 mv. to the Pt,

Ag, and Standard Cell terminals respec¬
tively. Variations in the internal resistance
of the dry cell can be compensated for by
adjusting resistor R2 when the switch is
thrown to the left. In this position the volt¬
age of the standard cell opposes a fraction
of the voltage from the battery. Resistor B
is adjusted until no current flows through
the galvanometer. In actual practice it has
been found that the voltage from a flashlight
dry cell does not change significantly, even
when the circuit is left open for 2 days. It
would have been satisfactory to omit the
standard cell and variable resistor B entirely.
The potential at the chloride electrodes could
be checked before each series of measure¬
ments, as is described below.

The electrodes are constructed as shown
in the insert of Figure 2. Short pieces of
silver or platinum wire are attached to stiff
bronze wire with hard solder. The wire elec¬
trode is then sealed in a straight glass tube.
It is not difficult to seal platinum into soft
glass. Silver wire of 22 gage can also be
sealed into soft glass but some of the seals
break soon after sealing. After the glass is
sealed onto the electrode wire, both the glass
tubing and the bronze wire can be bent in
the ordinary manner, since bronze wire
softens at red heat.

The stiff bronze wire is passed through a
hole drilled in a small radio jack plug as
indicated in the diagram. The four electrode
leads — Ag, Cu, and two Pt — are brought to
jacks on a mounting plate at the left end of
the box under the end of the burette. When
either pair of electrodes is plugged into its
corresponding jacks, the electrode wires just
reach the center of the titration vessel. The
Ag and Cu jacks are further differentiated
by color to reduce the danger of inserting an
electrode in the wrong terminal.

The micrometer-syringe burette consists
of a 1.5 cc. glass hypodermic syringe fitted
in a frame to which is attached a 1-inch
machinist's micrometer graduated in 1,000
parts. The burette assembly is described fur-

110

PACIFIC SCIENCE, Vol. 1, April, 1947

Fig. 1. Analysis kit for chloride and oxygen. Chloride reagents and syringes to the right; oxygen
reagents and syringe to the left. E, electrode holder. The chloride electrodes are just above the slide
rule. P, micrometer burette with syringe to hold sodium thiosulfate reagent. The silver nitrate syringe
is above chloride electrodes. N, galvanometer. Z, toggle switch. A, standard cell. T, rubber bulb ia
bottle of distilled water for washing. The syringes and slide rule fit in clips inside the lid of the box_
Most of the equipment remains in the box during the analysis.

Chloride and Oxygen Analysis Kit — Dean and Hawley

111

ther in Dean and Fetcher (1942: 237; see
also Dean: unpublished). For the oxygen
determination a solution of thiosulfate is
used in the burette and a 2 -inch 2 2 -gage
hypodermic needle bent downward at 90°
serves as the delivery tip. The chloride de¬
termination requires a strong solution of
silver nitrate, which corrodes all available
metals, so that a glass tip must be used. In
this apparatus we used a bent glass tube
attached to the burette by a number 0 one-
hole rubber stopper.3

3 The Ace Glass Company, Vineland, N. J., is
now able to supply ground glass tips to order
which will fit on a standard hypodermic syringe.

The titration vessel is a shortened test tube
about 25 mm. in diameter and 70 mm. long.
It fits in a wire test tube holder which is
attached to the electrode jack assembly. The
burette tip and the two electrodes dip into
the solution in the titration vessel. In addi¬
tion there is a glass tube to introduce air
bubbles for stirring. Air is forced by an
aspirator bulb into a 1 -liter can which
serves as an air reservoir, and flows from
there through a thin rubber tube to the glass
nozzle of the air stirrer. The aspirator bulb
can be conveniently squeezed by the left
hand of the operator during a titration.

A reagent shelf over the air reservoir

H lo.ooo |—

12.00

MICRO AMPS

1 - I 10,000 1 - 1

P^standardA

S2

V^/ CELL

DETAIL OF
ELECTRODES

Bronze
wire in
glass

Radio

jack

|Pt or Ag
wire

Fig. 2. Wiring diagram. Si and S2 are two parts of a double-pole, double-throw telephone-type
toggle switch. Wires run from the Pt, Ag, and Cu terminals to the electrode holder.

112

PACIFIC SCIENCE, Vol. 1, April, 1947

holds five 60-cc. screw top bottles and two
10-cc. screw top vials. In addition, there are
several larger reagent bottles in the bottom
of the box and a bottle of distilled water for
washing electrodes and glassware. A glass
tumbler is fitted under the electrodes for a
waste jar. A rubber ear syringe is fitted with
a glass delivery tip bent at an angle of 135°.
This syringe can be filled from the distilled
water bottle and the water in it then directed
in a stream into the inverted titration vessel
or onto the electrodes to clean them.

The reagents are all contained in screw
cap bottles or vials. The screw caps are lined
with Polythene, which is resistant to all the
reagents used in this work. Two of the vials
are fitted with rubber disks from 10-ml,
penicillin bottles. The disks are held on by
screw caps which have been drilled with
14 -inch holes. The rubber disks on the vials
can be repeatedly perforated by a hypo¬
dermic needle and the reagents withdrawn
through the needle.

All the reagents and equipment used for
the chloride analysis are marked with black
paint and the various reagents are identified
by white letters on the top, as well as by con¬
ventional labels on the sides of the bottles.
The reagents and apparatus for the oxygen
determination are likewise marked with red
paint and white letters. The metal parts of
the syringe pipettes and the burette are of
brass which was first dulled by a dip in silver
nitrate and nitric acid, and then coated with
Glyptal varnish which was baked on. This
treatment produces a non-glaring, corrosion-
resistant surface. The syringes are attached
by clips to the inside of the top of the box.
A small box of filter paper squares, about 3
cm. on a side, which are used to wipe the
burette tips and the electrodes, is also at¬
tached inside the top of the box. A rubber
stopper is screwed inside the top at the right-
hand side in such a way that the lid will not
close until the switch has been turned off.
This arrangement insures that the battery
will not be left on when the box is closed.

CHLORIDE ANALYSES

The chloride analysis depends upon the
fact that the potential of a silver electrode
changes rapidly when all the chloride ions
have been precipitated by silver ions. Since
it is impossible to measure a single electrode
potential, another electrode which is not
sensitive to chloride ions must be present to
complete the circuit. We have used a copper
electrode in the presence of nearly saturated
copper sulfate. The potential between these
two electrodes is about 200 mv. at the end
point. The potentiometer circuit supplies
200 mv. between the Cu and Ag electrodes
and, at the end point, no current will flow
through the galvanometer.

Before the end point is reached the poten¬
tial between the electrodes will be less than
200 mv. and the galvanometer will be de¬
flected to the right. There is a large protec¬
tive resistance in series with the chloride
electrodes so that the galvanometer may be
left continuously in circuit. As the end point
is approached the galvanometer needle ap¬
proaches £ero and is deflected to the left as
soon as the end point has been exceeded.
The deflection produced by the addition of
one or two burette divisions of silver nitrate
is greatest at the end point. This fact can be
used to set the potentiometer to the cor¬
rect potential. The potentiometer is adjusted
until the galvanometer indicates zero. The
galvanometer deflection is then noted after
the addition of two burette units of silver
nitrate. This procedure is repeated until the
galvanometer deflection is a maximum. The
potentiometer is left at this setting for sub¬
sequent titrations.

The silver nitrate reagent contains 4 per
cent silver nitrate and 0.5 per cent nitric
acid. The burette is filled by bringing the
reagent bottle of silver nitrate, marked with
a T on a black screw cap, up under the
burette tip. The plunger of the syringe is
withdrawn by unscrewing the micrometer,
and silver nitrate is sucked into the syringe.

Chloride and Oxygen Analysis Kit — Dean and Hawley

Any air bubbles are displaced by rotating the
burette in its clamp until the tip is upwards
and then forcing a little of the silver nitrate
reagent out of the tip. The burette is filled
up to the 1,000 mark on the micrometer
and the tip rinsed with water.

Ten cc. of saturated copper sulfate is
placed in the titration vessel; a syringe with
tip broken off is used to make the transfer.
Commercial bluestone can be used for this
solution, although it may introduce an un¬
desirably high blank. Reagent grade copper
sulfate is perfectly satisfactory. The copper
sulfate solution is placed under the elec¬
trodes and the air stirrer is started. A small
residual concentration of chloride ions may
produce a galvanometer deflection to the
right. If it does, silver nitrate is added until
the galvanometer reads zero. The burette
reading is recorded. The chloride sample,
about 0.4 cc., is then introduced from a
precision syringe pipette (Krogh, 1935:
130; Dean: unpublished). Silver nitrate is
added until the galvanometer again indicates
zero and the second burette reading is
recorded.

Neither the exact volume relations of the
syringes nor the concentration of the silver
nitrate need be known exactly. All these are
evaluated as a single factor by titrating the
same volume of a known standard solution
of sodium chloride which contains exactly
20 gm. of chloride ions per liter.

When the galvanometer has been brought
to zero a second time and the burette read¬
ing has been recorded, the air stirrer is re¬
moved and wiped. The burette is tipped up
and the tip is wiped and the burette is re¬
filled. The electrodes and the burette tip are
rinsed and the apparatus is ready for the
next determination.

The errors are of the order of 1 burette
division or 1 part in 800. Greater precision
could be obtained by precipitating most of
the chloride in a larger sample with silver
nitrate from another pipette. A more dilute

113

solution of silver nitrate could then be used
in the burette. It might be advisable to carry
out such determinations in more dilute solu¬
tions to avoid too much clumping of the
silver chloride precipitate. The accuracy is
limited by the reproducibility of the syringe
pipettes. Krogh (1935: 130) reports an
accuracy of 1 part in 10,000 and we have
several syringes with an accuracy of 2 parts
in 10,000.

OXYGEN ANALYSES

Oxygen is determined by its reaction with
a solution of manganous hydroxide to form
manganic oxides. When all the oxygen has
been absorbed, the solution is made acid
and the manganic ions react with iodide ions
to liberate free iodine. The iodine is titrated
with sodium thiosulfate in the presence of
two bright platinum electrodes. A potential
of 10 mv. is applied between these electrodes
and a current will flow as long as there is
iodine in the solution to remove electrons
from the negative electrode. As soon as all
the iodine has been removed the current
ceases to flow, except for a very small resi¬
dual current. As the end point is approached,
the galvanometer deflection decreases from
the left and the end point is taken when the
galvanometer indicates one unit deflection to
the left.

A 10-cc. syringe pipette (Krogh, 1935:
132; Dean: unpublished) is fitted with a
2-inch 18-gage stainless steel needle. An
aluminum cam cemented to the plunger with
Varno cement will engage a stop on the
side of the syringe holder when the syringe
holds 10 cc. An additional 0.2 cc. can be
introduced by rotating the cam away from
the side stop and pulling back until the
plunger reaches an end stop.

The syringe is first rinsed with Solution I.
This solution is made up by dissolving 90
gm. of Nal and 40 gm. of NaOH in 55 cc.
of water (Pomeroy and Kirschman, 1945:
716) and all air bubbles are expelled. This

114

PACIFIC SCIENCE, Vol. 1, April, 1947

reagent is kept in a small rubber-capped vial
marked on each side with one white dot.
The needle of the syringe is inserted into
the vial and the vial is inverted while the
solution is drawn in and the air bubbles are
expelled. This leaves about 0.1 ml. of solu¬
tion in the dead space of the syringe. The
water sample is then drawn into the syringe
until the cam on the plunger reaches the side
stop. The needle is then inserted into a
second vial, marked with two white dots,
which contains Solution II. This solution is
made from 40 gm. of MnCl2, 10 cc. of
6N HC1, and enough water to make 100 cc.
This solution is drawn into the syringe by
moving the plunger from the side to the end
stop.

The manganous chloride reacts inside the
syringe with the sodium hydroxide which
was left in the dead space, to produce a light
fluffy precipitate of manganous hydroxide.
This precipitate absorbs oxygen, and in 4
minutes substantially all of the oxygen will
have been absorbed. The contents of the
syringe are then discharged below the sur¬
face of 1 cc. of 6N HC1 in the titration
vessel. The syringe is rinsed first with the
acid solution to dissolve any manganous
hydroxide remaining in the syringe, and then
with two small portions of water from a
spare titration vessel. The 6N HC1 is con¬
tained in a 60-cc. bottle fitted with a rubber
medicine dropper. One dropper full is about
1 cc.

The titration vessel now contains iodine
equivalent to the oxygen which was present
in the water. The iodine is titrated with a
sodium thiosulfate solution. This solution
must be made up fresh every week by dilut¬
ing to 60 cc. one medicine dropper full of
a 60 per cent solution of sodium thiosulfate
containing 1 per cent borax. This concen¬
trated solution is quite stable. Most of the
iodine is discharged with thiosulfate from
the burette before the air stirrer is started,

because the air might otherwise remove some
of the iodine.

The thiosulfate solution and the volume¬
tric apparatus are standardized by the use of
a standard solution of KI03. This solution
is 0.0004167 molar and is equivalent to
14 cc. (STP) of oxygen per liter. The dead
space of the syringe is first filled with Solu¬
tion I. The syringe is then rinsed with acid
contained in the titration vessel and filled
with the standard iodate solution. The iodate
reacts with the iodide and acid to liberate
iodine in the syringe. The iodine is titrated
with thiosulfate, and the burette difference
which is found corresponds to 14 cc. of
oxygen per liter. Blank runs on water which
had been freed of oxygen by hydrogen and
platinized asbestos showed that the end
point and other errors are less than 0.1 cc.
of oxygen per liter.

The oxygen concentration is calculated in
essentially the same manner as that used for
the chloride. The initial burette reading is
always taken as 1,000. This procedure in¬
troduces a small constant error of the order
of magnitude of 0.1 cc. of dissolved oxygen
per liter.

SUMMARY

A portable apparatus which is equipped
for the determination of chlorides and dis¬
solved oxygen in pond waters is described.
The chloride is determined by a new elec¬
trometric method. Oxygen is determined by
a modified Winkler method and the iodi-
metric end point is detected electrometrically.
The volumetric apparatus consists of preci¬
sion syringe pipettes and a micrometer
burette. Up to 25 gm. of Cl per liter can be
determined with an accuracy of 0.0 6 gm.
per liter. Up to 20 cc. of dissolved oxygen
per liter can be determined with an accuracy
of 0.1 cc. per liter. Greater precision over a
smaller range of chloride concentration is
possible, since the syringe pipettes can attain
an absolute accuracy of one part in 10,000.

Chloride and Oxygen Analysis Kit — Dean and Hawley

REFERENCES

115

Dean, R. B. Improved syringe pipet and microm¬
eter burette (unpublished).

- , and E. S. Fetcher. Micrometer burette.

Science 96: 237-238, 1942.

- , and R. L. Hawley. A new rapid electro¬
metric method for the analyses of chlorides (un¬
published).

Foulke, C. W., and A. T. Bawden. A new type
of- end-point in electrometric titration and its
application to iodimetry. Amer. Chem. Soc.
Jour. 48: 2045-2051, 1926.

Krogh, August. Syringe pipets. Indus, and Engin.
Chem., Analyt. Ed. 7: 130-131, 1935.

- Precise determination of oxygen in water

by syringe pipets. Indus, and Engin. Chem.,
Analyt. Ed. 1: 131-133, 1935.

Pomeroy, Richard, and H. D. Kirschman. De¬
termination of dissolved oxygen, proposed modi¬
fication of the Winkler method. Indus, and
Engin. Chem., Analyt. Ed. 17: 715-716, 1945.
Trevor, J. W. The micrometer syringe. Bio chem.
Jour. 19: 1111-1114, 1925.

A New Species of Carex (Cyperaceae) from Fiji:
Pacific Plant Studies 61

Harold St. John2

INTRODUCTION

Carex is the largest genus of the
Cyperaceae, having well over 1,000 species.
This genus of sedges is abundantly repre¬
sented in the Arctic and Temperate Zones
of the Northern Hemisphere. In the Tropics
there are but few species. The genus ex¬
tends to the continents of the Southern
Hemisphere, but the representation there is
meager. In the tropical Pacific, some islands
support a very few species, while the others
entirely lack the genus. In view of this dis¬
tribution, it is of interest to announce a
newly discovered species from Fiji.

A NEW FIJIAN CAREX

Carex vitiensis St. John, sp. nov. Fig. 1.

Rhizomatis breve caespitosis lignosis, culmis
9-10 dm. altis erectis nudis trigonis glabris striatis,
laminis multis 7-8 dm. longis 4-6.5 mm. latis pal-
lide viridibus longe acuminatis glabris striatis mar-
ginibus scabris, vaginis brunneis omnibus lamini-
feris, inflorescentiis 45-55 cm. longis 1-1.5 cm.
latis interrupts, nodis 6-8, bracteis foliaceis ocreis
anguste cylindricis inferis 4-6 cm. longis, laminis
inferis 3-4.5 dm. longis, nodis 5-l4-spiculiferis,
spiculis adscendentibus androgynis lateralibus 2-
6.5 cm. longis 1.5-3 mm. latis, pedunculis 1-11
cm. longis filiformibus scaberulis, floribus masculis
paucis ad apicem spicorum, squamis foemineis
2. 2-3. 2 mm. longis obovatis nervosis glabris costis

1 This is the sixth in a series of papers designed
to present .descriptions, revisions, and records of
Pacific island plants. The preceding papers were
published as Bernice P. Bishop Museum Occa¬
sional Papers: 17 (7), 1942; 17 (13), 1943; 18
(5), 1945; Amer. Fern Jour. 35: 87-89, 1945;
Torrey Bot. Club Bui. 73: 588, 1946.

2 Chairman, Department of Botany, University
of Hawaii. Manuscript received November 13,
1946.

viridibus marginibus hyalinis albidis apice mucro-
niferis, utriculis 4. 8-5. 2 mm. longis 1-1.2 mm.
latis trigonis breve hirsutulis viridescentibus lateris
plerumque valde trinervosis corporibus anguste
fusiformibus, stigmatis 3, stylo incluso, achaeneis
2.7-3 mm. longis lucidis stramineis trigonis cor¬
poribus anguste ellipticis lateribus canaliculatis ad
basim cuneatis ad apicem et basim styli durescen-
tibus contractis.

Forming dense clumps; rhizome short, densely
caespitose, horizontal or descending, woody; culms
9-10 dm. tall, 1.5-2 mm. in diameter, central,
erect, naked, sharply trigonous, glabrous, striate,
greenish; leaves numerous, shorter than the culms,
7-8 dm. long, 4-6.5 mm. wide, flat, pale green,
long tapering, glabrous, finely striate nerved, the
margins scabrous; leaf sheaths brown, prominent,
all leaf-bearing; inflorescence 45-55 cm. long,
1-1.5 cm. wide, elongate, interrupted, the nodes
6-8; the bracts with the base green, closely sheath¬
ing, long cylindric, the lowest one 4-6 cm. long,
the others shorter, the blades foliaceous, exceeding
their spikes, the lowest one 3-4.5 dm. long; spikes
5-14 at each node, slender peduncled, ascending
or the tips slightly diverging; peduncles 1-11 cm.
long, filiform, scaberulous; spikes androgynous,
the lateral ones 2-6.5 cm. long, with a short stami-
nate apex, the terminal one 2.5-5 cm. long with
one or a few pistillate flowers at base; pistillate
portion of spikes 1.5-3 mm. in diameter, slender
cylindric, loosely flowered, the ascending perigynia
partly imbricate; pistillate scales 2. 2-3. 2 mm. long,
obovate, the back green, the margins hyaline and
whitish, nerves numerous, parallel, close, glabrous
except for the midrib excurrent into a scabrous
awn to !/4 the length of the body; staminate
scales similar but somewhat narrower and straw-
colored; perigynia 4. 8-5. 2 mm. long, 1—1.2 mm.
wide, trigonous, short white hirsutulous through¬
out, especially on the angles and beak, greenish,
strongly nerved, usually with 3 nerves on each side
and with marginal nerves, the body slender fusi¬
form, tapering into a slender, rigidly bidentate
beak nearly as long, the body with the two outer
faces nearly plane, the inner side channeled; stig¬
mas 3; style included; achene 2.7-3 mm. long,
smooth, straw-colored, sharply trigonous, with
wing-like angles and concave faces, the body nar¬
rowly elliptic, tapering to the 0.3 mm. stipitate

116

New Species of Carex (Cyperaceae) from Fiji — St. John 117

Fig. 1. a, habit of type specimen, St. John 18,330 (X Vi)\ b, spike showing pistillate flowers below,
and staminate scales above (X 3); c, perigynium and stigmas (X 10); d, lateral view of achene
(X 10); e, transverse median section of achene (X 10); f, pistillate scale (X 10).

118

PACIFIC SCIENCE, Vol. 1, April, 1947

base, contracted to the indurated cylindric per¬
sistent 0.2 mm. style base which is straight or
slightly twisted.

Fiji islands; Viti Levu, Taunaisali, Wai-
nisavulevu-Nubulolo divide, the central pla¬
teau between the Wainimala and Singatoka
Rivers, clumps in swampy rain forest, 3,800
feet altitude, August 18, 1937, H. St. John
18,330 (type in Bishop Mus.).

There have been only three species of
Car ex known in the Fijian flora and none of
them appears to be closely related to this
new species.

The new C. vitiensis is a member of the
subgenus Eucarex and apparently is to be
placed within the ample limits of section
Elatae. No very close relative is known, but
it appears to be somewhat remotely related
to C. longebrachiata Boeck. (C. longijolm
R. Br., not of Thuill. or Host) of eastern
Australia. C. longebrachiata Boeck. has the
leaves exceeding the culms, coriaceous; the
spikes androgynous or gynecandrous or rarely
unisexual, the upper 1-2 strictly staminate,

the others pistillate, spikes rather densely
flowered, narrowly cylindric, often pendu¬
lous; the perigynium 6 mm. long, long at¬
tenuate to the base, broadest near the middle,
the beak about 1/4 of the total length; and
the achene obovate above the stipitate base,
densely punctate, the indurated base of the
style contorted. In contrast, the new C. viti¬
ensis has the leaves about 4/5 the length
of the culms, chartaceous; the spikes an¬
drogynous, only the terminal ones largely
staminate; spikes loosely flowered, very slen¬
der cylindric, erect or the tips slightly diverg¬
ing; the perigynium 4. 8-5. 2 mm. long, short
attenuate to the base, broadest 1/3 of the
way from the base, the beak about 1/3 of
the total length; and the achene narrowly
elliptic above the stipitate base, smooth and
shining, the indurated base of the style
straight or slightly twisted.

The specific name is derived from the
name of the island, Viti Levu, where the
plant grows.

NOTES

Facilities for Research in the Natural Sciences in the Hawaiian Islands

More than twenty agencies or institutions,
both governmental and private, possess facilities
for research in the natural sciences in the Ha¬
waiian Islands. The following inventory of these
facilities has been prepared from responses by the
heads of the various organizations. It is believed
that this alphabetic listing will be valuable not
only to scientists visiting Hawaii or corresponding
with agencies in this region, but also to residents
of these islands who have not previously been
able to examine a complete list of local research
facilities and opportunities. Similar inventories of
research facilities in the natural sciences in other
areas of the Pacific region are planned for future
issues of Pacific Science.

Bernice P. Bishop Museum

address: Bernice P. Bishop Museum, Honolulu
35, Hawaii. Director: Dr. Peter H. Buck.
purpose: Collection, preservation, and study of
Hawaiian and kindred Pacific material in eth¬
nology and the natural sciences; publication of
results of study. (Only the natural sciences
will be covered in the present listing. ) The
Museum is affiliated with Yale University, and
the Director is a professor on the Yale faculty.
subdivisions: Departments of Botany, Mala¬
cology (terrestrial and marine). Entomology,
and Marine Zoology; also large collections in
ichthyology, ornithology, and geology.
persons engaged in research: (botany) Dr.
Harold St. John, Marie Neal, Edward Y.
Hosaka, Dr. F. B. H. Brown (unofficially
attached) ; (malacology) Dr. C. Montague
Cooke, Jr., Yoshio Kondo, Wray Harris
(marine) ; (entomology) E. C. Zimmerman;
(marine zoology) Dr. C. H. Edmondson;
(ornithology) Paul H. Baldwin (absent on
leave). The Museum also has on its staff four
honorary consultants and 1 1 honorary associates
in the natural sciences.

opportunities for field research: The Mu¬
seum budget includes appropriations for field
expenses. Trips among the Hawaiian Islands
and special expeditions to other parts of the
Pacific area are carried out from time to time.
library: The Library has an estimated 25,015
books and 11,580 pamphlets. These are chiefly
on Pacific ethnology and natural history, with
emphasis on Polynesia. No important report
of an early voyage into the Pacific is lacking.

There are complete files of many scientific
publications from American institutions, as well
as excellent files from institutions in all other
parts of the world having interest in the Pacific.
New books on pertinent subjects are acquired
as they appear, and the Library attempts to
fill the needs of the members of the staff.

collections: The Museum is designated by
territorial law as the depository for natural his¬
tory collections of the University of Hawaii
and other territorial departments, bureaus, and
boards. In addition to a large area devoted to
ethnology (the Museum has the largest col¬
lection of Hawaiian artifacts in the world),
the following collections are available in the
natural sciences: (1 ) Botany. Herbarium con¬
tains what is probably the largest collection of
Polynesian plants; the total number for the
Pacific area is 150,000 specimens. (2) Mala¬
cology: Synoptic collection of land and ma¬
rine shells on exhibition; land shell collection
is well over 2,000,000 specimens and marine
shell collection over 50,000 (3) Entomology:
Collection numbers over 400,000 specimens.
(4) Ornithology: Contains many specimens of
extinct Hawaiian birds. (5) Ichthyology: Large
collection of fish; also fine set of colored casts
on exhibition.

publications: Four series are published by the
Museum. (1) Memoirs (quarto) ; 12 volumes
containing 39 papers published to date. (2) Bul¬
letins (royal octavo, over 50 printed pages);
188 published to date. (3) Occasional Papers
(octavo, under 50 printed pages); 18 volumes,
totaling 274 papers, published to date. (4) Spe¬
cial Publications; 37 published to date. List of
publications may be obtained from the Director.

RESEARCH fellowships: Two Yale University-
Bishop Museum Fellowships of $2,000 were
offered annually by Yale University and Bishop
Museum for research work on ethnology and
the natural sciences of the Pacific area. They
were discontinued during the war but will be
resumed in the near future.

research opportunities: Other opportunities
are offered by the Museum from time to time
as the need arises and the funds permit. Ma¬
terials in botany, entomology, and marine
zoology are sent out to specialists in America
and Europe for identification, and the reports
are published by the Museum.

119

120

PACIFIC SCIENCE, Vol. 1, April, 1947

research policy: The Museum offers research
facilities to visiting scientists to study its collec¬
tions. Office space is provided. The Museum
recently provided desk accommodation for 10
scientists and 20 typists who worked on the
reports of the United States Commercial Com¬
pany’s economic survey of Micronesia. During
the proposed Pacific Science Survey, the Mu¬
seum is prepared to give office accommoda¬
tion to research workers in ethnology and the
natural sciences, as well as free access to its
collections.

California Packing Corporation

address: California Packing Corporation, P. O.

Box 149, Honolulu 10, Hawaii.
purpose: Since the corporation is primarily a
production organization, research facilities are
confined to those necessary to growing and
canning fruits and vegetables. Research in agri¬
culture is carried on by Maxwell O. Johnson.
The corporation is also served by its main office
in San Francisco and by the Pineapple Research
Institute.

facilities: Laboratory facilities in Honolulu are
limited to those required for simple analysis
and control of canning operations.

Hawaii National Park

address: U. S. Department of the Interior, Na¬
tional Park Service, Hawaii National Park,
Hawaii. Superintendent: Frank R. Oberhansley.
purpose: The Park Service is primarily an op¬
erating organization; research by the field staff
is encouraged when time permits.
special subdivisions: Naturalist Department;
Hawaiian Volcano Observatory (for descrip¬
tion of this Observatory see listing below under
"Hawaiian Volcano Observatory”).

PERSONS ENGAGED IN RESEARCH: G. O. Fager-
lund, botany; Clifton J. Davis, entomology.
facilities: Offices, laboratories, and photographic
darkroom.

opportunities for field research: Year-
round access to area of contrasted physio¬
graphic, climatic, and ecological conditions.
library: A few hundred volumes on biology.
collections: Fairly complete classified collec¬
tion of plants, birds, and insects of the Park
area.

publication series: Mimeographed Natural
History Bulletins interpreting natural condi¬
tions, published occasionally.
research policy: The Service co-operates in
every way within its means to encourage in¬
stitutions and individuals to conduct research
in its areas.

Hawaiian Pineapple Company

address: (Dole) Hawaiian Pineapple Company,
Ltd., Honolulu 1, Hawaii.

purpose: Research pertaining to the growing
and processing of pineapples and other sub¬
tropical crops.

staff: Technically trained persons engaged in
research number 25. Staff includes Dr. F. P.
Mehrlich, Assistant Vice President in Charge
of Research; Dr. George E. Felton, Chemist;
Dr. R. O. Belkengren, Food Biochemist; Dr.
Dillon S. Brown, Horticulturist; Dr. Melvin
Levine, Bacteriologist; Kenneth Kopf, Geneticist.
facilities: Laboratories in Honolulu for chem¬
ical research, frozen food research, food tech¬
nology, bacteriology, and plant physiology. Lab¬
oratory at Wahiawa for horticultural, genetic,
agronomic, and plant physiology research. A
similar laboratory on Lanai. Pilot plants and
special processing lines of commercial equip¬
ment are available in the Honolulu factory.
opportunities for field research: Compre¬
hensive field research is carried on relative to
agronomy, horticulture, genetics, and plant
physiology.

library: The library contains about 500 volumes.

Hawaiian Sugar Planters’ Association

address: Experiment Station, Hawaiian Sugar
Planters’ Association, 1527 Keeaumoku Street,
Honolulu 4, Hawaii. Director: Dr. Harold L.
Lyon.

purpose: To investigate and solve the field and
factory problems of the Hawaiian sugar in¬
dustry.

subdivisions: Departments of Agriculture, Bo¬
tany and Forestry, Chemistry, Climatology,
Entomology, Genetics, Geology, Pathology,
Physiology and Biochemistry, and Sugar Tech¬
nology. Each department is headed by an out¬
standing scientist.

persons engaged in research: About 65
research workers.

facilities: Each department has at its disposal
all the facilities and equipment necessary for
research in its special field.
library: The library contains 25,860 volumes and
thousands of separates, bulletins, and pamph¬
lets, all properly classified.
collections: A very extensive collection of in¬
sects of the Pacific ocean area; museums of
sugar canes and cane diseases.
publications: The Hawaiian Planters’ Record , a
magazine now in its fiftieth volume. Members
of the organization are encouraged to publish
results of research in various scientific journals.
research fellowships: None offered at present.
research policy: The Station always welcomes
visiting scientists and strives to assist them in
every way possible. During the year 1946, the
Station gave assistance in the field, laboratory,
and library to sugar cane experts from Aus¬
tralia, China, Cuba, Mauritius, and Tanganyika.

NOTES

121

Hawaiian Tuna Packers
address: Hawaiian Tuna Packers, Ltd., P. O.

Box 238, Honolulu, Hawaii.
facilities: At the present time the company
does not have facilities for scientific research,
as this organization, primarily engaged in pro¬
cessing, is only slowly getting its activities back
to a prewar basis of operation, and hopes to
reach normal production by the latter part of
1947.

Hawaiian Volcano Observatory
(Branch of Hawaii National Park collaborating
with Hawaiian Volcano Research Association and
the University of Hawaii)

address: Hawaiian Volcano Observatory, Ha¬
waii National Park, Hawaii. Volcanologist in
charge: R. H. Finch.

purpose: To maintain measurements and obser¬
vations on the active volcanoes Kilauea and
Mauna Loa and to conduct research on the
physics and chemistry of volcanoes by seismic,
magnetic, or other available methods and to
devise and test new methods.
persons engaged in research: R. H. Finch,
Volcanologist; Dr. H. A. Powers, Seismologist;
B. J. Loucks, Instrument Maker; occasional
visiting specialists.

facilities : ( 1 ) Hawaii N ational Park N aturalist-
Observatory Building (laboratories, seismo¬
graph vaults, instrument shop, darkroom).
(2) University of Hawaii Kilauea Laboratory
(separate laboratory and office buildings, mis¬
cellaneous instruments, collections, records,
etc.). (3) Seismograph stations at other points
on the island of Hawaii.

opportunities for field research: Proximity
to Kilauea Crater and accessibility of Mauna
Loa. Opportunity for studies in petrology, areal
geology, volcanic processes, and varied geo¬
physical measurements.

library: Specialized collection of books, jour¬
nals, and reprints in volcanology and related
subjects. (See also below, "Hawaiian Volcano
Research Association, Library.”)
publication series: Hawaiian Volcano Observa¬
tory Bulletin, 1912-1929, and Special Reports
(both in collaboration with Hawaiian Volcano
Research Association); Volcano Letter, 1925 to
date (in collaboration with Hawaiian Volcano
Research Association and University of Ha¬
waii). A number of publications by staff mem¬
bers have appeared in various scientific journals.
research fellowships: See "Hawaiian Volcano
Research Association, Research Fellowships.”
Research policy: Cordial collaboration; desk
space and laboratory facilities afforded to visit¬
ing scientists.

Hawaiian Volcano Research Association
address: Hawaiian Volcano Research Associa¬
tion. President: L. P. Thurston; Secretary:

L. W. de Vis-Norton, 320 James Campbell
Building, Honolulu, Hawaii; Scientific Direc¬
tor: Dr. T. A. Jaggar, University of Hawaii,
Honolulu 10, Hawaii.

purpose: To sponsor research in physical pro¬
cesses of Hawaiian volcanoes. The Association
co-operates in research with the Hawaiian Vol¬
cano Observatory.

persons engaged in research: (At University
of Hawaii) Dr. T. A. Jaggar, Research Asso¬
ciate in Volcanology, and R. A. Okuda, Junior
Researcher; (at Hawaii National Park) staff of
Hawaiian Volcano Observatory.

facilities: (1) Kilauea Laboratory of the Uni¬
versity of Hawaii (instrument shop, collec¬
tions, files, instruments, maps, records, etc.);
(2) Hawaiian Volcano Observatory at Hawaii
National Park (see "Hawaiian Volcano Ob¬
servatory”); (3) Volcanology Laboratory,
Room 2, Home Economics Building, University
campus.

library: (At Hawaiian Volcano Observatory,
Hawaii National Park) Many volumes, record
books, reprints, journals, maps, instrument de¬
signs, and seismograms.

exhibits: (At Hawaii National Park) Volcanic
specimens, photographs, models, and exhibition
seismograph.

publication series: See "Hawaiian Volcano
Observatory.”

research fellowships: The Directors will re¬
ceive applications at any time from research
investigators holding doctorate degrees who de¬
sire to pursue specialized Hawaii studies in
seismology, volcanology, and volcanological
oceanography. Persons of advanced grade hold¬
ing fellowships from mainland colleges may be
assisted in travel expense and provided with
apparatus or use of facilities. Address inquiries
to: Dr. T. A. Jaggar, Scientific Director, Uni¬
versity of Hawaii, Honolulu 10, Hawaii.

research policy: To encourage highly skilled,
specialized research on Hawaiian volcanology,
seismology, and related phenomena. Desk space
and laboratory facilities are offered to visiting
scientists.

Honolulu Board of Water Supply

address: Honolulu Board of Water Supply, P. O.
Box 3410, Honolulu 1, Hawaii.

purpose: To maintain, expand, and improve the
Honolulu water supply.

subdivisions (engaged in other than service ac¬
tivities) : Division of Geology (Dr. C. K. Went¬
worth and two others) ; Division of Chemistry
(L. T. Bryson and one other) ; Division of
Bacteriology (J. M. Downer and four others ) .

facilities: Separate laboratories for the three
divisions, with supplementary darkroom, mi¬
croscope room, and storage and office space.

122

PACIFIC SCIENCE, Vol. 1, April, 1947

Equipment for general studies in areal geology
and petrography, chemical analyses of water
and other materials, special studies of corrosion
and other impairment, bacteriology of water,
and the like.

OPPORTUNITIES FOR FIELD RESEARCH: (1) Work
in geology includes field mapping of structures
and ground-water features, laboratory and field
experiments in hydrology, and collection of
data and mathematical studies in meteorology
and hydrology (the latter in collaboration with
the Division of Water Resources of the Board
of Water Supply). (2) Work in chemistry in¬
cludes analysis of daily samples, annual com¬
plete analyses of water from various sources,
and required special analyses, as well as tests
of fuel oil, metals, and construction materials;
studies of corrosion or other failure as needed.
(3) Work in bacteriology includes daily sam¬
pling at sources and at points in the distri¬
bution system, with special test programs to
maintain required and recognized standards for
potable water.

library: About 100 feet of shelves containing
technical books and journals on engineering,
water supply, and related technical subjects.
collections and exhibits: Working collections
of rocks and 'drill cores; photograph files; occa¬
sional charts and technical exhibits.
publications: Biennial reports containing rec¬
ords of quantities and costs, analyses, and other
pertinent data, with occasional technical ap¬
pendixes. A few supplementary reports have
also been published on geology, ground water,
water law, and similar subjects.
research policy: Cordial informal relations
with visiting engineers and scientists.

Libby, McNeill and Libby

address: Libby, McNeill and Libby, Pineapple
Division, P. O. Box 1140, Honolulu 7, Hawaii.
Manager of Research: Dr. O. C. Magistad.
purpose: This company conducts research both
on pineapple production and on processing.
staff: About 10 men devote full time to research
problems under supervisory staff of university
graduates in chemistry or agriculture.
facilities: In addition to main laboratory in
the Libby Cannery, field laboratories have been
provided on the islands of Oahu, Molokai, and
Maui.

Pacific Chemical and Fertilizer Company

address: Research Division, Pacific Chemical and
Fertilizer Company, P. O. Box 48, Honolulu 10,
Hawaii.

purpose: (1) To conduct investigations in the
chemical and agricultural fields related to the
company’s business. (2) To co-operate with
local research organizations such as the Experi¬
ment Station of the Hawaiian Sugar Planters’

Association, the Pineapple Research Institute,
and the University of Hawaii Agricultural Ex¬
periment Station on research problems of mu¬
tual interest. (3) To conduct or direct research
for clients on a contractual basis. (4) To act
as consultants in chemistry and agriculture.
staff: Four chemists, one chemical engineer,
one agricultural technologist, and five addi¬
tional employees. The division is under the
direction of Dr. John H. Payne.
facilities: Laboratory area of 5,000 square feet. '
library: Technical library of some 1,500 volumes.
publications: Papers are published (most com¬
monly in chemical periodicals) by staff mem¬
bers from time to time.

RESEARCH opportunities: The company offers
no research fellowship at present. Facilities are
available to visiting scientists.

Pineapple Research Institute of Hawaii

address: Pineapple Research Institute of Hawaii,
P. O. Box 3166, Honolulu 2, Hawaii. President
and Director: Dr. Eugene C. Auchter.
purpose: The Institute, which had its beginnings
about 1912, is supported by the pineapple in¬
dustry of Hawaii. It is located on land adjoin¬
ing the University of Hawaii and co-operation
with the University is arranged in the study of
problems of mutual interest. The purpose of
the Institute is to conduct research on all
problems encountered in the production of
pineapple plants and fruit.
subdivisions: Departments of Entomology, Plant
Pathology, Plant Physiology, Genetics, Chemis¬
try, Agricultural Engineering, Meteorology, and
Publications.

staff: About 40 scientists are engaged in re¬
search. Additional employees include business
organization, secretarial staffs, and farm labor.
Department heads include Dr. E. G. McKibben,
agricultural engineering; Dr. M. B. Linford,
pathology; Dr. J. L. Collins, genetics; Dr.
Walter Carter, entomology; Dr. G. T. Night¬
ingale, plant physiology; Dr. Harold E. Clark,
chemistry; L. B. Leopold, meteorology; and
Joyce Roberts, publications. Many of the asso¬
ciates hold doctorate degrees.
facilities: Well-equipped laboratories are avail¬
able in all departments.

opportunities for field research: Green¬
houses, shade houses, and an experiment station
of 100 acres (located at Wahiawa, Oahu) are
available. Research may also be carried on in
the pineapple fields of member companies.
library: The library contains 4,352 bound and
unbound volumes and several hundred special
pamphlets; 101 research journals and technical
publications are received regularly.
exhibits: Permanent exhibits in several branches
of research have been set up. An excellent
living collection of the various pineapple spe-

NOTES

123

cies, hybrids, and clones is established on the
grounds of the experiment station at Wahiawa.
PUBLICATIONS: Pineapple News, 1927-1936; Pine¬
apple Quarterly, 1931-1936. Recent policy has
been to issue reports of research to the pine¬
apple industry, and to publish papers giving the
results of technical investigations in various
American and foreign scientific journals.
research fellowships: Through an arrange¬
ment with the University of Hawaii it is pos¬
sible for graduate students to prepare their
theses under the direction of members of the
research staff and to work in the laboratories
or at the experiment station. Occasionally re¬
search fellowships or research assistantships are
offered.

RESEARCH opportunities: Outstanding scientists
from the Mainland and elsewhere are often
invited to come to the Institute to work on
problems of common interest. To such scien¬
tists all the facilities of the Institute are made
available.

research policy: The Institute arranges co¬
operative studies with various scientific agencies
both in the Hawaiian Islands and on the Main¬
land on problems of mutual interest.

Territorial Board of Agriculture
and Forestry

address: Territory of Hawaii, Board of Commis¬
sioners of Agriculture and Forestry, King and
Keeaumoku Streets, P. O. Box 3319, Honolulu
1, Hawaii. President: Colin G. Lennox.
purpose: The Board is primarily concerned with
law enforcement, but does carry on some re¬
search.

subdivisions: Division of Fish and Game, which
is conducting biological research in fisheries and
wild life; Division of Animal Industry, which
carries on bacteriological and pathological re¬
search on animal diseases; Division of Ento¬
mology, which carries on biological research in
entomology primarily as it concerns the intro¬
duction of insect parasites. Foresters carry on
field research from time to time.
persons engaged in research: About nine.
facilities: Laboratories well equipped for all
necessary investigations. Laboratory facilities
for fisheries and wild life research are avail¬
able at present through the Department of
Zoology, University of Hawaii.
library: The Division of Entomology Library
consists of about 1,000 volumes, including the
principal sets of entomological publications.
collections: Division of Entomology has spe¬
cial collections which include fruit flies of the
world, parasitic insects, and general collections
made in Africa, Asia, Australia, Mexico, Pana¬
ma, and Brazil. A permanent exhibit of eco¬
nomically important insects is presented in
display cases.

publications: Biennial Report; Division of
Forestry, Botanical Bulletins 1-6 (1911-1919);
Hawaiian Forester and Agriculturist, 1904-1933.
Staff members have prepared special reports,
bulletins, and articles.

research policy: No standing offer of research
facilities is made, but these have been used on
several occasions by visiting scientists with the
Board’s permission. Inquiries should be ad¬
dressed to the president of the Board.

Territorial Board of Health

address: Territory of Hawaii, Board of Health,
Kapuaiwa Building, Honolulu, Hawaii. Presi¬
dent: Dr. C. L. Wilbar, Jr.
purpose: General charge, supervision, and care
of the health and lives of the people of the
Territory.

subdivisions: Preventive Medicine; Local Health
Services; Medical Services; Central Administra¬
tion; and Sanitation (including industrial hy¬
giene, food and drugs, rodent and mosquito
control, housing). An active research program
is carried on by Dr. David S. Bonnet, Medical
Entomologist, for the control and prevention of
diseases parried by rodents and mosquitoes.
facilities: Adequate laboratories are available
in the health department offices on the various
islands.

opportunities for field research: Profes¬
sional and administrative personnel are busy
with ordinary routine duties, but existing pro¬
grams offer material for much research. *
publications: Annual Report.
research fellowships: From time to time fel¬
lowships are offered to members of the Board’s
medical, nursing, sanitation, laboratory, and
other staffs.

U. S. Bureau of Animal Industry

address: U. S. Department of Agriculture, Bu¬
reau of Animal Industry, 219 Federal Office
Building, Honolulu 2, Hawaii. Inspector in
Charge: Dr. A. H. Julien.
research policy: This bureau is primarily a
routine inspection agency, and no research pro¬
gram is carried on at present.

U. S. Bureau of Entomology and
Plant Quarantine

address: U. S. Department of Agriculture, Agri¬
cultural Research Administration, Bureau of
Entomology and Plant Quarantine, Division of
Foreign Plant Quarantine, P. O. Box 340,
Honolulu 9, Hawaii.

research policy: This division is primarily an
inspection agency, and no research program is
carried on at present. (See listing on Fruitfly
Investigations division below.)

124

PACIFIC SCIENCE, Vol. 1, April, 1947

U. S. Bureau of Entomology and
Plant Quarantine
(Fruitfly Investigations)

address: U. S. Department of Agriculture, Agri¬
cultural Research Administration, Bureau of
Entomology and Plant Quarantine, Fruitfly In¬
vestigations, University of Hawaii Campus,
P. O. Box 340, Honolulu 9, Hawaii. Entomolo¬
gist in Charge: J. W. Balock.
purpose: Research on the biology, ecology, and
chemical control of fruit flies; development of
methods of fruit treatment to eliminate risk
that living insects of economic importance will
be transported through channels of commerce.
staff: Normally, three professional workers, one
sub-professional worker, and one administrative
clerk.

facilities: Well-equipped laboratory for en¬
tomological research (refrigeration facilities,
vapor-heat room and equipment, constant tem¬
perature cabinets, micro-balance, etc.).
opportunities for field research: Field re¬
search conducted in co-operation with in¬
dividual farmers or on Bureau’s own research
plots.

library: Limited library on general entomology.
publications: Research results appear in scien¬
tific journals and U. S. Department of Agri¬
culture publications.

research policy: Facilities and laboratory space
are willingly made available to visiting scien¬
tists interested in these problems. Such facil¬
ities have also been made available to local
scientists when practicable.

U. S. Coast and Geodetic Survey

address: U. S. Department of Commerce, U. S.
Coast and Geodetic Survey, Pacific District
Headquarters, 244 Federal Office Building,
Honolulu, Hawaii. Supervisor, Pacific District:
Lt. Comdr. L. C. Wilder.
purpose: The principal functions of the Coast
and Geodetic Survey are the surveying of all
coastal waters under the jurisdiction of the
United States and the production of the naut¬
ical charts and coast pilot publications required
for the navigation of those waters; the com¬
pilation of aeronautical charts for air naviga¬
tion; and the accomplishment, throughout our
country and its possessions, of geodetic control
surveys which provide essential basic data for
nautical charting and topographic mapping
(see U. S. Coast and Geodetic Survey Special
Publication 216, "The United States Coast and
Geodetic Survey,” Washington, 1938). Re¬
search is carried on principally in the office in
Washington, D.C., from field data assembled
there; work is principally in seismology, mag¬
netism, tides, currents, and sea water salinities
and temperature, and in the development of
necessary instruments and equipment for carry¬

ing on field work. The principal work in the
area directed from Pacific Headquarters is to
obtain and make available nautical informa¬
tion, charts, and tide and current tables to the
seafaring profession and allied interests.
facilities: At Barbers Point, Oahu, is located the
Coast and Geodetic Magnetic and Seismological
Observatory, in charge of R. F. White, Geo¬
physicist. It is likely that this observatory will
be expanded to handle the correlating of va^
rious earthquake reports and possibly of those
on seismic sea waves. The Pacific District will
soon set up, in co-operation with other agencies,
a comprehensive series of tide gages which will
also record water salinities and temperatures.
library: Limited file of publications upon tides,
currents, seismology, magnetism, chart and map
making, surveying, and astronomy.

U. S. Geological Survey,

Division of Surface Waters

address: U. S. Department of the Interior, Geo¬
logical Survey, Water Resources Branch, Divi¬
sion of Surface Waters, 225 Federal Office
Building, Honolulu 2, Hawaii. District Engi¬
neer: Max H. Carson. (The Geological Survey
is associated with the Territorial Division of
Hydrography, of which Mr. Carson is Chief
Hydrographer. )

purpose: Collection of reliable records of flow
of the principal streams and ditches in the
Territory and their interpretation. The organi¬
zation also collects data on the artesian wells
in the Territory and administers the laws re¬
lating to the conservation of artesian waters.
(See also Ground Water Division, below.)
persons engaged in research: The Geological
Survey co-operates with the Division of Hy¬
drography of the Territory, the two organiza¬
tions functioning as a unit. The staff includes
several hydraulic engineers, two engineering
aides, and one engineering draftsman in Federal
pay; and one hydraulic engineer, two engineer¬
ing aides, and two clerical employees in Terri¬
torial pay.

facilities: A small hydraulics laboratory is sit¬
uated just below Nuuanu Reservoir No. 3 in
Nuuanu Valley. Here, previous to the war,
several models of streams were built and tested.
At present no laboratory work is being done,
but plans call for one engineer to spend full
time in laboratory research.
library: Consists of U. S. Geological Survey
Water-Supply Papers 1-1014, Professional Pa¬
pers 42—207, Bulletins 600—930, Annual Reports
17-25, Miscellaneous Mineral Reports, various
monographs, and more than 500 state and mis¬
cellaneous reports and textbooks.
publication series: Publications of the Survey
on Hawaii are a part of the series of U. S.
Geological Survey Water-Supply Papers. Num-

NOTES

125

bers of the series pertaining to Hawaii are 77,

318, 336, 373, 43 0, 44 5, 465, 485, 515, 516,

535, 555, 575, 595, 615, 635, 655, 675, 695,

710, 725, 740, 755, 770, 795, 815, 835, 865,

885, 905, 935, 965, 985.

RESEARCH policy: No special facilities are of¬
fered to visiting scientists.

U. S. Geological Survey,

Ground Water Division

address: U. S. Department of the Interior, Geo¬
logical Survey, Ground Water Division, 333
Federal Office Building, Honolulu 2, Hawaii.
District Geologist: Dr. Gordon A. Macdonald.
(See also Surface Water Division, above.)
purpose: Investigation of the geology and ground-
water resources of the Hawaiian Islands.
persons engaged in research: Dr. Gordon A.
Macdonald, District Geologist; Dan A. Davis,
Associate Geologist.

facilities: Laboratory- — none. Petrographic mi¬
croscope.

opportunities for field research: Excellent.
library: About 500 volumes, including U. S.
Geological Survey Professional Papers, Bulle¬
tins, Water-Supply Papers.
collections: Petrologic collections from the
Hawaiian Islands.

publication series: Bulletins of the Hawaii
Division of Hydrography.
research policy: All possible co-operation is
offered to provide facilities for visiting scien¬
tists.

U. S. Public Health Service

address: U. S. Public Health Service, 208 Fed¬
eral Office Building, Honolulu, Hawaii.
research policy: No local research program is
carried on at present.

U. S. Weather Bureau Office

address: U. S. Department of Commerce, LJ. S.
Weather Bureau Office, Federal Office Build¬
ing, Honolulu 1, Hawaii.
purpose: Weather forecasting for aviation and
shipping in the Pacific, as well as for local
interests. Collection, summarization, and pub¬
lication of Hawaii weather records.
subdivisions: Forecast Office (John Rodgers Air¬
port), Acting Official in Charge: Charles M.
Woffinden; Climatology Office (Federal Office
Building), Acting Official in Charge: W. F.
Feldwisch.

facilities: Original records available from ap¬
proximately 300 points in the Hawaiian Islands
(chiefly rainfall records). Analyzed weather
maps of Northern Hemisphere on file.
library: Approximately 100 volumes bearing on
meteorology, forecasting, and climatology. Sev¬
eral hundred volumes and various pamphlets

of published weather data, mostly from the
United States, but some from various other
parts of the world.

publications: Monthly and annual Meteorolo¬
gical Summary for Honolulu; monthly and
annual Climatological Data for the Hawaiian
Islands.

University of Hawaii

address: University of Hawaii, P. O. Box 18,
Honolulu 10, Hawaii. President: Gregg M.
Sinclair.

purpose: The University, a land-grant institu¬
tion of higher learning founded in 1907, offers
many opportunities for research in the physical
and biological sciences, both in the research
programs of faculty and students and in re¬
search in co-operation with many of the gov¬
ernmental and other agencies in the Hawaiian
Islands, such as the Bernice P. Bishop Museum,
the Pineapple Research Institute, the Hawaiian
Sugar Planters’ Association, and the U, S. Bu¬
reau of Entomology and Plant Quarantine.
(The work of the University of Hawaii Agri¬
cultural Experiment Station is separately re¬
ported below.)

subdivisions: The University departments which
carry on research in the natural sciences, with
the names of departmental chairmen, include:
Agriculture (Harold A. Wadsworth), Bacteri¬
ology (Dr. Floyd W. Hartmann), Botany (Dr.
Harold St John), Chemistry (Dr. Leonora N.
Bilger), Geology (Dr. Harold S. Palmer),
Physics (Dr. Willard H. Eller), and Zoology
and Entomology (Dr. Robert W. Hiatt). Cer¬
tain members of the University faculty are
assigned, as part of their duties, to carry on
research at the Bishop Museum.

persons engaged in research: In addition to
the department chairmen mentioned above, the
University faculty in the natural and phys¬
ical sciences includes: (Agriculture) Louis A.
Henke, Dr. William B. Storey; (Bacteriology)
Dr. Oswald A. Bushnell; (Botany) Dr. Charles
J. Engard, Dr. Donald P. Rogers; (Chemistry)
Dr. Earl M. Bilger, Dr. Frederick G. Mann, Dr.
Robert C. Brasted, Dr. Robert D. Bright, Dr.
Robert A. Spurr; (Physics) Dr. E. H. Bram-
hall, Iwao Miyake; (Zoology and Entomology)
Dr. Frederick G. Holdaway, Dr. Joseph E.
Alicata, Dr. Albert H. Banner, Dr. Harvey I.
Fisher, Dr. Pauline Heizer, Dr. Gordon B.
Mainland, Dr. Leonard D. Tuthill.

facilities: Laboratory facilities are available on
the University campus, in Gartley Hall for
chemistry and physics, and in Dean Hall for
botany, geology, zoology, and entomology. Agri¬
cultural and nutrition laboratories are found in
Gilmore Hall. Other research laboratories on
the campus are those of the U. S. Bureau of
Entomology and Plant Quarantine, the Pine¬
apple Research Institute, and the University

126

PACIFIC SCIENCE, Vol. 1, April, 1947

Agricultural Experiment Station. A large lab¬
oratory will be housed in the Agricultural En¬
gineering Institute now under construction, the
gift of the Hawaiian Sugar Planters’ Associa¬
tion. Off-campus laboratories include the Ma¬
rine Biological Laboratory and Aquarium at
Waikiki, the Astronomical Observatory at Kai-
muki, and the Kilauea Laboratory in Hawaii
National Park.

opportunities for field research: Field re¬
search trips are organized periodically, under
the supervision of University faculty members.
In the past these trips have covered not only the
outer islands of the Hawaiian Group but also
other islands of the Pacific region. As a part of
the program of the University’s Pacific Islands
Research Committee headed by Dean Paul S.
Bachman, in December, 1945, four faculty
members made a reconnaissance visit to Micro¬
nesia, which was followed in the summer of
1946 by teams of scientists who made field
surveys in Micronesia particularly in botany,
zoology and bacteriology, and parasitology (see
Pacific Science, January, 1946, p. 61-62).

library: The University Library contains 163,950
bound volumes and 378,829 unbound parts and
pamphlets, including the main publications in
natural science. A union list of serials in the
Library indicates the locations of all periodicals
found in all libraries in the Territory. Standard
scientific works are to be found on the Library
shelves; well represented are works on the
botany of the Pacific area, tropical agriculture,
marine biology, and chemistry; there are also
many volumes on voyages and scientific expedi¬
tions to the Pacific. The Library is an official
depository for Federal and Territorial govern¬
mental publications. Scientific works printed in
the Chinese and Japanese languages are in¬
cluded in the Oriental Collection. Visiting
scientists are accorded free use of all Library
facilities.

exhibits: Collections or permanent exhibits, by
act of the Territorial Legislature, are placed in
the Bishop Museum, and a considerable part of
the botanical and zoological exhibits in that
museum is the work of University scientists.
The herbarium there housed contains the most
nearly complete collection of Hawaiian plants
in existence, including some species now extinct.
Certain departments of the University arrange
displays on the campus from time to time.
Large-scale relief maps of the major islands of
the Hawaiian Group are to be found in the
main lobby of Gilmore Hall. A botanical tour
of the campus is described in a free illustrated
booklet, In Green Manoa Valley.

PUBLICATION SERIES: Research Publications (1927
to date); Occasional Papers (1934 to date). In
January, 1947, the University began publication
of Pacific Science, a quarterly devoted to the
biological and physical sciences of the Pacific

region. The Volcano Letter, since 1938, has
been published by the University for the Ha¬
waiian Volcano Observatory and the Hawaiian
Volcano Research Association. Proceedings of
the Hawaiian Academy of Science has been
issued jointly by the University and the Acad¬
emy since 1940.

RESEARCH fellowships: Fellowships are offered
in scientific fields, on a half-time basis, to quali¬
fied graduate students.

research policy: The University, as a fully ac¬
credited institution of higher learning, is de¬
sirous of promoting scientific research in every
possible way, and members of its faculty and
staff are allowed time and funds to carry on
such research. Co-operative research of many
sorts is carried on by arrangement with various
other institutions and agencies. It is the policy
of the University to extend its hospitality to
visiting scientists who wish to arrange for lab¬
oratory and library facilities.

University of Hawaii Agricultural
Experiment Station

address: University of Hawaii Agricultural Ex¬
periment Station, Box 18, Honolulu 10, Hawaii.
Director: Dr. John H. Beaumont.

purpose: To conduct research and experiments
bearing upon the establishment and mainte¬
nance of a permanent and efficient agricultural
industry in the Territory.

subdivisions: Departments of Agronomy, Horti¬
culture, Animal Husbandry, Poultry Husband¬
ry, Nutrition, Soils and Agricultural Chemistry,
Plant Physiology, Plant Pathology, Vegetable
Crops, Parasitology, Entomology, and Agricul¬
tural Engineering.

PERSONS ENGAGED in research: More than 40.
Department heads include Dr. J. E. Alicata,
Dr. H. F. Clements, Dr. W. A. Frazier, Rene
Guillou, J. W. Hendrix, L. A. Henke, Dr. F. G.
Holdaway, Carey D. Miller, J. C. Ripperton,
Dr. G. D. Sherman, and Dr. W. B. Storey.

facilities: Adequate laboratories in Soils and
Agricultural Chemistry, Plant Physiology, Nu¬
trition, Poultry Husbandry, Horticulture, and
Vegetable Crops. Field experiment stations are
located on the University campus at Honolulu,
at Poamoho on Oahu, at Haleakala on Maui,
and at Kona on Hawaii.

library: University of Hawaii Library and Sta¬
tion Library.

publication series: Annual Report, Bulletin,
Circular, Technical Bulletin, Progress Notes,
Technical Papers.

research fellowships: Fellowships are offered
occasionally, as need arises.

research opportunities: Graduate and under¬
graduate research is carried on in collaboration
with the University of Hawaii.

research policy: Facilities of the Station are
freely offered to visiting scientists.

JULY, 1947

No

^ U V J

VOL. I f)rj

PACIFIC

SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

IN THIS ISSUE: Fisher —Utinomi's Bibliographic a Micronesica :
Chordate Sections • Clements and Munson— Arsenic Toxicity
Studies in Soil and in Culture Solution • Wentworth— Factors
in the Behavior of a Ghyben-Herzberg System • Zetler—

Travel Times of Seismic Sea Waves to Honolulu • NOTES

Published by

THE UNIVERSITY OF HAWAII
HONOLULU, HAWAII

BOARD OF EDITORS

A. Grove Day, Editor-in-chief, Department of English, University of Hawaii
Ervin H. Bramhall, Department of Physics, University of Hawaii
Vernon E. Brock, Director, Division of Fish and Game, Territorial Board of
Agriculture and Forestry

Harry F. Clements, Plant Physiologist, University of Hawaii Agricultural
Experiment Station

Charles H. Edmondson, Zoologist, Bishop Museum, Honolulu, T. H,
Harvey I. Fisher, Department of Zoology, University of Hawaii
Frederick G. Holdaway, Head, Department of Entomology, University of
Hawaii Agricultural Experiment Station
Maurice B. Linford, Head, Department of Plant Pathology, Pineapple Research
Institute, Honolulu, T. H.

A. J. Mangelsdorf, Geneticist, Experiment Station, Hawaiian Sugar Planters’
Association, Honolulu, T. H.

G. F. Papenfuss, Department of Botany, University of California, Berkeley 4,
California

Harold St. John, Chairman, Department of Botany, University of Hawaii
Chester K. Wentworth, Geologist, Honolulu Board of Water Supply

SUGGESTIONS TO AUTHORS

Contributions to Pacific biological and physical
science will be welcomed from authors in all parts
of the world. Manuscripts should be addressed to
Dr. A. Grove Day, Editor, Pacific Science, Uni¬
versity of Hawaii, Honolulu 10, Hawaii. Use of
air mail for sending correspondence and brief
manuscripts from distant points is recommended.
Manuscripts will be read promptly by members of
the Board of Editors and by other competent
critics.

Manuscripts may run from 1 to 30 pages in
length. Authors should not overlook the need for
good brief papers presenting results of studies,
notes and queries, communications to the editor,
or other commentary.

Preparation of Manuscript

Although no manuscript will be rejected merely
because it does not conform to the style of Pacific
Science, it is suggested that authors follow the
style recommended below and exemplified in the
journal.

Title. Titles should be descriptive but brief. If a
title runs to more than 40 characters, the author
should also supply a "short title” for use as a
running head.

Manuscript form. Manuscripts should be typed
on one side of standard-size, white bond paper
and double-spaced throughout. Pages should be

consecutively numbered in upper right-hand cor¬
ner. Sheets should not be fastened together in any
way, and should be mailed flat. Inserts should be
either typed on separate sheets or pasted on proper
page, and point of insertion should be clearly indi¬
cated.

Original copy and one carbon copy of manuscript
should be submitted. The author should retain a
carbon copy. Although due care will be taken, the
editors cannot be responsible for loss of manu¬
scripts.

Introduction and summary . It is desirable to state
the purpose and scope of the paper in an intro¬
ductory paragraph and to give a summary of
results at the end of the paper.

Dictionary style. It is recommended that authors
follow capitalization, spelling, compoundings ab¬
breviations, etc., given in Webster’s New Inter -
national Dictionary (unabridged), second edition;
or, if desired, the Oxford Dictionary. Abbrevia¬
tions of titles of publications should, if possible,
follow those given in U. S. Department of Agri¬
culture Miscellaneous Publication 337.

Footnotes. Footnotes should be used sparingly and
never for citing references (see later). Often,
footnotes may better be incorporated into the text
or omitted. When used, footnotes should be con¬
secutively numbered by superior figures through-
[ Continued on inside back cover']

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

Vol. I JULY, 1947 No. 3

Previous issue published April 1 , 1947

CONTENTS

PAGE

Utinomi’s Bibliographica Micronesica: Chordate Sections. Harvey I. Fisher . . 129

Arsenic Toxicity Studies in Soil and in Culture Solution. Harry F. Clements and

Jerome Munson . 151

Factors in the Behavior of Ground Water in a Ghyben-Herzberg System. Chester

K. Wentworth . . . 172

Travel Times of Seismic Sea Waves to Honolulu. Bernard D. Zetler ... 185

Notes:

New Botanical Bibliography of Pacific Islands by E. D. Merrill .... 189

Scientists and the Fortieth Anniversary of the University of Haivaii ... 189

Cover drawing by A. S. MacLeod

r

Pacific Science, a quarterly publication of the University of Hawaii, appears in January,
April, July, and October. Subscription price is three dollars a year; single copies are one
dollar. Check or money order payable to University of Hawaii should be sent to Pacific
Science, Office of Publications, University of Hawaii, Honolulu 10, Hawaii.

Utinomi’s Bibliographic a Micronesica: Chordate Sections

Harvey I. Fisher1

A copy of Bibliographica Micronesica /
Scientiae Naturalis et Cult us, by Dr. Huzio
Utinomi, became temporarily available in
the Territory of Hawaii late in the summer
of 1946. This bibliography of 208 pages
was published in 1944 by the Hokuryukan
Publishing Company in Tokyo. A negative
microfilm was made by the University of
Hawaii Library, and later certain sections
were enlarged and printed photograph¬
ically.

An interest in the vertebrate animals of
Micronesia, especially those of Yap, led me
to have certain Japanese titles translated for
personal use. It soon became evident that
although the bibliography was not com¬
plete, it did include many significant titles
that had previously been overlooked by
workers in vertebrate zoology.

This bibliography has great interest at the
present time. Failure to consider scientific
papers by Japanese students may be at¬
tributed to language difficulties and to lack
of knowledge even of the existence of such
papers. Apparently few copies of this bib¬
liography can be found outside Japan.
Further, interest in the area concerned has
been increased greatly by the recent occupa¬
tion of Micronesia by Americans. Several
expeditions and survey parties have been
through the islands since the cessation of
hostilities. Interest in these new possessions
has grown among scientists of the post-war
period, and many scientific investigations
can be expected in these islands that were
formerly under Japanese mandate. In many

1 Department of Zoology and Entomology, Uni¬
versity of Hawaii. Manuscript received March 1,
1947.

[

branches of science it would be inadvisable
to start a study without some knowledge of
the work carried on by Japanese scientists
in the mandated islands.

Because of the above facts it seems desir¬
able to publish immediately all the titles
given by Utinomi, and to add translations
of the titles and publications cited in the
Japanese language. The present paper in¬
cludes only those sections dealing with chor¬
date animals, and constitutes pages 24 to 43
of the original publication, in addition to the
translated Preface and Explanatory Notes.

The list of titles is of course not exhaus¬
tive, but it is not the purpose of this pub¬
lication to add titles to Utinomi’s list. A
complete bibliography of the chordates in
Micronesia would take years of preparation
and research in many libraries. The imme¬
diate usefulness of the bibliography in its
present form, it is hoped, will excuse some
incompleteness.

Many difficulties were encountered. One
of the most serious was our inability to be
absolutely certain of the "accepted” English
translations of the Japanese journals and
books. The Japanese ideographs may often
be interpreted in several ways; each may be
literally correct but at the same time may
not be the exact wording of the journal’s
name. For example, the ideograph "Kaiho”
may mean either "journal” or "transactions.”
In one instance we found in the United
States Department of Agriculture Miscel¬
laneous Publications No. 337, on abbrevia¬
tions of titles of publications, that the
ff Journal of the Formosa Natural History
Society” was the accepted translation, but on

130

a copy of this journal the masthead reads
" Transactions of ... Consequently,
"Kaiho” has been used consistently as "trans¬
actions,” but in some cases it probably should
mean "journal.” "Dobutsu” means animals,
and "Dobutsugaku” means zoology, but
"Dobutsu” as part of a journal’s title may
signify zoology. Some English translations
of the names of Japanese journals are, how¬
ever, given in an appendix to the original
book (p. 183 ff.). Difficulty of course
exists also in translating titles of papers*, but
for these we have been less concerned in ob¬
taining exact wording, for we felt that a
more or less precise indication of content
would be sufficient to make the translation
markedly useful to others.

Owing to this uncertainty in some trans¬
lations, we have deemed it advisable to in¬
clude the Japanese, and in some instances
Chinese, characters where they were given in
the original. Titles of journals, books, and
reports are given in the original characters
and in the Hepburn system of Romanization,
as well as in explanatory English translation
if no accepted translation could be found.

All translations and explanations added by
the present author are enclosed in brackets.
Brackets in the original have been reduced
to parentheses. Volume numbers in the
original have been replaced by italics below.

All citations have been left in their orig¬
inal form if this form was understandable;
that is, we have not attempted to make uni¬
form the manner in which papers are cited.
The original publication contains many typo¬
graphical errors, misspelled words, and in¬
correct and omitted diacritical marks. When¬
ever it could be determined without question
that an inaccuracy was present, correction
was made. However, inability to check the
citation of many titles and their pagination
has made it impossible to remedy all such
errors.

The use of common names in titles has
proved very bothersome. It took, for ex-

PACIFIC SCIENCE, Vol. I, July, 1947

ample, considerable effort to make certain
that the two characters meaning "grave” and
"builder” referred to megapode birds only,
and not to various burrowing birds. In some
few places we have been forced to use the
descriptive common name implied by the
Japanese characters, in lieu of a positive
scientific name.

I wish to acknowledge gratefully the
wholehearted assistance and co-operation of
Mr. K. Sakimura of the Pineapple Research
Institute in Honolulu and of Miss M. Fukuda
and Dr. Y. Uyehara of the University of
Hawaii, all of whom worked on the trans¬
lations from the Japanese. Final decision
on the scientific terminology, however, re¬
mains the responsibility of the present writer.

Since other members of the faculty of the
University of Hawaii are now preparing
translations of sections of the bibliography
that deal with their specialized fields, it was
felt that this first redaction should include
Utinomi’s Preface and Explanatory Notes to
the original book. These are hereinafter
given in their entirety in a translation by Dr.
Uyehara.

PREFACE

The compilation of this bibliography, as
I have previously stated in the first volume
of the 1938-39 issue of Kagaku Nanyo
[South Sea Science ), is an attempt to collect*
every possible paper, both native and for¬
eign, on the Japanese-owned South Sea
islands. Thus, I may offer materials neces¬
sary for the survey of resources and for
scientific research on the South Sea islands
which constitute the only southern territory
of our country, and at the same time may
furnish information to those who wish to
engage in the study of the science of the
South Seas at the Palao Tropical Biological
Station.

Five years have elapsed since my an¬
nouncement of this compilation, but the pub¬
lication was unavoidably delayed because the

Utinomi’s Chordate Section — Fisher

131

compiler had to serve three years of that time
in war duties. However, current develop¬
ments do not permit me to procrastinate any
longer. Although the existing circumstances
differ greatly from those of the pre-war
period in which this project was started, the
value of our South Sea islands has not
changed in the least. Moreover, an exhaus¬
tive scientific study of these islands as a basis
for the development of this region should
never be neglected. Hence, I believe the pub¬
lication of this bibliography now is very
timely; and I will be very pleased if it should
in any way serve as a guide to scientific re¬
search in our South Sea islands. As we note
in this collection, the scientific studies made
by Japanese scholars since these . islands
became a territory of Japan are not few.
Thus, they are already contributing greatly
toward the advancement of South Sea sci¬
ence. The compilation of this bibliography
is but the fruit of the efforts of these schol¬
ars. I take this opportunity to pay my deep¬
est respects to them.

In this bibliography, all treatises obtain¬
able, from both native and foreign sources,
that deal with the sciences (botany, zoology,
geology-mineralogy, oceanography, geo¬
physics, medicine, anthropology-ethnology,
and geography) relating to the South Sea
islands have been included. I have made an
effort to list even popular literature, if con¬
sidered sufficient to serve as good references,
bringing the total number to almost four
thousand. The greater part of these have
been collected by the compiler himself.
Furthermore, to insure a greater degree of
perfection, various specialists have been
asked to assist in the gathering of literature
or in revision, as itemized below. However,
the scope of the collection is extremely great
and owing to lack of a better indexing
method on the part of the compiler, there
may be more than a few omissions. In filling
this gap, I appeal for the co-operation of
those who utilize this bibliography; at the

same time I hope they will add titles as they
see fit.

In presenting this bibliography to the
world, I should like to list the names of those
who have given me greatest assistance in the
compilation of this work, and to express my
deepest appreciation for their kind efforts.
Moreover, I have been accorded facilities by
numerous government offices, universities
and colleges, research institutes, libraries,
public organizations, and individuals. For
want of space, I have not listed their names
here; but deep gratitude is due them for
their kindness.

Botany:2 Hosokawa, Takahide; Tsuyama, Ta-
kashi (Phanerogamae) : Tagawa, Motoji (Pteri-
dophyta) : Horikawa, Yoshio (Bryophyta): Ya-
mada, Yukio (Algae).

Zoology: Kuroda, Naganari (Mammalia;
Aves): Momiyama, TokutarS (Aves): Okada,
Yaichiro; Oshima, Masamitsu (Reptilia; Amphi¬
bia): Abe, Toshiaki (Pisces): Tokioka, Takashi
(Protochordata): Kuroda, Tokubei; Baba, Kiku-~
taro (Mollusca): Hayashi, RySji (Echinoder-
mata) : Oshima, Hiroshi; Murakami, Shiro (Holo-
thuroidea; Ophiuroidea) : Miyake, Sadayoshi
(Crustacea): Sat5, Ikio; Takakuwa, Yoshioki;
Takashima, Haruo (Myriapoda; Arachnoidea) :
Ezaki, Teizo (Insecta): Takahashi, Ryoichi
(Hemiptera; Thysanoptera) : Tokunaga, Masaaki
(Diptera): Kano, Tadao (Coleoptera) : Naka¬
mura, Yamato (Lepidoptera) : Kobayashi, Shin-
jiro; Ofuchi, Matatsu (Oligochaeta) : Takahashi,
Keiz5; Okuda, Shir5 (Polychaeta) : Sato, Hayao
(Gephyrea): Ozaki, Yoshimasa; Ogata, Fujiharu
(Trematoda): Haneda, Yoshiine (Protozoa):
Abe, Noboru (Coelenterata; Zoological Miscel¬
lany).

Mineralogy and Coral Reef: Tayama, Riza-
bur5; Eguchi, Motoki.

Oceanography: Uda, Michitaka.

Geophysics: Kumagae, Naoichi (Magnetics):
Kawasaki, Hideo (Meteorology).

Medicine: Haneda, Yata; Medical officers of
the Nanyocho (South Sea Islands Government
Office).

Anthropology-Ethnology: Motoda, Shigeru;
Sugiura, Kenichi (Ethnology).

Geography: Wada, Shunji (Geography):
Wada, Seiji; Ban, Yoshii (Fishery) : Yoshino,
Tsuyoshi (Agriculture and Forestry).

2 Several readings or translations are possible
when Japanese names are written in Chinese char¬
acters. Hence, popularly accepted pronunciation
has been used here whenever actual translation
was not possible.-TRANSLATOR’s Note.

132

PACIFIC SCIENCE, Vol. I, July, 1947

Lastly, may I express my sincere apprecia¬
tion to the following persons, who have
given me the greatest facilities and authoriza¬
tion possible during all my work: Dr. Keita
Shibata, director of the Resource Science
Research Institute of the Ministry of Educa¬
tion, Dr. Shikiji Hatai, former director of
the Palao Tropical Biological Station, Dr.
Takashi Komai, professor at the Kyoto Im¬
perial University, and Dr. Teiz5 Ezaki, pro¬
fessor at the Kyushu Imperial University.
My sincere thanks are also due to Dr. Yai-
chiro Okada of the Resource Science Insti¬
tute and to Mr. Takemori Shintani of the
Society for the Promotion of Science in
Japan for their assistance in the publication
of this work; and to Mr. Ryotaro Fukuda,
president of the Hokuryukan, who has will¬
ingly undertaken the publication of a work
of this nature which involves much diffi¬
culty in view of the present situation.

June, 1943 The compiler

EXPLANATORY NOTES

1. This collection includes all scientific
works obtainable on the South Sea
islands, either from periodicals and ir¬
regular publications or in book form.
Since the subject of the collection lay
primarily in the field of natural sciences
and secondarily in cultural sciences,
thoroughness may be wanting in the lat¬
ter field.

2. The scope of this collection comprises
an area of the southwestern Pacific
Islands generally known as Micronesia,
limited primarily to the Japanese islands
( Marianas, Palao, Carolines, and Mar¬
shalls ) , but including also the adjacent
Gilbert and Wake Island groups.

3. The greater portion of the list includes
those items which had been published by
the end of 1942; however, the compiler
has tried his best to include also publi¬
cations of later dates.

4. The arrangement of the collection under
each heading follows an alphabetical
order, with the titles of papers appear¬
ing in complete form exactly as they are
found in the respective publications.
The order of each citation is: name of
author or compiler, the year of publica¬
tion, the title, the name of the journal
or magazine (abbreviated), the volume,
the number, and the page.

5. In case an abstract in a European lan¬
guage is appended to a publication in
Japanese, the corresponding title in the
European language appears in paren¬
theses after the Japanese title.

6. The volume is given in bold type [in
italics now] and the number in small
type. In instances in which the issue
number and the individual volume num¬
ber are likely to be confused, "no.” is
written before the former.

7. When the author or compiler’s name is

not mentioned, " - ” is used to

indicate such an omission, and the works
have been arranged in chronological
order at the end of every heading.

8. Page number in brackets [in parentheses
now] indicates that that page should be
referred to in relation to particular head¬
ings or subjects.

9. When a Japanese journal or magazine
has titles in both Japanese and a Euro¬
pean language, the title used is in Japa¬
nese if the paper is in Japanese, and in
the European language if the work is in
that language.

10. The complete names of the journals and
magazines used in abbreviated forms
are listed in the appendix.

11. Dates of publications are all according
to the Western calendar; and to facili¬
tate references, a table indicating Japa¬
nese chronology as compared to that of
the West is inserted at the end of the
book.

Utinomi’s Chordate Section— Fisher

133

[CHORDATE SECTIONS]

2. “f % II (Mammalia)

Anderson, K. 1908 Twenty new forms
of Piero pus. Ann. Mag. Nat. Hist., ser. 8,
2, pp. 361-370.

— - - 1912 Catalogue of the Chiroptera

in the Collection of the British Museum.
Second Edition. Vol. I. Megachiroptera.
(pp. 172-180, 275-279, 295-310, 797-
798) Brit. Mus. (Nat. Hist.), London.

[Asano, N.] 1938

fcgfc'tT . [On the Dugong of Palau]. SI
®J2^1|]#[Shokubutsu oyobi Dobutsu, or
Botany and Zoology], 6, 6, pp. 1047-
1051; 7, pp. 1219-1228.

Brandt, J. F. 1835 Prodomus descrip-
tionis animalium ab H. Mertensio in
orbis terrarum circummvigatione observa-
torum. Fasc. I. Petropoli. (pp. 42-61,
73-75)

Cuvier, G. 1821-1824 Osoements fos-
siles. 2 ed., IV. (p. 45)

Desmarest, A. 1822 Mammalogie. II,
Suppl. (p. 547) Paris.

Dobson, G. E. 1878 Catalogue of the Chi¬
roptera (in the Collection of the British
Museum) . ( pp. 63-64, 360-361 ) Brit.
Mus. (Nat. Hist.), London.

Freycinet, L. de 1825 Voyage autour du
monde execute sur les cojvettes de S. M.
FUranie et la Physicienne, pendant les
annees 1817-1820. Historique, 11, 1.
(p. 271) Paris.

Giebel, C. G. .1855 S'iugethiere. (p. 998)
Berlin.

Gray, J. E. 1870 Catalogue of Monkeys,
Lemurs, and Fruit-eating Bats in the Col¬
lection of the British Museum, (pp. 107-
109) Brit. Mus. (Nat. Hist.), London.

Hamilton-Smith, [?] 1827 Griffith’s
Animal Kingdom. (IV, p. 115; V, p.
311)

[Hirasaka, K.] 193.9 m C AM.
[Dugongs That Swim].

[Taiwan Suisan Zasshi, or Formosan Fish¬
eries Magazine], 286, pp. 1-4.

- — 1939 X [Dugongs of

Palau], [Kagaku Nanyo, or

South Sea Science], 2, 2, pp. 69-76.

— - — 1942 AMM^ [Tales of Du¬
gongs]. [Minzoku Taiwan, or

Formosan Ethnology], 2, 10, pp. 6-7.

Hombron, J. B. & Jacquinot, C. H. 1843
Voyage au Pole Sud et dans FOceanie sur
les corvette F Astrolabe et la Zelee; ex¬
ecute par ordre du Roi, pendant les annees
1837-1840. Zoologie, Atlas, III.

*#^r[HoRii, e.] 1916

[Report on the Zoological
Investigations or Survey of the South Sea
Islands, the New Territory]

[Nanyo Shinsenryochi Shisatsu
Hokoku, or The Survey Report on the
South Sea Islands, the New Territory],
pp. 234-266 (pp. 235-236);

(1927) [inin Tochi
Chiiki Nanyogunto Chdsa Shiryo, or Data
on the Investigation of the Newly Occu¬
pied South Sea Islands (1927)], 1, pp.
245-281. (pp. 246-247)

Jacquinot, C. H. & Pucheran, J. 1846
Voyage au pole sud et dans Foceanie sur
les corvette FAstrolabe et la Zelee; ex¬
ecute par ordre du Roi, pendant les annees
1 837-1 840. Zoologie, t. Ill. Mammiferes
et Oiseaux.

#FFK^f( Kish Ida, K.) 1924 m%MaB ffi
(Monograph of Japanese Mammals) .381,
pp. [Special Publica¬

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PACIFIC SCIENCE, Vol. I, July, 1947

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pt. II. Ibis, 13th ser., 2, pp. 132-161.

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PACIFIC SCIENCE, Vol. I, July, 1947

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Mearns, E. A. 1909 A List of Birds
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- - - 1934 The Fishes of Oceania - -

Supplement 2. Mem. Bernice P. Bishop
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— - 1941 The Fishes of the Groups

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- 1929 The Fishes of the

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Arsenic Toxicity Studies in Soil and in Culture Solution1

Harry F. Clements and Jerome Munson2

The problem of arsenic accumulation in
soils is one of comparatively recent import¬
ance. As agriculture became more intensive,
it became necessary to use poisons to combat
attacks of certain insects, fungi, and more
recently, weeds. Because arsenic is very
poisonous to plant enemies and because it
is comparatively cheap, it was only natural
that it should have found general use. The
arsenic so used has for the most part ac¬
cumulated in the upper soil layers, and
sooner or later becomes a menace to crop
production. This paper is concerned with
this problem particularly with reference to
Hawaiian soils.

HISTORICAL REVIEW

Early work. — Toward the end of the last century
and the beginning of the present century, some
attention was given to the possibility that arsenic
might poison the crop itself. In 1894, Lyttkens
conducted experiments at the Halmstead Experi¬
ment Station in Sweden which showed plainly that
the arsenic in the soil is a strong poison to plants.

Stoklasa (1898), however, showed that while
arsenic could not replace phosphorus as an essen¬
tial element, it had stimulative effects on the de¬
velopment of the assimilation organs of the oat
plant. The issue thus became confused. While
some presented evidence that arsenic was a poison
to plants, others presented equally convincing evi¬
dence that arsenic stimulated plant growth. Bouil-
hac (1899) reported that a number of fresh-water
algae absorb arsenic acid from arsenates without
apparent injury, and the growth of one appeared

1 Published with the approval of the Director of
the University of Hawaii Agricultural Experiment
Station as Technical Paper 156. Manuscript re¬
ceived February 15, 1947.

2 Dr. Clements is Professor of Botany and Plant
Physiologist, University of Hawaii Agricultural
Experiment Station. Mr. Munson was formerly a
graduate assistant in botany. University of Ha¬
waii; his master’s thesis, now in the University
Library, is incorporated in the present paper.

more favorably influenced by arsenic acid than by
phosphoric acid.

The fact that arsenic is a natural component of
most soils and not uncommon in plants tended to
reduce the force of criticism against the use of it.
Zuccari (1914), working in Italy, analyzed 20 soil
samples varying in physical and chemical compo¬
sition and taken from different depths in different
geological formations and at varying elevations;
these samples showed an arsenic content varying
from 0.187 to 6 parts per 100,000 of soil, being
largest in soils containing the most iron com¬
pounds and varying almost directly with iron con¬
tent.

Reichert and Trelles (1922) analyzed 20 soils
from different parts of Argentina and found all
but one to contain arsenic, varying in amounts
from 0.1 to 2.25 mg. per 100 gm. of soil.

Williams and Whetstone (1940) analyzed a
wide range of soils for arsenic and found the
range of naturally occurring arsenic to be between
0.3 and 40 ppm. Vegetation found growing on
these soils ranged from less than 0.1 ppm to 10
ppm.

Greaves (1934) analyzed, for total arsenic,
water-soluble arsenic, and various soluble salts,
50 orchard soils which had been in cultivation for
some time and which varied widely in chemical,
physical, and biological properties. Total arsenic
varied from 7.2 to 367.2 pounds per acre foot.
The water-soluble arsenic varied from 0.7 to 31.9
pounds per acre foot.

Greaves (1913*0 earlier reported that some vir¬
gin soils contain arsenic in appreciable quantities
which comes from the decay of native rocks. Many
cultivated orchard soils contain it in large quanti¬
ties, but he found no uniform relationship between
the total quantity in different soils and the water-
soluble arsenic of these soils. He considered that
the solubility of arsenic is governed largely by the
salts in the soil and the form in which the arsenic
is applied.

Thus, with comparatively high arsenic amounts
not uncommon as natural components of soils,
opinions on the danger from the use of compara¬
tively small additions continued to be diverse.

Headden (1909) was perhaps the first to recog¬
nize the serious dangers which may accrue to or¬
chard trees from continued use of arsenical insec¬
ticides. In fact, he set off a heated controversy
which has raged ever since. In other reports
(1908; 1910), he recognized three types of poison-

C151]

152

PACIFIC SCIENCE, Vol. I, July, 1947

ing, one corrosive, one systemic, and one which
was an arsenic-lime poisoning. He pointed out
that arsenical deposits in the soil are for the most
part insoluble, but warned that the soluble frac¬
tions had already passed the danger limits in cer¬
tain areas. He recognized that absorption of arse¬
nic by the roots takes place. He further pointed
out that salts such as NaCl, Na2S04, and Na2C03
are all capable of making lead arsenate soluble in
the soil solution and that lime salts do not prevent
the solution of arsenates.

Headden was joined in his efforts by Swingle
and Morris (1911), who reported that serious in¬
jury may result to apple trees from the applica¬
tion of insoluble arsenicals and that recent wounds
through the outer bark, functional lenticels, and
dormant buds permit the absorption of arsenic in
solution.

Greaves (1913) observed that water-soluble
arsenic may exist in soils to the extent of 82 ppm
without entirely stopping ammonification and ni¬
trification. He considered it improbable that lead
arsenate, zinc arsenite, or arsenic trisulfide would
ever be applied to agricultural soil in quantities
sufficient to'be injurious to soil bacteria.

Greaves and Anderson (1915) reported actual
stimulation of soil flora by soluble arsenic in soil
at 10 ppm. Toxicity began at 40 ppm and nitrogen
fixation was entirely stopped at 250 ppm. The
quantity of 10 ppm which they reported as causing
stimulation exceeds that found in most soils, and
they concluded that arsenic will stimulate instead
of retard bacterial activities of soil. Greaves
(1916) reported that arsenic cannot replace phos¬
phorus in the vital process of nitrogen fixing, but
it can, in some manner, liberate phosphorus from
its insoluble compounds.

Swingle (1920) investigated the effect of arsenic
on species of soil bacteria responsible for impor¬
tant chemical changes such as ammonification and
nitrogen fixation. Contrary to the work of Greaves,
Swingle’s results showed that all the arsenical
compounds used were germicidal, but in different
degrees.

Green and Kestell (1919) found bacteria which
are resistant to arsenic to be infrequent in soil and
air, but fairly frequent in feces. Of the twelve or
more resistant species examined, only two showed
any chemical activity toward arsenic: one, which
oxidizes arsenite to arsenate, and another, which
reduces arsenate to arsenite. No relationship was
discovered between arsenate reduction and nitrate
reduction.

McGeorge (1915^; 1915£) appears to have been
the first in Hawaii to recognize the possible dele¬
terious effects which may result from the use of
sodium arsenite as a herbicide, a practice then
coming into general use in the Territory. His re¬
searches involved a study of the effect of sodium
arsenite on the growth of millet, cowpeas, and
buckwheat and on the physical, chemical, and bio¬

logical activities in heavy red clay, brown clay,
and highly organic silt soils. He found that the
effect of sodium arsenite on plant growth depends
upon the resisting power of the plant and upon
the chemical and physical nature of the soil. In
small quantities, the compound stimulated plant
growth in most instances, but when added at the
rate of 0.1 to 0.25 per cent, it made plant growth
virtually impossible.

Sodium arsenite, he discovered, materially al¬
tered the mechanical condition of the soil, its ac¬
tion being primarily that of a deflocculating agent
checking the movement of water.

Sodium arsenite, McGeorge showed, was
strongly fixed by the soil, even resisting the wash¬
ing of heavy rains. An analysis of a sample of
soil from a tract of land sprayed three times a
year for 5 years, at the rate of 5 pounds of sodium
arsenite per acre per application, showed all the
arsenic to be present in the top 4 inches of soil.
The fixation of arsenites by the soil involved
chemical reactions consisting of replacement of
solution of iron, calcium, magnesium, and humus,
owing in part to a hydrolysis of the sodium arse¬
nite in solution and in part to a combination with
the dibasic and tribasic elements, to form the rela¬
tively insoluble arsenites and arsenates.

Brenchley (1914^; 191 4£) distinguished be¬
tween higher and lower forms of plant life in
their reactions to arsenic. In certain algae, stimu¬
lation may result from the presence of arsenic
compounds. Some fungi apparently are able to
live in the presence of arsenical compounds. On
higher plants, the toxic effect of arsenic was found
much more marked with arsenious acid and its
compounds than with arsenic acid and its deriva¬
tives. Using peas and barley, she could observe no
stimulation even with the smallest quantities used.

Morris and Swingle (1927) reported that the
addition of small amounts of soluble arsenical
compounds to potted plants caused serious injury
to most of the plants under test. The authors con¬
cluded that the incorporation of arsenical com¬
pounds in the soil is a dangerous practice, and
may cause considerable injury as the concentra¬
tion increases. They further concluded that beans
and cucumbers were very susceptible, whereas
cereals and grasses were more resistant.

To judge from the material so far presented, it
appears that the use of arsenic has several prob¬
lems associated with it. The preponderance of
opinion is that arsenic is harmful to higher plants
but that when the arsenic occurs in very small
concentrations, it may cause some stimulation.
Contrary opinion on the latter point is, however,
rather substantial. The stimulating effect of arsenic
on algae and fungi seems not uncommon, but its
favorable influence on soil organisms regarded as
useful to agriculture is at least questionable. While
the argument was going along on these somewhat
academic lines, great quantities of arsenic were

Arsenic Toxicity Studies — Clements and Munson

153

applied to orchards and fields. As Headden and
Swingle had warned, the continued application
and the resulting accumulations of arsenic in the
surface layers of soil and its retention there re¬
sulted in crop reduction or failure in Washington
as reported by Vandecaveye, Horner, and Keaton
(1936), in South Carolina as reported by Albert
and Paden (1931) and Cooper et al. (1931), and
in Louisiana as reported by Reed and Sturgis
(1936). During the past 15 years, experimental
work on arsenic has been directed to several prob¬
lems: the contamination of plant parts which are
used as animal or human food, the levels at which
arsenic in soil may cause a reduction in crop yields,
and methods which might be used to render a
poisonous soil suitable for renewed crop produc¬
tion.

Recent work: Arsenic in food. — Chorley and Mc-
Chlery (1935), interested in the poisoning of fowls
following their consumption of poisoned locusts,
reported that when arsenic is administered in
small quantities such as occur on sprayed grass¬
hoppers, a domestic fowl can tolerate compara¬
tively large doses over a long period without any
visible ill effects.

Franke and Maxon (1936) gave rats intraperi-
toneal injections of disodium acid arsenite and
disodium acid arsenate, as well as other chemicals.
The minimum fatal doses were defined as the
smallest doses which would kill 75 per cent or
more of the animals in less than 2 days. The mini¬
mum fatal doses of the arsenite were 4.25-4.75
mg. of arsenic per kilogram of weight, whereas for
the arsenate, the fatal dose was 14-18 mg. per
kilogram.

Groves, McCulloch, and St. John (1946) con¬
cluded from their studies that lead arsenate spray
residues are much less toxic to swine than has gen¬
erally been supposed. One pig consumed the spray
residue from 1,007 kilograms (over 1 ton) of
heavily sprayed apples which contained 114.8 gm.
of lead arsenate in the form of spray residue. The
pig gained in weight approximately as much as
the control pig, and no abnormalities were appar¬
ent in blood studies on it. Data showed that of
the edible portions of the pigs which were fed
large quantities of lead arsenate, only the livers
contained more lead than the 7.14 ppm legal limit,
and none of the organs analyzed contained more
than the limit of 3.57 ppm of As203.

The effect which spray residues on plant parts
and metabolized arsenic contained within plant
tissue may have on higher animals, including
humans, has been receiving increasing attention.
Talbert and Tayloe (1933), reporting results of
feeding spray chemicals to rats, concluded that if
it may be assumed that the spray chemicals have
an effect upon man similar to that which they
have on albino rats, it is their opinion that there
is little likelihood that a human being would con¬
sume as spray residue on apples, sprayed and

handled in the usual manner, enough arsenic
either at one time or over an extended period to
be injurious.

Coulson, Remington, and Lynch (1934) com¬
pared the bodies of rats fed for 3 to 5V2 months
on diets of varying arsenic content derived (1)
from natural shrimp and (2) from added arsenic
trioxide. The bodies of animals which had re¬
ceived the largest amount of arsenic, 17.9 mg. per
kilogram, in the form of shrimp contained at least
four times as much arsenic as the stock diet con¬
trols, whereas those which received approximately
the same quantity of arsenic in the form of arsenic
trioxide contained 55 to 65 times as much as the
controls. An even greater difference between the
storage of arsenic from the two forms was shown
during the first 3 months than after 5V2 months,
a fact suggesting that the rats receiving the inor¬
ganic arsenic had at some period during the first
3 months reached an equilibrium, after which no
further storage took place.

There was no retardation of growth in any of
the arsenic fed animals nor any observable differ¬
ences in their physical vigor or appearance, and
in none of them was there any histological evi¬
dence of injury to the spleen, liver, or kidney due
to the feeding of arsenic at the level employed.

These authors (1935) further emphasized the
difference between metabolized and inorganic ar¬
senic in foods by showing that arsenic as present
in shrimp was found to be far less available for
storage in young rats than when fed at the same
level as As203. During the first 3 months of their
feeding trial, 18 per cent of the As203 incorporated
in the diet at a level of 17.9 mg. per kg. was stored
as against 0.77 per cent for the same amount of
arsenic in shrimp. The total amount stored was
not significantly increased by feeding the element
for an added 9 months. There was no evidence of
toxicity from the arsenic in either form after 12
months of feeding.

In two human subjects studied by these authors,
the ingestion of shrimp in amounts furnishing
1,180 and 980 gammas of arsenic was followed
by rapid and complete elimination of the arsenic.
Inorganic arsenic, although excreted more slowly
than shrimp arsenic, was apparently eliminated
more completely by these subjects than by the rats.
These results are considered to be of interest not
only for the light which they throw on the meta¬
bolism of arsenic, but also as additional evidence
that the manner in which inorganic elements are
used in the body depends upon the source or form
in which these elements are presented.

It appears, therefore, that arsenic which has
been metabolized by an organism — that is, arsenic
which the organism has absorbed and made a part
of itself — is less dangerous to animals than is in¬
organic arsenic such as may appear on the leaves
of plants shortly after spraying. This latter arsenic
is quite poisonous to livestock, as most sections of

154

PACIFIC SCIENCE, Vol. I, July, 1947

an agricultural country recognize. In an effort to
determine how much arsenic plants might absorb
and remain normal in appearance, Machlis (1941)
grew bush beans and Sudan grass in culture solu¬
tion. He presented striking evidence that the bean
plant may absorb arsenic far in excess of legal
tolerance and yet show no reduction in growth.

To discover the factors which may or may not
make the use of arsenic-contaminated food safe
for animal or human consumption requires further
critical studies. On the basis of recent work it
appears, however, that there has been more emo¬
tion and less knowledge about this subject than
is needed for an understanding of it.

Recent work: Arsenic accumulation in soil , its
effect on crop production, and corrective measures.
— Swingle (1923), in an effort to test the effect of
prolonged application of arsenic on plant growth,
applied various arsenicals to plots of ground in
the spring of the year, and the effect on crop pro¬
duction was noted. After 7 years of such proce¬
dure, beans and cucumbers made little growth,
while wheat and timothy grew fairly well. No
further applications of arsenic were made for 6
years and at the end of that time it was found
that very little of the arsenical had been removed
by rains or irrigation. Furthermore, it was found
very difficult to get the land back into condition
for cropping.

Paden and Albert (1930), working in South
Carolina, reported a relationship between the un¬
productivity of certain soil types and the accumu¬
lation of soluble arsenic in the soil resulting from
heavy applications of calcium arsenate. Lime im¬
proved the growth of cotton in the poisoned soil.
Soils which were relatively low in iron and other
materials which would be expected to render ar¬
senic insoluble were found to be the most seriously
affected by the arsenates.

Albert and Arndt (1931) found the concentra¬
tion of soluble arsenic as measured by collodion
bag dialyzates to be a more reliable index of ar¬
senic than is the total arsenic present in the soil.
In greenhouse experiments, the addition of 1 ppm
of arsenic definitely retarded root and top growth
of cowpeas. It was observed that the concentra¬
tion of 1 ppm of soluble arsenic as measured by
the collodion bag test was not unusual in soil
which had been receiving doses of arsenates. Lim¬
ing and the use of fertilizer along with iron and
clay compounds of the soil played an important
role in rendering arsenates harmless to sensitive
crops.

Hurd-Karrer (1936) believed as a result of field
tests and culture solutions that phosphate applica¬
tion will reduce or prevent arsenic injury to plants
where the soil type is such as to retain the phos¬
phate in available form.

Heggeness (1940), growing tomatoes in culture
solution, found no evidence of toxic stimulation,
even the most dilute solution (Y2 ppm) reducing

the yield by 20 per cent. Arsenic toxicity in the
tomato was partially dependent on phosphate
availability.

Keaton (1938) studied the oxidation-reduction
potentials of arsenate-arsenite systems in sand and
soil mediums. In these soils arsenic was fixed by
absorption and combination, but it was observed
that a higher percentage of arsenate than arsenite
was fixed by these soils. The addition of iron to
the system when the original ratio of arsenate to
arsenite was unity increased the redox potential
independent of the medium used. In the two soils
studied the colloidal fraction possessed a greater
reducing capacity and a lower potential than the
soil from which it was extracted.

Keaton and Kardos (1940) attempted treat¬
ment of orchard soil to overcome toxicity of arse¬
nic residues. Their studies indicate a relationship
between the oxidation-reduction potentials of the
soils treated and the conditions of plant growth.
The addition of ferric oxide caused an increase in
the redox potential. Alumina produced no effect
in oxidation or reduction. Soils with a high colloid
content were characterized by low potential and
small percentage oxidation. They suggested that
poisoned soils be treated with some mild oxidizing
agent capable of arsenic fixation, and named iron
oxide as such an agent.

Kardos, Vandecaveye, and Benson (1940) re¬
ported that severely toxic soils have been rendered
productive by making applications of 3 to 4y2
tons of ferrous sulfate per acre.

EXPERIMENTAL WORK

In order to study certain phases of arsenic
toxicity, the experiments reported in this
paper were undertaken. Data will be pre¬
sented and discussed under the following
headings :

Part I. The comparative toxicity of triva-
lent and pentavalent arsenic on
bean ( Phaseolus vulgaris L.), Su¬
dan grass ( Sorghum vulgare Per-
soon var. sudanense [Piper]
Hitch.), and tomato ( Lycopersicon
esculentum Miller).

Part II. The effect of different phosphorus
levels on the toxicity of trivalent
and pentavalent arsenic on bean,
Sudan grass, and tomato.

Part III. The influence of repeated crop¬
pings of bean, Sudan grass, and
tomato on a red residual soil and

Arsenic Toxicity Studies — Clements and Munson

155

on a black alluvial soil treated with
various increments of arsenic.

MATERIALS AND METHODS

Culture solutions. — For Part I and Part II
mentioned above, water cultures were used.
A total of 30 crocks was used for each of the
three species — 10 crocks at each of three
phosphorus levels — with each crock having
a known concentration of arsenic in the form
of sodium arsenite in the study of the triva-
lent form and sodium arsenate in the study
of the pentavalent form.

The experiments were conducted in a
greenhouse of the University of Hawaii
Agricultural Experiment Station. Four-
gallon size glazed crocks were used for the
Sudan grass and tomato samples, and 2 -gal¬
lon size for the bean. Distilled water was
used throughout and continuous aeration
maintained. The plants were held in corks
set into lids which fitted over the crocks.

The tomato and Sudan grass seeds were
germinated on cheesecloth and transferred
when large enough to facilitate handling
without injury. The bean seeds were germi¬
nated in black sand and transplanted as soon
as was practicable.

All the plants were given complete nutri¬
ents for 2 weeks. The following solutions
were used:

0 SUDAN

TOMATO GRASS BEAN

Ca(N03)2 . 005M .005M .005M

KNOs . 005M .005M .005M

MgSOi . 002M .002M .002M

CaCl2 . .003M .003M -

KHsPCL . . 001M .001M .002M

Iron (as FeS04) and standard amounts of
copper, manganese, zinc, and boron were
added.

After 2 weeks passed the solutions were
changed, but the nutrients as listed above
were maintained with the exception of
the phosphate, which was added in the
amount required in the experiment. The
arsenic increments were added as planned,

and the plants were selected for uniformity
and thinned to three plants per crock for
the bean and Sudan grass and two plants for
the tomato. All solutions were changed
weekly, and the levels of all salts were
maintained as required in the experiment.

Plants which did not live until the end
of the experiment were removed when they
died. All others were allowed to grow for
3 to 4 weeks after the original treatment;
then they were harvested. Green weights of
the plant tops were taken, after which the
plants were dried and dry weights taken.
The material was ground in a Wiley mill
and stored for subsequent arsenic and phos¬
phate analyses.

Methods of analysis. — The method of Cas-
sil and Wichmann (1939) was used in
making the arsenic determinations reported
in this paper. For the phosphate determina¬
tions, the Truog and Meyer (1929) modi¬
fication of the Deniges method was used.

The dry plant material was digested as
follows: Two grams or less of the sample
were added to 125 cc. Erlenmeyer flasks, fol¬
lowed by 10 cc. of H2S04 and 20 cc. of
HN03. The flasks were allowed to stand for
several hours, then heated slowly. Five-cc.
increments of HNOs were added, and the
digestion continued until the solution was
clear and yellow. A final 5 cc. of HNOs was
added with 5 drops of perchloric acid and
heat was applied until dense white SOs
fumes were evolved. The flask was removed
and cooled, and the solution made up to
125 cc. From this, suitable aliquots were
taken for the arsenic and phosphate deter¬
minations.

DATA AND DISCUSSION
Part I. Comparative Toxicity of Trivalent

and Pentavalent Arsenic

In Tables 1, 2, and 3 are presented data
for experiments using the three species
treated with sodium arsenate (Na2HAs04.

156

PACIFIC SCIENCE, Vol. I, July, 1947

TABLE 1. TOMATO

Data for the accumulation of arsenic (as ppm

AS2O3) AND PHOSPHORUS (AS PER CENT OF DRY

weight), as well as dry weights of plants

GROWN IN SOLUTIONS CONTAINING VARIOUS IN¬
CREMENTS OF PENTAVALENT ARSENIC (AS SODIUM
ARSENATE) AT DIFFERENT PHOSPHATE LEVELS.

ARSENIC

IN

SOLUTION

DRY

WEIGHT

ARSENIC

IN

PLANT

PHOS¬

PHORUS

IN PLANT

ppm AS2O3

gm.

ppm AS2O3

% dry wt.

Low phosphorus level (P = 10 ppm)

0.0

27

trace

0.72

1

30

5.0

.67

2.5

34

10.6

.61

5

21

22.5

.75

10

18

42.2

.75

15

10

76.5

.72

20

10

81.3

.67

25

6

93.0

.50

30

3

106.3

.72

40

2

162.5

.60

Medium phosphorus level (P = 60

ppm)

0.0

28

0

.75

6

29

6.7

.72

15

19

17.2

.84

30

18

49.7

.84

60

9

74.1

.69

90

4

120.1

.63

120

2

193.7

.60

High phosphorus level (P = 120 ppm)

0.0

28

trace

.72

12

23

12.2

.78

30

19

26.6

.69

60

11

61.3

.64

120

6

103.9

.60

7 H20), and in Tables 4, 5, and 6 the data
for the same species treated with sodium
arsenite (NaAs02).

Since it will be shown later that the phos¬
phorus level has an important influence on
arsenic toxicity, comparisons between arsen¬
ate and arsenite must be made within series
of similar phosphorus concentration. Com¬
parisons of the arsenate and arsenite experi¬
ments at the medium phosphorus level
(P = 60 ppm) indicate marked differences
between the action of pentavalent and of
trivalent forms.

The tomato plants growing in 60 ppm of
pentavalent arsenic accumulated 74.1 ppm
of arsenic in their tops, yet were normal ex¬
cept in size. Tomato plants grown in solu¬
tions containing 90 ppm and 120 ppm of

pentavalent arsenic lived to the end of the
experiment despite accumulations of 120
ppm and 194 ppm of arsenic in their tops.
The lethal concentration was between 120
ppm and 150 ppm in the culture solution.
In the arsenite series, on the other hand,
concentrations of 7 ppm and 1 1 ppm in the
solution very nearly stopped growth, al¬
though the plants on analysis showed only
12.5 ppm and 24.1 ppm of arsenic respec¬
tively in their tops. Even the low level of
3 % ppm in the culture caused marked stunt¬
ing; the lethal concentration was 11 to 15
ppm. Approximately 10 times as much
pentavalent arsenic is required in the culture
solution and in the plant tissue as trivalent
arsenic to produce equivalent injuries to
tomato plants.

TABLE 2. SUDAN GRASS

Data for the accumulation of arsenic (as ppm
AS2O3) AND PHOSPHORUS (AS PER CENT OF DRY
WEIGHT), AS WELL AS DRY WEIGHTS OF PLANTS
GROWN IN SOLUTIONS CONTAINING VARIOUS IN¬
CREMENTS OF PENTAVALENT ARSENIC (AS SODIUM
ARSENATE) AT DIFFERENT PHOSPHATE LEVELS.

ARSENIC

IN

SOLUTION

DRY

WEIGHT

ARSENIC

IN

PLANT

PHOS¬

PHORUS

IN PLANT

ppm AS2O3

gm.

ppm AS2O3

% dry wt.

Low phosphorus level (P = 10 ppm)

0.0

40

trace

0.61

1

23

13.5

.53

2.5

21

,19.1

.56

5

11

47.4

.42

10

5

67.2

.44

15

2

88.8

.42

20

2

103.1

.53

25

2

139.1

.54

Medium

phosphorus level (P =

60 ppm)

0.0

37

trace

.72

6

29

29.4

.75

15

18

41.0

.75

30

8

74.1

.39

60

4

98.4

.56

90

2

513.1

.66

High phosphorus level (P = 120 ppm)

0.0

36

trace

.84

12

28

12.8

.91

30

20

49.7

.78

60

14

84.4

.59

120

6

142.3

.52

Arsenic Toxicity Studies — Clements and Munson

157

TABLE 3. BEAN

Data for the accumulation of arsenic (as ppm
AS2O3) AND PHOSPHORUS (AS PER CENT OF DRY
WEIGHT), AS WELL AS DRY WEIGHTS OF PLANTS
GROWN IN SOLUTIONS CONTAINING VARIOUS IN¬
CREMENTS OF PENTAVALENT ARSENIC (AS SODIUM
ARSENATE) AT DIFFERENT PHOSPHATE LEVELS.

ARSENIC

IN

SOLUTION

DRY

WEIGHT

ARSENIC

IN

PLANT

PHOS¬

PHORUS

IN PLANT

ppm AS2O3

gm.

ppm AS2O3

% dry wt.

Low phosphorus level (P = 10 ppm)

0.00

50

trace

0.64

.1

47

2.5

.55

.25

38 .

2.8

.51

.5

38

2.5

.50

1.5

28

10.3

.66

2.0

15

13.7

.58

2.5

11

20.0

.53

Medium phosphorus level (P =

60 ppm)

0.0

44

trace

.59

.6

48

1.6

.59

1.5

43

2.7

.60

3

35

4.1

.63

6

38

5.1

.66

9

26

7.2

.61

High phosphorus level (P = 120 ppm)

0.0

57

trace

.63

1.2

54

1.9

.59

3

38

2.7

.66

6

37

5.0

.63

12

16

38.0

.72

The Sudan grass plants growing in solu¬
tions containing 30 ppm and 60 ppm of
pentavalent arsenic were apparently normal
except for size, although they accumulated
74.1 and 98.4 ppm of arsenic in their tops.
Marked injury was noted at 90 ppm; the
lethal concentration was between 90 ppm
and 120 ppm. With trivalent arsenic, how¬
ever, a concentration as low as 3% ppm of
arsenic in the culture solution resulted in
cessation of growth, although the plant tops
showed only 14.9 ppm of arsenic. The lethal
concentration in the culture solution of the
trivalent arsenic was between 11 ppm and
15 ppm. Sudan grass can tolerate approx¬
imately 10 times as much pentavalent arsenic
in the culture solution and in their tissues as
trivalent arsenic.

The bean plants were very intolerant of
arsenic in any form. They grew quite nor¬

mally, however, in solutions containing 9
ppm of pentavalent arsenic, from which
they accumulated 7.2 ppm of arsenic in their
tops. The lethal concentration was between
1 2 ppm and 1 5 ppm of pentavalent arsenic.
On the other hand, growth was prevented
by a concentration of 2.25 ppm to 2.85 ppm
of trivalent arsenic, despite an accumulation
of only 4.9 ppm of arsenic in their tops. The
lethal concentration of trivalent arsenic in
the culture solution was between 2.85 ppm
and 3.6 ppm. Trivalent arsenic is roughly
four times as toxic to bean plants as is the
pentavalent form.

Comparisons made between the arsenate
and arsenite series at the high and low phos¬
phorus levels show results similar to those
discussed above for the intermediate phos¬
phorus level.

The two forms of arsenic differ not only
in lethal concentrations, but also in their
immediate action on plant tissues. Triva¬
lent arsenic has a violent action, causing
complete disintegration of the roots and
burning of the tops in 1 or 2 days in lethal
concentrations. Pentavalent arsenic, on the
other hand, often takes several days to pro¬
duce any response other than wilting, even
in concentrations that eventually prove
lethal.

Part II. The Effect of Different

Phosphorus Levels on the Toxicity of

Trivalent and Pentavalent Arsenic

It was suggested by Hurd-Karrer (1939)
that a relationship exists between arsenic
toxicity and phosphorus availability. She
used sodium arsenate in culture solutions at
different phosphorus levels on studies with
the oat plant and came to the conclusion that
"in general, the arsenic was definitely toxic
in the presence of less than four times as
much phosphorus but non-toxic when there
was more than four times as much.”

Based on that observation, investigations
were undertaken to compare the effect of

158

PACIFIC SCIENCE, Vol. I, July, 1947

phosphorus level on trivalent and penta-
valent arsenic to determine whether the de¬
crease in toxicity was due to decreased
absorption of the toxic element in the pres¬
ence of large amounts of phosphorus or
whether it was due to some inhibitory effect
which phosphorus might have on arsenic
toxicity after absorption by the plant tissues.

Bean, Sudan grass, and tomato plants
were grown in culture solutions at three
phosphorus levels — 10 ppm, 60 ppm, and
120 ppm — and subjected to treatment with
sodium arsenate and sodium arsenite.

In Tables 1, 2, and 3 are recorded the
data for the tests with pentavalent arsenic.
It can be seen that an increase in the phos¬
phorus level markedly reduced the amount
of pentavalent arsenic absorbed, and re¬
sulted in better growth. For example, the
tomato plants grown in low-phosphorus
solutions containing 30 ppm of arsenic con¬
tained 106.3 ppm of arsenic in their tops,
as compared to 49.7 ppm and 26.6 ppm in
plants grown in medium- and high-phos¬
phorus solutions, respectively, containing
similar amounts of arsenic.

Similarly, increases in the phosphorus
level reduced the absorption of arsenic by
Sudan grass. The analyses of plant tops
showed 88.8 ppm, 41.0 ppm, and 20.0
ppm of arsenic from the low-, medium-,
and high-phosphorus solutions containing
15 ppm of arsenic.

The relationship of the phosphorus level
to arsenic absorption by the bean is not so
decisive. The bean has a very small range
of tolerance to arsenic and, as a result, the
magnitude of the differences between the
various cultures is correspondingly small.
The beans showed, for example, 10.3 ppm,
2.7 ppm, and 2.0 ppm of arsenic in their
tops when grown in low-, medium-, and
high-phosphorus solutions, respectively,
containing 1.5 ppm of arsenic.

As might be expected, a reduction in the
absorption of the toxic element by the plant

resulted in better plant growth. The con¬
trols showed no significant differences in the
dry weights at the three phosphorus levels,
indicating that 10 ppm of phosphorus were
adequate for the three plants studied. Analy¬
sis of the plant tops showed no significant
differences among plants grown in low-,
medium-, and high-phosphorus levels.
Furthermore, the dry weights seemed to cor¬
relate with the arsenic concentration in the
plant, irrespective of the phosphorus level
of the culture solution. These considerations
would seem to indicate that the phosphorus
in the culture solution is effective in inhibit¬
ing the absorption of pentavalent arsenic by
the plant but not in reducing the toxicity of
the element within the plant.

Hurd-Karrer (1939) suggested that
arsenates would be non-toxic if the P : As
ratio were more than 4:1. Although a high
phosphorus level greatly reduced the toxic
effects of a given concentration of arsenic in
the culture solution by^reducing its absorp¬
tion by the three plants studied here, it did
not prevent injury. For example, tomatoes
growing in solutions in which the P : As
ratio was 10 : 1 (120 ppm : 12 ppm) had
a dry weight of 23 grams as compared to 28
grams in the control — a reduction of 18 per
cent. Sudan grass in solutions containing
120 ppm of phosphorus and 12 ppm
of arsenic (P : As = 10 : 1) weighed 28.5
grams as compared to, 36 grams in the con¬
trol — a reduction of 21 per cent. Bean
plants growing in solutions containing 6
ppm of arsenic and 120 ppm of phosphorus
(P:As = 20:l) weighed 37 grams as com¬
pared to 57 grams in the control — a reduc¬
tion of 35 per cent. A high phosphorus level
reduces but does not prevent the absorption
of pentavalent arsenic. The degree of injury
depends on the amount of the toxic element
absorbed.

The data for the studies made with tri¬
valent arsenic are recorded in Tables 4, 5,
and 6. The results indicate that the action

Arsenic Toxicity Studies — Clements and Munson

159

TABLE 4 TOMATO

Data for the accumulation of arsenic (as ppm

AS2O3) AND PHOSPHORUS (AS PER CENT OF DRY
WEIGHT), OF PLANTS GROWN IN SOLUTIONS CON¬
TAINING VARIOUS INCREMENTS OF TRIVALENT AR¬
SENIC (AS SODIUM ARSENITE) AT DIFFERENT PHOS¬
PHORUS LEVELS.

ARSENIC

IN

SOLUTION

DRY

WEIGHT

ARSENIC

IN

PLANT

PHOS¬

PHORUS

IN PLANT

ppm AS2O3

gm.

ppm AS2O3

% dry wt.

Low phosphorus level (P 10 ppm)

0.00

29

trace

0.66

.25

30

trace

.67

.50

31

trace

.69

.75

30

trace

.66

1.00

28

1.6

.77

1.75

26

1.3

.76

2.50

20

3.9

.78

3.25

10

4.7

.70

4.00

9

10.9

.81

5.00

4

12.0

.81

Medium phosphorus level (P =

60 ppm)

0.00

32

trace

.75

1.00

29

trace

.77

2.00

24

3.3

.75

3.25

9

6.7

.84

7.00

1

12.5

.69

11.00

1

24.1

.50

High phosphorus

level (P rr 120 ppm)

0.00

33

trace

.85

1.00

28

trace

.88

3.25

7

7.8

.88

7.25

3

14.7

.84

of phosphorus on the absorption of trivalent
arsenic is quite different from its action on
pentavalent arsenic. From a given concen¬
tration of trivalent arsenic, the tomato plants
absorbed approximately the same amount of
the toxic element, irrespective of the phos¬
phorus level, and showed equal degrees of
injury. In the studies with Sudan grass, the
medium- and the high-phosphorus levels
reduced the absorption of arsenic over that
absorbed from the low level of phosphorus,
with a corresponding reduction in injury.
In the studies with the bean plants, the phos¬
phorus level had some effect on the absorp¬
tion of arsenic as shown in the analysis of
the plant material, although it should be
noted that the differences are so small as to
be insignificant.

These studies reveal, therefore, that the
form in which the arsenic occurs is an im¬
portant factor in determining the effect of
phosphorus. Hurd-Karrer (1937) in her
studies on the antagonism of related ions
found that sulfates more effectively reduced
the absorption of selenium from selenates
than from selenites. From these results she
suggested that "by analogy, phosphates
would be expected to have less effect on the
toxicity of arsenite than on that of arsenate"
(1939). The studies reported in this paper
confirm that supposition.

Part III. Toxic Levels of Arsenic
in Certain Hawaiian Soils

It is well known that arsenicals, when ap¬
plied to soil, are far less available to plants

TABLE 5. SUDAN GRASS
Data for the accumulation of arsenic (as ppm
AS2O3) and phosphorus (as per cent of dry

weight), of plants grown in solutions con¬
taining VARIOUS INCREMENTS OF TRIVALENT AR¬
SENIC (as sodium arsenite) at different phos¬
phorus LEVELS.

ARSENIC

IN

SOLUTION

DRY

WEIGHT

ARSENIC

IN

PLANT

PHOS¬

PHORUS

IN PLANT

ppm AS2O3

gm.

ppm AS2O3

% dry wt.

Low

phosphorus

level (P = 10 ppm)

0.00

63

trace

0.44

.25

63

5.2

.36

.50

48

7.8

.59

.75

48

11.9

.45

1.00

32

18.6

.56

1.75

17

20.7

.56

2.50

5

23.1

.73

3.25

4

22.7

.48

4.00

2

42.5

.55

5.00

3

46.9

.88

Medium phosphorus level (P =£

60 ppm)

0.00

63

trace

.94

1.00

39

8.6

.58

2.00

24

10.9

.53

3.25

4

14.9

.56

7.00

3

27.5

.67

11.00

2

91.2

.81

High phosphorus

level (P = 120 ppm)

0.00

68

trace

.98

1.00

45

4.1

.88

3.25

18

12.8

.99

7.50

8

32.5

.77

15.00

2

87.5

.81

160

PACIFIC SCIENCE, Vol. I, July, 1947

TABLE 6. BEAN

Data for the accumulation of arsenic (as ppm
As203) and phosphorus (as per cent of dry
weight), as well as dry weights, of plants

GROWN IN SOLUTIONS CONTAINING VARIOUS IN¬
CREMENTS OF TRIVALENT ARSENIC (AS SODIUM
ARSENITE) AT DIFFERENT PHOSPHORUS LEVELS.

ARSENIC

IN

SOLUTION

DRY

WEIGHT

ARSENIC

IN

PLANT

PHOS¬

PHORUS

IN PLANT

ppm AS2O3

gm.

ppm AS2O3

% dry wt.

Low phosphorus level (P = 10 ppm)

0.000

42

trace

0.53

.038

49

trace

* .47

.075

58

trace

.44

.113

49

0.9

.44

.150

41

1.2

.50

.263

27

0.9

.69

.375

25

2.1

.63

.488

26

2.1

.56

.600

23

3.9

.63

.750

30

3.7

.50

Medium phosphorus level (P =

60 ppm)

0.00

36

trace

.70

.3

3 4

trace

.70

.49

27

trace

.63

1.05

27

1.3

.63

1.65

15

4.8

.69

2.25

6

4.9

.69

2.85

5

7.0

.66

High phosphorus level (P = 120 ppm)

0.00

31

trace

.63

.15

31

trace

.72

.49

30

trace

.59

1.13

15

trace

.66

2.25

5

3.3

.75

than they are when applied to culture solu¬
tion. The degree to which arsenicals are
fixed by the soil is a characteristic of the soil
(Crafts and Rosenfels: 1939). The objec¬
tives of this part of the work are: first, to
determine the growth reaction of certain
crop plants to arsenic levels in two Hawaiian
soils; second, to determine the amount of
arsenic which these plants withdraw from
the soil; and third, to determine whether or
not it is practicable to use certain crop plants
to lower the arsenic levels of those soils
which have been rendered sterile to other
crop plants.

Methods. — Two soils were used — one a red
residual clay and the other a black alluvial
soil. The red is a residual soil taken from

the mountain slopes above Kailua, Oahu. It
is an infertile soil which requires heavy fer¬
tilization for crop production. The black
alluvial soil, taken from a papaya orchard
near Kailua, is, on the other hand, ex¬
tremely fertile. In fact, it was recommended
to us by Dr. L. A. Dean, Soils Chemist, as
being a soil whose available phosphorus
level was so high that no more phosphorus
could be fixed by it.3 The growth of plants
in the two soils reflected not only the differ¬
ence in chemical composition, but also the
difference in physical qualities, the black
soil being very well adapted to pot work.
These soils contained 14.7 ppm of native
arsenic. This amount is added to the incre¬
ments of soil arsenic shown in the accom¬
panying tables.

The soil was dried thoroughly, screened,
and ground in a plate mill. Samples of 500
grams each were weighed into No. 2 cans,
after which arsenic as sodium arsenite was
added in concentrations varying from 10
ppm to 3,000 ppm following the methods
described by Crafts and Rosenfels (1939).
The soil was allowed to dry thoroughly,
after which it was removed, pulverized again,
mixed, and returned to the can.

With the red soil, triplicate cans were
used for each arsenic concentration for each
of the three species, making a total of 225
cans. With the black soil, only one container
was used for each of the two species used
for each concentration of arsenic.

Tomato and Sudan grass seedlings were
started on cheesecloth and transplanted to
the cans when a few days old. Bean seeds
were germinated in black sand and trans¬
planted to the cans as soon as possible. With
tomato and bean, two plants per can were
used, and with Sudan grass five plants.

Drainage was provided by punching holes
in the cans. A complete nutrient solution

3 The phosphorus content of the two soils as ex¬
tracted with 0.002N H2SO4 was 24 ppm for the
red soil and 250 ppm for the black soil.

Arsenic Toxicity Studies— Clements and Munson

16!

*SO/L - CULTURE CAM MUM BCR

Fig. 1. Green weights and arsenic content of tomato plants grown in red soil with arsenic content
ranging from 15 ppm (Can 1) to 3,014 ppm (Can 24) of As203. The difference in levels of the two
green-weight curves is due to the season in which the plants were grown and is not related to treat¬
ment. (For As203 increments, see Table 7.)

was added weekly to the soil. Each crop
was allowed to grow 31 days from the time
of transplanting, after which the plant tops
were collected and green weights obtained.
The material was dried, ground in a Wiley
mill, and stored for future analysis.4

After the removal of one crop, the soil
was allowed to dry thoroughly, after which
each can was emptied and the soil pulver¬
ized, mixed, and returned. Roots were in
each case returned to the bottom of the can.

Results and discussion: Tomato. — The data

4 All the arsenic analyses made in connection
with these soil studies were made by the Chemis¬
try Department of the Experiment Station, Ha¬
waiian Sugar Planters’ Association. The depart¬
ment is directed by Dr. F. E. Hance, to whom sin¬
cere thanks are due.

presented in Table 7 and shown in Figure 1
for tomato plants in red soil reveal the rela¬
tive tolerance of young plants to soil arsenic
up to approximately 514 ppm. At higher
levels the ability of the plant to grow is dras¬
tically reduced to what would be growth
failure in a commercial field.

At levels below 514 ppm, growth of the
tomato is approximately at uniform levels,
no matter what the level of arsenic in the
soil. While there may be some slight evi¬
dence of stimulation, it is very inconclusive.
The arsenic content of the plant tissues
varies between 2 and 3 ppm for soil levels
of arsenic between 15 and 3 14 ppm. Between
this point and approximately 414 ppm of
soil arsenic, the arsenic level rises gradually

162

PACIFIC SCIENCE, Vol. I, July, 1947

TABLE 7. TOMATO-RED SOIL

Yield Data and Arsenic Content of Tissues.

CAN

NUMBER

As203

CONTENT

OF SOIL

SERIES I

AVERAGE OF 4 LATER SERIES

Green weight
of plants

As203 content
of tissues

Green weight
of plants

As203 content
of tissues

ppm

gm. per can

ppm

gm. per can

ppm

1

15

25.7

1.8

12.8

0.9

2

25

30.0

1.6

12.5

1.3

3

34

29.7

1.6

13.1

1.1

4

45

27.7

2.4

12.3

1.3

5

54

» 26'7

1.6

13.4

1.5

6

65

25.3

1.8

13.2

1.1

7

74

27.7

2.4

14.0

1.5

8

85

27.0

2.4

12.8

1.7

9

95

28.0

2.6

13.7

1.5

10

104

33.0

2.6

15.7

1.3

11

115

29.0

2.6

14.0

1.8

12

214

30.0

2.6

13.9

2.1

13

314

28.7

3.4

12.2

2.9

14

414

26.7

3.7

13.1

3.0

15

514

26.7

5.8

13.7

3.0

16

614

23.6

6.9

11.3

4.9

17

714

8.7

21.7

7.3

4.1

18

814

4.4

52.3

5.9

4.9

19

915

5.3

29.6

6.6

6.1

20

1014

2.0

6.3

8.1

21

1514

3.1

5.4

9.0

22

2014

1.6

4.8

19.9

23

2515

0.9

3.2

26.9

24

3014

0.8

.

2.7

52.7

TABLE 8. TOMATO-BLACK SOIL

Yield Data and Arsenic Content of Tissues.

CAN

NUMBER

As203

CONTENT

OF SOIL

SERIES I

AVERAGE OF 2

LATER SERIES

Green weight
of plants

As203 content
of tissues

Green weight
of plants

As203 content
of tissues

ppm

gm. per can

ppm

gm. per can

ppm

1

15

29.0

2

25

30.0

1.6

38.3

0.8

3

34

32.0

1.3

39.8

1.5

4

45

27.0

1.8

41.0

1.5

5

54

29.0

2.1

39.4

1.7

6

65

26.0

2.9

40.2

2.1

7

74

32.0

5.0

38.7

1.2

8

85

28.0

2.4

41.2

2.0

9

95

29.0

2.1

37.5

1.5

10

104

30.0

2.1

39.9

1.6

11

115

29.0

5.8

34.0

1.8

12

214

30.0

2.6

33.0

2.6

13

314

31.0

3.4

39.0

2.1

14

414

28.0

4.0

38.3

2.1

15

514

29.0

5.8

34.4

3.7

16

614

31.0

5.8

35.8

5.8

17

714

28.0

6.9

31.5

9.1

18

814

24.0

8.5

34.7

4.8

19

915

25.0

8.7

38.8

11.1

20

1014

30.0

9.2

31.5

15.8

21

1514

27.0

29.6

35.9

11.9

22

2014

25.0

64.2

29.5

30.5

23

2515

22.0

103.8

28.5

35.1

24

3014

20.0

76.1

32.3

50.0

Arsenic Toxicity Studies — Clements and Munson

163

Fig. 2. Green weights and arsenic content of tomato plants grown in black soil with arsenic content
ranging from 15 ppm (Can 1) to 3,014 ppm (Can 24) of As203. The difference in levels of the two
green-weight curves is due to the season in which the plants were grown and is not related to treat¬
ment. (For As203 increments, see Table 8.)

from 3 ppm to 4 ppm. From higher levels
of soil arsenic the plant absorbs increasing
amounts of the poison, and this heavier ab¬
sorption is reflected in greatly reduced
growth.

Although the level of soil arsenic which
might be described as critical for the tomato
is the same for the later crops of plants as
it is for the first crop, there is a striking dif¬
ference in the amounts of arsenic absorbed
by the later crops. Thus, the first crop in
Can 18 contained about 52 ppm of arsenic,
while the later crops in Can 18 absorbed
about one tenth as much. A very much
higher level of soil arsenic was necessary for
the later crops to absorb 52 ppm. Two pos¬
sible hypotheses suggest themselves. First,

the fixation of arsenic by the soil may be a
function not only of the nature of the soil
but of time. Second, the soluble arsenic
level of the soil may be reduced sufficiently
by the first crop to reduce its absorption by *
later crops. However, the fact that growth
in the first series as well as that in the later
series is reduced in the same cultures does
not lend support to the latter hypothesis.

Tomato: Black Soil. — The growth of
young tomato plants in black soil, as shown
in Table 8 and Figure 2, is very much better
than it is in the red soil. The black soil is
not only very fertile, but it possesses physical
qualities which make it a better soil for pot
work. The most striking contrast between
the two soils is that in the black soil there

164

PACIFIC SCIENCE, Vol. I, July, 1947

is no sharp curtailment of growth even in
the cans having over 3,000 ppm of soil
arsenic. Although the tomatoes growing in
the cans at the upper concentration levels
were slightly inferior in size to those in the
lower levels, they were otherwise normal in
every respect. They showed no premature
drying or yellowing of leaves, their roots
were as extensive as the others, and yet the
arsenic levels in the plant tissues were very
high. In Can 24, the first crop of plants
appeared perfectly normal with good color,
despite the fact that the crop contained 76.1
ppm of arsenic. Like those in the red soil,
though to a lesser degree, later crops of
tomatoes in black soil failed to extract as
much arsenic from the high arsenic soil as
did the first crop.

Sudan Grass: Red Soil. — Sudan grass
plants, as shown in Table 9 and Figure 3,
grew uniformly well in all cans up to Can
10, in which the arsenic content of the soil
was 104 ppm. There is no suggestion of
stimulation in the cultures having lower con¬
centrations of arsenic. At the higher levels
of soil arsenic, Sudan grass extracted sub¬
stantially higher amounts of the poison than
did the tomato. The first crop of grass ab¬
sorbed larger quantities of arsenic at the
higher levels than did later crops. In fact,
each succeeding crop of Sudan absorbed less
and less arsenic from the soil, but in each
crop the depression of growth occurred at
the same level of soil arsenic, a fact again
pointing to the idea that the actual arsenic
level in the plant is not causal, though it is

Fig. 3. Green weights and arsenic content of Sudan grass grown in red soil with arsenic content
ranging from 15 ppm (Can 1) to 3,014 ppm (Can 24) of As203. The difference in levels of the two
green-weight curves is due to the season in which the plants were grown and is not related to treat¬
ment. (For As203 increments, see Table 9.)

Arsenic Toxicity Studies — Clements and Munson

165

TABLE 9. SUDAN GRASS-RED SOIL
Yield Data and Arsenic Content of Tissues.

CAN

NUMBER

AS2O3

CONTENT

OF SOIL

SERIES I

AVERAGE OF 2

LATER SERIES

Green weight
of plants

As2Os content
of tissues

Green weight
of plants

As203 content
of tissues

ppm

gm. per can

ppm

gm. per can

ppm

1

15

20.8

0.5

15.7

1.5

2

25

19.0

1.6

15.7

1.5

3

34

18.7

0.8

15.5

1.3

4

45

19.0

1.6

15.1

1.6

5

54

19.0

1.3

16.0

1.3

6

65

18.3

2.6

15.8

3.8

7

74 -

18.7

4.5

15.6

2.4

8

85

20.0

2.1

16.3

3.0

9

95

19.7

2.6

14.8

3.7

10

104

18.3

16.6

2.9

11

115

17.3

4.0

11.9

3.6

12

214

12.7

8.2

11.9

4.6

13

314

11.3

10.3

12.2

6.5

14

414

8.7

14.5

6.1

6.3

15

514

4.7

23.2

6.3

12.3

16

614

2.2

31.4

4.9

14.1

17

714

0.3

1.7

21.8

18

814

0.6

1.0

35.5

19

915

0.2

0.9

28.1

20

1014

0.2

0.8

46.1

21

1514

0.1

0.4

22

2014

0.2

23

2515

0.2

24

3014

0.1

TABLE 10. SUDAN GRASS-BLACK SOIL

Yield Data and Arsenic Content of Tissues.

CAN

NUMBER

As2Oa

CONTENT

OF SOIL

SERIES I

AVERAGE OF 5

LATER SERIES

Green weight
of plants

As2C>3 content
of tissues

Green weight
of plants

As203 content
of tissues

ppm

gm. per can

ppm

gm. per can

ppm

1

15

25.3

2.4

36.8

1.3

2

25

22.6

5.5

33.2

1.2

3

34

23.8

5.0

31.6

1.8

4

45

25.8

4.2

32.4

1.6

5

54

23.2

3.4

33.6

1.2

6

65

24.6

3.4

32.8

2.2

7

74

27.3

34.5

1.3

8

85

25.8

3.7

31.1

1.6

9

95

25.2

1.3

34.1

2.1

10

104

24.6

3.4

34.9

2.1

11

115

22.7

5.8

36.3

2.4

12

214

26.4

2.1

34.3

5.4

13

314

25.6

0.4

29.6

7.7

14

414

20.5

9.2

30.0

10.7

15

514

17.7

16.1

27.4

13.5

16

614

23.7

14.0

29.0

12.9

17

71 4

13.5

18.5

27.1

17.6

18

814

10.5

17.4

24.6

17.4

19

915

6.8

16.1

24.4

24.6

20

1014

10.9

24.0

17.6

27.3

21

1514

4.8

28.5

10.4

29.4

22

2014

0.9

44.6

2.4

56.1

23

2515

0.1

1.0

86.5

24

3014

0.2

0.7

166

PACIFIC SCIENCE, Vol. I, July, 1947

Fig. 4. Green weights and arsenic content of Sudan grass grown in black soil with arsenic content
ranging from 15 ppm (Can 1) to 3,014 ppm (Can 24) of As2Oa. The difference in levels of the two
green-weight curves is due to the season in which the plants were grown and is not related to treat¬
ment. (For As203 increments, see Table 10.)

associated with the depression of growth.

Sudan Grass: Black Soil. — The growth of
Sudan grass in black soil, as shown in Table
10 and Figure 4, was much more luxuriant
than in the red soil. Furthermore, in the
black soil normal growth occurred at much
higher levels of soil arsenic, despite high
tissue levels of arsenic. Unlike the curve for
the tomato crops, there was a decided break
in that of the Sudan grass series. Above
Can 16 (614 ppm of soil arsenic) growth
was progressively more difficult, and at the
soil arsenic level of 2,014 ppm (Can 22)
there was no growth.

Although Sudan grass in culture solution
appeared as tolerant to arsenic as was the
tomato, it is quite clear that in soils, Sudan

grass is much less tolerant of arsenic than is
the tomato. This difference in the soil seems
only partly related to the ability of Sudan
grass to extract higher levels of arsenic from
a given soil. (Compare the arsenic levels in
Figures 2 and 4 from Cans 13 to 20, in
which growth of Sudan grass was still appre¬
ciable. ) It is also partly related to a differ¬
ence in the manner in which the plants hold
the arsenic within their tissues. Thus, when
the tomatoes growing in the black soil had
over 20 ppm within their tissues, their
growth was nearly normal. At the same tis¬
sue levels the growth of Sudan grass was
nearly stopped. Thus, tolerance to soil
arsenic involves root tolerance as well as tis¬
sue tolerance. Probably, root tolerance is

Arsenic Toxicity Studies — Clements and Munson

167

determined in part by the nature of the root
structure and in part by the intimacy as well
as nature of the contact between the root
surface and the soil surface bearing the
arsenic. Tissue tolerance, on the other hand,
may be related in part to protoplasmic struc¬
ture and in part to the form in which the
arsenic is held within the protoplasm after
it is absorbed.

Bean: Red Soil. — The bean plant (see
Table 11 and Figure 5), which in culture
solution was the most susceptible of the three
plants to arsenic injury, showed a tolerance
to soil arsenic only slightly below that of the
tomato, but considerably above that of Sudan
grass.

For. the bean, as for the tomato and Sudan
grass, although the level of soil arsenic at
which growth was sharply curtailed was the
same for the first crop as for later crops, the
actual amount of arsenic absorbed was
greater in the first crop than in later crops.

Also, the differences in the amounts of
arsenic absorbed by the various crops are not
all related to the differences in growth made.

GENERAL DISCUSSION

It is apparent that production of crops in
heavy Hawaiian soils which have been con¬
taminated with the herbicide sodium arsen-
ite will be affected variously depending on
the particular crop. Furthermore, crops which
may be looked upon as tolerant to arsenic
when grown in culture solution may become
relatively susceptible to fixed arsenic in the
soil. In culture solutions, the bean plant was
by far the most susceptible of the three plants
used, whether the arsenic was trivalent or
pentavalent. The tomato was the most re¬
sistant toward pentavalent arsenic, but was
about equal to Sudan grass in resistance to
trivalent arsenic. In soil cultures, however,
Sudan grass was considerably less resistant
to sodium arsenite than either of the other

TABLE 11. BEAN-RED SOIL

Yield Data and Arsenic Content of Tissues.

CAN

NUMBER

AS2O3

CONTENT

OF SOIL

SERIES I

AVERAGE OF 2 LATER SERIES

Green weight
of plants

AS2O3 content
of tissues

Green weight
of plants

As2Os content
of tissues

*

ppm

gm. per can

ppm

gm. per can

ppm

1

15

18.7

0.5

24.3

1.5

2

25

18.0

1.1

23.7

1.5

3

34

19.0

1.8

25.9

1.8

4

45

17.3

2.4

22.8

2.2

5

54

20.7

2.9

23.5

5.0

6

65

24.0

7.9

27.5

1.8

7

74

24.7

9.0

28.0

2.0

8

85

26.7

6.9

25.2

3.2

9

95

22.7

4.8

25.9

2.1

10

104

22.3 *

3.4

23.0

1.7

11

115

22.3

3.7

22.8

2.1

12

214

22.0

5.5

26.7

2.5

13

314

14.0

6.3

22.7

3.7

14

414

18.0

10.3

21.2

6.9

15

514

11.0

14.0

14.2

6.6

16

614

7.7

26.4

9.5

6.9

17

714

6.3

11.1

10.8

18

814

9.0

24.6

6.8

12.8

19

915

6.0

. 35.4

6.1

12.9

20

1014

2.5

4.2

12.8

21

1514

2.3

3.2

15.7

22

2014

1.1

2.0

25.1

23

2515

0.7

1.6

32.7

24

3014

0.2

1.5

46.5

168

PACIFIC SCIENCE, Vol. I, July, 1947

/ Z 3 4 £ & 7 6 9 70 77 7Z 73 74- 7& /& /7 76 79 ZO Z7 ZZ Z3> Z4-

30/ L - COL TL/RB CA A/ /VOM 3 BR

Fig. 5. Green weights and arsenic content of bean plants grown in red soil with arsenic content
ranging from 15 ppm (Can 1) to 3,014 ppm (Can 24) of As203. The difference in levels of the two
green-weight curves is due to the season in which the plants were grown and is not related to treat¬
ment. (For As203 increments, see Table 11.)

two. In the red soil, the arsenic concentra¬
tions which were toxic to Sudan grass, bean,
and tomato were approximately 110, 250,
and 550 ppm, respectively. In pounds per
acre foot of dried soil, these figures become
roughly 220, 500, and 1,100 pounds, respec¬
tively.

Perhaps the fact that Sudan grass is a vig¬
orous feeder especially of fixed phosphorus
in such soils is related to its greater sensitiv¬
ity to soil arsenic, which probably is sim¬
ilarly fixed. Such an observation has sup¬
port in the data presented for red soils,
which demonstrate that Sudan grass ex¬
tracted higher levels of arsenic from Cans
10 to 16 than did either of the other two
plants. Both tomatoes and beans when

grown on Hawaiian soils are fertilized
heavily with phosphates in order to obtain
good growth. The large grasses, however,
seem to be able to take their phosphorus even
though it is highly fixed.

It is doubtful whether arsenic which is
applied to soils as trivalent arsenic remains
trivalent after it has been in the soil for
some time. Although no direct pertinent
data are available from this study, indirect
data may be obtained from the arsenic levels
attained by the Sudan grass and tomato
plants growing on the black soil. Tomato
plants in black soil (Can 22) which were
apparently normal contained up to 104 ppm
of arsenic. Sudan grass plants in black soil
in Cans 21 and 22 contained between 30

Arsenic Toxicity Studies — Clements and Munson

169

and 86 ppm of arsenic. Yet when these
were grown in culture solution, only very
much lower levels of trivalent arsenic were
tolerated. The amounts of arsenic absorbed
from the soil were more in line with the
amounts of pentavalent arsenic absorbed
from culture solution. It cannot be pre¬
sumed, however, that after arsenic is ab¬
sorbed in the pentavalent form, it remains
as an inorganic compound after it becomes
a part of the plant’s metabolism.

The claim which has been made by many
that arsenic at proper levels is stimulating to
plant growth is not substantiated by the data
presented here. Neither in water culture nor
in soil culture is there any certain evidence
of such stimulation. What slight gains from
arsenic there may be are infinitesimal com¬
pared with the losses which are certain to
come after the arsenic content passes critical
levels.

The arsenic which was added to the soil
cans was not greatly reduced in amount
either by the growth of the plants or by
drainage which was provided. Even the
large amounts of arsenic contained in the
tomato plants grown in the high arsenic cans
represent very small proportions of the total
amount of arsenic in the soil. To calculate
the time needed to extract the arsenic from
the soil through continued use of tomato
plants, assuming the highest extraction ob¬
served in these tests, would require some¬
thing over 100 crops. It is far better to stop
the use of arsenic before critical levels are
reached. It is apparent from this work, as
well as from that of others, that no matter
how large or how small the annual incre¬
ments to the soil may be, substantially all of
the arsenic remains in the tilled layer. Re¬
ducing the increment of arsenic applied
merely prolongs the time of grace.

One observation needing to be brought
into sharp focus is that, as shown in all fig¬
ures in the text, the point of sterilization for
a given crop is very much higher than the

point at which crop production begins to
suffer curtailment because of accumulated
arsenic.

In red soil, Sudan grass began to suffer
growth curtailment at about 115 ppm
As2Os, the tomato at about 614 ppm, and the
bean at about 314 ppm. From these respec¬
tive points on, the increasing curtailment
varies for each crop. For the tomato the
further drop is precipitous, less so for the
bean, and still less so for Sudan grass.

Sobering is the report from Queensland
by Kerr (1939) that soil arsenic at the level
of 600 ppm resulted in complete growth
failure of sugar cane. In times of low prices
for agricultural produce, even a 5 per cent
curtailment of production due to soil arsenic
may well mean the difference between profit¬
able and unprofitable operation.

Studies carried on elsewhere have yielded
some treatments which may be useful in cor¬
recting arsenic toxicity. The use of heavy
phosphate applications, lime, iron oxide, and
organic matter (perhaps filter cake) have
shown promise of reducing the toxicity of
arsenic excesses. None of these treatments
reduces the arsenic content of the soil.
Furthermore, most of these treatments are
costly. Whether a single treatment is effec¬
tive for any length of time remains to be
determined.

There is only one permanent solution to
the problem of arsenic accumulation so far
as present-day information is concerned, and
that is the cessation of arsenic applications.
Substitution of other herbicides or other
weed-control practices which at the moment
may seem somewhat more costly may be the
cheapest in the long run. Certainly there
can be no reconciliation of a program of
arsenic applications to the soil with any long-
range view of agriculture.

SUMMARY

1. Studies made with plants treated with
sodium arsenate and sodium arsenite in cul-

170

PACIFIC SCIENCE, Vol. I, July, 1947

ture solutions show trivalent arsenic to be ap¬
proximately 1 0 times as toxic to Sudan grass
and tomato plants as the pentavalent form
and approximately four times as toxic to
bean plants as the pentavalent form. The
trivalent form acts more quickly and vio¬
lently on plant tissues.

2. Studies on the relationship of the phos¬
phorus level to the toxicity of pentavalent
arsenic show that an increase in the phos¬
phorus level materially reduces the absorp¬
tion of arsenic by bean, Sudan grass, and
tomato plants. The phosphorus was found
to have little or no effect on the toxicity of
the element after it has been taken into the
plant.

The phosphorus had little, if any, effect
on the absorption of trivalent arsenic from
culture solution by bean, Sudan grass, and
tomato plants.

3. Results are presented for several crops
of Sudan grass, tomato, and bean plants in a
re-cropping experiment with red and black
soils treated with increments of sodium
arsenite. It was found that as time elapsed,
more and more of the arsenic was fixed by
the soil, a fact indicated by a reduction in the
amount of arsenic found in the plant tops.
Growth curtailment, however, was observed
to take place each time at the same levels of
soil arsenic, irrespective of the levels of
arsenic absorbed.

It was found that the plant species varied
in the ability to withdraw arsenic from the
soil medium, tomato and bean being low
and Sudan grass high in ability to withdraw
the element.

4. Sudan grass and tomato plants were
grown in a black alluvial soil treated with
sodium arsenite, and the results were com¬
pared to those in the red soil experiment.
Marked differences were found in the re¬
sponse of the plants to a certain concentra¬
tion of arsenic in the two soils.

5. Whereas in culture solution Sudan
grass was as tolerant to arsenic as the tomato
and much more so than the bean, in soil,
Sudan grass was less tolerant to arsenic than
either.

6. The removal of soil arsenic by crops
which are tolerant to arsenic will at best be
a very slow process.

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Greaves, J. E. The occurrence of arsenic in soils.
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Groves, Kermit, E. C. McCulloch, and J. L.
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selenites to wheat. Amer. Jour. Bot. 24: 720-
728, 1937.

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Cane Growers’ Quart. Bui. 6: 189, 1939.

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Machlis, L. Accumulation of arsenic in the shoots
of Sudan grass and bush bean. Plant Physiol.
16: 521-544, 1941.

Morris, H. E., and D. B. Swingle. Injury to
growing crops caused by application of arsenical
compounds to the soil. Jour. Agr. Res. 34: 59-
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Paden, W. R., and W. B. Albert. Effect of the
addition of arsenical compounds to soils. So.
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1930.

Reed, J. F., and M. B. Sturgis. Toxicity from
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Reichert, F., and R. A. Trelles. Presence of
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Stoklasa, J. Concerning the substitution of ar¬
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1898.)

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25-2 6, 1923.

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on the effects of arsenical compounds upon
apple trees. Phytopathology 1: 79-93, 1911.

- Arsenical injury through the bark of fruit

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Truog, E., and A. M. Meyer. Improvements in
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and arsenic. Indus, and Eng. Chem., Analyt. Ed.
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Vandecaveye, S. C., G. M. Horner, and C. M.
Keaton. Unproductiveness of certain orchard
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lation. Soil Sci. 42: 203-216, 1936.

Williams, K. T., and R. R. Whetstone. Arsenic
distribution in soils and its presence in certain
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Factors in the Behavior of Ground Water in a Ghyben-Herzberg System

Chester K. Wentworth1

INTRODUCTION

The hydrostatic relationship between
fresh ground water and sea water along
coasts and in many islands has been recog¬
nized for about 60 years, since the work of
Badon Ghyben (1889) and of Herzberg
( 1901 ) . It has been studied in various parts
of the world but perhaps nowhere are there
more data concerning it than in Hawaii. In
the course of the Pacific war the occurrence
of ground water on many islands has been of
crucial importance and the concept of a
Ghyben-Herzberg lens has become widely
circulated.

Rain falling on the surface of an ideally
permeable circular island in the ocean is in
part absorbed into the ground. This water
percolates downward and accumulates at the
surface of the salt water at sea level. The
fresh water builds up to a height above sea
level determined by its amount and by the
permeability of the island rock, and also
presses downward until it extends about 40
times as far below sea level as it does above
sea level. The upper surface of such a
ground-water body can be shown to be a
domed one, and the lower surface is deeply
curved because of the ratio of 1 to 40. The
fresh- water body thus approximates the
form of a double convex lens, with the cir¬
cular edge coinciding with the circular coast
(Fig. 1). This is the Ghyben-Herzberg
lens, and the model on which the theory
rests. In many places, owing to differences
in rock structure, only a portion or sector of
the lens will be developed, but the principle
applies equally well.

1 Geologist, Board of Water Supply, Honolulu,
Hawaii. Manuscript received February 6, 1947.

The Ghyben-Herzberg balance is not by
any means universal along ocean coasts and
one may presume that it is well exemplified
along only a small fraction of continental
coasts. Otherwise it would probably be bet¬
ter known. Its somewhat limited occurrence
is due to the requirement of rocks within a
certain range of permeability, sufficient rain¬
fall, and lack of specialized structure in the
rocks. The rock structure must in the main
be fairly homogeneous and be isotropically
permeable if a well-characterized Ghyben-
Herzberg system is to develop.

Fig. 1. Section through an ideal permeable
island in the ocean, showing the form of the fresh¬
water body known as the Ghyben-Herzberg lens.
Owing to scale limitations the bottom of the lens
is shown only about 10 times as thick as the top
part, instead of the 40-fold relation that occurs in
nature.

In Hawaii, the principle of balance has
become well known because of its remark¬
able development in the Honolulu-Pearl
Harbor areas and in a few other localities on
Oahu (Alexander, 1908; Andrews, 1909;
Palmer, 1927: 17-20). It is not present or
at least is scarcely demonstrable along about
half of the windward coast because the rock
is of low permeability and unsuitable struc¬
ture. It is also much less developed on most
of the other islands of Hawaii. Practical im-

[ 172 ]

Factors in a Ghyben-Herzberg System — Wentworth

173

portance of the lens is great because in it a
much greater thickness of fresh or near¬
fresh water is accumulated at sea level than
would otherwise be the case. In fact the
fresh water is perched on salt water. Be¬
cause of the wide extent of such possible
perching, these basal water bodies lying in
Ghyben-Herzberg balance are the largest
ground water resources of the region.

In rocks not favorable for developing
such a lens, there may be springs of fresh
water at sea level. These springs are mostly
small and unimportant and are occasioned
by the cutting of the sea against the land, by
an action such as that which produces valley-
side springs. In Hawaii the basal water
which is in Ghyben-Herzberg balance may,
or may not, be in part artesian. Where a
cap rock exists, the thickness of the lens is
increased by its retardation; but the cap rock
is not essential. The development of hydro¬
static balance occurs even where the lighter
water of the lens is somewhat brackish. Such
a system may be of true Ghyben-Herzberg
form even where the land water carries a
considerable fraction of sea water. But for
practical use only the more perfect systems
where the upper water has less salt than cor¬
responds to about 1 to 2 per cent of sea water
are of interest. Salinity of the Honolulu
city supply corresponds to approximately
1/400 sea water.

Often water supply systems are planned
where there is some indication of fresh land
water at the coast. Here it is supposed that
only drilling or tunneling may be needed to
secure a potable supply. Often such excava¬
tion after prolonged test may show that
despite some degree of Ghyben-Herzberg
functioning the resulting developed water is
too saline for the proposed use. Or, the
water may grow progressively more saline
and show that the hydrostatic system is not
sufficiently stable to stand the disturbance of
even a moderate draft of water. Engineers
and others who have accepted the principle

are naturally disappointed and in turn ques¬
tion it after such adverse tests. But in the
writer’s view it is not surprising that the
fresh-water lens in certain areas is either
lacking or fails to meet the extremely severe
test of yielding potable water continuously.
Rather, it continues as one of the natural
wonders that such systems as that at Hono¬
lulu and a few other places have been de¬
veloped in course of geological time, in such
perfection and with such ability to withstand
modification through artificial development
of the water. These systems are the excep¬
tions rather than the rule.

Observations in various parts of Hawaii
and a growing knowledge of basal water
conditions on other Pacific islands permit
some broad generalizations concerning the
conditions essential to an effective Ghyben-
Herzberg system. An attempt is made in this
paper to outline the requisite conditions.

In the course of compiling this discussion,
the manuscript has been read and criticized
by Charles V. Theis, Arthur M. Piper, L. H.
Herschler, and Gordon A. Macdonald, each
of whom has made valuable suggestions.
The writer is indebted particularly to the lat¬
ter, with whom he has had many most profit¬
able discussions on this and related problems
over a period of years.

OUTLINE OF FACTORS

In outlining the factors affecting salinity,
attention is first drawn to the fact that the
Ghyben-Herzberg lens consists of a lighter
liquid, floating on a heavier liquid and
miscible with it. If the two liquids were not
miscible, they could maintain their separate
character and their common boundary indef¬
initely even without being restrained in a
permeable aquifer. However, as they are
miscible, if the permeable aquifer were not
present the two liquids would become mixed
and diffused in a short time so that the
lighter lens would be destroyed. On the

174

PACIFIC SCIENCE, Vol. I, July, 1947

other hand, if the rock at sea level is not
permeable, there is no opportunity for a
condition of hydraulic balance to become
established. The fresh water cannot adjust
itself to the salt water in relation to sea level
and thus no Ghyben-Herzberg lens will be
formed.

The first requisite is then a suitable degree
of permeability. This must be small enough
to prevent the general mixing which would
destroy such a system and large enough to
permit fresh water under the existing head
differences to move against sea water. Only
then can fresh water progressively displace
salt water in forming a Ghyben-Herzberg
lens. It will be seen presently that the per¬
meability which proves effective is relative to
other factors, and can be better discussed
after these have been listed.

Next to suitable permeability is an infil¬
tration of rainfall of sufficient amount and
continuity to build and maintain a fresh
basal ground-water body about a foot or
more above sea level. According to the rain¬
fall and recharge, there is built a surcharge
above sea level adequate to discharge the
average daily or annual amount to the peri¬
phery of the island. The maintenance of
this surcharge causes the downward accu¬
mulation of fresh water until an approxi¬
mate Ghyben-Herzberg lens is produced.
The miscibility of the fresh and salt water
tends to destroy the lens or the sharpness of
its margin. The rate of addition of fresh
water must be sufficient to overcome this
tendency.

The third requirement is a sufficiently
small fluctuation in both ground-water heads
and in sea- water levels to minimize the ef¬
fects of mixing and the spread of the zone
of mixing through reversal of movement.
Fluctuations in ground-water head are due
to seasonal and other variations in rain¬
fall and recharge. Chief changes in sea level
that we need to consider are those due to
tides. It seems fairly certain that small

islands, which with tidal range of 2 feet
show moderate stability of the Ghyben-
Herzberg lens, if subjected to a 15- or 20-
Toot tidal range would show serious disturb¬
ances of the lens.

It is difficult to give any categorical spe¬
cifications, but the best-known Ghyben-
Herzberg water bodies in Hawaii do not have
seasonal or annual fluctuations of water table
exceeding perhaps one fifth of the total
height of water table above sea level. It
seems likely that any annual change such as
half or two thirds of the mean value would
tend to prevent growth of a lens from which
any potable water could be taken. Both in
their accomplishment and also in their effect
on water quality, the amplitude of such fluc¬
tuations is obviously related not only to the
infiltration changes but also to the per¬
meability.

A fourth factor, and in some ways the
most important of all, is that of comparative
uniformity and regularity of permeability,
and freedom from large and long openings
crossing the boundary between fresh and
salt water. Much of the effectiveness of the
Ghyben-Herzberg mechanism depends on
the maintaining of a fairly smooth and
orderly boundary between the two liquids, as
closely analogous as possible to the mathe¬
matically definable boundary between im¬
miscible liquids. It is fairly evident that the
actual condition is somewhat remote from
this and, with fluctuating directions of move¬
ment, that existing large fissures or tubes
must carry long filaments of one sort of
water into the realm of the other, and vice
versa. Such conditions seem to explain the
observed vagaries in the composition of
water from different wells in the same dis¬
trict and even at the same depth. Despite
such irregular and fluctuating interpenetra¬
tions, a broad regularity of boundary is
shown in the more stable Ghyben-Herzberg
systems such as those of Oahu. Certainly if
changing heads tend to move the salt-fresh

Factors in a Ghyben-Herzberg System — Wentworth

175

boundary up and down alternately, any large
openings which cross the boundary will be
much more destructive in promoting inter¬
penetration of one kind of water by the
other than small openings. It appears clear
that heterogeneous permeability will be more
likely to produce an irregular and disorderly
boundary than a homogeneous or regular
permeability.

A fifth factor of importance is the effec¬
tiveness of a cap rock along the coast. Such
a barrier not only promotes the building of
higher heads of fresh water but indeed
creates a condition somewhat akin to a
U-tube so that at their two upper surfaces
the fresh- and salt-water bodies are effec¬
tively separated. The first and most ad¬
vanced intermixing of salt and fresh water
would normally take place at the coastal
margin. Here changing head differences
would be exerted across the shortest dis¬
tances between fresh ground water and free
sea water. Evidently a barrier along the
coast would have marked protective value.
In the Honolulu area the thickness and
width of the cap rock are such as to interpose
a distance of several thousands of feet in
most places between the water table and free
sea water, and this barrier is of tremendous
importance.

The factors mentioned above are, in sum¬
mary: (1) suitable permeability, (2) ade¬
quate infiltration, (3) limited fluctuation,
(4) regularity of permeability, (5) an ef¬
fective cap rock. Some aspects of their inter¬
relationships will now be discussed.

PERMEABILITY

It would be difficult to over-emphasize
the importance of time in the inter-relation¬
ship of the several factors. The first factor,
permeability, is of course a rate of discharge
through a specified cross-section, and infil¬
tration is expressed as an amount per unit
of time and per unit of area. It is the lag in

the dissipation of infiltrated water through
permeable rocks which causes the initial ac¬
cumulation of ground water and determines
the ultimate head at which balance between
gain and loss will be reached. In general,
on islands of similar geometrical form, infil¬
tration proceeds through areas that are pro¬
portional to the squares of linear dimensions,
whereas for the same heads, the areas
through which discharge to the sea takes
place are proportional to perimeters, hence to
the first powers of linear dimensions. Hence
derives the tendency to build higher water
tables on larger islands, thus restoring some
degree of equality with this second dimen¬
sion. It is also true that larger islands have
longer radii and greater likelihood of con¬
tinuous discharge even with discontinuous
rainfall and infiltration.

Permeability that is too great (relative to
infiltration and other factors) will result in
a water table so low that no permanent pres¬
sure against salt water will be maintained
and no permanent Ghyben-Herzberg lens
will exist. On the other hand, if permeabil¬
ity is too low the amount of water infil¬
trated will be small, and the exerting of a
systematic pressure against the sea water is
less likely to be established. With the higher
water tables due to less permeable rocks
there is less likelihood that rocks of reason¬
ably uniform permeability will extend to the
depth below sea level requisite for a func¬
tional system. Moreover, even if balance
exists, the lesser permeability precludes the
detectable response by which we might recog¬
nize it. Thus on various less observable
grounds the Ghyben-Herzberg condition
vanishes also with reduced permeability.

It is possible, too, that in some islands the
rock near sea level and slightly below at the
coast is less permeable than the general mass
below sea level and farther inland. Such
difference within a favorable range of mag¬
nitudes would promote the Ghyben-Herzberg
condition.

176

INFILTRATION

It is hardly practicable to suggest a numer¬
ical definition of adequate infiltration, but
some limiting data may be offered. On some
of the larger islands of Hawaii, where the
discharge through the shore perimeter in
some sections reaches values of 5, 10, or even
20 million gallons daily per shore-line mile,
with other conditions favorable, Ghyben-
Herzberg conditions are conspicuous and
stable. These conditions imply some intake
area with annual rainfall of 100 inches or
more. In many other areas of Hawaii, on
leeward coasts or on smaller islands, where
rainfall is mostly under 50 inches and
ground-water discharge is 1, y2, or 1/10
M.G.D. per shore-line mile, the Ghyben-
Herzberg condition is either missing or non-
demonstrable on the scale of practical water-
supply operations. However, it should be
remembered that under wartime or expedi¬
tionary conditions a vestigial fresh- water
lens may be of great temporary value even
though it may fail on continuous mechan¬
ized development.

The size of the island is important here;
on an island 2 miles in diameter the dis¬
charged ground-water fraction required to
equal 1.0 M.G.D. per shore-line mile is 42
inches over the area, whereas on one 10
miles in diameter 42 inches over the area
will give 5 M.G.D. per shore-line mile.
From experience in Hawaii the latter would
quite likely have the Ghyben-Herzberg con¬
dition; the former very likely would not,
unless the permeability or cap rock were
especially favorable.

It may now be practicable to say that the
required permeability assumed to be fairly
homogeneous throughout the whole thick¬
ness is such as will require the water table
to build up 2 to 10 feet or more and remain
at nearly constant levels perennially. With
lesser heads some degree of concentration of
fresh water may be found but it is less likely

PACIFIC SCIENCE, Vol. I, July, 1947

to be stable against mechanized exploita¬
tion.

FLUCTUATION

The permissible annual fluctuation so far
as we can estimate at present is somewhat
less than one half of the mean water-table
head, and in most cases less than one fourth
of that head. No data are at hand, for any
system of the magnitude of that of Hono¬
lulu, in which greater fractions of fluctua¬
tion are known. For systems with head of
5 feet or less it seems indicated that fluctua¬
tion from 2.5 to 7.5 feet of head would be
fatal to useful Ghyben-Herzberg stability.
No data are available to determine what
ratio of short-term fluctuations might be
tolerable, but naturally the tolerable limiting
amplitude would be lower than for the
longer term ones, and in the nature of the
case they are materially less.

REGULARITY OF PERMEABILITY

By regularity of permeability we mean
homogeneity of distribution and sizes of
interconnected openings. A formation con¬
sisting of well-sorted sand or gravel would
throughout its mass have regular permeabil¬
ity. A formation consisting of moderately
permeable material but broken by large,
irregularly spaced fissures or caverns would
have irregular permeability. The adverse
effect of irregular permeability would lie in
introducing large and changeable irregular¬
ity of pattern in the three-dimensional net¬
work of surfaces of equal pressure and hence
of lines of flow, of velocities, and of salini¬
ties. It is evident that in formations perme¬
able enough to meet the Ghyben-Herzberg
requirement, large irregularities will pro¬
mote intermixing and tend to effacement of
the zone of balance, which without frictional
retardation can only be stable between im¬
miscible liquids. In a sense, large openings
of such length and direction as to lead across

Factors in a Ghyben-Herzberg System — Wentworth

177

the zone of mixing are to be regarded as
short circuits which would produce poten¬
tial disturbance analogous to that in an elec¬
trical network. Moreover, the movement of
saline water toward fresh, or vice versa,
taking place during any phase would leave
residues of great importance. It therefore
appears that, increasingly, irregularity of per¬
meability would tend strongly toward efface-
ment of the salt-water-fresh-water contact
on which the Ghyben-Herzberg lens de¬
pends, just as would increase of general
permeability.

It appears that this variation in permeabil¬
ity, with some large openings going outside
the favorable range of permeability, may be
a very large factor in explaining the great
differences in the Ghyben-Herzberg condi¬
tions on different coral limestone islands, or
on different parts of the same island. Not
only initial differences due to structure of
the calcareous accumulation, but also fissures
developed near sea level by the action of
fresh water probably are important here.
Such an interpretation has been suggested
by the writer’s observations in the Marianas
during early stages of military operations
there and has also been emphasized by
others.2

CAP ROCK

It is not difficult to show why an effective
cap rock is so very significant in permitting
the growth of some of the larger Ghyben-
Herzberg lenses. It is an essential part of
that theory that in a steady condition of
dynamic balance, with the thickness of the
lens neither increasing nor decreasing, lines
may be drawn from various points of the
top of the lens (the water table) passing
downward and outward in a wide curve to
emerge in the ocean surface, along which
hydrostatic pressures are in balance. If the

2 Piper, Arthur M. Letter dated December 5,
1946.

position of the water table is changed by
fluctuations of recharge or of loss, move¬
ment will tend to take place along these lines
in accordance with the size of openings and
length of path. It is evident that those paths

Fig. 2. Schematic sections through the margin
of a Ghyben-Herzberg lens, under the three con¬
ditions indicated in the text. For convenience in
drawing, the mean line of balance is based on a
ratio of 10 : 1, rather than the true ratio of ap¬
proximately 40 : 1.

178

PACIFIC SCIENCE, Vol. I, July, 1947

nearer to the shore line are much the shorter
and that under fluctuating conditions of un¬
balance, water movement in larger openings
across the zone of contact would take place
here more rapidly than elsewhere (see Fig.
2 ) . Along a shore unprotected by a cap rock,
the thin edge of the Ghyben-Herzberg lens
would be especially vulnerable to disturb¬
ance or destruction during marked fluctua¬
tions. There, particularly, the adverse effect
of large fissures or other irregularity of per¬
meability is certain to be great.

From these considerations, the value of a
cap rock as a barrier between fresh and salt
water is readily seen. As stated elsewhere,
the interposition of the cap rock between
fresh and salt water in effect completes the
physical pattern of a U-tube. It tends to
raise the head of basal water and to truncate
and thicken its shore margin. This barrier
has the result of eliminating the thin edge
of the lens, with its dangerous sensitivity to
plus and minus fluctuations. It should not
be overlooked, also, that in most places the
Ghyben-Herzberg condition is first recog¬
nized and is most useful in the shore zone
where the water is most accessible and often
most needed. It is possible that more
complete and more extensive exploration
will demonstrate the interior existence of
Ghyben-Herzberg lenses in some islands
where the condition is not well shown at the
shore; such discoveries would confirm the
contentions of this paper.

PARTS OF THE LENS

In the preceding sections, the five factors
controlling the establishment and growth of
the Ghyben-Herzberg lens have been dis¬
cussed. Attention is now directed to the
parts of the lens and to the nature of its
lower boundary. It has been accepted that
the lower surface of the lens is a zone of
transition from fresh to salt water, and that
since the two liquids are miscible the zone

will have thickness. The perfect condition
of a sharp boundary can only obtain with
immiscible liquids. If the permeability is
too great for the amount of infiltration, or
if there is great irregularity of permeability,
the mixing and mutual interpenetration of
the two waters will be promoted and the
zone of transition will be thickened. Such
mixing may go so far that no part of the
lens is free from salt contamination.

GROWTH OF DIFFUSION ZONE

Another effect, that of fluctuation or alter¬
nate movements of the zone of transition,
may not be so readily discerned. With im¬
miscible liquids and no matrix such as rock,
the contact surface would move up or down
according to relative pressures and with little
or no deformation. However, in rock with
miscible liquids, the migration of the zone
downward into rock formerly filled with salt
water would first involve driving out some
of the salt water. But it would also result in
assimilation of some of the salt water re¬
maining longer in smaller openings. If
such a process continued, the water composi¬
tion at any one place would tend to approxi¬
mate more and more that of the advancing
water and retain less and less of the quality
of the water originally displaced. However,
there would commonly, after any short time,
be some residue of the displaced water. If
we assume that at any given time the local
composition is a function of the composi¬
tions of the two waters and of the time dur¬
ing which Water A has moved into the realm
of Water B, a process analogous to the
exponential law of rinsing, some interesting
consequences appear. If, for example, the
two waters have an initially sharp boundary,
and that boundary is moved under hydro¬
static changes of sign from one realm to the
other, in equal amounts successively, the
most immediate effect is the spreading of
the boundary so that it is no longer sharp but
assumes a graded transition form.

Factors in a Ghyben-Herzberg System — Wentworth

179

Fig. 3. A numerical model showing growth of the transition zone under the assumption of progressive
rinsing, analogous to the exponential theory of rinsing. The unmodified fresh and salt waters are
shown by the respective horizontal and vertical hatching. The transition zone is shown by the growing
band of figures, the successive values being parts per thousand of salt water.

Figure 3 is a numerical model showing
the effect of moving the junction between
two types of water to and fro in a permeable
medium having some storage capacity. At
the beginning of the test period, the junction
is assumed to be sharp, as shown by the re¬
spective patterns (Fig. 3). Each successive
column of figures represents the composition
in successive, equal periods of time. The
fresh water is assumed to move by 10 suc¬
cessive units of motion against the salt
water, thence to retreat by 20 units to a posi¬
tion 10 units on the other, or fresh, side of
the initial line of balance, and finally to
return by 10 units to that line and thus com¬
plete one full cycle. With each unit of
movement, the water in a given position is
assumed to be made up of 9 parts of oncom¬
ing water and 1 part of residual water. It
is immaterial for the discussion whether the
residue be assumed as 10 per cent or some
other figure.

The figures represent parts per thousand
of salt water. Above the transition zone all
the water is fresh, taken as zero parts. Below
it, the water is of sea-water composition,
taken as 1,000 parts. A certain raggedness
appears at the margins, owing to limiting
the calculations to the nearest thousandth.
So far as practicable the accumulation of
values of the next digit has been anticipated
in computing the marginal figures.

It is evident from Figure 3 that changes
take place both at the advancing margin and
at the following margin. The composition
at the advancing margin is changed toward
that of the water being invaded, and that
change migrates into the advancing front.
The same direction of change is reflected
through the zone, and the compositions in
the following margin also change toward
that of the water being invaded. The rates
of these changes are determined by the exist-
ing gradient of composition at various points
in the transition zone.

After the first reversal, the form of the
composition diagram becomes nearly sym¬
metrical (Fig. 4). With continued fluctua¬
tion the transition zone becomes progres¬
sively wider and the rate of change of com¬
position within it is slower. The fresh water
is more deeply penetrated by a graded fringe
of saline composition and the salt water
more deeply penetrated by a graded fringe
of freshened water. This effect is indicated
in the progressive flattening of the transition
curves of Figure 4, as well as by the march
of the figures in Figure 3.

It is possible that with a symmetrical
series of fluctuations a limit of width would
in time be reached , in a regularly permeable
aquifer. However, in any natural aquifer
and particularly with unsymmetrical fluctua¬
tions and with movement under new head

180 PACIFIC SCIENCE, Vol. I, July, 1947

STAGES

ALTERNATING MOVEMENTS OF MEAN LINE

0 100

COMPOSITION DIAGRAMS

TIME -

Fig. 4. Schematic diagram showing widening of the transition zone with successive cycles of alter¬
nate invasion of one type of water by the other. The lower part shows five composition diagrams,
commencing with the sharp boundary at the left. The general sigmoid form of the composition line
may be verified by plotting the figures given in the right-hand column of Figure 3.

differences through larger openings in dif¬
ferent zones, we cannot doubt that there
will be slow and persistent growth in the
width of the zone because of this rinsing
effect. Thus we suppose that, setting any
given limits at the two margins, the thick¬
ness of the transition zone will become
progressively greater as alternations are re¬
peated, or as the movement assumes greater
amplitude.

The composition diagram developed on
this assumption of rinsing is a symmetrical
curve starting as an asymptote to the zero
line of salinity, passing through an inflexion
at the 50 per cent point, and terminating as
an asymptote to the line of 100 per cent
salinity of sea water. After any series of
complete cycles the zone has widened, but
the line of equivalent density or balance is

found in the position formerly occupied by
the initial sharp boundary.

REVERSIBILITY

We seem now able to resolve the often
discussed problem of reversibility of the
process of saline encroachment. This has
vexed many people in Hawaii and seems to
be clarified by the above discussion. From
that line of reasoning, we may say at once
that if we consider the position of the 50
per cent line, or the equivalent position of
total salt against total fresh water, the pro¬
cess of saline encroachment does appear to
be completely reversible. After a period of
reversed movement equal in duration and
intensity to the earlier saline encroachment,
assuming each movement completed, we

Factors in a Ghyben-Herzberg System — Wentworth

181

would expect to find the middle of the zone
of transition or diffusion at the same level it
occupied initially. However, we are only
theoretically interested in the middle of this
zone; for practical purposes our interest in
an operating Ghyben-Herzberg system is
concentrated on that fringe nearest the fresh
water, where the salinity is equivalent to the
order of 1 per cent of sea water or less. It
appears that any movement of the zone of
transition causes it to widen. Thus the cen¬
ter of the zone may return to a former posi¬
tion after equal and alternate movements,
but the near edge of the zone, judged by any
defined standard, because of the previously
stated principle of widening will not retract
as readily as it advances.

Thus it appears that in the practical sense,
in relation to exploitation of potable or agri¬
culturally useful water, the encroachment of
saline water, under the natural plus artificial
and somewhat aggravated fluctuations, will
take place more readily than the reverse
process of elimination under a conservation
program. Thus there is an element of irre¬
versibility, despite the difficulty which vari¬
ous workers, including the present writer,
have had in seeing why the salt water could
not be driven back by an equal period of
reversed movement. Some have postulated
trapping of salt water in pockets. It must,
however, be pointed out that in a hydraulic
system where water may move either way,
trapping is not restricted to pocketing
against the direction of gravity but could
equally well work the other way with reverse
direction of water movement. It is con¬
cluded here that no special theory of trap¬
ping, especially trapping in one direction, is
needed, nor is any theory of directionality
required. The condition seems fully ex¬
plained by the concept of symmetrical thick¬
ening, due to rinsing, plus the fixing of
practical interest' in a position on the near,
or upper, side of the zone of transition.

DIFFERENCES IN SYSTEMS

We need now to offer acceptable explana¬
tions of the differing qualities of water in
various systems. It has been found in vari¬
ous places in Hawaii that the main part of
some of the larger Ghyben-Herzberg sys¬
tems may for many years furnish water of
surprising constancy of salinity. There are
three most evident sources of sodium chlor¬
ide: (1) from normal rock weathering, (2)
from salts left on the land from salt spray
or from more saline irrigation waters, (3)
from admixture in the aquifer with intrud¬
ing sea water.

We are well acquainted with marked in¬
creases due to the third factor; there is an
equal amount of evidence on aquifers which
over a period of many years, and even with
considerable reduction of head and increase
of draft, continue not to be affected by (3)
but represent a stable and not wholly defined
combination of ( 1 ) and ( 2 ) . This is true
of some aquifers where the draft of water
is from points several hundred feet below
sea level. Obviously such points are in a part
of the Ghyben-Herzberg lens that is not yet
affected by saline encroachment (from 3)
and cannot be a part of the transition zone.
That in course of time, through thickening
of the zone, they might become so is, unfor¬
tunately, one of the practical lessons we
learn.

On the other hand, we find aquifers in
which the Ghyben-Herzberg lens at any
level, from the top downward, yields water
of high" salinity, often increasing markedly
with draft, and appears to be deriving it
from normal and induced admixture of sea
water. In such places we can only conclude
that the whole lens is a part of the zone of
transition. It appears therefore that while
some lenses have a considerable fraction of
water not affected by the adjacent salt water,
others do not. For convenience we may call
the upper part the fresh- water core.

182

PACIFIC SCIENCE, Vol. I, July, 1947

MAINTAINING THE CORE

The fresh-water core is a unit through
which passes annually the amount of water
added to the water table. Some of this water
passes down the slope of the water table to
escape near the coast, but there is no ques¬
tion that there is considerable deeper circu¬
lation. However, except for the migration
of the zone of transition with fluctuations of
rainfall and draft, and the effect of a few
large openings with unbalanced pressures,
the water does not move through the transi¬
tion zone. More properly we can assume
that the water in the fresh core circulates
past the upper fringe of the transition zone.
Presumably this circulation in some lenses
is sufficiently active to offer considerable
resistance to the thickening of the transition
zone. This would operate by rinsing away
any slight increases in salinity that might
persist if the invading fringe of saline com¬
position were penetrating a truly static body.

It appears that in some of the thicker and
more functional Ghyben-Herzberg lenses,
the transition zone has not become thick
enough to reach the top of the lens. The cir¬
culation in this upper fresh-water core is
active enough to provide an adequate rins¬
ing action against the upper fringe of the
transition zone. Hence the integrity and
water quality of the fresh-water core are well
maintained. In other lenses, usually much
thinner and perhaps developed under less
favorable conditions, the transition zone has
either thickened to encompass the whole
thickness of the lens or perhaps has always
had some such thickness. Here there is no
fresh-water core, at least not at the coast or
where exploration has penetrated.

Experience in finding small amounts of
water of low salinity at the very top of such
water bodies by no means invalidates the
distinctions set forth above. Undoubtedly
in wet weather a certain amount of rain
water would move down the slope of the

water table with only slight mixing with the
prevailing ground water; such would be the
source of a fresh-water layer, but the layer
would be insufficient in amount. Not un¬
commonly water bodies which will yield
fresh water in small samples are found on
drilling and pumping to yield only water of
considerably higher salinity. For this reason
preliminary or bailed samples sometimes
lead to hopes that are later not realized.

It is not intended here to treat the various
complexities of water development from the
Ghyben-Herzberg lens, but the few ele¬
ments which enter into the problem may be
mentioned. We start by emphasizing that
the lens is a storage body or gland through
which water is moving, and that under bal¬
anced conditions the inflow and outflow
are equal in amount.

Because the upper surface slopes toward
the ocean and the rocks are permeable, there
is a steady loss proportional to the head
differences through various openings, whose
locations are usually not known in detail. So
long as this head is maintained these open¬
ings, if not plugged, will discharge the same
amount of water. If water is to be taken by
man, the head must be lowered until the loss
from various openings has been reduced by
the amount taken artificially. The lowering
of head makes water temporarily available
from storage at the upper surface; if we fol¬
low the Ghyben-Herzberg principle we can¬
not doubt that in due course the bottom of
the lens must shrink to reach a new equi¬
librium. This will yield, over a long period,
very large amounts of water. This condition
appears to explain the remarkable stability
against draft which is shown by some large
systems (Wentworth: 1942).

EFFECT OF DRAFT ON QUALITY
We have seen that the water of the lens
is in motion, toward points of outflow,
which in hydraulic terms are called sinks.
When a well or shaft is dug and water is \

Factors in a Ghyben-Herzberg System — Wentworth

183

taken therefrom, this becomes a new point
of outflow, or sink. The quality of the
water at any given point is determined by
the rates and amounts of flow induced under
hydraulic conditions from the several acces¬
sible sources. Availability of water from
those sources depends on the sizes and num¬
ber of openings connecting them with the
sampling point and on the proportionate
hydraulic flow induced by draft at the
sampling point. When the draft from a well
is applied to the pre-existing flow pattern,
that pattern is changed and the well com¬
petes for water against the other flow lines.
The quality of water drawn depends on the
flow pattern set up under the new conditions
and on the compositions of the several
waters available.

The larger the intake surface (as in a long
tunnel rather than a small well ) , the smaller
the drawdown required and the smaller the
disturbance of pre-existing flow lines re¬
quired to get the water. In such case the
salinity of the water drawn will probably be
less modified from that sampled from the
aquifer under the original flow conditions.
Particularly, the smaller the drawdown, the
smaller the likelihood of inducing flow from
lower parts of the lens, which may be more
saline.

In general, near the coast, the salinity of
water taken from a well increases with in¬
crease of depth and with increase of draft.
It also is greater at low regional heads than
at high, and, as stated above, is greater at
high drawdown than at low. Two wells of
similar depth and location often show quite
marked differences, owing to different,
though often unknown, openings in the for¬
mations they penetrate. Occasionally wells or
shafts are dug which encounter openings
that are thought to run inland and in which
the water becomes less saline as draft is in¬
creased; the reverse is far the more common
situation. These exceptions do not invali¬
date any of the recognized principles; they

merely emphasize the complexity of the
hydrologic conditions involved, and our
inability in most cases to make specific pre¬
dictions.

A Ghyben-Herzberg lens is of great eco¬
nomic value when it has a fresh-water core
of such size and stability as to permit the
desired draft of fresh water and yet continue
to maintain the freshness of the core. Where
these conditions of stability exist and can be
maintained, the amounts of water which can
be taken out and the capacity of the system
to sustain short period overdraft are truly
astonishing. A great deal of exploration and
usually much full-scale operation will be
required in most such systems before the safe
capacity or other conditions of operation can
be determined. The chief requisite in any
given case is a body of data that covers a
sufficient range of facts and of time, to¬
gether with recognition of the principles in¬
volved. The present paper is offered as an
elementary formulation of those principles
as they appear at present.

SUMMARY

The Ghyben-Herzberg lens is essentially a
gland, into which water moves from rainfall
and out of which it moves through natural
leaks and artificial discharge. Because of
contact with salt water, with which it can
mix, the lens of fresh water is in an inter:
mediate condition of equilibrium. To sur¬
vive, it must not be stagnant; water must
move through it. It can be destroyed by too
little source water or by too rapid escape of
water. In rocks that are too impermeable
the dynamic equilibrium may not be set up.
The formation and survival of such a lens in
a given place depends on the values and
mutual relations of the factors of permeabil¬
ity, rainfall, fluctuations in level, regularity
of permeability, and presence or absence of
a cap rock. Change of one of these condi¬
tions, such as fluctuation, may change the
balance on which such a lens depends.

184

PACIFIC SCIENCE, Vol. I, July, 1947

REFERENCES

Alexander, W. D. Letter to the editor. Pacific
Commercial Advertiser (Honolulu), Oct. 9,
1908.

Andrews, Carl B. Structure of the southeastern
portion of the island of Oahu, Hawaiian Islands.

19 p., 18 fig. [Thesis, Rose Polytechnic Institute,
1909.]

Badon Ghyben, W. Nota in verband met de
voorgenomen put boring nabij Amsterdam. K.
Inst. Ingen. Tijdschr., 1888-89 : 8-22, 2 pi. The
Hague, 1889.

Brown, J. S. A study of coastal ground water,
with special reference to Connecticut. 101 p.,

20 pi., 7 fig. U. S. Geol. Survey Water-Supply
Paper 537, Washington, D. C., 1925.

Herzberg, Baurat. Die wasserversorgung einiger
Nordseebader. four. Gasheleuchtung und Was¬
serversorgung., Jahrg. 44. Munich, 1901.

Palmer, H. S. The geology of the Honolulu arte¬
sian system. Rpt., Honolulu Sewer and Water
Commission, supplement. 68 p., 21 fig. Hono¬
lulu, 1927.

Stearns, H. T., and K. N. Vaksvik. Geology and
ground-water resources of the island of Oahu,
Hawaii. 479 p., 33 pi., 34 fig. Ter. of Hawaii
Div. Hydrog., Bui. 1. Honolulu, 1935.

Wentworth, Chester K. Storage consequences
of the Ghyben-Herzberg theory. Amer. Geophys.
Union Trans. 23: 683-693, 1942.

- The specific gravity of sea water and the

Ghyben-Herzberg ratio at Honolulu. 24 p., 5
fig. Hawaii Univ. Occas. Paper 39. Honolulu,
1939.

Fig. 1. Chart published by the U. S. Coast and Geodetic Survey showing seismic sea wave travel
times to Honolulu is here reproduced on a small scale with the omission of some detail. (To accom¬
pany "Travel Times of Seismic Sea Waves to Honolulu,” by Bernard D. Zetler, Pacific Science,
vol. 1, July, 1947.)

:• ; , v •

; ■ . ' a , :

- •

Travel Times of Seismic Sea Waves to Honolulu

Bernard D. Zetler1

The seismic sea wave which struck the
Hawaiian Islands on April 1, 1946, has
again focused attention on the necessity for
adequate protective measures against similar
disasters in the future. The problem is obvi¬
ously complex, involving rapid location of
the epicenter, the detection of the sea wave
as it moves toward the Hawaiian Islands, a
quick method of determining the time the
wave will reach the islands, and finally an
adequate means of providing security for
people and property. The purpose of this
study was the preparation of a chart of the
Pacific Ocean which would show the travel
time to Honolulu of a seismic sea wave from
the plotted position of an earthquake epi¬
center (see Fig. 1, insert sheet). Given the
time of the disturbance, the arrival time of
the wave at Honolulu becomes immediately
available.

Oceanographers have long accepted the
concept that the velocity of a seismic sea
wave is a function of the depth of water and
they have expressed it mathematically as
v = \/gd, where v is the velocity of the
wave, g the acceleration of gravity, and d
the depth of the water. However, this
formula for velocity has been considered by
some authorities to be a rough approxima¬
tion; it was believed that the actual velocity
would always be somewhat slower.

The results of the computations made in
the course of the study by Green (1946)
created more confidence in the accuracy of
travel times computed by means of this
formula. These computations were not in¬
fluenced by the recorded arrival times; the
times to several of the more distant places

1 Mathematician, U. S. Coast and Geodetic Sur¬
vey, Washington, D. C. Manuscript received
March 17, 1947.

[ 1

were computed before it was known that the
sea wave had been recorded on the gages.
Table 1 of that report lists 12 places whose
distances from the epicenter vary from 1,610
to 8,066 statute miles. A comparison of
observed with computed travel times to these
places shows an average variation of 1.2 per
cent, which is not consistently in one direc¬
tion.

It was decided that the procedures used
in the above project could be adopted in the
preparation of a chart which would show
the travel time of any seismic sea wave to
Honolulu. A series of ^-hour curves would
be drawn on the chart such that each would
represent the length of time a sea wave
would take to reach Honolulu from an epi¬
center at any point on the curve.

Points to be used as epicenters of sea
waves were selected in various directions
from Honolulu, and travel times were com¬
puted, using soundings from large-scale
charts, along arcs of great circles between
these points and Honolulu. Half-hour inter¬
vals were plotted along each arc with numer¬
ical time values increasing with distance
from Honolulu. The time curves were drawn
by connecting the respective l^-hour points.

Although observed travel times were
available for a number of sea waves which
had previously been recorded on the Hono¬
lulu tide gage, it was considered desirable to
use computed rather than observed data in
the construction of the chart. However, the
positions of the epicenters of recorded waves
were included among the points from which
travel times were computed in order to make
available a comparison of observed and com¬
puted travel times (see Table 1).

In preparing the time data along any par¬
ticular path, a great circle course between
5 ]

186

PACIFIC SCIENCE, Vol. I, July, 1947

the epicenter and the entrance to Honolulu
Harbor was plotted on a chart of the Pacific
Ocean. The path was transferred to larger
scale nautical charts and then divided into
sections of 120 nautical miles each, except
when rapid changes in depth required sec¬
tions of shorter length. The average depth
in each section was taken and the time re¬
quired to pass over it was computed as
follows:

Let d represent the mean depth, in fath¬
oms, of the 120-mile section and t the travel
time of the sea wave over that section.

Since v = V gd = 8.23 V d nautical miles
per hour* /= 120/8.23 V^=l4.58/V<7
hours.

The times thus computed were added
cumulatively, increasing with distance from
Honolulu, and the y>-hour points were de¬
termined by interpolation.

The dividing of a path into small sections
increases the precision of the determinations
over those which are obtained by using a
mean depth over a whole distance. The 120-
mile points were usually 0.25 to 0.35 hour
apart, thus allowing for reliable interpola¬
tion of 1^-hour points which would have
been impossible were the path to be consid¬
ered as a whole.

When a great circle course for a seismic
sea wave is first laid off on large-scale
charts, there are several details to be con¬
sidered. If the path crosses a large unit of
land, the portion of the wave front which
reaches Honolulu will have to go around
that land. Because the part of the wave
front in deeper water will advance more
rapidly, the path is considered the combina¬
tion of arcs of two great circles joined in
the deep water off the coast. If the path of
a wave involves crossing a large section of
shoal water, then the time of the wave front
which diverges somewhat from the great
circle course, but which travels over a deeper
course, must be considered. An excellent
example of the latter was found in the com¬

putation of the travel time of the wave from
the Aleutian Trench on April 1, 1946, to
Sitka, Alaska. The travel time along a great
circle course, through shoal water most of
the way, was computed as about 7 hours.
By considering a path going to the south¬
east for about 90 miles and then moving
along a great circle course from that point to
Sitka, the time was calculated to be about
3 hours, which was in almost perfect agree¬
ment with the observed time.

The latter example is an extreme case.
More frequently there was found a condi¬
tion in which part of a great circle course
covered an area less deep than that covered
by an adjacent path. Despite the additional
distance covered in diverging from a great
circle course to get into deeper water, the
total travel time may be less than that along
the original great circle path. Shallow areas
of this type caused irregularities in the time
curves and necessitated the computation of
additional paths and the consideration of the
bathymetric pattern.

The problem of how to treat the compu¬
tation of the travel time of a wave whose
path lies in a deep channel with compara¬
tively shallow water on both sides is a
troublesome one. If the formula were fol¬
lowed rigorously in such a case, the front of
the wave would gradually have to become
more and more pointed as the wave moved
forward, the shallow water on either side
slowing down that portion of the wave front
passing over it. The concept of such a
pointed wave front did not seem reasonable,
and the velocity was computed as just slightly
faster than that over the shallow area. The
observed travel times from several epicenters
lying in deep channels were found to com¬
pare favorably with the times thus computed.

Some of the epicenters used in comparing
observed with computed travel times were
found to plot on land near ocean deeps.
There have been some differences of opinion
among seismologists whether the true epi-

Travel Times of Seismic Sea Waves — Zetler

central positions are on or off shore. The
method used in this study necessitated a
given depth of water to make a computation
possible, and therefore the times were com¬
puted to the ocean deeps which are near the
plotted epicenters. The comparison of ob¬
served and computed travel times would
seem to indicate that this procedure is reason¬
able.

Three seismic sea waves which originated
near the Japanese island of Honshu have
been recorded on the tide record at Hono¬
lulu. The observed travel times exceed the
computed times by 14, 23, and 49 minutes.
The first two differences are sizable but not
necessarily serious. There are several fac¬
tors which can contribute to a small con¬
sistent variation. Accurate computations of
travel time require adequate and reliable
soundings. Some areas in the Pacific are in¬
adequately surveyed, and the accuracy of
some soundings in other areas is question¬
able. For the 120-mile travel path sections
which contain large variations in depth, the
procedure was to use a mean depth after
rejecting occasional extremely shallow
depths. This technique and the resulting
mean depths are somewhat subjective and
could lead to small inaccuracies in travel
time.

The seismic disturbance off the south coast
of Honshu on December 7, 1944, created a
sea wave which arrived at Honolulu 9 hours
and 9 minutes later, whereas the computed
travel time for the path is 8 hours and 20
minutes. The epicenter of the December 20,
1946, sea wave is approximately 170 nautical
miles west of the 1944 epicenter, the water
between the two is relatively shallow, and
great circle paths from the two to Honolulu
would virtually coincide. The difference of
only 3 minutes between the observed travel
times cannot be explained by some of the
difficulties mentioned above. It seems prob¬
able that some of the 49-minute difference
must be attributed to tide gage operation,

187

tide record interpretation, the seismological
determination of epicenter, or a combina¬
tion of these factors.

A serious misinterpretation of a tide mari-
gram can take place if the waves are small in
amplitude. The first wave is usually smaller
than those immediately following it and may
not be recognizable because of the seiche.
Therefore a time of arrival which appears
to be definite may refer to the second wave
rather than the first. With an observed
travel time greater than its true value, the
time difference, computed minus observed,
is large negatively.

Besides giving the travel time of a seismic
sea wave from an epicenter, the chart may
also be used in conjunction with apparatus
which detects the sea wave as it moves
toward the Hawaiian Islands. Midway
Island, for example, lies approximately on
a great circle arc between Honshu and Hono¬
lulu. If a wave were received at Midway
from an epicenter near Honshu, a warning
could be sent to Honolulu. The travel time
difference between Midway and Honolulu
of about 2 hours and 50 minutes is readily
obtained from the chart. The necessary
condition for using the chart in this fashion
is that the outpost lie close to the great circle
course from the epicenter to Honolulu.
There are a number of other islands in the
Pacific Ocean whose positions make possible
their use as detector outposts for waves from
other directions.

Travel time to other places in the Ha¬
waiian Islands can be estimated with the aid
of the time curves near the islands. For
example, a seismic sea wave originating near
Chile would reach Hilo about a half-hour
before it reached Honolulu, but a sea wave
from southeast Alaska would arrive at both
places at approximately the same time.

Table 1 gives the geographic position of
epicenter, the Greenwich time of the seismic
disturbance, the location of the gage which
recorded the receipt of the seismic sea wave

188

PACIFIC SCIENCE, Vol. I, July, 1947

TABLE 1

Comparison of Observed and Computed Travel Times of Seismic Sea Waves

(Mean variation of computed travel times with respect to observed times — 2.3 per cent.)

EARTHQUAKE

LOCATION

OF

TIDE GAGE

OBSERVED

TRAVEL

TIME

TIME

DIFFERENCE

(COMPUTED

MINUS

observed)

Epicenter

Greenwich Time

Near

Lat.

Long.

Year

Month

Day

Hour

deg.

deg.

h. m.

h. m.

min.

Colombia

1 N..

80 W.

1906

Tan.

31

15 33

Honolulu

12 57

—27

Chile

33 S.

72 W.

1906

Aug.

17

00 41

Honolulu

15 12

— 2

Kermadec Is.

29.2 S.

177.0 W.

1917

May

1

18 27

Honolulu

8 01

— 6

Kuril Is.

46.5 N.

151.4 E.

1918

Sept.

7

17 16

Honolulu

6 36

+ 4

Tonga Is.

21.2 S.

172.5 W.

1919

April

30

07 17

Honolulu

6 36

— 4

Chile

29.0 S.

71.0 W.

1922

Nov.

11

04 32

Honolulu

14 58

+ 2

Kamchatka

54.0 N.

161.0 E.

1923

Feb.

3

16 02

Honolulu

6 18

+ 6

Kamchatka

55.7 N.

162.5 E.

1923

April

13

15 31

Honolulu

6 44

— 8

California

34.9 N.

121.0 W.

1927

Nov.

4

13 51

Hilo

5 08

— 3*

Mexico

16.2 N.

97.2 W.

1928

June

17

03 19

Hilo

8 30

0*

Aleutian Is.

51 N.

170 W.

1929

Mar.

7

01 35

Hilo

4 45

— 9=’.:

Solomon Is.

10.6 S.

161.7 E.

1931

Oct.

3

19 13

Honolulu

7 46

—11

Solomon Is.

10.6 S.

161.7 E.

1931

Oct.

3

19 13

Hilo

8 19

— 29*

Mexico

19.2 N.

104.2 W.

1932

June

3

10 37

Honolulu

7 42

+ 13

Japan

39.1 N.

144.7 E.

1933

Mar.

2

17 31

Honolulu

7 33

— I4f

Alaska

55.5 N.

157.3 W.

1938

Nov.

10

20 19

Honolulu

5 01

— 1

Chile

31.5 S.

71.4 W.

1943

April

6

16 07

Honolulu

15 31

—24

Japan

33 N.

137 E.

1944

Dec.

7

04 36

Honolulu

9 09

— 49f

Aleutian Is.

53.5 N.

163.0 W.

1946

April

1

12 29

Honolulu

4 34

— 4

Japan

33.5 N.

133.7 E.

1946

Dec.

20

19 19

Honolulu

9 32

— 23f

* Computed times to Hilo were estimated with the aid of near-by time curves,
t See text for a discussion of these differences.

in the Hawaiian Islands, the observed travel
time, and the difference between computed
and observed travel times. The computed
travel times to Hilo were estimated with the
aid of near-by time curves. A number of
other seismic sea waves have reached the
Hawaiian Islands and the available mari-
grams were examined. These were not in¬
cluded in the table because the amplitudes,
as recorded by the gages, were too small to
permit the determination of the first rise or
fall of the waves. The natural seiche con¬
dition at both Honolulu and Hilo was the
primary reason for this difficulty.

REFERENCES

Bodle, R. R. Note on the earthquake and seismic
sea wave of April 1, 1946. Amer. Geophys.
Union Trans. 27: 464-46 5, 1946.

British Association for the Advancement
of Science, Comn. on Seismol. Invest. Rpt.

(through 1917); Internatl. Seismol. Sum. (from

1918).

Green, C. K. Seismic sea wave of April 1, 1946,
as recorded on tide gages. Amer. Geophys.
Union Trans. 27: 490-500, 1946.

Heck, N. H. List of seismic sea waves. Union
Geodesique et Geophysique Internatl., Ann.
Com. pour l’ Etude des Raz de Maree, no. 4:
20-41, 1934.

International Seismological Summary. See
British Association.

Jaggar, T. A. The great tidal wave of 1946. Nat.
Hist. 55: 263-268, 1946.

Macdonald, G. A., F. P. Shepard, and D. C.
Cox. The tsunami of April 1, 1946, in the Ha¬
waiian Islands. Pacific Sci. 1: 21-37, 1947.
Powers, H. A. The tidal wave of April 1, 1946.

Volcano Letter (Honolulu), no. 491: 1-4, 1946.
United States Coast and Geodetic Survey.
Manual of tide observations, iv + 92 p., 1 pi.
Spec. Pub. 196, Washington, D. C., 1941.

- United States earthquakes, 1939. iii + 69

p., 1 pi. Serial 637, Washington, D. C., 1941.

- United States earthquakes, 1943. iv + 49

p., 2 pi. Serial 672, Washington, D. C, 1945.

- United States earthquakes, 1944. iv + 43

p., 2 pi. Serial 682, Washington, D. C., 1946.

NOTES

New Botanical Bibliography of Pacific Islands by
E. D. Merrill

Because of its general interest to scientists,
attention is called to a botanical bibliography just
published which was prepared by Dr. E. D. Mer¬
rill of the Arnold Arboretum, Harvard University.

The present edition, which follows two earlier
ones published in 1924 and 1937, contains 3,850
author entries of works printed between 1773 and
1846. The area covered is primarily the islands
of the Pacific lying between 30° N. and 30° S.
latitude, and extending from Juan Fernandez and
Hawaii in the east to the Carolines, Marianas,
New Caledonia, and the Santa Cruz Islands in the
west. The bibliography is enriched by concise
abstracts for each entry. The classified subject

index, added by Dr. E. H. Walker of the Smith¬
sonian Institution, is of great value, giving ready
reference to all articles published on any par¬
ticular botanical subject or area.

The full reference to the volume follows:
Merrill, Elmer D. A botanical bibliography of
the islands of the Pacific; Walker, E. H. A
subject index to Elmer D. Merrill’s t(A botanical
bibliography of the islands of the Pacific.” U. S.
Natl. Mus., Contrib. U. S. Natl. Herbarium 30
(1). 404 p. Washington, D.C., 1947. Published
and sold by the Supt. of Documents, Govt.
Printing Office, Washington 25, D.C. Price
$1.00.

Scientists and the Fortieth Anniversary of the
University of Hawaii

The University of Hawaii, at a celebration on
March 13-25 of the fortieth anniversary of its
founding in 1907, presented to the campus and
community of Hawaii many programs of interest
to scientists. To carry out the theme of "The
Pacific Era and Higher Education,” a number of
eminent men were brought to Hawaii by President
Gregg M. Sinclair to give addresses; the program
also offered exhibits, panel discussions, and the
installation at the University of a chapter of Sigma
Xi, national scientific fraternity.

Participants in the program included Dr. Karl
T. Compton, president of the Massachusetts Insti¬
tute of Technology; Dr. Charles Seymour, presi¬
dent of Yale University; Dr. Howard L. Bevis,
president of Ohio State University; and Dr. Har¬
low Shapley, director of the astronomical observa¬
tory at Harvard University, national president of
the American Association for the Advancement of
Science, and national president of the Society of
the Sigma Xi.

Dr. Compton, during the period, spoke on the
following topics: "Importance of Technological
Progress to the Pacific Society,” "Five Atomic
Bombs and the Future,” and "From the Threshold
of the Atomic Age.” Dr. Seymour spoke on
"Science and Men in the Pacific.” Dr. Shapley
spoke on "The Expanding Universe” and "Science

[ 1

in This Day,” and together with Dr. Compton
acted as installing officer for the new Sigma Xi
chapter. Other events included a luncheon of the
Volcano Research Association, an official visit to
the Bishop Museum, and addresses by the visitors
before both houses of the Territorial Legislature.

Natural science exhibits on the campus included
research carried on at the University Agricultural
Experiment Station, bacteriological specimens and
films, greenhouse demonstrations of nutritional
deficiencies in plants, physical and chemical opera¬
tions and instruments, and exhibits by the Pine¬
apple Research Institute, the U. S. Bureau of Ento¬
mology and Plant Quarantine Fruitfly Laboratory,
and the Hawaiian Sugar Planters’ Association Ex¬
periment Station. The Bishop Museum offered a
special exhibit of ethnographic, botanic, and zo¬
ologic collections from Pacific islands.

Charter members of the new chapter of Sigma
Xi installed on March 19 were: Joseph E. Alicata,
Earl J. Anderson, Harry L. Arnold, Eugene C.
Auchter, John H. Beaumont, Earl M. Bilger,
Leonora N. Bilger, David D. Bonnet, Robert C.
Brasted, Elizabeth D. W. Brown, Forrest B. H.
Brown, Peter H. Buck, George O. Burr, Oswald
A. Bushnell, Walter Carter, Edward L. Caum,
Harold E. Clark, Harry F. Clements, J. L. Collins,
Charles Montague Cooke, Jr., Arthur Lyman Dean,

? 3

190

PACIFIC SCIENCE, Vol. I, July, 1947

Charles H. Edmondson, Willard H. Eller, John F.
Embree, Charles J. Engard, George E. Felton,
Harvey 1/ Fisher, Theodore W. Forbes, William A.
Frazier, William O. French, David T. Fullaway,
Herbert E. Gregory, Rene Guillou, Christopher J.
Hamre, Ernestine K. Hamre, Floyd W. Hartmann,
Constance E. Hartt, Pauline Heizer, Robert W.
Hiatt, Frederick G. Holdaway, Kenneth R. Kerns,
Alvin R. Lamb, Maurice B. Linford, Katharine
Luomala, Maude F. Lyon, Harold L. Lyon, Eugene
G. McKibben, Joseph P. Martin, F. P. Mehrlich,

Alice A. Nightingale, Harold S. Palmer, John H.
Payne, Cyril E. Pemberton, Henry N. Peters,
Charles F. Poole, Donald P. Rogers, Harold St.
John, G. Donald Sherman, Christos P. Sideris,'
Janet M. Smith, Jared G. Smith, Carl H. Spiegel-
berg, Robert A. Spurr, Edward J. Stirniman,
William B. Storey, Otto H. Swezey, Richard
Kwock Tam, William E. Vinacke, Kenichi Wata-
nabe, Ernest C. Webster, Chester .K. Wentworth,
Juliette Wentworth, John Mason Young, and
Elwood C. Zimmerman.

OCTOBER, 1947

t

B4.CIFIC

SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

IN THIS ISSUE: Abbott — Brackish-Water Algae from Hawaii
• Wentworth — Prediction of Rainfall in Hawaii • Bartsch —
Corculum of Pacific and Indian Oceans • Fisher — Skeletons of
Recent and Fossil Gymnogyps • Finch — Mechanics of Explo¬
sive Eruption of Kilauea in 1924 • Hiatt — Ghost Prawns in
Hawaii • Neal — Manilkara Found on Oahu • NOTES

Published by

THE UNIVERSITY OF HAWAII
HONOLULU, HAWAII

00

Mi iw,

NOV 5

BOARD OF EDITORS

A. Grove Day, Editor-in-Chief, Department of English, University of Hawaii

Ervin H. Bramhall, Department of Physics, University of Hawaii

Vernon E. Brock, Director, Division of Fish and Game, Territorial Board of
Agriculture and Forestry

Harry F. Clements, Plant Physiologist, University of Hawaii Agricultural
Experiment Station

Charles H. Edmondson, Zoologist, Bishop Museum, Honolulu, T. H.

Harvey I. Fisher, Department of Zoology, University of Hawaii

Frederick G. Hold away, Head, Department of Entomology, University of
Hawaii Agricultural Experiment Station

Maurice B. Linford, Head, Department of Plant Pathology, Pineapple Research
Institute, Honolulu, T. H.

A. J. Mangelsdorf, Geneticist, Experiment Station, Hawaiian Sugar Planters’
Association, Honolulu, T. H.

G. F. Papenfuss, Department of Botany, University of California, Berkeley 4,
California

Harold St. John, Chairman, Department of Botany, University of Hawaii

Chester K. Wentworth, Geologist, Honolulu Board of Water Supply

SUGGESTIONS TO AUTHORS

Contributions to Pacific biological and physical
science will be welcomed from authors in all parts
of the world. Manuscripts should be addressed to
Dr. A. Grove Day, Editor, Pacific Science, Uni¬
versity of Hawaii, Honolulu 10, Hawaii. Use of
air mail for sending correspondence and brief
manuscripts from distant points is recommended.

Manuscripts will be read promptly by members of
the Board of Editors and by other competent
critics.

Manuscripts may run from 1 to 30 pages in
length. Authors should not overlook the need for
good brief papers presenting results of studies,
notes and queries, communications to the editor,
or other commentary.

Preparation of Manuscript

Although no manuscript will be rejected merely
because it does not conform to the style of Pacific
Science, it is suggested that authors follow the
style recommended below and exemplified in the
journal.

Title. Titles should be descriptive but brief. If a
title runs to more than 40 characters, the author
should also supply a “short title” for use as a
running head.

Manuscript form. Manuscripts should be typed
on one side of standard-size, white bond paper
and double-spaced throughout. Pages should be

consecutively numbered in upper right-hand cor¬
ner. Sheets should not be fastened together in any
way, and should be mailed flat. Inserts should be
either typed on separate sheets or pasted on proper
page, and point of insertion should be clearly indi¬
cated.

Original copy and one carbon copy of manuscript
should be submitted. The author should retain a
carbon copy. Although due care will be taken, the
editors cannot be responsible for loss of manu¬
scripts.

Introduction and summary . It is desirable to state
the purpose and scope of the paper in an intro¬
ductory paragraph and to give a summary of
results at the end of the paper.

Dictionary style. It is recommended that authors
follow capitalization, spelling, compoundings, ab¬
breviations, etc., given in Webster’s New Inter¬
national Dictionary (unabridged), second edition;
or, if desired, the Oxford Dictionary. Abbrevia¬
tions of titles of publications should, if possible,
follow those given in U. S. Department of Agri¬
culture Miscellaneous Publication 337.

Footnotes. Footnotes should be used sparingly and
never for citing references (see later). Often,
footnotes may better be incorporated into the text
or omitted. When used, footnotes should be con¬
secutively numbered by superior figures through-
[ Continued on inside back cover ]

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

Vol. I OCTOBER, 1947 No. 4

Previous issue published August 9, 1947

CONTENTS

PAGE

Brackish-Water Algae from the Hawaiian Islands. Isabella A. Abbott ... 193

Cycles in Rainfall and Validity in Prediction of Rainfall in Hawaii. Chester K.

Wentworth . 215

The Little Hearts ( Corculurn ) of the Pacific and Indian Oceans. Paul Bartsch . 221

The Skeletons of Recent and Fossil Gymnogyps. Harvey I. Fisher . . . . 227

The Mechanics of the Explosive Eruption of Kilauea in 1924. R. H. Finch . . 237

Ghost Prawns ( Sub-Family Luciferinae ) in Hawaii. Robert W. Hiatt . . . 241

A Manilkara Found on Oahu, Hawaii. Marie C. Neal . 243

Notes:

Opportunities for Financing of Research in the Pacific Under the Fulbright Act 245
Editor's Comments . 246

Index to Volume I . 251

Cover drawing by A. S. MacLeod

Pacific Science, a quarterly publication of the University of Hawaii, appears in January,
April, July, and October. Subscription price is three dollars a year; single copies are one
dollar. Check or money order payable to University of Hawaii should be sent to Pacific
Science, Office of Publications, University of Hawaii, Honolulu 10, Hawaii.

Brackish-Water Algae from the Hawaiian Islands1

Isabella A. Abbott2

Marine algae from the Hawaiian Islands
(in older literature, the Sandwich Islands)
have been collected or studied by a few phy-
cologists. Brackish-water species, however,
although essentially marine in relationship,
have not been systematically studied for this
area.

The present study of the brackish-water
species was begun late in 1943, in connection
with a survey co-operatively conducted by
the University of Hawaii and the Territorial
Board of Agriculture and Forestry, on fish¬
ponds bordering the ocean. At first, two
ponds on the island of Oahu were selected
for study. In the following year, however,
more data and investigation of other ponds
seemed desirable. Three additional ponds
on Oahu and eight on the island of Molokai
were selected and included in the survey.
The study of algae, as well as that of animal
life, was thus extended to include all these
fishponds.

It soon became apparent that the variety
of algal forms present in the chosen ponds
constituted so large a problem that it would
be best to enlist the aid of certain special¬
ists. Myxophyceae were sent to Francis
Drouet of the Chicago Natural History

1 Research Paper 7, Cooperative Fisheries Staff
of the Territorial Board of Agriculture and For¬
estry and the University of Hawaii. Manuscript
received February 28, 1947.

2 The author completed most of the work on
this paper while on the Department of Botany
staff, University of Hawaii, and finished the

.manuscript at the University of California, Berk¬
eley, with the aid of the excellent algal herbarium
(cited as U. C. Herb.) and library at the latter
institution.

Museum and diatoms were sent to Paul
S. Conger of the U. S. National Museum,
where earlier collections of Hawaiian dia¬
toms are deposited. In the present report are
included the bulk of the Chlorophyceae, and
all Phaeophyceae and Rhodophyceae.

In the course of this study, a preliminary
account of some of these algae by J. T.
Conover, made under the direction of G. F.
Papenfuss at the University of California,
was received. Since the majority of the spe¬
cies in the notes of Conover had already
been determined, there was little in his ac¬
count that requires special mention here.

The present writer reports 9 genera of
green algae, 1 of brown, and 1 1 of red. The
algae studied are in the writer’s herbarium;
duplicate sets will be deposited in the her¬
barium of the Bishop Museum, and else¬
where if quantity permits.

Species listed seem to include those previ¬
ously known only from the marine, or only
from fresh-water, habitats, with the excep¬
tion of one species of Polysiphonia. This
genus, to the writer’s knowledge, is known
only from strictly saline or brackish-water
habitats; it was found in a fresh-water
pond, and was accompanied by such a well-
known fresh-water genus as Spiro gyra.

The writer is indebted to Charles Engard,
who collected nearly all the specimens of
algae. Members of the University fisheries
staff, especially Yoshinori Tanada, were
extremely helpful with habitat data. Thanks
are extended to Kazue Watanabe, who exe¬
cuted most of the drawings.

NOV 5

(

194

PACIFIC SCIENCE, Vol. 1, October, 1947

HABITAT

The fishponds already mentioned are dis¬
cussed by Hiatt (ined.) from the standpoint
of construction and as a habitat for fish and
invertebrates. These ponds are usually en¬
closures along the seacoast but may at times
be exposed directly to the sea. They are en¬
closed by a stone or mud wall oceanward.
Into some ponds fresh- water streams enter
on the land side. The ponds were handed
down among the natives for generations, and
were used mainly for the purpose of raising
fish for the kings and chieftains of the vari¬
ous islands until 50 years ago, when they be¬
came somewhat commercialized. Several of
the ponds under study, however, are still
held within the original families who were
given the ponds by chieftains.

The continued utilization of these fish¬
ponds no doubt has been encouraged by the
desire of various racial groups in the Ha¬
waiian Islands for certain kinds of table fish
which live in brackish water, at least dur¬
ing part of their life history. It was found
by the zoologists in the 1946 biological sur¬
vey that the two most desirable fish, mullet
( Mugil cep halos ) and milkfish or awa
( Chanos chanos ), fed largely on micro¬
benthos, and the milkfish fed also on larger
algae if they were available ( Hiatt, loc. cit. ) .
Identification of the dominant algae was
made in order to help "farm” the fishponds
intelligently, as it was found that the food
chain ultimately rested with some of these
forms.

These ponds furnish a type of habitat the
study of which may give additional ecolog¬
ical information. They are not usually more
than 6 feet in depth, and thus would allow
the penetration of light sufficient for the
growth of algae. However, the water is
usually turbid and stirred by the wind. The
bottom, consisting of mud, or more rarely
mud and sand, is frequently moved by the
ebb and flow of the tide through the gates
from the adjoining ocean. In most ponds,

therefore, the algae must of necessity be at¬
tached to the walls or on halophytic phanero¬
gams [Bat is maritima, Halophila ovalis (R.
Br.) Hooker; see Fig. 1].

Fig. 1. Halophila ovalis (R. Br.) Hooker.
Habit of a portion of a plant from Molii Pond.
Natural size.

Most of the algae show an affinity for the
more marine (sea wall) portions of the
ponds. They are found abundantly in such
localities, and but sparingly in other regions
of the ponds. The fresh-water species are
restricted to the mouths of streams where
there is little contact with the ocean, or to
fresh-water ponds which are apparently fed
by artesian wells. Diatoms are abundant
epiphytically, and in some ponds form a
thick mat on the floor of the pond, mixed
with other small algae and the larvae of cer¬
tain animals.

No seasonal variation in the appearance
and disappearance of various forms was
noted.

Brackish-Water Algae from Hawaii — Abbott

OCCURRENCE OF ALGAE

The ubiquitous Enteromorpha appeared
in all stations of the ponds, in water ranging
in salinity from that of the open ocean to
almost fresh. Two distinct species, showing
much variability, were recognized. The two
species of Ulva present, on the other hand,
showed distinct preference for the more
saline portions. Cladophora, that trouble¬
some entity, was present in all situations;
such distribution would lead one to believe
that variations of marine and fresh- water
forms are to be found mixed in the ponds.
Other green algae did not occur in great
quantity, as did the three just mentioned.

True fresh-water algae (Spiro gyra, des-
mids) were recorded from the fresh- water
ponds.

The only brown alga found was Ectocar-
pus indicus, which occurred as a common
epiphyte on other algae as well as on other
plants. The variation shown by this species
led to a critical examination of all species
which might be growing in this area. The
results, which are included with the species
description, require placing in synonymy
two species, one of which was so placed
quite independently of Boergesen (1941).

The red alga collected most frequently
(from all but two ponds) was a species of
Polysiphonia, described here as new to sci¬
ence. Another species of Polysiphonia found
abundantly in fresh water could not be iden¬
tified with the foregoing species, nor with
any other, as it lacked reproductive organs.
This seems to be a new record for the occur¬
rence of this genus in fresh water (chlorin-
ity values were 2.48 to 2.54 parts per thou¬
sand). This species was found in a fresh¬
water pond, strangely called Salt Lake, which
is apparently fed by subterranean wells. An
old connection with the sea has apparently
been sealed off. Other red algae found in
some quantity are Erythrotrichia carnea,
Centroceras clavulatum, and species of C era-

195

mium. The red algal species were nearly in
contact with the ocean and thus may be
thought to be marine, with the exception of
the fresh-water Polysiphonia.

KEY TO THE ALGAE

A simple key is presented to aid the inter¬
ested reader with some botanical training to
identify as far as genera the algae found in
the fishponds. Technical terms have there¬
fore been reduced to a minimum.

PartI. Chlorophyceae. The grass-green
algae.

1. Plants with uninucleate cells . 2

1. Plants multinucleate or with multi-

nucleate cells . . . 4

2. Parenchymatous . 3

2. Filamentous, unbranched, free-
floating, with spiral chloroplasts

. Spiro gyra, p. 196

3. Tubular plants, one layer thick in

section .. . Enteromorpha, p. 196

3. Foliose plants, membranous, with
simple to cleft margins, two layers
thick in section . Ulva, p. 197

4. Filaments unbranched . 5

4. Filaments profusely branched . 7

5. Entangled . 6

5. Attached, rhizoids only from the

basal cell, cells often bulbous .

. Chaetomorpha, p. 197

6. Floating, or if attached with short
rhizoidal branches along the en¬
tire attaching length, with few to
several nucl eL.Rhizoclonium, p. 197
6. Matted, completely multinucleate,
non-septate except at the location

of reproductive organs .

. V aucheria, p. 198

7. Plants in bushy soft tufts, the
branches lateral, opposite, alternate,
or fascicled, cut ofif from the main

axis . . Cladophora, p. 198

7. Branches not cut ofif from the main

axis

.8

196

PACIFIC SCIENCE, Vol. 1, October, 1947

8. Branches radial, the plant erect,
without trabeculae.-Bfyfl/uxr, p. 198

8. Branches opposite mainly (in
some marine species whorled),
the plants with rhizomatous base
and erect leaf -like portions, with
trabeculae . Caulerpa, p. 199

Part II. Phaeophyceae. The brown algae.
Plants epiphytic, branches secund, with
sporangia pedicellate or sessile, lat¬
eral or terminal . Ectocarpus, p. 199

Part III. Rhodophyceae. The red algae.

1. Plants epiphytic, filamentous, mostly
microscopic, with little differentia¬
tion of reproductive parts, asexual
reproduction mainly by monospores.. 2

1. Plants erect, filamentous to com¬
pressed, wiry to firm and cartila¬
ginous, asexual reproduction by tet-

raspores . 3

2. Thalli erect or creeping, uniseri-
ate; in well-developed specimens
3 to 4 cells thick at base, chro-

matophore stellate .

. Erythrotrichia, p. 200

2. Thalli erect, arising from a pseu-
doparenchymatous base, or from
a single cell, branches chiefly lat¬
eral, chromatophores parietal .

. Acrochaetium, p. 203

3. Axis flattened . 4

3. Axis terete . 5

4. Branching pinnate, tetrasporangia
cruciate, in sori in swollen lateral

branches . Gelidium, p. 203

4. Branching dichotomous or lat¬
eral, tetrasporangia cruciate, scat¬
tered in the thallus .

. Grateloupia, p. 205

5. Plants wiry, sparingly branched, in

matted tufts . W urdemannia, p. 204

5. Plants flaccid, bushy, with many

branches . 6

6. Tetrasporangia zonate, branches

frequently terminating in a hook

. . . Hypnea, p. 206

6. Tetrasporangia cruciate, branches

free . 7

7. Plants large, the thallus cylindrical
and parenchymatous.-Crraci/draz, p. 206

7. Plants mainly epiphytic, essentially

filamentous . . 8

8. With cortications, the superficial
cells shorter than the central cell„9

8. Without cortications, the super¬
ficial cells as long as the central

cell . . ............10

9. Plants corticated at the nodes, or but

little beyond..... . Ceramium, p. 208

9. Cortication throughout of small
rectangular cells in a longitudinal

series . . . Centroceras, p. 207

10. Colorless hairs lateral, spiralling,
male and female reproductive
structures borne in connection
with the hairs ..Polysiphonia, p. 212

10. Hairs in threes, terminal, tetra¬
sporangia borne on the deter¬
minate branches ..Taenioma, p. 210

DESCRIPTIONS OF SPECIES

CHLOROPHYCEAE

Spirogyra Link, 1820: 5

Two species are distinguished in this
genus on the basis of number of chloroplasts.
One species is from Salt Lake, and the other
from a fresh-water pond adjacent to Uala-
pue. Neither species can be identified be¬
cause of the absence of fertile material.

Enteromorpha Link, 1820: 5

Enteromorpha jlexuosa ( Wulfen) J. Agardh,
Till Alg. Syst. (3) : 126, 1883.

Plants tufted, to 14 cm. in height, usually
less, crenelated or simple, with little or gen¬
erally no branching. Cells usually arranged
longitudinally in a straight series (Setchell
and Gardner 1920: 256). Specimens com-

Brackish-Water Algae from Hawaii — Abbott

pare favorably with those in U. C. Herb.,
and one in herbarium of Bishop Museum.
Hawaiian specimens examined: Tilden 558
(Bishop); Reed 111, 198, 312, 452, 487,
515 (U.C.).

Found in Kuapa Pond at various stations,
Kihalaeo Pond, Kupeke, Keawanui, Niau-
pala, Ilae, and Molii.

Reported from the Hawaiian group by
Lemmermann (1905), Reed (1907), Rock
(1913), and MacCaughey (1918).

Enteromorpha intestinalis (Linn.) Link,
Epistola, p. 5, 1820.

Plants to 20 cm. in height, much branched,
the branches whorled, alternate, or opposite.
Cells angular in surface view. Some of the
specimens from the ponds are in agreement
with Setchell Hawaiian Algae 1 6, deter¬
mined by Collins (U.C.).

In all the major ponds.

Reported from the Hawaiian Islands by
Reed, Setchell (1905), Rock, and Mac¬
Caughey.

Ulva Linnaeus, 1753:1163

Ulva Lactuca Linn., Sp. Plantarum 2: 1163,
1753.

Plants to 15 cm. in height, expanded, with
scalloped margins, attached to rocks, twigs,
wood, and other algae. Plants are usually
light green in color. Base with many rhi-
zoids.

In all the major ponds.

Reported from the Hawaiian Islands by
Lemmermann, Reed, Rock, and Mac¬
Caughey. Abundant in marine habitats.

Ulva jasciata Delile, Flore d’Egypte, p. 153,
1813 (see Fig. 2).

Dark green plants to 20 cm. in height,
cleft and digitate, attached to rocks, forming
large patches. Plants not as membranous as
U. Lactuca. Rhizoids are few.

197

Collected in Kihaloko, Keawanui, Uala-
pue, Niaupala, all on Molokai Island.

Reported by Setchell, Reed, Tilden
(1901^), Reinbold (1907), Rock, and Neal
(1930).

Fig. 2. Ulva jasciata Delile. Habit of a plant
from Keawanui Pond. */4 natural size.

Chaetomorpha Kiitzing, 1845: 203

Chaetomorpha aerea (Dillwyn) Kiitzing,
Sp. Alg., 379, 1849.

Tufts of unbranched filaments to 4 mm.
in height. Cells inflated, to 120 ^ in width,
the walls thickened and stratified.

Molii on Batts maritima, Kupeke on a
bivalve, Keawanui on pond wall.

Chaetomorpha antennma has been re¬
ported by Reed, and I have examined her
collections of this species. None are of the
Ch. aerea type. Others reporting other spe¬
cies of Chaetomorpha from Hawaii are
Chamberlain (I860), Reinbold, Lemmer¬
mann, and MacCaughey. Setchell, Mac¬
Caughey, and Rock also report Ch. antennma.

Rhizoclonium Kiitzing, 1843: 261
Rhizoclonium sp.

Only one undeveloped specimen, whose
determination to species is not possible.

Ualapue Pond, entangled on Polysiphonia.
Newly reported here for the islands.

198

PACIFIC SCIENCE, Vol. 1, October, 1947

Vaucheria De Candolle, 1801: 17

Plants matted, floating, or on the bottom
of shallow parts of the ponds. Filaments
entangled, coenocytic, with spherical oogonia
or antheridia. These structures are stalked
or (in ours) sessile.

Three types of plants are found in the
ponds, one with no reproductive organs.
Determination is thus incomplete for this
specimen. The genus is newly reported from
the Hawaiian Islands.

Vaucheria dichotoma (Linn.) C. Agardh,

Synop. Alg. Scand., 47, 1817.

Large, free-floating masses, in the sum¬
mer with oogonia and antheridia. Oogonia
sessile, globular; antheridia with a terminal
opening, sessile, shaped much like oogonia
but slightly more elongate.

Kuapa Pond. In abundance.

Vaucheria Thuretii Woronin, in Bot. Zeit.,

157, 1869.

Filaments 60-80 /* in diameter, in dense
patches. Plants monoecious, with sessile
spherical oogonia; antheridia curving to a
lateral pore.

Keawanui Pond. Rare.

Vaucheria sp.

A sterile specimen, with measurements
smaller than the two species just mentioned,
was found in Molii Pond attached to a mol-
lusk.

Cladophora Kiitzing, 1843: 262

Plants bushy, to 15 cm. in height, with
prominent lateral branches cut off from the
main axes. Each cell multinucleate with
many chloroplasts.

A most variable genus, with representa¬
tives by far the most commonly found in
brackish water. Identification cannot be
made with certainty without a large amount
of comparative material. Lacking this, de¬
termination must be left at the genus.

In all the ponds, but in abundance espe¬
cially in the more saline ones.

Compared with the published illustra¬
tions of Brand (1905), these specimens
show much variation, and relationships can¬
not be established without examining the
specimens used by him.

Bryopsis Lamouroux, 1809^: 133
Bryopsis pennata Lamouroux var. secunda

(Flarvey) Collins and Hervey, Amer.

Acad. Arts and Sci., Proc. 53: 62, 1917.

Only one plant, 2 cm. in height, was col¬
lected, in the more saline portions of Kea¬
wanui Pond. The main branches arise from
a rhizoidal base, and branch in a pinnate
manner, the pinnules opposite each other.
A few branches show the secund type of
orientation. The plant is dark green.

Distribution: Florida, Bermuda, Bahamas
and islands of the West Indies to Aruba
Island, Netherlands West Indies (for the
species) .

A rather exhaustive comparison has been made
of this specimen with other Bryopsis specimens
from this area, and also from other parts of the
world, with emphasis on tropical forms. These
comparisons have been made in the University of
California Herbarium.

With certain specimens from Hawaii (Reed
1173, 1150, 1068 ) our specimen is in good agree¬
ment. Close comparison is also possible with
plants from Samoa and Tahiti (Setchell 1087,
Tutuila, and $ etch ell and Barks 5185, Tahiti, both
as B. Harveyana), from Formosa (legit Y.
Yamada), and to some extent specimens from
Dwarka (Boergesen 5534 as B . plumosa ), the Gulf
of California (Turner’s Island, Dawson 688 as
B. plumosa var. pennata ), and from New Guinea
(Kdruhach 29, det. Grunow, as B, Harveyana).
Specimens on sheets 341389, 341498, and 341390
which have been determined as B. Harveyana by
Setchell show little agreement with our specimens.
These plants are from the Malay Peninsula. Like¬
wise little similarity can be detected between the
Hawaiian specimens and those of the Atlantic
determined as B. plumosa (exemplified by Setchell
150 from Woods Hole), or of the Caribbean
forms of this species.

The North American specimens seem to be
Boergesen’s variety typica of B. plumosa. Boerge-
sen’s varieties pennata, Harveyana, and Leprieurii
of B. plumosa (1911: 147; 1913: 115) show more

Brackish-Water Algae from Hawaii — Abbott

similarity to variety typica than do our specimens
to any of these. Boergesen (1946: 341) has raised
B. Harveyana to specific rank. Setchell’s B. Har¬
veyana of Samoa (1924) and Tahiti (1926)
should not be included in synonymy with B.
plumosa var. Harveyana, as those specimens are
quite different from Boergesen’s specimens.

It is obvious from the literature that there is
great confusion as to species limits in this genus.
Lacking critical specimens, I am following the
interpretation of Taylor (1928) and of Collins
and Hervey (1917). At best, this practice is not
too satisfactory, as the Pacific plants by and large
are somewhat different from the Atlantic and
Caribbean forms. A large suite of specimens and
comparison with type material are desirable.

Br y op sis plumosa has been reported from the
Hawaiian Islands by Chamberlain, Tilden, and
MacCaughey. I have examined the Tilden speci¬
men (453) in American Algae Century V
(190l£), and find it to be identical with ours. In
all probability, the records are in agreement with
the material mentioned here.

Caulerpa Lamouroux, 1809^: 136
Caulerpa Sertularioides (Gmelin) Howe in

Torrey Bot. Club Bui. 32: 576, 1905.

Plants to 4 cm. in height, creeping parts
clinging tightly to sand and rock particles,
the upper portions flattened laterally, with
opposite or nearly opposite "leaflets.”

At one station, adjacent to the main
regions of Kuapa Pond, near the connections
to the sea.

The genus is common in marine habitats
in the Hawaiian Islands; this species and one
related closely to it are among the more
prominent members of the genus in Hawaii.
Reported from the Hawaiian group by
Eubank (1946).

PHAEOPHYCEAE

Ectocarpus Lyngbye, 1819: 130

Plants tufted, filamentous, arising from a
single basal cell, or from a group of cells,
with or without rhizoids at the base, attached
to twigs, wood, or other algae. Plants in
this series to 6 cm. in height, but more
usually 1-2 cm. Reproduction by plurilocu-
lar or unilocular sporangia.

199

In the sense of Hamel (1939: 66-67)
the species which is described below would
more properly fit in the segregated genus
Feldmannia, which differs in a few char¬
acters — mainly in that it has discoid chro-
matophores, whereas the limits of Ectocarpus
are such as include only those members of
this complex which have ribbon-like chro-
matophores.

One of the characters used to separate
Feldmannia further is the strong ramifica¬
tions of filaments near the base, a character
not shown by our plants. Sporangia are typ¬
ically pedicellate in Feldmannia, a condition
only infrequently occurring in our speci¬
mens.

Giffordia, in the sense of Hamel, is char¬
acterized by discoid chromatophores also,
but sporangia are always sessile, and the
plant has intercalary growth. These are not
constant characters in our plants.

It would thus seem best to retain the spe¬
cies below in the genus Ectocarpus, sensu
latiore, until European workers who have
authentic material can establish further char¬
acters for the separations so badly needed in
this complex.

In making a survey of literature pertinent
to Hawaiian material, some confusion was
encountered with Ectocarpus Duchassaingia -
nus Grunow (1870), recorded for the Pa¬
cific from Samoa (Setchell, 1924). From
the literature, this species seemed identical
with Ectocarpus indie us Sonder, recorded
earlier by Weber- van Bosse (1913: 129)
from Malaya. The writer has examined ma¬
terial used by Setchell in his study, and has
concluded that his plants are synonymous
with E. indicus Sonder.

When these studies were completed, re¬
prints of Boergesen’s instructive Mauritius
papers were received, and it was interesting
to find that he had come to the same conclu¬
sions regarding E. Duchassaingianus , which
he based on specimens from the Danish
West Indies (Virgin Islands).

200

Examination of a large number of speci¬
mens collected in the fishponds and in
marine habitats, as well as those deposited
in the herbarium of the Bishop Museum,
Honolulu, and the herbarium of the Univer¬
sity of California, Berkeley, has led the
writer to believe that there is much varia¬
tion in specimens as well as in interpretation
of Ec to car pus in die us. These notes, and
those that directly follow, are a result of the
examination of a large series. Such studies
have led the writer to consider Ectocarpus
Mitch ellae in the sense of Saunders (1901,
in Tilden 1901&) and E. Sargassi Saunders
as identical with the material in the present
investigation. These specimens are from
Hawaii and are distributed by Tilden
(American Algae Century V, nos. 439, 440a,
and 440b). I have examined both Bishop
Museum and University of California speci¬
mens of the exsiccatae and find them iden¬
tical with E. indicus.

Ectocarpus indicus Sonder, in Zollinger, H.

Verzeichn. . . . indischen Archipel. . . .

1842-48, p. 3 (not as usually cited: in A.

Moritzi, Syst. Verzeichn, 1857) (see Fig.

3 a-d).

Ectocarpus Duchassaingianus Grunow, Alg.

Novara, 1870, p. 45.

Ectocarpus Sargassi Saunders, in Tilden

American Algae Century V, nos. 440a and
440b, 1901 h.

Plants tufted, branching primarily dicho¬
tomous with many lateral branches, the main
branches about 10 n wide. Sporangia at¬
tached, sessile or stalked, on the inner sur¬
face of the branches, oval to oval-clavate,
distributed throughout the plant. Plants ris¬
ing from a creeping base.

Abundant in Kuapa Pond, usually on
Bat is maritima; Wailupe, Molii Ponds, on
Oahu Island. Also found in Keawanui, Ku-
peke, Ualapue, Niaupala Ponds, Molokai
Island.

PACIFIC SCIENCE, Vol. 1, October, 1947

In the marine habitat, commonly occur¬
ring on species of Sargassum.

Specimens examined ( in U. C. Herb. ) : as
Ectocarpus indicus, Potts 1173a, determined
by Setchell, from Samoa; Lindauer 28, det.
Lindauer, from New Zealand; Nasr 259
from Egypt (Red Sea); Li 124 ex herb.
Tseng, det. Setchell, from China; K. G.
Iyengar 83, det. Gardner, from Bombay; as
E. Duchassaingianus: Tilden 32, det. Tilden,
from Tahiti; Boergesen 1093, 1250, det.
Boergesen, from the Virgin Islands3; Hamel
45, det. Hamel, from the French Antilles;
Taylor 39308, 39602, det. Taylor, from the
Netherlands West Indies.

Tilden 440a, 440 h, as Ectocarpus Sargassi
Saunders, det. Saunders, in Tilden American
Algae Century V; 439 as E. Mitchellae, det.
Saunders. These specimens are from the
Hawaiian Islands. (Exsiccatae from Bishop
Museum and University of California Her¬
baria examined.)

Distribution: Throughout warmer seas.

Previously reported by Reed, MacCaughey,
Lemmermann, and Neal.

RHODOPHYCEAE

Erythrotrichia Areschoug, 1847: 209
Erythrotrichia carnea (Dillwyn) J. Agardh,

Till Alg. Syst. (6): 15, 1883.

Plants attached singly, or in small loose
tufts, to 4 cm. in height, the uniseriate fila¬
ments attached to other algae by a single
disk-shaped basal cell which may become
lobed. The lower, older parts of the plant
may be two or three cells in width. Chro-
matophore stellate, with a prominent pyre-
noid. Reproductive structures not seen.

Found in all major fishponds, epiphytic
on Enteromorpha spp., Grateloupia, Poly-
si phonia, and Gelid ium. Erythrotrichia car¬
nea is here reported from the Hawaiian

3 These specimens as well as others collected by
Boergesen in the Virgin Islands and identified as
E. Duchassaingianus (1913: 159) were trans¬
ferred by him to E. indicus (1941: 16).

Brackish-Water Algae from Hawaii — Abbott

201

Fig. 3. Ectocarpus indicus Sonder. A. Portion of plant with plurilocular sporangia. Scale B (divi¬
sions 50 microns). B. Portion of plant showing chromatophores. Scale B (divisions 50 microns). C.
Basal portion. Scale A (divisions 100 microns). D. Portion of plant with mature unilocular sporan¬
gium. Scale B (divisions 50 microns).

202

PACIFIC SCIENCE, Vol. 1, October, 1947

Fig. 4. Acrochaetium robustum Boergesen. A. Habit of plant with
monosporangia. Note single basal cell. Scale B (divisions 50
microns). B. Portion of plant with part of multicellular base. Scale
C (divisions 10 microns). Acrochaetium s erratum Boergesen: C.
Portions of plants showing monosporangia. Scale C (divisions 10
microns).

Brackish-Water Algae from Hawaii — Abbott

Islands for the first time. It occurs com¬
monly on many marine algae in this area.

Distribution: Occurring commonly in the
tropics, epiphytic on littoral and sublittoral
algae; extending into temperate waters as far
north as England (type locality).

Acrochaetium Nageli, 1861: 402

The treatment of this genus is based on
the work of Papenfuss (1945), who has
seen our specimens.

Acrochaetium robustum Boergesen, Mar.

Alg. D. W. I., 2 (1): 40-43, 1915 (see

Fig. Aa-b).

Plants tufted, to 1 cm. in height, epiphy¬
tic, with a simple or branched basal cell
partly to wholly endophytic. Branching pre¬
dominantly lateral, sometimes alternate, but
at the base dichotomous. Cells with a parie¬
tal chromatophore and a single pyrenoid,
and with pit connections readily seen. Cells
to 4 fx in width, and twice as long. Mono¬
sporangia prominent, sessile or pedicellate,
appearing laterally near the tips of the
fronds, singly or in pairs. Terminal spo¬
rangia are often seen. Sexual organs not
seen.

Found in Kuapa Pond, epiphytic on Batis
maritima, and a piece of coniferous wood.
Not the same species as other marine speci¬
mens from this area. The genus is newly
reported from the Hawaiian Islands.

Distribution: Danish West Indies, Japan.

Acrochaetium seriatum Boergesen, Mar. Alg.

D. W. I., 2 (1): 32-35, 1915 (see Fig.

Ac).

Plants to 5 mm. in height, tufted, soft,
epiphytic on Ulva fas data. Branching
chiefly lateral, occasionally alternate. Erect
portions arising from a multicellular creep¬
ing base. Monosporangia borne singly or in
clusters, sessile or pedicellate. In Keawanui
Pond, Molokai Island, on Ulva fas data.

Distribution: Danish West Indies, Ma¬
cassar.

203

Both these species of Acrochaetium were
collected in the more saline portions of
Kuapa and Keawanui. In all probability
they are not true brackish- water species.
Species of Acrochaetium occur frequently on
larger algae in the marine environment in
this area.

Gelidium Lamouroux, 1813: 40

Gelidium pusillum (Stackhouse) Le Jolis in
Soc. Sci. Nat. Cherbourg, Mem. 10: 139,
1863 (see Fig. 5).

Fig. 5. Gelidium pusillum (Stackh.) Le Jolis.
Habit of portion of a plant. X 4.

Plants small, associated loosely in clusters,
with a creeping base bearing erect parts to
3 cm. in height. Branching irregularly pin¬
nate or alternate, with both branches and
axis markedly flattened. Rhizines occupy a
large central area of the branches with small
cortical cells in three rows on the outside.

Tetrasporangia in small bulbous lateral
branchlets, sunken below the surface. Tetra-
spores in cruciate groups when mature, the

204

young tetrasporangia frequently showing
only one division. Sexual plants not seen.

Found in Molii Pond, Oahu; Ilae, Kea-
wanui, Kupeke, and Ualapue Ponds, Molo¬
kai, infrequently occurring, and with many
epiphytic diatoms and Erythrotrichia earned.

Distribution: In the tropics, northward to
England (type locality) .

This species is a variable one, and perhaps
these plants represent a distinct entity, but
for the present it seems best to identify them
with the species. All species of Gelidium
are named "limu loloa” or "limu ekaha-
kaha” by the Hawaiians. The species here
noted has been questioned by Reed. Mac-
Caughey lists the species. It does not seem
to be either of the Gelidium spp. listed by
Neal.

Gelidium pusillum var. conchicola Piccone

and Grunow in Piccone, Contribuzione

all’algologia Eritrea, Nuovo Giornale Bot.

Itah, 16: 316, 1884.

Plants smaller than the species, to 3 mm.
forming tufts, but inconspicuous. Blades
flattened especially at the tips. Rhizomes
with small colorless rhizoids. The plant is
sterile.

Fragment of plants from Molii Pond, as¬
sociated with Cladophora sp. and Polysi-
phonia. Newly reported from the Hawaiian
Islands.

Distribution: Bermuda, Florida, Colom¬
bia.

These plants agree favorably with de¬
scription and figures of Taylor (1928).

Wurdemannia Harvey, 1853: 245

Genus incertae sedis. Taylor, 1928, 1941,
1943, 1945, lists the genus with the Geli-
diales, but in 1940 lists it in the Cryptone-
miales (Rhodophyllidaceae) . Feldmann and
Hamel, 1934, list it in the Gelidiales.
Harvey (1853) and Boergesen (1919-20)
list it as uncertain.

PACIFIC SCIENCE, Vol. 1, October, 1947

W urdemannia miniata (Draparnaud) Feld¬
mann et Hamel in Revue Generate de Bo-
tanique 46: 545, 1934 (see Fig. 6; Fig.
lb).

W urdemannia setacea Harvey, Nereis Bor.
Am. (2): 245, 1853.

Fig. 6. W urdemannia miniata (Drap.) Feld¬
mann et Hamel. Habit of plant. xiy2.

Plants in thickly matted dark red tufts to
4 cm. in height, attached to the substratum
by numerous disk-shaped holdfasts from
which run several cylindrical rhizomes. Main
axis erect, sparingly branched, terete, and of
firm consistency. The young branches show
large medullary cells which become smaller
toward the outside. Superficial cells are
somewhat square and contain many chro-
matophores. In surface view, these cells are
small and irregularly shaped.

Tetrasporangia are found in sori near the
tips of the branches with few modified
sterile filaments. The sporangia are zonately
divided. Sexual plants are not known.

Found in Molii, Oahu Island; Ilae and
Keawanui, Molokai Island.

Distribution: Mediterranean (Montpellier,
type locality). Harvey’s plant was collected
at Key West. It is also known in the Atlantic
from the Brazilian coast northward to Ber¬
muda.

Reported here for the first time from the
Hawaiian Islands, and for the Pacific.

When sexual plants are known, this spe¬
cies will probably be segregated from the
Gelidiales because of its zonate tetrasporan¬
gia.

Brackish-Water Algae from Hawaii — Abbott

Grateloupia C. Agardh, 1820: 221

Grateloupia filtcina (Wulfen) C. Agardh,
Syst. Alg., 241, 1824.

Grateloupia filicina forma hawaiiana Mazza,
Nuova Not., 26: 75, 1915.

Plants purplish-red, cartilaginous, 14 cm.
in height, erect, the fronds attached in
groups by a single holdfast; branching with
a percurrent main axis, the whole plant as¬

205

suming a pyramidal shape. Branches to 3
mm. in diameter, linear, acuminate. Me¬
dulla composed of spherical cells of mod¬
erate size in transverse section, occasionally
showing stored dextrin particles, surrounded
on the outer surface by two layers of com¬
pact small cells. Plants collected are sterile.

Found in Keawanui, near pond gate in
quiet water, with much epiphytic Erythro-
trichia carnea.

Fig. 7. A. Gracilaria coronopifolia J. Agardh. Cross section of thailus. Scale A (divisions 100 mi¬
crons). B. W 'urdemannia miniata (Drap.) Feldmann et Hamel. Cross section of thailus. Scale C
(divisions 10 microns). C. Hypnea nidulans Setchell. Cross section of thailus. Scale B (divisions 50
microns). D. Hypnea nidulans Setchell. Cross section of portion of fertile branch showing tetra-
sporangia. Scale C (divisions 10 microns).

206

PACIFIC SCIENCE, Vol. 1, October, 1947

Distribution: Widely occurring in warm
waters, into colder regions.

This is an alga of choice eating qualities,
much desired by Hawaiians. It grows well
in various localities on the islands of Ha¬
waii and Maui especially, and is occasionally
seen in certain localities on the island of
Oahu, where it is said to have been planted
for the late Queen Liliuokalani. The alga
is known on Hawaii as "limu huluhulu-
waena,” and on Maui as "limu pakeleawaa.”
If it were not so well known, it would be
difficult to place the plants collected, because
they are sterile.

Grateloupia filicina has been reported
from the Hawaiian area by Reed, Rock,
MacCaughey, and Setchell. G. dichotoma,
reported by Chamberlain, has not been sub¬
stantiated.

Specimens examined (in Herbarium
Bishop Museum) : Drew 641, Waikiki;
Rock, Apr., May, 1908, Waikiki; Bailey in
1876, Lanai Island. Tilden 507, Kapaa
(Kauai Island), does not seem to be this
genus.

Gracilaria Greville, 1830: 121
Gracilaria coronopifolia J. Agardh, Sp. Alg.

2 (2): 592-593, 1852 (see Fig. la; Fig.

9>-

Gracilaria No. 1, No. 2, Neal, Hawaiian

Marine Algae, 68, Fig. ISb, 1930.

Plants erect, to 15 cm. in height, pink at
tips, white below, with one or more fronds
attached to a single holdfast. Branching
frequent, dichotomous, arcuate at tips, with
a corymbose aspect. Branches cylindrical in
transverse section with a large medullary
region of colorless rounded cells slightly
thickened, surrounded by a narrow cortical
layer and a layer of smaller superficial cells.
Female plants with prominent pink to red
cystocarps in the upper branches. Cysto-
carps with thick outer covering and a small
ostiole. Male and tetrasporangial plants not
seen.

Found in Keawanui and Ilae ponds,
Molokai.

Distribution: "ad Wahoo [Oahu] Insu-
larum Sandwicensium.” Apparently ende¬
mic to the Hawaiian Islands, and found
quite commonly in the marine habitat. Re¬
ported by Reinbold.

The "limu manauea” of the Hawaiians
(the "ogo” of the Japanese) is characteris¬
tic of sandy, sheltered areas about the
islands. Two other species of Gracilaria are
listed in the literature from Hawaii: G. con -
fervoides by Chamberlain and MacCaughey,
and G. euchemoides b y Chamberlain.
Neither of these species has been collected
by the writer. G. coronopifolia has been
mentioned by Chamberlain, Lemmermann,
Reed, Rock, MacCaughey, and Setchell, and
questioned by Neal. Neal’s two species (No.
1 and 2) are this species. Other forms from
the islands have been studied but it is difficult
to place them with certainty.

Hypnea Lamouroux, 1813: 131
Hypnea nidipca J. Agardh, Sp. Alg. 2(2):
451, 1852 (see Fig. 8).

Fig. 8. Hypnea nidifica J. Agardh. Habit of
a portion of a plant. X 2.

Brackish-Water Algae from Hawaii — Abbott

Plants erect to 20 cm. in height, or more
usually matted and attached epiphytically to
other algae, with slender branches alternate
or whorled and closely beset with very short
branchlets, the ultimate tips simple and
straight. Plants red to reddish-green when
fresh. In section, the main axis shows a large
medullary layer surrounded by cortical cells
in radial rows of smaller cells, and a some¬
what firm superficial layer of angular cells.
Tetrasporangia circling the base of the short
branchlets, the cortical cells forming distinct
radial filaments in these areas. Sporangia
zonately divided. Male and female plants
not seen.

Found in Ilae Pond, growing with Cera-
mium spp., and Polysiphonia. In the Ha¬
waiian area, common in sandy shallow
waters.

Distribution: "ad insulis Sandwich.” Re¬
ported by Reinbold from the Hawaiian
Islands, and from Malaya by Weber- van

Bosse (1928: 453).

Hypnea nidifica seems more closely re¬
lated to H. cornuta in habit than to H. cervi -
cornis, a comparison made by J. Agardh.
The anatomy of the frond is closer to the
next species than to the two species just
mentioned. Tetrasporangia are like those of
most other species.

The species has been listed by Chamber-
lain, Lemmermann, Reed, Setchell, Rock,
MacCaughey, and Neal. Hypnea armata has
been listed by Reed, Rock, and MacCaughey.
Hypnea cornuta, H. pannosa, and H. divari-
cata mentioned by Chamberlain have not
been verified.

Hypnea nidulans Setchell, Veg. Tutuila

Island, Carnegie Inst. Wash. 20: 161-

162, 1924 (see Fig. 7c-d) .

Plants smaller than H. nidifica, much
twisted and branched, growing in small tufts
among other algae. It is chiefly differen¬
tiated from H. nidifica in having tetraspo¬
rangia in nemathecia near the tips of the

207

fertile branchlets. In cross section, the cor¬
tical cells are smaller, with small uniform
superficial cells. Only tetrasporangial plants
were collected.

Found in Molii Pond. Newly reported
from the Hawaiian Islands.

Distribution: Samoa (type locality) ,
Setchell 1084, type, in U.C. Herb., Setchell
and Parks 3138 in Herb. Bishop Museum.
Also reported by Boergesen ( 1943 : 62 ) from
Mauritius and by Weber-van Bosse (1928:
454) from Malaya.

Fig. 9. Gracilaria coronopifolia J. Agardh.
Habit of a portion of a cystocarpic plant. Natural
size.

Centroceras Kiitzing, 1841: 731
Centroceras clavulatum (C. Agardh) Mon-

tagne, Flore d’Algerie, 140, 1840-1850.

Plants filamentous, usually in matted
tufts, stiff and brittle, or floating and en¬
tangled with other algae; dark purplish-red
in color, or pinkish where exposed. Fish¬
pond specimens are 6-8 cm. in height with
irregular branching and short internodes at
the tops of the branches, the tips sometimes
forcipate. Spines at the nodes most prom¬
inent in the upper parts of the filaments,
4 to 6; in the older parts usually 2 or de¬
ciduous. Spines are of two or more cells.
Cortical cells in section, 28 to 40 in the
nodal portions. Tetrasporangia sub-external,
formed in a horizontal row of 4 to 8 at the
nodes, tetrahedrally divided.

208

Found abundantly in portions of several
fishponds open to the sea: Molii on Oahu
Island; Ilae, Keawanui, Kupeke on Molokai
Island.

Widely distributed throughout tropical
waters. It is one of the most frequently en¬
countered of shallow-water algae in the
Hawaiian area. Tilden 401 (American
Algae, Century V) in Herbarium University
of California as Ceramium diaphanum is
Centroceras clavulatum. This specimen is
from the Hawaiian Islands.

The structure of plants of this species is
very variable, but the species is readily sepa¬
rated from the next (Ceramium) in being
more brittle and usually larger, and in pos¬
sessing continuous corticating cells.

It is listed from the Hawaiian Islands by

PACIFIC SCIENCE, Vol. 1, October, 1947

J. Agardh, Chamberlain, Setchell, Reed,
and Rock, and, as Ceramium clavulatum C.
Agardh, by Lemmermann and MacCaughey.

Ceramium (Roth) Lyngbye, 1819: 117
Ceramium is a large and complex genus.
The limits of many of the species, especially
the tropical ones, are as yet ill defined. Con¬
sequently, it seems best not to assign specific
names to either of the following species,
especially since not all necessary plants were
found.

Ceramium sp. (1) (see Fig. 10 a-b).

Plants 4-8 cm. in height, dark pink to red,
lying in soft tangled mats among other algae.
Base of plants sometimes with rhizoids, the
upper ends usually free. Branching irregu¬
larly dichotomous, with prominent forcipate

Fig. 10. Ceramium sp. (1). A. Nodal region showing cortication. Scale C (divisions 10 microns).
B. Tips of two plants, in the above figure showing forcipate nature. Scale C (divisions 10 microns).

209

Brackish-Water Algae from Hawaii — Abbott

210

tips in some plants, and only a suggestion in
others.

Specimens collected are sterile.

Found in Ilae Pond, Molokai Island.

Ceramium sp. (2) (see Fig. lla-b).

Plants 2-3 cm. in height, dark pink,
tufted, erect, branching with a percurrent
main axis, the tips dichotomous and occa¬
sionally forcipate. Outer edges of the
branches markedly dentate. Nodal bands
close in the younger parts, in the older parts
quite separated. Mature nodes marked by
three rows of flattened cells below the sep¬
tum, and smaller, angular cells above. Tetra-
sporangia 70 /x in diameter, sub-exposed,
with an upgrowth of smaller cortical cells
over them. Sporangia usually single at the

PACIFIC SCIENCE, Vol. 1, October, 1947

node, rarely two. Spermatangial and carpo-
gonial plants not seen.

Found in Ilae Pond, Molokai Island, and
Molii Pond, Oahu Island, in the more saline
portions adjacent to the sea.

Neither of these species of Ceramium
seems to be one of those reported previously
from the islands: C. Kuetzingianum by Mac-
Caughey and Ceramium sp. by Neal. Cera¬
mium diaphanum by Tilden is Centroceras
clavulatum.

Taenioma J. Agardh, 1863: 1256

A detailed account of this genus, and par¬
ticularly the following species, may be found
in Papenfuss (1944) and Tseng (1944).

Taenioma perpusillum (J. Agardh) J.
Agardh, Sp. Alg. 2 (3): 1257, 1863.

Fig. 12. Polysiphonia aquamara Abbott. Type specimens. Scale A (divisions 100 microns). A. Habit
of cystocarpic plant showing young and mature cystocarps. B. Shape of cystocarps from another
branch. C. Basal portion of tetrasporangial plant. D. Habit of plant showing tetrasporangia (cover
cells have been omitted).

Brackish-Water Algae from Hawaii — Abbott

211

Fig. 13. Polysiphonia sp. Habit of sterile plants from fresh-water pond (Salt Lake). Illustration
on left. Scale A (divisions 100 microns); on right. Scale B (divisions 50 microns).

212

PACIFIC SCIENCE, Vol. 1, October, 1947

Plants in small tufts, epiphytic on other
algae, or on small sessile animals, filamen¬
tous and soft. The determinate branches of
this species end in three hairs as opposed to
two terminal hairs in T. macrourum.

Specimens collected were sterile.

Keawanui Pond, Molokai Island, with
Polysiphonia.

This species is uncommon in this area.
Reported from the Hawaiian Islands by
Chamberlain and Papenfuss.

Polysiphonia Greville, 1824: 90
Section Oligosiphonia

Polysiphonia aquamara sp. nov. (see Fig.

12 a-d).

Plantae penicillatim fasciculatae, ad 8 cm.
longitudine sed plerumque minores, roseo-pur-
pureae, molles, erectae, basi rhizoideis permultis
praedito nec repente, ramis adscendentibus, alter-
nis, origine spiralibus. Trichoblasti in articulos
omnes insertae; cellulae pericentrales 4, haud
corticatae; tetrasporangia e ramulis secondariis
ultimis, plantae apicem versus locatis, orta, in
serie recta vel tortuosa. Plantae cystocarpicae
validiores, cystocarpis alternis, late urceolatis.

In Batidem maritimam vel plantas alteras
phanerogamicas, raro in lapides vel conchas obso-
letas, crescentes.

Plants tufted, without a prostrate creeping por¬
tion, to 8 cm. in height, usually smaller, reddish-
purple, soft. Base composed of many rhizoids.
Erect filaments arise from the base with branches
ascending, alternate and spiraling in a counter¬
clockwise directon, slightly divergent to with
branching chiefly at the tips. Trichoblasts fre¬
quently appearing whorled, inserted in the young
parts on every segment alternately, in the older
parts leaving scar cells. The branches of the tri¬
choblasts are 2 to 3 ranked, 50-200 m in length, in
the male plants covering V3 the length of the tips
of the axis. Pericentral cells 4, non-corticated.

Tetrasporangia borne in ultimate branches near
the tip of the plant, in a straight series or tortu-
lose, or when in the main branches occasionally
alternate. Spores dark brown to blackish, sporan¬
gia 4-partite, 20 n (usually less) in diameter.

Antheridial clusters borne on the basal un¬
branched portion of a trichoblast, alternate, sub-
cylindrical, with antheridia blunt to round. Cys-
tocarpic plants stouter than others, the cystocarps
alternate, broad urn-shaped, with pyriform carpo-
spores.

Type: Abbott 1535- legit C. J. Engard,
from Station 7, Kuapa Pond, Oahu Island,
epiphytic on Batis maritima. This number
includes tetrasporangial, cystocarpic, and
spermatangial plants, of which I designate
the tetrasporangial as the type specimen.
These plants were collected April 7, 1944.

This new species of Polysiphonia is very
abundant in the major fishponds, occurring
on Batis maritima, rocks, and dead shells.
It resembles P. subtillisima, which has also
been found outside the strictly marine en¬
vironment, but is distinguished from it in
that it has no prostrate creeping portion.

Polysiphonia sp. (see Fig. 13).

A species of Polysiphonia found in fresh
water (Salt Lake Pond, Oahu) ; grows abun¬
dantly on the pond walls, attached to phane¬
rogams, or to rocks. It has four pericentral
cells. Infrequent trichoblasts are found near
the tips of the plants. Although repeated
collections were made, no fertile material
was found. The salinity in this pond was
found to be nearly that of fresh water (2.5) .

To the best of my knowledge, this is the
first report of the genus in fresh water.

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12 (5): 61-110, pi. 7-22, 1926.

- - and N. L. Gardner. The marine algae

of the Pacific Coast of North America: II.
Chlorophyceae. Calif. Univ., Publ., Bot. 8 (2):
139-374, pi. 9-33, 1920.

Sonder, O. G. Algen. In H. Zollinger, Sys¬
tematise hes V erzeichniss der im indisc hen Ar chi-
pel in den Jahren 1842-1848 gesammelten sowie
der aus Japan empfangenen Pflanzen, vol. 1.
xii -f- 160 p., pi. Ill, Zurich, 1854. [Not seen;
cited by Boergesen 1941: 43.]

Taylor, W. R. Marine algae of Florida, with
special reference to the Dry Tortugas. Tortu-
gas Laboratory, Papers, Carnegie Inst. Wash.
Papers 25. v + 219 p., 37 pi. Washington,
D. C., 1928.

- Marine algae of the Smithsonian-Hartford

Expedition to the West Indies, 1937. U. S. Natl.
Mus., Contrib. U. S. Natl. Herbarium 28
(3): 549-561, pi. 20, 1940.

- Tropical marine algae of the Arthur

Schott Herbarium. Field Mus. Nat. Hist., Chi¬
cago, Bot. Ser. 20 (4) : 87-104, pi. 1-2, 1941.
- Marine algae from Haiti collected by H.

H. Bartlett in 1941. Michigan Acad. Sci., Arts,
and Letters, Papers 28: 143-163, 4 pi., 1943.

- — Pacific marine algae of the Allan Hancock

Expeditions to the Galapagos Islands. In Allan
Hancock Pacific Expeditions, vol. 12. iv -j- 528
p., 100 pi. Los Angeles, 1945.

Tilden, J. E. Collection of algae from the Ha¬
waiian Islands. Thrum’s Hawaiian Almanac
and Annual for 1902, 106-113. Honolulu,
1901(^).

- American Algae. Century V. Exisiccatae.

No. 401-500. Minneapolis, 1901 (£).

Tseng, C. K. Notes on the algal genus Taenioma.

Madrono 7 (7): 215-226, 1944.

Weber-van Bosse, A. Liste des algues du Siboga:

I. Siboga Expeditie Monog. 5 9(a): 1-186, pi.
1-5, fig. 1-52. Leiden, 1913.

- Liste des algues du Siboga: IV. Siboga

Expeditie Monog. 59(^) : 393-533, pi. 11-16,
fig. 143-213. Leiden, 1928.

Woronin, M. Beitrag zur Kenntniss der Vau-
cherien. Bot. Ztg. 27 (10): 153-160, pi. 1-2,
1869.

Cycles in Rainfall and Validity in Prediction
of Rainfall in Hawaii

Chester K. Wentworth1

In the present paper it is proposed first
to discuss a recent book titled Correlation of
Cycles in Weather, Solar Activity, Geomag¬
netic Values, and Planetary Configurations
(Johnson, 1946). This discussion is fol¬
lowed by an application of a method of
analysis therein described to the 56-year
series of rainfall data for the Honolulu in¬
take area known as the Honolulu Rainfall
Index (Board of Water Supply, 1947: 180).
The conclusion is reached that this method
does not result in prediction of the rainfall
for a future year with sufficient accuracy to
be of practical utility.

The treatment by Johnson represents an
enormous amount of labor in computing
and compiling data. It carries much food
for thought and exemplifies methods that
other investigators will find useful for ap¬
plication to specific problems. The present
discussion deals chiefly with the question of
whether the data presented show the capa¬
city of the method to predict future rainfall
quantities with useful accuracy. Some atten¬
tion is given to the general method and the
suggested correlations between various other
physical phenomena, but the present writer
does not claim competent knowledge in most
of these fields.

The method used by Johnson in the
analysis of cycles, or periodicities, is that
previously used by Dinsmore Alter and
described by him in 1937 and in several
other papers cited by Johnson. This pro¬
cedure is comparatively simple and has

1 Geologist, Honolulu Board of Water Supply.
Manuscript received March 25, 1947.

doubtless been devised and used by various
students having occasion to do such work.
It consists essentially of a process of finding
that constant interval between pairs of years
such that the average difference in rainfall
between the two members of a pair is a mini¬
mum. For any given interval, the average
error or difference between the first year’s
rainfall and the next year’s rainfall is called
the A index, presumably meaning the
"Alter index.” The interval that is found
to give the lowest A index is the period that
is assumed to give the most reliable estimate
of future values.

Johnson has carried the method much
beyond this point by subjecting the A differ¬
ences based on any one period to similar
analysis to find a second period, and so on to
several periods of diminishing importance.
He has also made some use of an index he
calls the J index, which appears to be the
mean cumulative departure, resulting from
the fact that the true cycle is a fractional
period.

It is not difficult to detect a periodic qual¬
ity in data on rainfall and other natural
phenomena. For any given span of data, the
most suitable period found in a first analysis
will inevitably result in some improvement
of estimate over that based on the arithmetic
mean of experience, as defined by the stand¬
ard deviation or probable error. Johnson
says, in his introduction, that an analysis
based on a 20-year period for the rainfall of
the Kualapuu, Molokai, area gave "very
close agreement, estimated roughly in prac¬
tical value as 88 %, if fulfilled in the future.

215

216

PACIFIC SCIENCE, Vol. 1, October, 1947

In an effort to solve the remaining 12 %, the
writer became entangled in an investigation
which led far afield.”

Moreover, any other systematic relation¬
ship found in random values for a given
quantity will, if applied, result in more
accurate estimates than the use of simple
averages without such additional terms. For
example, various stations considered by
Nakamura in his study of rainfall variation
on Oahu have a probable departure of the
rainfall of any given year from the mean of
about 30 per cent of that mean (Nakamura,
1933) . This figure is true of stations having
moderate rainfall, such as that of Kualapuu
or U. S. Weather Bureau at Honolulu. This
means that, by naming the mean annual
rainfall, anyone can predict, on the average,
within a range of 30 per cent. One means
of improving estimate is offered through a
recent study of geographic variation (Went¬
worth, 1946). In this paper it is shown
that if the rainfall of near-by, similar sta¬
tions for the same year is known, the prob¬
able error of estimate is cut down, on Oahu,
to about 14 per cent. This relationship is of
no use for future prediction, but is offered
simply as an example of refinement of esti¬
mate due to added knowledge.

Parallel to this, Johnson says his errors
were reduced to 12 per cent by use of his
data on periods. This seems reasonable,
and it is shown below that harmonic analysis
of the Honolulu Rainfall Index can furnish
a basis for somewhat similar improvement of
average estimate for a near-by, future year,
over that represented by the simple annual
mean or normal.

So far, the studies made by Johnson ap¬
pear to be significant. Beyond this point, in
the search for the remaining 12 per cent, it
is less clear either that the continued har¬
monic analysis of residuals is worth the
greatly protracted labor involved, or that
similarities of period found between rainfall
and other physical phenomena are truly the

result of cause and effect. Certainly, if
causation is at work and if the records of
such correlated sunspots, magnetic values,
or planetary positions are of long duration
or can be reliably projected into the future,
the correlation with rainfall would be of
much value.

The writer is not sufficiently acquainted
with these other phenomena to judge inde¬
pendently whether the similarities in period
do or do not represent a causal relationship.
The latter half of Johnson’s book constitutes
an interesting and exhaustive search for re¬
lationships between rainfall and other phe¬
nomena and there is abundant evidence of
cyclical variations which can be to some
extent defined. However, candor compels us
to conclude that true causal relationship has
not been proved, and it does not appear that
use of data from planetary or other changes
will yield predictions superior to those based
on one or two empirical rainfall periods
alone.

It is noteworthy that in a lecture given in
Honolulu in 1946, Dr. C. G. Rossby ex¬
pressed the view that useful predictions for
more than 1 month in advance are not now
practicable. This eminent authority had just
completed a tour of conference and inspec¬
tion in Hawaii, under the sponsorship of the
Hawaiian Sugar Planters’ Association and
the Pineapple Research Institute, and al¬
though he recommended renewed and more
extensive study of the problem, he was not
optimistic of success by available methods in
the foreseeable future.2 In line with this
statement, the paper by Johnson, while it
probes interesting possibilities, did not, in
the estimate of the present writer, contain
anything to show that prediction of the rain¬
fall of a single future year could be made

2 After this paper was completed, the writer
had opportunity to see a manuscript report by
H. Landsberg, prepared in collaboration between
the U. S. Weather Bureau and the University of
Chicago, in which a similar view was expressed.

Prediction of Rainfall in Hawaii — Wentworth

with useful accuracy. For this reason it was
decided to examine the Honolulu Rainfall
Index by a similar method for a further
check (see Table 1).

TABLE 1

The Honolulu Intake Rainfall Index^

YEAR

DECADES

IN

DECADE

1890

1900

1910

1920

1930

1940

0

132

92

122

82

108

70

1

85

101

134

108

94

79

2

91

137

85

91

117

109

3

91

110

102

124

79

91

4

100

113

100

81

94

74

5

100

107

106

81

94

64

6

66

98

134

60

109

86

7

61

123

109

154

116

8

117

88

115

83

103

9

81

95

70

87

117

* This index is now called the "intake” index, to distin¬
guish it from a "residential” index used for correlation
with domestic consumption on lawn irrigation and the like.

This index is the average of the per¬
centages of the mean for 10 stations in the
intake area of the Honolulu water supply.
It is much more representative of the rain¬
fall which provides water supply than is the
record of the official U. S. Weather Bureau
station, located in coastal Honolulu in the
low rainfall belt.

Our purpose here is to determine whether
the values of the Honolulu index, taken on
an annual basis, show systematic cycles in
such a way that the rainfall of even 1 or 2
years in advance can be predicted wifh useful
validity. No attempt has been made here
to correlate cycles of annual rainfall with
any other natural cycles. Obviously, no value
can be attached to any method of prediction
unless it follows a definite routine which
can be carried through uniformly by any two
persons who follow the adopted rule.

The first operation, similar to that used by
Johnson, and others previously, is to calcu¬
late the mean differences between annual
rainfall indexes separated by each of the
intervals considered as a possible cycle. For

217

example, the differences between the indexes
of the pairs 1890 and 1895, 1891 and 1896,
1892 and 1897, carried through to 1940 and
1945, are computed and averaged, without
regard to sign. This value represents the
average and also the most probable amount
by which the rainfall of any given year will
differ from one 5 years in the past or 5 years
in the future. The same calculations were
made for a 4-year interval, for 6 years, and
so on, up to 20 years, with results shown in
Table 2.

The 4-year difference is the mean of 52
values and that for 20 years is the mean of
3 6 values. It is evident that the means have
only a moderate range, that for a 1 6-year
interval being the lowest, 20.8 per cent.
Moreover, there is no marked superiority of
this interval over the next, and so on. Thus
the cyclical character is not strong.

In order to estimate how much of the
variation in annual rainfall is periodic, we
first need a measure for random variation.
Taking the whole series of 56 years, we find
the standard deviation to be 20.3 per cent.

TABLE 2

Mean Variations in the Honolulu Rainfall
Index for Cycles from 4 Years to 20 Years

LENGTH OF

CYCLE

MEAN

DIFFERENCE

FOR YEARS

ONE INTERVAL

apart

RANK,
LOWEST TO

HIGHEST

Interval

Per cent

4

22.4

6.0

5

21.6

3.5

6

24.3

12.0

7

22.5

7.5

8

24.0

10.5

9

21.8

5.0

10

26.5

16.0

11

24.9

13.0

12

21.6

3.5

13

24.0

10.5

14

22.5

7.5

15

25.0

14.0

16

20.8

1.0

17

25.9

15.0

18

28.2

17.0

19

23.7

9.0

20

21.2

2.0

218

From this is derived the probable error of
estimate if any given year in the future is pre¬
dicted to have a rainfall equal to the mean
of the whole period.

P. E. = 0.6745 (20.3) = 13.7 per cent

This means that the odds are equal, that
there is a fifty-fifty likelihood that the rain¬
fall in any future year will not differ from
the mean by more than 13.7 per cent. This
is a basis of prediction that is sound and
easily arrived at. Now we must inquire
whether, by use of cycles shown in the
record, we can reduce this probable error of
estimate or prediction.

Taking the 1 6-year cycle as most promis¬
ing, we say that the rainfall of any given
year is most likely to be similar to that of a
year 1 6, or 32, or 48 years earlier. For such
a group of years, the best representative
value is the average, and for each year, the
deviation from that group average is the
error that would have resulted from using
the average. Hence we take the whole series
of deviations of the rainfall of each year
from the average of the years separated from
it by multiples of 16. The root-mean-square
of this series is determined. This is the
standard deviation of the estimates based on
use of the mean of the 1 6-year cycle. For any
single series there are only 4 years, and it is
granted that the average can be much dis¬
torted by a very exceptional year. However,
by including the whole series in the root-
mean-square calculation we derive a stand¬
ard deviation comparable to the standard
deviation about a single mean. This turns
out to be 14.4 per cent. The corresponding
probable error is 9.7 per cent. This suggests
that by using the mean of the 1 6-year cycle,
rather than the mean of the whole series, the
fifty-fifty likelihood is that the estimate will
not be in error by more than 9.7 per cent.
Here we have an apparent improvement of
about one third of the expected error in¬
volved in using the over-all mean. This is

PACIFIC SCIENCE, Vol. 1, October, 1947

an improvement which can be used, even
though it is not startling.

The following table shows the standard
deviations and probable errors of variations
from the over-all mean and from the group
means for a few of the best and a few of
the worst cycles. In offering these data, the
writer makes no claim to having discovered
a cycle that will have general value or that
will be found useful for any other series
than the Honolulu Rainfall Index. For any
other series a similar procedure will indicate
whether any cycle appears to have at least
short-term prediction value.

TABLE 3

Measures of Estimate

CYCLE

STANDARD

DEVIATION

PROBABLE

ERROR

Variations from arithmetic
mean of 56 years, no

Per cent

Per cent

cycle . . .

Variations from group

20.3

13.7

mean on 16-year cycle... .
Variations from group

14.4

9.7

mean on 20-year cycle....
Variations from group

15.5

10.5

mean on 10-year cycle....
Variations from group

19.2

12.9

mean on 18-year cycle....

18.6

12.5

The above values indicate the probable
error of any given estimate based on the use
of the cycle named, assuming that the cycle
average is correct. However, this is an un¬
warranted assumption since the cycle aver¬
ages (such as the mean of the years 1890,
1906, 1922, and 1938) are at most based on
four values for the 1 6-year cycle and only
three for the 20-year cycle. We therefore
must examine the source of these averages.
For the 16 groups of years spaced by 16
years, we find that the probable errors of
the means range from the maximum of 8.6
per cent to 1.6 per cent, with a mean of 4.95
per cent. For the 20 groups of years spaced
by 20 years, these same values are 11.0 per
cent, 1.1 per cent, and 5.45 per cent.

Prediction of Rainfall in Hawaii— Wentworth

Even where the individual group shows
a smaller standard deviation, the groups are
so small that the measure is not reliable.
We have no basis for assuming any mean to
be closer than average probable errors for the
whole series. In Table 4, as an example, an
attempt is made to apply the 16- and 20-
year cycles, the most promising two cycles,
to the prediction of the index for the years
from 1939 to 1948 inclusive.

In this table, the actual index up to date
is given in column 2. The indexes predicted
by the 1 6-year cycle alone are given in col¬
umn 4, and those predicted by the 1 6-year
cycle followed by the 20-year cycle are
shown in column 6. Columns 3, 5, and 7
show errors of prediction by use of the mean
(column 3), by use of the 1 6-year cycle
(column 5), and by use of the 20-year cycle
after the 1 6-year (column 7). These show
that, on the average, there is a moderate im¬
provement of prospective accuracy of esti¬
mate by using the 1 6-year cycle, and a still
smaller improvement by using the 20-year
cycle in addition. However, anyone desiring
to use such data for practical purposes should
have his doubts aroused by noting that in two
of the predicted years, 1946 and 1947, the

219

predicted values in column 4 are quite dif¬
ferent from those in column 6 and of oppo¬
site deviation from normal. In both 1946
and 1947, there is marked change on using
the 20-year cycle. It is easy to trace this
effect, since these dates are 20 years after
the phenomenally low and high years of
1926 and 1927. The indexes for these 2
years are so aberrant that in much statistical
procedure they would be rejected. The 20-
year average is based only on two pairs —
1906 and 1926, 1907 and 1927— and the
effect of the latter year in each case is dis¬
proportionate. The figures for 1947 and
1948 are not offered as predictions but
simply as working data.

The conclusion drawn from the table and
other data presented is that by progressive
use of cycles, the prospective average errors
of estimation can be made slightly smaller
than those shown by predicting the mean.

However, the practical utility of any such
improvement is marred by the knowledge
that it is gained through the introduction of
group averages having probable errors that
we cannot reasonably assume to be less than
approximately 5 per cent. The problem
comes down to whether an estimate having

TABLE 4

Prediction by 16-Year and 20-Year Cycles

1

2

3

4

5

6

7

INDEX BY

ACTUAL

DEVIATION

INDEX BY

DEVIATION

16- AND 20-

DEVIATION

YEAR

INDEX

FROM

16- YEAR

FROM

YEAR

FROM

MEAN

CYCLE

ACTUAL

CYCLES

ACTUAL

1939

117

17

112

5

102

15

1940

70

30

83

13

65

5

1941

79

21

87

8

85

6

1942

109

9

98

11

10 4

5

1943

91

9

120

29

119

28

1944

74

26

77

3

79

5

1945

64

36

79

15

71

7

1946

86

14

108

22

85

1

1947

94

116

1948

114

119

Average deviation,

8 years .

20.2

12.8

9

220

a probable deviation of 13.7 per cent from a
mean of probable error of 1.8 per cent is
significantly improved by using an estimate
having a probable deviation of 9-7 per cent
from a mean of probable error of not less
than 5 per cent. Statistical treatment would
probably show a slight advantage with the
latter procedure in the long run, for the
simple reason that it is based on more com¬
plete use of the data. But for practical pur¬
poses any statistical, long-run gain is can¬
celled by the evident risk of an aberrant
estimate for a given year, as shown in Table
4. If numbers of simultaneous checks for
estimates were available, such aberrant esti¬
mates might be faced, but predictions of an¬
nual rainfall are checked one at a time, with
an unavoidable concentration of interest on
the one result.

With the present span of data, the
averages of rainfall cycles are so subject to
disturbance by aberrant years that any given
year’s prediction can be in error by 20 per
cent or more.

PACIFIC SCIENCE, Vol. 1, October, 1947

It appears that there is at least a psycho¬
logical difference between estimate and pre¬
diction. For every practical purpose we are
forced to the conclusion that the cycle analy¬
sis does not yield predictions of useful ac¬
curacy or reliability. To offer for a given
future year, on the present basis, a definite
prediction for public guidance in practical
matters would be an unwarranted presump¬
tion.

REFERENCES

Alter, Dinsmore. A simple form of periodo-
gram. Ann. Math. Statis. 8: 121-126, 1937.
Board of Water Supply, Honolulu, Hawaii,
Bien. Rpt. 11. 180 p. Honolulu, 1947.
Johnson, Maxwell O. Correlation of cycles
in weather, solar activity, geomagnetic values
and planetary configurations, viii + 149 p.
Phillips and Van Orden, San Francisco, 1946.
Nakamura, Winters T. A study of the varia¬
tion in annual rainfall of Oahu island, based
on the law of probabilities. U. S. Monthly
Weather Rev. 41: 354-360, 1933.
Wentworth, Chester K. Geographic varia¬
tion in annual rainfall on Oahu. 14 p., 4 fig.
Hawaii Univ. Res. Pub. 22. Honolulu, 1946.

The Little Hearts (Corculum) of the Pacific and
Indian Oceans

Paul Bartsch1

Form, coloration, and rarity place the
Little Hearts among the most attractive bi¬
valve mollusks of the Pacific and Indian
Oceans. They have been prized objects of
collectors since the early days of molluscan
history.

It was a pleasing surprise to receive for
determination recently, from Dr. Asela
Franco, 25 specimens of Little Hearts
with copious notes on others in her col¬
lection. The surprise was justified, for
during the cruise of the U. S. Bureau of Fish¬
eries Steamer "Albatross,” covering most of
the Philippine Archipelago, I took only two
specimens, on a reef at Tilig, Lubang Island.
Dr. Franco says: "Across from Cebu City
separated only by a small channel is the small
island of Mactan, where Magellan was
killed. East and south of Mactan are several
still much smaller islands. During low tide
one can walk from one to another of these
islands in some places. The heart shells are
found west, south, and east of Mactan, or in
the waters between Cebu City and Mactan
and between Mactan and the neighboring
small islands on the south and east sides.
Not only heart shells are found in these
places, but most of my shells were collected
there. It is my favorite collecting locality,
as it is near the city. All these smaller
islands mentioned belong to Cebu.

"Heart shells are collected during low
tide, not beyond about two feet of water.
They are usually found among fine broken
corals or in sandy places, the dorsal side

1 Associate, Division of Mollusks, United States
National Museum. Manuscript received February
26, 1947.

being buried a few centimeters deep. Some¬
times they are found flat down on the pos¬
terior side, perhaps because of the current
during high or low tide. Rarely one could
see them among seaweeds, and they are
never attached to any corals or stones. They
are not found in groups or bunches, and
both colored and white ones or different
types may be seen in the same region. More
heart shells are collected during the months
of May and June.”

Dr. Franco’s sending is particularly rich
in color markings, a fact which is helpful in
interpreting what some of the names be¬
stowed by the early writers embraced. Most
of the early specific names were based upon
coloration. That coloration was not a con¬
stant but a variable feature was then un¬
known, and this fact was responsible for the
list of synonyms here noted.

Dr. Franco’s collection, combined with the
42 lots in the National Museum from vari¬
ous localities, enables me to revise the genus
and bring the nomenclature up to date.

Corculum Roding

1798. Corculum Roding, Mus. Bolt., p. 188.

1811. Cardissa Megerle von Muhlfeldt, Mag. Ges.

Naturf. Fr. Berlin 5ter Jahrg., p. 52.

1870. Type designation by von Martens Cardium

cardissa L. Zool. Rec. for 1869, p. 586.

Little Hearts are members of the family
Cardiidae. They have a thin shell which is
anteriorly-posteriorly compressed, the two
valves of which when viewed anteriorly or
posteriorly present a heart-shaped outline.
The lateral edge of the shell, i.e., the middle
of the valves, curves forward or back de-

221

222

pending upon the species in question and is
so depressed that the anterior and posterior
sides almost touch within. The outer edge
may be spinose, denticulate, or smooth. The
left umbone is always curved behind the
right. The anterior surface of the shell bears
curved radiating cords which interlock at
their ventral margin. They may be smooth
or denticulated. These cords may be crossed
by lesser concentric cords or threads. The
sculpture of the posterior surface is weaker
than that of the anterior. Here, too, a heart-
shaped escutcheon is present.

KEY TO THE SPECIES OF Corculum

Margin of the shell strongly dentate.

Anterior side decidedly dished . monstrosum

Anterior side not decidedly dished.

Anterior and posterior sides about equally
convex.

Shell large (up to 66 mm. hlgh)....cardissa

Shell small (less than 25 mm. high) .

. dionaeum

Anterior side much more convex than

posterior side . obesum

Margin of the shell not dentate.

Anterior side convex . humanum

Anterior side concave.

Posterior side smoothish . levigatum

Posterior side radially ridged . aselae

Corculum monstrosum (Gmelin)

(Plate 2, Figure 3)

1782. Cardium monstrosum Chemnitz, Conch.
Cab., vol. 6, pi. 14, figs. 149-150 (non-bino¬
mial).

1791. Cardium monstrosum Gmelin, Linn. Syst.

Nat., ed. 13, vol. 1, pt. 6, p. 3253, no. 29.

1798. Corculum dolorosum Roding, Museum Bol-
tenianum, p. 189.

1798. Corculum injlatum Roding, ibid., p. 189.
1819. Cardium inversum Lamarck, Anim. s. Vert.,
vol. 6, p. 16, no. 46.

Chemnitz, while giving a good description
and figures of this species in 1782, was a
non-binomialist and does not figure in no¬
menclature. Gmelin based his name upon
Chemnitz’s statements and figures, and estab¬
lished the specific name C. monstrosum .

PACIFIC SCIENCE, Vol. 1, October, 1947

Roding, when he created the generic name
Corculum, cited both Gmelin’ s and Chem¬
nitz’s Cardium monstrosum as the basis for
his Corculum dolorosum. This, therefore, is
an absolute synonym of C. monstrosum
Gmelin. Roding also lists Corculum injla-
tum, a new name, which he bases upon
Cardium monstrosum Gmelin and Favannes
Conchyliologie, pi. 51, f. El. I am unable
to recognize anything in our collection
that corresponds completely with Favan-
nes’ drawing, which I am inclined to believe
places the escutcheon on the wrong side. To
dispose of the name Corculum injlatum
Roding, I am here designating his first cita¬
tion "Gmel. Cardium monstrosum” as type
of Corculum inflatutn Roding, which adds
this name to the list of synonyms of C. mon¬
strosum.

To this list must also be added the Car¬
dium inversum Lamarck, which was like¬
wise based on Chemnitz’s figures 149-150.

This species is differentiated from the rest
by its extremely convex, comparatively
smooth posterior side and denticulated outer
margin.

Shell soiled or yellowish white. The an¬
terior surface is convex along its ventral
margin excepting at the outer edge, which is
very strongly upturned, lending this surface
a deeply dished aspect. It is marked by
radiating cords which are separated by spaces
a little wider than the cords. The cords are
sharply spinose, the spines becoming weak¬
ened on the outer cords. The margin is
strongly dentate. In addition, the surface is
marked by fine threads paralleling the cords
and stronger, very regularly disposed incre¬
mental lines. The posterior surface is very
convex. It bears three poorly developed,
widely separated, finely nodulose, radiating
cords adjacent to the escutcheon. Outward
from these, merely indicated radiating cords
are present. Lines of growth and radiating
finer threads are scarcely indicated.

Corculum of Pacific and Indian Oceans — Bartsch

U.S.N.M. 545544 (Franco 1/). The specimen
figured measures: height, 36.7 mm.; length, 20
mm.; diameter, 35.3 mm. Dr. Franco states that
she has one measuring: height, 49 mm.; length,
47 mm.; diameter, 28 mm.

In addition to the figured specimen, the U. S.
National Museum has:

U.S.N.M. 7675, 2 specimens from the U. S.
Exploring Expedition, with no locality.

U.S.N.M. 545545 (Franco 2a), 1 specimen.

U.S.N.M. 545546 (Franco le), 1 specimen.

U.S.N.M. 545547 (Franco Id), 1 specimen.

U.S.N.M. 544829a (Hirase 2813), 2 specimens
from Osima, Osumi, Japan.

Corculum cardissa (Linnaeus)

(Plate 1, Figure 3)

1705. Cartissae, Hertjes, etc. Rumphius Am-

boinsche Rareteitkamer, pi. 42, fig. E, p. 132,

in part.

1758. Cardium cardissa L., Syst. Nat., p. 678, no.

59.

Linnaeus adopted Rumphius’ name,
placing it in his genus Cardium as Cardium
cardissa. He cites Rumphius’ plate 42, figure
E. Rumphius states that the most and best
are found on Nussalaut (now called Noesa-
laoet Island) and a few on Hitoe Island.

Rumphius, in addition to the one figured,
recognized two other forms which he de¬
scribed in his text and which have subse¬
quently received names, as will be stated in
the following pages.

The striking features of this species are its
large size and the strong, slender, almost
spinose denticles of the outer edge of the
valves. The two sides are about equally con¬
vex on the early shell. In adult shells the
outer edge curves anteriorly, leaving the
valves between the outer edge and the middle
concave, or dished. The anterior surface is
marked by rather broad^ depressed radiating
cords which are about as wide as the spaces
that separate them. These cords bear nodules
which are quite regularly distributed in ver¬
tical as well as horizontal series. They are
best developed in the early part of the shell.
In the spaces separating the radiating cords
fine threads are present, which are crossed
by closely spaced transverse threads. The

223

posterior valve bears the horn-colored, heart-
shaped escutcheon near the umbones which
is followed usually by four or more broadly
triangular ridges bearing regularly distrib¬
uted cusps on their crests. Between these
cords and the denticulated outer edge,
weakly developed and more closely spaced
cords are present. The spaces between the
cords show fine threads paralleling the cords,
while regularly, closely spaced, stronger
threads cross them.

The coloration of the anterior surface may
be soiled white, or yellowish white, or yellow
unicolor, or slightly rayed.

One specimen, U.S.N.M. 545548 (Franco’s 3e),
has a dark brown umbonal pit. The posterior
side usually shows a watered-silk effect, the pat¬
tern being arranged in both radiating and con¬
centric series.

One specimen, U.S.N.M. 545549 (Franco’s 3 a),
has a series of bright red spots arranged in radi¬
ating series.

U.S.N.M. 545550 (Franco’s 3) has the outer
edge of the basal part rose-red on both sides.

The specimen figured, U.S.N.M. 545551
(Franco’s la), measures: height, 56.2 mm.;
length, 27.8 mm.; diameter, 52.7 mm. Our largest
specimen, U.S.N.M. 7675, one of a series of
14 obtained by the U. S. Exploring Expedition
bearing the label "Pacific Islands,” measures:
height, 66.5 mm.; length, 31.2 mm.; diameter,
63.2 mm.

U.SN.M. 151464, 1 specimen from India.2
U.S.N.M. 521687, 5 specimens from Nicobar
Islands.2

U.S.N.M. 7675, 14 specimens from the Pacific
Islands (Exploring Expedition).

U.S.N.M. 75908, 1 specimen from the Indo-
Pacific.

U.S.N.M. 2544, 2 specimens (no locality).
U.S.N.M. 75596, 2 specimens from the Indo-
Pacific.

U.S.N.M. 120185, 1 specimen from the Indo-
Pacific.

U.S.N.M. 17465, 1 specimen from the East

Indies.

2 The Indian Ocean specimen measures: height,
42 mm.; length, 17.2 mm.; diameter, 38.8 mm.
The largest Nicobar specimen measures: height,
38 mm.; length, 15.7 mm.; diameter, 34.1 mm.
It is quite possible that the specimens from the
Indian Ocean may represent a smaller race that
may require a name in the future.

224

PACIFIC SCIENCE, Vol. 1, October, 1947

U.S.N.M. 32046, 1 specimen from the Indo-Pa-
cific.

US.N.M. 543532 (Franco lb), 1 specimen from
Cebu, P. I.

U.S.N.M. 543553 (Franco Ig), 1 specimen from
Cebu, P. I.

U.S.N.M. 543554 (Franco 3/), 1 specimen from
Cebu, P. I.

U.S.N.M. 543555 (Franco 1), 1 specimen from
Cebu, P. I.

U.S.N.M. 543556 (Franco 4), 1 specimen from
Cebu, P. I.

U.S.N.M. 543557 (Franco 4a), 1 specimen from
Cebu, P. I.

U.S.N.M. 543558 (Quadras Coll.), 1 specimen
from Balagnan Island, Surigao District, Min¬
danao, P. I.

U.S.N.M. 248248, 2 specimens from Tilig (reef),
Lubang Island.

U.S.N.M. 1074, 1 specimen from Loochoo Island.
U.S.N.M. 344827, (Hirase Coll. 2811), 3 speci¬
mens from Riukiu Islands, Japan.

Corculum dionaeum (Broderip and Sowerby)
(Plate 2, Figure 2)

1828. Cardium dionaeum Broderip and Sowerby,
Zool. Jour., vol. 4, p. 367.

1836. Cardium unimaculatum Broderip and Sow¬
erby, Proc. Zool. Soc. London, pp. 84, 85.
1845. Cardium dionaeum Reeve, Conch. Icon.,
pi. 21, fig. 122.

The small size of this species is its most
characteristic feature. Its range, too, is far
from that of the other forms here noted.
The presence of a color mark or marking
noted for C. unimaculatum, which appears
to be its only difference from C. dionaeum,
does not seem to warrant separating the two.
They appear from the literature to have the
same distribution. Broderip and Sowerby in
describing the species state that it was col¬
lected by Lieut. Belcher during Beechey’s
voyage on some island in the south Pacific.
Reeve cites Anaa as its habitat. Broderip
and Sowerby also cite Anaa as the place in
which Cuming collected unimaculatum.

Shell small, usually white, sometimes with
red about the escutcheon or various other
marking. The anterior surface is marked
with radiatingly curved, rather heavy cords
which become broader and flatter toward
the outer margin. They are nodulose, the

nodules being gradually reduced in strength
as the cords widen. Microscopic radiating
lines are present on the cords and in the
spaces that separate them. The entire sur¬
face is also marked by closely spaced, wavy
incremental lines. The posterior side bears
the rather long escutcheon below which are
three nodulose cords and beyond this is a
series of low, flattened, broad, radiating
ridges separated by mere impressed lines.
These ridges are of about the same strength
as those in the equivalent part of the anterior
surface. Here, too, the fine radial and incre¬
mental sculpture is present.

The specimen figured, U.S.N.M. 75955,
was collected by Pease in the Paumotu
Islands. It measures: height, 21.8 mm.;
length, 9-5 mm.; diameter, 18.2 mm.

In addition I have seen the following speci¬
mens:

U.S.N.M. 128480, 5 specimens from Anaa

Island.

U.S.N.M. 76814, 4 specimens from Paumotu

Islands.

U.S.N.M. 76120, 2 specimens from Paumotu

Islands.

U.S.N.M. 42325, 4 specimens from Paumotu

Islands.

U.S.N.M. 363437, 1 specimen from Mangaia

Island, Cook Islands.

U.S.N.M. 423441, 1 specimen from Lifu.
U.S.N.M. 32046a, 2 specimens from the Indo-
Pacific.

U.S.N.M. 17468, 1 specimen from the Pacific
islands.

Corculum obesum, new species
(Plate 1, Figure 2)

The distinctive characters of this species
are the extreme obesity of the anterior sur¬
face combined with the concave ventral side,
the margin being denticulated as in C.

cardissa.

This species appears to be less in size than
C. cardissa. The anterior surface is very
greatly elevated, whereas the posterior is only
slightly elevated near the escutcheon and
concave from there to the outer margin. The
anterior surface is marked by radiating nodu-

Corculum of Pacific and Indian Oceans — Bartsch

lose spiral cords separated by spaces as wide
as the cords bearing fine threads. Slender,
closely spaced, incremental threads cross the
radiating sculpture. The posterior surface
has the heart-shaped escutcheon which is
bordered by three low, broad, strongly nodu¬
lose ridges which extend over the elevated
part of the shell. The concave part is marked
by low flat cords separated by slight depres¬
sions, which grow wider from within out¬
ward. Wavy incremental lines render the
cords feebly nodulose. The outer margin of
the shell is strongly denticulated. The color
of the specimens before me is white anteri¬
orly, with a water-silk effect posteriorly.

The type, U.S.N.M . 543559 (Franco 1 h), comes
from Cebu. It measures: height, 36.1 mm.; length,
18 mm.; diameter, 29.9 mm.

U.S.N.M. 152449 contains 2 specimens from
Yokohama, Japan. US.N.M. 127623 contains 1
from Okinawa Island. U.S.N.M. 74469 contains
a young specimen referable here, obtained by the
U. 8. Exploring Expedition; it bears the locality
"East Indies.” Another young specimen, U.S.N.M.
128487, has no locality data.

Corculum humanum Roding
(Plate 2, Figure 4)

1782. Car din m humanum Chemnitz, Conch. Cab.,
vol. 6, pp. 153-154, pi. 14, figs. 145-146 (non¬
binomial).

1782. Car dm m roseum Chemnitz, Conch. Cab.,
vol. 6, pp. 154-155, pi. 14, figs. 147-148.

1798. Corculum humanum Roding, Museum Bob
tenianum, p. 189.

1819. Cardium junoniae Lamarck, Anim. s. Vert.,
vol. 6, p. 17.

Roding based his name upon Cardium
humanum Chemnitz. Chemnitz, being a non-
binomialist, has no status in nomenclature;
the species therefore dates from Roding.
Chemnitz states that his specimen came from
the Nicobar Islands. He also says that it is
present in the Greater and Lesser Moluccas.

Lamarck included C. humanum in his
Cardium junoniae, in which he also placed
Cardium roseum Chemnitz; I agree with

225

him. His C. junoniae therefore is a pure
and simple synonym of C. humanum.

In this species the shells attain a large
size. The margin is not spinose or denticu¬
late. The anterior side is convex and the
posterior is dished with the margin bent in¬
ward (not outward as in C. aselae) .

Shell large, white, yellowish, unicolor or
variously spotted or streaked with bright red
or suffused or washed with paler shades of
red, yellow, or orange. The anterior side is
well elevated and marked by strong, rather
broad nodulose radiating cords between and
upon which moderately strong parallel
threads are present. The strong cords widen
and weaken gradually edgeward. The incre¬
mental sculpture consists of very regular,
somewhat flattened threads which are sepa¬
rated by spaces almost as wide as the threads.
The posterior side is deeply concavely dished
for the outer two thirds of its surface. The
convex area adjoining the escutcheon bears
4 spinose cords; between these and the outer
edge low flattened cords are present, which
gradually grow wider from within outward.
Fine threads and incremental lines reticulate
the entire surface.

The specimen figured, U.S.N.M. 543560
(Franco 2), measures: height, 43 mm.; length, 18
mm.; diameter, 38.9 mm.

In addition, I have seen the following speci¬
mens referable here:

U.S.N.M. 168710, 2 specimens from the Indian
Ocean.

U.S.N.M. 488017, 1 specimen from the U. S. Ex¬
ploring Expedition with Pacific islands as
locality.

U.S.N.M. 17466, 1 specimen from the East Indies.
U.S.N.M, 168709, 2 specimens from the China
Seas.

U.S.N.M. 543561 (Franco 3 g), 1 specimen from
Cebu.

U.S.N.M. 543562 (Franco 3 h), 1 specimen from
Cebu.

U.S.N.M. 543563 (Franco 3 /), 1 specimen from
Cebu.

U.S.N.M. 543564 (Franco 3k), 1 specimen from
Cebu.

226

PACIFIC SCIENCE, Vol. 1, October, 1947

U.S.N.M. 90301 , 1 specimen from the Philippines.
U.S.N.M. 304240 , 1 specimen from Japan.
US.N.M. 344828 (Hirase 2812), 3 specimens
from Osima, Osumi, Japan.

Corculum levigatum, new species
(Plate 1, Figure 1)

This species resembles C. monstrosum,
but differs from it in being flatter, with the
margin not denticulate. The anterior side is
moderately elevated at the ventral edge. The
outer margin is decidedly upturned, a shape
which lends the side of the shell a dished
appearance. The surface is marked by strong
radiating cords, the inner of which are
weakly nodulose, while the outer are devoid
of them. These cords are separated by
spaces equaling the cords. Both the cords
and the spaces between them are marked by
fine threads that parallel them, and by very
regular transverse incremental lines which
are closely spaced and only a trifle stronger
than the threads. The posterior surface is
marked by low, broad radiating cords sepa¬
rated by shallow spaces a little narrower
than the cords. Of these cords, those near
the escutcheon show only traces of denticles.
The finer radial sculpture is almost absent,
while the incremental markings are very
regularly spaced threads separated by spaces
as wide as the threads.

The type, U.S.N.M. 343365, was obtained by
the U. S. Exploring Expedition and bears no spe¬
cific locality label. It measures: height, 47.5 mm.;
length, 20.1 mm.; diameter, 43.4 mm.

In addition to the type I have seen the follow¬
ing specimens:

U.S.N.M. 7673, 2 additional specimens from the
same source.

U.S.N.M. 17467, 1 specimen from the East
Indies.

U.S.N.M. 2544, 1 specimen obtained by the U. S.

Exploring Expedition in the East Indies.
U.S.N.M. 168709a, 1 specimen from China.
U.S.N.M. 344829 (Hirase 2813), 1 specimen from
Osima, Osumi, Japan.

Corculum aselae, new species
(Plate 2, Figure 1 )

This species has the anterior side mod¬
erately dished, but the extreme edge in
mature shells is bent posteriorly; the pos¬
terior side is also rather strongly ridged
radiately. In some features it recalls C.
levigatum, but the latter has the posterior
side uniformly dished and much smoother.

The color of this species is extremely
variable, ranging from white through yellow
and orange to rose. These shades may ap¬
pear in more or less solid tints or in inter¬
rupted or continuous rays or bands.

The anterior side is convex on the inner
half, then gradually becomes concave on the
rest of its surface. It is marked by strong
radiating ridges which are about as wide as
the spaces that separate them. The inner of
these cords are strongly nodulose, but the
nodules gradually become weaker outwardly
until they are scarcely perceptible on the outer
cords. Fine threads paralleling the ridges are
present in the spaces between the cords, while
weak incremental lines cross them. The pos¬
terior side is well elevated adjacent to the
escutcheon and here bears four low, rather
broad, weakly nodulose radiating ridges.
Outside this area the shell is less convex,
with the outer edge slightly upturned. This
surface is marked by depressed radiating
ridges separated by mere impressed lines,
and by very regular slender, closely spaced
incremental threads.

The type, U.S.N.M. 543566 (Franco 3 d),
measures: height, 44.2 mm.; length, 17.8 mm.;
diameter, 38.3 mm. (Near Cebu City, Cebu.)

In addition to the type, we have the following
specimens:

U.S.N.M. 543567 (Franco 3c).

US.N.M. 543568 (Franco 3 h).

U.S.N.M. 543569 (Franco 3m).

US.N.M. 543570 (Franco 1c).

U.S.N.M. 344829 (Hirase 2813), 1 specimen col¬
lected at Osima, Osumi, Japan.

■NR

Plate I. Corculum of the Pacific and Indian Oceans, (natural size)

\

Plate II. Corculum of the Pacific and Indian Oceans, (natural size)

The Skeletons of Recent and Fossil Gymnogyps

Harvey I. Fisher1

A study of the skulls of the cathartid
vultures (Fisher, 1944: 272-29 6) revealed
certain fundamental differences in the con¬
formation and proportions of skulls of
Recent Gymnogyps californianus and fossil
Gymnogyps from the tar pits of the Rancho
La Brea Pleistocene.' Even though some of
these differences were slight morphologi¬
cally, their presence in areas of minimal vari¬
ation such as the occipital ring of bones, the
basitemporal area, and the posterior palatal
region indicated that the Recent and fossil
condors were not identical. Further, the
magnitude of other differences and the
absence of overlap in the ranges of certain
significant measurements demonstrated that
two species probably were involved.

Before that study was made, it was be¬
lieved that the fossils from the Pleistocene
of Rancho La Brea belonged to californianus,
as did all Recent birds of this genus. There
remained, however, the species am plus,
from the Pleistocene deposits of Samwel
Cave in northern California. Further study
of am plus showed it to be conspecific with
the Rancho La Brea specimens because the
characters by which it had first been dif¬
ferentiated were shown to be identical in
specimens from the two deposits. As a
result, the name californianus was restricted
to the living form of Gymnogyps, and
am plus applied to all the Pleistocene speci¬
mens known from western North America.

It has been suggested that the two stocks
of condors represent subspecies of the same
species, i.e., chronological subspecies. There
may be a chronocline with the mean size of

1 Department of Zoology and Entomology,
University of Hawaii. Manuscript received
February 13, 1947.

the individuals of the population decreasing
from Pleistocene to Recent times. If this be
true for this species, as has been found for
other warm-blooded species, the fact might
illustrate Bergmann’s rule in a temporal
rather than a geographic way. Although this
present analysis of the skeleton tends to bear
out the belief that two subspecies are in¬
volved, it is still necessary to consider the
major specific differences in the skulls, and
to remember that the skeletal elements
studied in this investigation are in general
more plastic and more subject to the influ¬
ence of external environmental conditions
than are the bones of the skull. Thus, one
would expect to find in these elements a
greater similarity between the two series of
specimens.

At present the only criteria by which
these two species can be segregated are skull
characters. Except in the oil deposits of
southern California, fossil deposits seldom
contain complete skulls of birds. Hence, it
seemed desirable to obtain some means of
separating the two species on the basis of
other bony elements. For that reason, pri¬
marily, this study was undertaken. Second¬
arily, it seemed pertinent to test the often-
repeated assumption that "Pleistocene forms
are more variable than Recent forms.” It
has been noted previously that paleontolog¬
ical series of a species often do show consid¬
erable variability. This variability may be
explained in two ways : ( 1 ) the fossil species
actually is more variable; (2) the series is
inadequate and heterogeneous because of age
and sex factors that cannot be determined
and because the series often comes from
various localities and from deposits of dif¬
ferent ages.

227

228

PACIFIC SCIENCE, Vol. 1, October, 1947

The abundant series of G. am plus from
the Rancho La Brea provided ample speci¬
mens for an adequate series from one local¬
ity of known age; no sub-Recent specimens
were included. The Pleistocene tar pits may
have operated over a period of several thou¬
sands of years, but of all the known avian-
bearing horizons these are perhaps the most
accurately restricted chronologically. The age
factor was minimized by selecting well-
ossified and completely fused bones for both
the Recent and fossil series. Sexual varia¬
tion is of minor importance in these condors
because it was found ( Fisher, 1946: 547)
that the sexes of the Recent species do not
vary significantly in their skeletal measure¬
ments; hence, in the present study the sexes
were lumped. With these factors causing
variability eliminated, any variation in the
am plus series could result only from a cer¬
tain inherent variability in the homogeneous
stock, or possibly from inclusion of hetero¬
geneous stocks trapped in the oil at various
times during the Pleistocene. To measure
this variability the coefficient of variation
was calculated for each skeletal element
studied.

Measurements and ratios of the several
bones are presented in the tables. In these
can be found some 3,000 measurements that
were made with dial calipers. The range, the
mean and its error, the standard deviation
(o-) and its error, and the coefficient of
variation (V) are given in Tables 1 to 14.
Slight discrepancies are present in the third
place of some of the figures because many
of the computations were of necessity made
with only a slide rule. The magnitude of
the standard deviations was used to check
the adequacy of the sample when compared
to the observed range and mean. This check
is based upon the fact that an observed
range equal to 6a approximates the real
range and that an observed range of 4a prob¬
ably includes about 95 per cent of the real

range, whereas an observed range of 2a is
about equal to 68 per cent of the actual
range. The means were compared in various
ways. For example, the formula d/a was
used in comparing the means of large
samples, and the familiar "t” test was used
in comparing smaller samples. In a few
instances both tests were applied to a single
element. The value a d may be obtained in
two ways (Simpson and Roe, 1939: 192-
193) . Formula 1 :

is designed to test the hypothesis that both
the means and the variances are likely to be
the same in the populations from which the
two samples are taken. Formula 2:

is used to determine the possibility that the
two series differ significantly in the mean of
a variate, and it assumes that the sample
estimates of variance are near the true popu¬
lation values. Needless to say, a significant
test from either formula indicates that the
populations are likely to be different. Be¬
cause some workers prefer the first formula
and others the second and because it is of
considerable interest to note the varying
results when the two formulae are applied
to the same data, both formulae were used
in this study, and the results of all the sig¬
nificance tests are given in Tables 15 and 16.

In addition to this statistical analysis, a
careful qualitative comparison was made of
each element available for the two species.
This later comparison, however, proved
valueless, for no skeletal elements other
than those of the skull could be identified
on the basis of qualitative characters alone.
Thus the present paper is essentially a study
of absolute and relative measurements, their
variance, and their importance in identifying

Skeletons of Recent and Fossil Gymnogyps — Fisher

229

the species under study. However, the data
presented may also be of use in future studies
of the tempo and mode of evolution as
defined and studied by Simpson (1944).

Acknowledgments: I wish to express my
gratitude to Dr. George G. Simpson for
many helpful suggestions, and to acknowl¬
edge the use of facilities at the Los Angeles
County Museum and the Museum of Verte¬
brate Zoology, Berkeley, California, where
much of the work was done.

DISCUSSION OF MEASUREMENTS

In Table 1 the measurements of the wing
demonstrate that the mean wing length of
am plus is 4 per cent greater than in calif orn-
ianus ; the maximum length in am plus is 8
per cent greater, but the minimum lengths
are about the same. It should be noted,
however, that the entire range of wing
lengths in calif ornianus is within the limits
of the range of am plus. Consequently, the
species could not be separated on the basis
of wing length, even if associated skeletons

TABLE I

Measurements of the Wing (mm.)*

NO.

spec¬

imens

MEAN

MAX.

MIN.

Humerus . . . .

S 11

267

27 4

262

( 60

276

292

260

Ulna .......... ............

I 8

313

320

305

1 31

322

345

304

Metacarpus ............

f 3

132

133

131

(118

139

148

129

Digit II, phalanx 1

\ «

53.4

54.0

52.1

l 74

54.8

60.5

51.3

Digit II, phalanx 2

1 i

43.9

44.8

43.4

l 26

50.2

52.7

47.0

Total length . .

f

809

826

794

l

842

899

791

* First row ^of figures under each category pertains to
calif ornianus, the second row to amplus.

of the fossil were available. A statistical
study of the intramembral proportions of the
wing shows no significant differences be¬
tween the two species.

In no raw measurement (Table 2) of in¬
dividual humeri do the species differ; in the
series the fossil form is always largest, but
all the ranges overlap considerably. Signifi-

TABLE 2

Measurements (mm.) and Proportions of the Humerus*

Total length .

Greatest proximal width

Greatest distal width .

Length bicipital crest .

Length deltoid crest .

NO.

SPEC¬

IMENS

RANGE

MEAN

STANDARD

DEVIATION

COEFFICIENT

VARIATION

1 11

262-274

267 ± 1.2

3.93 ± .84

1.47± .31

( 60

260-292

27 6± .95

7.32 ± .67

2.65 ± .24

I 13

48.2-53.3

50.3± .42

1.51± .30

3.00± .59

1 52

50.2-57.5

53.7 ± .25

1.82 ± .18

3.39± .33

f 11

45.0-49.0

47.0

( 59

46.4-54.6

49.8

f 13

47.1-52.2

49.2

( 50

48.0-55.9

51.6

{ 12

97-119

105± 1.87

6.47 ± 1.32

6.16± 1.32

l 59

113-129

120 ± .57

4.4l± .41

3.67 ± .34

proportions of humerus

Proximal width : length . .

11

18.2-19.6

18. 6± .13

.44 ± .09

2.33± .50

1 52

18.1-20.6

19.5± .03

.20± .02

1.03± .10

Distal width : Ienpth

I 11

16.7-18.6

17.6

( 59

17.3-19.1

17.8

Bicipital length : length

1 11

1 7.9-19.0

18.3

( 50

17.6-20.1

18.8

Deltoid length : length .

11

36.5-42.7

39.2 ± .64

2.12± .45

5.4 ±1.2

l 59

40.4-46.8

43. 6± .16

1.2 ± .11

2.75± .25

* Hrst row of figures under each category pertains to calif ornianus, the second row to amplus.

230 r'

cance tests on humeral length, proximal
width, and length of deltoid crest (Table
15) demonstrate that the means of the two
series are significantly different. The ranges
of all intramembral ratios calculated for the
humerus overlap, but the significance tests
show that the means are statistically differ¬
ent. The bone is in all dimensions heavier
in the fossil. With the exception of the co¬
efficient of variation for length of the deltoid
crest, all these coefficients are greater in
am plus than in calif ornianus. The relatively
greater V for this crest in the Recent species
may be an indication that the length of the
crest is now undergoing a change. Through¬
out Table 2 the standard deviations for
measurements made on calif ornianus indi¬
cate that the sample probably was sufficiently
large to include 80 to 95 per cent of the
actual range. On this same basis the sample
of am plus was distributed over the range
that would be expected for 90 per cent of
the whole population.

Table 3 shows that absolute measurements
of the ulna for the two series overlap in
every instance. Significance tests on ulnar
length indicate that there is 1 chance in 20
that the two series could have come from
the same population; further, the means are
significantly different. Tests (Table 15) on

PACIFIC SCIENCE, Vol. 1, October, 1947
TABLE 4

Length of the Metacarpus (mm.)*

Number of specimens . j 3

I 118

Range . \ 131-133

I 129-148

Mean . f 132±.6l

\ 139±.37

Standard deviation . { 1.05±.43

l 4.06±.26

Coefficient of variation . . . { -80

\ 2.92±.19

* First row of figures under each category pertains to
calif ornianus, the second row to amplus.

distal and proximal widths of the ulna re¬
vealed that means of the two series are statis¬
tically distinct. Raw ratios of a single ulna
cannot be used to identify the species, but
again the means of ratios for the two series
are significantly different. As regards the
ulna, the coefficient of variation in amplus
is larger than in calif ornianus in one instance,
smaller in three, and about equal in another.
Although the sample for the Recent condor
was small, it probably includes about 80 per
cent of the expected range of variation of
the entire population.

The available number of metacarpal ele¬
ments of the modern condor was too small
for reliable treatment, but the metacarpus
seems to be significantly longer in the fossil
(Tables 5 and 15).

TABLE 3

Measurements (mm.) and Proportions of the Ulna*

NO.

SPEC¬

RANGE

MEAN

STANDARD

COEFFICIENT

IMENS

DEVIATION

VARIATION

Length .

( 8

305-320

313±1.75

4.96±1.24

1.58± .40

l 31

304-345

322±2.2

12.2 ±1.55

3.79 ± .48

Proximal width .

f 8

28.9-32.7

31. 1± .46

1.30± .33

4.17± 1.05

\ 25

32.2-36.7

34.3 ± .28

1.4l± .20

4.11± .58

Distal width .

S 8

16.1-25.0

20.6±:1.39

3.92 ± .98

19.0 ±4.75

\ 28

23.6-27.9

25.8± .10

.54± .07

2.09 ± .28

PROPORTIONS OF ULNA

Proximal width : length .

{ .

9.3-10.3

9.94 ±

.11

.31± .08

3.12 ± .78

\ .

9.9-11.0

10.5 ±

.04

.2 1± .03

2.00± .28

Distal width : length .

f .

5. 2-7.9

6.58±

.41

1.16± .29

17.7 ±4.42

i .

7.4-8. 6

7.95 ±

.05

.28± .04

3.52 ± .47

* First row of figures under each category pertains to calif ornianus, the second row to amplus.

Skeletons of Recent and Fossil Gymnogyps — Fisher

231

TABLE 5

Length of the Phalanges of the Wing (mm.)*

NO.

SPEC¬

IMENS

RANGE

MEAN

STANDARD

DEVIATION

COEFFICIENT

VARIATION

Digit I .

f 5

38.7-40.5

39.7 ± .30

.66 ± .21

1.66 it .52

1 17

39.1-42.6

4l.0± .26

1.06 ± .18

2.59— .44

Digit II, phalanx 1 . .

\ 6

52.1-54.0

53.4± .23

.56± .16

1.05± .30

l 74

51.3-60.5

54.8± .24

2.03— .17

3.70 it .30

phalanx 2 . . . .

{ 4

43.4-44.8

43.9— .31

.62 ± .22

1.40 ± .50

l 26

47.0-52.9

50. 2± .32

1.64± .23

3.27 it .45

Digit III . . . . . . .

I 5

34.0-36.2

35. 0± .35

.79— .25

2.2 6± .71

1 17

33.3-36.7

34.8 ± .22

.90 ± .15

2.59 =t .44

* First row of figures under each category pertains to calif ornianus, the second row to amplus.

TABLE 6

Measurements (mm.) and Proportions of the Sternum*

NO.

SPEC¬

RANGE

MEAN

STANDARD

COEFFICIENT

IMENS

DEVIATION

VARIATION

Length . . . . .

f 8

148-168

158±2.20

6.26± 1.5

3 .96 it .99

l 5

160-176

168±2.10

4.79— 1.51

2.85 ± .90

Keel length . . .

| 8

126-148

139±2.50

7.20±1.8

5.18±1.30

( 9

139-153

147± 1.68

5.05±1.19

3.43 it .81

Keel height . . .

\ 8

31-36

33.5 it .47

1.34 ± .34

4.00 it 1.00

X 12

32.7-39.8

35. 6± .52

1.79— .37

5.03 ±1.03

Anterior width . .

! 7

73-80

75.5±1.08

2.85 ± .76

3.77 ±1.01

l 6

76-82

79.4 ± .85

2.07 it .60

2.6l± .75

PROPORTIONS OF THE STERNUM

Keel length : length . .

1 8

83.1-91.7

87.7± 1.10

3.10±

.78

3.53 ± .88

l 5

87.1-94.2

89.8± 1.19

2.65±

.84

2.95 ± .94

Keel height : keel length .

f 8

22.6-26.2

24.2 ± .45

1.28±

.32

5.29±1.32

l 9

23.1-26.9

24.6± .46

1.39±

.33

5.65±1.33

Anterior width : length .

{ i

45.9-49.3

45.3

47.6± .49
45.3

1.30±

.35

2.73 ± .73

* First row of figures under each category pertains to calif ornianus, the second row to amplus.

TABLE 7

Measurements of the Coracoid (mm.)*

NO.

SPEC¬

IMENS

RANGE

MEAN

STANDARD

DEVIATION

COEFFICIENT

VARIATION

Length . . . . . .

Depth shaft . . . .

Smallest width shaft . . . .

Width of hear!

( 22
l 99

1 24
\ 100

\ 24

X 99

\ 15

X 100

101-113

111-128

9.3-13.9

10.9-15.8

13.4-17.4

14.8- 18.9

19.9- 21.6
20.0-23.5

109 ± .63
117± .36

12. 1± .24
13.2 ± .09
16.1

16.7

20.7

21.9

2.96± .45
3.58± .25
1.19± .17
.94 ± .07

2.72 ± .41
3.06 ± .22
9.83 ±1.42
7.12± .50

* First row of figures under each category pertains to calif ornianus, the second row to amplus.

232

PACIFIC SCIENCE, Vol. 1, October, 1947

Digit I of the wing is riot separable in the
two series; the ranges overlap, but the means
of total length are statistically different.
Length of phalanx 1 of digit II may or may
not be a good criterion to separate the spe¬
cies, for tests (Table 15) show that the
means and variances could have come from
a single stock, although the tests for differ¬
ences in the means only show them to be
significantly different. Phalanx 2 of digit II
shows a highly significant difference in
mean length between the two series. The
length of digit III is in no way significantly
different in the series under study (Tables
5 and 15). In all measurements of length
of digits the coefficient of variation is greater
in am plus, but this fact results at least in part
from the few specimens of calif ornianus that
were available. In the latter series the range
of the sample is perhaps equivalent to about
70 per cent of the actual range, but in
am plus the sample is representative of more
than 90 per cent of the whole population.

Relatively few specimens of the sternum
were available for study (Table 6), but
statistically the series of calif ornianus may be
representative of about 85 per cent of the
actual stock. The series for am plus exhibits
about 85 to 95 per cent of the probable
range of variation of the total population.
In every measurement of the sternum the
ranges overlap. Thus, raw measurements
are of no value in distinguishing the two spe-

TABLE 8

Spread of the Furculum (mm.)

calij ornianus ampins

Number of specimens . . 8 10

Range . 81-102 92-114

Mean . 96.0 ±2.5 100±2.3

Standard deviation . . . 7.1 ±1.8 7.3 ±1.6

Coefficient of variation . 7.5± 1.9 7.3±1.6

cies, although the fossil is larger in each
instance. Significance tests (Table 16) on
the means of sternal measurements show sig¬
nificant differences. Ratios showing intra-
membral proportions of the sternum do not
vary significantly between the two species
(Tables 6 and 16) .

To judge from the extent of the standard
deviation, the samples of the coracoids avail¬
able for the two series are adequate to repre¬
sent about 95 per cent of the total popula¬
tion. In no raw measurement can individual
coracoids of the two forms be distinguished.
In the fossil the coracoid is largest in every
dimension, but comparable ranges overlap.
Results of the significance tests show that
the two series differ significantly in their
means. The coefficient of variation for cora-
coidal length is greater in am plus, but the V
for depth of shaft is larger in calif ornianus.

Although the mean spread of the furcu¬
lum is greatest (Table 7) in am plus, the
means do not differ significantly (Tables 15
and 16) .

The series of scapulae for californianus
probably are representative of 95 per cent of

TABLE 9

Measurements (mm.) and Proportions of the Scapula*

NO.

SPEC¬

IMENS

RANGE

MEAN

STANDARD

DEVIATION

COEFFICIENT

VARIATION

Length .

Width middle of blade .

{ 23
\

105-120

119-132

9.1-11.9

10.7-12.6

115 ± .78
124± 1.10

10. 7± .13

3.74± .55
3.78± .81
.60 ± .09

3.25± .48
3.05± .65
5.6l± .83

l 11

11. 3± .18

.58± .12

5.13±1.09

PROPORTION

Width : length . .

7.6-10.3

9.3

( u

8.2-10.2

9.2

* First row of figures under each category pertains to californianus, the second row to amplus.

Skeletons of Recent and Fossil Gymnogyps — Fisher

233

TABLE 10

Measurements of the Leg (mm.)*

NO.

SPEC-

MEAN

MAX.

MIN.

IMENS

Femur .

19

138

147

132

100

147

159

136

Tibiotarsus . .

J

f 5

210

213

208

l 71

229

244

212

Tarsometatarsus

r 6

114

117

113

[ 100

123

134

113

Digit III (total)....

j

100

108

102

120

98

98

Phalanx 1 . . .

J

\ 3

42.5

43.6

41.9

<

1 72

43.6

48.0

40.8

Phalanx 2 .

J

i 3

30.8

31.3

30.0

1

[ 72

34.3

37.8

30.6

Phalanx 3 .

J

i 3

26.4

26.6

26.0

1

i 72

30.2

33.7

26.3

Total length .

J

562

579

551

1

i

607

657

559

* First row of figures under each category pertains to
calif or nianus, the second row to amplus.

the true population. The series for amplus
includes about 90 per cent of the variants
in that species. The means of scapular length
and width are greater (Table 9) in the fos¬
sil, and tests showed the difference in scapu¬
lar length to be highly significant (Table
15). There is no significant difference in
the ratios of width to length of the scapula.
The coefficients of variation for both mea¬
surements are greater in calif ornianus than
in amplus.

Leg lengths as shown in Table 10 demon¬
strate a greater difference between the spe¬

cies than do the wing lengths. Mean leg
length in amplus is 8 per cent greater than in
calif ornianus; maximum length is 15 per
cent greater in the fossil form, and minimum
length is about the same in two species.
Ratios of the lengths of the various elements
to total leg length show that there is no
apparent difference in the proportions of the
leg in the two species.

Femora of californianus that were mea¬
sured are characteristic of about 85 to 95 per
cent of the total population. The series of
femora for amplus includes about 95 per
cent of the expected range of the actual pop¬
ulation (Table 11). Individual femora of
either species cannot be identified on the
basis of any femoral measurement taken in
this study. In each dimension amplus is
larger, and within the series the differences
of the means are highly significant (Table
15). All coefficients of variation ( with one
exception) are larger in the Recent condor
than in the fossil.

The tibiotarsi of the two series are in¬
separable on the basis of raw measurements
(Table 12). Total length and length of
fibular crest are greater in amplus. It is in¬
teresting to note that the distal end of the
tibiotarsus is relatively slender in amplus, as
is the distal end of the ulna. Table 15 shows
that the differences in mean length of the
tibiotarsi are highly significant. The ab¬
normally low coefficients of variation for

TABLE 11

Measurements of the Femur (mm.)*

NO.

SPEC¬

IMENS

RANGE

MEAN

STANDARD

DEVIATION

COEFFICIENT

VARIATION

Length .

I 19

132-147

138±1.00

4.35± .71

3.15± .51

[ 100

136-159

147 ± .40

3.91± .28

2.66 it .19

Proximal trans. diameter .

f 19

29.4-34.9

31.1± .33

1.43 ± .23

4.60 it .75

\ 100

31.9-38.2

34.5 ± .13

1.30 it .09

3.77 it .27

Distal trans. diameter . . .

( 20

33.0-37.2

34.3± .26

1.16± .18

3.38 it .53

I 99

34.4-40.3

37.0 it .12

1.2 lit .09

3.2 7 it .23

T niampfpf choft

f 19

14.8-17.4

15.7

■L'vaji UiaUiu ICJL jllai L - - ..........

l 99

14.6-18.4

16.2

* First row of figures under each category pertains to californianus , the second row to amplus.

234

PACIFIC SCIENCE, Vol. 1, October, 1947

TABLE 12

Measurements of the Tibiotarsus (mm.)*

NO.

SPEC¬

IMENS

RANGE

MEAN

STANDARD

DEVIATION

COEFFICIENT

VARIATION

Length .

S 5

208-213

210± .90

1.94± .61

.92 ± .30

l 71

212-244

229 ± .90

7.29— .61

3.18± .27

Proximal width .

{ ,.8

26.2-29.4

27.3

Distal width . . .

I 6

23.4-23.5

24.3

l 66

22.3-26.2

24.1

Length fibular crest . . .

f ,6

43.3-47.0

45.1

1 69

42.3-58.8

51.2

* First row of figures under each category pertains to californianus, the second row to amplus.

californianus are probably the result of the
small sample.

The range of the length of the tarsometa-
tarsus in californianus is included in the com¬
parable range in amplus (Table 13). How¬
ever, the sample of the Recent form is small;
it is likely that it represents less than 70 per
cent of the probable range of the whole pop¬
ulation. In no absolute measurement are all
individuals of the two series separable. In
every dimension the fossil bone averages
larger, and these differences are highly sig¬
nificant (Table 15). The coefficient of vari¬
ation is greater in amplus for each of the
three measurements, but this probably is a
reflection of the small series for the modern
condor.

There were so few pedal elements avail¬
able for californianus that statistical analysis
was not undertaken. The only statements
that can be made regarding the phalanges of

the foot are that each phalanx is apparently
longer in the fossil and that the coefficients
of variation in amplus are of a magnitude
similar to that found in most zoological
series of Recent materials. The greater abso¬
lute lengths of the phalanges (as exemplified
by those of digit III) in amplus were found
to be the same relative length as in califor¬
nianus when their means were compared to
mean total length of leg.

SUMMARY

In this study it has been found impossible
to distinguish the two species, californianus
and amplus, of the cathartid genus Gymno-
gyps on the basis of any qualitative charac¬
ters in skeletal elements other than those of
the skull. Further, it is not possible to seg¬
regate all individuals of the two species on
the basis of absolute size of skeletal ele¬
ments. In no measurement made in this

TABLE 13

Measurements of the Tarsometatarsus (mm.)*

NO.

SPEC¬

IMENS

RANGE

MEAN

STANDARD

DEVIATION

COEFFICIENT

VARIATION

Length .

i 6

113-117

114± .71

1.75 ± .51

1.54± .44

\ 100

113-134

123± .40

3.96± .28

3.22± .23

Diameter through cotyla .

1 6

25.5-28.0

26.6± .38

.94 ± .27

3.53±1.02

\ 86

26.5-31.7

28.5± .13

1.16± .09

4.07 ± .31

Diameter through trochlea .

{ 6

28.3-30.2

29.4± .31

.75 ± .22

2.55 ± .74

l 97

29.5-34.5

32.2± .17

1.64 ± .12

5.09— .37

* First row of figures under each category pertains to californianus, the second row to amplus.

Skeletons of Recent and Fossil Gymnogyps — Fisher

235

TABLE 14

Length of Phalanges of Foot (mm.)*

NO.

SPEC¬

IMENS

RANGE

MEAN

STANDARD

DEVIATION

COEFFICIENT

VARIATION

f 3

22.9-24.5

23.6

l 24

22.2-27.9

25. 1± .28

1.35 ± .19

5.38± .78

f 3

30.2-31.0

30.6

l 72

29.7-35.1

33.0 ± .16

1.40 ± .12

4.24± .35

phalanx 2

S 3

24.6-25.2

24.8

1 49

24.8-29.9

27.3± .19

1.35 ± .14

4.95 ± .50

Digit III, phalanx 1

f 3

41.9-43.6

42.5

1 72

40.8-48.0

43.6 ± .35

2.95± .25

6.77 ± .56

phalanx 2

S 3

30.0-31.3

30.8

1 72

30.6-37.8

34.3 ± .17

1.4l± .12

4.11± .34

phalanx 3

j 3

26.0-26.6

26.4

1 72

26.3-33.7

30.2 ± .19

1.61 ± .13

5.33± .44

Digit IV, phalanx 1

S 3

23.0-24.0

23.4

l 60

22.6-26.9

24.4 ± .13

1.00 ± .09

4.10± .37

phalanx 2

15.9

15.9

1 io

16.5-18.7

17.7± .21

.66 ± .15

3.73 ± .83

phalanx 3

( l

12.9

12.9

1 6

13.0-14.5

13.8± .21

.51± .15

3.69 ±1.07

phalanx 4

f 2

16.3-16.4

16.4

l 3

18.2-18.9

18.6

* First row of figures under each category pertains to calif ornianus, the second row to ampins.

TABLE 15

Results of Significance Tests*

DIFF. IN

MEANS

AND

VARI¬

ANCES

P

DIFF. IN

MEANS

ONLY

P

Humeral length . . .

3.95

0.0001

5.88

0.0001

Proximal width humerus .

6.27

.0001

6.96

.0001

Length deltoid crest . .

9.88

.0001

7.66

.0001

Proximal width humerus : humeral length .

10.17

.0001

6.75

.0001

Length deltoid crest : humeral length .

9.87

.0001

7.67

.0001

Ulnar length .

2.04

.05

3.20

.001

Proximal width ulna .

5.72

.0001

5.94

.0001

Distal width ulna . . .

6.79

.0001

3.73

.001

Proximal width ulna : ulnar length..... .

5.95

.0001

4.79

.0001

Distal width ulna : ulnar length .

5.75

.0001

3.32

.001

Metacarpal length . . .

3.01

.001

9.81

.0001

Digit I of wing.......... . . .

2.57

.01

3.28

.001

Digit II, phalanx 1, wing . . .

1.66

.09

4.21

.0001

Digit II, phalanx 2, wing .

7.64

.0001

14.14

.0001

Digit III of wing .

0.45

.66

0.48

.63

Coracoidal length . . .

9.76

.0001

11.03

.0001

Coracoid, depth of shaft . . .

5.04

.0001

4.29

.0001

Furcular spread .

1.12

.24

1.13

.23

Scapular length . . .

6.62

.0001

6.67

.0001

Femoral length . . . . . . . .

8.86

.0001

8.36

.0001

Proximal width of femur . . .

9.99

.0001

9.59

.0001

Distal width of femur . . . .

9.27

.0001

9.43

.0001

Tibiotarsal length . . . . . . .

5.59

.0001

14.94

.0001

Tarsometatarsal length . . . . . . .

5.48

.0001

11.04

.0001

Diameter through cotyla of tarsometatarsus .

3.78

.0001

4.73

.0001

Diameter through trochlea of tarsometatarsus .

4.07

0.0001

7.92

0.0001

* Formula 1 (p. 228) was used for the test of difference in both means and variances, and Formula 2 (p. 228) was used
to test for significant differences in the means. Specific values are given only for those probabilities of a magnitude greater
than 1 in 10,000. All others are simply listed as less than 1 in 10,000.

236

PACIFIC SCIENCE, Vol. 1, October, 1947

study is there a gap between ranges of com¬
parable measurements that would make this
segregation feasible. In all elements where
there is a difference in average size, am plus
is the larger, except for the distal widths of
the ulna and tibiotarsus. Extremely large
or small specimens can be allocated in most
instances to am plus and calijornianus, re¬
spectively.

Ratios designed to show the proportions
of the bones cannot be used as a means of
differentiation for individual bones. In gen¬
eral, the bones are sturdier in the fossil form,
but the ranges of comparable ratios in the
two series consistently overlap.

Despite this inability to separate the two
series, the significance tests made on the
means of 34 different measurements and
ratios demonstrate conclusively that we are
dealing with two distinct forms which, in
most measurements, have significantly dif-

TABLE 16

Results of Significance (t) Tests on Means

t P

Sternal length . 2.84 <.02

Keel length . 2.54 >.02

Keel height . . 2.68 <.02

Keel, anterior width . 2.55 >.02

Keel length : total length . 1.21 >.10

Keel height : keel length . 0.58 >.10

Width scapular blade . 2.58 >.02

Spread of furculum . 1.10 >.10

ferent means. Only 5 of the 34 tests failed
to indicate significant differences between
the means and the two populations repre¬
sented by them. These 5 tests were on length
of digit III of the wing, furcular spread, and
two ratios on sternal proportions. However,
the significance test on both the means and
variances shows also that length of ulna,
length of digit I of wing, and length of
phalanx 1 of digit II are not significantly
different.

Gymnogyps am plus was found to be no
more variable than G. calijornianus. The co¬
efficient of variation was calculated for 33

identical measurements and ratios in each
species. In 18 ratios and measurements the
V was greater in calijornianus than in
am plus; in 15 instances the reverse was true.
The mean V for am plus was 3.56 and that
for calijornianus was 4.30. The relatively
small samples of several elements for cali¬
jornianus in all probability reduced the co¬
efficient of variation disproportionately in
this species, although the V for two ele¬
ments was abnormally high. In one series
(phalanges of the foot) where it seemed im¬
practical to calculate the coefficient of varia¬
tion for calijornianus, all the coefficients in
the large series for am plus were similar to
those usually found in zoological series.
Because coefficients of variation of linear
measurements and of ratios are not com¬
parable, the Vs were separated. In the
Recent species the average V for linear
measurements was 4.00, compared to 3.69
for the fossil. The Vs for the ratios were
5.45 for calijornianus and 2.98 for amplus.
To make the comparison even more reliable,
only those pairs of Vs based on samples of
10 or more linear measurements were in¬
cluded in the last examination of these co¬
efficients; the average V was 4.32 in cali¬
jornianus and 3.78 in amplus. Further, in
only three of the pairs in this analysis was
the V higher in the Pleistocene species.
Therefore, for Gymnogyps at least, Pleisto¬
cene and Recent specids show no major dif¬
ferences in total variability.

REFERENCES

Fisher, Harvey I. The skulls of cathartid vul¬
tures. Condor 46: 272-296, 1944.

- Adaptations and comparative anatomy

of the locomotor apparatus of New World
vultures. Amer. Midland Nat. 35: 545-727,
1946.

Simpson, George G. Tempo and mode in evo¬
lution. xviii + 237 p. Columbia Univ. Press,
New York, 1 944.

- — , and Anna Roe. Quantitative zoology.

xvii + 414 p. McGraw-Hill, New York, 1939.

The Mechanics of the Explosive Eruption of Kilauea in 1924

R. H. Finch1

Kilauea Volcano staged explosive erup¬
tions in May, 1924, after having exhibited
molten lava almost continuously for more
than 100 years. The explosions occurred
during a time of rapid retreat of the lava
column in Halemaumau (the active vent in
Kilauea caldera), of a general sinking of the
mountain top, and of enlargement of Hale¬
maumau by engulfment from a diameter of

1,500 feet to 3,000 feet. Detailed accounts
of the explosions have been published in
several journals (Jaggar, 1924: 30-37; Jag-
gar and Finch, 1924: 353).

The explosive eruptions in 1924 did not
establish a precedent for Kilauea, as the sur¬
face ash deposits indicate eight or nine quite
recent explosive periods (Finch, 1924: 1 )
and there are at least two ash beds in the
walls of Kilauea caldera. The material in
the surface ash beds indicates that the bulk
of the ash was produced by a series of mag¬
matic explosions (Wentworth, 1938: 101) .
The last explosive eruption of Kilauea prior
to 1924 was about 1790. Though the ma¬
terial thrown out in 1790 was largely acces¬
sory, the presence of bombs had led to the
assumption that all Kilauea explosions were
magmatic.

In 1924 the lack of juvenile material was
not especially conspicuous during the first

1 Volcanologist, Hawaiian Volcano Observa¬
tory, Hawaii National Park. Published by per¬
mission of the Director, National Park Service,
Department of Interior. Manuscript received May
21, 1947.

few explosions, since a large part of the
ejected blocks as well as sand and dust were
red-hot when they left the Halemaumau rim.
The large proportion of hot rock was not
surprising, as shortly before the explosions
Halemaumau had contained a lava lake

1.500 feet in diameter and the conduit was
filled to an unknown depth with hot material
except for a surface layer of landslide ma¬
terial, some of which was hot when it fell.
In the beginning of the explosive period,
members of the Observatory wore gas masks
when making investigations, but it was soon
found that none of the usual volcanic gases
was present in an appreciable amount. A
careful examination failed to find any juven¬
ile material in the explosion debris. Some¬
what reluctantly, then, we were forced to
acknowledge that we were dealing with
phreatic explosions.

Jaggar ( 1924: 35) has presented some of
the evidence pointing to the conclusion that
the 1924 explosions were due to steam and
suggested that a talus plug might act as a
seal in promoting a build-up of pressure.
The pressure required to produce explosions
of the magnitude observed was considerable
— a 14-ton block landed on the Halemau¬
mau rim over 1,300 feet above the bottom,
and an 8 -ton one was hurled a distance of

3.500 feet. The depth from which these
blocks came and the duration of their accele¬
ration (length of the "gun barrel”) is not
known. The pressure required to make an
appreciable showing in the bottom of Hale-

237

m

238

PACIFIC SCIENCE, Vol. 1, October, 1947

maumau had to exceed the pressure of the
talus plug above the seat of the explosions.
If the explosions took place 500 feet above
sea level, that would mean that they occurred
about 1,900 feet below the bottom of Hale-
maumau. The pressure of 1,900 feet of talus
material with a specific gravity of 2.0 would
be about 1,600 pounds per square inch.

To produce the explosions there must have
been either a gradual build-up of pressure
or a rapid generation of the same. If the
pressure build-up were gradual, then an ef¬
fective seal was required. A seal made up of
talus blocks even with an appreciable per¬
centage of dust would appear to be inade¬
quate. There were no indications of fusing
or cementing of the talus plug, and because
the plug developed above the retreating lava
column, there wras no molten lava to fill in
the voids between the blocks and help de¬
velop a seal. A slow build-up of pressure
would necessarily limit the amount of water
available for any one explosion. The pres¬
sure build-up would soon prevent the
further entry of water. A rapid generation
of steam pressure, as in a "flash boiler/'
would appear to be the most plausible ex¬
planation of the explosions. Any explana¬
tion of the explosions must consider the
periods of quiet between explosion and
series of explosive bursts (Jaggar, 1924:
44), often less than a minute apart.

The rapid sinking of the lava column
from a depth below the Halemaumau rim of
about 500 feet on May 1 to a depth exceed¬
ing 1,300 feet in the middle of the explo¬
sive period, as well as the increase in diam¬
eter of the crater from 1,500 to 3,000 feet
just prior to and during the explosions, indi¬
cates a progressive rupturing of the conduit
walls. A common interval between the ex¬
plosions was 6 to 8 hours. Finch (1943:
1-3) has suggested that intermittent ruptur¬
ing of the conduit walls and the resulting in¬
troduction of water may have been influ¬

enced by surging in the retreating lava
column.

The lack of chlorides in the material ejected
indicates that the steam came from fresh
ground water. Stearns (1946: 44) has sug¬
gested that the explosions may have been
caused by ground water trapped between
dikes at relatively high levels. G. A. Macdon¬
ald of the U. S. Geological Survey has sug¬
gested (oral statement, 1943) that ground
water confined between dikes may stand as
much as 1,500 feet above sea level in the vici¬
nity of Kilauea caldera. The lag in time,
often 2 or 3 minutes, between the premoni¬
tory symptoms of an explosion as shown by
peculiar earthquakes both recorded and felt
and the appearance of the explosion cloud in
Halemaumau indicates that the seat of the
explosions may well have been but compara¬
tively little above sea level — say, 500 feet. Es¬
timates of the depth of engulfment (Jaggar,
1924: 118) make 500 feet above sea level
seem a more probable seat of the explosions
than 1,500 feet. The relation between the
amount of material engulfed and the diam¬
eter of the conduit indicated a withdrawal
to at least 300 feet below sea level. With
the greater depth, more and larger trapped
pockets of water might be expected. At or
slightly above sea level the large body of
basal ground water would be available.
However, some of the first small explosions
may have been due to water trapped about
as high as Macdonald suggests. The retreat
of the lava column and progressive engulf¬
ment indicate the possibility of a consider¬
able vertical range in the seat of the explo¬
sions.

Collapsing in Halemaumau and harmonic
tremor (Finch, 1943: 1-2) indicated that
there was a retreat of the live lava column
both before and during the explosions. A
retreat of the lava column would leave the
conduit walls and much of the material
within at a temperature of about 1700° F.

Explosive Eruption of Kilauea in 1924 — Finch

Jaggar (1924: 59) suggested that shatter¬
ing during engulfment may have allowed
ground water to gain access to red-hot intru¬
sive bodies and have caused some of the
explosions.

The withdrawal of the lava and sudden
rupturing of the conduit walls provided a
means whereby water could suddenly gain
access to the hot volcano system and set up
a "flash boiler” mechanism. Next comes the
question as to the possibility of suddenly in¬
jecting a sufficient quantity of water to pro¬
duce the explosions. The Olaa well (Dun¬
can, 1942) with a drawdown of 8 feet by
continuous pumping showed an inflow of
80 gallons a second. If a void with the same
capacity as that of the sump below the water
surface were suddenly created, the amount
of water rushing in during the first half
second, say, might well be several times that
of any half second after pumping had been
in progress for some time. Likewise the dis¬
charge from water trapped in a dike com¬
partment, when the wall separating it from
a volcano conduit is suddenly ruptured,
might greatly exceed that of the next half
second. A considerable vertical range in the
shattered conduit would speed up the inflow
of water, for the velocity of inflow varies
roughly as the square root of the depth below
the surface of the water table.

The potency of volcanic heat in producing
steam blasts, when the conditions that keep
water out of the system are upset or dis¬
turbed, is easily understandable. A cubic
foot of water at a pressure of 1 atmosphere
and a temperature of 1700° F. would yield
over 5,000 cu. ft. of steam, or with a con¬
stant volume would produce a pressure of
80,000 lb. per sq. in. It should be noted,
however, that only a small part of the heat
of the rock would be available for any one
explosion. The available heat in a "flash
boiler” system would be limited to a surface
layer about y8 inch thick.

239

With the temperature of the rock known
(about 1700° F.), by assuming some surface
area it is easy to compute the available heat
and then calculate the amount of water that
could suddenly be converted to high-pressure
steam. Suppose that an explosion were ini¬
tiated by the formation of a crack where the
hot portion of the wall separating the con¬
duit from trapped ground water was 30 ft.
thick. The surface of the hot rock in the
walls of such a crack if 5 to 10 ft. wide and
70 ft. high, together with fragmental mate¬
rial within the crack, could easily exceed
6,000 sq. ft. This surface area and a thick¬
ness of y8 inch would give a volume of 62.5
cu. ft. With a specific gravity of 2.5, such a
volume of rock would weigh about 10,000
lb. The number of available B.T.U. above
212° F. in 10,000 lb. of basalt with an ini¬
tial temperature of 1700° F., if we assume
the specific heat is 0.30, is 4,470,000. Tak¬
ing the final temperature of the boiler sys¬
tem as 900° F. — on several occasions the
dust of the explosion was observed to glow
as it cleared the crater rim — the pressure
generated could exceed 40,000 lb. per sq. in.
This is many times the minimum require¬
ment mentioned above. The temperature of
the steam as it was produced may have been
above 900° F., with a resulting greater pres¬
sure.

After the explosion pressure had dissipated
in the case outlined above, another injection
of water could produce another explosion in
the same place. Progressive shattering of the
conduit wall in either a vertical or horizon¬
tal direction could account for the series of
explosion bursts that were noted on several
occasions. The rocks of each explosion were
largely confined to one sector of the ground
around Halemaumau. All sectors were event¬
ually covered with explosion debris. This
fact would indicate that the shattering was
more or less piecemeal, first on one side of
the pit, then on another. The amount of

240

PACIFIC SCIENCE, Vol. 1, October, 1947

condensation of water vapor in the explosion
clouds was never striking. The explosions
must have been produced by relatively small
amounts of steam.

The explosions would cease with the ces¬
sation of rupturing of the conduit walls. The
cessation in 1924 may have been due to the
lava column’s becoming stationary. There is
also the possibility that the conduit walls be¬
come structurally stronger at the level of the
Ghyben-Herzberg ground-water lens, which
is assumed to be about 32 feet above sea level
in the vicinity of Kilauea.

The above calculations are not intended to
indicate actual dimensions and volumes, but
rather to show that a small rock surface and
a small volume of water would suffice to ac¬
count for any of the 1924 explosions.

REFERENCES

Duncan, George. The dug well at Olaa Mill.

Volcano Letter (Honolulu) 477: 1-2, 1942.
Finch, R. H. The surface ash deposits at Kilauea
Volcano. Volcano Letter (Honolulu) 478: 1-3,
1942.

- Lava surgings in Halemaumau and the

explosive eruptions in 1924. Volcano Letter
(Honolulu) 479: 1-3, 1943.

Jaggar, T. A. Discussion. Hawaii. Volcano
Observ. Bui. 12 (5): May, 1924.

- and Finch, R. H. The explosive eruption

of Kilauea in Hawaii, 1924. Amer. Jour. Sci.
8: 353-374, 1924.

Stearns, H. T. Geology of the Hawaiian Islands.
Hawaii Div. Hydrog. Bui. 8. 106 p. Honolulu,
1946.

Wentworth, C. K. Ash formations of the
island Hawaii. 3rd Special Report of the Ha¬
waii. Volcano Observ. 183 p., 10 pi., 16 fig.
Honolulu, 1938.

Ghost Prawns (Sob-Family Luciferinae) in Hawaii

Robert W. Hiatt1

Specimens of the ghost prawn, Lucifer
faxonii Borradaile, have been found to con¬
stitute a significant item of the diet of the
local baitfish "nehu,” Anchoviella purpurea
(Fowler). The prawn was especially abun¬
dant in nehu taken from the Ala Wai Canal
in Honolulu and less abundant in nehu
caught in a fish pond supplied with tidal
water from the West Loch of Pearl Harbor.
Plankton tows subsequently made in these
localities showed the prawn to be very abun¬
dant in the Ala Wai Canal from January
to August and to be less abundant during
this period in Pearl Harbor. Plankton tows
also disclosed the presence of very small
numbers of this species (8 specimens in
6,700 gallons of water strained in June) in
fish ponds on the leeward shore of the island
of Molokai.

These findings represent the first records
for this species in the Hawaiian Archipelago
and the first record for the sub-family Luci¬
ferinae since Bate (1888: 468) cited Hawaii
as a locality in which L. typus H. Milne-
Edwards ( = L. reynaudii Bate) was col¬
lected by members of the '’Challenger” Ex¬
pedition. The name reynaudii applied to the
Hawaiian specimens by Bate has been con¬
signed by Hansen (1919: 49), with some
reservation, to the synonomy of L. typus H.
Milne-Edwards because these specimens fall
within the known range of variation for the
latter species. Thus, it is probable that two
of the six recognized species of this ano¬
malous and widely distributed group reside
in Hawaiian waters.

1 Department of Zoology and Entomology, Uni¬
versity of Hawaii. Manuscript received February
6, 1947.

Only two other authenticated records, one
for each species mentioned above, are known
for the Central Pacific area. Edmondson
(1923: 35) collected two specimens which
he designated as L. reynaudii H. Milne-
Edwards from surface tows in the lagoon
at Fanning Island, about 1,400 miles south
of Oahu. Upon re-examining the speci¬
mens, Edmondson (1925: 5) corrected his
earlier identification, and referred them to
L. faxonii Borradaile. These specimens have
been rechecked recently and compared to the
specimens taken in the Ala Wai Canal, and
are undoubtedly faxonii. All the known
records for this species in the Central Pacific
were of specimens collected from waters
less saline than ocean water, and those for
Hawaii were obtained from specimens col¬
lected from brackish, estuarine localities.
However, records elsewhere ( Hansen, 1919:
63) indicate that this species is pelagic as
well as an inshore inhabitant. L. faxonii is
widespread in the waters about the Nether¬
lands Indies and the West Indies, and in the
middle Atlantic from latitude 33° N. to
latitude 23° S.

L. typus ( = L. reynaudii Bate) was taken
in Hawaii at an undisclosed location by mem¬
bers of the "Challenger” Expedition, and
Edmondson {doc. cit. ) collected 1 male and
10 female specimens in a surface tow at night
a few miles southwest of Wake Island, about
2,000 miles west and north of Oahu. These
specimens have been checked and the male
is undoubtedly typus, as evidenced by the
long eyestalks, the position of the posterior
margin of the large ventral protuberance on
the telson, and the character of the petasma.
However, all the females are immature;

241

242

none has spines on the sixth abdominal seg¬
ment, nor is the protuberance present on the
telson. Thus, it is impossible to determine
definitely whether they belong to L. orien¬
tals Hansen or to L. typus. Since they were
taken in the same tow with the male speci¬
men identified as typus, it is probable that
they are conspecific. The distribution of
typus in the Pacific and Indian Oceans is
confused since orientalis and typus, both
long eyestalked species, were not separated
until 1919 (Hansen, op. cit.). Thus, all
records of long eyestalked species in these
ocean areas prior to 1919 must be discarded
as without specific value. L. typus is very
abundant in the Atlantic from latitude 34°
N. to latitude 40° S., and has been found in
the Pacific and Indian Oceans near the
Philippines, the South China Sea, the Bay
of Bengal, and the Netherlands Indies, as
well as at Wake Island and the Hawaiian
Islands. This species, so far as is known, is
not found inshore or in brackish water.

The known Hawaiian species may be sep¬
arated by the following key:

A. Distance between the labrum and the in¬
sertion of the eyestalks somewhat or only
a little longer than the eyestalks with
eyes (the basal short joint of the stalks

PACIFIC SCIENCE, Vol. 1, October, 1947

included). In the male, the terminal
portion of the petasma moderately
thick, and the processus ventralis shaped

as a somewhat broad plate...... .

- . - . L. typus H. M.-Edw.

B. Distance between the labrum and the in¬
sertion of the eyestalks more than twice
as long as the eyestalks with eyes. In the
male, the terminal portion of the petasma
tapers gradually from the base to the
acute end, and the processus ventralis is
needle-like, tapering to the very acute
end.. . L. jaxonii Borrad.

REFERENCES

Bate, C. Spence. Report on the Crustacea Ma-
crura collected by H. M. S. "Challenger” during
the years 1873-76. Challenger Rpt, Zool., vol.
24. xc + 942 p., 150 pi., 76 fig. London, 1888.

Edmondson, C. H. Crustacea from Palmyra and
Fanning Islands. Bernice P. Bishop Mus. Bui.
5. 43 p., 2 pi., 3 fig. Honolulu, 1923.

- , W. K. Fisher, H. L. Clark, A. L. Tread¬
well, and J. A. Cushman. Marine zoology of
tropical Central Pacific. Bernice P. Bishop Mus.
Bui. 27. ii + 148 p., 11 pi., 11 fig. Honolulu,
1925.

Hansen, H. J. The Sergestidae of the " Siboga ”
Expedition. Uitkomsten op Zoologisch, Bota-
nisch, Oceanographisch en Geologisch Gebied,
Monog. 38. 65 p., 5 pi., 14 fig. Leiden, 1919.

A Manilkara Found on Oahu, Hawaii

Marie C. Neal1

Among his studies of the Sapotaceae, Dr,
H. J. Lam has made at least two references
to a species of Manilkara found on Oahu,
Hawaii. In one reference ( Buitenzorg ]ard.
Bot. Bui, Ser. Ill, 7: 241, 1925) he pub¬
lished the description of it as a new species,
M. emarginata H. J. Lam, without illustra¬
tion. In another reference ( Blumea , 4: 342,
1941) he repeated the description, includ¬
ing it with descriptions of 14 other species,
and listing it as one of three incompletely
known, as flowers were lacking. In the first
reference, Dr. Lam states: ”1 discovered in
the collections of the Buitenzorg Herbarium
a specimen from Oahu, Sandwich Islands,
which seems to belong to this genus \Manil-
kara'] and seems worthy to be described
here.” The type specimen was in fruit, and
it was collected by H. M. Curran, a forester
of Manila, in April, 1911. But contrary to
Dr. Lam’s belief, this specimen evidently
was not collected from a native. tree of Oahu,
but was probably taken from an introduced,
cultivated tree in a public garden on Oahu
that is noted for its varied collection of in¬
troduced trees. No native species of Manil¬
kara is known in Hawaii.

Recently I rediscovered this species, and
very likely the same tree, in Foster Gardens,
Honolulu, Oahu, on February 11, 1947,
when it was bearing both flowers and fruit.
It was called to my attention by Mr. Gordon
Pearsall, who wished to have it identified.
It is a single tree, seemingly the original in¬
troduction and only specimen in Hawaii. It
is about 8 meters high, with a spread of 15
meters at the crown; the diameter of the
trunk is about 50 cm. near the base, where
a few large branches rise at angles of about

1 Botanist, Bernice P. Bishop Museum, Hono¬
lulu. Manuscript received April 9, 1947.

45°. The bark of trunk and lower branches
is gray and rough, and is deeply and regu¬
larly furrowed. The upper branches are
nearly smooth and bear numerous branch-
lets or twigs. As each twig tip bears a clus¬
ter of leaves about 5 to 1 5 cm. long, the leafy
canopy is dense. The leaves are shiny, dull
green above, lighter beneath, oval to obovate,
emarginate. The sap of the tree is milky. The
comparatively thick, yellowish, white-milky
pulp of the small, brown, one-seeded fruit
has a pleasing sweetish flavor.

So far as leaves and fruit are concerned,
the only parts of the tree available to Lam,
this tree fits well into his description of M.
emarginata. As many of the plants in Foster
Gardens were brought from different parts
of the world, especially from the Far East,
Malaya, and Malaysia, and records were not
always preserved, it has not been possible to
locate the home of each plant. As this
Manilkara is well grown, it may have been
collected as a seed in 1865-1866 by the
botanist Dr. William Hillebrand, who at
that time collected plants in and near Hong
Kong and in Java. Or it may have been
brought by some later collector.

The genus includes about 74 species, ac¬
cording to Dr. Lam, known from Central
America, Africa, Asia, and islands of the
Pacific. M. emarginata is said by Lam to be
related, apparently, to M. kauki (L.) Dub.,
the distribution of which is southeastern
Asia, Malaysia, the Philippines, and north¬
ern Australia. But analysis of the flowers in¬
dicates a much closer relationship to M. hex-
andra (Roxb.) Dub., distributed in central
India, Ceylon, Siam, Indo-China, and
Hainan. The possibility is thus suggested
that one or more of these regions is the home
of M. emarginata. In fact, the descriptions

243

244

PACIFIC SCIENCE, Vol. 1, October, 1947

of these two species agree so nearly that I
recommend reducing M. emarginata to a
synonym of M. hexandra, although M. hex¬
andra is described as having slightly longer
petal appendages and styles, and slightly
shorter petals and fruit than M. emarginata.
The somewhat variable leaves of both species
are similar in shape and size and minute re¬
ticulations (see Blumea, 4: 332, 340, and
Fig. 5, 1941).

To supplement Dr. Lam’s original de¬
scription of M. emarginata, here reduced to
a synonym of M. hexandra, I figure a leaf,
flower, and fruit (Fig. 1). I describe the
flower, which was not seen by him, as fol¬
lows:

Commonly 3 (1 + ) flowers develop at leaf axils
on pedicels that are green and scaly and about
1 cm. long. The 6 sepals are in 2 rows, reddish
brown, deltoid, reflexed in fruit. The 3 outer
sepals are 3 mm. long, thick, smooth within, cov¬
ered outside with tan scales, and edged with fine
tomentum. The 3 inner sepals are thinner,
smaller, and narrower than the outer ones, smooth
within, covered outside and edged with fine to¬
mentum. The corolla is smooth, with tube 0.7 5 ±

mm. long. The 6 petals are thin, light brown
tinted when dry, narrow oblong, 4 mm. long, the
edges revolute, each petal accompanied by 2 ovate
appendages nearly equaling it. Stamens 6; fila¬
ments nearly 3 mm. long, widest at base and
tapering to narrow tip; anthers nearly 2 mm. long;
staminodes 6, rounded, edged commonly with 2 or
3 teeth, which may or may not have long-tailed
tips. Ovary 1 mm. high, tomentose, 10-celled;
disk smooth, nearly 1 mm. high; style 3.7 mm.
long, smooth: stigma a cleft disk, slightly wider
than the style tip.

The discovery of this tree in Foster Gar¬
dens and its identification with a species of
the Eastern Hemisphere has a further sig¬
nificance. Other specimens collected by H.
M. Curran on Oahu in April, 1911, and dis¬
tributed to some herbaria may also have come
from Foster Gardens. This may be true of
a Vitex collected by Curran and described
by Lam as V . hawaiiensis H. J. Lam ( Buiten -
zorg Jard. Bot. Bui., Ser. Ill, 3: 39-60,
1921). Later [idem, 5:175, 1922), Lam
withdrew this name and stated that he be¬
lieved the plant to be a Mexican species,
V. mollis Kunth.

Fig. 1. Manilkara hexandra. A, leaf; B, flower and bud; C, part of corolla, outside: 3 petals and
3 pairs of appendages; D, 1 petal, 2 pairs of appendages, with pistil; E, part of corolla, inside: petal,
1 pair of appendages, 1 stamen, 2 staminodes; F, pistil, from side and in cross section; G, fruit, seed
from side and front.

NOTES

Opportunities for Financing of Research in the Pacific
Under the Fulbright Act

Millions of dollars will become available in
1948 that could be used for financing scientific re¬
search in Australia, New Zealand, the Dutch East
Indies, French Oceania, and other Pacific and
Asiatic countries under the terms of the Fulbright
Act passed by the 79th Congress. These funds
derive from income to the United States govern¬
ment through the disposal of surplus property to
the nations involved. Grants to qualified applicants
are to be made by a Board of Foreign Scholarships
which, it is expected, will be in operation in
Washington, D. C., by January, 1948.

The legislation covering this program is found
in Public Law 584, 79th Congress, approved
August 1, 1946, which amends the Surplus Prop¬
erty Act of 1944 to designate the Department of
State as the disposal agency for surplus property
outside the continental United States, its terri¬
tories and possessions. Pertinent passages from
this Act follow:

"In carrying out the provisions of this section,
the Secretary of State is hereby authorized to
enter into an executive agreement or agreements
with any foreign government for the use of cur¬
rencies, or credits for currencies, of such govern¬
ment acquired as a result of such surplus property
disposals, for the purpose of providing, by the
formation of foundations or otherwise, for (A)
financing studies, research, instruction, and other
educational activities of or for American citizens
in schools and institutions of higher learning
located in such foreign country, or of the citizens
of such foreign country in American schools and
institutions of higher learning located outside the
continental United States, Hawaii, Alaska (in¬
cluding the Aleutian Islands), Puerto Rico, and
the Virgin Islands, including payment for trans¬
portation, tuition, maintenance, and other expenses
incident to scholastic activities; or (B) furnishing
transportation for citizens of such foreign coun¬
try who desire to attend American schools and
institutions of higher learning in the continental
United States, Hawaii, Alaska (including the
Aleutian Islands), Puerto Rico, and the Virgin
Islands, and whose attendance will not deprive
citizens of the United States of an opportunity to
attend such schools and institutions: Provided,
however, That no such agreement or agreements
shall provide for the use of an aggregate amount
of the currencies, or credits for currencies, of any

one country in excess of $20,000,000 or for the
expenditure of the currencies, or credits for cur¬
rencies, of any one foreign country in excess of
$1,000,000 annually at the official rate of ex¬
change for such currencies, unless otherwise
authorized by Congress, nor shall any such agree¬
ment relate to any subject other than the use and
expenditure of such currencies or credits for cur¬
rencies for the purposes herein set forth: Provided
further, That for the purpose of selecting students
and educational institutions qualified to partici¬
pate in this program, and to supervise the ex¬
change program authorized herein, the President
of the United States is hereby authorized to ap¬
point a Board of Foreign Scholarships, consisting
of ten members, who shall serve without compen¬
sation, composed of representatives of cultural,
educational, student and war veterans groups, and
including representatives of the United States
Office of Education, the United States Veterans’
Administration, State educational institutions, and
privately endowed educational institutions: And
Provided further, That in the selection of Ameri¬
can citizens for study in foreign countries under
this paragraph preference shall be given to appli¬
cants who shall have served in the military or
naval forces of the United States during World
War I or World War II, and due consideration
shall be given to applicants from all geographical
areas of the United States.”

Amounts available under the Act in Pacific coun¬
tries, according to a State Department release of
July 17, are as follows, for a 20-year period: Aus¬
tralia, $5,000,000; France, $5,000,000; Netherlands
Indies, $7,000,000; New Zealand, $2,300,000;
Siam, $4,000,000; United Kingdom, $20,000,000;
Burma, $3,000,000; Philippines, $2,000,000; and
China, $20,000,000.

Members of the Board, announced by the White
House in July, are: General Omar Bradley, Vet¬
erans Administration; John W. Studebaker, U. S.
Commissioner of Education; Francis Spaulding,
New York Commissioner of Education; Helen C.
White, University of Wisconsin; Lawrence Dug¬
gan, Institute of International Education; Ernest
Lawrence, University of California; Sarah Bland-
ing, Vassar College; Walter Johnson, University
of Chicago; Charles Johnson, Fiske University;

245

246

PACIFIC SCIENCE, Vol. 1, October, 1947

and Martin P. McGuire, Catholic University of
America.

Interesting comments upon the opportunities
under the Act appear in a personal letter written
to Senator Fulbright on March 10, 1947, by
Harold J. Coolidge, executive secretary of the
Pacific Science Board, National Research Council.
His remarks follow:

"The more I study your bill, the more I become
convinced that Public Law 584 of the 79th Con¬
gress can open a new chapter in the field of inter¬
national scientific relations.

"As you know, we are faced in this country
with large numbers of young scientists, many of
whom are keen to undertake fundamental re¬
search which can best be carried out in a foreign
country. To such students the possibility of work¬
ing on ecological problems in New Guinea or
zoogeography where Wallace did his field work
represents nothing more than a naturalist’s dream,
far from any hope of realization. In many fields
of science, fundamental research came to a stand¬
still during the war, for obvious reasons.

"I feel that there are two important kinds of
help required to enable the qualified student to
engage in field research. First, the assurance of a
friendly reception and extension of certain facili¬
ties by the government of the foreign country
concerned. This is a matter that can usually be
arranged with the assistance of the State Depart¬
ment, and should not present great difficulties for
Americans in most countries in the post-war
world. Secondly, the highly difficult problem of
finding funds to finance travel and field or labora¬
tory research in the foreign country where the
research is to be undertaken.

"The Fulbright Bill makes it possible through
the Board of Foreign Scholarships, with the assist¬
ance of the State Department, to solve this serious
problem in a way that should not only greatly
benefit the student, as well as the foreign country
involved, but should likewise assure the possibility
of great strides in the advancement of fundamen¬

Editor’s

Comment from readers has been aroused by the
section from Utinomi’s bibliography on Micro¬
nesia printed in the July issue. Favorable remarks
have been made concerning the value of these
items to scientists now becoming interested in these
new American island possessions in the Pacific.
Readers have likewise raised the question, "How
was it possible to print both Japanese and Euro¬
pean languages side by side in the journal?” . . .
Frankly, the task of reproducing all these char¬
acters on the printed page was an exacting typo¬
graphic problem, which could not have been solved

tal scientific knowledge and the training of com¬
petent men, particularly in the fields of the nat¬
ural and related social sciences.

"You may remember my discussing with you
fellowship needs in the Pacific Area. The Pacific
Science Conference which met in Washington last
June recommended that 'the continuing organiza¬
tion [Pacific Science Board] arrange for research
fellowships at varying financial grades for compe¬
tent graduate students, and for grants-in-aid to
established scholars, including local inhabitants,
in the several fields of science involved, as a part
of the mechanics of staffing research.’ . . .

"Instead of the implementation of this recom¬
mendation being a distant vision, it now looks as
if it might be made a firm reality through pro¬
posed operations under the provisions of your
Bill. It is to be sincerely hoped that the money
for use of fellowships under this Bill will not be
diverted to bricks and mortar, or, as sometimes
rumored, to the meeting of government expenses
in foreign countries not directly related to the
basic purposes of the splendid and far-reaching
program which you had in mind.

"It is also hoped that the money being spent by
industrial firms on applied science, particularly
scientific technology, will not reduce the oppor¬
tunity that awaits many hundreds of other Amer¬
ican scientists in foreign fields, and that can only
be opened to them by those who administer the
funds made available for 'studies, research, and
other educational purposes’ under the Fulbright
Bill.

"I sincerely hope that provisions will be made
to ensure the participation of well-known scien¬
tists on your Board of Foreign Scholarships.

"Once more I wish to congratulate you on the
importance to international science and educa¬
tion of Public Law 584.”

Senator Fulbright, in acknowledging this letter
on March 13, said, in part: "I am in accord with
your views about the possibilities of the bill
which I introduced.”

Comments

without the energetic aid of the printers. The
services of two composing shops were needed, one
to set up the Japanese and Chinese ideographs,
and one to set up the translations and other pas¬
sages in roman characters. A reproduction proof
of all the ideographs was taken. Then, as the
roman matter was set up, the English compositor
had to leave proper space for each of the several
hundred oriental passages to be inserted. Page
proofs of the roman passages were made, and the
ideographs were then pasted, one after another,
in the vacant spaces. Finally, full-page line cuts

NOTES

247

of each of the eighteen pages of mixed matter
were photographed and etched, and printed by
letterpress along with the rest of the issue. Verily,
a tedious and time-consuming process! ... A
request has been made from Washington that the
January paper by Macdonald, Shepard, and Cox
entitled “The Tsunami of April 1, 1946, in the
Hawaiian Islands” be reprinted in the Annual
Report of the Smithsonian Institution. This illus¬
trated “tidal wave” study has aroused wide in¬
terest, and the results are obviously of national
importance. . . . Most of the Pacific Science
papers appearing in 1946 are still available in
separate reprint form, and individual copies of
any previous article may be obtained free of charge
by writing to the Office of Publications and Pub¬
licity, University of Hawaii, Honolulu 10, Hawaii.
. . . As Pacific Science completes herewith its
first year of publication, subscriptions are being

received from many parts of the world, along with
comments indicating that the journal is fulfilling
a special need by publishing research papers deal¬
ing with the biological and physical sciences in the
Pacific Area. Charter subscribers should now re¬
new the journal for the year 1948. . . . Dr. A.
Grove Day, under whose direction as Editor-in-
Chief Pacific Science was designed, inaugurated,
and published for the past year, will return to his
own teaching and research at the University of
Hawaii after the current issue is printed. . . . The
new editorial staff of the journal will consist of
the following men: Dr. Leonard D. Tuthill of
the Department of Zoology and Entomology, as
Editor-in-Chief; Dr. O. A. Bushnell of the Depart¬
ment of Bacteriology, as Assistant Editor; and
Thomas Nickerson, University Publications Edi¬
tor, as Managing Editor.

/

Index

aalii, 19, 48
Abbott, Isabella A.:

Brackish-Water Algae from the Hawaiian
Islands, 193-214
Acacia Koa, 1 4, 18, 19, 47, 99
Acanthocephalids, immature, 77
Acridotheris tristis, 70
Acrochaetium, 196, 203
robustum, 202, 203
seriatum, 202, 203
Aedes aegypti, 77, 80
albopictus, 77

Aelurostrongylus abstrusus, 77, 78
Agaricus, 93
alaea laau, 11, 12

Aleurites moluccana, 11, 15, 47, 99
triloba, 10
Algae

brackish-water, from Hawaiian Islands,
193-214

brown, 193, 195, 199-200
fresh-water, 195, 211
green, 193, 195, 196-199
key to Hawaiian brackish-water, 195-196
marine, 193, 194
red, 193, 195, 200-211
salt-water, 193, 194
Alicata, Joseph E.

Parasites and Parasitic Diseases of Domestic
Animals in the Hawaiian Islands, 69-84
Alphitobius diaperinus, 70, 79
Ammophirus insularis, 70, 79
Amphibia (in Bibliographic a Micronesica),
143-144

Amphipoda, 70, 71, 79, 80
Analysis of oxygen and chloride in pond
waters, 108-115
anaplasmosis, 69
Anoplocephala magna, 76, 81
perfoliata, 76, 81
Anchoviella purpurea, 241
Ancylostoma caninum, 77, 78, 80
Anoplura, 79, 80
Anthropological Sciences, 55
ants, 71, 80

Aphodius livius, 72, 79
spp., 74, 83
Arcyria denudata, 94
nutans, 94

Arhythmorhynchus sp., 77, 78
arsenic

accumulation in foods, 153-154
accumulation in soils, 151-171
in Hawaiian soils, 152, 159-169
Arsenic: phosphorus ratio in soils, 151-159

Arsenic Toxicity Studies in Soil and in
Culture Solution, 151-171
Arthropoda, 70-83
Artiodactyla, 79, 80, 82, 83
Artocarpus incisus, 96, 99, 100, 102, 106
Ascaridia galli, 70, 79
As car is suum, 74, 82
As car ops strongylina, 74, 83
Atractomorpha ambigua, 70, 79
auricular myiasis, 73
Auricularia, 94, 102, 104
Auric ularia adnata, 104, 105
ampla, 101-104
auricula- judae, 93, 103
auricularis, 102
cornea, 103
mesenterica, 104, 105
ornata, 103, 104-105
pel tat a, 105

Aves (in Bibliographica Micronesica), 136-143
awa, 194

Balantidium coli, 74, 82
Baldwin, Paul H., and Fisher, Harvey I.:
Notes on the Red-billed Leiothrix in
Hawaii, 45-51
Bartsch, Paul:

The Little Hearts (Corculum) of the
Pacific and Indian Oceans, 221-226
Basidiomycetes, 95-106
Bassia, 10

Batis maritima, 194, 197, 200, 203, 212
bean, toxicity of arsenic for, 151-171
beetles, 70, 71, 72, 74, 79, 83
Bennett’s Locality for Sandalwood, 9
Bibliographica Micronesica, Chordate
sections, 129-150

bibliography, botanical, of Pacific Islands
(notice of), 189
bird malaria, 51, 70

Bishop Museum (research facilities), 119-120

biting louse, 76

Bixa Orellana, 11, 12

blackhead of turkeys, 70

bladderworm, 73, 75, 76, 77, 78

Blattella germanica, 71, 72, 79, 80

blowfly, 73, 76

Board of Agriculture and Forestry, 123
of Health, 123

of Water Supply (Honolulu), 121
Boletus Katui, 93
sanguineus, 93
Bos taurus, 72-7 4, 78-79
bot fly, 76

251

252

PACIFIC SCIENCE, Vol. 1, October, 1947

botanical bibliography of the islands of the
Pacific (notice of), 189
Botryobasidium isabellinum, 106
Bourdotia, 96, 99
Bovicola bovis, 73, 79
caprae, 76, 81
bovine coccidiosis, 72

Brackish- Water Algae from the Hawaiian Islands,
193-214

Bryopsis, 196, 198
Harveyana, 198, 199
pennata var. secunda, 198
plumosa, 198, 199
plumosa var. Harveyana, 198, 199
var. Leprieurii, 198
var. pennata, 198
var. typica, 198, 199
Bufo marinus, 70

Bunostomum phlebotomum, 72, 79
burrowing roach, 70

California Packing Corporation
(research facilities), 120
candle nut tree, 10
canine coccidiosis, 77
Cams familiaris, 75, 76, 77, 80-81, 83
Capra hire us, 76-77, 81
Cardiidae, 221
Cardissa, 221

Cardium cardissa, 221, 223
dionaeum, 224
humanum, 225
inversum, 222
junonae, 225
monstrosum, 222
roseum, 225
unimaculatum, 224
Car ex longebrachiata, 118
longifolia, 118
new species of, 116-118
vitiensis, 116-118
Carica Papaya, 100
Carpophilus dimidiatus, 70, 79
Casuarina, 47
cat, 51
flea, 78

parasites of, 77
cattle, 72, 73, 78
cattle grub, 73
Caulerpa, 196, 199
Sertularioides, 199
cecal fluke, 71
Centroceras, 196, 207

clavulatum, 195, 207, 208, 210
Ceramium, 195, 196, 207, 210
diaphanum, 208, 210
Kuetzingianum, 208
sp. (1), 208-210
sp. (2), 209-210
Ceratiomyxa fruticulosa, 94

Ceratobasidium cornigerum, 9 6
Chaetomorpha, 195, 197
aerea, 197
antennina, 197
Chanos chanos, 194

chart showing travel times of seismic waves
to Honolulu (insert), 184-185
Cheilospirura hamulosa, 70, 79, 83
chickens, parasites of, 69-71, 79-80
Chinese dove, 70

Chloride and Oxygen Analysis Kit for Pond Waters,
108-115

Chlorophyceae, 193, 195, 196
Choanotaenia infundibulum, 71, 80
Choerostrongylus pudendotectus, 74, 83
Chordate sections, Bibliographica Micronesica,
129-150

Chrysomia megacephala, 73, 76, 79, 82
rufifacies, 73, 79
Cibotium Chamissoi, 17, 49
Citrus spp., 105
Cladophora, 195, 198, 204
Clements, Harry F., and Munson, Jerome:
Arsenic Toxicity Studies in Soil and in
Culture Solution, 151-171
coccidia, 69, 70, 72, 74, 77, 78, 79, 80, 82
coccidiosis
avian, 69, 79
bovine, 72, 78-79
canine, 77, 80
rabbit, 78, 82
swine, 74, 82

Cocos nucifera, 95, 97, 99, 100, 102, 106
Coleoptera, 70, 71, 79, 80
Columba livia domes tica, 69-71, 82
Columbicola columbae, 71, 82
Comatricha Typhoides, 94
Conocephalus saltator, 70, 71, 79, 80
Convolvulus Batatas, 10
Cooperia pectinata, 72, 79
punctata, 72, 76, 79, 82 •

C opr is incertus, 71
minutus, 71

Corculum (Little Hearts) of Pacific and
Indian Oceans, 221-226
Corculum, key to species, 222
Corculum aselae, 222, 225, 226
cardissa, 222, 223, 224
dionaeum, 222, 224
dolorosum, 222
humanum, 222, 225
inf latum, 222
levigatum, 222, 226
monstrosum, 222, 226
obesum, 222, 224
Cordia "myxa,” 105
Cordyline terminalis, 7, 47
Coriolus versicolor, 104
Corticium botryosum, 106
suecicum, 99

Index

253

Cox, D. C, et air.

The Tsunami of April 1, 1946, in the
Hawaiian Islands, 21-37
Cribraria tenella, 94
crop worm, 70
Cryptonemiales, 204
Ctenocephalides fells, 77, 78, 80
Culex quinquefasciatus, 77, 80
Curcuma longa, 11, 12
Cyathodes, 10

Cyathostomum coronatum, 75, 81
Cycles in Rainfall and Validity in Prediction of
Rainfall in Hawaii, 215-220
Cylicocercus catinatus, 75, 81
gold's, 75, 81
pater at us, 75, 81
Cylicocyclus leptostomus, 75, 81
nassatus, 75, 81

Cylicodontophorus btcoronatus, 75, 81
euproctus, 75, 81
Cylicostephanus calicatus, 75, 81
longibursatus, 75, 81
minutus, 75, 81

Cylicosternus asymetricus, 75, 81
Cyperaceae, 116-118
Cyrtosperma, 93

Dactylosternum abdominale, 70, 79
Damage by Tsunami of April 1, 1946, 34
Dean, R. B., and Hawley, R. L.:

A Chloride and Oxygen Analysis Kit for
Pond Waters, 108-115
Dendrophilus punctatus, 71, 80
Dermanyssus gallinae, 71, 80
Dermaptera, 70, 80
Dermestes vulpinus, 70, 71, 72, 79, 80
desmids, 195
Dianella, 10
sandwicensis, 11
diatoms, 193, 194, 204
Dicranopteris linearis, 17, 48
Dictydium cancellatum, 94
Dictyocaulus viviparus, 72, 79
Diospyros Hillebrandii, 11
Dipylidium caninum, 77, 78, 80
Diroflaria immitis, 77, 80
diseases, parasitic, of domestic animals, 69-84
cat, 77-78
cattle, 72-75, 78-79
chicken, 69-71, 79-80
dog, 77, 80-81
goat, 76-77, 81
guinea fowl, 69-71, 81
horse, 75-7 6, 81
peafowl, 69-71, 82
pigeon, 69-71, 82
rabbit, 78, 82
sheep, 76-77, 82
swine, 74-75, 82-83
turkey, 69-71, 83

Dispharynx spiralis, 70, 71, 79
Dodonaea viscosa, 19, 48
dog, 51

parasites of, 75, 76, 77, 80, 81
Dolomitization in Semi-arid Hawaiian Soils, 38-44
domestic animals, parasites and parasitic
diseases of, 69-84
dooe dooe, 15
dourine, 69
dove, Chinese, 70

ear canker, 74
tick, 73, 76
Earth Sciences, 55
earthworms, 74, 83

Echidnophaga gallinae ea, 71, 77, 78, 80, 81
Ectocarpus, 196, 199

Duchassaingianus, 199, 200
indicus, 195, 199, 200, 201
Mitchellae, 200
Sargassi, 200
Eimeria bovis, 72, 78
bukidnonensis, 12, IS
cylindrica, 12, IS
debliecki, 74, 82
scabra, 74, 82
spinosa, 74, 82
stiedae, IS, 82
tenella, 69, 79
zurnii, 12, IS
Elaeocarpus bipdus, 11
Elatae, 118
enaena, 11

English sparrow, 70, 71
Enteromorpha, 194, 195, 196
flexuosa, 196-197
intestinalis, 197
spp., 200

Eomenacanthus stramineus, 71, 80
epiphytes, 93

Epitragus diremptus, 70, 71, 79, 80
Equus cab alius, 75-76, 81
Erythrina sandwicensis, 19
Erythrotrichia, 196, 200
carnea, 195, 200, 204, 205
Euborellia annulipes, 70, 71, 80
Eucalyptus, 47
E.ucarex, 118
Eugenia malaccensis, 15
sandwicensis, 16
Eulota similaris, 71, 77, 78, 80
Euphorbiaceae, 11
Euxestus sp., 70, 79
Exidia ampla, 101
cornea, 103
Exidiopsis cerina, 96

explosive eruption of Kilauea in 1924, 237-240
eyeworm, 70

254

facilities for research in Hawaiian Islands,

119-126

Factors in the Behavior of Ground Water in a
Ghyben-Herzberg System, 172-18 4

Fannia sp., 73, 79

Fasciola gigantica, 69, 72, 75, 76, 79, 81, 82, 83
hepatic a, 72, 76
Fault at Waimea, Oahu, 85-91
Feldmannia, 199
Fclicola subrostrata, 78
Felis dome Stic a, parasites of, 77-78
Fiji, new species of Car ex from, 116-118
filarid, skin, 72
Filariodes osleri, 77, 80
Finch, R. H.:

The Mechanics of the Explosive Eruption
of Kilauea in 1924, 237-240
Fisher, Harvey I.:

Bibliographica Micronesica, Chordate
Sections, 129-150

The Skeletons of Recent and Fossil
Gymnogyps, 227-236

Fisher, Harvey I., and Baldwin, Paul H.:
Notes on the Red-billed Leiothrix in
Hawaii, 45-51

fish-ponds (of Hawaii), 193, 194
fleas
cat, 78

sticktight, 71, 77, 78
flies, 73, 75

flukes, 69, 71, 72, 75, 76, 79, 80, 81, 82, 83

Fo?nes amhoinensis, 93

food, accumulation of arsenic in, 153-154

Fossaria ollula, 72, 79, 81, 82, 83

fresh-water algae, 195, 211

fruitfly investigations, 124

Fujimoto, Charles K., et al.\

Dolomitization in Semi-arid Hawaiian
Soils, 38-44
Fuligo septica, 94

Fulbright Act and financing research in
Pacific, 245-246
Fungi Imperfecti, 106

Fungi of the Marshall Islands, Central Pacific
Ocean, 92-107

Galera sp. ( conjertae aff.), 93
G alius gallus, 69-71, 79-80
Gardenia Remyi, 47
Gastrophilus intestinalis, 76, 81
nasalis, 76, 81

Gastropoda, 78, 79, 80, 81, 82, 83
Gelidiales, 204
Gelidium, 196, 200, 203
pusillum, 203

pusillum var. conchicola, 204
geology, ground- water, 172-184
Geology (Waimea, Oahu), 85-91
Ghost Prawns (Sub-Family Luciferinae) in Hawaii,
241-242

PACIFIC SCIENCE, Vol. 1, October, 1947

Ghyben-Herzberg system, factors affecting,
172-184
Giffordia, 199
ginger, 15
gizzard worm, 70
Glaziella aurantiaca, 95
vesiculosa, 95

Gloeotulasnella calospora, 96
Gnaphalium, 10
sandwicensium, 11
goats, parasites of, 76-77, 81
Gongylonema ingluvicula, 70, 79
pulchrum, 72, 79
Goniocotes gigas, 71, 80
hologaster, 71, 80, 83
Goniodes stylifer, 71, 80, 83
Gonocephalus seriatum, 70, 71, 79, 80
Gracilaria, 196, 206
conjervoides, 206
coronopifolia, 205, 206, 207
euchemoides, 206
No. 1, 206
No. 2, 206

Grateloupia, 196, 200, 205
dichotoma, 20 6
filicina, 205, 206
j fili c in a forma Hawaiiensis, 205
ground water, factors affecting, 172-184
guava, 47, 48
Guepinia Spathularia, 101
guinea fowl, 69-71, 81
gullet worms, 72
Gyalocephalus capitatus, 75, 81
Gymnogyps, Recent and Fossil, Skeletons of,
227-236

Gymnogyps amplus, 227-236
calif ornianus, 227-236

Habronema microstoma, 75, 81
muscae, 75, 81

Haematopinus adventicius, 75, 83
eurysternus, 73, 79
suis, 75

Haemonchus contortus, 72, 76, 79, 82

Haemoproteus columbae, 70, 82

hala tree, 10

Halophila ovalis, 194

halophytes, 194

hau, 47, 48

Hawaii National Park (research facilities), 120
Hawaiian hawk, 51
Hawaiian Islands

algae, brackish-water, of, 193-214

eruption of Kilauea in 1924, 237-240

facilities for research in, 119-126

fault at Waimea, Oahu, 85-91

ghost prawns in, 241-242

Manilkara on Oahu, 243-244

parasitic diseases of domestic animals in, 69-84

prediction of rainfall in, 215-220

Index

255

red-billed Leiothrix in, 45-51
sandalwood on Oahu, 5-20
tsunami of April 1, 1946, in, 21-37
Hawaiian Pineapple Company
(research facilities), 120
Hawaiian soils, toxic levels of arsenic in,

152, 159-169

Hawaiian Sugar Planters’ Association
(research facilities), 120

Hawaiian Tuna Packers (research facilities), 121
Hawaiian Volcano Observatory
(research facilities), 121
Hawaiian Volcano Research Association
(research facilities), 121
Hawley, R. L., and Dean, R. B.:

A Chloride and Oxygen Analysis Kit for
Pond Waters, 108-115
head maggot, 76
height of waves

factors influencing, 28-34
in open sea, 25
on Hawaiian shores, 27
Hawaii, 33
Kauai, 27
Maui, 31
Molokai, 30
Oahu, 29

Helicomyces roseus, 106
Hemitrichia serpula, 94
stipitata, 94
vesparium, 94
Heterakis gallinae, 70, 79
Heterochaetella, 99
dubia, 97

Heterodoxus longitarsus, 77, 81
Hiatt, Robert W.:

Ghost Prawns (Sub-Family Luciferinae)
in Hawaii, 241-242
Hibiscus, 105

Arnottianus, 47, 99
tiliaceus, 47
Hirneola, 102
ampla, 101

Histomonas meleagridis, 70, 83
history

of sandalwood in Hawaii, 6
of tsunamis in Hawaii, 22
History, Present Distribution, and Abundance of
Sandalwood on Oahu, Hawaiian Islands, 5-20
hoawa, 11
hog, 51

mange mite, 75
hookworms, 77
horn fly, 73

horse, parasites of, 75-7 6, 81
hydrostatics, 172-184
Hymenolopis carioca, 71
exigua, 69, 71, 80
Hyostrongylus rubidus,

Hypholoma jaluitensis, 93

Hypnea, 196, 206
armata, 207
cervicornis, 207
cornuta, 207
divaricates, 207
nidifica, 206, 207
nidulans, 205, 207
pannosa, 207

Hypochnus isabellinus, 106
Hypoderma lineata, 73, 79

iliahi, 5, 6, 9, 12, 18, 19
indigo, 18

international co-operation, recommendations
for, 54

Ipomoea batatas, 11, 12
ironwood, 47
Isopoda, 71, 79

Japanese hill robin, 45

kalia, 11
kamani tree, 47

Kanehiro, Yoshinori, et air. i
Dolomitization in Semi-arid Hawaiian
Soils, 38-44
kangaroo lice, 77
karia, 10, 11
kealia, 11, 12

key to brackish-water algae of Hawaiian
Islands, 195-1 96
ghost prawns in Hawaii, 242
little hearts (Corculum) of Pacific and
Indian Oceans, 222
Kilauea volcano, 237
koa, 14, 18, 19, 47
kopiko, 19
kukui, 11, 15, 47

laau ala, 6, 18
Lachnea jaluitensis, 93
lajiling kijilik (rat’s ear fungus), 102
lama, 10, 11
lauhala, 9, 10, 12
lau, hula hula, 12
lau, keo keo, 12
Leiothrix lutea, 45-5 1
distribution in Hawaii, 46
eggs, 50
flocking, 48
food, 51

habitat in Hawaii, 47
hatching, 50
nesting, 48
young, 50

Libby, McNeill and Libby (research facilities),
122

256

PACIFIC SCIENCE, Vol. 1, October, 1947

lice, 71, 73, 75, 76, 77, 78
biting, 76, 78
kangaroo, 77
sucking, 76
lichens, 93

limu ekahakaha, 204
huluhuluwaena, 206
loloa, 204
manauea, 206
pakeleawaa, 206
Linognathus africanus, 76, 81
Lipeuris caponis, 71, 80
gallipavonis, 71, 83
heterographus, 71, 80
Litargus bait eat us, 70, 79
Little Hearts (Corculum) of Pacific and Indian
Oceans, 221-226
liver flukes, 69, 72, 75, 76
control of, 72

loss of life from tsunami of April 1, 1946, 35
Lualualei Valley, soil of, 39, 43
Lucifer faxonii, 241-242
orientalis, 242
reynaudii, 241
typus, 241, 242

Luciferinae (ghost prawns) in Hawaii, 241-242
Lu cilia s eric at a, 73, 79
lumma, 10, 11

Lycopersicon esculentum, toxicity of arsenic
for, 151-171
Lynchia maura, 71
Lyperosia irritans, 73
Lyponyssus bursa, 71, 80

Macdonald, G. A., et air.

The Tsunami of April 1, 1946, in the
Hawaiian Islands, 21-37
Macrodrili, 83

Makaha Valley, soil of, 39, 40

Malacca apple, 15

malaria, bird, 70

mamake, 11

mamani, 48

mamati, 10

Mammalia (in Bibliographic a Micronesica) ,
133-136
mange mites, 77

Manilkara emarginata, 243, 244
hexandra, 243
kauki, 243, 244

Manilkara Found on Oahu, Hawaii, 243-244

Marasmius c allop us var. jaluitensis, 93
Pandanicola, 93

Marshall Islands, fungi of, 92-107
Mechanics of the Explosive Eruption of Kilauea in
1924, 237-240
medicine, 58

Megninia cubitalis, 71, 80
Meleagris gallopavo, 69-71, 83
Melophagus ovinus, 76, 82

Menopon gallinae, 71, 80, 81, 83
phaeostomum, 71, 81, 82
Merrill, E. D.:

Botanical Bibliography of the Islands of the
Pacific (notice of), 189
Merulius Spathularia, 101
Messerschmidia argentea, 99, 104, 105
Metasphaeria fur, 93
Jus, 93
spp., 93

Metastrongylus elongatus, 74, 83
meteorology, 57

Metrosideros collina ssp. polymorpha, 18, 19, 47
mice, 77, 78
microbenthos, 194
Micronesia

bibliography of, 129-150
Marshall Islands, fungi of, 92-107
University of Hawaii Expedition to, 60-62, 92
U.S. Commercial Company Survey of, 62
milkfish, 194
mites, 71, 75, 77, 81
body, 71, 80
fowl, 71, 80
hog mange, 75, 83
mange, 77, 81
red, 71, 80
wing, 71, 80
mongoose, 51
Morinda citri folia, 11, 12
mosquitoes, 77
Mugil cep halos, 194
mullet, 194

Munson, Jerome, and Clements, Harry F.:
Arsenic Toxicity Studies in Soil and in
Culture Solution, 151-171
Musca dome Stic a, 71, 75, 80, 81
Mus mus cuius, 77, 78
myiasis, auricular, 73
mynah bird, 70, 71
Myxophyceae, 193
Myxomycetes, 94-95

National Research Council, 52
naupaka kuahiwi, 11, 19
Neal, Marie C:

A Manilkara Found on Oahu, Hawaii, 243-244
nehu, 24 1

Nematodirus spathiger, 76, 82
Nephrolepis exalt at a, 16

New Species of Carex (Cyperaceae) from Fiji,
116-117
noni, 11, 12
Notes

Editor’s comments, 247
facilities for research in natural sciences
in the Hawaiian Islands, 119-126
Micronesian expedition of University of
Hawaii, 60-62

new botanical bibliography of Pacific
Islands, 189

Index

257

opportunities for financing research in Pacific
under the Fulbright Act, 245-246
Pacific Science Conference, recommendations
of, 52-59

scientists and the fortieth anniversary of
the University of Hawaii, 189-190
survey of Micronesia by U.S. Commercial
Company, 62

Notes on the Red-billed Leiothrix in Hawaii, 45-51
nouputa, 10, 11

Numidia meleagris, 69-71, 81

Oahu, fault at Waimea, 85-91
oceanography, 57, 185-188
Ochrosia parviflora, 106
Oesophagostomum dentatum, 74
radiatum, 72, 79
Oestrus ovis, 76, 82
ogo, 206
ohava, 10, 11
ohe, 19
ohia ha, 19
ohia lehua, 18, 19, 47
oi, 47, 48, 49

Oidium tomentosum, 106-107
olena, 11, 12
Oligosiphonia, 212
olive-green creeper, 51
olopua, 19

Orchestia platensis, 70, 71, 79, 80
oreina, 11
oreyna, 10
Oriental blowfly, 76

Ornithostrongylus quadriradiatus, 70, 82
Orthoptera, 70, 71, 79, 80
Oryctolagus cunicularis, 78, 82
Osmanthus sandwicensis, 19
Osteomeles anthyllidifolia, 10, 11
Otobius megnini, 73, 76, 79, 82
Ovis aries, 76-77, 82, 83
Oxya chinensis, 70, 79, 80
Oxydema fu si forme, 70, 79
oxygen analysis, 108-115
Oxyspirura mansoni, 70, 79
Oxyuris equi, 75, 81

Pacific Chemical and Fertilizer Company
(research facilities), 122
Pacific Foundation War Memorial, 52
Pacific Islands, botanical bibliography of
(notice), 189

Pacific Islands Research Committee
(University of Hawaii), 60
Pacific Science Conference, recommendations
of, 52-59

Pacific Science Congress, 53
Pacific Science Survey, 53
palm foxtail, 47, 48
Palmer, Harold S.:

Fault at Waimea, Oahu, 85-91

Palorus ratzeburgi, 70, 79
Pandanus, 10
pulposus, 105
sp., 100, 101, 105, 106
papaya, 51

Parascaris equorum, 75, 81
Parasites and Parasitic Diseases of Domestic
Animals in the Hawaiian Islands, 69-9 1
parasites of
cat, 77-78
cattle, 72-75, 78-79
chicken, 69-71, 79-80
dog, 77, 80-81
goat, 76-77, 81
guinea fowl, 69-71, 81
horse, 75-7 6, 81
peafowl, 69-71, 82
pigeon, 69-71, 82
rabbit, 78, 82
sheep, 76-77, 82
swine, 74-75, 82-83
turkey, 69-71, 83
Paroreomyza bairdi mana, 51
Passer domesticus, 70
Pavo cristatus, 71, 82
peafowl, 71, 82
Pekin nightingale, 45
Pellicularia isabellina, 106
lembospora, 106, 107
vaga, 106
Peniophora, 97
pallidula, 97
pubera, 99
Sambuci, 99

pepeiaoakua (ghost ear fungus), 102
Perichaena depressa, 94
Phaeophyceae, 193, 195, 196, 199
phanerogams, 194

Phaseolus vulgaris, toxicity of arsenic for,
151-171

Pheidole spp., 80
vinelandica, 71, 80
Pheretima spp., 74, 83
phosphorus: arsenic ratio, 157-159
Phy corny cetes, 95
Physaloptera praeputialis, 77, 78
Physarella oblonga, 94
Physarum tenerum, 94
viride, 94
Wingatense, 94
Phytolacca, 10
sandwicensis, 11
pigeon fly, 70, 71
pigeon, parasites of, 69-71, 82
Pineapple Research Institute of Hawaii
(research facilities), 122
pineapples, 18
Pipturus albidus, 11
piroplasmosis of cattle, 69

Pisces (in 'Bibliographic a Micronesica') , 144-150
Pithecolobium Saman, 99

258

PACIFIC SCIENCE, Vol. 1, October, 1947

Pittosporum, 12
Plant Sciences, 57
Plasmodium vaughani, 51
Pleurotus, 102
Schwabeanus, 93
poina, 10

point of sterilization in soil, by arsenic, 169
pokeawi, 10

Polyporus Kamphoeveneri, 93
Sanguineus, 93
xanthopus, 93

Polysiphonia, 193, 195, 196, 197, 200, 204, 207,
211

aquamara, 210, 212
sp., 211, 212
subtillissima, 212
Polystictus sanguineus, 93
pond waters, analysis of, 108-115
chlorine, 112-113
kit (apparatus), 109-112
oxygen, 113-114
popolo, 11
poporo-tumai, 10
Porcellio laevis, 71, 79
Postharmostomum gallinum, 71, 80
Poteriostomum imparidentatum, 75, 81
poultry ascarid, 70

poultry, parasites of, 69-71, 79, 81, 82, 83
Pratella, 93

Probstmayria vivipara, 75, 81
Protochordata (in Bibliographic a Micronesica'),
150

protozoa, 69, 72, 74, 75, 76, 77, 78, 79, 80, 82, 83
pro ventricular worms, 70, 80
Psathyra Schwabeana, 93
Psathyrella disseminata, 93
Pseudolynchia canariensis, 70, 71, 82
Psidium Cattleianum, 51
Guajava, 47, 99
Psoroptes communis, 78, 82
Pterolichus obtusus, 71, 80
Public Health, 58
pukeawe, 11, 19, 48
Pulex irritans, 80
Pycnoscelus surinamensis, 70, 79
Pyrus anthylidijoliae, 10

rabbit, parasites of, 78, 82
Raillietina cesticillus, 71, 80
tetragona, 71, 80
rainfall, prediction of, 215-220
rat, 51

Rattus rattus alexandrinus, 78
norvegicus, 78
rattus, 78

Reptilia (in Bibliographic a Micronesica ),

143-144

research in Hawaiian Islands, facilities for,
119-126

Reynoldsia sandwicensis, 19

Rhipicephalus sanguineus, 77, 81
Rhizoclonium, 195, 197
sp., 197

Rhodophyceae, 193, 195, 196, 200-211
Rhodophyllidaceae, 204
Rogers, Donald P.:

Fungi of the Marshall Islands, Central
Pacific Ocean, 92-107
roundworms, 70, 72, 74, 75, 76, 77, 78,

79, 80, 81, 82, 83
Rubus rosaefolius, 47, 51
rumen fluke, 72, 79

Sac char um officinale, effect of arsenic on, 169
St. John, Harold:

The History, Present Distribution, and Abun¬
dance of Sandalwood on Oahu, Hawaiian
Islands, 5-20

A New Species of Carex (Cyperaceae)
from Fiji, 116-117
Samanea Saman, 99
Sandalwood ( see also Santalum ), 5-20
distribution on Oahu, 13
history of, in Hawaii, 6
location of original forests, 17
oil, 5

rate of growth, 16
Santalum ( see also Sandalwood), 5
album, 5

austrocaledonicum, 6
boninense, 6
ellipticum, 6
fernandezianum, 6
Freycinetianum, 6-19
haleakalae, 6
hendersonense, 6
insulare, 6
lanaiense, 6
lanceolatum, 5
Macgregorii, 5
obtusifolium, 5
ovatum, 5
paniculatum, 6, 18
papuanum, 5
Pilgeri, 6
pyrularium, 6
Yasi, 6

Sarcoptes scabiei suis, 75, 83
Sargassum, 200
Scaevola, 10

Gaudichaudiana, 11, 19
glabra, 11

Scaphophorum Agaricoides, 93
Schizophyllum Alneum, 93
commune, 93
Sebacina, 99

caesio-cinerea, 99
cinerea, 96
dubia, 96-97
farinacea, 97-100, 106

Index

259

Galzinii, 99, 100
petiolata, 98-100
Pululahuana, 100
umbrina, 100
seismic sea waves, 22

(travel times of, 185-188)

Septobasidium, 94, 105

sp. ( bogoriense affin.), 105-106
Setaria palmi folia, 47
sheep, parasites of, 76-77, 82
sheep tick, 76
Shepard, F. P., et al.\

The Tsunami of April 1, 1946, in the
Hawaiian Islands, 21-37
Sherman, G. Donald, et al.:

Dolomitization in Semi-arid Hawaiian Soils,
38-44

Sigma Xi, 189-190
Siphonaptera, 78, 80
Sip honia irritans, 73
Sitophilus oryzae, 70, 79

Skeletons of Recent and Fossil Gymnogyps, 227-236
skin filarid, 72
slugs, 77

snails, 71, 72, 77
land, 77
limnaeid, 72

soil, accumulation of arsenic in, 151-171
dolomitization in Hawaiian, 38-44
Hawaiian, toxic levels of arsenic in, 159-169
Sophora chrysophylla, 48

Sorghum vulgar e, toxicity of arsenic for, 151-171
sow bug, 71

sparrow, English, 70, 71
Sphaeria fur, 93
profuga, 93
spinose ear tick, 73, 7 6
Spirogyra, 193, 195, 196
Stachytarpheta cayennensis, 47, 49
staghorn fern, 48
Stemonitis fusca, 94
splendens, 94

Stephanoplaria stile si, 72, 79
Stephanuris dentatus, 74, 83
Stereum hirsutum, 104
stomach worms, 72, 74, 76, 77
Stomoxys calcitrans, 73, 75, 76, 81
Straussia Mar ini ana, 19
strawberry guava, 51
Streptopelia chinensis, 70
Strongyloides papillosus, 72, 79
sp., 74, 79, 83

Strongylus e dentatus, 75, 81
equinus, 75, 81
vulgaris, 75, 81
Stypella minor, 100-101
Styphelia T am eiameiae, 11, 19, 48
Subulina octona, 71, 77, 78, 80
Subulura brumpti, 70, 79
sucking louse, 76

Sudan grass, toxicity of arsenic for, 151-171

sugar cane, 18

effect of soil arsenic on, 169
Sus scrofa domestica, 74-75, 77, 82-83
sweet potatoes, 10

swine, parasites of, 74-75, 77, 82-83

Taenia hydatigena, 75, 76, 77, 80, 82, 83
s a gin at a, 73, 79
taeniaeformis, 77, 78, 79
Taenioma, 196, 210
macrourum, 212
perpusillum, 210

tapeworms, 71, 73, 75, 76, 77, 78, 79, 80, 81, 82,
83

taria, 10

Tenebroides nana, 70, 79
T er min alia Catappa, 47
T etrameres am eric ana, 70, 71, 80
Tetramorium caespitum, 71
spp., 80

Thelaphora cinerea, 96
Thelaphoraceae, 97
thimbleberries, 47, 51
ti, 7, 47, 48
ticks, 73, 74, 76, 77
sheep, 76
spinose ear, 73, 74
tidal waves, 22
toads, 70

tomato, toxicity of arsenic for, 151-171
Tomentella isabellina, 106
Tournefortia sp., 105
Toxascaris leonina, 77, 80

toxicity of arsenic in soil and in culture solution,
151-171

Toxocara canis, 77, 80

Travel Times of Seismic Sea Waves to Honolulu,
185-188
tree ferns, 49

Tribolium castaneum, 70, 79, 80
Trichinella spiralis, 74, 83
trichinosis, 74-75
Trichodectes latus, 77, 80, 81
Trichostrongylus axei, 75, 81
instabilis, 76
Trichuris ovis, 72, 79
suum, 74, 83
vulpis, 77, 80

Triodontophorus brevicauda, 75, 81
sen at us, 75, 81

tsunami ( see also seismic sea wave)
damage by, 34
definition of, 21
history of, in Hawaii, 22
loss of life from and injury by, 35
mitigation of disasters resulting from, 36
Tsunami of April 1, 1946, in the Hawaiian Islands,
21-37

Tubuliferae, 97
tui tui, 10

260

PACIFIC SCIENCE, Vol. 1, October, 1947

Tulasnella allantospora, 95-96
sphaerospora, 96
violea, 96

turkey, parasites of, 69-71, 83
turmeric, 11

Typhaea stercorea, 70, 79

uala, 11, 12
uki, 10, 11
Ulva, 195, 197
fas data, 197
Lactuca, 197

U. S. Bureau of Animal Industry (research
facilities), 123

U. S. Bureau of Entomology and Plant
Quarantine (research facilities), 123
Fruitfly Investigations (research facilities), 124
U. S. Coast and Geodetic Survey (research
facilities), 124

U. S. Commercial Company, 1946 Survey of
Micronesia, 62

U. S. Geological Survey (research facilities),
124-125

Ground Water Division (research facilities),
125

Surface Waters Division (research facilities),
124-125

U. S. Public Health Service (research facilities),
125

U. S. Weather Bureau Office (research facilities),
125

University of Hawaii (research facilities), 125
Agricultural Experiment Station (research
facilities), 126

Micronesian expedition, 60, 92
scientists and the fortieth anniversary, 189-190
ure, 10

U rtica argentea, 10

Utinomi’s Bibliographic a Micronesica:

Chordate Sections, 129-150
uulei, 11
uwara, 10

V aucheria, 195, 198
dichotoma, 198
sp., 198
Thuretii, 198
Vitex hawaiiensis, 244
mollis, 244

Waianae Valley, soil of, 39, 40
Waimea, Oahu, fault at, 85-91
water (ground water), 172-184
wave crests (interval between), 24
Wentworth, Chester K.:

Factors in the Behavior of Ground Water in a
Ghyben-Herzberg System, 172-184
Cycles in Rainfall and Validity in Prediction of
Rainfall in Hawaii, 215-220
white hibiscus, 47
wiliwili, 19
worms

bladder, 73, 75, 76, 77, 78

cecal, 70

eyeworm, 70

gizzard, 70

gullet, 72

hookworm, 77

proventricular, 70, 80

roundworms, 70,_72, 74, 75, 76, 77, 78, 79, 80,
81, 82, 83

stomach, 72, 74, 76, 77

tapeworms, 71, 73, 75, 76, 77, 78, 79, 80, 81,
82, 83

W urdemannia, 196, 204
mini at a, 204, 205
setacea, 204

Xylaria aurantiaca, 95

Zetler, Bernard D.:

Travel Times of Seismic Sea Waves to
Honolulu, 185-188
Zoological Sciences, 59

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Barber, H. H., and I. M. Kolthoff. A specific
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Bennett, George. Gatherings of a naturalist in
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Van Vorst, London, I860.

Coulter, John Wesley. A gazetteer of the Ter¬
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Res. Pub. 11. Honolulu, 1935.

Rock, Joseph F. The sandalwoods of Hawaii;
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No. 1

VOL. II JANUARY, 1948

PACIFIC

SCIENCE

li .

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

IN THIS ISSUE: Hartman — The Polychaetous Annelids
of Alaska • Fisher — Avian Introductions in Hawaii •
NOTES: Alicata — Parasites of Domestic Animals in Mi¬
cronesia ° Fisher — Laysan Albatross Nesting off Oahu •
Hiatt — The Hawaii Marine Laboratory • News Notes

Published by

THE UNIVERSITY OF HAWAII
HONOLULU, HAWAII

BOARD OF EDITORS

Leonard D. Tuthill, Editor-in-Chief
Department of Zoology and Entomology, University of Hawaii

O. A. Bushnell, Assistant Editor
Department of Bacteriology, University of Hawaii

Ervin H. Bramhall

Department of Physics, University of Hawaii

Vernon E. Brock

Division of Fish and Game, Territorial Board of
Agriculture and Forestry
P. O. Box 3319, Honolulu 1, Hawaii

Harry F. Clements

Plant Physiologist, University of Hawaii Agricultural
Experiment Station

Charles H. Edmondson

Zoologist, Bishop Museum, Honolulu 35, Hawaii

Harvey I. Fisher

Department of Zoology, University of Hawaii

Frederick G. Hold away

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Department of Botany, University of California
Berkeley 4, California

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Department of Botany, University of Hawaii

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P. O. Box 3410, Honolulu 1, Hawaii

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■

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

Vol. II JANUARY, 1948 No. 1

Previous issue published October 17, 1947

CONTENTS

PAGB

The Polychaetous Annelids of Alaska. Olga Hartman . 3

The Question of Avian Introductions in Hawaii. Harvey I. Fisher .... 59

Notes:

Observations on Parasites of Domestic Animals in Micronesia.

Joseph E. Alicata . 65

Laysan Albatross Nesting on Moku Manu Islet, off Oahu, T. H.

Harvey I. Fisher . 66

Preliminary Note on the Oceanographic Program of the Hawaii Marine

Laboratory. Robert W. Hiatt . 67

News Notes . 68

Pacific Science, a quarterly publication of the University of Hawaii, appears in January,
April, July, and October. Subscription price is three dollars a year; single copies are one
dollar. Check or money order payable to University of Hawaii should be sent to Pacific
Science, Office of Publications, University of Hawaii, Honolulu 10, Hawaii.

The Polychaetous Annelids of Alaska

Olga Hartman1

INTRODUCTION

This PAPER is based on the polychaetous an¬
nelids obtained by the Alaska King Crab Investi¬
gation, which was sponsored by the United
States Fisheries Commission. The annelids were
taken almost entirely from southern and south¬
western Alaska, from Port Ashton west to the
western end of the Alaska Peninsula (Map I).
In addition, some were obtained from scattered
stations eastward to Icy Straits, near Pleasant
Island, and to the north and west in the Bering
Sea, north of St. Lawrence Island (Map II).
The studies were conducted from September to
November, 1940, and from February to August,
1941. Bathymetric ranges were largely limited
to shallow depths, including shore to 60 fathoms,
but three stations represented by polychaetes
were in depths of 100 to 150 fathoms. A general
report of the investigation has been published
(Investigation Staff, 1942).

The intertidal areas of Alaska have heretofore
been little explored for the annelid fauna. Ex¬
cept for the vast collections of the U.S.S. "Alba¬
tross” made by the Alaskan Salmon Commission
during the summer of 1903 (of which much
was dredged from deep water), the published
records are quite limited, although they extend
over many years (1821 to 1943).

The collections of the present investigation
comprise 99 species, including six new to
science. There are 45 new records for Alaska,
including three species from material in the
authors collection. These, together with 168
species (and seven questionable names) previ¬
ously recorded, bring the total number for

1 Allan Hancock Foundation, University of Southern
California. Manuscript received April 15, 1947.

Alaska to 213 species plus the seven question¬
able names. These results are discussed in detail
below.

The various stations are listed and the known
ecological data of each are given at the end of
the systematic treatment.

The bibliographic citations in the systematic
section include the original description and such
others as aid in the ready identification of the
species. For most of the species originally de¬
scribed from the eastern Pacific the citations are
Complete unless they are synonymized elsewhere.
In all cases it is possible to consult all references
for Alaska and environs from the data given.

Acknowledgments: The collections on which
the present work is based were obtained under
the direction of Dr. Waldo L. Schmitt. The
preliminary sorting of collections was performed
by the staff of the Invertebrate Division of the
United States National Museum, to whom
thanks are extended. The two maps and figures
1, 8, and 12 were done by Mr. Anker Petersen.
The author is deeply indebted to the adminis¬
tration of the Allan Hancock Foundation for
allowing time and facilities to conduct these
studies.

The specimens are deposited in the United
States National Museum; a duplicate set is in
the collections of the Allan Hancock Founda¬
tion, University of Southern California.

HISTORICAL RESUME

The literature on the polychaete fauna of
Alaska is enriched largely through the results of
expeditionary ventures that had other objec¬
tives. Chronologically considered, the data are
as follows. The first annelids were described by

3

4

PACIFIC SCIENCE, Vol. II, January, 1948

Chamisso and Eysenhardt (1821) from collec¬
tions made during a Russian voyage around the
world. Two new species, Nereis heteropoda
and Sternaspis elegans, were reported from Un-
alaska. Both remain indeterminable, although
the first was redescribed by Grube (1855).

A later Russian expedition resulted in the de¬
scription of three new species by Grube (1851),
Nereis vexillosa, Polynoe cirrata, and Cirratulus
borealis , from Sitka. The latter two have
since been referred to Harmothoe imbricata
(Linnaeus), and, questionably, Cirratulus cirra-
tus (Muller), respectively. The same author
(Grube, 1855) added two more to the list,
Polynoe vittata and Polynoe tuta, which are now
known as Arctonoe vittata (Grube) and Holo-
lepidella tuta ( Grube ) .

The International Polar Expedition to Point
Barrow, Alaska, in 1885, resulted in the
description of Arenicola glacialis Murdoch
(1885*0 and the addition of five other species
(Murdoch, 1885£). These are Melaenis loveni
Malmgren, Phyllodoce groenlandica Oersted
(here called Anaitides ) , Cast alia multipapillata
Theel (here called Psammate) , Aricia arctica
Murdoch (questionably referred to Scoloplos
armiger Muller), and Brad a granulata Malm¬
gren.

Johnson (1901), in reporting on the anne¬
lids from Puget Sound, Washington, named one,
Polynoe insignis (Baird) (here called Halo-
sydna brevisetosa Kinberg), from Kadiak Island,
Alaska.

Moore (1902) described numerous species
from northern Greenland. Four of these, Gat-
tyana senta Moore (here called Eunoe ), Gat-
tyana ciliata Moore, Lagisca multisetosa Moore,
and Eunoe truncata Moore (here called Herma-
dion ), were later (Moore, 1905*z: 525) said to
have come from Icy Straits, Alaska, rather than
from Greenland.

In 1899 the Harriman Alaska Expedition ex¬
plored the waters about Alaska. Among the
polychaetes, only the families Sabellidae and
Serpulidae were investigated (Bush, 1904). As a
result, 18 species were newly recorded from

Alaska, and five others, whose identity cannot be
established (see Hartman, 1942, for revision
and summary), were reported.

The U.S.S. "Albatross,” operated by the Alas¬
kan Salmon Commission in the summer of 1903,
explored the waters from Port Townsend, Wash¬
ington, to Shelikof Strait, Alaska. The poly¬
chaetes were ably described by Moore (1905 to
1908). As a result, 97 species of annelids were
reported from Alaska, 81 for the first time.
However, five of these, Brada pilosa Moore, Ster¬
naspis f oss or Stimpson, Pseudopotamilla ano-
culata Moore, Pseudopotamilla brevibranchiata
Moore, and Pseudopotamilla reniformis (Leuc-
kart), are regarded as synonyms, and one other,
Nereis ( Alitta ) virens Sars, includes two species.

McIntosh (1910) added one new record,
Lumbriconereis fragilis (Muller) (here called
Lumbrineris) , from Alaska but the record is not
clear. It appears in his monograph of British
annelids.

Nine new records were added by Treadwell
(1914) among a total of 24 species from
Alaska. A re-examination of the material on
which these records were based has shown the
following misidentifications: Nereis procera
Ehlers is Nereis pelagica Linnaeus, Naineris
longa Moore is Naineris dendritica (Kinberg),
Cistenides hyperborea Malmgren is Cistenides
brevicoma (Johnson), Eudistylia polymorph a
(Johnson) is Eudistylia vancouveri (Kinberg),
and Schizobranchia nobilis Bush is Schizo-
branchia dubia Bush. The type collection of
Scolecolepis alaskensis Treadwell from Shuma-
gin Islands, Alaska, has been examined and the
species is here newly referred to the genus
Nerine.2

2 The prostomium lacks anterior horns; it is pro¬
longed forward as a snout. Branchiae are present from
the second setigerous segment and continued through
a long region but absent from a considerable posterior
portion. They are fused for their entire length with
the postsetal lamella. The posterior end is a flat, short
lobe. The first segment is biramous and has slender
pointed setae both above and below. Hooded hooks
are present in both notopodia and neuropodia, together
with a tuft of pointed setae, after about the fifty-ninth
segment. The prostomium may lack eyes. These char¬
acters are those of Nerine.

Annelids of Alaska — HARTMAN

5

A single species was recorded from Alaska by
Ditlevsen ( 1917 ) as Harmothoe aspera Hansen,
but this is questionably referred to Lagisca mul-
tisetosa Moore, and was earlier reported. The
same year Essenberg (1917) described a new
species, Euphrosyne multibranchiata, from Ko¬
diak Island, Alaska.

Treadwell (1921) added Nereis ( Cerato -
nereis) alaskensis as a new species, but it has
been referred to Ceratonereis paucidentata
(Moore) (Hartman, 1938^: 13). He listed 13
additional species in 1925 and 1926. Of these,
five were new records. Subsequently (Tread¬
well, 1943) he added Neosabellides alaskensis
Treadwell.

Other recent additions are three new species
added to the list by Hartman (1938 b) and 17
new records added by Berkeley (1942) in a
report of 49 species from Alaska.

Between 1929 and 1934 the State Hydro-
logical Institute of the Union of the Socialist
Soviet Republics (Russia) made extensive
faunal investigations in the northern part of the
Japan Sea. The polychaetes that were collected,
together with some others taken from this re¬
gion, comprised a total of 272 species (Annen¬
kova, 1937 and 1938, with summary in the
latter). Among these, 61 species are identical
with those known from Alaska. These are in¬
dicated in the systematic list below by the letter
/. Annenkova stated (1938) (translation from
the Russian) that about 40 per cent of the ob¬
served fauna of the northern Japan Sea are
species native to the northern part of the At¬
lantic Ocean, and that only about 10 per cent
are common to the Alaska-California fauna. She
adds, however, that more intensive collecting is
necessary before sound conclusions can be
drawn. The list below indicates that nearly 30
per cent of the species are the same.

Annenkova (1938: 142-144) recognized 12
groups of species. These groups are : ( 1 ) Arctic-
boreal species, numbering about 25; (2) Arctic
species, about 22; (3) species common to the
Arctic, Atlantic, and Bering Sea, about 18;
(4) boreal species common to the north At¬

lantic, about 22; (5) subarctic species with
interrupted distribution, about 8; (6) species
common to Japan and the western Bering Sea,
6 named in a list; (7) Japan-Okhotsk species,
18 named in a list; (8) species common to
California, Alaska, and the north Japan Sea,
about 16; (9) subtropical Japanese species, 9 or
10; (10) Indo-Pacific species, 9; (11) endemic
species, 17; and (12) cosmopolitan species, 13-
In summary, Annenkova states that 36 per cent
are warm- water species, 23 per cent are Arctic-
boreal, 11 per cent are western Pacific, and 11
per cent are Arctic species.

For the present, six groups may be recognized
from the Alaskan species, although it is likely
that no actual barriers exist; rather, with more
intensive studies, many species now having a
restricted range will probably be found more
widely dispersed geographically.

The nearest affinities indicated by the tabula¬
tion of this material are with the eastern north
Pacific coast, which has nearly 40 per cent repre¬
sented; with the north Atlantic, which has nearly
37 per cent; and with the north Japan Sea,
which has nearly 30 per cent.

The six groups recognized here are designated
a to / in the following list, in which the 220
species known from Alaska are systematically
arranged by families. Of these 220 species 7 are
doubtful. In the list the reference following the
complete name indicates the first record for
Alaska. The names without such citation are
here recorded for the first time.

The bracketed letters a to f and J indicate:

a — Alaska only, including 35 species
b — Alaska and Arctic, including 4 species
c — Alaska and either British Columbia, Wash¬
ington, or Oregon, including 38 species
d — Alaska to California, including 89 species
e — Alaska to California, and Japan or China,
including 38 species

/ — Alaska and north Atlantic, including 78
species

7 — refers to the species occurring in Annen¬
kova’s (1938) list.

6

PACIFIC SCIENCE, Vol. II, January, 1948

List of the Species of Polychaetes
Known from Alaska

Family APHRODITIDAE, one species

Aphrodita japonica Marenzeller (Moore,
1908) ie]

Family POLYNOIDAE, 27 species
Antinoe macrolepida Moore (Moore, 1905)

id]

Antinoe sarsi Kinberg (Chamberlin, 1920)
( as Antinoella by Annenkova ) [/, /]
Arctonoe pulchra (Johnson) (Moore, 1908)

id]

Arctonoe vittata (Grube) (Grube, 1855)

y, n

Enipo cirrata Treadwell (Treadwell, 1925)

M

Eunoe barbata Moore (Treadwell, 1925)

w

Eunoe depressa Moore (Moore, 1905) [ c, /]
Eunoe nodosa (Sars) (Moore, 1910) if, J]
Eunoe senta (Moore) (Moore, 1905) id]
Ev amelia triannulata (Moore) id]

Gattyana amondseni (Malmgren) (Moore,
1908) [/]

Gattyana ciliata Moore (Moore, 1905) ic, ]]
Gattyana cirrosa (Pallas) (Berkeley, 1942)

if, n

Gattyana imbricata Treadwell (Treadwell,
1926) ia]

Gattyana iphionelloides (Johnson) ic]
Halosydna brevisetosa Kinberg (Johnson,
1901) id]

Harmothoe hirsuta Johnson (Moore, 1908)

id]

Harmothoe imbricata (Linnaeus) (Grube,
1851) ie,f,J]

Hermadion truncata ( Moore ) ( Moore, 1905 )

ic]

Hololepida magna Moore (Moore, 1905)

ic]

Hololepidella tut a (Grube) (Grube, 1855)

ic,J]

Lagisca lamellifera (Marenzeller) (Moore,
1910) id] (= L. multisetosa papillata
Moore (1908))

Lagisca multisetosa Moore (Moore, 1905)

id]

Lagisca rarispina (Sars) (Moore, 1908) [/}
Lepidonotus caeloris Moore (Moore, 1903)
id]

Lepidonotus robustus Moore (Moore, 1905)

ia]

Melaenis loveni Malmgren (Murdoch, 1885)

if]

Family POLYODONTIDAE, one species
PeisidAce asp era Johnson (Moore, 1908) id]

Family SIGALIONIDAE, one species

Pholoe minuta Fabricius (Moore, 1908) [/,

/]

Family CHRYSOPETALIDAE, one species

Paleanotus chrysolepis Schmarda id]

Family EUPHROSINIDAE, five species
Euphrosine arctia Johnson (Moore, 1908)
id]

Euphrosine bicirrata Moore (Moore, 1905)
id]

Euphrosine heterobranchia Johnson (Tread¬
well, 1914) ic]

Euphrosine hortensis Moore (Moore, 1905)
id, J]

Euphrosine multibranchiata Essenberg (Es-
senberg, 1917) ia, J]

Family SPINTHERIDAE, one species
Spinther alaskensis new species ia]

Family PHYLLODOCIDAE, nine species
Anaitides citrina (Malmgren) (Moore, 1908)
[/]

Anaitides groenlandica (Oersted) (Murdoch,
1885) [ c,f,J]

Anaitides medipapillata (Moore) (Tread¬
well, 1926) id]

Anaitides mucosa (Oersted) (Moore, 1908)
id, f]

Eteone calif ornica Hartman id]

Eteone spetsbergensis Malmgren ic, f, J]
Eulalia viridis (Muller) ie, f, J]
Notophyllum folio sum (Sars) [/]
Notophyllum imbricatum Moore (Moore,
1906) ie, /]

Family ALCIOPIDAE, one species

Callizona angelini (Kinberg) (Moore, 1908)

M

Family HESIONIDAE, two species

Psammate aphroditoides (Fabricius) (Cham¬
berlin, 1920) [ b]

Psammate multipapillata (Theel) Murdoch,
1885) ib]

Family SYLLIDAE, 13 species

Autolytus alexandri Malmgren (Chamberlin,
1920) if]

Autolytus magnus Berkeley ic]

Autolytus prismaticus (Fabricius) (Cham¬
berlin, 1920) ib, c, /]

Pionosyllis gigantea Moore (Moore, 1908)
id]

Annelids of Alaska — HARTMAN

7

Piono syllis magnifca Moore (Moore, 1906)

id, n

Syllis quaternaria Moore (Moore, 1906) [a]
Trypanosyllis gemmipara Johnson id, J]
Typo syllis alternata ( Moore ) ( Moore, 1 908 )

id, n

Typo syllis armillaris ( Muller ) ( Moore, 1908 )

if, n

Typo syllis collaris new species ia]

Typo syllis elongata (Johnson) id}

Typosyllis pulchra (Berkeley) ic]

Typosyllis stewarti (Berkeley) ic]

Family NEPHTYIDAE, seven species
Nephtys assimilis Malmgren (Moore, 1908)
[/} (or possibly Nephtys homhergi Au-
douin and Edwards)

Nephtys ciliata (Muller) (Moore, 1908)

ie, f, n

Nephtys coeca (Fabricius) (Moore, 1908)

ie, f, J]

Nephtys malmgreni Theel (Moore, 1908)

if, n

Neyhtys punctata Hartman (Hartman, 1938)

id}

Nephtys rickettsi Hartman (Hartman, 1938)

id]

Nephtys schmitti Hartman (Hartman, 1938)

id]

Family NEREIDAE, 11 species

Ceratonereis paucidentata (Moore) (Moore,
1908) id]

Cheilonereis cyclurus (Harrington) ie, J]
(= Nereis schischidoi Izuka (Annenkova,
1938))

Neanthes brandti (Malmgren) (Moore,
1908) M

Neanthes virens (Sars) (Moore, 1908) ie,

f,n

Nereis neoneanthes new species ic]

? Nereis heteropoda Chamisso and Eysen-
hardt (Chamisso and Eysenhardt, 1821)

M

Nereis pelagica Linnaeus (Moore, 1908) ie,

f,n

Nereis procera Ehlers id]

Nereis vexillosa Grube (Grube, 1851) ie, J]
Nereis zonata Malmgren [/, /}

Platy nereis agassizi (Ehlers) (Moore, 1908)

ie, n

Family SPHAERODORIDAE, one species
Sphaerodorum minutum (Webster and Bene¬
dict) (Chamberlin, 1920) [/}

Family GLYCERIDAE, two species

Glycera capitata Oersted (Moore, 1908) ie,

f,n

Hemipodus borealis Johnson id]

Family GONIADIDAE, four species

Glycinde picta Berkeley (Berkeley, 1942) ic]
Glycinde wireni Arwidsson (Moore, 1908)

w

Goniada annulata Moore (Moore, 1905) id]
Goniada maculata Oersted (Berkeley, 1942)

ie, f, J]

Family ONUPHIDAE, three species

Nothria conchylega (Sars) (Berkeley, 1942)

ie, f, J]

Nothria geophiliformis (Moore) (Moore,
1908) ie]

Nothria iridescens (Johnson) (Moore, 1908)

id]

Family EUNICIDAE, one species
Eunice longicirrata Webster (Moore, 1908)

id,f]

Family LUMBRINERIDAE, seven species
Lumbrineris bicirrata Treadwell id]

? Lumbrineris fragilis (Muller) (McIntosh,
1910) [/,/]

Lumbrineris heteropoda Marenzeller (Moore,
1908) ie, J]

Lumbrineris latreilli Audouin and Edwards

id,f]

Lumbrineris similabris Treadwell (Tread¬
well, 1926) ic]

Lumbrineris zonata (Johnson) id]

Ninoe simpla Moore (Moore, 1908) ia]

Family ARABELLIDAE, two species

Drilonereis filum Claparede (Berkeley, 1942)

if]

Drilonereis nuda Moore id]

Family DORVILLEIDAE, one species
Dorvillea pseudorubrovittata Berkeley ic]

Family ORBINIIDAE, four species

Haploscoloplos alaskensis new species id]
Haploscoloplos elongata (Johnson) id]
Naineris dendritica (Kinberg) (Treadwell,
1914) id]

Scoloplos armiger (Muller) (? Murdoch,
1885; verified by Berkeley, 1942) ie, f, J]

Family PARAONIDAE, one species
Aricidea heteroseta new species id]

8

Family SPIONIDAE, nine species

Anaspio boreus Chamberlin (Chamberlin,

1920) M

Laonice cirrata (Sars) \e, f, J]

Nerine alaskensis (Treadwell), new com¬
bination (Treadwell, 1914) ia]

Polydora giardi Mesnil id, /]

Polydora socialis (Schmarda) id, /]
Prionospio cirrifera Wiren (Berkeley, 1942)

tel

Prionospio malmgreni Claparede ic, f, J]
Scolecolepides arctius Chamberlin (Chamber¬
lin, 1920) M

Spio filicornis (Miiller) (Chamberlin, 1920)

le,f,n

Family CHAETOPTERIDAE, one species
Chaetopterus variopedatus (Renier) (Berke¬
ley, 1942) [e,n

Family CIRRATULIDAE, four species
Acrocinus heterochaetus Annenkova [a, J ]
Cirratulus cirratus (Muller) (Grube, 1851)
{>,/,/}

Cirratulus robustus Johnson (Treadwell,
1914) id]

Tharyx hamatus new species ia]

Family ARENICOLIDAE, two species

Arenicola glacialis Murdoch ( Murdoch, 1885 )

M

Arenicola pusilla Quatrefages (Treadwell,
1914) id]

Family OPFIELIIDAE, seven species

Ammotrypane aulogaster Rathke (Moore,
1908) Id, f]

Armandia bioculata Hartman id]

Armandia brevis (Moore) (Moore, 1906)
id]

Ophelia limacina (Rathke) (Berkeley, 1942)

id,n

Travisia brevis Moore (Berkeley, 1942) id]
Travisia forbesii Johnston (Moore, 1908) [/]
Travisia pupa Moore (Moore, 1906) id']

Family SCALIBREGMIDAE, one species
S calibre gma inf latum Rathke (Moore, 1908)

te fl

Family FLABELLIGERIDAE, five species
Brada granulata Malmgren (Murdoch, 1885)

[fl

Brada villosa (Rathke) (Moore, 1906) ic, f]
Flabelligera infundibularis Johnson (Tread¬
well, 1914) id]

Stylarioides papillata (Johnson) (Moore,
1908) id]

PACIFIC SCIENCE, Vol. II, January, 1948

Stylarioides plumosa (Muller) (Berkeley,
1942) fa f]

Family CAPITELLIDAE, three species
Capitella capitata (Fabricius) id, /}
Heteromastus filiformis (Claperede) id, f]
Notomastus giganteus Moore (Moore, 1906)

M

Family MALDANIDAE, nine species
? Asychis lacera Moore id]

Asychis similis (Moore), new combination
(Moore, 1906) id]

Clymenella tentaculata Moore (Moore, 1906)

M

Maldane sarsi Malmgren (Moore, 1908) id,

n

Maldanella robusta Moore (Moore, 1906)

tel

Nicomache lumbricalis (Fabricius) (Moore,
1906) id, f, J]

Nicomache personata Johnson ic]
Notoproctus pacificus (Moore) (Moore,
1906) id]

Petaloproctus tenuis borealis Arwidsson ic]

Family OWENIIDAE, one species

Owenia occidentalis (Johnson) (Treadwell,
1914) id]

Family SABELLARIIDAE, two species
Idanthyrsus ornamentatus (Chamberlin)
(Berkeley, 1942) ie, J] (Reported as
I. armatus Kinberg)

Sabellaria cementarium Moore (Moore, 1906)

ie,J]

Family STERNASPIDAE, two species

? Sternaspis elegans Chamisso and Eysenhardt
(Chamisso and Eysenhardt, 1821) ia]
Sternaspis scutata Ranzani (Moore, 1908)

te, f, n

Family PECTINARIIDAE, five species

Amphictene auricoma (Muller) (Moore,
1908) ic,f]

Cistenides brevicoma (Johnson) (Treadwell,
i9!4) id]

Cistenides granulata (Linnaeus) (Chamber¬
lin, 1920) ic,f,J]

Cistenides hyperborea Malmgren (Berkeley,
1942) [/,/}

Pectinaria belgica (Pallas) (Berkeley, 1942)

tafl

Family AMPHARETIDAE, 15 species

Amage anops (Johnson) (Berkeley, 1942)

te, n

Annelids of Alaska — HARTMAN

9

Ampharete arctica Malmgren (Moore, 1908)

fo f, n

Ampharete hrevihranchiata Treadwell (Tread¬
well, 1926) {a}

Ampharete eupalea Chamberlin (Chamberlin,

1920) M

Ampharete gruhei Malmgren (Berkeley,

1942) u,n

Ampharete johanseni Chamberlin (Chamber¬
lin, 1920) [a]

Ampharete reducta Chamberlin (Chamberlin,
1920) [a]

Amphicteis alaskensis Moore (Moore, 1905)

M

Amphicteis glabra Moore (Moore, 1905) {d}
Lysippe labiata Malmgren (Berkeley, 1942)

Melinna cristata (Sars) (Moore, 1908) [c,

f,n

Melinna denticulata Moore (Moore, 1905)

M

Neosabellides alaskensis Treadwell (Tread¬
well, 1943) W

Sabellides octocirrata Sars (Berkeley, 1942)

u n

Sarny tha sexcirrata (Sars) (Chamberlin,

1920) [d,f]

Family TEREBELLIDAE, 16 species
Amphitrite cirrata (Muller) (Moore, 1905)

ic, n

Eupolymnia crescentis Chamberlin [i]
Eupolymnia heterobranchia (Johnson)
(Moore, 1908) M

Eupolymnia nesidensis japonica (Marenzel-
ler) W

Leaena abranchiata Malmgren [ a, f, /]

Leaena nuda Moore (Moore, 1905) (or pos¬
sibly Lanas s a) [, a ]

Neoamphitrite robusta (Johnson) (Moore,
1908) M

Neoleprea spiralis (Johnson) [e, J]

Nicolea zostericola ( Oersted ) [a, f, /]

Pista cristata (Muller) (Moore, 1908) [e,

f.n

? Pista fas data (Grube) (Moore, 1908) [c,

n

Polycirrus sp. (Moore, 1908)

Spinosphaera sp.

Terebellides stroemi Sars (Moore, 1908) [e,

f,n

Thelepus crispus Johnson (Treadwell, 1914)
Thelepus hamatus Moore (Moore, 1905) \d}

Family SABELLIDAE, 16 species

Chone gracilis Moore (Moore, 1906) [c]
Chone infundibuliformis Kroyer (Bush,
1904) [*,/,/]

Euchone analis (Kroyer) (Berkeley, 1942)

ic, f, n

Eudistylia polymorpha (Johnson) (Bush,
1904) (as Bispira by Annenkova) \e, /}
Eudistylia tenella Bush (Bush, 1904) {a}
Eudistylia vancouveri (Kinberg) (Bush,

1904) M

Megalomma splendida (Moore) (Moore,

1905) Id}

Myxicola aesthetica (Claparede) (Bush,
1904) W/J

Myxicola infundibulum (Montagu) (Bush,

1904) W,/]

Potamilla neglecta (Sars) (Bush, 1904) {d,

n

Pseudopotamilla intermedia Moore (Moore,

1905) M

Pseudopotamilla occelata Moore (Moore,
1905) M

Sabella crassicornis Sars (Bush, 1904) [d, /}
Sabella media (Bush) (Bush, 1904) [d~\
Schizobranchia dubia Bush. (Bush, 1904) [a]
Schizobranchia insignis Bush (Bush, 1904)

M

Family SERPULIDAE, 11 species and 5 ques¬
tionable names

Chitinopoma occidentalis (Bush), new com¬
bination (Bush, 1904) {d\

Crucigera irregularis Bush (Bush, 1904) [c]
Crucigera zygophora (Johnson) (Bush,
1904) ld,n

Paradexiospira violaceus (Levinsen)

Serpula vermicularis Linnaeus (Bush, 1904)

fo f, n

Spirorbis ( Laeospira ) borealis Daudin (Bush,
1904 ) Id, f }

Spirorbis quadrangularis Stimpson (Moore,
1908) [a,fi

Spirorbis semidentatus Bush (Bush, 1904)

M

Spirorbis spirillum Linnaeus (Moore, 1908)

fo /, /]

Spirorbis tridentata Levinsen (Moore, 1908)

u,n

Spirorbis 'variabilis Bush (Bush, 1904) M
? Spirorbis abnormis Bush (Bush, 1904) [a]
? Spirorbis incongruus Bush (Bush, 1904)

M

? Spirorbis lineatus Bush (Bush, 1904) M
? Spirorbis rugatus Bush (Bush, 1904) [a]
? Spirorbis similis Bush (Bush, 1904) [a}

PACIFIC SCIENCE, Vol. II, January, 1948

FIG. 1. Spinther alaskensis new species: a, entire animal in dorsal view, X 6.4; b, portions of
typical notopodial ridges from the dorsal region to show the alternation of setae in anterior and
posterior margins, enlarged; c, neuropodium from a median segment, in anterior view, showing
the acicular sac and developing setae torn from the body wall, enlarged.

Annelids of Alaska —HARTMAN

11

Except for the pelagic families of polychaetes,
the only ones not known to be represented are
the following: Pareulepidae, Palmyridae, Am-
phinomidae, Typhloscolecidae, Pilargiidae, Ly-
saretidae, Apistobranchidae, Magelonidae, Diso-
midae, and Ctenodrilidae. With the exception
of the Amphinomidae, these are small families
which rarely occur in collections. The amphi-
nomids are largely tropical.

DISCUSSION

Family APHRODITIDAE

Genus Aphrodita Linnaeus

Aphrodita japonica Marenzeller
Aphrodita Japonica Marenzeller, 1879: 111—
112, pi. 1, fig. 2; Moore 1908: 338-339;
Hartman, 1939: 21-22, pi. 1, fig. 1-5; Berke¬
ley, 1924: 187.

Collections. Stations 116-40 (1); 131-40
(1); 135-40 (2); 138-40 (1); 139-40 (1);
140-40 (3).

Some are very large, measuring 4.5 inches
long and 2.25 inches wide. The lateral hairs are
long but only slightly opalescent. This species
is known to range throughout the northern
Pacific and southward; present records are from
Alaska in 28 to 48 fm.

Family POLYNOIDAE

Genus Arctonoe Chamberlin

Arctonoe fragilis (Baird)
Lepidonotus fragilis Baird, 1863: 108.

Polynoe fragilis Johnson, 1897: 179-181, pi. 7,
fig. 36, 45, pi. 8, fig. 52.

Arctonoe fragilis Hartman, 1938c: 116; Berke¬
ley, 1942: 188.

Collections. Stations 24-40 ( 2 ) ; 25-40 ( 1 ) ;
34-40 (1).

This species is widely known through the
northeast Pacific; it is free-living or associated
with echinoderms. The present records are from
Canoe and Pavlof bays, in 25 to 150 fm.

Arctonoe vittata (Grube)

Fig. 2 a-f.

Polynoe vittata Grube, 1855: 82-83.

Polynoe lordi Johnson, 1897: 175-177, pi. 7,
fig. 35, 44, pi. 8, fig. 51.

Halosydnoides vittata Okuda, 1936: 565-568,
fig. 4, 5.

Arctonoe vittata Hartman, 1939: 29-30, pi. 3,
fig. 33-37; Berkeley, 1942: 188.

Collections. Stations 20-40 (2); 21-40 (1);
20-40 to 22-40 (2); 34-40 (1); 35-40 (1);
59-40 (1); 70-40 (1); 71-40 (2); 80-40
(1); 83^0 (1); 129-40 (1); 131-40 or 132-

40 (1); BT 70-41 (1); C 44-41 (2); C 71-

41 (1); L 4-41 (4).

These collections include individuals with at
least three color patterns. One group is pale
with a broad dark band across the dorsum an¬
teriorly; the individuals come from stations
20-40 to 22-40, 59-40, 80-40, 83-40, 129-40
and C 71-41. Another group has elytra more
or less intensely mottled, as shown by Okuda
(1936: 566); these individuals come from sta¬
tions 21-40, 34-40, 35-40, 71-40 and BT 70-
41. A third group has elytra with a narrow dark
edge on the inner and posterior margins; the
individuals are from station L 4-41. In addition,
there are some individuals that are entirely pale
(preserved), coming from stations 70-40, 71-
40, 131-40 or 132-40 and C 44-41. For each
of these groups the characteristic setae from the
notopodium and neuropodium of the second
segment (first setigerous segment) have been
examined and found to agree in details as shown
in figures of notosetae (Fig. 2 a, 2 c) and neuro-
setae (Fig. 2 h, 2 d) from stations 71-40 and
80-40, and two notosetae (Fig. 2 e, 2 f) from
station L 4-41.

A. vittata is frequently associated with mol-
lusks, echinoderms or other chaetopods, or it

12

PACIFIC SCIENCE, Vol. II, January, 1948

Fig. 2. Species of Arctonoe and Spinther. a-f, Arctonoe vittata (all figures are distal ends of notosetae and
neurosetae from the first setigerous segment, X 425) : a, notoseta and b, neuroseta from station 71—40; c, noto-
seta, and d, neuroseta from station 80-40; e, worn notoseta, and f, unworn notoseta from station L 4-41.
g—j, Spinther alaskensis: g, neuropodial hook from a median segment, X 100; h, j, entire notopodial setae from a
median segment showing variation in tips, X 425; i, bifid notopodial seta, X 812.

may be free living. The present records are from
known ranges in western and southern Alaska,
in 5 to 150 fm., and from the Bering Sea in
33 fm.

Genus Lepidonotus Leach

are none on the present individuals or on those
of the original collection ( loc . tit.).

L. rohustus was first described from Shelikof
Strait, Alaska, in 48-65 fm. The present records
are from Canoe Bay and Cold or Pavlof Bay, in
15 to 40 fm., not far from the type locality.

Lepidonotus robustus Moore

Lepidonotus rohustus Moore, 1905^: 544-546,
pi. 36, fig. 32-35.

Collections. Stations 5 1-40 ( 3 ) ; 61-50 ( 1 ) ;
Canoe or Pavlof Bay ( 1 ) .

These individuals agree closely with Moore’s
description (1905: 544). It now seems doubt¬
ful that this species is actually the same as
L. helotypus (Grube) (1877: 49) originally
described from China, although earlier I ( 1938c:
109) followed Seidler (1924: 56) in referring
it to the latter. L. helotypus is said to have
small, dark, nailheaded spines on elytra. There

Genus Halosydna Kinberg

Halosydna brevisetosa Kinberg
Halosydna brevisetosa Kinberg, 1857: 18, pi. 5,
fig. 25; Berkeley, 1942: 189.

Polynoe brevisetosa Johnson, 1897: 167-170,
pi. 6, fig. 24, pi. 7, fig. 31, 40, pi. 8, fig. 46.

Collections. Station 51-40 (1).

This well-known species is common through¬
out the northeast Pacific, from Alaska south to
southern California. It is sometimes commensal,
in tubes of other chaetopods, where it attains
comparatively great size. The present record is
from Canoe Bay in 25 to 40 fm.

Annelids of Alaska— HARTMAN

13

Genus Hololepida Moore

Hololepida magna Moore
Hololepida magna Moore, 1905^: 541-544, pi.

35, fig. 24-29; Moore, 1908: 320; Berkeley,

1923: 214.

Collection. Station L 13-41 (1).

This is known only from southern Alaska and
British Columbia; it ranges in depths of 13 to
230 fm.

Genus Hololepidella Willey
Hololepidella tuta (Grube)

Polynoe tuta Grube, 1855: 82.

Harmothoe tuta Johnson, 1901: 394-396, pi.

2, fig. 18, 19, pi. 3, fig. 20-22.

Hololepidella tutta Annenkova, 1937: 147, pi.

3, fig. 15, 16.

Polyeunoa tuta Berkeley, 1942: 188.

Collections. Stations 35-40 (fragment); 59-
40 ( short fragment) ; 60-40 ( 1 ) ; D 3-41 ( 2 ) .

Originally described from Sitka, Alaska, this
has been widely reported from both sides of the
north Pacific ( see synonymy above ) . The pres¬
ent records are from southwest Alaska in 20 to
30 fm.

Genus Harmothoe Kinberg
Harmothoe imbricata (Linnaeus)
Harmothoe imbricata Johnson, 1897: 181, pi.

7, fig. 37; Fauvel, 1923: 55, fig. 18; Berkeley,

1923: 215; Berkeley, 1942: 187.

Collections. Stations 12-40 (1); 24-40 (1);
27-40 (1); 33-40 (1); 34-40 (1); 47-40
(1); 51-40 (about 15); 59-40 (1); 61-40
(1); 72-40 (1); 89-40 (1); 97-40 (3); 108-
40 (1); CT 12-41 (1); D 7-41 (1); D 15-41
(12); L 2-41 ( 1 ) ; L 3-41 (2); L 20-41 (3);
Lazy Bay ( 1 ) ; Seldovia ( 1 ) .

Color patterns are highly variable. Some in¬
dividuals have elytra more or less heavily
mottled with gray to russet pigment; others have
the first pair pale, the rest dark; a few in¬
dividuals are striped longitudinally, the elytra
dark brown on the inner portions and pale on

the outer parts. In all individuals, notosetae are
coarser than neurosetae; the latter are distinctly
bidentate at the distal ends. The dorsum is com¬
pletely covered by the imbricated elytra. On
the prostomium, anterior eyes are far forward,
under the prostomial peaks and directed an¬
teriorly. The elytra are thick and leathery in
texture, sparsely fringed along their outer free
margins, oval in shape. Dorsal cirri are smooth
except for a few short, scattered papillae. Neph-
ridia are very long and slender; they are clearly
seen on the ventral side of parapodia.

H. imbricata is common in colder waters of
the north Pacific, south at least to central Cali¬
fornia. The present records are from southwest
Alaska and Bering Sea, from shore to 150 fm.

Genus Evarnella Chamberlin
Evarnella triannulata (Moore)
Harmothoe triannulata Moore, 1910: 346-350,
pi. 29, fig. 18-22.

Evarne triannulata Berkeley, 1923: 215.
Evarnella triannulata Berkeley, 1942: 188.

Collections. Stations 2 1-40 (1); 33-40 (1);
51-40 (6); 58-40 (1); 59-40 (1); 60-40
(4); 61-40 (5); 72-40 (1); 89-40 (1); 93-
40 (1); 100^0 (1); 128-40 (4); A 8-41
(1); C 5-41 (2); D 7-41 (2); D 8-41 (2);
D 11-41 (4); L 18-41 (1); Sand Point (1).

The body consists of 39 or 40 segments;
elytra number 15 pairs and cover most of the
body except the last five or six short segments.
Elytra are oval, the margin nearly entire except
for a delicate fringe; they are more or less
completely and closely covered by fine, slen¬
der, minute, dark spines; at the posterior mar¬
gin there are a few larger, long, dark-brown,
pendant-like papillae. The prostomium has broad
peaks; the four eyes are large, the anterior pair
is lateral in position, just in front of the middle
half of the prostomial length. Neurosetae have
a long main tooth and a small, short subterminal
secondary tooth.

This species was originally described off
southern California in 38 to 238 fm.; it has
since been reported from British Columbia, in

14

PACIFIC SCIENCE, Vol. II, January, 1948

15 to 20 fm. (Berkeley, 1923) and 555 meters
(Berkeley, 1942). The present records are from
southern Alaska and Bering Sea, in 1 3 to 68 fm.

Genus Eunoe Malmgren
Eunoe depressa Moore
Eunoe depressa Moore, 1905^: 5 3 6-5 38, pi.
34, fig. 17, 18, pi. 35, fig. 19, 20.

Collections. Stations 33-40 (1); 34-40 (6);
58-40 (1); 61-40 (2); 70-40 (1); 80-40
(3); 97-40 (1); 98-40 (1); 103-40 (1);
BT 70-41 (1); C 147-41 (1); C 150-41 (1);
D 7-41 (4); D 15-41 (1); D 16-41 (1).

The size of these individuals varies greatly,
ranging from a few mm. long (juvenile) to
comparatively gigantic; a large one measures
68 mm. long, without the everted proboscis,
which is 28 mm. long; width is 25 mm. with¬
out, 35 mm. with, parapodia. Elytra are thick,
leathery in texture and have entire margin; they
are pale but are mottled with rust-colored pig¬
ment. Their surface is covered with low, flat¬
tened nodules of varying sizes, and a few larger
ones are more or less limited to the posterior
half of the elytral surface. The prostomium has
four dark eyes; the anterior pair is lateral, near
the middle of the prostomial length.

Some collections have data recording com¬
mensalism with hermit crabs; a small, probably
juvenile, individual from station D 16-41 comes
from the body cavity of a large Alaskan king
crab, Paralithodes camtschatica (Tilesius). The
large body size of some adults is perhaps corre¬
lated with a commensalistic habit.

This species has remained unknown except
through its original account, based on collections
from Alaska, in 8 to 19 fm.; some of these were
labeled hermit crab messmates (Moore, 1905:
538). The present records are from south¬
western Alaska in 5 to 150 fm., and from
Bering Sea in 15 to 60 fm.

Genus Gattyana McIntosh
Gattyana cirrosa (Pallas)

Aphrodita cinhosa Pallas, 1766: 95-97, pi. 8,
fig. 3-6.

Gattyana cirrosa Moore, 1908: 337; Fauvel,
1923: 49-50, fig. 17; Berkeley, 1942: 187.

Collections. Stations 33-40 (4); 35-40 (2);
51-40 (1); 61-40 (1); 128-40 (4); 131-40
(1); Mist Harbor (1); Pavlof Bay (1).

This species has been widely reported from
the colder waters of the Northern Hemisphere,
including both sides of North America. The
present records are from southwestern Alaska in
15 to 50 fm.

Gattyana iphionelloides (Johnson)
Harmothoe iphionelloides Johnson, 1901: 391—
392, pi. 1, fig. 2-7.

Gattyana iphionelloides Berkeley, 1945: 321.

Collection. Station 106-40 (1).

This species is known from Washington
(Johnson), western Canada (Berkeley), and a
reef in Alitak Bay, Alaska; it is intertidal.

Genus Hermadion Kinberg

Hermadion truncata (Moore)

Harmothoe ( Eunoa ) truncata Moore, 1902:

272-274, pi. 14, fig. 21-28.

Hermadion truncata Moore, 1908: 332-333;
Berkeley, 1923: 215; Berkeley, 1945: 323.

Collection. Station 91-40 (1).

Originally the type locality was given as
northern Greenland (Moore, 1902) but was
later (Moore, 1908) corrected to Icy Cape,
Alaska. Berkeley (1923) has recorded the spe¬
cies from British Columbia. The present record
is from Baralof Bay, in 24 fm.

Family POLYODONTIDAE

Genus Peisidice Johnson

Peisidice aspera Johnson
Peisidice aspera Johnson, 1897: 184-185, pi.
9, fig. 56-59, pi. 10, fig. 63; Moore, 1908:
338; Berkeley, 1923: 216; Berkeley, 1942:
189.

Annelids of Alaska — HARTMAN

15

Collections. Stations 20-40 to 22-40 (1);
60-40 (2); 61-40 (2); 70-40 (6).

This species is known to range widely through
the northeastern Pacific The present records are
from southwestern Alaska, in 15 to 40 fm.

Family SIGALIONIDAE

Genus Pholoe Johnston

Pholoe minuta (Fabricius)

Aphrodita minuta Fabricius, 1780: 314.
Pholoe minuta Moore, 1908: 338; Fauvel,

1923: 121-122, fig. 44; Berkeley, 1942: 189.

Collections. Stations 5 1-40 ( 1 ) ; 60-40 ( 1 ) .

The present records are within the known
range; they come from Canoe Bay and Leonard
Harbor, in 20 to 40 fm.

Family CHRYSOPETALIDAE

Genus PALEANOTUS Schmarda

Paleanotus chrysolepis Schmarda
Paleanotus chrysolepis Schmarda, 1861: 163,

pi. 37, fig. 326-329; Berkeley, 1942: 27.
Heteropale hellis Johnson, 1897: 163-164, pi.

6, fig. 20-23; Berkeley, 1923: 212.

Collection. Station 60-40 (2).

This record is in the known range, from
southwestern Alaska in 15 to 40 fm.

Family SPINTHERIDAE

This family is known through a single genus,
Spinther Johnston. The body is broad, flat, and
sole-like. The entire dorsum is covered with
transversely prolonged notopodial ridges; these
are continued around the front to encompass
the prostomial parts so that those of the two
sides merge at the middle front. The ventral
side of the body is papillated or smooth. The
prostomium is a tiny, inconspicuous lobe, with
or without eyes; it is set some distance back,

between the notopodial ridges and immediately
over the ventral mouth. There is a small, con¬
ical or subspherical, median prostomial antenna
that largely covers the prostomium. The pro¬
boscis is a voluminous, unarmed, rosette-like,
eversible organ.

Parapodia are biramous. Notopodia are long,
transversely arranged, dorsal ridges; they are
provided with many spine-like setae that are
arranged in transverse rows; they have entire or
bifid tips. Neuropodia are long, lateral exten¬
sions of the body wall; they may or may not
have a distal extension (called a cirrus); they
are armed with one to several, strong, falcate,
composite, yellow hooks that are encased in an
embedded bundle of slender acicula (Fig. 1 c).
The anal aperture is dorsal, near the posterior
end of the body.

In so far as known, all species occur on the
surface of sponges, in shallow to moderate
depths.

Genus Spinther Johnston

Spinther Johnston, 1845
Oniscosoma Sars, 1851
Cryptonota Stimpson, 1854

Type S. oniscoides Johnston

Three species, S. oniscoides Johnston, S. mini-
aceus Grube, and S. arcticus Wiren, were recog¬
nized and described in von Graff’s revision
( 1888), but the first of these with some reserve
because of the ambiguity surrounding John¬
ston’s description. Riddell (1909: 101-108)
clarified this doubt, after a study of topotypes of
S. oniscoides, and showed that von Graff’s first
species is actually S. citrina (Stimpson), and
that S. oniscoides Johnston, constitutes a fourth
species. Since then, a fifth species, 5'. australi-
ensis Augener (1913), has been described.

Confusion still prevails (see Fauvel, 1923:
140) concerning the specific names of von
Graff, especially since the rules of nomencla¬
ture were not applied in the choice of acceptable
names. Thus, although von Graff showed con¬
clusively that Oniscosoma arcticum Sars ( 1851 )

16

PACIFIC SCIENCE, Vol. II, January, 1948

is the same as Spinther miniaceus Grube
(I860), he retained the second name since it
was deemed undesirable to retain the first for
a form that may be southern in distribution.
However, a specific name may not be rejected for
inappropriateness (Article 32, Int. Rules Zodl.
Nomen. ) . Furthermore, the name Spinther
arcticus Wiren ( 1883 ) was retained for another
species which differs from the older S. arcticus
(Sars). Again, the rules of nomenclature dic¬
tate that a specific name is to be rejected as a
homonym when it has been previously used for
another species of the same genus (Article 35).
S. arcticus (Sars) therefore has priority over
S. miniaceus Grube, and the latter becomes a
synonym of the former. S. arcticus Wiren is
a homonym and to be rejected. I propose
S. wireni, new name, for the latter species.

The name S. major Levinsen (1883) was
proposed to replace S. arcticus Hansen (1882)
(not Sars nor Wiren). It was shown by
von Graff (1888) to be the same as S. onis-
coides [= S. citrina (Stimpson)].

The known species of Spinther are thus:

( 1 ) S. arcticus ( Sars ) , which includes S. mini¬
aceus Grube, from western and southern Europe;

(2) 5'. australiensis Augener, from southwestern
Australia; (3) S. citrina (Stimpson), which

includes 5. oniscoides von Graff (not Johnston),
from eastern Canada and the New England
states; (4) 5. oniscoides Johnston, from Ireland;
and (5) 5. wireni, new name, wdiich includes
5. arcticus Wiren, not Sars, nor Hansen, from
Bering Sea. A sixth species, 5. alaskensis, is
newly described below.

Key to Species of Spinther

1. Ventrum papillate..... . 2

1. Ventrum smooth or only wrinkled . 4

2. Distal end of neurosetal shaft crenulate;
notosetae largely entire at tip, a few

bifid . . . S. citrina (Stimpson)

2. Distal end of neurosetal shaft trilobate;
notosetae entire except for a very few,

very slender, bifid ones...... . .

. . . S. alaskensis new species

2. Distal end of neurosetal shaft entire . . 3

3. Notosetae bifid only.. ..5. oniscoid.es Johnston

3. Notosetae bifid and entire, in equal num¬

ber and about equally thick.... .

. . . . . 5. wireni new name

4. Neurosetae with a large lateral tooth .

. . . S. australiensis Augener

4. Neurosetae without a lateral tooth . .

. . . . . . .5. arcticus (Sars)

Chart Showing Comparison of Characters for Species of Spinther

Name of
Species

Parapodial

Cirri

Neurosetae

Number

of

Segments

Length

IN MM.

Ventrum

Shape

Distal end

OF SHAFT

Notosetae

S. oniscoides
Johnston

papillate

present

falcate

smooth

smooth

bifid only
(Riddel,

1909: 103)

20-25

4-13

S. citrina
(Stimpson)

papillate

present

falcate

smooth

crenulate

largely entire,
a few bifid, the 2
equally thick
(Graff, 1888: 17)

30-48

11-26

S. wireni
new name

papillate

present

falcate

smooth

smooth

bifid and entire in
equal number and
equally thick

43-52

20-25

S. alaskensis
new species

papillate

present

falcate

smooth

trilobed

entire except for
very few, very
slender bifid ones

46-47

ca. 28

S. arcticus
(Sars)

smooth

absent

falcate

smooth

entire

bifid only

12-24

1-9

S. australiensis
Augener

smooth

absent

falcate,

lateral

tooth

? entire

bifid only

15-31

4. 5-7.5

Annelids of Alaska — HARTMAN

17

Spinther alaskensis new species
Fig. 1 a-c, 2 g-j.

Collections. Stations 5 1-40 ( 1 ) ; 66-40 (2).

The largest individual measures 28 mm. long
and 20 mm. wide. Number of segments is 46
or 47. The first two segments, preceding the
prostomium, and the last six or seven, following
the pygidium, are short and incomplete mid-
dorsally. The dorsum is slightly arched and
nearly uniformly covered with notopodial ridges
except for a narrow, median, longitudinal stripe
(Fig. 1 a). The ventrum is flat and solelike;
it is more or less completely covered with seg-
mentally arranged rows of spherical papillae,
strewn thickly over the neuropodial bases and
less so medially; the segmental intervals are
smooth. A broad, median region has similar
papillae but much sparser and irregularly dis¬
persed.

The prostomial appendage is a spherical
(Fig. la) (station 66-40) or elongate (station
51-40) papillar lobe. The prostomium is low
and has two pairs of large, oval, reddish-brown
eyespots, located in the groove where the lobe
and body join; the anterior eyes are the larger
but the two of a side nearly merge with each
other.

The notopodial ridges ( dorsal lamellae ) have
wavy fore and hind margins; the waves cor¬
respond in their distribution with that of the
setal fascicles, the concavity lacks setae, the con¬
vexity has them. The anterior and posterior
margins of the lamellae are about equally de¬
veloped. The first two pairs of notopodia, in
front of the prostomium, are directed forward
(Fig. l a); the third pair is in line with the
prostomium and farther back they are lateral in
position. The most posterior are directed back
so as to surround the anal region. The noto¬
podial setae are arranged in alternating series
(Fig. 1 b) along the membranous margins of
the ridges. In median parapodia, where they
have their maximum development, there are
about 16 sets of setal fascicles in a notopodium;
they consist of eight sets in the anterior margin

and alternate with the same number of sets in
the posterior margin. Each set consists of a
fan-shaped fascicle of about nine setae, but
some setae are embedded and visible only by dis¬
section. The total number of setae in a ridge
approximates 140 to 200. The symmetry of this
pattern on a well-preserved individual is a
striking feature and may signify a unique struc¬
tural character in the functioning of the in¬
dividual, perhaps to aid the flow of water
forward and back.

Notopodial setae are of two kinds (Fig. 2
h-j). They consist of thick, bluntly pointed,
straight, acicular spines numbering eight to
ten in each fan-shaped fascicle and a few, very
slender, delicate rods that are slightly curved and
distally bifid (Fig. 2 i) . The thickness of the
latter is only one-fourth to one-sixth that of the
larger spines; they are thus easily overlooked.

Neuropodia are long and thick; they taper
distally and are provided with a thick, conical,
superior lobe (Fig. 1 c) that is somewhat post-
setal in position. Their bases are strongly
wrinkled as though capable of great lateral ex¬
tension, and the ventral side is covered with
spherical papillae that resemble those on the
ventral side of the body. One, seldom two, large
composite, yellow hooks project from the distal
end of the neuropodium. These large hooks are
flat and knifelike; their appendage is strongly
curved. The distal end of the shaft, at its longest
part, is weakly trilobed (Fig. 2 g). The em¬
bedded part of the setal bundle, by dissection,
is seen to consist of several developing, com¬
posite hooks, in various stages of growth; they
are surrounded by an enveloping acicular
bundle. The acicula are long, slender, tapering,
pale rods (Fig. 1 c).

The proboscis, partly everted in one in¬
dividual, but observed also by dissection, is a
soft, voluminous, rosette-shaped sack, as is
typical of the genus.

S. alaskensis belongs to the group of species
in which the ventrum is papillose. It has noto¬
podial setae with tips that are both entire and
bifid. To this group belong also S. citrina

18

PACIFIC SCIENCE, Vol. II, January, 1948

(Stimpson) and S. wireni, mihi. In the latter,
the bifid and entire notosetae are about equal in
number and equally thick; in the former they
are also about equally thick. In S. alaskensis,
the bifid setae are not only very few in number
but are much more slender and very incon¬
spicuous. S. alaskensis attains greater size than
other species of the genus, but this character,
as also the approximate segmental count, may
have little significance (see chart on p. 16).

Holotype in the U. S. National Museum.

Type locality. Canoe Bay, Alaska, in 25 to
40 fm.

Distribution. Southern Alaska.

Family PHYLLODOCIDAE

Genus Notophyllum Oersted

Notophyllum imbricatum Moore
Fig. 3 a-c.

Notophyllum imbricatum Moore, 1906^: 217-

219, pi. 10, fig. 1-3; Moore, 1908: 329;

Berkeley, 1924: 287; Berkeley, 1942: 190.

Collection. Station 12-40 (2).

Dorsal cirri are thick, deeply and broadly im¬
bricated, and uniformly drab green; they are
slightly narrowed distally; sixteenth and post¬
median ones are shown in Figure 3 c, b. Neu¬

ropodia taper distally and are notched at the
end of the acicular lobe (Fig. 3 a). There are
two pairs of nuchal lappets or there may be an
additional tiny pair on the inner side.

This is known only from the northeast Pa¬
cific; the present record is from the shore of
Canoe Bay.

Notophyllum foliosum (Sars)

Fig. 3 d-f.

Phyllodoce foliosa Sars, 1835: 60-61, pi. 9,
fig. 26.

Notophyllum foliosum Fauvel, 1923: 170-171,
fig. 16.

Collections. Station L 18-41 (1); Sand Point

(1).

This is smaller than N. imbricatum ( above ) .
There is a single pair of broad nuchal lappets.
Dorsal cirri are imbricated; they are suffused
and streaked with dark pigment, most intense
at the outer periphery. Dorsal cirri are more
broadly rounded distally than in N. imbricatum;
sixteenth and postmedian ones are shown in
Figure 3 f, e. Posterior neuropodia lack a deep
incision (Fig. 3 d).

This has remained unrecorded from the
Western Hemisphere. The present records are
from Bare Island in 13 to 15 fm, and from
Sand Point.

Fig. 3. Species of Notophyllum. a-c, Notophyllum imbricatum: a, postmedian neuropodium in anterior
view, setae omitted, X 75; b, postmedian dorsal cirrus, X 15; c, sixteenth dorsal cirrus, X 15. d-f, Notophyl¬
lum foliosum: d, postmedian neuropodium in anterior view, setae omitted, X 60; b, postmedian dorsal cirrus,
X 15; /, sixteenth dorsal cirrus, X 15.

Annelids of Alaska — HARTMAN

19

FIG. 4. Eteone calif ornica (station 47-40) : a, anterior end in dorsal view, enlarged (the prostomial eyes are
deep seated) ; b, ninth parapodium in posterior view, X 99; c, far posterior parapodium in posterior view, X 99;
d, anal cirri, enlarged.

Genus ANAITIDES Czerniawsky

Key to Species of Anaitides

1. Ventral cirri taper distally to a sharp

point . A. mucosa

1. Ventral cirri distally blunt or rounded.... 2

2. Proboscis with six rows of papillae on a

side, with 12 or 13 in each row .

. . . . . A . groenlandica

2. Proboscis with only four rows of papillae
on a side and about four papillae in
each row . . . A. citrina

Anaitides groenlandica (Oersted)
Phyllodoce groenlandica Oersted, 1843: 192—
193, fig. 19, 20, 22, 29-32.

Anaitides groenlandica Bergstrom, 1914: 141-
143, fig. 41; Berkeley, 1924: 287; Berkeley,
1942: 189.

Collections. Stations 9-40 (1); 51-40 (5);
D 14-41 (14); Canoe or Pavlof Bay (1).

This is a large, robust species. The proboscis
has six rows of papillae on a side, with 12 or
13 in each row, at the middle of the series.
Dorsal cirri are large and the distal ends are
truncate.

Originally described from Greenland, this has

been reported from both sides of northern
North America. The present individuals were
dredged from Canoe Bay and the Bering Sea.

Anaitides mucosa (Oersted)
Phyllodoce mucosa Oersted, 1843: 31, fig. 25,
79, 83, 89; Moore, 1908: 328.

Anaitides mucosa Bergstrom, 1914: 143-144,
fig. 43.

Phyllodoce citrina Berkeley, 1924: 287.
Phyllodoce ( Anaitides ) mucosa Berkeley, 1945:
324.

Collection . Station 131-40 (1).

Dorsal cirri are trapezoidal in shape; ventral
cirri are broad basally, acutely pointed distally.
This species has been recorded previously from
Alaska (Moore, 1908: 328). The present record
is off Cape Chiniak, in 32 to 35 fm.

Anaitides citrina (Malmgren)
Phyllodoce citrina Malmgren, 1866: 95-96, pi.
13, fig. 24; Moore, 1908: 328; Fauvel, 1923:
150, fig. 52.

Anaitides citrina Bergstrom, 1914: 140-141,
fig. 41.

Phyllodoce ( Anaitides ) citrina Berkeley, 1945:
324.

20

PACIFIC SCIENCE, Vol. II, January, 1948

Collections. Stations 12-40 (1); 35-40 (1,
juvenile).

The proboscis has only four rows of papillae
on a side, with about four papillae in each row.
This species has been recorded from Afognak
Island, Alaska (Moore, 1908), and western
Canada (Berkeley, 1945); the present records
are from the vicinity of Pavlof Bay.

Genus Eulalia Savigny

Eulalia viridis (Muller)

Fig. 5 a.

Nereis viridis Muller, 1771: 156.

Eulalia viridis Fauvel, 1923: 1 60, fig. 57; Berke¬
ley, 1924: 288; Berkeley, 1942: 189.

Collections. Stations 21-40 (1); 20-40 to
22-40 (4); 24-40 (1); 47-40 (1); 51-40
(3).

Fig. 5. Species of Eulalia and Eteone. a, Eulalia
viridis (station 24-40) : 118th or twenty-second last
parapodium in anterior view showing long dorsal cir¬
rus, X 61. b, Eteone spetsbergensis (station D 14-
41) : postmedian parapodium, X 48.

The number of segments is about 140, length
is 32 mm. The proboscis is closely covered with
papillae. Dorsal cirri are considerably prolonged,
fully three to four times as long as wide (Fig.
5 a); ventral cirri extend distally beyond the
setal lobes.

This cosmopolitan species has been reported
previously from the northeast Pacific (Berkeley,
1924 and 1942). The present records are from
southwestern Alaska, taken in shallow dredging.

Genus Eteone Savigny

Eteone spetsbergensis Malmgren
Fig. 5 b.

Eteone spetsbergensis Malmgren, 1866: 102,
pi. 15, fig. 38; Bergstrom, 1914: 202-204,
fig. 77; Berkeley, 1945: 325.

Collection. D 14-41 (1).

A single, large, robust individual, 60 mm.
long, with eggs, has the 15 last segments re¬
generated. The dorsum is pale and has a broad,
longitudinal, reddish-brown stripe on either
side, most intense in the anterior half; the pig¬
ment does not cover the dorsal or ventral cirri.
Throughout the body the dorsal cirri are broader
than long and are asymmetrical; the ventral
cirri are distally blunt (Fig. 5 b shows a para¬
podium from a postmedian segment).

This Arctic species has been reported from
the Bering Sea by Bergstrom (1914), and from
western Canada by Berkeley (1945); the pres¬
ent collection comes from the Bering Sea, in
36 fm.

Eteone californica Hartman
Fig. 4 a-d.

Eteone californica Hartman, 1936^: 131, fig.
49-51.

Collections. Stations 47-40 (4); 60-40 (1);
108-40 (7); Dolgai Harbor (3).

The color (preserved) is dull green, most
intense on dorsal and ventral cirri. These in¬
dividuals are fully four to five times as large as
some from the type locality (San Francisco
Bay, California ) . The prostomium has two em¬
bedded, dark eyes, and a tiny nuchal papilla at
its posterior margin (Fig. 4 a). The proboscis
terminates distally in 14 soft, subglobular papil¬
lae; its proximal portion is smooth. The first
setigerous segment is smaller than those follow¬
ing and it lacks dorsal cirri. In the anterior
region the ventral cirri extend distally about as
far as the setigerous lobe (Fig. 4 b), but by the
twenty-fourth segment the ventral cirri are
shorter and come to be inconspicuous (Fig. 4

Annelids of Alaska — HARTMAN

21

c) in postmedian segments. The anal ring has
a pair of short, blunt processes (Fig. Ad).

This species is known from central California,
shore, and western Mexico (Rioja, 1941: 687);
the range is hereby extended to southwestern
Alaska, from shore to 25 fm.

Family SYLLIDAE

Genus Typosyllis Langerhans

Key to Species of Typosyllis

1. Prostomium partly covered by a nuchal

collar (Fig. 6 a) . . . T. collaris

1. Prostomium without a nuchal collar . 2

2. Setae distally bidentate... . . . . . 3

2. Setae distally entire......... . . . . . 5

3. Anterior dorsum with a pigment pattern
consisting of broken transverse lines. .

. . . . . J T . armillaris

3. Dorsum without pigment pattern (pre¬

served) . . . 4

4. Superior setae in anterior segments with

long appendage; articulation of setae
complete.. . : . . . T. alternate*

4. Superior setae in anterior segments

with shorter appendage; articulation
of setae incomplete . T. elongata

5. Larger, 100 mm. long or longer; dorsal

cirri short . T. stewarti

5. Smaller, about 30 mm. long; dorsal cirri

long . . . . . T. pulchra

Typosyllis alternata (Moore)
new combination

Syllis alternata Moore, 1908: 323-325, fig. a-f;
Berkeley, 1923: 206; Berkeley, 1938: 37-38.

Collections. 35-40 (1); Lazy Bay (4).
Setae are entirely composite; this species is
therefore referred to the genus Typosyllis. There
are four atokous individuals, and a female
epitokous stolon from Lazy Bay. Antennae and
dorsal cirri are clearly articulate throughout. In
the anterior region, dorsal cirri are about equally
long, but thereafter they alternate long and

short, the number of articles ranging from 25
to 18. Ventral cirri are fairly long throughout.
Setae are of a single kind, distally bidentate, but
those in the superior part of the fascicle, espe¬
cially in anterior segments, tend to have long
appendages. Superior and inferior setae from a
postmedian segment are similar to anterior
setae but somewhat thicker and have a shorter
appendage. Acicula are yellow and distally
knobbed.

A female epitokous individual consists of head
and 26 setigerous segments; swimming setae
are present on all except the first segment. The
prostomium is bilobed and has four reddish
eyes. The anterior ones are much the larger
and oblong in shape; they are antero-ventral in
position and have elongate lenses; the posterior
eyes are circular in shape.

Syllis harti Berkeley (1941: 36) from British
Columbia, Canada, also appears to be a Typo¬
syllis, since it is provided with only composite
setae. It bears resemblance to T. alternata, but
in the former the ventral cirri are even longer
than in the latter, and the dorsal cirri have 30
to 40 articles each.

T. alternata was originally described from
Alaska and has been reported from the north¬
east Pacific, south to southern California (Moore,
1923: 256) and from western Mexico (Rioja,
1941: 691). The present specimens are from
southwestern Alaska.

Typosyllis elongata (Johnson)
new combination

Pionosyllis elongata Johnson, 1901: 403-405,

pi. 6, fig. 67-70, pi. 7, fig. 71.

Syllis elongata Berkeley, 1923: 206; Berkeley,

1938: 41; Berkeley, 1942: 190; Rioja, 1941:

688.

Collections. Stations 12-40 ( 1 ) ; 20-40 to
22-40 (1); 24-40 (1); 51-40 (5); 108-40
(4); D 11-41 (1).

This was originally described in the genus
Pionosyllis because the palpi are partly fused at
their bases. Berkeley (1938: 38) has ques¬
tioned the value of this character and I agree

22

with her conclusion. Since all setae are com¬
posite (or presumably so), and dorsal cirri are
articulate, it is herewith referred to Typosyllis.
Setae are few in parapodia. Anteriorly there are
about 12 in a parapodium; the dorsal one to
three often lack their appendage, thus resem¬
bling simple setae, but this lack is believed to
be due to loss through wear. Posteriorly each
parapodium has only about six setae, and the
dorsal ones similarly lack a distal appendage.
In anterior segments the superior setae are
slender and have longer appendages than those
in the inferior part of the fascicle, or those in
median and posterior segments. Their tips are
bidentate, with an accessory tooth that is very
slender and long; when worn, such setae appear
to have an entire tip. The cutting edge has a
row of fine spines; the outer side of the shaft
has a few fine spines at its thickened portion.
True ypsiloid (simple) setae are lacking (see
also Berkeley, 1938: 41). In far posterior seg¬
ments, the setae are similar to those in median
segments but more slender. In all setae the
articulation tends to be incomplete.

T. elongata differs from T. alternata (see
above) in that the setal articulation is incom¬
plete in the former, complete in the latter; the
appendage of anterior setae is shorter in the
first than in the second; the superior seta
often resembles an ypsiloid one in the first,
whereas it retains its appendage in the second.
T. elongata was originally described from Wash¬
ington south to California but has since been
reported from various parts of the northeast
Pacific (see synonymy above). The present
records are from southern Alaska to the Pribilof
Islands in the Bering Sea, from shore to 125 fm.

Typosyllis armillaris (Muller)

Nereis armillaris Muller, 1776: 2626.

Syllis armillaris Moore, 1908: 323; Berkeley,
1923: 206.

Syllis ( Typosyllis ) armillaris Fauvel, 1923:
264-265, fig. 99.

Collections. Stations 60-40 (5); 61-40
(10); 70-40 (4); L 18-41 (5).

PACIFIC SCIENCE, Vol. II, January, 1948

Originally described from Greenland, this
species has been reported from cosmopolitan
areas, especially in boreal and arctic seas. The
present records are from Leonard Harbor in
20 to 25 fm., Cold Bay in 15 to 35 fm., and
northwest side of Bare Island, in 13 to 15 fm.

Typosyllis pulchra (Berkeley)

Syllis pulchra Berkeley, 1938: 34-35, fig. 1.
Typosyllis pulchra Hartman, 1944: 250.

Collection. Station 108-40 (3).

This species is pigmented chocolate brown
dorsally. Composite setae have entire tips.
Originally described from western Canada, it
has been found in central California ( Hartman,
1944: 250). The present record is Alitak Bay,
shore.

Typosyllis stewarti (Berkeley)
new combination

Syllis stewarti Berkeley, 1942: 191.

Collection. Station 108-40 (1).

A single large, much-coiled individual has a
strongly arched dorsum; it is chocolate brown
above, pale below; cirri and antennae are also
pale. Antennae and dorsal cirri are short but
distinctly moniliform. The median prostomial
antenna is inserted far back at the posterior
margin of the prostomium. Parapodia are in¬
conspicuous though fleshy; there are about five
yellow acicula in each; they terminate distally
in a blunt tip. Setae are composite and of a
single kind; therefore, the species is considered
a Typosyllis. The appendage is short and fal-
cigerous; it has a smooth tip and the cutting
edge has a few long spines. The uppermost ones
resemble those below. Anterior setae resemble
those in posterior and median segments but
those in the middle of the body are thicker and
larger than those elsewhere.

T. stewarti has heretofore been known only
through a single individual from Vancouver
Island, Canada (Berkeley, 1942); the present
record is Alitak Bay, shore.

Annelids of Alaska — Hartman

23

Typosyllis collaris new species
Fig. 6 a-c.

Collection. Station D 8-41 (4).

Several small individuals are colorless except
for the four red eyes; the largest one measures
12 mm. long for 45 segments but is posteriorly
incomplete. The body is short and plump. The
prostomium is broader than long; it has a long
median, and two shorter lateral, antennae and
the four eyes are in trapezoidal arrangement
(Fig. 6 a); the posterior portion is somewhat
overlain by a nuchal lobe (hence the specific
name) which extends forward to the posterior
margin of the posterior eyes. The prostomial
antennae are articulate, most distinctly so in
their distal halves and decreasingly so to near
their bases; the median one is more than twice
as long as the paired ones (Fig. 6 a). The palpi
are free from one another except at their bases;
they are broadly subrectangular and about as
wide as long.

The proboscis terminates distally in 10 soft,
widely spaced papillae; on its dorsal side at the
anterior end there is a conspicuous semitranslu-
cent greenish tooth, equitriangular in shape.

The proportions of prostomial antennae, tenta¬
cular cirri, and anterior dorsal cirri are shown
in Figure 6 a.

The parapodia are short and plump but taper
slightly distally. The anterior ones resemble the
posterior, except that setae tend to be more
numerous in front. Dorsal cirri are articled in
their distal halves but are more or less smooth
at the base. In anterior parapodia the postsetal
lobe is slightly prolonged but the setal lobe is
blunt. In median and postmedian parapodia
( Fig. 6 b) the presetal lobe is somewhat drawn
out and shorter than the postsetal one.

Setae are entirely composite and resemble
one another throughout. The superior and in¬
ferior ones in any fascicle, as also those in
anterior and posterior segments, are similar to
one another, but the appendage of the first is
slightly longer than that in the last. They num¬
ber 25 to 30 in anterior segments and decrease
to 15 to 18 in posterior segments. The shaft is
distally spinous; the appendage is short, boldly
bidentate at its free end and there are a few
long spines along the cutting edge (Fig. 6 c).
Acicula are pale yellow, slightly knobbed dis¬
tally, and number usually two in a parapodium.

Fig. 6. Typosyllis collaris new species: a, anterior end in dorsal view, enlarged; b, postmedian parapodium
dn anterior view, only some of the 18 setae shown, X 89; c, seta from a median parapodium, X 956.

24

Anal cirri are long, slender though coiled, and
closely articled.

T. collaris is unique among species of the
genus Ty posy llis in having a nuchal collar;
tentacular and dorsal cirri are articulate in their
distal halves but tend to be smooth at the base;
setae have a spinous shaft and a bidentate ap¬
pendage; ventral cirri are short throughout. In
some respects this recalls SyMs cucullata Mc¬
Intosh (1908: 191) from the Isle of Wight,
which also has a nuchal collar and short com¬
posite setae but in this the setae are distally
entire and the nuchal collar is not so wide as in
T. collaris. Also, the prostomial antennae are
much longer. Since setae are presumably en¬
tirely composite in S. cucullata, it is perhaps
also to be referred to the genus Typosyllis.
Holotype in the U. S. National Museum.
Type locality. Bering Sea in 42 fm.
Distribution. Alaska.

Genus Trypanosyllis Claparede
Trypanosyllis gemmipara Johnson

Trypanosyllis gemmipara Johnson, 1901: 405-
406, pi. 7, fig. 72-76; Johnson, 1902: 302-
315, fig. 7-17; Moore, 1908: 328; Berkeley,
1923: 207; Berkeley, 1938: 42; Berkeley,
1942: 191.

Collection. Station 82-40 (1).

The last 10 or more segments are very short,
taper strongly distally, and are immediately
preceded, on the ventral side, by a thick cluster
of 30 or more buds of varying sizes. This
species has been reported from the northeast
Pacific. The present record is from near Big
Koniuji Island in 25 to 30 fm.

Genus Autolytus Grube
Autolytus magnus Berkeley
Autolytus magnus Berkeley, 1923: 210, pi. 1,
fig. 3, 4; Berkeley, 1938: 47; Berkeley, 1945:
318.

Collection. Station 20-40 to 22-40 (1).
This is a single, atokous, much-coiled in-

PACIFIC SCIENCE, Vol. II, January, 1948

dividual. The nuchal epaulettes are long, sin¬
uous lappets that extend from the posterior
margin of the prostomium and diverge outward,
following the inner bases of parapodia; they
extend back through the fifth setigerous seg¬
ment. The prostomium is trapezoidal, widest
anteriorly, and longer than wide; it has four
red eyes near the ectal margins. The three an¬
tennae are thick, long, and wrinkled but not
articulated; the median one exceeds the paired
ones in length and all are longer than the peris-
tomial cirri. The anterior dorsal cirri are only
about half as long as the paired prostomial
antennae. Palpi are fused medially and only
about half as long as the prostomium when
seen from the dorsum.

Parapodial lobes are thick, the setae disposed
in close, thick fascicles in anterior and median
segments but diminish in number farther back.
Anterior parapodia have five or six acicula in
each; median parapodia have about four each;
they are yellow, taper distally and terminate in
acute points. Setae are entirely composite, the
shaft distally spinous, the appendage bidentate
with strong, secondary tooth.

A. magnus has been described through the
Sacconereis ( = epitoke female) (Berkeley,
1923) and the Polybostrichus (= epitoke
male) (Berkeley, 1938, 1945) stages, both
from British Columbia. The present atokous
form is from Canoe Bay, in 15 to 40 fm.

Family NEPHTYIDAE

Genus Nephtys Cuvier

Nephtys caeca (Fabricius)

Nereis caeca Fabricius, 1780: 304-305.
Nephtys caeca Johnson, 1901: 401-402; Moore,
1908: 341; Berkeley, 1924: 290; Berkeley,
1942: 192.

Collections. Stations 35-40 (1); 51-40 (1);
A 61-41 (7); Lazy Bay (10).

Some individuals measure nearly 8 inches
long (preserved). Branchiae are not present

Annelids of Alaska — HARTMAN

25

before the fifth or sixth segments and the first
few pairs are small; the last four or more seg¬
ments are abranchiate, but the last 10 or more
pairs of branchiae are small and papillar. The
proboscis has 22 rows of papillae at its distal
end; the proximal surface (on the everted pro¬
boscis) is covered with low, wartlike elevations.

N. caeca is a cold-water species, common in
the northeast Pacific, rarely occurring south
to central California (Hartman, 1938: 148).
The present collections are from southwestern
Alaska, shore to 40 fm.

Family NEREIDAE

Genus Neanthes Kinberg

Neanthes brandti (Malmgren)

Alitta brandti Malmgren, 1866: 183.

Nereis virens Johnson, 1901: 398, pi. 3, fig.

26-30; Moore, 1908: 344.

Neanthes brandti Hartman, 19 44£: 252.

Collection. Lazy Bay, off Alitak Bay (5).
This species is distinguished from N. virens
(Sars) chiefly in the greater dentition of the
proboscidial armature. It occurs commonly
throughout the northeast Pacific, south to south¬
ern California. The present record is within
the known range.

Genus Cheilonereis Benham

Cheilonereis cyclurus (Harrington)

Nereis cyclurus Harrington, 1897: 210-220,
pi. 16, fig. 1-3, pi. 17, fig. 1-7, pi. 18, fig.
1-5; Johnson, 1901: 400, pi. 4, fig. 46, pi. 5,
fig. 48-52; Moore, 1908: 343-344; Berkeley,
1924: 292.

Cheilonereis cyclurus Hartman, 1940: 219.
Collection. Station 107-40 (1).

The single individual comes from Alitak Bay,
in 30 fm. It has previously been reported from
Alaska (see synonymy above).

Genus Nereis Linnaeus
Nereis procera Ehlers

Nereis procera Ehlers, 1868: 557-559, pi. 23,

fig. 2; Johnson, 1901: 400-401, pi. 4, fig.
47, pi. 5, fig. 53-59; Moore, 1908: 343;

Berkeley, 1924: 291-292; Berkeley, 1945:

326.

Collection. Station 24-40 (2).

This species has been recorded from various
parts of the northeast Pacific. The present col¬
lection comes from Canoe Bay, trawled from
125 fm.

Nereis zonata Malmgren

Nereis zonata Malmgren, 1867: 164, pi. 6, fig.

34; Fauvel, 1923: 338-339,“ fig. 130.

Collection. Station 51-40 (3).

Length of a complete individual is 70 mm.
for 73 setigerous segments. In one specimen
the body cavity contains large ova. The peris-
tomial tentacles are short, the longest reach back
to the anterior end of the first setigerous seg¬
ment; the shortest are about as long as the
prostomial antennae. Parapodial ligules do not
change greatly from anterior to posterior re¬
gions, but the dorsal portion changes gradually.
In postmedian and far posterior segments the
differences are most notable; here the dorsal
ligule is approximately quadrangular, the inser¬
tion of dorsal cirrus carried outward. Still far¬
ther back, the length of dorsal and ventral cirri
comes to be increasingly great. Anal cirri are
tapering and about as long as the last seven
setigerous segments.

Homogomph falcigerous notosetae are first
present from about the thirty-second segment;
they have a long appendage with denticulations
on one side. The proboscis is provided with a
pair of thick, dark-brown jaws with five blunt
teeth on the cutting edge. On the maxillary
ring, area I has two small cones in tandem; area
II has about 24 larger and smaller cones in
irregular crescentic arrangement; area III has
about 20 larger and smaller cones in an oval

26

PACIFIC SCIENCE, Vol. II, January, 1948

patch; area IV has about 25 cones in a crescent,
the largest ones on the side toward area III. On
the oral ring, area V has none, area VI has seven
or eight small circular cones; areas VII and VIII
(continuous) have seven larger pointed cones
in a row on the maxillary side and about six
to eight irregular rows of many tiny cones;
those on the oral side are gradually smaller. The
largest cones are on areas II and IV and those
on the maxillary side of VII.

The proportions of dorsal ligule to other
parts of the parapodium, especially in far pos¬
terior parapodia, differ somewhat from those
described (Malmgren, 1867) for individuals
from the type locality, in that in the present
case the dorsal ligule is relatively larger, but in
other respects there is agreement.

Originally described from Greenland, this
species has been reported from both sides of
the Atlantic. The present record is Canoe Bay,
25 to 40 fm.

Nereis vexillosa Grube
Nereis vexillosa Grube, 1851: 4-6, pi. 2, fig. 1,
5, 6; Johnson, 1901: 399, pL 3, fig. 31, 32,
pi. 4, fig. 33-38; Berkeley, 1924: 290-291.

Collections. Stations 3-40 (18); 47-40 (3);
108-40 (4); Dolgai Harbor (5); Lazy Bay,
off Alitak Bay (2); head of Lazy Bay (10).

This species is well known in the north
Pacific, from Alaska south to central California.
The present records are from shore stations in
southwestern Alaska.

Nereis pelagica Linnaeus
Nereis pelagica. Linnaeus, 1761: 508; Moore,
1908: 342; Fauvel, 1923: 336-337, fig. 130;
Berkeley, 1924: 291; Berkeley, 1942: 192.

Collections. Stations 12-40 (7); 31-40 (2);
33-40 (4); 34-40 (1); 51-40 (12); 52-40
(1); 61-40 (2); 70-40 (1); 108-40 (3);
128-40 (1); C 5-41 (2); D 15-41 (3);
Mitrofania Bay (2 male epitokes).

This species is apparently common through¬
out littoral areas of the northeast Pacific. In

some individuals the parapodial ligules are
opaque white, in others fuscous to dusky black,
as typical for N. neonigripes Hartman (1936).
In all, however, the proboscidial parts are much
alike and the proportions of parapodia are
similar. Homogomph falcigerous notosetae are
present from a premedian segment and are
continued posteriorly to the end. In the two
male epitokous individuals from Mitrofania
Bay, there are only 16, instead of 17, anterior
setigerous segments, and natatory parapodia are
continued posteriorly to the end. I am now
inclined to regard N. neonigripes Hartman
( 193 6b: 471-472) from California as a variety
of N. pelagica, if not merely a color phase. In
living individuals the dark parapodial ligules of
the latter are striking but they tend to fade out
in fixed specimens.

N. pelagica has been widely reported from
all seas.

Nereis neoneanthes new species
Fig. 7 a-d.

Collections. Station C 150-41 (1), western
Oregon (1).

A single incomplete individual was removed
from within a sabellid tube; it is over 100 mm.
long but the body is soft and slightly macerated.
There are over 131 segments; the body is long
and slender. The prostomium has four circular,
embedded eyes in the usual arrangement. The
paired palpi have long bases that project for¬
ward beyond the paired antennae; their pal-
podes are subspherical. The peristomial ten¬
tacles are short, simple, and tapering; the longest
reaches back to about the third setigerous seg¬
ment, the shortest is almost twice as long as a
prostomial antenna. The peristomial ring is
nearly twice as long as the first setigerous ring.

The proboscis is unique in having many
paragnaths. The oral ring has many rounded
paragnaths, all similar in appearance but those
on the maxillary side increase in size gradually.
The paragnaths cover most of the oral ring of
areas VII and VIII and are only slightly sepa¬
rated by a narrow space from those of paired.

Annelids of Alaska— HARTMAN

27

areas VI. Each area VI has six or eight similar
cones, and area V has three slightly larger cones
than those on area VI. The largest paragnaths
of the oral ring are those on the maxillary side
of area VI. On the maxillary ring, area I has a
single small cone, area II has a narrow crescent
of four or five small cones; III has an oval patch
of about 18 cones, including four larger ones
on the oral side and others of varying sizes; the
largest are about equal to those on the maxillary
side of area VII; area IV has a crescent of about
14 in approximately two rows intermediate in
size between those of III and II. Jaws are trans¬
lucent, dark, horny, brown, and thick; there
are 12 short, crenulate, slightly oblique teeth
along the cutting edge.

The first and second parapodia are uniaci-
cular and uniramous as typical of the genus.
From the third, slender dark acicula occur singly
in each lobe and are continued so throughout.
Anterior parapodia have nearly equal dorsal,
middle, and ventral ligules; there is no ligule
in the notoacicular portion; the neuroacicular
lobe is shorter than the parapodial ligules; the

dorsal cirrus is inserted at the base of the dorsal
ligule and extends distally about as far; the
ventral cirrus is shorter than the ventral lobe.

Parapodial ligules change little proceeding
back except for the dorsal one; already in the
anterior third of the body it increases in width,
surpassing the others in size and by the middle
it extends distally beyond the others. The inser¬
tion of the dorsal cirrus is gradually outward
and comes to be about midway on the dorsal
ligule (Fig. la).

Setae are of the usual kind. Notopodia are
provided with only homogomph spinigers in
anterior segments; from about segment 40 there
are one or two homogomph falcigers (Fig. 7
c) accompanying the spinigers and they finally
replace the latter. At their greatest develop¬
ment the falcigers number five or six in median
segments, but the number decreases further
back. Under low magnification these hooks ap¬
pear dusky. Neuropodia are provided with
homogomph spinigers superiorly, and with
heterogomph falcigers (Fig. 7 b) and hetero-
gomph spinigers below the aciculum.

FIG. 7. Nereis neoneanthes new species ( a-c from station C 150-41, d from Oregon) : a, postmedian
parapodium, X 44; b, neuropodial falcigerous hook from same parapodium, X 700; c, notopodial falcigerous
hook from same parapodium, X 700; d, far posterior parapodium, X 88.

28

PACIFIC SCIENCE, Vol. II, January, 1948

A more complete, well-preserved individual
is in the collections of the Allan Hancock Foun¬
dation and comes from 35 miles west of Depoe
Bay, Oregon, dredged in 60 to 74 fm. This is
100 mm. long and has over 225 segments. The
body is similarly long and slender, and tapers
posteriorly. No color remains except for small,
paired brown spots dorsally, at the sides, within
the parapodial bases. The parapodial ligules
are opaque white. The everted proboscis has
about the same paragnathal formula as that de¬
scribed for the individual from Moffet Point.
Notopodial falcigers are first present from seg¬
ment 46. A far posterior parapodium is shown
in Figure 7 d .

N. neoneanthes is characterized in having
paragnaths on all areas of the proboscis, those
of areas VII and VIII are most numerous; para-
podia change posteriorly such that the dorsal
ligule comes to be broad and long, carrying the
dorsal cirrus to about midway its length. Aci-
cula are black and occur singly in parapodial
lobes; homogomph falcigers have a short ap¬
pendage with blunt tooth distally and fine teeth
on the cutting edge. The body form is long and
slender. In the last-named respect N. neone¬
anthes resembles N. procera Ehlers, but the two
differ in their proboscidial arrangement and
parapodial parts. In its high paragnathal count,
it approaches N. eakini Hartman, but the latter
has different parapodial parts.

Holotype in the U. S. National Museum.

Type locality. Off Moffet Point, Alaska, in
60 fm.

Distribution. Alaska and Oregon, in 60 to
74 fm.

Genus Platynereis Kinberg

Platynereis agassizi (Ehlers)

Nereis agassizi Ehlers, 1868: 542-546, pi. 23,

fig. 1; Johnson, 1901: 399-400, pi. 4, fig.

39-45; Berkeley, 1924: 292.

Platynereis agassizi Moore, 1908: 344.
Platynereis dumerilii agassizi Berkeley, 1942:

192.

Collections. Stations 27-40 (1); 33-40 (3);
97-40 (1); 129-40 (2); L 11-41 (4); L 20-
41 (15).

The individuals here recorded are smaller
than those typical for central California, ap¬
proximating only one-half to two-thirds the size
of the latter. P. agassizi has been widely re¬
ported from parts of the northeast Pacific. The
present records are from southwestern Alaska,
in 14 to 48 fm.

Family GLYCERIDAE
Genus Glycera Savigny

Glycera capitata Oersted

Glycera capitata Oersted, 1843: 196-198, fig.
87-88, 90-94, 96, 99; Berkeley, \921a\ 411;
Berkeley, 1942: 193.

Collections. Stations 33-40 (1); 34-40 (2);
51-40 (3); A 61-40 (1); CT 12-41 (1);
D 3-41 (1); Lazy Bay (1); Seldovia (1).

Originally known from Greenland, this spe¬
cies has since been reported from arctic and
boreal seas. The present records are from south¬
western Alaska, in 17 to 150 fm.

Genus Hemipodus Quatrefages

Hemipodus borealis Johnson

Hemipodus borealis Johnson, 1901: 411-412,
pi. 10, fig. 104; Hartman, 1940: 244, pi. 43,
fig. 121.

Collections. Crab Bay, summer 1932, col¬
lected by E. F. Ricketts (1); Sitka, August
1932, collected by E. F. Ricketts (2).

The ringed prostomium and uniramous para-
podia characterize this as a Hemipodus. It is
the only known representative of the genus from
Alaska. Its range is southward to western Mex¬
ico, in littoral sandy zones.

Annelids of Alaska — HARTMAN

Genus Glycinde F. Muller

Glycinde picta Berkeley
Glycinde picta Berkeley, 1921a: 412; Berkeley,

1942: 194.

Collections. Stations 35-40 (2 anterior frag¬
ments); 97-40 (1).

The parapodial change, from uniramous to
biramous condition, occurs at the twenty-ninth
segment in all individuals, instead of the twenty-
fifth to twenty-seventh segment, as described
by Berkeley (1921a: 412). Where notopodia
are developed they have an elongate dorsal cir¬
rus and a broadly rounded presetal lamella.
They are provided with five or six yellow,
simple, hooded hooks. Neuropodia have a dor¬
sal, triangular postsetal lobe and a longer, though
similar, presetal lobe throughout the length of
the body; the presetal lobe is the longer. Neuro-
setae are composite spinigers with heterogomph
articulation. The prostomium has two eyes on
the basal ring and two less distinct spots on the
distal ring. The proboscis is provided distally
with two large, dentate jaw pieces on the dorsal
side, and a circlet of many tiny, quadricuspidate
pieces laterally and ventrally.

This species is known only from British
Columbia and Alaska. The stations here re¬
corded are southwestern Alaska, in 18 to 30 fm.

Family LUMBRINERIDAE
Genus Lumbrineris Blainville

Lumbrineris bicirrata Treadwell

Lumhrinereis bicirrata Treadwell, 1929: 1-3,

fig. 1, 2.

Lumbrineris bicirrata Hartman, 1944: 156.

Collections. Stations 52-40 (1); 59-40 (2).

The original collection came from Friday
Harbor, Washington; present records are from
Canoe Bay in 40 fm., and between Inner Iliasik
and Goloi Island, in 20 to 30 fm.

Lumbrineris latreilli Audouin and Edwards
Lumbrineris latreilli Audouin and Edwards,

29

1833: 242-244; Berkeley, 1942: 195; Hart¬
man, 1944: 158.

Lumbriconereis latreilli Fauvel, 1923: 431-432,
fig. 171.

Collections. Station 12-40 (1); 108-40 (4);
L 18-41 (1).

This species is cosmopolitan in distribution.
The present records are from southwestern
Alaska, from shore to 15 fm.

Lumbrineris zonata (Johnson)
Lumbriconereis zonata Johnson, 1901: 408-409,
pi. 9, fig. 93-100.

Lumbrineris zonata Hartman, 1944: 146-147
(with synonymy).

Collection. Sitka, August 1932, collected by
E. F. Ricketts (1).

This species was originally described from
Washington but has since been recorded from
other parts of the northeast Pacific to Lower
California. The present record is the most
northern one.

Family ARABELLIDAE

Genus Drilonereis Claparede

Drilonereis nuda Moore

Drilonereis nuda Moore, 1909: 254-256, pi. 8,
fig. 21-23; Hartman, 1944: 178-179, pi. 13,
fig. 297-302 (with synonymy).

Collection. Sitka, August, 1932, collected by
E. F. Ricketts (1).

Previously recorded from central California,
south to Panama, this marks the most northern
record for the species and genus.

Family DORVILLEIDAE

Genus Dorvillea Parfitt

Dorvillea pseudorubrovittata Berkeley

Dorvillea pseudorubrovittata Berkeley, 1921a:
409-410; Hartman, 1944: 189.

30

PACIFIC SCIENCE, Vol. II, January, 1948

Collection. Station 70-40 (4).

The articulation and proportion of palpi and
antennae agree well with those in the original
description. The species is known only through
the original account from British Columbia
(Berkeley, 1927 a) ; the present individuals are
from Cold Bay, in 15 to 35 fm.

Family ORBINIIDAE

Genus Haploscoloplos Monro

Key to Species of Haploscoloplos

1. Subpodal lobe (Fig. 8 b) present .

. H. alaskensis

1. Subpodal lobe absent . 2

2. Thorax consists of 20 segments; branch¬

iae present from sixteenth segment;
thoracic postsetal lobe short, papiilar

. H. elongata

2. Thorax consists of 14 segments; branch¬
iae present from eleventh segment;
thoracic postsetal lobe long (Fig. 8 e)
. H. sp.

Haploscoloplos elongata (Johnson)

Scoloplos elongata Johnson, 1901: 412-413,
pi. 10, fig. 105-110; Berkeley, 1927^: 413.
Haploscoloplos elongata Hartman, 19 44£: 257.

Collection. 59-40 (3).

Thoracic neuropodia are provided with only
pointed setae; acicular spines and subuluncini
are absent; this is therefore referred to Haplo¬
scoloplos. Specific characters include the fol¬
lowing. The prostomium is pointed, triangular.
Branchiae are first present from the sixteenth
setigerous segment; they are small through three
to five segments, but broad and laterally fim¬
briated farther back. The thorax consists of
20 setigerous segments in which the last one is
transitional. Thoracic notopodia and neuropo¬
dia have a short, simple, postsetal lobe at the
mid-length of their ridges; at first they resemble
papillae but by the transitional segment they are
larger and triangular.

Subpodal lobes, ventral cirri, and intercirri
are absent. Setae are long, pointed, and spinous
along their free length; those in thorax and
abdomen, in notopodia and neuropodia, re¬
semble one another except for their relative
thicknesses and lengths. In addition, three to
five furcate setae occur in abdominal notopodia,
accompanying the pointed ones.

The dorsum, between the bases of the larger
branchiae, is marked with a reticulated pigment
pattern that persists in alcohol. The branchial
tips and the broad neuropodial flanges have a
punctate dark pigment. It is this pigmented
feature, together with the similarity of the pro¬
boscis, that first prompted the identity of these
individuals with Johnsons description of the
species. In other respects there is likewise agree¬
ment, but the original description is lacking in
important details.

Scoloplos acmeceps Chamberlin (1919: 15),
from California, is another species since it has
acicular spines in thoracic neuropodia; it is a
true Scoloplos Blainville.

H. elongata was originally recorded from
Puget Sound, Washington; the present in¬
dividuals come from between Inner Iliasik and
Goloi Island, in 20 to 30 fm.

Haploscoloplos alaskensis new species
Fig. 8 a-c.

Collections. Stations 35-40 (1); 60-40 (1);
Lazy Bay (6).

One individual, posteriorly incomplete, meas¬
ures 30 mm. long for 146 segments. No color
remains (preserved). The prostomium is an¬
teriorly pointed, equitriangular and conical, or
only slightly depressed; there are no visible
eyespots. The first segment is achaetous and
apodous. The thorax includes the first 16 setig¬
erous segments; dense fascicles of pointed setae
occur in both notopodia and neuropodia. The
seventeenth segment marks the beginning of
the abdominal region; neuropodial lobes are
longer and slenderer than those in front, and
the setae are disposed in prolonged slender tufts.

Annelids of Alaska — HARTMAN

31

Fig. 8. Species of Haploscoloplos. a—c, Haploscoloplos alaskensis new species : a, far posterior
parapodium in anterior view, X 90; b, seventeenth parapodium in posterior view, X 140;
c, furcate notopodial seta, X 3950. d-f, Haploscoloplos sp. (Greenland) : d, fiftieth parapodium
in posterior view, X 156; e, tenth parapodium in posterior view, X 156; f, furcate notopodial
seta, X 3950.

32

PACIFIC SCIENCE, Vol. II, January, 1948

Branchiae are first present from the eleventh
or twelfth setigerous segment; the first are very
small but they elongate and widen rapidly
through six segments and come to be large at
the beginning of the abdominal region. Pre¬
served, many branchiae lack fringe at their outer
margins, but a few retain a delicate, close pube¬
scence that extends distally to a subterminal
enlargement (Fig. 8 a).

In the first 14 segments parapodia have an
elongate, triangular postsetal lobe along the mid¬
length of both notosetal and neurosetal ridges.
These lobes increase in size gradually through
the thoracic region. From the thirteenth setig¬
erous segment another small lobe, resembling
a ventral cirrus, makes its appearance; it is at
the lower edge of the neurosetal ridge; it in¬
creases in size through four segments, to the
sixteenth one and is continued back to the
twenty-second parapodium, where it merges
with the superior part of a broad parapodial
flange. At the sixteenth setigerous segment a
subpodal lobe (Fig. 8 b), located some distance
below the lower podal one, makes its ap¬
pearance. It is usually simple, rarely bifid. This
subpodal lobe moves gradually more ventrally
in position; it is continued into the anterior
abdominal region, through the twenty-third
parapodium, but is absent thereafter. Intercirri
are absent. Abdominal parapodia have the pro¬
portions shown in Figure 8 a.

Setae are numerous in thoracic segments;
those in neuropodia are the denser and disposed
in transverse series; notopodial setae form a
tuft. All taper to fine points and are spinous
along their free length. Abdominal setae are
similar but fewer in number and slenderer than
those in front. Furcate setae (Fig. 8 c) are
present in notopodia. Acicula are pale yellow,
straight and slender; they terminate distally in
a point; in abdominal segments they number
three or four in neuropodia and five or six in
notopodia.

H. alaskensis is characterized in having 16
thoracic setigerous segments. Branchiae are pres¬
ent from the eleventh or twelfth segment and

continued posteriorly to or near the end. Podal
lobes, resembling ventral cirri, occur on para¬
podial segments 15 to 22. Subpodal lobes, widely
separated from the podal lobes, are on para-
podal segments 16 to 23. Among the several
species of Haploscoloplos , only one, H. pana-
mensis Monro (1933: 1045) from Pacific Pana¬
ma, has subpodal lobes, but in it they are not
continued into the abdominal region and they
have a position proximal to the podal lobe, not
widely removed from it, as in H. alaskensis.

Holotype in the U. S. National Museum.

Type locality. Lazy Bay, Alaska.

Distribution. Southern Alaska.

Haploscoloplos sp.

Fig. 8 d-f.

Collection. Murchison Sound, Greenland, in
60 fm., coll. Capt. R. A. Bartlett (1-).

One incomplete individual in the collections
of the National Museum, though not from
Alaska, seems worth recording since it also
comes from a far northern locality ( Greenland ) .
It may represent an undescribed species, but the
material is too imperfect to ascribe a specific
name. It consists of 50 anterior segments and
measures 16 mm. long; greatest width in the
thorax is about 1.5 mm. The prostomium is
acutely pointed in the front, somewhat de¬
pressed and longer than wide; it lacks eyespots.
Branchiae are first present in the thorax, from
the eleventh setigerous segment; they are very
small at first but increase in size gradually so
that by the first abdominal segment they are
larger than the postsetal lobes. The transition
from thorax to abdomen is at the fifteenth seti¬
gerous segment, where neuropodia change char¬
acter abruptly, from ridges to tufts.

Thoracic and abdominal parapodia are pro¬
vided with only simple, pointed setae through¬
out. Embedded acicula are slender, yellow, and
few in number. Furcate setae (Fig. 8 /) occur
with pointed setae in abdominal notopodia;
they have a smooth stalk. Thoracic setae are in
full spreading fascicles, densest in neuropodia.

Annelids of Alaska — Hartman

33

They are bounded behind by a low fleshy ridge,
from which a long, slender lobe arises at about
the mid-length of the ridge (Fig. 8 e).

Abdominal notopodia have a long, simple,
postsetal lobe. Neuropodia have a longer, pre-
setal process and a shorter, postsetal ( or slightly
ventral) one (Fig. 8 d). There are no ventral
cirri but a short flange (or ridge) is present at
the lower end of neuropodia. Intercirri, podal
and subpodal lobes are lacking.

This specimen differs from other species of
the genus most noticeably in the slenderness of
its thoracic postsetal lobes, and abdominal noto-
setal process, also the bilobed neuropodial
process (Fig. 8 d).

This represents the first record of a Haplo-
scoloplos from Greenland.

Family PARAONIDAE

Genus Aricidea Webster

Aricidea heteroseta new species
Fig. 9 a-d.

Collection. Lazy Bay (2).

A slender, threadlike though much-coiled in¬
dividual measures not over 25 mm. long and
about 1 mm. wide; it consists of 110 segments
(posteriorly incomplete). The prostomium is
flat, depressed, trapezoidal in shape and about
as long as wide; the greatest width is behind
the insertion of the median antenna. At its an¬
terior end it is broadly rounded but when seen
from the ventral side it appears slightly incised
medially. The dorsal surface is plain except for
the attachment of antennae, and the slightly
crescentic nuchal slits near the posterior portion
(Fig. 9 a). The median antenna is smooth and
tapering; it extends back to the middle of the
first segment.

Branchiae are present from the fourth to
thirty-fourth setigerous segment; they number
30 pairs. They are abruptly absent thereafter.
The first are already large but they increase in
size gradually so that the two of a pair overlap

medially. They are broad and straplike; they
taper to a point distally and are fimbriated at
their lateral margins but the tip is smooth ( Fig.
9 b).

Parapodia have a prolonged, postsetal, noto-
podial lobe; this is small at first but increases in
size through the anterior branchial region and
diminishes thereafter. In the anterior branchial
region it is broad, auricular, and has the propor¬
tions shown in Figure 9 b; it gradually dim¬
inishes in width in the posterior branchial
segments so that it comes to be slender, cirri-
form, and is continued so to the end of the
pieces. Setal lobes are most conspicuous in the

Fig. 9. Aricidea heteroseta new species : a, anterior
end in dorsal view, X 34; b , ninth parapodium in
anterior view, X 1 19; c, far posterior parapodium,
X 119; d, acicular neuroseta from the same parapo¬
dium, X 652.

34

PACIFIC SCIENCE, Vol. II, January, 1948

Annelids of Alaska — HARTMAN

35

34

PACIFIC SCIENCE, Vol. II, January, 1948

Annelids of Alaska — HARTMAN

35

Map I. Locality map of southwestern Alaska showing most of the area in which collections were made by the Alaska King Crab Investigation.

36

first 18 segments and diminish in size and full¬
ness thereafter. Neuropodia are larger and fuller
than notopodia throughout. In anterior seg¬
ments both branches of parapodia are provided
with only tapering, pointed setae, those in neu¬
ropodia thicker and shorter (Fig. 9 b). In the
postbranchial region, notopodia come to be very
inconspicuous and are provided with only a
few slender setae, but their postsetal lobes con¬
tinue long and slender (Fig. 9 c). In post-
branchial neuropodia, setae consist of two kinds
arranged in a single transverse series; they in¬
clude a few long, slender setae, longest superi¬
orly, which alternate with thicker, shorter,
acicular setae with a distal arista (Fig. 9 d).
The posterior end is unknown.

A. heteroseta is characterized in having a
simple prostomial lobe in which the median
antenna is short; branchiae are present from the
fourth to thirty-fourth segments and number
30 pairs; anterior branchial segments have an
auricular postsetal, notopodial lobe; posterior
neuropodia are provided with two kinds of setae
including long, slender, and shorter, acicular
ones with an arista. The specific name refers to
the last named character.

Holotype in the U. S. National Museum.

Type locality. Lazy Bay, Alaska.

Distribution. Southern Alaska.

Family SPIONIDAE

Genus Laonice Malmgren
Laonice cirrata (Sars)

Nerine cirrata Sms, 1851: 207.

Laonice cirrata Fauvel, 1927: 38, fig. 12; Berke¬
ley, 1936: 474; Berkeley, 1942: 196.

Collection. Station 139-40 (1).

This species has been recorded from areas in
the northern Pacific. The present locality is off
Hallo Bay, in 28 to 40 fm.

PACIFIC SCIENCE, Vol. II, January, 1948

Genus Spio Fabricius

Spio filicornis (Muller)

Nereis filicornis Muller, 1776: 218.

Spio minus Chamberlin, 1920: 16B, pi. 3, fig.

1-4; Hartman, 1938^: 13.

Spio filicornis Fauvel, 1927: 43-44, fig. 15.
Spio filicornis pacifica Berkeley, 1936: 475-
476.

Collection. Lazy Bay (3 anterior ends).

No color remains. The prostomium is an¬
teriorly truncate, with rounded margins and not
narrowed forward. Hooded crotchets are first
present from the seventeenth setigerous seg¬
ment. The variety pacifica Berkeley (1936:
475) appears to be the same, since it is said to
differ in color only.

This species was first described from Green¬
land but has been reported from many geo¬
graphic regions. The present lot is from
southwestern Alaska, intertidal.

Genus Prionospio Malmgren

Prionospio malmgreni Claparede
Prionospio malmgreni Claparede, 1879: 73-76

pi. 22, fig. 3; Fauvel, 1927: 61-62, fig. 21;

Berkeley, 1921a\ 414.

Collection. Station 72-40 (1).

The posterior eyes are large, not small; in
other respects this individual agrees with the
descriptions.

This species has been reported from British
Columbia (Berkeley, 1921 a) \ the present col¬
lection is from Cold Bay, in 15 to 50 fm.

Genus Polydora Bose
Polydora giardi Mesnil

Polydora giardi Mesnil, 1896: 195, pi. 13, fig.

1-12; Fauvel, 1927: 50-51, fig. 17; Hartman,

1941: 309, pi. 48, fig. 3; Rioja, 1941: 727.

Collections. Stations 20-40 to 22-40 (4+)
26-40 (many, in shell fragments); 61-40
(many, in shell fragments); L 18-41 (1).

Annelids of Alaska — HARTMAN

37

The range is hereby extended in the Pacific,
from Mexico and central California to south¬
western Alaska, in 13 to 100 fm.

Polydora socialis (Schmarda)
Leucodore socialis Schmarda, 1861: 64, pi. 27,
fig. 209.

Polydora socialis Hartman, 1941: 290, 310, pi.
48, fig. 41, 42.

Collection. Station L 18-41 (1).

The single individual is posteriorly incom¬
plete. The dorsum is marked with paired dark
spots. The prostomium is distinctly bifid at its
anterior end and has four tiny eyespots in
trapezoidal arrangement. The nuchal ridge ex¬
tends back through 11 setigerous segments and
lacks a median papilla. Dorsal lamellae of the
first setigerous segment are conspicuous and ex¬
ceed in size those farther back. The first segment
has both notosetae and neurosetae. Branchiae are
first present from the eighth segment and con¬
tinued posteriorly through a long region.

Originally described from Chile, this species
las since been reported from southern Cali¬
fornia. The present record is far to the north,
in Kupreanof Strait, 2 miles northwest of Bare
Island, in 13 to 15 fm.

Family CH AET OPTERID AE
? Chaetopterus sp.

Collection. Station C 5-41 (empty tubes).

Several empty tubes, recalling those of the
cosmopolitan species, Chaetopterus variopedatus
Renier, come from Icy Straits, east end of
Pleasant Island, in 7 fm.

Family CIRRATULIDAE

Genus ClRRATULUS Lamarck

Cirratulus cirratus (Muller)
Lumbricus cirratus Muller, 1776: 214.

Cirratulus cirratus Fauvel, 1927: 94, fig. 33.

i

Cirratulus cingulatus Johnson, 1901: 422-423,
pi. 14, fig. 145-148; Berkeley, 1942: 197.

Collections. Stations 61-40 ( 2 ) ; 108-40 ( 2 ) .

This species is known from the northern
Pacific (see synonymy above). The present
records are from Cold Bay, in 15 to 30 fm.,
and Alitak Bay, shore.

Genus Tharyx Webster and Benedict

Tharyx hamatus new species
Fig. 10, a-e.

Collection. Station 108-40 (2).

The larger one measures about 12 mm. long
for over 100 segments. Both individuals are
dark slate-colored, but the prostomium and
tentacles are pale. The body rings are short and
closely crowded. The prostomium is approxi¬
mately equitriangular in shape and somewhat
depressed conical; it lacks visible eyespots.
Nuchal organs are present but not conspicuous
(Fig. 10 a). The peristomium or anterior apo-
dous region (since it probably consists of three
coalesced segments) is three to four times as
long as, and much thicker than, the prostomium.
The paired palpi are thick; each has a longi¬
tudinal groove, and is inserted dorsolaterally so
that the palpal bases are not quite touching each
other.

The lateral tentacles are most numerous in
the anterior region but continued on some seg¬
ments in a posterior region. They originate
immediately above the notopodial ridge through¬
out. Parapodia are reduced to mere ridges.
Setae in the anterior region are pointed and
slender; they extend laterally for a distance
equaling about half the width of the body at
their origin. Hooks are present in neuropodia
already before the middle of the body, but in
notopodia not until thereafter. In the posterior
fourth of the body the neuropodia are pro¬
vided with only hooks, numbering six to eight
in a single transverse series (Fig. 10 h)\ the
corresponding notopodia have both pointed
setae (Fig. 10 e) and similar hooks, alternating
with one another, together equaling eight to

38

PACIFIC SCIENCE, Vol. II, January, 1948

ten. All hooks are clearly bidentate at their
distal end, the notopodial (Fig. 10 d) some¬
what finer than the neuropodial (Fig. 10 c)
ones. The tube is fragile, dark, and cindery; it
is irregular in shape and occupies crevices of
serpulid tubes.

T. hamatus is characterized in having some
segments with both notopodia and neuropodia
provided with bidentate hooks; tentacular cirri
originate immediately above the notopodial
ridge. Three other species of Tharyx have been
described from the northern Pacific region;
they are T. multi fills Moore (1909: 267) and

Fig. 10. Tharyx hamatus new species (station 108-
40) : a, anterior end in left lateral view, X 35;
h, posterior neuropodium showing distal ends of neuro-
setae, X 105; c, neuroseta from the same parapodium,
X 1125; d, two notosetae from the same parapodium,
X 1125; e, distal end of a pointed notoseta, X 1125.

T. gracilis Moore (1923: 187) from California,
and T. parvus Berkeley (1929: 307) from
British Columbia. The first and third of these
species have only pointed setae in notopodia
and neuropodia; T. gracilis has pointed setae in
notopodia and blunt spines in neuropodia.
T. hamatus differs from all of these most clearly
in that parapodia have bidentate hooks in some
segments.

Holotype in the U. S. National Museum.

Type locality. Alitak Bay, Alaska, shore.

Distribution. Alaska.

Genus Acrocirrus Grube

This is a small genus, characterized by the
presence of composite neuropodial hooks. The
anterior end is provided with a pair of thick,
grooved palpi and a few pairs of tentacular
cirri. Notopodial setae are simple, slender, and
distally pointed. Color of the body is usually
dark.

Seven species have been described in the
genus; five originate in the north Pacific. They
are (1) A. crassiftlis (Moore, 1923: 188) from
southern California, (2) A. heterochaetus An¬
nenkova (1934: 326) from Bering and Japan
seas, (3) A. mur oranensis Okuda (1934: 202)
from Japan, (4) A. uchidai Okuda (1934:
197) from Japan, and (5) A. validus Maren-
zeller (1879: 148) from Japan. Among these,
only one, A. heterochaetus, is known to have
the eleventh segment modified and provided
with heavy, simple hooks; it agrees therein with
the type of the genus, A. frontifilis Grube (see
Fauvel, 1927: 104) from the Mediterranean
Sea.

Acrocirrus heterochaetus Annenkova
Fig. 11, a-c.

Acrocirrus heterochaetus Annenkova, 1934:

326-327, fig. 7 [in Russian].

Collections. 20-40 to 22-40 ( 1 ) ; 24-40

(1).

The color (preserved) is dark slaty brown;
there is no pigment pattern. The prostomium is

Annelids of Alaska — HARTMAN

39

short and terminates at its anterior end in a
short palpode medially; it has a pair of dark
eyespots dorsally. The palpi are thick and their
bases occupy much of the frontal region. The
tentacular cirri are similarly thick, few in num¬
ber, and inserted on a short anterior region.
Notopodial tufts are first present from the
second setigerous segment. The eleventh seg¬
ment is modified and about twice as long as the
segments proximal to it; its notopodial tuft is
normal, but its neuropodia are modified and
each is provided with a single heavy, simple,
yellow hook (Fig. 11 b) (in one case two
hooks are present). These hooks are much like
the shafts of the composite hooks except that
they are much thicker. This modified segment
seems to be a device used in tube construction
or anchorage within the tube.

Parapodial ridges have transverse series of
small papillae that are continued only slightly
beyond the parapodial bases. Notopodial tufts
are similar to one another throughout; usually
each has about six slender, pointed setae. The
latter have close transverse rows of fine spines
along the free length (Fig. 11c). Neuropodia,
except in the eleventh setigerous segment, are
provided with only composite hooks; they have
a falcate appendage that terminates distally in
a single strong tooth and a delicate sheath
(Fig. 11 a)\ the appendage and shaft are not
completely separated at the articulation.

A. heterochaetus was known only through its
original description, based on individuals from
Bering and Japan seas, from sublittoral zones
to 74 meters. The present records are from
Canoe Bay, in 15 to 40 fm.

Family ARENICOLIDAE
Genus Arenicola Lamarck

Arenicola pusilla Quatrefages
Arenicola pusilla Quatrefages, 1865: 266; Ash¬
worth, 1912: 114-123, pi. 7, fig. 15, pi. 8,
fig. 18, pi. 10, fig. 12-25, pi. 13, fig. 44, pi.
14, fig. 49; Berkeley, 1932^: 315; Berkeley,
1942: 198.

Collections. Lazy Bay (25); head of Lazy
Bay (1).

This species has been reported from Alaskan
waters; the present records are from south¬
western Alaska, intertidal.

Family OPHELIIDAE

Genus Armandia Filippi

Armandia bioculata Hartman
Armandia bioculata Hartman, 1938c: 105-106,
fig. 51-54.

Collections. Stations 47-40 (5); 51-40 (5).

The body consists of 29 or 30 setigerous
segments. Branchiae are present from the second
segment to the penultimate (or rarely last)
segment, numbering 28 (or 29) pairs. Thepros-
tomium is broadly rounded anteriorly and termi¬
nates in a small palpode. Nuchal organs are
more or less conspicuous, and partly everted in
some individuals. Two dark eyespots, one above
the other, are deeply embedded in the tissue of
the prostomial lobe.

Lateral eyespots are dark brown, present from
segments 7/8 to 17/18 and number 11 pairs.
They vary in size and shape among themselves;
most are large and circular, others are mere
specks, a few are somewhat oval. Branchiae
are long and cirriform; the first and last pairs
are smaller than the others. Parapodia are
simple, rounded, with a tiny postsetal lamella in
neuropodia. The pygidium terminates on its
ventral side in a pair of longer cirri, and laterally
in four or five similar, though smaller, cirri; a
median unpaired filament projects for a longer
or shorter distance from within the pygidial
funnel.

These indivdiuals are referred to A. bioculata
largely because of the nature of pygidial struc¬
tures and the distribution of branchiae. A. brevis
(Moore) (1906: 354), from Icy Cape, Alaska,
differs in that the pygidium, which was presum¬
ably perfect, was described without terminal

40

cirri; the lateral eyespots were described as con¬
spicuous, hemispherical in shape, and indis¬
tinctly faceted, with some small black or dark
brown ones.

The range of A. bioculata, originally described
from California, is thus extended to Canoe Bay,
Alaska, shore to 40 fm.

Fig. 11. Species of Aero cirrus and Nicomache.
a-c, Acrocirrus heterochaetus (station 24—40) : a, one
of four neurosetal hooks from a median region,
X 338; b, heavy hook from the modified eleventh
setigerous segment, X 102; c, portion of one of six
notosetae showing the spinous character, X 850. d-g,
Nicomache per sonata (Lazy Bay) : d, heavy acicular
hook from the fourth setigerous segment, X 335;
e, rostrate uncinus from a postmedian neuropodium,
X 335; f, same, in frontal view, X 335; g, posterior
end in right lateral view, X 12.

PACIFIC SCIENCE, Vol. II, January, 1948
Family SCALIBREGMIDAE

Genus SCALIBREGMA Rathke

Scalibregma inflatum Rathke
Scalibregma inflatum Rathke, 1843: 184-186,
pi. 9, fig. 15-21; Moore, 1908: 336; Cham¬
berlin, 1919: 392; Fauvel, 1927: 123-124,
fig. 44; Berkeley, 1930: 68.

Collection. Station 59-40 (1).

This species has been reported from both
sides of North America; the present record is
from between Inner Iliasik and Goloi Island,
in 20 to 30 fm.

Family FLABELLIGERIDAE
Genus Flabelligera Sars

Flabelligera infundibularis Johnson

Flabelligera infundibularis Johnson, 1901: 417,
pi. 12, fig. 124-127; Chamberlin, 1919: 398;
Berkeley, 1930: 69.

Collections. Stations 51-40 (1); 61-40 (1).
This was originally described from Puget
Sound and later reported from Alaska (Cham¬
berlin, 1919). The present records are from
Canoe Bay and Cold Bay, in 15 to 40 fm.

Genus STYLARIOIDES delle Chiaje

Stylarioides papillata (Johnson)
Trophonia papillata Johnson, 1901: 416, pi. 12,
fig. 122-123; Moore, 1908: 356.

Stylarioides papillata Berkeley, 1942: 198.

Collections. Stations 35-40 (1); 128-40
(1); A 8-41 (1).

The present records are within the known
range; they come from Pavlof Bay, Shelikof
Strait, and Orca Bay in Prince William Sound,
in 35 to 48 fm.

Annelids of Alaska — HARTMAN

41

Genus Brada Stimpson

Brada granulata Malmgren

Brada granulata Malmgren, 1867: 194, pi. 1$,
fig. 71; Murdoch, 1885&: 155.

Collection. Station 51-40 (1).

A single individual of this Arctic species is
represented. Murdoch (1885: 155) has previ¬
ously reported it from Alaska at Point Barrow,
in about 3 fm. The present record is Canoe
Bay, in 25 to 40 fm.

Family CAPITELLIDAE

Genus Heteromastus Eisig

Heteromastus filiformis (Claparede)
Capitella filiformis Claparede, 1864: 509, pi. 4,
fig. 10.

Heteromastus filiformis Fauvel, 1927: 150-152,
fig. 53; Hartman, 1947: 427-428.

Collection. Head of Lazy Bay (1).

The anterior end through the fifth setigerous
ring is thicker than the rest of the anterior re¬
gion and its surface is reticulated. The thorax
consists of 12 segments; the first is achaetous,
the second to sixth segments have only capillary
setae, and the seventh to twelfth segments are
provided with long-handled hooks. The pos¬
terior abdominal segments have inflated para-
podial ridges, but there are no other branchial
structures. The proboscis is covered with fine
papillae.

This species has been reported from both
sides of the north Atlantic Ocean; the present
record is the first from Alaska; the single indi¬
vidual comes from shore at the head of Lazy
Bay.

Genus Capitella Blainville

Capitella capitata (Fabricius)
Lumbricus capitatus Fabricius, 1780: 279.
Capitella capitata Chamberlin, 1920: 25B; Fau¬
vel, 1927: 154-155, fig. 55; Berkeley, 1929:
312; Hartman, 1947: 405.

Collection. Station 47-40 (1).

The single individual comes from shore in
Canoe Bay. The species has previously been re¬
ported from the north Pacific (see synonymy
above ) .

Family MALDANIDAE

Genus Nicomache Malmgren

Nicomache per sonata Johnson
Fig. 11, d-g.

Nicomache personata Johnson, 1901: 419-420,
pi. 13, fig. 134-139; Berkeley, 1929: 314.

Collections. Stations 61-40 (2); head of
Lazy Bay (14).

The antero-dorsal region is spotted with
black. There are 22 setigerous segments. The
first four segments are provided with single,
heavy, yellow spines in neuropodia, or the
fourth may have three spines in each parapo-
dium; the fifth has two to four spines, and
farther back there is an increasing number of
setal structures; a seventh may have eight spines.
They are at first acicular, but by the fourth
segment they are slightly rostral (Fig. 11 d)\
in the next two or three segments they are
transitional and in the posterior segments are
strongly beaked (Fig. 11 e, /).

There is a single, ante-anal, apodous, and
achaetous segment. The pygidium has a circlet
of 19 or 20 short, triangular papillae (Fig. 11 g)
usually about equally spaced and similar to one
another, or some may be bifurcate or reduced to
mere elevations.

Typically the rostrate hooks have a strong,
recurved, beaklike tooth and about six smaller
teeth above (Fig. 11 e); the fringed region is
extensive (Fig. 11 /). The long, hairlike dorsal
setae are already present from the fourth setig¬
erous segment and are continued posteriorly at
least to the eighteenth one; they extend far out
from the sides of the body (Fig. 11 g). The
tube is thick, coarse, hard and dark, and it con-

PACIFIC SCIENCE, Vol. II, January, 1948

42

sists of cindery or coarse sand particles cemented
together.

Originally described from Washington, this
species has since been reported from British
Columbia ( Berkeley, 1929 ) . The present records
are from Cold and Lazy Bays, in depths to
30 fm.

Nicomache lumbricalis (Fabricius)
Sabella lumbricalis Fabricius, 1780: 374.
Nicomache carinata Moore, 1906^: 242-244,
pi. 11, fig. 36-39.

Nicomache lumbricalis Berkeley, 1942: 199.

Collections. Stations 62-40 (1); ? 35-40
(posterior end).

A single anterior end and a posterior frag¬
ment were taken. The anterior end lacks spots
but is colored uniformly reddish brown. The
first three setigerous segments have one (or
two) large yellow spines in neuropodia. There
are no transitional rostrate hooks; the fourth
setigerous segment has series of 10 to 12 much
smaller rostrate hooks, continued posteriorly.
There are two preanal achaetous segments.

Berkeley (1942: 199) has referred N. cari¬
nata Moore to this species — a conclusion which
seems justified. The present records are within
the known range, including southern Alaska
south to Vancouver Island, Canada.

Genus Petaloproctus Quatrefages

Petaloproctus tenuis borealis Arwidsson
Petaloproctus tenuis borealis Arwidsson, 1907:

118-122, pi. 3, fig. 85-90, pi. 8, fig. 268-272;

Berkeley, 1942: 199.

Collection. Station 70-40 (1).

A single tiny individual, only 11 mm. long
and very slender, is probably a juvenile; it is
partly surrounded by a sandy tube. The anal
plaque is large and entire; the margin is not
crenulate. There are 2 1 setigerous segments and
two preanal, achaetous ones. The first three
neuropodia are provided with heavy, acicular
spines.

Originally described from Scandinavia, this
species has been reported from British Colum¬
bia by Berkeley ( 1942 ) ; the present individual
is from Cold Bay, in 15 to 35 fm.

Genus Asychis Kinberg

? Asychis lacera (Moore)
new combination

Maldane lacera Moore, 1923: 235-237.

Collection. Station 139-40 (1-, anal plaque
missing).

A single incomplete individual is question¬
ably referred to this species. The first setigerous
segment has a conspicuous collar, deepest on
the ventral side. The cephalic plaque is crenu¬
late along its margin and has about 12 lobes on
the dorsal margin. The first setigerous segment
lacks neuropodia but the second one has a
transverse series of rostrate hooks, as do those
farther back.

This species remains unknown except through
the original account, based on a single complete
individual, 55 mm. long, dredged from southern
California in 549 to 585 fm. Through the
courtesy of the administration of the U. S. Na¬
tional Museum, I have been able to examine the
type. The prostomial keel is only weakly ob¬
servable, hence it is here referred to Asychis
Kinberg. Both cephalic and anal plaques are
crenulate' at their outer margins. The cephalic
plaque has deep lateral incisions and 12 nearly
equitriangular lobes around its dorsal margin.
The lateral portions are set off from the entire
ventral lobe, and the shorter lateral lobes have
three to five irregular, crenulate lobes. The anal
plaque has deep, circular incisions; its ventral
margin has four pointed, triangular lobes; the
dorsal edge has seven shallow, triangular lobes.
There is a conspicuous, sheathing, complete
collar at the anterior margin of the first setig¬
erous segment, highest on the ventral side; it
has a lateral incision just above the first setal
fascicle.

The single fragment in the collection is re¬
ferred to this species largely because of the deep,

43

Annelids of Alaska — HARTMAN

sheathing collar and the crenulate cephalic
plaque.

This species has remained unknown except
through the single record from deep water off
southern California; the present record is off
Hallo Bay, in 28 to 40 fm.

Family SABELLARIIDAE
Genus Idanthyrsus Kinberg

Idanthyrsus ornamentatus Chamberlin

? Sabellaria saxicava Baird, 1863: 109.
Idanthyrsus ornamentatus Chamberlin, 1919:

262-263, pi. 3, fig. 2-5; Hartman, 1944:

337, pi. 31, fig. 34.

Pallasia johnstoni Berkeley, 1930: 74-75.

Collections. Stations 13-40 (10); 126-40
(3); 131-40 (1); 139-40 (1); D 11-41 (1).

The tubes are firmly concreted and broadly
attached to the substratum. They tend to be
sinuous or slightly coiled. They are attached to
pelecypod shells, living or broken fragments,
also the carapace of decapods and other hard
objects.

1. ornamentatus is distinguished from nearly
related species, I. armatus Kinberg and I. pen-
natus (Peters), most conspicuously in the shape
of the outer opercular spines (Hartman, 1944).
Sabellaria saxicava Baird is incompletely known
but originates from Vancouver Island, hence is
questionably referred to this species. Berkeley
(1930: 74) considered I. ornamentatus and
Sabellaria saxicava probably synonymous, a con¬
clusion which I consider also likely, but she
referred them to Pallasia johnstoni (McIntosh)
which has been referred to I. pennatus (Peters)
(see Johansson, 1927: 88-90). The last named
is a nearly related species, but distinguishable
in the structure of the outer opercular spines.

Previous records are from northern California
to southern Alaska. The present records are
from Canoe Bay, in 30 fm., north side of Sheli-
kof, in 65-80 fm., off Cape Chiniak, in 32-35
fm., off Hallo Bay, in 28-40 fm., and Walrus
Island, Bering Sea, in 31-33 fm.

Family AMPHARETIDAE

Genus Amphicteis Grube

Amphicteis alaskensis Moore

Amphicteis alaskensis Moore, 1905c: 846-849,
pi. 44, fig. 1-4.

Collection. Station 59-40 (1).

The single individual measures 32 mm. long.
The original collection comes from Kadiak Bay,
and southeast Alaska, in 41 to 48 fm. The
present record is from between Inner Iliasik
and Goloi Island, in 20 to 30 fm.

Genus Ampharete Malmgren

Ampharete grubei Malmgren
Ampharete grubei Malmgren, 1866: 363, pi. 19,
fig. 44; Moore, 1923: 201; Fauvel, 1927:
227-228, fig. 79; Berkeley, 1942: 201.

Collection. Station D 3-41 (1).

This species has been recorded from both
sides of northern North America and the north
Atlantic Ocean (see synonymy above). The
present record is from Castle Bay, off Chignik
Bay, in 21 to 52 fm.

Family TEREBELLIDAE

Genus Amphitrite Muller

Amphitrite cirrata Muller
Amphitrite cirrataMsillts, 1776: 188; Chamber-
lin, 1920: 22B; Berkeley, 1942: 202.
Amphitrite palmata Moore, 1905c: 858-859,
pi. 44, fig. 19-22.

Amphitrite radiata Moore, 1908: 350.

Collections. Stations 60-40 (2); C 5 — 41
(1); D 7-41 (1); D 11-41 (2); head of
Lazy Bay ( 2 ) .

Individuals attain a length of about 150 mm.
There are 17 thoracic setigerous segments. Each
of the three branchial segments has a pair of
lateral lappets. Thoracic notopodia are provided
largely with smooth capillary setae, but the

44

shorter ones are very finely toothed at the tip.
Branchiae consist of many simple filaments that
are palmately arranged. The tube is thick-walled
and made up of sand and mud that is closely
packed.

Amphitrite palmata Moore (1905c: 858),
from Alaska, is here considered to be identical;
since the specific name palmata was preoccupied
by Malmgren, the specific name radiata , was
later proposed (Moore, 1908: 350).

The present records extend from Icy Straits,
east end of Pleasant Island, west and north to
the east end of Walrus Island, Bering Sea; the
bathymetric range is shore to 38 fm.

Genus Neoleprea Hessle

Neoleprea spiralis (Johnson)

Amphitrite spiralis Johnson, 1901: 426-427,
pi. 16, fig. 169-171.

Neoleprea spiralis Hessle, 1917: 193; Berkeley,
1942: 205.

Terebella spiralis Berkeley, 1929: 308-309.

Collection. Station 12-40 (1).

There are 40 thoracic setigerous segments.
The two pairs of branchiae are dendritically
branched. The anterior end lacks thoracic lap¬
pets. Thoracic setae are distally toothed.

Originally recorded from Washington, this
species has been further reported from British
Columbia (Berkeley, 1929, 1942); the present
record extends the range to Canoe Bay, Alaska,
shore.

Genus Eupolymnia Verrill

Eupolymnia crescentis Chamberlin

Eupolymnia crescentis Chamberlin, 1919: 265-
266, pi. 3, fig. 6-7.

Collection. Station 128-40 (14).

The peristomial fold is provided with num¬
erous small eyespots. This species was first
described from northern California; the present
record is from Shelikof Strait, off Hallo Bay,
an 35 to 48 fm.

PACIFIC SCIENCE, Vol. II, January, 1948

Genus Neoamphitrite Hessle

Neoamphitrite robusta (Johnson)

Amphitrite rohusta Johnson, 1901: 425-42 6,
pi. 16, fig. 164-168; Moore, 1908: 350.
Neoamphitrite rohusta Hessle, 1917: 184;

Berkeley, 1942: 202.

Terebella rohusta Berkeley, 1929: 308.

Collections. Stations C 5-41 (1); D 3-41

(1).

Both records are in the vicinity of the known
range in the northeast Pacific. The localities are
Icy Straits in 7 fm., and Castle Bay, in 21 to
52 fm.

Genus NlCOLEA Malmgren

Nicolea zoster icola (Oersted)

Terehella zostericola Oersted, 1844: 68.
Nicolea zostericola Fauvel, 1927: 261-262, fig.
90.

Collection. Station D 15-41 (1).

There are 15 thoracic setigerous segments;
uncini are present from the second of these.
Branchiae number two pairs and are dendrit¬
ically branched; they are inserted on the two
segments preceding the first setigerous segment.
The peristomium has a row of eyespots. Nephri-
dial papillae are present on the second branchial
segment and on the third and fourth setigerous
segments; they have long lobes externally.
Thoracic dorsal setae are smooth and capillary;
uncini are avicular.

In so far as I am aware, this is the first record
from the north Pacific; it originates 20 miles
north of St. Lawrence Island, in the Bering Sea,
in 15 to 16 fm.

Genus Thelepus Leuckart

Thelepus hamatus Moore

Thelepus hamatus Moore, 1905: 856-858, pi.
44, fig. 16-18; Berkeley, 1929: 309.

Annelids of Alaska— HARTMAN

45

Collection. Station 80-40 (1).

A single small individual, posteriorly incom¬
plete, measures only 16 mm. for 35 setigerous
segments, but the body cavity contains large,
yolk-laden eggs. The body is broadest in the
anterior thoracic region. Ventral scutigerous
plates are very broad through the first four
setigerous segments and occupy the space across
the ventrum so that parapodia are dorso-lateral
in position. The peristomium has a band of
dark brown eyespots. Branchiae are on the two
presetal segments and consists of a few short,
tentacular filaments, each inserted separately
from the others. Pointed dorsal setae are smooth
and limbate; they are continued posteriorly
through at least 35 segments. Uncini are first
present from the third setigerous segment; they
have the form originally shown, with a strong
main tooth and two or three lesser teeth above,
surmounted by more numerous small denticles.

This species was first made known from the
Behm Canal, Alaska, in 130 to 193 fm.; later
Berkeley (1929: 309) recorded it from British
Columbia. The present record is from Larsen
Bay, on the east side of Nagai Island, in 5 to
25 fm.

Genus Leaena Malmgren

Leaena abranchiata Malmgren

Leaena abranchiata Malmgren, 1866: 385, pi.

24, fig. 64; Hessle, 1917: 197.

Collection. Station D 3-41 (1). 4

The single individual has 10 thoracic setig¬
erous segments; uncini are present from the
second one. Thoracic dorsal setae are broadly
bilimbate. Uncini occur in double rows only in
the tenth to sixteenth uncinigerous segments.
There is a conspicuous, high, dorsal membrane
across the segment preceding the first setigerous
one.

Apparently this is the first record from the
north Pacific; it comes from Castle Bay, off
Chignik Bay, in 21 to 52 fm. It has been pre¬
viously reported from the north Atlantic.

Genus Spinosphaera Hessle
Spinosphaera sp.

Collection. Station 61-40 (one fragment).

A single, incomplete fragment consists of
about 55 setigerous segments and measures 16
mm. long. There are over 48 thoracic segments.
Branchiae and peristomial eyespots are absent.
Thoracic dorsal setae are limbate; some have a
dentate tip and a subapical spinous region, as
typical of the genus. Uncini are avicular. The
record is from Cold Bay, in 15 to 30 fm.

Genus Polycirrus Grube
Polycirrus sp.

Collection. Station 60-40 (1).

A single individual that cannot be definitely
referred to any known species has 13 anterior
thoracic setigerous segments which lack uncini;
this is followed by a region of over 50 seg¬
ments provided with uncini. The collection was,
made in Leonard Harbor, in 20-25 fm.

Family SABELLIDAE

Genus Sabella Linnaeus
This genus has come to include two groups
of species; one (to which the genotype S. peni-
cillus Linnaeus belongs) lacks tentacular paired
eyespots and the tentacular radiole is not angu-
late on its external margins; the other (to which
S. crassicornis Sars belongs) has paired ten¬
tacular eyespots and the tentacular radiole is
strongly angulate on its external margins. Since
the genera Demonax Kinberg and Parasabella
Bush were erected for species which belong to
the genotypic group Sabella, these designations
must be regarded as synonyms of Sabella Lin¬
naeus. It may be desirable to erect a new name
for the group to which S. crassicornis Sars be¬
longs. (See also Hartman, 1942: 78.)

Sabella media (Bush)

Parasabella media Bush, 1904: 200-201, pi. 27,
fig. 3-5, pi. 33, fig. 34-36, pi. 34, fig. 3, pi.

46

PACIFIC SCIENCE, Vol. II, January, 1948

36, fig. 13, 14, pi. 37, fig. 30; Berkeley, 1930:
70; Berkeley, 1932^: 315.

Parasabella maculata Bush, 1904: 201, pi. 28,
fig. 8, 9, pk 33, fig. 8, 12, 33, pi. 34, fig. 2,
pi. 36, fig. 12, 15, 16, 21, 22.

Sabella media Hartman, 1942: 79-80, fig. 159,

160.

Collection. Canoe Bay (1).

The single collection is from Alaska, from a
previously reported locality.

Sabella crassicornis Sars

Sabella crassicornis Sars, 1851: 202-203; Jo¬
hansson, 1927: 119-121; Hartman, 1942:
78-79.

Sabella elegans Bush, 1904: 194-195, pi. 26,
fig. 2, pi. 27, fig. 6, pi. 33, fig. 20, 21, pi. 34,
fig. 1, 4, 5, 10; Moore, 1908: 359; Berkeley,
1930: 70.

Sabella leptalea Bush, 1904: 195-196, pi. 27,
fig. 6.

Collections. Station 60-40 (2); Seldovia

(2).

This species is common in cold waters of both
northern Pacific and Atlantic oceans; the present
records are near the known range; they come
from Leonard Harbor in 20 to 25 fm. and from
the beach at Seldovia.

Genus POTAMILLA Malmgren
Potamilla neglecta (Sars)

Sabella neglecta Sars, 1851: 203.

Aspeira modesta Bush, 1904: 202-203, pi. 25,
fig. 3, pi. 36, fig. 27-31, 33-35.

Potamilla neglecta Johansson, 1927: 143-145;
Hartman, 1942: 81.

Collection. Station D 15-41 (1).

This has been reported from Kadiak, Alaska
(Bush, 1904); the present record is from 20
miles north of St. Lawrence Island, Bering Sea,
in 15 to 16 fm.

Genus PSEUDOPOTAMILLA Bush

Pseudopotamilla intermedia Moore
Pseudopotamilla intermedia Moore, 1905 b\
562-564, pi. 37, fig. 15-22; Hartman, 1938;*:
19, 25, 26.

Pseudopotamilla reniformis Moore, 1908: 359-
360; Berkeley, 1930: 70; Berkeley, 1932;*:
315.

Collections. Stations 20-40 to 22-40 (frag¬
ment); 60-40 (many).

The type locality is Afognak Bay, Alaska, in
14 to 19 fm. (Moore, 1908: 359). Present
records are from Canoe Bay, in 15 to 40 fm.,
and Leonard Harbor, in 20 to 25 fm.

Pseudopotamilla occelata Moore

Pseudopotamilla occelata Moore, 1905£: 559-
562, pi. 37, fig. 8-14; Berkeley, 1930: 70;
Hartman, 1938;*: 19, 20, 25, 26, 27, pi. 2,
fig. 6, 7.

Pseudopotamilla brevibranchiata Moore, 1905 A :
555-559, pi. 37, fig. 1-7.

Collections. Stations 61-40 (2); Mist Har¬
bor (2).

These records, Cold Bay, in 15 to 30 fm., and
Mist Harbor, are within known ranges.

Genus SCHIZOBRANCHIA Bush

Schizobranchia insignis Bush
Schizobranchia insignis Bush, 1904: 206-207,
pi. 24, fig. 1, 2, pi. 27, fig. 1, pi. 28, fig. 5,
pi. 35, fig. 2, 12, 13, 15, 16, 26, 27; Hartman,
1942: 82-83; Berkeley, 1942: 206.
Schizobranchia nobilis Bush, 1904: 207, pi. 24,
fig. 3, pi. 28, fig. 7, pi. 33, fig. 22, pi. 35,
fig. 1, 3-6, 8, 10, 11, 23; Berkeley, 1930: 71.

Collections. Stations 61-40 (1); under the
dock at Sand Point (1).

Both individuals have eight thoracic setig-
erous segments; tentacular radioles are split
three to five times. These records are within
the known range.

Annelids of Alaska — HARTMAN

47

Schizobranchia dubia Bush

Schizobranchia dubia Bush, 1904: 208-209, pi.
28, fig. 1, pi. 29, fig. 1, pi. 33, fig. 7, pi. 36,
fig. 1-3, 17-20, pi. 37, fig. 28; Hartman,
1942: 83-84.

Collections. Stations 20-40 to 22-40 (many);
24-40 (many); 51-40 (many); 52-40 (tubes
only); 61-40 (many); 108-40 (many); C 150—
41 (many).

In most instances there are many tubes clus¬
tered together; they are interwoven but not
connected to one another at their bases. The
tubes have a strong, tough chitinized base and
are usually covered on the outside, especially
distally, with fine, dark-colored sand grains.
Thoracic setigerous segments usually number
only six, though rarely seven or five. Tentacular
radioles are bifurcated once or twice, or rarely
three times.

This species was originally described from
Alaska; the present collections are from Canoe
Bay in 15 to 125 fm., Cold Bay in 15 to 30
fm., Alitak Bay on the reef, and off Moffet Point
in 60 fm.

Genus Myxicola Koch

Myxicola infundibulum (Montagu)

Amphitrite infundibulum Montagu, 1808: 109-
110, pi. 8.

Myxicola pacifica Johnson, 1901: 431-432, pi.
19, fig. 193-198.

Myxicola conjunct a Bush, 1904: 217-218, pi.

26, fig. 1, 4, pi. 38, fig. 1-11.

Myxicola infundibulum Hartman, 1938^: 19,
pi. 1, fig. 5-11, pi. 2, fig. 1; Hartman, 1942:
86.

Collections. Stations 12-40 (1); D 11-41

(1).

This cosmopolitan species has been reported
from Alaska ( see synonymy above ) . The pres¬
ent collections are from Canoe Bay, shore, and
east of Walrus Island, Bering Sea, in 31 to
33 fm.

Genus Chone Kroyer
Chone sp.

Collections. Stations 36a-40 (1); 108-40

(1).

Two tiny individuals, measuring less than
10 mm. long, whose specific identity has not
been determined, come from Pavlof Bay in
27 fm. and Alitak Bay, reef.

Family SERPULIDAE

Genus Serpula Linnaeus

Serpula vermicularis Linnaeus
Serpula vermicularis Linnaeus, 1767: 1267;
Chamberlin, 1919: 269; Berkeley, 1930: 73;
Berkeley, 1932 a: 316; Hartman, 1942: 16;
Berkeley, 1942: 206.

Serpula Columbiana Johnson, 1901: 432-433,
pi. 19, fig. 199-204.

Serpula splendens Bush, 1904: 230-232, pi.
24, fig. 3, pi. 29, fig. 2, pi. 30, fig. 2, 3,
pi. 33, fig. 31, pi. 35, fig. 18, pi. 37, fig. 31,
pl. 39, fig. 33.

Collection. Seldovia ( 1 ) .

This species is well known from the northeast
Pacific, including Alaska.

Genus CRUCIGERA Benedict

Crucigera zygophora (Johnson)

Serpula zygophora Johnson, 1901: 433-434, pl.
19, fig. 205-208.

Crucigera zygophora Bush, 1904: 233, pl. 29,
fig. 5, pl. 31, fig. 2, pl. 33, fig. 3, pl. 39, fig.
8-13, 15, 17, 20; Berkeley, 1942: 207; Hart¬
man, 1942: 87-88.

Crucigera formosa Bush, 1904: 233-234, pl.
28, fig. 3, 4, pl. 31, fig. 1, pl. 33, fig. 4, pl. 39,
fig. 6, 7, 10, 11, 14.

Collections. Stations 12-40 (12 and tube
clump); 13-40 (1); 21-40 (1+), 20-40 to
22-40 (clump); 24-40 (1); 70-40 (several);
82-40 (clump); 109-40 (clump); C 160-

48

PACIFIC SCIENCE, Vol. II, January, 1948

41 (clump); L 18-41 (clump); Canoe Bay
( clump ) ; Mist Harbor ( clump ) .

These localities are within the known range
in Alaska. Depths range from shore to 125 fm.

Crucigera irregularis Bush

Crucigera irregularis Bush, 1904: 234, pi. 25,
fig. 5, pi. 29, fig. 4, pi. 33, fig. 13, pi. 39,
fig. 1-5; Berkeley, 1930: 73; Berkeley, 1942:
73.

Collections. Stations 60-40 ( 1 ) ; 84-40 ( 1 ) ;
88-40 (1); 139-40 (1); A 8-41 (1); D 11-
41 (1).

Originally described from Juneau, Alaska,
this species has been reported from British
Columbia (Berkeley, 1942); the present collec¬
tions come from southern and southwestern
Alaska, in 15 to 39 fm., and from Walrus
Island, Bering Sea, in 31 to 33 fm.

Genus Chitinopoma Levinsen, emended

Type C. groenlandica (Morch)

The operculum is soft and conical; it is
covered by a slightly convex, chitinous plate.
The tube is calcareous, elongate, more or less
irregularly sinuous, and provided with a strong,
longitudinal Carina. The tentacular crown has
six or seven pairs of radioles, each provided
with long paired pinnae and a longer or shorter
free distal end. The thoracic collar is high; it
consists of a pair of dorsal lobes and a broad,
ventro-lateral portion; a thoracic membrane is
absent. Thoracic setigerous segments number
seven. The collar ring is provided with a setal
fascicle, including special setae with a finlike
expansion, and geniculate setae. Other thoracic
segments have notopodia and neuropodia. The
seta! formula is as follows: thorax with smooth
geniculate and slender setae in notopodia, and
flat uncinial plates with about nine teeth in
neuropodia. Abdominal segments are numerous,
each is provided with uncinial plates in noto¬
podia and only one or two geniculate setae in
neuropodia.

The type of the genus, C. groenlandica
(Morch), was originally named Serpula trique-
tra Fabricius (1780) (not Linnaeus) and
originated in Greenland. Morch, in a revision
of the family (1863: 375), referred this spe¬
cies to Hydroides norvegica var. groenlandica.
Malmgren (1867: 120) raised the variety to
rank of species, questionably in the genus
Hydroides. Later, Levinsen (1884: 203) erected
the genus Chitinopoma, with new species,
C. fabricii, and referred Serpula triquetra Fabri¬
cius to it. As shown by Bush (1904: 229) (in
footnote), Levinsen s name is superfluous and
must give way to Malmgren’s name.

In a key to the Serpulidae, Bush ( 1904: 224)
placed Chitinopoma Levinsen in the same cate¬
gory with Hyalopomatopsis St. Joseph, and dif¬
ferentiated them thus:

Chitinopoma with uncinial plates trapezi-
form, with appressed teeth, the lowest
larger than the others. Operculum with
concave horny plate.

Hyalopomatopsis with uncini somewhat sim-
iliar to those in Spirorbis, the teeth longer.
Operculum with a chitinous or horny cap.

Actually, these statements reveal no real dif¬
ferences, since the opercular caps are convex in
both cases and the shape of uncinial plates is
not materially different. Both have a soft oper¬
culum with horny, convex cap, and both have a
high thoracic collar and no thoracic membrane.
However, the type of Hyalopomatopsis, H. mar-
enzelleri (Langerhans), has only six thoracic
setigerous segments; Chitinopoma has seven;
abdominal setae are geniculate in the latter, not
so in the former. Capillary setae are present
only in the second to seventh thoracic setigerous
segments in Chitinopoma, and presumably pres¬
ent in all segments in Hyalopomatopsis (see
Chamberlin, 1919: 475). The type of Hyalopo¬
matopsis originates from abyssal depths in the
middle east Atlantic Ocean and may be known
through only a single species. A second species
which has been attributed to it, H. occidentalis
Bush (1904: 229), is now referred to Chiti¬
nopoma ( see page 50 ) .

Annelids of Alaska — HARTMAN

49

Fig. 12. Chitinopoma occidentals: a, thoracic uncinial plate from left side, seen in posterior view,
X 1342; b, larger collar seta, seen from the side, X 748; c, smaller collar seta from the same fascicle,
seen from the side, X 895; d , anterior end of body, showing operculum with attached tentacular
crown, in dorsal view (opercular cap turned to right to show asymmetry), X 19.4; e, anterior end
of tube, showing circular aperture, X 15.6; /, entire tube attached to shell fragment, in dorsal view,
X 4.4.

50

PACIFIC SCIENCE, Vol. II, January, 1948

Chitinopoma may be nearly related to Micro-
serpula Dons for which the single known spe¬
cies, M. inflata Dons, has been recorded from
Arctic seas (Brattstrom, 1945). In the latter,
the tube is said to have ovicels distally.

Chitinopoma occidentalis (Bush)
new combination

Fig. 12, a-f.

Hyalopomatopsis occidentalis Bush, 1904: 229-
230, pi. 40, fig. 3, 22, pi. 44, fig. 2, 4, 8.

Collections. Stations 13-40 (several); 21-40
(3); 20-40 to 22-40 (many); 24-40 (many);
25-40 (several); 35-40 (several); 47-40 (sev¬
eral); 61-40 (several); 70-40 (several); 82-
40 (few); 84-40 (several); 100^10 (several);
Mist Harbor (1+).

White calcareous tubes are attached to shell
fragments, other serpulid tubes, gastropod shells,
living rock oysters, branchiopods, carapace of
crabs, and other hard surfaces. The tubes are
hard, smooth, and broad; they have a strong,
median keel with a sharp notch above, at the
aperture (Fig. 12 /); the aperture is circular
(Fig. 12 e). Tubes seldom cover one another
unless crowded; they are sinuous or irregularly
twisted; they measure 30 to 50 mm. long.
Larger individuals (fixed in the tube) are 15 to
20 mm. long. The body consists of a well-
developed tentacular crown, seven thoracic, and
50 or fewer abdominal, segments.

The operculum replaces the dorsal radiole on
the left side. It has a smooth, straight stalk and
extends distally beyond the outer ends of the
radioles. The expanded, distal portion, seen from
the dorsum is symmetrical, but seen from the
side (Fig. 12 d) is asymmetrical, the greater
convexity on the ventral side. It is a soft, pale
vesicle surmounted by a slightly chitinized, con¬
vex cap. The tentacular crown has six radioles
on the left side, seven on the right one. Ten¬
tacular pinnae are longest on the distal third of
the radiolar length but surpassed in length by
the long, free distal filament of the rachis.

The collar is a high, thin membrane; it con¬
sists of a pair of long, dorsal lobes that com¬
pletely conceal the peristomium; they are sepa¬
rated from other parts of the collar by deep,
dorso-lateral clefts. Laterally and ventrally, the
collar consists of a still longer piece that extends
forward to conceal over half of the radiolar
length. When the collar is pushed back, a pair
of deep-seated, ocular spots is visible in the
fleshy base of the peristomium. There is no
indication of a thoracic membrane.

Collar setae are arranged in a lengthwise,
horizontal series, slightly dorsal to the other
notopodia. They have two kinds of setae; an¬
teriorly there are six or seven larger, pointed
setae with broad, thick expansion (Fig. 12 b)\
immediately behind are a comparable number
of slenderer ones that are also crenulate at the
outer margin (Fig. 12 c).

Second to seventh thoracic setigerous seg¬
ments resemble one another, but the last one
has a greatly reduced parapodium. Notopodia
are each provided with two kinds of setae, in¬
cluding six to eight heavier, smooth, slightly
limbate ones in front, and a comparable num¬
ber of slenderer ones farther back. Neuropodia
have vertical rows of uncinial plates. These
plates, where best developed in middle thoracic
segments, number about 50 in a ridge. The
largest ones are at the ventral end of the series,
decreasing gradually in size dorsally, so that the
uppermost one is only about half as long as the
lowest. The last thoracic neuropodium has only
about 15 uncinial plates.

The structure of the uncinial plates is such
that their three-dimensional arrangement is dif¬
ficult to ascertain in any one view. As typical
of other serpulids, they lie in a closely appressed
row. It is difficult to dissect them out in¬
dividually, since there is a strong tendency for
all of them to hold together and to spring back
after depression. The reason for this is only
partly that they are bound together by fleshy
fibrils and muscles. Actually, each plate (Fig.
12 a) is provided with a locking mechanism
which interlaces with the plates on either side;

Annelids of Alaska — HARTMAN

51

all hooks of a series are thus held together
firmly so that they function as units. One may
liken the plate to a hand in which the broad,
embedded part is the palm, and the distal free
teeth the fingers. The teeth of the plate are
extended at an angle slightly obtuse to the
plate, or one might say that palm and fingers
are at somewhat greater than right angles to
each other. As a result, one cannot see both
palm and fingers in full outline at any one time.
The teeth, numbering usually nine (less often
10) are nearly equal to one another in size;
they are long and slender, with tapering, slightly
curved tips; they project from the fleshy lobe of
the neuropodium. In addition to these nine
teeth, the lowest end of the plate has two addi¬
tional hooks, a smaller distal one, corresponding
to the thumb of a hand, which is directed back¬
ward and ventrally, and a longer, more out¬
stretched one, comparable to an index finger,
which is directed forward and ventrally. These
two hooks constitute the locking mechanism,
interlacing with similar hooks on the plates
proximal to them.

Abdominal parapodia have similar, though
smaller, uncinial plates in notopodia, and one or
two toothed setae in neuropodia. The abdo¬
men consists of 45 to 50 segments and termi¬
nates posteriorly in a pair of short, subspherical
papillae below the anal aperture.

C. occidentalis was originally described from
Prince William Sound and later reported from
Alaska (Moore, 1908: 362). Hyalopomatopsis
occidentalis Moore (1923: 254), off Santa Rosa
Island, California, may be another species (or
genus) since the tube has not only a median
carina, but the surface is wrinkled with trans¬
verse growth lines, and the collar setae are
different.

C. groenlandica Levinsen has also been re¬
ported from the northeast Pacific (Pixell, 1912:
790, and Berkeley, 1930: 74). This differs from
C. occidentalis in that the tube has not only a
median carina, but also fine transverse striations
and the special collar setae have a finlike por¬

tion that is set off from the main blade by a
constriction.

The present collections are from south and
southwestern Alaska, from shore to 125 fm.

Genus Dexiospira Caullery and Mesnil

Dexiospira spirillum (Linnaeus)
Serpula spirillum Linnaeus, 1767: 1264.
Spirorbis spirillum Moore, 1908: 362; Pixell,
1912: 796-797, pi. 88, fig. 8; Berkeley, 1930:
74; Berkeley, 1932: 316; Berkeley, 1942;
207.

Spirorbis ( Dexiospira ) spirillum Fauvel, 1927:
392-393, fig. 132.

Collections. Stations 9-40 (many); 10-40
(6); 25-40 (many); 52-40 (10).

Tubes are attached to algae or hard objects.
This species has been reported from inter¬
tidal regions of the northeast Pacific, including
Alaska. The present records are well within
known ranges.

Genus Laeospira Caullery and Mesnil

Laeospira borealis (Daudin)

Spirorbis borealis Daudin, 1800: 145; Borg,
1917: 22-26, fig. 5-11; Hartman, 1942:
92-93.

Spirorbis asperatus Bush, 1904: 245, pi. 28,
fig. 10, pi. 30, fig. 4, pi. 41, fig. 4-6, 8, 10,
11, 19, 31, 32, pi. 43, fig. 1-3, 7, 13, 26.

Collection. Station 25-40 (several).

These individuals come from Canoe Bay, in
25 fm. Previous records are from Sitka and
Prince William Sound, Alaska (Bush, 1904).

Genus PARADEXIOSPIRA Caullery and Mesnil

Paradexiospira violaceus (Levinsen)
Spirorbis violaceus Levinsen, 1884: 202; Bush,
1904: 242-243, pi. 41, fig. 1, 2; pi. 42, fig,
8-12.

Spirorbis ( Paradexiospira ) violaceus Fauvels
1927: 391-392, fig. 132,

52

PACIFIC SCIENCE, Vol. II, January, 1948

Collections. Stations 20-40 to 22^0 (many ) ;
.24-40 (3); 25-40 (several).

Tubes are attached to hard objects such as
shell fragments, stones, carapace of crabs. Bush
(1904) has reported the species from Sitka,
Alaska, and British Columbia. Present records
are from southern Alaska in 15 to 125 fm.

LIST OF STATIONS

Sta. 3-40. Sept. 6. Port Ashton, Alaska. Shore,
at high tide.

Sta. 9-40. Sept. 13. Canoe Bay, northern part.
Shore, at high tide.

Sta. 10-40. Sept. 16. Canoe Bay, southwest
shore. Shore, at high tide.

Sta. 12-40. Sept. 17. Canoe Bay, north shore.
Sta. 13-40. Sept. 17. Canoe Bay, northwest
corner. Gill net, in 30 fm.

Sta. 20-40. Sept. 21. Canoe Bay, middle part.
Otter trawl, in 40 fm.

Sta. 21-40. Sept. 21. Canoe Bay, northwest part.
Otter trawl, in 35 fm.

Sta. 20, 21, 22-4*0. Sept. 21. Around Canoe Bay.

Otter trawl, in 15-40 fm.

Sta. 24-40. Sept. 23. Canoe Bay, northwest part.
Trawl, in 125 fm.

Sta. 25-40. Sept. 23. Canoe Bay, northwest part.
Trawl, in 25 fm.

Sta. 26-40. Sept. 23. Canoe Bay, northwest part.
Trawl, in 100 fm.

Sta. 27-40. Sept. 23. Canoe Bay, entire western
part. Trawl, in 14-36 fm.

Sta. 31-40. Sept. 24. Canoe Bay. Trawl, in 34-
36 fm.

Sta. 33-40. Sept. 25. Pavlof Bay, northern part.
In 18 fm.

Sta. 34-40. Sept. 25. Pavlof Bay, middle section.
In 150 fm.

Sta. 35-40. Sept. 25. Pavlof Bay, entrance to
bay by Cape Tolstoi. Sticky bottom.

Sta. 36a-40. Sept. 25. Pavlof Bay, in 27 fm.

Sta. 47-40. Sept. 29. Canoe Bay, mid-northern
shore. Shore, at low tide.

Sta. 51-40. Oct. 2. Canoe Bay. In 25-40 fm.
Sta. 52-40. Oct. 2. Canoe Bay. In 40 fm.

Sta. 58-40. Oct. 8. Volcano Bay. In 25-30 fm.
Sta. 59-40. Oct. 8. Between inner Iliasik and
Goloi Island. In 20-30 fm.

Sta. 60-40. Oct. 10. Leonard Harbor. In 20-
25 fm.

Sta. 61-40. Oct. 10. Cold Bay. In 15-30 fm.

Sta. 62-40. Oct. 11. Cold Bay. In 15-20 fm.

Sta. 66-40. Oct. 15. Canoe Bay, northwestern
part. In 35-45 fm.

Sta. 70-40. Oct. 17. Cold Bay. In 15-35 fm.

Sta. 71-40. Oct. 17. Cold Bay. In 15-25 fm.

Sta. 72-40. Oct. 17. Cold Bay. In 15-50 fm.

Sta. 80-40. Oct. 21. Larsen Bay, on east side of

Nagai Island. In 5-25 fm.

Sta. 82-40. Oct. 22. Off southwest shore of Big
Koniuji Island. In 25-30 fm.

Sta. 83-40. Oct. 22. Off south side of Big Ko¬
niuji Island. In 25-55 fm.

Sta. 84-40. Oct. 24. Stepovak Bay. In 15 fm.

Sta. 88-40. Oct. 24. Stepovak Bay. In 90 fm.

Sta. 89-40. Oct. 25. Unga Strait or Stepovak
Bay. In 37-47 fm.

Sta. 91-40. Oct. 26. Baralof Bay, Squaw Har¬
bor. In 24 fm.

Sta. 93-40. Oct. 29. Spitz Island. In 55-68 fm.

Sta. 97-40. Oct. 21. Alitak Bay. In 18-30 fm.

Sta. 98-40. Oct. 31. Alitak Bay. In 42 fm.

Sta. 100-40. Oct. 31. Alitak Bay. In 30 fm.

Sta. 103-40. Nov. 1. Alitak Bay. In 20-45 fm.

Sta. 106-40. Nov. 2. Alitak Bay. In 15-30 fm.

Sta. 107—40. Nov. 2. Alitak Bay. In 30 fm.

Sta. 108-40. Nov. 3. Alitak Bay. From reef.

Sta. 109-40. Nov. 4. South entrance to Olga
Bay. In 40 fm.

Sta. 116-40. Nov. 6. Off Cape Alitak. In 40 fm.

Sta. 126-40. Nov. 14. North side of Shelikof.
In 65-80 fm.

Sta. 128-40. Nov. 15. Shelikof Strait, off Hallo
Bay. In 35-48 fm.

Sta. 129-40. Nov. 15. Shelikof Strait, off Hallo
Bay. In 48 fm.

Sta. 131-40. Nov. 16. Off Cape Chiniak. In
32-35 fm.

Sta. 131-40 or 132-40. Nov. 16. Off Cape
Chiniak. In 32-35 or 45-77 fm.

Sta. 135-40. Nov. 20. Off Hallo Bay. In 40-
48 fm.

Sta. 138-40. Nov. 21. Off Hallo Bay. In 28-
40 fm.

Sta. 139—40. Nov. 21. Off Hallo Bay. In 28-
40 fm.

Sta. 140-40. Nov. 21. Off Hallo Bay. In 28-
40 fm.

Sta. A 8-41. Mar. 14. Orca Bay, off Sheep Point,
Prince William Sound. In 36 fm., blue mud.

Sta. A 61-41. May 8. 40 mi. above Port Moller.
In 17-18 fm., sand, gravel.

Sta. BT 70-41. Aug. 18. Bering Sea, off Black
Hill. In 33 fm., gray sand.

® DI5.lt

Annelids of Alaska — HARTMAN

53

U

3

s2

54

Sta. C 5-41. Feb. 28. Icy Straits, east end of
Pleasant Island. In 7 fm., over gray glacier
mud.

Sta. C 44-41. Mar. 28. Albatross Bank. In 60
fm., line gravel and white sand.

Sta. C 71-41. Apr. 11. Pavlof Bay, north end.
In 60 fm., coarse sand and mud.

Sta. C 147 — 41. May 31. Off Cape Leontovich.
In 60 fm., fine gray sand.

Sta. 150-41. June 1. Off Moffet Point. In 60
fm., fine gray sand.

Sta. C 160-41. June 8. Canoe Bay. In 60 fm.,
soft mud.

Sta. CT 12-41. May 30. Off Black Hill. In 105
fm., gravel.

Sta. D 3-41. June 24. Castle Bay, off Chignik
Bay. In 21-52 fm.

Sta. D 7-41. July 17. Bering Sea, 57° N, 163°
48' W, in 38 fm.

Sta. D 8-41. July 18. Bering Sea, 58° 34' N,
165° 17' W, in 42 fm.

Sta. D 11-41. July 24. Bering Sea, 12 14? mi.
east of Walrus Island, Pribilofs. In 31-33 fm.

Sta. D 14-41. Aug. 5. Bering Sea, 62° 25' N,
173° W, in 36 fm.

Sta. D 15-41. Aug. 7. Bering Sea, 20 mi. north
of St. Lawrence Island, in 15-16 fm.

Sta. D 16-41. July 18. 50 mi. northwest of
Amak Island, Bering Sea, in 50 fm.

Sta. L 2-41. Mar. 19. Olga Bay, south end of
Kodiak Island. In 25-30 fm.

Sta. L 3-41. Mar. 19. Lazy Bay, Alitak. In 15
fm.

Sta. L 4-41. Mar. 19. 9 mi. SSW of Cape Ali¬
tak, 56° 48' N, 154° 30' W. In 35 fm.

Sta. L 11-41. Mar. 22. Three Saints Bay, Kodiak
Island. In 20-30 fm.

Sta. L 13-41. Mar. 23. Port Hobson, north end
of Sitkalidak Island. In 15 fm.

Sta. L 18-41. Apr. 2. Kupreanof Strait, south
side, 2 mi. NW of Bare Island. In 13-15 fm.

Sta. L 20-41. Apr. 5. Raspberry Strait, across
from Port Wakefield. In 2-30 fm.

Canoe Bay or Pavlof Bay. Sept., 1940.

Canoe Bay. Sept. 23, 1940.

Dolgai Harbor. Oct. 6, 1940. Low tide, off
water works.

Sand Point. Oct. 25, 1940.

Mist Harbor. Oct. 27, 1940.

Pavlof Bay. Nov., 1940.

Mitrofania Bay.

Lazy Bay, off Alitak Bay. Nov. 12, 1940. Dig¬
ging in sand and gravel at low tide.

Head of Lazy Bay. Jan. 22, 1941.

Seldovia. Mar. 18, 1941.

PACIFIC SCIENCE, Vol. II, January, 1948
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179-259, 2 pi.

Moore, J. P., and K. J. Bush. 1904. Sabellidae
and Serpulidae from Japan with descriptions
of new species of Spirorbis. Acad. Nat. Sci.
Phila., Proc. 56: 157-179, 2 pi.

Muller, O. F. 1771. Von Wurmern des siissen
und salzigen Wassers. 156 p. Heinick
Mumme und Faber, Copenhagen.

- 1776. Zoologiae Danicae prodromus

sen animalium Daniae et Norvegiae indi-
genarum characters, nomina et synonyma
imprimis popularium. xxxii -f- 274 p.
Havniae.

Murdoch, J. 1883 a. Description of seven new
species of Crustacea and one worm from
Arctic Alaska. U. S. Natl. Mus., Proc. 7:
518-522.

- 1885A Report of the International

Polar Expedition to Point Barrow, Alaska
(1885). Vermes. Chaetopoda. 152-156.

0RSTED, A. S. 1843. Annulatorum danicorum
conspectus. Fasc. 1 : Maricolae. 1-52, 7 pi.

- 1844. De regionibus marinis. Ele¬
ment a topographiae historico naturalis freti
Oresund. Havniae.

Okuda, S. 1934. The Polychaeta genus, Acro-
cirrus, from Japanese waters. Hokkaido Imp.
Univ., Faculty Sci., Jour. ser. 6, zool., 2: 197—
209.

- 1936. Japanese commensal polynoids.

Annot. Zool. Jap. 15: 561-571, 7 fig.

Pallas, P. S. 1766. Miscellanea Zoologica qui-
bus novae imprimis atque obscurae animalium
species des crib untur et observationibus iconi-
busque illustrantur. 224 p., 14 pi. Hagae
Comitum.

Pixell, H. 1912. Polychaeta from the Pacific
coast of North America. Pt. 1. Serpulidae
with a revised table of classification of the
genus Spirorbis. Zool. Soc. London, Proc.
1912: 784-805, 3 pi.

QUATREFAGES, J. L. A. 1865. Histoire naturelle
des anneles marins et d’eau douce. Paris.

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wegena. Nova Acta Acad. Leop. Carol. Nat.
Cur. 20: 1-264, 12 pi.

Riddell, W. 1909. Spinther oniscoides John¬
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Rioja, E. 1941. Datos para el conocimiento de
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Sars, M. 1835. Beskrivelser og lagttagelser
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The Question of Avian Introductions in Hawaii

Harvey I. Fisher1

A UNIQUE FACT about the avifauna of the
Hawaiian Islands is that seldom is a native bird
seen in the urban areas. However, birds of a
few species are abundant in the same areas;
these birds are exotics introduced for various
"good” reasons. The importation of birds has
not been limited to forms that would be re-
tricted to the habitat provided by the lower
coastal areas which, with few exceptions, are
also the urban sections. Hawaiian mountains
and forests have many established foreign birds,
and native birds in most regions are relatively
rare.

In perhaps no other similarly circumscribed
area in the world have as many exotic species
been introduced successfully. Bryan (1944: 84)
records 232 species for Hawaii; of these, 94 are
exotics of which 53 are probably established and
the remainder of unknown status. He notes that
so little field work has been done that it is im¬
possible to know the fate of many of the ex¬
otics. In addition, it is worthy of mention that
in many instances no one even knew at the
time what was being liberated here; the records
of the territorial agency concerned often simply
state "500 small birds,” or list little-used col¬
loquial names, or those used by dealers in birds.
Furthermore, the home locale of the bird is not
generally recorded; the port of embarkation for
Hawaii is the only information we have that
sheds light on the native region of the species,
and such information is of little use. In those
instances in which closely related species, and
subspecies of one species, have been imported
(as in doves), and where interbreeding may
have occurred, we probably never will be able

1 Department of Zoology and Entomology, Univer¬
sity of Hawaii, Honolulu, T. H. Manuscript received
July 9, 1947.

to unravel the situation enough to determine the
breeding stock first liberated. Had all pertinent
information been recorded, the ornithologist
now and in the future might have been able to
note the adaptive changes in the structure, food
habits, and behavior of the birds occasioned by
the environmental conditions in Hawaii. This
knowledge would in turn aid us in evaluating
past and possible future importations, and it
would have been of interest and value to orni¬
thologists all over the world.

Importations in the past have been for the
most part a result of the activities of a few or¬
ganizations and several individuals. At least one
group in Hawaii was organized primarily for the
purpose of introducing and establishing song¬
birds in these islands. There is no question that
these groups and individuals believed they were
"improving” the natural attractiveness of the
islands. One may question, however, the benefits
derived from these activities. In addition to the
species purposely liberated, several kinds of
birds have first come in as cage birds and later
escaped. Among these are the Chinese Thrush
(T roch alopter um canorum ), which escaped in
Honolulu during a large fire in 1900, the Straw¬
berry Finch ( Amandava amandava ) , and the
House Finch, or Linnet ( Carpodacus mexi -
canus). These are, of course, inadvertent libera¬
tions, but the effects are often the same as in
purposeful liberations. Care must be exercised
in permitting the entry of cage birds.

Much of the fervor for exotic birds is based
upon the assumption that Hawaii is an "avian
desert.” In part this is true today if one con¬
siders only the native species and the lower,
urbanized parts of the islands. It is not and
never has been true of the higher, forested areas
where some of the native Hawaiian species still

59

60

PACIFIC SCIENCE, Vol. II, January, 1948

exist in considerable numbers. Furthermore, it
is probable that the importation of foreign
species has contributed not a little to this ap¬
parent paucity of the avifauna by driving the
native species farther into the mountains and
perhaps by aiding materially in the extinction of
some species.

This desire to have birds in the Hawaiian
Islands has gone beyond the concept of filling a
vacancy in the lowlands and has proceeded to
extremes as shown by the following statements
of a former official of the Territorial Board of
Agriculture and Forestry in the Paradise of the
Pacific (49 (1): 28, 30, 1937): "We want to
fill the Islands with birds of all desirable species
that will survive here” and, "Hawaii can be
transformed into a universal aviary, a bird haven
where every known species of birds, not in¬
jurious to Hawaii, will propagate and thrive
under the conditions of their natural habitats.”
It is true that these statements are qualified by
the terms "desirable” and "not injurious,” but
there is no positive way to predict the "in¬
jurious” phases of an exotic birds behavior in
its new home. Yet, this same official indicates
quite correctly (op. cit., p. 6) that what is de¬
sirable at one time may not be desirable at a
later date by saying in regard to species wanted
that ". . . things are different in the 1930’s than
in the 1850’s.” He might also have said that
birds desirable to one economic group are not
so to another, and that birds esthetically desir¬
able may not be so desirable economically, and
wice versa. No mention was made of the desire
on the part of some to try to preserve and in¬
crease the populations of native birds, many of
which are peculiar to these islands and, if lost
bere, would be extinct.

The movement for more and more birds has
also been supported by unfounded generalized
statements ( Aviculture , 3:333-334, 1931; and
4:70-71, 1932) that exotic birds are not detri¬
mental to the native species in Hawaii. To my
knowledge no comprehensive study has ever
been made of the interaction of an exotic and a
native bird in Hawaii!

Whether the past introductions of birds were
beneficial or detrimental is beside the point; we
must now accept the successful exotics as a part
of the avifauna; but at the same time extreme
care should be exercised in the importation of
additional species. It is not enough that a bird
is desired by a group or an individual, and that
it appears innocuous in its native ecological
niche. No one can foretell definitely and accu¬
rately the overall activities of a species trans¬
planted to a new region, nor can anyone foresee
all the multitudinous implications of such trans¬
planting. However, as Me A tee (1925: 160)
states ". . . when we consider animals and plants
not strictly domesticated, successful introduc¬
tions have almost invariably had regrettable
consequences.”

Birds that are omnivorous in their original
home need special investigation before being
transplanted. Such birds are often opportunists,
as regards food, and thus feed on the most abun¬
dant and easily obtained food item. This item
may be beneficial insects, a particular farm
crop, or some other article more "valuable” than
the bird itself, even to those who had clamored
for its importation. A species predominantly
insectivorous in its native region may for various
reasons change its food habits so that its major
item of food in its new home is grain, or per¬
haps fruit. Or, if it fed almost exclusively on a
single insect or group of insects in its native
home, it may find that this insect or group is not
as easily obtained as some other insect; it may
then feed on the most abundant food supply.
Consequently, such a bird, when imported for
insect control, particularly of a specific insect,
may be a failure. Yet, it may become firmly
rooted here. Perhaps the search starts again,
and another species is tried, and another, with
varying results, until we have a polyglot fauna
such as is found in Hawaii at the present time.

The breeding potential, or the ability of a
bird to reproduce successfully, is another im¬
portant consideration when a bird is moved into
a new region. We need not concern ourselves
with those which cannot breed successfully. A

Avian Introductions — Fisher

61

species transported to a more equable climate
may show an increased length of breeding sea¬
son; it may nest several times in a year and may
be more successful in raising its young. The
Kentucky Cardinal ( Richmondena cardinalis)
is a good example; in the Territory of Hawaii
this bird breeds throughout the year, and a
single pair has been known to rear three broods
in a year, compared to one or sometimes two
broods in continental United States. Its num¬
bers are increasing remarkably. Other species
show similar trends. Not only is the longer
season important in raising the breeding poten¬
tial. In their native countries most of the species
are partly limited by the amount of food avail¬
able and by the number of species using them
as prey. With an abundant food supply for
many different species, as in Hawaii, and the
possibility of birds adapting their food habits to
the more easily obtained foods, there is a dis¬
tinct probability of enormous increases in the
populations of certain species. This may not be
desirable.

In the islands there is no avian predator to
help control the populations; the only hawk
( Buteo solitarius ), which does not feed on birds
except occasionally, is limited to the island of
Hawaii, and is present in such small numbers
that it is ecologically unimportant. The Short¬
eared Owl (Asio flammeus sandwich ensis) is a
possible predator. It is thought to feed primarily
on mice, but its food habits are largely un¬
known. The only other vertebrate animal species
available to prey on young or adult birds or
their eggs are: feral cats, dogs and hogs, the
native rat ( Rattus hawaiiensis ) , which is not
numerous, several exotic species of the family
of Old World mice and rats which are cosmo¬
politan, and the many mongooses ( Herpestes )
which have to a great extent failed to accom¬
plish the purpose for which they were imported,
that is, to keep the rat populations at a minimal
level. There are those who maintain the mon¬
goose is the major factor keeping the rats under
control, but field observations indicate that large
numbers of rats and of mongooses are present

in the same areas. The tendency of rats to spend
more time in trees may have been occasioned
by the activities of the mongoose. Because of
this tendency to live in trees, the rats are now
more of a menace to the tree-nesting birds;
formerly their predatory activities were more
or less restricted to ground-nesting birds. Aside
from this, the mongoose is probably the most
important control on ground-nesting birds in
Hawaii. This theory is attacked by those who
have examined stomachs of the animal on the
grounds that they do not usually find feathers
or remains of eggs in the stomachs. A study of
the feeding habits of the mongoose has shown
the inadequacy of such an argument because the
mongoose never eats the shell of an egg, and in
eating birds it makes only one or two small
openings in the carcass, works through these,
and leaves the outside of the bird ruffled but
practically intact. Considering this method of
feeding, one could not expect to encounter
numerous feathers in the stomach of the mon¬
goose. The workers who studied the stomach
contents do not mention in their unpublished
studies the fate of the ground-nesting birds of
Jamaica when the mongoose was introduced,
but they do cite the success of the mongooses in
helping to control the rats there.

Whether or not the mongoose fulfilled the
promises of its importers, we now have rats and
mongooses in abundance on the main islands,
except on Kauai, Niihau, and Lanai, where the
mongoose was never liberated. Both animals
exert a depressing effect on the population of
birds.

Avian diseases also aid in controlling num¬
bers of birds, but we know practically nothing
of these in Hawaii. Avian malaria has been
found in the exotic Red-billed Leiothrix ( Leio -
thrix lutea lutea) in Hawaii (Fisher and Bald¬
win, 1947: 51). We cannot say definitely that
this disease was first introduced with imported
birds, but the evidence seems to point that way;
nor can we say definitely that the native birds
have been adversely affected by malaria. For
most animals imported into the Territory a

62

PACIFIC SCIENCE, Vol. II, January, 1948

period of quarantine is imposed to give some
modicum of insurance against the introduction
of diseases communicable to animals and man.
No such provisions are present for avian impor¬
tations as evidenced by the recent arrival
(March, 1947) and almost immediate release
of a number of Mexican Buntings. Holding
wild birds in custody might result in the loss of
some individuals, but it would be preferable to
subjecting native birds and already established
exotic birds to the danger of unknown diseases.
At the very least, representative fecal and blood
smears should be made, and a few birds of each
shipment should be sacrificed for autopsy and
pathological study. It seems insufficent to allow
the entry of birds on the basis of a clean bill of
health as testified by the importer or by a
veterinarian in the country of export.

Except in one or two areas in these islands,
the weather is never a major factor in decreas¬
ing already established populations. In other
parts of the world, inclement winter weather
conditions may act as a severe check, especially
on small birds, by increasing greatly the mor¬
tality rates.

The interaction of the exotics and the native
flora and fauna is a matter of concern and is
unpredictable. Not only this interaction but also
that between the various exotic species estab¬
lished here should be investigated. One example
may emphasize the significance of the interlock¬
ing activities of exotics in Hawaii. Lantana
Camara was brought in as an ornamental plant.
Mynahs ( Acridotheres tristis ) were imported
especially to aid in controlling the army worm
(. Laphygma exempta ). Various species of doves
were established for sundry reasons. Lantana in
its native Mexico is not a pest and mynahs and
doves are not undesirable in their original
homes. The individual importations of these
three organisms seemed harmless. Each impor¬
tation was made for specific reasons, and each
might not have been so detrimental had not
the others also occurred. This is what happened
in Hawaii. The mynah fed on army worms as
was expected, but it also began to feed on

lantana berries, so much so that correlative fluc¬
tuations in the abundance of berries and mynahs
were observed. The army worm is now seem¬
ingly a secondary food item, subject to selection
on the basis of relative abundance. Although
doves may feed on lantana berries to only a
limited extent, it is enough to have been a
factor in spreading the seeds. The seeds pass
through the digestive tract of birds and are
viable. Experimentation with various other
seeds has shown that viability is often actually
increased by the passage of a seed through a
bird. This might be true with lantana. It is
known that other factors were of importance
in spreading lantana, but in this review we are
interested in birds primarily, and they did have
a part in the dissemination of the seeds. As a
result of all factors lantana spread widely and
became one of the most noxious of plants in
the Territory. To curb the spread of lantana
certain insects were brought in. They were suc¬
cessful in part, but it is reported that in areas
where the lantana was eradicated or greatly
reduced another undesirable exotic plant took
over.

The result of establishing the mynah in Ha¬
waii is similar to that obtained in Fiji, as re¬
ported by Stoner (1923: 328-330). In Fiji
the mynah was also imported to control in¬
jurious insects, but, as in Hawaii, it soon began
eating more easily obtained foods, and, in addi¬
tion, was harmful to the native birds. There¬
fore, by 1923, the mynah was considered a pest
and was no longer protected. One important
aspect of Stoner’s observation was that the
mynah in the Fijian area was heavily parasitized.
It is also replete with parasites in Hawaii. Thus,
there is the possibility that species already pres¬
ent in a region may be infected when the
mynah is transplanted there, subject, of course,
to the degree of host specificity of the parasites.
Moreover, on the offshore bird islands of Oahu
the author and his students have found the
mynah pecking open the eggs of Sooty and
Noddy Terns.

Avian Introductions — FISHER

63

The person responsible for introduction of
the mynah is also credited with the importation
of the Rice-bird ( Munia punctulata) , which is
now numerous on all the islands. It became a
pest in the rice fields, but this is a minor dis¬
credit now because of the decline of rice as an
important crop in the islands and not because
of any foresight at the time it was introduced.

The Kentucky Cardinal, imported because of
its brilliant color and song, is now considered
detrimental by some fruit growers. This situa¬
tion, in contrast to the usual situation in con¬
tinental United States, is probably brought about
by the relatively large populations of cardinals,
discussed previously, and the relatively small
acreages devoted to fruits in the Territory. It
simply means that in Hawaii more birds are
present per unit of fruit grown. Consequently,
the percentage of damaged fruit is greater. One
cannot, however, condemn the bird entirely on
this evidence, for it may aid in controlling cer¬
tain insects.

The European Skylark ( Alauda arvensis)
was considered by many to be a pest when it
became established in New Zealand. Although
this bird is present in the Hawaiian Islands, it
seems unlikely that it will become a major
nuisance as long as it continues to frequent open
grassland; on most of the islands the area of
suitable habitat is relatively small and is not in
juxtaposition to truck farms. However, on the
island of Kauai this species is regarded as a
scourge to newly planted lettuce in the truck
farming country. If the skylark should become
more and more reliant on seeds of cultivated
plants, or if the truck farms should expand to
include part of the range of the skylark, or
even extend to the edge of the range, it seems
likely that the species will be on the pest list.

The Red-billed Leiothrix was established for
its pleasing song. As mentioned previously, it
has been found to be a carrier of avian malaria.
It is now considered undesirable by various fruit
and vegetable farmers.

The Linnet ( Carpodacus mexicanus ) has al¬

ready acquired the Hawaiian name of "Ai-
nikana” ( papaya eater ) because of its proclivity
to eat papaya, a staple fruit of the islands. At
present, however, the papayas damaged are
usually those considered overripe for human
consumption.

The ecological balance of the various or¬
ganisms in an area, which is established by
natural processes over long periods of time, has
been so altered in Hawaii that years will be
necessary for any sort of equilibrium to be
reached, and it will never be attained if in¬
discriminate introductions do not cease. Most
of the importations have been of this category.
Little care or thought has been given to the
complexities of an organism’s behavior, other
than that it is esthetically pleasing, or that it
might destroy a certain plant or animal as part
of its activity.

We in Hawaii must change our concepts re¬
garding importations. We must not accept birds
on the simple basis that someone wants them
and that no objection has been raised. We
should express our concern about continued,
unstudied introductions, especially in view of
past experiences here and elsewhere. The re¬
sponsibility for proving the "need” for a certain
species should lie with the importer, and he
should bear the burden of a complete investiga¬
tion by competent authorities over a period of
time. This would prevent many introductions
and would of course make all importations dif¬
ficult. I feel that our general attitude should be
that we want no more exotics.

However, if importations for transplanting
and for cage purposes are to continue, a definite
program of control should be set up. Such a
program might include the following points:

1. The first step toward importation should be
a study of the various aspects of the species
in its native habitat. If the species has been
established in other areas foreign to it, the
effects of introduction there should be noted.
In many instances much information could
be gained simply by a survey of the litera¬
ture on the species involved.

64

PACIFIC SCIENCE, Vol. II, January, 1948

2. As soon as a shipment arrives, representative
specimens of both sexes should be prepared
as study skins for use in positive identifica¬
tion and for future reference.

3. For all birds imported, there should be
recorded: (a) The native home of the
birds. This should be specific, listing coun¬
try, province or district, and nearest town,
and the date on which the birds were first
removed from this area. If, as is true quite
frequently with cage birds, the birds have
been raised in a country other than their
native home, this fact should be recorded.
(b) The number of males and females
should be noted, if it is possible to sex the
birds on the basis of external characters.

4. Immediately after arrival in the Territory,
all birds in any one shipment should be
placed in quarantine, as are other animals,
for a certain period of time, to provide some
protection against introducing parasites and
disease.

5. At the end of the period of quarantine a
general pathological examination should be
performed on a sample of the importation,
and fecal and blood smears made.

6. At the time of release to the importer for
liberation, the exact locality at which the
birds are to be freed should be put on
permanent record. The date of liberation,
the valley, ridge or town, and the island of
release should be noted.

7. In some cases it might be best at first to
limit the liberation of a species to one island
in the archipelago. The effects of the species
on the plants and animals of that island and
its own success in surviving and reproducing

could be studied for a few years. Later, if
desired, it could be moved to other islands.
This procedure might save some of the
islands from the effects of a new pest species.

REFERENCES

Bryan, E. H., Jr., and J. C. Greenway, Jr.
1944. Contribution to the ornithology of
the Hawaiian Islands. Harvard Univ. Mus.
Compar. Zool., Bui. 94 (2): 80-142.

Bryan, William A. 1947. The introduction
of birds into Hawaii. Hawaii. Ent. Soc., Proc.
2 (4): 169-175.

Caum, Edward L. 1933. The exotic birds of
Hawaii. Bernice P. Bishop Mus., Occas.
Papers 10 (9): 1-55.

Fisher, Harvey I., and Paul H. Baldwin.
1947. Notes on the Red-billed Leiothrix in
Hawaii. Pacific Sci. 1: 45-51.

Henshaw, H. W. 1901. Introduction of for¬
eign birds into the Hawaiian Islands with
notes on some of the introduced species.
Hawaiian Almanac and Annual 132-142.

Isenberg, A. H. 1931. Transplanting foreign
birds to the Hawaiian Islands, and other
notes. Aviculture 3: 333-334.

- 1932. Exotic birds not detrimental

to native birds in the Hawaiian Islands. Avi¬
culture 4: 70-71.

Locey, F. H. 1937. Introduced game birds
of Hawaii. Paradise of the Pacific 49 ( 1 ) :
27-30.

McAtee, W. L. 1925. Introduction upon
introduction. Auk 42: 160.

Stoner, Dayton. 1923. The mynah. — A
study in adaptation. Auk 40: 328-330.

NOTES

Observations on Parasites of Domestic Animals in Micronesia

As A PART OF THE MlCRONESIAN EXPEDITION
of the University of Hawaii in the summer of
1946, observations were made by the writer on
parasitic diseases of man and of some of the
important economic animals on the islands of
Ponape and Guam, and on Moen (Truk archi¬
pelago). Brief reports have already been pub¬
lished by the writer on observations of parasites
of man (Jour. Parasitol. 32: 12-13, 1946) and
on murine leptospirosis (Science 105: 236,
1947). The present note summarizes some of
the observations made on parasites taken mostly
from domestic animals.

Parasites of Cattle. Many skin lesions asso¬
ciated with extensive tick infestation were noted
on 12 cows examined in the Net and U districts
of Ponape. The ticks have been identified as
Boophilus annulatus australis (Fuller), carriers
of bovine piroplasmosis or "Texas fever." Re¬
ports ( Civil Affairs Handbook, Office Chief
Naval Operations U. S. Navy, OPNAV P22-5,
p. 129. 1944) indicate that on Ponape many
cattle have in the past died of this disease. The
examination of the feces of six of the above-
mentioned cows failed to reveal presence of
liver fluke eggs. A verbal report from Mr. Oliver
Nampei, resident of the island, indicated that
liver flukes had been noted in cattle on Ponape.
A search by the writer in several fresh-water
streams in the Net and U districts for lymnaeid
snails, known carriers of liver flukes, yielded
negative findings.

The examination of one bull near Agana,
Guam, revealed a moderate infestation of ticks
identified as Boophilus annulatus australis (Ful¬
ler) and Amblyomma cyprium Neumann. Liver
flukes, Fasciola hepatica Linn., are known to
occur in cattle on Guam. The identification was
made by the writer in 1940 from specimens
submitted by Mr. A. I. Cruz of the Guam De¬
partment of Agriculture. The intermediate host
for these flukes on Guam is believed to be the
fresh- water snails Fossaria ollula (Gould),
which were collected by the writer in a swampy
area near Agana. This snail has been experi¬
mentally proved to be a suitable carrier for

65

F. hepatica (Hawaii Agr. Expt. St a. Rpt. 1946:
99, 1947).

Parasites of Swine. On the island of Ponape,
the post-mortem examination of a pig, approxi¬
mately 1 Vi years old, revealed several immature
kidney worms, Stephanurus dentatus Diesing,
within nodules in the mesenteric and perirenal
fat. The liver of the pig showed considerable
discoloration and fibrosis, conditions commonly
seen in kidney-worm infection caused by the
young migrating worms. The large intestine of
the animal also showed a moderate infection of
nodular worms, Oesophagostomum dentatum
(Rudolphi).

On the island of Guam, two pigs, butchered
at the slaughterhouse of the U. S. Commercial
Company, showed infections of lungworms,
Metastrongylus elongatus (Dujardin) , and nodu¬
lar worms, Oesophagostomum dentatum (Ru¬
dolphi). In addition, one of the pigs harbored
a moderate infection of kidney worms, Step¬
hanurus dentatus Diesing. The liver of the pig
showed many white spots and young kidney
worms. The butcher at the slaughterhouse stated
that in his estimation about 50 per cent of the
hogs slaughtered there showed similar kidney-
worm lesions.

The limited observations made on Guam and
Ponape indicate that kidney-worm infection in
pigs may be widespread in Micronesia. Kidney
worms produce serious damage to pigs, and
therefore represent a group of parasites of con¬
siderable economic interest. Since the natives
depend on swine as one of the major sources of
animal protein, it is desirable that proper meas¬
ures be taken to control this parasite. A method
which has been suggested by the U. S. Depart¬
ment of Agriculture (Farmers Bui. 1787) for
kidney-worm control in the southern states con¬
sists in raising pigs on pastures surrounded by
a dry, bare area which allows the action of the
sun to destroy the developing eggs and larvae of
the parasite. This method does not appear ade¬
quate for many areas of Micronesia where rain¬
fall is common and the sky is cloudy or overcast
more than three-fourths of the time. It appears

66

PACIFIC SCIENCE, Vol. II, January, 1948

that the pasture rotation system might prove
more suitable for Micronesia. In addition, young
pigs should be raised on sheltered, dry or con¬
crete floors and after weaning kept away from
older hogs on places that have not been used by
older hogs. To prevent the spread or increase
in the incidence of the infection, it is important
that infected hogs are not shipped to new areas.
Piglets to be shipped to new localities should
come from litters that have been raised on con¬
crete floors or dry areas.

Parasites of Chickens. On the island of
Ponape the following parasites were found in
four adult chickens: (1) proventricular round-
worms, letrameres sp.; (2) cecal worms, Het-
erakis spp. (probably H. gallinae (Gmelin) and
H. lingnanensis Li); (3) tapeworms, Amoe-
hotaenia sp. (probably A. sphenoides (Rail-
liet) ) and Raillietina sp. (probably R. echino-
bothrida (Megnin) ) ; (4) lice, Lipeurus caponis
(Linn.), Menopon gallinae (Linn.), and Oxy li¬
peurus angularis Peters; (5) mites, Pterolichus
ohtusus Robin and Megninia cubitalis (Meg¬
nin). The chicken mite, M. cubitalis, was also
collected from several chickens on the island of
Guam.

Parasites of Dogs. Post-mortem examination
was made on one dog on Ponape. The small

intestine of this animal showed extensive in¬
flammation associated with a large number of
hookworms, Ancylostoma caninum Ercolani. A
few tapeworms, identified as Dipylidium sp.,
were also found in the small intestine. The dog
also showed a light infestation of fleas, Cteno-
cep halides felis (Bouche).

Parasites of Rats. Several mites, Laelaps
echidninum Berlese, were collected from a few
rats trapped on Ponape. No fleas were found on
18 rats which were examined. Of the kidneys
of 22 rats trapped on Moen and 18 rats from
Ponape, 3 and 2, respectively, showed, in stained
sections, presence of leptospirae morphologically
identical to those of Leptospira icterohaemorrha-
giae (Inada and Ido).

The writer wishes to acknowledge the as¬
sistance of individuals who identified some of
the parasites reported above, as follows: ticks
from cattle, C. N. Smith; roundworms and tape¬
worms of chickens, E. E. Wehr; lice from
chickens, C. F. W. Muesebeck and E. W. Staf¬
ford; mites from chickens, E. W. Baker; fleas
from a dog and mites from rodents, C. E. Pem¬
berton; lymnaeid snails from Guam, H. A.
Rehder. — Joseph E. Alicata, University of Ha¬
waii Agricultural Experiment Station, Honolulu,
Hawaii.

Laysan Albatross Nesting on Moku Manu Islet, off Oahu, T. H.

On February 23, 1947, a young albatross was
found among the Red-footed Boobies ( Sula
s. rubripes ) and Sooty Terns ( Sterna f. oahu-
ensis) nesting on Moku Manu, which is ap¬
proximately three-fourths of a mile off Mokapu
Peninsula on the northeastern side of Oahu. At
this time the bird was covered with down and
identification was impossible. By May 10 the
young bird had assumed the characteristics of a
young Diomede a immutabilis, and on July 12,
when the picture was made, there could be little
doubt of its identity. On July 29 the bird could
not be found. Presumably it had left the island,
as do the young at this season on other islands.
This is the first recorded instance of the Laysan
Albatross nesting in the eastern end of the Ha¬
waiian archipelago. Its most easterly nesting
ground has heretofore been thought to be the
Bird Islands, east of Necker Island in the Ha¬
waiian Islands and some 400 miles to the north¬
west of Moku Manu.

Although successive trips were made to the
nest site on March 20, April 19, May 10, 17, 31,
June 14, July 12 and 29, 1947, adult albatrosses
were never observed on the island. A single

adult Diomedea immutabilis was observed May
31 resting on the water just outside the reef at
Kaneohe Bay, some 3 miles from Moku Manu.
On June 14 another adult was observed flying
above Mokulua Islet, off Lanikai, Oahu. These
are the first inshore records of the Laysan Alba¬
tross in the waters surrounding the eastern end
of the Hawaiian chain. — Harvey 1. Eisher,
Department of Zoology and Entomology, Uni¬
versity of Hawaii, Honolulu, Hawaii.

Preliminary Note on the Oceanographic Program
of the Hawaii Marine Laboratory

Dr. Harald U. Sverdrup, director of the
Scripps Institution of Oceanography of the Uni¬
versity of California, was invited to come to the
University of Hawaii for the period November
14 to 22 to advise upon a program of ocean¬
ographic investigation to be undertaken at the
new Hawaii Marine Laboratory. Plans for the
erection of the laboratory building are now com¬
pleted and the physical plant should be ready
for use during the latter part of 1948. Details
on the laboratory and its attendant docks, ma¬
rine railway, outdoor tidal ponds, and aquarium
tanks together with the operating policies will
appear later. The entire physical plant and the
endowment for its operation have been donated
to the University of Hawaii by Edwin W.
Pauley and four associates, Harold Pauley, Allen
Chase, Poncet Davis, and Samuel B. Mosher,
co-owners of the small island, Moku-O-Loe, in
Kaneohe Bay, Oahu, on which the laboratory
will be situated. The University of California
will cooperate with the University of Hawaii in
many of the activities of the laboratory, which
will be equipped for biological as well as ocean¬
ographic research.

Plans outlined for oceanographic research at
the laboratory emphasize those aspects of ocean¬
ography peculiar to this sub-tropical region of
the central Pacific. Broadly speaking, the ocean¬
ographic program may be called a study of the
ecology of the near-shore and off-shore waters
around the islands of Hawaii, and the study is
readily divided into three phases: (1) Ecology
of the near-shore waters, i.e., primarily the
waters inside the reefs; (2) ecology of the
transition region beyond the reef; (3) ecology
of the oceanic waters.

These subdivisions are established for the
following reasons: The very near-shore waters

67

have their endemic faunal and floral populations
but these waters are also the nursery grounds for
much of the food of pelagic fishes. Within many
of the near-shore areas there is a rapid exchange
of water with the outside, and it appears pos¬
sible that a considerable amount of the organic
matter present off the reef area is produced on
and inside the reef. The region directly beyond
the reef is another ecological subdivision, dis¬
tinct in character from the oceanic water at
greater distances from the shore. How far the
near-shore effects reach is unknown. Similarly,
it is not known if the off-shore waters are uni¬
form or if variable conditions related to external
influences such as variable winds or upwelling
are commonly encountered.

Kaneohe Bay is well suited for an intensive
study of near-shore waters. The bay inside the
reef has an area of about 15 square miles, and
is connected with the outside waters by several
channels. This study will involve (1) a pro¬
gram of sounding for the purpose of construct¬
ing a chart of the bottom topography, ( 2 ) the;
collection of bottom samples for the preparation,
of a chart showing the character of the bottom
with particular reference to its biological sig¬
nificance, (3) the recording of tidal observa¬
tions in order to reduce observed depths to a
common level and possibly to give some indica¬
tion of the transport of water across the reef,,
(4) a study of the exchange of water between
the bay and the open sea, (5) a study of the
temperature distribution for the purpose of re¬
lating possible variations to the pattern of cir¬
culation and to the effect of radiation, (6) a
study of transparency by regular observations,
( 7 ) a chemical investigation of salinity, oxygen,
phosphates, nitrates, and, intermittently, calcium,
alkalinity, pH, and other characteristics, all of'

68

PACIFIC SCIENCE, Vol. II, January, 1948

which will be related to the pattern of circula¬
tion and used in connection with a study of the
productivity of the region. The biological work
accompanying this physical and chemical in¬
vestigation will involve, initially, a great deal of
systematic work which will be accompanied by
studies of the production of organic matter and
the relation of the organisms to their physical,
chemical, and biological environment.

The transition zone will be examined by ex¬
tending part of the work outlined for Kaneohe
Bay to the waters just beyond the bay. Addi¬
tional investigation of this area will include
bathythermograph observations.

To provide a basis for planning off-shore
oceanographic work, the Scripps Institution will
examine and discuss the results of numerous

bathythermograph observations which have been
made around the Hawaiian Islands. The sea¬
sonal and geographical variations in the char¬
acter of the mixed top layer will be given
special consideration. The currents around the
islands will be analyzed as far as possible. Such
a study may disclose certain critical localities
which should receive special attention, and the
oceanic work may then be planned to take such
features into account.

It is believed that for some time the oceanic
work will be strictly exploratory. Routine ob¬
servations, including temperature, salinity, minor
constituents, transparency, and biological fea¬
tures, will be made of the character of the sur¬
face layer and of the waters at or directly below
the thermocline.

News Notes

During his visit to Honolulu to advise the
University of Hawaii on establishing the Hawaii
Marine Laboratory, Dr. Harald U. Sverdrup,
director of the Scripps Institution of Ocean¬
ography, spoke on "Wind, Sea, and Swell” at a
public meeting, sponsored by the University and
the Hawaii Chapter of the Society of the Sigma
XL He also conducted a symposium for in¬
terested scientists on the subject "Ocean Cur¬
rents.”

Aboard the auxiliary schooner "Albatross,” the
Swedish Deep-Sea Oceanographic Expedition
visited Honolulu from November 28 to Decem¬
ber 12, 1947. The Expedition, under the leader¬
ship of Dr. Hans Pettersson, possesses equip¬
ment and personnel for making physical, chem¬
ical, geological, and some biological observations
during its world tour. It is following, in part,
the course of the "Challenger” Expedition of
1873-76, and, in the Pacific, has already made

stops at the Galapagos Islands, Tahiti, and Ha¬
waii. From Honolulu the Expedition will go to
Kusaie and Ponape, and from there southward
to the Netherlands East Indies. While in Hono¬
lulu the scientists and the crew of the "Alba¬
tross” were entertained by local scientists and by
Scandinavian members of the community.

Dr. Thomas A. Jaggar’s monumental work,
The Origin and Development of Craters, has
been released as Memoir 21 of the Geological
Society of America. This book records, as the
author states, his "experience in measuring
physical processes at Hawaiian craters and in
mapping changes at the craters themselves,” and
uses this experience as "a reasonable theme
whereon to base a new approach to field vol¬
canology.”

JAGGAR, Thomas A. The Origin and- Development
of Craters. Geological Society of America, Memoir
21. xvii + 508 p., 87 pi., 14 fig. Waverly Press,
Inc., Baltimore, 1947.

APRIL, 1948

)

No. 2

VC~. ~

RVCIFIC

SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

IN THIS ISSUE: Martin — Galapagos Fungi • Hiatt
— Hawaiian Decapod Crustacea • Leopold — Diurnal
Weather Patterns • St. John — Flora and Vocabu¬
lary of Pingelap • Schaefer — Morphometry of
Yellow fin Tuna • Yamashina — The Marianas Mal¬
lard • Brock — A New Blennoid Fish # NOTES-

Published by

THE UNIVERSITY OF HAWAII
HONOLULU, HAWAII

BOARD OF EDITORS

Leonard D. Tuthill, Editor-in-Chief
Department of Zoology and Entomology, University of Hawaii

O. A. Bushnell, Assistant Editor
Department of Bacteriology, University of Hawaii

Ervin H. Bramhall

Department of Physics, University of Hawaii

Vernon E. Brock

Division of Fish and Game, Territorial Board of
Agriculture and Forestry
P. O. Box 3319, Honolulu 1, Hawaii

Harry F. Clements

Plant Physiologist, University of Hawaii Agricultural
Experiment Station

Charles H. Edmondson

Zoologist, Bishop Museum, Honolulu 35, Hawaii
Harvey I. Fisher

Department of Zoology, University of Hawaii

Frederick G. Holdaway

Entomologist, University of Hawaii Agricultural
Experiment Station

Maurice B. Linford

Plant Pathologist, Pineapple Research Institute
P. O. Box 3166, Honolulu 2, Hawaii

A. J. Mangelsdorf

Geneticist, Experiment Station, Hawaiian Sugar
Planters’ Association
P. O. Box 2450, Honolulu 4, Hawaii

G. F. Papenfuss

Department of Botany, University of California
Berkeley 4, California

Harold St. John

Department of Botany, University of Hawaii

Chester K. Wentworth

Geologist, Honolulu Board of Water Supply
P. O. Box 3410, Honolulu 1, Hawaii

Thomas Nickerson, Managing Editor, Office of Publications and Publicity, University of Hawaii

SUGGESTIONS TO AUTHORS

Contributions to Pacific biological and physical
science will be welcomed from authors in all parts
of the world. Manuscripts may be addressed to the
Editor-in-Chief, PACIFIC SCIENCE, University of
Hawaii, P. O. Box 18, Honolulu 10, Hawaii, or to
individual members of the Board of Editors. Use of
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scripts from distant points is recommended.

Manuscripts will be acknowledged when received
and will be read promptly by members of the Board
of Editors or other competent critics. Authors will be
notified of the decision reached as soon as possible.

Manuscripts of any length may be submitted, but it
is suggested that authors inquire concerning possibili¬
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before sending their manuscript. Authors should not
overlook the need for good brief papers presenting
results of studies, notes and queries, communications
to the editor, or other commentary.

Preparation of Manuscript

Although no manuscript will be rejected merely
because it does not conform to the style of Pacific
SCIENCE, it is suggested that authors follow the style
recommended below and exemplified in the journal.

Title. Titles should be descriptive but brief. If a title
runs to more than 40 characters, the author should
also supply a "short title” for use as a running head.

Manuscript form. Manuscripts should be typed on
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cannot be responsible for loss of manuscripts.

Introduction and summary. It is desirable to state the
purpose and scope of the paper in an introductory
paragraph and to give a summary of results at the end
of the paper.

Dictionary style. It is recommended that authors fol¬
low capitalization, spelling, compoundings, abbrevia¬
tions, etc., given in Webster’s New International Dic¬
tionary (unabridged), second edition; or, if desired,
the Oxford Dictionary. Abbreviations of titles o(
publications should, if possible, follow those given in
U. S. Department of Agriculture Miscellaneous Publi¬
cation 337.

Footnotes. Footnotes should be used sparingly and
never for citing references (see later). Often, foot-

{ Continued on inside back cover ]

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

VOL. II APRIL, 1948 No. 2

Previous issue published January 2, 1 948

CONTENTS

PAGE

Additions to Galapagos Fungi. G. W. Martin . 71

Records of Rare Hawaiian Decapod Crustacea. Robert W. Hiatt .... 78

Diurnal Weather Patterns on Oahu and Lanai, Hawaii. Luna B. Leopold . 81

Report on the Flora of Pingelap Atoll, Caroline Islands, Micronesia, and
Observations on the Vocabulary of the Native Inhabitants: Pacific Plant

Studies 7. Harold St. John . 97

Morphometric Characteristics and Relative Growth of Yellow fin Tunas (Neo-

thunnus macropterus) from Central America. Milner B. Schaefer . 114

Notes on the Marianas Mallard. Yoshimaro Yamashina . 121

A New Blennoid Fish from Hawaii. Vernon E. Brock . 125

Notes:

Fishes Taken in Wellington Harbour. W. J. Phillipps . 128

Transfer of Hawaiian Volcano Observatory. Chester K. Wentworth . 131

Interbreeding of Laysan and Black-footed Albatrosses. Harvey I. Fisher . 132

The Poisoning of Bufo marinus by the Flowers of the Strychnine Tree.

Vernon E. Brock . . 132

Pacific Science, a quarterly publication of the University of Hawaii, appears in January,
April, July, and October. Subscription price is three dollars a year; single copies are one
dollar. Check or money order payable to University of Hawaii should be sent to Pacific
Science, Office of Publications, University of Hawaii, Honolulu 10, Hawaii. Reprints
of major articles are available and requests for these will be filled in so far as possible.

Additions to Galapagos Fungi

G. W. Martin1

In CONNECTION with the investigations on
tropical deterioration conducted by the Quarter¬
master Corps of the United States Army, oppor¬
tunity was afforded for a brief visit to South
Seymour Island, in the Galapagos group, in
early September, 1945, in company with Dr. E.
S. Barghoorn and Mr. R. T. Darby.

Traveling by plane from the Canal Zone, we
also made short stops at Salinas, Ecuador, and
Talara, Peru. In all three areas samples of tex¬
tiles, chiefly tentage, paulins, sandbags, and
camouflage cloth, which had been exposed in the
course of service, were collected. All of these
regions are extremely arid, and it seemed worth
while to attempt to learn what fungi had been
able to attack fabrics under such conditions.
That deterioration had occurred was abundantly
evident from the state of the material sampled
and, while the relative importance of biological
agencies as compared with chemical and physi¬
cal factors in causing such deterioration is diffi¬
cult to evaluate, the suggestion is very strong
that in fabrics in contact with or near the soil,
the bulk of the deterioration is due to fungi.

Since cultures were to be made from all sam¬
ples, a supply of previously sterilized test tubes,
bottles, and heavy paper folders was carried, and
all samples were placed in such sterilized con¬
tainers at the time of collection. In each local¬
ity, a few hours were available for miscellaneous
collections, and these also were placed, when¬
ever it was suspected that cultures might profit¬
ably be made, in such sterile packets. The fol¬
lowing account treats only of those samples
taken in the Galapagos.

1 Professor of Botany, State University of Iowa, Iowa
City, Iowa. Manuscript received May 13, 1947.

South Seymour Island is small, roughly trian¬
gular in shape, about 5 miles long and miles
wide in the southern portion, separated from
the much larger Indefatigable (Santa Cruz)
Island to the south by a narrow strait scarcely
Vi mile wide. It is relatively low, although in
the southeast it fronts the sea with precipitous
cliffs arising abruptly for 200 feet. The surface
is extremely irregular, with volcanic boulders of
every size making progress difficult, except on
the excellent roads.

Svenson (1946) has recently published an
extensive account of the vegetation of all three
areas visited, and more than a casual mention
of particular features connected with the fungi
would be superfluous. The average annual rain¬
fall on South Seymour Island is less than 4.5
inches, virtually all of it falling in the first 4
months of the year. Yet, despite this and the
numerous goats roaming the island, vegetation
was surprisingly abundant in early September.
The two most conspicuous plants are Bursera
graveolens (HBK) Triana & Planch., a small,
pale-barked tree, and a columnar-trunked Opun-
tia, presumably O. insularis Stewart, but there
are numerous other woody species, including the
dark-green Scutia spicata (Willd.) Weberb.,
looking like a juniper or yew at a short dis¬
tance, and several legumes, one of which was in
bloom at the time, its bright yellow flowers at¬
tracting numerous bees. Everywhere there is
evidence of what must be a rather abundant
growth of grass in the rainy season.

The only extensive account of Galapagos
fungi appears to be that of Bonar (1939), who
cites the scanty earlier reports (four species
under five names) and reports 59 species and

171]

72

PACIFIC SCIENCE, Vol. II, April, 1948

varieties represented in the material he studied,
one of which is duplicated in an earlier report,
making a total of 62 species or varieties from
the archipelago. None of the species reported
by Bonar was recognized in the collections here
noted.

All Myxomycetes and a majority of the other
fungi were developed in moist chambers in Iowa
City. The collections were removed from their
sterile containers or wrappings, put into sterile
Petri dishes with flamed forceps, wet with ster¬
ile carbon water, and incubated at room temper¬
ature. In several dishes Myxomycetes fruited in
3 to 5 days after wetting. Molds usually ap¬
peared a little later. On the other hand, several
species of Myxomycetes were slow in appearing
but, once started, continued to develop over a
considerable period.

In the listing which follows, species marked
with an asterisk are those which developed in
moist chambers. The numbers given are my own
collection numbers. All specimens are deposited
in the herbarium of the State University of
Iowa. Where material permits, portions will be
distributed to other institutions. A number of
species not listed here are in the hands of vari¬
ous specialists for study.

Acknowledgments: I am indebted to Dr. H.
K. Svenson for determination of host species, to
Dr. G. R. Bisby for determining the Hysterogra-
phium and for comments on other specimens
examined by him, to Dr. L. E. Wehmeyer for
describing and illustrating the new Phaeopelto-
sphaeria, and to Dr. D. P. Rogers for determin¬
ing the Sebacina.

MYXOMYCETES

*Arcyria cinerea (Bull.) Pers.

On dead wood of Bursera, 6314 , 63222; on
thorns of Scutia spicata, 6318. This is the small,
slender, long-stalked phase of this common spe¬
cies which is often encountered in the tropics.

2 Two numbers listed as occurring on the same sub¬
stratum indicate two different collections.

*Badhamia affinis Rost.

On wood of Bursera , 6329. The early fruit-
ings, which began to appear the third day after
wetting, were typical. Later fruitings tended to
be smaller, with smaller spores, relatively longer
stalks, and a somewhat physaroid capillitium,
but the manner of appearance was such as to
suggest that all arose from the same plasmo-
dium, although the plasmodium itself was not
observed.

*Badhamia gracilis Macbr.

On dead stems of Opuntia, 6326. Cacti and
yuccas are favorite substrata for this species.

*Clastoderma Debaryanum Blytt
On wood of Bursera, 6311.

*Comatricha elegans (Racib.) Lister
On wood of Bursera, 6313, 6324.

*Cribraria languescens Rex
On wood of Bursera, 6310. Originally wet on
October 17, 1945, the wood on which this grew
produced this species later in the same month.
Still later, it bore six additional species of Myxo¬
mycetes (6311 to 6316) but no more C. lan¬
guescens until it was allowed to become com¬
pletely dry in January, 1947. It was again wet
with sterile carbon water about March 1, and
by March 6 a typical fruiting had matured.

*Cribraria violacea Rex
On wood of Bursera, 6312.

*Echinostelium minutum deBary
On wood of Bursera, 6313.

*Perichaena corticalis (Batsch) Rost.

On wood of Bursera, 6316, 6323. This spe¬
cies appeared shortly after the wood was wet
and continued to develop singly or in small
clusters for a period of about 3 months. The
majority of the sporangia are characterized by
a prominent circumscissile ridge marking the
line of dehiscence, and this often joins with a
coarse and prominent reticulation on the upper
surface. The spores were at first bright ochra-

Galapagos Fungi — MARTIN

73

ceous in mass, but have tended to become duller
with age. They are uniformly warted and 10-
11 jit in diameter. The plasmodium was dingy
on emergence, becoming dull rose just before
transformation. Numerous mounts have re¬
vealed no trace of capillitium. P. corticalis var.
liceoides G. Lister (1911: 251) was erected for
forms with scanty or no capillitium and with
few granular deposits in the wall of the peri-
dium. Examination of a large series of collec¬
tions from numerous localities shows that these
two characters vary independently and suggests
that the varietal name is superfluous. This is,
of course, even more true of the specific names
which the varietal name was intended to super¬
sede.

*Perichaena depressa Libert

On thorns of Scutia, 6317. Typical, except
that the majority of the fructifications are soli¬
tary and very small, correlated with the small
thorns on which they developed. All are strongly
flattened, with the circumscissile dehiscence
characteristic of the species, and there are sev¬
eral small clusters. This material was wet on
November 28, 1945. The Perichaena began to
appear about a month later, and a few sporangia
were still developing as late as April, 1947. This
period of over 16 months is, in my experience,
by far the longest time during which any col¬
lection has produced myxomycete fructifications.
Also on goat dung, 6309; larger and more clus¬
tered.

* Perichaena vermicular is (Schw.) Rost.

On wood of Bursera, 6327.

*Stemonitis pallida Wingate

On thorns of Scutia , 6319. On wood of Bur¬
sera, 6328.

ASCOMYCETES

*Ascophanus argenteus (Curr.) Boud.
On goat dung, 6282.

*Ascophanus carneus (Fries) Boud.

On goat dung, 6331.

Gloniopsis sp. ( Fig. 2 a-e )

On the dead wood of Bursera there were nu¬
merous elongate black bodies suggesting hyster-
othecia. These were extremely abundant, occur¬
ring on perhaps a majority of the dead branches
seen. It was not until they were examined mi¬
croscopically that it was recognized that three
species were involved. Two were the Hystero-
graphium and the Phaeopeltosphaeria listed be¬
low; the third was a Gloniopsis. The hystero-
thecia (Fig. 2 a) are black, fusoid, and striate,
and most of them appear to be raised well above
the general surface of the wood. A cross section
(Fig. 2b, c) shows that the base is composed of
scarcely altered wood flanked on either side by
a black stromatic layer representing a continua¬
tion of the walls of the hysterothecium. The
subhymenial layer is distinctly thinner than the
hymenium. The latter is composed of densely
compacted, gelatinous, apparently unbranched
paraphyses penetrated by scattered asci in vari¬
ous stages of development, only a few at a time
bearing mature spores. The asci (Fig. 2d) are
short-cylindrical and for the most part 4-6-
spored. The ascospores (Fig. 2e ) are oval, hy¬
aline, muriform or somewhat irregular in their
septation, and extremely variable in size, the
great majority ranging from 25-31/* in length
by 11-1 8/* in width. One ascus was seen con¬
taining but two ascospores, one of which meas¬
ured 58X20/*. A number of species of Gloniop¬
sis with large spores are listed in Saccardo. Of
these G. somala Baccarini (see Saccardo, 1928:
1119), from Italian Somaliland, could represent
this species, and the specimens are provisionally
filed under Baccarini’s name. On dead limbs and
branches of Bursera, 6245, 6254.

Hysterographium mori (Schw.) Rehm
On dead wood of Bursera, 6252, 6255. De¬
termined by G. R. Bisby. Dr. Bisby notes that
No. 6252 approaches H. guaranicum Speg., as

74

PACIFIC SCIENCE, Vol. II, April, 1948

described, but does not believe that it is suffi¬
ciently distinct to be worthy of recognition.

Phaeopeltosphaeria irregularis Wehmeyer,
sp. nov. (Fig. 1)

In superficie caulis maculas dense dispersas,
ellipticas, 1-1.5 mm. longas, 0.5 mm. crassas,
tumidas, nigricantes f ormans; ostiolo centrali pa-
pilliformi vix erumpenti; perithecia 300-550/a
diametro, 200-350/a alta, singula in lignum sub
maculis clypeiformibus immersa; pariete crasso,
prosenchymatoso, ab ligno adjacenti separate;
asci late cylindrici, 90-9 5 /a longi, 12.5/a crassi,
saepe 6-7-spori; paraphyses numerosae, fili-
formes, persistentes, 1 /a diametro; sporae uni-
seriatae, subglobosae vel ellipsoidales, 10.5-18/a
longae, 7-9/* crassae, olivaceae, varie septatae,
1-cellulae vel muriformes, cum .1-3 septis trans-
versalibus, ad septa constrictae, cellulis ali-
quibus verticaliter 1-septatis.

Appearing on the surface as thickly scattered,
elliptic, raised, blackened spots, 1-1. 5X0.5 mm.,
with a central, barely erumpent, papillate
ostiole; perithecia 300-5 50 X 200-350/*, im¬
mersed singly in the wood, beneath a clypeus-
like blackening of the surface tissues; wall
10-20/a thick, prosenchymatous, free from the
surrounding wood tissue which is somewhat
blackened; asci stout-cylindric, 90-95 X 12.5/a,
with a claw-like base, and often with only 6 to
7 spores; paraphyses numerous, persistent, fili¬
form, 1/a in diameter; spores uniseriate, subglo-
bose to ellipsoid, 10.5-18 X 7-9/*, olive-brown,
variously septate, one-celled to muriform with
one to three transverse septa and one or more

Fig. 1. Phaeopeltosphaeria irregularis: a, radial
section of a perithecium showing the clypeate blacken¬
ing of the surface; h, ascospores, illustrating variation
in septation; c, ascus with ascospores.

cells with vertical septa, somewhat constricted
at the septa.

Galapagos: South Seymour Island. On
dead, decorticated wood of Bursera graveolens,
September 6, 1945, 6251 , type.

The genus Phaeopeltosphaeria Berk and Pegl.
(1892:139) was based upon P. caudata, on
woody stems, which might be considered as a
Peltosphaeria with brown spores or a Pleospora
with a clypeate blackening about the peri¬
thecium. It has fusoid spores which are much
larger than those of P. irregularis . Phaeopelto¬
sphaeria panamensis Stev. and King. (Stevens,
1927:50) seems to be the only subsequently
described species. Its spores are described
merely as "muriform, fusiform; olivaceous or
straw-colored; 16x5/*,” but the figures (81-83 )
given show them to be more irregularly 3-
septate than in this species and with tapered
rather than rounded ends. It is also found on
leaves of Chaetochloa, and the authors state that
it resembles the spots of Phyllachora Chaeto-
chloae Stev. on this host. On the basis of this
latter statement, Petrak (1929:387) claims that
P. panamensis is a Pleospora [ Pleospora pana¬
mensis (S. & K.) Petr.] and probably parasitic
in the Phyllachora stroma.

The collection from the Galapagos Islands is
quite distinct from either of these described
species. It is true that there is a variable degree
of blackening of the tissues above the peri¬
thecia of certain species of Pleospora , but if the
genus Phaeopeltosphaeria is to be recognized at
all, this collection is a typical species.

BASIDIOMYCETES
Sebacina petiolata Rogers

On dead wood of Bursera, 6246. Recently
described (Rogers, 1947:99) from Cuba,
Hawaii, and the Marshall Islands. The Gala¬
pagos collection, determined by D. P. Rogers,
was growing at the base of an old dead trunk of
Bursera beneath the soil level and was revealed
only when the trunk was pulled over. On the

Galapagos Fungi — MARTIN

75

basis of examination with a hand lens it was
recognized in the field as probably a Sehacina
and so entered. Early in October it was soaked
and put in a moist chamber and a scanty but
adequate spore-print was secured. Rogers
describes the spores as "evenly oblong to ellip¬
soid-oblong, 9-11X6-7. 5/a, or ellipsoid-sub-
globose, 7-9 X 6-8/a.” This description is in
close agreement with that of spores found in
mounts from the dried specimen. The spores
from the spore-print are almost all globose,
10-11/a in diameter.

The only other Basidiomycetes collected are
a unique, rough-spored Coprinus isolated from
goat dung and a small Pleurotus which appeared
on Bursera wood. Both were secured in pure
culture. The Coprinus fruits readily in culture,
and has been referred to Dr. A. H. Smith for
detailed study. The Pleurotus has thus far failed
to form fructifications.

FUNGI IMPERFECTI

* Helicospor ium guianensis Linder

Referred to this species on the basis of the
yellow color of the conidia in mass; the slender
conidiophores, 4.5/a in diameter below, with a
tendency to slightly swollen, rounded tips; the
branching, bladder-like projections on which the
spores are borne; and the size of the spores.
Differing from the species as described (Lind¬
er, 1929: 280) in the branching of the conidio¬
phores, which is much like that of H. aureum
(Cda.) Linder, from which species it differs,
however, in the more slender conidiophores and
the character of the spore-bearing branches.
Further study may reveal that such forms merge
by imperceptible degrees into H. aureum , but
for the present it seems permissible to maintain
the distinction. On thorns of Scutia spicata.

* Memnoniella echinata (Riv.) Galloway

This widespread species occurred in several
cultures and was particularly abundant on dead
Opuntia stems.

* Tetracrium incarnatum sp. nov. (Fig. 2 f-h)

Sporodochiis pulvinatis, pallide cinnamomeis
vel incarnatis, 0.4-0.8 mm. diam.; conidiophoris
elongatis, tenuatis, basibus 5/a diam., apicibus 2/a
diam., protrudentibus usque ad 80/x; conidiis 3-
8-digitatis, ramis radiatio-cylindraceis, plurisep-
tatis, 45-50/a longis, 3.5-4.5/a latis.

Sporodochia pulvinate to subglobose, at first
white, becoming pale cinnamon, pallid ochra-
ceous or flesh-colored (close to pale ochraceous
buff of Ridgway), 0.4-0.8 mm. in diameter;
conidiophores slender, protruding from body of
sporodochium 60-80/a, 5/a in diameter at base,
tapering to 2/a at apex just below constriction
marking junction of spore; conidia digitate, of
3-8 multiseptate, subparallel, cylindrical arms,
45-50/a in total length, the arms 3.5-4.5/a in
diameter.

GALAPAGOS: South Seymour Island. On dead
stem of Opuntia sp. collected September 5, 1945,
moistened October 25, 1945, developed January,
1947, 6333 , type.

After the appearance of the Myxomycetes
already noted, the material in the moist cham¬
bers became covered with various molds which
soon disappeared and were replaced by a dense
growth of the Memnoniella, which appeared to
cover the substratum completely. It was not
until January, 1947, that the sporodochia of the
Tetracrium were noted, although they may have
appeared earlier. They tended to form at the
tips of spines or other projections. The con¬
spicuously protruding conidiophores made the
sporodochia appear, under the binocular, as
though covered with glandular hairs. Further
examination showed that the Memnoniella had
been almost completely replaced by a Curvularia.

The genus Tetracrium was established by
Hennings (1902: 116) for a fungus from Bra¬
zil occurring on orange leaves covered with in¬
sect larvae. Hennings believed the fungus at¬
tacked the larvae first and then spread to the
leaves and twigs. Although a few mites were
present in the chambers, there was no evidence
of any connection between them and the fungus
here described. Hennings assigned his genus to
the Mucedinaceae. Hohnel (1911:405) re¬
examined Hennings’ material and found that it

76 PACIFIC SCIENCE, Vol. II, April, 1948

Fig. 2. a—e, Gloniopsis sp.: a, habit, from above, X 18; b, cross section showing crust on wood, X 18;
c, cross section with more massive walls, X 100; d, ascus, X 400; e, ascospore, X 1000. f-h, Tetracrium
incarnatum: f, young conidiophore with characteristic tip which will develop into basal cell of spore; g, later
stage with arms of conidium formed but not yet septate; h, four mature conidia; f-h , all X 1000.

was associated with the perithecia of a Putte-
mansia, of which it was obviously the conidial
stage. He transferred the genus Tetracrium from
the Mucedinaceae to the Tuberculariaceae, not¬
ing the number of arms in the conidia, described
as four by Hennings, varied from two to seven.

Saccardo (1906: 560) compiled the name as
Tetracium, and his error is copied by Clements
and Shear (1931). T. Aurantia Henn., the type,
differs from the present species in several re¬
spects, notably in color (white to chalky), in
the much larger spores, and in the very short

Galapagos Fungi — MARTIN

77

conidiophores. Hohnel established a second spe¬
cies, T. coccicola, based on the conidial stage of
Ophionectria coccicola (Ell. & Ev.) Berl. &
Vogl. as described and illustrated by Zimmer-
mann (1901:874). In this species the three
arms are very long, up to 240/x according to
Zimmermann, and at right angles to each other.
Seaver (1909: 198) describes the conidia of the
same species, using the name Scoleconectria coc¬
cicola (Ell. & Ev.) Seaver, as having three to
five arms, each up to 150/* long. It is certainly
distinct from the Galapagos fungus.

The species was readily secured in pure cul¬
ture. It grows rather slowly and fruits sparsely
on most of the ordinary culture media, but
forms good sporodochia on weak malt-extract
agar and on agars prepared from soil-grass de¬
coction and dung decoction.

REFERENCES

Berlese, A. N., and V. Peglion. 1892. Micro-
miceti Toscani. Contribuzione alia flora mi-
cologia della Toscana. Nuovo Gior. Bot. ltd.
24: 97-172.

Bonar, Lee. 1939. Fungi from the Galapagos
and other Pacific coastal islands. Calif. Acad.
Sci., Proc. (IV), 22: 195-206.

Clements, F. E., and C. L. Shear. 1931. The
genera of fungi. 49 6 p., 58 pi. Wilson, New
York.

Hennings, P. 1902. Fungi S. Paulenses I. a cl.
Puttemans collecti. Hedwigia 41: 104-118.

Hoehnel, Franz von. 1911. Fragmente zur
Mykologie XIII. Akad. der Wiss. Wien,
Math. -Nat. Kl. Sitzher., Abt. 1, 120: 379-
484.

Linder, David H. 1929. A monograph of the
helicosporous Fungi Imperfecti. Mo. Bot.
Gard., Ann. 16 (3): 227-388.

Lister, Arthur. 1911. A monograph of the
Mycetozoa. (ed. 2, rev. G. Lister). 302 p.,
200 pi. Trustees of the British Museum,
London.

Petrak, F. 1929. Mykologische Notizen. X.
Ann. My col. 27: 324-410.

Rogers, Donald P. 1947. Fungi of the Mar¬
shall Islands, Central Pacific Ocean. Pacific
Sci. 1: 92-107.

Saccardo, Pier Andrea. 1882-1931. Sylloge
fungorum omnium hucusque cognitorum.

1906. vol. 18. Supplementum universale,
Part VII.

1928. vol. 24 (sec. II). Supplementum
universale, Part X.

Seaver, Fred J. 1909. The Hypocreales of
North America — II. Mycologia 1: 177-207.

Stevens, Frank Lincoln. 1927. Fungi from
Costa Rica. III. Biol. Monog. 11: 1-102.

Svenson, Henry K. 1946. Vegetation of the
coast of Ecuador and Peru and its relation to
the Galapagos Islands. Amer. Jour. Bot. 33:
394-498.

Zimmermann, A. 1901. Einige javanische, auf
Cocciden parasitierende Ascomyceten. Centbl.
f. Baku, Abt. 2, 7: 872-87 6.

Records of Rare Hawaiian Decapod Crustacea

Robert W. Hiatt1

Within the past year four crustaceans which
are rare or previously unknown in Hawaii have
been collected. Since these records are of im¬
portance to both taxonomists and students of
animal distribution, it has been deemed worth
while to make them available. The specimens
are deposited in the collection of the University
of Hawaii Marine Laboratory.

Suborder NATANTIA
Family Rhynchocinetidae

Rhynchocinetes rigens Gordon

Known previously only from the island of
Madeira (Gordon, 1936: 76) and from Ber¬
muda (Burkenroad, 1939: 310; Gurney and
Lebour, 1941: 113), this shrimp is found on and
just off the reef on the south shore of the island
of Oahu, Hawaiian Islands. This species, the
only one of the six recognized members of the
genus known to occur in the Atlantic, has not
been found in the Pacific previously. Since it
has been found in Hawaii, which is the eastern
limit of the most widely distributed Indo-
Pacific species, it is reasonable to suppose that
it occurs in many other Pacific regions west of
Hawaii, but thus far it has not been recorded.
Perhaps the late discovery of this species in an
area in which so much collecting has been done
may be attributed to the fact that here, as in
Bermuda, this species occurs as a sedentary, noc¬
turnal, littoral form and thus would not be eas¬
ily collected unless suitable collecting gear were
used. The smallest Hawaiian specimens were
taken with the aid of a light and a dip net; the
larger ones were taken from a fine-meshed fish

1 Department of Zoology and Entomology, Univer¬
sity of Hawaii. Manuscript received July 3, 1947.

trap set in approximately 6 fathoms off Dia¬
mond Head, Oahu. One female specimen col¬
lected on February 19, 1947, was ovigerous.

Hawaiian specimens ranged in length from
55 to 115 millimeters compared to a range of
from 80 to 97 millimeters for the specimens
from Madeira.

The vertical striae on the carapace are well
developed in both sexes but are most conspicu¬
ous in the males. The third tooth behind the
articulation of the rostrum is largest in all the
Hawaiian specimens, as it is in the holotype.
All Hawaiian specimens have 9 ventral teeth
on the rostrum, whereas the holotype has 8. No
significance may be attached to this feature,
however, because the ventral teeth on the rostra
of the paratypes vary from 7 to 11. The dorsal
teeth of the rostrum are arranged similarly to '
those of the holotype. In the Hawaiian females
the articulation of the rostrum extends less than
half way to the strong lateral ridge, whereas in
the holotype it extends almost to the ridge. In
males from both localities the articulation ex¬
tends to the ridge.

No significant differences were noted between
the appendages of the holotype and those of the
Hawaiian specimens except in the first pleopod
of the females. The Hawaiian specimens have
a broad, scale-like protuberance extending from
the disto-lateral area of the protopodite as far
as the basal one-fourth of the exopod. The lat¬
eral and distal margins of this protuberance are
thickly clothed with setae.

The branchial formula of Hawaiian speci¬
mens is identical with that of the Madeiran
specimens, which differs from the branchial
formula of R. typus M.-Edw. in that the arthro-
branch corresponding to the fourth pereiopod

C78]

Decapod Crustacea — Hi ATT

79

(on XIII.) is absent. The total number of
arthro- and pleurobranchs on IX. to XIV. is
thus 10 instead of 11. Well-developed epi-
podites occur on all the pereiopods with the ex¬
ception of XIV.

Rhynchocinetes rugulosus Stimpson

Although this species is known from the
Hawaiian archipelago through a small specimen
dredged up in 1902 (Rathbun, 1906:911) by
the "Albatross” at French Frigate Shoals in 17
fathoms, and another small specimen taken at
Laysan Island in 1923 (Edmondson, 1925:6),
a specimen taken off Oahu is the first instance
of its occurrence among the main Hawaiian
Islands. The species is apparently widespread
throughout the Indo-Pacific region, having
been first known from Port Jackson, Australia
(Stimpson, 1860:36). Later it was collected at
Lord Howe Island (McCulloch, 1909:310), the
Kermadec Islands (Chilton, 1910:548), the
Loyalty Islands (Borradaile, 1916:85), and in
Japan (Kemp, 1925:263; Kubo, 1936:1887).

One male specimen 80 millimeters in length
was taken in a small-meshed fish trap off Dia¬
mond Head, Oahu, along with two large males
of jR. rigens. This specimen is considerably
larger than Stimpson’s type; it has 5 teeth dor-
sally near the tip of the rostrum as compared
to 3 for the type specimen, and it has 13 teeth
below as compared to 12 in the type. These
differences are probably within the range of
variation for these variable structures. In other
morphological aspects no differences from the
type are apparent.

Two of the six species comprising the un¬
usual genus Rhynchocinetes , in which the ros¬
trum is articulated with the carapace, are now
known to occur in Hawaii. They may be dis¬
tinguished as follows:

1. Two teeth on carapace behind rostral
articulation; no tooth on either side of
fourth or fifth abdominal somites above
posterior edge of pleuron; rostrum with¬
out lateral ridge, articulation with cara¬
pace complete. . . . R. rugulosus Stimpson.

2. Three teeth on carapace behind rostral
articulation; a tooth on each side of fourth
and fifth abdominal somites above pos¬
terior edge of pleuron; rostrum with
strong lateral ridge, articulation with cara¬
pace incomplete. . . . R. rigens Gordon.

Family Gnathophyllidae

Hymenocera elegans Heller

A female specimen measuring 55 millimeters
in length was taken at a depth of 4 fathoms in
a crevice in lava rock encrusted with coral
( Porites lohata ), located about 200 feet toward
Kohala from the new dock at Kawaihae on the
island of Hawaii. This record represents the
second in Hawaiian waters for this bizarre
shrimp, and the first record for the island of
Hawaii. Edmondson (1935:17) collected the
first Hawaiian representative in 1934 among
the branches of a Routes sp. coral head on a
shallow reef in Kaneohe Bay, Oahu. The color
pattern of the present specimen was almost
identical with the one described by Edmondsoa

Suborder REPTANTIA
Family Latreillidae

Latreillopsis hawaiiensis Edmondson

This species, described by Edmondson (1932:
2 ) from a single specimen, has been col¬
lected in Hawaii for the second time. On Janu¬
ary 28, 1947, one of these giant, deep-sea crabs
became entangled in the fish lines of a fisherman
who was bottom-fishing to a depth of 500
fathoms, 5 miles directly off Kewalo Basin,
Oahu, and it was subsequently brought to my
attention. Since the type specimen was taken
at 30 fathoms and the present specimen was
collected at 500 fathoms, it is apparent that the
bathymetric range of this unusual crab is great.

The greatest length of this male specimen is
126 mm.; the greatest width is 111 mm., almost
identical with the type specimen in size. The
rostral spine and the supraocular spines are worn

80

PACIFIC SCIENCE, Vol. II, April, 1948

down greatly. Other morphological features
check with the original description.

Two species of lepadid barnacles thickly
clothe certain parts of the exoskeleton. The
barnacles are grouped closely in the exhalant
branchial grooves and on the mouth parts.
Many others are attached to the pereiopods and
to the carapace and abdomen. These same bar¬
nacles occur on the type specimen, but are much
less abundant.

Two pereiopods, the right third and fifth,
are severed at the fracture plane and the frac¬
tured surface contains only the darkened dia¬
phragm which indicates that the appendages
were probably lost during the ascent of the
animal to the surface. The remaining legs show
no signs of recent regeneration. Round punc¬
tures at which the exoskeleton has been broken
through are present on the merus of the right
cheliped, on the carpus of the left cheliped, on
the merus of the left fourth pereiopod, and on
the merus of the right second pereiopod. Sim¬
ilar punctures were present on the type speci¬
men.

REFERENCES

Borradaile, L. A. 1916. British Antarctic
("Terra Nova”) Expedition, 1910. Zoology
3: 75-110, 16 fig. Nat. Hist. Rpt. Brit. Ant¬
arctic Exped. 1910.

Burkenroad, Martin D. 1939. Some re¬
marks upon non-peneid Crustacea Decapoda.
Ann. Mag. Nat. Hist. XI, 3: 310-318.

Chilton, Charles. 1910. The Crustacea of
the Kermadec Islands. New Zeal. Inst., Trans.
43: 544-573, 4 fig.

Edmondson, Charles H. 1925. Marine
zoology of tropical central Pacific. Bernice
P. Bishop Mus., Bui. 27: ii+148, 11 fig., 11

PL

- 1932. A giant Latreillopsis from Ha¬
waii. Bernice P. Bishop Mus., Occas. Papers
9(24): 1-9, 1 fig., 1 pi.

- 1935. New and rare Polynesian

Crustacea. Bernice P. Bishop Mus., Occas.
Papers 10(24): 1-40, 11 fig., 2 pi.

Gordon, Isabella. 1936. On the macruran
genus Rhynchocinetes, with description of a
new species. Zool. Soc. London, Proc. 1936:
75-88, 7 fig.

Gurney, Robert, and Marie V. Lebour.
1941. On the larvae of certain Crustacea
Macrura, mainly from Bermuda. Linn. Soc.
London, Jour. 41 (277) : 89-181, 26 fig.

Kemp, Stanley. 1925. Notes on Crustacea
Decapoda in the Indian Museum. Indian
Mus., Rec. 27(4): 249-343, 24 fig.

Kubo, I. 1936. On a Japanese shrimp of genus
Rhynchocinetes, R. rugulosus Stimpson. Bot.
and Zool., 4: 1887-1892, 2 fig.

McCulloch, Allan R. 1909. Studies in Aus¬
tralian Crustacea. No. 2. Austral. Mus., Rec.
7: 305-314, 2 fig.

Rathbun, Mary J. 1906. The Brachyura and
Macrura of the Hawaiian Islands. U. S. Fish
Comm. Bui. 1903: 827-930, 79 fig., 24 pi

Stimpson, W. 1860. Prodromus description^
animalium evertebratorum, quae in Expedi¬
tion ad Oceanum Pacificum Septentrionalem,
a Republica Federata missa, Cadwaladaro
Ringgold et Johanne Rodgers Ducibus, ob-
servavit et descripsit. Acad. Nat. Sci. Phila.f
Proc. 1860: 22-47.

Diurnal Weather Patterns on Oahu and Lanai, Hawaii

Luna B. Leopold1

INTRODUCTION

Weather forecasting in the Pacific area
has been predominantly aimed at serving air¬
plane operations. On the other hand, little or
no developmental work has been done to pro¬
vide bases for weather forecasts for agriculture.
In Hawaii the two largest industries are agri¬
cultural-— -the growing of sugar cane and pine¬
apples. Techniques and organization to provide
both long- and short-range forecasts would be
of considerable aid to these industries.

Though a long-continued interest in weather
on the part of agriculturists in Hawaii is shown
by a large number of rain gages and tempera¬
ture measurements, detailed analyses involving
additional critical data are required to give
accurate pictures of the variations of meteoro¬
logical elements over the diverse topographic
areas of the islands. A better over-all descrip¬
tion of these factors is an early step in the
development of a sound basis for local fore¬
casting.

Hawaii, lying in the trade-wind zone, experi¬
ences relatively small, day-to-day variations in
weather when compared with a continental
locality in the belt of westerlies. Yet great dif¬
ferences in ecologic habitats are found within
very short distances. Day-to-day synoptic
changes are subtle and difficult to follow, owing
to the wide expanses of ocean where no upper
air data are obtainable. Particularly in such cir¬
cumstances, the study of diurnal fluctuations may
contribute to a better understanding of the man¬
ner in which the great local differences originate.

Figures 1 and 2 show the location of stations
on Oahu and Lanai which are discussed in this
paper. The profiles drawn in the direction of

^Meteorologist, Pineapple Research Institute of Ha¬
waii. Manuscript received September 11, 1947.

the prevailing trade wind, ENE-WSW, provide
some picture of the topography. On Oahu, the
two ranges of mountains are oriented nearly
perpendicular to the trades, and, therefore, pro¬
vide barriers causing large differences in oro¬
graphic rainfall. These have been discussed in
connection with the mean annual isohyets of
Voorhees (1929) and by Nakamura (1933),
Wentworth (1946), and others.

Acknowledgments: The writer acknowledges
with thanks the help of M. H. Halstead and
Gretchen Hastings, who assisted with the tabu¬
lation. C. K. Stidd contributed in helping with
the drafting of the figures. Charles M. Woffin-
den and the staff of the U. S. Weather Bureau
co-operated by making the special radiosonde
ascents.

DIURNAL RAINFALL PATTERNS

As described previously, rainfall in Hawaii
results primarily from orographic effects on the
trade winds, from frontal passages, and from
easterly waves. Most "kona” storms are actually
related to frontal passages.

Kona weather is a local term which often is
erroneously used to imply a condition of south
wind. A better translation for the word kona is
"leeward,” and in terms of weather, it implies
a cessation of the normal northeasterly trade
wind. This ordinarily causes a strengthening of
onshore sea breezes.

On the southern coast of Oahu, for example,
northeasterly winds blow more or less continu¬
ally during ordinary trade-wind weather in spite
of a tendency for an onshore sea breeze to de¬
velop in the afternoon. When the trade wind
decreases, the sea breeze asserts itself.

On even more sheltered leeward coasts an
afternoon sea breeze is the rule, and a decrease

181]

82

PACIFIC SCIENCE, Vol, II, April, 1948

in trade wind is accompanied by an increase in
the onshore flow.

The trade-wind decrease is commonly asso¬
ciated with the passage of a front or pressure
trough. In such instances, the wind direction
over the ocean tends to change to south or
southwest preceding the pressure trough, and
veering to westerly in the area behind the
trough. The tendency for reversal in wind
direction from the northeasterly trades accounts

for an inflow of warmer air from lower latitudes.

It can be seen, therefore, that "kona” weather
cannot be interpreted as "south-wind weather”
except in particular localities, and it follows
that "leeward” is a more correct interpretation
of the term.

Frontal storms are those related to pressure
troughs in the westerly winds aloft. This west¬
erly circulation is strongest in winter. In sum¬
mer the trade winds reach higher levels as the

Fig. 1. Map of Oahu showing location of stations and day and night wind directions.

Diurnal Weather Patterns — LEOPOLD

83

strength of the easterly circulation around the
Pacific high-pressure cell increases. Pressure
troughs in this system of easterly winds also
move in the direction of the wind, in this case
from the east. These disturbances, called "east¬
erly waves,” provide, in summer, decks of high
clouds as well as an increase in height and size
of the normal fair-weather low clouds of the
ocean.

The importance of convective rain in the Ter¬
ritory has not been sufficiently emphasized.
Rain or showers from cumulus or cauliflower
clouds are often seen over the open ocean or
over the lower and drier portions of the Ha¬
waiian Islands, particularly in the afternoon.

Though well-developed thunderstorms are
rare in the area, convective clouds built up suf¬
ficiently to yield rain are experienced during
periods when the temperature inversion aloft
becomes weak or disappears. The stability of
the layer through the temperature inversion and
the dryness of the air above the inversion
ordinarily limit the height of development of
cumulus clouds.

Convective storms are sufficiently important
to bear a local name, "naulu,” used on the
islands of Maui, Lanai, and Molokai. It refers
to a type of rain which occurs primarily in the
afternoon, and which is characterized by short
duration and high intensity.

Fig. 2. Map of Lanai showing stations and wind directions.

84

PACIFIC SCIENCE, Yol. II, April, 1948

Preparation of Diurnal Rainfall Curves

Because different synoptic situations are re¬
lated to these rainfall types, separation of geo¬
graphical areas over which these types occur is
of some interest. In order to make such separa¬
tion, diurnal rainfall curves were prepared from
recording rain-gage records for six of the sta¬
tions on Oahu and the two stations on Lanai.
These recording gages, with the exception of
that at Honolulu, were installed by the U. S.
Soil Conservation Service about 1941, and are
maintained by the sugar and pineapple planta¬
tions.

Certain lapses in record made it impossible
to analyze exactly the same record period at all
gages, but in general, the period January 1,
1945, to December 31, 1946, was used in all
cases except Honolulu. The Honolulu quanti¬
ties had been counted and tabulated for the
period 1923 to 1941 by H. P. Parker of the
U. S. Weather Bureau, who kindly allowed the
writer to use some of his tabulations for this
analysis.

From the original recorder charts, the rain¬
fall amounts for stations other than Honolulu
were tabulated hour by hour for the period of
record. No attempt was made to adjust the
chart total to the total recorded in the stand¬
ard rain gage which, in most installations, is set
up adjacent to the recorder. It became apparent
at an early stage that a short period of record at
gages in dry localities would not yield statis¬
tically significant results comparing number of
occurrences of various amounts of rain in indi¬
vidual hours. Therefore, all individual occur¬
rences in a given hour were lumped, regardless
of amount of rain which fell during the hour.
"Traces” of rain do not show up on a Ferguson
reconnaissance type gage, so the minimum
amount which was counted as an occurrence
was 0.01 inch. A seasonal breakdown was
chosen of four periods of 3 months each, start¬
ing with December, January, and February as
the winter period.

It was desired to determine whether there

was a significant difference in rain occurrence
at various parts of the day. In Figure 3A is the
histogram of rainfall occurrence for individual
hours at Kawaihapai. This station has a mean
annual rainfall of less than 30 inches. The total
number of rainfall occurrences in the 6 winter
months of the analyzed record (December, Jan¬
uary, and February for 2 years) was 110. It can
be seen that the small number of occurrences
necessitated grouping to bring out significant
differences between various times of day.

For each seasonal group the number of occur¬
rences was computed for all combinations of 8
consecutive hours, i.e., 00002 to 0800, 0100 to
0900, etc. For purposes of statistical analysis,
the day was broken into three 8-hour periods,
one of which provided the maximum number
of occurrences in any 8-hour combination. The
graphs in Figure 3B show the resulting break¬
down for Kawaihapai. The data in Figure 3B
are the same as in Figure 3 A, merely grouped
into 8-hour totals.

Statistical Test for Significance

To determine whether the maximum number
of occurrences in an 8 -hour period was signifi¬
cantly different from the number in the other
periods, a chi-square test was used. Assuming
that the three 8 -hour periods had equal chance
of rain occurrence, the chi-square computation
determined the number of occurrences which
might occur in 8 consecutive hours once in 100
times as a result of pure chance, and again once
in 20 times. If a significantly greater number
of rain occurrences was noted in a given 8-hour
period than might be expected in random trials,
then the original hypothesis of equal chance for
all hours appears untenable. In such cases we
can reasonably assume that a causal factor
operates to provide the observed distribution.
Obviously the statistical procedure does not tell
us whether the period which the data represent
was a good sample of a much longer period.

2 Hour of day in this report is in local standard time
(LST). Times are shown on basis of a 24-hour clock,
thus 1300 is 1:00 P.M.

Diurnal Weather Patterns — Leopold

85

NUMBER OF RAINFALL OCCURRENCES AT VARIOUS HOURS

FIG. 3 A KAWAIHAPAI

INDIVIDUAL HOURS JAN. 1945 - JAN. 1947

J1

_ _ INDIVIDUAL HOURS JAN. 1945 - JAN. 1947 Jj.

AAAjj "Aj A

>H SIGNIFICANCE LEVEL

1 r

FIG. 3B KAWAIHAPAI '

8 HOUR SUMS JAN. 1945 - JAN. 1947

~1

LEVEL

..

I

24 0

12

24 0

12

24 0

~~t — i f

FIG. 3C LEILEHUA

MAY I, 1943- 15

r

12

60

40

20

24

HIGH SIGNIFICANCE 1% LEVEL

SIGNIFICANCE^ LEVEL

.A

80

60

40

20

150

100

O 12 24 0 12 24 0 12 24

DEC.-JAM-FEB. MAR.- APR. -MAY JUNE- JULY- AUG.

HOUR OF DAY

Fig. 3. Number of rainfall occurrences at various hours at three stations on Oahu.

The maximum number of occurrences which
might, by pure chance, fall into a time period
of 8 consecutive hours is plotted as a long-
dashed line in Figure 3B at the ordinate value
computed. The minimum number which might
fall in an 8 -hour group by chance is similarly
represented at the computed lower ordinate. In

the same way the number of occurrences ex¬
pected once in 20 trials is shown by short-
dashed lines at appropriate ordinate values.
These computed numbers required for signifi¬
cance (adjusted X2 = 3.841) and high signifi¬
cance (adjusted X2 = 6.635) obviously depend
in part on the total number of rainfall occur-

86

PACIFIC SCIENCE, Vol. II, April, 1948

fences, and are therefore different for each of
the seasonal graphs.

Inspection of Figure 3B shows that no 8-hour
period had a significantly low or high number
of occurrences in either the winter or fall
periods. In spring and summer, the lower sig¬
nificance limits were just reached but no highly
significant groups were found

On the other hand, at Leilehua, Figure 3C,
highly significant maxima were reached in the
hours 1100 to 1900 in winter, 1000 to 1800 in
summer, and 2200 to 0600 in fall. The low
significance line was reached in the 1000 to
1800 period in spring. None of the 8-hour
groups of Leilehua showed a highly significant
minimum number of occurrences.

Comparing the two stations, it appears that
Kawaihapai has much more uniform distribu¬
tion of rainfall occurrences throughout the day
than Leilehua. The former shows no consistent

tendency for rain in any part of the day. Lei¬
lehua, however, has a strong and significant
tendency to receive a maximum number of rain¬
fall occurrences in the afternoon during all
seasons except fall.

The other stations were analyzed in a similar
manner, and the results are summarized in the
following table. In explanation of Table 1,
Kawailoa Girls’ School, for example, showed a
highly significant rainfall maximum between
the hours of 2300 and 0700 in spring. At that
season a highly significant deficiency of rain¬
fall occurred between the hours of 1500 and
2300. In summer a rainfall maximum occurred
between the hours of 2200 and 0600, while no
8-hour period showed a highly significant defi¬
ciency. Winter and fall had no 8-hour periods
which showed a large enough or a small enough
number of rain occurrences to reach the high
significance limits.

TABLE 1. HOURS OF RAINFALL MAXIMUM AND MINIMUM AT VARIOUS STATIONS

STATION

DISTANCE

FROM

CREST

ELEVATION

OF CREST

ELEVATION

OF STATION

HOURS OF HIGHLY SIGNIFICANT NUMBER OF RAIN OCCURRENCES

Rain occurrences

Winter

Spring

Summer

Fall

Kawailoa ]
Girls’ [

School J

miles

3.0*

feet

3,000

feet

300

(Maximum number
(Minimum number

None

None

2300-0700

1500-2300

2200-0600

None

None

None

Waimea

4.4

1,000

360

| Maximum number
\ Minimum number

None

None

2100-0500

1300-2100

2100-0500

0500-1300

2200-0600

None

Honolulu

5.2

2,500

50

(Maximum number
(Minimum number

0000-0800

0800-1600

0000-0800

0800-1600

0000-0800

0800-1600

2300-0700

0700-1500

Opaeula )
No. 8 j

6.0

1,800

690

( Maximum number
(Minimum number

2300-0700

None

None

None

2300-0700

None

None

None

Leilehua

10.7+

2,800

920

(Maximum number
(Minimum number

1100-1900

None

None

None

1000-1800

None

2200-0600

None

Kawaihapai

13.3+

1,000

200

(Maximum number
(Minimum number

None

None

None

None

None

None

None

None

No. 537 )
Lanai J

2.7

2,000

1,450

(Maximum number
(Minimum number

None

None

None

None

1400-2200

0600-1400

0900-1700

None

No. 5519 ]

Lanai )

3.7

2,750

1,350

(Maximum number
(Minimum number

None

1600-2400

1000-1800

None

1100-1900

1900-0300

0400-1200

None

* On windward side of Koolau Range,
t Situated 2 to 3 miles windward of Waianae Range.

Diurnal Weather Patterns— Leopold

87

In addition to showing the breakdown of
highly significant 8-hour periods given in
Table 1, the Honolulu record was sufficiently
long to show significance in individual hours.
Figure 3D shows the occurrences hour by hour
for Honolulu. This provides a somewhat more
complete picture of the diurnal curve of rain¬
fall occurrence for a station receiving predom¬
inantly trade-wind orographic or nocturnal rain.

At appropriate ordinate values a long-dashed
line shows the number of occurrences necessary
in an individual hour for high significance.
That is, if we assume all hours have an equal
chance for rainfall, there is less than one chance
in 100 that random drawing would provide an
individual hour with a larger number of occur¬
rences. Similarly, the lower dashed line shows
the lowest number an individual hour should
receive in 100 trials of random drawing.

Comparison of Diurnal Rainfall Curves at
Various Stations

It is evident that certain stations have a rain¬
fall maximum during the night, which is gen¬
erally associated with a minimum in the after¬
noon. Stations having nocturnal maxima are
Honolulu, Kawailoa Girls’ School, Waimea, and
Opaeula No. 8. The opposite case is a maxi¬
mum in the afternoon and a minimum during
nighttime hours. This group includes No. 537
Lanai, and Leilehua. Kawaihapai has no sig¬
nificant difference between hours. No. 5519
Lanai shows an afternoon maximum in spring
and summer.

The four Oahu gages which lie near the
Koolau Range, including Kawailoa Girls'
School on the windward side, show nocturnal
maxima. The two gages at some distance from
the Koolau Range-— that is, Leilehua and Kawai¬
hapai — showed either no significant difference
between hours or an afternoon maximum.

The difference is the dominance of orographic
rain from the trade winds as against convective
showers. The latter are an afternoon phenome¬
non while trade-wind orographic precipitation
occurs primarily at night.

Lanai, as an island, is much drier than Oahu
and shows other peculiarities due to the fact
that it is partly in the rain shadow of the much
higher island of Maui. Both recording gages
on Lanai have afternoon rainfall maxima.
Though they are relatively close to the moun¬
tains, which are just upwind, apparently the
overall rainfall deficiency on Lanai due to the
rain-shadow effect diminishes the importance of
orographic trade-wind showers. Opaeula No.
8 lies twice as far from the mountain crest as
the Lanai gages, yet still shows nocturnal
maxima. It will be noted, however, that Opaeula
No. 8 had no highly significant hours of rainfall
minima. This should probably be interpreted
as an indication of diminishing importance of
orographic rains owing to the distance from
the crest.

Again, Leilehua lies only 2 miles upwind of
the crest along the central mass of the Waianae
Range. The station nevertheless shows pre¬
dominantly afternoon rainfall maxima. Kawai¬
hapai lies some distance downwind of the lower
northerly nose of the Koolau Range. It is 2.7
miles upwind of an 1,800 foot crest of the
Waianae Range. Lack of highly significant
periods of rainfall indicates the mixed effect of
orographic and oceanic rain with convective
showers. The explanation might lie in the few
total occurrences of rain, but this was tested by
increasing the length of record and the results
still gave no significant hours.

The diurnal rainfall characteristics of Lei¬
lehua and the Lanai gages emphasize the im¬
portance of the desiccation of air by the barrier
over which the air is forced to rise. The eleva¬
tion of the band of maximum rainfall on East
Maui and on the volcanoes of Hawaii clearly
demonstrates this principle. Yet the dominance
of afternoon or convective rainfall even at the
base of the windward slopes of the Waianae
Range and so close to leeward of the Lanai
Range is a little surprising.

The statistical results presented above from
the limited number of recording rain gages are
borne out by the experience of many observers.

88

PACIFIC SCIENCE, VoI. II, April, 1948

WIND DIRECTION

The diurnal wind patterns for Oahu and
Lanai are shown in simplified form geograph¬
ically in Figures 1 and 2. The full arrows show
the normal daytime surface wind direction, and
the dotted arrows the night winds. The regime
at Waianae is from non-instrumental observa¬
tions only, but the wind changes shown on the
map for other places are all established by re¬
cording wind vanes.

The locations of good records of wind direc¬
tion are unfortunately distributed. A plethora
of records exists in the Honolulu-Pearl Harbor
area, though space does not allow all locations
to be shown on the map of Figure 1. The data
clearly establish the fact that there is usually
no diurnal change in wind direction in the
Honolulu area. This is true also at Aiea and
Pearl Harbor. Yet only a short distance away at
Waipahu a northwest night wind clearly shows
up as the usual thing.

All stations in the southern part of Oahu have
a southerly afternoon sea breeze when the large-
scale pressure gradient is weak and wind speeds
low. These conditions characterize what is
locally called "kona weather” on Oahu, when a
general tendency for light southerly wind per¬
sists.

Kaneohe, on the windward side of Oahu,
shows no diurnal change in wind direction.
Wheeler Field has the most variable wind, but
during the day it is usually northeasterly, and
in the night, northwesterly, north, or sometimes
southerly. Waianae has a very definite afternoon
sea breeze sufficiently strong on most days to
give a westerly onshore wind, in direct opposi¬
tion to the tendency for a trade wind.

It appears that sea-valley and land-mountain
winds affect the wind direction over most of the
westerly third of Oahu. Complete reversal of
wind direction occurs only on the protected lee
coast which lies down-trade wind of both the
two mountain ranges. A nocturnal land-moun¬
tain wind prevails both at Waipahu and Waia-
lua.

A definite sea-breeze front can be found even
on the small island of Lanai nearly every sum¬
mer day. Owing to the low mountain range and
the generally dry weather, little vegetation
grows on the lower parts of Lanai. The leeward
plateau is nearly completely planted to pine¬
apple. This gives rise to strong surface heating
and results in a well-marked sea-breeze regime.
The map in Figure 2 shows the daytime wind
directions and the mean position of a standing
sea-breeze front. Directly above this front is a
daytime cloud which can be seen on many an
afternoon, its position moving a little from day
to day depending on the strength of the sea
breeze. This cloud is certainly the result of ris¬
ing air above the place where the sea and trade
winds meet. A similar cloud line above the
meeting place of sea and trade winds is charac¬
teristic of West Molokai.

The diurnal rainfall regime directly related
to wind patterns is very clear on the big island
of Hawaii. Along the Hamakua coast of the
windward or northeast side of the island, there
is a definite mountain wind at night from the
west, and in early morning a small cloud bank
parallels the coast just offshore. The writer has
noted the sea breeze begin at 1100 at the eleva¬
tion of 4,000 feet near Umikoa, and with this
wind clouds blow in from the coast and rain
begins about noon. The rain or drizzle ends in
the late afternoon and it clears up about night¬
fall, remaining clear at Umikoa all night. The
same phenomenon was noted by Dutton ( 1883 )
in the Kona districts of Hawaii.

SURFACE WIND SPEEDS

Average diurnal curves of wind speed for
various stations are presented in Figure 4.
Kaneohe Naval Air Station is located on a pen¬
insula projecting windward from the island
mass, and approximately represents conditions
over the open ocean to the windward of the
islands. It is apparent that all stations show a
maximum wind speed in the afternoon, but the
island stations show slightly lower maximum

Diurnal Weather Patterns — -LEOPOLD

89

WIND SPEED

SUMMER

FIG. 40

FIG. 4 C HEIGHT OF TEMPERATURE INVERSION

„ — - °^

- *

. JUNE 27, 1!

^ J

^^HYI

POTHETICAL 1

CURVE

o /0”',

MEAN

X

o

-

■ /

/

0

/

+ /

/

/

t

V ©

V

\

V

©

+/!

*Z*JUNE 26, <-7

1

V

MEAN

FOR

24 DAYS WITH INVERSION

MAY 12

TO JULY 9, 1947

X

MEAN

FOR

NOVEMBER 1946

O

MEAN

FOR

SEPTEMBER 1946

m

MEAN

FOR

AUGUST 1946

+

MEAN

FOR

NOVEMBER 1946 AT WEATHER

SHIP “BIRD DOS , 50°N, I40°W LOCAL

TIKE FOR THAT LONGITUDE
MEAN FOR SEPTEMBER 1946 AT WEATHER
SHIP "BIRD DOS’

HOUR OF DAY LOCAL STD TIME

Fig. 4. Diurnal changes of surface wind speed and of the height of the temperature inversion.

speeds than the open ocean (Kaneohe). The
night winds at island stations are of much lower
speed. It is quite clear, therefore, that the
strong diurnal change in wind experienced by
island stations is the result of a nocturnal reduc¬
tion in speed relative to that over the open
ocean.

At Waialua the daytime wind is of moderate
speed from the east-northeast. The nocturnal
wind is light and from the southeast. The vector
which must be added to the trade wind to pro¬
duce the observed night velocity is from the
southwest or from the Waianae Range.

At Waipahu the northwest night wind must
be caused by a vector from the west added to

the trade wind. This westerly component again
comes from the direction of the Waianae Range.

Honolulu experiences no diurnal change in
direction but lower speeds at night than during
the day. A sea-breeze component would tend to
reduce the daytime wind speed while a land
breeze should increase the nighttime speeds.
Therefore, the real winds must be still greater
during the day and less at night than the ob¬
served winds.

Part of the diurnal speed changes at Kaneohe
must be due to sea-land-breeze effects which
would tend to produce the observed winds, with
greater speeds during the day than at night.

Surface friction would account for the slightly

90

PACIFIC SCIENCE, Vol. II, April, 1948

lower daytime speeds over the land than over
the ocean (Kaneohe).

These observations indicate that the diurnal
speed changes cannot be attributed to the sea-
land-breeze regime alone. The effect of sea-
valley and land-mountain winds is apparendy
strongest in the vicinity of the Waianae Range.
This is particularly apparent by the comparison
of Waipahu and Aiea. Part of the explanation
probably lies in the fact that the Waianae
Range, in the rain shadow and much drier than
the Koolau Range, is covered in its lower
reaches by a more sparse and xerophytic vege¬
tation cover than are the moist foothills of the
Koolau. This could give rise to greater heating
and cooling effects on the lower layers of air.

The higher wind speed during the day is
ordinarily attributed to transfer of momentum
from higher wind speeds aloft to lower levels
through increased daytime mixing as a result
of greater instability. In the case of the island
stations, a very stable layer is produced at night
immediately above the ground surface as a result
of radiative heat loss. The development of a
nocturnal surface temperature inversion is ap¬
parent in many individual Honolulu soundings3
and shows up in the mean sounding for January
presented in Figure 5. Obviously this near¬
surface stability will be much more pronounced
over land than over water, and explains the large
reduction of nocturnal speeds over the land.

The depth of the layer of low nocturnal
speeds also points to stability as the explanation.
Figure 6 shows the wind speeds at the four pibal

8 A sounding is a measurement of pressure, tempera¬
ture, and relative humidity aloft. It is plotted in the
form of a graph of temperature vs. pressure as in Fig¬
ure 5. Relative humidity at various pressures is usu¬
ally entered on the graph. A sounding is taken by
release of a small radio transmitter (radiosonde)
carried aloft by a balloon filled with helium. Signals
indicating the three types of measurements are trans¬
mitted by the radio and picked up by a radio receiver
on the ground. The words "radiosonde observation’'
are sometimes abbreviated to "raob.” Such observa¬
tions are made twice daily by the Weather Bureau at
Honolulu.

or rawin4 scheduled times at Honolulu. It is
apparent that the diurnal change aloft is smaller
but in opposite phase to that of surface wind.
The mean thickness of the ground inversion is
roughly the same as the depth through which
the large diurnal wind-speed changes occur.

FIG. 5. Soundings above Honolulu.

Since the strong diurnal change in speed is
measured at all stations, even at Wheeler Field
(elevation 850 feet), and is observed at still
higher localities in central Oahu, it appears that
the nighttime speed reduction is a near-surface
phenomenon. It must occur in a layer some¬
what less than 500 feet thick, just above the
general land surface regardless of the topog¬
raphy. Insufficient observations are available to
determine whether this is true for mountain
crests.

4 "Pibal” is the abbreviation for pilot balloon.
"Rawin'' means upper wind observations by means
of radar. In each case, a balloon filled with helium is
released, rising at a constant rate. Its position in space
is tracked by observing it from the ground, through a
telescope in the case of a pilot balloon, or by a radar
set in case of a rawin. By plotting its position at
evenly spaced time intervals, the movement can be
computed and thus the wind speed and direction at
each level can be determined. Upper air winds are
measured four times each day by the Weather Bureau
at Honolulu, twice by pilot balloon (visual tracking
of the balloon) , and twice by rawin.

Diurnal Weather Patterns — -Leopold

THE TEMPERATURE INVERSION AND ITS
DIURNAL CHANGE

As an example of a summertime sounding,
the September 1 4, 1946, radiosonde flight of
1730 LST (0400 Greenwich time) is plotted in
Figure 5. The base of the subsidence inversion,
marked on the graph, was at an elevation of
5,380 feet above mean sea level. The tempera¬
ture increased 1.6 °C. through the inversion
which is exactly the mean magnitude for 44
soundings in the summer of 1947. Together
with the accompanying rapid decrease in mois¬
ture at or near the same level, the inversion is
apparently quite sufficient to limit the top of
trade-wind clouds. Although direct correlation
of cloud tops with the temperature inversion
has not been made locally by means of airplane-
meteorograph soundings, it is common observa¬
tion that the low-cloud tops in the area are at
quite a uniform level over the ocean even when
they are cumuliform in character. Moreover, the

20

15

1500 FT.

1

1 u

20

15

i

a:

* ,0

1000 FT

£ 20
UJ

a

*0

15

Q

Z

> 1 o

i— ' *

500 FT.

. . J|,

20

15

10

i 5

>

SURFACE

/

<

) 6 12 18 24

HOUR

Fig. 6. Wind speed at Honolulu — surface and
aloft. Averages from pilot balloons for August, 1946.

91

writers own observations indicate that the
height of cloud tops over the mountains is not
much different than the general level of the
cloud tops over the open ocean. The writer
made a rough check with crude thermometers
on the slope of Haleakala, the 10,000-foot
mountain of East Maui. On the road to the
summit, an inversion of temperature of 3°C.
was easily distinguished, and it represented very
closely indeed the cloud tops prevailing at the
time.

Until more careful measurements are made,
it may be assumed that, as in the well-authen¬
ticated situation in Southern California (Nei-
burger, Beer, and Leopold, 1945), the tempera¬
ture inversion is the top of the low cloud deck,
and that the additional height of orographic
clouds over the mountains is not of a large
magnitude.

The co-operation of the.U. S. Weather Bureau
was solicited to obtain additional upper air data
by special radiosonde flights. During a 2 -day
period, June 26 and 27, 1947, four extra flights
were made, which, in addition to the regularly
scheduled radiosondes, provided one ascent
every 6 hours. The heights of isotherms and the
inversion for the 2 days of special ascents are
shown in Figure 7 plotted as a time-height
cross section. On these particular days the sub¬
sidence inversion did not continue unbroken
through the night, but as shown to be common
in Southern California, disappeared at one level
and reformed nearly simultaneously at another.
The nearly isothermal layer at 7,500 feet, 1800
LST June 26, already indicated the beginning
of the inversion which was well established at
7,700 feet at 0530 the following day. On the
26th the inversion base reached its maximum
height at 1030, and on the 27th at about 1500.

The height of the inversion base for these 2
days has been replotted on Figure 4C. The
mean heights of the inversion base for the
months of August, September, and November,
1946, are plotted on the same Figure 4C at
scheduled radiosonde times, 0530 and 1730 LST.

92

PACIFIC SCIENCE, VoL II, April, 1948

Unfortunately, two soundings a day were not
flown during 1946 until November. The mean
heights for a period during the summer of 1947
are also plotted.

Using the average inversion heights at sched¬
uled radiosonde times at Honolulu, the times of
maximum and minimum heights for the 2 days
of special observations can be used to approxi¬
mate a mean diurnal curve of inversion height
which has been drawn on Figure 4C.

Fig. 7. Time-height section above Honolulu Air¬
port, June 25 to 28, 1947.

The heights of the inversion at the weather
ship "Bird Dog” (lat. 30°N., long. 140° W.)
indicate a gradual slope upward to the west from
California to Honolulu, as was noted by Von
Ficker (1937) for the Atlantic and discussed
by Neiburger ( 1945). Using the data on inver¬
sion height at Los Angeles collected by the Cali¬
fornia Stratus Investigation of 1944 (Neb
burger, Beer, and Leopold, 1945), the mean
slope from California to ship "Bird Dog,” a
distance of 1,250 miles, is 1/1600. From "Bird
Dog” to Honolulu, a distance of 1,300 miles, the
slope is 1/3100. These slopes corroborate well
the estimate made by Neiburger and check Von
Ficker’s measurements over the Atlantic.

The stratus ship just off the coast at Los
Angeles showed the inversion base to have its
maximum height about 0630 local standard

time for that longitude, and minimum about
1300. The weather ship "Bird Dog” showed a
mean height of the inversion base higher at
0630 than at 1830, the local times of her radio¬
sonde flights. Without intermediate soundings,
it is impossible to estimate exactly when the
maximum and minimum occur at that location.
The hypothetical mean curve for Honolulu in¬
dicates that the maximum height occurs about
1100 LST and the minimum about 2200. Since
a radiosonde released at the Weather Bureau
Station at the Honolulu Airport drifts west-
southwest with the trade wind over the ocean
during its entire flight, the sounding represents
lee-side conditions more oceanic than insular.
A general check on a daytime maximum inver¬
sion height is provided by observations of
Powers and Wentworth (1941) on the slopes
of Mauna Kea. From Pohakuloa, above the
cloud deck, they noted a daytime increase in the
height of the cloud top, which receded to lower
levels at night.

The diurnal variation in the height of the
temperature inversion was attributed by Nei¬
burger ( 1944, 1945) primarily to the result of
sea-land-breeze effects together with lesser ef¬
fects of advection and surface heating. He pos¬
tulated that the sea breeze increased the wind
speed over the coast causing the inversion over
the coastline to lower during the day, the air
escaping inland through the mountain passes.
Nighttime land-mountain breezes flowing
toward the ocean caused the inversion gradually
to rise during the night. This explanation fits
less well the conditions over Honolulu than Los
Angeles. First, the sea and land breeze does
not appear to be stronger than on the coast of
the continental land mass. The island is small,
and its opposite sides lie within short horizon¬
tal distances. Yet there are no direct indications
of large variations in the inversion over these
short distances. Second, the inversion is consid¬
erably higher over Honolulu than over Cali¬
fornia, which would tend to minimize the
diurnal height changes resulting from sea-land-

Diurnal Weather Patterns — Leopold

93

breeze effects. Yet the magnitude of the diurnal
change in height, admitting a very incomplete
record at Honolulu, appears to be greater at
Honolulu than at Los Angeles. Third, the dif¬
ference in inversion height between scheduled
radiosonde times at the ship '‘Bird Dog” is
greater than that for the west coast and less than
the difference at Honolulu. It is in the same
phase as that at Honolulu. These heights and
those of California stations are summarized in
the following table.

Thus it appears from limited data that the
diurnal curve of the temperature inversion does
not fit easily into the explanation which appears
reasonable for the California coast. There is
apparently a diurnal change in height over the
open ocean, felt by near-coast and insular loca¬
tions, which is not explained by sea-land-breeze
effects.

The same kinds of diurnal temperature
changes in the atmosphere aloft above the inver¬
sion were noted at Honolulu as were described
for California (Leopold and Beer, 1947), i.e.,
an appreciable warming during the daytime.
There is still some question whether this
measured diurnal temperature change in the free
air aloft is real or whether insolational heating
of the radiosonde provides an appreciable por¬

tion of the increase. If these cyclical tempera¬
ture changes prove to be real, they might indi¬
cate a diurnal cycle of vertical motion quite un¬
related to sea-land-breeze effects which would
affect the height of the inversion.

RELATION BETWEEN VARIOUS DIURNAL
PHENOMENA

Loveridge’s (1924) curves for the diurnal
variation of rainfall for Honolulu, 1905-23, are
well verified by Parker’s tabulations (see p. 84)
for the same stations for 1923-41. The occur¬
rence of trade-wind rains primarily at night has
been discussed by Loveridge (1924) and Jones
(1939). The former noted the out-of -phase
relation of surface wind speed at Honolulu and
the rainfall. With the night wind speed ex¬
plained by stability in the lower layers, the ques¬
tion is raised concerning the relation of rainfall
to the temperature inversion. It might be argued
that the lower cloud tops at night which should
accompany a lower nighttime inversion would
reduce the tendency for precipitation at the
same hours. The effect of cloud height alone is
probably minimized by the fact that none of the
clouds providing the trade-wind rain reach into
freezing temperatures.

Wind speeds aloft actually increase some-

TABLE 2. DIURNAL CHANGES OF TEMPERATURE INVERSION—LOS ANGELES TO HONOLULU

SAN

CLEMENTE

ISLAND

STRATUS

SHIP

U.C.L.A.

SANTA

ANA

SHIP

“BIRD DOG”

HONOLULU

Miles from local coast

60 off shore

10 off shore

5 inland

8 inland

0

Maximum height in
feet .

2,300

1,600

1,850

1,600

8,400

Minimum height in
feet .

1,700

1,100

1,200

750

6,300

Time of maximum
height .

0630

0630

1100

0800

1100

Time of minimum
height .

1130

1300

2200

1930

2200

Difference in height
0530-1730 LST feet

300

400

200

650

550*

540f

700

* Ship "Bird Dog” data for November, 1946; f '’Bird Dog” for September, 1946.
Notes: Honolulu data for November, 1946, and May-June, 1947.

California data for "ship period,” 2 weeks in September, 1944.

Times are local standard for respective longitudes.

94

PACIFIC SCIENCE, Voi. II, April, 1948

what at night and are directly out of phase with
inversion heights, as can be seen by comparison
of Figure 4C and Figure 6. Though the aver¬
age increase is small, it appeared in September
data as well as in the August data presented in
the figure. That these changes of wind speed
are related to the lower inversion at night is in¬
dicated by a simple calculation of hydraulics.
Using the shape of the curve of Figure 4C and
the plotted points showing mean heights of the
inversion base for individual months, the maxi¬
mum and minimum heights of the inversion
were estimated for August and September, 1946.
The diurnal curves of wind speed at various
levels aloft presented in Figure 6 for August,
1946, were similarly computed for September,
1946.

Assuming no compression, the mean wind
speed below the inversion should increase as the
inversion decreases in height if energy is to be
conserved. The ratio of wind speeds should be
the same as the ratio of inversion heights if the
inversion is a surface through which parcels of
air do not pass. Table 3 presents these ratios.

The diurnal curves of wind speed could not
be drawn for elevations above 2,000 feet be¬
cause clouds limited the height of pilot-balloon
observations. Nevertheless, the ratios are suffi¬
ciently close, considering the limitations of the
data, to indicate that the diurnal variations of
wind speeds aloft are the result of diurnal
changes of inversion height as would be ex¬
pected from theoretical considerations.

Loveridge (1924) and Jones (1939) attrib¬
ute the nocturnal rainfall at Honolulu to the
radiative cooling at the top of the clouds.

Soundings from Honolulu Airport are too far
from the mountain crest to be representative of
conditions in the orographic clouds. However,
some nocturnal cooling at all levels seen in the
Honolulu soundings is probably also true Over
the mountains with additional cooling near
cloud tops. Cooling at all levels implies a lower
lifting condensation level or a lower cloud base
at night than in the daytime. This is verified
by observation. Higher nocturnal wind speeds
aloft probably mean more turbulence and larger
droplet size. All these factors would tend to
provide a nocturnal maximum of rainfall.

In so far as the city of Honolulu is concerned,
many rain showers result from the blowing of
rain droplets considerably leeward of the edge
of the cloud producing them. Higher wind
speeds would again tend to a nocturnal rainfall
maximum in the city.

SUMMARY

The importance of convective shower activity
in areas leeward of the main zone of orographic
rainfall has not hitherto been brought out.
Afternoon maxima of rainfall are observed in
the center of the leeward plateau of Lanai and
along the west edge of the Wahiawa saddle of
Oahu. This implies that convective showers are
an important source of moisture in many of the
drier parts of the islands where only a moderate
part of the moisture has been dropped from the
air as a result of prior orographic lifting. It is
likely that too much moisture has been extracted
to give many local convective showers in the
Lualualei area, though no gages were available
there for analysis. Such areas must depend on

TABLE 3. RATIOS OF MAXIMUM/MINIMUM INVERSION HEIGHTS AND RATIOS OF WIND

SPEEDS, HONOLULU

MAXIMUM AND

RATIO MAXIMUM/

RATIO OF MAXIMUM/

PERIOD

MINIMUM INVERSION

MINIMUM INVERSION

MINIMUM WIND SPEEDS

HEIGHT IN FEET

HEIGHT

AVERAGE 500-2,000 FEET

August, 1946 ....

;

o o
o o

1.39

1.23

September, 1946 . . .

f 8,000 l

l 6,000 j

1.33

1.44

Diurnal Weather Patterns-— LEOPOLD

95

the less frequent kona storms for important
sources of rainfall

, The subsidence inversion of temperature oc¬
curs at higher elevations over Honolulu than
over Los Angeles. The inversion has a diurnal
change in height similar to certain coastal sta¬
tions in Southern California. The local sea breeze
shows up only on the lee or well-protected parts
of Oahu and Lanai. Because of the small size of
the islands and the considerable height of the
inversion, diurnal changes in this height as a
result of convergence in the sea breeze seem
a less likely explanation for Honolulu than for
Los Angeles.

Diurnal changes in surface wind speeds are
consistent over the islands. The nocturnal speeds
are very much less than those over the open
ocean. This is apparently explained quite ade¬
quately by nocturnal stability in the lower layers.

Wind speeds aloft increase slightly at night
and the magnitude of this increase corresponds
to that which would be expected by the changes
in height of the inversion.

REFERENCES

Dutton, C. E. 1883. Hawaiian volcanoes.

U. S. Geol. Survey, Ann. Rpt. 81-219.

Jones, S. B. 1939 The weather element in

Hawaiian climate. Assoc. Amer. Geog., Ann.
29: 29-57.

Leopold, L. B., and C. G. P. Beer. 1947. The
coastal sea breeze in relation to diurnal tem¬
perature changes in the lower atmosphere.
Amer. Met. Soc., Bui. 28 (8) : 371-380.

Loveridge, E. H. 1924. Diurnal variations of
precipitation at Honolulu. U. S. Monthly
Weather Rev. 52: 584-585.

Nakamura, W. T. 1933. A study of the vari¬
ations in annual rainfall of Oahu based on the
law of probabilities. U. S. Monthly Weather
Rev. 61: 354-360.

Neiburger, M. 1944. Temperature changes
during formation and dissipation of west
coast stratus. U. S. Weather Bur., Res. Paper
19, Washington.

— - C. G. P. Beer, and L. B. Leopold.

1945. The California stratus investigation of
1944 . 84 p., 35 fig. U. S. Weather Bur.,
Washington.

Powers, W. E., and C. K. Wentworth. 1941.
Air movements in the Mauna Kea-Mauna
Loa saddle, Hawaii. Amer. Met. Soc., Bui.
23: 6-13.

Von Ficker, H. 1937. Die Passatinversion.
Veroffentl. d. Met. Inst., Univ. Berlin, Bd. 1,
Heft 4.

Voorhees, J. F. 1929. A quantitative study of
the rainfall of the island of Oahu. App. A,
Sup. to Dept. Honolulu Sewer and Water
Comm., Rpt. 293-302. Honolulu.

Wentworth, C. K. 1946. Geographic varia¬
tion in annual rainfall on Oahu. 14 p., 4 fig.
Hawaii Univ., Res. Pub. 22. Honolulu.

96

PACIFIC SCIENCE, Vol. II, April, 1948

Fig. 1. Map of Pingelap Atoll.

Report on the Flora of Pingelap Atoll, Caroline Islands, Micronesia, and
Observations on the Vocabulary of the Native Inhabitants:

Pacific Plant Studies 71

Harold St. John2

INTRODUCTION

The scientific literature concerning the bot¬
any of the Caroline Islands, Micronesia, is al¬
ready of considerable extent. It includes check
lists and ecological accounts of most of the high
islands, a check list of Micronesia, and a floris-
tic treatment of the woody plants. In the Caro¬
line Islands only five island groups contain high
islands. These are Palau, Yap, Truk, Ponape,
and Kusaie. They have extensive floras, and as is
natural, these have received the most intensive
botanical investigation.

The atolls and low coral islands are much
more numerous in the Carolines than are the
high islands. These single coral islands or island
clusters are 43 in number. Strange as it appears,
no detailed report has been published on the
flora of any one of these low islands. During
the Christmas period of 1945 the writer led a
four-man mission from the University of Hawaii
on a 3 weeks’ scientific reconnaissance of Micro¬
nesia. It was made possible by the courtesy and
assistance of the United States Navy, which pro¬
vided transportation by airplane and ship, and
other facilities.

While returning from Kusaie to Ponape on
board the navy vessel LCI 567, it was possible
to make a brief stop on December 27, 1945, at
Pingelap Atoll, which lies about halfway be¬
tween the two larger islands. It was stormy

1 This is the seventh of a series of papers designed
to present descriptions, revisions, or records of Pacific
island plants. The preceding papers were published
as: Bernice P. Bishop Mm Occas. Papers 17(7),
1942; 17(13), 1943; 18(5), 1945; Amer. Pern Jour.
35* 87-89, 1945; Toney Bot. Club, Bui 73: 588,
1946; Pacific Sci. 1: 116-118, 1947.

2 Chairman, Department of Botany, University of
Hawaii. Manuscript received September 5, 1947.

during the night voyage, and this bad weather
delayed the landfall from dawn to midmorning.
The sky was murky and one rain squall after
another drove across the sea, greatly reducing
visibility. Nevertheless, the miraculous radar
enabled the navigators to pick up and locate the
island and approach with assurance, till it
loomed up a mile ahead as a low dark line on the
gray sea. Circling the south end the vessel ap¬
proached and lay to off the western shore of the
larger and southernmost islet, Pingelap Island,
just opposite the single village.

Ready and eager to get ashore, the writer
climbed down a rope ladder and dropped into
the first boat to come alongside. It was a trim
and slim two-man outrigger canoe. It was large
enough so that even with an extra passenger
there were still several inches of freeboard, and
the trip to the shore was made without bailing.
The reef was a shelving one, extending far out,
but submerged enough so that the canoe easily
floated all the way to the beach. Of the two
paddlers, the one in charge was a sturdy, elderly,
white-haired man named Soas. Both were eager
for the cigarettes offered them, but the driving
rain prevented their being lighted.

Our ship was the second to visit the island in
4 years. Three months before, a United States
Navy ship had repatriated some seventy-five of
the men who had been working for the Japanese
armed forces as forced agricultural laborers on
the plantations on Ponape. They returned in
want of new clothing and goods, to find their
families and neighbors in similar need. Many
men, women, and youths had for clothes only
a few ragged bits of cloth. The most needy were
clothed in girdles of leaves. Soas appeared with

197}

98

PACIFIC SCIENCE, Vol. II, April, 1948

ragged shorts partly covered by a skirt of leaves,
but later he changed to a better garment.

As the little canoe grounded on the beach, a
host of people advanced. It seemed as if an
interminable function of handshaking was im¬
minent, but it was possible to limit it to a few of
the elders. A chief controlled them and lined
them up. The group of some two hundred sang
a hearty song of welcome. It was a warm and a
stirring reception. Lest the impression be
gained that the reception was a great personal
triumph, it should be made clear that curiosity
was enough to bring many to the beach, and the
hope that the stranger landing on the beach was
a trader bringing cloth, thread, knives, and other
goods, was a strong motive to bring out the
people. Unfortunately, the few articles, knives,
cloth, chewing gum, and cigarettes, carried by
the botanist were only enough for the guide and
his family.

The village ( see map, Fig. 1 ) stretched along
a single straight street starting from the
bombed church at the south end and running
northward parallel with the west beach. Most
of the homes were frame structures, but a num¬
ber were of thatch, as were all the outbuildings.
Fruit trees and ornamental shrubs and herbs were
numerous in the village, which was well kept
and attractive. Not many other food plants were
cultivated in the village. Fecundity of the people
was evident, for small children appeared in
swarms, and had to be carefully dodged when
one walked in the village.

East of the north end of the village and about
midway across the island was a large swampy
depression, at least 300 feet wide and 600 feet
long. This had been converted into a "lepuel”
or wet garden. Each family controlled a lot in
it, for wet-land agriculture. At a glance it ap¬
peared to be a solid growth of Cyrtosperma
Chamissonis, the most important starchy food
crop. The plants were of fair size in the black,
wet muck, reaching about 8 feet in height.
These were not more than 2 years old. If al¬
lowed to grow to maturity at 3 years, they more

than double in height, and produce an enor¬
mous corm. Soas said that the largest corms were
2 feet in diameter and 5 feet long and so heavy
that two men were needed to carry one. Infre¬
quent in the patch were plants of Colocasia
esculenta, Musa paradisiaca, and Saccharum offi-
cinarum.

The whole island was wooded. Native plants
were not rare, but the forest stand was of Cocos
nucifera, which had been planted to produce
copra for trade with foreigners. These coconut
trees made an even canopy which, at 73 feet,
dominated the scene.

Second in importance as a starchy food was
Tacca Leontopetaloides, called "mugamuk.” This
was stated to be planted in garden patches. It
also occurred widespread throughout the coco¬
nut plantation, where it was self-sown. From
small tubers it persisted and apparently it spread
also by seed, which may have been broadcast. It
was certainly not indigenous, and did not grow
on the top of the beaches or in any close prox¬
imity to them.

The shallow margin of the lagoon was note¬
worthy, for there were several large patches of
mangroves, both Rhizophora mucronata and
Sonneratia alba.

PINGELAP LANGUAGE

No published or other record has been found
of any compilation or study of the Pingelap lan¬
guage. One would expect it to be similar to the
tongues spoken on either one of the adjacent
large islands, Kusaie or Ponape.

The writer’s boatman, named Soas, spoke a
little English, so he became guide and informant
during the brief but vigorous exploration of
Pingelap Island. Even without an interpreter or
a common language, it is possible to obtain the
native names of plants from a good informant.
The writer has succeeded in doing so on numer¬
ous Pacific islands. Soas furnished the names
for every plant collected and for others merely
observed. The crowds of bystanders were asked,
and they confirmed many of the names Soas sup-

Flora of Pingelap— St. John

99

plied. As is characteristic among unspoiled Poly¬
nesians, Melanesians, and Micronesians, from
childhood on, every person knows the name and
uses of essentially every plant in the flora. For
several of the plants information was obtained
as to their economic or ethnic uses. For some of
the economic or crop plants, such as Cyrto -
sperma , Colocasia, and Pandanus, this informa¬
tion was extensive and detailed. The natives
recognize, name, and keep distinct numerous
cultivated varieties. Names for these varieties,
as well as for the species themselves, were ob¬
tained. Together, they make a total of 80
names, and as examples of the Pingelap lan¬
guage have some importance. Each one has
been studied to determine whether it appears
in identical or modified form as a descriptive
word or phrase in the languages of the Mar¬
shalls, Kusaie, Ponape, or Truk. Surprisingly,
this study shows little in common with any of
these languages, and the few identities seem
mere coincidences. For instance, "mesawsol” is
a cultivated variety of Colocasia in Pingelap,
while in Kusaie the word "meza-oual” (Lesson,
1839:516) means the last quarter of the moon.
Some community of significance is possible but
seems improbable. This was the only word that
seemed to suggest an identity.

ALPHABET

The plant names were recorded as heard. No
preconceived theory of the language was used
or convention adopted that one letter should
represent several different sounds. The words
were written down as they sounded to an
American. No difficulty was experienced in re¬
cording the consonants, but a little was with the
vowels. As indicated in the following table,
vowels without any mark represent a long vowel,
while the short vowels were marked, as in a.

Vowels used in Recording of Pingelap
Vocabulary
a— -as a in father
a— as a in h^zt
e— as a in s^y
e — as e in bet

i — as ee in keep
i — as i in bit
o— as o in snow
6— -as o in pop
u — as u in r^le
u—~ as u in duck

ETHNOBOTANY

The flora of Pingelap Island, as here re¬
corded, includes 57 species, falling into the fol¬
lowing groups:

Indigenous . 32

Crop plant and cultivated or introduced
economic trees ...................................... 12

Ornamentals . . 10

Adventive weeds . . 3

Total 57

Of the 57 Pingelap plants all but five are now
known to occur on the Marshall Islands. The
ones lacking are nos. 19, 21, 23, 26, and 29 of
the list which follows. Though the Marshalls
lie some distance to the east, the nearest,
Ujelang Atoll, being 243 miles to the north,
and the most remote, Pokaakku, being 805
miles away, still they are all atolls or coral
islands, and, as is well known, have a flora
mostly of wide-ranging species.

Of the 57 Pingelap plants, 42 are also known
on Kusaie. The species missing there are nos.
8, 10, 12, 17, 21, 23, 26, 28, 32, 43, 44, 45, 48,
50, and 52. Four of these are cultivated orna¬
mentals, and one an introduced weed, so their
absence is not significant. Then, too, the flora
of Kusaie is not as completely known as that of
some of the other Caroline Islands.

Of the 57 Pingelap plants, 41 are also known
on Ponape. The species missing are nos. 8,
10, 12, 16, 17, 19, 23, 25, 26, 27, 28, 32, 34,
45, 48, and 50. These include three cultivated
ornamentals and one introduced weed.

Of the 57 Pingelap plants, 46 are also known
on Truk. The species missing are nos. 6, 7, 8,
17, 24, 25, 26, 27, 36, 48, and 52. These include
two cultivated ornamentals and three weeds.

It is quite possible that more intensive explo¬
ration and collecting on Kusaie and Ponape will

100

PACIFIC SCIENCE, Vol. II, April, 1948

reveal the existence of some of these species,
reducing the list of the missing ones.

Some of the plants of Pingelap Atoll have
vernacular names that are unique and local.
Other plants, common to several of the islands,
have names that are identical or so similar that
they are doubtless linguistic variants. These
are distinguished by italic type in the tabu¬
lation that follows.

Names preceded by an asterisk were recorded
in the field by the writer. The others are com¬
piled from sources listed in the bibliography.
Those names were obtained over a span of
more than a century and were recorded by
French, German, Japanese, and American ex¬
plorers who all used their own orthography, yet
the homology of their rendering of the vernac¬
ular names is striking and significant.

TABLE OF VERNACULAR NAMES ON PINGELAP AND ADJACENT ISLANDS

PLANT SPECIES

ISLANDS AND LIST OF VERNACULAR PLANT NAMES

Pingelap

Ponape

Truk

Kusaie

Marshall Is.

Asplenium nidus

♦seilik

ngok, nuk, nuk

moilukluk,

♦mueyliklik

♦kartep,
karatup, ardap

Nephrolepis

biserrata

♦pues

rawtil

emere, amare

♦bairik

Polypodium

Phymatodes

*kiteu

kiteu, kitiu,
kitheu, ketdu

onnum, chichi,
chiji, sichon

kemkem, kilm,
klim

♦kino, ♦kwino

Vittaria elongata

♦lit

♦wujoet

Pandanus sp.

*kipai

kipar, kipal, taip

kepar, fadj , fach

me-ale, *muang

♦bop, bob

Thalassia

Hemprichii

*walat

oldt

♦wujoet in
loidjit

Eragrostis

amabilis

♦rosakai

♦wujoet,

♦wujoTch,

♦wujues,

♦ujoet,

♦ujoich, ujos

Lepturus repens

♦ro sakai

♦wujoet,
♦wujoich bugur,
♦wujues,

♦ujoet,

♦ujoich,
ujoj, ujuj

Saccharum

officinarum

*seu

tseu, won

wou, uou

*taoh, ta

*to, *dau

Thuarea involuta

♦mokarak

unnom, unom

♦wujoet,

♦wujoich,

♦wujues,

♦uyoet,

♦ujoet,

♦ujoich,

♦ujotch, ujuj,
ujos maroro,
♦kakumkum

Cyperus

javanicus

*sdpasdp

use

nikaunoim,
amana, moirer

sapasap, *ujoet,
♦wujoet,

♦wujoet in ion
buil

Fig. 3. South end of lagoon, showing seedlings and forest of mangrove, Rhizophora mucronata.

PHOTO BY H. ST. JOHN.

FIG. 4. Guide Soas holding penduncle,
and standing breast high to the leaf of
the starchy vegetable Tacca Leontopeta-
loides or "mugamuk.”

PHOTO BY H. ST. JOHN.

Fig. 5. Wet-land garden or "lepuel” of Cyrtosperma Chamissonis or "Muiang.

PHOTO BY H. ST. JOHN.

Fig. 6. Village street on Pingelap Island, showing plants, from the left, Pandanus sp. or "kipai”; Nephrolepis
bisenata or "pucs"; Carica Papaya or "kaineap”; and Cocos nucifera or "ni.” PHOTO BY H. ST. JOHN.

Fig. 7. Family of natives, showing the guide Soas (man at right) and his relatives. PHOTO BY H. ST. JOHN.

Fig. 9. Natives on beach at landing. PHOTO BY H. I. FISHER.

Flora of Pingelap-— St. John

101

TABLE OF VERNACULAR NAMES ON PINGELAP AND ADJACENT ISLANDS ( Continued )

PLANT SPECIES

ISLANDS AND LIST OF VERNACULAR PLANT NAMES

Pingelap

Ponape

Truk

Kusaie

Marshall Is.

Fimbristylis

cymosa

*rosakai

fedil, puker

*berelitchman,

*pererlitchman,

perelejman,

peri j man,

berej isman,

*dilitchman,

*drelisman,

*drelitchman,

*merelijiman,

*malelitchmar,

*uioet

Cocos nucifera

*ni

ni

nu

*nu (drinking
nut), non,
kwanu,

*kaenu

*ni

Colocasia
esculenta
var. antiquorum

*sawa

sawa, tsaua

sawa, sarawai,
onni

*katak, taka

*katak

Cyrtosperma

Chamissonis

*maiang

mwong

pashon, fanan,
pula

*pashok

*iaratz, *iaratch,
iaraj

C. Chamissonis

var.

*muiang an
Ngatik

mwang en Natik

C. Chamissonis
var.

*stmidm

simiten

*simenton

Crinum

asiaticum

*kiep

kiop, kiaup, kip,
sip

*kiep, *guiep,
gib

Zephyranthes

rosea

*kiep

Tacca Leontopet-
aloides

*mugamuk

mokomok,

mokamok,

mokintok

mokomok,

mokumok,

makmok,

mokemok

*mokmok

*magamuk,

* mag amok,

*makamuk,

makmok,

makmok,

mokemok,

mokamok

Dioscorea sp.

*kep

kep, kaapwalap

ep, ampul

*ohkani

*mata

Musa paradisiaca

*wis (on

Woleai I.
called : wiss)

oio, ut

uch, udj

*ousch, eusr,
oune

*kebrang,

kebreng,

kabrang,

kabiran,

*binana

(—banana)

M. paradisiaca
var.

* Taiwan

taiwang

* T aiwan banana

M. paradisiaca
var.

*Amerika

Amerika

M. paradisiaca
var.

Hakatan

nakatan

*lakatan

102

PACIFIC SCIENCE, Vol. II, April, 1948

TABLE OF VERNACULAR NAMES ON PINGELAP AND ADJACENT ISLANDS ( Continued )

PLANT SPECIES

ISLANDS AND LIST OF VERNACULAR PLANT NAMES

Pingelap

Ponape

Truk

Kusaie

Marshall Is.

Peperomia

ponapensis

*warin

(P.spp.)
rebij rege,
*rabitchidraga

Artocarpus

incisus

*mai

mai, mai

mai

*mos, mosse,
mo-us, mohs

*me, ma

A. incisus var.

*me pa

mei pa

me pwo

meipa

A. incisus var.

*me si

met se

me si

Ficus sp.

*kawain

neen, nin

awan, auon,
aiiwdn, aoan,
au

shra, *konyah

*tebero, tepero

Pilea microphylla

*re

tabalok

Pipturus

argenteus

*oroma

arome (on
Nomwin I. ;

Hall Is.
called :
aroma)

alko

arame, *arme,

* armai

Ceodes

umbellifera

*mas

Mirabilis Jalapa

*pesikulok

*emenawa,

*emen aur,
*emmen aur,
*ulitch, ulij

Bryophyllum

pinnatum

*lamalam

Derris trifoliata

*kainipil

kan arai

unenipot, up

Vigna marina

*nimelitop

woniika

^margnejojo,
*margonejojo,
*margonejoyo,
markinichojo,
marlap, chojo

Acalypha grandis
var. genuina

*kurulong

manau

manou, monou

wut

Euphorbia Atoto

*pelepel

*builbuil,

* builibuilikar ,
*berrul,

*berol, peml,
*berau,

*perau,

*peiralo

Phyllanthus

Niruri

*limaimeir

limairpwong

negamaur,

nikammoiir,

amoesis

mar kaueue
(nameless on
most islands)

Allophylus

timorensis

*kitak

ngo

*kitak, *kutak,
*kudak,

*kutal

Flora of Pingelap— ST. JOHN

103

TABLE OF VERNACULAR NAMES ON PINGELAP AND ADJACENT ISLANDS ( Continued )

PLANT SPECIES

ISLANDS AND LIST OF VERNACULAR PLANT NAMES

Pingelap

Ponape

Truk

Kusaie

Marshall Is.

Triumfetta

procumbens

*konop

kiuin, kun,
liodot

*adat, *adat,
*atat, hatat,
*hadat

Sida fallax

*kao

sioi le

*kio, *keo,

*guio

Thespesia

populnea

*penne

pone, pona

pono, pond,
pona, polio,
bolle, okuran,
likokon

*banoh, pakeena

Calophyllum

Inophyllum

*sepang

isyo, luak, itsau

ijau, legitu,
regits, rekich,
rakich, rekit,
wangu, mosur

itu, eet, *etuh

luech, luej ,

*luet, *lues,
*luguez,

*luguetz

Carica Papaya

*kaineap

momiap

kippwau, kipau,
kipuau

*es

*keinapu,

*keinabu,

keinabbu,

kinapu,

kenabu,

keinapu,

*geinapu

Pemphis acidula

*ka-i-ni

ngi

engi

kasugel

*kungi, *gungi,
kinik

Sonneratia alba

*kosa

koto, kawtaw,
kwat

saras, salas,
sales, taras

folofol, flofol

eroeak

Barringtonia

asiatica

*wi

mi, ul

kun, gun, guon,
azan

kaenal, *pospus,
poasi-poasi

*wup, *wuep,
wop, *ob

Rhizophora

mucronata

*ak

ak, aak

ak, addo, chia,
tia

Terminalia

Catappa

*tepdp

tipap, thipdp,
thipwopu

as, asas

sarf

*kodel, kotal,
kutil

Terminalia

litoralis

*zmn

sin

un

kiking, *kugung,
kukung,

*kung,

*ekkung,

ekun

Jussiaea suffruti-
cosa var.
ligustrifolia

*kuri

teleurak,

deleurak

likeinenpul,

aunenipuin,

aunenipwin,

nigaulen

nen kut a kut

Plumeria

acutifolia

*po maria
(—Plumeria)

sour

*meria, *mei ria
(=Plumeria)

104

PACIFIC SCIENCE, Vol. II, April, 1948

TABLE OF VERNACULAR NAMES ON PINGELAP AND ADJACENT ISLANDS ( Continued )

PLANT SPECIES

ISLANDS AND LIST OF VERNACULAR PLANT NAMES

Pingelap

Ponape

Truk

Kusaie

Marshall Is.

Asclepias

curassavica

*kimeme

*kepok
(“kapok) ,

*ialu

(=yellow)

Messerschmidia

argentea

*sesen

titm

emoloset,

amaloset,

amoneset,

chen

sasran,

*shawshon

*krin, kirin,
*kurin, *girin,
kidrin

Clerodendrum

inerme

*ilau

Haw

ulo, apuech,
apuoch,
apwoch, etiu,
pucherik

*wuletch,

*wules, ulij

Premna

integrifolia

*sokuk

topuk, tupuk,
awr

lior,

nior, umukau,
umukaii

*fienket, fiankig

*kar, *gar

Pseuderanthe-
mum atropur-
pureum

*sarinairam

kaiwak

*jemla wulues

Guettarda

speciosa

*eles

ith

mosor

*wudilonaro,

*wutilumar,

*wutilomar,

*wudilomar,

♦wudinakatche,

wut

Ixora

carolinensis

*kalesu

katiu

atiu, achiu,
achen

galusa, kalce

*kajiru, *gajiru

Morinda

citrifolia

*obul

’umpul, woipel,
weipul, kirikei

nobur, nopur,
alin, arin, nen

*ee, e, hi

*nin, nen

Scaevola

frutescens

*ramek (on
Woleai Is.,
called :
ramakaa)

rarnuk, inuk,
enat

not, nat,
amaloset

>

kusros

*kunnat,

*kunnat,

*gunnat,

konnat,

*kennat,

ka-na-ta,

*kanun,
mar kinat

Wedelia biflora

*kisuwell

ngkau

atiwot, atuat,
atuot, eadiat

agaia, *ekeh

*morijetch,

♦marijetch,

*marjatch,

marajej,

morigides,

*markibuebue,

*margkiwewe,

*margiwewe,

markueue,

*buTlibuili-

kath

Flora of Pingelap— St. John

105

COMPARISON OF VERNACULAR PLANT NAMES

Analysis of these tabulations seems to reveal
the linguistic affinities of Pingelap.

Kusaie is the closest high island, being only
166 nautical miles to the southeast by east.
Actually, there is one other small island, Mokil,
situated 60 miles northwest by west, but nothing
is known of its flora, the people, or their lan¬
guage. Also, Ponape is situated 168 miles north¬
west by west from Pingelap, so it is not much
farther away than Kusaie.

Plants with Similar Names on Kusaie and
Pingelap

8 cultivated food crops
1 timber tree, probably cultivated
1 indigenous tree
1 cultivated ornamental shrub

1 indigenous

12 total (out of the 42 species in common)

Degree of Similarity: Four, or 33.3 per cent,
with identical names. These are all cultivated
food plants. Eight, or 66.6 per cent, with
altered names. All, or 100 per cent, are much
modified.

The Marshall Islands are remote, at least 243
nautical miles away, but are similar coral islands.
They include in their flora 52 of the species,
that is, all but five of the Pingelap plants.

Plants with Similar Names on the Marshall
Islands and Pingelap
5 cultivated food plants
3 cultivated ornamentals

2 perhaps cultivated, perhaps indigenous

3 indigenous

13 total (out of the 52 species in common)

Degree of Similarity: Four, or 30 per cent, with
identical names. These contain 1 cultivated
food plant, 1 cultivated ornamental, 1 sedge
cultivated or native, and 1 indigenous tree.
Nine, or 70 per cent, with altered names. Of
these, 3 (or 23 per cent) are much modified.

Truk, situated 600 miles to the west, is a
group with several volcanic islands in a lagoon
surrounded by an atoll ring with coral islets.

Plants with Similar Names on Truk and
Pingelap
11 cultivated food plants
1 cultivated ornamental
1 tree, medicinal, probably cultivated
1 shrub, cultivated ornamental or native
1 timber tree, probably cultivated
6 indigenous

1 shrub, native or cultivated, an economic
fiber plant

22 total (out of the 46 species in common)

Degree of Similarity: Four, or 18 per cent, with
identical names. Three of these are cultivated
food plants. Eighteen, or 82 per cent, with
altered names. Of these names, 4 (or 22 per
cent) are much modified.

The island of Ponape is fairly close, lying 168
miles to the northwest by west.

Plants with Similar Names on Ponape and
Pingelap

10 cultivated food plants

1 timber tree, probably cultivated
1 tree, fish poison, probably introduced
1 tree, food plant and ornamental, intro¬
duced

1 tree, food plant and medicinal, probably
introduced

11 indigenous

25 total (out of the 40 species in common)

Degree of Similarity: Six, or 24 per cent, with
identical names. These contain 4 cultivated
food plants, 1 fish poison, 1 indigenous tree.
Nineteen, or 76 per cent, with altered names.
Of these, 1 (or 5 per cent) is much modified.

Comparison of these data indicates that the
Pingelap plant names have most in common
with those of Ponape. There are 25 plants in
common with similar vernacular names; of
these, 6 are identical, and of the 19 modified
names, 18 are but slightly modified. There are
11 indigenous plants in the list, twice the num¬
ber for any other island or group. Even these
identities are small, as 34 of the Pingelap plants
have different names, yet the resemblances are
strongest between these two islands. It seems
evident that the vocabulary of Pingelap, though
quite distinct, has a small but definite similarity
to that of Ponape.

106

PACIFIC SCIENCE, Vol. II, April, 1948

Trukese is next in affinity, there being 22
plants with names in common. But, it is seen
that only 6 are certainly indigenous, that only 4
(or 18 per cent) have precisely identical names,
and that among the list of altered names about
22 per cent are much modified.

The Marshall Islands have little claim to a
close relationship, as the number of related
plant names is small, only 13. Of these, only 3
are indigenous plants, and 70 per cent of the
names are altered. Of these, 23 per cent are
much modified.

In Kusaie, of the plants in common, 33.3 per
cent have identical names, but the grand total
is only 12 species and varieties. Eight of the
names are much modified. The total of 12 spe¬
cies and varieties is so small that the high per¬
centage of identities, based on only three species,
is not significant.

All in all, comparisons of these Pingelap
plant name? indicate that the vocabulary of
Pingelap, at least in these names for common
objects, shows some affinities to that of Ponape.
With the other surrounding islands — Truk,
Kusaie, and the Marshalls — the words in com¬
mon are few and the affinities slight.

CATALOGUE OF THE FLORA OF PINGELAP

The specimens were all collected by H. St.
John on December 27, 1945. The vernacular
Pingelap name is given in quotation marks,
followed by the author’s collection number for
the species. The names of indigenous species are
printed in bold roman type, while those of ad-
ventives and cultivated plants are in bold italics.

POLYPODIACEAE

1. Asplenium nidus L.

"Seilik,” 21,477. Occasional, epiphyte on
moist tree trunks.

2. Nephrolepis biserrata (Sw.) Schott
"Pues,” 21,484. On ground or tree trunks,
in moist forest.

3. Polypodium Phymatodes L.

"Kitiu,” 21,479. Common on ground or
trees.

4. Vittaria elongata Sw.

'Tit,” 21,466. On mossy bases of coconut
trunks, in moist woods. A common fern of
the tropical Pacific, occurring from Africa,
India, Burma, Malaya, and on the high
islands from Sumatra and the Philippines
through Malaysia and Australia, and Poly¬
nesia to the Marquesas. Apparently this col¬
lection is the first to be reported for the
species on an atoll or a low coral island. No
such record was known to Wagner (1945:
74-76), though on page 76 he reported it
on Guam, with five other fern species on a
coconut trunk "in a shady bushy location on
the wooded side of a limestone hill.”

PANDANACEAE

5. Pandanus sp.

"Kipai.” One kind was collected, but unfor¬
tunately it was lost on the airplane trip back
to Honolulu. Pandanus trees of good size
were common both on the beaches and in
the interior. They are important to the in¬
habitants, furnishing edible fruit, timber
from the trunks, and thatch from the leaves.
My native informant, Soas, could not re¬
member all, but furnished the names of the
following species or kinds:

(1) "Asibuirek”

(2) "Nanagaisal”

(3) "Sonumei”

(4) "Nanagasak”

( 5 ) "Aisesewil”

(6) "Maukosokosok”

(7) "Muisamuis”

(8) "Esies”

(9) "Suioibueibuei”

(10) "Arawa-an,” or "Arawan”

(11) "Muisigel”

(12) "Meikilikil”

(13) "Luaramuk”

(14) "Tobodin”

Flora of Pingelap — St. John

107

No. (1) resembles the varietal names
"Ajbirik” of Ailuk Atoll, "Ajibuiruk” of
Majuro Atoll, and "Agiwirok” of Jaluit
Atoll in the Marshall Islands. No. (10) re¬
sembles "Eruan” of Ailuk Atoll, Utirik Atoll,
Me jit Island, Majuro Atoll, Namu Atoll,
Jaluit Atoll, and Likiep Atoll, and "Erwan”
of Wotje Atoll, and Jaluit Atoll. No. (13)
resembles "Loarmai” of Ebon Atoll. No.
(14) resembles "Tibitin” of Majuro Atoll,
and "Tabatin” or "Tabawdin” of Ebon Atoll.

HYDROCHARITACEAE

6. Thalassia Hemprichii (Ehrenb.) Aschers.
"Walat,” 21,458- Small plants in sand, sub¬
merged in shallow sea water off outer beach.
The young plants were sterile, but by exami¬
nation of the foliar anatomy, identification
was reasonably certain.

GRAMINEAE

7. Eragrostis amabilis (L.) Wight & Arn.
"Rosakai.” Not collected, but observed.

8. Lepturus repens R. Br.

"Rosakai” (the name of any grass or grass¬
like plant). Observed, but not collected.

9. Saccharum officinarum L. — Cultivated.
"Seu.” Observed, but not collected. Small
clumps were grown by the huts or houses in
the village, and occasional plants were seen
in the extensive wet field or "lepuel” for
Cyrtosperma culture. The vernacular names
of the five cultivated varieties were:

(1) "Kala”

(2) "Sowesasa”

(3) "Palau”

(4) "Teimos”

(5) "Ieseng”

lO.Thuarea involuta (Forst. f.) R. & S.
"Mokarak,” 21,465. Repent on sands near
beach.

CYPERACEAE

11. Cyperus javanicus Houtt.

"Sapasap,” 21,471. Edge of fresh swamp.
Used to perfume coconut oil.

12. Fimbristylis cymosa R. Br.

"Rosakai,” 21,491. In woods by lagoon
beach.

PALMAE

13. Cocos nucifera L. — Cultivated and spon¬

taneous.

"Ni.” Observed, but not collected, abundant.
Various growth stages of the nut are named,
as:

"Pen,” the drinking nut, or two-thirds
grown

"Aring,” the ripe nut
"Par,” the sprouted nut

ARACEAE

14. Colo cast a esculenta (L.) Schott var. anti¬

quorum (Schott) Hubb. & Rehd. — Cul¬
tivated.

"Sawa.” Observed, but not collected. A few
plants were seen, but they were grown in
the Cyrtosperma "lepuel” almost as speci¬
mens. It is a crop of very minor importance.
The seven following varieties were distin¬
guished by name:

(1) "Bokor”

(2) "Tawang” [PTaiwan]

(3) "Mesawsol”

(4) "Koso”

(5) "Sawa Pingelap”

(6) "Sauk”

(7) "Pemeru”

15. Cyrtosperma Chamissonis (Schott) Merr.

— Cultivated.

"Muiang.” Observed, but not collected. The
most important food crop. Near the center
of the island was a large swamp that possi¬
bly had been enlarged by digging. It seemed
to be 300 feet wide and more than 600 feet

108

PACIFIC SCIENCE, Vol. II, April, 1948

long. Within this low, wet garden, or
"lepuel,” each family controlled a plot and
each year fertilized it with leaves and trash.
The "lepuel” was thickly planted to "Mui-
ang.” The natives reported that the crop
matured in 3 years, but could be harvested
any time after 1 year. If allowed to grow to
maturity, the corm attained large size, as
much as 2 m. in length and 6 dm. in diam¬
eter and a weight so heavy that two men
were needed to carry it. Five varieties were
cultivated, and were known by the follow¬
ing vernacular names:

( 1 ) "Muiang An Ngatik”

(2) "Muian Sdontol”

( 3 ) "Nane Pakeleman”

(4) "Simidin”

(5) "Seriseng”

The name of No. (1) refers to Ngatik
Island, a coral island 200 miles westward of
Pingelap.

AMARYLLIDACEAE

1 6. Crinum asiaticum L. — Cultivated.

"Kiep.” Observed, not collected. Seen only
as a cultivated plant in the village.

Yl.Zephyranthes rosea (Spreng.) Lindl. —
Cultivated.

"Kiep.” An ornamental, by the houses; ob¬
served, not collected.

TACCACEAE

18. Tacca Leontopetaloides (L.) Ktze. — Rev.
Gen. PI. 704, 1891.

Leontice Leontopetaloides L., Sp. PL 313,
1753.

T. pinnatijida Forst., Char. Gen. 70, t. 35,
1776.

"Mugamuk,” 21,480. Cultivated and spon¬
taneous. Commonly planted, also persisting
and spreading in the woods, abundant. Stem
fibers used in plaiting hats. The tubers are
an important source of starchy food. They
are grated, the pulp washed in three changes

of sea water, washed in fresh water, then
discarded. The starch which accumulates as
a sediment is dried and preserved for use as
food.

DIOSCOREACEAE

19. Dios corea 1 korrorensis R. Knuth
"Kep,” 21,481. Cultivated in village, the
stems climbing on a tree. Tuber edible, said
to attain a maximum size of 1 m. in length,
and 3 dm. in diameter. The plant was ster¬
ile, but the vegetative parts match well those
of D. korrorensis.

MUSACEAE

20. Musa paradisiac a L- — Cultivated in village

and in the wet "lepuel.”

"Wis.” Numerous plants were seen and they
were vigorous and productive. They repre¬
sented both the subsp. normalis Ktze. and
the subsp. sapientum (L.) Ktze., and in¬
cluded the following named varieties:

(1) "Latin”

(2) "Amerika”

(3) "Usigaras”

(4) "Iyeman”

(5) "Lakatan”

(6) "Taiwan”

(7) "Kutkut”

(8) "Panilo”

(9) "Manila”

(10) "Wuseak”

Obviously Nos. (2), (3), and (9) bear
names indicating foreign origin, and per¬
haps No. ( 1 ) does also. No. (5) is the ver¬
nacular name on Jaluit Atoll for Pandanus
Lakatwa Kanehira. No. (2) is reported by
Lt. Comdr. S. H. Elbert in his Trukese dic¬
tionary to be the "Lady Finger” variety.

PIPERACEAE

21. Peperomia ponapensis C. DC.

"Warm,” 21,475. On coral stone wall in
moist forest, near lagoon beach. Medicinal,

Flora of Pingelap— St. John

109

the pounded, fleshy leaves being used as a
poultice for boils. Few of the Pacific atolls
support Peperomia, so this locality record is
noteworthy.

The specimen was submitted to and iden¬
tified by Dr. T. G. Yuncker as P. ponapensis.
The collector had also studied this species,
deciding that it was the most similar one,
but that the plants from Pingelap differed in
having the leaves averaging smaller, mostly
not 3-5 cm. long; and the fruit smaller,
0. 5-0.6 mm. in diameter. These differences
are still evident, but in a genus with so many
microspecies, he has no desire to add an¬
other, so accepts the determination as P.
ponapensis. Ponape is an adjacent high
island, lying 168 miles west by north.

MORACEAE

22. Artocarpus incisus (Thunb.) L. f., Suppl.
411, 1781. Cultivated.

Rademachia incisa Thunb., Vet. Akad. Stock¬
holm, Handl. 37: 254, 1776.

A. communis Forst., Char. Gen. 101, 1776.
Sitodium-altile Parkinson, Jour. Voy. En¬
deavour 45, 1773.

A. altilis (Parkinson) Fosberg, Wash. Acad.

Sci., Jour. 31: 95, 1941.

"Mai.” Observed, but not collected. Abun¬
dant and vigorous, the trees attaining a
height of 20 m. or more.

Variety with Seed-bearing Fruit:

(1) "Mei sabarek”

Varieties with Seedless Fruit:

( 1 ) "Meipa”

(2) "Mei si”

There is confusion concerning the bino¬
mial for the breadfruit, and it is not asserted
that these words will settle it, but they are
given in justification of the name adopted.
Rademachia incisa Thunb., published in
1776, was little used, while Artocarpus com¬
munis Forst. of 1776 provided the accepted
generic name and the combination Artocar¬
pus incisus (Thunb.) L. f., made in 1781,

furnished the binomial that won almost uni¬
versal acceptance (as A. incisa') for more
than a century. Then Merrill (1906: 43)
readopted A. communis Forst., listing A.
incisa (Thunb.) L. f. as a synonym. In sev¬
eral later publications he continued this
usage, without giving an interpretation, but
he apparently preferred communis because
it was published in the genus Artocarpus.
The International Rules of Botanical No¬
menclature of that date or of this current
date do not include a rule validating this
choice, while Art. 4 and 5 apply, as in cases
of doubt, authorizing the following of es¬
tablished custom, as in this case, the choice
of A. incisus (or incisa). A final basis
would be that of strict priority, but no one
has yet been able to establish the exact dates
within the year 1776 for the two publica¬
tions by Thunberg and by Forster. If biblio¬
graphic research can establish the exact
dates, this matter will be finally settled.

Another detail in question is that of the
gender of the generic name Artocarpus. The
name was published by Forster (1776: 101-
102, t. 51, 51a) and the two Greek roots
were given — artos, bread, and karpos, fruit
— from which the name was derived. The
single species A. communis was listed, but
with no clear indication of the gender. The
termination us would ordinarily be mascu¬
line, but the practice of making genera fem¬
inine was so general, especially with trees,
that one cannot now be certain that the
Forsters decided to make the genus mascu¬
line. The specific name they used, commu¬
nis , is either masculine or feminine. There
is no other evidence in the Forster book, in
the index or text, that gives any indication.
Corner states (1939: 282) that Forster sub¬
sequently used A. incisus, but the writer
notes that G. Forster changed to A. incisa F.
( = Forst. f.) (1786: 23), though he should
have credited the combination to L. f. or at
least to L. f. emend. Forst. f., as based on a
change of gender.

110

PACIFIC SCIENCE, Vol. II, April, 1948

The two simultaneously published genera
and binomials, Rademachia incisa Thunb.
and Artocarpus communis Forst., were first
united in 1781 by the younger Linnaeus
(Linne, 1781: 411-412). His choice, which
we must accept as final (Int. Rules, Art. 56),
was the genus Artocarpus . Though this gen¬
eric name was adopted from the Forsters, he
chose the specific name from Thunberg and
the present International Rules validate such
a choice. Since he was the next author to
publish a specific name in the genus Arto¬
carpus, he also could determine the gender
of the name. He used the form Artocarpus
incisus (Thunb.) L. f., which would have
been decisive, establishing the generic name
as masculine, had he not for the second
species used the feminine one A. integrifolia
L. f., which he coined anew for Rademachia
integra Thunb., a substitution now illegal,
but which indicated his use of a conflicting
gender. Thus, the younger Linnaeus only
added new confusion to the problem of the
gender of the name.

The next author to publish on Artocarpus
appears to have been Lamarck (1789:207-
210), who treats the genus in detail, includ¬
ing five species, three of them new. Of the
five names, A. Philippensis is either mascu¬
line or feminine; A. jaca (=A. Jaca) is a
name based both on the generic name Jaca
and the vernacular name "Jak,” so it does
not truly indicate the gender, but the three
other specific names, incisa, heterophylla,
and hirsuta, are all feminine, so this choice
by Lamarck, perhaps following G. Forsters
second usage, may be accepted as determin¬
ing the gender of the generic name, and
nearly all subsequent botanists did so accept
it. It so stood until 1935 when the third
International Rules of Botanical Nomencla¬
ture included a new mandatory rule, 72(2),
applying to the gender of generic names:
". . . all other modern compounds ending in
the Greek masculine carp os (or carpus) are

masculine.” This rule changes the gender of
Artocarpus to masculine. It is so used by
Corner (1939:280) but he does not dis¬
cuss the gender and he also uses the femi¬
nine name A. integrifolia L. f., as he did in
his earlier paper in the same journal (1939:
71, 80).

It does not seem to be generally realized
that Artocarpus is now established as a mas¬
culine genus, and the name for breadfruit
is A. incisus (Thunb.) L. f. The writer is
fully aware of the name A. altilis (Parkin¬
son) Fosberg, based on the earliest name for
the breadfruit, Sitodium-altile Parkinson, but
the validity of this name is still in doubt,
and even Fosberg himself has proposed
(1939:230-231) that Sitodium be made a
rejected name and Artocarpus a conserved
name. The issue is much involved, so for
the time being, the long-established name
Artocarpus incisus (Thunb.) L. f. will be
retained for the breadfruit tree.

23. Ficus sp.

"Kawain,” 21,464 . Young tree, 3 m. talL
Fruit cooked and eaten; bark fiber used for
fish line. Probably indigenous.

URTICACEAE

24. Pilea microphylla (L.) Liebm. — Adven-

tive weed.

"Re.” Observed, but not collected. Common
in village on stone walls and foundations.

25. Pipturus argenteus (Forst. f.) Wedd.
"Oroma,” 21,486. In woods. Tree 8 m. tall,
by 2 dm. in diameter. The bast fiber is used
for making fish nets.

NYCTAGINACEAE

26. Ceodes umbellifera J. R. & G. Forst.
"Mas,” 21,487 . Tree 10 m. tall, by 3 dm. in
diameter, in the village. There is an islet in
the Ulithi Atoll named Mas Island.

27. Mirabilis Jalap a L. — Cultivated.
"Pesikulok.” Observed, but not collected. An
ornamental, commonly cultivated in the vil¬
lage!

Ill

Flora of Pingelap — St. John

CRASSULACEAE

28. Bryophyllum pinnatum (Lam.) Kurz —

Cultivated.

"Lamalam,” 21,482. Ornamental, introduced
by the Germans. Found by village street.
The vernacular name may be geographic,
alluding to the Lamaram Islands, lying 120
miles west by north.

LEGUMINOSAE

29. Derris trifoliata Lour.

"Kainipil,” 21,470. Vine, climbing on trees
near beach.

30. Vigna marina (Burm.) Merr.

"Nimelitop,” 21,460. Vine, trailing or climb¬
ing, top of beach.

EUPHORBI ACE AE

31 .Acalypha grandis Benth. var. genuina
Muell. Arg. — Cultivated.

"Kurulong.” Observed, not collected. An
ornamental, grown in the village beside the
houses or as a hedge plant, introduced dur¬
ing the German rule.

32. Euphorbia Atoto Forst.

"Pelepel,” 21,467. Tufted, erect; leaves glau¬
cous beneath. In grassy thicket.

33. Phyllanthus Niruri L.

"Limaimeir,” 21,485. Common in open
places or in forest. Medicinal, used for treat¬
ing dysentery.

SAPINDACEAE

34. Allophylus timorensis (DC.) BL
"Kitak,” 21,478. Young tree, 7 m. tall, the
flowers white.

TILIACE AE

35. Triumfetta procumbens Forst. f.

"Konop,” 21,476. Trailing on sand in open
woods. The flexible stems provide a firm,
shiny fiber much used, when dyed, in plait¬
ing belts, mats, etc.

MALVACEAE

36. Sid a fallax Walp. — Cultivated in village.
"Kao,” 21,456. Shrub 2 m. tall.

37. Thespesia populnea (L.) Soland.

"Penne,” 21,473. Tree 8 m. tall, by 3 dm. in
diameter; flowers fading red. By lagoon
beach. Wood of good quality, used for ax
handles, etc.; bark fiber used for making fish
nets.

GUTTIFERAE

38. Calophyllum Inophyllum L.

"Sepang,” 21,461. Tree 15 m. tall, by 1 dm.
in diameter. Top of beach, only a few trees
seen, apparently introduced by the natives.
Wood used for canoe hulls, etc. Fruit medi¬
cinal and a source of oil. The vernacular
name may be geographic, referring to Sai¬
pan Island.

CARICACEAE

39. Carica Papaya L. — Cultivated.

"Kaineap.” Observed, not collected, com¬
mon. The vernacular name resembles "Kei-
napu” of Namu, Likiep, Ailuk, Utirik, Me jit,
Majuro, and Ebon in the Marshalls; and
"Keinabu” in Aur Atoll.

LYTHRACEAE

40. Pemphis acidula Forst.

"Ka-i-ni,” 21,474 . Tree 7 m. tall, by 2 dm.
in diameter. On lagoon beach. Wood used
for handles, pipes, etc.

SONNERATIACEAE

41. Sonneratia alba Sm.

"Kosa,” 21,468. Tree 8 m. tall, by 2 dm. in
diameter, the roots with knees. A mangrove,
growing in the shallow salt water of the
lagoon. Wood good, used for tool handles.

LECYTHIDACEAE

42 . Barringtonia asiatica (L.) Kurz

"Wi.” Observed, but not collected. Prob¬
ably introduced by the natives because of its
value as a fish poison.

112

PACIFIC SCIENCE, Vol. II, April, 1948

RHIZOPHORACEAE

43. Rhizophora mucronata Lam.

"Ak,” 21,469 . Tree 8 m. tall, by 3 dm. in
diameter, with prop roots. A mangrove tree,
growing in shallow salt water of lagoon.
Good timber.

COMBRETACEAE

44. Terminalia Catappa L. — Cultivated.
"Tepop.” Observed, but not collected.

45. Terminalia litoralis Seem.

"Win,” 21,459 . Tree 9 m. tall, by 7 dm. in
diameter; flowers greenish; fruit 15-18 mm.
long, ellipsoid, crimson, edible. Wood used
for tool handles.

ONAGRACEAE

46. Jussiaea suffruticosa L. var. ligustrifolia

(HBK.) Griseb. — Introduced weed.
"Kuri.” Observed, but not collected. Seen
growing in the low, wet Cyrtosperma patch.

APOCYNACEAE

47. Plumeria acutifolia Poir. — Cultivated.

"Po maria.” This vernacular name for the
introduced ornamental tree is clearly only
the natives’ method of pronouncing Plume¬
ria.

ASCLEPIADACEAE

48. Asclepias curassavica L. — Introduced weed.
"Kimeme,” 21,490. In the village.

BORAGINACEAE

49. Messerschmidia argentea (L. f.) I. M.

Johnston

"Sesen,” 21,457. Tree 8 m. tall, by 3 dm. in
diameter, common.

VERBENACEAE

50. Clerodendrum inerme (L.) Gaertn.
"Ilau,” 21,472. Shrub, the arching branches
2-4 m. long. In forest near lagoon beach.

51. Premna integrifolia L.

"Sokuk,” 21,488. Tree 8 m. tall, by 2 dm.
in diameter; flowers white; fruit black. In
moist woods.

ACANTHACEAE

52. P sender anthemum alropurpureum (Bull)

Radik. — Cultivated.

"Sarinairam,” 21,489. Cultivated in village;
introduced by the Germans. Shrubs 1-4 m.
tall; flowers white, with rose-magenta spots
in the throat.

RUBIACEAE

53. Guettarda speciosa L.

"Eles,” 21,492. Tree 7 m. tall, by 2 dm. in
diameter. The logs used for canoe hulls.
The white, fragrant flowers used as orna¬
ments in the hair or used to perfume coco¬
nut oil.

54. Ixora carolinensis (Val.) Hosokawa, aff.

var. typica Fosb. — Cultivated.

"Kalesu,” 21,463. Cultivated in the village
as an ornamental. Shrub 5 m. tall. This does
not match any of the several varieties de¬
scribed by Fosberg, but comes closest to the
var. typica.

55. Morinda ci trifolia L. — Growing away from

the village, but apparently not native.
"Obul.” Observed, but not collected. Though
the fruit is bitter, slimy, and nauseating, the
natives use it as an edible fruit and as a
medicine.

GOODENIACEAE

56. Scaevola frutescens (Mill.) Krause
"Ramek,” 21,462. Shrub 8 m. tall, by 2 dm.
in diameter; flowers white; fruit white.

COMPOSITAE

57. Wedelia biflora (L.) DC.

"Kisuwell,” 21,483. Half scandent shrub. In
moist woods, common.

Flora of Pingelap — ST. JOHN

113

REFERENCES

Bryan, Edwin H., Jr. Revegetation of certain
Marshall Islands. In the Bishop Museum
Library. [Manuscript.]

Corner, E. J. H. 1938-39. Notes on the sys-
tematy and distribution of Malayan Phanero¬
gams II and III. Straits Settlements Gard. Bui.
10(1): 56-81, 2 pL, 5 fig., 1938; 10(2):
239-329, 1939.

Elbert, S. H. 1947. Trukese-English and Eng-
lish-Trukese dictionary, vi+337. U. S. Naval
Military Government, 14th Naval District.
[Plant names, pp. 297-302.}

Forster, Johann Georg Adam. 1786. De
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commentatio botanica. 80 p. Havde & Spener,
Berolini.

Forster, Johann Reinhold, and Johann
Georg Adam Forster. 1776. Characters
generum plantarum, quas in itinere ad insulas
Maris Australis collegerunt, descripserunt, de-
linearunt, annis MDCCLXX1I-MDCCLXXV .
viii+150, 75 pi. B. White, T. Cadell, & P.
Elmsly, Londini.

Fosberg, F. Raymond. Micronesian plant
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[Manuscript.]

— - — 1939. Nomenclature proposals for

the 1940 Botanical Congress. Amer. Jour.
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Gulick, Luther H. 1880. A vocabulary of
the Ponape dialect, Ponape-English and Lng-
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Hambruch, PAUL. 1932. Ergebnisse der Siid-
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von Dr. G. Thilenius, II Ethnographie, B:
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Karte. Friederichsen, de Gruyter & Co., Ham¬
burg. [Die Pflanzenwelt, pp. 349-356.}

Kanehira, Ryozo. 1933. Flora Micronesica.
vi+468+37, 21 pi., 211 fig. Japanese South
Seas Bureau. [In Japanese.}

- 1935. An enumeration of Microne¬
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Koidzumi, Gen-ichi. 1915. The vegetation
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257.

Kraemer, Augustin, and Hans Nevermann.
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Gruyter & Co., Hamburg.

1932. Bd. 5, Inseln um Truk, xxvi+452, 229
Abbild., 30 Taf., 10 Karten. [Flora, pp.
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1935. Bd. 6(1), Inseln um Truk, vi+291,
118 Abbild., 24 Taf., 3 Farbtaf.

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Lamarck, Jean Baptiste Pierre Antoine
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Lesson, P. 1839. Voyage autour du monde en-
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Linne, Carl von (the son). 1781. Supple-
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Merrill, Elmer D. 1906. The flora of the
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Jour. 35(3): 74-7 6.

Morphometric Characteristics and Relative Growth of Yellowfin Tunas
(Neothunnus macropterus) from Central America

Milner B.

The problem of whether the yellowfin tuna
stocks inhabiting different parts of the Pacific
Ocean are genetically independent is of inter¬
est both from taxonomic and economic view¬
points. Several species of yellowfin tuna have
been described from the Pacific Ocean, sup¬
posedly differentiated from each other by the
relative lengths of the fins. The determination
of the validity of these species requires exam¬
ination of series of fish of all sizes, and of both
sexes, to determine whether some of the descrip¬
tions may not be based on sex-connected or size-
connected variations of a single species. Since
the yellowfin tuna is one of the most valuable
commercial varieties, it is of considerable im¬
portance to determine whether the groups en¬
countered in different parts of the Pacific are
all members of one large stock which is, there¬
fore, entirely open to exploitation at any point
in its range, or whether there are a number of
separate stocks, in which case the exploitation of
one would have no effect on the exploitation of
the others.

Approach to the problem by methods of mor¬
phometric analysis requires the examination of
series from each of a number of different locali¬
ties. Since these fish are of large size, the few
specimens in the various museums are insuffi¬
cient for the purpose. The only practical pro¬
cedure is to make the counts and measurements
in the field. The area to be covered is so very
large that it is impractical, at present at least,
for one person to visit all the various localities.
Therefore, it seems desirable that the data for

1 South Pacific Investigations, U. S. Fish and Wild¬
life Service. Published by permission of the Director,
U. S. Fish and Wildlife Service. Manuscript received
October 6, 1947.

Schaefer1

the fish from a given locality be put on record
as soon as obtained, for subsequent comparison
with those of other places as they become
available.

MORPHOMETRIC DATA

From late January until late June, 1947, the
factory ship "Pacific Explorer” was anchored in
the Gulf of Nicoya, on the Pacific coast of Costa
Rica, for the purpose of freezing a cargo of
tunas from the adjacent oceanic fishing grounds.
The author was aboard this vessel and the fishing
boats supplying the mother ship as an observer
for the Fish and Wildlife Service until March
7, when he was relieved by J. C. Marr. Between
January 22 and April 15 morphometric measure¬
ments and counts were made by the author and
Mr. Marr on a series of 46 yellowfin tuna from
the waters off Costa Rica. These were made on
recently caught, unfrozen fish, either aboard the
fishing vessels or aboard the mother ship. Data
for each fish are tabulated in Table 1. All
measurements are in millimeters. Specimens
were selected according to size, so as to give a
fairly even representation throughout the range
of sizes encountered in this fishery. Our speci¬
mens ranged in length from 542 to 1,571 mm.
Since we are interested in determining the mor¬
phometric characteristics of fish of different
sizes, the arbitrary selection of fish by sizes is
justified because "the regression function does
not depend on the frequency distribution of the
independent variate” (Fisher, 1934:127).

Sex was determined for most of the fish
measured. For fish over about 650 mm. this
was quite easy, because these larger fish were
undergoing development of the gonads during

C114]

Morphometry of Tuna — Schaefer

115

TABLE 1. MORPHOMETRIC DATA FOR COSTA RICAN YELLOWFIN TUNA
(Specimens above horizontal line measured by author, those below by J. C. Marr.)

m

WEIGHT J

TOTAL LENGTH

HEAD LENGTH

SNOUT TO INSERTION

FIRST DORSAL

SNOUT TO INSERTION

SECOND DORSAL

SNOUT TO INSERTION

ANAL

GREATEST DEPTH

LENGTH PECTORAL FIN

PECTORAL INSERTION TO

INSERTION FIRST DORSAL

LENGTH OF BASE OF

FIRST DORSAL

LENGTH OF LONGEST

(FIRST) DORSAL RAY

LENGTH OF SECOND

DORSAL FIN

LENGTH OF ANAL FIN

LENGTH OF LONGEST

DORSAL FINLET

DIAMETER OF IRIS

MAXILLARY LENGTH

FIRST DORSAL FIN RAYS

DORSAL FINLETS

ANAL FINLETS

GILL RAKERS

kilos

mm.

mm.

mm.

mm.

mm.

mm.

mm.

mm.\mm.\ mm.

mm.

mm.

mm.

mm.

mm.

no.

no.

no.

no.

F

10.8

835

228

253

456

505

215

219

121

194

99

101

99

27

31

89

14

9+1

9+1

F

19.1

1,041

275

297

524

605

256

146

246

118

34

32

107

14

9+1

9+1

11+21

F

21.3

1,080

284

313

564

629

270

283

154

260

130

188

204

34

34

109

14

9+1

9+1

8+25

18.1

1,030

282

300

530

612

258

270

242

116

169

189

31

35

106

14

9+1

9

9+20

8.3

719

203

223

334

441

188

195

168

89

83

88

23

29

81

13

9+1

9+1

9+22

....

10.8

854

235

249

447

513

207

248

131

205

98

132

141

28

30

92

14

9+1

10

9+21

M

70.3

1,571

405

440

805

917

410

346

232

389

189

305

309

53

44

153

13

10

10

8+19

....

31.3

1,173

306

340

607

686

321

300

185

281

144

242

203

48

34

119

14

9+1

9+1

9+22

6.4

703

209

207

393

434

170

201

103

169

97

85

95

22

30

80

14

9+1

9+1

10+21

F

1,475

389

425

746

845

372

345

217

337

166

255

313

44

40

147

13

10

9+1

10+21

F

22.2

1,084

293

314

566

632

271

280

151

261

170

216

33

33

114

13

10

9+1

10+21

F

30.8

1,166

312

347

611

673

305

297

185

275

142

189

232

41

38

122

13

9+1

9

10+20

M

1,411

372

405

705

782

357

335

214

318

158

308

397

46

41

143

13

10

9+1

10+21

M

1,242

329

352

641

702

321

291

190

302

147

265

284

41

37

125

14

9+1

9+1

9+20

M

1,189

310

344

614

693

306

304

177

288

146

211

245

39

36

119

14

9+1

9

9+21

F

1,057

290

319

557

625

268

275

160

251

126

172

192

33

36

111

14

9+1

9

9+20

M

823

229

251

442

492

221

237

125

206

100

115

116

29

30

89

14

9+1

9

10+21

M

9.1

770

226

250

429

466

209

217

127

191

85

101

103

26

31

86

14

8+1

8+1

9+22

M

26.3

1,144

311

336

602

669

293

301

174

275

132

230

243

38

35

120

14

9+1

8+1

11 + 22

F

22.7

1,090

295

323

570

635

272

294

163

261

140

209

216

37

35

114

13

9+1

9+1

10+20

M

3.2

542

165

177

304

339

147

152

85

132

61

55

59

17

27

63

14

9+1

8+1

11+21

M

12.2

887

247

265

480

529

225

259

125

206

108*

130

128

27

96

13

9+1

9+1$

9+21

M

29.9

1,132

312

338

601

664

299

313

181

256

147

221

231

37

119

13

9+1

9+1

10+21

F

6.4

698

205

213

381

423

181

196

108

164

83

88

92

23

77

14

8+2

7+1

10+20

M

13.1

897

251

264

484

530

223

245

133

214

104

131

137

27

94

14

9+1

8+1

9+20

F

4.5

637

185

201

348

393

161

184

98

142

72

77

77

20

73

13

9+1

9+1$

10+21

M

13.1

883

247

263

475

522

225

243

135

205

98

127

130

29

95

14

9+1

8+1

11+20

M

5.4

662

190

204

366

409

170

191

99

74

75

80

22

73

13

9+1

8+1$

9+20

M

15.9

973

266

279

515

575

230

268

143

223

115

155

167

31

102

14

9+1

8+1$

8+20

F

15.4

922

264

275

499

556

238

262

145

201

111

132

160

30

101

13

8+2

8+1$

11 + 22

F

15.0

931

257

270

487

553

234

245

128

198

108

119

132

29

96

13

8+2

8+1$

10+21

M

15.9

918

261

284

494

558

240

257

145

206

111

143

166

30

94

14

9+1

9+1$

9+21

M

15.9

963

270

288

516

579

241

268

149

210

112

131

137

29

34

102

13

9 ( + 1 ?)

8+1

9+21

M

15.4

957

262

279

510

567

245

262

146

216

112

139

164

28

32

102

14

8+2

8+1

11 + 21

F

154

943

257

276

498

552

236

264

141

199

108

143

146

30

34

98

13

8+2

8+1

10+21

M

16.8

976

265

287

510

572

247

274

147

202

115

153

176

32

35

105

13

8+1

9( + l?)

11 + 21

M

29-0

1,233

323

358

644

706

329

313

194

271

152*

231

274

40

38

120

14

9+1

8+1

9+21

M

3.6

574

169

182

325

352

152

159

93

136

65

63

62

18

30

62

14

8+2

8+1

10+21

M

19.1

996

267

296

524

583

257

268

157

224

122

157

173

32

34

110

14

8+2

9+1

11 + 21

M

25.9

1,114

296

322

575

642

281

285

169

246

130

166

173

37

38

116

14

9+1

8+2

9+21

M

15.4

931

257

274

496

561

242

260

141

219

108

133

137

30

33

97

14

8+2

8+2

9+22

F

16.8

97 6

270

293

522

584

235

274

141

224

119

139

155

32

35

102

14

8+2

8+2

10+22

F

6.4

693

204

220

391

428

178

198

109

161

82

94

22

32

80

14

7+2f

8+1

11+21

M

3.6

618

180

193

343

381

155

173

92

136

72

67

70

20

29

68

13

8+2

8+2

10+20

M

5.0

638

186

199

360

399

167

188

102

153

83

77

75

21

29

73

14

8+2

8+1

9+20

F

4.5

624

184

195

349

387

164

168

97

143

81

71

68

20

29

71

14

8+2

8+2

11+21

* Second dorsal ray is the longest. f Sixth finlet missing. $ Doubtful whether +1 or +2.

116

PACIFIC SCIENCE, Vol. II, April, 1948

this season, and many of them were in ad¬
vanced stages of maturing of the sexual products.
The smallest fish, under about 650 mm., which
are believed to be fish only a year old, had un¬
developed gonads and the determination of sex
was difficult.

Measurements were made in millimeters with
the same slide-calipers used by Godsil and
Byers (1944), or with dividers for short dis¬
tances such as diameter of iris or length of
maxillary. The various measurements and
counts were made in the exact manner described
by these authors in their Appendix II except as
noted below.

Pectoral insertion to insertion first dorsal was
measured with the tip of the fixed arm of the
caliper at the insertion of the first dorsal fin, the
sliding arm being brought to the anterior ter¬
mination of the dorsal margin of the pectoral
fin.

Length of longest (first) dorsal ray was
measured with dividers from the juncture of the
ray (with the fin extended) and the contour of
the body to the tip of the ray. The longest ray
was the first ray in all cases except two, which
are indicated in the table.

I refer to the ’'length” of the second dorsal
and anal where Godsil and Byers use the term
"height.”

Length of longest dorsal finlet (the 5th or
6th) was measured with dividers from the an¬
terior margin of the finlet to the tip of the pos¬
terior filament. The number of dorsal finlets
and number of anal finlets were counted from
posteriorly forward and if the last, most an¬
terior, finlet (the last two in some cases) was
attached to the second dorsal (or anal fin) by
a membrane, it is recorded separately from the
count of the free finlets. Thus if there were 9
finlets, all free, the record is 9; if there were 9
free finlets and one attached by a membrane to
the fin, the record is 9+1. This seems not to
be a very good character because of the diffi¬
culty of distinguishing between attached finlets
and the posterior end of the fin itself. It may be
noted that our counts tend to average higher

than those of Godsil and Byers, even when the
difference in method of counting is taken into
account. There also seems to be some difference
between the author and Mr. Marr as to when a
finlet is considered attached.

The character "length of base of first dorsal”
would be more accurately called "first dorsal in¬
sertion to second dorsal insertion” since, follow¬
ing Godsil and Byers, that is the measurement
taken for this character.

Weights were usually taken with a spring
balance reading in pounds. Fish over 100
pounds were weighed in pounds on a platform
scale. Some small fish were weighed on a small
spring balance, read to 0.1 kilo. Because the
readings recorded in pounds to the nearest
pound were later translated to kilos, the ac¬
curacy is slightly less than indicated; individ¬
ual weights may be in error by as much as 0.25
kilo.

RELATIVE GROWTH

The measurement data in Godsil and Byers’
paper (1944) are recorded in terms of body
proportions, that is, in terms of the times a
given measurement is contained in the body
length, or in the head length, depending on
the character. Body proportions have also been
used to characterize supposed species, for ex¬
ample, by Jordan and Evermann ( 1926) and by
Nichols and La Monte (1941). Where data
from fish of different sizes are compared, this
is unsatisfactory unless the ratio of the dimen¬
sion under consideration bears a constant ratio
to the dimension, such as total length, which is
employed as a standard. Where such a constant
ratio does not exist, it is necessary to compare
only fish of the same size or, which is more effi¬
cient, to compare the regression of the given
character on fish length (or head length).
Nichols and La Monte have recognized this in
the case of the length of the second dorsal and
anal fins and have combined a fish size with a
ratio of fin length to body or head length in
drawing up species descriptions, and have made

Morphometry of Tuna — SCHAEFER

117

some attempt to take this into account in their
key.

In order to establish the morphometric char¬
acteristics of the stock of yellowfin tuna off
Costa Rica, for subsequent comparison with
stocks from other parts of the Pacific, I have
computed for each dimension measured the
linear mean-square regression on the total
length, or on the length of head in the cases
of length of maxillary and diameter of iris.
Where the rate of increase of the character
measured is not proportional to the rate of in¬
crease of the total length, that is, where the
original variables do not yield a linear regres¬
sion, a transformation of variables has been
made so that the new variables yield a linear
relationship. This was necessary in three cases.
The rate of increase of length of second dorsal
and of anal fins is greater than that of total
length, while the rate of growth of the pectoral
fin is less than that of total length, over the
range of sizes examined.

The statistics describing the regressions are
tabulated in Table 2. The linear mean-square

regression is completely specified in each case
by the means of the two variables, the number
of specimens, the regression coefficient, and the
standard deviation from regression. The latter
is also called the standard error of estimate by
some authors. Where the regression of the orig¬
inal variables is linear we have also tabulated
the value of the y intercept in order to facili¬
tate determination of whether the dependent
variable may be considered to be in constant
proportion to the independent variable.

Over the range of sizes considered, all the
characters measured, with the exception of the
lengths of the pectoral, second dorsal, and anal
fins, bear a linear relationship to the length of
the fish. That is, the rate of increase of each of
the dimensions, with these exceptions, is pro¬
portional to the rate of increase in total length.
The proportion of the dimension considered to
the total length will be constant in a given case,
however, only if, in addition, the y intercept of
the regression line is zero. If the intercept dif¬
fers from zero, the value of the proportion will
vary with the size of the fish. Only for the re-

TABLE 2. STATISTICS DESCRIBING REGRESSIONS OF BODY PROPORTIONS OF YELLOWFIN

TUNA FROM COSTA RICA

INDEPENDENT

VARIABLE X

DEPENDENT VARIABLE y

X

y

ry.x

b

a

N

Total length

Head length .

931.6

261.4

4.39

0.2350

37.8

46

Total length

Snout to insertion first dorsal fin....

951.6

282.2

5.35

0.2635

31.5

46

Total length

Snout to insertion second dorsal fin

951.6

503.0

11.45

0.4768

49.4

46

Total length

Snout to insertion anal fin .

951.6

563.0

7.58

0.5351

53.8

46

Total length

Total length

Greatest body depth .

Pectoral insertion to insertion first

951.6

243.5

7.60

0.2555

0.4

46

dorsal .

955.1

144.5

5.83

0.1469

4.2

44

Total length

Length base first dorsal... .

958.1

222.4

10.15

0.2358

-3.5

45

Total length

Length longest (first) dorsal ray .

948.7

112.8

5.22

0.1178

1.2

45

Total length

Length longest dorsal finlet .

951.6

30.9

2.00

0.03361

-1.1

46

Log total length

Log total length

Length pectoral fin . . .

2.9640

253.5

7.52

445.9

1.694

45

Log length second dorsal fin .

2.9640

2.1361

0.0362

45

Log total length

Log length anal fin .

2.9668

2.1711

0.0414

1.832

44

Length second dorsal fin

Length anal fin .

151.2

164.9

17.62

1.150

-9.0

44

Length of head

Diameter of iris .

266.8

33.7

1.303

0.06038

17.6

35

Length of head

Length of maxillary .

261.4

100.3

2.17

0.3781

1.5

46

Log total length

Log weight (kilos) . .

2.9538

1.1222

0.0266

2.940

93

Logarithms are to the base 10.
x_— mean of values of x.
y = mean of values of y.

ry.x = standard deviation from regression (standard error of estimate),
b = regression coefficient of y on x.
a = y intercept of regression line.

N = number of specimens.

118

PACIFIC SCIENCE, Vol. II, April, 1948

gression of greatest body depth on total length,
and the regression of length of longest first dor¬
sal ray on total length do the intercepts fail to
differ significantly from zero. For the pectoral
insertion to insertion of first dorsal, and length
of base of first dorsal, the regressions on total
length have y intercepts differing significantly
but yet so slightly from zero that the expression
of these measurements as percentages of total
length would result in a negligibly small error
from this source. This is also true for the re¬
gression of length of maxillary on length of
head. For the remaining characters, the size of
the fish has a considerable effect on the dimen¬
sion expressed as a percentage of total length,
and the same is true for diameter of iris ex¬
pressed as a percentage of head length.

The lengths of the second dorsal and anal
fins are in proportion to the 1.69 power and
1.83 power of the total length, respectively.
This very rapid increase of fin length with fish
length follows the equation

y=cxh . . . (1)

where y is the fin length, x is the total length,
b is the value indicated in Table 2, and c is an
arbitrary constant depending on the units of
measurement. (Here, where the measurements
are in millimeters, £=5.45 X^O-4 for the anal
fin and c=1.30X10-3 for the second dorsal.)
The standard deviation from regression, con¬
verted from logarithms as expressed in Table 2
to percentages, amounts to 8.7 per cent for the
second dorsal fin and 10.0 per cent for the anal
fin. If the deviations were randomly assorted
by fish size we would expect to find in about
one case in 100 a fish with second dorsal fin
varying as much as 23 per cent from the aver¬
age for a given size of fish, and a fish with anal
fin varying as much as 29 per cent from the
average for a given size of fish. Examination
of the data, however, has shown that the devia¬
tions are not entirely randomly assorted, but
that they are to some degree related to size of
fish, the variability, on a logarithmic plot, being
somewhat greater for the larger fish. This

means that at the larger sizes, say over about
a meter in total length, the variation may be
expected to be somewhat greater than the
numerical values indicated, while for small fish
it will be less.

The deviations from the average for a given
size are not attributable to the sex of the fish
in any case. No sexual dimorphism of fin lengths
or other measurements has been found from our
data.

The exponents b in (1) for anal fin length
and second dorsal fin length are so nearly equal
that the regression of the latter on the former
is linear or nearly so. The least-squares fit to
this regression indicates that, on the average, for
any given size in the range investigated the anal
is somewhat longer than the second dorsal.

The pectoral fin grows more slowly than the
length of the fish over this range of sizes. It
was found that, for this range, the relationship
between fin length y and total length x may be
expressed in the form

y = 446 log10 x — 1068 ( 2 ) .

There is no recognizable difference between
the measurements of the two observers with the
single exception of the character "length of base
of first dorsal.” Examination of the data indi¬
cates a tendency for the measurements of this
distance by Mr. Marr to be a little smaller than
those of the author. The statistics of the linear
mean-square regressions, computed from the
data of the two observers, are:

Schaefer Marr

Number of specimens.. 21 24

Mean total length (x

in Table 2) . 1047.6 879.8

Mean length of base of
first dorsal (y in
Table 2) . 250.0 198.1

Standard deviation

from regression . 7.35 7.29

Regression coefficient
(b in Table 2)

0.2330 0.2090

Morphometry of Tuna- — SCHAEFER

119

The slopes of these regressions do not differ
significantly from those of the pooled data, the
statistics for which may be found in Table 2.
The levels of the lines however, that is, the
values of x for a given value of y, are less in
the case of Mr. Marr’s measurements and
greater in the case of the author’s measure¬
ments than would ordinarily be expected from
random sampling from a population having the
values estimated from the pooled data. The
probabilities, in each case, lie between 0.02 and
0.01. There seems, therefore, to be a real dif¬
ference between the two sets of measurements.
Since the measurements by the two persons were
made on different groups of fish, there is a pos¬
sibility that this difference represents an actual
difference between the two groups of tunas.
However, no difference between the two is
shown by the data on other dimensions; in par¬
ticular no difference between the distances of
snout to insertion of first dorsal, snout to inser¬
tion of second dorsal or body depth, one of
which should reflect any actual difference in
the distance between the insertions of the first
and second dorsal fins. Therefore, it seems most
likely that the difference represents a differ¬
ence in the measuring by the two observers, al¬
though it is not apparent just how this arose.

LENGTH-WEIGHT RELATIONSHIP

In addition to the data recorded in Table 1,
the lengths and weights were determined for a
number of specimens over the same size range,
a total of 93 in all. The regression of logarithm
of weight on logarithm of length is described
in Table 2. This relationship corresponds to the
equation

W=2.74X10-8 L2-940 . (3)

where L is the total length in millimeters and
W is the weight in kilos.

TAXONOMIC NOTES

The yellowfin tuna off Costa Rica appear to
be assignable to the species Neothunnus wid¬

er opterus (Temminck and Schlegel). The speci¬
mens examined by me agree closely enough with
the descriptions of that species in Kishinouye’s
monograph (1923) and in Godsil and Byers’
paper ( 1944). The latter concluded that all the
yellowfin tunas of the Pacific examined by them,
including specimens from Costa Rica, were
members of this single species. Nichols and
La Monte’s data (1941) on length of second
dorsal and anal fins for N. macropterus, which
they consider to be a synonym of N. albacora,
given for various fish lengths, fall within the
expected limits of variation of the values esti¬
mated for those same lengths from the regres¬
sions in our Table 2. The only exception to
this is their Portuguese specimen 5 feet long,
which had dorsal and anal lobes contained
"2.6 to 2.8 times in the length.” From our data,
it is estimated that at this length only about 1
per cent of specimens would have second dor¬
sals contained less than 3.9 times in the length,
or anals contained less than 3.2 times in the
length.

Frade (1929, 1931) has found that there are
rather distinct anatomical differences between
the Portuguese yellowfin, N. albacora, and the
Pacific yellowfin, N. macropterus. He has found
that the air bladder of N. albacora has a large
dorsal diverticulum, which is not present on the
air bladder of N. macropterus according to
Kishinouye’s description, which is confirmed by
the study of Godsil and Byers (1944). Kishi¬
nouye’s Figure Q (page 373) also shows the
cutaneous artery of N. macropterus arising at
the level of the 9th vertebra, whereas that of the
Portuguese yellowfin was found by Frade to
arise at the level of the 8th. This difference is
not confirmed by Godsil and Byers, however,
who also found the cutaneous artery arising at
the level of the 8th vertebra in their specimens
of N. macropterus.

Decision as to whether or not the variety of
yellowfin tuna from the Hawaiian Islands,
having very long anal and second dorsal fins at
larger sizes, is distinct from the more common
variety, and, if so, whether the difference is spe-

120

PACIFIC SCIENCE, Vol. II, April, 1948

cific or only racial, must await examination of
a series from the type locality. This fish was
described from Hawaii by Jordan and Ever-
mann (1926) as Neothunnus itosibi. Nichols
and La Monte ( 1941 ) state that "at a weight of
around 100 pounds the fin lobes . . . from the
tropical Pacific compare in development with
the maximum obtained by the Common Yellow-
fin. At about 4 feet 5 to 9 inches, weighing 140
pounds or less, the lobes are contained 2.1 to
2.8 times in standard length . . .” These anal
and second dorsal fins seem to be significantly
longer than we would expect to find in N.
macropterus, from Costa Rica at least. How¬
ever, it is also possible that the variability of
fin length is greater among the members of the
Hawaiian stock. Godsil and Byers’ Hawaiian
specimens do not help with the solution of this
problem since they were of very small size,
537-573 mm.

It is of interest to note that Frade (1931)
has found that in the vicinity of Portugal there
exist for the same size two types of N. alb ac or a:
". . . comme pour N. macropterus du Pacifique,
il existe pour la meme taille deux types de
N. albacora: l’un a 2e dorsale et anales longues,
correspondant a N. macropterus forma itosibi
et 1’autre a 2e dorsale et anales courtes, cor¬
respondant a N. macropterus forma macrop-
terus.”

Beebe and Tee-Van (1936) consider that
". . . the various nominal forms of the yellow-
finned tuna belong to the same species, and that
the forms typified by the large Allison’s tuna
represent but large-finned specimens of the
smaller short-finned individuals.” Walford
(1937) found that among yellowfin tunas ex¬
amined in California canneries . . the dorsal
and anal fins were of all lengths, intergrading to
such an extent that it is impossible to separate
them into two groups. In general, the largest,
consequently the oldest fish had the longest
fins.”

It seems well established that wherever yel¬
lowfin tuna occur, both short-finned and long-
finned individuals are encountered, and that this
character is correlated with the size of the fish.
Whether racial differences in the regressions of
fin length on fish size will be found between
localities we cannot say at this stage of our
knowledge. Within the Costa Rican stock, how¬
ever, as represented by the present sample, there
seems to be but a single race of yellowfin tuna.

REFERENCES

Beebe, William, and John Tee-Van. 1936.
Systematic notes on Bermudian and West
Indian tunas of the genera Parathunnus and
Neothunnus. Zoologica [New York] 2 1 ( 14) :
177-194.

Fisher, R. A. 1934. Statistical methods for re¬
search workers. Ed. 5, xii-f- 319 p. Oliver &
Boyd, London.

Frade, F. 1929. Sur quelques thons peu connus
de l’Atlantique. Soc. Portug. de Sci. Nat. Bui.
10(20): 227-234.

- - - — 1931. Donees biometriques sur trois

especes de thons de l’Atlantique orientale.
Cons. Perm. Int. pour VFxplor. de la Mer,
Rapp, et Proc.-Verb. 70: 117-126.

Godsil, H. C, and R. D. Byers. 1944. A sys¬
tematic study of the Pacific tunas. Calif. Div.
Fish and Game, Fish. Bui. 60: 1-131.
Jordan, D. S., and B. W. Evermann. 1926.
A review of the giant mackerel-like fishes,
tunnies, spearfishes and swordfishes. Calif.
Acad. Sci., Occas. Papers 12: 1-113.
Kishinouye, K. 1923. Contributions to the
comparative study of the so-called scombroid
fishes. Tokyo Imp. Unin., Col. Agr. Jour.
8(3): 293-475.

Nichols, J. T., and F. R. La Monte. 1941.
Yellowfin, Allison’s and related tunas. Ichthy.
Contrib. Int. Game Fish Assoc. 1(3) : 27-32.
Walford, L. 1937. Marine game fishes of the
Pacific Coast from Alaska to the equator.
xxix-f- 205 p., 69 pi. Univ. California Press,
Berkeley.

Notes on the Marianas Mallard1

Yoshimaro Yamashina

The Marianas Mallard, Anas oustaleti, is
one of the most interesting species of Micro-
nesian birds. Found only on Saipan, Tinian,
and Guam, three small islands of the Marianas,
it is very scarce both in museum collections and
in its native habitat. Special precautions, there¬
fore, should be taken to protect it from ex¬
termination.

The Marianas Mallard was first reported by
Bonaparte (1856) as Anas boschas a. Freycineti
( nom . nud.) based on the single specimen pre¬
served in the Paris Museum, which was said to
have come from "Les Isles Malouines.” Later
Salvadori (1894) described the same specimen
as Anas oustaleti, but its exact abode was not
known until Oustalet (1896) reported six spe¬
cimens from Guam. Afterwards, Hartert ( 1898)
added Saipan to its range, and Phillips (1923)
added Tinian. Its habits and status, however,
were unknown until Japanese investigators
studied them after 1931.

Anas oustaleti is found in two places in
Saipan. One is a lagoon covered with man¬
groves, north of Garapan village, where Hyo-
jiro Orii, my collector, obtained 11 specimens
in 1931 and Takeo Kozima later took four more
for me. The second place is a small fresh-water
pond called Charan-Kanoa, south of Garapan
village, surrounded by a rich growth of aquatic
plants.

Tinian Island, south of Saipan, is at present
the main habitat of this duck. Hagoi Pond, in
the northern part of that island, is a small body
of fresh water surrounded by about 40 acres

1 Submitted with the approval of the Chief, Natural
Resources Section, General Headquarters, Supreme
Commander for the Allied Powers, Tokyo, Japan.
Manuscript received November 4, 1947.

of marsh. My collectors obtained nine speci¬
mens there between 1931 and 1933. Kuroda
obtained 10 more specimens from this locality
in 1936, and in 1940 he received four live birds
which he kept in his aviary in Tokyo for several
years. According to Kuroda, his collector ob¬
served two flocks of Anas oustaleti there at the
time, each containing 50 to 60 ducks, the largest
flocks of this duck ever seen.

In Guam, where Anas oustaleti was first dis¬
covered, the Marianas Mallard seems to be least
plentiful in comparison with the other two
islands. According to Phillips (1923) only a
few ducks remain in the Talafofo Valley.

In general habits Anas oustaleti resembles
Anas poecilorhyncha (including superciliosa) ,
which is widely distributed from New Zealand
through Micronesia and the Philippines to
China and Japan. The duck resides in lagoons
and fresh-water ponds throughout the year, and
according to Phillips (1923) breeds in Guam
in January and February, at the end of the rainy
season. On Saipan and Tinian eggs and duck¬
lings have been collected in June and July,
which are in the dry season (Kuroda, 1942).
Saipan natives speak of their breeding from
January to March. Thus, this duck seems to
breed at all seasons of the year. Kuroda ( 1942 )
described a nest with seven eggs taken July 4,
1941, at Hagoi Pond, Tinian Island. It was
found among the rushes, and was made of dead
leaves, stems, and roots, and lined with down.
The eggs were grayish white with a pale green¬
ish tinge, and measured 61.6 X 38.9 mm.

Anas oustaleti has two types of plumage, one
resembling Anas platyrhynchos, and the other
Anas poecilorhyncha. Therefore, certain earlier

[121]

122

PACIFIC SCIENCE, Vol. II, April, 1948

investigators, who saw only the platyrhynchos
type, were convinced that Anas oustaleti is a
near relative of Anas platyrhynchos (Salvadori,
1894), while others, who saw only the poeci-
lorhyncha type, thought it nothing but a sub¬
species of Anas superciliosa {—Anas poecilo-
rhyncha superciliosa) (Hartert, 1930). I was
inclined to think, after obtaining a series of spe¬
cimens from Saipan Island, which includes adult
males of both types, that Anas oustaleti must
have two color phases, a platyrhynchos type and
a poecilorhyncha type. Kuroda later proved this
conjecture correct by observing the moult of
living specimens (Kuroda, 1941, 1942). The
descriptions of both types are as follows:

A. PLATYRHYNCHOS type:

Adult male in nuptial plumage: Whole head
is dark green, except at the sides where buff
feathers are plentifully intermingled, a dark
brown streak through the eye, and faint white
ring on the lower neck. Feathers on scapulars
and sides of body are as those of Anas poeci¬
lorhyncha. Sides of body are vermiculated but
some brown feathers are found even in the full
nuptial plumage. Upper breast is dark reddish
chestnut with dusky spots. Upper and under tail
coverts are as in Anas platyrhynchos. Speculum
is as that of Anas platyrhynchos, but the tips
of the greater coverts are buff instead of white.
Central tail feathers are more or less curled
upward. Base of bill is black, tip is olive color.
Iris is dark brown. Feet, reddish-orange, webs
darker.

Adult male in eclipse plumage: Resembles
the eclipse plumage of Anas platyrhynchos.

A. POECILORHYNCHA type:

Adult male in nuptial plumage: Resembles
Anas poecilorhyncha pelewensis from the Palau
Islands and Truk Island, but sides of head are
browner, superciliary stripes and ground color
of cheeks are more buffy. Feathers on upper
breast and sides of body are more broadly edged
with brown. Speculum is usually violet-purple
as in the platyrhynchos type, but in two speci¬

mens from Saipan and Tinian, respectively, it is
dark green as in Anas poecilorhyncha pelewen¬
sis. Tips of the secondaries are usually white,
but sometimes very faint as in Anas poecilorhyn¬
cha pelewensis, and in one specimen from Sai¬
pan they are buffy. Bill is olive color with a
black spot in the center of the upper mandible.
Iris, dark brown. Feet, dark orange, darker in
joints and webs.

Adult male in eclipse plumage: Same as the
nuptial plumage.

Thus it is apparent that the platyrhynchos
type is composed of characteristics 90 per cent
peculiar to Anas platyrhynchos, and 10 per cent
similar to Anas poecilorhyncha, while the poe¬
cilorhyncha type reverses the percentages, being
90 per cent similar to poecilorhyncha and 10
per cent to platyrhynchos. Therefore, Anas ous¬
taleti may conceivably have originated from a
compound of Anas platyrhynchos and Anas poe¬
cilorhyncha stock.

Having compared a number of both types, I
find the platyrhynchos type to be less numerous
than the poecilorhyncha. In the 50 known spe¬
cimens of Anas oustaleti (36 of them formerly
in Japanese collections) only six specimens were
the platyrhynchos type. However, the ratio be¬
tween the numbers of both types varies on each
island. On Saipan, at the northernmost end of
its distribution, four of the six known adult
males were platyrhynchos type, while on Tinian,
only one platyrhynchos type was discovered
among 24 specimens. On Guam, the platyrhyn¬
chos type seems to be very rare, only one having
been reported among about a dozen specimens.

There have been two hypotheses advanced as
to the origin of Anas oustaleti. Phillips (1923)
and Hartert (1930) thought it probably stem¬
med from Anas superciliosa (=Anas poecilor¬
hyncha superciliosa) , but this hypothesis does
not explain the occurrence of the platyrhynchos
type. Kuroda (1941-1942) and Delacour and
Mayr ( 1945 ) supposed that Anas oustaleti must
have descended from Anas platyrhynchos stock
which arrived long ago from the north, chiefly

Marianas Mallard-— YAMASHINA

123

because Anas oustaleti resembles Anas platyr-
hynchos, especially in the color of the speculum.
However, this assumption seems hardly to fit the
case, because it overlooks the remarkable simi¬
larity of the poecilorhyncha type to Anas poeci-
lorhyncha superciliosa. Furthermore, the specu¬
lum of Anas oustaleti does not always resemble
that of Anas platyrhynchos, being sometimes
dark green without conspicuous white edgings,
and hence more like that of Anas poecilorhyn¬
cha superciliosa.

On the other hand, it is well known that
Anas platyrhynchos can easily be crossed with
Anas poecilorhyncha (including superciliosa) ,
producing offspring very similar to the platy¬
rhynchos type of Anas oustaleti. In the male of
F1 offspring of this cross in nuptial plumage
the head is green except on the sides, where
buff feathers are mixed plentifully, with a dark
stripe through the eye. The mantle is as that of
Anas poecilorhyncha, but breast, sides of body,
wing, and tail are similar to those of Anas
platyrhynchos, the central tail feathers curling
backward. The female of this cross is inter¬
mediate between the parental species, showing
the dark stripes on the cheeks found only in
Anas poecilorhyncha. A particularly interest¬
ing fact is that the speculum of Anas platyrhyn¬
chos acts as dominant over that of Anas poe¬
cilorhyncha. The platyrhynchos type speculum
in Anas oustaleti might be derived in this way,
and the occasional occurrence of the poecilo¬
rhyncha type speculum may be explained by
the formation of a homozygous condition gov¬
erning the recessive gene of the poecilorhyncha
type speculum. From these evidences we may
deduce that Anas oustaleti originated from
hybridization between a local race of Anas
poecilorhyncha, probably formerly resident on
the Marianas, and Anas platyrhynchos stock
which straggled there occasionally from the
north. The- scarcity of the platyrhynchos type in
the southern islands strengthens this supposi¬
tion. The opportunity for hybridization should
occur more rarely in the south, and thus more

frequent back-crossing of the hybrid with the
indigenous Anas poecilorhyncha on Tinian and
Guam explains the superabundance there of the
poecilorhyncha type. As the hybridization
should have taken place more frequently to the
north in Saipan, the ratio of occurrence of the
platyrhynchos type is logically higher there.

There is no evidence of previous occurrence
of Anas poecilorhyncha (including superciliosa)
in the Marianas Islands, but it seems logical,
because Anas poecilorhyncha is found in the
neighboring Caroline and Palau Islands, as well
as in the Philippines and Japan. Neither is
Anas platyrhynchos recorded from the Marianas,
but its winter range reaches in the east to the
Hawaiian Islands and in the west to Borneo
and South India, and it winters frequently just
north of the Marianas in the Bonin and Volcano
Islands. Therefore, it is not unreasonable to
suppose that Anas platyrhynchos has occasion¬
ally straggled to the Marianas, remained and
produced hybrids with an Anas poecilorhyncha
stock which was formerly indigenous there.2

According to recent cytogenetical investiga¬
tions, the interspecies hybrid shows more or less
sterility due to a dissimilarity of parental chro¬
mosome constitutions. This sterility, however,
can be reduced by pure breeding after back-
crossing, which forms new homozygotes hav¬
ing a chromosome constitution different from
either of the parental species. The possibility
of species formation in this manner was sug¬
gested by Danforth and Sandnes ( 1939). In the
Marianas Islands, the habitat of this duck is
restricted to very small ponds or lagoons found
in each island. Consequently, the population is
very small, and pure blood lines may be carried
in each population occupying a certain pond or
lagoon. The phenomenon suggested by Dan¬
forth and Sandnes (1939) seems to be realized
by these circumstances.

2 1 believe Anas wyvilliana of the Hawaiian Islands
and Anas laysanensis of Laysan Island originated from
Anas platyrhynchos stock alone, because no dichro¬
matic phases have been reported in either species and
there is no evidence that a second species ever occurred
in these islands.

124

PACIFIC SCIENCE, Vol. II, April, 1948

Speciation by hybridization either in natural
or under artificial conditions is often reported
in the plant kingdom, but it is exceedingly rare
among animals. Anas oustaleti could have de¬
veloped only in the special environment of the
Marianas Islands, and is therefore noteworthy
from the viewpoint of evolution.

REFERENCES

Bonaparte, C. L. 1856. {Paris} Acad, des
Sci., Compt. Rend. 43: 649.

Danforth, C. H., and G. Sandnes. 1939.
Behavior of genes in intergeneric crosses.
Effect of two dominant genes on color in
pheasant hybrids. Jour. Hered. 30: 537-542.
Delacour, J., and E. Mayr. 1945. The family
Anatidae. Wilson Bui. 57: 3-55.

Hartert, E. 1898. On the birds of the Mari¬
anne Islands. Novitates Zool. 5: 66, 69.

— - 1930. List of the birds collected by

Ernst Mayr. Novitates Zool. 36: 112.

Kuroda, N. 1941. [A study of Anas oustaleti.']
Tori, 11 (51-52): 99-119. [In Japanese.}

— - — 1942. [A study of Anas oustaleti.

Pt. II.} Tori, 11 (53-54): 443-448. [In
Japanese.}

OUSTALET, M. E. 1896. Les mammiferes et les
oiseaux des lies Mariannes. Nouv. Arch. Mus.
d’Hist. Nat. 2 (2): 49-50.

Phillips, J. C. 1923. A natural history of the
ducks. Vol. 2: 53-54. Houghton Mifflin,
Boston.

Salvadori, T. 1894. Anas oustaleti and Nyroca
innotata, spp. nov. Brit. Ornith. Club , Bui.
4 (20): 1.

A New Blennoid Fish from Hawaii

Vernon E. Brock1

The genus Petroscirtes is one of the genera
which are particularly characteristic of the Indo-
Australian ichthy-fauna of the western tropical
Pacific. Aside from a single dubious record,
discussed below, Petroscirtes has not been re¬
ported from Hawaiian waters. However, the
discovery of the blennoid fish described here
establishes Petroscirtes, without doubt, as a part
of the Hawaiian ichthy-fauna.

Petroscirtes ewaensis new species

Holotype. United States National Museum
No. 133821, 87.4 mm. in standard length. Taken
off Ewa Beach, Oahu, T. H., January 3, 1947.
Caught in the open end of a pipe brought up
from the bottom in 30 feet of water.

Dorsal 46. Anal 30. Pectoral 12. Pelvic (I?)
3. Body scaleless, slender, somewhat compressed,
greatest depth at vent. Snout pointed, mouth
inferior, gape extending approximately to middle
of eye. Hind border of eye about midway in
head length. Head 5.8 in standard length, depth
7.0; greatest body width 13.2; dorsal fin base
1.2, anal fin base 1.7, pectoral length 7.7, from
snout tip to vent 2.7. Snout 3.4 in head length,
eye 4.8, large canine tooth 5.4, interorbital 3.3,
fifth dorsal fin ray 3.1. Single row of 45 in¬
cisors across front of lower jaw, with large
curved canine at each side; canine received into
socket in upper jaw when mouth closed. Single
row of about 31 incisors across front of upper
jaw; no canines present. Row of 4 short ten¬
tacles across base of chin. Row of 8 pores on
top of snout from about midway between fore
margin of eye and snout tip back to upper hind-

1 Director, Division of Fish and Game, Territorial
Board of Agriculture and Forestry, Honolulu, Hawaii.
Manuscript received June 7, 1947.

border of opercle; 1 pore just before dorsal
origin; 5 or 6 pores along preopercle margin;
2 pores forward above and below and 1 be¬
hind and below eye. Dorsal origin about mid¬
way between hindborder of eye and edge of
opercle. Dorsal, low and long, none of the rays
elevated, not joined to caudal. Origin of pelvics
about one-half their length ahead of gill open¬
ing.

In life, general color a rich brick-red with two
lateral, black-edged, iridescent blue bands. The
upper band on dorsal part of side about its
width below dorsal base, about equal to diameter
of pupil of eye, extending from snout tip to just
above eye, then back to caudal fin base and for
a short distance out on caudal. The lower band,
wider than diameter of eye and about as wide
as interspace between it and upper band, origin
on tip of snout as fine line, expands and curves
beneath eye, widest about opposite vent then
tapered to point on basal part of caudal. Nar¬
row, dark-edged, iridescent blue band from
snout tip along top of head to dorsal insertion.
Dorsal and anal brick-red; end of rays narrowly
tipped with blue. Caudal reddish, narrow blue
line on upper and lower edge extending pos¬
teriorly for less than half its length. Pelvic rays
faintly darkened near tips. Pectoral clear.

Color faded in preservative. Two lateral bands
a slate blue with dark brownish-black margins;
red interspaces faded to a dead, light grayish-
white.

The name ewaensis is given in reference to
Ewa, the district of Oahu adjacent to the place
of capture.

Discussion: The discovery of this species in
Hawaiian waters serves both to demonstrate fur-

U25]

New Blennoid Fish-— BROCK

127

ther the basic affinity of the Hawaiian ichthy-
fauna with that of the Indo-Australian faunal
province for which, of course, a redundancy of
proof exists, and to indicate the marginal posi¬
tion of the Hawaiian ichthy-fauna in that
province. The genus Petroscirtes has member
species widely distributed throughout the west¬
ern tropical Pacific, but aside from one dubious
record [Fowler’s remarks on Petroscirtes fila¬
ment osus (Valenciennes) (1928: 428)} no
members of the genus have yet been reported
from Hawaiian waters. The discovery of P. ewa-

ensis establishes the existence of Petroscirtes in
the Hawaiian portion of the Indo-Australian
fauna province, but as it is a new species and
hence never reported from any other portion of
the province, this fact may be another bit of
evidence of the relative isolation of the Ha¬
waiian portion of that province.

REFERENCE

Fowler, Henry W. 1928. The fishes of
Oceania, iii— (— 540 p., 82 fig., 49 pi. Bernice
P. Bishop Mus. Mem. 10. Honolulu.

NOTES

Fishes Taken in Wellington Harbour

This record of the fishes of Wellington Har¬
bour is the result of a series of notes collected at
intervals during a period of over 20 years. My
first paper ( New Zeal. Jour. Sci. and Technol.
1(5): 268-271, 1918) supplies a record of the
edible fishes of Wellington; and to this, addi¬
tions have been made in subsequent papers.
This summary of fish fauna deals exclusively
with fishes recorded from the harbour, and does
not by any means claim to be a complete record;
for we doubtless have many aquatic visitors who
do not remain for any length of time.

The climate of Wellington is very variable;
but the temperature of the harbour water re¬
mains relatively constant unless the summer is
consistently warm, when all sorts of northern
species such as John Dory, Zeus faber, and
northern mullet, Mugil cephalus, invade our
waters. It also is well to remember that we are
dealing with shoals of fishes which are anything
but consistent in their appearance and are at
the mercy of a variable and ever-changing food
supply, so that species recorded in a given year
may not be seen again for a comparatively long
period. The residual fish fauna of the harbour
is, I think, small. Herring, Agonostomus fors-
teri, are the most common species, and next to
these the spotties, Rseudolabrus celidotus, which
are found in abundance in many localities.
Shoals of larger fishes, notably kahawai, Arripis
trutta, and barracouta, T by r sites atun, prey on
these smaller fishes in the summer months.

It is probable that the relative abundance of
fish life in the harbour is on the decrease. The
traffic of large vessels at the wharves appears to
have reduced the number of fishes in that area;
and the fishing grounds off Seatoun and Worser
Bay are either fished out or now partly deserted.

1. Geotria australis Gray. Lamprey.

In the "piharau stage” this lamprey lives in
Wellington Harbour until it is well over 1 foot
long, and then migrates up the Hutt River to
spawn. Several specimens of a very beautiful

green-blue colour have been taken by the Petone
fishermen off Somes Island and forwarded to
the Dominion Museum for identification.

2. Notorhynchus pectorosus ( Garman ) .

Seven-gilled Shark.

This species appears in the harbour at irregu¬
lar intervals. It is probable that it enters only
in pursuit of small fishes. Only two specimens
have been examined. (Phillipps, New Zeal.
Dominion Mus. Rec. 1(2): 5, 1946).

3. Eulamia brachyurus (Gunther). Whaler.

This is a species that is rarely taken in the
harbour. In the Dominion Museum there is a
cast of a specimen a little over 9 feet long from
off Somes Island.

4. Raja nasuta Muller and Henle. Skate.

Sometimes small skates are taken by fisher¬
men off Rona Bay.

5. Callorhynchus milii Bory. Elephant Fish.

Some years ago a small elephant fish was
taken off Rona Bay and sent to the Museum for
identification. This species, so abundant in
Otago Harbour, is almost unknown north of
Cook Strait.

6. Sardinops neopilchardus ( Steindachner ) .

Pilchard.

A note on the occurrence of this species in
Wellington Harbour is given by me in New
Zeal. Jour. Sci. and Technol. 7(3): 191, 1924.
Shoals enter the harbour in the winter, about
August, and are indicated by the diving of gan-
nets.

7. Gonorhvnchus gonorynchus (Linnaeus).

Sand Fish.

This species is sometimes taken off Rona Bay
and specimens are occasionally forwarded to the
Museum for identification. The sand fish may
be quite common as it escapes the drag net by
burrowing.

[128]

NOTES

8. Leptocephalus conger (Linnaeus). Conger

Eel.

The conger eel is not uncommon off East¬
bourne in February.

9. Hippocampus abdominalis Lesson. Sea¬

horse.

Several specimens have been taken close to
Oriental Bay.

10. Coelorhynchus australis (Richardson).

Javelin Fish.

Occasional examples have been taken off
Rona Bay and forwarded to the Museum.

1 1. Macruronus novae-zelandiae ( Hector ) .

Whiptail.

Occasionally, young are taken in the early
summer months.

12. Merluccius gayi (Guichenot). Whiting.

Occasional examples have been taken in the
harbour.

13. Physiculus bachus (Bloch and Schn. ) .

Red Cod.

One or two small specimens have been re¬
ported, but the species is rare.

14. Zeus faber Linnaeus. John Dory.

Specimens are occasionally taken off Somes
Island in the summer months.

15. Rhombosolea plebeia (Richardson). Sand

Flounder.

This species is taken off Seatoun and Rona
Bay throughout most of the year.

16. Rhombosolea leporina Gunther. Yellow-

belly.

This flounder is caught with sand flounders,
but never in any quantity.

17. Peltorhampus novae-zeelandiae Gunther.

Sole.

This species is taken in small numbers off
Rona Bay.

18. Pelotretus flavilatus Waite. Lemon Sole.
Small numbers are taken off Rona Bay.

19. Agonostomus forsteri (Cuv. and Val.)

Yellow-eyed Mullet.

129

This mullet is popularly called "herring”; and
in North Auckland it is called "sprat.” Young
are common throughout the year in both brack¬
ish and salt water.

20. Mugil cephalus Linnaeus. Northern Mullet.

Small numbers are taken at Rona Bay during
periods of warm weather.

21. Seriolella brama (Gunther). Warehou.

Warehou are taken just inside the harbour
"Heads” in the spring months.

22. Polyprion oxygeneios (Bloch and Schn.).

Hapuku or Groper.

Mr. J. Patterson informs me that he saw small
groper taken off Wellington wharves in the
1890’s. They now are rare inside the harbour
entrance.

23. Longirostrum platessa (Cuv. and Val.).

Trevally.

The trevally is taken during summer in sev¬
eral parts of the harbour, but mostly near the
entrance.

24. Trachurus novae-zelandiae Richardson.

Horse Mackerel.

This species is fairly common in warm wea¬
ther but disappears during winter. Mr. J. Pat¬
terson informs us that in the 1890’s this was a
common species around Wellington wharves.

25. Seriola lalandi Cuv. and Val. Kingfish.

The kingfish apparently follows pilchards and
other fishes into the harbour. It is caught by line
fishermen off Kaiwarra and Ngahauranga, and
taken by Petone fishermen off Somes Island. It
is a difficult fish to catch, and small specimens
are the rule.

26. Arripis trutta (Forster). Kahawai.

In the summer months this species is not un¬
common in the harbour, where with kingfish it
follows shoals of smaller fishes.

27. Pagrosomus auratus (Forster). Snapper.

At intervals throughout part of the year snap¬
per may be taken in southern parts of the har¬
bour.

28. Scorpis violaceus (Hutton). Maomao.

The first maomao described was taken in
Wellington Harbour (Hutton, New Zeal. Inst.,

130

PACIFIC SCIENCE, Vol. II, April, 1948

Trans, and Proc. 5: 261, 1873). On March 9,
1941, Mr. B. P. R. Phillipps, fishing at Kaiwarra,
caught the second reported specimen. This was
a small maomao, 147 mm. long, and 60 mm.
high at the anal origin. There were traces of
blue on the body and red along the lateral line.
The radial formula is D.10-j-28; A. 3+28;
V.l+5. It is quite possible that this species is
a great deal more common in Wellington Har¬
bour than is indicated by the paucity of reported
specimens.

29. Dactylopagrus macropterus (Forster).

Tarakihi.

This species sometimes is taken in southern
portions of the harbour.

30. Latridopsis ciliaris (Forster). Moki.

Moki sometimes are caught in the harbour
but are said to be not nearly so common as they
were 40 years ago.

31. Pseudolabrus celidotus (Forster). Spotty.

The spotty is more or less common in rocky
localities around the edge of the harbour.

32. Parapercis colias (Forster). Blue Cod.

Small numbers are taken by amateur fisher¬
men in certain years only. About January, 1946,
a small example (about 1 foot long) was taken
off Wellington wharves by Mr. B. P. R. Phil¬
lipps, the only specimen I have seen taken in
recent years.

33-Thyrsites atun (Euphrasen). Barracouta.

Formerly, the barracouta was common in the
harbour, but is not fished for now to any extent.

34. Jordanidia solandri (Cuv. and Val.). Hake

or Southern Kingfish.

The hake is taken during February and some¬
times in March, but is not common.

35. Acanthoclinus quadridactylus (Bloch and

Schn. ) . Taumaka.

This species is rare in the harbour. Some
years ago Mr. W. O. S. Phillipps caught a large
specimen when line-fishing at Pipitea Point.

36. Hemerocoetes waitei Regan.

This species has occasionally been taken in
drag nets in the vicinity of Eastbourne.

37. Tripterygion varium (Forster). Cock-a-

bully.

A type which eventually may prove to be new
has D.7+23+15; C.2+11+2; A.29; P.19;
V.2; and 30 raised scales in the lateral line.
However, much comparative material on this
species is required before we can determine the
limits of its variation.

38. Congiopodus leucopaecilus (Richardson).

Pig Fish.

Occasional small examples have been sent
into the Museum from Rona Bay fishermen.

39. Chelidonichthys kumu (Lesson and Gar-

not). Red Gurnard.

Small numbers were taken off Seatoun and
Rona Bay.

40. Diplocrepis puniceus ( Richardson ) .

Sucker.

This species has been recorded at Seatoun by
Dr. W. R. B. Oliver.

41. Trachelochismus littoreus (Forster).

This is another species recorded at Seatoun
by Dr. Oliver.

42. Cantherines scaber (Forster). Leather

Jacket.

The common leather jacket occasionally is
taken in the harbour, small specimens arriving
at the Museum for examination every few years.

— W. J. Phillipps , Dominion Museum, Well¬
ington, New Zealand.

Transfer of Hawaiian Volcano Observatory

The Hawaiian Volcano Observatory, at
Kilauea Volcano, Hawaii, has been transferred
from the National Park Service to the United
States Geological Survey. The change was made
on December 28, 1947, by order of the Secre¬
tary of the Interior. The larger part of the equip¬
ment of the Observatory will be moved from its
present location in the Park Naturalist Building
to the now unused Uwekahuna Museum on the
rim of Kilauea Crater about 2 miles farther
west. The museum building will be remodeled
to house the shop, laboratories, and office of the
Observatory, which will continue its study of
volcanic processes, and will serve as a training
station in connection with the Geological Sur¬
vey’s studies of volcanic regions in the Pacific
and elsewhere. The Volcano Observatory is dedi¬
cated to the measurement and observation of
volcanic behavior and carries on studies in seis¬
mology, geomagnetism, chemistry, physics, and
other fields related to volcanic activity. Students
of volcanology and local friends of the Observa¬
tory will applaud these plans to continue and
strengthen the work of this scientific institution
which has been one of the pioneer contributors
to the understanding of volcanoes during its
nearly 40 years of existence.

The Hawaiian Volcano Observatory was es¬
tablished in 1912 as an outgrowth of prelimi¬
nary visits by Dr. T. A. Jaggar and Dr. R. A.
Daly. The bulk of the interest and support for
the work before 1912 came from the Massa¬
chusetts Institute of Technology, with some col¬
laboration by the Geophysical Laboratory of the
Carnegie Institution and others. Dr. Jaggar was
the moving spirit in establishing the Observa¬
tory in 1912, and was its director from 1912
until his retirement in 1940. In 1919 the Ob¬
servatory was transferred from the Massachu¬
setts Institute of Technology to the U. S.
Weather Bureau, and thence in 1924 to the
U. S. Geological Survey. In 1935 it was again
transferred, this time to the National Park Serv¬
ice, and thus it came under the administrative
jurisdiction of the Superintendent of the Hawaii
National Park, which had been created in 1916.
During his earlier visits to Hawaii, Dr. Jaggar
had enlisted the interest of various residents of
the Territory, and in 1911 the Hawaiian Vol¬
cano Research Association was organized. The
preliminary studies and the regular work of the
Observatory have had substantial, at times vital,
support through subscription by various mem¬
bers of this local organization. Many of the

special projects of the Observatory, as well as
fellowships, special or part-time assistance, and
expenditures for housing and other equipment
have been made possible through the continued
interest and generosity of the Research Associa¬
tion. From time to time, the University of Ha¬
waii has supported volcanic studies at Kilauea,
and for a number of years it has maintained two
buildings near the present Observatory as office
and laboratory quarters. Since 1940, Dr. Jaggar
has been Research Associate in Volcanology on
the University staff and has divided his time
between Honolulu and Kilauea.

The office and other facilities of the Ob¬
servatory occupied a building located on the
northeast rim of Kilauea Crater from 1912 until
1940. There were supplementary buildings
near-by and over 20 years ago the Research As¬
sociation provided a fireproof building to house
important accumulated records. In the early part
of 1940, the Volcano House, hotel for visitors
to the volcano and the Park, was destroyed by
fire. Plans for a new hotel, developed by Park
officials, called for its contsruction on ground
overlapping the existing Observatory, and for a
new observatory and naturalist building not far
away. The construction of the latter building
and the moving of the equipment had not been
completed when the Pacific war broke out. Be¬
cause of its disordered state of incompleted
moving, the immediate occupation of the partly
completed Observatory building by various units
of the armed forces, and the curtailment of nor¬
mal Park activities in favor of war work, the
Observatory had truly a difficult time during
much of the war period.

Ruy H. Finch became volcanologist and di¬
rector in 1940 after many years of earlier work
at Kilauea and in related volcano work in Cali¬
fornia. With only meager assistance he was
able to continue the most essential seismometric
and other observations and in some measure to
protect the equipment and records against a
variety of natural enemies and the hazards of
war-time confusion. From the necessarily low
level of operation maintained during the war,
the return to an effective program of study has
been partly accomplished. It is expected that the
present transfer to the Geological Survey, with
its growing program in the Pacific area and its
fundamental interest in the study of volcanoes,
will result in a period of renewed growth and
accomplishment by the Hawaiian Volcano Ob¬
servatory.-^ — Chester K. Wentworth.

[131]

Interbreeding of Laysan and Black-footed Albatrosses

Rothschild (The avifauna of Laysan and the
neighboring islands, p. 292, 1893) recorded a
single specimen of albatross which he regarded
as a cross between Diomedea nigripes and D.
immutabilis. In May, 1945, on Midway Island,
I noted a nigripes and an immutabilis feeding
the same partly fledged young. Because of this,
I made a particular search for further evidence
of interbreeding when I again visited Midway
in December, 1946.

On Eastern Island of Midway on December
30, 1946, an apparent hybrid was observed for
40 minutes, and kodachrome pictures were made.
This bird was nigripes in color except for white
underparts on the body extending forward to
the anterior part of the breast. Its behavior,
however, was like that of immutabilis; the head
was carried high in walking — not extended for¬
ward and low as is customary for nigripes. It was
standing in a mixed group on a sand dune near
the beach.

In two different places on the beach of Sand
Island, Midway, a nigripes and an immutabilis

The Poisoning of Bnfo marinus by

On May 9, 1947, a paralyzed toad, Bufo
marinus, was picked up beneath a large
strychnine tree, Strychnos nux-vomica, growing
on the grounds of the Territorial Board of
Agriculture and Forestry. This toad would be¬
come convulsed, with the fore and hind legs
stiffly extended in a muscular spasm, whenever
it attempted to move or was touched. Any
stimulus above or along the spine was especially
effective in causing such a spasm.

The animal was anesthetized by an injection
of sodium pentobarbital and the stomach dis¬
sected out. On opening the stomach, the follow¬
ing material was found: one snail, two ants, two
cockroaches, one small beetle, one small uniden¬
tified leaf, and ten flowers of Strychnos nux-
vomica.

were present at a single nest isolated from the
rest of a nigripes colony. In both instances a
Laysan Albatross was incubating an egg, and
the Black-footed Albatross was resting along¬
side. A subsequent visit to one of the nests
revealed a nigripes incubating the egg and an
immutabilis standing near-by.

Still another mixed pair was observed, on the
beach on December 31, rubbing the bills together
and stroking the feathers of the neck. Occa¬
sionally one or the other would execute some
movements of the dance typical of its species,
but the couple never did perform at the same
time. No nest was present here.

It seems likely that interbreeding of these
species is more frequent than is usually believed.
This may not be surprising in view of the sim¬
ilarity in size, structure, and habits, and the over¬
lap of nesting colonies on crowded islands.—
Harvey 1. Fisher, Department of Zoology and
Entomology , University of Hawaii, Honolulu,
Hawaii.

the Flowers of the Strychnine Tree

It would appear probable, judging both from
the symptoms and the presence of Strychnos
nux-vomica flowers in the stomach, that the toad
was suffering from strychnine poisoning. Arnold
reports the fatal poisoning of Bufo marinus in
this fashion and speculates on the reason for
the ingestion of the Strychnos flowers by the
toad (Arnold, Harry L., Poisonous plants of
Hawaii, 71 p., 24 pi. Tongg, Honolulu, 1944).
It is possible that such ingestion by the toad
reported here was adventitious while feeding
upon insects, since the area beneath the strych¬
nine tree was thickly carpeted with blossoms
fallen from it. — Vernon E. Brock, Director, Divi¬
sion of Fish and Game, Territorial Board of
Agriculture and Forestry, Honolulu, Hawaii.

[132}

JULY, 1948

NO. 3

a y o » j

VOL. II

E4.CIFIC

SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

IN THIS ISSUE: Hiatt — Biology of Pachygrapsus
crassipes • Wagner — A New Fern from Rota •
Bridge — Occurrence of Schist on Truk, Eastern
Caroline Islands • NOTES • News Notes

Published by

THE UNIVERSITY OF HAWAII
HONOLULU, HAWAII

BOARD OF EDITORS

Leonard D. Tuthill, Editor-in-Chief
Department of Zoology and Entomology, University of Hawaii

O. A. Bushnell, Assistant Editor
Department of Bacteriology, University of Hawaii

Ervin H. Bramhall

Department of Physics, University of Hawaii

Vernon E. Brock

Division of Fish and Game, Territorial Board of
Agriculture and Forestry
P. O. Box 3319, Honolulu 1, Hawaii

Harry F. Clements

Plant Physiologist, University of Hawaii Agricultural
Experiment Station

Charles H. Edmondson

Zoologist, Bishop Museum, Honolulu 35, Hawaii
Harvey I. Fisher

Department of Zoology, University of Hawaii

Frederick G. Holdaway

Entomologist, University of Hawaii Agricultural
Experiment Station

Maurice B. Linford

Plant Pathologist, Pineapple Research Institute
P. O. Box 3166, Honolulu 2, Hawaii

A. J. Mangelsdorf

Geneticist, Experiment Station, Hawaiian Sugar
Planters’ Association
P. O. Box 2450, Honolulu 4, Hawaii

G. F. Papenfuss

Department of Botany, University of California
Berkeley 4, California

Harold St. John

Department of Botany, University of Hawaii

Chester K. Wentworth

Geologist, Honolulu Board of Water Supply
P. O. Box 3410, Honolulu 1, Hawaii

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[ Continued on inside back cover ]

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

VOL. II JULY, 1948 No. 3

Previous issue published April 1, 1948

CONTENTS

PAGE

The Biology of the Lined Shore Crab, Pachygrapsus crassipes Randall.

Robert W. Hiatt . 135

A New Bern from Rota, Mariana Islands. W. H. Wagner, Jr. . 214

A Restudy of the Reported Occurrence of Schist on Truk, Eastern Caroline

Islands. Josiah Bridge . 216

Notes:

On the Occurrence of Bauxite on Truk. Josiah Bridge . 223

Holes in the Webs of Shearwaters. Frank Richardson . 224

Seventh Pacific Science Congress. Gilbert Archey . 225

Further Information Concerning the Fulbright Act . . . 226

News Notes . 228

Pacific Science, a quarterly publication of the University of Hawaii, appears in January,
April, July, and October. Subscription price is three dollars a year; single copies are one
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of major articles are available and requests for these will be filled in so far as possible.

Pachygrapsus crassipes Randall. (Dorsal view of female, X 2.5.)

The Biology of the Lined Shore Crab,
Pachygrapsus eras sipes Randall

Robert W. Hiatt1

INTRODUCTION

The conspicuous lack of knowledge of the
natural history of littoral Brachyura (true crabs)
of the west coast of North America has for a
long time indicated the necessity for an investi¬
gation into the biology of a common and well-
known decapod, such as the subject of this
study, Pachygrapsus eras sipes Randall (Frontis¬
piece). This investigation has been designed
to fulfill this need in part and has been pursued
primarily from the ecologic and behavioristic
points of view. A considerable portion of this
paper has been devoted to the description of
the molt and intermolt cycle of P. crassipes.
Some recent data which have appeared in the
literature have dealt with the extremely sig¬
nificant molt and intermolt cycles of the can¬
croid and oxyrhynchoid crabs, but the grapsoid
group, to which P. crassipes belongs, has been
neglected. Therefore, the present study, which
includes comparable work on this grapsoid crab,
materially extends the knowledge of these sub¬
jects within the group Brachyura. Included here,
also, is a brief discussion treating the problem
of the transition in the Brachyura from a
marine habitat to a semi-terrestrial or amphib¬
ious existence. Among littoral Brachyura on
the northern California coast, P. crassipes ranges
highest along the shore and exhibits certain
patterns both physiologically and behavioris-
tically illustrative of this significant landward
movement.

A primary aim has involved recording of
field observations which have logically pre¬

1 Department of Zoology and Entomology, Univer¬
sity of Hawaii. Manuscript received October 1, 1947.

ceded laboratory investigations. Such field ob¬
servations have provided the natural basis for
the present laboratory studies, making the latter
far more interesting and significant. On the
other hand, clues to certain behavior patterns
were exhibited among the captive crabs prior
to their observation in the natural environment.
In addition, the correlation of laboratory data
with field observations has helped to illuminate
certain behavior patterns, not only of P. cras¬
sipes , but of Brachyura in general, which have
heretofore been incompletely known. It has
not been the intention of the author to study
exhaustively the multitude of morphological
and behavioristic phenomena associated with
this species. Rather, an attempt was made to
become better acquainted with the behavior
of the crab in question, with the underlying
phenomena involved, and thus with the Brachy¬
ura in general.

Most of the knowledge of the natural history
of decapod crustaceans stems from a series of
studies upon those with economic value,
although for diverse reasons these studies have
subordinated natural history data in favor of
securing information on life histories. The early
literature is filled with abbreviated note-type
accounts, but the more extensive investigations
of H. C. Williamson on Cancer pagurus Linne,
reported at the turn of the century (1899),
were the first to provide substantial knowledge
concerning any one decapod crustacean. Shortly
thereafter Hay (1905) published "The Life
History of the Blue Crab,” in which he con¬
tributed much to the natural history of this
species. Pearson (1908), in addition to his
own investigations on C. pagurus, included

136

PACIFIC SCIENCE, Vol. II, July, 1948

Williamson’s earlier findings, and in so doing
provided the first comprehensive study of a
crab. In 1909 Herrick summarized many years
of research in a classic monograph on the
American lobster, Homarus americanus, which
for the past 35 years has stood unparalleled in
the field. In 1915 the natural history and be¬
havior of the fiddler crab were presented by
Schwartz and Safir, and in 1918 Churchill
amalgamated all the information on the life
and natural history of the blue crab, Callinectes
sapidus Rathbun, into a single authoritative
report. In 1930 MacGinitie published an account
of the natural history of the mud shrimp,
Upogebia pugettensis (Dana), which was fol¬
lowed by a similar paper (1934) on Callianassa
calif orniensis Dana. These investigations pro¬
vided the first consequential studies of any west
coast decapod crustacean which was unimpor¬
tant economically. A series of studies through¬
out the past quarter century on the life history,
growth, and migration of the Pacific edible crab,
Cancer magister Dana, by Weymouth (1917),
by the same author in collaboration with Mac-
Kay (1934, 1935, 1936), and by the latter
author (1942), has furnished considerable sig¬
nificant information. More recently, Broekhuy-
sen (1936) published an account of the devel¬
opment, growth, and distribution of a com¬
mon European shore crab, Carcinides maenas
(Linne). This was followed in 1941 by his
similar investigation of the South African shore
crab, Cyclograpsus punctatus M. Edwards. Other
literature of a more specific nature will be dis¬
cussed where pertinent in the text of this report.

In the voluminous literature dealing with the
decapod Crustacea, only a few brief notes on
the habits of the crabs of the genus Pachygrapsus
can be found, and these furnish extremely few
data on P. crassipes.

Acknowledgments: The writer is especially in¬
debted to the late Dr. S. F. Light, of the Depart¬
ment of Zoology at the University of California,
whose valuable assistance and helpful criticisms
were most useful in the pursuance of this study.
Special credit is also due to Dr. L. E. Noland,

of the Department of Zoology at the University
of Wisconsin, for many suggestions. I should
like to express my gratitude to Dr. E. Yale
Dawson, who provided identification of the algal
species mentioned in this paper. The tempera¬
ture data of surface water in San Francisco Bay
used in this study were furnished the writer
through the courtesy of Captain C. D. Wash¬
burn, Jr., of the Branch Hydrographic Office,
U. S. Navy, San Francisco, to whom I am indeed
grateful. Without the material aid extended by
the Department of Zoology at the University
of California, this study would have been im¬
possible, and much credit is due to the Univer¬
sity of Hawaii for extending many facilities
necessary to bring this research to a successful
conclusion.

To my wife, Elizabeth, I owe an immeasurable
amount of gratitude for her constant encourage¬
ment and generous assistance both in the field
and in the laboratory. Dr. D. C. Matthews, of
the Department of Zoology at the University
of Hawaii, and Mrs. Matthews gave generously
of their time to assist in critically reviewing the
manuscript.

TAXONOMY

Synonymy

Pachygrapsus crassipes Randall. 1839. Acad.
Nat. Sci., Phila., Jour. 8: 127 (type
locality, Sandwich Islands; this locality is
probably erroneous; type in Mus. Phila.
Acad. Nat. Sci.); de Man. 1890. Notes
Leyden Mus., 12: 86, pi. 5, fig. 11; Rath-
bun. 1917. U. S. Natl. Mus., Bui. 97: 241,
pi. 59.

Grapsus eydouxi Milne-Edwards. 1853. Ann.
Sci. Nat., ser. 3, Zool., 20: 170 (type
locality, Chile; this also is probably an
erroneous locality record; type in Paris
Mus. ) .

Leptograpsus gonagrus Milne-Edwards. 1853.
Ann. Sci. Nat., ser. 3, Zool., 20: 173 (type
locality unknown; type in Paris Mus.).

Biology of Pachygrapsus crassipes — HlATT

137

Stimpson ( 1857), Kingsley (1880), de Man
(1890), Rathbun (1917), and Boone (1927)
concur in the opinion that Leptograpsus gona-
grus H. Milne-Edwards, for which the type
locality is unknown, is identical with P. crassipes
Randall. The meager description of L . gonagrus
by Milne-Edwards (1853) agrees with the spe¬
cific characters of P. crassipes. Furthermore,
Milne-Edwards substituted the name Leptograp¬
sus for Pachygrapsus. Stimpson (op. cit.) ad¬
vanced several reasons for retaining the name
Pachygrapsus for the group designated Lepto¬
grapsus by Milne-Edwards.

Kingsley (1880), Rathbun (1917), and
Boone (1927) believe that Grapsus eydouxi
Milne-Edwards is likewise synonymous with P.
crassipes Randall. Here again it is obvious that
the description of G. eydouxi as set forth by
Milne-Edwards (1853) agrees with that of P.
crassipes. It seems rather incongruous, however,
that he should place two apparently similar
specimens in different genera in the same re¬
vision. The type locality for G. eydouxi is cited
as Chile. If this is correct, G. eydouxi is not
synonymous with P. crassipes because the latter
crab does not occur on the Latin American
coast (see p. 138).

Systematic Position

P. crassipes is one of eleven species of a genus
distributed in a nearly cosmopolitan manner.
Its members inhabit both the Pacific and
Atlantic coasts of America, the western coast
of Africa, coasts bordering the Mediterranean,
and the Indo-Pacific, and Central Pacific shores.
Four genera of the family Grapsidae have been
reported from the California coast; they may be
readily distinguished from each other (see key
in Schmitt, 1921: 269). The seven species of
Pachygrapsus recorded from North and South
America may be identified by the use of the key
in Rathbun (1917: 241).

Description

Although a moderately complete, generalized,
morphological description of P. crassipes is

given by Rathbun (1917: 242), no information
is made available by this or other authors with
regard to sexual dimorphism. Measurements
of the length and width of the carapace of sev¬
eral hundred crabs show that the males average
larger than the females. The mean width of
the carapace of adult males which ranged in
size from 16.0 to 45.8 millimeters was found
to be 36.8, with a standard deviation of 11.2.
Similar measurements on adult females which
ranged from 15.0 to 39.7 millimeters showed
the mean width to be 30.9, with a standard
deviation of 4.7. The mean length of the cara¬
pace of adult males was found to be 31.6 milli¬
meters, with a standard deviation of 9.6. Similar
measurements of females showed the mean
length to be 26.4 millimeters, with a standard
deviation of 3.9. These differences in size be¬
tween the sexes do not become apparent until
the crabs reach a carapace width of 22 milli¬
meters (see Fig. 10), after which the carapace
of female crabs becomes relatively narrower and
shorter than that of male crabs of the same age
and belonging to the same intermolt period.
The usual brachyuran sexual dimorphism in
the abdomen is apparent. Sexual differences in
the chelipeds are pronounced, those of the male
being approximately 8 per cent longer. The
propodite and dactylopodite of this appendage
exhibit the greatest difference; those of the
males exceed those of the females by approxi¬
mately 10 per cent.

Type Locality

The assertion by Randall (1839) that the
type locality of P. crassipes is the Sandwich
Islands (Hawaiian Islands), a region outside
the normal geographic range of this species
(Fig. 1), has made obligatory an investigation
of this anomaly. Randall states that the type
specimen of P. crassipes was collected on the
shores of the Hawaiian Islands by Nuttall. How¬
ever, frequent and thorough searches for decapod
Crustacea in the Hawaiian Islands during the
past century, in addition to recent explorations
by myself, have failed to disclose this species.

138

PACIFIC SCIENCE, Vol. II, July, 1948

Further, Randall states that the type specimen
of Meto grapsus mess or (Forskal) ( = Pacby -
grapsus parallelus Randall) was found on the
coast of Oregon by Nuttall during the same
voyage. It, in turn, has not been known to
occur on the American coast, but it is common
on the Hawaiian Islands. Inasmuch as the names
are correctly affixed on the type specimens,
Stimpson (1857) made a rather likely sugges¬
tion; namely, that the labels giving the collec¬
tion localities of the two species in Nuttall’s
collection were in some way accidentally ex¬
changed. Holmes (1900) also points out that
several other species reported from the Pacific
coast by Randall have not since come to light.

DISTRIBUTION

Geographical Distribution

The authenticated geographical range of P.
crassipes extends along the western coast of
North America from north latitude 45° (New¬
port, Oregon) to north latitude 24°20' (Santa
Margarita Island, Baja California), and along
the coast of Japan and Korea from north lati¬
tude 34° to 37°.

A number of records outside the above local¬
ities had to be considered in an attempt to
establish the authentic geographical range. For
example, Rathbun (1917) cites Chile as a
collection locality. Correspondence with Dr.
C. E. Porter, an active carcinologist in Chile,
has established the fact that P. crassipes does
not exist along Latin American shores. There¬
fore, either Grapsus eydouxi (see Synonymy,
p. 136) is not synonymous with P. crassipes or
the type locality of the former is incorrect.

Rathbun (1902) also cites the Galapagos
Islands as another locality of collection. The
specimen upon which the record is based was
taken at 12 fathoms, a depth in decided con¬
trast to the well-known habits of the species
elsewhere. This, together with evidence sup¬
plied by Garth (1946), indicates that P. cras¬
sipes is not a member of the Galapagan fauna
and that the specimen catalogued from there,

which I have seen and know to be P. crassipes,
was in some manner mixed with specimens col¬
lected from the west coast of America on the
same expedition.

P. crassipes is associated geographically with
but one other congener, P. transversus (Gibbes),
and with it only toward the southern boundary
of its range on the east and west coasts of Baja
California and the west coast of Mexico. Here¬
tofore, the two species have been thought to
overlap in the Galapagos and Cocos Islands.
A great difference in size enables one to dis¬
tinguish quickly between these two forms. The
length of the carapace of a large specimen of
P. crassipes is approximately 40 millimeters, the
width approximately 44 millimeters. P. trans¬
versus, on the other hand, is only half as large;
the length of a large specimen is about 15 milli¬
meters, the width about 20 millimeters. The
most striking diagnostic character is the width
of the front, which in P. crassipes is half or
almost half as wide as the carapace; whereas in
P. transversus the front is distinctly more than
half as wide as the carapace.

The geographic distribution of P. crassipes
is completely disjunctive. The individuals on
the coast of Japan are separated from those on
the west coast of America by the wide expanse
of the Pacific Ocean (Fig. 1). To account for
this distributional phenomenon, it is necessary
to assume that the migration between the two
continents was somehow accomplished without
leaving tangible evidence of the process. The
physical characteristics of its habitat (p. 142)
offer little assistance to the solution of the dis¬
tributional pattern, largely because this species
possesses a wide substrate tolerance and is some¬
what eurythermal. Moreover, although rocky
shore line is common along the coast of Korea
and Japan, the animals there are restricted to
a narrow band of latitude 2.5° to 3° in extent.

The isothermic pattern for surface water and
air temperature at sea level during both summer
and winter in Japan and America (see Schott,
1935) fits a wide latitudinal spread on the
western coast of North America but is strongly

Biology of Pachygrapsus crassipes — HIATT

139

135® 150* 165* ISO* 185® 150* 135® 120® 105® 00®

convergent in the region of Japan. This isother¬
mic pattern seems to offer a distinct clue as to
why the latitudinal distribution of this species
is restricted in the Orient. However, more crit¬
ical examination shows that the tolerable tem¬
perature range for this species as exhibited on
the Pacific coast of America is almost nowhere
fully occupied in the Oriental region.

That this species could, but does not, possess
a greater latitudinal distribution in the Orient
indicates that either ( 1 ) control of its distribu¬
tion is accomplished by factors (food supply,
parasites, etc) which have not been taken into
consideration because of insufficient knowledge
of the Oriental region, or (2) the presence of
this species in the Orient has been of relatively
recent occurrence. The latter hypothesis seems
unwarranted, however, for reasons cited below.

Extremely rapid invasions of new areas are
known to have been accomplished by this and
other species of Crustacea. Stebbing (1888,
1893, 1906), Alcock (1899, 1900), Fulton
and Grant (1901), Chilton (1911), and Pan¬
ning (1939) have recorded numerous observa¬

tions of Crustacea which, by means of trans¬
oceanic shipping, have attained new frontiers.
Some of these transportations have been suc¬
cessful, e.g., Eriocheir japonicus as described by
Panning (1939). It is certain that at present,
and for a long time past, the dispersal of P.
crassipes over its geographical range could not
have occurred by a littoral migration along the
north Pacific shores, presumably because of the
unfavorable temperatures. The low winter tem¬
perature along the Oregon coast probably de¬
limits the northern extent of this species; other
ecological requirements seem satisfactory farther
northward. If a littoral migration of this type
had taken place when the climate was temperate
along the north Pacific shores, one would expect
to find some manifestation of morphological
change accompanying the long isolation of the
populations. However, the American and Jap¬
anese types are apparently identical.

Although the currents in the Pacific are
favorable for the transportation of pelagic fauna
between American and Japanese coasts, it is
highly improbable that this species was dis-

140

PACIFIC SCIENCE, Vol. II, July, 1948

persed in this manner. The first crab stage,
which is no longer free-swimming, occurs ap¬
proximately 5 weeks after hatching. The time,
therefore, is far too short for a successful ocean
crossing by larvae. The possibility of the use
of natural rafts as vehicles cannot be dismissed
lightly in view of the habits and habitat of this
form. However, both these hypotheses seem
less plausible than the suggestion to follow.

Since P. crassipes was not mentioned in the
faunal accounts of Japan until 1890, it is not
unreasonable to speculate on the probability
that marine traffic between the west coast of
America and Japan has supplied the medium
of dispersal. The extensive reproductive season
of this species would insure the presence of a
quantity of zoeae swarming in the harbors.
These zoeae may easily have been taken into
the ballast tanks of ships which were filled on
the California coast and subsequently emptied in
Japanese waters. Inasmuch as the ecological
requirements of the niche are fulfilled in Japan,
it seems feasible that the species would become
established; in fact, a constant intermixing of
this nature may explain the uniformity of the
species in the two widely separated areas.

Habitat and Ecologic Niche

P. crassipes is a eurytopic species which
ranges through the depositing shore and eroding
shore subbiochores; consequently, it is neces¬
sary to designate the characteristics of the habitat
in general, and to subdivide this general habitat
into the three biotopes and fasciations which
the writer considers most significant. Generally
speaking, the habitat of this crab is that region
of the strand extending from upper low (0.0
tide level) to highest high intertidal zone (6.0-
foot tide level in the San Francisco Bay region )
where there is a hard substrate containing
crannies, crevices, or holes, free from loose
stones, sand, or mud, and supporting a more
or less luxuriant growth of ulvaceous or fila¬
mentous algae.

With respect to the vertical distribution on
the strand, the present data show that this

species may be located in crevices in the adtidal
zone (at Pacific Grove, California) as low as
the - 0.7-foot tide level. However, the indi¬
viduals at this level were secluded in refuges
at the base of nearly vertical rocky promontories
which extend several feet out of the water at
high tide. It is assumed that they had followed
the receding water level, since the crevices they
usually occupy are located in the intertidal zone
higher up the rock. Such vertical migrations
of a few feet are common, in contrast to a
lack of extensive horizontal movements. Where
no rocks protrude at high tide, the crabs are
absent from crevices at the - 0.7-foot tide level,
thus furnishing evidence for the preceding as¬
sumption. The greatest proportion of crabs are
to be found in crevices and tide pools at the
lower edge of the supratidal zone. Relatively
fewer are found higher in the supratidal zone
(from the +5.0- to the +8.0-foot tide level at
Pacific Grove ) and the area slightly above,
except at night. Additional information on
this phase of distribution is presented in the
section on foods and feeding habits.

The first and most important biotope is char¬
acterized by large rock promontories. These
rocks are generally continuous and unbroken
by tide channels from points high on the shore
to the low-tide level. Their multiplicity of
crevices affords the crabs refuge from danger
during the periods of the day in which activity
is minimal. The greatest concentration of crabs
is found in the crevices of these rocky coasts.
Tide pools of varying depths are frequently
encountered, all of which provide activity cen¬
ters for the crabs. Much of the coast line in the
region of central California is of this type;
Plate 1, Figure 1, illustrates a typical rocky
habitat at Pacific Grove, California.

Microscopic algal forms associated with this
biotope at Pacific Grove consist in the main of
the following types: around the edges of the
tide pools and in damp crevices are found
juvenile sporelings of some ulvaceous alga, sev¬
eral species of non-colonial diatoms, an abun¬
dant supply of Oscillatoria sp., a sparse supply

Biology of Pachygrapsus crassipes — HIATT

141

of Lyngbya (two species), and a sparse supply
of Cladophora sp. It is the foregoing group of
plants which form the greater part of the food
consumed by P. crassipes in central California.

In addition to the minute algae the following
macroscopic forms are significant in the life of
this crab:

IJlva lactuca L. : This species is sparsely represented
•in the first biotope on the tops and sides of the
rocks just below high-tide level; it is of less im¬
portance here than in the second biotope to be
described.

Vucus furcatus A g. : An abundant species which sup¬
plies this animal both cover and food.

Enteromorpha sp.: A common species which pro¬
vides considerable food. Longer fronds furnish
effective cover for small crabs (many of the
small, recently molted individuals may be col¬
lected under this alga).

Grateloupia Setchelii Kylin and Rbodoglossum af¬
fine (Harv.) Kylin: Both species are common in
high tide pools and are consistently used by the
animals for food.

Endocladia muricata (Harv.) J. A g.: This species
grows on the rocks in and immediately below
crevices near high-tide level. It is a relatively
important source of food.

The algal forms listed below are, as a group,
less important than the above types. Algal fronds
broken from plants lower in the strand or from
pelagic types may often be found floating in
the high tide-pools; these floating pieces are
often seized by the crabs and partially devoured;
thus, they are important integers in the life
equation of this animal.

Cryptopleura lobulifera (J. Ag.) Kylin: Found
floating in tide pools at high-tide level.

Laminaria Sinclarii (Harv.) Farlow: Found floating
in tide pools at high-tide level.

Macrocystis pyrifera (L.) C. A. Ag.: Found float¬
ing in tide pools at high-tide level.

Pelvetiopsis limitata (Setchell) Gardner: On rocks
and in tide pools just below high-tide level.

Gelidium sp.: Growing in rock crevices at high-
tide level.

Gigartina leptorhynchos J. Ag.: Growing in tide
pools at high-tide level.

Iridophycus sp. : Growing on rocks just below high-
tide level.

Egregia Menziesii (Turn.) Aresch.: Growing on
rocks just below high-tide level.

Cladophora gr amine a Collins: Growing in crevices
containing sand just below high-tide level.

Gastroclonium Coulteri (Harv.) Kylin: Growing
on rocks just below high-tide level.

Agardhiella Coulteri (Harv.) Setchell : Growing on
rocks just below high-tide level.

Corallina officinalis L.: Growing in tide pools just
below high-tide level.

Gigartina Harvey ana (Kuetz.) Setchell and Gard¬
ner: Growing on rocks in tide channels just above
low-tide level.

Gigartina corymbif era (Kuetz.) J. Ag.: Growing on
rocks in tide channels just below low-tide level.

Cryptopleura violacea (J. Ag.) Kylin: Growing on
rocks in tide channels just above low-tide level.

The second and less common biotope is found
along the outer coast, as well as along man¬
made shore lines which are met most commonly
in estuaries and bays. This type is characterized
by the presence of large, relatively stable bould¬
ers, which occupy an area of the strand coincid¬
ing with the intertidal range of this species.
On the outer coast these boulders generally rest
upon a solid substrate of smooth rock or upon
other boulders. Along estuaries, e.g., the Oak¬
land Estuary in the San Francisco Bay, granite
boulders have been arranged two or three deep
along the shore, extending from low- to high-
tide level; very few project sufficiently high
to be in a position comparable to the splash
zone of the outer coast. At the upper limit of
these boulders along the estuarine strand the
terrain becomes flat, and the boulders gradually
decrease in number and size. The crevices
requisite for the existence of this species are
formed by the contiguous surfaces of the
boulders. The crab population in this habitat
shows a progressive decrease from the large
boulders placed low on the strand to the smaller
ones farther shoreward. This gradient in popu¬
lation may be attributed to the fact that the
boulders extending shoreward are smaller,
stacked in a more shallow layer, and tend to be
more deeply imbedded in mud or sand; such
conditions manifestly reduce the number of
available refuges. In these areas where environ¬
mental conditions for P. crassipes are met along
a rocky embankment, shoreward of which occurs
a narrow, relatively muddy tidal flat, the usual
linear order of grapsoid habitation on the strand
becomes inverted. In this terrain one frequently
encounters both Hemigrapsus nudus Dana and
H. oregonensis Dana farther shoreward than P.
crassipes; whereas in the first biotope both these
species frequent comparatively lower fasciations
in the littoral zone than the subject of this study.

142

PACIFIC SCIENCE, Vol. II, July, 1948

This invasion may be attributed jointly to the
decreased number of suitable refuges for P.
crassipes and the muddy nature of the substrate
in the high, flat zone which is suitable habitat,
especially for H. oregonensis.

The algal species contributing most to the
general welfare of P. crassipes in this estuarine
habitat is Ulva lactuca, whose short fronds or
very young sporelings form the principal food
for this crab. Ulva is abundant on the large
boulders from middle high- to high-tide level;
the plants become progressively smaller as the
high-tide level is approached.

The third and least common biotope in which
P. crassipes is found is the muddy shore of bays
and estuaries. Along these, restricted to high-
tide level, are numerous individuals which take
refuge in small holes in the mud bank. Extended
observations have furnished evidence that the
holes are not dug by the crabs, but are excavated
by wave action which has removed the loose
mud and sand from the netted roots of a dense
stand of pickleweed {Salicornia ambigua
Michx. ) which grows shoreward from high-tide
level. The splash zone, here an area extending
back on a slight incline from the bank, is cov¬
ered by a dense growth of Ulva lactuca which
grows on the hard mud between the stems of
Salicornia. This growth of Ulva, the lone macro¬
scopic algal species present, covers a flattened
area extending approximately 5 feet back from
the top of a bank on a level with the high-tide
mark. The habitat described above is typical
of portions of the shores of Bolinas Bay, Bodega
Lagoon, and Elkhorn Slough (PI. 1, Fig. 2).

The vertical section of the bank varies from
6 to 18 inches in height; the crest of the bank
corresponds to the "upper high” intertidal zone,
and the base is on a level with the "middle high”
zone. Consequently, the vertical range of the
crab habitat is restricted when compared with
other types of terrain outlined above. From the
base of the bank the intertidal zone consists of
a down-sloping stretch of mud containing gravel
and sand. At the ebb of a - 0.5 -foot tide this
slope averages approximately 25 feet in breadth,

and a medium high tide completely submerges
it. The receding tide serves to deposit on this
slope a myriad of organisms killed by the exten¬
sive clam digging operations which occur at
many points along these bays and estuaries. A
wealth of animal food in addition to the abun¬
dant algal supply is thereby provided. P. cras¬
sipes has never been observed to venture more
than 4 to 5 feet away from the refuge holes in
the banks; hence it offers little, if any competi¬
tion to H. oregonensis, which likewise occupies
holes in the bank, but characteristically forages
over the entire expanse of the submarine slope.
The latter species accepts any available organic
matter and seems to forage typically as a true
scavenger.

P. crassipes may well be considered a eury-
topic species because of its wide distribution
along the entire width of the strand. The popu¬
lation is greatest at the typical high fasciation,
and follows a decreasing gradient in abundance
farther landward as well as farther seaward. In
the first biotope mentioned, the habitat condi¬
tions consist of those deep crevices in or near
tide pools at the high-tide level, with an abun¬
dant supply of filamentous or ulvaceous algae
at hand. The habitat characteristic of the second
biotope may be designated as those crevices
under or between large boulders which ordin¬
arily have a dense growth of Ulva or other
minute forms of algae on their exposed surfaces.
The topography characteristic of the third bio¬
tope consists of shallow holes in hard mud
banks, immediately above which is an abundant
supply of ulvaceous algae.

The primary topographic units described
above apparently provide all the factors requi¬
site for the welfare of the species: a hard sub¬
stratum (rock or solid, packed mud), ample
food which consists primarily of the more
minute species of algae and secondarily of dead
animal tissues, protection from wave action and
predators by seclusion in crevices, and insurance
against desiccation by close association with tide
pools or damp crevices into which the sun’s
rays do not penetrate.

Biology of Pachygrapsus crassipes — HlATT

143

The trio of closely associated grapsoid crabs
along the central California coast (P. crassipes ,
H. nudus, and H. oregonensis) exhibit distinct
preferences for certain types of substrate, not¬
withstanding the fact that the habitats preferred
overlap to a certain minor degree. In order to
ascertain more specifically the underlying causes
for this habitat preference, an analysis of the
preferred physical features of the environment
of each species was made, after which certain
pertinent structural, physiological, and behavior¬
istic comparisons among the crabs were made.
It was found that H. oregonensis prefers a
muddy, silty substrate; H. nudus is found more
commonly on a sandy or gravelly bottom; and,
as indicated above, P. crassipes shows a prefer¬
ence for the hard, non-silty substrate. A clue to
this distributive pattern may possibly be found
in a study of the structures concerned with ex¬
ternal respiration, inasmuch as it is necessary
that a constant supply of clean water be car¬
ried through the branchial chamber. An exam¬
ination of the abundance and position of setae,
both in the vicinity of the incurrent openings
to the branchial chamber and on the mastigo-
branch of the third maxilliped, was made.

A thick mat of very fine setae on the branchi-
ostegite immediately above the coxae of the
pereiopods is easily observed in H. oregonensis.
The dorsal side of each coxa likewise bears a
thick tuft, which, with the setae of the branchi-
ostegite, forms a fine strainer covering the incur¬
rent openings to the branchial chamber. Func¬
tional evidence for this setal sieve is apparent
since a heavy deposit of silt covers the struc¬
ture. The mastigobranch of the third maxilliped
is exceedingly plumose, and serves to sweep the
gill surfaces free of silt which does filter through
the incurrent sieve.

The setae on the branchiostegite and pereio-
podal coxae of H. nudus, although comparatively
fewer in number, are far heavier than those of
the preceding species. They arise in clusters of
three and four on the branchiostegite and pro¬
ject ventrolaterally; those near the ventral border
of the branchiostegite extend out to enmesh

with those which originate on the base of the
coxae. The stiff nature of these setae seems an
adaptation for withholding large particles such
as sand grains. The setae are heaviest on the
ventral border of the branchiostegite above
coxae 3, 4, and 5; those setae farther cephalad
are less rigid, though more numerous. The
mastigobranch of the third maxilliped is heavily
plumose.

P. crassipes, on the other hand, exhibits fewer
and finer setae. The branchiostegite, with the
exception of the ventral border, is virtually de¬
void of setae. Those setae located on the ventral
border of the branchiostegite enmesh with the
relatively sparse, fine coxal setae to cover the
opening to the branchial chamber. The mastigo¬
branch of the third maxilliped has comparatively
fewer setae than the two species previously
described.

It is evident that these morphological dif¬
ferences in the rigidness and abundance of setae
participate in establishing the character of the
environment selected. The fine, dense mat of
setae covering the incurrent channels of the
branchial chamber in H. oregonensis is well
adapted for straining out particles of a silt-like
nature. Therefore, among other contributing
factors, this structural adaptation enables this
species to subsist on the soft, muddy banks of
estuaries where it is especially common, and a
habitat wherein the remaining two crabs under
discussion cannot become established. H. ore¬
gonensis is rarely found associated with P. cras¬
sipes (except in the third biotope mentioned)
probably because the former species cannot
withstand the extended periods of desiccation
which the latter type undergoes in its position
high in the intertidal zone. However, in those
estuaries wherein the fasciation of P. crassipes
is lower than the highest tide level, and where
sufficient cover is available to maintain rela¬
tively moist refuges, H. oregonensis does become
a competitor for places of refuge; but, because
of the diverse feeding habits the two species
show little interspecific competition for sub¬
sistence. Associations of this type are to a

144

PACIFIC SCIENCE, Vol. II, July, 1948

certain extent competitive in that the maximum
number of each species present is probably not
as high as it would be if only one species were
present. It seems to be essentially an ecological
compromise in which both species suffer quan¬
titatively in the struggle for subsistence — further
evidence of the great halobiotic pressure of the
strand.

The rigid, clumped setae of H. nudus seem
well adapted to withhold large grains of sand;
an examination of the setae substantiates this
assumption. It is apparent that the sparse but
heavy setae are ineffective in straining out sus¬
pended silt; consequently, if the animals were
placed on a muddy substratum respiratory dif¬
ficulties would undoubtedly occur. This species
and H. oregonensis are, therefore, only infre¬
quently found associated. Where a close associa¬
tion exists, the substratum will normally be
transitional between sand and silt. Both species
in such an association are doubtlessly near the
frontiers of their respective tolerances. H. nudus
is often found associated with P. crassipes where
the latter occupies a position relatively low in
the intertidal zone. However, at the high-tide
level where P. crassipes is most abundant, very
few individuals of H. nudus are found. Under¬
lying causes for the virtual absence of H. nudus
from areas with comparatively long exposures
to air are obscure. No significant difference
between the abilities of the two species to with¬
stand desiccation was found. However, a clue
to the inherent preferential differences with
respect to habitat may be gained from the
hydrotactic responses exhibited by each type.

P. crassipes is negatively hydrotactic; whereas,
H. nudus and H. oregonensis are positively
hydrotactic. This fact may readily be demon¬
strated by placing representatives of the three
species in an aquarium which is tilted to pro¬
vide deep water at one end and a dry area at
the opposite end. The crabs will move to their
preferred stations immediately; the latter two
species submerge in the deep end, and the
former species takes a position at the dry end.

This behavior pattern of P. crassipes was utilized
in the laboratory where the aquaria were tilted
to simulate tide-pool conditions — deep water
at one end and none at the other. The crabs
spent most of each day at the shallow or dry
end. Those reared in large tanks containing
projecting rocks were observed upon the rocks
the greater part of the time until they were
disturbed, in which case they would dart under¬
neath the rocks. This reaction to disturbance
is typical of their normal littoral life, which will
be discussed later. This negative hydrotactic
response of P. crassipes is a prominent factor
underlying its success, because this species alone,
among littoral crabs, wanders onto the rocks
high above the water to reach the uncontested
food supply found there.

Preliminary experiments designed to test dif¬
ferences in the ability of these three local shore
crabs to withstand desiccation were performed
as follows: Representatives of each species of
equivalent sizes in stage Q (see section on
Intermolt Cycle, p. 146) were deposited in
individual battery jars and placed in the same
room to insure comparable external environ¬
mental conditions. Observations were then made
at 15 -minute intervals from 4:00 A.M. to 10:00
P.M. The interval of time which elapsed be¬
tween the initiation of the experiment and the
subsequent death of each animal was recorded.

It was found that H. oregonensis possessed
much less ability to withstand desiccation than
either H. nudus or P. crassipes. All individuals
of H. oregonensis were moribund in 6 hours
and all representatives of this group succumbed
in 10 hours. None of the specimens of H.
nudus and P. crassipes was moribund until 13
hours had elapsed, most of them lasting for a
minimum of 18 hours. All were dead in 24
hours. No differences in tolerance to desicca¬
tion were apparent between P. crassipes and
H. nudus.

A second experiment along similar lines was
performed to test further the relative abilities
of P. crassipes and H. nudus to withstand desic¬
cation. This experiment was performed at the

Biology of Pachygrapsus crassipes — HlATT

145

TABLE 1

A Comparative Summary of the Number of Gills, Volume of Gills in Relation to Body
Volume, and Habitats of Selected Littoral Brachyurans on the Coast of Central California.

SPECIES

NUMBER OF
SPECIMENS
TESTED

NUMBER
OF GILLS

PERCENTAGE OF GILL
VOLUME TO BODY
VOLUME

HABITAT

Range

Mean

Cancer antennarius . .

4

18

4.9-5.1

5.0

Low-tide zone and below;
crevice dweller

Cancer magister . .

1

18

4.4

Below low-tide mark;
sandy substrate

Pugettia productus . . .

4

18

3. 1-4.0

372

On algae in tidal zone
and off shore

Hemigrapsus nudus .

3

16

27-2.8

2.77

Rockweed belt; sand
below rocks

Hemigrapsus oregonensis ....

2

16

23-2.5

2.4

Tidal zone; muddy shores

Pachygrapsus crassipes .

5

18

23-2.5

2.36

High-tide mark and
above; rock crevices

seashore where normal environmental condi¬
tions prevailed. Ten specimens of P. crassipes
and 12 specimens of H. nudus, all in stage C„
were collected at Pacific Grove and placed in
dry aquaria. The aquaria were concealed high
on the rocks under a large boulder which ex¬
cluded the direct rays of the sun. Observations
were made at 15 -minute intervals from 4:00
A.M. to 10:00 P.M.

The results of these tests were comparable
with those obtained in the laboratory, except
that death did not ensue as rapidly.

Under these seashore conditions, crabs of a
size similar to those utilized in the laboratory
lived approximately twice as long. A direct
relationship between the size of the crab and
the tolerance to desiccation was found; the
smaller crabs became inactive and died sooner
than the larger ones. Crabs of both species
having a carapace width of approximately 40
millimeters lived about 48 hours. One specimen
of H. nudus having a carapace width of 57 milli¬
meters lived 70 hours. This apparent relation¬
ship between tolerance to desiccation and size
seems to explain, in part at least, why smaller
crabs of both species are invariably found in
the most moist area of the habitat. These data
demonstrate that the littoral horizon selected by
each of the species concerned is not determined

by its ability to withstand long periods of desic¬
cation.

It is highly probable that some physical as
well as physiological changes would occur in
the respiratory mechanism of a crab which
remained exposed to air for relatively extended
periods. Further, this change should be cor¬
related with the amount of exposure endured;
therefore, a proportionate decrease in gill vol¬
ume and surface to reduce desiccation would
seem a reasonable supposition in view of the
fact that ability to resist desiccation is generally
dependent upon the surface area of external
membranes. Pearse (1929^) has shown that
as crabs become adjusted to a terrestrial life
there is a gradual reduction in the number and
volume of the gills. In order to ascertain the
status of the gill-volume-body-volume ratio of
P. crassipes in contrast to other Brachyura, a
series of crabs was collected from habitats
ranging from below low-tide level to above
high-tide level.

The proportionate volume of the gills com¬
pared to the volume of the body was secured
in the following manner: The crabs were
hardened 24 hours in 1 per cent chromic acid
and subsequently rinsed with tap water; pereio-
pods were removed at the fracture plane; the
carapace was removed, and the gills were cut

146

PACIFIC SCIENCE, Vol. II, July, 1948

free from their attachments; all parts were
placed on blotting paper and allowed to drain
for 5 minutes; the gills were then dropped into
one graduate partially filled with water, and
the remaining body parts into another graduate
similarly filled; the volume was recorded as
the difference in the column of water. The
results are summarized in Table 1.

Unfortunately, crabs more highly adapted to
a semi-terrestrial existence than P. crassipes were
not available for this study; consequently, infor¬
mation was secured from statements concerning
gill reduction in the near-terrestrial types made
by Pearse (1929 a). He has shown that some
fiddler crabs ( Uca minax, U. pugnax, and U.
pugilator) have but 12 gills. Therefore, with
the exception of P. crassipes, it may be assumed
that in general crabs adapted to lengthy expo¬
sures to the air will have the gill number re¬
duced. It follows that a reduction in the num¬
ber and volume of gills is undoubtedly a critical
factor in the selection of a habitat; i.e., H. nudus
and H. oregonensis can withstand relatively long
periods of desiccation because of a gill reduc¬
tion, and consequently are found in the "middle
high” intertidal zone, where they are exposed
for many hours each day. P. crassipes, on the
other hand, presents an enigma in that the
usual number of gills (18) for strictly aquatic
forms is not altered; however, the total volume
of the gills in relation to the total volume of
the body is the least of any of the animals
examined; and the anterior arthrobranch of the
second thoracic segment, the gill which is lost
in both species of Hemigrapsus, is minute.
Apparently, evolution with regard to the gills
in P. crassipes has tended toward the reduction
in size of all gills in lieu of reduction in num¬
ber. The proportional percentage of gill volume
to body volume in this species is somewhat less
than in either species of Hemigrapsus, but seems
inadequate to account for the tolerance of the
former type to a drier environment. The slight
difference between the gill-volume-body-volume
relationship in the three grapsoid types corre¬
sponds with their similarity in resistance to

desiccation. In fact, in the Puget Sound area
where P. crassipes does not occur, H. nudus
apparently ranges in the tidal horizon which
would normally contain P. crassipes; however,
the individuals of the former species at this
higher elevation are smaller than those within
the optimal environmental frontiers for the
species (Ricketts and Calvin, 1939).

Sex Ratio

Sexual records from collected specimens were
kept for 10 months of the year, not only to
ascertain the relative numbers of the sexes,
but to secure information on the supposed
tendency of ovigerous females to sequester them¬
selves. Virtually all the specimens were col¬
lected at night with the aid of a flashlight;
hence no sexual discrimination occurred while
collecting. A total of 2,224 animals was col¬
lected, of which number 1,141 were males and
1,085 were females. The nearly even sexual
distribution (males exceed females by approxi¬
mately 3 per cent), together with the fact that
the females equal or exceed the males during 3
widely separated months, appears to indicate
that monthly inequalities in sex ratio are insig¬
nificant. Furthermore, during August, when
1,264 crabs were collected at random, the sex
ratio was virtually even, indicating that the
ovigerous females at the height of the breeding
season (28 per cent of all females collected
were ovigerous) do not exhibit sequestering
habits; indeed, the proportionate number of
females actually increased during the breeding
season.

INTERMOLT CYCLE

The phenomenon of molting and filling of
the haemocoelic spaces by imbibition of water
has been considered by most workers to be a
recurring event in the life of Crustacea with
the intervening period between actual molts a
more or less static one. However, the profound
transformations occurring within the Crustacea
from the conclusion of one molt to the termina¬
tion of the one following are a continuous series.

Biology of Pacbygrapsus crassipes — HIATT

147

Since P. crassipes is available and easily col¬
lected along most of the west coast of the United
States, and since it readily adapts itself to lab¬
oratory confinement, it will doubtless become an
important experimental animal. Consequently,
it would be advantageous in controlled experi¬
mental studies to know the precise intermolt
condition of the animals employed. The chief
objective in this study of the intermolt cycle
was to ascertain the internal changes and to
select some external morphological feature
which would enable the worker to identify the
internal or physiological condition.

It is important to note that the changes dur¬
ing the intermolt cycle from its initiation to
its termination do not differ with the size and
age of the crab. The crab increases progressively
in age throughout the intermolt cycle, and its
completion is manifest by exuviation which is
the result of an increase in size, but the physio¬
logical level at the onset of the following inter¬
molt cycle is exactly comparable to the like
part of the preceding cycle. These transforma¬
tions, largely concealed during the intermolt
cycle, are exhibited after ecdysis. Therefore, we
may define the intermolt cycle as a series of
transformations occurring during the period
between the molts, not including the preceding
or the impending molt.

The literature relating to the intermolt cycle
contains many expressions which designate vari¬
ous externally recognizable physiological con¬
ditions, such as "soft crab,” "hard crab,” "turgid
crab,” and "limp crab.” All of these terms refer
more or less directly to the amount of water
or haemocoelic fluid contained in the tissues
and haemocoelic spaces. Olmstead and Baum-
berger (1923) were the first investigators to
analyze scientifically these intermolt conditions.

Early workers recognized a series of physio¬
logical changes for one or more periods within
the intermolt cycle, but none considered the
entire cycle representative of a continuous suc¬
cession of events. The search for knowledge
relative to the formation of the integument led

several investigators to discover isolated anato¬
mical and physiological phenomena with regard
to the intermolt cycle. The most notable of
these findings were (1) the discovery of the
accumulation of calcareous reserves on the
walls of the stomach; (2) the formation of
setae; (3) the secretion of chitin. All of these
phenomena were detected by Braun in 1875
while studying the macruran, Astacus fluviatilis.
Unfortunately, they were discovered in studies
confined to stages immediately preceding the
molt and no attention was directed to earlier
transformation; hence he recognized no seria-
tion. The meticulous investigations of Vitzou
(1882), accomplished during observations on
the histogenesis of the brachyuran integument,
did not impress upon him that a seriation oc¬
curred during endysis. Paul and Sharp (1916)
found the initial clue to seriation through bio¬
assays of the ratio of hepatopancreas weight to
body weight in the crab, Cancer pagurus. Elm-
hirst (1923) employed the traditional "pre¬
molt,” "post-molt,” and "intermolt” intervals.
Later, Yonge (1932) discovered a series of
stadia correlated with the amount of reserve
calcium in the stomach walls of the lobster,
Homarus gammarus, but this character is not
available for use in studies on Brachyura, be¬
cause, with the exception of certain land crabs
(Stebbing, 1893), nothing equivalent has been
discovered in the Brachyura. Most of the seria¬
tion described prior to the studies of Olmstead
and Baumberger (1923, 1928) is founded upon
the interval of time elapsing after ecdysis. It is
at once apparent that this method is restricted
in utility because ( 1 ) it is inadequate for com¬
parative studies between species in view of the
established interspecific differences with respect
to the time lapse during the intermolt interval;
(2) it is inapplicable for intraspecific com¬
parisons because the intermolt interval varies
directly with age; (3) crabs reared under dis¬
similar environmental conditions show incon¬
sistent intermolt intervals.

Olmstead and Baumberger, recognizing the
need for a more accurate estimation of the

148

PACIFIC SCIENCE, Vol. II, July, 1948

intermolt condition of crabs utilized for experi¬
mental purposes, subdivided the total cycle into
six stages: (1) "newly molted,” (2) "soft,”
(3) "paper shell,” (4) "hard,” (5) "pillans,”
and (6) "about to molt.” Each term describes
the character of the integument which in turn
is affected by the amount of water in the tissues
and haemocoelic spaces and the impregnation
of the integument with calcareous materials.
Although this system provides a rapid means
of grouping crabs, it possesses certain disadvan¬
tages. For example, the stage "pillans,” the dura¬
tion of which is 3 or 4 days at the maximum in
P. crassipes, is confused with the latter part of
the period "hard” and the early part of the
period "about to molt.” Furthermore, the stage
"hard” includes several easily distinguishable
stadia which are of considerable importance in
the cycle.

Recently, the solution to the problem of
securing a near universal method for both inter-
and intraspecific comparisons during the inter¬
molt period was proffered by Drach (1939),
and it is upon his studies on Maia squinado
Herbst, Cancer pagurus Linne, and Carcinides
maenas (Linne) that this study of the intermolt
cycle of P. crassipes is based.

Drach sought to establish a series of mor¬
phologically determinable "stadia” throughout
the intermolt cycle in order to construct a stand¬
ardized key to the changes which occur. Several
criteria for the establishment of these stadia
were selected: (1) each stadium should corre¬
spond to a definite internal transformation; ( 2 )
the key characters for the extremes of any
stadium must be easily recognizable, differing
considerably from those of the stadium preced¬
ing or following; (3) a sufficient number of
stages should be established so that only a small
fraction of the animals examined falls into any
one classification; (4) these stadia must be so
established that all animals collected may be
successfully and rapidly classified. This approach
to the problem seems valid because it considers
the continuous activity throughout the entire
intermolt cycle and is directed away from former

designations which imply stages of both mor¬
phologic and physiologic inactivity. The advan¬
tages of this system may be briefly summarized:
(1) it is usable for both wild and captive
specimens; (2) it is valid for comparison be¬
tween different intraspecific age classes; ( 3 ) it is
adapted for comparative interspecific use be¬
cause of the high degree of uniformity between
the higher Crustacea with regard to endysis.

The present study supplements the informa¬
tion contributed by Drach for the cancroid and
oxyrhynchoid types by extending the knowledge
of the transformations occurring throughout
the intermolt cycle to a third cosmopolitan
group, the grapsoid type.

The classification presented here for P. cras¬
sipes is a modification of that set forth by Drach.
Throughout the classification of the cycle, ex¬
ternal morphological signs have been employed
where possible, and each has been checked by
utilizing some character concerned in the genesis
of the integument. The hardening of the exo¬
skeleton is diagnostically accurate, but the major
portion of the cycle occurs during the time in
which the exoskeleton is hard; hence, other
indicative characters must be employed. The
precocious development of the spines of the
new integument furnishes the diagnostic char¬
acter utilized after the old integument becomes
totally rigid. To facilitate the interspecific com¬
parative aspect, Drach’s method of designating
the various periods and stages is adhered to.

The cycle is composed of four major divisions
designated A, B, C, and D. Each major period
is subdivided into stages with numerical indices
corresponding to the order of succession. A
major period, during which minor but distinct
transformations occur, frequently will have cor¬
respondingly more stage indices than a major
period during which but few transformations
occur. For example, A, a very brief period, ex¬
hibits two distinct integumental transformations
designated as stages A1 and A2; while C, a
relatively long period, exhibits four major in¬
tegumental transformations, C15 C2, C3, and C4.
The major periods are equivalent in all species,

Biology of Pachygrapsus crassipes — Hi ATT

149

but the supplementary stages which are diag¬
nosed in the initial portion of the intermolt
cycle by differential sclerotization of the integu¬
ment vary with the shape of the carapace of
the crabs concerned. The curvature and thick¬
ness of the carapace differ for each species,
hence, each must be studied individually to
determine the adaptability for any standardized
series of intermolt stages. This procedure was
followed in the present study; the sequence of

sclerotization is shown in Table 2. It was noted
that differences between these data and those
presented by Drach (1939) for M. squinado
were slight; consequently only slight modifica¬
tion was necessary to adjust the criteria involved
in designating comparable stadia. This succes¬
sion of rigidity in the various areas of the
integument was found to be constant regardless
of the size of the animal or the temperature
of the water.

TABLE 2

Summary of Post-exuvial Sclerotization and Loss of Flexibility in Various Integumental
Regions of a Specimen of P. crassipes with a Carapace Breadth of 28.5 Millimeters. Signs: 44,
Integument Completely Soft; +, Integument Partially Rigid but Easily Depressed; — , Integu¬
ment Completely Rigid.

DAYS

AFTER

MOLT

CARAPACE

THORACIC

STERNITES

CARPUS AND

PROPODUS OF

CHELAE

MERUS AND

CARPUS OF

WALKING LEGS

STAGE

Proto-

gastric

Area

Meso- and
(Jrogastric
Area

Anterior

Branchial

Area

Posterior

Branchial

Area

Branchio-

stegites

Cardiac

Area

1

44

44

44

44

44

44

44

44

44

Ai

2

4

44

44

44

44

44

44

44

44

Aa

3

4

44

44

44

44

44

44

44

44

Aa

4

—

44

44

44

44

44

44

4

44

Aa

5

—

44

44

44

4-+

44

44

4

44

Aa

6

—

4

4

44

44

44

44

4

44

Bi

7

—

4

4

44

44

44

44

4

44

Bt

8

—

4

4

44

44

44

44

4

44

Bx

9

—

4

4

44

44

44

4

4

44

Bx

10

■ —

4

4

44

44

44

4

44

Ba

11

—

4

4

44

44

44

4

—

44

Ba

12

—

4

4

44

44

44

4

—

44

Ba

13

—

—

4

4

44

44

4

— .

44

Cx

14

—

—

—

4

4

4

4

—

44

Cx

15

—

—

—

4

4

4

4

—

44

Cx

16

—

—

—

4

4

4

—

44

Cx

17

—

—

—

4

4

4

—

—

4

Cx

18

—

—

—

—

4

4

—

—

4

Ca

19

—

—

—

—

4

4

—

—

4

Ca

20

— •

—

—

—

4

4

—

—

4

Ca

21

—

—

—

—

4

4

—

—

4

Ca

22

—

—

—

—

4

4

—

—

4

Ca

23

—

— -

—

—

4

4

—

—

4

Ca

24

—

—

—

—

4

4

—

—

4

Ca

25

—

— ■

—

— .

4

4

• —

—

4

Ca

26

—

—

—

—

4

4

—

—

4

Ca

27

—

—

—

—

4

4

—

—

4

Ca

28

—

__

—

—

4

4

—

—

4

Ca

29

— -

—

—

—

4

4

—

—

4

Ca

30

—

—

—

—

4

4

—

• —

4

Ca

31

—

—

—

■ —

4

4

—

—

4

Ca

32

—

—

—

—

4

4

—

—

4

Ca

33

—

—

—

—

4

4

—

—

4

Ca

34

—

—

—

—

—

—

—

—

—

Cs

150

PACIFIC SCIENCE, Vol. II, July, 1948

.Prior to a consideration of the diagnostic
characters for each intermolt stage, a brief
summary of the morphologic pattern of the
integument is presented, since a knowledge of
integumentation is necessary in order to imple¬
ment definitions of stadia. The integumentary
pattern was studied and accurately described as
early as 1860 (Williamson), but we owe our
present extensive knowledge of its histological
composition to the work of Vitzou (1882).
The integumentary strata are described briefly,
beginning with the outer stratum and progress¬
ing inward. The laminar association of these
strata may be observed in Figure 5.

1. A thin layer without definite morphologic
structure (improperly designated cuticle),
which should be termed epicuticle to be
consistent with the corresponding forma¬
tion in insects (Wigglesworth, 1933).

2. A series of parallel strata, unequally calci¬
fied, occasionally pigmented, which con¬
stitutes the pigmented layer.

3. A series of parallel strata, greatly calcified,
generally devoid of pigment, thicker than
the pigmented layer, and providing from
three- fourths to four-fifths of the entire
integument.

4. An internal layer consisting of a very thin
lamella, which is densely chitinized but
non-pigmented, and known as the mem¬
branous layer.

It has been known since the work of Vitzou
(1882) that the epicuticle and pigmented layer
are secreted by the epithelium before the molt
and, therefore, underly the old integument;
whereas, the principal and membranous strata
are not secreted until after ecdysis.

The time intervals indicated in the following
description of intermolt stadia are always the
result of the function of several integrating
factors. Important among these are the size
of the individuals, the temperature of the water,
and the adequacy of the food supply. The
writer is acutely aware of the misconceptions
stemming from data derived from captive ani¬

mals; however, the variable factors in the en¬
vironment were held to a minimum wherever
possible. For example, the food supply, which
in nature is abundant, was completely adequate
in the laboratory insofar as could be ascer¬
tained; the mean daily temperature of the lab¬
oratory water varied from 15.2° C. in mid- April
to 16.3° C. in June. The eight animals employed
in this study ranged in size from 22.2 to 37.4
millimeters in carapace breadth. Differences in
the intermolt interval ranged from 50 days for
a crab 22.2 millimeters in width to 77 days for
a crab 34.4 millimeters in width; the mean
interval was 68 days. The above-mentioned
intervals are somewhat longer than those oc¬
curring in nature for similar sized crabs. For
this reason, the proportional percentage interval
of each stage in relation to the entire intermolt
interval is utilized. Although slight errors may
occur in case of differential prolongation of
certain stages in captive animals, the presenta¬
tion utilized seems adequate for all practical
purposes.

Period A. Duration: 1V^ to 3 days; 4.0 per
cent of the total interval. This is the period
which immediately follows the molt. The cara¬
pace is completely soft and will depress at the
slightest pressure of a finger. This period is
subdivided into two stages:

Stage At. Duration: 8 to 12 hours; 1.5 per
cent of total interval. During these early
post-molt hours the integument has the
consistency of a soft membrane. Movement
is possible, but the animal moves only when
disturbed and never elevates the body above
the substrate. The color is brilliant in con¬
trast to the pre-molt crab. Water is still
being absorbed; weight and size are there¬
fore indeterminate (Fig. 6). The turgidity
of the gastric area, due perhaps to the filling
of the stomach with water during imbibi¬
tion, may be felt by passing the finger over
this area of the carapace.

Stage A2. Duration: 1 to 2 days; 2.5 per cent
of the total interval. The carapace now has

Biology of Pachygrapsus crassipes — HIATT

151

a parchment-like consistency. The crab can
elevate the body upon its legs and move
with considerable agility. Water absorption
has terminated and the weight is virtually
constant henceforth. The color of the cara¬
pace is similar to A4. No food is ingested
as yet.

Period B. Duration: 4 to 6 days; 8.0 per cent
of the total interval. The carapace is somewhat
rigid, although deformable everywhere save in
the protogastric area. The meso- and urogastric
areas, together with the branchial areas, are
somewhat rigid; the remaining regions, includ¬
ing the branchiostegites and sternal areas, remain
plastic. The appendages are flexible and may be
bent without breaking. Feeding is resumed by
the larger crabs on the sixth day following the
molt. This period is likewise divisible into two
stages, depending upon the degree of rigidity
of the chelae:

Stage Br Duration: IVz to 3 days; 3.0 per
cent of the total interval. The integument
of the merus and propodus of the chela is
relatively supple; it may be depressed with¬
out breaking. Most of the animals begin to
take food during this stage.

Stage B2. Duration: 2Vi to 4 days; 5.0 per
cent of the total interval. The integument
of the merus and propodus now resists pres¬
sure; if great pressure is applied it will
break.

Period C. Duration: 32 to 50 days; 67.0 per
cent of the total interval. This period, the most
extensive of the cycle, is initiated when the in-
tegumental areas become rigid, although some,
exhibiting less convexity, may retain their plas¬
ticity throughout the first two stages of this
period. Complete sclerotization is achieved by
the onset of Stage C3, making it mandatory that
other diagnostic criteria for integumentation be
employed beyond this point. This period con¬
tains four stages with a fifth stage a likelihood:

Stage Q. Duration: AVz to 5 days; 8.0 per
cent of the total interval. Except for some
elasticity in the anterior and posterior

branchial areas together with the concave
intestinal region, the carapace is completely
rigid. The branchiostegites and sternites
are still flexible, as are the carpus and merus
of the ambulatory legs which may be bent
in the flattened direction. The broad sides
of the pereiopods may be squeezed together
very easily.

Stage C2. Duration: 7 to 10 days; 14.0 per
cent of the total interval. With the excep¬
tion of the intestinal area the carapace is
entirely rigid. The branchiostegites, ster¬
nites, and ambulatory legs are less flexible
but have not attained rigidity. It is rather
difficult to distinguish between stages Q
and Co, inasmuch as these differences are a
matter of degree. The recognition of each
stage must be gained through experience.

Stage C3. Duration: IV2 to 11 days; 15.0 per
cent of the total interval. At the initiation
of this stage the entire exoskeleton becomes
rigid, making it easily distinguishable from
Stage C2, and the internal membranous
layer has not yet been formed, thereby dif¬
ferentiating this stage from Stage C4. At
the termination of Stage C2 the principal
layer is completely synthesized, whereas the
membranous layer becomes totally synthe¬
sized at the termination of Stage C3. There¬
fore, the four integumental strata are com¬
plete at the onset of Stage C4.

Stage C4. Duration: 15 to 22 days; 30.0 per
cent of the total interval. This is the most
extensive stage of the cycle and is primarily
characterized by the completion of the in¬
tegumentary skeleton. The internal mem¬
branous layer, when present in its early
stages of genesis, is not closely adherent to
the more external principal layer synthe¬
sized previously. By cracking a portion of
the carapace and cautiously elevating it, the
shiny membranous layer is seen to overlie
the epidermis. The distinction between
Stages C3 and C4 is perhaps the sharpest be¬
tween any two successive stages. For rapid
diagnosis the following procedure has been

152

PACIFIC SCIENCE, Vol. II, July, 1948

found to be more useful and more easily
accomplished than chipping away the up¬
per layers of the carapace: With the for¬
ceps, grasp one of the dactyls about midway
of its length and break it by bending it first
to one side and then to the other. After
breaking, exert an outward tug in an at¬
tempt to withdraw the broken distal half
from the proximal portion. The withdrawn
portion of the dactyl consists of the three
upper strata; the fourth layer (membra¬
nous layer), if present, will remain attached
to the proximal half of the dactyl because it
is not closely adherent to the three upper
strata now withdrawn. The sac formed by
the internal membranous layer retains the
form of the removed dactyl. Caution should
be exercised in the operation to avoid de¬
stroying the membranous layer. Stage C4
is verified if the sac is present; if it appears
absent, the stage is either C3 or C4T.

Stage C4T. Duration: From Stage C4 until
the death of the crab. In this stage the in¬
ternal membranous layer has become closely
adherent to the principal layer and is not
recognizable except upon histological ex¬
amination.

This stage is common in most of the
genera of larger Brachyura in which ecdysis
occurs until the attainment of a certain
maximum size; growth, therefore, ceases at
this point. Its occurrence in P. crassipes
seems substantiated by the evidence pre¬
sented herewith. MacGinitie (1937) de¬
scribed a specimen covered with barnacles
and even small Mytilus edulis. He omits
size data, but from a knowledge of the dur¬
ation of the early stages of the intermolt
cycle this animal was probably in the C4T
stage. On August 5, 1940, three large males
of P. crassipes (44.2, 47.2, and 47.8 milli¬
meters in carapace breadth) were collected;
each supported one specimen of Balanus
glandula upon its carapace. One of the bar¬
nacles had a diameter of Va inch and the
others were smaller. All three crabs were

apparently in the C4T stage. Moreover, a
large male in the C4 stage (width, 47.6 mil¬
limeters) was collected on August 15, 1940.
Nearly 8 months later this crab was still
in the C4 stage with no evident signs of an
impending molt. In field collections only
one male over 44 millimeters in width was
found in a stage later than C4, but many
were designated as Q; likewise, no females
over 40.0 millimeters in breadth were in a
stage later than C4. The evidence would
seem to insure the validity of this stage, but
so few crabs are collected near the maxi¬
mum size that this stage must be generally
uncommon and most crabs apparently die
or are destroyed by predators, etc., before
reaching a maximum size.

Period D. Duration: 10 to 16 days; 21.0 per
cent of the total interval. During this period
the integument undergoes a series of transfor¬
mations preparatory to the impending molt, dur¬
ing the course of which the future spines are
formed by the secretion of the epicuticular and
pigmented layers of the new, developing integu¬
ment. The termination of this period is marked
by resorption of the lime salts of the integument
in both localized and generalized areas. At the
onset of the period a diminution of activity,
which is accentuated as ecdysis approaches, is
apparent. There are four easily identified stages
in this period:

Stage D4. Duration: 3^2 to 6 days; 8.0 per
cent of the total interval. This stage is
marked by the formation of spines which
are secreted several days before the initial
chitinous stratum of the pigmented layer is
deposited. When the dactyl is broken and
examined, soft spines which push out the
internal membranous layer overlying them
can be observed. It is necessary to manipu¬
late the membranous sac to bring the spines
into view, inasmuch as their extreme soft¬
ness causes them to be closely appressed to
the surface of the sac. This character is the
only clear-cut distinction between C4 and

Biology of Pachygrapsus crassipes — Hi ATT

153

Dl5 there being no important changes in
the old integument. However, toward the
latter portion of this stage gelatinization of
the membranous layer is initiated. This may
be easily demonstrated by chipping off a
segment of the carapace. The broken piece
will become freed very easily and will re¬
veal the gelatinous layer below. Muscular
insertion on the old integument is not im¬
paired since no great diminution in mus¬
cular activity is yet apparent.

Stage D2. Duration: 31/2 to 6 days; 8.0 per
cent of the total interval. This stage is char¬
acterized by secretion of part of the pig¬
mented layer. Diagnosis is based upon the
chitinization of the spines, which are hard
in this stage. When the dactyl is exsheathed
the new dactyl appears similar to the old
removed portion, as though the old integu¬
ment were a glove covering the new.

Stage D3. Duration: lVi to 314 days; 4.0 per
cent of the total interval. Resorption has
now progressed sufficiently far to be recog¬
nized. Slight pressure on the epimeral line
will result in its splitting along its entire
length. The membranous layer is com¬
pletely gelatinous, and resorption is virtu¬
ally completed in all parts of the exoskele¬
ton. The activity of the crab is much
reduced.

Stage D4. Duration: 12 to 15 hours; 1.0 per
cent of the total interval. This abbreviated
stage immediately precedes exuviation. Re¬
sorption is now complete, and the epimeral
line splits along its entire length. The ac¬
tivity of the crab entirely ceases because the
muscles are now inserted on the supple,
new integument. The carapace is elevated
posteriad and exuviation ensues.

In an effort to test the validity of the identifi¬
cation of the various intermolt stages, a com¬
parison was made between the intermolt condi¬
tion of animals collected in the field and the
proportional percentage of time represented by
the individual stages of the entire cycle as ascer¬

tained from captive specimens. A field exami¬
nation to ascertain the intermolt stage of 574
individuals was made in 4 days during August,
1940. Frequencies of crabs within the various
intermolt stages, together with the proportional
interval for any given intermolt stages as deter¬
mined by laboratory methods, are indicated in
Figure 2. A marked correlation between field
and laboratory data is apparent, thus serving to
confirm in part the criteria for the divisional
diagnoses of the intermolt cycle set forth above.

Several phenomena which normally would
be discussed in other sections of this paper are
briefly considered at this time because of their
direct association with various intermolt stages.
Throughout the examination of the several hun¬
dred specimens mentioned in the paragraph
above, it was noted that ovigerous females con¬
stantly fell into stages C3 and C4. The data
indicate that of a total of 80 ovigerous females,
23 were classified as stage C3, 55 were placed

FIG. 2. A comparison between the frequency of
occurrence of the intermolt stages in wild P. crassipes
examined during August and the proportional extent
of the individual stages with respect to the time re¬
quired for the entire intermolt interval ascertained
from captive crabs during the same seasonal period.
The solid line represents the frequency of wild crabs
in the various intermolt stages; the dashed line indi¬
cates the proportional amount of the intermolt interval
assigned to the various stages.

154

PACIFIC SCIENCE, Vol. II, July, 1948

in stage C4, while stages D4 and D2 each con¬
tained one berried crab. It seems apparent,
therefore, that ova are extruded onto the abdom¬
inal endopoditic setae during stage C3 and are
normally hatched prior to the attainment of
stage D4. In view of the facts that the incuba¬
tion period of the ova has been found to range
from 25 to 30 days (see p. 201 ), that the normal
intermolt interval in female crabs ranging from
16 to 40 millimeters in carapace breadth varies
from 23 to 40 days (Table 3), and that stages
C3 and C4 combined comprise approximately
one-half of the entire intermolt cycle, it is quite
evident that ovigerous females undergo an
abnormally prolonged intermolt interval.

Behavior patterns of P. crassipes are obviously
dependent to a greater or less extent upon the
intermolt stage at any given time. It was ob¬
served that certain crabs in the communities
under observation tended to remain secluded
in crevices throughout the daylight hours. Dur¬
ing August, approximately half of the tide-
pool population remained secluded, never enter¬
ing into the active life within the pool or its
environs. A reasonable explanation of this se¬
questering habit is found in Figure 7, wherein
it is shown that more than 55 per cent of the
total population of crabs was in a relatively
soft condition ( stages At- C2 ) . During diurnal
hours crabs apparently seclude themselves imme¬
diately preceding ecdysis and remain concealed
until stage B4 is attained. Evidence for this
inference was secured from the fact that diurnal
collections of crabs in tide pools has failed
to include crabs in stages D4 to A2. This con¬
cealment during daylight hours is almost cer¬
tainly an act of self-preservation. A crab in
stages D4 to A2 is virtually defenseless inasmuch
as the chelae are supple and the muscular inser¬
tions are in a state of reduced efficiency because
of the flexible integument.

Nocturnal collections, on the other hand,
consisted of crabs in all stages of the intermolt
cycle. The absence of light probably provides
the animals with increased freedom of move¬
ment and safety because the vision of predators

as well as of the crabs themselves, which are
cannibalistic on occasions (see p. 179), is con¬
siderably reduced. Night, therefore, is the most
opportune time to make collections of this
species. A representative sampling of all inter¬
molt stages may be made at this time, whereas
it is virtually impossible to do so during day¬
light hours.

It is the writer’s firm conviction that many
types of experimental data may be accurately
secured through the use of comparatively few
animals if the intermolt cycle is correctly diag¬
nosed. Therefore, by extending the knowledge
of the intermolt cycle to the grapsoid group, a
framework upon which to base both experi¬
mental and natural history data is provided.
Observations in both the laboratory and field,
particularly the former, will in most cases be
inaccurate and misleading if the intermolt stage
of the animals, which represents a clue to their
physiological state, is not diagnosed and con¬
sidered.

ECDYSIS AND ASSOCIATED PHENOMENA

One of the most significant features in the
life history of the Arthropoda is the act of
molting, an incident in, and an expression of,
growth. Nowhere among the Arthropoda is the
process so striking and abrupt as it is in the
higher Crustacea. Notwithstanding the volumi¬
nous aggregation of literature concerning the
life histories of crustaceans, few descriptions
of the actual process of ecdysis exist, and none
has been published upon the exuviation of
grapsoid crabs. One of the objectives of the
present study has been to describe meticulously
ecdysis in P. crassipes. In addition, considerable
statistical data pertinent to various aspects of
the molt and its effect on the crab have been
accumulated and set forth below. From the
above-mentioned data, an attempt has been
made to ascertain the age groups in the exist¬
ing crab population, as well as the age-size
relationship.

Reaumur (1712, 1719) was the first investi¬
gator to describe exuviation of a decapod crus-

Biology of Pachygrapsus crassipes — Hi ATT

155

tacean in detail, the exuviation of the river
crayfish ( Astacus fluviatilis) . Later, Couch
(1837) was amazed to find a perfect exuvia of
the lobster ( Astacus etiropeus ) and was respon¬
sible for the initial inference that after a cer¬
tain size is reached the lobster discontinues the
molting act. His description of exuviation ac¬
complished by the common edible crab, Cancer
pagurus, in 1843 was the first account of ecdysis
within the Brachyura. Salter (I860) read be¬
fore the Linnaean Society an account of ecdysis
in the lobster and described in considerable
detail each precasting and casting activity, and
emphasized that the observation was indeed
rare. That he was entitled to this assumption is
indicated by my experience with P. crassipes
(see p. 160). Several later descriptions of the
molting act of the lobster (Packard, 1886; Her¬
rick, 1909; and Elmhirst, 1923), together with
the description of ecdysis in M. squinado by
Drach (1939), have served to provide a basis
for comparison between the brachyuran and
astacuran modes of exuviation.

Exuviation

Two distinct phases of ecdysis are apparent:
(1) a passive phase (no significant muscular
activity), initiated in P. crassipes approximately
1 day prior to the molting act and manifest
by water absorption which serves to increase
body volume and which exerts sufficient pres¬
sure to separate the epimeral line or pleural
groove on the branchiostegites; (2) an active
phase, which is identified by actual muscular
activity significant to the molting act. The
active phase is initiated immediately after the
split along the pleural groove, and its termina¬
tion is manifest by the complete exuviation.

Crabs used in the study of ecdysis were reared
in individual aquaria tilted to provide deep sea
water at one end and none at the other, thus
closely simulating the natural water-air rela¬
tionship of the normal environment. Many of
the crabs resided in the aquaria in excess of 8
months, and in that interval successfully com¬
pleted their fourth molt. The laboratory diet

consisted of small pieces of fresh liver offered
once daily, after which the water was changed.
To facilitate observation of the molting act as
well as other behavior traits, a shelf with a
glass bottom was constructed directly above
the author’s desk, thus providing a means of
constant observation for most of the day. It was
found that observation from the ventral aspect
provided a maximum opportunity to view the
details of ecdysis.

Both behavioristic and morphological changes
are manifest prior to the impending molt. For
4 or 5 days preceding the molting act crabs
above 10 millimeters in width cling to a corner
of the aquarium and ordinarily refuse food, but
exceptions were encountered in which a small
quantity would be eaten until 2 days prior to
the molt. This abstinence from feeding is con¬
spicuous because the animals are normally vora¬
cious, actually leaping to seize suspended food.
Crabs below 10 millimeters in width regularly
consume food until 2 days before ecdysis with¬
out exhibiting signs of inertness, a characteristic
undoubtedly stemming from the shorter inter-
molt interval undergone by small individuals.
The abstinence from food and the general
seclusiveness seem to be natural phenomena
common to both astacuran and brachyuran
types. Elmhirst (1923) and Herrick (1909)
described identical reactions in the lobster, while
Broekhuysen (1941) indicates similar habits for
Cyclograpsus punctatus M. Edw. on the South
African coast. Muscular activity in P. crassipes
virtually ceases; the crab will move but slightly
if disturbed the day preceding the molt. The
diminished activity preceding the molt was like¬
wise noted by Hay (1905) for Callinectes
sapidus, and undoubtedly results from the tran¬
sitory condition of muscle insertions as they
change from the old integument to the new.
It is apparent that movements by crabs in this
condition would be decidedly ineffectual.

A trio of morphological signs indicative of
the impending molt are relatively precise in
this species. The first, and most perceptible
change, is that in the pigmentation, which, al-

156

PACIFIC SCIENCE, Vol. II, July, 1948

though less pronounced than in some astacuran
forms (the lobster, Herrick, 1896, and Elm-
hirst, 1923; and the crayfish, Braun, 1875), is
distinctive. Like C. punctatus, P. crassipes be¬
comes drab, losing the characteristic luster of
the integument so apparent prior to stage D3.
In addition to the pigmentary deposition in the
new integument during D3 and D4, the opacity
is probably also the result of the dissolution of
the deeper layers of the old integument. A sec¬
ond morphological indicator of the impending
molt is the resorption of the lime salts in cer¬
tain areas of the integument; namely, the
branchiostegites, the sterna, and the basi-ischi-
opodite and meropodite of the chelae (Fig.
3, R. ) . The epimeral lines become desclerotized,
allowing the upper and lower halves of the
branchiostegite to be forced apart as a result
of the increased pressure of the haemocoelic
fluid. This desclerotization occurs several days
prior to the molt and may be discovered by
applying slight pressure on the pleural groove.
Approximately 3 days before the molt, the
pressure will result in a precocious schism along
the groove. A third morphological sign of the
premolt phase is the friability of the carapace.
Among specimens collected in the field, many
had carapaces which cracked as they were
grasped. Upon examination they were found to
be in stage D3. This significant character was
utilized by Olmstead and Baumberger (1928)
as the criterion upon which the stage "pillans”
was based. Crabs of economic importance are
aptly designated "peelers” in this stage because

C., carpus; CO., coxa; D., dactylus; F.P., fracture plane;
M., merus; P., propodus; R., resorption line.

the old integument may be picked off to reveal
the lustrous new integument underneath.

Fig. 4. A diagrammatic indication of the expansive
movements occurring during the passive phase of
ecdysis. Diagram A exhibits the relationship of the
old and new integuments prior to ecdysis. Diagram B
shows the relationships at the termination of the pas¬
sive phase. AR., arch of branchial chamber; AR'., new
arch of branchial chamber; BR., gill; CP., old carapace;
CP'., new carapace; E.L., epimeral line; EP., epimeron;
EP'., new epimeron; N.I., new integument; PL., pleural
wall; PL'., new pleural wall; ST., sternal plastron.

On several crabs, the separation of the notal
and pleural halves of the branchiostegites oc¬
curred a day prior to the active phase of the
molting act. As the rift between the two halves
of the branchiostegite widens, the gap between
the two becomes continuous with the transverse
lacuna between the posterior edge of the cara¬
pace and the first abdominal tergite. The ele¬
vated carapace resembles the lid of a box hinged
at the anterior end of the animal, and under¬
neath the old integument may be seen the new,
somewhat wrinkled one. The expansive move¬
ments within the body of a crab undergoing
ecdysis are diagrammatically indicated in Figure
4. The arrows indicate vertical and lateral ex¬
pansion. The new, underlying integument is
thin and wrinkled prior to the splitting of the
epimeral line (Fig. 4 A), after which the old,
thicker carapace is elevated allowing the new
integument to stretch to the size to which it
will eventually harden (Fig. 4 B).

Late in the passive phase the epimeral split
extends anteriorly almost to the mouth. Some
inconsistencies with respect to the origin of the

Biology of Pacbygrapsus crassipes — Hi ATT

157

schism appear in the literature. Pearson ( 1908,
on Cancer pagurus) and Hay (1905, on Colli-
nectes sapidus) contend that the epimeral frac¬
ture appears first at the anterior end of the
groove near the oral region and extends pos¬
tered to connect each end of the gap between
the posterior edge of the elevated carapace and
the first abdominal tergite. Churchill (1918,
on C. sapidus) refutes Hay and suggests that
the converse is true. This study on P. crassipes
definitely corroborates Churchill’s findings.
Many of the crabs which succumbed during the
passive phase of the molt show the fracture in
several stages of its splitting process; each frac¬
ture is oriented from the posterior to the anterior
direction. Indeed, one may gently elevate the
carapace and observe the actual splitting an¬
teriorly.

Resorption of lime salts from the sternal
integument is not apparent externally until the
onset of the active phase of ecdysis. A split
between the fused post-oral cephalothoracic and
the fourth thoracic sterna, together with a schism
along the mid-sternal line at the junction of the
right and left sterna, is visible at the time the
limbs are withdrawn. Slight pressure on these
desclerotized areas prior to the molt likewise
serves to detect the impending molting act.

When the elevated carapace reaches an angle
of approximately 30° as a result of increased
body expansion, the pereiopods are moved
slightly, simulating walking movements, and a
depression appears to one side of the cardiac
area in the new integument. This initial depres¬
sion disappears, but concurrently a new depres¬
sion is developed on the opposite side of the
cardiac area; each individual depression persists
approximately 10 seconds and recurs rhyth¬
mically for a period of about 7 minutes in an
average-sized female crab of 24.6 millimeters
in width. A smaller crab with a carapace
breadth of 7.2 millimeters was observed to com¬
plete its active phase of exuviation in 3 minutes
and 10 seconds. Drach (1939) reports that the
pulsations in M. squinado vary from 10 to 15
minutes. He further states that the alternating
depressions are the result of muscle contraction

in the coxopodites and basipodites of the
thoracic appendages, and that these particular
muscular movements are the principal ones
occurring during the active phase. The muscles
mentioned take origin on the endopleurites and
endosternites, neither of which has appreciable
rigidity at ecdysis; consequently, these newly
formed skeletal elements offer less resistance
to the contraction of the muscles than their
insertions on the new integument of the coxo¬
podites and basipodites which are still en-
sheathed by the old integument. Inasmuch as
the endoskeleton and exoskeleton are continuous
integumental structures, one would expect that
particular part of the carapace (cardiac area)
to be moved concurrently with the adjacent
pleural wall. Since all of the muscles inserting
on the coxa and basis of the various pereiopods
take their origins in the same area of the pleural
wall, and since the pleural wall is bound to
the cardiac area of the carapace by a non-
sclerotized portion of the integument, it is
obvious that the depression occurring in the
cardiac area during ecdysis will take place in
that area only regardless of which legs are
being withdrawn from the old integument. The
hydrostatic pressure within the haemocoelic
cavity forces the depression to disappear when
the muscles on that particular side become
quiescent.

The resultant accomplishment of these pul¬
sations mentioned above is the withdrawal of
the fourth and fifth pereiopods from the old
integument. When the coxae and basipodites
of the remaining legs are withdrawn, a rotation
of the freed segments occurs in a manner sug¬
gestive of an attempt to release the distal podo-
meres from their sheaths. As the podomeres
are successively withdrawn they quickly become
distended with haemocoelic fluid and are thereby
stiffened to assist in exerting a tug on those
remaining in the old sheath.

The fifth pair of walking legs is the first
pair to be withdrawn because withdrawal activ¬
ity starts there, and also, perhaps, because they
are the shortest pair. However, all undergo
simultaneous exuvial progress which ultimately

158

PACIFIC SCIENCE, Vol. II, July, 1948

results in complete consecutive withdrawal of
all pereiopods from the fifth pair to the chelae.
Special difficulties in withdrawal are presented
by the chelae because of the enlarged distal
podomeres. Special lines of absorption on the
coxa, basi-ischium, and merus have been men¬
tioned previously (Fig. 3, R.). This resorbed
line fractures to release a roughly triangular
flap of integument. This flap is hinged at the
junction between the merus and basi-ischium
and the anterior portion of the line of absorp¬
tion on the merus. The lines of absorption on
the basi-ischium and coxa fail to fracture but
become very soft and pliable.

The frequency with which the animals under¬
going ecdysis lose one or more limbs illustrates
the critical nature of exuviation in the general
welfare of the individual. A cursory examina¬
tion of several exuviae which contained chelae
of crabs unable to withdraw them from the old
exoskeleton revealed that the lines of absorp¬
tion were hard, with resorption insufficiently
advanced to effect the fracture, which indicated
that resorption of calcium salts in the old integu¬
ment was apparently impeded in some manner.
The severence of appendages through the in¬
ability to withdraw them successfully has been
recognized previously and designated "exuvial
mutilation”; however, a more precise expression,
'exuvial autotomy,” suggested by Drach ( 1939)
seems more appropriate.

After the successful withdrawal of the pos¬
terior four pairs of appendages, the old sternal
integument is separated from the new. An
elongation of the body, together with an upward
and backward movement of the crab, combines
to exert the necessary pull on the abdomen,
which is subsequently withdrawn. The limbs
are then directed forward. An ejection of
amber fluid from the mouth occurs, after which
a few seconds of quiescence ensues. After this
brief period of inactivity, rapid movements of
the limbs occur in a forward-backward motion.
This activity is repeated two or three times,
and serves to remove the buccal appendages
and the chelae. The amber ejection is probably
the material in the stomach which must be

expelled with the old epithelium. The chelae
are decidedly wrinkled as they are exsheathed
because of the forced expulsion of the haemo-
coeiic fluid to enable the bulbous distal end to
be withdrawn through the small coxal opening.
However, they become turgid immediately after
ecdysis.

An examination of the exuvia shows that
the apodemes and branchial filaments are pre¬
served in approximately their original morpho¬
logical position. Since it is not possible to
observe the exsheathing of the apodemes and
gills this phenomenon must be interpreted
through critical study of the morphological
characteristics of the exoskeleton.

The apodemes are invaginations of the in¬
tegument, each lamina being formed by two
thicknesses of sclerotized integument united by
their morphologically external surfaces. Prior
to the impending molt new strata are secreted
and these strata (the epicuticle and pigmented
layer) of the new integument lie internal to
the old integumental strata of the apodemes
(Fig. 5); during the withdrawal of the ap¬
pendages the new apodeme is slipped off the old,
which is rigidly attached to the exuvia. Mechan-

Fig. 5. A diagrammatic section through an apodeme
to show the integumental strata. The arrow indicates
the direction of withdrawal at ecdysis. E., epicuticle;
E/, new epicuticle; M.L., membranous layer; P.L., pig¬
mented layer; P.L/, new pigmented layer; PR.L., prin¬
cipal layer.

Biology of Pachygrapsus crassipes — Hi ATT

159

ically, the casting off of the old apodemes is
facilitated because they are generally directed
toward the basal portion of the appendage;
therefore, the withdrawal of the new appendages
presents no special difficulty. A comparison
between the apodemes of a skeleton and those
of an exuvia shows that those apodemes not
directed posteriorly are either resorbed in stages
D3 and D4 or are secondarily oriented in a
posterior direction after the sternal schism.

The apodemes which partition the two sides
of the thorax offer more difficulty to successful
exuviation than those of the pereiopods. They
consist of invaginations of the endosternites and
pleural walls and serve as the attachment for
the origins of the muscles of the basipodites
and coxopodites. It was stated above that when
the body is elevated above the old sternal floor,
this elevation, in conjunction with anteropos¬
terior elongation, serves to disengage the apo¬
demes which are directed posteriorly after the
sternal schism. A comparison of the endo-
skeletal structure of a normal animal with that
of an exuvia shows that the normal structure
has been altered during ecdysis. This altera¬
tion is manifested in partial or total resorption
in certain endoskeletal areas. Without this re¬
sorption, molting would not be possible because
the transverse apodemes together with their
ramifications would hinder the withdrawal of
tissues without injury. The occurrence of the
sternal schism in P. crassipes serves to reduce
effectively the degree of resorption which would
otherwise be requisite providing the sternum
remained entire.

The structural association of the phyllobran-
chiate gills with the proximal portion of the
gill-bearing thoracic appendages makes it im¬
perative that the withdrawal of the gills coincide
with the onset of the active phase of ecdysis.
Inasmuch as the brachyuran gill is composed
of two series of lamellae inserted on a median
raphe, the lamellae must flatten and be drawn
through the median raphe over the orifice of
insertion on the skeleton. The branchial aper¬
tures of the skeleton are small, but the edges
are rounded so that the gills pass through them

without great difficulty. Subsequent to the com¬
plete withdrawal of the new gill from its old
integument, it remains wedged between the
old and new epimeral walls until the body is
elevated to a position above the old exuvia and
until the proximal podomeres are moved to the
upper level of the old pleural wall. It is evident
that the abnormal torsion undergone by the
gills during ecdysis might conceivably provide
respiratory difficulties which would designate
ecdysis as a critical interval in the life of the
animals. Supportive evidence for the above
statement was secured upon the compilation of
mortality data for crabs confined to the labora¬
tory. Of a total of 78 fatalities among laboratory
animals, 61 (78.2 per cent) occurred prior to
complete withdrawal of the pereiopods. There¬
fore, these deaths took place when the gill
lamellae were in an abnormal, contorted con¬
dition.

One of the features of ecdysis which invites
discussion is the origin of the mucilaginous
material which covers the inner surface of the
exuvia and the external surface of the new
exoskeleton. Its apparent function of lubrica¬
tion of the frictional surfaces during ecdysis is
manifestly significant. Its initial appearance in
P. crassipes occurs when the rift between the
posterior edge of the carapace and the first
abdominal tergite shows a tenuous membrane
stretched across it. With further separation be¬
tween the carapace and the first abdominal ter¬
gite the membrane fractures. There is little
doubt that this mucilaginous layer is probably
the same as that first described by Reaumur
(1712), who recognized its apparent function.
Vitzou (1882) suggested that the mucilage
was secreted by the epithelium, in which case
it would have had to penetrate the two new
integumental strata prior to reaching its defini¬
tive location between the old and new integu¬
ments. Herrick (1896) subsequently proved
Vitzou’s contention erroneous because this layer
bore the impress of a mosaic of cells, and, in
view of its cellular composition, postulated
that it was either the first secreted portion of the
new integument or the internal membranous

160

PACIFIC SCIENCE, Vol. II, July, 1948

stratum of the old. Yonge (1932) provided
proof to substantiate the second possibility set
forth by Herrick. Furthermore, he asserted that
the internal membranous layer was dissolved
by the action of cells which passed into it from
the epithelium prior to and during ecdysis.
Drach (1939) agreed with Yonge concerning
the stratum involved but found no basis for
the mode of gelatinization suggested by that
investigator. After a meticulous histological
study of the internal membranous layer during
stages D4 to D4, Drach found that the lympho¬
cytes were irregularly grouped and insufficient
in numbers to effect a uniform transformation
of this stratum into a jellified condition. He
maintained, moreover, that the homogeneity of
this jellification presupposed the intervention
of some chemical agent secreted by the epithelial
cells. In the Insecta an enzyme, chitinase, causes
an analogous resorbing action. Wigglesworth
( 1933 ) stated that chitinase is a glandular secre¬
tion of the epidermis; therefore, it is not un¬
reasonable to assume that in the Crustacea a
comparable enzyme is secreted by epithelial
cells or glands which induces the gelatinization
so significant to the success of exuviation.

The relatively high exuvial frequency coupled
with the extreme infrequency of observations
of the molting act seem to present at least two
implications. First, ecdysis must occur very
rapidly, or second, ecdysis must take place
during times when the animals are not under
observation. It is the writer’s opinion that the
initial hypothesis may be discounted (see p.
155). The latter supposition, however, is ade¬
quately supported by data compiled from ani¬
mals confined to laboratory aquaria. Of the 14 1
molts which occurred among captive crabs,
all but three took place between 8:00 P.M. and
5:00 A.M.; thus, 97.9 per cent of the molts
occurred during nocturnal hours. Broekhuysen
(1941) and MacKay (1942) also noted that
most of their captive specimens underwent
exuviation at night. Early suggestions which
coupled molting with moon and tidal effects
are contradictory, devoid of evidence, and un¬
tenable in the present investigation; therefore,

these conjectural causes are discarded forthwith
in favor of the evidence presented below. It has
been pointed out previously that crabs in stages
A1 and A2 are extremely susceptible to injury
and possible fatality if attacked by predaceous
species during the first 6 to 10 post-exuvial
hours. The reduced vision of littoral predators,
including cannibalistic P. crassipes and the ubi¬
quitous Norway rat ( Rattus norvegicus) , is cer¬
tainly advantageous to the comparatively de¬
fenseless, recently molted crabs. The several noc¬
turnal hours following the molt undoubtedly
provide substantial, but quantitatively unknown,
protection for this species. Moreover, the ele¬
vated littoral position preferred by P. crassipes
markedly exposes them to the predation of
numerous gulls. Therefore, if molting were
undergone during the day to any appreciable
extent, the mandatory position in the tide
pools, at least for a portion of the early, post-
exuvial hours, would place these crabs in certain
danger which would be magnified because of
the subnormal locomotory powers of the soft
crabs.

Although considerable variability occurs in
the time required for maximum post-exuvial
expansion, an indication of typical expansion
with respect to the time intervals involved may
be secured from Figure 6, which illustrates these
data derived from three distinct sizes of crabs
selected to emphasize the comparative aspect.
Pre-exuvial carapace breadths were: crab A, 6.3
millimeters; crab B, 25.0 millimeters; and crab
C, 35.8 millimeters. The first post-exuvial
measurement of crab A was made immediately
after the molt and revealed a 4.7 per cent size
increment with respect to initial width at that
time. Crabs B and C were not measured until
2 hours after the completion of ecdysis. Regard¬
less of the initial carapace width, the greater
part of post-exuvial expansion occurs prior to
the third hour after ecdysis. Actually, consider¬
able expansion occurs before exuviation is com¬
pleted, because swelling of the body accom¬
panies the active phase of exuviation. However,
the expansion is not measurable until ecdysis
terminates.

Biology of Pachygrapsus crassipes — HIATT

161

Fig. 6. Typical post-exuvial expansion for three
distinct sizes of crabs. See text for details.

The impregnation of carbonates in the exo¬
skeleton of P. crassipes begins at the tip of the
chelae and becomes apparent about 3 6 hours
after ecdysis. However, the insufficient scleroti-
zation proximal to the tips prevents the average¬
sized crabs (32.0 millimeters) from employing
these structures defensively until the third or
fourth day after the molt. The pinch is ineffec¬
tive on the fourth day but becomes progressively
more effective until normalcy is nearly achieved
on the twelfth day. Inasmuch as the carpus and
merus of the ambulatory legs are the final por¬
tions of the integument to become sclerotized
(Table 2), the ability of P. crassipes to employ
these pereiopods effectively before the broad¬
ened sides have attained rigidity is attributable
to their shape and direction of movement. The
edges of these flattened podomeres are oriented
approximately perpendicular to the substrate,
and the long axis of the appendages assumes a
lateral position which forms a right angle with
the main axis of the animal. This orientation
is significant because the direction of locomotion
is likewise lateral (see p. 177). The legs are
incapable of bending on their edges, but flex
readily on their lateral surfaces prior to total
sclerotization. Locomotion in the early post-
molt interval is possible and effective because
of the shape and orientation of the ambulatory
appendages.

Seasonal Exuvial Periodicity

The recorded data indicate that ecdysis in
Crustacea inhabiting temperate areas occurs

more frequently during the warmer period of
the year. Herrick (1909), Churchill (1918),
and Broekhuysen (1936, 1941) concur in this
belief; the former two authors further stated
that most of the molts occur on the Atlantic
coast of America from June to October, while
the latter worker graphically illustrated a more
extended molting season for the South African
C. punctatus. The molting incidence of P. cras¬
sipes simulates the more extensive seasonal type
illustrated by C. punctatus.

The seasonal exuvial periodicity of P. cras¬
sipes under normal environmental conditions
was ascertained through collections of crabs
which were subsequently examined for their
intermolt stage. Although sex was disregarded
in the above analysis, it must be pointed out
here that slight abnormalities in the molting
incidence are exhibited by ovigerous females
(see p. 154). These deviations from the normal
cycle of successive molts are, however, insuf¬
ficient to alter the general chain of events.
Ecdysis occurs most frequently during August,
September, and October; but considerable exu¬
viation is apparent in all except the winter
months, November through February. The exu¬
vial periodicity presented here is particularly
apparent to the frequent intertidal-pool visitor.
The early morning receding tide discloses a
far greater number of exuviae in the pools dur¬
ing the warmer months of the year than during
the cooler winter season. The rather lengthy
span of relatively high molting frequency is
probably associated with the small range of
temperature characteristic of the geographical
range of this species.

The comparatively significant role of tempera¬
ture fluctuations in the physiology of poikilo-
thermal species, together with the fact that
temperature is one of the most widely fluctuat¬
ing physical characteristics in the environment
of these littoral animals, provides the basis for
a comparison between thermal fluctuation and
seasonal molting periodicity. The temperature
of the surface water was selected as that most
appropriate for this purpose. The figures pre¬
sented were secured from the U. S. Coast and

162

PACIFIC SCIENCE, Vol. II, July, 1948

Geodetic Survey report for the year 1939. The
readings are surface temperatures taken at Fort
Point, San Francisco Bay. The figures correspond
closely to those published by Sumner et al.
(1914), and to those taken just outside the
Golden Gate by the U. S. Weather Bureau
during the years 1915 to 1924. A marked
coincidence is noted between the exuvial peri¬
odicity and the mean surface temperatures for
the annual period (Fig. 7). It is significant
that the apices of thermal and exuvial activity
fail exactly in line. Ecdysis seems to be per¬
ceptibly slowed when the surface temperature
drops below 14° C., indicating that the physi¬
ological processes, presumably those participat¬
ing in endysis, are slackened in pace. Supportive
evidence for this thesis was secured from sev¬
eral captive crabs of similar size upon which

a careful record was kept concerning their
intermolt development through’ the cooler
months. The normal intermolt interval for
crabs of 24 to 30 millimeters in width varies
from 45 to 60 days during the relatively warm
midsummer months. However, the captives
which molted in November and early December
had only reached either stage C4 or D4 more
than 3 months later. The intermolt records of
these captive crabs show that endysis is slack¬
ened to approximately one-half the pace typi¬
cally exhibited during warmer seasons.

To check the findings described above, the
exuvial frequency of captive crabs together
with the temperature fluctuations of the lab¬
oratory water were recorded monthly. These
data are combined with the field data shown
in Figure 7.

Fig. 7. A comparison between ocean surface and laboratory water temperature and the annual exuvial
incidence in wild and captive P. crassipes, respectively. The surface temperature readings represent the monthly
mean temperatures recorded at Fort Point, San Francisco Bay, 1940.

Biology of Pachygrapsus crassipes — Hi ATT

Variation in Post-exuvial Size Increment

Considerable variation in the size increment
after ecdysis seems apparent both between dif¬
ferent species of crabs and between different
size groups within a species. In addition, a
slight differential variation in size increments
is apparent between the sexes. Williamson
(1903) and Pearson (1908) investigated the
post-exuvial size increment in C. pagurus and
presented statements which implied that the
size increment at each successive molting period
was constant. When the data presented by the
above authors were plotted by Olmstead and
Baumberger (1923) it was shown that their
conclusions were erroneous. Further, the latter
workers demonstrated that the smaller and pre¬
sumably younger specimens increased to a
relatively greater extent than did the older and
larger specimens. Broekhuysen (1941) and
MacKay (1942) found a similar condition in
C. punctatus and C. magister, respectively, while
Gray and Newcombe (1938) showed the con¬
verse to be true for C. sapidus , but to a lesser
extent in males than in females.

Individual variability, both with respect to
animals of similar size and animals of different
sizes, has been conspicuous throughout the
present investigation of the molting charac¬
teristics of P. crassipes. An examination of
Figure 8 will show that variability in post-
exuvial size increment for both wild and cap¬
tive crabs above 10 millimeters in carapace
breadth may reach as high as 200 per cent.
Smaller captive animals exhibit a variation in
excess of 400 per cent. It is also apparent that
there is a proportionate decrease in the per¬
centage of post-exuvial width increments with
age. However, when actual width increments
instead of ratios were plotted, it was seen that
young crabs (below 15 millimeters in breadth
of the carapace) increase in size at ecdysis up
to a maximum of 2 millimeters; middle-sized
crabs (between 15 and 35 millimeters in
breadth of the carapace) show increases up to
4 millimeters following the molt; while very
large crabs (above 35 millimeters in breadth

163

Fig. 8. Scatter diagram comparing the variation in
percentage of the post-exuvial width increment in
wild and captive crabs above 10 mm. in breadth of
the carapace. The regression lines were fitted by the
method of least squares.

of the carapace) show much less growth at the
molt and seldom increase in size more than 2
millimeters in breadth of the carapace.

Of great significance is the demonstrated fact
(Fig. 8) that the laboratory environment, in
spite of the fact that it was meticulously regu¬
lated, is dissimilar to the natural environment:
this is shown by the generally smaller growth
increment (size for size) for captive crabs as
compared with wild crabs. The mean growth
increments for all captive crabs and for all wild
crabs above a carapace breadth of 10 millimeters
were computed and the standard "t” test was
applied to the two sets of data. The difference
between the means of the two sets of data was
found to be highly significant statistically ("t”
= 7.27: V for 0.01 =2.8). The regression
lines shown in Figure 8 illustrate this point
graphically. The great similarity in structure
and response of the various species of the higher
Crustacea suggests a strong possibility that the
debilitating effects of laboratory life will affect
them all. It is of great importance, then, that
experimentalists treat results obtained in the
laboratory with reserve and caution before the
precise effects of confinement are well known.

164

PACIFIC SCIENCE, Vol. II, July, 1948

A great number of the captive crabs main¬
tained in the laboratory were injured specimens
(individuals which had lost one or more perei-
opods). The post-exuvial size increment data
for these individuals were recorded separately
and compared with the data derived from nor¬
mal captive animals. It was found that, in gen¬
eral, the magnitude of variation in size incre¬
ments of injured crabs is less than that for
normal captives; in addition, the individual
post-exuvial size increments are significantly
less. Underlying factors responsible for this
phenomenon seem at this time to be largely
conjectural. MacGinitie (1937) infers that
mutilation may stimulate molting in Crangon
calif orniensis as well as in other macrurous
forms. There are, however, no data presented
to support this inference. Several controlled
experiments on regeneration in P. crassipes indi¬
cate that both acceleration and slackening of the
intermolt cycle may occur, depending upon the
degree of integumental development at the time
of mutilation (see p. 194). Mutilation prior
to stage C3 lengthens the intermolt interval in
P. crassipes. Limbs severed during the early C4
stage show an accelerated regeneration, but the
intermolt interval is normal. It would seem
that an accelerated intermolt cycle would be
advantageous after the loss of one or more
appendages because it is certain that maximal
efficiency in activities concerned with general
welfare would be dependent upon the entire
complement of appendages. However, further
study is required for a satisfactory solution to
this phase of the problem.

Comparative studies on the post-exuvial width
increments of male and female crabs indicate
that no significant variation between the sexes
occurs until an initial width of approximately
25 millimeters is attained. The foregoing in¬
formation was secured by plotting initial and
post-exuvial widths of both sexes from recent
molts and exuviae collected in the normal
habitat (Fig. 9). The ratio of initial to final
width remains approximately identical for both
sexes until two rather distinct ratios become

apparent, with the digression taking origin in
those crabs about 25 millimeters in initial width.
The smaller post-exuvial size increments of
females after attainment of sexual maturity show
why representatives of this sex are invariably
smaller than males in a given intermolt cycle
after the tenth to twelfth molt (Table 3), and,
in addition, show why the largest specimens
collected are always males (Fig. 9).

Fig. 9. Sexual dimorphism in post-exuvial width
increments for P. crassipes. These data are derived
from measurements of recently molted wild crabs for
which the exuviae were found. The lines have been
fitted by eye.

Exuvial Frequency

Prior to a consideration of the number of
exuvial stages during the life span of P. cras¬
sipes, it is necessary to point out that reproduc¬
tive activity, to a certain extent, alters the usual
exuvial succession in females. There is evi¬
dence to indicate that coition occurs by the
time the female reaches stage A2 (see p. 199).
It has been shown previously in this paper that
the ova are extruded onto the pleopods during
stage C3, and are subsequently hatched during
stage C4. Approximately 25 days are required
for an average-sized adult female to reach stage
C3 at which time ova would be extruded onto
the pleopods. About 30 days are required for
the incubation period, which is concluded
toward the end of stage C4. Some 10 to 15 days

Biology of Pachygrapsus crassipes — HIATT

165

are then required for the changes undergone
during period D. A minimum total intermolt
interval of 65 to 70 days is therefore required
for the breeding females. Non-ovigerous cap¬
tives of similar size require an interval of but
40 to 50 days. This delayed intermolt interval
of ovigerous females, coupled with the fact
that exuvial incidence as well as spawning fre¬
quency is highest at this season, indicates that
the usual temporal sequence of ecdysis is inter¬
rupted to such an extent that the total number
of expected molts is decreased by one each year
subsequent to maturity.

On the basis of the data set forth in Figure 9
it is possible to calculate the number of inter¬
molt intervals which will occur in crabs from
3.4 millimeters or more in initial width to the
maximal adult size. For example, let us assume
the initial width of a given crab to be 5.5 milli¬
meters. By placing a line along the ordinate of
the 5. 5 -millimeter point, it will be found that
the growth curve will be intersected at a point
which corresponds to 6.6 millimeters on the
abscissa. Then, assuming the latter width to
be the next initial width, the corresponding
post-exuvial width (8.0 millimeters) may be
similarly located. The growth curve, therefore,
enables one to calculate the entire sequence of
molts from the first crab stage to the termina¬
tion of growth. The calculated size increments
based on the curves shown in Figure 9 check
accurately with the size increments of wild
crabs which underwent ecdysis in the field. The
post-exuvial size increments of recently molted
wild crabs in representative size categories pro¬
vide the following data which closely adhere
to the expected size increments as calculated
from the growth curve: small crabs of undeter¬
mined sex increased in size from 4.8 to 5.8
millimeters; male crabs increased from 19.7
to 22.6 millimeters, and from 33.6 to 37.4 milli¬
meters; and female crabs increased from 7.7
to 9.0, 21.0 to 24.0, and 30.0 to 32.4 millimeters.

Calculations made in the manner outlined
above demonstrate that females with an initial

width of 3.4 millimeters undergo 21 additional
molts before attaining maximal size, as con¬
trasted to 18 for males of corresponding initial
width. These data are presented together with
other pertinent data in Table 3. The growth
curve for females, presented in Figure 10, takes
into account the fact that female crabs appar¬
ently undergo a greater total of molts than do
males. The curves, of course, represent an
estimation of the number of molts. Further¬
more, it is apparent from the discussion above
that considerable individual variation may and
does present itself; nevertheless, on a basis
of the recorded observations, the curves indicate
the type of growth that characterizes this species.

Age

The interval between hatching and the attain¬
ment of the adult or true crab stage must be
known in order to present accurately an age-
size scale. It was impossible to determine this
span for P. crassipes because of laboratory
deficiencies. However, Hart (1935) investigated
the larval development of Hemigrapsus nudus
and H. oregonensis, and the intervals given for
these two near relatives probably approximate
the corresponding developmental interval for
P. crassipes. Hart described one prezoeal stage,
five zoeal stages, and one megalopal stage for
both species of Hemigrapsus. Moreover, it was
shown that the larval stages of the congeners
were virtually identical both in structure and
in time of development. Therefore, if the larval
development of P. crassipes is similar to that
of its two near relatives, seven molts would
occur in the interval between the emergence
from the egg and the attainment of the first
crab stage. These data, coupled with those
presented in Table 3, indicate that for a male
to attain the apparent maximal size of 47.0
millimeters ( the largest size recorded for males )
it must successfully undergo about 25 molts;
a female, to attain the maximal size of 44.0
millimeters ( the largest size recorded for
females), must undergo about 28 molts.

166

PACIFIC SCIENCE, Vol. II, July, 1948

TABLE 3

Summary of the Estimated Number of Molts, the Estimated Number of Days in the Inter-
molt Interval between Each Successive Exuviation, and the Size Increments at Each
Exuviation for Both Sexes of P. crassipes from Hatching to the Attainment of Maximum Size.

INTERMOLT

INTERVAL

MOLT

NUMBER

FEMALES

MALES

Number
of Days

Size

Increment*

Number
of Days

Size

Increment*

Larval stagesf— . . . .

35

35

1st to 2nd.... . . .

1

18

3.4- 4.4

18

3.4- 4.4

2nd to 3rd . . .

2

19

4.4- 5.5

19

4.4- 5.5

3rd to 4th . . . . .

3

21

5.5- 6.8

21

5.5- 6.8

4 th to 5 th .

4

22

6.8- 8.2

22

6.8- 8.2

5th to 6th . .

5

23

8.2- 9.7

23

8.2- 9.7

6th to 7 th . . . . .

6

24

9.7-11.4

24

9.7-11.4

7 th to 8 th .

7

25

11.4-13.2

25

11.4-13.2

8th to 9th . . .

8

26

13.2-15.1

27

13.2-15.1

9th to 10th.... .

9

27

15.1-17.2

29

15.1-17.2

10th to 11th . . .

10

27

17.2-19.3

31

17.2-19.3

11th to 12 th . .

11

28

19.3-22.0

33

19.3-22.0

12th to 13th .

12

29

22.0-25.0

35

22.0-25.2

13th to 14th . .

13

30

25.0-27.8

37

25.2-28.2

14th to 15th .

14

31

27.8-30.3

40

28.2-31.3

15 th to 16th .

15

32

30.3-32.7

44

31.3-34.7

16th to 17 th . . . . . . .

16

33

32.7-34.8

45

34.7-38.5

17 th to 18th . . .

17

36

34.8-36.7

47

38.5-42.5

18th to 19th .

18

39

36.7-38.4

50

42.5-46.0

19th to 20th... .

19

40

38.4-40.0

20th to 21st .

20

41

40.0-41.5

21st to 22nd... . . . . . . .

21

45

41.5-42.8

Total . . . . . .

652

652

* The size increments are based upon the data set forth in Figure 9. All measurements represent the greatest carapace width
in millimeters.

t The intervals shown for larval stages are based on those found by Hart (1935).

Hart (1935) found that the time interval
from hatching to the first crab stage in H. nudus
and H. oregonensis spanned 4 to 5 weeks. Each
successive zoeal stage required more time than
the one which preceded it. The present data,
set forth in Table 11, indicate that the inter-
molt interval between the first crab stage and
the second crab stage varies between 14 and 20
days, with a progressive increase in the inter-
molt interval with age. Captive crabs larger
than 7 millimeters in width were reared under
conditions slightly unfavorable to optimum
growth; hence the intermolt interval of captive
specimens spans a somewhat longer period than
that characteristic of wild crabs. The chief
value of the figures obtained from captive ani¬
mals resides in the fact that they are maximal;
consequently, they aid materially in computing
intermolt intervals. Significant assistance was
derived from the information on intermolt

intervals of P. crassipes secured from wild speci¬
mens by Olmstead and Baumberger (1923).
When the intermolt span for captive crabs was
compared with that furnished by the above
investigators for crabs of identical widths, it
was found that the time involved for captive
specimens was increased approximately one-
third. A scale of intermolt intervals has been
synthesized for both males and females and is
presented in Table 3.

The usual method of plotting size-frequency
data, to ascertain the age groups of which a
population is composed, was employed on a
collection of several hundred crabs taken during
a 1-week period in August, and was found to
yield confusing results. Histograms were drawn
for each distribution, and the modal groups
comprising the distributions were superimposed
with a number of normal curves having dif¬
ferent standard deviations and different areas,

Biology of Pachygrapsus crassipes — HlATT

167

in an attempt to identify and separate the homo¬
logous groups in this series of size frequencies.
The method employed was that developed by
O. E. Sette and used by Brock (1943) in his
studies on the Oregon albacore fishery. Although
the plotted data had trimodal characteristics, the
differences in height between the modal and
adjacent intermodal classes were slight and
variable. It was impossible to fit any set of
normal curves to the data.

Factors responsible for the absence of clearly
defined age groups are undoubtedly associated
with (1) the frequent exuvial periodicity for
all age classes; (2) the variability in post-exu-
vial size increments; (3) the extensive breed¬
ing season.

Inasmuch as the number of molts and their
corresponding intermolt intervals are approxi¬
mately known, it was possible to designate
graphically the size-age relationship. Since there
is a period from November to March during
which ecdysis is unlikely to occur, and since
breeding females apparently lose one molt each
year because of the extended ovigerous period,

these modifications were incorporated into the
size-age scale presented in Figure 10. The fre¬
quency with which ovigerous females were col¬
lected during each month of the year shows
that June represents the mid-point in the breed¬
ing season; consequently, the curves for size
and age stem from that period.

Several factors pertinent to growth and age
are exhibited in Figure 10. First, three distinct
age groups are represented, with crabs of both
sexes attaining the supposed C4T stage during
the third year. Measurements of several thou¬
sand individuals collected during August, 1940,
showed that the frequency of large male crabs
fell abruptly at 40 millimeters width with the
largest specimen taken measuring 47.0 milli¬
meters, while the frequency of large female
crabs descended rapidly from the 30-millimeter
size with the largest specimen measuring 44
millimeters. Second, the crabs of both sexes
about 13 millimeters in width may be con¬
sidered to represent maximal size for first-year
individuals; crabs between this size and 30
millimeters represent second-year individuals;

AGE IN DAYS

Fig. 10. Size and age curves for P. crassipes.

168

PACIFIC SCIENCE, Vol. II, July, 1948

and crabs above this width seem to be third- or
perhaps fourth-year specimens and show no
evidence of impending ecdysis. Third, the
curves indicate that size-age dimorphism first
becomes apparent in crabs with a carapace
width between 20 and 25 millimeters. Further,
the data set forth in Figure 1 1 present evidence
to show that this dimorphism occurs just sub¬
sequent to the onset of sexual maturity in
females. These data were secured from 80
ovigerous females collected from August 3 to
10, 1940. In addition to establishing corrobora¬
tive evidence with respect to size dimorphism,
the data presented in Figure 11 show that the
size span for ovigerous females is too extensive
to fall within any single age group, but does
coincide well with the data presented in Figure
10. Fourth, it is apparent that the size diver¬
gence between sexes, after sexual maturity in
females, is associated with a successive propor¬
tional decrease in post-exuvial size increment.
Notwithstanding all the variables attendant
upon the growth of crabs, the life span and
growth characteristics set forth above are, in
general, characteristic of this species.

Comparative information on age investiga¬
tions of other species of Brachyura is particu¬
larly significant because virtually all workers
have followed different avenues of approach,
yet nearly all obtained comparable results. Aside
from the comparatively lengthy life span of 8
to 10 years for Cancer magister (MacKay,
1942), those known are very similar to that
of P. eras sipes. Hay (1905) and Churchill

>0

9

e

16 16 20 22 24 26- 28 30 32 34 36 33 40 42 44

WIDTH IN MM.

Fig. 11. Size frequency of ovigerous P. crassipes
collected during August, 1940.

(1918) found a 3-year life span for Callinectes
sapidus with mating and spawning initiated in
the second year, and Broekhuysen ( 1936, 1942)
discovered a similar span for Carcinides maenas
and Cyclograpsus punctatus.

GENERAL HABITS AND BEHAVIOR
Visual, Chemical, and Tactile Senses

Observations and simple experiments per¬
formed with both captive and wild crabs indi¬
cate that the visual and tactile senses of P. cras¬
sipes are well developed, while the chemical
sense seems to operate on a somewhat lower
functional grade. The complex integration and
mutual cooperation of all three senses are of
such nature that an estimation of the degree of
participation in an observed reaction can only
be arbitrary at best.

Although visual acuteness among the Crus¬
tacea is doubted by many investigators, an
observer of P. crassipes could scarcely fail to
note their alertness. The stalked eyes are capable
of considerable movement. At rest the normal,
undisturbed animal will frequently move the
eyes forward, backward, and laterally. These
movements suggest a critical surveillance of
the environment in an attempt to achieve the
most acute images possible. When frightened,
the animals lower the eyestalks laterally into a
groove for their reception between the orbital
and suborbital areas of the carapace. The faceted
surface of the lowered eye receives protection
from the orbital spine.

It is generally believed that the sharpness of
any image recorded by this type of eye will
depend upon the number of facets which per¬
ceive the object. Inasmuch as the number of
facets engaged by an object varies inversely
as the square of the distance, P. crassipes must
be somewhat myopic. If the number of facets
engaged by the object provides the lone cri¬
terion for visual acuteness, the rapid reaction
of P. crassipes to persons walking at a distance
of several yards is somewhat enigmatic. It
would seem, therefore, that a moving object

Biology of Pachygrapsus crassipes — HlATT

169

stimulating a number of facets in rapid suc¬
cession imparts an impulse to the central ner¬
vous system which results in a more extensive
reaction than that resulting from the perception
of stationary objects. Slight movements within
a few feet of the animals elicit instantaneous
responses; whereas considerable movement at a
distance is required to obtain a response of
similar magnitude. The greatest degree of visual
activity in the faceted eye is achieved not only
at short distances but, in addition, on that area
of the eye which has the least curvature, since
it is obvious that the greatest number of light
rays emanating from an object will strike com¬
paratively more ommatidia if the faceted surface
is flattened. My observations coincided with
the predicted response founded on the fore¬
going principles. Figure 12 is presented to
demonstrate the curvature and extent of the
faceted surface of the eyes of P. crassipes in
several planes of vision. It is seen that the
least faceted surface occurs on the medial por¬
tion of the eye ( D ) ; this is to be expected be¬
cause the opposite eye receives visual stimuli
from the identical angle. The dorsal surface
(A) is strongly convex and slightly faceted.
These data indicate that vision dorsad would
be poor, a fact substantiated on numerous occa¬
sions while observing crabs from a vantage
point directly above. Space does not permit a
detailed presentation of these observations. The
posterior region of the eye (E) is likewise
strongly convex and provided with relatively
few facets. Vision in a posterior direction does
not appear to be acute, but it is difficult to
effect successful observations because the pos¬
terior end of the crab is usually directed toward
a sheltering area in the habitat. It would seem
that the compensatory movements of the eyes
(rotation of the eyestalk to direct highly faceted
areas of the eye to new positions), which will
be discussed below, serve to span the posterior
direction effectively.

The anterior portion (C) of the faceted sur¬
face is much greater in comparative area and
exhibits less convexity than the regions dis¬

cussed above. Vision in the forward direction,
therefore, would be expected to be good. This
supposition has proved valid many times. The
slightest movement of a person at a distance up
to 30 yards elicits immediate response. Com¬
pensatory movements probably increase visual
acuity in the forward direction because the ani¬
mals seem somewhat more responsive to visual
stimuli than one would expect from the number
of facets located in the anterior portion of the
eye. The outer or lateral (B) surface of the
eyestalk is comparatively flat and completely
covered by ommatidia. Observations indicate
that visual acuity is most pronounced in the
lateral direction, a predicted result based upon
an examination of the morphological features
of the eye. Further, it appears highly significant
that the most effective visual surface lies in
the plane of normal locomotion.

The compensatory movements of the eye-
stalks, such as those described for other Crus¬
tacea (Bethe, 1897; Cowles, 1908), occur in
P. crassipes when they are tilted either from
left to right or in an anteroposterior direction.

c

Fig. 12. Comparative amount of curvature and
faceted surface on various aspects of the eye of P.
crassipes. A, dorsal surface; B, outer or lateral sur¬
face; C, anterior surface; D, medial surface; E, pos¬
terior surface.

170

PACIFIC SCIENCE, Vol. II, July, 1948

When the anterior end of the crab is tilted
downward from the horizontal plane, the eye-
stalks are gradually lowered to their normal
reclining position. As the crab is tilted back¬
ward and upward the eyestalks move outward
and come finally to describe a 45° angle with
the sagittal plane of the body; in this position
the eyestalks are farthest removed from the
depressed position. This movement serves to
bring the best visual surface, the lateral area
( B ) , into a position to perceive in a horizontal
forward direction; whereas, when the animal
is held so that the longitudinal axis is parallel
to the horizontal substrate, this lateral faceted
region is directed laterally. Through the em¬
ployment of this rotary compensatory move¬
ment, the crab assumes an oblique position as
it attains the top of a rock with steep sides;
the dactyls of the ambulatory legs of the ad¬
vancing side are anchored on the top edge,
while the legs on the downhill side are placed
just below the top of the rock. Hence, the
crab is oriented to facilitate a survey of the
upper rock surface with the advance eye before
exposing the entire body from above. Par¬
ticularly successful observations of this behavior
pattern are encountered by approaching the
rocky shore without undue disturbance. Crabs,
previously located on the top surface, will
crawl over the edge of a rock and leave visible
to the observer only the dactyls and propodus
of the ambulatory legs of one side, as though
they were hanging from the top of the rock.
By approaching with caution, following the
initial disturbance of the animals on the upper
rock surface, the crabs may be seen clinging to
the edge of the rock in a position to observe
movements on its top surface. There is little
doubt that these compensatory movements con¬
tribute considerably to more successful vision
which is, of course, closely associated with the
general welfare of the animals in their com¬
paratively precarious surroundings.

Repeated attempts were made to induce
compensatory movements of the eyestalks by
moving objects of several colors and sizes in

arcs at varying distances in many planes, but
all met with failure. It is difficult to compre¬
hend this lack of eye rotation inasmuch as it
transcends the supposition that rotation of the
eyes serves to achieve acuteness in perception.
In most vertebrate animals similar experiments
are successful in inducing compensatory rota¬
tion.

Upon tilting a crab to the right from its
normal horizontal position, the left or upper
eye becomes lodged in its groove as if com¬
pletely withdrawn; and the right or lower eye
moves to an erect position almost perpendicular
to the carapace. The reverse behavior is charac¬
teristic when animals are tilted to the left.
Crabs observed on steep rock walls in a normal
sideways orientation display this differential
eyestalk movement; the upper eye is depressed,
directing the anterior surface laterally and the
lateral, well-faceted surface posteriorly. The
lower, erect eyestalk commands a better view
than if it were in its normal position. Its posi¬
tion provides increased posterior vision without
decreasing anterior vision to any appreciable
extent, while the lateral vision remains normal.
The compensatory reaction suggests an adapta¬
tion to increase posterior vision when the ani¬
mal’s back is not to the wall, as it were, while
ascending steep rocks.

Crabs which remain motionless in shallow
pools or near the edges of deeper pools charac¬
teristically have the faceted surface of the eyes
protruding above the water level. Perception
of objects outside the pool would, of course,
be enhanced because the bending of light rays
upon entering the water is thereby obviated.

Rather typical responses toward the preda¬
cious gulls are exhibited by these crabs.
Throughout the observations of crabs in tide
pools and the surrounding rocks, rapid move¬
ments for concealment were frequently noted.
Gulls in flight close at hand were the stimulat¬
ing agent. On several occasions the shadow of a
flying gull was sufficient to elicit a hiding re¬
sponse. Whether or not these responses to
shadows were effected by the actual sight of

Biology of Pachygrapsus crassipes - — Hiatt

171

the gulls, the sudden change in light intensity,
or a combination of both, is of interest. To test
the effect of moving shadow alone, the writer
undertook to cast shadows over the crabs from
a place of concealment. The animals became
startled but rarely concealed themselves in the
near-by crevices. It would seem, therefore, that
perception of the gulls themselves, rather than
the casting of the shadows, was the stimulus
contributing to the concealment response.

Several isolated observations seem to sub¬
stantiate further the keen perception of P. cras¬
sipes. The following excerpts from my field
notes will serve to describe visual acuity in this
species:

The sardine, which was placed near a crevice
into which a crab had withdrawn, drew several
flies, one of which settled at the entrance to
the crevice; the crab moved very slowly toward
the fly, as if to stalk it. When within 3 inches
of the fly, the crab, with a very rapid thrust of
the right cheliped, succeeded in capturing the
fly. A series of quick movements of the chela
followed as if an attempt were being made to
kill the insect; the fly was subsequently con¬
veyed to the mouth and eaten.

This activity occurred just 2 feet below my
observation post which was a collapsible blind
utilized to facilitate close-up observation. Its
construction consisted of a detachable frame
and large-mesh wire netting through which
fronds of algae were woven.

A second observation further substantiates
the visual ability suggested above.

A crab was seen to emerge from a tide pool
and rapidly pursue a rock slater ( Ligia occiden¬
tals Dana) which had ventured close to the
edge of the pool. The crab had apparently per¬
ceived its objective prior to its emergence from
the pool because it moved slowly out of the
pool, and walked cautiously toward the rock
slater. The crab did not alter its stalking pace
until the rock slater took flight. Both prey and
predator ran with extreme rapidity in and out
of crevices in the rock wall before the rock
slater succeeded in eluding the crab.

The constant behavior pattern of P. crassipes
in response to food and materials tossed into

tide pools and near uncovered ledges under
which crabs were concealed has provided the
basis for some simple experiments which were
designed to diagnose partially the relative sig¬
nificance of the three senses involved in the
procurement of food. Before outlining the ex¬
periments and discussing the data derived there¬
from, a preliminary account of the behavior
of this crab in response to food, as recorded in
field observations, will be helpful in setting
forth the objectives of the experiments to follow.

Without exception, providing all the ani¬
mals involved were present, the initial response
to food materials tossed into a middle high tide
pool along the central California coast was
made by sculpins ( Oligocottus maculosus Jor¬
dan). Hermit crabs followed closely behind
the sculpins. Upon rare occasions small P. cras¬
sipes would reach the food before the hermit
crabs. It was frequently observed that food
thrown into a pool failed to attract the atten¬
tion of the crabs unless it was first located by
other species.

The response of P. crassipes to floating food
was rather constant. The following is an excerpt
from my field notes:

A fragment of cantaloupe tossed into a pool
containing P. crassipes within submerged crev¬
ices elicited no response until the breeze had
forced the morsel toward the edge of the pool,
at which time a crab emerged and walked along
the bottom of the pool directly below the food.
As the food reached the water’s edge the crab
climbed toward it, seized it with a cheliped, and
pulled it to the bottom of the pool.

Similar responses were noted regularly. It is
significant that the food was constantly moving,
and that the crabs at no time offered to swim
to the surface to grasp it.

Food tossed near exposed crevices which
sheltered crabs was found soon after it fell.
In one instance a crushed black turban snail
(Tegula funebralis Adams) was tossed on a
rock surface approximately 1 foot from a
sequestered crab. The crab moved out, seized
the turban, and returned to its crevice in 5
seconds. This behavior pattern, because of the

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PACIFIC SCIENCE, Vol. II, July, 1948

rapidity with which the activity occurred, dem¬
onstrates with certainty that visual perception
was employed. Indeed, an identical response
could be induced by tossing pebbles near the
edge of the crevice; the crabs emerged, touched
the pebbles with the advanced ambulatory legs,
but failed to pick them up. Response time
varied from 5 to 30 seconds, and on no occa¬
sion was any substance ignored.

Food lying on the rocks failed to attract the
crab’s attention, providing accidental contact
was not made. One crab reached its crevice
high on the rocks subsequent to the deposition
of an odorous sardine within 3 inches of the
refuge opening. A second crab emerged from
the pool and passed within 6 inches of the
sardine but ignored it completely. The age of
the sardine provided no adverse effect upon
its palatability because it was located and eaten
during the extensive nocturnal wanderings char¬
acteristic of this species.

The retarded response of P. crassipes toward
securing food placed in the tide pools suggests
that this species often depends upon the visual
stimulation provided by feeding activity of other
tide-pool residents. To test this suggestion,
observations were made in a pool which har¬
bored no forms other than P. crassipes. Food
materials were cautiously introduced into the
pool with the aid of a long pole manipulated
from behind the blind. The interval which
elapsed between the introduction of the food
and the initial response toward obtaining it
was indicative of the proclivity of the crabs
toward recognizing its presence. Between 7
and 8 minutes elapsed before emergence of
the smaller crabs from crevices to seek the food.
After discovery, frequent combats occurred.
The larger crabs appeared soon after the smaller
ones had located the food. It would seem that
the movements of the smaller crabs played a
role in stimulating the larger individuals to
seek the food material.

Activity mediated chiefly through chemical
stimulation was manifest on several occasions.
During one observation period several fragments
of abalone ( Haliotis rufescens Swainson) were

carefully placed in one end of a tide pool in
which crab activity was slight. After 7 minutes
the food was removed. Apparently no crabs
had observed its deposition, inasmuch as no
investigational activity ensued. Shortly there¬
after, several small crabs left their crevices and
began what simulated a random search. Shortly
after the emergence of small crabs, larger indi¬
viduals ventured forth. Within a short time
most of the crab population of the pool was
concentrated in the recently baited end. The
high degree of pugnacity demonstrated by larger
crabs seemed to indicate that they were perhaps
strongly stimulated by the meat juices diffusing
through the water of the pool. All crabs meticu¬
lously examined the substrate with the distal
portion of the dactyls. Moreover, all individuals
exhibited extreme complacency when the writer
approached sufficiently to obtain close-up photo¬
graphs of the group. Under normal conditions
the similar approach of a person would instantly
send them deep into their refuge places. It
seemed that the stimulus to search for the
origin of the meat juices superseded the natural
behavior of concealment upon the approach
of possible danger.

The foregoing field observations suggest that
the chemical sense is ineffective in inducing
rapid responses, inasmuch as the animals neg¬
lected to search for the food until it had been
present for several minutes. However, it seems
likewise apparent that upon adequate chemical
stimulation, the chemical sense is of considerable
significance in regulating the activity of the
crabs. It has been suggested above that moving
food, food being eaten by other individuals,
or food tossed sufficiently close to a crab to
stimulate its visual centers elicits an immediate
response. It would, therefore, seem that the
procurance of food, both in and out of water,
is mediated primarily through visual stimuli.

These field data seem contrary to those set
forth by Nagel (1894) and Bethe (1896,
1897 ) for C. maenas. Both authors concur that
this species, in its search for food, is aided by
the chemical sense; Bethe infers that the chem¬
ical stimuli are the principal ones involved, the

Biology of Pachygrapsus crassipes — HlATT

173

eyes contributing little or nothing to food
procurement. Therefore the experiments out¬
lined below were devised to test more specifi¬
cally the relative utilization of the three senses
in the normal habitat.

Experiment 1 : A fragment of abalone viscera was
wrapped in several layers of cheesecloth to make it
invisible to the crabs, yet to provide ample oppor¬
tunity for the diffusion of meat juices through the
water. Care was taken to keep the outside of the
parcel free from meat juice. The parcel was secured
by a string and lowered into the pool with a long
pole. A small crab, originally 8 inches from the
parcel, moved to it in 30 seconds. It picked at the
cloth for 25 seconds and returned to its refuge. Eight
minutes later a second and larger crab visited the
parcel. Shortly thereafter several other large crabs
reached the vicinity of the parcel. Increased activity
followed, climaxed by periodic boxing contests (see
p. 180). One large crab approached the parcel,
turned around, and backed up to it with the fifth
pereiopods extended in a tactile-like manner. After
a few seconds the crab walked away from the parcel
and repeated the backing-up procedure. The parcel
was pulled about the pool by the large crabs, which
alternately fought their competitors and examined
the bait for a period of 20 minutes before its removal
from the pool.

The foregoing experiment was repeated with a
rock substituted for the meat and wrapped as before.
A small crab reached the parcel in 5 seconds, gave it
a cursory examination and soon departed. Another
approached the parcel 70 seconds later, climbed upon
it to examine the surface with the dactyls, but soon
moved away. A third crab arrived shortly thereafter,
pinched the parcel, and left it in 4 seconds. During
the remaining 15-minute interval no other crabs
came to the parcel.

After the parcel containing the stone was removed,
another parcel containing abalone viscera was intro¬
duced. It was significant to note that the first small
crab reached the parcel after 1 minute had elapsed.
This individual pinched the parcel a single time and
immediately departed. Within the first 7 minutes
three crabs reached the parcel but left after a hasty
examination. Shortly thereafter, a large individual
approached, backed up to the parcel, and meticulously
examined it with the dactyls of the fifth pereiopods.
Several more crabs approached, examined the parcel,
and engaged in frequent contests; but they notably
seized each lull in the fighting to examine the parcel
further. Considerable milling about with frequent
tugging at the parcel ensued for the remainder of the
15-minute period.

Experiment 2 : A parcel containing abalone viscera,
similar to that of the preceding experiment, was
lowered upon the rocks bordering a tide pool. Al¬
though the parcel was within the usual range of many
crabs, only two approached it; these abandoned it
after a cursory examination. Other crabs walked close
by, but none attempted to investigate the parcel in a
15 -minute period.

The above procedure was repeated with a rock sub¬
stituted for the meat. The parcel was placed near
several crabs on the rocks but all ignored it. After
3 minutes a small crab approached it, and after 6
minutes a larger individual reached the parcel; how¬
ever, both retreated after a hasty investigation. No
other crabs ventured near the parcel in a 20-minute
interval.

Experiment 3: A pebble smeared with abalone
intestinal chyme was tossed into a pool. In 45 seconds
a small crab seized the pebble in one cheliped and
picked at it with the other, frequently bringing the
latter appendage to the mouth. Within 30 seconds
several crabs emerged and milled about the baited
region. Pugnacity was intense, but the foremost ac¬
tivity centered on a thorough examination of the
substrate with the pereiopodal dactyls and the chelae.

The procedure was repeated, with a clean pebble
substituted for the smeared one. In 10 seconds a small
crab located the pebble but abandoned it almost
immediately. No further activity occurred near the
second pebble for a 15-minute period.

Experiment 4: An abalone fragment was tossed
into a pool. The splash stimulated the crabs to seek
refuge. None responded positively to the food for
5 Vi minutes until a large crab emerged slowly from
a near-by refuge and traveled in the general direction
of the meat. The animal seemed to grope its way
toward the meat in a manner which suggested the
reception of chemical rather than visual stimuli. After
this crab discovered the meat and removed it, several
others congregated at the baited region and milled
about it. The substrate was meticulously examined
with the tips of the chelae which were frequently
elevated and drawn to the mouth, thereby exhibiting
evidence of strong stimulation by the juices in the
water.

The same procedure was repeated but the abalone
meat was first immersed in CS2. After 30 seconds had
elapsed, a crab seized the bait, conveyed it out of the
pool to a position on the rock wall, and devoured
it as readily as normal flesh.

Experiment 5: A fragment of abalone was tossed
onto the rocks within sight of four crabs. The meat
was seized by one individual in 55 seconds.

The same procedure was repeated with a piece of
abalone previously immersed in CS2. It was tossed

174

PACIFIC SCIENCE, Vol. II, July, 1948

onto a rock 6 inches from a crab; the crab seemed
frightened and returned to the water, completely
ignoring the meat. However, 4 minutes later, the
crab returned, seized the meat, and readily devoured it.

Other pieces of abalone saturated with CS2 were
tossed near crabs on a dry ledge. On each occasion
the fragments were eaten in a normal manner. More¬
over, one crab conveyed a fragment of abalone into
the blind employed for these observations, an act
which provided an unusual opportunity to test the
reaction of the crab to odors registered as repugnant
by humans. Carbon bisulphide was actually poured
over this piece of meat as the crab ate it; the crab
at no time exhibited any visible sign of adverse stim¬
ulation.

Experiment 6: The eyes of two large crabs were
covered with a mixture of shellac, lampblack, and
plaster of Paris. Subsequent to the application of this
paint, the crabs would not react to shadows or move¬
ments. One crab was allowed to wander over the
rocks until it remained stationary in a suitable crevice.
A fragment of pungent abalone viscera was held
within 1 centimeter of the oral field for 3 minutes.
No response whatsoever was elicited. No antennal
movement occurred although Bethe (1896, 1897)
noted movement under similar circumstances for C.
maenas\ antennal movement occurs when the identical
procedure is followed with P. crassipes immersed in
water (see below). The meat was immediately
grasped if it was brought into contact with the
chelae. When it touched the dactyl of the second
ambulatory leg, it was thrust under the body to the
chelae and subsequently eaten.

The above experiment suggests that this species
apparently does not react to odors. However, upon
contact of the chelae and dactyls with the meat, the
manipulation of the morsel into eating position was
rapid. Whether or not the fleshy consistency of the
food provided the correct stimulus for this behavior
cannot be irrevocably demonstrated; however, some
clarification was effected in a later experiment.

The identical procedure was repeated with a blinded
crab in an aquarium. Whenever crabs are submerged,
the antennules, antennae, and oral appendages move
continually. It was necessary, therefore, to note the
increase in movement of one or all of these structures
to ascertain whether or not a response was elicited
from an introduced stimulus. A fragment of abalone
intestine was held 4 inches from the posterior end
of the crab. After 15 seconds the antennules, antennae,
and chelipeds moved far more rapidly than normally,
registering stimulation by the food. The water was
changed, and a new piece of meat was held 1 inch
from the posterior end; increased appendicular move¬
ments were noted in 10 seconds. This procedure was
repeated with the meat suspended immediately above
the carpus of the ambulatory legs; a response was

noted in 1 5 seconds. The procedure was again repeated
with the meat suspended 1 inch in front of the oral
area; a response was noted after 45 seconds.

It is apparent that the direction of the respiratory
current is responsible for the differential response to
the food. The respiratory current enters the bran¬
chial chamber laterally and leaves it anteriorly via
a trough, thereby conveying chemical stimuli to the
receptors on the antennules as the current emerges
from the respiratory chamber. These facts would
seem to clarify, in part, the factors underlying the
backward approach to food sought by these crabs,
inasmuch as detection of food via chemical means is
accomplished more rapidly from the lateral or pos¬
terior than from the anterior areas.

Experiment 7: Ten blinded crabs were placed in
individual aquaria; a fragment of art gum eraser was
placed in contact with the dactyl of the second ambu¬
latory leg of each. The second walking leg was em¬
ployed here not only because it is the longest, but
because field observations indicated that it was the
one most frequently utilized by the crabs to test
objects immediately below them. The crabs gathered
the bits of eraser beneath them, grasped the frag¬
ments with the chelae, and conveyed them to the oral
area, whereupon they were immediately rejected.

The above procedure was repeated with a small
clean pebble substituted for the eraser. The pebble
was completely disregarded in all cases; the animals
exhibited a slightly disturbed response and moved
an inch or two away from the pebble.

The experiment was again repeated, with pieces of
liver substituted for the pebbles. The gathering re¬
sponse was similar to that shown the eraser, but the
liver was eaten after it had been conveyed to the
mouth.

Although the above experiments are not con¬
clusive, they do supplement characteristic field
behaviorisms which aid in affixing a partial
evaluation of the comparative utility of the
senses in the procurement of food. It is appar¬
ent that all three senses are employed; more¬
over, the degree of individual participation of
the senses seems to depend entirely upon the
conditions which prevail at any given time.
Vision was definitely relied upon in portions
of all experiments except where the eyes were
rendered opaque. Visual response to food stimuli
seemed apparent early in each experiment, which
implies that movement of the food material
was significant. Instantaneous response to food
materials some distance away can be regarded

Biology of Pachygrapsus crassipes — Hi ATT

175

as depending entirely upon visual stimuli, be¬
cause without exception the diffusion of food
juices through the water required several min¬
utes (Experiments 1, 3, 4, and 6). Furthermore,
it seems, from Experiments 1, 3, 4, and 6, that
the threshold of chemical stimulation for this
species is relatively high because responses
which suggested stimuli of chemical nature were
invariably delayed several minutes after the
introduction into the pool of some food sub¬
stance. In Experiment 6 the juices were visible
in the water about the crab for several seconds
before response was shown.

The tactile sense of this species appears to
be rather highly developed. Except in Experi¬
ment 5, the participation of this sense was
clearly revealed. The apparent prodding by
the chelae and distal podomeres of the remain¬
ing pereiopods discloses that these crabs rely
greatly upon this sense in their food procure¬
ment. Moreover, it is employed equally well
in and out of water. In addition to explaining
the frequently observed backward approach to
food under water on the basis of the orientation
of the respiratory current, the elevation of the
fifth pereiopods to be employed as testing
structures brings forth the supporting role
played by the tactile receptors. It is significant
to note that this backward approach to food
has never been observed out of water, suggest¬
ing that in this type of reaction under sub¬
merged conditions, tactile sensations are sub¬
ordinate to chemical ones. The data derived
in Experiments 6 and 7 provide proof that food
is identified by the dactyls, although the latter
experiment seems to indicate that these struc¬
tures are not selective chemically. Experiment 7
provides evidence that the dactyls do distinguish
texture of substances; and inasmuch as their
chemical reception is slight or wanting, the
major role played by the dactyls in food procure¬
ment may possibly be concerned with texture
differentiation. Food materials may be selected
in this manner because they feel soft and fleshy.
At any rate, the spongy texture of the art gum
eraser was sufficient to release all the reactions
attendant to food manipulation.

Nocturnal foraging, the chief type involved in
food procurement by this species, presents an
entirely different aspect of comparative sensory
participation. Apparently vision is completely
in disuse at this time. Motion near crabs at
night elicits no response; however, if the mov¬
ing object is slightly illuminated, instantaneous
responses occur. Furthermore, the nocturnal
behavior of this species tends to limit the utility
of the chemical sense because the crabs emerge
from the tide pools and forage high on the
rock surfaces. Experiments 2, 5, and 6 indicate
that odors elicit virtually no response; conse¬
quently, the chemical sense is reduced to the
sense of taste alone, which would seem to serve
no significant function in the actual searching
for food.

The tactile sense becomes exceedingly im¬
portant at night in lieu of the non-effective
visual and chemical senses. Inasmuch as the
major constituent of the diet is algae, primarily
of the ulvaceous type, and since the foraging
area is generally adjacent to diurnal refuges,
the tactile sense apparently meets the require¬
ment of food procurement. Fishermen frequently
leave sardines and other bait materials high on
the rocks upon the termination of their activi¬
ties. These food caches are frequently located
during the nocturnal wandering of the crabs,
notwithstanding the fact that this food is often
deposited in supralittoral areas which are sev¬
eral feet higher than the normal upper horizon
of the diurnal range. Frequently, too, these food
caches are unnoticed although crabs often wan¬
der near them. It seems reasonable to assume
that those found were located accidentally, rather
than through the medium of a sense of smell.

Additional information concerning the tac¬
tile sense was derived through observations on
normal and blinded animals. Two crabs, one
with opaqued eyes and one in a normal con¬
dition, were released on a rock IV2 feet from
the edge of a tide pool; the rock sloped gently
to the pool. The normal individual ran rapidly
to the water, submerged, and concealed itself
under a rock. The blinded crab wandered slowly

176

PACIFIC SCIENCE, Vol. II, July, 1948

and aimlessly about the rock surface, going
directly away from the water at the start. As it
walked along, the advancing ambulatory legs
were held out straight from the body when not
employed for support. After 10 minutes of this
slow, groping ambulation, the animal reached
the water’s edge. Upon entering the pool the
crab passed over the algal film at the edge and
immediately proceeded to forage by scraping
it with the tips of the chelae, exactly as normal
crabs would do.

Two additional blinded crabs were released
at the locus mentioned above. One groped about
the roCk, settling momentarily in any shallow,
sun-swept crevice encountered, but finally estab¬
lished itself in a larger crevice 1 hour after its
wanderings had begun. At this time it was but
5V2 feet from the point of release. The second
crab paralleled the activity of the first but soon
became oriented in a direction leading to the
pool. The advancing pereiopods were employed
as tactile structures. After the crab entered the
pool, it descended along a ledge with the fifth
pereiopods held high upon a perpendicular wall
and walked with the remaining three pairs. The
advancing appendages were alternately utilized
for support and tactile organs. The chelae were
elevated and spread at a wide angle, apparently
to provide defense for any exigency. The fore¬
going observations seem to indicate that al¬
though the eyes are the chief means of orienta¬
tion, tactile perception can be substituted al¬
though with considerable forfeiture of efficiency.

Notwithstanding the rather high development
of the tactile sense, this species apparently does
not hear sounds within the human range of per¬
ception. Crabs only 5 feet away from the blind
exhibited no response to noises made by shout¬
ing, clapping hands, or the pounding together
of rocks. Moreover, loud shouting at night
within 2 feet of foraging individuals failed to
elicit a response. Further observations pertain¬
ing to the role played by the various senses are
incidental to the main theme of certain other
aspects of this study and will be set forth where
pertinent.

Diurnal and Nocturnal Distribution

Throughout the diurnal hours, members of
this species tend to remain secluded in crevices,
regardless of their position in or out of tide
pools. If undisturbed, they may wander about
in the tide pools or over short distances on the
rocks. At the first sign of danger they dash
into crevices and often literally fall down the
rocks to conceal themselves in the deep crevices
below.

The movement from the refuges to the higher
rock surfaces begins after dusk at a time when
artificial light is required to observe the crabs.
Many crabs align themselves along the edges
of the tide pools to scrape the algal mat. Others
climb high on the rocks in search of the rich
supply of minute algae which grows within the
splash zone. It is a rare occurrence to find a
crab concealed in a crevice at night unless some
disturbance has stimulated it to seek refuge.
Artificial illumination does not disturb their
foraging habits. However, if motion occurs in
close proximity to them in the presence of faint
illumination, they will move rapidly toward
the protective crevices. Virtually all the collec¬
tions which have provided data for this study
were made during the night, because a true
representation of sizes and intermolt stages in
the population could be obtained only at this
time. The observed night population is many
times greater than that observed throughout
diurnal periods.

This species is not unique in its nocturnal
habits. Bateson (1889) states that most Crus¬
tacea are more active by night than by day,
while Hara (1933) reports that the land crabs,
Varuna literata and Sesarma tetragonum , are
considerably more active at night. Drezwina
(1908), while studying phototropic response
in Pachygrapsus marmoratus and C. maenas,
found that P. marmoratus invariably chose the
darkened end of an aquarium, while C. maenas
preferred the illuminated end.

P. crassipes was found to prefer the darkened
end of an aquarium if the water was constant
in depth throughout the darkened and illumi¬
nated halves. When an aquarium was tilted to

Biology of Packygrapsus crassipes — Hi ATT

177

provide deeper water at one end and none at
the other, this species preferred the deep end
regardless of its darkened or illuminated charac¬
ter. The foregoing data, secured from dozens
of laboratory animals reared over a period of 3
years, suggest that although this species is nega¬
tively phototropic, the hydrotropic responses
are dominant.

Locomotion

Locomotion of Brachyura has been investi¬
gated by List (1897), Bethe (1897), and
Cowles (1908). The initial two workers con¬
fined their studies to those species which are
essentially restricted to an aquatic life and are
not particularly adapted to locomotion on land.
Cowles, on the other hand, limited his observa¬
tions to Ocypoda arenaria, a land crab which
travels with relatively great speed. The present
data for P. crassipes describe locomotion in an
animal which is neither strictly aquatic nor
essentially terrestrial, but which occurs on the
strand in a position between those types pre¬
viously investigated. Bethe (1897) observed
that C. maenas ordinarily travels sideways, but
that to a very limited extent it can move forward
and backward; in any event, he does not concur
with List (1897) that Brachyura, in general,
travel in an oblique direction.

Like C. maenas, P. crassipes normally runs
and walks in a direction perpendicular to the
sagittal plane of the body. It is not unusual,
however, for the crabs to travel in other direc¬
tions. Unlike C. maenas, P. crassipes moves
obliquely sideways with little decrease in speed
and travels directly forward with ease. It has
been previously stated that this species can
move backward when approaching food in a
tide pool or when backing into a refuge. When
speed is mandatory, the movement is invariably
sideways; but when foraging is the chief objec¬
tive, the crabs can and do move in any direc¬
tion. Pugnacious crabs have been observed to
approach each other by walking directly forward
for distances up to 2 feet. The chelae are flexed
in front of the oral area during the approach
to combat.

The sequence of movement of the ambula¬
tory legs is 2, 3, 1, and 4. Often, however, the
third and first legs move concurrently. Varia¬
tions occur in the sequence of movement, but
the order mentioned above is invariably adhered
to during normal walking. It is significant that
the second ambulatory leg, which is the largest,
initiates movement; its length probably permits
it to be utilized both as an ambulatory and
tactile organ.

Crabs taken by surprise when the rocks under
which they are concealed are overturned, resort
to speed and skillful dodging to escape capture.
The ease with which they move in any direc¬
tion is exceedingly advantageous. Under no
circumstances do they feign death, a behavior
pattern characteristic of the closely associated
H. nudus. It is a general hypothesis that crabs
achieve greater agility and speed as they become
adapted to exposed conditions. For example,
members of the land-crab genus Sesarma have
been estimated to run approximately 10 miles
per hour; whereas C. magister, an aquatic spe¬
cies, is exceedingly sluggish on land. A study
of the speed of P. crassipes was considered sig¬
nificant because of the unique transitory posi¬
tion of this crab between the littoral and ter¬
restrial areas. Although an accurate speed test
for P. crassipes is difficult to make, several large
crabs were placed on the starting line of a
course laid out on a flat rock. Each crab was
frightened and caused to run while a stop watch
was employed to record the times involved.
Many resorted to dodging tactics, but the writer
was successful in obtaining some time records
over very short distances. The distances involved
were too short to provide accuracy, but an indi¬
cation of the possible speed of movement was
obtained. The most rapid time clocked was 0.9
second over a 4-foot course, indicating a speed
of approximately 3 miles per hour. For all
practical purposes this speed seemed to coincide
with field estimates of speed. Therefore, P. cras¬
sipes runs approximately a third as rapidly as
Sesarma. Despite the comparative slowness of
P. crassipes in contrast to Sesarma, additional

178

PACIFIC SCIENCE, Vol. II, July, 1948

evidence for the hypothesis set forth above for
brachyurans concerning the correlation of speed
with habitat may be obtained by comparing
the speed of the former with that of its more
sluggish, near relative, H. nudus, located at a
lower level on the strand.

Swimming is rarely undertaken by P. cras-
sipes and is poorly executed because of the lack
of structural adaptation for this method of
locomotion. Infrequently, crabs were observed
to leave the top of a submerged rock by giving
a powerful kick; the ambulatory appendages
then move at a very rapid rate until the animal
attains the opposite bank. At no time has swim¬
ming been successfully undertaken over dis¬
tances in excess of 1 to 2 feet. Movement
through the water is retarded, notwithstanding
the fact that the appendages are moved swiftly.
On one occasion a crab swam forward by mov¬
ing the limbs in an anteroposterior direction
instead of the normal lateral flexing. This
method seemed more efficient than the usual
lateral swimming because the broad surfaces
of the legs provided greater resistance to the
water.

Food and Feeding Habits

The food of P. crassipes consists, in order of
decreasing importance, of (1) live algae, both
matted and frondal, (2) detritus left by reced¬
ing tides and fishermen, and ( 3 ) living littoral
animals.

By far the most important method of food
procurement is the scraping of the minute
algal mat from the bottom and sides of tide
pools, damp crevices, and the tops and sides of
boulders. The excavated tips of the chelae ( Fig.
3) are highly adapted to this method of forag¬
ing. The algae most commonly eaten are listed
on page 14 1.

Nocturnal foraging is preferred and large
numbers of crabs are found at night high on
the rocks in the splash zone where they scrape
the algal film. Nocturnal foraging is not pecu¬
liar to P. crassipes and seems to be prevalent
among Brachyura in general; most land crabs
(Cowles, 1908; Andrews, 1909; Cott, 1929;

Hara, 1933), some spider crabs (Milligan,
1915), and undoubtedly many others are more
active nocturnally. Diurnal feeding, however,
is common with P. crassipes , particularly on
warm days. The crabs do not venture far from
the pools or refuges, and are content to forage
on the algal film growing adjacent to their
places of concealment. When the tide recedes,
they frequently engage in consuming the short
fronds of Ulva which cover the boulders in
many areas of the strand.

The lack of pugnacity among these com¬
paratively belligerent animals during their for¬
aging on live algal food seems to indicate that
intraspecific competition for this type of sub¬
sistence is negligible. It has been mentioned
previously that crabs frequently appear to be
so completely absorbed with foraging activities
that they are unaware of approaching danger
from predators.

Other workers who have recorded observa¬
tions on this species (Hewatt, 1936; Ricketts
and Calvin, 1939) have placed emphasis on
detritus as the principal source of nutriment.
Inasmuch as their observations were divided
among all the littoral forms and perhaps directed
toward these crabs only at infrequent intervals,
it is probable that they were impressed with
this type of food material which seems, from
far more extensive study, to be secondary in
importance. While it is true, as Ricketts and
Calvin (1939) state, that these crabs and the
beach hoppers are the most active scavengers
in this particular ecological association, it is
evident from the information at hand that food
secured from this source contributes but a minor
portion of the total diet. All available carrion
is quickly consumed, but the amount of carrion
deposited on the rocks by the receding tide is
almost negligible; indeed, observations of crabs
eating food of this nature are infrequent.

The least significant food source (Pearse,
1931, to the contrary), but the most interest¬
ing from the point of view of behavior, is other
littoral animals. Variations as to the extent and
type of predacious behavior were disclosed
among crabs in a given locality as well as be-

Biology of Pachygrapsus crassipes — HIATT

179

tween widely separated populations. A single
record of a most interesting behavior pattern
was taken during observations on crabs in a
tide pool at Moss Beach, California. The fol¬
lowing excerpt from my field notes serves to
describe this activity.

A large male walked slowly by a black turban
(Tegula funebralis) in a forward direction; it
stopped beside the turban, quickly side-stepped
laterally and grasped the shell, overturned it,
and seized the turban behind the operculum
with a cheliped before it could withdraw. The
crab held the turban for several minutes, making
repeated attempts to pull it from its shell. After
attempts to extricate the animal met with fail¬
ure, the free cheliped was employed to tear the
flesh behind the operculum. A few seconds
later the crab repeated its stalking behavior on
a hermit crab in a black turban shell, but the
hermit crab withdrew before the crab could
grasp it. The opercular function of the hermit
crabs large cheliped was dynamically illustrated.

Observations made along the Monterey coast
disclosed repeated attempts by P. crassipes to
crush the shells of the littorines which are
present in vast numbers in the high-tide pools.
All attempts met with failure. Recently molted,
partially devoured H. nudus were found in two
tide pools at Monterey. Circumstantial evidence
seemed to incriminate P. crassipes inasmuch as it
was the predominant inhabitant of the pools,
particularly since it is known that it is canni¬
balistic on recently molted individuals.

Indisputable records of cannibalistic behavior
in this species— 17 in the laboratory and 3 in
the field— were secured. The soft parts of the
body are eaten; the pereiopods and branchial
chambers are undisturbed. Without exception,
the ravaged individuals were recently molted
animals, no later than stage Ax. Recently molted
animals in the laboratory aquaria were invaria¬
bly victims, if hard crabs were present in the
same aquarium. It was early discovered that
molt and intermolt data had to be secured from
isolated individuals in the laboratory. The lack
of more frequent records of cannibalism is un¬
doubtedly correlated with nocturnal ecdysis and
the rocky habitats with abundant refuges to

which the crabs retire immediately after exu¬
viation.

Stomach analyses were made on 50 specimens
of this species collected early in the morning.
The crabs were preserved in 10 per cent formalin
and examined shortly thereafter. The major
constituents detected in order of decreasing
quantity were as follows: algal tissues, diatoms,
and striated muscle fragments. The latter item
was found in 17 stomachs; several partially
devoured sardines left adjacent to the pool by
fishermen were undoubtedly the source. Had
the fish been absent, it is doubtful that any
animal food would have been found in the
stomachs. These data serve to substantiate the
field observations presented above. Pearse
(1931) cites food consumption figures of 51
per cent flesh and 49 per cent plant material
for P. crassipes in Japan. If these data are
normal, the food habits of the Japanese repre¬
sentatives differ widely from those on the Amer¬
ican coast.

P. crassipes along the central California coast
may be designated as essentially an herbivore,
ordinarily a grazing herbivore, less commonly a
plant scavenger, while facultatively a carnivore,
chiefly an animal scavenger, and less frequently
a predator. An organism subscribing to such
generalized food habits can easily withstand
periodic deficiencies in one or more food
sources; this, coupled with the fact that the
high littoral zone offers an uncontested food
supply, has been a major factor underlying the
success of this species.

Use of the Chelipeds

The chelae are perhaps the most important
and useful appendages of P. crassipes, and as
such merit special consideration here. The sev¬
eral podomeres articulate at varying angles,
enabling the crab to describe an extensive arc
about itself. Sexual dimorphism with respect
to the chelae is negligible, those of the male
being only slightly larger. Although very few
of the normal activities of this crab are ordina¬
rily undertaken without the use of the chelae,
several crabs which had autotomized the chelae

180

PACIFIC SCIENCE, Vol. II, July, 1948

were observed both in the laboratory and field
to be undertaking the necessary activities with¬
out the use of these appendages. Under these
circumstances the first pair of ambulatory ap¬
pendages was utilized to perform several of
the duties usually identified with the chelae.
The duties of these appendages are varied but
their major accomplishments are food procure¬
ment and protection, with reproductive activi¬
ties associated with them to a lesser extent.

Food procurement is achieved in several ways
with the aid of these structures. The spaded
tips are adapted for scraping the algal mat; the
toothed inner borders of both the dactylus and
finger of the propodus are employed to hold
and tear flesh and algal fronds. Upon discovery,
fleshy food is invariably grasped by the chelae
and conveyed to the oral region. Two methods
of employing the chelae during scavenging were
commonly observed: (1) one cheliped holds
the material while the other conveys bits of
it to the mouth; (2) both chelae hold the food
in juxtaposition to the mouth parts while the
mandibles chew off bits from the periphery
of the food material. When the latter method is
employed on a disc-shaped fragment of flesh or
algal frond, the bites of the mandibles produce
a scalloped pattern on the periphery of the
food item.

It was commonly observed that crabs dragged
their food out of the tide pools to higher
sections of the rocks. Frequently, other crabs
would contest the food and a veritable tug of
war would ensue, each animal grasping the food
with one or both chelae.

In addition to the prehensile use of the
chelae, this species apparently employs them to
aid in food selection. Crabs which slowly walk
over the substrate often drag the chelae and
frequently halt to scrape the substrate with one
or both of these appendages. An examination
of the setae on the propodus and dactylus reveals
the presence of both the long and the short
type of setae, which suggests that both tactile
and chemical stimuli are perceived. At least
one other crab is said to utilize the chelae in
the selection of food (Pearse, 1912). There is

little doubt that the chelae of P. crassipes have
functional tactile receptors, but observations
fail to establish any clue concerning their
capacity to function for chemical perception.

Both chelae are employed with equal facility
in conveying food to the mouth when they are
identical, or nearly so, in length. Observations
were made on five crabs for periods ranging
from 2 to 16 minutes to ascertain the number
of movements of each cheliped per minute. No
consistent preference was shown for either
cheliped by these crabs; and out of a total of
626 movements by all five crabs, 320 were
made by the right cheliped, while 306 were
made by the left cheliped.

The use of either cheliped in securing frag¬
ments of Mytilus on a shell apparently depends
upon the side from which the food material
is most easily taken. One cheliped is employed
to hold the shell, the other is utilized to tear
the flesh and convey it to the mouth. However,
exceptions were apparent in those animals
which had recently regenerated an undersized
cheliped. Invariably, the food was held with
the large cheliped, while the flesh was torn
away and conveyed to the mouth by the diminu¬
tive one, a unilateral type of feeding which is
characteristic of male fiddler crabs (Pearse,
1912; Schwartz and Safir, 1915). The pugnacity
of P. crassipes is shown through the activity of
the chelae. A substantial portion of the diurnal
activity of this species is concerned with fre¬
quent clashes between individuals of the same
and opposite sexes, with combats between males
predominant. If the combatants differ markedly
in size, the larger crab usually displays little
interest in the contest and soon wanders off,
even though he may be hotly pursued by his
smaller antagonist.

During a contest the crabs face each other
and frequently box with flexed chelae by push¬
ing at the antagonists chelae with the flattened
external surfaces. If this activity fails to result
in the retreat of one combatant, the chelae are
extended and rapid thrusts are directed at the
opposing crab. If the contest is between two
males of comparable size, the thrusting of chelae

Biology of Pachygrapsus crassipes — HlATT

181

may continue for several seconds; one of the
crabs usually withdraws from the contest. The
proximity of food accentuates the pugnacity,
and contests occur among all sizes and sexes.
Crabs of this species have never been observed
to lock their chelae on their adversaries in any
manner, although it seems likely that this be¬
havior might occur as it does in the fiddler
crabs (Pearse, 1912). These conflicts, although
frequent, result in little if any dismemberment
of individuals; no casualties have been observed,
although innumerable contests have been wit¬
nessed.

Defense against predacious enemies is accom¬
plished both by flight and by action of the
chelae. The latter behavior is manifest when
the crabs are picked up; the chelae provide
vigorous opposition to predators through their
strong, vise-like grip. In addition to pinching
action, the articulation between podomeres en¬
ables the cheliped to undergo considerable tor¬
sion, contributing greatly to the discomfort of
the predator.

Crabs which have been cornered by an adver¬
sary take a stance designated by Bethe (1897)
as the "Aufbaumreflex.” The animal elevates
itself upon the dactyls; the chelae are raised
and spread widely; and the body proper is held
well above the substrate. This behavior is
primarily a bluff to deter the antagonists, be¬
cause the crab will take flight if an opportunity
for concealment is at hand; however, lacking
that opportunity, the defiant animal will fiercely
thrust the chelae at the attacker.

Chelae seldom participate in the reproduc¬
tive activity of P. crassipes. Throughout copula¬
tion the chelae of both sexes are generally
flexed against the oral field; infrequently, those
of the males assist in grasping the female. No
nuptial activity is assigned to the chelae although
such activity has been suggested for male fiddler
crabs (Alcock, 1892). The sole indication of
a nuptial activity was a slight up-and-down
movement of the flexed cheliped of a male
which became separated from a female during
copulation.

Activity under Varied External Conditions

The diurnal activities of this species in a
tide pool attain a maximum intensity on bright,
sunny days; their movements perceptibly slacken
on dull and cloudy days. To place the measure¬
ment of activity on an objective basis, a record
was made of the number of complete move¬
ments made by the chelae from the substrate to
the mouth while the crabs were scraping the
algal mat. The animals selected for observation
were scraping and feeding on the algal mat
immediately below the surface of the water
near the perimeter of a tide pool. Temperatures
were taken of the water near the crabs. Move¬
ments of the chelae of each crab were recorded
over 4-minute intervals.

These data, together with extensive field
observations on territorial associations, indicate
that activity of the crabs is correlated with the
temperature; activity seemed significantly slack¬
ened below 65° F. and accelerated when tem¬
peratures increased beyond this figure. On foggy
days during which the sun was continually
obscured, the temperature of the tide pools
was generally lower than that of the air and
crab activity within the tide pools was slight.
The rapid rise of tide-pool temperature during
periods when the sun is unobscured tends to
benefit the crab population by providing condi¬
tions under which this species is most active.

The conspicuous inactivity of crabs under
shaded ledges was in unequivocal contrast to
the intense activity which occurred in adjacent
tide pools. On a relatively warm day tem¬
peratures of the outside air, of air under a
ledge in which crabs were concealed, and of
tide-pool water were found to be 60°, 58°, and
84° F., respectively. It would seem, therefore,
that this temperature differential would satis¬
factorily account for the differences in activity
between tide-pool and under-ledge crabs. Obser¬
vations which extended over several consecutive
hours showed that even during the warmer days
of the year the crabs under ledges exhibited only
a minimum of movement.

182

PACIFIC SCIENCE, Vol. II, July, 1948

Tide pools remained warmer than the air for
the first few nocturnal hours, which accounts
partially for the intense activity in the pools
in the early evening. Later in the night, the
crabs wandered out of the tide pools and ledges
to forage areas higher on the strand. The air
temperatures were, at this time, relatively low;
consequently, the foraging animals moved quite
slowly. The foregoing data indicate that tem¬
perature must be added to the factors which
exert a regulatory effect on these crabs.

In addition to temperature changes, the
periodic fluctuation of activity created by the
changing tide must be considered in any dis¬
cussion concerning the behavior of a littoral
species. The subsistence of P. crassipes in the lit¬
toral zone subjects the animal to regular
changes of exposure and submersion, with the
result that the sum total of its activities tends
to be placed on a rhythmical basis. Its elevated
position on the strand has succeeded in remov¬
ing some of the restrictions on activity and in¬
activity imposed upon species located lower in
the littoral area; nevertheless, the tidal influence
was clearly manifested. The relative inactivity
of the crabs during high tide, and the activity
shown throughout the hours of exposure, fur¬
nished evidence of a behavior pattern which
seldom deviated from the standard; indeed, it
tended to be stereotyped. The life of this crab
seems to approach a monotony of repetitions, a
seldom-changing series of actions and reactions.

The Influence of the Ocean on P. crassipes

Drezwina (1908) studied the influence of
the ocean on C. maenas in some detail; she con¬
cluded that the crabs showed a tropic response
to the sea, which she designated "hydrotropism.”
The wind, slope of the land, and amount of
light were demonstrated to have no bearing
on the response which repeatedly occurred in
the same manner without regard to external
conditions. None of the crabs traveled in a
direction opposite to the sea, and the crabs with
painted eyes oriented themselves no differently
from normal ones. Time did not permit a study

of these behaviorisms in great detail for P. cras¬
sipes , but the results of some simple field obser¬
vations tend to confirm the findings of Drez¬
wina with some modification.

Sixty-one crabs were released high on the
beach behind a rocky outcrop which shielded
the sea from the sight of the crabs. The writer
stood 10 feet to seaward from the point of re¬
lease. Many small rocks were strewn about the
area, both in a direction away from the sea
and toward it. Approximately one-half of the
crabs immediately started toward the ocean,
disregarded rock refuges in the vicinity, trav¬
eled around the rocky outcrop, and went down
the inclined beach for several feet before con¬
cealing themselves in ledges situated there. After
a momentary stop, most of them continued to¬
ward the water until damp crevices or small
tide pools were reached. Of the remaining half,
many traveled parallel to the ocean and halted
momentarily at each rock encountered. Although
the water was visible, several crabs traveled
at least 20 feet in a lateral direction before they
moved toward the ocean. The remaining crabs
went away from the sea and took refuge below
rocks higher on the strand, a behavior in notable
contrast to that of C. maenas (Drezwina, 1908).
In some individuals the stimulus for conceal¬
ment apparently overcame the desire to seek
the ocean water. Within an hours time the
crabs high on the beach gradually moved from
rock to rock in a direction toward the sea. At
the close of a 2 -hour observation period, all
had either reached the edge of the water or were
secluded in moist crevices. Although the innate
tendency of all crabs to react in a like manner
was evident, the response was more pronounced
in some individuals than it was in others. It
is barely possible that some of these crabs,
which were collected during the night on their
high foraging area, were representatives of low
tide-pool areas, while others may have chosen
a higher level in the rocks as their usual habitat.
Perhaps concealment in the high, dry crevices
would suffice for the latter crabs, whereas the
former would have a stronger tendency to seek

Biology of Pachygrapsus crassipes — IilATT

183

water. A more thorough investigation of these
phenomena is required to classify the differen¬
tial responses.

To test further the influence of the ocean
on this species, 25 normal crabs and 25 blinded
crabs were released in groups of 5 from a locus
in a shallow depression atop a large rock sur¬
face which was several feet higher than the
splash zone. The ocean bordered the rock on
the outer and lateral sides. The air was vir¬
tually calm, and the day was bright and warm.
The wanderings of the crabs for a period of 10
minutes were recorded (Figs. 13 and 14). The
solid lines represent uphill travel, and the dashed
lines represent downhill travel. The concentric
areas delimit distances of 3, 6, and 12 feet and
serve to indicate the distance traveled by the
crabs. With the exception of the landward side,
the ocean was approximately equidistant from
the point of release.

These data show that one of the patent
behaviorisms of these crabs is to seek a con¬
cealing crevice. The normal crabs immediately
sought the crevices; five gradually attempted to
make their way toward the ocean. The slope of
the rock seemed inconsequential inasmuch as
the animals did not deviate from the general
direction of travel. It is significant that none
of the normal crabs left crevices to travel in a
direction counter to the ocean.

The data for blinded crabs compared favor¬
ably with those secured for normal ones. In

Fig. 13. The movements of normal P. crassipes
within a 10-minute period subsequent to their release
high on the rocks. (See text for details.)

Fig. 14. The movements of blinded P. crassipes
within a 10-minute period subsequent to their release
high on the rocks. (See text for details.)

contrast to normal crabs, Figure 14 reveals
that the blinded ones traveled in all directions
from the locus. Moreover, they traveled in un¬
certain paths, frequently reversing and recross¬
ing. Those which located crevices were content
to remain. Those which walked around to the
top side of the large crevices wandered about
aimlessly and failed to locate a refuge. Crabs of
both groups eventually traveled toward the
ocean; none ventured away from it.

The response of this species toward wave
action is virtually stereotyped. Its behavior
is not that of extreme fear, such as Schwartz
and Safir (1915) suggest for Uca, although it
bears an overt resemblance to such behavior.
The response varied slightly, depending on
whether the animals were located in tide pools
or in rock crevices. Upon the initial indication
of wave action, crabs in a tide pool curtailed
their activity to a minimum. As the wave action
became more intense, the animals concealed
themselves deep in their crevices, and did not
emerge until the tide receded.

Crabs under ledges withdrew at the first splash
of a wave, moved back as far as possible, and
wedged themselves in by pressing the carapace
and appendages against the rock. At no time
were crabs observed in vulnerable locations as
the tide approached their refuges. Crabs which
had climbed up to high, less-protected areas

184

PACIFIC SCIENCE, Vol. II, July, 1948

returned as the tide receded to the deep crevices
below. The foregoing behavior pattern seems
contrary to their previously mentioned penchant
for remaining out of water. Crabs which follow
the receding tide return as the tide ebbs and
begins to rise. It seems apparent that the deep
crevices just below high-tide level offer for
this species the most desirable refuge against
wave action.

Relationship of P. crassipes to Certain Other

Species in the High Littoral Zone

General interrelationships with respect to
the biota of the strand at Monterey Bay, Cali¬
fornia, have been set forth by Hewatt (1936).
The present observations serve to supplement
those recorded by Hewatt for the littoral species
more or less closely associated with P. crassipes.
Although this species is generally found in a
fasciation relatively uncontested by other forms,
its extensive vertical range brings it into inti¬
mate association with several other brach-
yurans which provide potential competition.

The association of P. crassipes and H. nudus
is perhaps the most important because of the
overlapping refuge places in certain regions
throughout much of their geographical ranges.
It has been found that a differential interspecific
tolerance exists in different types of biotopes.
At Monterey both species were commonly found
just below high-tide level, under rocks which
rest on a solid substrate, or on coarse gravel. In
addition, both occurred in tide pools which lie
below high-tide level. Although there was no
interspecific antagonism displayed on the boul¬
der beach, the individuals which occupied the
tide pools frequently engaged in combats for
certain desirable crevices. At no time were the
two species found in juxtaposition. The rela¬
tively larger chelae of H. nudus generally en¬
abled these crabs to drive off P. crassipes of com¬
parable size.

Extensive collecting soon demonstrated that
H. nudus remained in refuges which were lower
on the strand and generally cooler than those of
P. crassipes. Inasmuch as temperature appears
to be a significant factor in the distribution of

littoral animals, the temperature was recorded
in the localities which seemed to contain most
of each of the two species of crabs at midday.
The air temperature during this investigation
was 69.5° F.

Temperatures of 10 rock crevices containing
P. crassipes ranged from 58.6° F. to 61.2° F.,
with a mean temperature of 60.0° F. Tempera¬
tures of the moist sand under 12 rocks harbor¬
ing H. nudus ranged from 57.6° F. to 59.8° F.,
with a mean temperature of 58.0° F. Thus, the
differential of temperature between the rock
crevices and surface sand below rocks harbor¬
ing P. crassipes and H. nudus , respectively, is
slight and probably does not account for the
different locations selected by each crab. Mid¬
day temperatures of eight high tide pools
frequented only by P. crassipes ranged from
68.2° F. to 84.5° F., with a mean temperature
of 72.0° F.; whereas temperatures of eight lower
tide pools containing mostly H. nudus ranged
from 56.5° F. to 59-3° F., with a mean tem¬
perature of 57.5° F. or 14.5° F. lower than the
average of the higher pools. Thus, the tempera¬
tures of tide pools may have considerable influ¬
ence upon the segregation of these species, H.
nudus being as characteristic of the zone of the
rockweeds (2.0 to 4.0 tide level) as P. crassipes
is of the naked zone higher up (over 4.0 tide
level). Although their zonation on the strand
overlaps to some extent, very few of the indi¬
viduals of one species ever have contact with
those of the other.

Foods consumed by H. nudus and P. crassipes
differ sufficiently to reduce to a minimum inter¬
specific competition for sustenance. The former
subsists primarily on detritus and infrequently
on algal fronds; whereas members of the latter
species sustain themselves on scrapings of the
algal film and on detritus. During the intensive
nocturnal feeding, P. crassipes climbs to the tops
and sides of boulders to eat young LJlva fronds,
while H. nudus remains below the boulders,
seldom attempting to climb them. Because of
dissimilar feeding habits, the association of these
species within the same tide .pool is not strictly
competitive.

Biology of Pachygrapsus crassipes — HlATT

Intimate association between P. crassipes and
H. oregonensis was rarely observed along the
outer coast, but was frequently observed in bays
and estuaries. Here, as in the association with
H. nudus, the propinquity of the species was
primarily for purposes of concealment in a
habitat which provided a minimum of refuge
places. In the third biotope (see p. 142) both
species occupied adjacent holes in the muddy
bank. On several occasions individuals of H.
oregonensis were observed to enter holes con¬
cealing P. crassipes. The hasty departure of the
former indicated that the two species rarely, if
ever, concealed themselves within the same
hole. Moreover, the holes which contained P.
crassipes were found to have only a single
occupant; those which concealed H. oregonensis
often sheltered from one to three individuals.

Both P. crassipes and the porcellanid crab
Petrolisthes cinctipes Randall are found under
rocks of the upper and middle tidal zones.
Undoubtedly, the concealing habits of both
species are responsible for their association
inasmuch as no specific, competitive interrela¬
tionship has ever been recorded. There can be
no competition for food because P. cinctipes is
a plankton feeder (MacGinitie, 1937). Associa¬
tions between P. crassipes and several other lit¬
toral types have been discussed elsewhere in
this paper.

Precocious Young

The extraordinary display of activity shown
by young individuals of this species is suffici¬
ently outstanding to merit special attention.
The young of many cursorial and defenseless
ungulate mammals are able to run within a
short time subsequent to birth, have the legs
developed out of all proportion to the body,
and usually exhibit mental precocity to a
marked degree. Similarly, the immature indi¬
viduals of P. crassipes are able to attain com¬
paratively great speed, have the legs relatively
longer than the adults, and likewise exhibit
a high degree of alertness. Although this species
is confined to a lower littoral stratum than are
the true land crabs, which exhibit all these

185

characters on a more highly evolved level
(Pearse, 1912, 1914; Cott, 1929), the long
periods of exposure of P. crassipes seem to have
provided sufficient stimuli for the manifesta¬
tion of those faculties ordinarily associated with
animals in less protected habitats. The eyes
of young crabs seem to be comparatively larger,
and the legs appear to be relatively longer than
those of adult crabs. In order to check these
characters, crabs of representative carapace
lengths (measured from the front to the pos¬
terior border), from the first crab stage to the
largest individual, were selected, and measure¬
ments of the long, second ambulatory leg and
the greatest length of the faceted surface of
the eye were recorded. The length of the eye
was measured along the greatest (longitudinal)
length of the faceted surface and does not in¬
clude the eyestalk and terminal style. From
these measurements ratios were computed be¬
tween the length of these structures and the
carapace. These data are set forth in Table 4.

TABLE 4

Ratios between the Length of the Second
Walking Leg and Length of Faceted Eye-Sur¬
face to Length of the Carapace of Several
Individuals of P. crassipes Ranging from 3.3 to
38.3 Millimeters in Carapace Length. (Meas¬
urements in Mm.)

CARAPACE

SECOND WALK¬
ING LEG

EYE

x

43

8

t

c

•5

6JQ

6

x ’S

1-2 s

o «j

bjn u

X

&

r- X

3-2 8

<-4-c O CX,

swO

1-4

CQ

3

3

«oo

X

J3

MOO

3.7

3.3

5.0

1.82

0.7 6

0.23

9.1

7.6

13.3

1.75

1.3

.17

15.2

13.4

24.2

1.80

1.7

.13

20.5

17.8

30.6

1.71

1.9

.11

24.0

21.0

35.0

1.66

2.1

.10

31.1

25.9

43.6

1.68

2.4

.09

37.6

31.7

50.4

1.58

2.5

.08

45.8

38.3

61.0

1.59

2.8

0.07

It is apparent, by reference to columns 4 and
6 that the length of the second pereiopod and
of the faceted surface of the eye, respectively,
as compared with the length of the carapace,

186

PACIFIC SCIENCE, Vol. II, July, 1948

is relatively greatest in the youngest crabs. Fur¬
ther, the ratio gradually decreases as the size
of the crab increases, reaching the final and
lowest value in the largest crabs.

Despite the fact that the visual perceptive
surface of the eyes of small crabs is compara¬
tively greater than that of larger crabs, we
cannot conclude that their apparent precocity
results from better vision. Indeed, the larger
individuals, which possess considerably flatter
faceted areas that enable them to direct more
ommatidia toward an object, are certain to
perceive objects with greater facility. Undoubt¬
edly the precocious development of the eyes
substantially contributes to the alertness of the
younger individuals but their bolder habits with
respect to potential danger, and their ability
invariably to reach food tossed into a tide pool
before the larger crabs, probably result from
nervousness peculiar to young crabs, which are
merely activated by visual stimuli. It is possible
that their lack of experience and associated
inhibitions contribute to their behavior. The
older crabs seem to proceed with far more
reserve and caution.

The comparative agility of young P. crassipes
and H. nudus or H. oregonensis was well illus¬
trated when collections were made by rapidly
turning over rocks along the shore. The imma¬
ture specimens of P. crassipes were easily segre¬
gated by turning over small rocks and grasping
the most rapidly moving crabs which were
invariably this species. The small, sluggish crabs
under rocks were generally young H. nudus or
H. oregonensis . The nimbleness characteristic
of young P. crassipes is in marked contrast to
the sluggishness of the other two species. Al¬
though the cursorial attainments of these small
crabs are far less effective than those of true
land crabs, young P. crassipes are definitely segre¬
gated behavioristically from the more phleg¬
matic marine forms.

Intraspecific Territoriality

Early observations on the general behavior of
P. crassipes disclosed apparent territorial rela¬
tionships among individuals in tide pools. In¬
vestigators of fossorial land crabs (Pearse,

1912; Cowles, 1908) have shown that these
animals return to their burrows from distant
points on the beach. The behavior was con¬
sidered to represent a homing instinct rather
than a territorial behavior, although Pearse
states that fiddler crabs would contest the pres¬
ence of any crab at the mouths of their burrows.
Studies on territorialism among the Crustacea
in general have been neglected, with the excep¬
tion of terrestrial isopods (Allee, 1926, 1938;
Miller, Ph.D. thesis in the Library of the Uni¬
versity of California) and the littoral isopod
genus Ligia (Miller, op. cit.) which achieve
some degree of social aggregation. No critical
investigations of territorial behavior among the
Brachyura exist; therefore the present study
was undertaken both to satisfy this need in part,
and to learn certain behavior patterns required
for a comprehensive understanding of this
crab. Inasmuch as data will be presented in
detail at a later date, it will suffice to give here
only a brief resume of these findings.

It was not anticipated that territorialism, if
present in this species, would be as well defined
as that found among many vertebrates. The
proportionately enormous crab population, the
relatively small habitable area, and the tre¬
mendous biotic pressure upon littoral species
tend to complicate interspecific associations and
to make these associations obscure and com¬
paratively difficult to study.

It was apparent from a 3 -year period of
extensive observations on this species that true
territorialism with respect to life-sustaining ac¬
tivities as exhibited by many vertebrates does
not exist. However, the refuges are characteris¬
tically defended with brisk determination; each
crab selects the crevice best suited to it and
thenceforth defends it from all intruders. The
success achieved in the defense of an area is
dependent upon the size and pugnacity of the
individual. Two of similar size will fight
viciously for a desirable crevice until the most
virile of .the pair succeeds, a behavior pattern
reminiscent of the "peck order” first described
for birds by Allee (1938). Large crabs appar¬
ently defend a small area immediately adjacent

Biology of Pachygrapsus crassipes — Hi ATT

187

to their refuges, whereas small individuals seem
content to defend only the crevice itself.

On several occasions a crab was found in the
same refuge for 2 successive days, indicating
that some degree of stability is attained. The
smaller crabs tend to remain in restricted areas
and show more evidences of territoriality than
do the larger crabs. However, the underlying
factor here may not be innate territorialism but
rather self-preservation, inasmuch as the crev¬
ices accepted by young crabs are sufficiently
small to exclude the larger individuals which
wander about the pool. The data secured seem
to show that the virility order together with
the number of desirable refuges in a tide pool
combine to regulate the distribution of the
animals.

The gregariousness of grapsoid crabs lower
on the strand (H. nudus and H. oregonensis) in
contrast to the individualism of P. crassipes,
together with the independent existence of fos-
sorial land crabs, may indicate that territorialism
is progressively more highly developed as crabs
become adapted to a terrestrial existence. It is
likely that this trend toward territoriality is
associated with the comparatively spacious
Lehensraum extending landward from the
strand, in contrast to the narrow stratified zones
so characteristic of the littoral area.

Although these crabs live together in large
numbers they neither exhibit co-operation with
one another, nor manifest any tendency toward
such communal existence as that displayed by
many terrestrial arthropods. The writer has
found no evidence that these animals associate
for mutual aid either in foraging for food or
for defensive purposes. Crabs appear to have
their own refuges and are responsible for their
own interests. In this respect they agree with
other crustaceans, for although they possess an
endless variety of structural adaptation suited
to a multitude of habitats and modes of life,
very few have taken advantage of the oppor¬
tunities offered by a communal association
among members of the same species.

DEFENSIVE MUTILATION AND REGENERATION
Defensive Mutilation

The ease with which P. crassipes severs its
pereiopods, together with the high frequency
of collected specimens found with regenerating
appendages, makes it imperative that an investi¬
gation of these phenomena be made to evaluate
their relationship to the general welfare of the
animals. It is generally accepted by investigators
of defensive mutilation that the automatic sev¬
erance of an appendage from the body is a
reflex act. However, not until recently (Wood
and Wood, 1932 ) was the mechanism whereby
this is accomplished fully determined, although
it had been frequently discussed for more than
a century. To comprehend the interpretations
presented in the literature and the observations
made on P. crassipes, it is necessary to clarify
the definitions of the terms which have been
applied to self-mutilation and closely allied
phenomena. Since the term autotomie was intro¬
duced by Fredericq (1883), investigators have
included under it the following closely asso¬
ciated but distinct phenomena:

Autopasy results when an outside agent
is responsible for the severance of an ap¬
pendage at a pre-formed breakage plane
(Pieron, 1907).

Autotilly is the separation of an appendage
at a pre-formed breakage plane with the
assistance of mouth parts, chelae, or other
pereiopods of the animal itself (Wood
and Wood, 1932).

Autotomy refers to the reflex severance of
an appendage without aid from any source
other than from the appendage severed
(Fredericq, 1883).

Autophagy is the act of consuming a part
of the body, usually after severance from
the remainder of the animal (Wood and
Wood, op. cit.).

The failure of early investigators in this
field to distinguish between the foregoing types
of self-mutilation has resulted in the publica-

188

PACIFIC SCIENCE, Vol. II, July, 1948

tion of descriptions of at least five distinct "self-
mutilation mechanisms,” each different and only
two being partly correct (MacCulloch, 1825;
Fredericq, 1882, 1883; Demoor, 1891; Wiren,
1896; and Paul, 1915 b). Space does not permit
a review of the historically significant mech¬
anisms of self-mutilation; the reader is referred
to Wood and Wood {op. cit.) for a synoptic
resume.

A knowledge of the morphological features
of the appendages which are directly involved
with self-amputation is a prerequisite for under¬
standing the mechanism for severance in P. cras-
sipes. Figure 15 serves to illustrate these features
which, with their descriptions, will clarify the
mechanism in this species.

i.

Fig. 15. Autotomy in P. erassipes; A, the pereiopod
in normal position; B, the limb has been elevated
until processes X and Y become contiguous; C, further
levation has resulted in severing the appendage at the
fracture plane. B., basis; CO., coxa; F.P., fracture plane;
I., ischium; X, protuberance on coxa; Y, protuberance
on ischium.

The coxa (Fig. 15, co.) is a short, stout
cylinder articulating with the sternum and
epimeron and moving in an anteroposterior di¬
rection. A process which extends distally on the
posterior side (x) is important in autotomy.

The basis (Fig. 15, B.) is an extremely short
cylinder with a smaller diameter than the coxa.
It articulates with the coxa and moves in a
dorsoventral direction; the wide arthrodial
membrane which connects the coxa and basis
permits extensive movement. The basis, being
narrow, slips in under the dorsal, distal edge of
the coxa during extreme elevation, a condition
essential for autotomy. The distal end of the
basis is marked by a groove which encircles
the appendage and separates the basis from the
ischium, the two podomeres being fused.

Inserting on the anterodorsal edge of the
proximal rim of the basis is the long slender
tendon from which the fibers of the anterior
levator basis (autotomizer muscle) arise. The
muscle has widely distributed origins: from
the anterior surface of the endopleurite, from
the endosternite, and a few fibers from the
epimeron and the dorsal surface of the coxa
(Fig. 16, A.L.B.) .

The ischium (Fig. 15, 1.) is larger and wider
than the basis with which it is immovably fused.
On its dorsoposterior surface a protuberance
(y), which is important in the mechanism of
autotomy, extends medially toward the body.
None of the more distal segments are necessary
for autotomy because it will occur in a normal
fashion when they are removed.

The mechanism of autotomy set forth by
Wood and Wood ( 1932 ) has been corroborated
in all respects by this study on P. erassipes. Their
observations on dead animals have been re¬
peated on this species and were found identical
The rapidity with which the limb is autotomized
makes it impossible to observe the process;
therefore it is imperative that the observations
be performed on a preserved specimen or on a
complete appendage detached from the body
with the autotomizer muscle intact.

To disclose the autotomizer muscle, windows
were cut in the dorsal side of the coxa and

Biology of Pachygrapsus crassipes — HlATT

189

epimeron, and the wide arthrodial membrane
between the coxa and ischium was removed
(Fig. 16). By inserting forceps through the
epimeral window the muscles inserting thereon
could be grasped, and by tugging on each muscle
it was possible to determine which was respon¬
sible for elevating the ischium— -the anterior
levator basis. The posterior levator basis, which
inserts on the posterodorsal edge of the basis,

CO.

Fig. 16. The basal podomeres and autotomizer
muscle of the left fifth pereiopod of P. crassipes. The
integument on the dorsal side of the coxa and the
arthrodial membrane between the coxa and basis have
been removed to show the insertion of the anterior
levator basis. A.L.B., anterior levator basis'. ; BA., basis;
CO., coxa; F.P., fracture plane; IS., ischium; M., merus;
X, cut edge of coxal integument; Y, tendon of anterior
levator basis.

is short and arises from the proximal edge of
the coxa, hence it is not easily grasped through
the epimeral window. This leaves only the
anterior levator basis available for such an
operation. By grasping the tendon (the fibers
themselves tear too easily) with the forceps
and holding a finger on the upper side of the
merus to serve as the resistance, it was possible
to simulate autotilly by pulling on the tendon.
The break occurred at the fracture plane and
proceeded with a minimum of effort. When
the appendage was pulled or twisted without
also pulling on the autotomizer muscle, breaks
occurred at the arthrodial membranes between
the podomeres. The foregoing observations
serve to emphasize two significant points con¬
cerning mutilation: (1) the weakest point in
the leg is not the fracture plane, and (2) the
autotomizer muscle must be stimulated to con¬
tract before self-amputation can occur. Fred-
ericq’s application of a resistance (some force
other than the crab’s own body; e.g., placing a

finger on the dorsal edge of the merus) influ¬
enced him to believe that some external force
was indispensable, and it is the type of amputa¬
tion which he described as autotomie. It has
been shown above that this type of amputation
is more aptly designated autotilly. Later work¬
ers discovered that the appendages could be
severed without any form of external resistance.

Amputated appendages were commonly
found in the aquaria if the water was stagnant
or contaminated with decaying food. Inasmuch
as no external agents were available, the crab
must have shed its appendage or appendages
by autotilly or autotomy. P. crassipes does em¬
ploy autotilly and it is, perhaps, the method
most generally used. Autotilly can be readily
demonstrated by injuring the distal podomeres
(with the exception of the dactylus). The limb
anterior to the mutilated one is moved pos¬
teriorly and over the injured member; the
injured member is straightened and moved
upward while the anterior limb provides the
resistance. The leg breaks at the fracture plane
and autotilly occurs. It is accomplished hastily,
the entire procedure occupying less than 2
seconds in healthy individuals.

Figure 15 illustrates the method whereby
the morphological features of the coxa and
basi-ischium, together with the autotomizer
muscle, are alone responsible for the mechanics
of autotomy. Autotomy may also be demon¬
strated on dead or preserved animals, although
it is difficult to perform. Among living animals
this method is perhaps the least utilized and
has never been observed in this species; how¬
ever, considerable circumstantial evidence set
forth below serves to denote its presence. The
procedure follows: The autotomizer muscle con¬
tracts, forcing the appendage dorsally; the basis,
of less diameter than the coxa, slips inside the
latter structure, allowing protuberance Y on the
ischium to contact protuberance X on the coxa.
The meeting of these two structures provides
the resistance required to sever the limb when
the autotomizer muscle is further contracted.
The fracture originates at the dorsal side and
travels ventrally.

190

PACIFIC SCIENCE, Vol. II, July, 1948

Autotomy is difficult to perform because of
the maximal contraction which the autotomizer
muscle must exert on the rim of the basis in
order to elevate the limb sufficiently high, to¬
gether with the additional tension required to
autotomize the appendage. Proof of the occur¬
rence of this method was secured when two
captive crabs severed all of the legs with the
exception of the chelae and the fifth pair.
Pereiopods 2, 3, and 4 probably were autotil-
lized, but the fifth pair is articulated in a manner
which prohibits the chelae from contacting it.
Three days after the last autotilly, the fifth pair
of appendages of one crab was severed. Five
days after the last autotilly in the second crab,
one of the fifth pair was severed. The fore¬
going behavior suggests two significant points;
namely, (1) that autotomy of the type illus¬
trated in Figure 15 must have occurred, and
(2) that the long interval between autotilly
and autotomy may possibly be attributed to
the poor condition of the animals after the
wholesale severance of most of their appendages.
It is possible that the interval between autotilly
and autotomy was a period of convalescence,
during which the animals regained the vigor
necessary to accomplish this type of mutilation.
Several of the captive crabs severed pereiopods
2, 3, and 4 from one side; but few severed the
fifth pair if those anterior had been cast pre¬
viously.

In the laboratory a unique behaviorism de¬
signed to contribute to autotomy was observed
during ecdysis. A crab which was having con¬
siderable difficulty in withdrawing the ambu¬
latory appendages was observed to sever three
limbs (right pereiopods 2, 3, and 4) by auto¬
tilly. The second pereiopod of the left side
was likewise autotillized, but in an attempt to
sever left pereiopods 3 and 4 (left pereiopod
5 was successfully exsheathed) the animal ele¬
vated the right side of the body until it de¬
scribed an angle of approximately 75° with the
substrate. All podomeres of the left pereiopods
rested on the bottom of the aquarium. In this
manner the crab, in its weakened condition,
brought the processes x and Y (Fig. 15) to¬

gether and effected autotomy.

Autopasy is especially prevalent in this species
because of its scurrying behavior. Crabs will
sever a grasped appendage in a fraction of a
second. To test the number of successive auto-
pasies which an individual crab will voluntarily
undergo when held by the tip of an ambulatory
leg, 20 individuals were collected at random.
Each animal was placed on the substrate in its
normal habitat and a dactyl was held. The first
ambulatory leg grasped was severed almost
immediately by each of the individuals. An
assistant rapidly re-collected the crabs. At no
time did the crabs offer to employ the chelae
in defense. In the second test another leg was
held which was likewise severed; however, more
time elapsed before autopasy ensued; each animal
required from 2 to 10 seconds to complete the
process. The re-collected crabs were put to a
third test. At this time 11 of the 20 specimens
attempted to defend themselves by employing
the chelae; when this defense was frustrated,
16 of the 20 cast the third appendage. The
time varied between 30 seconds and 2 minutes.
After re-securing the animals a fourth leg was
held. Without exception, each crab made vio¬
lent slashes with the chelae. Three crabs severed
the fourth leg in 45 seconds, 2 Vi minutes, and
3 minutes and 10 seconds, respectively. None
of the remaining 17 animals would cast the
fourth leg. All of them became placid and
reluctant to move.

The reluctance of the animal to sever its
chelae was evident. Without exception, when
a cheliped was grasped, the free chela was
immediately used to defend the animal. Whether
this behavior was associated with the ability
of the animals to move the chelae in the lateral
and forward area or to some instinctive behavior
to protect these valuable appendages is not
known. When the free cheliped was success¬
fully parried, autopasy occurred without excep¬
tion. The second cheliped was cast but once in
20 tests.

Some investigators contend that autotomy
cannot occur while the crab is in a soft-shelled
condition; however, in P. crassipes autotomy,

Biology of Pachygrapsus crassipes — HlATT

191

autotilly, and autopasy all occur in crabs in the
A1 stage. It was soon discovered that the limbs
of newly molted individuals could not be
grasped or held in any manner because auto¬
pasy is effected rapidly at this time. Despite
utmost care, autopasy has occurred on several
occasions while newly molted animals were
being measured. Moreover, appendages are
frequently found in aquaria containing recently
molted crabs, which indicates that autotilly is
common prior to complete sclerotization.

Although cannibalism is prevalent when
newly molted crabs are located in an aquarium
with hard-shelled individuals, none of the latter
has been observed to consume portions of a
severed appendage, nor have any severed ap¬
pendages been found from which portions had
been eaten. It seems conclusive, therefore, that
autophagy is lacking in this species. However,
crushed appendages tossed into a tide pool
are readily consumed. The lack of autophagy
may perhaps be partially explained by the fact
that the sclerotized integument precludes con¬
sumption of the potential food; however, it is
strange that the soft appendages of a recently
molted crab are not utilized.

Regeneration

Although sufficient time to pursue an exhaus¬
tive investigation into the subject of regenera¬
tion in P. crassipes was lacking, a cursory survey
of this important activity of these crabs was
made. The present study embraces the phenom¬
enon of regeneration through three rather dis¬
tinct avenues of approach: (1) Meticulous
periodic examinations of several regenerating
appendages were undertaken to disclose the
events which occur during this process; (2)
data were secured on regeneration and inter-
molt stages of a select group of crabs, not to
provide a statistically accurate account of the
regeneration by intermolt stages, but to find a
few salient facts concerning regeneration, which,
when correlated with the information now avail¬
able on the intermolt cycle, might provide a
framework for further investigation of this oft-
occurring phenomenon in this species; and ( 3 )

to establish the frequency with which this
species undertakes regeneration in its normal
environment, the data for regenerating append¬
ages on hundreds of collected specimens were
analyzed.

Historically it is found that regeneration in
decapod crustaceans has drawn the attention of
biologists for more than two centuries. Reaumur
(1712) first described the regeneration of legs
of crabs, lobsters, and crayfish. Later, informa¬
tion on morphological development (Brooks,
1882; Herrick, 1896; Perkins, 1927; Yu, 1932;
and a host of others) was combined with con¬
siderable experimental data chiefly presented by
Zeleny (1905) and Emmel (1907) to formu¬
late our present extensive knowledge of this
phenomenon.

Before presenting the regeneration data for
P. crassipes , a brief synopsis of the internal
morphological transformations which occur
from the instant of pereiopodal mutilation to
the onset of regeneration of the new limb will
be helpful toward an understanding of the
external limb transformations which occur in
this species. Our present histological and cytolo-
gical knowledge of regeneration in decapod
Crustacea stems primarily from macruran and
anomuran types (Reed, 1904; Emmel, 1910;
Paul, 1915 a), with the latter worker contribut¬
ing, in addition, some information on the brach-
yuran group. In general, all three groups follow
the same fundamental pattern and differ only in
detail. The following account is fundamentally
based on regeneration in the hermit crab as
described by Paul (op. cit.).

At the fracture plane the sclerotized strata
of the integument are discontinuous for a part
of the circumference of the leg. The columnar
epithelial cells are greatly enlarged at this point
and possess processes which extend inward,
meeting those of the opposite side. These fibers
mat together to form the diaphragm. The appen¬
dicular artery passes through a foramen, which
it completely fills; but the foramen for the
nerve is a funnel-like prolongation of the dia¬
phragm which fits loosely about the nerve,
thereby affording the venous blood a return

192

PACIFIC SCIENCE, Vol. II, July, 1948

passageway. Paul has demonstrated that upon
severance of the appendage both the artery and
nerve retract from the diaphragm, and blood
extravasates from the ruptured distal end of
the artery. The increased pressure in the haemo-
coelic space forces together the funnel-like flaps
which completely occlude the foramina. There¬
fore this mechanism supplants, in part at least,
the earlier hypothesis (Reed, 1904) that a
blood clot formed over the foramina, thus stop¬
ping the flow of blood. Actually some blood
does pass to the outside of the diaphragm at
autotomy because a layer of clotted blood covers
the developing papilla for several days in P.
crassipes, and has likewise been reported in
other species.

The subsequent papilla formation is accom¬
plished by the proliferation of cells from the
free edges of the columnar epithelium. Emmel
(1910) and Paul (1915^) have shown that
blastematic cells which participate in regenera¬
tion of both the artery and nerve emanate
from the epidermal layer, while the free ends
of the severed artery and nerve contribute but
little in the ensuing regeneration. The initial
structure replaced is the diaphragm, a safety
feature in the welfare of the crab; furthermore,
differentiation within the papilla occurs from
the fracture plane distally, never vice versa.
Within a few days after autotomy, the epithe¬
lial cells form a layer over the stump and begin
to proliferate at the center, initiating formation
of the papilla, which is conspicuous a few days
following autotomy. A significant feature de¬
scribed by both the above authors and sub¬
stantiated by the present investigation is the
lack of sclerotization of the regenerating ap¬
pendage. This phenomenon is undoubtedly
associated with the vast size increment from its
papillary form to its functional manifestation
after ecdysis. Throughout the entire papillary
development the appendage is encased in a
pliable sheath which is cast off during exuvia¬
tion.

The initial information secured concerning
regeneration in P. crassipes was derived from a
study of 25 regenerating appendages on eight

male and two female captive animals, each with
one to five severed pereiopods. These animals,
which were selected at random, ranged from 20.8
to 30.4 millimeters in width. Their intermolt
stage frequencies were: Q, one; early C4, three;
late C4, three; D2, two; and D3, one. Several
pereiopods were severed at ecdysis and thus pro¬
vided an opportunity for study of regenerative
progress from stage Ax. Daily observations
included a check on the intermolt stage com¬
bined with a measurement and description of
developing papillae. When observations were
concluded 60 days later, five of the crabs were
still living, three had succumbed during ecdysis,
and two had died of other causes. Two crabs
had molted and successfully regenerated their
appendages, and several others were in advanced
intermolt stages. Although the study was inter¬
rupted before adequate data concerning the
relationships between regeneration and the
intermolt cycle were secured, sufficient informa¬
tion was collected to clarify, in part, some of
the nebulous data accumulated by pioneer in¬
vestigators who were unaware of the uncom¬
promising association between endysis and re-
generational development. A description of the
morphological features of regeneration in P.
crassipes will be set forth first, followed by a
synoptic account of its association with integu-
mental development.

Three days subsequent to autotomy the blood
clot appeared completely black. The time inter¬
val varied, however, from 2 to 4 days, and
coincided with the latter part of stage A2 or
the early part of stage Bx. This clot was pushed
outward from below by the proliferating epi¬
thelial cells until it was fractured on about the
eighth day (but again the period varied from
6 to 10 days). The papilla was conspicuous
2 or 3 days later. The intermolt stage in which
the papilla fractured the clot varied from B4
to Q. The interval between autopasy and the
growth of the papilla to a length of 1 milli¬
meter required about 11 to 16 days, although
one individual developed to this degree in but
9 days.

Biology of Pachygrapsus crassipes— Hiatt

193

The earliest indication of podomeric differen-
tation was manifest when the papilla reached
approximately 1 millimeter in length. Three de¬
pressions appeared at this time: two longitu¬
dinal, one on each side; and one horizontal,
immediately below the apex. The longitudinal
furrow marks the separation of the merus on
one side and the carpus and propodus on the
other; the horizontal depression denotes the
articular area which separates the carpus from
the propodus. The dactylus became visible a
few days later, appearing as another longitu¬
dinal furrow at the base of the papilla on the
side containing the propodus. These depres¬
sions deepened, and another near the base of
the merus indicated the division between this
podomere and the basi-ischium. Regeneration
in the chelipeds was slightly different from that
in the ambulatory appendages. The papilla of
the former was larger and capitate. Only two
longitudinal furrows were conspicuous laterally,
and the furrow which marks the separation of
the dactyl from the propodus differed from
that found in the ambulatory legs. The dactyl
of the cheliped is in apposition to the extended
finger of the propodus, therefore the furrow is
single, longitudinal, and perpendicular to the
furrows which indicate the junction of the
merus and propodus. The remaining podomeres
developed similarly to those of an ambulatory
appendage.

External differentiation of the podomeres was
complete in approximately 18 to 21 days from
time of severance, by which time the animal
reached the C2 stage. Crab No. 10 attained
this stage in 13 days. The precocious develop¬
ment of the papilla of crab No. 10 is signifi-
cent because autotomy occurred in early C4, and
complete regeneration ensued prior to the first
post-autotomal molt. Of the five C4 crabs from
which limbs were removed, only crab No. 10
accomplished complete regeneration by the
first molt.

Pigmentary deposition in the podomeres
took place in the sequence of podomeric differ¬
entiation; it began on the merus and propodus,

later progressed to the carpus and dactylus, and
reached the basi-ischium last. Pigment was
apparent about 24 to 27 days after severance
of the appendage; crab No. 10 exhibited pig¬
mentation as early as the sixteenth day. At the
onset of pigmentation the papillae varied from
1.5 to 2.0 millimeters in length.

Appendages severed during stage A1 required
between 30 and 33 days to develop papillae 4
millimeters in length; crab No. 10 reached this
stage in 18 days. At this time, the crabs which
underwent ecdysis before papillary development
had reached stage C3 or early C4.

Upon the attainment of stage D4 the papillae
which exhibited complete podomeric differen¬
tiation reached a length of about 7 millimeters.
Of four completely developed pereiopodal pa¬
pillae, three measured 7.1 millimeters and one
measured 7.2 millimeters immediately prior to
ecdysis. During the investigation two crabs
had papillae which grew in an anomalous man¬
ner. Both these papillae were severed at ecdysis
and new, normal papillae were later observed.
It is evident, therefore, that normal regenerative
processes may be modified. However, the ab¬
sence of malformed appendages in wild crabs,
combined with the exuvial autotomy of anoma¬
lous papillae, suggests that aberrantly formed
limbs rarely, if ever, succeed in reaching a func¬
tional condition.

The present study on captive animals, in
addition to extensive collection of wild crabs
with regenerating limbs, has shown that two or
more severed limbs will attain the same degree
of regeneration before ecdysis regardless of when
autotomy occurred before stage C4. However,
the data at hand are insufficient to ascertain
whether or not regeneration is accelerated when
mutilation occurs in an intermolt stage prior
to C4. One would surmise that acceleration
was inevitable, and the information available
strongly suggests this phenomenon.

To obtain information concerning regenera-
tional development and the intermolt cycle of
wild crabs, 203 crabs were collected and exam¬
ined during March, 4941. Of this number, 58
had nearly mature papillae (i.e., papillae with

194

PACIFIC SCIENCE, Vol. II, July, 1948

a length in excess of 4 millimeters were con¬
sidered nearly mature ) . A comparison of mature
papillae and intermolt stages is indicated in
Table 5. It is shown that papillae may closely
approach maturity by the termination of stage
C3, which fact, when combined with laboratory
data, indicates that severance probably occurred
at ecdysis or shortly thereafter. However, most
do not attain maturity until stage C4 is reached.
The laboratory data likewise confirm this ob¬
servation. The diminished representation of
mature papillae in period D is directly pro¬
portional to decreased frequency with which
animals in period D are found in nature.

TABLE 5

Comparison between the Intermolt Stage
and Frequency of Nearly Mature Papillae in
Wild Specimens of P, crassipes.

INTERMOLT

STAGE

NUMBER OF
CRABS
EXAMINED

NUMBER OF
NEARLY MATURE
PAPILLAE

c3 . .

33

3

G .

96

32

Dx . . .

40

13

D2 .

27

8

Da .

7

2

Total .

203

58

The foregoing observations on regenerative
progress and integumental synthesis have dis¬
closed some facts which not only effect a partial
clarification of certain heretofore nebulous con¬
cepts of regeneration, but suggest new lines of
approach toward the solution of pertinent prob¬
lems. These data indicate that within intermolt
stage C4 there is a critical period before which
regeneration of a complete appendage will
occur prior to the molt, and after which com¬
plete regeneration will not occur prior to the
molt. Only one appendage of six severed dur¬
ing the C4 stage in the captive animals com¬
pleted its regeneration before the molt, and
this one was severed during a very early C4
stage. None of the five appendages severed
during period D advanced beyond the blood-
clot stage; whereas all of the 19 appendages
severed in early intermolt stages underwent
complete development; and those crabs which

successfully underwent ecdysis had completely
regenerated limbs. Stage C4, it will be recalled,
marks the conclusion of the synthesis of the
old integument, and all subsequent stages are
concerned with its partial dissolution. It seems
likely, therefore, that any regeneration which
occurs prior to the complete synthesis of the
old integument probably retards that synthesis
until the regenerating member slackens its
development in the latter stages. Meager sub¬
stantiation of this hypothesis was secured from
crab No. 10, which, at the onset of regeneration,
was in an early C4 stage. Complete regeneration
ensued prior to ecdysis, requiring 29 days from
C4 to D4 — a longer period than that required
by all other crabs in the experiment, although
several were larger and would be expected to
require even additional time if other conditions
were equivalent. It seems reasonable to assume
that experiments on regeneration may be
performed with considerable accuracy if the
investigator is cognizant of the transformations
which occur during the intermolt cycle and is
able to ascertain them without error. It is clear
that the inability of former investigators to
diagnose the intermolt period accurately has
clouded the results of most of the experimental
investigations on regeneration in decapod Crus¬
tacea.

Measurements were made to ascertain the
growth of the regenerating limbs on captive
crabs for at least three successive molts. The
graphic analyses of pereiopodal growth (Fig.
17) indicate that three molts are undergone
before regenerating limbs reach the size of the
corresponding limb on the opposite side of the
animal. After the first post-regenerative molt
the limb extends to about three quarters of its
normal size (PI. 2, Fig. 1).

The probability of appendicular regeneration
occurring from some point other than the
fracture plane in P. crassipes was also investi¬
gated. Morgan (1902) demonstrated that this
phenomenon could occur in hermit crabs, not
only from points distal to the fracture plane
but also from regions proximal to it. Although
no controlled experimentation on P. crassipes

Biology of Pachygrapsus crassipes — Hi ATT

195

Fig. 17. Comparative growth of normal and regen¬
erating appendage of P. crassipes at ecdysis. A, growth
of the third pair of pereiopods; B, growth of the
chelae. The percentages indicate the size of the regen¬
erating limb compared with the opposite normal limb
at a certain intermolt interval. L.P., left pereiopod;
R.P., right pereiopod.

was undertaken, sufficient observations on both
captive and wild crabs have been made to de¬
scribe this feature in some detail. With the
exception of the dactyls, regeneration from any
point in the appendage other than the fracture
plane has never been recorded in an examina¬
tion of well over 3,000 crabs in collections and
of countless numbers observed in the field. It
seems probable, therefore, that under normal
conditions this species always severs the limbs
at the fracture plane. The regeneration of por¬
tions of the dactyls is common in nature, and,
because of their utility in diagnosing intermolt
stages (p. 152), several crabs which had regen¬
erating dactyls were available for laboratory
observation over extensive periods. The papilla
of the dactyl grows approximately to its normal
size; but, since the regenerating portion of the
dactyl is invested by an external membrane
similar to that covering the papilla at the
fracture plane, the papilla displays no spines.
The investing membrane is cast at ecdysis and
the new dactyl emerges in its normal form.
In wild crabs the frequency of regenerating
dactyls is highest for the chelae, doubtless the
result of the vigorous activity involving the
chelae which often leads to the fracture of the
distal portion of this podomere. One such

example is shown in Plate 2, Figure 2. It is
significant and distinctly advantageous to the
crab that an injury to the dactyl does not result
in autotilly or autotomy. It was found that
injury to all podomeres except the dactyl stim¬
ulated virtually immediate amputation at the
fracture plane. It has long been known that
the thoracic nerve must be stimulated to effect
both autotilly and autotomy; and since branches
of this nerve do not extend into the dactylus
(Pearson, 1908), injury to this distal podomere
is insufficient to activate the mechanism of am¬
putation.

Of the several thousand crabs handled during
this study, approximately 30 per cent had one
or more regenerating appendages. Such a high
frequency of severed appendages implies a
relatively precarious existence which is reflected
in the ease of amputation and relatively fre¬
quent molts. If, for example, comparisons are
made between P. crassipes and those brachyuran
species residing lower on the strand — or below
low-tide level— with respect to type of habitat,
activity, predation, and capability of self-ampu¬
tation, marked differences are manifest imme¬
diately. Hundreds of crabs ( Cancer magister
Dana) brought into wholesale markets by
fishermen have been examined for regenerating
appendages. Relatively few regenerating append¬
ages were found; indeed, to find one was rather
rare. Autopasy is less easily accomplished in
this cancroid species and much more time is
required for this activity than for the process
involved in P. crassipes. Thus the infrequent
regeneration in C. magister is probably asso¬
ciated with the difficulty of self-amputation,
which in itself may be an adaptation to the
infrequent molts and the comparatively more
placid existence of this crab. Other cancroid
crabs ( C. productus Randall and C. antennarius
Stimpson) which occur in the intertidal zone
were examined similarly and found to exhibit
characteristics comparable to C. magister.

Differences between P. crassipes and the more
sluggish and less-exposed cancroid crabs signifi¬
cantly portray many aspects of the evolutionary

196

PACIFIC SCIENCE, Vol. II, July, 1948

changes within the Brachyura, which are, per¬
haps, the result of intense biotic pressure along
the littoral area. It is certain that the cancroid
crabs mentioned above could not subsist high
in the littoral zone. P. crassipes, on the other
hand, has become adapted to its exposed littoral
horizon by achieving, among other things, the
ability of facile self-amputation and rapid regen¬
eration. These features have undoubtedly con¬
tributed to its success in reaching and establish¬
ing itself in an area rich in food materials,
notwithstanding the fact that it is exposed to
countless dangers.

PREDACEOUS AND PARASITIC ENEMIES
OF P. crassipes

The countless dangers encountered by P. cras¬
sipes include relatively few predators but each
kind occurs in rather high frequency. Among
the vertebrate predators, the gulls (p. 170) are
perhaps the most important. The flight reflex
exhibited by P. crassipes when a gull or its
shadow passes by provides some circumstantial
evidence pointing to the significance of these
birds, although their successful predation on the
crabs has never been noted. Frequently gulls
have been observed dipping suddenly over the
rocks where the crabs reside. The absence of
protective coloration in this species would seem
to make them effective targets for this winged
predator. The numerous crab skeletons found
on wharves below gull perches substantiate
the predatory nature of the gulls with respect
to the shore crabs. Rats which frequent the
littoral area during the night probably prey
upon these nocturnally active crabs, and the
inability of the crabs to detect danger at night
would seem to make them easy victims. This
crab does not escape the extensive littoral pre¬
dation of man. Although the crabs are dis¬
regarded as a food source, small boys and even
adults have frequently been observed in attempts
to extricate the crabs from their crevices.
Though most of the attempts fail, the crabs
are often severely mutilated. To escape capture,
P. crassipes will back deep into a crevice and

elevate the carapace until the rough striae on
the protogastric lobes are pressed tightly against
the rock. It is virtually impossible to extricate
a crab in this position because the striae serve
as an effective resistance against the rock, mak¬
ing the animal practically immovable within the
limits of the strength of the body parts. The
chelae are freed for defense in this position, and
continued molestation will eventually result in
the surrender of these appendages. Usually the
exoskeleton is crushed before the animal can
be withdrawn from its wedged position.

In general, it may be stated that P. crassipes
is not easy prey for vertebrate predators— except
at night — for a number of reasons: (1) Their
alertness when exposed and their agility when
disturbed often enable them to escape from
predators; (2) they possess an exceptional
ability to amputate the appendages that are
grasped; (3) during diurnal hours they tend
to remain in the vicinity of their refuges.

Among the invertebrate predators, the larger
sea anemones ( Bunodactis elegantissima and
Anthopleura xanthogrammica) are known to
consume small crabs. Portions of P. crassipes
have been observed protruding from the stomo-
daea of these species. It is a common occurrence
in the high-tide pools at Monterey to find bits
of P. crassipes integument among a Bunodactis
bed, and feeding experiments conclusively indi¬
cate that anemones will readily utilize the crabs
for food. None of the carapaces of the devoured
crabs taken from anemones measured, or would
have measured, over 20 millimeters in breadth.
Large crabs thrown on a Bunodactis bed invaria¬
bly elude attempts at capture. Smaller crabs
are much less successful.

It is necessary to consider the crabs them¬
selves as potential enemies of each other. Can¬
nibalism is frequent and is a perpetual threat
to recently molted individuals.

The larger animal ectoparasites common to
littoral crabs are exceedingly sparse on the
P. crassipes found along the coast of central
California. No macroscopic ectoparasites were
ever found on collected specimens. A detailed

Biology of Pachygrapsus crassipes — Hi ATT

197

examination was made of 150 individuals col¬
lected at Monterey. These were packed in ice
for 3 hours, and subsequently preserved in 10
per cent formalin. The branchiostegites and
gills were removed; both, plus the epimeral wall,
were washed with a stream of water; the water
and contents were centrifuged, and the material
was searched for parasites exclusive of proto¬
zoans. No parasitic species were encountered.
Only one reference to parasitism on this species
in California occurs (Baker, 1912), and this
reference concerned an undescribed parasitic
isopod located in the branchial cavity. Pearse
(1931) indicates that Japanese specimens of
P. crassipes have a high degree of infestation
with the rhizocephalan, Sacculina. However,
since P. crassipes does not occur abundantly
within the more northerly range of Sacculina
and Peltogaster along the western American
coast, a low incidence of infection might occur
in the area of overlapping ranges, but infected
lined shore crabs have never been found.

REPRODUCTION

Sexual Dimorphism and External Genitalia

Only slight sexual dimorphism occurs in this
species except for the abdomen, which is tri¬
angular in the male and subcircular in mature
females. Sexual variation of the abdomen be¬
comes macroscopically apparent in crabs as
small as 6 millimeters in carapace breadth. In
females a progressive abdominal transformation
ensues from a triangular shape in young crabs
to the attainment of the subcircular form at
maturity.

Extensive sexual differences are noted when
the abdomens are extended. The first and second
abdominal appendages of the male are modified
for use as the intromittent organ; the initial
pair is the larger and has a tubular form,
whereas the second pair is considerably smaller
and is located directly posterior to the tubular
pair. The second pair functions as plungers or
pistons working inside the tubular appendages;
together they accomplish the transfer of the
spermatophores into the genital apertures of the
female. To effect the transportation of sper¬

matophores from the coxal aperture on the fifth
pereiopod to the intromittent organ, a sheath
which arises from the coxa encloses the terminus
of the vas deferens and extends to the base of
the first intromittent appendage. This sheath
functions as a funnel to convey the genital prod¬
ucts into the intromittent organ. The flexed
abdomen covers the coxal opening and sheath.

When the abdomen of a female is extended,
the oviducal apertures on the third thoracic
sternite and four pairs of large abdominal pleo-
pods are apparent (PL 2, Fig. 6). The copu-
latory structures of the male are inserted into
the vulvae during impregnation. Inasmuch as
the intromittent organs of the male are restrict¬
ed to movement in an anteroposterior direction
and the vulvae of the females are immovable
(except perhaps for slight stretching in stage
Ax just after ecdysis), it is evident that a suc¬
cessful spermatophoric transfer could hardly
be achieved between crabs differing greatly in
size. The ova issue through the vulvae and
become adherent to the endopoditic setae of
each pleopod, thereby forming the egg mass
(PI. 2, Fig. 3).

Age and Size at Sexual Maturity

The age and size at sexual maturity were
ascertained by a comparison between the size
range of the total number of ovigerous females
collected on the coast of central California,
together with a microscopic examination of the
ovaries of 50 selected females whose carapaces
ranged in width from 10 to 24 millimeters.
The size range of ovigerous females collected
during the most important spawning period is
presented in Table 6. The most diminutive
ovigerous female examined among wild speci¬
mens measured 16.9 millimeters in breadth.
The records secured from dissections of females
selected for size during the height of the spawn¬
ing season are set forth in Table 7. It is ob¬
vious, from the data provided in Tables 6 and 7,
that sexual maturity in females is achieved when
the carapace breadth measures approximately
15 millimeters at about the eleventh or twelfth
month after hatching.

198

PACIFIC SCIENCE, Vol. II, July, 1948

TABLE 6

Summary of the Size Frequencies of Ovi-

GEROUS P. crassipes COLLECTED DURING THE FOUR

Important Spawning Months.

MONTH

:

NUMBER OF
FEMALES
COLLECTED '

NUMBER OF
OVIGEROUS
FEMALES

RANGE OF

WIDTH

(MM.)

PERCENTAGE OF

OVIGEROUS

FEMALES

May ..........

20

5

28.5-41.5

25

June . .

151

61

18.7-39-2

40

July ............

362

112

17.8-39.7

31

August ......

626

144

16.9-44.0

23

Fifty selected male crabs were examined to
ascertain their size and age at sexual maturity.
The abdomens of the male crabs were extended,
after which the genital sheaths about the vas
deferens were excised and transferred to glass
slides and examined microscopically for the
presence of spermatozoa. A summary of the
information secured is presented in Table 8. A
comparatively precocious sexual maturity is evi¬
dent for males. Adulthood is attained in approxi¬
mately 7 months, at which time the crabs meas¬
ure about 12 millimeters in breadth. Despite
this early maturity, the males do not participate
in reproductive activities until the second year
unless they were hatched within the first 3
months of the year.

Most adult females become ovigerous some¬
time between April and September (Table 6).
However, concrete evidence exists to show that
berried females infrequently occur during the
remaining months. Ricketts and Calvin (1939)
note that berried females may be taken in such
divergent months as February and June. More¬
over, the writer has collected the megalops stage
of this crab at Monterey during March, which
indicates that hatching occurred in the latter
part of January and, inasmuch as incubation
required at least a month, the female involved
became ovigerous in the latter part of Decem¬
ber. Off-season spawning is not uncommon
among the decapod Crustacea. Herrick (1909)
has noted it in the lobster, and Broekhuysen
( 1936, 1941 ) has found it to be characteristic
of two other shore crabs, C. maenas and C.
punctatus.

Copulation and Impregnation

Pre-nuptial pairing or exhibitionism is lack¬
ing in this species, although the former is preva¬
lent among the larger Brachyura (Williamson,
1903; Hay, 1905; Churchill, 1918). Moreover,
the actual onset of copulation has been noted
but once during this study, although several
captive and wild crabs have been found in coitu
less than a minute after they had been observed

TABLE 7

Record of the Age and Ovarian Development of 50 Selected Females of P. crassipes Col¬
lected on June 11, 1941.

BREADTH

(MM.)

NUMBER OF
CRABS

AGE*

(MONTHS)

EGGS+

OVARIAN

DEVELOPMENT

20.0-24.0 . . .

4

14-16

+

eggs large; ovary gravid

18.0-19.9 ................................

4

13

+

eggs large; ovary gravid

17.0-17.9 . . .

6

12

+

eggs medium-sized;
ovary well-developed

16.0-16.9 . . .

5

12

+

eggs medium-sized;
ovary well-developed

15.0-15.9 . . .

7

11

+

5 ovaries developed;

2 undeveloped

14.0-14.9 . . .

7

10

—

ovary undeveloped

13.0-13.9 ................................

6

8-9

— -

ovary undeveloped

12.0-12.9 . . . .

5

7

— -

ovary undeveloped

11.0-11.9 . . .

3

6

—

ovary undeveloped

10.0-10.9 . . .

3

5

—

ovary undeveloped

* Ages were determined from Figure 10, page 167.

t A plus sign (+) indicates that eggs were present; a minus sign ( — ) , that they were absent.

Biology of Pachygrapsus crassipes — HIATT

199

as separated individuals. The actual copulatory
position is, therefore, assumed with rapidity.
The infrequent observations of copulation, when
compared with the vast number of hours of
observation both in the laboratory and field,
would indicate that copulation occurred chiefly
at night or that it persists for a very brief inter¬
val. The present data strongly favor the former
supposition.

TABLE 8

Size and Age at Adulthood (Determined
by Presence of Spermatozoa) of 50 Male P.
crassipes Taken on June 13, 1941.

BREADTH

(MM.)

NUMBER
OF CRABS

AGE*

(months)

SPERMA¬

TOZOA

22.0-26.0

6

15-16

many

20.0-21.9

8

14

many

16.0-19.9

7

11-14

many

14.0-15.9

8

10-11

few to many

12.0-13.9

8

7-10

none to few

10.0-11.9

7

5-6

none to few

8.0-9.9

6

4-5

none

* Ages were determined from Figure 10, page 167.

Several close-up observations of copulating
pairs have disclosed a constant positional pat¬
tern. Contrary to all descriptions of the copu¬
latory act in other Brachyura, in this species
it is the male which lies on its carapace below
the female. The second, third, and fourth perei-
opods of the male are placed between those of
the female, and the dactyls are hooked over
the lateral edge of her carapace. The fifth perei-
opods of the male are looped around the pos¬
terior side of the corresponding appendages of
the female, and the dactyls are hooked at the
posterolateral edges of her carapace. The am¬
bulatory appendages of the male, therefore,
serve to grasp the female. The chelae are gen¬
erally flexed in their normal folded position;
however, at intervals those of the male are
employed to grasp the female, and occasionally
are utilized for protection from the thrusts of
the female’s chelae directed toward the male’s
oral area. The abdomen of the female com¬
pletely covers that of the male, that of the latter
being below the abdomen of the female. In

this position the ventral surfaces of each mem¬
ber are contiguous, and the action of the penes
is difficult to observe.

Although difficult to follow from above or
below, impregnation was observed upon two
occasions from the side. The abdomen of the
male moved slightly forward and back with the
penes inserted only a short distance into the
vulvae.

Although the infrequent observations of
copulation in this animal suggest brevity, the
data secured for P. crassipes indicate extreme
variability and comparatively long duration.
In one field observation, in which impregnation
appeared successful, the pair were in coitu for 45
minutes. Upon another occasion captive crabs
were observed in copulation for 6 minutes (the
onset was unnoticed) prior to separation. After
the separation the male partially flexed and
extended its abdomen arhythmically and, when¬
ever the male approached its mate, the abdomen
oscillated with increased tempo. In addition,
this male repeatedly elevated and depressed
its right cheliped. It seemed apparent that im¬
pregnation had not been completed. A copula¬
tory act by another captive pair occurred and
lasted for 15 minutes. When the crabs sep¬
arated, they stood with their flexed chelae con¬
tiguous and the abdomens of both moved brisk¬
ly; however, copulation was not resumed. Copu¬
lation by a third pair lasted for 20 minutes.
The animals frequently moved about the aqua¬
rium while the female supported the suspended
male. Another pair were in coitu for 5 minutes
prior to separation. Several hours later the pair
was again observed in copulation which per¬
sisted 4 minutes. It is highly significant to note
that all the females mentioned above, com¬
parable to most accounts in the literature, were
in stage Alt while the males were in C3 or
above. On the other hand, Broekhuysen (1941)
indicated that copulation in C. punctatus
occurred only between hard-shelled crabs.

Some anomalous copulatory behavior observed
both in captive and wild crabs aided in apprais-

200

PACIFIC SCIENCE, Vol. II, July, 1948

ing the mating phenomenon. A hard-shelled
male was placed in an aquarium with a newly
molted female and her exuvia. Twenty minutes
later the male grasped the exuvia with the
chelipeds and turned over on its back in the
normal copulatory position. Throughout the
struggle to arrange the exuvia in the correct
position, the abdomen of the male oscillated
rapidly back and forth. This abortive struggle
persisted for 5 minutes after which the male
righted itself and moved away. Although the
male was willing to copulate, a fact deduced
from its behavior with the exuvia, no attempt
was made to engage the recently molted female.
The endopoditic setae of the exuvia above were
examined and found to contain empty egg cap¬
sules (PL 2, Fig. 5), which indicated that the
non-copulating female described above had been
ovigerous during the preceding intermolt inter¬
val. On another occasion an attempt at copula¬
tion was observed between a pair of crabs under
a ledge high above the adjacent tide pool. The
animals separated 5 minutes after they were
discovered. The female was collected and found
to be ovigerous. The foregoing anomalous be¬
havior patterns seem to indicate that the males
exhibit but slight discrimination toward their
mating partners.

Copulation in this species occurs at least
once yearly and is suspected to take place twice
annually in some crabs. Recently molted females
whose exuviae bore empty egg capsules on the
endopoditic setae invariably exhibited empty
spermathecae; subsequent impregnation, there¬
fore, would be required during the following
breeding season, or later in the same breeding
season, providing a second batch of eggs were
extruded. In many other Brachyura possessing
far more extended intermolt intervals, it has
been found that one impregnation suffices for
the fertilization of the eggs of several egg
batches which are expelled within a single
intermolt interval (Williamson, 1903; Churchill,
1918; Broekhuysen, 1936, 1941).

Extrusion and Incubation of Ova

Although the interval between impregnation
and spawning in P. crassipes has not been ascer¬
tained, its direct association with the intermolt
cycle enables one to estimate the interval with
considerable accuracy. Inasmuch as eggs are
never extruded prior to stage C3, the interval
between impregnation and extrusion of eggs
will depend directly upon the size of the female
and will vary from approximately 16 days for
small-sized, mature females to 25 days for large
females.

Fertilized ova are expelled from the body of
the female through the vulvae located on the
thoracic sternum. They become attached to the
fine endopoditic setae of the four pairs of pleo-
pods located on the second to fifth abdominal
metameres. These pleopodal endopodites are
clothed for the most part with long, fine setae
which are simple throughout, with the excep¬
tion of a few which branch at the base. With
the exception of the proximal segment, these
setae arise in distinct bundles from each endo¬
poditic segment (Fig. 18). The tufts arise only
on the medial, lateral, and posterior surfaces of
the endopoditic segments; the anterior surface
is naked.

The general appearance of the egg mass has
suggested its usual designation, the "sponge.”
Eight tufts or clumps of eggs are present, which
correspond to the eight abdominal endopodites.
These tufts are so crowded with ova that they
become contiguous and appear as one mass. The
sponge of a female measuring 32.0 millimeters
in breadth is a rounded mass approximately 12
millimeters long by 40 millimeters wide by 12
millimeters deep (PL 2, Fig. 3). The eggs
appear black to the unaided eye, but when
viewed microscopically with transmitted light
they are deep brown. The abdomen is forced
back by the sponge until it describes a 60°
angle with the sternal surface of the thorax and
is curved behind the egg mass. A basket-like
cavity which shields the egg mass is formed by
plumose setae of the pleopodal exopodites (Fig.
18) and by the lateral margin of the abdominal
tergites.

Biology of Pachygrapsus crassipes — HlATT

201

Fig. 18. Front view of the second abdominal pleo-
pod of P. crassipes from right side. END., endopodite;
EX., exopodite; PROT., protopodite.

The number of eggs extruded at one time
seems to vary directly with the size of the crab.
To estimate the number of ova contained in a
sponge, an ovigerous female 36.5 millimeters
in carapace breadth was selected. Each of the
eight endopodites which bore the sponge was
removed and compared. All contained approxi¬
mately the same number of eggs; consequently
the eggs adherent to one endopodite were
counted and the sum multiplied by eight, which
yielded a product of 48,604. Although this
figure is an estimate, it can be reasonably
assumed that the sponge of an average-sized
female will contain approximately 50,000 ova.
This number is considerably less than that esti¬
mated for Callinectes (Smith, 1885; Paulmier,
1901; Churchill, 1918) and slightly less than
Williamson’s (1903) estimate for C. pagurus.
No information concerning this subject for
crabs of a size comparable to P. crassipes is

recorded. Inasmuch as the size of ova differs
but slightly in all the crabs noted above, it
seems probable that the enormous number of
eggs which comprise the egg mass of the larger
Brachyura is correlative with the size of the
animals.

Excellent opportunities for critical study on
the incubation of the ova were presented by
captive females which expelled their ova in
aquaria. Observations were made at 3 -day inter¬
vals throughout the entire incubation period.
A few eggs were removed from the setae,
examined grossly, and measured. Inasmuch as
these data were approximately uniform for four
females under observation, they were averaged
and are presented in summary in Table 9.

The foregoing data show that the incubation
period was approximately 29 days. Other ovi¬
gerous females under observation carried eggs
from 26 to 31 days. The significance of this
incubation interval to the intermolt cycle has
been discussed on page 153. Although a uni¬
formity in size of eggs was apparent through¬
out development, it was found that when first
expelled, the ova were misshapen and abnor¬
mally lengthened. Immediately after expulsion,
the eggs actually shortened. It was further evi¬
dent that during incubation the ova lengthened
proportionately more than they broadened. This
differential growth is undoubtedly a manifesta¬
tion of the extensive elongation of the larval
crab in an anteroposterior direction.

All of the captive ovigerous females exhibited
irregular abdominal movements. These move¬
ments, which were accomplished by moving the
abdomen backward and forward in jerky
arhythmic beats, probably served to aid in main¬
taining efficient aeration of all the eggs in the
mass. The wafting motion separates the eggs
and thus permits the water to circulate among
them. It is virtually impossible for water to
circulate in the densely packed, stationary
sponge. Ovigerous females in tide pools like¬
wise wafted the abdomen, signifying that the
movement is not merely an adjustment to tepid
and less-aerated, laboratory sea water. The rate
and periodicity of this wafting procedure were

202

PACIFIC SCIENCE, Vol. II, July, 1948

TABLE 9

Summary of the Incubation Time, Size, and Gross Development of the Ova of Captive P.
crassipes during June, 1940. (Mean Aquarium Water Temperature, 16.2° C)

WIDTH

(MM.)

LENGTH

(MM.)

DAYS

AFTER

EXPULSION

GROSS DESCRIPTION OF DEVELOPMENT

285

320

0

uniformly dark brown; many zygotes, others in 4-8-cell stage, all
blastomeres large; no perivitelline membrane apparent

286

311

3

uniformly dark brown; blastomeres filling capsule; homogeneous;
no perivitelline membrane apparent

288

310

5

uniformly dark brown; blastomeres smaller and more numerous;
slight withdrawal of blastomeres from one section of the capsule
reveals perivitelline space

291

314

8

uniformly dark brown; blastomeres very numerous; further asym¬
metry in blastomeric bulk reveals greater perivitelline space

294

316

10

uniformly dark brown; differentiation progressing; no pigment
deposits present; blastomeres occupy about 0.6 of egg capsule

298

328

13

differentiation advanced; no pigment deposits; yolk cells in one part
of cell mass only

312

351

18

highly differentiated; heart not beating

320

358

20

highly differentiated; heart beating; structures plainly visible; few
ova on bottom of aquarium

324

368

24

29

no ova hatched; heart beat stronger, irregular (approximately 82
beats per minute)

all ova on bottom of aquarium; many hatched

highly variable and apparently had no rela¬
tionship to the degree of embryological devel¬
opment.

Both captive and wild ovigerous females were
noted to thrust the chelae into the egg mass
frequently, and to move the ova about in a
manner which resembled a preening behavior.
When the chelae were inserted into the sponge,
the abdomen oscillated violently. Frequently,
material which appeared to be ova was con¬
veyed to the mouth. The writer believes that
this preening or cleansing behavior is not asso¬
ciated with actual ingestion of the eggs, inas¬
much as they thickly covered the bottom of the
aquaria subsequent to this activity. It is pos¬
sible that accidental plucking of the eggs resulted
from attempts to remove foreign particles from
the egg mass. Most of the ova observed on the
bottoms of the aquaria after this cleansing
activity had a short portion of the funicle
attached, indicating that they had been severed
from the setae. Frequently, broken setae with
several attached ova were found. The severed
eggs were invariably viable regardless of the
developmental level.

The behavior of wild, ovigerous females with
respect to submersion proved somewhat un¬
usual and contrary to expectation. They fre¬
quently wandered up a rock surface and
remained motionless in the direct sun for
lengthy periods — 1 hour and 15 minutes on one
occasion, and 5 minutes less than that on an¬
other. Both these females were observed on
the same day during which the air temperature
was 72° F. It was further noted that in collec¬
tions, the proportion of females increased dur¬
ing the breeding season, a fact which indicates
that the ovigerous condition does not necessi¬
tate a sequestered existence. It seems likely,
therefore, that the embryos demand no more
than a minimum of submersion; further, the
proportionately greater number of berried
females found on the rocks than in tide pools
may be associated with some biological require¬
ment such as increased warmth to permit the
maximum tempo of embryonic development.

The expulsion of more than one batch of
eggs a year has been recorded for several of
the Brachyura (Churchill, 1918; Broekhuysen,
1936, 1941). Although no direct evidence of a

Biology of Pachygrapsus crassipes- — Hi ATI

203

second sponge within a single breeding season
is available for P. crassipes, certain circumstan¬
tial evidence is sufficiently significant to remove
the occurrence from doubtful rank. First, exam¬
inations were made of the ovarian development
of berried crabs; and second, an examination
of the ovarian development of recently molted
female crabs, which might have been berried
during the previous intermolt interval, was
undertaken. Information on previous egg-bear¬
ing of recently molted females was secured
through an examination of the endopoditic
setae of the exuviae. The data concerning the
ovarian development of ovigerous females are
summarized in Table 10.

The gravid ovaries of many berried females
during the height of the breeding season con¬
tribute significant evidence to the support of
the writer’s supposition that a second batch of
eggs would be extruded within the 3 to 4 months
remaining in the breeding season. However,
it is to be noted that a few of the ovigerous
females which, at the time of collection, had
carried the expelled eggs about halfway through
the incubation period exhibit very undeveloped
ovaries. It is probable that these females would
not extrude a second sponge in the time which
remained in the spawning season.

Nine females in stage A2 and their exuviae
were collected in June, 1941, for an examina¬
tion of their present ovarian development and
of the endopoditic setae of the exuvia of each,

to ascertain whether or not they had been ovi¬
gerous during the preceding intermolt interval.
Only one had been ovigerous in the preceding
intermolt interval and the ovary in this newly
molted crab was ripe and gravid. Six of the
others had small, undeveloped ovaries; two had
ovaries which were nearly ripe, with ova of
medium size. The outstanding feature of the
data presented in Table 10, when associated
with the data set forth immediately above, is
the disclosure that an entire intermolt period
may intervene between periods of sponge pro¬
duction. Inasmuch as the periods of ecdysis in
this species are relatively frequent, it is not
unreasonable that such a phenomenon should
occur within the extensive breeding season.
Exceptions to this behavior occur; e.g., it seems
certain that the animal which contained the
gravid ovary and which had a record of egg¬
bearing during the preceding intermolt interval
would have become ovigerous for the second
time in consecutive intermolt intervals. Whether
or not the remaining crabs had been ovigerous
earlier in the spawning season is conjectural.
Some of the larger specimens probably had been;
whereas the smaller individuals which con¬
tained, for the most part, relatively undeveloped
ovaries probably would not have become gravid
until the latter part of the spawning season.
It seems probable, therefore, that the periodicity
of gonadal development varies, not only between
individuals, but within a single individual in

TABLE 10

The Size, Intermolt Stage, and Ovarian Development of Ovigerous Females of P. crassipes
Collected during June, 1941.

CARAPACE

BREADTH

(MM.)

INTERMOLT

STAGE

CONDITION OF
INCUBATING EGGS

OVARIAN DEVELOPMENT

27.0

G

eyes visible

ripe, gravid

32.4

G

yolky blastomeres

ripe, gravid

31.9

G

blastomeres to one side

young, undeveloped

26.7

G

highly differentiated

ripe, gravid

30.3

G

highly differentiated

nearly ripe, gravid

26.5

G

eyes visible

ripe, gravid

36.0

G

yolky blastomeres

very young, undeveloped

26.4

G

highly differentiated

very young, undeveloped

24.0

G

heart beating

young, some development

32.2

G

undifferentiated

ripe, gravid

204

PACIFIC SCIENCE, Vol. II, July, 1948

different years. Further, the evidence at hand
shows with virtual certainty that some indi¬
viduals expel two batches of eggs within a single
breeding season. The few crabs which mature
a second batch of eggs near the close of the
regular breeding season are probably the ani¬
mals which participate in the off-season spawn¬
ing during the winter months.

Hatching and Subsequent Growth

The multitude of empty egg capsules which
adhere to each endopoditic seta of female exu¬
viae indicates that hatching occurs while the
eggs are adherent to the female. Ovigerous
females in the captive environment of the lab¬
oratory had plucked most of the sponge prior
to complete embryonic development. At the
termination of the normal incubation interval,
the floor of the aquaria was often covered with
loose, viable ova. Near hatching time the ex¬
ceedingly high mortality of these ova which
had been removed by the female’s chelae seems
to indicate that the movement of the abdomen
and the setae by crabs in the wild environment
may actually aid in the hatching process of the
prezoea from the enveloping capsule.

Hatching was observed microscopically. The
investing membrane split about halfway around
its periphery above the dorsal surface of the
embryo; the prezoeae emerged dorsal side first
in curled balls and soon began to move the
appendages and to flex the abdomen. The pre¬
zoeae hatched in the laboratory were non-
motile. Most of the prezoeae underwent ecdysis
the first day; others molted during the second
day. The first zoeae were likewise immotile,
and no subsequent larval development was
obtained because of inadequate facilities. For
an account of the zoeal stages of closely allied
species, the reader is referred to Hart (1935).

The megalops stage of P. crassipes has been
figured in outline fashion in Rathbun (1923,
PI. 34, Figs. 1 and 2 ) and in Johnson and Snook
(1935, Fig. 305), but an adequate description
has heretofore not been published. On March
8, 1941, several megalops of this species were
collected at Carmel, California. Two of these

were subsequently reared in the laboratory until
the sixth crab stage, thereby definitely disclos¬
ing their identity. The collections were made
by gently picking up rocks submerged in tide
pools at a medium-high tide level (4.0 feet)
and sweeping the surfaces with a soft brush
while the rocks were submerged in a tub of
sea water. Subsequently, the water was strained
through a plankton net of coarse mesh. The
transparency of specimens of the megalops
stage makes it impractical to search for them
on the rocks. The only clearly visible struc¬
tures are the few chromatophores and the gastric
mill. The megalopa crawl and swim readily.
Swimming is accomplished by rapid abdominal
movements similar to those of many macrurous
species. The dactyli are furnished with long
hook-like spines to facilitate the grasping of
rock and algal surfaces.

The following account, together with a photo¬
graph of the megalops of P. crassipes (PL 2,
Fig. 4), will fulfill the need for a description
of this phase of the life history: length of cara¬
pace, 5.6 millimeters; width of carapace, 2.7
millimeters; almost transparent, slightly yellow¬
ish in color; scattered black chromatophores on
the eyestalks, on the carapace above the gastric
region, and around the intestine; three chro¬
matophores on each merus of the ambulatory
legs, two on each coxa, two on each propodus,
and one on each dactyl; front wide, with center
turned down to form the rostrum; chelae well
developed; swimmerets broad and flattened with
long plumose setae.

The megalops of P. crassipes may be distin¬
guished from that of H. nudus and H. oregonen -
sis by the greater size- — nearly twice as broad
and long as H. nudus and twice or more as
broad and long as H. oregonensis. In addition,
H. oregonensis has no plumose setae on the
smoothly rounded posterior margin of the telson,
whereas P. crassipes has two long, median setae
and several shorter ones. Setae similar to those
described above for P. crassipes are found on the
telson of H. nudus; therefore, distinction be¬
tween these two species must be made by size
or by characters not mentioned here.

Biology of Pachygrapsus crassipes — HIATT

205

TABLE 11

Summary of the Gross Growth Characteristics of Two Captive Male Specimens of P. crassipes
Reared from the Megalops Stage.

STAGE

CARAPACE WIDTH
(MM.)

PRIOR TO ECDYSIS

CARAPACE WIDTH
(MM.)

AFTER ECDYSIS

SIZE INCREMENT
IN MM.

PERCENTAGE
SIZE INCREMENT

DAYS BE¬
TWEEN MOLTS

No. 1

No. 2

No. 1

No. 2

No. 1

No. 2

No. 1

No. 2

No. 1

No. 2

First crab.......—..

3.7

3.6

?

?

Second crab........

37

3~6

4.5

4.4

65

6*8

21.7

22.2

14

20

Third crab..........

4.5

4.4

5.5

5.3

1.0

0.9

22.2

20.4

14

17

Fourth crab ........

5.5

5.3

6.4

6.1

0.9

0.8

16.4

15.1

20

16

Fifth crab . .

6.4

6.1

7.9

7.2

1.5

1.1

23.4

18.0

25

19

Sixth crab .

7.9

7.2

10.1

8.4

2.2

1.2

27.8

16.7

26

31

Totals .

6.7

4.8

111.5

92.4

99

103

The first crab stage of P. crassipes is likewise
much larger than that of either H. nudus or
H. oregonensis. It is nearly twice the width and
length of H. nudus and more than twice the
width and length of H. oregonensis. A descrip¬
tion of the first crab stage of P. crassipes follows:
carapace length, 3.7 millimeters; carapace width,
3.2 millimeters; front produced in two shallow
lobes, unlike the straight frontal margin of the
adult; the eyes proportionately very large, ex¬
tending past the sides of the carapace; the two
lateral teeth of the carapace distinct, and the
thoracic striae characteristic of the species faintly
indicated; the pereiopods provided with fine
setae, both plumose and simple, with bands of
dark and light areas along the ambulatory legs;
the antennae and antennules proportionately
larger and dissimilar to subsequent crab stages;
the dorsal aspect of the animal containing min¬
ute chromatophores which impart a stippled
effect.

The growth pattern of P. crassipes , from the
megalops stage up to and including the sixth
crab stage, was secured from observations on
two captive crabs collected during the megalops
condition. Both crabs were reared in large
refrigerator dishes containing sea water from
the locality in which they were collected. The
mean monthly temperatures during the rearing
period were: March, 14.8° C; April, 15.3° C.;
May, 15.8° C; and June, 16.2° G Food in the
form of minute polychaete worms ( Mercierella
enigmatica Fauvel) was plentifully supplied.

The data are summarized in Table 11. A re¬
markable constancy in total elapsed time with
respect to the attainment of certain molt stages
is immediately apparent. However, the varia¬
bility in size increment (19.1 per cent) signifi¬
cantly contributes additional evidence concern¬
ing individual variation in growth discussed at
some length on page 163.

THE TRANSITIONAL POSITION OF P. crassipes

BETWEEN A LITTORAL AND TERRESTRIAL
EXISTENCE

Specific and distinct habitats occur in all
parts of the earth; but nowhere are they more
apparent than along the seashore, where most
littoral animals occur in horizontal zones within
a vertically thin intertidal area. The high littoral
zone occupied by P. crassipes is significant in
this respect because its upper limit on the
strand is adjacent to true terrestrial conditions;
few species of the Brachyura range higher.
Therefore, P. crassipes is a species which has
almost acquired a terrestrial existence, and it
occupies a position on the strand higher than
that of most littoral crabs. Certain morphological
transformations in this species, which are cor¬
related with the attainment of a terrestrial
habitat, have been studied by Pearse (1931). A
physiological study by Jones (1941) on the
osmo-regulatory features of this species has
contributed additional evidence of its transi¬
tional character. The present investigation is
concerned with several additional features which

206

PACIFIC SCIENCE, Vol. II, July, 1948

are closely associated with the attainment of a
terrestrial habitat. The results of these studies
provide evidence to show that this species is
one which is physiologically, morphologically,
and behavioristically transitional between a
purely marine and purely terrestrial crab.

It is generally acknowledged that adjustment
to terrestrial conditions requires the ability to
resist desiccation. In this regard it was shown
earlier that, although 18 gills are present, the
volume of the gills in relation to the volume
of the body of P. crassipes is the lowest of any
of the crabs tested (see p. 146). Pearse ( 1929£)
has shown that crabs which are more highly
adapted to a terrestrial life generally have fewer
gills and these, by comparison, constitute a
smaller percentage of total body volume. It was
shown previously (see pp. 144 and 145 ) that this
species withstands desiccation longer than closely
allied but lower inhabitants on the strand. It
was also shown earlier (see p. 184) that P. cras¬
sipes is found typically in places of greater
temperature fluctuation than other crabs on the
strand in central California. Although the desic¬
cation tests fail to demonstrate a significant
difference between P. crassipes and H. nudus, the
latter species is invariably found in places of
lower temperature. The high littoral position
of P. crassipes unquestionably demonstrates its
adaptation to a range of temperature fluctua¬
tions wider than that of the other littoral crabs
of the region.

It has been pointed out that in the past the

development of speed by animals has often been
associated with aridity (Jehu, 1923). This
hypothesis gains support from those crabs which
have achieved the greatest success in conquering
terrestrial existence. Relatively great speed is
common to all land crabs in contrast to that of
crabs of the lower littoral belt. To examine this
hypothesis with respect to P. crassipes, the podo-
meric length of the second ambulatory leg of
this species was compared with that of H. nudus
(Table 12). Although all the podomeres of
P. crassipes are longer than the corresponding
ones of H. nudus, the two distal podomeres,
especially the dactylus, show a proportionately
greater difference in length than do the others.
P. crassipes , therefore, is adapted morphologi¬
cally for speed in a manner analogous to cur¬
sorial ungulates whose legs, especially the distal
segments, exhibit a tendency to lengthen.

The regulation of osmotic pressure within
the body is of great importance to marine or¬
ganisms for entering terrestrial fresh-water
habitats. Pearse (1931) has shown that crabs
living continually in, or closely associated with,
the ocean have the densest body fluids; those
which live on land have the least saline fluids.
Arthropods then, like vertebrates, in becoming
adjusted to a life on land, lose some of the salt
content of the blood. This loss suggests that
such animals are developing mechanisms for
maintaining the blood at a more or less constant
salinity, regardless of changes in the surround¬
ing medium. The salt content of the body

TABLE 12

A Comparison of Podomeric Length of the Second Walking Leg between Specimens of P.
crassipes AND H. nudus OF EQUIVALENT CARAPACE BREADTH.

WIDTH OF
CARAPACE
(MM.)

MEASUREMENTS (IN MM.) OF

THE SECOND WALKING LEG

SPECIES

I, II, III
Coxa, Basis,
Ischium

IV

Merus

V

Carpus

VI

Propodus

VII

Dactylus

Total

H. nudus .... .

27.1

5.0

12.5

8.2

6.0

7.5

39.2

P. crassipes .

27.1

5.7

13.9

8.6

7.7

8.8

44.7

Ratio of

P. crassipes
to H. nudus .

1:1

1.14:1

1.11:1

1.04:1

1.28:1

1.15:1

1.13:1

Biology of Pachygrapsus crassipes-— HlATT

207

fluids of P. crassipes is considerably reduced in
comparison to brachyurans which range lower
in the littoral belt (Pearse, 1931), but it is
higher than that of crabs which have achieved
a terrestrial or an almost terrestrial existence.

P. crassipes has achieved both hyperosmotic
and hypoosmotic regulation (Jones, 1941).
Therefore this species can regulate against the
increasing salt concentration of the water in the
gill chamber during periods of exposure to
the air and can maintain a constant body salinity.
A provision of this type is requisite to an active
terrestrial existence. H. nudus and H. oregonen-
sis, on the other hand, do not regulate hypo-
osmotically toward a salinity greater than that
of the ocean water (Jones, op. cit .), a fact
which discloses physiological evidence pertinent
to their restriction to lower and more shaded
positions on the strand. Both these species
exhibit hyperosmotic regulation in brackish
water to a greater magnitude than was found for
P. crassipes. This condition partially explains
why the latter species never occurs in water as
brackish as that in which the two former species
occur. Jones has shown, further, that hypo-
osmotic regulation occurs only among the Grap-
sidae and is exceedingly variable within the
group, ranging from complete absence (H.
nudus and H. oregonensis) to high development
( Uca crenulata, and all land crabs).

Homeostasis is definitely correlated with the
more highly advanced physiological grades
found in the vertebrates among birds and mam¬
mals. Therefore it is not unreasonable to assume
that stability of the internal physiological mech¬
anism of arthropods is conducive to increased
activity on the part of the possessor. Further¬
more, those crabs which have highly developed
hypoosmotic regulation (Uca, Grapsus, Pachy¬
grapsus, etc. ) have effected the greatest progress
toward the conquest of the terrestrial habitat;
hence they remain exposed to air for the great¬
est length of time.

Certain near-terrestrial Brachyura have evolved
homing instincts reminiscent of purely terres¬
trial groups. Ocypoda arenaria , when molested,

will return directly to its burrow from con¬
siderable distances (Cowles, 1908); the same
was noted for fiddler crabs (Pearse, 1914).
This behavior is unquestionably associated with
their fossorial habit. Although P. crassipes does
not burrow, several animals have been observed
to wander short distances away from their base
and later to return to the identical point of
departure. To test the possibility of a homing
instinct in this species, an experiment was
devised in which 21 crabs were collected from
a tide pool. The carapaces were marked with
large numbers, and the crabs were released at
four different loci 25 feet away from the pool.
The releasing loci were selected to present four
different types of routes home, varying from
simple slopes to complex crevice formations.
The crabs were released at night and observa¬
tions were begun the following afternoon.
Seven of the crabs were found back in the
original pool; four of six releases deposited in
difficult terrain returned; only two of five re¬
leased on a moderately difficult terrain returned;
and none returned from the locus separated from
the pool by smooth, level rock. Several marked
crabs were found in crevices near the releasing
loci. It seems apparent, from the foregoing
results, that homing behavior is nearly, if not
totally, lacking in P. crassipes. The crabs seemed
to be opportunists and secluded themselves in
the best possible location. The crabs which
returned to the tide pool probably found it
during their usual nocturnal wandering.

Algal types which grow highest in the littoral
belt may be significant in luring this species
farther toward land. The abundance of this
food, coupled with the fact that it forms the
major portion of the diet, is unquestionably
one of the reasons for the success of the species
in maintaining itself in this relatively exposed
area. It has been suggested by Pearse (1929*0
that crabs become herbivorous as they approach
terrestrial life; those which attain almost true
terrestrial existence are entirely herbivorous.
The crabs on the coast of central California are
herbivores, with algae comprising the predom-

208

PACIFIC SCIENCE, Vol. II, July, 1948

inant food supply; consequently this species
adheres to the food type characterestically util¬
ized by terrestrial species.

The reduction of gill volume relative to body
volume, the ability to withstand desiccation, the
eurythermal tolerance, the high locomotor de¬
velopment, the osmo-regulatory development,
and the herbivorous food habits, all combine
to substantiate the thesis that this species has
progressed from a purely marine habitat toward
a terrestrial existence. With these data at hand,
it is possible to discuss with considerable assur¬
ance the route by which this species may be
attaining a terrestrial life. Two routes to land
are open: one via brackish to fresh water
through estuaries and bays, and the other via
the littoral area without first encountering de¬
creased salinity. Inasmuch as hypoosmotic regu¬
lation is more highly developed than hyperos¬
motic regulation, it is certain that the adapta¬
tion to land has progressed through the littoral
area rather than via the estuarine route, notwith¬
standing the occurrence of P. crassipes in the
more saline areas of bays and estuaries. In con¬
trast, the absence of hypoosmotic regulation, in
conjunction with the highly developed hyperos¬
motic regulation, indicates that H. nudus and
H. oregonensis are progressing toward a terres¬
trial existence via the estuarine route.

Barrell (1916) and Lull (1917) minimize
the number of animals which have attained land
directly through the intertidal zone; but, if we
may judge from the multitude of species now
partially adjusted to land, there must have been
many in the past. Inasmuch as this species is
attaining or has achieved morphological, physi¬
ological, and behavioristic adaptations which
enable it to withstand the drier conditions at¬
tendant to a near-terrestrial habitat, and inas¬
much as these adaptations coincide with those
found in true land crabs, it is evident that
this species, at the present time, has progressed
toward a terrestrial existence via the littoral
route.

Barrell (op. cit .) also maintains that the
rarity of passage of crustaceans, gastropods, and

vertebrates from a truly marine to a truly terres¬
trial mode of life through the apparently open
path of the tidal zone, contributes evidence that
an unused food supply could not alone operate
as a cause sufficient to induce this change. How¬
ever, the supposition that one or another factor
is the primary cause for migration from sea
to land is a somewhat limited point of view.
The preservation of species depends upon the
procurance of food, protection from enemies,
reproduction, and continuous adjustment to
continually changing environment; hence one
lure is not sufficient cause for a migration of
this magnitude. The utilization of algal types
which occur high on the strand was only pos¬
sible after the crab was adapted to it, so that
their utilization did not interfere with other
exigencies of life. P. crassipes has, then, made
progress toward the attainment of a terrestrial
habitat, the main prerequisites being aerial
respiration, water conservation, swiftness, and
internal stability.

SUMMARY

1. The present investigation of Pachygrap>sus
crassipes Randall, the lined shore crab, is con¬
cerned especially with its geographical and eco¬
logical distribution, intermolt cycle, molting,
general habits and behavior, defensive mutila¬
tion and regeneration, predaceous and parasitic
enemies, reproduction, and growth. In addition,
information is advanced pertaining to the level
attained by this species in the transition from a
purely marine toward a terrestrial habitat.

2. The authenticated geographical range of
this species follows well-defined isotherms and
extends along the western coast of North Amer¬
ica from latitude 45° N. to latitude 24° N.,
and along the coast of Japan and Korea from
latitude 37° N. to latitude 34° N. Probable
avenues of dispersal to the Orient from America
are discussed, and locality records outside the
established range are considered.

3. The general littoral habitat of P. crassipes
includes three distinct biotopes. Each of these
biotopes provides the fundamental ecological

PLATE 1

Fig. 1. A view of terrain type 1 at Pacific Grove, California. Note tide pool in foreground and nu¬
merous crevices.

FIG. 2. A view of the mud bank along Bolinas Bay. Note the holes in the bank which serve P. cras-
jvper as refuge places. Ulva lactuca is abundant about the stems of Salicornia sp. shown above the vertical

PLATE 2

Fig. 1. A photograph which indicates the comparative smallness of the newly regenerated cheliped
(left side) after the first post-regenerational ecdysis.

Fig. 2. A photograph which shows the manner of papilla formation and regeneration of a dactyl located
on a cheliped.

Fig. 3. A photograph which discloses the "sponge” of an ovigerous female. Note the comparative size
of the ova, and the basket-like arrangement provided by the exopoditic and tergal setae.

Fig. 4. A photograph which shows the megalops of P. crassipes in dorsal view. Note particularly the
comparatively large eyes and long ambulatory appendages bearing hooked dactylar spines.

Fig. 5. A photomicrograph which shows empty egg capsules attached to the endopoditic setae of the pleo-
pods of a recent exuvia. Close examination will reveal the twisted funiculi.

Fig. 6. A photograph which shows the ventral aspect of a female crab with the abdomen extended. Note
the vulvae near the mid-line of the sixth sternite and the four pairs of abdominal pleopods.

Biology of Pachygrapsus crassipes — Hi ATT

209

requirements of this eurytopic species, viz., a
hard substrate containing refuge places, gen¬
erally devoid of loose stones, sand, or mud,
and possessing a more or less luxuriant growth
of ulvaceous or filamentous algae. The inter¬
specific relationship between the three grapsoid
crabs which range on the highest littoral hori¬
zon are described with respect to morphological,
physiological, and behavioristic differences.

4. A study of the seriation of morphological
transformations during the intermolt cycle ad¬
vances this knowledge to the grapsoid group.
External morphological characters are desig¬
nated as indicators of internal change. These
characters should enable investigators to diag¬
nose accurately the stages within the intermolt
cycle. A framework upon which to base accu¬
rate observations on both experimental and
behavioristic data is thereby provided. The
utility of such criteria to disclose periodicity
within a single intermolt interval is conclusively
confirmed by an examination of several hundred
wild specimens.

5. Exuviation is described in detail and its
critical relationship to the life of the crab is
made apparent. The incidence of molting is
clearly demonstrated to be associated directly
with the temperature of the water; the exuvial
frequency is highest in summer months and
relatively low from November to March. Post-
exuvial expansion is nearly completed in 3
hours. The individual size increment after ecdysis
is highly variable, and the percentage of size
increment varies inversely with the initial size
and the age. Comparative studies on the post-
exuvial size increments of male and female
crabs indicate that no significant variation occurs
until an initial width of approximately 25 milli¬
meters is attained. Thereafter, females exhibit
a smaller post-exuvial size increment. Male
crabs have been estimated to undergo 18 molts
and females 21 molts before attaining maximum
size, which is reached in a minimum of 3 years.

6. Visual, chemical, and tactile perception
are well developed in this species. Odor has
no stimulatory effect. When submerged food

is sought, visual perception generally precedes
either chemical or tactile, but chemical percep¬
tion becomes predominant after a brief interval
of time. Vision attains predominance during
diurnal aerial activity; the tactile sense gains
ascendancy after dusk. Although this species
does not respond to sound vibrations within
man’s range, the tactile sense is a significant
factor in regulating the activities of this crab.

7. This species is versatile with respect to
directional locomotion, and is transitional be¬
tween purely aquatic and terrestrial brachyurans
with respect to speed.

8. P. crassipes may be designated essentially
an herbivore, ordinarily a grazing herbivore,
less commonly a plant scavenger, while facul¬
tatively a carnivore, chiefly an animal scavenger,
and less often a predator.

9. Field data show that periodic changes in
daily temperature, tides, exposure, and illumina¬
tion have created a behavior pattern in these
animals which seldom deviates from the stand¬
ard and which approaches a monotony of repeti¬
tions.

10. The precocity and boldness of the young
crabs of this species are probably attributable
to their lack of experience and inhibitions,
rather than to the relatively large, faceted eye-
surface. The older and larger crabs with a
greater faceted eye-surface indulge in the varied
activities with more reserve and caution.

11. Advanced territorial relationships do not
exist for P. crassipes. Refuges are defended but
no individual forage areas are protected. There
is no tendency toward social aggregation.

12. Defensive mutilation in this species
occurs frequently and is easily achieved. The
mechanism underlying this phenomenon is de¬
scribed in detail, and comparisons are made
with other brachyuran species.

13. Experimental results show that a point
is reached during the final synthesis of the
integument (intermolt stage C4) before which
regeneration of a complete appendage will
ensue prior to ecdysis, and after which regenera¬
tion is inhibited prior to ecdysis. Three sue-

210

PACIFIC SCIENCE, Vol. II, July, 1948

cessive molts are required to bring a regenerat¬
ing limb back to normal size. With the excep¬
tion of the dactyl, regeneration from any point
in the appendage other than the fracture plane
has never been recorded in examinations of
the several thousand crabs observed. Easy self-
mutilation, rapid regeneration, and compara¬
tively brief intermolt cycles apparently con¬
tribute greatly to the success of this species in
reaching and establishing itself in an area rich
in food materials, notwithstanding the fact
that it is exposed to constant predation.

14. Predaceous enemies include gulls, rats,
man, sea anemones, and, because of their canni¬
balistic tendencies, the crabs themselves. Exter¬
nal parasites on this species are a rarity.

15. The usual brachyuran sexual dimorphism
is present and the sexes appear in virtually
equal numbers. Females were found to mature
in 11 or 12 months, at which time they had a
carapace breadth of about 15 millimeters. Male
crabs are more precocious; mature males appear
about 7 months subsequent to hatching, at
which time they measure about 12 millimeters
in carapace breadth.

16. Copulation and impregnation are de¬
scribed in detail. Most of the adult females
become ovigerous between April and Septem¬
ber; however, a few expel eggs during the
winter months. Evidence seems conclusive that
a portion of the females extrude two batches
of eggs within a single breeding season. The
incubation period requires approximately 1
month. The gross development of the incubat¬
ing larvae and the behavior of the berried
females are described.

17. The megalops stage of P. eras sipes is
described and compared with other closely asso¬
ciated grapsoid species.

18. An account of the growth of two care¬
fully reared crabs from the megalops to the
sixth crab stage shows considerable individual
variation in growth and a remarkable constancy
with respect to total elapsed time between the
post-megalopal molt and the attainment of the
sixth crab stage.

19. Morphological, physiological, and be¬
havioristic adaptations exhibited by this species
disclose that it has made progress toward the
attainment of a true terrestrial habitat, because
the main prerequisites ( aerial respiration, water
conservation, swiftness, and homeostasis) have
been partially achieved. Further, these adapta¬
tions — osmo-regulation in particular — seem to
indicate that the apparently open path from
the ocean to a terrestrial habitat via the inter¬
tidal zone is being traversed.

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— — 1929^. The ecology of certain estuarine
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425-427.

Randall, J. W. 1839. Catalogue of the Crus¬
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Townsend, from the west coast of North
America and the Sandwich Islands, with de¬
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Biology of Pachygfdpsus crassipes — HIATT

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- — 1923. The brachyuran crabs collected

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Reed, M. A. 1904. The regeneration of the
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A New Fern from Rota, Mariana Islands

W. H. Wagner, Jr.1

About 75 species of ferns are known from the
Mariana Islands. Of these the majority are com¬
mon and wide-ranging. Only a few species are
endemic or generally considered rare.

The only previous records of the cosmopoli¬
tan and very large genus Lastrea in the Marianas
are of the common species with much-dissected
fronds: L. Torresiana (Gaudichaud) Moore
{—Poly stichum Torresianum Gaudichaud,
Freyc. Voy. Bot. 33. 1827) from Guam, and
the similar L. ornata (Wallich) Copeland
( =Phegopteris ornata Fee, Hosokawa, Nat.
Hist. Soc. of Formosa, Trans. 26: 233. 1936)
from Alamagan, Sarigan, and Anatahan. The
following is a species of Lastrea with bipinnati-
fid fronds, known from eight complete fronds
obtained from two separate plants on the island
of Rota, collected by D. F. Grether of the Uni¬
versity of Wisconsin.

Lastrea Gretheri n. sp.

Fronde pinnata, chartacea, pubescente, utraque facie
glandulifera, 3 7 cm. alta, 1 0 cm. lata; stipitibus 1 1 cm.
aids, minute pubescentibus, sulcatis, stramineis, sed
basi atrocastaneis, et paleis atrocastaneis pubescentibus
linearibus acuminatis vestitis; pinnis ad alam 1 mm.
latam pinnatifidis, fere sessilibus, 5.5 cm. longis, 1.0
cm. latis; pinnis basalibus non brevioribus; segmentis
5-6 mm. longis, plerisque oppositis, oblongis, obtusis,
integris, marginibus deflexis; venulis 6-8-paribus, li-
beris, simplicibus; soris parvis, 3-5-paribus, ad seg-
menta submarginalibus, confertis; indusio crasso, cas-
taneo, persistente, glanduloso; sporis nigris.

A fern of the habit of Cyclosorus dentatus (Forsk.)
Ching. Rhizome not taken but form of stipe bases
indicating a creeping rootstock. Scales of stipe-bases
linear-acuminate 3-7 mm. long, 0.5- 1.0 mm. broad
at base, shiny dark-brown, provided with numerous
needle-like pale hairs, 0.1 mm. long. Fronds 4-5,
oblong-lanceolate, 37 (33-41) cm. tall, including the
stipe, 10 (9-12) cm. broad at the middle. Stipes 11
(9-13) cm. long, 1.5 mm. thick in the middle, drying
deeply sulcate, atrocastaneous in lower 3-6 cm., shad¬
ing into pale straw-color, densely beset with pale hairs.

1 Department of Botany, University of California,
Berkeley. Manuscript received October 28, 1947.

Rachis drying quadrate-ridged, and deeply sulcate, 1.3
(1.2-1. 5 mm.) thick in the middle, densely clothed
with white hairs except on ventral face where they
are red-brown. Pinnae 26 (25-28) pairs, opposite or
slightly subopposite except at tip, all fertile, lower
several pairs pointing downward or at right angles to
the rachis, the lowest of same size or but slightly
smaller than those above. Largest pinnae 5.5 (5.0-
6.0) cm. long, 1.0 (0.9-1. 2) cm. broad at base, nearly
sessile. Segments deflexed along margins, mainly op¬
posite, rounded oblong, entire, 5-6 mm. long, 2.5 mm.
broad at base, usually 18 pairs, costal wing 1 mm.
wide. Pinna-tips 5 (4-6) mm. long, 2-3 mm. broad
at base, narrowing to a rounded point. Lamina char-
taceous, soft in the living state, gray-green, abundantly
covered with pale hairs which are longer on the costa
dorsally, and of brown color on the costa ventrally,
and with numerous glands with yellow to orange-
brown expanded tips on both surfaces. Veins entirely
free, mostly opposite and simple, 6-8 pairs. Sori sub¬
marginal, crowded, about 0.5 mm. in diameter, 3-5
pairs, confined to segments as in Cyclosorus interruptus
(Willd.) Ching, and dark-brown. Indusia thick, per¬
sistent, red-brown, reniform, with numerous globular
shining dark-brown glands grouped near the sinus and
smaller, much paler ones on wings. Spores jet black.

Type: Growing on bare coral-limestone rock
in a crevice in a rather exposed situation on a
bank along a road at 800 ft. altitude on the
north slope of the plateau of Rota, Mariana
Islands. July 28, 1946. D. F. Grether 4468.
University of California Herb. no. 736319.
Duplicates are deposited at the U.S. National
Herbarium and the Bernice P. Bishop Museum.

Of Pacific ferns, this new species is most simi¬
lar to Lastrea Harvey i (Mettenius) Carruthers,
of Melanesia and Polynesia; the present plant
differs from it in smaller size, somewhat thicker
texture, lack of strongly dwarfed basal pinnae,
much more abundant hirsuteness, and in the
presence of numerous glands. Of other related
plants, L. euaensis Copeland of Tonga is nearer
to L. Harveyi than to the present species. L.
Margaretae (E. Brown) Copeland, of Rapa,
differs in the thicker texture, dwindling pinnae,
and broad gray-brown scales of the stipe-bases.

C214]

New Fern from Rota — WAGNER

215

FIG. 1. Lastrea Gretheri W. H. Wagner: a, habit of frond; b, details of pinnules; c, stipe-base scale showing
distribution of hairs; d, hair from lamina; e, forms of glands; /, form of indusium with distribution of dark
and light glands (these commonly much fewer, or some replaced by hairs).

A Restudy of the Reported Occurrence of Schist on Truk,
Eastern Caroline Islands1

Josiah Bridge2

INTRODUCTION

The islands of the western Pacific Ocean are
commonly grouped into two main divisions:
continental and oceanic. The former are be¬
lieved to be the remnants of a continental area;
the latter are thought always to have been
islands. The distinction is based upon several
facts, among which are the types of rocks of
which the islands are composed, the major
structure lines of the ocean basins, and the
structures of the islands themselves. The loca¬
tion of the eastern border of the former Mela¬
nesian or Australasian continent has been the
subject of considerable speculation. A number
of authors have presented "lines” showing the
distribution of different rock types, structural
data, and other geologic information in the
western Pacific islands, and from these have
made various deductions about the former extent
of the Melanesian continent. Among these are
the "border of the South Pacific Basin” (Mar¬
shall, 1912), the "andesite line” (Born, 1933),
the "metamorphic-plutonic” and "probable
boundary of the former Melanesian continent”
lines (Ladd, 1934), and the "Sial line” (Stearns,
1945; 1946). Inasmuch as many of the islands
in the vicinity of these "lines” were closed to
all except Japanese geologists from about 1916
until quite recently, most of the evidence for
their position and the subsequent classification
of the islands on either side of them as con¬
tinental or oceanic has been derived largely
from a study of the older literature. It is the
purpose of the present paper to revise a por¬
tion of the "metamorphic-plutonic” and "con¬
tinental boundary” lines of Ladd in accordance
with some recent observations made on Truk,

1 Published by permission of the Director, U. S.
Geological Survey.

2 Geologist, U. S. Geological Survey. Manuscript
received February 11, 1948.

one of the key island groups determining the
position of these lines.

Acknowledgments: The writer is deeply in¬
debted to Miss Jewell J. Glass and C. S. Ross
for the petrographic examinations and descrip¬
tions of the various rocks and for the photo¬
micrograph (Fig. 2). Thanks are also due to
G. A. Macdonald, H. S. Ladd, and J. I. Tracey,
all of whom have read and criticized the manu¬
script and have offered many helpful suggestions.

PREVIOUS WORK

Ladd (1934: 51, Fig. 6) gave a brief de¬
scription of the former Melanesian continent
and outlined on a map of the Pacific Ocean
south of lat. 10° N. the probable eastern boun¬
daries of several types of rocks, as well as the
probable eastern boundary of the former Mela¬
nesian continent. His map, with some modifi¬
cations, is reproduced as Figure 3 in this paper.

In the original figure, one of the lines, desig¬
nated the "metamorphic-plutonic line,” starts
just east of Yap, extends southeastward around
the Truk group, then turns abruptly south to
New Ireland and thence southeastward along the
northeastern edge of the Solomon Islands. A
second line, hereafter referred to as the "con¬
tinental line,” marks the "probable former
boundary of the Melanesian continent and true
structural boundary of the Pacific Basin” and
lies just east of and roughly parallels the "meta¬
morphic-plutonic line.” Both lines were drawn
so as to include Truk among the continental
islands and to exclude Ponape and Kusaie, high
volcanic islands lying east of Truk and geo¬
logically similar to it.

The inclusion of Truk among the continental
islands was based on the reported occurrence
of schist on one of the islands of the group
(Daly, 1916). Dalys paper, in turn, was not
based on direct observation but on information

1216]

TK24 TK23 TK22 TK21

Fig. 1. Looking west from the lighthouse on the eastern end of Moen Island showing
localities on Mt. Witipon from which trachytic material was collected. The approximate
extent of this material is shown by the light color and the absence of trees. Mt. Tonaachau
at right. PHOTO BY JOSIAH BRIDGE.

Fig. 2. Photomicrograph of trachytic material from specimen TK-24. Crossed nicols;
X 86. Photomicrograph by c. s. ross.

Schist on Truk — BRIDGE

217

contained in papers by various authors. In
Tables I and II, Daly credited the reporting of
amphibolite schist on Truk to Kramer (1908).
Kramer’s statement is short and explicit and
may be translated as follows:

The Truk atoll is a combination of high
islands surrounded by an atoll ring. Everywhere,
insofar as I have examined the high islands, I
found only volcanic rocks, of which feldspar
basalt was by far the most abundant type. The
only exception to this is Mt. Vidiboen, a bare
mountain 275 m. high on the northeastern part
of the large island of Vela. The eastern slope
of this mountain rises so gradually that one can
easily climb it on horseback, in fact a wagon
road could be built to the summit without great

difficulty. This mountain is unforested, and its
rock is amphibolite schist similar to the rock
on Yap. Dr. Klautzsch confirms my identifica¬
tion.

Stearns (1945, 1946) has drawn the "Sial
line” so as to include the Caroline submarine
plateau and all of the high, volcanic islands
rising above it, on the Sialic or continental side.
He states (1945: 615) that “The Sial line
lies a little to the east of the andesite line of
Gutenberg” and adds that "the line is the prob¬
able boundary of the continental platform of
Australasia” and that "these islands [west of the
Sial line] might be called Sialic islands from
the character of their basement, as all are com-

EXPLANATION

mm mm mm mm mm Limit Of PaleOZOiC rocks

mm Limit of Mesozoic rocks

Limit of. metomorphic or
Plutonic rocks (revised)

•••••••••• Proboble boundary

Melanesian Continent (revised)

aaMoaa Boundary of metomorphic and plutonic
areas as formerly recognized
oooooooooo Boundary of Melanesian Continent
as formerly recognized

Fig. 3. Map of the Pacific Ocean south of lat. 10° N., showing the general distribution of various types of
rock, the probable eastern boundary of the former Melanesian continent, and the true structural boundary of
the Pacific Basin. (After Ladd, 1934.)

PACIFIC SCIENCE, Vol. II, July, 1948

218

FlG. 4. Sketch map of the eastern part of Moen Island, showing Mt. Witipon and the localities from which
rock samples were collected.

posed of, or believed to be underlain by, con¬
tinental rocks.” He offers no evidence in sup¬
port of these statements, and for the reasons
to be given, the writer prefers the arrangement
shown in Figure 3.

FIELD WORK

In August, 1946, the writer spent 10 days
in Truk studying the geology and mineral re¬
sources of the islands in- connection with the
Economic Survey of the former Japanese Man¬
dated Islands. This survey was conducted by
the U. S. Commercial Company, a subsidiary of
the Reconstruction Finance Corporation, and
was made at the request of the U. S. Navy.3
At the time of the visit the writer was not

3 The geologic reports resulting from this survey
have been placed in open file in the offices of the U. S.
Geological Survey, Washington, D. C., and Honolulu,
T. H. A microfilm of the entire report has been de¬
posited in the Library of Congress.

familiar with Kramer’s report and consequently
made no special search for metamorphic rocks.
Later, during the preparation of the report, his
attention was called to Kramer’s statement, and
he was at once impressed with the similarity
of the description of the topography and vege¬
tation of Mt. Vidiboen on Vela Island with
conditions observed on the eastern part of Moen
Island.

Subsequent inquiry has established the fact
that the two islands are identical. According
to Karl J. Pelzer (oral communication, 1947),
a geographer with the Economic Survey who
was stationed on Truk for 4 months in the spring
and summer of 1946, Vela is one of the many
names which have been applied to Moen
(Wona) Island, and Vidiboen is clearly a
variant of Witipon, an 890-foot peak which
dominates the eastern end of this island (Figs.

Schist on Truk — Bridge

219

1 and 4). The general form of the mountain
agrees quite well with Kramers description, the
upper slopes are only sparsely forested, and
there is a wagon road along the eastern spur
which reaches the summit. This is the only
one of the three high peaks on Moen which is
reached by a road. The slopes of the upper
half of Witipon are covered with a dense
growth of coarse grass, ferns, low brush, and
scattered trees, and stand in sharp contrast to
the densely forested slopes of the other peaks
on the island (Fig. 1). Thus there appears to
be little doubt of the correct identification of
the region described by Kramer.

In the course of the investigations, Witipon
was climbed on two different occasions, rock
samples were collected at several localities ( Fig.
4), and it is believed that a representative
suite was obtained. The lower part of the moun¬
tain is composed of a series of basalt flows.
Two distinct types, one porphyritic, the other
non-porphyritic, may be recognized in hand
samples. These basalts are capped by what
appears to be a thick bed of well-stratified tuff,
but which has actually been determined as a
trachytic flow. This flow is at least 300 feet
thick. The entire series dips north at low
angles, commonly less than 5 degrees.

It is believed that this trachytic flow, repre¬
sented in the collections by specimens TK-21,
22, and 24, 4 is the rock which Kramer identi¬
fied as schist. The fine, parallel, flow structure,
and the glistening, micaceous-appearing, freshly
broken surfaces, particularly those which are
parallel to the flow lines, give this rock a
schistose appearance, so that superficially it
resembles the chlorite and amphibolite schists
found on Yap. However, it is much lighter in
color, and its mineralogic composition and tex¬
ture as seen in thin sections ( Fig. 2 ) are totally
different. According to G. A. Macdonald (per¬

4 The specimens described in this article, together
with rocks collected from other islands visited on this
trip, are now in the laboratories of the U. S. Geo¬
logical Survey, Washington, D. C. They will eventu¬
ally be deposited in the U. S. National Museum. A
duplicate set is already on deposit in the Bernice P.
Bishop Museum in Honolulu.

sonal communication, 1948), the Hawaiian
trachytes and oligoclase andesites commonly
have well-developed, thin, tabular feldspar plates
oriented parallel to the flow planes. The sur¬
faces of these plates often exhibit a micaceous-
appearing sheen, caused by the parallel orienta¬
tion of innumerable tiny feldspar grains, caus¬
ing the rock to resemble a fine-grained mica
schist. This explanation applies equally well to
the Truk specimens.

No other occurrence of this trachytic material
was found, either on Moen or on the other
islands which were visited in the Truk group.
However, this does not rule out the possibility
of other occurrences, for the examination was at
best a hasty reconnaissance and many of the
large islands were not visited.

Mt. Chukumong (Teroken, on H. O. Chart
6047), 1,214 feet, the highest peak on Moen
Island, lies about 1.25 miles west of Witipon.
If the regional strike and dip of the trachytic
material continue westward without apprecia¬
ble change, this material should cap the summit
of Mt. Chukumong, and should also appear in
the slopes of Mt. Tonaachau (794 feet), the
conspicuous peak on the northwestern tip of
Moen Island (Fig. 1). No definite information
about the kind of rock on its summit is avail¬
able. However, the upper slopes are covered
with a dense forest, totally unlike the grassy
upper slopes of Witipon, and this suggests that
the summit may be underlain by a different type
of rock. Mt. Tonaachau was climbed from two
different sides and only basalts and basaltic
agglomerates were seen. Macdonald (personal
communication, 1948) says that trachytic flows
which he has observed on other oceanic islands
are commonly extremely localized, and that no
special structural assumptions are necessary to
account for the local distribution of this ma¬
terial.

The western and northwestern shores of the
island were examined and no indications of
metamorphic rock were found. It is possible,
but in the writer’s opinion highly improbable,
that such rocks occur along the southern and
southeastern coasts, which were not studied.

220

PACIFIC SCIENCE, Vol. II, July, 1948

PETROGRAPHY

The specimens collected on Moen Island have
been examined by Miss Jewell J. Glass of the
U. S. Geological Survey, and the following
quotations are taken from her report:

TK-19, TK-20, black, porphyritic volcanic
rock. Identified as: Porphyritic basalt. This
rock is fresh and represents a typical example
of porphyritic olivine basalt. The phenocrysts
consist of unaltered crystals of olivine, augite,
and bytownite. The groundmass consists of a
net-work of feldspar (plagioclase) laths, fine¬
grained augite and olivine, with abundant octa-
hedra of magnetite.

TK-17, TK-18, dense, black volcanic rock.
Identified as: Fine-grained basalt. Different in
texture but the same in composition as TK-19
and TK-20 (described above). The ground-
mass consists of closely packed laths of plagio¬
clase, fine grains of olivine, swarms of black
granules of magnetite and a few phenocrysts of
augite. In TK-18 the olivine plates have altered
to iddingsite.

TK-24, pale gray, fine-grained rock contain¬
ing a scattering of feldspar phenocrysts. Identi¬
fied as: Tr achy tic rock . This is an extrusive
igneous rock. In thin section it shows, in the
crystallized groundmass, typical trachytic struc¬
ture. The groundmass consists essentially of
minute lath-shaped feldspar crystals having a
distinct parallel arrangement, or flow pattern,
and a sprinkling of tiny granules of magnetite.
The tabular feldspar phenocrysts show a zonal
banding and indications of multiple twinning.
The irregular optical properties of the feldspar
indicate that it has undergone some internal
structural change. Minute flakes of biotite and
chlorite are scattered throughout the mass, and
a few remnants of resorbed biotite crystals
remain. A small percentage of a brownish ma¬
terial occupies spaces between the feldspar
laths. Some of it resembles glass, but most of
it is birefracting and appears to be fine-grained
aggregates of ferromagnesian minerals. Occa¬
sionally a minute grain of augite is observed.

TK-21. This specimen is the same type of
rock as TK-24. However it has been considerably
altered by weathering. A crushed sample of the
more altered portion of the rock contains a
platy or apparently micaceous mineral, roughly
hexagonal in outline, and which is an alteration
product of undetermined composition.

TK-22, cream-colored, soft, chalk-like ma¬

terial. Identified as: Completely altered prod¬
uct of the trachytic rocks described above. Ther¬
mal analysis was not conclusive, neither was the
x-ray pattern. Suffice it to call the material
clay until more work can be done.

TK-23 consists of reddish-tan, irregularly
shaped, nodular material, associated with clays
(TK-23 -a). These overlie the trachytic material
and are believed to have been derived from it.
The nodules have been determined to be
bauxite.5

CONCLUSIONS

The placing of the continental line involves
two general classes of factors: (1) differences
in the kind and composition of the rocks form¬
ing the islands and (2) differences in struc¬
tures. The latter is possibly the more important
and may be further subdivided into: (a) broad
major structures reflected by the larger features
of the ocean bottom and (b) lesser structural
features which may be observed in the rocks
exposed on the various islands in question.
The importance of any set of criteria varies
greatly from place to place, and all of the
above factors have been used in the present
relocation of the lines.

In view of the identification of Kramer’s
material as a trachytic flow and of the observa¬
tions made on Moen and on many of the adja¬
cent islands, the occurrence of metamorphic
rocks on Truk now seems highly improbable.
The rocks collected on Truk are quite similar
to those found on Ponape and Kusaie, and
quite different from those observed on the high
islands to the west and southwest, a fact already
noted by Yossii (1937: 74, Tables I, II), who
lists trachyte from Ponape, but not from Truk
or Kusaie. This suggests that these three island
groups (Truk, Ponape, and Kusaie) are the
deeply eroded summits of a group of volcanoes
rising above the submarine Caroline plateau,
and that they are more nearly analogous to the
Hawaiian volcanoes than to the various island
groups lying to the west and southwest. There
is, therefore, no petrologic reason for including

5 See "On the Occurrence of Bauxite on Truk” on
page 223 of this issue.

Schist on Truk — Bridge

221

the islands of this group with the continental
islands.

Structural evidence also favors the exclusion
of the Caroline submarine plateau from the
continental area. Ladd’s original lines (Fig. 3),
drawn to outline occurrences of metamorphic
and plutonic rocks, cross this plateau between
Truk and Ponape, thereby placing Truk and
the western half of the plateau in the con¬
tinental area and leaving Ponape, Kusaie, and
the eastern half in the oceanic area, while
Stearns (1945, 1946) places the entire Caroline
group in the continental area. A study of exist¬
ing bathymetric charts (Int. Hydrographic
Bureau, 1940) shows little in the configuration
of the ocean bottom to favor such a division.
The Caroline plateau is shown as a number
of irregular elevations rising above the 5,000-
meter contour and not separated from the basins
on either side by deep trenches or other indica¬
tions of major structural features. However, it
must be recognized that most of the bathy¬
metric charts now available have been con¬
structed from insufficient data and that the
topographic features shown on them are at best
only approximations. The most reliable chart
of recent date (H.O. chart 5485) does not
extend far enough east to include the eastern
Carolines. A comparison of this chart with
earlier ones shows many important refinements
in ocean-bottom topography which have been
made possible by the inclusion of new data.

The "metamorphic-plutonic” and 'continen¬
tal boundary” lines, as here drawn ( Fig. 3), for
the most part follow major structural features.
These are the West Caroline and Palau trenches,
profound deeps with maximum depths of 4,500
and 3,500 fathoms, which lie just east of and
parallel to the Yap and Palau ridges (H.O.
chart 5485). Southwest of the Palau Islands
the lines swing abruptly southeast along the
northeast base of the submarine ridge on which
the Solomon and Admiralty Islands are located
and eventually join the continental lines as
drawn by Marshall (1912) and Ladd (1934).
No well-marked structural trench is present
along the base of this ridge, the average depths

being 2,000 to 2,500 fathoms, with only a few
points exceeding the higher figure. However,
there is a marked break in slope at the base
of this ridge, which is in direct prolongation
with the trend of the great Mindanao trench,
and this suggests that it might be the diminish¬
ing extension of a major structural line. The
northeastern slope of the Admiralty-Solomon-
Island ridge is similar to the northern front of
the Caroline plateau, having about the same
degree of slope although the latter descends
to somewhat greater depths. On the basis of
the known configuration of the ocean bottom
as shown on the existing bathymetric charts
there is little choice between the continental
line as here drawn and the "Sial line” of Stearns
(1945, 1946). The distinction must be made
on other evidence.

Final reasons for placing the "metamorphic-
plutonic” and "continental lines” so as to ex¬
clude the Caroline submarine plateau from the
continental area are the absence of extensive
deformation and evidence of recent uplift in
the rocks of the high islands which rise above
it, a striking contrast to conditions on the high
islands lying to the west of the "continental
line.”

On Truk, Ponape, and Kusaie there are no
metamorphic rocks, and there is little or no
evidence of extensive folding or faulting. Such
dips as may be seen in the various lava flows
are gentle, and may well be original. There are
no strongly elevated and tilted limestone ter¬
races, such as are common in the Mariana
Islands and to a lesser extent in the islands of
the Palau and Admiralty groups. In fact, the
occurrence of limestone on Truk, Ponape, and
Kusaie is limited to the low, discontinuous, 5-
to 7-foot bench which partially surrounds most
of the high islands. On Truk, Ponape, and
Kusaie there are indications of topographic
benches at relatively high elevations. These
have been interpreted by Tayama (1939) as
terraces of marine origin, older than the sur¬
face upon which the Plio-Pleistocene limestones
of the Mariana, Palau, and Admiralty groups
were deposited. The absence of metamorphism

222

PACIFIC SCIENCE, Vol. II, July, 1948

and deformation in the volcanic series and the
absence of highly elevated and tilted limestones
of Plio-Pleistocene age suggest that the plateau
has been a relatively stable unit since the middle
part of the Tertiary period.

In the southern Mariana Islands the Tertiary
and early Quaternary limestones have been
raised as much as 1,500 feet above the present
sea level (Rota). Locally the strata have been
highly tilted and the surfaces of marine ter¬
races have been offset and tilted by faulting
(Guam, Rota, Tinian, Saipan). Moreover, the
volcanic rocks underlying these limestones have
been strongly folded as well as faulted (Guam,
Saipan). All evidence at hand indicates that
diastrophic forces were active in the Mari¬
ana Islands throughout the Tertiary, and that
they are still going on. In the Palau Islands
the uplift of the Plio-Pleistocene limestones
amounts to at least 700 feet (Urukthapel
Island ) , and in the Admiralty group, limestones,
presumably of the same age, have been raised
at least 200 feet (Manus). The volcanic rocks
upon which these limestones rest have been
moderately folded and faulted, but there is no
evidence of regional metamorphism in the rocks
seen in either of these groups.

The Yap ridge appears to be older, and, at
the present time, more stable than either the
Mariana or Palau ridges. No raised limestones
have been found on Yap, and in this respect the
group resembles the high islands of the eastern
Carolines. The position of Yap with respect
to major structural features, together with the
fact that most of the rocks exposed on its
islands are well metamorphosed and are of
typical continental varieties, is responsible for
placing it west of the continental line.

Note: After this paper was transmitted my
attention was called to two other papers, one
botanic (Selling, 1947), one geologic (Bryan,
1945), which contain maps showing the posi¬
tion of one or more lines in the western Pacific
Ocean. A third and highly important paper
(Hess, 1948) appeared while this article was
in press. Although no mention of these papers
is made in the text they are included in the list
of references.

REFERENCES

Born, A. 1933. Der Geologische Aufhau der
Erde. Handbuch der Geophysik. Bd. 2, Berlin.
Bryan, W. H. 1945. The relationship of the
Australian continent to the Pacific Ocean —
now and in the past. Roy. Soc. N. S. Wales,
Jour, and Proc. 78(1,2) : 42-62.

Daly, R. A. 1916. Petrography of the Pacific
islands. Geol. Soc. Amer., Bui. 27: 325-344.
Gutenberg, B., and C. F. Richter. 1939.
Structure of the crust. Continents and oceans.
Chap. XII in Physics of the earth — VII.
Internal constitution of the earth, x-J-413
p. McGraw-Hill, New York.

Hess, Harry H. 1948. Major structural fea¬
tures of the western North Pacific: An inter¬
pretation of H. O. 5485, Bathymetric Chart,
Korea to New Guinea. Geol. Soc. Amer., Bui.
59: [in press].

International Hydrographic Bureau.
1940. Carte generale bathymetrique des

oceans. Feuille A-III, A'-III. 3rd Ed. Scale
1:10,000,000. Monaco.

Kramer, A. 1908. Aus den Schutzgebieten der
Sudsee, Studienreise nach den Zentral- und
West-Karolinen. Mitt, aus den Deut. Schutz¬
gebieten. 21 (3): 169-186.

Ladd, H. S. 1934. Geology of Vitilevu, Fiji.
Bernice P. Bishop Mus. Bui. 119- iii+263,
44 pi. Honolulu.

Marshall, P. 1912. President’s address, Sec¬
tion C, Geology, Austral. Assn. Adv. Sci.
13: 90-99.

Selling, Olof H. 1947. Further studies in
Schizaea. Svensk Bot. Tidskr. 41(4): 431-
450.

Stearns, H. T. 1945. Late geologic history of
the Pacific Basin. Amer. Jour. Sci. 243 (pt.
2): 614-626.

- - — 1946. An integration of coral reef

hypotheses. Amer. Jour. Sci. 244 (pt. 1) :
245-262.

Tayama, R. 1939. Terraces of the South Seas
Islands under the Japanese Mandate. Imp.
Acad. Japan, Proc. 15: 139-141.

U. S. Hydrographic Office. 1944. Truk,
northern half. H. O. Chart No. 6047. Scale
1:55,600. Washington, D. G
— — — 1946. Bathymetric chart, Korea to New
Guinea, including the Philippine Sea. H. O.
Chart No. 5485. Scale 1:4,075,440. Wash¬
ington, D. C.

Yossn, M. 1937. Distribution of igneous and
metamorphic rocks in the South Seas Islands
under Japanese Mandate. Imp. Acad. Japan,
Proc. 13: 74-77.

NOTES

On the Occurrence of Bauxite on Truk

In a paper appearing elsewhere in this issue
(Pacific Set. 2(3): 216) mention is made of
the identification of bauxite on Moen Island
in the Truk group. Although the Japanese
had probably discovered this material on these
islands in the course of their comprehensive
search for mineral deposits throughout the
former mandated area, the writer knows of no
published reports on the subject. The purpose
of this note is to record this occurrence and
to give a brief description of the deposit.

The Truk group lies in lat. 7° 25' N., long.
151° 30' E., and consists of about 15 large
and small volcanic islands rising out of a
lagoon about 30 miles in diameter and sur¬
rounded by a reef ring. The total area of the
islands within the lagoon is about 37 square
miles.

The only bauxite found to date in the Truk
group is located on Moen, the second largest
island in the group (area about 7.3 square
miles). The deposit appears to be confined
to a small area of about 200 acres on the
summit of Mt. Witipon (Takeun on some
maps), an 890-foot peak which dominates the
eastern end of Moen (see p. 218, Fig. 4, this
issue). The material, represented in the col¬
lections (see p. 220, this issue) by specimen
TK-23, consists of reddish-tan, irregularly
shaped, vesicular nodules, which are from 2
to 3 inches in diameter, and which superficially
resemble small, dried sponges. Freshly broken
surfaces are mottled in shades of red and
yellow, and have a fine, granular texture. These
nodules occur in a pale, yellowish-buff, fine¬
grained clay, but are most abundant at the
surface as a result of concentration during the
erosion of the clay. The clay itself is the
weathered residue of a fine-grained trachytic
flow (see p. 219, this issue) and is quite dif¬
ferent from the gray and deep-red clays derived
from the weathering of the basalts which cover
most of the island.

Early in 1947 a sample of one of these nodules
was tested on a portable thermal analysis unit
(Hendricks, Goldich, and Nelson, Econ. Geol.

41: 64-75, 1946) and was estimated to con¬
tain about 60 per cent gibbsite and 20 per cent
kaolinite. From this it was estimated that the
sample contained about 42.5 per cent alumina.
This was determined from the gibbsite fraction
alone, and no allowance was made for the
alumina in the supposed kaolinite.

Recently a chemical analysis of the same
nodule was made in the laboratory of the U. S.
Geological Survey ( W. W. Brannock, analyst),
and the results were as follows:

Insol. 9.37

A1203 53.08

Fe203 7.26

Ti02 0.66

Loss on ignition 29.68

An X-ray determination of the same material
made in the laboratory of the Geological Sur¬
vey, by George Switzer, showed that the pre¬
dominant mineral present was gibbsite, mixed
with some halloysite. Boehmite is not present,
and kaolinite occurs in very small amounts. A
re-examination of the curve produced by the
portable thermal analysis machine shows that
it carries a low peak at 140° and a strong peak
at 610°. These peaks agree closely with those
in the curve for halloysite given by Hendricks,
Goldich, and Nelson ( op. cit., Fig. 5 ) and indi¬
cate that the second mineral should have been
interpreted as halloysite. Twenty per cent of
halloysite would add about 7.9 per cent of
A12Os and 9.3 per cent of Si02 to the sample,
and, considering the very rough nature of the
estimate, the new totals agree very well with
the analysis.

Using the standard assumption that each
unit of per cent of Si02 makes 1.1 per cent of
alumina unavailable in the Bayer process, the
available A1203 of the sample is reduced to
42.78 per cent and is therefore not of com¬
mercial grade. The low iron content of this
bauxite is noteworthy, especially when com¬
pared with analyses of bauxites from other
islands in the Pacific, notably Bintan in the
Dutch East Indies and Palau in the western

[223]

224

PACIFIC SCIENCE, Vol. II, July, 1948

Carolines, which average at least 16 per
cent Fe203.

The associated clays were also tested on the
portable thermal analysis unit and were found
to contain less than 5 per cent gibbsite. Two
samples of clay derived from the weathering
of basalt were also tested by this method. One
contained a very small amount of gibbsite, the
other none.

The discovery of bauxite on Truk was not
unexpected, for bauxite deposits of some sort
are known to occur on most of the high vol¬
canic islands in the western Pacific. Deposits
have long been known to occur on Ponape and
Kusaie, the two islands which geologically are
most nearly comparable to Truk. To date, the
trachytic material from which this bauxite has
been derived has not been found by the writer
on any of the other bauxite-bearing islands, and
in this respect the deposit is unique. In fact,
this parent material has not been identified on
any of the other islands in the Truk group itself.
This does not mean that it does not occur, but

merely that the reconnaissance made in 1946
was very brief, and that not all of the islands
in the group were visited or studied in any
detail. Yossii (Imp. Acad. Japan, Pro c. 13: 74-
77, 1937) reports trachyte from Ponape, but
not from any other island in the former South
Seas Mandate. Bauxite deposits, some of them
rather low in iron, occur on Ponape, but their
relation to the parent rock was not observed
by the writer.

Although the presence of bauxite on Truk
is interesting from a scientific standpoint, the
deposit is too small and the bauxite is of too
low a grade to be of commercial importance
at the present time. No further examination
of the islands for bauxite alone appears to be
warranted; however, a study of the occurrence
and extent of the deposits should be a part of
any general geological survey of the group. —
Josiah Bridge, Geologist, U. S. Geological Sur¬
vey. (Published by permission of the Director,
U. S. Geological Survey.)

Holes in the Webs of Shearwaters

DURING the COURSE of examining young and
a few adult Wedge-tailed Shearwaters (Puf-
finus paciftcus cuneatus) on Rabbit and Moku
Manu Islands just off Oahu, in the fall of 1947,
23 of 38 birds checked at random were found
to have holes in the webs of their feet. These
holes varied in number and size from several
large ones (see photograph) to one or more
very small ones. As many as seven holes were
found in a single foot. Sometimes, as is also
shown in the photograph, the holes were mar¬
ginal and formed gaps on the edges of the webs.

The holes were first found in young a few
weeks old (smaller young were not observed).
They rarely appeared in the feet of 25 marked
young which were observed from the age of
about 1 month to 3 months. Once formed in
these older young, the holes remained essen¬
tially unchanged. Apparently the webs became
tougher and thicker with age and less easily
torn. Large holes, in particular, seemed to have
been acquired during the first few weeks after
the young hatched.

The explanation of the holes has not been
conclusively determined. Small scabs have been
noted on the feet of a number of young birds. I
have picked off such scabs, as a bird might, and

Fig. 1. Wedge-tailed Shearwater’s foot. Life size.

thus made or disclosed small web perforations.
(Of 6 pin pricks made in the webs of 2 young,
1 small permanent hole resulted. ) Further peck¬
ing or scratching at such a small hole might
well lead to a larger one, for a tear in the web
soon rounds out as the edges contract and heal.

The cause of the original small scabs, if they
may be taken as the starting point in hole de¬
velopment, seems most likely to be the bites of
the large hippoboscid flies ( ? Olfersia sp. ) that
very commonly infest the young. Small carni¬
vorous ants are present and might cause irrita¬
tion that would be pecked. Fighting between

NOTES

225

young is not a possible explanation because the
young seldom, if ever, leave their burrows dur¬
ing the first month or two. Nor does it seem
likely that the parents are involved, since the
feet of the young shearwaters are kept under
the body and hidden by down.

Review of much of the literature on shear¬
waters has revealed no mention of holes in their
webs although I know others have observed the
condition here. It is possible that this is a local

condition in these Hawaiian populations, where
the species or large numbers of the parasitic fly
might explain the frequency of holes in webs.
The result is probably of no disadvantage to the
adult birds in their swimming except in those
few cases in which perforation progresses to an
extent such as shown in the photograph. —
Frank Richardson, Department of Zoology and
Entomology, University of Hawaii, Honolulu,
Hawaii.

Seventh Pacific Science Congress

FEBRUARY 2 TO 23, 194?

To be held at AUCKLAND and CHRISTCHURCH, NEW ZEALAND. Under the auspices of
the Royal Society of New Zealand, and with the assistance of the Government of New Zealand.

ORGANIZATION

The Pacific Science Association has ac¬
cepted the invitation of the Royal Society of
New Zealand to hold the Seventh Pacific Science
Congress in New Zealand in 1949. The forth¬
coming Congress has the same scope and pur¬
pose as those previously held, i.e., Honolulu
( 1920 ) , Sydney and Melbourne ( 1923 ) , Tokyo
and Kyoto (1926), Batavia and Bandoeng
(1929), Vancouver and Victoria (1932), and
Berkeley, Palo Alto, and San Francisco (1939).

The Royal Society of New Zealand has ap¬
pointed an Organizing Committee charged with
the general arrangements for the Congress, with
responsibility for perfecting the program and for
local arrangements in Auckland and Christ¬
church.

MEMBERSHIP

The Pacific Science Association has a mem¬
bership of 46 countries; i.e., those within or bor¬
dering the Pacific or having territorial responsi¬
bilities therein, and those which have carried out
research in the Pacific. Each country holds mem¬
bership through a Representative Institution, i.e.,
its National Academy or National Research
Council, or organization of similar status.

Personal membership of the Congress is under
three categories:

Official Members

( 1 ) Official delegates representing the consti¬
tuent countries of the Pacific Science As¬

sociation, restricted in number as deter¬
mined by the Association.

(2) Delegates otherwise accredited to the
Congress by the governments of the par¬
ticipating countries or by agencies of
those countries corresponding to the
Royal Society of New Zealand.

( 3 ) Officers and honorary officers of the Con¬
gress, members of the Organizing Com¬
mittee, chairmen and secretaries of stand¬
ing committees of the Pacific Science As¬
sociation, and chairmen of committees
and sections.

(4) Persons specially invited to attend by the
President of the Congress.

Members

Members include those persons, other than
official members, who are accredited to the
Congress by scientific societies, universities,
and other research organizations. Members
will be accorded full privileges of the Con¬
gress.

Participants

This category includes all other registered at¬
tendants at the Congress, including ladies ac¬
companying official members and members.
Participants will be accorded full privileges
of the Congress except that official members
and members will be given preference in
housing accommodation and on excursions.
The registration fee for New Zealand partici¬
pants will be £ 1 : 1:0.

226

PACIFIC SCIENCE, Vol. II, July, 1948

To permit the Organizing Committee to make
suitable provisions, each Representative Institu¬
tion is requested to send to the Secretary of the
Congress at the earliest possible date the names
and addresses of all official members, members,
and participants who are expected to attend the
Congress.

PUBLICATIONS

Handbooks and Program will be issued to all
members; the Proceedings of the Congress will
be issued to Representative Institutions and offi¬
cial members. Members and participants may
purchase the Proceedings at cost price.

PROGRAM ORGANIZATION
Ten divisions representing different sciences
have been organized and divisional meetings for
discussion of papers of divisional interest will
be held. [The ten divisions under which the
meetings will be organized are: ( 1 ) Geophysics
and Geology; (2) Meteorology; (3) Oceano¬
graphy and Marine Biology; (4) Zoology; (5)
B.otany; (6) Soil Resources and Agriculture;
(7) Anthropology; (8) Public Health and Nu¬
trition; (9) Social Sciences; (10) Organization
of Research.] The Program will, however, give
chief place to symposia on special topics of in¬
terest to two or more divisions. A first draft of
the program has been sent to Representative In¬
stitutions; the final program will be arranged
after hearing their views. [The final program
will be published in the October number of
Pacific Science if it is available in time-Ed.]

OFFICERS OF THE CONGRESS

Patron

His Excellency the Governor-General of New
Zealand, Sir Bernard Freyberg, V.C., G.C.M.G.,
K.C.B., K.B.E., D.S.O.

Honorary Presidents

The Right Hon. P. Fraser, C.H., Prime Minis¬
ter of New Zealand.

The Hon. T. H. McCombs, Minister of Scien¬
tific and Industrial Research.

Honorary Vice-Presidents

His Worship the Mayor of Auckland, J. A. C.
Allum, Esq., GB.E.

His Worship the Mayor of Christchurch, E. H.
Andrews, Esq., C.B.E.

Herbert E. Gregory, President of the First
Pan-Pacific Science Congress.

H. M. Tory, President of the Fifth Pacific Sci¬
ence Congress.

Ross G. Harrison, President of the Sixth Pa¬
cific Science Congress.

President

R. A. Falla, President of the Royal Society of
New Zealand.

Secretary -General

Gilbert Archey, Director, Auckland Institute
and Museum, Auckland. Cable Address: con¬
gress AUCKLAND.

The Secretary-General will welcome inquiries
or suggestions from intending members or par¬
ticipants, or from scientific institutions. — Gil¬
bert Archey , Secret ary -General , Seventh Pacific
Science Congress.

Further Information Concerning the Fulbright Act

The following summary of information,
available to date, concerning the implementa¬
tion of the Fulbright Act has been received
from Harold J. Coolidge, Executive Secretary of
the Pacific Science Board of the National Re¬
search Council.

The Fulbright Act (Public Law No. 584)
authorizes the Department of State to use a por¬
tion of the foreign currencies resulting from the
sale of surplus property abroad for purposes of
educational interchange and activities with for¬
eign countries.

At present agreements have been signed with
only two countries — China and Burma; but ne¬
gotiations are in progress with the following:
Australia, Austria, Belgium, Czechoslovakia,
Egypt, Finland, France, Greece, Hungary, Iran,
Italy, the Netherlands, the Netherlands East In¬

dies, New Zealand, Norway, the Philippines,
Siam, Turkey, the United Kingdom, and it is
expected that other countries may be added to
the list.

It should be stressed that since the money
available is only in foreign currencies and is not
convertible to American dollars, individual ar¬
rangements must be made for each American
participating in the program for such dollar bal¬
ances as he will require to meet his family needs
and other obligations in the United States dur¬
ing the period of his absence abroad.

While the term educational interchange may
be interpreted very broadly, the following ampli¬
fication will serve as a more useful guide to the
types of activities envisaged:

Aid in international reconstruction by assist¬
ing foreign countries to secure the services of

NOTES

227

Americans with specialized knowledges and
skills and to assist the peoples of these countries
to understand the American people, their
achievements, and their ideals.

Provision for Americans to study, teach, and
conduct research abroad in connection with
American schools or with institutions of higher
learning, and to add to the American store of
knowledge of foreign areas, peoples, and cul¬
tures.

Opportunities for a limited number of foreign
students to study in American institutions abroad
and to assist foreign students and teachers to
engage in educational activities in the United
States by paying for their transportation wher¬
ever foreign currencies can be used for this pur¬
pose.

Under the terms of the Fulbright Act, a Board
of Foreign Scholarships is charged with the re¬
sponsibility of selecting individuals and institu¬
tions which will participate under the act and
with the supervision of the exchange program.
The Board is composed of individuals represent¬
ing a wide range of educational and cultural
interests in addition to representatives of the
government agencies most concerned.

The Board has delegated responsibility for
preliminary screening of applicants for grants
to three agencies:

1. The Institute of International Education for
those wishing to study in foreign institu¬
tions, primarily at the graduate level;

2. The Office of Education for those wishing
to teach abroad in national elementary and
secondary schools;

3. The Conference Board of Associated Re¬
search Councils for those wishing to teach,
lecture, or offer technical instruction in con¬
nection with institutions of higher learning
or to pursue studies and research abroad at

the post-doctoral level. The Conference
Board will also screen applicants for teach¬
ing positions in American elementary and
secondary schools abroad.

For discharging this responsibility, the Con¬
ference Board has established a Committee on
International Exchange of Persons with offices
at the National Academy of Sciences Building,
Washington, D. C. All inquiries concerning the
exchange of professors, lecturers, specialists, and
research scholars at the post-doctoral level, and
inquiries concerning opportunities for teaching
in American primary and secondary schools
abroad including requests for application forms
should be addressed to:

The Executive Secretary,

Committee on International Exchange of
Persons,

Conference Room of Associated Research
Councils,

2101 Constitution Avenue,

Washington 25, D. C.

Inquiries relating to graduate student exchanges
should be addressed to:

Institute of International Education,

2 West 45th Street,

New York 19, N. Y.

All inquiries relating to national primary and
secondary school teaching should be addressed
to:

The Office of Education,

4th and Independence Ave. S.W.,
Washington, D. C.

Inquiries relating to exchanges other than those
concerned with the Fulbright Act should be ad¬
dressed to:

The Division of International Exchange
of Persons,

Department of State,

Washington 25, D. C.

228

PACIFIC SCIENCE, Vol. II, July, 1948

News Notes

A conference on the subject of conservation
in the U. S. Trust Territory in the Pacific was
held in Honolulu April 19 and 20 under the
sponsorship of the Pacific Science Board of the
National Research Council. More than a score
of persons representing the administrative or¬
ganization of the Trust Territory, the U. S. Navy,
the National Park Service, and other govern¬
mental and scientific departments and organiza¬
tions participated. The phases of the subject
considered were: (1) archeological sites and
monuments, (2) natural vegetation reserves,
(3) native participation in conservation, (4)
soil and water conservation and land use.

The conclusions of the conference were sum¬
marized and forwarded to the Pacific Science
Board for its consideration at the meeting sched¬
uled for May 21, 1948, in Washington, D. C.

A summary of the proceedings and recom¬
mendations of the Washington meetings will
appear in an early number of Pacific Science.

* * *

Pacific Discovery: This new "journal of nature
and man in the Pacific world,” published by the
California Academy of Sciences, is designed to
make manifold science interesting and intelli¬
gible to layman and scientist alike. With its
first attractive issue it achieves a welcome entry
into the wide, open areas of the Pacific and
of the press. Don Greame Kelley as editor and
art director, Robert C. Miller as managing
editor, and a group of associate editors drawn
from the faculties of the University of Cali¬
fornia, Stanford University, the University of
Washington, and from the scientific staff of
the California Academy of Sciences assure
varied interests and technical guidance for
forthcoming issues of the journal.

Pacific Discovery: published bi-monthly by the Cali¬
fornia Academy of Sciences. Editorial, subscrip¬
tion, and advertising offices: Golden Gate Park,
San Francisco 1 8, California. Subscriptions: Three
dollars per year; fifty cents for single copies.

The United States Geological Survey has re¬
sumed publication of the Geophysical Abstracts
after a 4-year interval, during which they were
issued by the U. S. Bureau of Mines. The Geo¬
physical Abstracts are published quarterly as
an aid to those engaged in geophysical research
and exploration. The bulletin covers world
literature on geophysics contained in periodicals,
books, and patents. It deals with exploration
by gravitational, magnetic, seismic,- electrical,
radioactive, geothermal, and geochemical meth¬
ods and with underlying geophysical theory
and related subjects. Copies may be purchased
singly or by annual subscription from the Super¬
intendent of Documents, Government Printing
Office, Washington 25, D. C. For subscription,
the Superintendent of Documents will accept
a deposit of $5.00 in payment for subsequent
issues. When this fund is near depletion the
subscriber will be notified. The deposit may
also be used to cover purchase of any other
publication from the Superintendent of Docu¬
ments. The present price of each copy of the
Geophysical Abstracts is 20 cents.

* * #

Japanese Ornithology and Mammalogy dur¬
ing World War II. — Report No. 102 of the
Natural Resources Section, General Headquar¬
ters, Supreme Commander for the Allied Powers,
47 p., Tokyo, 1948.

This annotated bibliography of 204 technical
papers published in Japan from January, 1941,
to November, 1947, is the tenth report pub¬
lished by SCAP dealing with zoological sub¬
jects, in a series that will bring to the attention
of scientists outside of Japan many significant
and valuable papers by Japanese scientists. The
bibliography of ornithology and mammalogy
was compiled by Dr. O. L. Austin, Jr., assisted
by Dr. Masauji Hachisuka for the avian section,
Dr. Haruo Takashima for the mammalian sec¬
tion, and Nagahisa Kuroda for the section on
periodicals.

NO. 4

.II ,\S\V\ OCTOBER, 1948

PACIFIC

SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

IN THIS ISSUE: Pettersson — Swedish Deep-Sea Expedition •
Lincicome and McConnaughey — A New Nematode of the Genus
Pseudophysaloptera • Bushnell — Scientific Institutions in the
Pacific Area • Schaefer and Marr — Juvenile Euthynnus lineatus
and Auxis thazard • St. John — Plant Records from the Caroline
Islands • Fisher — A New Echiuroid Worm and Key to Genera
of Echiuridae • Powers — Explosive Eruptions of Kilauea •
Copeland — Origin of Polynesian Flora • NOTES • INDEX

Published by

UNIVERSITY OF HAWAII
HONOLULU, HAWAII

BOARD OF EDITORS

Leonard D. Tuthill, Editor-in-Chief
Department of Zoology and Entomology, University of Hawaii

O. A. Bushnell, Assistant Editor
Department of Bacteriology, University of Hawaii

Ervin H. Bramhall

Department of Physics, University of Hawaii

Vernon E. Brock

Division of Fish and Game, Territorial Board of
Agriculture and Forestry
P. O. Box 3319, Honolulu 1, Hawaii

Harry F. Clements

Plant Physiologist, University of Hawaii Agricultural
Experiment Station

Charles H. Edmondson

Zoologist, Bishop Museum, Honolulu 35, Hawaii

Harvey I. Fisher

Department of Zoology, University of Hawaii

Frederick G. Holdaway

Entomologist, University of Hawaii Agricultural
Experiment Station

Maurice B. Linford

Plant Pathologist, Pineapple Research Institute
P. O. Box 3166, Honolulu 2, Hawaii

A. J. Mangelsdorf

Geneticist, Experiment Station, Hawaiian Sugar
Planters' Association
P. O. Box 2450, Honolulu 4, Hawaii

G. F. Papenfuss

Department of Botany, University of California
Berkeley 4, California

Harold St. John

Department of Botany, University of Hawaii

Chester K. Wentworth

Geologist, Honolulu Board of Water Supply
P. O. Box 3410, Honolulu 1, Hawaii

Thomas Nickerson, Managing Editor, Office of Publications and Publicity, University of Hawaii

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never for citing references (see later). Often, foot-
[ Continued on inside back cover ]

PACIFIC SCIENCE

A. QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

Vol. II OCTOBER, 1948 No. 4

Previous issue published July 1, 1948

CONTENTS

PAGE

The Swedish Deep-Sea Expedition. Hans Pettersson . 231

A New Nematode of the Genus Pseudophysaloptera from an Okinawan Shrew.

David R. Lincicome and Bayard H. McConnaughey . 239

A List of Scientific Institutions in the Pacific Area. O. A. Bushnell . . . 243

Juvenile Euthynnus lineatus and Auxis thazard from the Pacific Ocean off

Central America. Milner B. Schaefer and John C. Marr .... 262

Plant Records from the Caroline Islands, Micronesia: Pacific Plant Studies 8.

Harold St. John . 272

A New Echiuroid Worm from the Hawaiian Islands and a Key to the Genera

of Echiuridae. Walter K. Fisher . 274

A Chronology of the Explosive Eruptions of Kilauea. Howard A. Powers . 278

The Origin of the Native Flora of Polynesia. Edwin Bingham Copeland . 293

NOTES:

On the Herding of Prey and the Schooling of the Black Skipjack,
Euythunnus yaito Kishinouye. Robert W. Hiatt and Vernon
E. Brock . . . 297

An Addition to the Fish Fauna of the Hawaiian Islands. Vernon E.

Brock . 298

United States Committee on the Oceanography of the Pacific .... 299

The Pacific Oceanic Fishery Investigation . 300

Western Regional Conference for UNESCO . 300

News Notes . 301

INDEX . 303

Pacific Science, a quarterly publication of the University of Hawaii, appears in January,
April, July, and October. Subscription price is three dollars a year; single copies are one
dollar. Check or money order payable to University of Hawaii should be sent to Pacific
Science, Office of Publications, University of Hawaii, Honolulu 10, Hawaii. Reprints
of major articles are available and requests for these will be filled in so far as possible.

The Swedish Deep-Sea Expedition

Hans Pettersson1

INTRODUCTION

Cut off from active work at sea during the
Second World War, Swedish oceanographers de¬
voted their efforts largely to improving the
technique of deep-sea oceanography. The raising
of long and undisturbed sediment cores from
great depths appeared to us to be an especially
desirable development. Very little advance had
been made in coring methods during the half
century which elapsed between the cruises of
the "Challenger” and of the "Meteor.” In the
mid-thirties C. S. Piggot of the Carnegie Insti¬
tution, Washington, D. G, succeeded in obtain¬
ing cores between 1 and 3 meters long from
great depths by firing a coring tube vertically
downward into the sea bottom with an explod¬
ing charge (Piggot, 1936: 207). The chief
difficulty in this method was the friction be¬
tween the column of sediment and the interior
wall of the coring tube. Recently an attempt
to overcome this difficulty was made by utilizing
the high pressure at great depths to operate
a "vacuum core sampler” (Pettersson and Kul-
lenberg, 1940, 1941). An undisturbed core 14
meters in length was raised by means of this
instrument in the Gullmar Fjord on the west
coast of Sweden in 1942. Shortly afterward Kul-
lenberg developed the "piston core sampler,”
by means of which an undisturbed core 20.3
meters in length was raised from the Gullmar
Fjord in 1945 (Kullenberg, 1947). In this
device, by trigger action, the entire heavy steel
coring tube with attached weights is auto¬
matically released on approaching the bottom
and sinks down into the sediment. The main
wire cable from which the coring tube is
suspended is stopped abruptly as the coring
tube and weights are released to sink into the

1 Director, Oceanografiska Institutet, Goteborg,
Sweden. Manuscript received May 3, 1948; trans¬
mitted from the Seychelles.

D

sediment. A piston originally at the lower end
of the coring tube and attached by an internal
wire to the main cable is kept motionless while
the coring tube descends around it. Because
of the high pressure, the immovable piston
immobilizes the column of sediment beneath
it as it is cut out by the corer. A view of the
piston core sampler suspended alongside the
research vessel is shown in Figure 1.

In the spring of 1946 the Swedish govern¬
ment permitted the research vessel "Skagerak”
to be used on a trial cruise to the western
Mediterranean during which the piston core
sampler proved its worth. Cores 8 to 15 meters

Fig. 1. The Kullenberg piston core sampler about
to be hauled aboard the "Albatross.”

OCT 2 6 194®

232

in length were raised from depths between
1,600 and 3,600 meters. In many of these cores
numerous zones of coarse particles, largely of
pyroclastic origin (volcanic ash), were found
intercalated in the ordinary sediment, thus giv¬
ing an unrivaled record of the volcanic activity
in the vicinity during historic and prehistoric
time (Pettersson, 1946). A few of these cores
have since been subjected to various physical,
chemical, and biological analyses. In addition
to the standard analyses, the cores were exam¬
ined for pollen and radium content ( Pettersson,
et al, 1948). Study by Dr. Fred Phleger of the
Woods Hole Oceanographic Institution of the
Foraminifera contained in different layers of
three cores from the Tyrrhenian Sea indicates
that considerable change in the temperature of
the surface water has occurred, which inci¬
dentally makes it highly probable that the span
of time required for the deposition of these
organisms extends well back into the last
glaciation (Phleger, 1947).

Another Swedish invention tested during
the same cruise was developed by Professor W.
Weibull of the Bofors Armament Works. This
device records, by means of hydrophones and an
oscillograph on board the ship, the time lag
between echoes from the surface of the sedi¬
ment and from transition layers below this
surface. The echoes are initiated by exploding
depth charges at depths of 100 meters or less.
With this instrument a maximum thickness of
the sedimentary carpet was found in the center
of the Tyrrhenian Sea where, below a water
layer of 3,600 meters, the sediment appears to
have a thickness of nearly 3,000 meters (Wei¬
bull, 1947).

THE PRESENT EXPEDITION

The interest evoked in our native city of
Goteborg by these new tools of deep-sea re¬
search, and by the promise they give of a new
grip on the unsolved problems of the ocean
bed, made it possible for me to obtain, from
private donors, the extensive financial backing
required for an all-Swedish circumnavigating

PACIFIC SCIENCE, Vol. II, October, 1948

cruise (Pettersson, 1947: 399). Altogether
about 2 million Swedish kroner (more than
one-half million dollars) have been given to
the Royal Society of Goteborg (Goteborgs
Kungl. Vetenskaps och Vitterhetssamhale),
which nominated a committee for planning and
organizing the cruise. The great difficulty of
finding a ship suitable for the expedition was
happily overcome through the generous offer
of the great Brostrom Shipping Combine of
Goteborg to lend us their new training ship,
the 1,450-ton motor schooner "Albatross,” at
net running costs for the duration of a 15-
month cruise. They also gave permission to
install nine laboratories, work shops, refriger¬
ation machinery, and cold storage rooms, be¬
sides cabins and a mess room for the scientific
and technical staff (air conditioned for work in
the tropics) all set up within the space nor¬
mally used for cargo. The "Albatross” is shown
in Figure 2. Thanks to wholehearted co-oper¬
ation from great Swedish industries, a specially
constructed, electric deep-sea winch with an
electric power station of 140 kilowatts and
other necessary equipment were completed in
time. A view of a portion of this winch is
shown in Figure 3. The refitting of the ship
was carried out at the Lindholmen Shipyard in
Goteborg where the "Albatross” had been built.
The shipyard work was done at a fraction of the
normal cost, thanks to the generosity of the
owner of Lindholmen.

The scientific and technical staff is com¬
prised of ten men. Besides the author they are:
Dr. Borje Kullenberg, oceanographer and in¬
ventor of the piston core sampler; Dr. Nils
Jerlow, oceanographer and specialist on sub¬
marine light and on the transparency of sea
water; Dr. Fritz Koczy, oceanographer and
specialist on radioactivity and submarine pho¬
tography; Leif Bruneau, chemist; Dr. Gustaf
Arrhenius, geologist; Viggo Wenzel, depth-
charge soundings and short-wave specialist; A.
Jonasson, chief mechanic; and K. Pettersson,
assistant in the sediment work. Dr. John

Deep-Sea Expedition — PETTERSSON

233

Fig. 2. The ’’Albatross” under full sail.

Eriksson, surgeon to the expedition, was in cursions on shore. The ship is most ably corn-
charge of the biological work during the first manded by Captain N. Krafft and officers. The
part of the cruise and headed botanical ex- total number of non-scientific personnel is 24,

234

PACIFIC SCIENCE, Vol. II, October, 1948

FIG. 3. The deep-sea winch aboard the "Albatross.” Only the power drums are shown in the
photograph; the 8,000 meters of steel cable are stored on a third great spool which is not shown.

including 12 apprentices between 17 and 21
years of age.

Since our sounding gear is very heavy (ap¬
proximately 1,500 kilograms), and is carried
by a steel-wire cable 8,000 meters in length, a
very powerful electric winch capable of raising
and lowering a load of over 10 metric tons at a
maximum speed of 100 meters a minute was
specially built for the cruise. In order to work
such heavy gear undisturbed by wind waves
and ocean swell, the route to be followed was
chosen so as to fall mainly within or near the
region of equatorial calms where, fortunately,
the sea bottom offers problems of special inter¬
est. Our original plan was to begin with a
tour of the north Atlantic Ocean down to the
equator. Because of unavoidable delay in start¬
ing, postponed from the beginning of March
to the beginning of July, and the necessity of
avoiding the hurricane season in the West
Indies and the southwest monsoon in the west¬
ern Indian Ocean, it was necessary to take a
short cut from Madeira to Martinique and from

there to Cristobal, where we arrived on August
19. After necessary readjustments had been
carried out on our deep-sea winch, in which
we were most generously assisted by the Naval
Command of the Canal Zone, we passed through
the Panama Canal on August 26, and on the
following day we left Balboa on a WSW course.

IN THE ATLANTIC

On our way across the Atlantic we had op¬
portunities for testing our gear. Between Ma¬
deira and Cristobal eight cores varying in
length from 10 to 15 meters were obtained.
Most of them consisted of typical red clay,
which in the lower levels was considerably
tougher than in the upper layers. The greatest
depth from which a core was raised was nearly
6,000 meters. As long as we had very calm
weather or were running before a moderate sea
and swell our deep-sea echo sounder, specially
constructed for the cruise by the Marine In¬
struments of London, gave legible records down
to 6,000 meters. The record often became

Deep-Sea Expedition — PETTERSSON

235

illegible when we moved against head-winds
or an adverse swell because of air bubbles inter¬
fering with the ultrasonic beam. Even with
this limitation, which results largely from the
shallow draught of the "Albatross,” the echo-
graph was most useful for our coring operations.
The bottom profile showed an astonishing rug¬
gedness even at great depths. Perfectly even
surfaces extending over greater distances than
a few nautical miles were rarely met in the open
Atlantic. A curious rise or fall of the bottom
profile by "steps,” 20 to 50 fathoms and often
considerably more in height, suggestive of faults
across our course is quite frequent in the
records. Professor Weibull, who personally
followed the cruise to Cristobal, found dis¬
tinct echoes from a "bottom below the bottom”
corresponding to a thickness varying between
300 and 2,500 meters, i.e., values comparable
to those found in the Mediterranean from the
"Skagerak” in 1946.

IN THE PACIFIC

From Balboa we steered toward the Gala¬
pagos group (see map), raising sediment cores
and taking a few oceanographic series under
way. In the Bight of Panama a few hauls in
750 and 1,500 meters, respectively, were made
with our large ring net 2 meters in diameter
in which catches of weird-looking bathypelagic
fishes and invertebrates were brought up.

The cores taken here were greenish-gray
and rich in organic remains. In one case there
was a distinct smell of sulphuretted hydrogen
from the lower parts of the core. Near the Gala¬
pagos the sediment was of a light greenish color,
was rich in foraminiferal tests, and in one case
zones of dark volcanic ash were found in the
lower levels.

Our purpose in stopping at the Galapagos
was twofold. While the men aboard the ship
investigated the cool upwelling water south of
the islands, which is rich in plankton, and took
sediment cores from the bottom there, five of us
went ashore on uninhabited James Island where
we spent five unforgettable days collecting

plants for our great Swedish authority on the
Pacific island flora, Professor Carl Skottsberg
of Goteborg.

From James Island the "Albatross” followed
a course to the WNW which afforded means
for a complete oceanographic section across the
Equatorial Countercurrent, where the two lines
of divergence (upwelling water) and the line
of convergence (descending water) were quite
distinct. The oxygen minimum in intermediate
water layers was also very conspicuous. An
almost total lack of oxygen, less than 0.02 cc.
per liter, was found in a depth of 150 meters as
far north as latitude 14° 13' N, longitude 120°
25' W. Close to the 18th parallel north, the
"Albatross” turned south, crossing the Equa¬
torial Current System a second time during
which we repeated the observations by water
sampling in different depths and raising cores
from the sea bottom. The sediment was mainly
of the red clay type, but near the equator cal¬
careous sediments appeared which were rich
in Foraminifera. Curious signs of stratification
were also manifest. A great impediment to our
coring operations, even at great depths, was
the frequent occurrence of hard bottom which
the core samplers could not penetrate. In a
couple of cases this led to partial or total loss
of the corer. In two cases at least, fragments of
basaltic rock caught in the bit of the core
sampler proved the obstacle to have been a
lava bed covered by a thin veneer of sediment.
Similar difficulties occurred also near Nukuhiva
in the Marquesas. A botanical excursion was
made to the high Tovii Plateau of Nukuhiva
where bore kernels were taken from one of
the rare peat bogs known in the tropics for
future analytical examination for pollen. Be¬
tween Nukuhiva and Tahiti, where we arrived
on October 24, our crop of sediment cores was
very meager because of the hard bottom.

The sounding of sediment thickness by depth
charges gave results different from those found
previously in the open Atlantic Ocean, in the
Mediterranean, and in the Caribbean Sea. Vir¬
tually no definite signs of reflecting layers
situated at more than 200 meters below the

236

PACIFIC SCIENCE, Vol. II, October, 1948

surface of the sediment were obtained. On the
other hand, distinct reflexes from depths vary¬
ing between 60 and 180 meters were repeatedly
obtained. In one case where the corer had been
stopped by a superficial layer of lava, distinct
reflexes from depths similar to those mentioned
in the sentence above were obtained, which
indicates that the superficial layer of lava is
evidently not impenetrable to the acoustic waves
recorded by our hydrophones.

After a pause of 10 days in Papeete the
"Albatross” set out on November 2 on a
northerly course for Hawaii. Oceanographic
series across the Equatorial Current System
were made here also, with results very similar
to those made previously. While under way
we had occasion to repeat several of the stations
made over 70 years earlier by the famous
"Challenger” Expedition, with results corrobo¬
rating theirs but with a technique of coring
which afforded a depth of penetration more
than 20 times as great. The cores, varying
from nearly pure radiolarian ooze to nearly pure
globigerina ooze, displayed highly interesting
stratificatons, with white calcareous sediment
alternating with a dark brown or a red clay type.
It is tempting to ascribe these variations in
the composition of the sediment to climatic
fluctuations, acting either directly on the sur¬
face temperature of the water and its plankton,
or possibly causing a displacement of the whole
current system as the polar ice caps grew and
varied with inevitable effects on the atmos¬
pheric circulation. Here also, especially near
the Hawaiian Islands, hard bottom interfered
with coring operations and led to loss of instru¬
ments.

In Honolulu, when we arrived on November
28, we were most cordially received by munici¬
pal authorities, by our colleagues at the Bernice
P. Bishop Museum and at the University of
Hawaii, and also by the Scandinavian colony
represented by the Swe-Nor-Den Society mak¬
ing our stay an unmitigated pleasure. We had
occasion to lay our results before our colleagues
at a round-table conference at the University
of Hawaii, and profited greatly from discussions

there and from most stimulating suggestions
and advice from Dr. T. A. Jaggar, the greatest
volcanologist of our time.

From Honolulu we cruised SW until the
equator was crossed, changed our course to
WNW, crossed the equator again, and stopped
at Kapingamarangi Atoll where submarine light
measurements were carried out in the sheltered
waters of the lagoon. At the beginning of this
segment of our cruise the occurrence of hard
bottom again made coring operations difficult,
and the bottom profile presented the same
rugged and hummocky features as in the eastern
and central Pacific. Farther west the bottom
became more even and more amenable to both
coring and to sediment soundings by depth
charges. However, adverse winds, to which the
"Albatross” is rather susceptible because of her
low-powered auxiliary Diesel engine, made our
progress slower than we expected.

In general, the cores taken along our west¬
erly course near the equator were of calcareous
sediment, frequently stratified with brownish-
gray clay alternating with whitish-gray sedi¬
ment. Still farther west a greenish color alter¬
nated with gray or brown tones. Our last days
in the Pacific Ocean were devoted to hydro-
graphic series and soundings in the southern¬
most part of the Mindanao trench. Here a core
raised from the maximum depth of 7,700 meters
had a length of only 4 meters, the core sampler
having been stopped by a layer of coarse sand
rich in volcanic particles! An attempt to raise
a core from a still greater depth with an extra
wire cable attached to the lower end of our
8,000 meter steel cable failed when the extra
wire snapped after the corer was on its way
up. The instrument and its precious content
were lost.

Faint but legible echograms were taken at
cross sections over the trench in latitude 5° 20'
N. The results prove that the depths given in
the Snellius charts are several hundred meters
too great. Attempts to measure the sediment
thickness in the trench failed because the
ruggedness of the bottom profile produced
spurious echoes which obscured any deeper

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Deep-Sea Expedition — PETTERSSON

237

echoes from transition layers below the surface
of the sediment.

Steering south from the Mindanao Deep we
finally reached the idyllic harbor of Ternate
on January 26, 5 months less a day after our
start from Balboa. Our cruise across the Pacific
was finished.

SUMMARY OF RESULTS

During our Pacific cruise 57 cores with an
integral length of nearly 500 meters have been
taken, most of which display more or less dis¬
tinct stratification. In most, but not in all, cases,
the sediment profiles raised by means of the
long piston core sampler have been supple¬
mented by cores from the surface layers taken
by means of a short core sampler of Dr. Phle-
ger’s construction and kindly loaned by him for
the duration of the cruise.

According to the technique developed by Dr.
Kullenberg, the cores are extricated from thin
lining tubes 70 centimeters in length inside
the steel coring tube and, after a cursory exami¬
nation on board, are wrapped first in briophane
and then in pergament paper. These wrapped
cores are introduced into aluminum tubes filled
with molten paraffin wax and stoppered, after
which they are placed in cool storage, between
5° and 8° G, pending analysis after our return
to Sweden. Experience shows that cores pre¬
served in this manner remain practically un¬
changed indefinitely.

Sediment soundings by depth charges have
been carried out at 75 different positions, where
in most cases, two depth charges were set to
explode at different depths (500 and 2,500,
4,500, or 6,500 meters). Only between Balboa
and the Galapagos were reflexes recorded from
depths below the surface of the sediment sen¬
sibly deeper than 300 meters. For this striking
difference between our results in the Atlantic
and in the Pacific, no explanation can yet be
offered.

Four complete hydrographic sections have
been made across the Equatorial Countercurrent
System, yielding results similar to those indi¬
cated above. When worked up from the dy¬

namic point of view these sections, supple¬
mented by frequent bathythermograph sound¬
ings between the stations, will afford valuable
material for the study of the Equatorial Counter-
current and the accompanying strips of conver¬
gence and divergence.

At a few deep stations large-volume water
samples of 25 liters were taken from different
depths for analysis for radium (by the BaS04
Mitreissreaktion) and uranium (by fluor¬
escence). Such analyses are necessary for con¬
firmation of the ionium precipitation hypo¬
thesis set forth by the author in 1936
(Pettersson, 1937), and subsequently substan¬
tiated through the work of C. S. Piggot and
W. D. Urry (1939: 405; 1941), and also in
order to utilize measurements of radium in the
sediment for determining the rate of sedimen¬
tation in the upper layers (Pettersson, 1943,
1945).

Optical studies on sea water were carried
out, partly by direct measurements of sub¬
marine daylight in different spectral ranges
down to depths of 100 meters, and partly by
photographic methods down to greater depths
by means of a pressure-tight submarine camera.
In addition ultraviolet components have for the
first time been measured near the surface in
the open sea by means of a special technique
(Johnson, 1946). Finally, the amount of
suspended particles at different depths was
measured by means of the Tyndall method,
using samples obtained with specially prepared
water bottles. Interesting results which show
the occurrence of distinct maxima of such par¬
ticles at certain depths were found by these
measurements.

The records from our echo sounder have al¬
ready been referred to. When worked up
systematically, results of value both from a
bathymetrical and from a morphological view¬
point may be expected. An interesting obser¬
vation made repeatedly when crossing the bands
of convergence bordering the Equatorial Coun¬
tercurrent was extra reflections on the echo-
gram in depths between 80 and 150 fathoms.
These reflections indicate densely packed or-

238

PACIFIC SCIENCE, Vol. II, October, 1948

ganic matter, presumably shoals of fish or other
bathypelagic organisms. It is possible that a
kind of intermediate fishing grounds along
the equator may be made accessible to exploi¬
tation along commercial lines through a future
development of the midwater trawling net.

Preliminary measurements of the geothermal
gradient in deep-sea deposits were made by
means of a geothermometer of special con¬
struction. The conditions on the bottom as
well as those prevailing at the surface, i.e., wind,
waves, swell, and currents, necessarily severely
restrict the use of this instrument for it has to
be plunged down into the deposit to a depth
of about 12 meters and remain there until
equilibrium with the surrounding temperature
has been attained. Only two successful shots
could be made in the Pacific Ocean,, both giving
unexpectedly high values for the geothermal
gradient there (between 20 and 30 meters per
degree centigrade). A discussion of these re¬
sults, which indicate a more intense flow of
geothermal heat upwards through the ocean
bed than one has, for general reasons, been in¬
clined to assume, must be postponed until more
data have been obtained.

In summary it may be said that the new
technique developed in Sweden for investi¬
gating the deep-ocean bed and its sediments has
been found to work well in oceanic depths, and
it promises a new and fruitful field of work
on the border line between oceanography and
submarine geology. When our cores have been
examined by different kinds of analyses, a
task which will require several years to accom¬
plish, new light will be thrown on problems
regarding the deep-sea bottom, its morphology,
and its sediments and their chronology. For
the latter study both foraminiferal analysis and
radioactive age determinations are available.

The investigation of the deep-ocean bed, its
stratigraphy, and its fauna manifestly calls for
co-ordinated efforts on an international scale. In
this future work Honolulu, with its famous
Bishop Museum and its University of Hawaii,
appears destined to become the foremost center
of research in the central Pacific Ocean.

REFERENCES

Johnson, Nils G. 1946. On anti-rachitic
ultra-violet radiation in the sea. Oceanog.
Inst . Goteborg, Meddel. 8: 1-16.

Kullenberg, B. 1947. The piston core-sam¬
pler. Svenska Hydr.-Biol. Kom . Skrifter III
(Hydrografi), 1(2): 1-46.

Pettersson, H. 1937. Das Verhaltnis Thor¬
ium zu Uran in den Gesteinen und im Meer.
Akad. d. Wiss. Wien , Anzeiger Nr.l6: 1-2.

- • 1943. Manganese nodules and the

chronology of the ocean floor. Oceanog. Inst.
Goteborg, Meddel. 6: 1-43.

- - — 1945. Iron and manganese on the ocean

floor. Oceanog. Inst. Goteborg, Meddel.
7: 1-37.

— — — - 1946. Oceanographic work in the
Mediterranean. Geog. Jour. 107 (3,4) : 163-
166.

— — - 1947. A Swedish deep-sea expedition.
Roy. Soc. London, Proc. B, 134: 399-407.

— - and B. Kullenberg. 1940. A vacuum

core-sampler for deep-sea sediments. Nature
145: 306.

- — - - — 1941. Vakuumlodet. Oceanog.

Inst. Goteborg, Meddel. 5: 1— 16. [In Swed¬
ish.}

- — et aL 1948. Three sediment cores from

the Tyrrhenian Sea. Oceanog. Inst. Goteborg,
Meddel. 15: 1-94.

Phleger, Fred B., Jr. 1947. Foraminifera
of three submarine cores from the Tyrrhen¬
ian Sea. Oceanog. Inst. Goteborg, Meddel.
13: 1-19.

PlGGOT, C. S. 1936. Core samples of the ocean
bottom. Smithsn. Inst., Ann. Rept. 1936:
207-216, 6 pi.

- — and Wm. D. Urry. 1939. The radium

content of an ocean bottom core. Wash.
Acad. Sci., Jour. 29: 405-410.

- - - 1941. Radioactivity of ocean

sediments. III. Radioactive relations in ocean
water and bottom sediment. Amer. Jour. Sci.
239: 81-91.

— — — - — — 1942. Radioactivity of ocean
sediments. IV. The radium content of sedi¬
ments of the Cayman Trough. Amer. Jour.
Sci. 240: 1-12.

Urry, Wm. D., and C. S. Piggot. 1942. Radio¬
activity of ocean sediments. V. Concentra¬
tions of the radio-elements and their sig¬
nificance in red clay. Amer. Jour. Sci. 240:
93-103.

Weibull, W. 1947. The thickness of ocean
sediments measured by a reflexion method.
Oceanog. Inst. Goteborg, Meddel. 12: 1-17.

A New Nematode of the Genus Pseudophysaloptera
from an Okinawan Shrew

David R. Lincicome and Bayard H. McConnaughey1

INTRODUCTION

Two SHREWS, identified as Suncus murinus
riukiuanus (Kuroda), collected on Okinawa
Shima, Ryukyu Islands, were examined for
parasites in August, 1945. A large number of
nematodes were taken from the stomach, peri¬
toneal cavity, connective tissue in axillae of the
hind legs, and the pericardial cavity. Two
species are probably represented, one of which
forms the basis of this report. It was taken
from the stomach of each of the two hosts.
The writers are indebted to Dr. Frank N.
Young 2 for collection and identification of the
shrews.

Specimens were studied largely as wet whole
mounts in glycerine or lactophenol. The pat¬
tern of the caudal papillae in the male was
determined by dissection of more than a dozen
specimens. The terminal portion of the body
was severed just anterior to the bursa and re¬
moved to a glass slide in lactophenol. After
removal of the membranous bursa the tail
was flattened with ventral side uppermost and
held in place by a cover glass. The papillae
were then examined under the microscope.

The worms are apparently members of the
genus Pseudophysaloptera Baylis 1934 (family
Physalopteridae Leiper, 1908, subfamily Phy-
salopterinae Railliet, 1893) and have been here¬
tofore unrecognized in the literature. The name
Pseudophysaloptera riukiuana is therefore pro¬
posed.

1 Department of Medical Microbiology, University
of Wisconsin, Madison, Wisconsin, and Department
of Zoology, University of California, Berkeley, Cali¬
fornia, respectively. Manuscript received March 22,
1948.

2 Assistant professor of zoology, University of
Florida.

DESCRIPTION OF SPECIES
Pseudophysaloptera riukiuana n. sp.

General: Body white or opaque; cuticula in¬
flated at anterior end to form a collar in both
sexes (see Fig. 1, d)\ two large labia each
bearing two prominent submedian papillae and
median amphid (see Fig. 1, d); three bluntly
rounded teeth on the internal surface of each
lip, these teeth appearing to originate from a
common base in oblique en face views. The
cuticular surface reflected over the lips appears
denticulated as it becomes minutely folded.
Esophagus very long and composed of two
parts: anterior portion short, muscular; pos¬
terior long, glandular. Cervical papillae not
observed. Nerve ring located at level of caudal
half of the anterior part of esophagus.

Female: Body length, 14.5-28.0 mm.; body
width, 0.65-1.2 mm. Collar width, 0.27-0.44
mm. Distance to nerve ring, 0.32-0.97 mm.
Distance to excretory pore, 0.7-1. 8. mm. Eso¬
phagus: total length, 3.38-5.0 mm.; length and
width of anterior part, 0.47-0.88 X 0.13-0.22
mm.; width of posterior part, 0.2-0.43 mm.
Prominent, saddle-shaped constriction of body
marking site of vulvar opening, located in the
anterior half, third, or fourth of body. Uterus
didelphic in type (see Fig. 1, c). Eggs 46-52 n
in length and 26-28 /;, in width, with thick
hyaline shells and embryonated in utero.

Male: Body length, 9.5-14.0 mm.; body width,
0.43-0.7 mm. Collar width, 0.22-0.38 mm.
Distance to nerve ring, 0.32-0.42 mm. Distance
to excretory pore, 0.47-0.65 mm. Esophagus:
total length, 2.49-3.91 mm.; length and width
of anterior part, 0.41-0.61 X 0.09-0.18 mm.;
width of posterior part, 0.14-0.50 mm. Caudal

[239]

240

papillae (see Fig. 1, a) all sessile; 6 pairs on
ventrolateral surface posterior to ano-genital
opening, 1 large, unpaired, immediately anterior
to the ano-genital pore, 2 additional pairs
anterior and lateral to this opening. No spicules
observed either in free specimens or those in
copulo.

DISCUSSION

At the time he recognized the genus Pseudo-
physaloptera in 1934, Baylis described P.sori -
cina from a species of Crocidura collected in
the Tanganyika Territory, Africa, which, to the
writers’ knowledge, is the only species that has
been ascribed to the genus.

P. soricina, however, has been recorded from
other hosts by other investigators. In 1937
Chen reported this organism from "Suncus
coerulus” a shrew from South China. Baylis
again recorded P. soricina in 1944 "from Suncus
coeruleus kandianus” in Ceylon. Later Crusz
(1946) described the worm from the musk
shrew, " Suncus coeruleus ” in Ceylon.

The first American finding of this helminth
was made by Morgan (MS.), who reported
it in the Masked Shrew, Sorex p. personatus,
and in the Smoky Shrew, Sorex f. fumeus, in
Tennessee, Kentucky, Wisconsin, and Iowa. A
few females tentatively assigned as P. soricina
by Morgan were also collected from the Short¬
tailed Shrew, Blarina brevicauda brevicauda.

Specific differences in the genus Pseudophy-
saloptera apparently are chiefly concerned with
the male organism. The pattern of the caudal
papillae on the ventral surface offers a basis for
the separation of P. soricina and P. riukiuana.

Baylis records four pairs of caudal papillae,
all of which are typically sessile, post-anal, and
lateral. Baylis has kindly sent one of us ( D.L. )
a pair of the cotype specimens for examination
and comparison. We have confirmed four pairs
of post-anal papillae (see Fig. 1, b), but be¬
lieve that there may be an additional lateral
pair slightly anterior to the ano-genital open¬
ing. This pattern is in contradistinction to the
six lateral post-anal pairs, the single, large

PACIFIC SCIENCE, Vol. II, October, 1948

ventromedian pre-anal, and the two lateral pre-
anal pairs of papillae in the species at hand.

The expanded cuticular collar at the anterior
end of both sexes may be of significance. Baylis
(1934) states, ’'The head (fig. 4) has the same
structure as in Physaloptera, consisting of two
hemispherical lateral lips, followed by a wider
neck, the cuticle of which forms a collar.” At
the same time, Baylis ( op. cit., fig. 4, page 347 )
shows little evidence of what the writers in¬
terpret as a "collar” although a relatively small
proportion of the extreme anterior end is shown.
Examination of a cotype female loaned by Dr.
Baylis reveals no prominent inflation such as is
present on P. riukiuana (see Fig. 1, e).

There are other differences in body length,
distance to nerve ring, and total length of
esophagus that may be related to the degree of
contraction of the worms at fixation. The forms
under consideration here were generally well
fixed and well relaxed. Both specimens loaned
by Baylis were in a marked degree of contrac¬
tion.

Chen (1937) and Crusz (1946) have both
recorded P. soricina from shrews in China and
Ceylon, but their descriptions seem to offer
some doubt as to their specific identifications.
The papilla pattern of the male in both in¬
stances is described as "numerous.” Crusz’s
Figure 5 (page 63) shows at least a dozen
large and small papillae in scattered and irre¬
gular distribution on each side over the ventral
surface. The number and distribution of the
caudal papillae of the male are markedly dif¬
ferent here from P. riukiuana. With reference
to this point, in a personal communication
under date of March 1, 1948, Baylis stated, "I
have examined one of Mr. Crusz’s specimens . . .
I am not at all convinced that all the small
papillae’ shown in his figures are really papillae
at all. I think many of them are subcuticular
structures

As further evidence of a possible difference
between P. soricina of Chen and Crusz and the
present material, the absence of spicules might
be cited. Chen (1937: 428) states, "Spicules

New Nematode — LlNClCOME and McConnaughey

241

e

Fig. 1. Anatomical details of Pseudophysaloptera species (all drawings made with aid of camera lucida).
a. Lateral view of posterior end of male Pseudophysaloptera riukiuana. (Five post-anal and a single pre-anal pa¬
pillae are shown. The full pattern [2 pre-anal pairs, 1 single pre-anal and 6 post-anal pairs] could not be
demonstrated in every male because of the opaqueness of this area of the body.) b, Lateral view of posterior
end of cotype male Pseudophysaloptera soricina Baylis 1934. c. Lateral view of region of vulva of female
Pseudophysaloptera riukiuana showing didelphic uterus, d, Lateral view of anterior end of male Pseudophysa¬
loptera riukiuana. e, Lateral view of anterior end of cotype female of Pseudophysaloptera soricina Baylis 1934.

A, amphid; AG, ano-genital opening; BR, bursa; C, collar; E, esophagus; G, gut; L, lip; LP, labial papilla;
P, post-anal papilla; PAP, paired pre-anal papilla; PP, single pre-anal papilla; T, labial tooth; UB, uterine
branch; V, vulva.

242

PACIFIC SCIENCE, Vol. II, October, 1948

(?) indistinct, in the form of two barely dis¬
cernible structures, equal, pointed at both ends,
0.15 mm. long.” Crusz (1946) on the other
hand states, "A very distinct pair of spicules
present in one worm; slender, well ‘chitinized/
unequal, pointed anteriorly, and blunt or prob¬
ably broken off posteriorly. Right spicule 0.231
mm. long, left spicule 0.190 mm. long. No
trace of any spicules in the other two worms.”
In the same personal communication referred
to above, Baylis has "carefully examined, with
Mr. Crusz, the specimens from which he made
his drawing showing ‘spicules/ The structures
he shows as such are certainly there, and in the
position in which they are shown. They appear
to be unquestionably internal, and not (as I
had half expected) lying on the surface of the
preparation . . . My own suggestion is that they
do not belong to this specimen at all, but to
some other nematode (possibly Castronodus
strassenii Singh, 1934 which occurs in nodules
of the stomach of the same host), which had
somehow inserted them and had them broken
off.”

There is no evidence of the presence of

spicules in any of the males in the present col¬
lection.

REFERENCES

Baylis, H. A. 1934. On a collection of cestodes
and nematodes from small mammals in Tan¬
ganyika Territory. Ann. and Mag. Nat. Hist.
X, 13: 338-353.

- - 1944. Notes on some parasitic nema¬
todes. Ann. and Mag. Nat. Hist. XI, 11:
793-804.

Chen, H. T. 1937. Some parasitic nematodes
from mammals of South China. Parasitology
29(4): 419-434.

Crusz, Hilary. 1946. Contributions to the
helminthology of Ceylon. II. Notes on some
parasitic nematodes, with a description of
Anisakis tursiopis sp. nov. Ceylon Jour. Sci.
(B) 23 (2): 57-66.

Morgan, B. B. MS. The Physalopterinae
(Nematoda) of North American Vertebrates.
[Manuscript Thesis, University of Wisconsin
Library, Madison, Wisconsin.}

A List of Scientific Institutions in the Pacific Area

O. A. Bushnell1

INTRODUCTION

This list of scientific institutions in the Pacific
area is presented with the frank hope that, in
its way, it will help in the dissemination of
knowledge and the exchange of information
upon which the friendship of nations and the
peace of the world are built. The Editors of
Pacific Science believe that most scientists, as
should all men, seek peace as earnestly as they
do truth; and they believe, too, that, with the
world as it is now, scientists and teachers — no
matter what their disciplines may be — bear
much of the burden of hope which the people
of the world have placed upon their leaders in
their yearning for peace.

The opportunity to exchange ideas and per¬
sons, freely and without constraint, is one of
the most important elements in understanding
either a people or a nation. Any program or
action, then, which can facilitate this exchange,
is a useful one. Many such programs have al¬
ready been initiated, and many more of them
are contemplated. The admirable policy of ex¬
changing teachers and students among nations;
the programs and aspirations of UN, UNESCO,
WHO, FAO; the programs of governments
(such as that of the Fulbright Act), of inter¬
national congresses (such as the Seventh Pacific
Science Congress in New Zealand next Feb¬
ruary), of universities and research foundations,
of churches and of organized and private char¬
ities; the quiet work of individuals, all are con¬
tributing, as wars and treaties never can, to
understanding and to peace. The mere presenta¬
tion of this list of scientific institutions in the

1 Department of Bacteriology, University of Hawaii;
Assistant Editor, Pacific Science. Manuscript received
May 29, 1948.

Pacific area is a very small step toward this
goal, but — if only because no such catalogue
has ever been published before — it is felt that
the list will be of value to those individuals
and institutions throughout the world who may
be aided by it in making their larger progress
toward that goal.

In the compilation of this list, information
was sought direct from sources in each of the
countries concerned. Letters of inquiry, asking
for the names, addresses, and special qualifica¬
tions of scientific institutions in each of the
countries in the Pacific hemisphere "at which
visting scientists might find the facilities (and
the invitation) to pursue research in their par¬
ticular subjects,” were sent to the Embassies,
Legations, Consulates, and Information Services,
in the United States, of each of the Pacific coun¬
tries. Identical letters were sent to the Em¬
bassies, Legations, Consulates, and Information
Services of the United States in each of the
countries concerned. Similar requests were sent
to the Office of the Director General of
UNESCO, Paris, and to the Executive Secretary
of the United States National Commission for
UNESCO, Department of State, Washington,
D.C.

The co-operation received in response to these
letters of inquiry was amazing and heartening.
Answers were received from almost every office
addressed, and the information which was re¬
quested was freely given by every country, with
one expected and regrettable exception. The
list as it is published can be assumed to present,
along with the facts which were sought, the
good will and the interest of the countries
represented in it.

[243}

244

The list is a selected one, compiled from the
many answers received to the letters of inquiry.
It probably presents all of the major scientific
institutions now functioning in the countries in
and around the Pacific Ocean and undoubtedly
presents some of the minor institutions as well.
Because it is difficult to appraise national vanity
(even among scientists, often among diplo¬
mats!), and because of my own ignorance of
the stature of these institutions in faraway
countries, it is possible that some institutions
which are unworthy of the distinction have been
added to the list, and it is probable that some
which should have had their place in the list
have been left out. For these errors of judg¬
ment or omissions of fact I alone must be held
responsible, the while I hope that increasing
knowledge of the countries of the Pacific, the
dissemination and use of this list, and the co¬
operation of scientists throughout the world will
help to correct them.

The countries included in the list are ar¬
ranged alphabetically, and the institutions in
each country are also arranged according to
their alphabetical order. Each entry presents
the name of the institution (in SMALL CAP¬
ITALS), its address (in roman type), and —
where this information is known, or where it
is not already evident from the name of the
institution — the special qualifications of the
institution, or some descriptive summary of its
function (in italics). For example:

BERNICE P. BISHOP museum, Honolulu 35,
Hawaii. Collection, preservation, and study
of Hawaiian and Pacific material in eth¬
nology and the natural sciences...

In the cases of the major universities and
some of the research institutions which are
equipped with facilities to do research in most
of the fields of science, it has not been thought
necessary to name each of the departments of
the institutions. The word General suggests
their eminence and their versatility.

On the other hand, the mention of some

PACIFIC SCIENCE, Vol. II, October, 1948

specialty for an institution does not necessarily
mean that its work is confined entirely to the
field of that specialty. The specialty listed
means only that, in the opinion of the scientists
in that institution, or of the cultural attache
evaluating it, or of myself endeavoring to cat¬
alogue it, the word used is the one that seems
best to describe the work for which the insti¬
tution is distinguished. It does not mean that
no other work, either in related or in different
fields, is done there.

The fact that an institution is placed in this
list does not mean, furthermore, that it is ready
to accommodate visiting scientists immediately.
In every instance, a scientist who wishes to
pursue research in some institution other than
his own must first make personal and definite
arrangement with the institution of his choice
in order to he certain that the institution pos¬
sesses facilities which he might use and that he
would have the invitation of the institution ( or
of the country in which the institution is lo¬
cated) to share its facilities. This injunction is
particularly true for institutions in those coun¬
tries tvhich have recently been afflicted with
war, or which are still troubled with civil wars,
but it applies to all institutions, wherever they
may be. The further difficulties of exchange of
persons, of funds, of equipment, are so variable
that they cannot be mentioned here, but must
be left to the patience and perseverance of the
individual scientist to surmount.

Spokesmen for many of the countries whose
scientific institutions are represented in this
list, and for many of the institutions themselves,
have stated that they are very much interested
in helping visiting scientists and that they
would welcome visitors — "always provided, of
course, that space and equipment can be found
for them.” This is the difficulty in almost every
country: the welcome and the good will are
readily offered, but the great demands made by
their own people upon these institutions place
a premium upon laboratory space and equip¬
ment that all too often cannot meet even the
local needs.

Scientific Institutions — BUSHNELL

245

Acknowledgments: The assistance of the staffs
of the many diplomatic offices and information
services and of UNESCO, who helped to pro¬
vide the information requested of them, has
already been acknowledged in private letters to
them, but public acknowledgment is recorded

here. They are too many to name separately,
and, in many cases, so anonymous as to be
unknown at all, but their assistance has been so
valuable that the list should stand rather as
testimony to their work than to my own.

AUSTRALIA

AUSTRALIAN CHEMICAL INSTITUTE, 39 Martin
Place, Sydney, New South Wales.

AUSTRALIAN INSTITUTE OF ANATOMY, Can¬
berra, A. C. T.

AUSTRALIAN INSTITUTE OF MINING AND MET¬
ALLURGY, 309 Little Collins Street, Mel¬
bourne, Victoria.

AUSTRALIAN MUSEUM (SYDNEY), College
Street, Sydney, New South Wales.

AUSTRALIAN NATIONAL MUSEUM, Canberra,

A. C. T.

AUSTRALIAN NATIONAL RESEARCH COUNCIL,
Science House, 157 Gloucester Street, Sydney,
New South Wales.

BAKER INSTITUTE, Alfred Hospital, Melbourne,
Victoria.

Medical research.

BUREAU OF MINERAL RESOURCES, GEOLOGY,
AND GEOPHYSICS, Chancery House, 485
Bourke Street, Melbourne, C. 1., Victoria.

BUREAU OF SUGAR EXPERIMENT STATIONS,
William Street, Brisbane, Queensland.

COMMONWEALTH FORESTRY BUREAU, Canberra,

A. C. T.

COMMONWEALTH METEOROLOGICAL BUREAU,
Victoria and Drummond Streets, Carlton, Vic¬
toria.

COMMONWEALTH OBSERVATORY, Mt. Stromlo,
Canberra, A. C. T.

COMMONWEALTH SERUM LABORATORIES, 45
Poplar Street, Parkville, Melbourne, Victoria.

COMMONWEALTH X-RAY AND RADIUM LAB¬
ORATORY, University of Melbourne, Carlton,
N. 3, Victoria.

COUNCIL FOR SCIENTIFIC AND INDUSTRIAL RE¬
SEARCH (C. S. I.R.), 314 Albert Street, East
Melbourne, C. 2, Victoria. (Inquiries are to
be addressed to the Secretary.)

The C. S. I. R. is the largest Commonwealth
Government organization, and is exten¬
sively subdivided as follows:

ATOMIC PHYSICS RESEARCH, University
of Melbourne, Carlton, N. 3., Victoria.

BUILDING MATERIALS RESEARCH, Gra¬
ham Road, Highett, S. 21, Victoria.

COMMONWEALTH RESEARCH STATION,
Merbein, Victoria.

dairy research section, Lorimer
Street, Fishermen’s Bend, S. C. 8, Vic¬
toria.

DIVISION OF AERONAUTICS, Lorimer
Street, Fishermen’s Bend, S. C. 8, Vic¬
toria.

DIVISION OF ANIMAL HEALTH AND PRO¬
DUCTION, Park Street and Fleming-
ton Road, Parkville, N. 2, Victoria.

DIVISION OF BIOCHEMISTRY AND GEN¬
ERAL NUTRITION, University Grounds,
Adelaide, South Australia.

DIVISION OF ECONOMIC ENTOMOLOGY,
P. O. Box 109, Canberra City, A. C. T.

DIVISION OF ELECTROTECHNOLOGY, Na¬
tional Standards Laboratory, City
Road, Chippendale, New South Wales.

DIVISION OF FISHERIES, Box 21, P. O.
Cronulla, New South Wales.

DIVISION OF FOOD PRESERVATION AND
TRANSPORT, Private Bag, Homebush,
New South Wales.

DIVISION OF FOREST PRODUCTS, 69
Yarra Bank Road, South Melbourne,
S. C. 4, Victoria.

DIVISION OF INDUSTRIAL CHEMISTRY,
Lorimer Street, Fishermen’s Bend, S.
C. 8, Victoria.

DIVISION OF METROLOGY, National
Standards Laboratory, City Road,
Chippendale, New South Wales.

DIVISION OF PHYSICS, National Stand¬
ards Laboratory, City Road, Chippen¬
dale, New South Wales.

DIVISION OF PLANT INDUSTRY, P. O.

Box 109, Canberra City, A. C. T.

246

PACIFIC SCIENCE, Vol. II, October, 1948

DIVISION OF RADIOPHYSICS, National
Standards Laboratory, City Road,
Chippendale, New South Wales.

division of soils, Waite Agricultural
Research Institute, Private Mail Bag,
Adelaide, South Australia.

FLAX RESEARCH, Graham Road, Highett,
S. 21, Victoria.

INFORMATION SERVICE, 425 St. Kilda
Road, Melbourne, S. C. 2, Victoria.

IRRIGATION RESEARCH STATION, Private
Mail Bag, Griffith, New South Wales.

MIN ERAGR APHIC INVESTIGATIONS, c/o
Geology School, University of Mel¬
bourne, Carlton, N. 3, Victoria.

ore dressing investigation, c/o Met¬
allurgy School, University of Mel¬
bourne, Carlton, N. 3, Victoria.

radio research board, c/o Electrical
Engineering Department, University
of Sydney, Chippendale, New South
Wales.

SECTION OF MATHEMATICAL STATIS¬
TICS, c/o University of Adelaide,
North Terrace, Adelaide, South Aus¬
tralia.

SECTION OF METEOROLOGICAL PHYSICS
RESEARCH, 572 Flinders Lane, Mel¬
bourne, C. 1, Victoria.

SECTION OF PHYSICAL METALLURGY,
University of Melbourne, Carlton, N.
3, Victoria.

SECTION OF TRIBOPHYSICS, University
of Melbourne, Carlton, N. 3, Victoria.

WOOL TEXTILE TECHNOLOGY, 314 Al¬
bert Street, East Melbourne, C. 2,
Victoria.

WALTER AND ELIZA HALL INSTITUTE FOR MED¬
ICAL research, Royal Melbourne Hospital,
Parkville, Melbourne, N. 2, Victoria.

INSTITUTE OF AGRICULTURE OF THE UNIVER¬
SITY OF WESTERN AUSTRALIA, Nedlands,
Western Australia.

INSTITUTE OF MEDICAL RESEARCH OF NEW
SOUTH wales, Royal North Shore Hospital,
St. Leonards, New South Wales.

KANEMATSU MEMORIAL INSTITUTE OF PATH¬
OLOGY, Sydney Hospital, Sydney, New South
Wales.

MARINE BIOLOGICAL STATION, UNIVERSITY OF
SYDNEY, Port Jackson.

MELBOURNE WOMEN’S HOSPITAL, Swanston
Street, Melbourne, N. 3, Victoria.

Pathology .

microbiological laboratory, 93 Macquarie
Street, Sydney, New South Wales.

MUNITIONS SUPPLY LABORATORIES, Maribyr-
nong, Victoria, with branch establishments at
Adelaide and Sydney.

MUSEUM OF TECHNOLOGY AND APPLIED SCI¬
ENCE (formerly known as the Sydney Tech¬
nological Museum ) , Harris Street, Broadway,
Sydney, New South Wales.

NATIONAL HEALTH AND MEDICAL RESEARCH
COUNCIL, Canberra, A. C.T. (Inquiries are
to be addressed to the Director-General of
Health.)

NATIONAL MUSEUM (VICTORIA), Swanston
Street, Melbourne, Victoria.

NATIONAL UNIVERSITY OF AUSTRALIA, Can¬
berra, A. C.T. ("This University is now in
the course of being built. Details of organi¬
zation are not yet available, but research de¬
partments of medicine and physics are al¬
ready being established.”) [C.S.I.R., Letter,
February 20, 1948.}

PERTH MUSEUM, Beaufort Street, Perth, West¬
ern Australia.

QUEENSLAND museum, Gregory Terrace and
Bowen Bridge Road, Brisbane, Queensland.

SCHOOL OF MINES, North Terrace, Adelaide,
South Australia.

SCHOOL of MINES, Ballarat, Victoria.

SCHOOL of MINES, Kalgoorlie, Western Aus¬
tralia.

SOUTH AUSTRALIAN museum, North Terrace,
Adelaide, South Australia.

TASMANIAN museum, Macquarie Street, Ho¬
bart, Tasmania.

UNIVERSITY OF Adelaide, North Terrace, Ade¬
laide, South Australia.

General.

UNIVERSITY OF Melbourne, Carlton, N. 3,
Victoria.

General.

university of Queensland, George Street,
Brisbane, Queensland.

General.

UNIVERSITY OF SYDNEY, Parramatta Road, Syd¬
ney, New South Wales.

General.

UNIVERSITY OF TASMANIA, Hobart, Tasmania.

General.

UNIVERSITY OF WESTERN AUSTRALIA, Ned¬
lands, Western Australia.

General.

Scientific Institutions — BUSHNELL

247

STATE GOVERNMENTS AND DEPARTMENTS:
"Each of the six State Governments of Aus¬
tralia maintains scientific departments or sub¬
departments which deal largely with scientific
problems arising within the State, and which
also carry out a considerable amount of routine
and other scientific work associated with the
laws and regulations of the State. Thus, in each
of the States, there is a Department of Agricul¬
ture; a Forest Service; a Department of Main
Roads; a Mines Department; a Lands Depart¬
ment; and Government Chemical Laboratories,
etc. These bodies can render assistance in spe¬
cialized fields, but direct approach to them
would be desirable in all cases.” [C.S.I.R., Letter,
February 20, 1948.]

Mr. J. E. Cummins, Officer-in-Charge of the
Information Service of the Council for Scientific
and Industrial Research, writes that "it is not
possible to express an opinion as to whether
each of the institutions or departments [in¬
cluded in this list for Australia] would be able
to provide accommodation for visiting scien¬
tists. Unfortunately, the facilities for research
in some of the scientific institutions are very
limited, and, depending upon the time of visit

and the period of this, a definite decision could
not be made as a general statement. However,
I am sure that all of the Australian research in¬
stitutions listed would be only too pleased to
do their utmost to provide assistance.

"In the specific case of the C.S.I.R _ the

Executive Committee of the Council have in¬
formed me that they would be glad to welcome
visiting scientists in any of their laboratories
and would be only too pleased to make facilities
available for them. May I suggest, therefore,
that . . . intending visitors should make direct
enquiries before visiting Australia and while
their plans are still in the formative stage.”

This information was received from the In¬
formation Service of the Council for Scientific
and Industrial Research of the Commonwealth
of Australia, 425 St. Kilda Road, Melbourne,
S. C. 2, through the assistance of the Scientific
Liaison Office of the Australian Embassy, Wash¬
ington, D. G; from the British Consulate, Hono¬
lulu, T. H.; from the British Information Serv¬
ices, New York; from the American Embassy,
Canberra, Australia; and from UNESCO, Paris.
February-April, 1948.

BRITISH MALAYA

BOTANICAL gardens, Singapore.

COLLEGE OF AGRICULTURE, Kuala Lumpur.
INSTITUTE FOR MEDICAL RESEARCH, Kuala
Lumpur.

KING EDWARD VII COLLEGE OF MEDICINE, Singa¬
pore.

raffles college, Singapore.

General.

RAFFLES MUSEUM AND LIBRARY, Singapore.
Anthropology, archaeology, botany, and
zoology of Malaysia.

RUBBER RESEARCH INSTITUTE, Kuala Lumpur.

This information was received from the Amer¬
ican Consulate General, Singapore. March, 1948.

CANADA

FISHERIES EXPERIMENTAL STATION, 898 Rich¬
ards Street, Vancouver, B. C.

Fisheries technology.

PACIFIC BIOLOGICAL STATION, Nanaimo, B.C.

Fisheries, marine biology, oceanography.
UNIVERSITY OF alberta, Edmonton, Alberta.
General.

UNIVERSITY OF BRITISH COLUMBIA, Vancouver,

B. C.

Agriculture, bacteriology, biophysics, geol-
ogy, geophysics, marine algae and oceanog¬
raphy, nuclear physics, preventive medicine,
spectroscopy.

UNIVERSITY OF MANITOBA, Winnepeg, Mani¬
toba.

Botany, chemistry, geology, zoology.
UNIVERSITY OF SASKATCHEWAN, Saskatoon,
Saskatchewan.

Physical chemistry, medicine, physics.

This information was received from the Uni¬
versity of British Columbia, Vancouver, B. G,
and from the National Research Council of
Canada, through the Canadian (N. R. G) Sci¬
entific Liaison Office, Washington, D. C. Jan-
uary-April, 1948.

248

PACIFIC SCIENCE, Vol. II, October, 1948

CANAL ZONE

CANAL ZONE BIOLOGICAL AREA ( formerly
Barro Colorado Island Biological Laboratory),
Balboa, Canal Zone.

Biological research.

CANAL ZONE BRANCH, BUREAU OF ENTOMOL¬
OGY, U. S. DEPARTMENT OF AGRICULTURE,
Balboa, Canal Zone.

CANAL ZONE EXPERIMENT GARDENS, Pedro
Miguel, Canal Zone.

Botanical research.

This information was received from the
American Embassy, Panama, R. P., and from
UNESCO, Paris. January- April, 1948.

CHILE

AMERICAN SOCIETY OF AGRICULTURAL SCI¬
ENCES, CHILEAN CHAPTER, Departamento de
Genetica Fitotecnica, Ministerio de Agricul-
tura, Santiago.

INSTITUTO BACTERIOLOGICO DE CHILE, Bor-
gono 1470, Santiago.

INSTITUTO BIOLOGICO Y ESTACION EXPERI¬
MENTAL DE LA SOCIEDADE NACIONAL DE
agricultura, F. Lazcano 1502, Santiago.

INSTITUTO PANAMERICANO DE INGENIERIA DE
MINAS Y geologia, Casilla 9228, Calle Aus-
tinas 1111, Santiago.

MUSEO araucano, Temuco.

MUSEO NACIONAL DE HISTORIA NATURAL,
Quinta Normal, Santiago.

OBSERVATORIO ASTRONOMICO, El Bosque, Cis-
terna, Santiago.

UNIVERSIDAD CATOLICA DE CHILE, Santiago.
Astronomy, biological and physical sciences,
mathematics, medicine.

UNIVERSIDAD DE chile, Santiago.

Biological and physical sciences, mathe¬
matics, marine biology, medicine, seis¬
mology.

UNIVERSIDAD DE CONCEPCION, Concepcion.
Mathematics, medicine, physical sciences.

This information was received from the Em¬
bassy of Chile, Washington, D. C; from the
American Embassy, Santiago; and from UNES¬
CO, Paris. February-April, 1948.

CHINA

ACADEMIA SINICA, Nanking.

Astronomy, geology, meteorology.

ACADEMIA SINICA, Shanghai.

Botany, chemistry, engineering, mathemat¬
ics, medicine, physics, psychology, zoology.

AURORA UNIVERSITY, Shanghai.

Medicine, science and engineering; Le
Musee Heude.

CATHOLIC UNIVERSITY OF PEIPING, Peiping.
Anthropology, biology, chemistry, physics,
science.

CHINA GEOGRAPHICAL RESEARCH INSTITUTE,
Nanking.

CHINESE ASSOCIATION FOR THE ADVANCEMENT
OF science [formerly Science Society of
China}, 235 Shensi Road (South), Shanghai.

CHINESE CHEMICAL society, c/o National Cen¬
tral University, Nanking.

CHINESE MEDICAL ASSOCIATION, 4l Tzeki Rd.,
Shanghai.

CHINESE PHYSICAL SOCIETY, c/o National
Academy of Peiping, Peiping.

CHINESE PUBLIC HEALTH ASSOCIATION, 179
Pei Hsia Road, Nanking.

GEOLOGICAL SOCIETY OF CHINA, 942 Chu-
kiang Road, Nanking.

LINGNAN UNIVERSITY, Canton.

Agriculture, biology, chemistry, medicine,
physics, science and engineering.
NATIONAL ACADEMY OF PEIPING, Peiping.
Botany, chemistry, geology, physics, physi¬
ology, zoology.

NATIONAL ACADEMY OF PEIPING, Shanghai.

Materia medica, radium.

NATIONAL ACADEMY OF PEIPING, Wukang,
Shensi.

Botanical survey of northwest China.
NATIONAL AMOY UNIVERSITY, Amoy.

Fisheries, oceanography, science and engi¬
neering.

Scientific Institutions- — BUSHNELL

249

NATIONAL ANWHEI UNIVERSITY, Anking.

Agriculture , science.

NATIONAL CENTRAL UNIVERSITY, Nanking.

General.

NATIONAL CHANGCHUN UNIVERSITY, Chung-
chun.

Agriculture, engineering, medicine.
NATIONAL CHEKIANG UNIVERSITY, Hangchow.

Agriculture, engineering, science.
NATIONAL CHIAO TUNG UNIVERSITY, Shanghai.
Engineering, science.

NATIONAL CHI NAN UNIVERSITY, Shanghai.
Science.

NATIONAL CHUNG CHEN UNIVERSITY, Nan-
chang, Kiangsi.

Agriculture, engineering, medicine, science.

NATIONAL CHUNGKING UNIVERSITY, Chung¬
king.

Arts and science, engineering, medicine.
NATIONAL FUH TAN UNIVERSITY, Shanghai.
Agriculture, science.

NATIONAL HONAN UNIVERSITY, Kaifeng.

Agriculture, engineering, medicine, science.
NATIONAL HUNAN UNIVERSITY, Changsha.
Engineering, science.

NATIONAL KWANGSI UNIVERSITY, Kweilin.

Agriculture, science and engineering.
NATIONAL KWEICHOW UNIVERSITY, Kweiyang.

Agriculture, arts and science, engineering.
NATIONAL LANCHOW UNIVERSITY, LandlOW.
Arts and science, medicine, veterinary med¬
icine.

NATIONAL NAKAI UNIVERSITY, Tientsin.
Science.

NATIONAL NORTHEASTERN UNIVERSITY, Muk¬
den.

Agriculture, engineering, science.
NATIONAL NORTHWESTERN UNIVERSITY, Sian,
Shensi.

Medicine, science.

NATIONAL PEI YANG UNIVERSITY, Tientsin.

Engineering, science.

NATIONAL PEKING UNIVERSITY, Peiping.
General.

NATIONAL SHANGHAI MEDICAL COLLEGE,
Shanghai.

Biochemistry , medicine, pathology, pharma¬
cology.

NATIONAL SHANSI university, Taiyuan.

Engineering, medicine.

NATIONAL SHANTUNG UNIVERSITY, TsingtaO.
Agriculture, engineering, medicine, ocean¬
ography, science.

NATIONAL SUNYATSEN UNIVERSITY, Canton.
General.

NATIONAL SZECHWAN UNIVERSITY, ChengtU.

Agriculture, science and engineering.
NATIONAL TAIWAN UNIVERSITY, Taipei.

Agriculture, engineering, medicine, science.
NATIONAL TSING HUA UNIVERSITY, Peiping.
General.

NATIONAL TUNG CHI UNIVERSITY, Shanghai.

Engineering, medicine.

NATIONAL WUHAN UNIVERSITY, Wuchang.
General.

NATIONAL YIN SHIH UNIVERSITY, Kinghua,
Chekiang.

Agriculture, arts and science, engineering.
NATIONAL YUNAN UNIVERSITY, Kunming.

Agriculture, engineering, medicine, science.
NATURAL SCIENCE SOCIETY OF CHINA, c/o Na¬
tional Central University, Nanking.

PEIPING UNION MEDICAL COLLEGE, Peiping.

Medicine, public health.

UNIVERSITY OF NANKING, Nanking.

Agriculture, science.

UNIVERSITY OF SHANGHAI, Shanghai.

Science.

YENCHING UNIVERSITY, Peiping.

Science.

The information for this selected list was
received through the American Embassy at
Nanking, China, from the office of Dr. Han
Ch’ing-lien, Director of the Department of Cul¬
tural Relations of the Ministry of Education in
Nanking. The very great number of scientific
institutions in China makes it impossible to
include all of them in this presentation: only
the major institutions — most of the national
universities and some of the established private
universities and scientific associations — have
been listed here. The special qualifications noted
for them are generalizations based upon the
colleges, institutes, or departments which Dr.
Han’s office has indicated are found at the
respective institutions. In the cases of the
largest universities, which possess most of the
facilities and curricula offered by large univer¬
sities the world over, the word General has been
used to denote this fact.

Further information about the institutions
listed here, or about the numerous smaller col¬
leges and provincial universities, societies, and
institutes throughout China which are not con¬
sidered in this list, can be obtained from the
helpful office of Dr. Han Ch’ing-lien in Nan¬
king.

250

PACIFIC SCIENCE, Vol. II, October, 1948

COLOMBIA

CENTRO NACIONAL DE INVESTIGACIONES DEL
CAFE, Chinchina, Caldas.

Coffee technology.

COLLEGIO DE SAN JOSE, Medellin, Antioquia.

General; Museum of Natural Sciences.

DEPARTAMENTO DE tierras, Ministerio de la
Economia Nacional, Bogota.

Forestry and soil conservation.

DEPARTAMENTO TECNICO DE LA FEDERACION
NACIONAL DE CAFETEROS, Bogota.

Coffee technology.

divisiones TECNICAS, Ministerio de Higiene,
Bogota.

Public health.

ESCUELA NACIONAL DE MINAS, Medellin, An¬
tioquia.

Mining and associated sciences.

ESCUELA NORMAL SUPERIOR, Bogota.

General.

ESTACION AGRICOLA EXPERIMENTAL DE AR-
mero, Armero, Tolima.

Agricultural research.

ESTACION AGRICOLA EXPERIMENTAL DE PAL¬
MIRA, Palmira, Valle.

Agricultural research.

ESTACION AGRICOLA EXPERIMENTAL DE SE¬
VILLA, Sevilla, Magdalena.

Agricultural research.

FACULTAD DE AGRONOMIA DE LA UNIVERSIDAD
NACIONAL, Medellin, Antioquia.

Agronomy, agriculture.

GRANJA GANADERA DE MONTERIA, Monteria,
Bolivar.

Veterinary parasitology.

INSTITUTO DE BIOLOGIA, Departamento Na¬
cional de Agricultura, Institute Ciencias Na-
turales de la Ciudad Universitaria, Apartado
Postal 2535, Bogota.

Botany, entomology, phytopathology.

INSTITUTO DE CIENCIAS NATURALES DE LA
UNIVERSIDAD NACIONAL DE COLOMBIA, Ciu¬
dad Universitaria, Apartado Postal 2535,
Bogota.

Botany, entomology, ornithology.

INSTITUTO DE ESTUDIOS ESPECIALES "CARLOS
FINLAY,” Bogota.

Medical entomology, virology.

INSTITUTO DE LA SALLE, Apartado 475, Bogota.

Natural sciences.

INSTITUTO DE MALARIOLOGIA, Bogota.

Malariology, parasitology.

INSTITUTO GEOFISICO DE LOS ANDES COLOM-
bianos, Apartado 270, Bogota.

Geophysics, meteorology.

INSTITUTO NACIONAL DE HIGIENE "SAMPER
MARTINEZ,” Bogota.

Chemotherapeutics, pharmacology, public

health.

LABORATORIO CENTRAL DE INVESTIGACION DE
lepra, Bogota.

Leprosy.

LABORATORIO CUP (CESAR URIBE PIEDRA-
HITA ) , Bogota.

Bacteriology, chemistry, parasitology, phar¬
macodynamics.

LABORATORIO QUIMICO NACIONAL DE ANALI-
sis E INVESTIGACION, Ministerio de Minas y
Petroleos, Bogota.

Geochemistry, geophysics.

OBSERVATORIO ASTRONOMICO NACIONAL, Bo¬
gota.

Astronomy, meteorology.

UNIVERSIDAD de ANTIOQUIA, Apartado 217,
Medellin, Antioquia.

General.

UNIVERSIDAD NACIONAL DE COLOMBIA, Ciudad
Universitaria, Bogota.

General.

This list was selected from a Directorio Cienti-
fico Colombiano, prepared by Dr. Enrique Perez
Arbelaez at the direction of the Ministry of
National Education, Colombia, and sent to us
by the American Embassy, Bogota, in January,
1948.

Scientific Institutions — BUSHNELL

251

COSTA RICA

DEPARTAMENTO NACIONAL DE AGRICULTURA,
San Jose.

Agriculture, botany.

INSTITUTO INTERNACIONAL DE CIENCIAS AGRI¬
COLAS, Turrialba.

Agricultural research.

INSTITUTO NACIONAL GEOGRAFICO, San Jose.

UNIVERSIDAD DE COSTA RICA, San Jose.
General.

This information was received from the Em¬
bassy of Costa Rica, Washington, D. C.; from
the American Embassy, San Jose; and from
UNESCO, Paris. February-April, 1948.

ECUADOR

UNIVERSIDAD CATOLICA, Quito.
UNIVERSIDAD CENTRAL, Quito.
UNIVERSIDAD DE CUENCA, Cuenca.
UNIVERSIDAD DE GUAYAQUIL, Guayaquil.
UNIVERSIDAD DE LOJA, Loja.

This information was received from the Em¬
bassy of Ecuador, Washington, D. C., and from
the American Embassy, Quito. January, 1948.

FIJI ISLANDS

AGRICULTURAL department, Fiji Government,
Suva. (Address the Director of Agriculture.)
["Office accommodation and library facili¬
ties ... for one or two visiting scientists
who are not concerned with laboratory
studies . . .”]

medical DEPARTMENT, Fiji Government, Suva.
(Address the Director of Medical Services.)
CENTRAL LEPER HOSPITAL, Makogai.
PATHOLOGICAL LABORATORY, Suva.
["Facilities . . . can readily be expanded
to meet the requirements of special
research projects. Visiting research
workers would be welcome so far as
limitations of space allow . . .”]
tuberculosis hospital, Tamavua.

This information was received from the Director
of Agriculture and from the Director of Med¬
ical Services, Fiji Government, Suva, through
the American Consulate, Suva; and from the
British Consulate, Honolulu, T. H. February-
March, 1948.

GUATEMALA

DIRECCION GENERAL DE AGRICULTURA, De-
partamentos de Sanidad Vegetal y Plantas
Medicinales, Guatemala City.

INSTITUTO DE MINERIA E HIDROCARBUROS,
Guatemala City.

Mining, soil science.

JARDIN BOTANICO, Paseo "La Reforma,” Gua¬
temala City.

OBSERVATORIO NACIONAL METEOROLOGICO, La
Aurora.

Meteorology.

SOCIEDAD DE GEOGRAFIA E HISTORIA, Guate¬
mala City.

This information was received from the Guate¬
malan Embassy, Washington, D. C, and from
UNESCO, Paris. January- April, 1948.

252

PACIFIC SCIENCE, Vol. II, October, 1948

HONDURAS

ASOCIACION MEDICA HONDURENA, Teguci¬
galpa.

Medical research.

ASOCIACION DE QUIMICA Y FARMACIA DE HON¬
DURAS, Tegucigalpa.

Chemistry and pharmacology.

LANCETILLA EXPERIMENT STATION OF THE
TELA RAILROAD COMPANY, La Lima.

USD A and United Fruit Company research
in rubber and other plant crops.

SOCIEDAD DE ANTROPOLOGIA, Tegucigalpa.
Anthropology.

SOCIEDAD DE GEOGRAFIA E HISTORIA DE HON¬
DURAS, Avenida Cervantes, Tegucigalpa.
Geography and history.

This information was received from the Hon¬
duran Ministry of Education, through the Amer¬
ican Embassy, Tegucigalpa, and from UNESCO,
Paris. January-April, 1948.

INDO-CHINA

ecole francaise d’extreme-orient, 26 Bou¬
levard Carreau, Hanoi.

Anthropology, archaeology, ethnology.

INSTITUT DE BIOLOGIE, Saigon.

INSTITUT OCEANOGRAPHIQUE DE NHATRANG,
Nhatrang, Annam.

INSTITUT PASTEUR, in Dalat, Hanoi, Nhatrang,
and Saigon.

Medicine, public health.

INSTITUT DE RECHERCHES AGRONOMIQUE DE
SAIGON, 58 Rue du Docteur Angier, Saigon.

STATION D’ESSAI D ’AGRICULTURE, Bias.
UNIVERSITE DE HANOI, 11 Boulevard Bobillot,
Hanoi.

General.

UNIVERSITE DE saigon, Saigon.

Medicine, natural and physical sciences.

This information was received from the Em¬
bassy of France, Washington, D. C., and New
York; from the French Consulate, Honolulu,
T. H.; and from the American Consulate at
Hanoi, Indo-China. February, 1948.

JAPAN

AGRICULTURAL EXPERIMENT STATION, Minis¬
try of Agriculture and Forestry, Tokyo.

AGRICULTURAL RESEARCH INSTITUTE, Ofuna.

APPLIED SCIENCE RESEARCH INSTITUTE, Kyoto.

brewing experiment station, Ministry of
Finance, Tokyo.

CANNING INSTITUTE OF JAPAN, Tokyo.

CHROMOSOMES RESEARCH INSTITUTE, Yama-
nishi-ken.

ELECTRONIC SCIENCE RESEARCH INSTITUTE,
Osaka.

FERMENTATION RESEARCH INSTITUTE, Ministry
of Commerce and Industry, Chiba.

FERTILIZER RESEARCH INSTITUTE, Tokyo.

FISHERIES EXPERIMENT STATION, Tokyo.

FOOD CHEMISTRY RESEARCH INSTITUTE, Osaka.

FOOD RESEARCH institute, Ministry of Agri¬
culture and Forestry, Tokyo.

FORESTRY EXPERIMENT STATION, Tokyo.

FUEL RESEARCH INSTITUTE, Ministry of Com¬
merce and Industry, Saitama-ken.

FURUKAWA PHYSICAL AND CHEMICAL RE¬
SEARCH INSTITUTE, Tokyo.

GEOLOGICAL SURVEY, Tokyo.

GOVERNMENT CHEMICAL AND INDUSTRIAL RE¬
SEARCH laboratory, Ministry of Commerce
and Industry, Tokyo.

GOVERNMENT HYGIENIC LABORATORY, Ministry
of Welfare, Tokyo.

GOVERNMENT RESEARCH INSTITUTE FOR CERA¬
MICS, Ministry of Commerce and Industry,
Kyoto.

HAKODATE FISHERIES COLLEGE, Hakodate.

HATTORI FOUNDATION FOR BOTANICAL RE¬
SEARCH, Miyazaki-ken.

HIROSHIMA COLLEGE OF SCIENCE AND LITERA¬
TURE, Hiroshima.

General sciences, marine biological station,
theoretical physics institute.

Scientific Institutions — BUSHNELL

253

HITACHI CENTRAL LABORATORY, Tokyo.

Chemical, electrical, and mechanical tech¬
nology.

HOKKAIDO FISHERIES EXPERIMENT STATION,
Yoichi.

HOKKAIDO FORESTRY EXPERIMENT STATION,
Sapporo.

HOKKAIDO INDUSTRIAL RESEARCH INSTITUTE,
Sapporo.

HOKKAIDO UNIVERSITY, Sapporo.

Agricultural sciences, medicine, techno¬
logical sciences; Akkeshi Marine Biological
Station, Algological Research Laboratory,
Institute of Low Temperature Research.

HORTICULTURAL EXPERIMENT STATION, Shi-
zuoka-ken.

INDUSTRIAL MATERIALS RESEARCH LABORA¬
TORY, Fukuoka-ken.

IWATA INSTITUTE OF PLANT BIOCHEMISTRY,
Tokyo.

KEIO UNIVERSITY, Tokyo.

Engineering, medicine, science.

KIHARA INSTITUTE FOR BIOLOGICAL RESEARCH,
Kyoto.

KOBAYASHI BACTERIOLOGICAL LABORATORY,
Fukushima-ken.

KOBAYASHI INSTITUTE OF PHYSICAL RESEARCH,
Tokyo.

KYOTO MEDICAL COLLEGE, Kyoto.

KYOTO UNIVERSITY, Kyoto.

Agriculture, engineering, medicine, science;
Beppu V olcanological Hot Springs Insti¬
tute, Otsu Hydrobiological Station, Seto
Marine Biological Station; Institutes of
Chemical Research, Engineering Research,
and Wood Research.

KYUSHU UNIVERSITY, Fukuoka.

Agriculture, engineering, medicine, science;
Amakusa Laboratory for Marine Biology,
Hikosan Biological Research Laboratory,
Institute for Wood Research.

MEGURO RESEARCH INSTITUTE, Tokyo.

Biology, chemical technology.

NAGOYA RESEARCH INSTITUTE OF SCIENCE AND
INDUSTRY, Nagoya.

NAGOYA UNIVERSITY, Nagoya.

Engineering, science; marine biological sta¬
tion.

NATIONAL RESEARCH COUNCIL, Tokyo.

Special committees for all fields of research.

NATURAL RESOURCES RESEARCH INSTITUTE,
Tokyo.

NIHON UNIVERSITY, Tokyo.

Agriculture, engineering.

NOGUCHI RESEARCH INSTITUTE, Tokyo.
Medical microbiology.

NOHARA RESEARCH INSTITUTE, Nagoya.
Biochemistry, chemical technology.

OCEANOGRAPHIC CHEMISTRY INSTITUTE,
Kyoto.

OHARA INSTITUTE FOR AGRICULTURAL RE¬
SEARCH, Okayama.

OSAKA INDUSTRIAL RESEARCH LABORATORY,
Ministry of Commerce and Industry, Osaka.

OSAKA UNIVERSITY, Osaka.

Engineering, medicine, science; Institute of
Scientific and Industrial Research.

PHYSICAL AND CHEMICAL RESEARCH INSTI¬
TUTE, Tokyo.

PHYSICAL INSTITUTE FOR RADIO WAVES, Tokyo.

SANKYO company, Tokyo.

Biochemistry, chemical technology.

SCIENTIFIC FISHERIES INSTITUTE, Hakodate.
SERICULTURE EXPERIMENT STATION, Tokyo.
TEA EXPERIMENT STATION, Shizuoka-ken.

TOHOKU UNIVERSITY, Sendai.

Engineering, medicine, science; Asamushi
Marine Biological Station, Mukaiyama Ob¬
servatory, Oceano chemistry Laboratory; In¬
stitutes of Agricultural Research, Glass,
Metallurgy, and Scientific Measurements.

TOKUGAWA BIOLOGICAL INSTITUTE, Tokyo.

TOKYO COLLEGE OF AGRICULTURE AND FOR¬
ESTRY, Tokyo.

TOKYO INSTITUTE OF TECHNOLOGY, Tokyo.

TOKYO university, Tokyo.

Agriculture and fisheries, enginering, med¬
icine, science; Botanical Garden, Misaki
Marine Biological Station; Institutes of As¬
tronomy, Earthquake Research, Radiation
Chemistry, Physiographic Research, Science
and Technology.

TOKYO UNIVERSITY OF LITERATURE AND SCI¬
ENCE, Tokyo.

Science; Shimoda Marine Biological Lab¬
oratory, Sugadaira High Land Biological
Laboratory.

YAMASHINA ORNITHOLOGICAL INSTITUTE,
Tokyo.

WASEDA UNIVERSITY, Tokyo.

Science and engineering.

254

PACIFIC SCIENCE, Vol. II, October, 1948

This information was taken from Report No.
12 [Index to Japanese Research and Develop¬
ment during Period July-Dee ember, 1 946, pub¬
lished April, 1948] of General Headquarters,
Supreme Commander for the Allied Powers,
Economic and Scientific Section. This Index
"lists the detailed subjects of research projects
pursued in Japan during the last half of 1946
in each field of science and technology, together
with the names of the laboratories which did
the work . . The selected list presented here
by Pacific Science is considerably abridged, and
shows only the major institutions in Japan —
as best as can be judged from the SCAP Index
— at which scientific research may be carried on.
Many other institutions — like the prefectural
experimental stations and colleges — are not

listed here; and it is possible, according to the
preface of the SCAP report, that some of the
finest small laboratories in Japan have been
omitted, unintentionally, because information
about them was either unobtainable or because
awareness of their quality was lost when their
appraising was reduced to the statistics of the
report.

Requests for information about institutions
or facilities in Japan, and concerning their ac¬
cessibility to visiting scientists, should be sent
to General Headquarters, Supreme Commander
for the Allied Powers, Economic and Scientific
Section: attention Scientific and Technical
Division, APO 500, c/o Postmaster, San Fran¬
cisco, California.

KOREA

(South Korea Interim Government)

PUSAN FISHERIES COLLEGE, Pusan.

SEOUL NATIONAL UNIVERSITY, Seoul.

Arts and science, engineering, medicine,
veterinary medicine.

SEOUL NATIONAL UNIVERSITY, Suwon.
Agriculture.

SEVERANCE MEDICAL COLLEGE, Seoul.
Medicine, nursing, public health.

This information was received from the Depart¬
ment of Education, South Korea Interim Gov¬
ernment, USAMGIK, APO 235-2, c/o Post¬
master, San Francisco, California. March, 1948.

MEXICO

INSTITUTO DE BIOLOGIA APPLICADA, Ignacio
Mariscal 155, Mexico, D. F.

INSTITUTO DE CARDIOLOGIA, Calzada de la Pie-
dad 300, Mexico, D. F.

INSTITUTO NACIONAL DE ANTROPOLOGIE HIS-
TORIA, Cordoba 73, Mexico, D. F.

INSTITUTO PANAMERICANO DE GEOGRAFIA E
historia, Observatoria 192, Tacubaya, D. F.

INSTITUTO DE SALUBRIDAD Y ENFERMEDADES
TROPICALES, Plan de San Luis y Prolongacion
Carpio, Mexico, D. F.

OBSERVATORIO ASTRONOMICO NACIONAL, Ta-
cubaya, D. F.

SOCIEDAD BOTANICA DE MEXICO, Morelos 61,
Mexico, D. F.

SOCIEDAD DE ESTUDIOS ASTRONGMICOS Y GEO-
fisicos, Av. Observatorio 192, Tacubaya,
D. F.

SOCIEDAD MEXICANA DE ESTUDIOS SOBRE TU¬
BERCULOSIS, Apartado Postal 2425, Mexico,
D. F.

UNIVERSIDAD NACIONAL AUTONOMA:

INSTITUTO DE BIOLOGIA, Casa del Lago, Cha-
pultepec, Mexico, D. F.

instituto de fisica, Tacuba 5, Mexico,
D. F.

INSTITUTO DE GEOGRAFIA, Lie. Verdad No.
3, Mexico, D. F.

instituto de geologia, Cipres 176, Mexico,

D. F.

INSTITUTO DE MATEMATICAS, Tacuba 5,
Mexico, D. F.

INSTITUTO DE QUIMICA, Santa Cruz Atenco,
Tacuba, D. F.

This information was received from the [Mexi¬
can] Commission for the Promotion and
Coordination of Scientific Research, through
the American Embassy, Mexico, D. F. February,
1948.

Scientific Institutions — BlJSHNELL

255

NETHERLANDS INDIES

ALGEMEEN PROEFSTATION DER ALGEMENE VER-
ENIGING RUBBERPLANTERS SUMATRA OOST-
KUST [CENTRAL EXPERIMENT STATION,
GENERAL ASSOCIATION OF RUBBER PLANT¬
ERS OF THE EAST COAST OF SUMATRA], Me¬
dan, Sumatra.

Rubber research.

ALGEMEEN PROEFSTATION VOOR DEN LAND-
BOUW [GENERAL AGRICULTURAL EXPERI¬
MENT station], Buitenzorg, Java, with a
branch station at Makassar, Celebes.

Agricultural technology , botany, phyto¬
pathology, soils.

BOSBOUWPROEFSTATION [GOVERNMENT FOR¬
EST RESEARCH INSTITUTE], Buitenzorg, Java.

CENTRAAL BUREAU VOOR TECHNISCHE ONDER-
ZOEKINGEN [CENTRAL TECHNOLOGICAL

laboratories], Karanganjar 55, Batavia,
Java.

CENTRALE VERENIGING TOT BEHEER VAN
PROEFSTATIONS VAN DE OVERJARIGE CUL¬
TURES IN NEDERLANDSCH-INDIE [ASSOCIA¬
TION OF COMBINED EXPERIMENT STATIONS
FOR PERENNIAL CROPS IN THE NETHER¬
LANDS indies], Central Office, Batavia, Java,
with experiment stations at Buitenzorg,
Djember, Klaten, and Malang, Java, and at
Medan, Sumatra.

Agriculture and agronomy, especially of
tobacco.

DIENST VAN DEN MIJNBOUW [DEPARTMENT
OF mines], Molenoliet Oost, Batavia, Java.

Head Office, and section for economic
geology.

DIENST VAN DEN MIJNBOUW [DEPARTMENT
OF mines], Wilhelmina Boulevard 7, Ban¬
doeng, Java.

Chemistry, engineering geology, geological
exploration, geological museum, paleonto¬
logy, and volcanology.

eijkman INSTITUTE [Central Laboratory of
the Public Health Service of the Netherlands
Indies], Oranje Boulevard 69, Batavia, Java.
Medicine, nutrition, public health.

GEWESTELIJK LABOR ATORIUM VAN DEN DIENST
DER VOLKSGEZONDHEID [REGIONAL LABORA¬
TORY OF THE PUBLIC HEALTH SERVICE],
Makassar, Celebes.

INSTITUUT VOOR LEPRA ONDERZOEK [LEPROSY
RESEARCH institute], Bandoeng, Java.

INSTITUUT PASTEUR, Pasteursweg, Bandoeng,
Java.

INSTITUUT VOOR DE ZEEVISSERIJ [FISHERIES
RESEARCH institute], Pasar Ikan, Batavia,
Java.

KON. BATAVIAASCH GENOOTSCHAP VAN KUN-
STEN EN WETENSCHAPPEN [ROYAL BATAV¬
IAN SOCIETY OF ARTS AND SCIENCES], Kon-
ingsplein West 12, Batavia, Java.

KON. MAGNETISCH EN METEOROLOGISCH OB-
SERVATORIUM [ROYAL METEOROLOGICAL

observatory], Engelsche Kirkweg 3, Bata¬
via, Java.

KON. NATUURKUNDIGE VERENIGING IN NEDER¬
LANDSCH-INDIE [ROYAL SCIENCE SOCIETY],
Koningsplein Zuid 11, Batavia, Java.

LABORATORIUM VOOR DE BINNENVISSERIJ
[LABORATORY FOR INLAND FISHERIES], Bui¬
tenzorg, Java.

LABORATORIUM VOOR SCHEIKUNDIG ONDER¬
ZOEK [LABORATORY FOR CHEMICAL RE¬
SEARCH], Buitenzorg, Java.

LABORATORIUM VOOR TECHNISCHE HYGIENE
[LABORATORY FOR TECHNICAL HYGIENE],
School weg, Bandoeng, Java.

S LANDS PLANTENTUIN [GOVERNMENT BO¬
TANICAL GARDENS], Batavia, Java.

Laboratorium voor het Onderzoek der Zee
{ Laboratory for Marine Biology }.

S LANDS PLANTENTUIN [GOVERNMENT BO¬
TANICAL gardens], Buitenzorg, Java.

Herbarium en Museum voor systematische
Botanie {Herbarium and Museum for sys¬
tematic botany }, Treub Laboratorium
{ Visitors’ Laboratory), Zodlogisch Museum
{ Zoological Museum).

S LANDS PLANTENTUIN [GOVERNMENT BO¬
TANICAL GARDENS], Tjibodas, Java.

Bergtuin Tjibodas { Tjibodas Mountain
Garden ).

MILITAIR HYGIENISCH INSTITUUT [MILITARY
HYGIENE institute], Hospitalsweg 24, Ba¬
tavia, Java.

NEDERLANDS INDISCH INSTITUUT VOOR RUB-
BERONDERZOEK [NETHERLANDS INDIES IN¬
STITUTE FOR rubber research], Buiten¬
zorg, Java.

NEDERLANDS INDISCH STERRENKUNDIGE VERE¬
NIGING [NETHERLANDS INDIES ASTRONO¬
MICAL society] Oude Hospital w eg 14,
Bandoeng, and Bosscha Sterrenwacht, Lem-
bang, Java.

PROEFSTATION VAN DE JAVA-SUIKERINDUSTRIE
[EXPERIMENTAL STATION FOR THE JAVA
sugar INDUSTRY], Passoeroean, Java.

TOPOGRAFISCHE DIENST [TOPOGRAPHICAL SUR¬
VEY], Goenoeng Sahari 89, Batavia, Java.

UNIVERSITEIT VAN INDONESIE [UNIVERSITY OF
INDONESIA], Bandoeng, Java.

Technical sciences and mathematics.

256

PACIFIC SCIENCE, Vol. II, October, 1948

UNIVERSITEIT VAN INDONESIE [UNIVERSITY OF
INDONESIA], Batavia, Java.

Medicine.

UNIVERSITEIT VAN INDONESIE [UNIVERSITY OF
INDONESIA], Buitenzorg, Java.

Agricultural sciences , veterinary sciences.
VEEARTSENIJKUNDIG INSTITUUT [VETERINARY
RESEARCH INSTITUTE], Buitenzorg, Java.
WATERLOOPKUNDIG LABOR ATORIUM [HYDRO¬
DYNAMICS laboratory], Hogeschoolweg,
Bandoeng, Java.

This information was received from Organisatie
Natuurwetenschappelijk Onderzoek i. o. (Or¬
ganization for Scientific Research in the Nether¬
lands Indies, prep, comm.), Koningsplein Zuid
11, Batavia, Java; from the book, Science and
Scientists in the Netherlands Indies , edited by
Pieter Honig and Frans Verdoorn, and pub¬
lished in 1945 in New York City by the Board
for the Netherlands Indies, Surinam, and
Curasao; and from the American Consulate
General, Batavia, Java. March, 1948.

NEW CALEDONIA

INSTITUT francais d’oceanie, Noumea, New
Caledonia.

Agricultural entomology, ethnology, geol¬
ogy, marine biology, phytopathology, phys¬
ical oceanography, and others to be added.
["The purpose of the Institute is . . . research
of a scientific nature in the French regions of the
Pacific, and a thorough collaboration in all
research with foreign scientists to corroborate

observations of general interest. The first pro¬
gram was formed at the suggestion of the
South Pacific Commission.” — From a letter
from the Director of the Institute.]

This information was received from the Di¬
rector of the Institute, through the American
Consulate at Noumea, New Caledonia. February,
1948.

NEW ZEALAND

AGRICULTURE department, Head Office, Do¬
minion Farmers’ Institute Building, Welling¬
ton, C. 1. (Communications should be ad¬
dressed to the Director-General of Agricul¬
ture. )

RUAKURA ANIMAL RESEARCH STATION,
Ruakura, South Auckland Province.
RUKUHIA SOIL FERTILITY RESEARCH
STATION, Rukuhia, South Auckland
Province.

SEED TESTING station, Palmerston
North.

WALLACEVILLE ANIMAL RESEARCH
STATION, Wallaceville, Wellington.
AUCKLAND MUSEUM, Auckland.

{ Facilities for visiting scientists are not
available at present.)

CANTERBURY MUSEUM, Christchurch.

Biology, ethnology.

CARTER OBSERVATORY, Wellington.

CAWTHRON INSTITUTE, Nelson.

Entomology, plant problems, soil.
DOMINION MUSEUM, Wellington.

Natural sciences.

HAWKES BAY MUSEUM, Napier.

Maori ethnology; Russell Duncan Library
of exploration in New Zealand, Pacific,
and Antarctic.

LEATHER AND SHOE RESEARCH ASSOCIATION,
111 Sydney Street, Wellington.

MARINE DEPARTMENT, Head Office, T. & G.
Building, Lambton Quay, Wellington.

MARINE DEPARTMENT, Fishery Research Lab¬
oratory, Clark’s Building, Wingfield Street,
Wellington, N. 1.

NEW ZEALAND FERTILISER MANUFACTURERS’
RESEARCH ASSOCIATION, (INC.), P. O. Box
2080, Auckland.

NEW ZEALAND POTTERY AND CERAMICS RE¬
SEARCH ASSOCIATION, (INC), 111 Sydney
Street, Wellington.

NEW ZEALAND WOOLEN MILLS RESEARCH
association, (INC.), c/o Prof. E. G. Soper,
Chemistry Department, University of Otago,
Dunedin.

OTAGO MUSEUM, Dunedin.

Ethnology of New Zealand and of Pacific
Islands, especially of New Hebrides.

SCIENTIFIC AND INDUSTRIAL RESEARCH DE¬
PARTMENT [Dominion Government]:

AGRONOMY DIVISION, Private Bag,
Christchurch.

APIA OBSERVATORY, Apia, Western
Samoa.

Research on terrestrial magnetism,
tidal observations, and seismology.

Scientific Institutions — BUSHNELL

257

ATOMIC PHYSICS SECTION, 37 Majori-
banks Street, Wellington.

AUCKLAND INDUSTRIAL DEVELOPMENT
LABORATORIES, P. O. Box 2225,
Auckland.

BIOMETRICS SECTION, 37 Majoribanks
Street, Wellington.

botany division, 8 The Terrace, Wel¬
lington.

DEFENSE DEVELOPMENT SECTION, P. O.
Box 1152, Christchurch.

DOMINION LABORATORIES:

Durham Street West, Auckland.

P. O. Box 1290, Christchurch.

P. O. Box 562, Dunedin.

Ill Sydney Street, Wellington.

dominion OBSERVATORY, Kelburn,
Wellington.

DOMINION physical laboratory,
Gracefield Road, Lower Hutt.

ECONOMIC SURVEYS, 117 Sydney Street,
Wellington.

ENTOMOLOGY DIVISION, Cawthron In¬
stitute, Nelson.

FATS RESEARCH LABORATORY, Sydney
Street, Wellington.

FLAX RESEARCH STATION, Foxton.

GEOLOGICAL survey, Head Office, 156
The Terrace, Wellington.

geological survey, Volcanological
Laboratory, Rotorua.

GRASSLANDS DIVISION, P. O. Box 16, Pal¬
merston North.

HEAD OFFICE, 117 Sydney Street, Wel¬
lington. (Address the Secretary.)

INDUSTRIAL PSYCHOLOGY DIVISION,
Bowen Street, Wellington.

INFORMATION BUREAU, 117 Sydney
Street, Wellington.

IONOSPHERE SECTION, Christchurch.

MAGNETIC OBSERVATORY, Hagley Park,
Christchurch.

PLANT CHEMISTRY LABORATORY, P. O.
Box 16, Palmerston North.

PLANT DISEASES division, Private Bag,
Auckland.

SOIL BUREAU, 54 Moles worth Street,
Wellington.

WILD LIFE SECTION, 117 Sydney Street,
Wellington.

STATE FOREST service, 23 Fitzherbert Terrace,
Wellington, N. 1. (Address the Director.)
UNIVERSITY of new ZEALAND, Bowen Street,
Wellington. (Address the Registrar.) [The
"governing body” of the Universities and Col¬
leges of the Dominion.]

AUCKLAND UNIVERSITY COLLEGE, Auck¬
land. (Address the Registrar.)
Architecture, botany, chemistry, ge¬
ology, marine biology, physics.
CANTERBURY AGRICULTURAL COLLEGE,
Lincoln, Canterbury. (Address the
Registrar. )

General.

CANTERBURY UNIVERSITY COLLEGE,
Christchurch. (Address the Rector.)
General; mountain botanical sta¬
tion.

MASSEY AGRICULTURAL COLLEGE, Pal¬
merston North. (Address the Prin¬
cipal. )

Agricultural research, dairy re¬
search, veterinary sciences.
UNIVERSITY OF OTAGO, Dunedin. (Ad¬
dress the Registrar.)

Bacteriology and public health,
dentistry, home science, medicine ,
mines and metallurgy.

VICTORIA UNIVERSITY COLLEGE, Wel¬
lington. (Address the Registrar.)
General.

This information was received from the Infor¬
mation Bureau of the Department of Scientific
and Industrial Research of the Dominion Gov¬
ernment of New Zealand, through the Legation
of New Zealand, Washington, D. C, and the
American Legation, Wellington, N. Z.; from
the Art Galleries and Museums Association of
New Zealand, Wellington; from the Registrars,
Principals, and Rectors of the several univer¬
sities and colleges making up the University of
New Zealand; and from UNESCO, Paris.
March-May 1, 1948.

NICARAGUA

COLLEGIO centro AMERICA, Granada. This information was received from the Amer-

INSTITUTO PEDAGOGICO DE VARONES, Managua, ican Embassy, Managua, D. N., Nicaragua.
D. N. March, 1948.

258

PACIFIC SCIENCE, Vol. II, October, 1948

PANAMA

GORGAS MEMORIAL LABORATORY, Apartado
1252, Panama. (Address the Director.)
Medical entomology.

INSTITUTO NACIONAL DE AGRICULTURA, Di-
visa. (Address the Director.)

Agricultural research.

MUSEO NACIONAL de panama, Apartado 662,
Panama. (Address the Director.)
Archaeology , entomology.

This information was received from the Amer¬
ican Embassy, Panama, Republic of Panama.
January, 1948.

PERU

ESCUELA SUPERIOR DE CIENCIAS PEDAGOGICAS,

Avenida Grau 429, Lima.

ESTACION EXPERIMENTAL AGRICOLA DEL
NORTE, Lambayeque.

ESTACION EXPERIMENTAL AGRICOLA DE LA
MOLINA (INSTITUTO DE ALTOS ESTUDIOS),
Lima.

FACULTAD DE CIENCIAS MEDICAS, Lima.
INSTITUTO DE GENETICA DEL ALGODON DE LA
SOCIEDAD NACIONAL AGRARIA, Apartado
350, Lima.

This information was received from the Peru¬
vian Embassy, Washington, D. G, and from
UNESCO, Paris. January-April, 1948.

THE PHILIPPINES

BUREAU OF ANIMAL INDUSTRY, Manila.
BUREAU OF FISHERIES, Manila.

BUREAU OF FORESTRY, Manila.

BUREAU OF HEALTH, Manila.

BUREAU OF HOSPITALS, Manila.

BUREAU OF MINES, Manila.

BUREAU OF PLANT INDUSTRY, Manila.
INSTITUTE OF science (formerly Bureau of
Science), Manila.

NATIONAL MUSEUM, Manila.

PHILIPPINE GENERAL HOSPITAL, Manila.
UNIVERSITY OF THE PHILIPPINES, Manila.
WEATHER BUREAU, Manila.

This information was received from the Insti¬
tute of Science, Manila, through the Philippine
Consulate-General, Honolulu, T. H. February,
1948.

EL SALVADOR

CENTRO NACIONAL DE AGRONOMIA, Santa
Tecla, Depto. La Libertad.

Agricultural research.

DIRECCION GENERAL DE SANIDAD, Calle Arce
No. 87, San Salvador.

Public health and tropical diseases.
UNIVERSIDAD AUTONOMA DE EL SALVADOR,
Ave. Cuscatlan y 2 a. Calle Poniente, San
Salvador.

Architecture and engineering, chemistry,
dentistry, medicine, pharmacy.

This information was received from the Em¬
bassy of El Salvador, Washington, D. C., and
from the American Embassy, San Salvador,
El Salvador. January, 1948.

SIAM

THE CHULALONGKON UNIVERSITY, Bangkok.
General.

DEPARTMENT OF AGRICULTURE EXPERIMENT
STATION, Bangkok.

Agricultural research.

LEPROSY CONTROL DEPARTMENT AND LEPER
hospital, Department of Public Health,
Bangkok.

PASTEUR INSTITUTE, Saladaeng, Bangkok.
Bacteriology, serology.

This information was received from the Royal
Siamese Embassy, Washington, D. G, and from
the American Embassy, Bangkok, Siam. Feb¬
ruary, 1948.

Scientific Institutions — BUSHNELL

259

No information available.

SIBERIA (U.S.S.R.)

UNITED STATES

[It would be impossible, of course, to name
every institution in the United States which
has some concern with science in the Pacific.
In this list only the major institutions in the
Pacific states of the United States and in the
territories of Alaska and Hawaii have been
noted. Information concerning the institutions
in Hawaii was taken principally from Pacific
Science 1: 1 19-126, 1947, and for Alaska and
the Pacific states from the catalogues of the
several institutions included in the list.]

ALASKA

UNIVERSITY OF ALASKA, Fairbanks.

General , especially archaeology, geophysics.

CALIFORNIA

CALIFORNIA ACADEMY OF SCIENCE, San Fran-

cisco.

General.

CALIFORNIA INSTITUTE OF TECHNOLOGY,
Pasadena.

General; Arms Laboratory of Geological
Sciences, Bridge Laboratory of Physics,
Chemical Engineering Laboratory, Experi¬
ment Station, Gates and Crellin Laboratory
of Chemistry, High-Potential Research
Laboratory, Hydrodynamics Laboratory,
Kellogg Laboratory of Radiation, Kerckhoff
Laboratories of the Biological Sciences,
Kerckhoff Marine Biological Laboratory
( at Corona del Mar), Mudd Laboratory
of the Geological Sciences, Robinson
Laboratory of Astrophysics, Seismological
Research Laboratory, and others.

COLLEGE OF THE PACIFIC, Stockton.

General; Pacific Marine Station of Biologi¬
cal Sciences at Dillon Beach.

POMONA COLLEGE, Claremont.

General; Marine Laboratory at Laguna
Beach.

STANFORD UNIVERSITY, Palo AltO.

General; Hopkins Marine Station at Paci¬
fic Grove; Natural History Museum.

STATE OF CALIFORNIA, DIVISION OF FISH AND
GAME, FISHERIES LABORATORY, Terminal
Island, San Pedro.

UNIVERSITY OF SOUTHERN CALIFORNIA, 3551
University Avenue, Los Angeles 7.

General; Allan Hancock Foundation for
Biological Sciences.

UNIVERSITY OF CALIFORNIA.

["The University of California is composed of
academic colleges, professional schools, divi¬
sions, departments of instruction, museums,
libraries, research institutes, bureaus, and foun¬
dations, and the University of California Press,
situated on eight different campuses throughout
the State, namely: Berkeley, Los Angeles, San
Francisco, Davis, Riverside, Mount Hamilton,
La Jolla, and Santa Barbara.” — Catalogue, Uni¬
versity of California, 1948.]

I. AT BERKELEY.

General; Anthropological Museum, Cali¬
fornia Museum of Vertebrate Zoology,
Crocker Radiation Laboratory, Institute
of Experimental Biology, Museum of
Paleontology.

II. AT LOS ANGELES.

General; Medical Department.

III. AT SAN FRANCISCO.

Dentistry, medicine, nursing, public health ;
Hooper Foundation for Medical Research.

IV. AT DAVIS.

Agricultural research, veterinary sciences.

V. AT RIVERSIDE.

Agricultural research; Citrus Experiment
Station.

VI. AT MOUNT HAMILTON.

Lick Observatory.

VII. AT LA JOLLA.

Scripps Institution of Oceanography.

U. S. FISH AND WILDLIFE SERVICE, Stanford
University, Palo Alto.

South Pacific Investigations in Ichthyology.

260

PACIFIC SCIENCE, Vol. II, October, 1948

HAWAII

BERNICE P. BISHOP MUSEUM, Honolulu 35.

Collection, preservation, and study of Ha¬
waiian and Pacific material in ethnology
and the natural sciences; departments and
collections of botany, entomology, mala¬
cology ( both terrestrial and marine),
marine zoology; large collections in ich¬
thyology, ornithology, and geology. Library,
chiefly on Pacific ethnology and natural
history.

CALIFORNIA PACKING CORPORATION, P. O.

Box 149, Honolulu 10.

Canning operations.

HAWAII MARINE LABORATORY, Coconut Island,
Oahu. {Communications should be addressed
to the Department of Zoology and Ento¬
mology, University of Hawaii, Honolulu 10.]
Marine biology, oceanography.

HAWAII NATIONAL park, U. S. Department of
the Interior, National Park Service, Hawaii
National Park, Hawaii.

Natural history.

HAWAIIAN PINEAPPLE COMPANY, LIMITED,
Honolulu 1.

Research in groiving and processing of
pineapples and other subtropical crops.

HAWAIIAN SUGAR PLANTERS' ASSOCIATION EX¬
PERIMENT station, 1527 Keeaumoku Street,
Honolulu 4.

Agriculture, botany and forestry, chemistry,
climatology, entomology, genetics, geology,
pathology, physiology and biochemistry,
sugar technology.

HAWAIIAN TUNA PACKERS, LIMITED, P. O.
Box 238, Honolulu.

Research in oceanic fisheries methods.

HAWAIIAN VOLCANO OBSERVATORY (a branch
of the U. S. Geological Survey collaborating
with the Hawaiian Volcano Research Associa¬
tion and the University of Hawaii), Hawaii
National Park, Hawaii.

Measurements and observations on active
volcanoes; research in the chemistry and
physics of volcanoes.

HAWAIIAN VOLCANO RESEARCH ASSOCIATION,
c/o University of Hawaii, Honolulu 10.
Research in physical processes of Hawaiian
volcanoes. (The Directors will receive ap¬
plications at any time from research in¬
vestigators holding doctorate degrees who
desire to pursue specialized studies in
Hawaii in seismology, volcanology, and
volcanological oceanography. Persons of
advanced grade holding fellowships from
other institutions may be assisted in travel

expenses and may be provided with appa¬
ratus or use of facilities. Inquiries should
be addressed to Dr. T. A. Jaggar, Scientific
Director, at University of Hawaii, Ho¬
nolulu 10, Hawaii.)

HONOLULU BOARD OF WATER SUPPLY, P. O.
Box 3410, Honolulu 1.

Bacteriology, chemistry, geology.

LIBBY, MCNEILL AND LIBBY, P. O. Box 1140,
Honolulu 7.

Pineapple production and processing.

PACIFIC CHEMICAL AND FERTILIZER COMPANY,

P. O. Box 48, Honolulu 10.

Agricultural chemistry.

PINEAPPLE RESEARCH INSTITUTE OF HAWAII,
P. O. Box 3166, Honolulu 2.

Agricultural engineering, chemistry, ento¬
mology, genetics, meteorology, plant path-
ology, plant physiology, as related to pro¬
duction of pineapple plants and fruit.

TERRITORIAL BOARD OF AGRICULTURE AND
FORESTRY, P. O. Box 3319, Honolulu 1.
Animal industry, entomology, fish and
game, forestry.

TERRITORIAL BOARD OF HEALTH, Honolulu.

Public health laboratories on major islands
of Haivaiian group.

U. S. BUREAU OF ENTOMOLOGY AND PLANT
QUARANTINE, University of Hawaii Campus,
P. O. Box 340, Honolulu 9.

Fruitfly investigations.

U. S. COAST AND GEODETIC SURVEY, Pacific
District Headquarters, 244 Federal Office
Building, Honolulu.

U. s. GEOLOGICAL SURVEY, Division of Surface
Waters, 225 Federal Office Building, Ho¬
nolulu.

u. S. GEOLOGICAL SURVEY, Ground Water Divi¬
sion, 333 Federal Office Building, Honolulu.

u. S. GEOLOGICAL SURVEY, Hawaiian Volcano
Observatory. See HAWAIIAN VOLCANO OB¬
SERVATORY.

U. S. WEATHER BUREAU, Federal Office Build¬
ing, Honolulu.

UNIVERSITY OF HAWAII, P. O. Box 18, Ho¬
nolulu 10.

General; Library; Marine Biological Labo¬
ratory; cooperative associations with U. S.
Bureau of Entomology and Plant Quar¬
antine, the Pineapple Research Institute,
the Hawaiian Sugar Planters’ Association
Experiment Station, the University of Ha¬
waii Agricultural Experiment Station, the
B. P. Bishop Museum, and many other
private and governmental agencies in Ha¬
waii.

Scientific Institutions — BUSHNELL

261

UNIVERSITY OF HAWAII AGRICULTURAL EX¬
PERIMENT station, P. O. Box 18, Honolulu
10.

Agricultural chemistry, agricultural engi¬
neering, agronomy, animal husbandry,
entomology, horticulture, nutrition, para¬
sitology, plant pathology, plant physiology,
poultry husbandry, vegetable crops.

OREGON

OREGON STATE COLLEGE, Corvallis.

General.

UNIVERSITY OF OREGON, Eugene.

General; Condon Museum of Geology,
Museum of Natural History, Oregon State
Museum of Anthropology.

UNIVERSITY OF OREGON, Portland.

Medicine, public health.

WASHINGTON

MUSEUM OF THE STATE OF WASHINGTON,
Seattle.

General, especially anthropology and ver¬
tebrate zoology.

STATE COLLEGE OF WASHINGTON, Pullman.
General.

UNIVERSITY OF WASHINGTON, Seattle 5.

General; Oceanographic Laboratories at
Seattle and Friday Harbor.

U. S. FISH AND WILDLIFE SERVICE, Mountlake
Laboratory, Seattle.

Juvenile Euthynnus lineatus and Auxis thazard from the Pacific Ocean off

Central America

Milner B. Schaefer and John C Marr1

In a previous paper (Schaefer and Marr, in
press), juvenile stages of two commercially
important tunas, Neothunnus macropterus
(Temminck and Schlegel ) and Katsuivonus
pelamis ( Linnaeus ) were described. They
were taken, with dip nets at night, under
flood lights, in the oceanic waters of the
Pacific off Costa Rica and northern Panama.
At some of the same stations where these
were taken, and at others, juveniles of two
other species of scombroid fishes which have
been identified as Euthynnus lineatus Kis-
hinouye, the black skipjack, and Auxis thaz¬
ard (Lacepede), the frigate mackerel, were
captured by the same means. Neither of these
is utilized by the American commercial fish¬
ery in the Pacific. The black skipjack of the
Asiatic side of the Pacific, E. yaito Kishinouye,
however, is of considerable commercial im¬
portance to the Japanese fishery and it may be
expected that E. lineatus will eventually be
similarly exploited. Commercial catches of
frigate mackerel in the middle Atlantic states
averaged slightly over 100 tons in 1942-44
(Fiedler, 1945; Anderson and Power, 1946,
1947).

Euthynnus lineatus Kishinouye 1920

Adults of this species are occasionally cap¬
tured in Central American waters incidentally
to the tuna fishery. Four specimens were
examined by us in the early spring of 1947.
Two of these were captured in a bait-net in
the Gulf of Nicoya, Costa Rica, on February
22; these fish, one male and one female, had

1 South Pacific Investigations, U. S. Fish and Wild¬
life Service. Published by permission of the Director
of the U. S. Fish and Wildlife Service. Manuscript
received March 25, 1948.

gonads in a very advanced stage of maturity.
A female with running-ripe eggs was taken on
a trolled feather jig off Quepos Point, Costa
Rica, on April 4. A ripe male was taken in a
purse seine haul off Cape Blanco, Costa Rica,
on April 29. It is thus apparent that this species
spawns in Central American waters during the
early spring. The capture of juveniles further
confirms this.

Juveniles were taken at the following sta¬
tions on the dates indicated: 08° 2 O' N., 84°
10' W., March 18, 1947; 8 specimens, 48 to
86 mm. total length. 09° 20' N., 85° 20' W.,
March 19, 1947; 10 specimens, 29 to 56 mm.
total length. 09° 10' N, 85° 20' W., March
20, 1947; 1 specimen, 61 mm. total length.
(All total lengths in this paper are from tip
of snout to tip of shortest median caudal ray.)

In Figures 1 and 2 representative specimens
of these juveniles are depicted. They are rela¬
tively less deep bodied than the juveniles of
Neothunnus of the same sizes, being similar in
this regard to Katsuwonus, from which, how¬
ever, they may easily be distinguished by the
pigmentation of the first dorsal fin. The entire
first dorsal is heavily pigmented in Euthynnus
of all sizes collected, while in Katsuwonus , up
to 44 mm. at least, there is only a light pig¬
mentation of the anterior margin and of the
distal edge of the fin.

The second dorsal fin remains completely
unpigmented in fish up to about 45 mm. total
length, at which size the fin begins to show
some pigment at the bases of the rays. In our
largest specimen, 86 mm. total length, the
second dorsal is fairly dark about half way to
the tips of the rays, the distal half remaining
unpigmented. The pigmentation of the head
and body is similar to that of Katsuwonus at

[ 262 ]

Juvenile Tunas — SCHAEFER and Marr

263

Fig. 1. Photograph of juvenile Euthynnus lineatus Kishinouye.

the smaller sizes, but the snout and anterior part
of the head is rather more extensively pig¬
mented in the smaller Euthynnus of these col¬
lections than in either Katsuwonus or Neo-
thunnus of similar sizes. On the smaller of our
specimens of Euthynnus the lateral pigmentation
does not extend far below the mid-line, but
there is a conspicuous series of black spots
along the bases of the anal fin and following
finlets. As the fish increase in size the pigmen¬
tation extends further down the sides, par¬
ticularly posteriorly, until at about 45 mm. the
pigmentation at the level of the anal fin ex¬
tends the entire depth of the fish, although
shading off ventrally. By the time the fish has
reached 55 mm. the entire head is rather dark,
the body is almost black dorsally, shading off

gradually to a white belly. In some specimens
of about 45 mm. to 60 mm. the pigment on the
upper sides is concentrated to form 8 or 9
extremely faint vertical bars. In the 86 mm.
specimen these are not visible. The peritoneum
bears numerous large dark spots dorsally which
are visible through the body wall in small
specimens. The caudal fin begins to exhibit
some pigmentation in fishes of about 50 mm.
total length.

In addition to the examination of specimens
preserved in alcohol, two specimens of 54 mm.
and 56 mm. were prepared for study of bony
parts by staining with alizarin and clearing,
after the method of Hollister (1934). The
pectoral fin in the two stained specimens has
29 rays and 27 rays, respectively. These rays

264

PACIFIC SCIENCE, Vol. II, October, 1948

Fig. 2. Juvenile Euthynnus lineatus; photographed against a white background to illustrate pigmentation of
dorsal fin.

cannot be counted accurately in unstained in¬
dividuals because of the small size of the most
inferior rays. The dorsal fin has 15 spines in
six specimens examined and 14 in a seventh,
the first or second anterior spines, which are
of nearly equal length, being the longest and
the remaining spines decreasing in length rapid¬
ly and successively. The first dorsal reaches
almost to the insertion of the second. The sec¬
ond dorsal fin rays, counted in 5 specimens, are
12 in number, and there are in each case 8 dorsal
finlets. The anal fin has 11 or 12 rays, and there
are 7 anal finlets. The rays of the second dorsal

and anal are difficult to count except in stained
material because of the shortness of the first one
or two rays. The finlets are connected to each
other and to the fin by a thin membrane which
extends less and less far toward the tips of the
finlets as the fish grow until in the largest speci¬
men of 86 mm. it is only a vestige between the
bases of the finlets.

The gill rakers of our smallest specimens are
very tiny projections on the gill arches and
are difficult to count accurately. On such a
specimen, of 32 mm. total length, they were
counted on the first gill arch as 7 + 20. As the

Juvenile Tunas — SCHAEFER and MARR

265

fish increase in size the gill rakers not only in¬
crease rapidly in length, but their number ap¬
parently increases. Counts of rakers on the first
arch on specimens of various sizes are as fol¬
lows:

48 mm. 8 + 25

52 mm. 9 + 25

61 mm. 10 + 26

86 mm. 11 + 27

The vertebrae, counting the urostyle, are
19 + 18 in one stained specimen and 20 + 17
in the other. They bear large inferior foramina
on the last three or four precaudal vertebrae
and on the first nine or ten caudal vertebrae.
The haemal canal is very large, being broader
than the body of the vertebrae beneath the
precaudal and anterior caudal vertebrae. The
lateral processes of the posterior caudal ver¬
tebrae are well developed in both specimens;
there are no lateral processes on the anterior
precaudal vertebrae. The first complete haemal
arch was found to be on the 15 th vertebra by
dissection of the 56 mm. specimen.

Specimens of all sizes in our collection have
about 20 to 30 conical, inwardly curved teeth
on each side of each jaw. The palatines each
bear a row of eight to ten rather large conical
teeth. The vomer bears four or five rather
small teeth which may be easily overlooked.

Our very smallest specimens, of about 30
mm., have visible the remnants of three spines
at the lower posterior angle of the preopercle.
In larger specimens the growth of the bone
has completely obliterated these. Smaller speci¬
mens than those in our collection may be pre¬
sumed, by analogy with Neothunnus, to have
more prominent and perhaps more numerous
preopercular spines.

The intestine of these specimens, as is char¬
acteristic of the Katsuwonidae, is relatively
straight and is not folded. It runs back along
the right inferior portion of the stomach. The
liver is in three lobes, the right lobe being very
much longer than the other two. In seven
specimens examined, the right lobe of the liver

extended posteriorly three-fourths or more of
the length of the body cavity.

That these specimens belong to either the
genus Euthynnus or Katsmuonus is indicated
by the morphology of the vertebral column.
The well-developed inferior foramina on some
precaudal as well as caudal vertebrae, forming
with the very large haemal arches the so-called
"trellis,” is characteristic of these genera ( Starks,
1910; Kishinouye, 1923).

These specimens have several characters
agreeing with Euthynnus but not Katsmuonus .
The very great elongation of the right lobe of
the liver is definitive. In Katsuwonus, both in
adults described by Kishinouye (1923: 363,
450, and 453, Fig. N) and by Godsil and
Byers ( 1944: 11, 30), and in juveniles described
by Schaefer and Marr (in press), the right
lobe, although much larger than the other two,
is not nearly as large as that of these specimens,
which corresponds to Kishinouye’s description
of the liver of Euthynnus. The first closed
haemal arch of Katsuwonus occurs on the 12th
vertebra according to both Kishinouye and
Godsil and Byers, whereas Kishinouye states
that Euthynnus has the first complete haemal
arch further back, on the 16th vertebra. The
low vertebral count of our specimens is also
definitive, although this was a matter of some
concern at first because it does not agree with
the literature. Kishinouye (1923: 338, 452)
found that Euthynnus has 39 vertebrae count¬
ing the urostyle, probably based on E. yaito
alone. He described E. lineatus from a single
specimen from Manzanillo, Mexico, but prob¬
ably did not dissect the fish to count the verte¬
brae. (Kishinouye: in the Suisan Gakkwai Ho,
III, 113, 1920. We have been unable to
examine this reference and our information is
from Kishinouye, 1923.) He differentiates this
species from E. yaito on the coloration and the
relative size of the head. In his discussions he
assumes all species of Euthynnus to have 39
vertebrae. This matter was cleared up through
the kind cooperation of C. R. Clothier of the

2 66

California State Fisheries Laboratory, who has
examined the skeletons of four adults identified
from Kishinouyes description as E. line atm.
He has sent us the following data regarding
these specimens, the first three of which were
from tuna-clipper landings from unknown
points somewhere south of the Mexican border,
and the fourth of which was captured off Espi-
ritu Santu Island in the Gulf of California:

Specimen number.......... . 12 3 4

Vertebra bearing 1st haemal

arch.................................... — - 16 15 16

Total vertebrae ( including

urostyle ...... 37 39 37 37

Abdominal vertebrae..... — - 21 19 20

Caudal vertebrae....... ............. 18 18 17

Three of these four specimens have 37
vertebrae, corresponding with our juveniles,
and disagreeing with Kishinouye. Mr. Clothier
has also advised us that, in addition, H. C
Godsil of the same laboratory, has examined
two specimens - from near Magdalena Bay each
of which had 37 vertebrae. It appears that E.
lineatus has 37 vertebrae as a rule. The verte¬
brae counts of our juveniles agree, then, with
those of adult E. lineatus identified from other
characters. All other characters examined
agree well enough with Kishinouyes descrip¬
tions, and it was therefore concluded that these
were juveniles of E. lineatus.

There are few references to juvenile Euthyn-
nus in the literature. Kishinouyes (1923: 388)
smallest specimens, 13 cm. in length, from the
Asiatic side of the Pacific, were larger than any
of ours. He described them as follows: "They
are very slender and have eight or more trans¬
verse bands on the side. These bands are nearly
vertical and fade toward the ventral median line.
When they grow to a total length of 19 cm.
the body becomes very broad, the thoracic spots
appear, the bands gradually disappear from the
ventral part and the dorsal part of the bands
becomes oblique.”

Ehrenbaum (1924) found among the Medi¬
terranean collections of the Danish Oceano¬
graphical Expeditions six young tunas, 5.9 to

PACIFIC SCIENCE, Vol. II, October, 1948

10.1 mm. in length, which he referred to

Euthynnus alliteratus ( Raf . ) . He differentiated
these specimens from those of Auxis thazard
only with great difficulty, on the basis of the
first dorsal rays. The identification seems doubt¬
ful, but, since none of our specimens of Eu¬
thynnus are as small as Ehrenbaum’s, we can¬
not verify it.

Auxis thazard (Lacepede) 1802

As is the case with many of the scombroids,
there is some doubt as to whether there is a
single cosmopolitan species of Auxis or whether
different species occur in different parts of the
world. The solution of this problem is be¬
yond the scope of this paper and our juveniles
are referred to Auxis thazard. No adult speci¬
mens were examined by us during this voyage,
but the capture of juveniles indicates that this
species spawns in Central American Pacific
waters during the early spring. Juveniles were
taken at the following stations on the dates
indicated: 08° 20' N., 84° 10' W.; March 18,
1947; 2 specimens, 67 and 68 mm. total length.
09° 43' N, 85° 54' W.; March 19, 1947; 52
specimens, 21 to 53 mm. total length. 08° 7'
30" N., 83° 8' 30" W.; May 7, 1947; 3 speci¬
mens, 23 to 41 mm. total length. 09° 43' N.,
85° 54' W.; May 17, 1947; 1 specimen, 42 mm.
total length. Through the courtesy of Dr. J. T.
Nichols, we were also able to examine 5 speci¬
mens, 22 to 31 mm. in total length, taken
under a light at night from the "Askoy” at
04° 01' N., 80° 26' W. on March 24, 1941.
Nichols and Murphy (1944: 241) suggest that
these may be young Euthynnus , but they are
undoubtedly Auxis .

Representative specimens of juvenile A.
thazard are shown in Figures 3 and 4. In the
smaller specimens the prominent areas of pig¬
mentation are on the upper and lower jaws,
above the snout, around the postero-ventral
margin of the orbit, on the upper operculum,
between the orbits, along the mid-line of the
body, along the bases of the dorsal and anal fins

Juvenile Tunas — SCHAEFER and MARR

267

FIG. 3. Photograph of juvenile Auxis thazard (Lacepede)

including the finlets, and around the posterior
end of the urostyle. Large chromatophores in
the peritoneum show through the body wall
along the upper half of the body cavity. None
of the fins or finlets bears pigment spots, with
the exception of the first dorsal. The first dor¬
sal bears a few scattered chromatophores, large¬
ly distributed along the spines. However, the
first dorsal in general appearance is virtually
colorless, as may be seen in Figure 4, especially
as contrasted with Euthynnus or Neothunnus.
With increasing size the local areas of pigmen¬
tation expand. In our largest specimens the
head is well pigmented. The dorsal half of the

body is uniformly dark. This dark area extends
below the mid-line on the caudal peduncle.
The chromatophores in the peritoneum are no
longer visible through the thickened body wall,
and the ventral surface is unpigmented. Even
in our largest specimens none of the fins is
heavily pigmented; the caudal is more pig¬
mented than the others.

Juvenile Auxis are rounder in cross section
than Euthynnus , being relatively less com¬
pressed laterally. The head length and length
of the caudal region in Auxis are less in com¬
parison to the total length than is the case in
Euthynnus. The maxillary in Auxis generally

268

PACIFIC SCIENCE, Vol. II, October, 1948

FIG. 4. Juvenile Auxis t hazard photographed against a white background, illustrating lack of pigmentation of
fins.

extends to a point between the anterior margin
of the iris and the vertical bisector of the iris,
whereas in Euthynnus it generally extends to a
point between the vertical bisector of the iris
and the posterior margin of the iris. In the
smaller specimens the lower jaw is shorter than
the upper, but the jaws become equal at about
50 mm. total length. The smallest specimens
have three spines at the angle of the preopercle,
but these are gradually grown over and dis¬
appear at about 35 to 40 mm. total length.
There are indications, on alizarin-stained speci¬

mens, that seven more spines are present in
smaller individuals. Similarly, a figure of a 17.5
mm. specimen of A. t hazard from the Medi¬
terranean (deBuen, 1932: Fig. 27) shows a
total of nine preopercular spines. The lateral
keel along the caudal peduncle starts to develop
at about 40 mm. total length.

The large dorsal interspace characteristic of
Auxis actually contains small spines or rays.
These are usually subcutaneous. Even in our
smallest specimens they are not visible unless
the fish is cleared and stained or unless the

Juvenile Tunas — SCHAEFER and Marr

269

specimen is shrunken from the preservative so
that the spines project above the dorsal profile.
In adult Auxis from Culion, Philippine Islands,
examined through courtesy of Dr. G. S. Myers
of the Stanford University Natural History
Museum, the interneural supports of the sup¬
pressed rays are well developed; the rays them¬
selves are vestigial, however, and can be found
only by staining and dissection. Fin ray counts
on our stained juvenile specimens are as fol¬
lows:

Total length of specimen......

mm.

25

mm.

31

mm.

46

First dorsal... . . .

11

11

11

Dorsal interspace.... . .

6

7

8

Second dorsal.... . . .

12

11

11

Dorsal finlets.... . .

8

8

8

Anal......... . . . .

14

13

13

Anal finlets .

7

7

7

Ventral . . . . .

7

7

7

Pectoral. . . .

22

23

25

Other, unstained, specimens

also

have eight

dorsal finlets and seven anal finlets. In the
smaller specimens the finlets are joined by a
membrane that extends completely or nearly
to their distal ends. With increasing size this
membrane becomes less prominent until in our
longest specimens it is wanting or exists only
between the bases of the finlets.

In all our specimens, each side of the upper
jaw bears about 20 small, conical teeth; each
side of the lower jaw about 25 small, conical
teeth; each palatine 6 or 7 teeth; and the
vomer none. The vomer and palatines are ex¬
posed. Starks (1910: 97), with reference to
Auxis thazard, and Kishinouye (1923: 460),
with reference to the genus Auxis , state that
there are no palatine teeth, and this is apparent¬
ly true in the adults. In the two adults from
Culion, Philippine Islands, the vomer and pala¬
tines are toothless and are not exposed. These
specimens are probably Auxis hvra Kishinouye.

The gill rakers on the most anterior gill arch
of our smallest specimens are very tiny pro¬
jections and are difficult to count. They are first
apparent near the angle of the arch. With an
increase in the size of the fish, the rakers near

the angle of the arch increase in length and
new rakers are added distally on each arm.
The full complement of rakers is apparently
attained at about 50 mm. total length, as
judged by the following counts on specimens
of various sizes:

21

mm.

4

+

18

24

mm.

5

+

21

27

mm.

6

+

24

32

mm.

9

+

26

42

mm.

9

+

30

46

mm.

10

+

31

52

mm.

12

+

36

54

mm.

11

+

35

68

mm.

11

+

31

Gill raker counts on the adult specimens re¬
ferred to above are 12 + 34 and 11 + 32.
Kishinouye (1923: 462-3) gives 9 + 30 as the
gill raker count for A. hira and 10 + 36 as the
count for A. mam Kishinouye. Possible varia¬
tion in these counts is not mentioned. He does
not list A. thazard, but A. mam is probably
a synonym of A. thazard, as he tentatively sug¬
gests in his synonymy.

The nasal rosette is visible only in cleared
specimens.

The vertebrae were counted as 20 + 19 (in¬
cluding the urostyle) in three cleared and
stained specimens. This agrees with Kishinouye
(1923: 452) and Frade and deBuen (1932:
70), but disagrees with Starks (1910: 97),
who counted 22 + 15 vertebrae. On a speci¬
men 46 mm. total length, small ventral fora¬
mina are present on the 9th to 15 th caudal
vertebrae. On smaller specimens of 25 and 31
mm. total length, these foramina are not dis¬
cernible, if present. In our specimens, the
pedicles (of Starks = epihaemal process of
Kishinouye) of the caudal vertebrae, bearing
the closed haemal arches, are extremely short
or non-existent. In a specimen of 25 mm. total
length, the haemal arches of the caudal ver¬
tebrae are long, wide, and elliptical. In a speci¬
men of 46 mm. total length, the haemal arch

270

PACIFIC SCIENCE, Vol. II, October, 1948

is still long, but less wide. In a specimen of
72 mm. total length, the haemal arch is still
long, but even less wide. In fact, the pedicle
is divided and contains a large part of the
haemal arch. In the two smaller specimens,
the haemal arch is certainly closed on the 18th
abdominal vertebra and is probably closed on
the 17th vertebra; it is definitely open on the
16th vertebra. In the 72 mm. specimen, the
haemal arch is open on the 17th vertebra and
closed on the 18th. Kishinouye (1923: 339)
states that the first closed haemal arch of
Auxis occurs on the 1st caudal vertebrae
(= 21st). In an adult specimen examined
through the courtesy of Dr. G. S. Myers, the
first closed haemal arch is on the first caudal
vertebra. This specimen is from Wakanoura,
Japan, and is referred to Auxis hira. We have
been unable to examine any adult Auxis from
the eastern Pacific in order to attempt to re¬
solve this apparent discrepancy. C. R. Clothier
has informed us that one specimen from the
vicinity of Sebastian Viscaino Bay and two
from the vicinity of Espiritu Santu Island, all
identified as A. t hazard , have a total of 39 verte¬
brae with the first haemal arch on the 21st
vertebra. It seems unlikely that haemal arches
that are closed would, with growth, become
open. Our specimens may, therefore, be juve¬
niles of an undescribed species. It is not
unlikely that the sides of the pedicle fuse with
an increase in size of the fish so that the haemal
arch becomes greatly reduced in size. The possi¬
bility exists, however, that it may remain
divided in this form.

The liver is divided into three lobes. The
right lobe is almost as long as the visceral
cavity and bears a prominent hepatic vein on
its outer surface; the other two lobes are small.
The stomach is long and lies above the rest of
the viscera. The caecal mass is considerably
shorter than the stomach. The intestine is rela¬
tively short, straight, and not folded. It runs
posteriorly along the right inferior side of the
stomach. There is no air bladder.

The vertebral count, the great length of the
right lobe of the liver, the presence of a caudal
keel, and the absence of an air bladder indicate
that these juvenile scombroids belong to the
family Katsuwonidae. The large interspace
between the dorsal fins and the low ray count
of the first dorsal, as well as the absence of the
elaborate "trellis” of the vertebrae, indicate
that they are Auxis , rather than Euthynnus or
Katsuwonus. In the larger specimens, gill raker
and fin ray counts will also serve to separate
Auxis from the other genera. Positive specific
allocation must await examination of adult
specimens from the area.

A specimen of A. thazard, 17.5 mm. long,
from the Mediterranean, was described and fig¬
ured by deBuen (1932: 36-38). This is some¬
what smaller than our smallest specimen and the
differences between them, such as differences
in numbers of preopercular spines, etc., are
probably due to the differences in size. Sparta
(1933: 16) mentions that specimens of Auxis
bisus Bonaparte ( = A. thazard Lacepede ) about
1 cm. in length have been collected under a
light at night in the Straits of Messina. Ehren-
baum (1924: 33-38) reported on 132 juvenile
A. thazard from the Mediterranean collections
of the Danish Oceanographical Expeditions.
His largest specimen was 12 mm. in length, or
about 9 mm. shorter than our smallest specimen.
Again, differences between his descriptions and
our specimens may probably be attributed to
differences in size. These specimens were ap¬
parently taken in May, July, August, and Sep¬
tember. He states that Sanzo found ripe ovar¬
ies in June and July. Sella (1924) describes
Mediterranean specimens of A. bisus Bonaparte
(= A. thazard Lacepede) ranging from 3 to 10
mm. These are much smaller than our speci¬
mens. He states that juveniles up to 10 to 15
mm. in length are found from the second half
of June to September 20. He also mentions
that before a length of 12 to 15 mm. is at¬
tained, about six rays are formed in the mem¬
brane connecting the first and second dorsal

Juvenile Tunas — SCHAEFER and Marr

271

fins and that these rays subsequently become
contained in the median furrow so that they
are no longer visible. This is in accord with
our findings. The dates of our collections indi¬
cate the possibility of a somewhat earlier spawn¬
ing season for Auxis off the Pacific Coast of
Central America.

We have been unable to examine papers by
Sanzo (1909, 1910) which apparently contain
information on larval and juvenile Auxis.

REFERENCES

Anderson, A. W., and E. A. Power. 1946.
Fishery statistics of the United States : 1942.
U. S. Fish and Wildlife Serv., St at. Dig. 11:
1-248.

- 1947. Fishery statistics of the United

States : 194 A U. S. Fish and Wildlife Serv.,
Stat. Dig. 14: 1-241.

deBuen, Fernando. 1932. Formas ontogeni-
cas de peces (nota primera). Inst. Espanol
de Oceanogr., Notas y Resumenes. II, 57:
1-38.

Ehrenbaum, E. 1924. Scombri formes. Kept.
Danish Oceanogr. Exp. 1908-1910. No. 8.
2 [Biology] (A.ll) : 1-42.

Fiedler, R. H. 1945. Fishery statistics of the
United States : 1941. U. S. Fish and Wildlife
Serv., Stat. Dig. 7: 1-174.

Frade, Fernando, and Fernando deBuen.
1932. Poissons scombri formes ( excepte la
famille Scombridae). Clef de classifications
princalement d’apres la morphologic interne.
Comm. Int. pour I’Expl. Sci. de la Mer Medi-
terranee, Rapp, et Proc. Verb., 1 (annexe A) :
69-70.

Godsil, H. G, and R. D. Byers. 1944. A sys¬
tematic study of the Pacific tunas. Calif. Div.
Fish and Game, Fish Bui. 60, 131 p., 76 fig.

Hollister, Gloria. 1934. Cleaning and dye¬
ing fish for bone study. Zoologica 12 (10):
89-101.

Kishinouye, Kamakichi. 1923. Contribu¬
tions to the comparative study of the so-called
scombroid fishes. Tokyo Imp. Univ., Col. Agr.
Jour. 8 (3): 293-475, 22 pi.

Nichols, J. T., and R. C. Murphy. 1944. A
collection of fishes from the Panama Bight,
Pacific Ocean. Amer. Mus. Nat. Hist., Bui.
83: (art. 4): 217-260, 4 pi.

SANZO, L. 1909. Uova e larve di Auxis bisus.
Monitore Zool. ltd. 20: 79-80.

— - 1910. Uova e larve di Scomberoidi.

Rev. mensile di pesca 1910: 201-205.

Schaefer, M. B., and J. C. Marr. In press.
Spawning of the yellowfin tuna (Neothun-
nus macropterus ) and skipjack (Katsuwonus
pelamis) in the Pacific Ocean off Central
America, with descriptions of juveniles. U. S.
Fish and Wildlife Serv., Fish Bui. 51: in press.

Sella, M. 1924. Caratteri differenziali dei
giovana stadi di Orcynus thynnus Ltkn., O.
alalonga Risso, Auxis bisus Bp. R. Accad.
Naz. dei Lincei, Cl. di Sci. Fis., Mat. e Nat.,
Rend. V, 33: 300-305.

Sparta, A. 1933. Osservazioni compiute nello
Stretto di Messina sul comportemento dei
pesci e cefalopodi all’azione di sorgenti lumi-
nose. R. Comitat. Tallasogr. ltd., Mem. 206:
1-22.

Starks, E. C. 1910. The osteology and mutual
relationships of the fishes belonging to the
family Scombridae. Jour. Morph. 21 (1):
77-99, 3 pi.

Plant Records from the Caroline Islands, Micronesia:
Pacific Plant Studies 81

Harold St. John2

After the close of active hostilities in the
Pacific in 1945, the United States Navy made it
possible for the University of Hawaii to send
a scientific reconnaissance party into the Caro¬
line Islands, even though thousands of Japanese
military prisoners and a few detachments of
hostile "hold-out” bands were still there. Plant
collections were made on Kusaie Island from
December 24 to 26, 1945; and in the Palau
Group, on Angaur Island, on January 2, 1946.
Below are listed the species taken which have
not previously been recorded from these islands
or for which the vernacular names used by the
natives have not been recorded. The specimens
are deposited in the Bishop Museum, Honolulu.

Plants from Kusaie

AN GIOPTERID ACE AE

Angiopteris Beecheyana de Vriese

A. eve eta sensu Hook, and Grev. non Hoffm.,
fide Hosokawa, Nat. Hist. Soc. Formosa,
Trans. 26: 44, 1936.

Kusaie Island: divide S. of Lele Harbor,
St. John 21,440, "taime.”

OPHIOGLOSSACEAE

Ophioglossum pendulum L.

Kusaie Island: divide S. of Lele Harbor,
St. John 21,443, "karkarweh.”

1 This is the eighth in a series of papers designed to
present descriptions, revisions, and records of Pacific
island plants. The preceding papers were published as
Bernice P. Bishop Mus., Occas. Papers 17 (7), 1942;
17 (13), 1943; 18 (5), 1945; Amer. Fern Jour. 35:
87-89, 1945; Torrey Bot. Club, Bui. 73: 588, 1946;
Pacific Sci. 1 : 116, 1947; Pacific Sci. 2: 96-113, 1948.

2 Chairman, Department of Botany, University of
Hawaii. Manuscript received September 1, 1947.

CYATHEACEAE

Cyathea nigricans Mett.

C. affinis sensu Kanehira, non Schrader.

Kusaie Island: divide S. of Lele Harbor,,
trunks 8 m. by 2 dm., St. John 21,441, "po.”

SELAGINELLACEAE

Selaginella Kanehirae Alston

Kusaie Island: Yekala Waterfall, St. John
21,437.

PANDANACEAE

Freycinetia ponapensis Mart.

Kusaie Island: Innemu River, St. John
21,449, "fuka.”

HYDROCHARITACEAE

Thalassia Hemprichii (Ehrenb.) Aschers.

Kusaie Island: Lele Harbor, in salt water
1-2 m. deep, and on outer reef, H. St. John
and H. 1. Fisher 21,433. First record for
Kusaie, though abundant in the shallow
water of the harbor.

LILIACEAE

Cordyline terminalis (L.) Kunth

Taetsia fruticosa (L.) Merr.

Kusaie Island: divide S. of Lele Harbor, in
moist native forest, St. John 21,442, "ing-
in-kal.” First record for Kusaie.

DIOSCOREACEAE

Dioscorea korrorensis R. Knuth

Kusaie Island: Lele Island, cultivated, St.
John 21,433, "ohkani.” First record from
Kusaie.

[272 3

Caroline Island Plants — St. John

273

RHAMNACEAE

Colubrina asiatica (L.) Brogn.

Kusaie Island: divide S. of Lele Harbor,

St. John 21,439 , "la.”

VERBENACEAE

Premna integrifolia L.

Kusaie Island: Innemu River, St. John 21,452,
"fienket.”

Plants from Palau

CAPPARIDACEAE

Capparis cordifolia Lam.

Angaur Island: coral sea cliff, St. John 21,500,
"i-il-lel-lameng-ngernger.’^

VERBENACEAE
Callicarpa cana L.

Angaur Island: coral limestone sea cliff, St.
John 21,502, "dup.” First record from
Angaur Island.

Clerodendrum inerme (L.) Gaertn.

Angaur Island: coral limestone sea cliff, St.
John 21,501, "kellel-lap-ni.” First record
from Angaur Island.

RUBIACEAE

Hedyotis albido-punctata (Merr.) Fosberg
Oldenlandia albido-punctata Merr.

Angaur Island: coral limestone sea cliff, St.
John 21,503, "ngesil.” First record from
Angaur Island.

A New Echiuroid Worm from the Hawaiian Islands and a Key to the

Genera of Echiuridae1

Walter K. Fisher2

Since echiuroid worms and their eggs pro¬
vide excellent material for biological experi¬
mentation, it is regrettable that the principal
Hawaiian species is without a name. This paper
is intended to supply the deficiency.

I collected the type series in 1902, during the
Hawaiian cruise of the "Albatross,” in tide
pools on the reef between Honolulu harbor and
Waikiki. The material was sent to the late
Professor H. B. Ward and only recently became
available for study. A long sojourn in weak
alcohol has caused deterioration of the speci¬
mens.

Genus Anelassorhynchus Annandale

Anelassorhynchus Annandale, 1922: 148. Type,

Thalassema branchiorhynchus Annandale and

Kemp.

Diagnosis : Resembling Thalassema s.s. in
having the longitudinal muscle layer of the body
of uniform thickness, without specialized longi¬
tudinal bands, but differing in having prolonged,
often coiled, lips to the ciliated funnel of the
nephridium.

The group contains about twelve species
from warm, shallow waters with the exception
of an undescribed species from off Monterey
Bay, California, 1083 fathoms, bottom temper¬
ature 38.5° F.

Anelassorhynchus porcellus new species

Diagnosis: Nephridia four, all behind setae;
a very long segment of intestine between end

1 Published with permission of the Secretary of the
Smithsonian Institution. Manuscript received April
13, 1948.

2 Professor Emeritus, Stanford University, Hopkins
Marine Station, Pacific Grove, California.

of foregut and beginning of siphon; anal vesicles
very long with a special basal inflated portion
attached to body wall by muscular frenula; no
caecum; setae small, without interbasal muscle;
proboscis fleshy, deciduous, the two margins
meeting at base to form lower lip; skin rather
thick in adult, with transverse verrucae, in con¬
tracted state; size 30 to 70 mm. in length, the
diameter variable.

Description: Body wall rather thick in large
specimens, slightly translucent in smaller ones;
skin usually closely wrinkled transversely, so
that the small, closely placed glandular swell¬
ings have a transverse alignment. In middle of
body, owing to stretching of skin, wrinkles
may disappear and glands flatten out so as to
be practically indistinguishable. (It is pretty
much a matter of accident in preservation.)

Setae small (3 mm. long), with well-marked
hook, placed rather close together, only a short
distance from mouth. No interbasal muscle
uniting inner ends of setae; basilaterals rather
weak.

Inner layer of muscles of body wall smooth;
middle, longitudinal layer undifferentiated as in
other species of the genus.

All four nephridia behind setae, variable in
size, sometimes greatly distended by sex prod¬
ucts. Lips of nephrostome prolonged and usual¬
ly spirally coiled.

Deflated anal vesicles very long and character¬
istically swollen at base, which is attached to
anterior side wall of cloacal cavity. Walls of
this cavity thin and attached to body wall by
many frenula which also involve the swollen
basal portion of the vesicles (Fig. 1, c). In
some specimens vesicles covered with tiny

[ 274 ]

Echiuroid Worm — FISHER

275

brown spots, perhaps the nephric elements,
although ciliated funnels cannot be recognized.

Alimentary canal very long and filled with
coarse "sand.” No caecum present. Foregut
without obviously differentiated subdivisions
recognizable from outward appearance. Seg¬
ment adjacent to ring blood-vessel (B2) some¬
times set off by a slight constriction. Very long
segment of the intestine, between B2 and be¬
ginning of siphon, apparently without ciliated
groove. (This presiphonal segment is generally
short.) In one specimen 50 mm. long, with
empty and relaxed intestine, presiphonal seg¬
ment can be traced for 40 mm. before a break
occurs. It is almost certainly longer. Foregut
25 mm. in length. In another specimen a seg¬
ment about twice as long as foregut is without
siphon, although siphon is recognizable over
an extensive portion of the midgut. Entire
intestine about ten times contracted body length;
in specimens examined in a most wonderful
snarl, complicated in all but one by sausage-like
swellings filled with sand and shell fragments.

Vascular system (Fig. 1, a) of usual Thalas¬
sema type, with well-developed ring vessel and
some slight variations in the details of the con¬
nection with the neurointestinal vessel (B3).
(It may be remarked that the length of the
neurointestinal varies from half the length
shown in Figure 1, a to five times that length,
depending entirely upon position of the end of
the forgut, which has great freedom of move¬
ment.) 'In one specimen, killed while passage
from stomach to intestine was distended by
sand, ring vessel ( B2 ) three times usual diameter
in preserved specimens.

Type: In the collection of the United States
National Museum.

Type locality: Honolulu; reef south of harbor,
in tide pools, June 6, 1902. Collected by W. K.
Fisher.

Specimens examined: Honolulu reef, 15;
Puako Bay, Hawaii, 2. I have also examined

several collected by R. W. Hiatt at Halape,
Hawaii, where they were found in sand under
rocks.

Discussion: This species is probably rather
closely related to Thalassema semoni Fischer
(1896: 338, Fig. 4; Amboina) but the descrip¬
tion lacks all details of the alimentary canal
and figures of importance: Proboscis lost; larger
of the two specimens 55 mm.; skin bluish-gray,
rather thin and translucent; musculature not
split into bundles; papillae cover skin almost
without intervals, though more concentrated
at posterior end; setae small; nephridia four,
with spiral tubes; anal vesicles thin, brown,
longer than half the body; they are attached
to body wall by muscles; caecum not observed.
Fischer states (1914: 19) that Thalassema
sabinmn Lanchester (1905: 40, Tale Sab, Sin-
gora) is a synonym of semoni. However, Pra-
shad (1919: PI. 11, Fig. 10) figures a dissec¬
tion of sabinum, which shows a well-developed
caecum and short anal vesicles. The proboscis
is adherent, being an important respiratory or¬
gan owing to the ecology of the species. This
figure also indicates that the presiphonal seg¬
ment of intestine is short. A. sabinus is ob¬
viously not close to porcellus and is probably
not the same as semoni.

No true Thalassema has been recorded from
the Hawaiian Islands. Other echiuroids which
I have examined from the islands are: Oche-
to stoma erythrogrammon Leuckart and Riippell,
Nawiliwili, Kauai (A. E. Verrill), and Halape,
Hawaii (R. W. Hiatt); Ochetostoma manjuo-
dense Ikeda, Halape, Hawaii (R. W. Hiatt);
Anelassorhynchus inanensis (Ikeda), Halape,
Hawaii (R. W. Hiatt). There are undoubtedly
other species present in these waters.

As the genera allied to Thalassema are rather
difficult to distinguish, the following synopsis
may prove useful.

Keys to other genera of Echiuroidea, prin¬
cipally Bonelliidae, and figures of anatomy will
be found in Fisher, 1946.

27 6

PACIFIC SCIENCE, Vol. II, October, 1948

Fig. 1. Anelassorhynchus porcellus: a, map of anatomy of anterior portion of body, X5, (mesenteries are
•omitted, the foregut (P,0,C} is capable of being straightened parallel to nerve cord so that the neurointes-
tinal vessel [B3] is then greatly lengthened, the nephridia [n} are usually greatly swollen by contained eggs
or sperm ) ; b, diagram of ring vessel surrounding end of foregut to show connection with B1 and B3; c,
cloaca with anal vesicles, X5, from a smaller specimen than a\ d, seta, X10, and its hook, enlarged; e, sketch
of anterior two-thirds of a typical specimen with detached proboscis, XL

AV, anal vesicles; B1— b\ dorsal, ring, neurointestinal, and ventral blood vessels, respectively; C, stomach; CF,
ciliated funnel or nephrostome; C, cloaca; I, intestine; N, nephridium; NC, nerve cord; O, oesophagus; P,
pharynx; S, seta.

Echiuroid Worm— -Fisher

277

KEY TO GENERA OF ECHIURIDAE

a1. Two circles of posterior setae.....................—. . . .. . ...JEchiurus Guerin-Meneville

a2. No posterior setae present

b1. No differentiated thicker bands in longitudinal muscle layer
c1. Nephrostome of nephridia without elongated spirally coiled lips

d1. The neurointestinal blood vessel in direct connection with dorsal vessel
directly by a ring vessel or indirectly by a ring vessel at end of foregut;
segment of intestine between ring vessel and beginning of siphon short and
with ciliated groove; nephrostome with inconspicuous lips; proboscis not

especially expanded at tip, often deciduous . T halassema Lamarck

d2. Neurointestinal vessel connected with dorsal vessel by numerous capillaries
in wall of gut; segment of intestine between stomach and beginning of siphon
very long (2 or 3 times body length) and with or without ciliated groove;
nephrostome with conspicuous flap-like lip; proboscis very deciduous, long,

slender, expanded at tip*. . . . . . Arhynchite Sato

c2. Nephrostome with elongated, spirally coiled lips... .Anelassorhynchus Annandale
b2. Longitudinal muscle layer with slight to pronounced differentiation into longi¬
tudinal muscle bands, 6 or more in number
c1. Nephrostome of nephridia with small circular lips; inner layer of muscles not

differentiated into separate transverse fascicles between longitudinal bands .

. . . . . . .Lissomyema Fisher

c2 Nephrostome with elongated spiral lips

d1. Differentiated muscle bands weak, the zones between not showing a fascicu¬
late arrangement of inner oblique muscles; in small specimens longitudinal

bands very faint or visible only in posterior region . Listriolobus W. Fischer

d2. Longitudinal muscle bands strongly developed; zones between crossed by
separated fascicles of innermost oblique layer

e1. Nephridia in 1 to 5 pairs; vascular ring vessel at beginning of midgut . .

. . . . . . . Ochetostoma Leuckart and Ruppell

e2. Nephridia, at least in male, in 6 to 14 groups of 1 to 4, the groups arranged

in pairs; vascular ring vessel at posterior end of pharynx . . . . . .

. . . . . . Ikedosoma Bock

* I have specimens of a new form from off California in which the proboscis is present. In the two
described species there seemed to be no trace of a proboscis scar.

REFERENCES

Annandale, Nelson. 1922. The marine ele¬
ment in the fauna of the Ganges. Bijdr.
Dierk pt. 22, Feest-nummer 70sten Ge-
boortedag van Dr. Max Weber, p. 143-154.

Fischer, Wilhelm. 1896. Gephyreen. Zoo-
logische Forschungsreisen in Australien und
dem Malayischen Archipel. Bd. 5. In: Den-
kschr. Med.—Naturwiss . Gesellsch. Jena .
8: 337-339, figs. 1-4.

— - - - 1914. Weitere Mitteilungen iiber die

Gephyreen des naturhistorischen (zoologi-
schen) Museums zu Hamburg. Hamburg
naturh. Mus., Mitt . 31 Jahrg., Beih. 2: 1-28,

1 pi.

Fisher, Walter K. 1946. Echiuroid worms of
the north Pacific Ocean. U . S. Natl. Mus.,
Proc. 96: 215-292, pi. 20-37.

Lanchester, W. F. 1905. On the sipunculids
and echiurids collected during the Skeat Ex¬
pedition to the Malay Peninsula. Zool. Soc.
London, Proc. 1905: 35-41, 1 pi.

Prashad, B. 1919. Zoological results of a tour
in the Far East. Echiuroids from brackish
water with the description of a new species
from the Andamans. Asiatic Soc. Bengal,
Mem. 6: 323-338, 1 pi.

A Chronology of the Explosive Eruptions of Kilauea

Howard A. Powers1

INTRODUCTION

The explosive eruptions of Kilauea, both
prehistoric and historic, have been much dis¬
cussed in the literature on Hawaiian volcanoes.
Some early general observations attributed all
of the surface ash deposits to the known explo¬
sive eruption of 1790 (Dana, 1891: 42-45;
Jaggar, 192 1 : 1 14-118) ; others recognized evi¬
dence of several different prehistoric explosions
(Hitchcock, 1911: 166-169; Sidney Powers,
1916). Later studies, with better exposures in
artificial cuts on the windward rim of the crater,
demonstrated a number of long intervals of
quiet between several eruptions (Finch, 1925;
Stone, 1926). This paper adds another, still
more detailed, chapter to the accumulated
knowledge, but much remains to be learned and
recorded before complete understanding of the
explosive phases of the volcano is attained.
Many of the thoughts expressed herein are the
result of discussion and exchange of ideas in
the field with many coworkers, whose contri¬
butions are gratefully accepted and acknowl¬
edged. T. A. Jaggar, R. H. Finch, E. G. Win¬
gate, A. E. Jones, all formerly with the Hawaiian
Volcano Observatory; J. E. Doerr, Jr., former
naturalist, Hawaii National Park; H. S. Palmer
of the University of Hawaii; C. K. Wentworth
of the Honolulu Board of Water Supply; and
G. A. Macdonald of the United States Geologi¬
cal Survey have all contributed to the accumu¬
lation and consideration of material which is
presented in this paper.

The erratic original deposition of the pyro-
clastics of Kilauea, controlled by combinations
of atmospheric elements and directed explo¬

1 Naturalist, Hawaii National Park; since July 1,
Seismologist, Hawaiian Volcano Observatory, U. S.
Geological Survey. Published by permission of the
Director, National Park Service, Department of the
Interior. Manuscript received December 23, 1947.

sions, has been amply discussed by Wentworth
(1938: chapter 3). Very few individual beds
of material could be traced with assurance as
horizon markers, even for the short distance
around the circumference of the crater, because
of this erratic deposition. Also, removal of
material by wind and water erosion during and
between eruptions has been tremendous. As a
net result of these two factors, there is no cross
section in the area which contains a complete
representation of all the explosion deposits.

Successful analysis of the stratigraphy and
chronology of these deposits thus depends as
much on appraising an erosion surface in the
section as upon correctly describing the beds
of pyroclastic materials. In this study, great
emphasis has been placed on recognition of
characteristics which can be used to distinguish
unconformities and erosion surfaces produced
during time intervals of perhaps a few hours
between explosions from those produced dur¬
ing time intervals measured in years or tens of
years between eruptions. The following charac¬
teristics of the present desert surface have been
used to identify similar long-exposed surfaces
in cross sections of the deposits: surface oxida¬
tion, encrustation, desert varnish on fragments,
concentration of coarser particles, thinly lam¬
inated dust deposits, and truncation of pre¬
viously consolidated layers (Plate IB and 1C).
Humus and other remains of plant growth are
self-evident where they exist.

On each of eight segments of the crater rim,
a complete composite section of the pyroclastic
deposits was built up by compilation from a
number of cross sections which were sufficiently
close together so that correlations between them
could be made with confidence. Particular at¬
tention was paid to recording depositional

[ 278}

Eruptions of Kilauea — POWERS

279

TABLE 1. GENERALIZED SECTION OF PYROCLASTIC DEPOSITS, KILAUEA RIM.

indicates humus layer.

ERUPTION

TYPE

MAXIMUM

THICKNESS

1924

Phreatic

18 inches

1815 ca.

Magmatic

48 inches

1790 ca.

Phreatic

45 inches

18-K

Phreatic

20 inches

17-K

Phreatic

40 inches _

Magmatic

21 inches

15-K

Phreatic

36 inches

14-K

Phreatic

24 inches

13-K

Magmatic

80 inches

12-K

Phreatic

10 inches

11-K

Phreatic

7 inches

10-K

Phreatic

12 inches

5-K to

Magmatic

175 inches

9-K

(about)

3lKlo~~

Magmatic

200 inches

4-K

(about)

Phreatic

11 inches

1-K

Magmatic

12 inches ^

Lava Flows

400 feet

5-U

Magmatic

8 inches

4-U

Magmatic

8 inches

3-U

Magmatic

8 inches

2-U

Magmatic

12 inches

1-U

Magmatic

18 inches

LOCATION OF
MAXIMUM
THICKNESS

THICKNESS ON
WINDWARD RIM

TYPE OF
SURFACE ON
WINDWARD RIM

S.W. Floor Kilauea
S.W. Rim Kilauea

1 inch

0

Forest

West Rim Kilauea
S.W. Rim Kilauea
S.W. Rim Kilauea

3 inches

7 inches

16 inches

Exposed

Exposed

S.E. Rim Kilauea

6 inches

S.W. Rim Kilauea
S.W. Rim Kilauea

S.E. Rim Kilauea

3 inches

0

10 inches

Exposed

Exposed

North Rim Kilauea
N.W. Rim Kilauea
S.W. Rim Kilauea

10 inches

5 inches

2 inches

Exposed

Exposed

S.E. Rim Kilauea

25 inches

S.E. Rim Kilauea

30 inches

N.E. Rim Kilauea

N.E. Rim Kilauea

1 1 inches

12 inches

? ?

North Rim Kilauea
N.W. Wall Kilauea
N.W. Wall Kilauea
N.W. Wall Kilauea
N.W. Wall Kilauea
N.W. Wall Kilauea

breaks, erosional breaks, and breaks represent¬
ing exposure as a surface for an important time
interval. The thickest “blanket” occurrence
(contrasted to pocket accumulations) in each
small area is stated as the thickness of individual
layers in the composite section; likewise the
greatest number of separate layers found in
each small area is used to represent a given
eruption series. Thus, the composite section
cannot be duplicated in any single section in
an area, but represents the maximum number
and blanket thickness of recognizable layers
in each area. Similarly, composite sections were
compiled for areas away from the rim in each
major direction.

Cross correlation of these composite sections,
depending as much on correlations of time-
interval patterns as on a few recognizable hori¬
zon layers, has yielded a composite stratigraphic
column and chronologic sequence of explosive

activity for Kilauea. This is summarized in
Table 1; the details of the most critical sections
and the writer’s correlation between sections are
presented in Table 2. Eruptions of the two
separate series have been numbered serially,
i.e., 1-U for the oldest Uwekahuna tuff erup¬
tion, and 1-K for the oldest Keanakakoi erup¬
tion.

An overestimate of the time represented by
some of the erosion surfaces may have yielded
too great a number of eruptions. On the other
hand, eruptions of the magnitude of the 1924
explosions might easily have been omitted en¬
tirely because of failure to discover remnant
patches of their deposits. On the present rim
surface, diligent search will find only a few
patches of 1924 ash remaining (Plate IB and
1C). Comparable remnant patches, buried in
the stratigraphic section, could easily be over¬
looked.

280

PACIFIC SCIENCE, Vol. II, October, 1948

Fig. 1. Kilauea Crater and vicinity drawn from pares of the Glenwood and Kilauea Crater Quadrangles,
U.S.G.S. topographic maps of Hawaii. Contour interval, 100 feet. Heavy contour lines are used on surfaces
of the Kilauea dome which are apparently undisturbed by faulting; light lines are used for surfaces disturbed
by faulting. Lettered localities are discussed in the text.

In the following text, it will be evident that
descriptions of materials are presentations of
fact, while appraisals of time intervals and cross
correlations are presentations of the writers
interpretation.

UWEKAHUNA FORMATION

The oldest pyroclastic material which can be
associated definitely with the central crater of
Kilauea, the Uwekahuna formation ( Stone,
1926: 27-28), is found in type locality in out¬
crops in the base of the northwest crater wall.
The thickest section has been buried by the

1919 lava flow, but it was photographed by
Jaggar in July, 1913 (Plate 2B), and cliff de¬
tails in the photograph can be identified now
in the field, making it possible to locate the
exact position of the buried outcrop (locality
A in Fig. 1 and Plate 2 A) and to determine
that the top of the tuff lies about 16 feet below
the present surface of the 1919 pahoehoe lava
at an estimated altitude of about 3,630 feet.
This deposit was described by Sidney Powers
(1916: 230) as "...exposed to a thickness of
only 17 feet and for a length of about 500 feet,
at the base of a cliff 170 feet in height.” The
present top of the cliff, above this deposit of

Eruptions of Kilauea — POWERS

281

Fig. 2. Generalized pattern of distribution of several eruption deposits, shown at one-half the scale used
in Figure 1. A, reticulite from eruption 1-K; B, composite vitric deposits from eruptions 3-K to 9-K, inclu¬
sive; C, vitric deposits from eruption 13-K; D, lithic deposits from eruption 15-K; E, lithic deposits from
eruption 17-K; F, lithic deposits from the 1790 eruption.

Uwekahuna tuff, has an altitude of about 3,920
feet, indicating that the tuff lies beneath nearly
300 feet of bedded lava flows in the wall of the
present crater, rather than the 170 feet sug¬
gested by Powers’ sentence.

The most extensive occurrence of the Uwe¬

kahuna tuff is an apparently continuous, nearly
horizontal deposit extending for about 5,000
feet (between localities B and C in Fig. 1 and
Plate 3A), interbedded in the lavas of the
northwest wall of Kilauea crater, and exposed
in outcrops between talus fans at the base

282

of the cliff. At B the tuff is 3 feet thick, lies
10 feet above the present crater floor at an
altitude of 3,560 feet, and ends abruptly to the
northeast against the edge of a rim block which
has slumped about 20 feet with respect to the
rim section in which the tuff is exposed —
enough displacement to bury the tuff layer
beneath the crater floor if it continues into the
slumped block. At C the tuff is exposed at the
base of the crater wall at an altitude of 3,590
feet, then follows an unconformity which trun¬
cates some lava layers in the cliff, rising steeply
to the southwest to an altitude of about 3,690
feet where it levels off for a short distance be¬
fore it disappears to the southwest behind a
rim block which has slumped about 300 feet
with respect to the main crater wall containing
the tuff outcrops (Plates 2 A and 3A).

A third occurrence of tuff, interbedded in
lavas of crater-rim age, is found at the bottom
of the largest tree mold under the surface
pahoehoe flow a third of a mile northwest of
that corner* of Kilauea crater (Doerr, 1933:
3-7). Here, a group of koa trees apparently
grew in soil formed on a surface blanket of
tuff. At least 20 of these trees were surrounded,
without being toppled over, by the last lava
flood which covered this part of the outer slope
of Kilauea, and molds of their trunks are pre¬
served in the congealed lava. The largest mold,
having a maximum diameter of 7 feet and a
depth of about 18 feet, is so located that it acts
as a sump to drain the surface runoff from
several acres, and erosion by water entering the
mold has removed the tuff layer to form a
cavern as much as 20 feet wide and 150 feet
long, floored by an underlying pahoehoe lava
surface and roofed by the bottom of the upper
lava flow. The tuff exposed along the sides of
the cavern ranges from 0 to 36 inches in thick¬
ness. It mantled the irregular surface of the
underlying pahoehoe lava, and had been greatly
eroded before burial by the surface flow. The
tuff lies at an altitude of slightly over 4,000 feet.
For reasons stated in the following paragraphs,

PACIFIC SCIENCE, Vol. II, October, 1948

this tuff is correlated with the Uwekahuna for¬
mation of the type locality.

The type deposit of the Uwekahuna tuff (in
the base of the northwest crater wall) ranges in
thickness from a few inches to over 7 feet. It
consists of several beds of fine to coarse vitric
pumice, intercalated with variable thicknesses
of lithic debris mantling an unmodified surface
of pahoehoe lava. A typical section (locality S
on Plates 2 A and 2C) is:

Inches

Bottom of overlying lava

Coarse vitric pumice . . . 3

Oxidized desert surface

Coarse pumice with a few lithic fragments . 6

Oxidized desert surface

Agglomerate of lithic dust and fragments . 15

Oxidized desert surface

Coarse pumice with a few lithic fragments . 4

Oxidized desert surface

Coarse vitric pumice . . . . . . 8

Oxidized desert surface

Fine vitric shards grading down to coarser . 12

Surface of underlying lava flow

Most outcrops show the five layers of vitric ma¬
terial separated by oxidized desert surfaces; the
amount, coarseness, and texture of the inter¬
calated lithic debris vary greatly from outcrop
to outcrop. The section on the outer slope of
the Kilauea dome at the bottom of the tree
mold consists of:

Inches

Bottom of overlying lava flow

Vitric pumice . . . . . . . 2

Oxidized surface

Medium vitric pumice . . . 5

Oxidized surface

Fine to medium, bedded vitric pumice . 6

Oxidized surface

Fine, bedded vitric pumice . 9

Oxidized surface

Very fine vitric ash and pumice . 3

Surface of underlying lava flow

Thus, five epochs of magmatic fountaining,
separated by times of quiet but with no inter¬
vening deposition of lithic debris, are repre¬
sented at the tree mold locality. If the variable
lenses of intercalated lithic debris are ignored
in the type section, the five vitric layers in the

TABLE 2. CORRELATION OF COMPOSITE SECTIONS

The terms "dust,” "sand,” "gravel,” and "rubble” refer to beds of crystal and lithic fragments, graded somewhat as to size of particle; the terms "ash,’
shard,” and "pumice” refer to beds of primary vitric fragments, graded similarly as to size. "Piso” denotes pisolitic structure of the fine materials, "ass’

_£ _ _ _ „ ■ ... ■ .1 _ _ _ r 1^1 l_s__£ - .. ... -f- - «. _ . 1 i * _ Lai. « ai. _ _ «- . e . . . . . - _ - 7

TABLE 2. CORRELATION OF COMPOSITE SECTIONS

The terms "dust,” "sand,” "gravel,” and "rubble" refer to beds of crystal and lithic fragments, graded somewhat as to size of particle; the terms "ash,”
"shard,” and "pumice" refer to beds of primary vitric fragments, graded similarly as to size. "Piso” denotes pisolitic structure of the fine materials, "agg"
refers to an unsorted aggregate of lithic fragments ranging from dust to large blocks, and "pum agg” refers to rare beds of vitric lapilli which contain a few
lithic blocks. Breaks between layers are represented by symbol lines as follows: breaks in deposition . desert surface ?$$$$$*; humus or soil surface sSSSS.

Eruptions of Kilauea — POWERS

283

type sections of the Uwekahuna tuff may be
correlated satisfactorily with the five vitric layers
found at the tree mold locality. Future study
may indicate that the lithic debris represents
deposits from phreatic eruptions; the present
evidence suggests that the composition and dis¬
tribution of the lithic lenses are too erratic even
to represent phreatic eruption deposits. At
present it is thought that they are talus debris
and outwash material interbedded with the
mantle deposits of pumice on the floor of an
old crater near the base of an old crater wall.

Of the tuff pictured in Plate 2B, Sidney
Powers (1916: 230) gives the following de¬
scription: "The beds are composed principally
of yellow ash with some rock-fragments 1-2
inches in diameter, lava droplets, thread-lace
scoria, and a few bombs 6 inches in length.”
Wentworth (1938: 88) discounts the possible
correlation of this tuff with the Pahala tuff.
Macdonald thinks it may be a continuation of
the tuff still exposed northeast of Uwekahuna
and it is so considered by the writer.

To the northwest and the northeast, beyond
the edge of Kilauea surface flows, a deposit of
partly consolidated crystal-vitric tuff is found
which possibly correlates in age with one of
these Uwekahuna magmatic eruptions. In the
Bird Park area (Fig. 1), this bed lies on pala-
gonitized Pahala tuff under the edge of an aa
flow from Mauna Loa, which in turn is older
than the lowest member of the Kilauea surface-
ash deposits. About two miles northeast of
Kilauea, at about 3,800 feet (locality D, Fig. 1 ),
a bed of coarse, slightly palagonitized vitric
shards lies on top of the completely palagoni¬
tized Glenwood tuff under a layer of unaltered
surface ash from Kilauea.

All of these tuff deposits are considered parts
of the same formation— the Uwekahuna tuff —
deposited on different parts of an older Kilauean
dome with a collapse caldera in the summit.
The tuff is considered to be the remnant de¬
posits of five epochs of intense lava fountaining
or magmatic explosion separated by intervals

of relative quiet measured in at least tens of
years.

Studies now in progress on the rate of refor¬
estation in given climatic zones may eventually
yield data making useful the state of reforesta¬
tion as a quantitative indicator of elapsed time
since a given area was a new volcanic surface.
At present, the use of forest growth as an
indicator of the age of a surface is not much
better than intelligent guessing, but one is
tempted to use it in evaluating the lapse of time
between the eruption of the pre-Uwekahuna-
tuff pahoehoe flow and the destruction of the
koa grove by the post-tuff flow at the tree mold
locality. Present climatic conditions between
the tree mold locality and an area just north¬
east of Keanakakoi are not widely different;
the average annual rainfall is approximately 60
to 70 inches at both places, with slightly more
fog at the tree mold area. Deposits from the
1790 eruption average over a foot thick on
the Kilauea rim just north of Keanakakoi, but
thin to 3 inches in less than a quarter of a
mile to the northeast. The present surface is
practically barren of tree growth north of Kea¬
nakakoi, but supports a dense forest of ohia and
amaumau fern a quarter of a mile to the north¬
east. In fact, the transition from dense growth
to barren surface occurs in a zone about 300
yards wide in which scattered trees and ferns
have obtained a foothold (Plate 1A). The
conclusion seems justified that the 1790 deposit
killed the surface vegetation up to a fairly
sharp line and that the advance of reforestation
over the new surface has progressed about 300
yards in 150 years. Judging from the present
relationship of the slopes of Kilauea to the
forested slopes of Mauna Loa, the tree mold
area could not have been closer than half a
mile (probably a mile) to living koa forest
after the eruption of the pre-Uwekahuna sur¬
face lava flow. The trees which formed the
molds hardly could have obtained foothold until
the end of the second Uwekahuna magmatic
explosion since its eruption of 9 inches of
pumice at this spot probably would have killed

284

PACIFIC SCIENCE, Vol. II, October, 1948

forest growth. The advance of the koa forest
across more than half a mile of barren surface
would seem to demand at least 300 years, and
growth of the tree with a diameter of 7 feet
would seem to require perhaps 300 years. The
time lapse between emplacement of the two
upper lava flows enclosing the Uwekahuna tuff
could well approach 1,000 years.

Above the lower outcrops of Uwekahuna
tuff, truncated by the present crater wall, are
at least 30 pahoehoe flows ranging in thickness
from 1 to 35 feet (Macdonald, in press) and
separated by flow contacts which show no evi¬
dence of prolonged erosion or soil accumulation.
This epoch of rapidly recurring eruptions of
fluid lava apparently filled the supposed old
crater, overflowed the old outer rim at least
one flow deep on the northwest (the tree mold
flow), and restored a constructional dome shape
to the summit of Kilauea. Reconstruction of
the probable shape of the dome by using present
surface contours in segments that have not been
faulted, and restoring the measurable amount of
slump in the down-faulted segments, indicates
that the probable summit of the dome was
elongated east-west across the north end of the
present caldera between Uwekahuna and the
east end of Kilauea Iki. On Figure 1, the con¬
tours of the unfaulted Kilauea surface are indi¬
cated by a slightly heavier contour line.

This epoch of rapidly recurring overflow ap¬
parently closed with the collapse of at least
part of the present caldera and with the start
of a series of explosive eruptions which de¬
posited the members of the Keanakakoi forma¬
tion (Wentworth, 1938: 92) on the surface
of the present caldera rim.

KEANAKAKOI FORMATION

The Keanakakoi formation includes all frag¬
mental deposits emplaced on the present rim of
Kilauea by explosive eruption of the summit
crater. In theory it should represent all of the
late explosive eruptions, but of necessity any
explosions too feeble to deposit an appreciable

layer of material on the rim exclude themselves
because they have left no visible record. The
present interpretation identifies deposits from
21 different explosive eruptions separated by
breaks representing time lapses to be measured
at least in years. Not less than seven of these
breaks were long enough for re-establishment of
a vegetative cover on the humid windward rim.
Ten of the eruptions involved only phreatic
explosions; 11 were magmatic explosions or in¬
tense lava fountaining.

The oldest layer of known pyroclastic material
on the surface of the Kilauea rim lavas is
the "reticulite” (Wentworth, 1938: 93) or
"thread-lace scoria” (Dana, 1891: 163). In
places on the northeast rim, thin patches of
lavender clay are found on the lava under the
reticulite, but it is not known whether this clay
represents the remnants of an earlier explosive
eruption deposit, or is merely surface detritus
which was mechanically eroded from the surface
of bare pahoehoe, concentrated into pockets,
and mixed with accumulated humus. Oxidation
of the glassy pahoehoe skin is found everywhere,
though it cannot be demonstrated that this oxi¬
dation occurred before deposition of the reticu¬
lite. The surface of the clay pockets is also
oxidized and contains humus, so it is quite
probable that a vegetative cover had at least
partially developed on the northeast rim prior
to the eruption of the reticulite. This deposit
can be found in blanket thicknesses of about
12 inches on the north and east rims of the
crater (Fig. 2A). Thin blanket deposits and
pocket accumulations have been found entirely
around the rim and as far away as the Bird
Park. It is found under the edge of the Kea-
moku aa flow from Mauna Loa.

It is difficult to appraise the time interval
between deposition of the reticulite and its
burial by the next eruption deposit. Charcoal
fragments are found on the surface of the
reticulite on the northeast rim. They may be
relics of vegetation burned by the deposition of
the reticulite, or they may represent vegetation

Eruptions of Kilauea-— POW ERS

285

that grew on the surface of the reticulite. The
reticuhte is a material which could support
vegetative growth and show very little decom¬
position or mixture with humus.

The second eruption was phreatic, and de¬
posits from it are found only on the north and
east rims. On the slumped rim-blocks below
Waldron’s Ledge along the Volcano House-
Halemaumau Trail, are 36 inches of mixed,
tan pisolitic clay and lithic fragments up to a
cubic foot in volume. The deposit on Byron’s
Ledge, at the north end, is similar in make-up,
but only 6 inches thick. On the Steaming Flat
(N. Rim 3,950 in Table 2), the deposit con¬
sists of 5 inches of pisolitic tan clay overlain
by 6 inches of layered dust and fine sand. This
11 -inch blanket on the Steaming Flat probably
would have destroyed existing vegetation. A
humus layer on its top indicates that some soil
was formed and a vegetative cover developed
again before the next eruption. Scattered pumice
is mixed in with the soil surface in many places,
and a thin pahoehoe lava flow covered part of
the north end of Byron’s Ledge during this time
interval.

Remnant deposits from the third and fourth
explosive eruptions range in thickness from
about 200 inches west of Keanakakoi (S. Rim
3,650 in Table 2) to 2 inches on the top of
Uwekahuna Bluff (N.W. Rim 4,075 in Table
2). A 12 -inch section on the Steaming Bluff
consists of:

Inches

Humus layer at old surface

Fine vitric shards....................................... . 2

Wind-blown vitric shards, dune-like................ 1

Fine vitric shards grading down to pumice...... 2

Depositional break

Pea-size vitric lapilli . 2

Depositional break

Fine vitric shards grading down to pumice...... 2

Depositional break
Fine vitric shards grading down

to coarse pumice...................................... 3

Humus layer, surface of second eruption deposit

The thickest section of these vitric deposits
(near Keanakakoi) is in a cliff which is in¬

accessible for detailed study. In the walls of
a gaping surface crack southeast of Cone Peak,
under the crack flow, the section is 108 inches
of alternating beds of fine, dun-colored pisolitic
vitric shards and coarser pumice lapilli up to
one-half inch in diameter. One mile east (local¬
ity E in Fig. 1 ) , the section consists of about 80
inches of bedded, dun-colored vitric shards;
another half a mile to the east, just northwest
of Ahua Kamokukolau, the section is only 6
inches of unconsolidated, apparently wind-
drifted, dun-colored vitric shards.

Following the deposition of these vitric beds
in the third and fourth eruptions, a complete
vegetative cover became re-established on the
windward rim, and pronounced erosion occurred
on the leeward rim (Plates ID and 4B).

A series of at least five magmatic explosive
eruptions next deposited beds of vitric shards
and pumice totaling over 20 inches on the
windward rim and over 130 inches on the lee¬
ward rim. A thin pahoehoe flow lies on top of
this series on the southeast rim of Kilauea near
Keanakakoi. The eruptions were separated by
time intervals long enough to produce exposed
surface features but too short to permit the
growth of vegetative cover on the windward
rim; however, recovery of the windward rim
vegetation followed the last of these eruptions.

The small size and fragmental shapes of the
shards making up a large part of these thick
vitric beds and the presence of pisolitic struc¬
tures, seem to indicate that the eruptions which
produced them were more violently explosive
than the lava fountains which have been ob¬
served at Kilauea (or Mauna Loa) in recent
years. On the other hand, the explosions were
less violent than some of the phreatic blasts
which lifted fine material into the upper air
above the trade winds, because the distribution
of the vitric beds (Fig. 2B) is typical of deposi¬
tion under trade-wind influence.

The extremely rapid thinning of these beds
of vitric ash in all directions is very striking.
Beds 100 inches in thickness half a mile due

286

PACIFIC SCIENCE, Vol. II, October, 1948

southwest from the present leeward rim of the
crater decrease to less than 10 inches in the
next half mile to leeward. The size of particles
decreases rapidly also; no particles reaching a
diameter of Vi inch are found in the beds of
the original blanket deposit at distances 2
miles to leeward. Pieces of frothy pumice con¬
siderably larger than this are found in pockets
as much as 4 miles distant, but their occurrence
indicates, without much question, that they have
been skipped and rolled downhill by wind
drifting. This very localized distribution of the
products of the greatest vitric ash eruptions
known to have come from the central crater of
Kilauea, raises a serious question as to the abil¬
ity of earlier Kilauea crater explosions to have
contributed directly and appreciably to the ash
deposits of Hilina Pali and similar areas over
10 miles from the central crater.

The next three eruptions, numbers 10, 11,
and 12, were relatively weak phreatic explosions,
as the lithic particles in their deposits are all
smaller than medium gravel. The earlier deposit
is thicker on the southwest rim, perhaps due to
trade-wind control during deposition, while the
latter two are thickest on the windward rim.
The time intervening between each of these
eruptions was insufficient for reforestation, but
the time following the third one was long
enough to produce a vegetative cover, as evi¬
denced in the section on the southeast rim
at the edge of the humid area. A good soil is
present on the deposit from this twelfth erup¬
tion on the northwest rim, but it represents a
telescoping of several ensuing time intervals
and uninterrupted growth of vegetation up to
the time of the seventeenth eruption.

The thirteenth eruption was essentially one
of intense fountaining, but locally several of the
explosions hurled out small amounts of wall-
rock fragments (probably from talus piles
within the craters) which are mingled with the
pumice. The deposit is localized (Fig. 2C):
80 inches of coarse and fine pumice with a few
lithic blocks are found on the south rim due
west of Keanakakoi; 11 inches of bedded

pumice with a few small lithic fragments are
found on the southeast rim; and a few inches
of pumice alone are found on the rest of the
circumference of the crater. A thin pahoehoe
flow, which issued from a concentric crack a
mile southwest of the crater and flowed over
several acres of surface, is correlated tentatively
with this magmatic eruption.

In detail, the thickest section of this deposit
consists of:

Inches

Desert surface

Fine vitric shards approaching dune sand . 4

Textural break

Bedded, fine to coarse pumice, some lithic

fragments . ..24

Textural break

Bedded, pisolidc vitric shards and coarse

pumice . 40

Textural break

Coarse pumice . . 12

Surface of underlying material

The fourteenth eruption was phreatic, and its
deposits remain only on the west and southwest
rim of the crater: 3 inches of lithic fragments
up to half an inch in diameter on the west rim,
and 24 inches of bedded lithic dust on medium
to coarse lithic fragments on the southwest
rim.

The fifteenth eruption was a phreatic explo¬
sion which deposited lithic blocks of over a
cubic foot in volume more than a quarter of
a mile beyond the present southwest rim and
on Uwekahuna Bluff (Fig. 2D). At the south¬
west rim, the deposit is 36 inches thick and con¬
sists of large to small lithic blocks mixed
through a matrix of pisolitic dust. Away from
the crater rim, the lithic fragments play out
rapidly from the thick layer of partly consoli¬
dated, fine pisolitic dust. At the contact of the
Kilauea slope against Mauna Loa at the foot
of Kipuka Puaulu (Bird Park), the section is
over 6 inches thick, consisting of an upper layer
of compact pisolitic dust, a middle layer of dust
and fine lithic sand, and a lower layer of piso¬
litic dust. This bed mantles the lower mile of
the Mauna Loa slope northwest of Kilauea. It

Eruptions of Kilauea — POWERS

is 6 inches thick at B.M. 3,633 where the belt
road climbs onto the edge of the Keamoku
flow 2 miles west of the crater. At F in Figure
1, east of Cone Peak, the section shows oxida¬
tion of the upper members and consists of 4
inches of dust to fine sand, 1 inch of pisolitic
dust, 2 inches of laminated dust on coarse sand,
and 1 inch of medium-sized lithic sand.

The time intervening between the thirteenth
and the sixteenth eruptions was long enough
for a vegetative cover to develop on the south¬
east rim. The fact that oxidation is deeper (in
the leeward sections) on the deposits of the
fifteenth eruption than on either the thirteenth
or fourteenth suggests that the greater part of
the time interval occurred between the fifteenth
and sixteenth eruptions.

The deposits of pumice, mixed with a few
lithic fragments, of the sixteenth eruption are
concentrated to thicknesses of 21 inches on the
south rim near Keanakakoi. At the edge of
the humid area (S.E. Rim 3,775 in Table 2)
4 inches of this pumice are topped with a good
humus layer. Elsewhere the surface shows much
erosion and oxidation of wind-blown surface
material and carries abundant drift pumice and
Pele’s hair.

The remnant rim deposits of the seventeenth
eruption range in thickness from 6 inches on
the northeast rim to 41 inches on the south¬
west rim (Fig. 2E) and are sufficiently well
preserved to suggest an appraisal of the explo¬
sive phases of the eruption. After an assumed
preliminary stage which left no record on the
rim, the first (recorded) phase was the most
violent phreatic explosion of the eruption and
was followed by a pause in which rains cleared
the air of dust. This is suggested by the basal
layer which is found as follows: 3 inches of
lithic fragments ranging from coarse sand up
to 3 -inch diameter (a few blocks exceeding a
cubic foot in volume) capped by a top film
of fine sand and dust on the northeast rim; 24
inches of lithic fragments ranging from a quar¬
ter inch up to 6 inches (a few blocks of as much
as 4 cubic feet in volume) capped by 1 inch of

287

pisolitic dust on the southwest rim; and inter¬
mediate thicknesses of similar materials on the
southeast and northwest rims. The second phase
included at least four explosions following one
another so closely that the air was not cleared
of fine dust between them. The deposit on the
northeast rim consists of four layers of coarse
to fine lithic sand totaling about 3 inches, and
on the southwest rim consists of four layers of
lithic lapilli (up to Vz inch in diameter) grad¬
ing into coarse lithic sand totaling about 15
inches in thickness. The third phase included
one explosion followed by rains localized on
the leeward side of the crater. This last explo¬
sion deposited a thin layer of lithic sand grad¬
ing into about an inch of non-pisolitic dust on
the windward rim and 6 inches of lithic lapilli
and coarse lithic sand capped by 4 inches of
pisolitic dust on the leeward rim.

No humus formed on these deposits on the
windward rim, but the surface features indi¬
cate wind work, crustation, and oxidation char¬
acteristic of exposure for an interval at least of
some years’ duration. Pumice, Pele’s hair, and
Pele’s tears scattered on the surface indicate
lava fountaining during the interval.

The deposits from the eighteenth eruption
range in thickness from 2 inches on the wind¬
ward rim to 20 inches on the leeward rim. A
possible separation of the explosions is again
suggested by the details of the section on the
southwest rim. At the start of the eruption,
four relatively small phreatic explosions de¬
posited a total of 10 inches on the southwest
rim made up of four layers of coarse lithic sand,
each capped by lithic dust, and a total of 2
inches of pisolitic lithic dust on the windward
rim. Two more phreatic explosions deposited,
respectively, a 4-inch and a 2 -inch layer of
coarse to fine lithic sand on the leeward rim
and a 1-inch layer of pisolitic dust mixed with
coarse lithic sand on the windward rim. Per¬
haps both of these phases occurred during trade-
wind rains. The last explosion deposited an un¬
sorted aggregate of lithic fragments in a matrix
of tan pisolitic dust all around the rim, ranging

288

PACIFIC SCIENCE, Vol. II, October, 1948

from 2 inches thick on the northeast to 12
inches on the south near Keanakakoi.

The surface of this deposit shows the effects
of prolonged exposure and wind erosion (Plate
4C) in every area, but no humus layer accumu¬
lated on the windward rim.

The remnant deposits on the crater rim from
the 1790 eruption (Finch, 1947: 1) range in
thickness from about 3 inches on the windward
rim to a maximum of about 44 inches on the
broad fault block forming the inner, western
rim of Kilauea (Fig. 2F). At this locality, the
deposits indicate at least three main explosive
phases. The lowest layer is 8 inches of lithic
fragments (up to 2 inches in diameter) grad¬
ing up into a thin layer of fine lithic sand. The
middle layer is 12 inches of unsorted, fine to
coarse, lithic fragments capped by a thin layer
of lithic dust. The upper layer, 24 inches
thick, consists of 18 inches of medium to large
lithic blocks (up to 6 inches) grading up into
4 inches of coarse to fine lithic sand, grading
into 2 inches of fine lithic dust. A mile south,
on the southwest rim, the deposit is about 12
inches of unsorted, fine to coarse, lithic frag¬
ments, and it continues in similar thickness and
composition around the south rim to the section
due north of Keanakakoi. From this point north¬
east along the rim, the deposit thins very rapidly
to 3 inches in less than a quarter of a mile
(Plate 1A). On all of the windward rim, the
deposit consists of unstratified material ranging
from dust to pebbles approximating a hen’s
egg in size, now mixed with humus as well as
some indistinguishable, fine material from his¬
toric eruptions. It makes up the bulk of the
present surface soil, though there has been no
chemical decomposition of the lithic fragments.
On the leeward desert rim, except where it is
covered by later deposits, coarse fragments from
this deposit have been concentrated and fine
material has been crusted to form the stony
pavement characteristic of the desert surface.

The thickness of this deposit decreases rapidly
in all directions away from the crater. A maxi¬
mum thickness of 6 inches can be found on the

outermost rim due west of Halemaumau at
3,800 feet altitude, on the broad slope east of
Cone Peak at 3,600 feet, and due south of
Keanakakoi at 3,600 feet. The size of fragments
decreases and the proportion of dust increases
with distance away from the crater rim. Many
blocks with a volume exceeding a cubic foot
and a few with a volume exceeding a cubic
yard are found on the rim around the south¬
westerly half; fragments up to 6 inches in
diameter are found to the southwest as far as
Cone Peak; fragments up to 2 inches in diam¬
eter are not rare to the southwest as far as the
3,400-foot contour; and many fragments reach
half an inch in diameter near Mauna Iki at
2,900 feet.

The deposit is made up almost entirely of
crystalline fragments and contains no juvenile
vitric pumice, though, in a small area, bread-
crusted blocks and cored bombs are common.
Near Keanakakoi, such fragments make up
nearly 1 per cent of the deposit and blocks and
cored bombs up to a cubic foot in volume are
found as much as half a mile from the crater
rim. They are very rare in the other quadrants
of the crater rim. The insignificant amount of
still-plastic material (at time of eruption) rep¬
resented by these bombs is considered to have
been "bench magma” left behind by the abrupt
withdrawal of the active magma column, and
the eruption is believed to have been made up
entirely of phreatic explosions.

Correlation of specific eruption layers in sec¬
tions somewhat distant from the rim of the
crater is almost entirely speculative. On wind¬
ward slopes an appreciable thickness of ma¬
terial was deposited by very few eruptions,
and any thin deposits lost their identity
by incorporation in the then-existing, surface
soil. To leeward, immediate attack by wind
erosion rapidly destroyed most thin layers of
ash, adding the coarser materials to the moving
sand dunes and transporting the fine material
entirely out of the area in which it was depos¬
ited. Beds of water-saturated pisolitic mud
which have been partly consolidated and sur-

Eruptions of Kilauea — POWERS

289

face-crusted during the initial drying out are
the only thin leeward layers which remain in
place for an appreciable length of time.

In the belt of young ohia forest a mile
northeast of the rim of Kilauea, the ash sec¬
tion and a possible correlation with the Keana-
kakoi series eruptions is:

Eruption

4 inches of surface humus grading into next
layer

2 inches of lithic sand and fine gravel . 1790

3 inches of pisolitic aggregate . 18-K

7 inches of lithic sand and gravel . 17-K

Humus layer

8 inches of lithic sand and gravel . 12-K

Humus layer

2 inches of pumice mixed with humus . 9-K

8 inches of vitric ash mixed with humus .

. . 8-K to 3-K

Humus layer

2 inches of fine clay . . . 2-K

8 inches of reticulite . . . . .....1-K

Two miles to the northeast, in a more mature
forest made up of ohia, tree fern, and koa, the
section is:

Eruption

Humus top layer

8 inches of lithic sand and fragments mixed

with humus . . . . . 1790 to 12-K

1 inch of pumice mixed with humus . 9-K

15 inches of vitric shards and ash . 8-K to 1-K

In the desert, 6 miles to the southwest of
Kilauea, at the locality of the human footprints,
there remain patches of two pisolitic layers,
probably representing beds of original deposi¬
tion, and much wind-transported, dune sand.
One section showed:

Inches

Shifting dune sand . 6

Crusted stony pisolite layer . . . . 2

Imprisoned dune sand . 36

Mud-cracked pisolitic layer . . . . . . 2

Imprisoned dune sand . . . 48

Pahoehoe lava surface

The lower pisolitic bed is made up of three
layers: at the bottom is half an inch of finely
laminated dust, next an inch of pisolitic dust,
and on top is another half an inch of finely

laminated dust (Plate 3B). The surface of this
bed is cut by mud cracks which extend down
part way into the pisolitic layer. The cracks
have been filled with fine drifted dust. In the
surface of the layer are found some of the
fossil footprints, impressed even into the middle
of the central pisolitic layer. The whole bed
has been consolidated enough to resist imme¬
diate removal by wind and rain, but it has
been and is being removed slowly by erosion.
In many places all of the upper laminated dust
and the middle pisolitic layers have been re¬
moved, leaving only a thin crust of the lower,
laminated, dust layer. In the section described
above, a trench was driven into the bank ex¬
posing a 10-foot section containing the two
footprint-bearing pisolitic beds with dune sand
between. In this 10-foot artificial exposure, the
lower bed ranged in thickness from 2 inches to
half an inch in places where wind scour had
removed the upper two-thirds of the bed prior
to its burial by the dune sand.

The upper footprint bed consists of mixed
sand, lithic fragments (up to a half inch in
diameter), and pisolitic dust and ranges in
thickness from 1.5 to 2 inches. Its surface is
protected by a thin crust, and the entire layer
is partly consolidated. The scattered remnants
of this deposit in the footprint area have not
yet been traced with any assurance into the
continuous layers nearer the crater which can
be correlated with eruptions with some con¬
fidence. It appears reasonable to assign the
upper, crusted footprint layer to the 1790 erup¬
tion, chiefly because it is the youngest and be¬
cause it is associated with the known presence of
Hawaiians in the area during the eruption. On
the other hand, it might be argued that the two
pisolitic layers should be correlated with the
two phreatic eruptions which made the thickest
deposits remnant on the southwest rim near the
crater, namely the seventeenth and fifteenth
eruptions. Possibly more detailed field work
may yield data on which a definite correlation
may be based. At present, it seems certain that
the two footprint-bearing layers are products of

290

PACIFIC SCIENCE, Vol. II, October, 1948

two different phreatic eruptions which prob¬
ably should be chosen from among the 1790,
the seventeenth, and the fifteenth eruptions.

Evaluation of the time span covered by the
eruptions of the Keanakakoi formation can be
no better than a guess at present; but perhaps
a guess, tempered by the impressions of rela¬
tive time intervals and periods of exposure that
one gets while "living with the problem” in the
field, is worth recording.

The present forest cover on the northeast
rim of Kilauea is probably the heaviest tree
growth which has developed in any period since
the emplacement of the upper lava flow of this
part of the crater rim. The deposits of the
1790 and the eighteenth eruption on this part
of the rim are too light to have killed well-
established trees, though each probably killed
small ground-covering vegetation. Deposits from
the seventeenth eruption were thick enough
probably to have killed all vegetation as far as
half a mile to the northeast of the rim. Judging
from comments of early visitors, a good ground
cover and at least a scattered stand of scrubby
ohia were established on the immediate rim as
long ago as 1825. Taking everything into con¬
sideration, my impression is that the present
reforestation of the rim since the seventeenth
eruption has required approximately 400 years.
The next heaviest vegetative cover formed on

the northeast rim was established in the time
interval between the twelfth and the seventeenth
eruptions. The humus layer indicates heavy
ground cover, and molds of small ohia trees
have been found. The "kill” of the twelfth
eruption probably extended about the same
distance to the northeast of the rim as did that
of the seventeenth. This time interval may
have been about 300 years. Much less time
seems to be required for the intervals between
the ninth and the twelfth, and the fifth and the
ninth eruptions, since the humus is thin. The
vegetative cover developed on the deposits of
the fourth eruption consisted largely of fern,
ohelo, and other low growth and probably
could have developed in about 200 years. A
similar period apparently would suffice for the
time between the second and third eruptions.
An estimate of perhaps 1,500 years is suggested
for the total time covering the explosive erup¬
tions of the Keanakakoi formation.

CONCLUSIONS

Detailed study of the disconformities and
unconformities between the beds in the pyro¬
clastic deposits of Kilauea crater and appraisal
of the relative time intervals which they repre¬
sent, combined with study of the lithology of
the beds, have facilitated a preliminary catalog-

Plate 1 A. Abrupt transition from heavy forest cover to barren surface, possibly the southeast limit of
"forest kill” by the 1790 eruption, perpetuated because local climatic conditions have inhibited reforestation.
Looking northeast across Keanakakoi Crater (foreground) and the east end of Kilauea (left background)
from locality P. Photograph by H. A. Powers, September, 1947.

PLATE IB. Remnant of the mantle of 1924 bedded lithic ash, originally plus 5 inches thick, as exposed by
the small excavation in the foreground. This area is subject to torrential rains, but the absence of an incised
drainage net indicates that runoff erosion is subordinate in effectiveness to wind planation and undercutting.
The present surface is planed across the bedding of 1924 ash at a low angle, and a desert pavement of residual
larger fragments is beginning to accumulate. Photograph by H. A. Powers, September, 1947, at locality M.

PLATE 1C. Remnant of 1924 ash, originally plus 3 inches thick in this area, showing results of wind ero¬
sion by undercutting along least resistant layers. Most of the remaining patches of the 1924 ash mantle are
in slight surface depressions where moisture persists longer than on the adjacent slopes, perhaps retarding re¬
moval of the patches by wind erosion. Photograph by H. A. Powers, September, 1947, at locality N.

PLATE ID. Keanakakoi formation pyroclastics mantling the inner face of the southwest wall of Kilauea
Crater exposed by a short inflowing ephemeral stream. The bedded vitric deposits of the third and fourth
eruptions (lower left) are truncated by an erosion surface equivalent to a good humus layer on the humid
northeast crater rim. On the unconformity are the bedded vitric deposits of the fifth to the ninth eruptions
(center and right center). The upper beds represent the seventeenth to 1790 eruptions mantling a major
erosion surface from which all deposits of the tenth to the sixteenth eruptions have been removed. Photo¬
graph by H. A. Powers, September, 1947, at locality T.

Plate 1b

Plate lc

Plate Id

PLATE 2A. Northwest wall of Kilauea Crater seen from near Keanakakoi. The Uwekahuna tuff formerly
was exposed at A, and at present is exposed at C and S. The rim block containing A has slumped a greater
amount from its original elevation than has the rim block containing C and S. Photograph by H. A. Powers,
September, 1947.

Plate 2B. Bedded Uwekahuna tuff at locality A buried by the 1919 pahoehoe flow on the crater floor
which filled against the cliff as high as the dashed line. Vertical solid line indicates the span of a 13-foot sur¬
veyor’s rod located against points of the cliff face which have not changed since the photograph was taken by
T. A. Jaggar, July, 1913.

PLATE 2C. Uwekahuna tuff, 314 feet deep, lying on the basal pahoehoe flow exposed in the cliff at S. Base
of tuff is 5 feet above the present crater floor. Inter-eruption surfaces have been spotted on the negative.
Photograph by H. A. Powers, September, 1947.

PLATE 3 A. Kilauea, northwest crater wall, seen from Keanakakoi, in which Uwekahuna tuff crops out a
few feet above the crater floor between talus fans from C to B. Photograph by H. A. Powers, September, 1947.

PLATE 3B, Lower mud-cracked stratum retaining human footprints, lying on bedded dune sand about 6
miles southwest of Kilauea. Photograph of footprints by T. A. Jaggar, July, 1921; of vertical section by H.
A. Powers, August, 1931.

PLATE 3C upper crusted stratum of stony pisolite retaining human footprints in the same locality. Photo¬
graph by T. A. Jaggar, July, 1921.

PLATE 3D. Pumice of about 1815 bedded on remnants of 1790 gravel not exceeding an inch in thickness,
lying on pre-1790 stony pisolite. Below the 12-inch rule is the pisolitic agglomerate of the fifteenth phreatic
eruption. Photograph by LI. A. Powers, September, 1947, at locality W.

Plate 4a

Plate 4b

Plate 4c

Eruptions of Kilauea — POWERS

291

ing of at least 2 6 separate explosive eruptions
of Kilauea. Each eruption appears to have been
caused either by phreatic or by magmatic ex¬
plosions; no single eruption seems to require
both phreatic and magmatic explosions to ac¬
count for the lithologic constituents in its de¬
posit.

The Uwekahuna formation seems to have
been deposited on the floor and outer slopes
of an earlier caldera by at least five magmatic
eruptions separated by appreciable time inter¬
vals.

The Keanakakoi formation seems to represent
at least 10 phreatic and 11 magmatic eruptions
which have occurred since the last major over¬
flows of lava which form the present crater rim.

Details of distribution of the Keanakakoi
deposits promise to be useful in working out
the details of the late structural history of
Kilauea. For example, the slumped rim blocks
in the northeast corner carry the thickest deposit
from eruption number 2-K, suggesting early
collapse of these blocks; and most of the de¬
posits on Byron’s Ledge are not appreciably
thicker than those on the adjacent southeast
rim, which is 100 feet higher. Perhaps this
indicates a late date for the collapse of Byron’s
Ledge.

The study of the early Keanakakoi magmatic
explosion deposits has a direct bearing on the
problem of the contribution of the Kilauea

summit crater to the older Pahala ash formation
of Hilina Pali and other areas 10 miles or more
distant from the summit crater.

The chronological table of explosive erup¬
tions may conceivably be useful as an actual
time scale when coupled with results from
studies of rate of reforestation. The most mature
forest east and north of Kilauea Iki apparently
was never killed by any explosive eruption dur¬
ing the Keanakakoi time and thus may repre¬
sent uninterrupted encroachment from the slope
of Mauna Loa as much as 5 miles distant across
the surface of the last rim overflow of Kilauea.
A less mature forest extending from the crater
rim to about a mile northeast of the rim may
represent the total development of vegetation
since the seventeenth eruption, which probably
killed all vegetation to that distance. The growth
immediately on the northeast rim may represent
the recovery from a partial kill of vegetation
in 1790.

The general picture sketched in by this study
suggests these long-period fluctuations in the
intensity of lava pressure:

(1) pre-Uwekahuna, high-pressure, dome¬
building phase

(2) low-pressure collapse and explosive
eruptions of Uwekahuna tuff

( 3 ) high-pressure, post-Uwekahuna resump¬
tion of dome-building

Plate 4A. Keanakakoi pyroclastics on the outer slope of the southwest Kilauea rim. Deposits of the fif¬
teenth, seventeenth, eighteenth, and 1790 eruptions mantle an erosion surface which truncates beds of the
fourteenth to the tenth eruptions. The erosion surface marked by the hammer truncates the vitric ash of erup¬
tions earlier than the tenth. Details of the section in upper left center show in Plate 4B, and details of the
upper beds at the head of the re-entrant behind the figure show in Plate 4C.

PLATE 4B. Remnants of two layers of 1790 gravels, separated by a depositional break, lie on a desert sur¬
face truncating the stony pisolite of the eighteenth eruption. Erosion surfaces have been emphasized by re¬
touching on the negative. Beds from the seventeenth eruption reach 8 inches in thickness at the right of the
section and lie on the eroded surface of beds from the fifteenth eruption, the lowest layer which is continuous
across the face of the section. A lens of pumice from the thirteenth eruption (marked by pocket knife at left)
lies between remnant patches of beds from the fourteenth and twelfth eruptions. The vitric deposits from the
third and fourth eruptions lie beneath the horizontal desert surface (not retouched), which extends across
the photograph just below the grass clump on the right.

PLATE 4C. Gravel of 1790, from 4 to 6 inches thick, lying on the eroded layer of stony pisolitic mud
from the eighteenth eruption, originally 3 inches thick in this section. Erosion occurred after consolidation of
the pisolitic mud, as it truncates the bedding within the layer, and has cut the bed to a remnant 1 inch thick
in the photograph. Beneath the stony pisolite layer is a desert surface eroded on deposits from the seven¬
teenth eruption about 10 inches thick including the basal layer of coarsest fragments.

Photographs by H. A. Powers, September, 1947, at locality U.

292

PACIFIC SCIENCE, Vol. II, October, 1948

(4) low-pressure collapse of present caldera

( a ) partial collapse and early Keanaka-
koi magmatic explosions

( b ) severe collapse and phreatic explo¬
sions reaching a maximum in the
fifteenth and seventeenth eruptions

(5) possible gradual resumption of high-
pressure dome-building suggested by the
important crater-filling activity since
1790

REFERENCES

Bibliography of publications appearing since 1938
is believed to be complete; literature prior to that
date, completely discussed by Wentworth (1938), is
not listed here unless specifically referred to in the
text.

Dana, James D. 1891. Characteristics of 'vol¬
canoes. xvi+399 p., 15 pi. Dodd, Mead, and
Company, New York.

Doerr, John E., Jr. 1933. Tree molds in the
Volcano Golf Course, Hawaii Natl. Park
Nature Notes 3: 3-7.

Finch, R. H. 1925. The ash deposits at Kilauea
Volcano. Volcano Letter 17: 1.

- - — 1942. The surface ash deposits at

Kilauea Volcano. Volcano Letter 478: 1-3.

- - 1947. Kilauea in 1790 and 1823. Vol¬
cano Letter 496: 1-3.

Hitchcock, C. H. 1911. Hawaii and its vol¬
canoes. viii+3 14 p. Hawaiian Gazette Co.,
Ltd., Honolulu-.

Jaggar, T. A. 1921. Fossil human footprints
In Kau Desert. Hawaii. Volcano Observatory
Monthly Bui. 9: 114-118.

Macdonald, Gordon A. In press. Petrography
of the island of Hawaii. U. S. Geol. Surv.

Powers, Sidney. 1916. Explosive ejectamenta
of Kilauea. Amer. Jour. Sci. 4 1: 227-244.

Stearns, H. T., and G. A. Macdonald. 1946.
Geology and ground-water resources of the
island of Hawaii. Hawaii Div. of Hydrogra¬
phy Bui. 9. xiii+363 p. Honolulu.

Stone, John B. 1926. The products and struc¬
ture of Kilauea. Bernice P. Bishop Mus. Bui.
33. 59 p., 2 pi. Honolulu.

Wentworth, Chester K. 1938. Ash forma¬
tions of the island Hawaii. 3rd Spec. Rpt.,
Hawaii. Volcano Observatory, viii+183 p.
Honolulu.

The Origin of the Native Flora of Polynesia1

Edwin Bingham Copeland2

My invitation to this symposium states that
"It is desirable that a man should report on
his own research, but no speaker should feel
confined to this range. What we want is an
evaluation of recent research in each field as
it has come from or applies to the Pacific Basin,
and then the speaker’s own estimate of whither
this may be leading.”

My own research has been restricted to the
ferns. If the Philippines be within the field of
interest, this research has covered a period of
44 years. Twenty years ago, on the invitation
of the Bishop Museum, I prepared and pub¬
lished in their bulletin series a fern flora of
Fiji. A few years later, I did the same for the
Society Islands. Comparing these two groups
of islands, it was evident that their ferns had
migrated eastward. Their common ferns are
mostly common farther west, even as far away
as Malaya, and their endemics are derived from
these common, wide-ranging species.

The conclusion that the ferns of Polynesia
were immigrants from the Malay region was at
that time almost inevitable. It is old and reason¬
able dogma that evolution, in a large way, takes
place on continents, and that islands draw thence
their population. So, Asia, which used to in¬
clude the Sunda Islands, seemed the natural
source of the flora of the neighboring islands
to the East — Celebes, and then New Guinea,
and thence the Solomons, and Polynesia, and
New Caledonia, and New Zealand.

This interpretation of nature was supported
by a mental aberration: The flora of Java was

1 Read by invitation to the Pacific Section, A.A.A.S.,
at San Diego, June 17, 1947.

2 Research associate in botany, University of Cali¬
fornia, Berkeley, California. Manuscript received
April 22, 1948.

the first in this part of the world to be well
known, and when we found in the Philippines
a plant already known in Java, we simply
assumed Javan origin. We used to speculate
on the route of immigration, and to be surprised
that this route seemed to be by Celebes oftener
than by Borneo.

There were items which ought to have dis¬
turbed our confidence. For instance, a con¬
siderable number of species in the highland of
Northern Luzon are common to China and
even to the Himalayas. On the map, this looks
like the beginning of a route from the continent
to Polynesia; and men have migrated eastward
from the Philippines. But not one of the plants
in question reaches even to Southern Luzon.

My eyes were opened when, beginning some
sixteen years ago, I made a monographic study
of the family Hymenophyllaceae, the filmy ferns.
Contrary to anticipation, or even suspicion,
this study led me unescapably to the conclusion
that this family is entirely of Antarctic origin.
I abstain from presenting the detailed evidence
for this conclusion, because it has already been
digested and published (Copeland, 1938). It
is as positive as it will be when illustrated by
fossil evidence — of which the first item, from
the island Chiloe, reached me in May of this
year.

The Antarctic origin of Hymenophyllaceae
being positively established, it struck me that
it would be very strange if some or many other
ferns did not have a similar history. This was
already recognized for a few genera and species,
which were regarded as remarkable in this
respect. A comprehensive study of distribution
showed me at once that these were not in
reality exceptional cases, but were the con-

[293]

294

PACIFIC SCIENCE, Vol. II, October, 1948

spicuous examples of a general rule — that not
less than half of all ferns were of Antarctic
ancestry. Before I was ready to publish, this
figure grew to three-quarters; and I have more
recently raised it to 90 per cent. These figures
apply to the ferns of the world. Except for a
possible half-dozen Hawaiian species of freak¬
ishly scattered apparent geographic affinity,
there is not one of the ferns of the Pacific
islands which I do not now regard as of reason¬
ably direct Antarctic origin.

Let me presume to hammer this home, be¬
cause, as to the ferns, I speak with some author¬
ity. Contrary to our old ideas, the rich fern
flora of Malaya was not the source of Poly¬
nesian ferns — not of one Polynesian fern. For
it to have been so, the sun would have had to
rise in the west. New Guinea is richer in
ferns, the richest land in the world, and is the
immediate source of the most of the ferns of
the Philippines, Malaya, and southeastern Asia.
For a brief period, clinging to a fragment of
earlier prejudice, I pictured Polynesia as like¬
wise populated from New Guinea; but this is
not so likely.

Fiji is a plausible center of radiation of
Polynesian ferns. I do not believe that it re¬
ceived its ferns directly from New Guinea,
but it may have done so from somewhere along
the path of northward migration — from the
general region of the New Hebrides, as a sug¬
gestion. Eastward migration in these latitudes
is against the prevailing wind and against the
direction of storms, which explains the rapid
drop in population from Fiji to Tahiti, and
eastward in general.

Only Rapa among oceanic islands shows some
indication of having any ferns not of ultimate
New Zealand origin. If Juan Fernandez and
the Galapagos are included in the subject of
discussion, their ferns are of either direct
Antarctic (as to some of the ferns of Juan
Fernandez), or American origin — ultimately
Antarctic, but by way of Graham Land and
Tierra del Fuego. Almost all Hawaiian ferns
are derived ultimately from New Zealand.

So much for the ferns, about which I speak
from personal knowledge. It is unthinkable
that what occurred with ferns should not have
happened with other plants, to the extent that
there was a source of supply in Antarctica, and
at least as far as they depend upon the wind
for dissemination.

Doctor Hooker’s original postulate of an
Antarctic origin of austral floras was based on
his observation of the distribution of flowering
plants (Hooker, I860). Skottsberg, 30 years
ago (1915: 142), published a list of plants
common to New Zealand and sub- Antarctic
America, and without a plausible northern
source. This list includes representatives of 49
families, including Leguminosae, Compositae,
Rubiaceae, sedges and grasses. As to the species
on Skottsberg’s list, Antarctic origin is little less
than certain. Polynesian plants, even if com¬
mon to New Zealand, were excluded from that
list. For today’s purposes, they should of course
be included. If they and their immediate rela¬
tives, presumably of more recent evolution, be
included, a list thus compiled will include more
than half of the flowering plants of Polynesia.
At this point, my conclusion is that all Poly¬
nesian ferns and more than half of Polynesian
flowering plants are of ultimate Antarctic
ancestry.

For the most of the remainder of the flower¬
ing plants, no other ancestry or origin can be
postulated with greater plausibility. The situa¬
tion as to them is simply that the evidence has
not yet been discovered, or is not yet digested
and understood. There are no important
families of flowering plants except Myrtaceae
(regarding Proteaceae as relatively unim¬
portant) for which a nearly completely austral
origin can be affirmed with the same confidence
as for the ferns. I suspect all Polynesian
Rubiaceae of Antarctic origin; also, excepting
a few ubiquists, all Cyperaceae and Gramineae.

As is true of ferns, the "flowering floras” of
Polynesia and Malaya have much in common,
and the time has been when men could conclude
therefore that the poorer flora of Polynesia was

Polynesian Flora — COPELAND

295

derived from the comparatively rich flora of
Malaya. Today, it is more generally recognized
that the better explanation is that the two floras
have a common origin of their common ele¬
ments, and New Guinea is looked to for the
common source. But again, taking a lesson
from the ferns, and bearing in mind the diffi¬
culty of direct eastward migration, it seems
more reasonable to look for a common outside
source than to suppose that one area was
colonized from the other. To the extent that
this common flora is of ultimate austral origin,
it would have had to back-track to the Solomons
and New Hebrides before entering Polynesia,
and to do this against the winds and the oceanic
currents; I cannot believe that on any major
scale this ever happened. So, of present land
areas, I regard New Caledonia and the New
Hebrides as the roughly approximate region
from which Fiji and thence Polynesia received
the bulk of their vegetation.

Some such history as I propose has been re¬
jected by some very respectable authorities.
Thus Diels, in the Setchell Festschrift (1936:
191 ), treating of groups of plants of distinctive¬
ly austral occurrence, wrote that "it is impossible
to infer that they originated in the south.”
This is evidently not true, because I do so infer,
and so have Hooker, and Christ (1910: 248),
and Skottsberg, and still others.

More disquieting to me, and not so summarily
to be dismissed, is Merrill’s recent (1945)
statement: "It has also been suggested that
scattered throughout this vast region are cer¬
tain types that were apparently derived from
ancient Antarctica. But this idea is purely
theoretical and is one that can scarcely be
proved.” As I am expounding this theoretical
and scarcely provable idea, I must of course
concede that direct proof is impossible. But
other proof can carry conviction. A frog in the
milk pail is not direct proof that the milk has
been diluted. None of us has seen any plant
emigrate from Antarctica, and no plants are
now so emigrating. In the absence of direct
evidence, the indirect evidence by present dis¬

tribution, coupled with the fact, unknown to
Hooker and to Christ, that Antarctica was at a
definite and not too remote past time a fit place
for plants, is the nearest possible approach to
direct proof that they did migrate thence, into
and across temperate lands.

Although migration from Antarctica has long
ceased (the last warm era there was Miocene,
say twenty million years ago), I suppose that
it goes on now along the old paths where the
climate now permits. Because Merrill has pro¬
vided dependable figures on the distribution of
species, I will use them to show how this mi¬
gration seems to have occurred more recently
in two families, Elaeocarpaceae and Orchidaceae.

The pertinent figures on the occurrence of
species of Elaeo carpus are:

New Guinea . 114

Philippines . 49

Borneo . 40

Java . 18

New Caledonia . 30

Fiji . 12

Samoa . 6

Rarotonga . 1

Hawaii . 1

Merrill concludes, in harmony with general
practice, that New Guinea is the focus of dis¬
tribution. As to all lands farther west, I read
his evidence in the same way. As to the rapid
decrease in number of species from group to
group in Polynesia, his figures agree with the
evidence of the ferns in showing the great diffi¬
culty of eastward migration in this region. But
his figure of 30 species in New Caledonia is
very significant to me. New Caledonia has one-
fortieth of the area of New Guinea, but has
more than one-fourth as many species of
Elaeocarpus. The winds and the currents are
always from New Caledonia to New Guinea,
and I find it hard to doubt that this is the direc¬
tion of migration. The genus is of minor im¬
portance in Polynesia, but what species there
are seem most probably to have come from the
west or southwest, not from the northwest, not
from New Guinea.

296

As to the orchids, now regarded as the largest
family of plants, Merrill presents these figures
on the number of species:

New Guinea . 2,000

Philippines . 900

Fiji . 125

Tahiti . 30

Marquesas . . 4

Hawaii . 3

New Zealand has 66 species, and an old
figure (Hooker’s) for Tasmania is 74 — large
enough to point strongly to Antarctic origin.

The overwhelming mass of tropical orchids
are epiphytes. I have compiled no figures, but
am advised that in the Philippines, and in all
that part of the world, the ratio of epiphytes
to terrestrial species is more nearly ten to one
than five to one. In New Zealand, there are
7 epiphytes and 59 terrestrial species; in Tas¬
mania, 1 epiphyte and 73 terrestrial species.
The paucity of epiphytes in the far south may
be explained by the climate. But Polynesia is
tropical. Its ratio of epiphytic to terrestrial
orchids is about three to one — not more than
half of what should be expected if colonization
had been from New Guinea. The obvious con¬
clusion seems to be that it was colonized from
farther south, and that adaptation to tropical
conditions remains incomplete.

PACIFIC SCIENCE, Vol. II, October, 1948

Conclusion: The ferns, and a considerable
part of the flowering plants, of Polynesia are
of austral origin. As to the bulk of the flower¬
ing plants, I cannot claim, against Merrill’s
recently published judgment, that the present
tendency of thought is in support of my views,
but have shown that some of his detailed figures
do support them.

REFERENCES

Christ, Hermann. 1910. Die Geographie der
Fame. 357 p., 1 pi., 3 maps. G. Fischer,
Jena.

Copeland, E. B. 1938. Genera Hymenophyl-
lacearum. Philippine Jour. Sci. 67: 1—1 10.
Diels, Ludwig. 1936. The genetic phytogeog¬
raphy of the southwestern Pacific area, with
particular reference to Australia. In: Essays
in geobotany in honor of William Albert
S etch ell. T. H. Goodspeed, Ed. xxv + 319 p.
University of California Press, Berkeley.
Hooker, J. D. 1860. On the origination and
distribution of species: -Introductory essay to
the flora of Tasmania. Amer. Jour. Sci. II,
29: 1-25, 305-326.

Merrill, E. D. 1945. Plant life of the Pacific
world, xv + 295 p. Macmillan, New York.
Skottsberg, Carl. 1915. Notes on the rela¬
tions between the floras of subantarctic Amer¬
ica and New Zealand. Plant World 18(5):
129-142.

NOTES

On the Herding of Prey and the Schooling of the
Black Skipjack, Euthynnus yaito Kishinouye

While participating in the Bikini Scientific
Resurvey in the northern Marshall Islands in
the summer of 1947, the authors observed three
medium-sized black skipjack ( Euthynnus yaito
Kishinouye) herding a closely packed school
of several hundred scads ( Decapterus sanctae-
helenae (Cuvier)) over a large coral head in
Rongerik lagoon. For fully a 3 -hour period the
aquatic life on and about this coral head was
studied by use of the skin-diving technique, and
during this time this unusual group passed back
and forth frequently, apparently unconcerned
about our presence. Since Kishinouye’s reference
to such herding activity by tuna ( Tokyo Imp.
Univ., Col. Agr. Jour. 8(3): 382, 459, 1923)
is couched in generalities, and since our ob¬
servations on the black skipjack are contrary to
statements made by this author, we place these
underwater observations on record to clarify
several moot points in the feeding and schooling
habits of this species.

The predilection of these pelagic fish for this
coral head is understandable on the basis of the
part such coral heads play in the general eco¬
logical features of a lagoon in the Marshalls.
The coral head, typical of many, rose about 75
feet from the lagoon floor to approximately 25
feet below the surface of the water. It was
roughly circular, about 75 yards across, with a
rather steep talus slope forming its sides. The
surface of the mound was covered with sand and
larger coral fragments, interspersed with groups
of living corals of numerous species and sparse
patches of algae. Ecologically such coral heads
seem to occupy a physical niche comparable to
an oasis on land, as each supports a more
luxuriant growth of organisms than does the
surrounding terrain. Numerous species of com¬
mon coral fish, several medium-sized carnivorous
fish, and many plankton-feeding fish habitually
remain on and about these mounds. Thus the
elements essential to food chains are more or
less isolated in the vicinity of these coral heads,
and there both scads and some species of tuna

habitually forage. The fact that these condi¬
tions prevail afforded us an opportunity for re¬
peated observations upon these pelagic species
from a unique vantage point.

The three tuna usually followed the school of
scads rather closely, with one tuna at each rear
flank of the school and the third lagging behind
them. Now and then the scads would turn off
to one side, at which time the tuna on that side
would move forward swiftly and herd them
back into line. It became obvious that the school
of scads was prevented from leaving the area
over the top of the coral head, being herded back
whenever it moved over deep water. On one
occasion a laggard scad was swiftly picked off by
the rearmost black skipjack; however, except for
this incident the tuna made no attempt to prey
upon the scads during our period of observa¬
tion. Scads are uncommon in the lagoons in the
northern Marshalls and probably do not consti¬
tute a large proportion of the diet of this species
of tuna. Studies made by Jack Marr and Os¬
good Smith of the U. S. Fish and Wildlife Serv¬
ice (unpublished data) on the food habits of
33 black skipjack caught outside of the lagoons
failed to show any scads in the diet of this
species. Scads were most frequently encountered
in the stomachs of the dog-tooth tuna ( Gymno-
sarda nuda Kishinouye) which ranges in the
same area.

In his report on tuna herding prey, Kishi¬
nouye {op. cit.: 382) states: "Bonitos, except
Euthynnus yaito, are said to be very clever in
making a school of small fish very dense, by
swimming around the school of the victims, and
devouring stray or forlorn individuals gradually.
On the contrary, tunnies and seerfishes swim
into a school of victims, and disperse them. The
feeding of fish seems not always the same
throughout the year. The striped bonito is said-
to decline to take bait in certain seasons, gen¬
erally in midsummer.” In his discussion of the
food and feeding habits of the black skipjack,
Kishinouye {op. cit.: 459) says that the species

[ 297 ]

298

PACIFIC SCIENCE, Vol. II, October, 1948

is voracious and that it feeds primarily on small
fish and medium-sized plankton. He also de¬
scribes their feeding method, whereby they dart
swiftly into a school of small fish and scatter
them in the same manner as do the pelamids
and tunas. Our observations on the feeding
methods of the black skipjack certainly do not
coincide with those set forth by Kishinouye.
However, it is entirely possible, as he states, that
the feeding habits of the black skipjack may
vary throughout the year. Undoubtedly their
habits vary depending upon the type of food
available at any particular time of the year.

Kishinouye also implies that Euthynnus yaito
is usually a solitary fish and is not found in
schools. Schools of this species, ranging from a
few fish to many hundreds or even thousands of
individuals, are often seen around the Hawaiian
Islands throughout the year; and, during the
summers of 1946 and 1947, very large schools
of this species were common in the northern
Marshalls. It is quite likely that such schools
exist in the Marshalls throughout the year;
except for the summer months, however, no
observations have been made. Chapman
{Calif. Fish and Game, 32(4): 165-170, 1946)
states that black skipjack were present in large
schools at Midway Island, Johnston Island, and
Palmyra Island in the fall of 1943. Most of the
observations reported by Kishinouye were made

at the periphery of the range of Euthynnus
yaito; in this region smaller schools and perhaps
even solitary individuals would more or less be
expected.

The only observations on the herding of prey
by fish other than the tuna or tuna-like fish were
reported by Gudger {Carnegie Inst. Wash.,
Pub. 252: 75-7 6, 1918), who cites three in¬
stances involving the great barracuda, Sphyraena
barracuda (Walbaum). In each case only a
single barracuda was concerned; but since this
species is rather solitary, it would have been
rare indeed to see it engaged in a co-operative
effort. In two cases the prey was herded into
very shallow water; in the other instance the
small fish remained around piles and swam
among the rocks, not attempting to escape. In
none of these observations on barracuda did the
process of herding seem to be so well defined
or so efficiently accomplished as it was in the re¬
lationship of the black skipjack to the scads.
Of great significance was the fact that these skip¬
jack were engaged in a co-operative effort. —
Robert W. Hiatt, Department of Zoology and
Entomology, University of Hawaii, and Vernon
E. Brock, Division of Fish and Game, Territorial
Board of Agriculture and Forestry, Honolulu,
Hawaii. Published by permission of Chief of
Staff, Armed Forces Special Weapons Project,
National Military Establishment.

An Addition to the Fish Fauna of the Hawaiian Islands

Spencer Tinker, Director of the Waikiki
Aquarium, called the writer’s attention to a large
and showy chaetodontid which he had added to
the aquarium collection as a species unknown
to him and to local fishermen. This species was
readily identified as Pomacanthodes imperator
(Bloch), heretofore not reported from Hawai¬
ian waters. P. imperator is a reef dweller char¬
acteristic of the Indo-Australian faunal region
of which Hawaii stands as the northeastern
frontier; hence, upon faunal grounds, the occur¬
rence of the species in Hawaii is not inexpli¬
cable. However, in view of the intensive shoal
'water fisheries in the Hawaiian area together

with the volume of ichthyological collecting and
observing that have occurred here in the past, it
would appear that the species is very rare in
local waters.

This specimen, 198 mm. in total length, was
taken on January 10, 1948, in 15 fathoms of
water by a trap fisherman off Ewa, Oahu. The
fish, which was injured in the trap, died on
January 13, 1948, and is presently preserved in
the fish collection at the University of Hawaii
Marine Laboratory at Waikiki. — Vernon E.
Brock, Director, Division of Fish and Game,
Territorial Board of Agriculture and Forestry,
Honolulu, Hawaii.

NOTES

299

United States Committee on the Oceanography of the Pacific

The first meeting of the United States Com¬
mittee on the Oceanography of the Pacific was
held on June 19 and 20, 1948, at the California
Academy of Sciences, San Francisco, California.
This committee was established in 1947 by the
National Research Council through the Pacific
Science Board to replace the former American
National Committee on the Oceanography of
the Pacific. Members of the reconstituted com¬
mittee are Thomas G. Thompson (chairman),
Director of the Oceanographic Laboratories,
University of Washington; Richard H. Fleming,
Chief, Oceanographic Division, Hydrographic
Office, Department of the Navy, Washington,
DC.; Robert C. Miller, Director, California
Academy of Sciences, San Francisco; Charles J.
Fish, Supervisor, Pacific Oceanic Biology Project,
Oceanographic Institution, Woods Hole; Oscar
E. Sette, Director, Pacific Oceanic Fishery In¬
vestigation, U. S. Fish and Wildlife Service,
University of Hawaii, Honolulu; Roger Revelle,
Associate Director, Scripps Institution of
Oceanography, La Jolla; and Robert W. Hiatt,
Chairman, Department of Zoology and Ento¬
mology, University of Hawaii, Honolulu. Har¬
old J. Coolidge, Executive Secretary of the
Pacific Science Board, met with the Committee.

The organization of American participation
in oceanography and marine biology at the
Seventh Pacific Science Congress to be held in
New Zealand during February, 1949, occupied
most of the agenda. It was decided that presen¬
tations at the Congress should be classified in
three categories: (1) over-all reports covering
the decade since the last Congress, (2) local
regional programs, and (3) the high seas Pa¬
cific Basin program. Suitable titles covering
these phases were formulated and scientists ac¬
tive in the various fields were asked to prepare
papers for presentation at New Zealand. Two
symposia, one concerning conservation of ma¬
rine resources and the other involving marine
biogeographical provinces in the Pacific Basin,
were recommended for inclusion in the pro¬
gram.

A second topic on the agenda dealt with the
need for standardization of gear and methods,
and methods and arrangements for correlation
and integration of programs involving the
oceanography of the North Pacific Basin. Since

the absence of Japanese scientists from the New
Zealand Congress precludes the possibility of
considering these items in detail, it was deemed
advisable to hold a conference on this subject
after the Congress at a time when investigators
from all nations who would participate in
oceanographic research in the North Pacific
Basin would be able to attend. A subcommittee
composed of Charles J. Fish ( chairman ) , Robert
W. Hiatt, and William C. Herrington was ap¬
pointed to consider the matter.

Continuation of the work between Congresses
was considered highly desirable, and to this end
a recommendation will be made to the Science
Congress that each country appoint a represen¬
tative to serve as a member of a "Standing
Committee on Oceanography.” It is hoped that
implementation of suggestions for co-ordinated
work made at each Congress will be facilitated
through such special channels.

A special report by R. W. Hiatt on the de¬
velopment of the Hawaiian oceanographic re¬
search and training program in relation to the
establishment of the Hawaii Marine Laboratory
and the initiation of the vast program of re¬
search on tunas undertaken by the U. S. Fish and
Wildlife Service was presented. Plans have been
formulated or are being formulated for co¬
operation with several mainland institutions in¬
cluding the University of California and the
Woods Hole Oceanographic Institution. C. J.
Fish reported on the progress of the Pacific
Oceanic Biology Project which was created by
the Office of Naval Research upon recommen¬
dation of the Pacific Science Conference and is
being carried out under contract with the Woods
Hole Oceanographic Institution. Plans being
developed for continuation after September,
1948, emphasize training of oceanic biologists
who are needed before field work can begin.
Arrangements have been made with Rhode
Island State College and its Narragansett Ma¬
rine Laboratory to provide academic training
and field experience for graduate students prior
to their assignment for study under specialists
at Woods Hole. Close co-operation with the
U. S. Fish and Wildlife Service and the Ha¬
waiian training program will be maintained. —
R.W.H.

300

PACIFIC SCIENCE, Vol. II, October, 1948

The Pacific Oceanic Fishery Investigation

On July 1, 1948, the sum of $1,000,000 be¬
came available to the U. S. Fish and Wildlife
Service, through Congressional appropriation.
This sum will implement H.R. 859 (known
widely as the Farrington Fisheries Program)
which provides " . . for the exploration, investi¬
gation, development and maintenance of the
fishing resources; and development of the high
seas fishing industry of the territories and island
possessions of the United States in the tropical
and subtropical Pacific Ocean and intervening
seas.” The bill authorizes the U. S. Fish and
Wildlife Service to secure laboratories, vessels,
and personnel to investigate the biologic, tech¬
nologic, and economic problems concerning the
tunas and tuna-like fishes. Oscar E. Sette, for¬
mer Chief, South Pacific Investigations of the
U. S. Fish and Wildlife Service with headquar¬
ters at Stanford University, has been appointed

director of this vast program. A new laboratory
and administrative headquarters will be con¬
structed on the campus of the University of
Hawaii. Berthing facilities and warehouse space
have been secured at Pearl Harbor. Although
plans have not been completely formulated at
this writing, an extensive oceanographic in¬
vestigation will parallel the fishing operations.
Additional personnel in the fields of ichthyology
and fisheries biology have been added to the
University staff to provide undergraduate and
graduate curriculums designed to train ichthy¬
ologists, fisheries biologists, and oceanographers
for work of this type in the Pacific. Close co¬
operation between the staff of the Fish and
Wildlife Service laboratory and the staff of the
University will be maintained to provide the
best theoretical and applied training for stu¬
dents at all academic levels. — R.W.H.

Western Regional Conference for UNESCO

The regional conference for UNESCO
(United Nations Educational, Scientific and Cul¬
tural Organization) held at San Francisco, May
13-15, 1948, was attended by nearly 3,000 dele¬
gates from eight western states, Hawaii, and
Alaska. The delegates represented many dif¬
ferent types of organizations. Some, probably a
small minority, were delegates from states or
territories and represented the local chapters of
national scientific organizations by request of
the national officers.

At the opening plenary session the aim of
UNESCO was stated to be the prevention of
war. A great many scientists in attendance were
startled by the abruptness and simplicity of this
statement. Many delegates, acquainted with
some of the work already done by UNESCO in
aiding the restoration of libraries, exchange of

technical and cultural literature, and the like,
had conceived UNESCO to be a specialized seg¬
ment in the United Nations pattern but were
not prepared for the phraseology of wide, popu¬
lar appeal, nor for the subsequent extremely
general discussions, dealing with various aspects
of internationalism in relation to people, which
were lacking in any working approach to the
business of particular fields of endeavor on the
international level.

While it was for many scientists a stimulating
experience to attend such a meeting as citizens,
it may well be queried to what extent represen¬
tation from various specialized scientific or¬
ganizations will be sustained unless some pre¬
pared opportunity is afforded for conferences
on problems of international scientific liaison
and collaboration. — C.K.W.

NOTES

301

News Notes

The Pacific War Memorial announces the
establishment of a Memorial Research Field Sta¬
tion on Koror Island, Palau Archipelago. The
former Japanese meteorological station has been
turned over to the Memorial by the Navy De¬
partment. This building is of poured concrete,
with a penthouse. Electric power is available.

The Memorial is anxious to make the facil¬
ities available for use by qualified scientists as
soon as possible. By agreement with the War
Memorial the Navy Department will provide
transportation for qualified Pacific War Memo¬
rial personnel.

Inquiries concerning use of the station should
be addressed to: Pacific War Memorial, 44

West 50th Street, New York, N.Y.

# * *

Insects of Hawaii. — This inclusive treatment
of the insect fauna of the Hawaiian Archipelago
is being written by Elwood C. Zimmerman, As¬
sociate Entomologist, Experiment Station, Ha¬
waiian Sugar Planters’ Association, and Curator
of Entomology, Bernice P. Bishop Museum,
Honolulu. The first five volumes of a probable

12 to 15 total have been completed. Volume 1
was released July 27, the remaining four volumes
will be available during the summer and fall.
While this is a treatment of the insects of Ha¬
waii, it will be extremely useful throughout the
entire Pacific Area.

Volume 1, which is introductory, covers the
geological history of the Archipelago and va¬
rious phases of the origin and development of
the endemic fauna and flora. The remaining
volumes are devoted exclusively to discussion
of the Insecta. The groups covered in these
volumes are:

Vol. 2. Apterygota-Thysanoptera
Vol. 3. Heteroptera
Vol. 4. Homoptera: Auchenorhyncha
Vol. 5. Homoptera: Sternorhyncha

Elwood C. Zimmerman, Insects of Hawaii. Pub¬
lished by the University of Hawaii Press. Price per
set, paper, $24.00. May also be purchased as single
volumes: Vol. 1, $3.50; Vol. 2, $5.50; Vol. 3,
$4.50; Vol. 4, $4.50; Vol. 5, $6.00. Orders may
be sent to University of Hawaii Press, P. O. Box 18,
Honolulu 10, Hawaii.

Index

Acalyploa grandis var. genuma, 102, 111
Acanthaceae, 112

Acanthoclinus quadridactylus, 130
Acridotberes tristis, 62
Aero cirrus, 38
crassifilis, 38
frontifilis, 38

heterochaetus, 8, 38-39, 40
muroranensis, 38
uchidai, 38
validus, 38

Addition to the Fish Fauna of the Hawaiian
Islands (Note), 298
Additions to Galapagos Fungi, 71-77
Agardhiella Coulteri, 141
Agonostomus forsteri, 128, 129
Alaska King Crab Investigation, 3
Alaska, polychaetous annelids of, 3-58
Alaskan king crab, 14
Alaskan Salmon Commission (1903), 3, 4
Alauda arvensis, 63
albatross, black-footed, 132
Laysan, 66, 132

"Albatross” Expedition (American, 1902-1903),
3,4,79,274

"Albatross” Expedition (Swedish, 1947-1948),
67, 231-238

albatrosses, interbreeding of, 132
Alciopidae, 6

algae, marine, 140-141, 178
Alicata, Joseph E. :

Observations on Parasites of Domestic Animals
in Micronesia, 65-66
Alitta, 4
brandti, 25

Allan Hancock Foundation, 3
Allop hylus timorensis, 102, 111
Amage anops, 8
Amandava amandava, 59
Amaryllidaceae, 108
Amblyomma cyprium, 65
Ammotrypane aulogaster, 8
Amoebotaenia sphenoides, 66
Ampharete arctica, 9
brevibranchiata, 9
eupalea, 9
grubei, 9, 43
johanseni, 9
reducta, 9

Ampharetidae, 8, 43
1 Amp hicteis alaskensis, 9, 43
glabra, 9

Amphictene auricoma, 8
Amphinomidae, 1 1
Amphitrite cirrata, 9, 43-44
infundibulum, Al
palmata, 43, 44
radiata, 43, 44
robusta, 44
spiralis, 44
Anaitides, 4, 19-20
citrina, 6, 19-20

groenlandica, 6, 19
medipapillata, 6
mucosa, 6, 19

Anas boschas a. Freycineti, 121
laysanensis, 123
oustaleti, 121-124
platyrhynchos, 121-124
poecilorhyncha, 121-124
pelewensis, 122
superciliosa, 121-124
wyvUliana, 123
Anaspio boreus, 8
Ancylostoma caninum, 66
andesite line, 216-218, 220-222
Anelassorhynchus, 274, 277
inanensis, 275
porcellus, n. sp., 274-277
Angiopteridaceae, 272
Angiopteris Beecheyana, 272
eve eta, 272

animals, domestic, in Micronesia, parasites of,
65-66
annelids,

list of species known from Alaska, 6-9
new species of, 17, 23, 26, 30, 33, 37
polychaetous, of Alaska, 3-58
Anthopleura xanthogrammica, 196
Antinoe macrolepida, 6
sarsi, 6
Antinoella, 6
Aphrodita cirrhosa, 14
japonica, 6, 1 1
minuta, 15
Aphroditidae, 6, 1 1
Apistobranchidae, 1 1
Apocynaceae, 112
Araceae, 107-108
Arabellidae, 7, 29
Archey, Gilbert:

The Seventh Pacific Science Congress, 225-226
Arctonoe fragilis, 11
pulchra, 6
vittata, 4, 6, 11-12
Arcyria cinerea, 72
Arenicola glacialis, 4, 8
pusilla, 8, 39
Arenicolidae, 8, 39
Arhynchite, 277
Aricia arctica, 4

Aricidea heteroseta, n. sp., 7, 33, 36
Armandia bioculata, 8, 39—40
brevis, 8, 39

Arripis trutta, 128, 129
Artocarpus altilis, 109, 110
communis, 109, 110
heterophylla, 110
hirsuta, 110
incisus, 102, 109-110
integrifolia, 110
Jaca, 110
Philippensis, 110
Asclepiadaceae, 112

[ 305 ]

306

Asclepias curas savica, 104, 112
Ascomycetes, 73-74
Ascophanus argenteus, 73
carneus, 73

Asio flammeus sandwichensis, 61
Aspeira modesta, 46
Asplenium nidus, 100, 106
Astacus eur opens, 155
fluviatilis, 147, 155
Asychis lacera, new comb., 8, 42
similis, 8

Autolytus alexandri, 6
magnus, 6, 24
prismaticus, 6
autopasy, 187-191
autophagy, 187-191
autotilly, 187-191
autotomy, 187-191
Auxis, 262, 266-271
bis us, 270
hira, 269, 270
maru, 269
t hazard, 262-271

avian diseases, effect on avifauna, 61
avian introductions in Hawaii, 59—64
avian malaria, 61

Badhamia affinis, 72
gracilis, 72

Balanus glandula, 152
barnacles, lepadid, 80
barracouta, 128, 130
Barringtonia asiatica, 103, 111
Basidiomycetes, 74-75
bauxite, on Truk, 223-224
Bikini Scientific Resurvey, 297
Biology of the Lined Shore Crab, Pacbygrapsus
crass ipes Randall, 135-213
Introduction, 135—136
Taxonomy, 136-138
Synonymy, 136-137
Systematic Position, 137
Description, 137
Type Locality, 137-138
Distribution, 138-146

Geographical Distribution, 138-140
Habitat and Ecological Niche, 140-146
Sex Ratio, 146
Intermolt Cycle, 146-154
Period A, 150-151
Period B, 151
Period C, 151-152
Period D, 152-153

Ecdysis and Associated Phenomena, 154-168
Exuviation, 155—161
Seasonal Exuvial Periodicity, 161-164
Exuvial Frequency, 164-165
Age, 165-168

General Habits and Behavior, 168-187
Visual, Chemical, and Tactile Senses,
168-177

Locomotion, 177—178
Food and Feeding Habits, 178-179
Use of Chelipeds, 179-181
Activity Under Varied External Conditions,
181-182

The Influence of the Ocean on P. crassipes ,
182-184

Relationship of P. crassipes to Certain

PACIFIC SCIENCE, Vol. II, October, 1948

Other Species in the High Littoral Zone,
184-185

Precocious Young, 185-186
Intraspecific Territoriality, 186-187
Defensive Mutilation and Regeneration,
187-196

Defensive Mutilation, 187-191
Regeneration, 191-196
Predaceous and Parasitic Enerflies of
P. crassipes, 196-197
Reproduction, 197-205

Sexual Dimorphism and External Genitalia,
197

Age and Size at Sexual Maturity, 197-198
Copulation and Impregnation, 198-200
Extrusion and Incubation of Ova, 200-204
Hatching and Subsequent Growth, 204-205
Transitional Position of P. crassipes between a
Littoral and Terrestrial Existence, 205-208
Summary, 208-210
References, 210-213
"Bird Dog,” weather ship, 92-93
birds, introduction into Hawaii, 59-64
Blarina brevicauda brevicauda, 240
blennoid fish, new species of, 125-127
Bonelliidae, 275
booby, red-footed, 66
Boophilus annulatus australis, 65
Boraginaceae, 112
Botany :

Additions to Galapagos Fungi, 71—77
New Fern from Rota, Mariana Islands, 214-215
Origin of the Native Flora of Polynesia,
293-296

Plant Records from the Caroline Islands,
Micronesia, 272-273

Report on the Flora of Pingelap Atoll, 96-113
bovine piroplasmosis, 65
Brada granulata, 4, 8, 41
pilosa, 4
villosa, 8
Bridge, Josiah:

A Restudy of the Reported Occurrence of Schist
on Truk, Eastern Caroline Islands, 216-222
On the Occurrence of Bauxite on Truk, 223—224
Brock, Vernon E.:

An Addition to the Fish Fauna of the Hawaiian
Islands (Note), 298

A New Blennoid Fish from Hawaii, 125-127
The Poisoning of Bufo marinus by the Flowers
of the Strychnine Tree (Note), 132
Brock, Vernon E., and Robert W. Hiatt:

On the Herding of Prey and the Schooling of the
Black Skipjack, Euthynnus yaito Kishinouye
(Note), 297-298
Bryophyllum pinnatum, 102, 111
Bufo marinus, poisoning of, 132
Bunodactis elegantissima, 196
Bursera graveolens, 71, 72, 73, 74, 75
BUSHNELL, O. A.:

A List of Scientific Institutions in the Pacific
Area, 243-261
Buteo solitarius, 61

Callianassa calif ornien sis, 136
Callicarpa cana, 273

Callinectes sapidus, 136, 155, 157, 1 63, 168, 201
Callizona angelini, 6
Calophyllum Inophyllum, 103, 111

INDEX

307

Calorhynchus milii, 128
Cancer antennarius, 145, 195
magister, 136, 145, 163, 168, 177, 195
pagurus, 135, 147, 148, 155, 157, 163, 201
pro duct us, 195

cannibalism (among crabs), 179, 191
Cantherines s caber, 130
Capitella capitata, 8, 41
filiformis, 41
Capitellidae, 8, 41
Capparidaceae, 273
Capparis cor difolia, 273

Carcinides maenas, 136, 148, 168, 172, 176, 177,
183, 198

Carica Papaya, 103, 111
Caricaceae, 111

Caroline Islands, 96-113, 216-222, 223-224,
272-273

Kusaie, plant records from, 272-273
Palau, plant records from, 273
Pingelap, flora of, 96-113
Truk, bauxite on, 223-224
Truk, bauxite occurrence of schist on, 216-222
Carpodacus mexicanus, 59, 63
Castalia multipapillata, 4
Castronodus strassenii, 242
cattle, in Micronesia, parasites of, 65
Central America, juvenile Euthynnus lineatus and
Auxis thazard off, 262-271
yellowfin tuna off, 114-120
Ceodes umbellifera, 102, 110
Ceratonereis, 5
paucidentata, 5, 7
Chaetocbloa, 74
Chaetopteridae, 8, 37
Chaetopterus sp., 37
variopedatus, 8
"Challenger” Expedition, 231
Cheilonereis cyclurus, 7, 25
Chelidonichthys kumu, 130
chickens, in Micronesia, parasites of, 66
Chitinopoma, emended genus, 48—50
fabricii, 48
groenlandica, 48, 51
occidentalis, new comb., 9, 48-51
Chone gracilis, 9
infundibuliformis, 9
sp., 47

Chronology of the Explosive Eruptions of
Kilauea, 278-292
Chrysopetalidae, 6, 1 5
Cirratulidae, 8, 37-39
Cirratulus borealis, 4
cingulatus, 37
cirratus, 4, 8, 37
robustus, 8

Cistenides brevicoma, 4, 8
granulata, 8
hyperborea, 4, 8
Cladophora graminea, 141
sp., 141

Clastoderina Debaryanum, 72
Clerodendrum inerme, 104, 112, 273
Clymenella tentaculata, 8
Cocos nucifera, 98, 101, 107
Coelorhynchus australis, 129
Colocasia esculenta, 98, 99
var .antiquorum, 101, 107
Colubrina asiatica, 273

Comatricha elegans, 72
Combretaceae, 112
Compositae, 112
Congiopodus leucopaecilus, 130
Conservation in U. S. Trust Territory of the
Pacific (Note), 228

continental boundary lines, 217-218, 220-222
control of avian introductions, proposed plan for,
63-64

convective storms, 83
Copeland, Edwin Bingham:

The Origin of the Native Flora of Polynesia,
293-296
Coprinus sp., 75
Corallina officinalis, 141
Cordyline terminalis, 272
core samplers, 231
crab, deep-sea, 79-80

lined shore, biology of, 135-213
Crangon calif orniensis, 164
Crassulaceae, 111
Cribraria languescens, 72
violacea, 72

Crinum asiaticum, 101, 108
Crocidura sp., 240
Crucigera formosa, 47
irregularis, 9, 48
zygophora, 9, 47-48
Crustacea, rare Hawaiian, 78-80
Cryptonota, 15
Cryptopleura lobulifera, 141
violacea, 14 1
Ctenocephalides felis, 66
Ctenodrilidae, 1 1
Curvularia sp., 75
Cyathea affinis, 272
nigricans, 272
Cyatheaceae, 272

Cyclograpsus punctatus, 13 6, 155, 156, 161, 163,
168, 198, 199
Cyclosorus dentatus, 214
interruptus, 214
Cyperaceae, 107
Cyperus javanicus, 100, 107
Cyrtosperma Chamissonis, 98, 99, 101, 107-108

Dactylopagrus macropterus, 130
decapod Crustacea, records of, in Hawaii, 78-80
Decapterus sanctae-helenae, 297-298
Demonax, 45
Derris trifoliata, 102, 111
Dexiospira spirillum, 51
Diomedia immutabilis, 66, 132
nigripes, 132
Dioscoreaceae, 108, 272
Dioscorea ? korrorensis, 108, 272
sp., 101

Diplocrepis puniceus, 130
Dipylidium sp., 66
Disomidae, 1 1

Diurnal Weather Patterns on Oahu and Lanai,
Hawaii, 81-95

dogs, in Micronesia, parasites of, 66
domestic animals of Micronesia, parasites of, 65-66
Dorvillea pseudorubrovittata, 7, 29-30
Dorvilleidae, 7
Drilonereis filum, 1
nuda, 7, 29

echinoderms, 1 1

308

PACIFIC SCIENCE, Vol. II, October, 1948

Echinostelium minutum, 72
Echiuridae, key to, 277
echiuroid worms, 274-277
Echiuroidea, 275
Echiurus, 277
Egregia Menziesii, 141
Elaeocarpaceae, 295
Elaeocarpus, 295
Endocladia muricata, 14 1
Enipo cirrata, 6
Enteromorpha sp., 141
Eragrostis amabilis, 100, 107
Enoch eir japonicus, 139
Eteone calif ornica, 6, 19, 20-21
spetsbergensis, 6, 20
ethnobotany, of Pingelap Atoll, 99-106
Euchone analis , 9
Eudistylia polymorpha, 4, 9
tenella, 9
vancouveri, 4, 9
Eulalia viridis, 6, 20
Eulamia brachyurus, 128
Eunice longicirrata, 7
Eunicidae, 7
Eunoe , 4, 14
barbata, 6
depressa, 6, 14
nodosa, 6
senta, 6
truncata, 4

Euphorbia Atoto, 102, 111
Euphorbiaceae, 111
Euphrosine arctia, 6
bicirrata, 6
heterobranchia, 6
hortensis, 6
multi branchiata, 6
Euphrosinidae, 6
Euphrosyne multibranchiata, 5
Eupolymnia crescentis, 9, 44
heterobranchia, 9
nesidensis japonica, 9

Euthynnus, 262, 263, 265, 266, 267, 268, 270
alliteratus, 266
lineatus, 262-271
yaito, 262, 265, 297-298
Evarne triannulata, 13
Ev amelia triannulata, 6, 13
explosive eruptions of Kilauea, 278-292

Farrington Fisheries Program, 300

Fasciola hepatica, 65

fern, new species from Rota, 214-215

ferns, 293-296

Ficus sp., 102, 110

Fiji, mynah bird in, 62

Fimbristylis cymosa, 101, 107

fish, blennoid, new species of, 125-127

Fisher, Harvey I. :

Interbreeding of Laysan and Black-footed
Albatrosses (Note), 132
Laysan Albatross Nesting on Moku Manu Islet,
off Oahu, T. H., 66

The Question of Avian Introductions in Hawaii,
59-64

Fisher, Walter K.:

A New Echiuroid Worm from the Hawaiian
Islands and a Key to the Genera of Echiuridae,
274-277

fisheries program in the Pacific, 300
Fishes Taken in Wellington Harbour (Note),
128-130

Flabelligera infundibularis, 8, 40
Flabelligeridae, 8, 40
fleas, 66
flukes, liver, 65

flora, of Caroline Islands, Micronesia, 272-273
of Pingelap, 96-113
of Polynesia, 293-296
Fossaria ollula, 65
Freycinetia ponapensis, 272
Fucus furcatus, 14 1

Fulbright Act, further information on, 226-227

Fungi Imperfecti, 75-77

fungi, of Galapagos Islands, 71-77

Galapagos Islands, fungi of, 71-77
Gastroclonium Coulteri, 141
Gattyana amondseni, 6
ciliata, 4, 6
cirrosa, 6, 14
iphionelloides, 6, 14
imbricata, 6
senta, 4

Gelidium sp., 141
Geology :

A Chronology of the Explosive Eruptions of
Kilauea, 278-292

A Restudy of the Reported Occurrence of Schist
on Truk, Eastern Caroline Islands, 216-222
On the Occurrence of Bauxite on Truk (Note),
223-224

Transfer of the Hawaiian Volcano Observatory
(Note), 131

Geophysical Abstracts, note on, 228
Geotria australis, 128
Gigartina corymbifera, 14 1
Harvey ana, 141
leptorhynchos, 141
Gloniopsis somala, 73
sp., 73, 76

Glycera capitata, 7, 28
Glyceridae, 7, 28-29
Glycinde picta, 7, 29
wireni, 7

Gnathophyllidae, 79
Goniada annulata, 1
maculata, 7
Goniadidae, 7

Gonorhynchus gonorhynchus, 128
Goodeniaceae, 112
Gramineae, 107
Grapsus, 207
eydouxi, 136, 137, 138
Grateloupia Setchelii, 141
Guam, 65-66

Guettarda speciosa, 104, 112
Guttiferae, 111
Gymnosarda nuda, 297

Haliotis rufescens, 172
Halosydna brevisetosa, 4, 6, 12
Halosydnoides vittata, 11
Haploscoloplos, key to genus, 30
Haploscoloplos alaskensis, n. sp., 7, 30-32
elongata, 7, 30
panamensis, 32
sp., 30-32

INDEX

309

Harmothoe aspera, 5
hirsuta, 6

imbricata, 4, 6, 13
iphionelloides, 14
triannulata, 13
truncata, 13
tut a, 13

Harriman Alaska Expedition (1899), 4
Hartman, Olga:

The Polychaetous Annelids of Alaska, 3-58
Hawaii :

decapod Crustacea, records of, 78-80
explosive eruptions of Kilauea, 278-292
Laysan albatross nesting in, 66
new blennoid fish from, 125-127
new echiuroid worm from, 274-277
question of avian introductions into, 59-64
weather patterns in, 81-95
Hawaii Marine Laboratory, oceanographic program
of, 67-68, 299

Hawaiian Crustacea, records of, 78-80
Hawaiian Volcano Observatory, transfer of, 131
Hedy otis albido-punctata, 273
Helicosporium aureum, 75
guianensis, 75
Hemerocoetes waitei, 130
Hemigrapsus nudus, 141, 143, 1 44, 145, 146, 165,
166, 177, 178, 179, 184, 185, 186, 187, 204,
205, 206, 207, 208

oregonensis, 141, 142, 143, 1 44, 145, 146, 165,
166, 185, 186, 187, 204, 205, 207, 208
Hemipodus borealis, 7, 28
Hermadion, 4, 14
truncata, 6, 14
Herpestes sp., 61
Hesionidae, 6
Heterakis gallinae, 66
lingnanensis, 66
Heteromastus filiformis, 8, 41
Heteropale bellis, 15
Hiatt, Robert W. :

Biology of the Lined Shore Crab, Pachygrapsus
eras sipes Randall, 135-213
The Pacific Oceanic Fishery Investigation
(Note), 300

Records of Rare Hawaiian Decapod Crustacea,
78-80

United States Committee on the Oceanography
of the Pacific (Note), 299
Hiatt, Robert W., and Vernon E. Brock:

On the Herding of Prey and the Schooling of the
Black Skipjack, Euthynnus yaito Kishinouye
(Note), 299

Hippocampus abdominalis, 129
Hololepida magna, 6, 13
Hololepidella tuta, 4, 6, 13
Homarus americanus, 136
gammarus, 147
hookworm, 66
Hyalopomatopsis, 48
marenzelleri, 48
occidentalis, 48, 50, 51
Hydrocharitaceae, 107, 272
Hydroides norvegica var. groenlandica, 48
Hymenocera elegans, 79
Hysterographium, 73
guaranicum, 73
mori, 73-74

Ichthyology :

An Addition to the Fish Fauna of the Hawaiian
Islands (Note), 298

Fishes Taken in Wellington Harbour, 125-127
Juvenile Euthynnus lineatus and Auxis thazard
from the Pacific Ocean off Central America,
262-271

Morphometry of Yellowfin Tunas, 114-120
New Blennoid Fish from Hawaii, 125—127
On the Herding of Prey and the Schooling of the
Black Skipjack, Euthynnus yaito Kishinouye
(Note), 297-298

The Pacific Oceanic Fishery Investigation
(Note), 300

Idanthyrsus armatus, 8, 43
ornamentatus, 8, 43
pennatus, 43
Ikedosoma, 277

"Insects of Hawaii,” notice of, 301
International Polar Expedition (1885), 4
Iridophycus sp., 141
Ixora carolinensis, 104
var. typica, 112

"Japanese Ornithology and Mammalogy during
World War II,” notice of, 228
Jordanidia solandri, 130
Jussiaea suffruticosa var. ligustrifolia, 103, 112
Juvenile Euthynnus lineatus and Auxis thazard
from the Pacific Ocean off Central America,
262-271

Katsuwonus, 262, 263, 265, 270
pelamis, 262
keys:

to species of, Anaitides, 19
Haploscoloplos, 30
Spinther, 16
Typosyllis, 21

to genera of Echiuridae, 277
kidney worms, 65

Kilauea, explosive eruptions of, 278-292
kona weather, 81-82, 88
Koror research station, establishment of, 301
Kullenberg piston core sampler, 231
Kusaie, 97, 98, 99, 105, 106, 272-273

Ladd, lines of, 216-218, 220-222
Laelaps echidninum, 66
Laeospira borealis, 51
Lagisca lamellifera, 6
multisetosa, 4, 5, 6
multisetosa papillata, 6
rarispina, 6

Laminaria Sinclairii, 141
Lamphygma exempta, 62
Lantana camara, 62
Laonice cirrata, 8, 36
Lastrea euaensis, 214

Gretheri, n. sp., 214-215
Harveyi, 214
Margaritae, 214
ornata, 214
Eorresiana, 214
Latreillidae, 79-80
Latreillopsis hawaiiensis, 79-80
Latridopsis ciliaris, 130

Laysan Albatross Nesting on Moku Manu Islet,
off Oahu, T. H. (Note), 66

310

Leaena abranchiata, 9, 45
nuda, 9

Lecythidaceae, 111
Leguminosae, 111
Leiothrix lutea lutea, 61, 63
Leontice Leontopetaloides, 108
Leopold, Luna B. :

Diurnal Weather Patterns on Oahu and Lanai,
Hawaii, 81-95
lepadid barnacles, 80
Lepidonotus caeloris, 6
fragilis, 11
helotypus, 12
robustus, 6, 12
Leptocephalus conger, 129
Leptograpsus gonagrus, 136, 137
Leptospira icterohaemorrhagiae, 66
Lepturus repens, 100, 107
lepuel, 98, 108
Leucodore socialis, 37
lice, 66

Ligia occidentalis, 171, 187
Liliaceae, 272

Lincicome, David R., and Bayard H.
McConnaughey:

A New Nematode of the Genus Pseudophy-
saloptera from an Okinawan Shrew, 239-242
lined shore crab, biology of, 135-213
lines of Ladd, 216-218, 220-222
Lipeurus caponis, 66
Lissomyema, 277

List of Scientific Institutions in the Pacific Area,
243-261

list, of species of polychaetous annelids from
Alaska, 6—9

of stations, for collection of Alaskan annelids,
52, 54

Listriolobus, 277

liver flukes, absence on Ponape, 65
presence on Guam, 65
Lon giro strum plates sa, 129
Lumbriconereis fragilis, 4
latreilli, 29
zonata, 29

Lumbricus capitatus, 41
cirratus, 37

Lumbrinereis bicirrata, 29
Lumbrineridae, 7, 29
Lumbrineris, 4, 29
bicirrata, 7, 29
fragilis, 7
heteropoda, 7
latreilli, 7, 29
similabris, 7
zonata, 7, 29
lungworms, 65
Lyngbya spp., 14 1
Lysaretidae, 1 1
Lysippe labiata, 9
Lythraceae, 111

McConnaughey, Bayard H., and David R.
Lincicome :

A New Nematode of the Genus Pseudophy-
saloptera from an Okinawan Shrew, 239-242
Macrocystis pyrifera, 141
Macruronus novae-zelandiae, 129
Magelonidae, 1 1

PACIFIC SCIENCE, Vol. II, October, 1948

Maia squinado, 148, 149, 155, 157
Maldane lacera, 42
sarsi, 8

Maldanella robust a, 8
Maldanidae, 8, 41-43
Malvaceae, 111

Mammalogy, Japanese, in World War II, 228
maps:

of eastern part of Moen Island (in Truk Atoll),
218

of Kilauea crater and vicinity, 280
of Pacific Ocean Area, showing:

distribution of Pachygrapsus eras sipes, 139
distribution of various types of rock, 217
route of Swedish Deep-Sea Expedition research
ship "Albatross” 1947-48, insert 236-237
of patterns of distribution of eruption deposits
at Kilauea, 281
of Pingelap Atoll, 96
of southwestern Alaska, showing:

collecting areas of Alaskan King Crab
Investigation, 34-35

stations at which annelids were collected, 52
Mariana Islands, 65-66, 121-124, 214-215
new fern from, 214-215
Marianas Mallard, notes on, 121-124
Marshall Islands, 99, 105, 106
Marr, John C, and Milner B. Schaefer:
Juvenile Euthynnus lineatus and Auxis thazard
from the Pacific Ocean off Central America,
262-271

Martin, G. W.:

Additions to Galapagos Fungi, 71-77
Megalomma splendida, 9
Megninia cubitalis, 66
Melaenis loveni, 4, 6
Melinna cristata, 9
denticulata, 9
Memnoniella echinata, 75
Memorial Research Field Station in Palau, 301
Menopon gallinae, 66
Mercierella enigmatica, 205
Merluccius gayi, 129
Messerschmidia argentea, 104, 112
metamorphic-plutonic line, 216-218, 220-222
Metastrongylus elongatus, 65
"Meteor” Expedition, 231
meteorology, 81-95
Metograpsus mess or, 138
Micronesia :

parasites of domestic animals of, 65-66
Pingelap Atoll, flora and vocabulary of, 96-113
plant records from, 272-273
University of Hawaii Expedition to (1945), 97
Microserpula inf lata, 50
Mirabilis Jalapa, 102, 110
mites, 66
Moen Island
bauxite on, 223
parasites of animals, 65
schist on, 218-220
mongoose, 61
Moraceae, 109-110
Morinda citrifolia, 104, 112
Morphometric Characteristics and Relative
Growth of Yellowfin Tunas ( Neothunnus
macro pterus') from Central America, 114-120
Mucedinaceae, 75-76
Mugil cephalus, 128, 129

INDEX

311

Munia punctulata, 63
Musa paradisiaca, 98, 101, 108
varieties, 101, 108
Musaceae, 108
Mycology :

Additions to Galapagos Fungi, 71-77
mynah bird, 62-63
Mytilus edulis, 152, 180
Myxicola aesthetica, 9
conjunct a, 47
infundibulum, 9, 47
pacifica, 47

Myxomycetes, 72-73, 75

Naineris dendritica, 4, 7
longa, 4

names, vernacular, of plants on Pingelap and
adjacent islands, 100-104
Natan tia, 78-79
Neanthes brandti, 7, 25
virens, 7

Neoamp hitrite robusta, 9, 44
Neoleprea spiralis, 9, 44
Neosabellides alaskensis, 5, 9
Nephrolepis biserrata, 100, 106
Nephtyidae, 7, 24-25
Nephtys assimilis, 7
7, 24-25
ciliata, 7
hombergi, 7
malmgreni, 7
punctata, 7
rickettsi, 7
schmitti, 7

Neothunnus, 262, 263, 265, 267
albacora , 119-120
itosibi, 119-120
macropterus, 114-120,262
forma itosibi, 120
macropterus, 120
Nereidae,. 7, 25-28
Nereis agassizi, 28
alaskensis, 5
armillaris, 22
24

cyclurus, 25
filicornis, 36
heteropoda, 4, 7
neoneanthes, n. sp., 7, 26-28
neonigripes, 26
pelagica, 4, 7, 26
procera, 4, 7, 25, 28
schischidoi, 7
vexillosa, 4, 7, 26
virens, 4, 25
viridis, 20
zonata, 7, 25-26
Nerine, 4

alaskensis, new comb., 8
cirrata, 36
new combinations:

Asychis lacera, 42-43
Chitinopoma occidentalis, 50-51
Nerine alaskensis, 8
Typosyllis alternata, 2 1
elongata, 21-22
stewarti, 22

New Echiuroid Worm from the Hawaiian Islands
and a Key to the Genera of Echiuridae,
274-277

New Fern from Rota, Mariana Islands, 214-215
New Nematode of the Genus Pseudophysaloptera
from an Okinawan Shrew, 239-242
new species:

Anelassorhynchus porcellus, 274-277
Aricidia heteroseta, 33—36
Haploscoloplos alaskensis, 30-32
Lastrea Gretheri, 214-215
Nereis neoneanthes, 26—28
Petroscirtes ewaensis, 125-127
Phaeopeltosphaeria irregularis, 74
Pseudophysaloptera riukiuana, 239-242
Spinther alaskensis, 17-18
Tharyx hamatus, 37—38
Tetracrium incarnatum, 75-77
Typosyllis collaris, 23-24
News Notes, 68, 228, 301
Nicolea zosterica, 9, 44
Nicomache carinata, 42
lumbricalis, 8, 42
per sonata, 8, 40, 41-42
Ninoe simpla, 7
nodular worms, 65
Notes :

Addition to the Fish Fauna of the Hawaiian
Islands, 298

Bauxite on Truk, 223-224
Fishes Taken in Wellington Harbour, 128-130
Herding of Prey and the Schooling of the Black
Skipjack, Euthynnus yaito Kishinouye,
297-298

Holes in the Webs of Shearwaters, 224-225
Information concerning the Fulbright Act,
226-227

Interbreeding of Laysan and Black-footed
Albatrosses, 132

Laysan albatross nesting near Oahu, 66
Oceanographic program of Hawaii Marine
Laboratory, 67-68

Pacific Oceanic Fishery Investigation, 300
Parasites of domestic animals in Micronesia,
65—66

Poisoning of Bufo marinus by Flowers of
Strychnine Tree, 132
Seventh Pacific Science Congress, 225-226
U. S. Committee on the Oceanography of the
Pacific, 299

Western Regional Conference for UNESCO,
300

Notes on the Marianas Mallard, 121-124

Nothria conchylega, 7
geophiliformis, 7
irridescens, 7
Notomastus giganteus, 8
Notophyllum folio sum, 6, 18
imbricatum, 6, 18
Notoproctus pacificus, 8
Notorhynchus pectorosus, 128
Nyctaginaceae, 110

Observations on Parasites of Domestic Animals
in Micronesia (Note), 65-66
oceanographic expedition, Swedish, 68
oceanographic program of Hawaii Marine
Laboratory, 67-68

312

PACIFIC SCIENCE, Vol. II, October, 1948

Oceanography :

Swedish Deep-Sea Expedition, 231-238
U. S. Committee on the Oceanography of the
Pacific (Note), 299
Ochetostoma, 277

erythrogrammon, 275
manjuodense, 275
Ocypoda arenaria, 177, 207
Oesophagostomum dentatum , 65
Okinawa, 239

Oldenlandia albido-punctata, 273
Olfersia sp., 224
Oligocottus maculosus, 171
Omphidae, 7
Onagraceae, 112
Oniscosoma, 15
arcticum, 15
Ophelia limacina, 8
Opheliidae, 8, 39-40
Ophioglossaceae, 272
Ophioglossum pendulum, 272
Ophionectria coccicola, 77
Opuntia insularis, 71, 72, 75
Orbiniidae, 7, 30-33
Orchidaceae, 295-296

"Origin and Development of Craters,” notice of,

68

Origin of the Native Flora of Polynesia, 293-296
Ornithology :

Avian Introductions in Hawaii, 59-64
Interbreeding of Laysan and Black-footed
Albatrosses, 132

Japanese, in World War II, 228
Laysan Albatross Nesting on Moku Manu, 66
Notes on the Marianas Mallard, 121-124
Oscillatoria sp., 140
Owenia occidentalis, 8
Oweniidae, 8

Oxylipeurus angularis, 66

Pachygrapsus crassipes, biology of, 135-213
marmoratus, 176
parallelus, 138
transversus, 138

Pacific area, weather forecasting in, 81
"Pacific Discovery,” notice of, 228
"Pacific Explorer,” 114
Pacific Oceanic Fishery Investigation, 300
Pacific Science Congress, notice of, 225-226, 243,
299

Pacific War Memorial Research Field Station in
Palau, notice of, 301
Pacific yellowfin tuna, 114-120
Pagrosomus auratus, 129

Palau Islands, Memorial Research Field Station in,
301

plants from, 27 2-27 3
Paleanotus chrysolepis, 6, 15
Pallasia johnstoni, 43
Palmae, 107
Palmyridae, 1 1
Pandanaceae, 106-107, 272
Pandanus, 99
Lakatwa, 108
sp., 100, 106

Paradexiospira violaceus, 9, 51-52
Paralithodes camtschatica, 14
Paraonidae, 7, 33-36

Parapercis colias, 130
Parasahella maculata, 46
media, 45

parasites, in mynah birds, 62

of domestic animals in Micronesia, 65-66
Pareulepidae, 1 1
Pectinaria belgica, 8
Pectinariidae, 8
Peisidice asp era, 6, 14-15
Pelotretus f lav ilat us, 129
Peltogaster sp., 197
Peltorhampus novae-zeelandiae, 129
Pelvetiopsis limitata, 141
Pemphis acidula, 103, 111
Peperomia ponapensis, 102, 108-109
Perichaena cortica{is, 72-73
var. liceoides, 73
depressa, 73
vermicularis, 73

Petaloproctus tenuis borealis, 8, 42
Petrolisthes cinctipes, 185
Petroscirtes ewaensis, n. sp., 125-127
filamentosus, 127
Pettersson, Hans:

The Swedish Deep-Sea Expedition, 231-238
Phaeopeltosphaeria, 73, 74
caudata, 74
irregularis, n. sp., 74
panamensis, 74
Phegopteris ornata, 214
Phillipps, W. J.:

Fishes Taken in Wellington Harbour (Note),
128-130

Pholoe minuata, 6, 15
Phyllachora Chaetochloae, 74
Phyllanthus Niruri, 102, 111
Phyllodoce citrina, 19
foliosa, 18
groenlandica, 4, 19
mucosa, 19

Phyllodocidae, 6, 18-21
Physiculus bachus, 129
"pibal,” 90

Pilea microphylla, 102, 110
Pingelap, flora of, 96-113
Pionosyllis elongata, 21
gigantea, 6
magnifica, 1

Pipturus argenteus, 102, 110
piroplasmosis, bovine, 65
Pista cristata, 9
f as data, 9

piston core sampler, 231

plant names, in Pingelap, 96-113

Plant Records from the Caroline Islands,

Micronesia: Pacific Plant Studies 8, 272-273
Platy nereis agassizi, 7, 28
dumerilii agassizi, 28
Pleospora panamensis, 74
Pleurotus sp., 75
Plumeria acutifolia, 103, 112
Polychaetous Annelids of Alaska, 3-58
Poly cirrus sp., 9, 45
Poly dor a giardi, 8, 36-37
socialis, 8, 37
Polyeunoa tuta, 13

Polynesia, origin of native flora of, 293-296

INDEX

313

Polynoe brevisetosa, 12
cirrata, 4
fragilis, 11
insignis, 4
lordi, 11
tuta, 4, 13
vittata, 4, 1 1
Polynoidae, 6, 1 1
Polyodontidae, 6, 14
Polypodiaceae, 106
Polypodium Phymatodes, 100, 106
Poly prion oxygeneios, 129
Polystichum Torresianum, 214
Pomacanthodes imperator, 298
Ponape, 65-66, 97, 98, 99, 105, 106
Porites lobata, 79
Potamilla neglecta, 9, 46
Powers, Harold A. :

A Chronology of the Explosive Eruptions of
Kilauea, 278-292
predators, on birds, 61
Premna integrifolia, 104, 112, 273
Prionospio cirrifera, 8
malmgreni, 8, 36
proventricular roundworms, 66
Psammate, 4
apbroditoides, 6
multipapillata, 6

Pseuderantbemum atropurpureum, 104, 112
Pseudolabrus celidotus, 128, 130
Pseudopbysaloptera, 239, 240
riukiuana n. sp., 239-242
soricina, 240, 241
Pseudopotamilla anoculata, 4
brevibranchiata, 4, 46
intermedia, 9, 46
occelata, 9, 46
reniformis, 4, 46
Pterolichus obtusus, 66
Puffinus pad ficus cuneatus, 224-225
Pugettia productus, 145
Puttemansia sp., 76

Rademachia incisa, 109, 110
integra, 110
radiosonde, 90

Raillietina echinobothrida, 66
rain-gauge records, 84
rain, oceanic, 87
orographic, 87
Raja nasuta, 128

rats, in Micronesia, parasites of* 66
Rattus hawaiiensis, 61
norvegicus, 161
"rawin,” 90

Records of Rare Hawaiian Decapod Crustacea,
78-80

Report on the Flora of Pingelap Atoll, Caroline
Islands, Micronesia, and Observations on the
Vocabulary of the Native Inhabitants: Pacific
Plant Studies 7, 96-113
Reptantia, 79-80

Restudy of Reported Occurrence of Schist on
Truk, Eastern Caroline Islands, 216-222
Rhamnaceae, 273

Rhizophora mucronata, 98, 103, 112
Rhizophoraceae, 112
Rbodoglossum affine, 141

Rhombosolea leporina, 129
plebeia, 129

Rhynchocinetes rigens, 78-79
rugulosus, 79
typus, 78

Rhynchocinetidae, 78-79
Richardson, Frank:

Holes in the Webs of Shearwaters (Note),
224-225

Richmondena cardinalis, 61
Rota, new fern from, 214-215
roundworms, proventricular, 66
Rubiaceae, 112, 273

Sabella, 45-46

crassicornis, 9, 45, 46
elegans, 46
leptalea, 46
lumhricalis, 42
media, 9, 45-46
neglecta, 46
penicillus, 45
Sabellaria cementarum, 8
saxicava, 43
Sabellariidae, 8, 43
Sabellidae, 4, 45-47
Sabellides octocirrata, 9
Saccharum officinarum, 98, 100, 107
Sacculina sp., 197
St. John, Harold:

Plant Records from the Caroline Islands,
Micronesia: Pacific Plant Studies 8, 272-273
Report on the Flora of Pingelap Atoll, Caroline
Islands, Micronesia, and Observations on the
Vocabulary of the Native Inhabitants: Pacific
Plant Studies 7, 96-113
Salicornia ambigua, 142
Samytha sexcirrata, 9
Sapindaceae, 111
Sardinops neopilchardus, 128"

Scaevola frutescens, 104, 112
S calibre gma inf latum, 8, 40
Scalibregmidae, 8, 40
Schaefer, Milner B.:

Morphometric Characteristics and Relative
Growth of Yellowfin Tunas ( Neothunnus
macropterus) from Central America, 114-120
Schaefer, Milner B., and John C. Marr:
Juvenile Euthynnus lineatus and Auxis thazard
from the Pacific Ocean off Central America,
262-271

schist on Truk, reported occurrence of, 216-222
Schizobrancbia dubia, 4, 9, 47
insignis, 9, 46
nobilis, 4, 46

scientific institutions in Pacific Area, list of,
243—261

Scolecolepides arctius, 8
Scolecolepis alaskensis, 4
Scoleconectria coccicola, 77
Scoloplos acmeceps, 30
armiger, 4, 7

Sc or pis violaceus, 129

Scripps Institution of Oceanography, 67, '68 '
Scutia spicata, 71, 72, 73, 75 '

Sebacina petiolata, 74-75
Selaginellaceae, 272 •'

314

PACIFIC SCIENCE, Vol. II, October, 1948

Selaginella Kanehirae, 272
Seriola lalandi, 129
Seriolella brama, 129
Serpula Columbiana, 47
spirillum, 51
splendens, 47
triquetra, 48
vermicularis , 9, 47
zygophora, 47
Serpulidae, 4, 9, 47-52
Sesarma tetragonum, 176, 177
shearwaters, holes in the webs of, 224-225
shrews, 239-240
Sial line, 216-218, 220-222
Sid a fallax, 103, 111
Sigalionidae, 6, 15
Sigma Xi, Hawaii chapter, 68
Sitodium-altile, 109, 110
Sonneratia alba, 98, 103, 111
Sonneratiaceae, 111
Sorex /. fumens, 240
p. personatus, 240
sounding, weather, 90
Sphaerodoridae, 7
Sphaerodorum minutum, 7
Sphyraena barracuda, 298
Spinosphaera sp., 9, 45
Spinther, 15—16
key to genus, 16

Spinther alaskensis, n. sp., 6, 10, 12, 16, 17-18
articus, 15, 1 6
australiensis, 15, 16
citrina, 15, 16, 17
major, 16
miniaceus, 15, 1 6
oniscoides, 15, 16
wireni, new name, 16, 18
Spintheridae, 6, 15-18
Spio filicornis, 8, 36
filicornis pacijica, 36
minus, 3 6

Spionidae, 8, 36-37
Spirorbus, 48
abnormis, 9
asperatus, 5 1
borealis, 9, 5 1
incongruus, 9
lineatus, 9
quadrangularis, 9
rugatus, 9
semidentatus, 9
similis, 9
spirillum, 9, 5 1
tridentata, 9
variabilis, 9
violaceus, 5 1

stations, list of, for collection of Alaskan annelids,
52,53

Stemonitis pallida, 73
Stephanurus dentatus, 65
Sterna f. oahuensis, 66
Sternaspidae, 8
Sternaspis elegans, 4, 8
fossor, 4
scutata, 8

storms, convective, 83
frontal, 82-83
Strychnos nux-vomica, 132

Stylarioides papillata, 8, 40
plumosa, 8
Sula s. rubripes, 66
Suncus caeruleus, 240
coerulus, 240
c. kandianus, 240
murinus riukiuanus, 239
Swedish Deep-Sea Expedition, 231-238
note on, 68

swine, in Micronesia, parasites of, 65-66
Syllidae, 6, 21-24
•Sy/Z/V alternata, 21
armillaris, 22
cuculata, 24
elongata, 21
21

pulchra, 22
quarternaria, 7
stewarti, 22

Trfccd Leontopetaloides, 98, 101, 108
pinnatifida, 108
Taccaceae, 108
Taetsia fruticosa, 272
Tegula funebralis, 171, 179
temperature inversion, 91-93
Terebella robusta, 44
spiralis, 44
zostericola, 44
Terebellidae, 9, 43-45
Tereb ellides stroemi, 9
Terminalia Catappa, 103, 112
litoralis, 103, 112
terns, 62, 66
Tetracium, 76
Tetracrium, 75-76
Aurantia, 16
coccicola, 77

incarnatum, n. sp., 75-77
Tetrameres sp., 66
"Texas fever,” 65
Thalassema, 274, 275, 277
branchiorhynchus, 274
sabinum, 275
semoni, 275

Thalassia Hemprichii, 100, 107, 272
Tharyx gracilis, 38

hamatus, n. sp., 8, 37-38
multi jilis, 38
parvus, 38
Thelepus crispus, 9
hamatus, 9, 44—45
Thespesia populnea, 103, 111
Travisia brevis, 8
forbesii, 8
8

Thuarea involuta, 100, 107
Thyrsites atun, 128, 130
Tiliaceae, 111

Trachelochismus littoreus, 130
Trachurus nouvae-zelandiae, 129
trade winds, 81-95
Trip ter ygion varium, 130
Triumfetta procumbens, 103, 111
Trochalopterum canorum, 59
Trophonia papillata, 40
tropical deterioration, investigation on, 71

INDEX

315

Truk, 65, 97, 99, 105, 106
bauxite on, 223-224

reported occurrence of schist on, 216-222
Trypanosyllis gemmipara, 7, 24
Tuberculariaceae, 76
tuna, Allison’s, 120
Hawaiian, 119-120
morphometry of, 114-120
Pacific, 114-120
Portuguese, 119-120
Typhloscolicidae, 1 1
Typosyllis, key to species of, 21
alternata, new comb., 7, 21
ar miliar is, 7, 21, 22
collaris, n. sp., 7, 21, 23-24
elongata, new comb., 7, 21-22
pule hr a, 7, 21, 22
stewarti, new comb., 7, 21, 22

Uca, 183,207
crenulata, 207
minax, 146
pugilator, 146
pugnax, 1 46

Ulva Lactuca, 141, 142, 178
UNESCO, 243, 300
United States, Army, 71

Committee on the Oceanography of the Pacific
(Note), 299

Fish and Wildlife Service, 114, 299, 300
Fisheries Commission, 3
Geological Survey, 228
National Museum, 3
Navy, 97

Soil Conservation Service, 84
Trust Territory of the Pacific, 228
Weather Bureau, 84, 90, 91
University of California, 67
University of Hawaii, and Hawaii Marine
Laboratory, 66, 78

Micronesian Expedition (1945), 65, 97
University of Southern California, Allan Hancock
Foundation, 3
Upogebia pugettensis, 136
Urticaceae, 110

vacuum core sampler, 231

Varuna liter ata, 176

Verbenaceae, 1 12, 273

Vigna marina, 102, 111

Vittaria elongata, 100, 106

vocabulary of Pingelap, observations on, 97-113

Volcanology :

A Chronology of the Explosive Eruptions of
Kilauea, 278-292

Wagner, W. H., Jr.:

A New Fern from Rota, Mariana Islands,
214-215

weather, forecasting in Pacific area, 81
patterns, diurnal, in Hawaii, 81-95
ship, "Bird Dog,” 92-93
Wedelia biflora, 104, 112
Wentworth, C. K.:

Transfer of Hawaiian Volcano Observatory
(Note), 131

Western Regional Conference for UNESCO
(Note), 300

wet garden (in Pingelap), 98

worms, in domestic animals in Micronesia, 65-66

Yamashina, Yoshimaro:

Notes on the Marianas Mallard, 121-124
yellowfin tuna, Hawaiian, 119-120
Pacific, 114-120
Portuguese, 119-120

Zephyranthes rosea, 101, 108
Zeus faber, 128, 129

Zimmerman, Elwood C., "Insects of Hawaii,”
notice of, 301
Zoology:

Addition to the Fish Fauna of the Hawaiian
Islands (Note), 298

Biology of the Lined Shore Crab, Pachygrapsus
crassipes Randall, 135-213
Fishes Taken in Wellington Harbour (Note),
128-130

Herding of Prey and the Schooling of the Black
Skipjack, Euthynnus yaito Kishinouye (Note) ,
297-298

Interbreeding of Laysan and Black-footed
Albatrosses, 132

Juvenile Euthynnus lineatus and Auxis thazard
from the Pacific Ocean off Central America,
262-271

Morphometric Characteristics and Relative
Growth of Yellowfin Tunas ( Neothunnus
macropterus) from Central America, 114—120
New Blennoid Fish from Hawaii, 125-127
New Echiuroid Worm from the Hawaiian
Islands and a Key to the Genera of Echiuridae,
274-277

New Nematode of the Genus Pseudophysalop-
tera from an Okinawan Shrew, 239—242
Notes on the Marianas Mallard, 121-124
Poisoning of Bufo marinus by the Flowers of
the Strychnine Tree (Note), 132
Records of Rare Hawaiian Decapod Crustacea,
78-80

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