Journal of the Washington Academy of Sciences

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JOURNAL

OF THE

WASHINGTON ACADEMY
OF SCIENCES

VOLUME 30, 1940

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ACTUAL DATES OF PUBLICATION, VOLUME 30

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JOURNAL

OF THE
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Vou. 30 JANUARY 15, 1940 No. 1

ANTHROPOLOGY .—Prehistoric culture waves from Asia to America.t
DiamMonp JENNESS, National Museum of Canada, Ottawa,
Canada. (Communicated by H. B. Collins, Jr.)

The recent excavations of Collins on St. Lawrence Island and at
other places around the Bering Sea? seem to bring out one very
important point, viz., that there has been no extensive migration
across Bering Strait, unless it be of Eskimo, since the early centuries
of the Christian Era. The Eskimo culture strata in that region show
no profound disturbance such as one would expect from an invading
horde, but rather a gradual change, stimulated to some extent by
Asiatic as well as strictly American influences, but not by the intru-
sion of an alien people. Now Nordenskiold and others’? have proved
that although a few Polynesians may on one or more occasions have
reached the shores of America, there has never been any transoceanic
migration large enough to affect profoundly the physical composition
of the aborigines in the New World or the evolution of their cultures.
We can rule out likewise any immigration by way of Kamchatka and
the Aleutian Archipelago, if for no other reason than that the archi-
pelago has yielded no traces of earlier remains than those of the
Aleutian Eskimo, who undoubtedly reached their home from Amer-
ica. Bering Strait, therefore, was the only route of ingress into this
hemisphere, and the forefathers of every known division of Indians
must already have crossed this strait by the beginning of the Chris-
tian Era.

This conclusion harmonizes well with the results of linguistic
studies. Hitherto we have utterly failed to link up any American
Indian language with any language or group of languages in the Old

1 Address of the retiring president of the American Anthropological Association, de-
livered at Chicago, IIl., on December 29, 1939.

2 Cotuins, H. B. Archaeology of St. Lawrence Island, Smithsonian Misc. Coll.
Ber. Re canraae: E. Origin of the Indian civilizations in South America. Com-
parative Ethnographical Studies 9. 1931.

Dixon, Rotanp B. The long voyages of the Polynesians. Proc. Amer. Phil. Soc.
74(3): 167-175. 1934.

1

JAN 22 1940)

2 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

World. Thus Rivet’s effort to connect the Hokan dialects of Cali-
fornia with Malayo-Polynesian, and some Patagonian languages with
Australian, has apparently found few supporters. Some day we may
succeed in Joining up Eskimo with the Ural-Altaic languages and in
proving that the Athapaskan or Déné tongues of North America
are genetically related to the Sinitic tongues of eastern Asia. Such
relationships, however, even if confirmed, must be exceedingly dis-
tant, for we know how little the Greenlandic Eskimo dialects have
diverged from the north Alaska ones, despite a separation of 1,000
years, and how small is the difference, after an equally long separa-
tion, between the Navaho Indian dialect in the Southwest of the
United States and the dialects spoken in the Mackenzie River Basin.
Probably much of the linguistic diversity among the Indians and
Eskimo took place in Asia before their entry into the New World;
but the fact that no American tongue is palpably related to any
Asiatic one strongly suggests that the inhabitants of the New World,
barring the Eskimo to whom I will return later, separated off com-
pletely from those of the Old more than 2,000 years ago.

On ethnological grounds, too, there seems no reason to question
this conclusion, because the traits that are common to Asia and
America, apart from a few that are concentrated near the bridge-head
at Bering Strait, are so widely diffused in both continents that they
evidently carry a very respectable antiquity. Even the resemblances
between the Palae-Asiatics and the Indians of the northwest coast of
America hardly demand a migration in post-Christian times. If there
was such a migration it is more likely to have been from America to
Asia by way of the Aleutians and Kamchatka, as Collins has shown,*
than from Asia to America; moreover, it was a relatively insignificant
migration that introduced into northeast Asia a few cultural traits
such as labrets, certain forms of stone lamps, a certain type of
house, and perhaps some folk tales, but failed to effect any far-
reaching changes. It can hardly account for the much deeper re-
semblances, e.g., in physical type and clothing, between the Palae-
Asiatics and some of the American Indians.

For the millennia that preceded the Christian Era, the millennia
that saw the peopling and subsequent isolation of America, arch-
eology, our safest guide, has afforded us hitherto only one or two un-
certain clues. The main props for our theories have come from eth-
nology, linguistics, and physical anthropology, none of which can

4 Couuins, H. B., op. cit., pp. 375-378. Also Culture migrations and contacts in |
the Bering Sea region. Amer. Anthrop. 39(3): 375-884. 1937.

JAN. 15, 1940 JENNESS: CULTURE WAVES FROM ASIA 3

furnish more than the vaguest indications of a time sequence. In
founding our theories on these disciplines we are building on shifting
sand, and we need not be surprised if the theories topple over when
the spades of the archeologists succeed in uncovering new and
possibly unexpected remains.

The geographical position of the Athapaskan Indian tribes along
the pathway from Bering Strait toward the Equator, the late date
(about 1000 a.p.) when their advance columns reached the Southwest
of the United States, the comparatively minor changes in their dia-
lects from the Mackenzie Delta to Arizona, and the still demonstrable
affinity of their language (if we may trust Sapir) to the Sinitic tongues
of eastern Asia, all suggest that their movement into America did
not long precede the Christian Era. There are faint indications that
they may have entered this continent, or at least have advanced
south from Alaska, in two waves, one considerably earlier than the
other; for it is noticeable that the most divergent or archaic-seeming
dialects (e.g., Haida and Tsetsaut) lie on the west side of the Rockies,
where the snowshoe and the wooden toboggan so omnipresent in the
Mackenzie River Basin seem to have been unknown in pre-European
times. There are faint indications, also, that their irruption into the
Mackenzie River Basin created a considerable displacement of other
peoples who were occupying this region at the time, or were located
on its outskirts. A recent botanical investigation by Dr. Raup sug-
gests that the grasslands of the Peace River area, perhaps too the
forest zone along the northern edge of the plains, was muskeg land
or tundra no longer ago than 2,000 or 3,000 years, and that the pres-
ent-day bison and moose were preceded by herds of caribou. Presuma-
bly, the Eskimo of those days extended much farther south than
they do now and were pushed eastward and northeastward by the
invading Athapaskans. I incline to think that it was at this period
the Caribou Eskimo were restricted to their present home on the
Barren Lands west of Hudson Bay; and that a kindred group of
Eskimo fugitives occupied the coasts of the eastern Arctic, where they
developed that mysterious Dorset culture, which extended in pre-
historic times from Newfoundland to Greenland.

In addition to driving out the Eskimo, the Athapaskans may have
dislodged some Algonkian tribes, as Birket-Smith believes, and
started them on a movement that carried them into the Labrador
Peninsula. Certainly the traditions of these Montagnais and Naskapi

5 Cf. Brrxet-SmituH, Kaj. Folk-wanderings and culture drifts in northern North
America. Journ. Soc. Americanistes Paris, n.s., 22: 26-29. 1930.

4 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

Indians bring them from the west, and the strange stone culture dis-
covered by Strong near Nain,® on the Labrador coast, despite its
distinctly Algonkian flavor, seems so alien to them that we may
ascribe it tentatively perhaps to some early group that was later
absorbed or destroyed by other Algonkians, as were the now-extinct
Beothuk Indians of Newfoundland.

I am more reluctant to follow Birket-Smith in attributing to the
Salish Indians of British Columbia an early home on the Canadian
plains from which they were driven westward by the Athapaskans,
because so many traits in Salish culture point to a southern rather
than a plains’ origin, and their language, even should it prove to be
Algonkian, as Sapir suggests, differs so widely from Blackfoot, Cree,
and other members of that linguistic stock that it surely indicates a
very long separation. Nevertheless, we must not overlook the pos-
sibility that the Salish Indians may have a dual origin, that they may
be an amalgamation of two groups, one of which came originally
from the south, and the other from the Canadian plains.

Shapiro and Seltzer’ have pointed out the marked resemblance in.
physical type between the northeastern Algonkians (including the
Hurons, who absorbed many Algonkians into their ranks), the
Chipewyan Indians of Lake Athabasea, and large groups of Arctic
Eskimo, particularly those in Coronation Gulf, Smith Sound, and
Seward Peninsula. Now we know that at least the Coronation Gulf
Eskimo, like those of Hudson Bay, dwelt inland only a few centuries
ago. Hence physical anthropology also seems to indicate that the
Eskimo and Algonkians formerly lived in such close contact, some-
where in the heart of Canada, that either the Eskimo freely took
Algonkian wives or certain Algonkian groups adopted the Eskimo
culture, and, under pressure from the invading Athapaskans, moved
northward to the Arctic coast.

It is idle to speculate on the Asiatic home of the Athapaskans or
the route they followed to Bering Strait. Even archeology may never
be able to throw light on this question, because the majority of their
tools and weapons had blades and points of bone rather than of stone,
and bone disintegrates very rapidly. This, at least, is true of Canada.
In Alaska there seems to be a greater wealth of stone implements,

6 Strong, W. D. A stone culture from northern Labrador and its relation to the
Eskimo-like cultures of the northeast. Amer. Anthrop. 32: 126-144. 1934. Cf. the
review by WINTEMBERG, W. J., in Geogr. Rev., Oct., 1930, p. 673.

7 SHapiro, H. L. The Alaskan Eskimo. Anthrop. Pap. Amer. Mus. Nat. Hist.
31(pt.6). 1931; Theoriginof the Eskimo. Proc. 5th Pacific Sci. Congr. 4: 2723-2732.
1933.

SeLTzER, Cart C. The anthropometry of the western and copper Eskimos.
Human Biol. 5(3): 312-370. 19383.

JAN. 15, 1940 JENNESS: CULTURE WAVES FROM ASIA 5

and there too we find pottery as far up the Yukon River as the mouth
of the Tanana. Both these traits, however, may well be due to Eskimo
influence, since no other American Athapaskan tribe was acquainted
with pottery unless it bordered on a pottery-using people.

We may assume, then, until evidence to the contrary is forthcom-
ing, that hordes of Athapaskan-speaking peoples crossed Bering
Strait some time in the first millennium B.c. and forced their way
southward, some by way of the Mackenzie River, others down the
western slopes of the Rockies until they reached Colorado, Arizona, —
and New Mexico. It was they, perhaps, who introduced bows and
arrows and the Mongoloid strain in Pueblo I remains, shortly before
the end of the first millennium a.p.; they, too, who introduced the
snowshoe and other important elements into America, as Birket-
Smith cogently argues—though I find it difficult to ascribe to an
Athapaskan invasion all the elements he includes in the snowshoe
complex, particularly hunting territories, which I suspect are post-
European, and moccasins, cradle-boards, bark vessels, and the use of
fatty substances for tanning skins, since these elements occur also
in the extreme south of South America.

Inseparably linked with the Athapaskan invasion are the Eskimo,
whom they partly dislodged, and the Indians of the northwest
Pacific coast, the ‘“‘totem-pole’’ Indians whose origin and culture still
remain a profound mystery.

Let us consider the latter first. Smith’s excavations in the shell-
heaps along this Pacific coast have yielded rather negative results,
although the forest growth proves that some of the heaps were aban-
doned at least 500 years ago and that their lowest levels must be
several hundred years older. They revealed that there was a long-
headed strain in the population that is absent in the modern Indians
of the region; also that a few implements had a somewhat restricted
range, being absent either in the more northern heaps, or in the more
southern. By and large, however, there was little or no indication of
any earlier culture than that which was still flourishing along this
coast in the nineteenth century, although it was originally somewhat
simpler, and more nearly related, apparently, to that found inland up
the Fraser River. Even in the first millennium a.p., then, it was ap-
parently well rooted in its present home.

As far as their physical type is concerned, the modern Indians
of this area are indistinguishable, Hrdli¢ka states, from the Gilyak
and other tribes on the Amur River in Siberia; but the affinities of
the earlier, long-headed strain are uncertain.

6 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

Linguistic studies indicate that Haida and Tlinkit are greatly
modified forms of Athapaskan or Déné, that Tsimshian is a Penutian
tongue related to some languages in California, while the three south-
ern languages, Kwakiutl, Nootka, and Salish, may ultimately prove
to belong to the Algonkian linguistic stock. This helps our present
enquiry very little, except that it suggests a pressure of Athapaskan
tribes in the north strong enough to impose the language but not to
alter the physical type; and since the Haida and Tlinkit languages are
so unlike each other, and so unlike other Déné tongues, it suggests
also that they originate from the earliest Athapaskan wave and have
undergone considerable changes since, partly owing to their isolation
and partly to the influence of neighboring tongues.

The evidence of ethnology is rather confusing. Several traits, nota-
bly weaving with loom and spindle, the sib and moiety system, and
the chewing of ‘“‘tobacco”’ with lime, suggest a linkage with the south
and middle America. Others may have developed locally, e.g., plank
houses, special types of twined basketry, the caste stratification, and
the peculiar style of art. A few traits lead us northward; thus the
whaling practices of the Nootka, and the decorated lamplike vessels
of stone that were made by the Coast Salish, find their nearest if not
their only parallels among the Eskimo. Several traits, however, carry
us beyond the Eskimo right into Asia. There is slat armor, dis-
tributed almost continuously through Bering Strait (where it was
used in the first millennium a.p.) to Japan and China; woven hats,
a definitely Asiatic trait; curved fish knives, which recall East Asiatic
curved knives as well as the Eskimo wlo; a musical style that seems
altogether different from that of other American Indians, but, ac-
cording to Barbeau,* so strongly Asiatic that certain songs practically
coincide with northeastern Siberian ones, while others closely re-
semble Chinese Buddhist chants; and a social organization based on
wealth rather than on descent or prowess as elsewhere in America,
an organization that expressed itself outwardly in a potlatch system
strongly reminiscent of Indonesia and Melanesia, and in totem poles
and grave monuments that, despite profound differences, instinctively
draw our eyes to the grave posts on the Amur River. Even the unique
art of this northwest coast may offer a clue, because, as Collins?’ has
pointed out, its eye designs resemble those of the mysterious Old
Bering Sea Eskimo and also the eye designs on Chinese Shang Dy-
nasty bronzes of the second millennium B.c. Finally, we have such

8 BARBEAU, Marius. The Siberian origin of our north-western Indians. Proc. —
oth Pacific Sci. Congr. 4: 2781-2784. 1933.
7 Copmins; Eos. (Op. cits. p929S:

JAN. 15, 1940 JENNESS: CULTURE WAVES FROM ASIA a

close parallels in mythology between the northwest Indians and the
Palae-Asiatic tribes of Siberia that Jochelson went so far as to postu-
late a backward movement of tribes from America into northeastern
Asia.

We might also add in this connection two other elements not found
on the Pacific coast itself but present among the Interior Salish In-
dians of the Fraser River Basin. One is the torpedo-shaped bark
canoe, known elsewhere only from the Amur River in Siberia. The
other is the semiunderground house, distributed all round the north
Pacific basin from China to the Southwest of the United States but
in so many forms that the genetic relationship of them all is still
uncertain.

We do not know, of course, the relative ages of all these traits com-
mon to Asia and the northwest coast of America. Some may be com-
paratively modern, others very old. Some may have spread by slow
diffusion, just as tubular pipes and the tobacco Nicotiana attenuata
spread northward and reached the Fraser River only a short time be-
fore European occupation; others, again, may have been carried by
a mass migration. Two or three of them, however, notably the social
organization based on wealth, the talent for sculpture, and the music
(if this is confirmed) appear so deeply seated that almost involun-
tarily we associate them with some invading people, a people whose
original home lay somewhere, perhaps, around the Amur River. Yet
whether such an invasion ever did take place, and, if so, whether it
preceded or followed the Athapaskan invasion, must remain unsettled
until we know more of the archeology of the north Pacific coast of
America and also of northeastern Asia.

Archeology has made more progress with the Eskimo, the other
people who appear to have been influenced by the Athapaskan inva-
sion. Here I should like to pay tribute to the magnificent work of
Danish scholars, not only the brilliant galaxy still living but to the
long line of their predecessors, from Hans Egede in the eighteenth
century, Henry Rink in the nineteenth, to the last and in some re-
spects the greatest of them, the late Knud Rasmussen. Thanks largely
to Danish researches, supplemented by those of the Smithsonian In-
stitution in Washington, we know that behind the modern Eskimo
cultures there are three ancient ones, the Old Bering Sea in the west,
the Thule, which originated in the west but spread over Arctic Amer-
ica to Greenland, and the Dorset, which was restricted to the eastern
Arctic. The origins of all three cultures still await the results of further
excavations in both the east and the west. Tentatively, however, I

8 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

should advance the following hypotheses, in the hope that they may
stimulate and guide the workers of the future.

In spite of suggestions to the contrary, I still believe that the third
culture, the Dorset, is a genuine Eskimo one that has absorbed certain
Indian traits, rather than an Indian culture that has Eskimoized
itself, for the reason that we know of no Eskimo culture except the
Thule that could have influenced it, and many of its non-Indian
traits are equally non-Thule. Of special significance is the fact that
a few of these traits seem to hark back to a very early Eskimo stage,
because we have no parallels to them except in the far west. Collins?®
has already pointed out that in the Dorset, as in the ancient Aleutian
and Old Bering Sea cultures of western Alaska, chipped stone imple-
ments immeasurably outnumber those implements of polished slate
that are so characteristic of Thule and later times; also that Dorset
art represents a fairly close approach to Old Bering Sea style I.

Let us examine these early eastern and western cultures more
closely. In both the Dorset and the Old Bering Sea, but not in later
remains, we find incurved side scrapers and trapezoidal knives of
chipped chert, small slate implements with rubbed edges" that may
have been boot creasers, and rubbing stones of polished crystalline
rock, quartz in the east and basalt in the west. From an Aleutian
shell-heap, again, Hrdlicka has brought back such typical Dorset
types as small leaf-shaped blades notched on each side of the base,
knives with curving edges like miniature hunting-knives, and points
with concave bases, the only difference being that the Aleutian speci-
mens are chipped from crude basalt instead of the more amenable
chert and quartz. One Aleutian knife (?) even has three notches on
each side of the base, as we find on a few Dorset specimens also.”

Now we must not forget that in addition to these special forms
known only from the east and the far west, the Dorset culture pos-
sesses many other old Eskimo traits, such as toggle harpoon heads,
eyed needles and tubular needle cases, chipped end-scrapers, polished
stone adz heads and adzlike scrapers, barbed bone fish-spear points,
stone lamps and pots, and even ‘“‘jumping stones.’’’® It is true that
some of these objects have peculiar shapes, but there are others hardly

LOC OLLINS eh) 1 Op. cite. puouor

11 In the Dorset culture implements of both slate and chert.

2 At Kachemak Bay de Laguna found the following Dorset types: planing adz
blade, chipped asymmetric knife blade with notched tang, one type of harpoon head,
needle, and dart heads barbed symmetrically and asymmetrically on both sides.
(FREDERICA DE Laguna, Archaeology of Cook Inlet, Alaska, p. 2138, 1934.)

13 “Jumping stones”? have been discovered in Newfoundland and, more recently,
inland from Hudson Bay, in the territory of the Caribou Eskimo.

JAN. 15, 1940 JENNESS: CULTURE WAVES FROM ASIA 9

distinguishable from Eskimo types elsewhere. Furthermore, the Dor-
set people possessed in full measure the skill of other Eskimo in carv-
ing bone and ivory, for in 1937 Rowley brought back from I[glulik,
on the northwest coast of Hudson Bay, some excellent figurines of
Dorset manufacture that rival in workmanship similar figurines from
any period of Eskimo history.

In view of all this, are we not justified in suspecting, not merely
that the Dorset culture is genuinely Eskimo, but that it has stemmed
from the same parent trunk as the ancient cultures of western Alaska?

If we accept this reasoning, then we must believe that the ancestors
of the Dorset people separated from the western Eskimo before the
flowering of the Old Bering Sea culture about the beginning of the
Christian Era. This would date their entry into Canada not later
than the first millennium B.c., and possibly even in the second mil-
lennium. The closing centuries of the second millennium B.c. appear
in fact the more probable if, as I have already suggested, it was an
invasion of Athapaskan tribes that pushed the Dorset people out to
the coast of the eastern Arctic; for the Athapaskans themselves, as
we have seen, must have crossed Bering Strait from Asia before the
Christian Era.

Collins" has already pointed out certain features in which the Dor-
set Eskimo influenced later cultures in the eastern Canadian Arctic
and in Greenland. He has shown that the modern form of the eastern
harpoon head, with its bifurcated base and line holes on the upper
surface, is probably derived from a Dorset type; also that Solberg’s
Stone Age culture in northwest Greenland represents a mixture of
Dorset and Thule elements. Even the physical characteristics of the
Greenland Eskimo may have been modified by Dorset admixture,
since Greenland Eskimo skulls (outside of Smith Sound) resemble
those of the old Birnirk Eskimo in the western Arctic (who were the
immediate successors, if not the actual contemporaries of the Old
Bering Sea people), more than they do Thule skulls from the eastern
Arctic, or the skulls of Thule descendants in Smith Sound, South-
ampton Island, and Barrow.»

Let us turn our eyes again to the western Arctic, where Collins
has so brilliantly deciphered the Old Bering Sea culture and traced
its evolution, or devolution, down to modern times. The Birnirk

1 Coniins, H. B. Op. cit.,-pp. 315, 336.

ORO fi. FiscHEr-M¢.iEr, Ke Skeletal remains of the central Eskimos. Rep. 5th

Thule Exped. 3(1). 1937; and Skeletons from ancient Greenland graves. Medd.
Gr¢gnland 119(4). 1938.

10 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

phase of Barrow appears in his sequence as the immediate successor
of the Old Bering Sea, marking the beginning of the Punuk culture.
It seems rather strange, however, that its characteristic harpoon
heads are made of bone instead of the usual ivory, and that they are
rarely if ever decorated. Like the later Thule-type harpoon heads
also present on St. Lawrence Island, they rest there uneasily as if
they were intruders and did not belong to the strict order of succes-
sion, Old Bering Sea, Punuk, and modern. One wonders, too, why the
Old Bering Sea art should have undergone a slow and gradual modi-
fication on St. Lawrence Island all through Punuk times, century
after century, whereas at Barrow it vanished completely in the Bir-
nirk stage. |

Can it be that the St. Lawrence material is slightly misleading?
The Birnirk (and its first-born child the Thule) is perhaps not a direct
offspring of the Old Bering Sea, but both may be offspring of some less
advanced culture that flourished on the northeast coast of Siberia
around the mouths of the Kolyma and Indigirka Rivers. From this
region the hypothetical parent culture may have sent its offspring
eastward. One branch crossed over Bering Strait and proceeded north
along the Alaskan coast to Barrow, blazing a trail that was followed
by trading parties in later centuries; it still maintained connections
with the south, however, since Geist?* found a whetstone of Kobuk
River nephrite in Old Bering Sea remains on St. Lawrence Island.
The other branch colonized the coastline of Siberia southward from
East Cape and either then or later established a few outposts on St.
Lawrence Island. Subsequently, I suspect, the southern colony on the
Siberian shore below Bering Strait, powerfully stimulated by a yet
more southern source (ultimately, it may be, from China), revolu-
tionized its style of art, and acquired perhaps some new elements,
such as pottery and the bow drill, that appear to have been unknown
in the still earlier period I hinted at before, the period when the an-
cestors of the Dorset people first crossed Bering Strait onto Alaskan
soil and pushed into the heart of North America.

The reader may say, perhaps, that, like a fecund rabbit, I have
already delivered too large a brood of unsubstantiated theories (some
of them possibly still-born). Nevertheless I hope he will pardon me
if I add one more progeny to the overabundant litter.

In Old Bering Sea remains on St. Lawrence Island Collins found
many dog skulls that had been broken for their brains; and the dogs

146 Gist and Rainey. Archaeological excavations at Kukulik, St. Lawrence Is- -
land, Alaska. Misc. Publ. Univ. Alaska 2: 190. 1936.

JAN. 15, 1940 JENNESS: CULTURE WAVES FROM ASIA _ 11

were of a smaller breed than those of Punuk and later times. Further-
more, objects associated with dog traction, such as toggles, flat bone
sled-shoes, and whip ferrules, did not appear until the close of the
Punuk period, or roughly 200 years ago, so that evidently the St.
Lawrence islanders throughout most of their history never used dogs
to drag their heavy-runner sleds. In the eastern Arctic, however,
Mathiassen found numerous dog-harness toggles in Thule remains
dating back 1,000 years or so; and the dogs that wore this harness
doubtless belonged to that rather large and heavy breed so prevalent
in the eastern Arctic today. Moreover, the Thule Eskimo, like the
modern, seldom or never ate them, for Mathiassen remarked no
broken dog skulls in any Thule site. No dog bones have yet been
recovered from an unmixed Dorset site, nor have we found any sled
toggles or harness toggles, though there are some flat, bone sled-
runners. We read in Frobisher’s Voyages,!’ however, that in the six-
teenth century the Eskimo of Frobisher Bay, in the heart of the
old Dorset range at the eastern entrance to Hudson Strait, kept two
distinct breeds of dogs, a smaller one for eating and a larger one for
dragging the sleds.

How are we to explain these facts? It seems to me quite possible
that dog traction was unknown to the earliest Eskimo who reached
America, not only to those who remained in Alaska, but to those too
who pushed eastward into Canada and later spawned the Dorset
people, and perhaps also the modern Caribou Eskimo. Both groups
alike, however, kept a small breed of dog for hunting and for eating.
Then, about the end of the Birnirk phase at Barrow, some time in
the first millennium a.p., a larger, sturdier breed of dog was intro-
duced into Arctic America from Siberia, where dog traction, if not
earlier than reindeer traction, arose as a substitute. About the same
_ time, too, whaling originated in the same region or was introduced
from Siberia also. Under the combined impulses of dog traction and
whaling certain bands of these north Alaskan Eskimo trekked east-
ward, carrying their Thule culture with them; and in the eastern
Arctic they encountered and merged with the Dorset people. Dog
traction then became general throughout the whole of the Arctic,
though St. Lawrence Island, being in a kind of back eddy, did not
receive it until rather late. The smaller breed of dog in the west and
east became extinct, but in Frobisher Bay a mixed group of Thule and
Dorset Eskimo retained and ate it down to the sixteenth century,
when it disappeared there also.

17 The three voyages of Martin Frobisher, pp. 136-137. Hakluyt Society, London, 1876.

12 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

You will note that I have pictured the original homeland of the
Eskimo, not in America, but in northeast Siberia about the mouths
of the Kolyma and Indigirka Rivers. It would not surprise me if it
were in this region, rather than in northern Alaska, that the Birnirk
culture evolved, and even the subsequent Thule. Yet it is probable
that the homeland as thus defined is far too narrow, that it should be
extended westward. Certainly in post-Christian times there were
Eskimo-like people far to the westward, on the Yamal Peninsula, for
example, at the mouth of the Ob, where Chernezov has: excavated
three of their earth lodges,'* probably also in northeast Russia, since
the kayak and bidarka are reported from that region as late as the
sixteenth century.'® If some antecedent to the Old Bering Sea, Birnirk
and Dorset cultures could be discovered on the Arctic coast of western
Siberia, it would vastly lessen the gap, both in time and space, be-
tween the historic Eskimo cultures and those of the epipalaeolithic
peoples of northern Europe to which they bear a considerable
resemblance.

In expounding his fertile theory of two culture layers in northern
Eurasia and North America, an earlier coastal or ice-hunting layer
and a later inland or snowshoe layer, Hatt justly signaled out the
Eskimo as belated survivors of the ice-hunting stage who had adapted
themselves to life on the seashore and to the hunting of sea mammals.
If, as I have attempted to show, this adaptation occurred on the
Arctic coast of Siberia, not later than the second millennium B.c. and
probably much earlier, than we should look for its inland predecessor
in that ‘“‘Siberian pocket’’ of which Zolotarev speaks, the Barstinsky
Steppes, the upper Irtish, Ob, and Yenesei regions, and the narrow
strip of territory extending to Yakutsk. Quite probably it will prove
to be but one of many cultures, closely alike, that extended during
epipalaeolithic and early neolithic times from the Baltic to eastern
Siberia. The snowshoe may have originated on the southern fringe
of this zone, perhaps near the Lake Baikal region. At all events, the
complex to which it gave rise seems to have contributed very little
to the Eskimo cultures until relatively recent times, if we disregard
the pressure exerted by its American carriers, the invading Atha-
paskans, on the Eskimo of eastern Canada. As far as we know today,
the snowshoe itself first appears among the Eskimo in the Thule-age
mound-dwellings at Wales, Alaska, which may not be older than six
or eight centuries.

18 Cf. ZoLoTAREV, A. The ancient culture of north Asia. Amer. Anthrop. 40:
15. 1988.
19 Cf. MacRitcuiz, D. Journ. Roy. Anthrop. Inst. 42: 493-510. 1912.

JAN. 15, 1940 JENNESS: CULTURE WAVES FROM ASIA 13

I have suggested that pottery, being unknown to the Dorset
people, reached Bering Strait after some of the Eskimo had already
entered America and wandered eastward. Richthofen*® has drawn
attention to the striking resemblances, particularly in decoration,
between pottery found at Krasnojarsk and other places in Siberia,
and pottery from the Algonkian or Woodland area in eastern Canada
and the northeast United States. Following up this observation,
McKern” suggests that ‘‘a culture closely related and directly parent
to the Woodland Pattern, with pottery but without agriculture, origi-
nated in Asia, came into America and inland by way of the Yukon
and Mackenzie Valleys, had a special development in a locale center-
ing just south of Lake Superior to become what is now classified as
the Woodland pattern, and diffused from that center west, south and
east to its maximum area limits, which are not as yet well defined.”

There are serious objections to this hypothesis. In the first place
we have no evidence that any of the pottery found in this section of
North America dates back beyond the Christian Era, and in more
than one place, e.g., at Lamoka, we have discovered the remains of
an earlier people who, like the Newfoundland Beothuk, did not use
pottery. Secondly, not a single sherd of pottery is known from the
Mackenzie River Basin or the upper reaches of the Yukon; and the
pottery on the lower Yukon was probably copied from the Eskimo
of the Punuk period. We have every reason to believe that no Atha-
paskan tribe ever made pottery unless, like the Sarcee, it was in close
contact with a pottery-using people. It is true that we have found
sherds at a few sites along the southeastern fringe of the Mackenzie
Basin—at Isle 4 la Crosse, Reindeer Lake, and Cree Lake—but only
within the range of Cree penetration, and the sherds themselves
resemble Woodland pottery from eastern Canada. In view of the
vast potteryless gap separating this Woodland area from the Alaskan
Eskimo, and the comparative lateness of pottery, apparently, in the
Woodland area itself, it would seem more reasonable to believe that
the latter acquired the idea of making pottery from the pottery-
making peoples bordering them on the south than to connect them
with the Krasnojarsk and other Siberian cultures so far removed
in space, if not also in time. As long as the highway from Asia to
America—that is to say, all Alaska outside the Eskimo area, and the

20 RICHTHOFEN, B. Frur. V. Zur Frage der archieologischen Beziehungen
zwischen Nordamerika und Nordasien. Anthropos 27: 123-151. 1982.

21 McKern, W.C. An hypothesis for the Asiatic origin of the Woodland culture
pattern. Amer. Antiquity 3: 138-148. 1937. Cf. also Fewxes, V. J. Aboriginal
potsherds from Red River, Manitoba. Ibid.: 143-155.

14 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

whole of northern and western Canada—yield no sherds, we should
cling to the theory that American pottery evolved quite inde-
pendently of pottery in the Old World.

Beyond the second millennium B.c. we enter a realm of twilight,
where ethnology almost ceases to flicker and archeology provides
only one or two faint gleams to light our path. From the Gobi
Desert in Mongolia, where Nelson” discovered a preneolithic micro-
lithic culture of Azilio-Tardenoisian character, and Afontova on the
Yenesei, where Von Merhart” suspects a microlithic station, we jump
to the Amur River, whence other microliths are reported,” and from
there to Fairbanks, Alaska, where microliths unearthed on the uni-
versity campus seem to Nelson identical with his Mongolian finds.”
Can it be that these mark a culture movement from Asia into
America, and not merely a culture movement, but a movement of
peoples?

Still more recently, and from Fairbanks also, it is reported that a
stone spearhead resembling a Yuma type was found embedded in a
small mastodon.”* Now Yuma points are closely related to the Folsom
complex, the oldest yet known in America, dating from a period when
the camel, the mammoth, the mastodon, and other animals now
extinct were still comparatively abundant. Because we have hitherto
discovered no trace of this complex outside of the United States and
the Canadian prairies, certain writers have suggested that it is a purely
North American development. As Nelson?’ points out, however, we
must continue to assume that its ancestry lies in the Old World until
we find in North America a still older and more primitive industry
from which it can be derived. This Yuma-like spearhead at Fairbanks
should encourage us to search for the complex farther north and west,
right into Asia itself.

The Fairbanks discoveries are intriguing from another standpoint.
They appear to disclose one of the stations on man’s journey from the
Old to the New World, thereby enabling us to map out his route.
Many Eskimo have journeyed from Bering Strait round the Arctic
coast of Alaska to the Mackenzie River Delta, following a route that
was probably open in early postglacial times also. It may, indeed,

22 BERKEY, C. P., and Neuson, N.C. Geology and prehistoric archaeology of the
Gobi Desert. Amer. Mus. Nov., no. 222. 1926.

23 Von Meruart, Gero. The palaeolithic period in Siberia: Contributions to the
prehistory of the Yenisei region. Amer. Anthrop. 25: 45-46. 1923.

24 Sovietskaya Archaeologiya, no. 1, quoted in Antiquity, Dec. 1937, p. 497.

25 Nevson, N.C. Notes on cultural relations between Asia and America. Amer.
Antiquity 2: 267-272. 1937.

26 Amer. Antiquity 3: 188. 1937.

27 Newtson, N. C. Amer. Antiquity 2: 320. 1937.

JAN. 15, 1940 FENNELL: NEW SPECIES OF VITIS 15

have been easier at that period than today, for the climate was per-
haps milder and the Mackenzie Delta not quite so far north. The dis-
coveries at Fairbanks seem to indicate, however, that some at least
of the early migrants passed up the Yukon Valley, crossed to the
eastern side of the Rockies (probably over the low divide at the head-
waters of the Liard River), and traveled down the eastern foothills
of the mountains into the United States. Some of the later migrants
_may have traveled down the western side of the Rockies also, but in
early postglacial times this route was probably blocked by ice.

BOTANY.—Two new North American species of Vitis. J: L. FEN-
NELL, U. 8. Plant Introduction Garden, Coconut Grove, Fla.
(Communicated by 8. F. BLAKE.)

This paper includes descriptions of a new grape of the aestivalis
series from Florida and of another of the rotundifolia group from Mex-
ico. The latter is of especial interest as the first member of the section
Muscadinia found in the tropics.

Vitis gigas Fennell, sp. nov. Fig. 1
FLoRIDA BLUE GRAPE

Rami saepius glandulari-spinulosi, innovationibus dense albido- vel
ferrugineo-tomentosis; folia magna saepius 3—5-lobata inaequilateralia, lobis
acutis vel acuminatis; thyrsi fructiferi plerumque ramosi conici; baccae
nigrae glaucae 10-14 mm diam.; semen ca. 6 mm longum 4 mm latum
pallide olivaceum, raphi canalem apice seminis paene explente, rostro et
chalaza obscure aurantiacis.

Series Aestivales. Vigorous high-climbing and heavy-foliaged vine with
large leaves and strong heavy canes; internodes medium to long, mostly
beset with few to many glandular prickles on strong growths; young wood
green or reddish, mostly with some pruinose bloom near nodes, but this
often obscured by the rusty wool; canes terete, finely striate, mostly re-
taining gray tomentum and black prickles into winter, then of a dark
buckeye color; diaphragm of full-sized dry canes typically about 3 mm thick;
tendrils intermittent, strong, often trifid, brown-woolly; growing tips and
young leaves covered with a dense white or cream or even rusty-colored
feltlike tomentum which mostly becomes rusty-floccose a foot or so back
from tip. Petioles slender, medium to long, usually 6-8 cm, often retaining
loose wool, round or sometimes shallowly grooved above; leaves large, com-
monly 6—7 inches long from tip of apex to tip of basal lobe, by 5—6 inches
broad, often larger, rusty, tan or gray-tomentose beneath; generally ovate,
mostly sharp-shouldered, but often 3- or sometimes 5-lobed, and then often
unequilateral; margin irregularly, sometimes deeply sinuate-toothed with
most of the principal veins ending in a slender cusplike point; basal sinus
broadly to narrowly U-shaped; apex of leaf acute and mostly long-tapering;
shoulder points mostly acute and often long-acuminate, occasionally

* Received September 15, 1939.

16 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

Fig. 1.—Vitis gigas, X23.

JAN. 15, 1940 FENNELL: NEW SPECIES OF VITIS ; 17

rounded and then mostly short-acuminate-tipped. Inflorescences 11-19 cm
long, conical to broadly so, mostly well compounded; peduncle slender and
moderately long, often with false tendril; pedicel slender and long, giving
cluster a somewhat lax appearance, at insertion of berry green and but
little warty. Berry spherical to slightly oblate, 10-14 mm, black, with
medium to heavy bloom, mostly blue in appearance when ripe; pulp green-
ish. Seeds plump, 6 mm long by 4 mm broad, pale olive or tan in color due
to the tan-colored scurf which holds tightly to the coffee-colored testa;
chalaza round to ovate, little sunken, and like the beak, of a dull orange
color; raphe and chalaza surrounded by a tiny groove on outer face; ventral
depressions pale tan in color, shallow, curving away from raphe toward top.

Fioripa: Sebastian River, near Roseland, Brevard County, July 20,
1938, J. L. Fennell 713 (type nos. 52252-52253, National Arboretum
Herbarium).

From present knowledge Vitzs gigas apparently has a very limited range,
though in colonies where found it constitutes the typical and abundant rep-
resentative of the Aestivalian series. In a few cases it takes close observa-
tion to distinguish by the foliage alone the Florida blue grape from some of
the natural hybrids of V. shuttleworthi or of V. sempsoni (V. cinerea flori-
dana). However, the spinulose pubescence of the internodes, the blue bloom
of the shoots and fruit, the much larger leaves, the green pulp of the berries,
and the different seeds are distinctive.

In a few respects, such as shape of leaf, Vztzs gigas is suggestive of some of
the midway forms of V. aestivalis and V. argentifolia as seen in Virginia or
Kentucky. The great vigor and rampant growth of the Florida species, how-
ever, as well as the different soil and climatic associations and its botanical
peculiarities, set the Florida blue grape apart as a distinct kind.

The species is characteristically found forming heavy canopies of foliage
over trees along the dry shelly banks of brackish waterways and lagoons.
Together with the rampant Dalbergia ecastaphyllum, it often forms almost
smothering mantles of growth that extend from the water’s edge to the tops
of the highest jungle trees. I have often seen half-grown boys clamber up
over these strong tangles of vines from the water to the very topmost

branches of the trees 50 feet or so above the river.
| Vitis gigas is found mostly in sweet soil having a high shell content.

Vitis popenoei Fennell, sp. nov. Fig. 2
‘“TOTOLOCHE” GRAPE; SouTH Mexican MuSCADINE GRAPE

Cortex adhaerens lenticellatus non fissilis; medulla nodis non interrupta;
cirrhi interrupti simplices; folia serrata glabrata longiora quam latiora
longe attenuata, in petiolis pubescentibus supra sulcatis; semen reticulatum
in facie exteriore e rostro ad apicem sulcatum.

Subgenus Muscadinia. Slender vine with tight, lenticellate, nonshredding
bark; pith of canes continuous; matured first year canes olive-brown in color,
lucid, sometimes finely puberulent especially near nodes, often somewhat
swollen above nodes; tendrils intermittent, simple, glabrous except near
base; young growth somewhat angled or squared. Mature leaves small to
medium, usually cordate-ovate with slenderly acuminate apex, not broader

18 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

than long (from insertion of petiole to apex), thin, rugose, lucid, glabrous on
both sides except for some pubescence on the nerves beneath, moderately to
coarsely serrate and often finely ciliate, the sinus narrowly to broadly
V-shaped, the under surface occasionally somewhat reddish, the shoulder
points sometimes prominent; lateral veins (not including the subbasal veins)
3 or 4 pairs; petioles slender, moderately long, grooved above, finely and

fie ANY
Pie ah
ANB oars alk
i: ) Win Ste i}
HD \~

I, “Nh

I
slik

Fig. 2.— Vitis popenoet, Xz3.

closely puberulent. Fruiting panicle not seen (said to be of small to medium
size); berry not seen (said to be about one-half inch in diameter, maroon red
to purple in color; skin thick; pulp green; ripe fruit has musky aroma
[Popenoe’s notes]). Seeds mostly about 5.5-6 mm long, about 4 mm broad,
ovoid to nearly oblong, often unequally developed; beak short; groove ex-
tending from beak to top of seed on outer face; chalaza elliptical to some-
times nearly round; raphe evident only on inner face of seed and then as
narrow thread; color of seed very dark and surface when dry reticulate-
wrinkled.

JAN. 15, 1940 GOODDING: NEW SPECIES OF MUHLENBERGIA 19

Mexico: Grown at Coconut Grove, Fla., from seed collected by Wilson
Popenoe (P.I. 119001) at Puerto Mexico, Isthmus of Tehuantepec, Vera-
eruz, Mexico. Type no. 56266, National Arboretum Herbarium, collected
July 5, 1939, by J. L. Fennell (no. 1008).

In general appearance Vztzs popenoer is somewhat more suggestive of V.
rotundifolia than of V. munsoniana owing to the mostly smaller-toothed
leaf margins and the thin, rugose leaf texture. It is at once distinguished from
either of these species, however, by the relatively longer leaves and by the
slenderly prolonged leaf apex. In this respect the foliage is somewhat sug-
gestive of that found on certain forms of V. cordifolia. Average leaves of
both V. rotundifolia and V. munsoniana measure broader than long (from
insertion of petiole to apex) in distinct contrast with those of V. popenoez,
which are mostly not broader than long.

In addition to its botanical differences V. popenoer occupies a distinct
climate and a range several hundred miles farther south than that occupied
by either of the other two known species of Muscadinia. It is the first and
only muscadine grape yet known to occur naturally within the tropics. A
study of its genetic relationship as regards the other muscadine and Euvitis
species might bring to light certain characters of horticultural or botanical
interest. i

BOTANY.—Two new species of Muhlenbergia.1 CHARLOTTE O.
Goopp1NnG, Rocky Mountain Herbarium, Laramie, Wyo. (Com-
municated by JASON R. SWALLEN.)

Recent study of the genus Muhlenbergia has revealed the presence
of two new species, both of which occur in the canyons of southern
Arizona.

Muhlenbergia xerophila C. O. Goodding, sp. nov.

Perennis; culmi dense caespitosi, glabri vel scaberuli, 45-90 em alti; ligula
obtusa, 1.5-3 mm longa; laminae involutae, 15-50 em longae, 1-1.5 mm
latae; panicula pallida, 15-35 em longa; glumae 2-3 mm longae, acutae vel
_acuminatae, scabrae ad apex; lemma 4 mm longum, scabrum; callum hirsu-
tum; arista 18-25 mm longa.

Plants perennial; culms densely caespitose, glabrous to very slightly
scabrous especially below the nodes, 45-90 cm tall or sometimes more;
sheaths rounded, very slightly scaberulous; ligule membranaceous, 2-4 mm
long, obtuse; blades involute, 1-1.5 mm wide, 15-50 em long; panicle open,
15-35 cm long about 4-5 em wide, pale, the branches up to 7 em long, fili-
form, flexuous; spikelets about 4 mm long; glumes equal to slightly unequal
2-2.5 or even 3 mm long, 1-nerved, acute to acuminate, the apex scabrous
to very short pubescent; lemma about 4 mm long, scabrous above with short
tufts of hairs on the callus, 3-nerved, the midnerve extending into a slender
awn 10-35 mm long.

* Contributions from the Department of Botany and the Rocky Mountain Her-
penn of the University of Wyoming, Laramie, Wyo., no. 175. Received September

20 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

This species differs from M. setifolia in having acute to acuminate glumes,
longer blades, which are not curled below, and a longer panicle; from M.
rigida in having as a rule a longer and narrower panicle, a shorter, broadly
rounded ligule, and longer acute to acuminate glumes, scabrous at the tip;
and from M. dubia and M. metcalfer: in having a wider panicle, a shorter
ligule, which is rounded instead of somewhat lacerated above, acute glumes,
scabrous at the apex, and long-awned lemmas.

Type: L. N. Goodding M 262, collected in Sycamore Canyon, Ariz.,
September 22, 1937. The type specimen is deposited in the Soil Conservation
Service Herbarium at Tucson, Ariz., with duplicate types in the Rocky
Mountain Herbarium, Laramie, Wyo., andin the U.S. National Herbarium.

SPECIMENS REFERRED TO THIS SPECIES: Szlveus 3477, collected in Syca-
more Canyon, Ariz., September 25, 1938; Szlveus 3489, collected in Box
Canyon, Ariz., September 25, 1938; Goodding A 9493, collected in Forest
Cabin Canyon, Baboquivari Mountains, Ariz., September 24, 1938; Hardies
and Proctor M 152, collected in Sycamore Canyon, Ariz., November 24,
1936.

Muhlenbergia dubioides C. O. Goodding, sp. nov.

Perennis; culmi dense caespitosi, striati; ligula truncata, 1-2 mm longa;
laminae involutae, 15-50 em longae, 1-2 mm latae; panicula pallida, an-
gusta, 15-35 em longae, 2—4 cm lata; glumae 2-3 mm longae, acute vel
erosae, scabrae ad apex; lemma 3.5-4 mm longum, scabrum; callum hirsu-
tum; arista 3-10 mm longa.

Plants perennial; culms densely tufted, more or less striate, 50-100 em
tall; sheaths rounded; ligule firm to membranaceous, truncate, 1-2 mm
long; blades elongate, involute, 15-50 cm long, 1-2 mm wide, scabrous above
and glabrous to scaberulous below; panicle 15-35 ecm long, 2—4 em wide,
pale, the branches appressed, the lower branches 2—8 cm long, rather densely
flowered; pedicels slender, scabrous; spikelets about 4 mm long; glumes
membranaceous, 2-3 mm long, subequal, faintly nerved, acute, more or
less erose at apex, scaberulous above; lemma 3.5-4 mm long, 3-nerved,
scaberulous above and often almost to the base, the callus with tufts of
hairs 1-1.5 mm long, the midnerve extending into a rather straight, sca-
brous awn 3-10 mm long. ,

This species resembles M. dubia in the appearance of the panicle but dif-
fers in having a shorter truncate ligule, acute glumes, and a hairy callus.
From M. xerophila this species may be separated by its narrow contracted
panicle, short-awned lemmas, and more truncate ligule.

Type: W. A. Silveus 3490, collected in Box Canyon, Ariz., September 25,
1936. The type specimen is deposited in the Soil Conservation Service
Herbarium in Tucson, Ariz.

Also referred to this species is H. W. Hardies and Kd Morris M 155,
collected in Bear Canyon, Santa Catalina Mountains, Ariz., November
28, 1936.

JAN. 15, 1940 CARLGREN: ACTINIARIA FROM ALASKA 21

ZOOLOGY.—Actiniaria from Alaska and Arctic waters... OsSKaAR
CARLGREN, Lund, Sweden. (Communicated by Watpo L.
SCHMITT.)

Capt. F. E. Lewis, of Balboa, Calif., owner and commander of the
M.S. Stranger, with the assistance of W. Williams as collector, in the
course of his 1937 expedition to the coasts of Alaska and north of
Bering Sound obtained an extensive series of marine invertebrates.
Included in the material were five species of actinians, which were
submitted to me some months ago by the U. 8. National Museum for
determination. Two of the species seem to be new to science. One of
them, Epiactis polaris, n. sp., is especially interesting, as it develops
its embryos in a circular brood chamber situated in the uppermost
part of the body—a type of brood chamber not previously known
from Arctic waters but described by me from southern oceans. Be-
cause there is a close relationship between the Actiniaria in Bering
Sea and the waters of north Japan, I have added here a species of
Edwardsia dredged in the Sea of Japan in a depth of 2,300 meters,
showing that a species of this genus, most of the members of which
live in very shallow water, can occur in so great a depth. Finally, I
avail myself of the opportunity to comment briefly on some Actiniaria
described by Uchida from the coast of Japan.

Family EDWARDSIIDAE
Edwardsia arctica Carlgren Fig. 1

Locality —Sea of Japan, 41° 38.5’ N., 132° 08’ E., 2,300 m., mud and rub-
ble, temperature at the bottom +0.18°, K. Derjugin, 15.10.1932, 2 speci-
mens.

Discussion.—The specimens were small, sterile, and very badly preserved.
The body was so strongly contracted that several tentacles were squeezed
out through the aboral part of the body, the invaginated part containing a
great deal of detritus, rendering the examination of the animals very dif-
ficult. I think, however, that they may be referred to Edwardsia arctica.
There is nothing about the temperature in which the specimens were taken
to refute such a supposition, as it was a little over zero. As a rule, the nema-
thybomes seemed to contain no nematocysts. By maceration, however, I
found some 46-48 by about 4.5u in size. The nematocysts of the tentacles
were 19-24 by 2.5u, the spirocysts about 12 by 1.5-27 by 5yu. There are
probably 16 tentacles. The appearance of the retractors (fig. 1, a) and parie-
tal muscles (fig. 1, b) of the perfect mesenteries and the distribution of the
latter on the column seem to agree fairly well with those in arctica (Carlgren,
Actiniaria, The Danish Ingolf Expedition 9, 1, fig. 33-38, p. 41. 1921).

1 Received July 26, 1939.

22 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

Fig. 1.—Edwardsia arctica Carlgren: a, Transverse section of retractor; 6, transverse
section of parietal muscle of perfect mesentery.

Family HALCAMPOIDIDAE
Halcampoides purpurea (Shed. )

Locality.—Punuk Island, Bering Sea, 15 fathoms, fine sand, mud, rocks,
15.1.19387, M.S. Stranger, W. Williams, several specimens.

Peachia parasitica (Agassiz)

Locality —Nash Harbor, Nunivak Island, Alaska, rocky gravel, fine sand
mud, 20.7.1937, M.S. Stranger, W. Williams, 1 specimen.

Discussion.—The fairly strongly contracted, sterile specimen, the length
of which was 1.8 cm, the breadth 1.3 cm, had 10 tentacles and 20 mesenteries
arranged typically. The conchula was provided with three lobes, one median
and cylindrical and two more conical. The nematocysts of the column were
22-29 by 3.5-5y, often a little curved; those of the tentacles 26-36 by 3.5—
5u, numerous; those of the actinopharynx partly 27-36 by 4.5-5y, partly
14-17 by 2.5y; those of the filaments partly 27-35 by 3.5—4y, partly 36.5—
49 by 5-5.5u, microbasic p-(penicilli-like) mastigophors; the spirocysts up to
about 26 by 2.5u. The color agreed very much with that of Bicidiopsis tubt-
cola Verrill. It was pale salmon in alcohol, and the tentacles were provided
with four brown cross bands.

Remarks.—Peachia (Siphonactinia) parasitica Agassiz, Bicidiopsis tubt-
cola Verrill, Bicidiopsis arctica Verrill, and probably Peachia quinquepunc-
tata McMurray are, I think, different development stages of one and the
same species—a form that reaches a considerable size before it gives up its
parasitic mode of life and develops the eight imperfect mesenteries. The dif-
ferent appearance of the conchula is of no great importance, as in Peachia the

JAN. 15, 1940 CARLGREN: ACTINIARIA FROM ALASKA 23

number of papillae increases with the age of the specimen. Verrill (Report
Canadian Arctic Expedition, pt. G, p. 125, 1922), proposed the genus Bici-
diopsis for Siphonactinia having 20 mesenteries and a mesogloeal sphincter.
This latter statement is certainly incorrect—there was no sphincter in the
present specimen. He mentions also that Bzcidiopsis sometimes had 18 ten-
tacles. If really so, the specimens may have been very abnormal or probably
belonged to Eloactis or Haloclava.

Family BUNODACTIIDAE |
Epiactis lewisi, n. sp. Fig. 2

Description.—Body cylindrical, smooth. Fairly close to the sphincter a
circular deep invagination forming a brood chamber in the female and con-
taining very large embryos very rich in yolk. Sphincter endodermal, circum-
script. Tentacles conical, rather long, probably 24. Longitudinal muscles
of the tentacles and radial muscles of the oral disk ectodermal. Actinophar-
ynx longitudinally suleated. Two well-developed siphonoglyphs. Twelve (6-++
6) mesenteries, all well developed, perfect, and fertile. Two pairs of direc-
tives. Not more mesenteries at the base than at the margin. Retractors of all
mesenteries strong, fairly concentrated, those of the mesenteries of the sec-
ond order, however, weaker than those of the first. Parietobasilar muscles
very well developed on a strong offshoot of the mesogloea. Ova very large
and rich in yolk. Nematocysts of the column 23.5—26 by 2.5; basitrichs,
those of the tentacles 29-31 by about 3y, those of the actinopharynx partly
36-38 by about 4y, few, partly 26-30 by about 4.5y, the latter microbasic
p-mastigophors, those of the filaments partly 30-34 by 4.5u, microbasic p-
mastigophors, partly 31-38 by about 3u, partly 14-18 by 2-2.5y, the latter
probably basitrichs.

I have figured the sphincter (fig. 2, a) and two mesenteries, one of the
first order and one of the second at the level of the lowermost part of the
actinopharynx (fig. 2, c). Because I did not wish to mutilate the single speci-
men I had, I have examined closely only two pairs of the mesenteries of the
second order. As these pairs were perfect and fertile, there is no reason to
suppose that the other mesenteries of the second order should be otherwise
constituted.

The brood chamber was formed by a deep circular invagination fairly
close to the sphincter. In the bottom of this invagination each embryo lay
in a special pocket (fig. 2, b). Because of their great size (0.2 by 0.25 em in
diameter), they were rather few in number. The embryos were probably in
the gastrula stage. Owing to their poor preservation, I can not decide whether
they were ciliated.

Color unknown.
Size in preserved state: Length 2 cm, breadth 1-1.1 cm.

Locality—68° 30’ N., 169° 20’ W., about 30 fathoms, fine gray mud, 7.7.
1937, M.S. Stranger, F. ae) Lewis, 1 specimen.

Remarks.—The species is closely related to Epiactis (Pseudophellia) arc-
tica (Verrill) and FE. marsupialis Carlgren, both of which, however, had the
brood chamber in the lowest part of the body and more mesenteries at the
base.

24 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 1

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Fig. 2.— Epiactis lewisi, n. sp.: a, Transverse section of sphincter; b, longitudinal
section of uppermost part of body, showing sphincter and brood chamber with em-
bryo; c, section of one mesentery each of first and second orders at level of lowermost
part of actinopharynx.

Cribrinopsis williamsi, n. sp. Figs. 3, 4

Description.—Pedal disk broad, diminishing in breadth upward. Column
provided with verrucae in its upper part. Margin with perforated, fairly well
developed pseudosphaerules. Fossa deep. Sphincter strong, circumscript,
with a rather short main lamella. Tentacles about 48, almost as long as the
body, conical, and, at least in somewhat contracted state, almost papillous.
Longitudinal muscles of tentacles ectomesogloeal, strong, radial muscles of
oral disk mesoectodermal. Two broad siphonoglyphs with well-developed
aboral prolongations. Mesenteries more numerous than the tentacles, at the
base about 72, two pairs of directives, about 24 pairs perfect. All mesenteries
were sterile. Retractors of the mesenteries recalling those of E'piactis polaris
but weaker, parietobasilar muscles as in this species. Nematocysts of the

JAN. 15, 1940 CARLGREN: ACTINIARIA FROM ALASKA 25

column 17-19 by 2-2.5yu, those of the pseudosphaerules 17-22 by 2-2.5un,
those of the tentacles 30-38 by 2.5-3u4, common, those of the acttnopharynx
partly 34-41 by 4—5y, partly 27.5-31 by 2.5—-3y, partly 14-19 by about 2.5y,
those of the filaments 24-29 by 4.5-5u, microbasic p-mastigophors; spiro-
cysts of tentacles about 19 by 2-38 by 4u.

tat Ve cpa
Since
u ay
iy aa
r Prot

Fig. 3.—Cribrinopsis williamst, n. sp.: a, Section of uppermost part of column,
showing sphincter (s) and one perforated pseudosphaerule (p); 6, cross section of
tentacle.

I have reproduced a section of the uppermost part of the column in Fig. 3,
a, showing the sphincter (s) and one perforated pseudosphaerule (p). On the
left side there is an invagination in the endoderm and mesogloea, indicating
the border of a verruca. The pseudosphaerules do not seem to contain more

Fig. 4.—Cribrinopsis williams, n. sp.: An individual tentacle.

26 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

nematocysts than other parts of the column; it was difficult, however, to get
good maceration preparations of them. Fig.4 shows the exterior of a tentacle,
Fig. 3, b a cross section. The main lamella of the sphincter in other places was
sometimes a little stronger, the folds more numerous.

Color unknown.

Size: Breadth of the pedal disk 1.3 cm, at the margin 0.9 cm, height of
the body, 1.3 cm, length of the inner tentacles 1.2 cm, of the outer ones 0.5
cm.
Locality — Humpback Bay, Alaska, 56° 11’ N., 181° 54’ W., 15 fathoms,
near shore, 19.8.1937, M.S. Stranger, W. Williams, 1 specimen.

Remarks.—As the specimen was sterile, the placing of the species in the
genus Cribrinopsis is somewhat dubious, but there is no objection to refer-
ring the species to this genus except for the presence of pseudosphaerules,
which I did not note in my description of Cribrinopsis similis (1.c., 1921, p.
156), the only known species. But, as in the genus Bunodactis, in which
pseudosphaerules sometimes occur, it is not surprising that they may be
present in Cribrinopsis. In fact, at first I was inclined to identify the speci-
men with szmilis, but, as the nematocysts were smaller, the tentacles con-
siderably longer, thinner, and more papillose, and pseudosphaerules were
present here, I think it is better to consider the present form as a distinct
species.

Family ACTINOSTOLIDAE

Stomphia coccinea (O. F. Miiller)

Localities —Off Codbora Bay, Victoria, British Columbia, 10 fathoms,
hard bottom, 9.8.1937, M.S. Stranger, W. Williams, 2 specimens; Hump-
back Bay, Alaska, 56° 11’ N., 131° 54’ W., 15 fathoms, near shore, 19.8.1937,
M.S. Stranger, W. Williams, 1 specimen; Cleveland Passage, Alaska, 57°
33’ N., 133° 30’ W., 12 fathoms, rocky, 16.8.1937, M.S. Stranger, W. Wil-
liams, 1 specimen; Teller, Alaska, 65° 16’ N., 166° 25’ W., 5 fathoms, mud,
some rocks, 11.7.1937, M.S. Stranger, W. Williams, 1 specimen; 68° 30’ N.,
169° 20’ W., about 30 fathoms, fine gray mud, 7.7.1937, M.S. Stranger,
F. E. Lewis, 1 specimen.

Most of the specimens were small. As to the determination of the last spe-
cies I am somewhat uncertain. It is possible that it is a young Actinostola.

COMMENTS ON CERTAIN SPECIES OF JAPANESE ACTINIARIA

Uchida (Sci. Rep. Tohuko Imp. Univ. (4), Biol., 13 : 293.1938) refers Buno-
dactis stella to Anthopleura, owing to the fact that the figure of this species
given by Verrill, 1864, indicates the presence of marginal sphaerules. I have
examined many specimens of stella from different localities and find no traces
of such formations. Verrill also refers the species to Bunodactis. Unfortunate-
ly, Uchida gives no notes about the nematocysts. The marginal sphaerules
proper always are provided with atrichous nematocysts. If atrichs are ab-
sent in Uchida’s A. stella, this species is probably identical with Verrill’s
form; if the contrary, not. Uchida (l.c., p. 309) set up Epiactis ritterz, de- —

JAN. 15, 1940 HAYES & FERGUSON: MACROSTOMUM RUEBUSHI 27

scribed by Torrey from Alaska, as synonymous with Epzactis prolifera
Verrill. This is, I think, not correct. The species I called Cnidopus ritteri
(Torrey), also taken on the coast of Alaska, is very different from prolifera
and, as far as I can see, is identical with Torrey’s species; the nematocyst
batteries at the base of Cnidopus probably consist of atrichs. It is question-
able if Uchida’s Eloactis mazelii (1.c., p. 288) is identical with the European
species. Some other species described by Uchida are not referred to the right
genus. Andwakia hozawaz is probably not Andwakia, as it seems to lack
tenaculi proper. Milne-Edwardsia akkeshi (Annot. Zool. Japon. 13: 571.
1932) is certainly not Milne-Edwardsia but belongs to a new genus, if it is
not identical with Drillactis, the anatomy of which is unknown. Phellia de-
cora (Annot. Zool. Japon. 17: 623.1938) is not a Phellia but a Telmatactis (or
possibly another genus previously described, see Stephenson, The British
sea anemones, 1935, and Carlgren, Vet.-Akad. Handl. Stockholm (8) 17:
67—-68.1938). Moreover, it raises the question as to whether the synonyms
enumerated by Uchida of decora are correct.

ZOOLOG Y.—Notes on the morphology of Macrostomum ruebushi var.
schmitti.1 WAYLAND J. Hayss, Jr., University of Virginia, and
FREDERICK F. FERGuSON, College of William and Mary. (Com-
municated by Waxtpo L. ScHmMIrTT.)

In October and November 1938, while investigating a pond and its
spring source situated beside Charlotte Pike, Nashville, Tenn., one
quarter of a mile west of the intersection of Old Hickory Boulevard
with the Pike, we discovered what appears to be a new variety of a
turbellarian worm, Macrostomum ruebushi var. schmitti. In this, the
type locality, the variety, of which we obtained many specimens and
studied about 50 mature individuals, was associated with Stenos-
tomum virginianum, S. tenuicaudatum, Dalyellia rossi var. tennesseensis,
Gyratrix hermaphroditus, and unidentified species of Provortex and
Geocentrophora. The new variety is named in honor of Dr. Waldo L.
Schmitt, curator of marine invertebrates, U.S. National Museum, in
gratitude for his kind assistance in our work upon Turbellaria. The
cotypes are deposited in the U. 8. National Museum as no. 20529.

Description.2—Sides of the dorsoventrally compressed body subparallel
for the greater part of the length of the animal, without lateral indentations
in the cephalic region, with slight gradual depressions marking off the
spatulate tail; body colorless except for dark eyes and enteric inclusions;

1 Received September 6, 1939.

* The name of this variety was first published in the program of the Seventeenth
Annual Meeting of the Virginia Academy of Sciences at Danville, Va., p. 18, May 6,
1939, without description. The characters of the variety were discussed by Mr. Hayes
in the paper that he delivered at that time before the Academy. The present account,
however, constitutes the first publication of the variety.

28 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

total length 1 to 1.7 mm. Epidermis of flattened, usually pentagonal cells
bearing an even coat of cilia 10 to lly long. Rhabdites, in groups of 1 to 12,
distributed over the entire epidermis but most abundantly at the extremities;
weak Rhabditenstrassen present anteriorly; Stabchen abundant, radially
arranged about the female gonopore; rhabdites 9.8 to 14y long, 1.4 to 2.1u
wide; Stabchen smaller. With groups of sensory hairs 30 to 50yu long and
lacking basal spines distributed among the cilia laterally and posteriorly, the
Jateral hairs stouter than the posterior ones; with semirigid spines 25y long
restricted to the anterior cephalic margin. Paired cerebral ganglia joined by
a broad commissure to form a crescent-shaped ‘‘brain’’ located equidistant
between the dorsal and ventral sides of the body. Paired eyes approximately
17u in greatest diameter immediately behind and dorsal to the “brain.”
Mouth bounded by ciliated lips located ventrally 145 to 190u from anterior
end. Pharyngeal glands directed laterally and then posteriorly, often ex-
hibiting in fixed and stained material large eosinophilic vacuoles bounded
by basophilic granules. Enteron saclike, considerably lobed, ciliated, ex-
tending dorsally above female gonopore almost to union of vasa deferentia.
Excretory system of two laterodorsal main-stems extending almost from one
end of the body to the other and connected posterior to the male gonopore
by a commissure; main-stems with paired lateral branches including paired
end-stems, which, at a level 100 or 130u behind the anterior end of the
enteron, pass mesially and dorsally over the enteron and, coiling, end in
paired dorsal openings. Flame cells with flagella about 10u long. Testes
obovate, smooth, located lateroventral and only slightly posterior to an-
terior end of enteron. Vas deferens extending caudally from each testis to
posterior end of enteron and there uniting with its counterpart in entering
the vesicula seminalis. False vesicula seminalis not observed. Vesicula
seminalis muscular, contractile, thin walled when filled with sperm, thick
walled when empty. Entrance from vesicula seminalis to vesieula granu-
lorum guarded by a sphincter. Vesicula granulorum with the proximal portion
ciliated, distal portion and the stilette usually filled with granular ma-

Figs. 1-6.—Macrostomum ruebushi var. schmitti: 1, Dorsal view of gross anatomy,
detail of epidermal spines, detail of sensory hairs, detail of rhabdites, and detail of
flame cell; 2, male sex apparatus and two views of penis stilette under varying pressure
beneath the cover glass (scheme of measurement: x=base, y=opening, z=total
length); 3, normal, living sperm; 4, living sperm swollen by contact with water and
three views of the body of the same sperm in different stages of movement; 5, longi-
tudinal section through the testis, showing the condensation of the nucleus during the
maturation of the spermatid; 6, somatic metaphase chromosome plate.

ABBREVIATIONS USED

DRO rag EN er ee SO PRE RAD tater Oa SU ncen c body =.00 ss. eo hve oe eer ovary
RRR UN a ee a ee eee ee, MICS Cilia ope. Se eee ee penis stilette
CTD er ORIEL Cte tg reac Le chromatingeranules pg, ec ee eee pharyngeal glands
Acta adh Png ae ND Te MEE Soca CVO PN, 22 eee dre eee protonephridium
CURA spre nay Lae chan eee T etna sn ete RO sagt eNnteron! pps eds ea eee excretory pore
CD iu Ra cae sent Mosel tas, Meade. «20a Ns epidermis’ rhiy. 2 Cass 5 no eee rhabdite
Ges eR ns oN tn Mates Sra ee ar feeler:* “(Shiosnee eed os eee sensory hairs
OGRE sah. Bie eaten female genital atrium sn, sn’, sn’’....nucleus in stages of con-
Ui LARP tue ach at female genital pore densation to form chromatin granule
GAEL gos ters ae Oa Zee an, a oe ae flagellum’: Sp). ov keeles. 60S 2 coer eee spine
RS yn cee OR RAM ES oe ganglion of brain’ “Ct... ceed oo tail
REE AE. SNR CREME, «fy 3a mem mouth) “te. 2) ee eee oe ee testis
MEG DPE eos AR ol See male gonopore!: vdieo vas deferens
eA hee ee ete it s 8. o. cay eee o MUClOUS): GH att eh hawk omer vesicula granulorum
OG ene Eke cds Se OVIGUCT US ca 22 Aces eee vesicula seminalis

1 UE cco Cee eg is So RRM oe EER opening

29

6
>

JAN. 15, 1940 HAYES & FERGUSON: MACROSTOMUM RUEBUSHI

(See opposite page for explanation.)

10:

Figs.

30 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 1

terial organized into bundles. Penis stilette a conical tube housed in a muscu-
lar tunic; the cuticular walls thickest near the base and there hardly a
micron thick, capable of slight change of shape under pressure; the base
widened, highly crenate, the apex flexed at an angle less than 90° and with
the opening lacking a distal lip or flange, terminal on the convexity of the
curve; average measurement? of base 18.5y, of opening 14.1y, of total length
62.9u. Male gonopore approximately 65u from the posterior end of body.
Mature sperm cell highly mobile, 35 to 40u long, 3.5 to 4u wide, composed
of body and of tail provided with axial filament, body with two flagella
about 20u long extending lateroposteriorly from the anterior portion, pos-
terior portion of body with a single, oblong, in life extremely hyaline granule
about 2u long, which after proper fixation always stains with Heidenhein’s
hematoxylin, Gentian violet, and Feulgen reaction, none of which treat-
ments stain the remainder of the cell. Female genital system typical for the
genus. Ovary about one-fourth as long as the body. Egg grayish. Female
gonopore about 200u from the posterior end of the body. Chromosome num-
ber, N=3, 2N =6. Chromosomes, as fixed in Allen’s B-15 fixative, with one
large pair about 3u long and the centromere submedian, one medium-sized
pair with the centromere displaced slightly toward one end, and one small
pair with submedian centromere.

Differential diagnosis.—Penis stillette not distally enlarged, the proximal
end truncated at right angles to the shaft and crenated, the distal end ob-
liquely truncated and with terminal opening not provided with a distal lip,
the shaft with a single flexure of less than 90° near the distal end and with a
total length averaging 63u; pharyngeal glands without striking pigmenta-
tion; sensory hairs 50y or less in length; enteron extending well beyond the
female genital atrium; mature sperm 35 to 40u long and characterized by
two flagella and by a single, nuclear-staining granule located in the posterior
part of the sperm body.

Remarks.—Macrostomum ruebushi var. schmitti possesses the specific
characters of Macrostomum ruebushi Ferguson (1939) but is distinct from
the species and its other varieties and from all other species of Macrostomum
heretofore described by the characters stated above in the differential diag-
nosis as may be shown by comparing these characters with those set forth
in the key to the genus in Ferguson (1939-1940).

LITERATURE CITED

FERGUSON, FREDERICK F. The morphology and taxonomy of Macrostomum virgini-
anumn.sp. Zool. Anz. 119: 25-32, 8 figs.

———. The morphology and taxonomy of Macrostomum beaufortensis n. sp. Zool.
Anz. 120: 230-235, 6 figs. 19387.

. A monograph of the genus Macrostomum QO. Schmidt 1848. Part I. Zool.

Anz. 126: 7-20, 2 figs., 4 maps. (Subsequent partsin same journal.) 1939-40.

and Jonss, E. R., Jr. Studies on the turbellarian fauna of the Norfolk area. I.

Macrostomum ruebushi var. kepneri n. var. Zeitschr. fiir Morph. und Okol.

1939. (In press.)

and StrrewaLt, M. A. The morphology and taxonomy of Macrostomum
phillipsi n. sp. Zool. Anz. 123: 42-45, 4 figs. 1938.

GraFF, L., von. Das Tierreich. TurbellariaII. Rhabdocoelida. Pp. 47-55. 1913.

Hayrs, W. J., Jr. The morphology and taxonomy of Macrostomum ruebushi var.
schmitti n. var. Prog. Virginia Acad. Sci., May 1939, p. 18. 1939.

3 Obtained by the measurement under oil immersion of a dozen mature specimens.

JAN. 15, 1940 AVERY: EGG-LAYING HABITS OF FAIRY SHRIMP ol

ZOOLOGY.—Studies on the egg-laying habits of the fairy shrimp.
JOHN LAWRENCE AVERY, George Washington University. (Com-
municated by D. B. Young.)

During the course of life-history studies on the fairy shrimp Hubran-
chipus vernalis, certain data concerning egg-laying by this form were
collected. It is the purpose of this paper to present these data.

In the female shrimp the eggs pass anteriorly through ducts on
each side of the body until they come to lie in the brood pouch, which
is a conspicuous saclike appendage located on the ventral surface of
the female just posterior to the last pair of phyllopods. Located within
the brood pouch is a horizontal paddlelike structure, pivoted at the
center, which is in continual motion as long as eggs are present in the
pouch. Its function is apparently that of keeping the eggs well aerated.

When they first enter the brood pouch the eggs are small, light-
gray spheres. After a day or more they become almost twice as large
and assume their final light-brown color. At the time of egg-laying,
all the mature eggs in the brood pouch are expelled. Of all the shrimps
under observation, only one (shrimp 6) expelled small, gray, appar-
ently unfertilized eggs.

All the shrimps used in the experiment were of the same species,
- H. vernalis, and were collected from the same pond. Whenever possi-
ble, immature shrimps were selected to ensure the collection of all
eggs laid during the life of the individual. Because of the difficulty
of obtaining suitable food, large numbers of these young shrimps died
before reaching the egg-laying stage. Only those that laid eggs are
considered. For comparative purposes, several young mature shrimps
were used. The mature and immature were separated on the basis of
size, development of the claspers in the male and of the brood pouch
in the female, and on general appearance. That these criteria were
sufficient was later shown by the fact that within eight days after
isolation all shrimps classified as mature had laid eggs.

In practice, one male and one female shrimp of approximately
the same size were placed together in a 500-cc beaker containing
either filtered pond water or conditioned tap water. Preliminary
trials indicated that a temperature of 50°-58° F. was satisfactory for
egg-laying. All shrimps were therefore kept within this temperature
range.

The eggs, being heavier than water, sink to the bottom of the
beaker. Examinations were made at frequent intervals and any eggs

1 Received September 11, 1939.

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syouiaa sndiysupiqngy AM ONIAWT-DOW NO VLVQ—'| ATAVY,

JAN. 15, 1940 AVERY: EGG-LAYING HABITS OF FAIRY SHRIMP oo

found were removed with a pipette and counted. The results are
shown in table 1.

These results show definitely that in EL. vernalis the eggs are not
retained in the brood pouch until released by the death and disinter-
gration of the female, as has been reported for some shrimps. Neither
have there been any cases observed in which the eggs hatched while
retained within the brood pouch.? Unsuccessful attempts were made
to hatch the eggs immediately after they had been laid. However,
the writer has observed eggs laid in March hatch in November.

Six distinct depositions of eggs were laid by one female shrimp
(shrimp 11) during the period between March 18 and April 20. An
average of three batches of eggs was laid by each of 14 shrimps. The
average number of eggs laid by each shrimp was 41.7 with approxi-
mately 14 eggs a laying. In the group collected as mature shrimps,
females laid an average of 13.75 eggs a batch. In the group collected
as immature shrimps the corresponding figure was 14.1. While there
was no significant difference in the number of eggs to the batch, the
individual shrimps in the group collected as mature specimens pro-
duced more batches, resulting in a greater average number of eggs to
the individual. The greatest number of eggs laid by a female at one
time was 51 (shrimp 29). However, during the spring of 1938 a ma-
ture shrimp was collected that laid 118 eggs during a 24-hour period.
Another shrimp collected during the same season laid 165 eggs in
two batches over a 5-day period. Such variations are probably caused
by environmental factors.

2 Moraan, A. H. Field book of ponds and streams. 448 pp. 19380. Putnam’s,
New York.

04 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

ORNITHOLOGY .—The starling’s family life and behaviors. H. A.
ALLARD, Washington, D. C.

In many parts of the United States and Canada the introduced
starling (Sturnus vulgaris) has become one of our most familiar and
abundant birds. While it is generally conceded that this bird is
highly versatile and adaptive in its behavior, there is much to learn
considering the daily activities of its summer family life, and the
various manifestations of its gregarious life throughout the winter-
time.

In February, 1929, the writer erected at the end of his barn an
observation box, so constructed with a glass top in a darkened attic
that one could at all times observe intimately every behavior of the
parent birds and the young within, since the eyes of the observer
were brought within 7-8 inches of the nest. This box was furnished
with an entrance hole slightly over 12 inches in diameter, to admit
the starlings readily. This is a matter of some importance since
starlings can not pass a hole 14 inches in diameter, which is suf-
ficiently large for the entrance of bluebirds. :

The following account is based mainly upon behaviors revealed by
a pair of starlings which appropriated this observation box.

THE NEST

The starling is primarily a cavity-nesting bird. Old previously
used nesting sites are preempted usually by the same pair of starlings
throughout the season. Even throughout the wintertime the parent
birds hold their respective boxes and, when not engaged in hunting
food, spend the hours from dawn till dark fighting off meddlesome
intruders of their kind, all the while indulging in their usual native
loquacity and varied mimicry.

In February and March, or even in January, the starlings renew
their nesting activities with an attendant increase in the power and
variety of their vocal mimicry and expressions. They begin a very
active inspection of the boxes and carry out and discard much of the
old nesting material.

The new, clean observation box erected for them was quickly
appropriated, and nest building began in March. Material, including
mostly straw and grass stems, together with a few feathers, was
very leisurely carried in by both sexes.

From the outset, the site of the nest depression itself was indicated

1 Received October 26, 1939.

JAN. 15, 1940 ALLARD: STARLING’S FAMILY LIFE 35

by a circular space bare to the bottom of the box, which elsewhere
was covered about an inch deep with a mat mostly of dead grass
stems and leaves. The circular space remained bare, neatly and care-
fully surrounded by the nest foundation, which was built up around
it. As this mat became deeper, the depression itself finally received
a layer of fine grasses and some feathers, to constitute the final nest
hollow destined for the reception of the eggs.

From time to time both birds added an occasional straw or feather
to the nest even long after the young had appeared. Occasionally a
green leaf was brought in, a behavior that is indulged in by a number
of birds, more especially by the larger birds of prey, but the purpose
of this green material is not well understood.

Throughout all the procedure of nest-building the male worked
as diligently and as devotedly as the female, for the starling’s home
activities involve an unfailing cooperation between the sexes at all
times.

INCUBATION AND BROODING

Five eggs were laid, but the exact dates of laying were not deter-
mined. Following laying the male was quite as punctilious and faith-
ful in the incubation of the eggs as the female.

The carrying of food to the brooding parent by its mate was never
observed at any time. This would seem quite unnecessary since both
parents share equally in the work of incubation and brooding. When
one leaves the nest the other is usually entering and at once takes its
place.

Frequently one or the other parent has arrived at the box entrance
and expressed its desire to enter with food. The bird within, however,
maintains its claim to the nest and not even its mate can enter until
it has left.

On April 21 four of the five eggs had hatched, one delaying hatch-
ing until April 22, and this belated bird was destined by these fateful
circumstances to become the runt of the family.

While the fledglings were very young and helpless the mother star-
ling, alone, brooded them at night. The male was never seen to do
this, although during the day he was quite as solicitous of the family
and did his full share of brooding at every opportunity. While the
birds were very tiny and naked and temperatures were low, the brood-
ing impulse was much more persistently shown. Later, as the young
birds became larger and covered with feathers, they were left at night.

When brooding took place, the parent bird settled down over the

06 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 1

young, always with the same characteristic procedure. This is done
with a gentle sidewise wiggle into the final rest position. It would
appear that this allows a better adjustment of the little birds to the
parent’s body, and is practiced by both male and the female.

FEEDING THE YOUNG

As previously stated, four eggs were hatched on April 21, and one
on April 22. The young remained in the nest just 21 days. During
this period development was rapid, from tiny, naked, blind, tottering
mites of life to practically fully grown and feathered adults. This ne-
cessitated continuously changing adjustments on the part of the
parents in supplying food to the young, in the sanitation of the nest,
in brooding, ete.

Feeding began at once, both birds bringing in tiny caterpillars and
other insects usually one at a time. These were usually crushed or
macerated and very deftly and always very gently tucked into the
tiny gaping throats. It was obvious that the parents seemed to realize
that the tiny nestlings were in a helpless stage and required very
dainty and gentle handling.

As the birds increased in size, there was a noticeable increase not
only in the size of the caterpillars and spiders brought to them, but
these were gathered by beakfuls, not singly as before. This change in
the manner of feeding came about rather suddenly after the young
birds were 6-7 days old. There was also a marked change in the
manner of feeding, the very gentle profferings of the first few days
giving place to more hurried and less solicitous jabs down the throats
of the growing birds. There was of necessity a very evident speeding
up of the entire round of family attention, involving more frequent
visits with food, larger beakfuls, larger insect material, until finally,
when the birds were half grown, hard-shelled June beetles, mulberries,
and other bulky material were fed in a very impetuous manner.

Even after the young birds are fully grown and have left the nest,
the starling families do not appear to disintegrate for some time.
Although the young birds are well able to glean their own food, they
may frequently be seen following the hard-worked parents around
beseeching food. At such times it is not unusual to see the parents
feeding them in the trees or on the lawns.

NEST SANITATION

It is obvious that where there are helpless young birds some degree
of sanitation must be practiced so long as they remain in the nest.

JAN. 15, 1940 ALLARD: STARLING’S FAMILY LIFE ot

This apparently becomes one of the great problems of the starling
in the restricted room of their boxes and cavities. However, they
make every effort to keep their young in a cleanly condition, and as
the young themselves become older, they, too, become imbued with
a sanitation impulse, which expresses itself in movements to or be-
yond the rim of the nest to deposit their excrement. The extent to
which nest sanitation succeeds depends upon many factors, such as
the industry of the parents, their mutual cooperation, the size of the
nest box, the character of the food, the season of the year, tempera-
ture, etc. In very small crowded quarters, nest sanitation becomes
much more difficult than in an airy, roomy box such as the writer’s
observation box.

Fecal deposition began as soon as feeding took place, and as the
alimentary canal became filled the act almost regularly followed the
feeding reaction. Both parents carried the fecal matter away, search-
ing very circumspectly among the young for such material, picking
up even the smallest particles. On a very few occasions this excre-
ment was eaten, but this seems to be a very unusual behavior on the
part of the starlings. The parents seemed to be quickly aware of fecal
deposition and even appeared to watch a suspicious bird in the ex-
pectancy of this act, oftentimes, after each feeding.

Throughout the entire day, food was brought in usually with every
visit, and excrement carried out on leaving. Only rarely did a parent
bird enter without food or leave without fecal material. It was inter-
esting to observe that when a parent bird brooded, it was instantly
aware of the deposition of feces beneath it, probably from the feel
of this extrusion upon its feet at times, and straightway it looked
beneath and bore it away, to return quickly with food.

The starlings are early risers, and when the female had remained in
the box at night to brood the very young birds, she left them at very
early dawn even before it was light enough for her to seek food.
However, during this pre-feeding period she busied herself industri-
ously by carrying out fecal matter that had been deposited during
darkness. Likewise in the evening, when the dusk had put an end to
feeding operations, both birds continued to carry out fecal material
so long as they could see.

At first the young were too weak and helpless to do more than
defecate in the nest. Within four or five days from hatching, however,
they began to evince a tendency toward nest sanitation of their own
initiative. The young birds were making efforts to reach the side of
the nest and scrambled over one another to take this position. After

38 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

the sixth day many depositions took place at the very edge of the
nest, and very soon these were placed beyond and well away from the
nest rim. This tendency finally became an impelling mood, and near
the beginning of the third week of their existence the young had es-
tablished a defecating zone far beyond the nest and toward the open-
ing of the box. This is shown in Fig. 1. The actual factors that

Re) at ion
To

Entrance hole

Rear of bos feces

Fig. 1.—Sketch showing the deposition of the feces in a well-marked fecal zone
outside the starling’s nest at the beginning of the second week of existence by the
young nestlings. It is evident that attention to personal cleanliness and nest sanitation
has arisen in the behavior of the young birds in the course of their rapid development.
With the strength and control of the muscular system has come a conscious or sub-
conscious psychic manifestation of sanitation, working most admirably in the direction
of comfort and economy of the family association.

determined the location of this zone are not known, but there is
strong reason:to believe that orientation was associated with the
entrance hole and was dependent upon light or temperature condi-
tions or both. Whatever the correct explanation, it is obvious that
the young themselves, as soon as they were able, were governed by
a sanitation impulse, and the adoption of this fecal zone greatly
simplified the labors of the parents and resulted in better nest
sanitation. |

MAKING THE BED

From the time the young appeared, and until they had become
almost independent of the nest, the parent birds concerned them-

Jan. 15, 1940 ALLARD: STARLING’S FAMILY LIFE 39

selves very much with the making up of the nest, as it seemed. This
operation was performed with equal attentiveness by both parents.
The parent stood astride the nestlings or among them and jabbed
its beak everywhere deep down beneath their frail little bodies,
evidently poking holes into the consolidated mat, perhaps to aerate
it or to hasten drying. Whatever this procedure meant to themit
was done very frequently and thoroughly and with such vim that
one wondered how the tiny nestlings escaped being injured at times.

This shaking up of the bed was also performed by the brooding
mother during the night, when she remained with the young for the
first few days after they had appeared. Oftentimes at dawn when
she finally left the box to feed or to seek food for the young, it was
evident that the nest had been entirely remade during the night. This
was shown by an obvious deepening of the nest bed and a bringing
in of new and clean straw material and feathers from the mat within
reach.

As one or the other of the parents has taken its position over the
young birds to brood them, it has more than once been seen to reach
out and seize a straw and poke it beneath its body. At other times it
has picked up a loose feather and dropped it over its body so that it
fell loosely over the head or body. Such material probably sooner
or later falls into the nest and constitutes a new and clean lining.

REACTIONS TO EXTRANEOUS MATERIAL
INTRODUCED INTO THE BOX

During the writer’s study of the starlings’ family life, various ex-
periments were carried out. When the parents had gone, a great
variety of material was dropped into the box, including pieces of
flowers, green leaves, green shoots, bits of banana, fragments of
paper, chewed wads of paper, wood shavings, prune seeds, and such
animal material as May beetles, snails in the shell, cutworms, chrysa-
lids, and angleworms, all of which had been immobilized by crushing
to prevent their escape. Likewise small strawberries were dropped
into the box. Much of this constituted the normal material fed to
half-grown birds.

The green leaves were scrutinized critically but were usually al-
lowed to remain, together with such material as small pieces of paper
and dry wood shavings. All other material, of whatever nature, in-
cluding May beetles, strawberries, etc., even though constituting the
natural food when brought in by the parents, was unhesitatingly
discarded. Such material as paper, bits of leaf material, etc., that

40 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

could be incorporated into the nest appeared to pass as nest material
and was allowed to remain. Food substances of whatever nature,
however, appeared to fall into the category of waste material com-
parable to fecal matter and were usually very promptly discarded.

Strangely enough, if the parent bird dropped an insect morsel
while attempting to feed a young bird, it invariably picked this up
and offered.it again. On the contrary, insect material found in the box
or nest was never directly fed. In one instance a roach dropped into
the box was quickly carried out by the returning male, and when he
reappeared he offered a ground roach to the young. Whether this
was the same roach previously dropped into the box can not be
established, as these insects were sometimes brought to the young.

The placental structure of a large green pepper with the seed
attached was dropped into the box in the absence of the parents.
This object was nearly as large as the entire dimensional size of the
young. It was casually inspected by the first bird to enter the box
but was untouched. However, when accidentally knocked into the
nest by one of the parents returning later with food, it was finally
carried out of the box, but not without some effort owing to its weight
and bulkiness.

SOCIAL RELATIONSHIPS

The starlings at all times are more or less gregarious in their
relations. Even when the family burdens of the summertime break
up their great winter aggregations, they remain friendly to one
another and never entirely lose the helpful social spirit. Unlike many
other more individualistic and independent birds, they appear to
evince no definite territorial claims and forage and feed wherever
they choose, oftentimes together.

The starling is very largely a ground feeder. As it scrutinizes the
grass and ground debris it has a very characteristic habit, peculiar
also to the grackle and perhaps to various other blackbirds. As it
thrusts its beak into the grass and ground debris, it spreads the
mandibles apart with every thrust, either to enlarge the hole its
beak has made or perhaps to feel with its tongue. Whatever the
purpose of this behavior it is an invariable racial trait. The starling
never scratches in the ground debris as do the chewink and many
sparrows and never tosses the loose debris about with its beak as does
the brown thrasher.

Although the starling is preeminently a ground feeder, preferring
lawns and open fields for its foraging, it is inordinately fond of mul-

JAN. 15, 1940 ALLARD: STARLING’S FAMILY LIFE 41

berries and very readily distinguishes the ripe black fruits from the
red green ones. To this degree its color sense is highly developed.

In the wintertime when the gregarious mood has reached its height,
the flocking impulse is a marked feature of all its behaviors. It flies
from its city rendezvous to the country to feed by day, and returns
in the evening, often aggregated into enormous flocks, whose precise
and synchronized maneuverings are marvelous to behold.

In the wintertime the starlings are wont to feed in similar forma-
tions, which move very systematically over a field as the birds walk
along. However, with their keen insight those in the rear of the for-
mation realize that the new territory just ahead of the advancing
front line is likely to afford the choicest morsels. These then con-
stantly arise and fly ahead and descend to become in their turn
briefly the front line, for they too, will soon be replaced by others
flying up from the rear. Thus the foraging flock moves along
smoothly, partly by walking and partly by flight, enabling all the
members to have their chance in the scrutiny of new ground, which
otherwise would be denied those always remaining in the rear.

The starlings cling very tenaciously to the boxes and nesting sites
they have preempted and will fight almost to the death any inter-
lopers of their own kind, or even other birds. The writer once found
a pair of females locked so tightly together by beak and claw, as the
result of a combat, that both birds lay helpless in the box, and when
picked up could only be disengaged with much effort. How long they
had lain thus is not known.

In the writer’s trees the flickers fought bravely to oust starlings
from their boxes, but they were no match for their smaller and more
agile adversaries. On several occasions a flicker struck at a starling
with great force, but his blows were too slow and deliberate to reach
his more active enemy. On several occasions the starlings showed
tendencies to gang up against the larger bird and were frequently
seen to cling tenaciously to their feathers in their attacks.

The story was entirely different with the little screech owls that
nested next door to the starlings. The latter never ventured into the
owls’ quarters, although on several occasions the little owls at dusk
were seen peeping into the starlings’ homes. In spite of their hatred
of the owls, both birds carried on their economies in close proximity,
but it was plainly obvious that the starlings merely accepted the
inevitable quietly, since there was nothing else to do.

Occasionally a small hawk appeared; then the starlings became

42 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 1

yon Park, Va. 1929-30, in relation to the sunrise curve

Broken line shows the normal daily mean temperature at Washing-

starlings at L
IK,

ival of the

perature at time of arriva

, for 49 years.

Fig. 2.—Time of dawn arr

(E.S.T.) and tem
oval, IDE (Op

JAN. 15, 1940 ALLARD: STARLING’S FAMILY LIFE 43

alert and even gathered into bands and flew about ready to persecute
it with their disconcerting maneuverings.

During the summer when the female remained with the tiny young
at night, the males appeared to roost nearby in pine trees. Later when
neither parent remained with the young birds, the males and females
appeared to repair to their respective roosts somewhere near at hand.
In autumn, when the flocking urge is at its height, both parents,
apparently still mates throughout the winter season, fly to and from
the city where they spend their nights.

The time of leaving the nesting box at night and of the arrival at
dawn depends upon the factor of light intensity, and these curves
follow closely the curves for the seasonal time of sunrise and sunset
(Figs. 2-4). During the winter season, however, those light intensities
that send the birds to roost are far higher than those that drive
them to roost in summertime, as if they were very loath to leave their
young. It is highly probable here, however, that we have marked
seasonal differences in physiological sensitivity of the eye to light,
owing to the very different hormonal organization expressing itself
at the two seasons. |

DISCUSSION

We have followed some of the more striking behaviors of a family
of starlings from the time of nest building to the maturity of the
young birds and the final abandonment of the nest by these exactly
21-22 days from the hatching of the eggs. It is evident that there are
many outstanding behaviors in the starlings’ life history concerned
with nesting, incubation, feeding, and sanitation that are of much
interest.

In the first place, a remarkably close cooperation between the male
and the female obtains in all the essential activities connected with
the preparation for and the raising of the family.

The male aids in nest building, incubates the eggs, feeds the young,
and attends to the cleanliness of the nest with as much interest and
punctiliousness as does the female. There appears to be one phase of
duty, however, that he does not normally assume. The female, alone,
appears to remain with the tiny, helpless young for the first few nights
of their existence.

The writer has heard it stated that starlings are filthy birds, but
this can not apply indiscriminately to all starling parents. It is prob-
ably nearer the truth to say that starlings do their best to maintain
clean nests for their families.

44 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

S40
AM Ps BX,
‘ ws a S.

Mean te m|perature fh /
iva 1929-193@Y

“April “May june

Fig. 3.—Mean time of arrival of the starlings at Lyon Park, Va., derived from the
data of 1929 and 1930. The two-year temperature mean at the time of arrival is also
shown. The correlation between sunrise and the two-year mean for the first arrival for
the years 1929-30 as obtained was 0.962 +0.009. The correlation between the two-year
mean of first arrival and the two-year temperature mean was found to be —0.482 +10.

Starlings: departure a
: nQSs :
;

y evevl 83

Lee
S

Fig. 4.—Departure of the starlings from trees of the writer at Lyon Park, Va.,
at sundown, 1930, in relation to sunset (E.S.T.).

JAN. 15, 1940 ALLARD: STARLING’S FAMILY LIFE 45

Should one or the other parent die, there is every reason to believe
that the remaining parent would assume the care of the family to the
best of its abilities. In such an event it is probable that the usual
degree of cleanliness of the nest could not be so successfully main-
tained as when two conscientious mates were working industriously
toward the same end. |

Nest sanitation is a real necessity for the health and comfort of the
young birds, and many factors must modify the degree of success
attained, one of no small importance being the roominess of the cavity
holding the nest.

It is obvious that the young themselves very soon evince tendencies
toward cleanly habits, which has finally expressed itself in the adop-
tion of a definite fecal zone entirely without the nest. We have, here,
the foreshadowing of the localized or latrine concept, which some
animals have evolved to a high degree.

Where sanitary conditions have not been maintained, for one
reason or another, conditions may become so filthy as to cause the
death of the young birds. In one instance a young bird that had left
its nest nearby was scarcely able to use its legs from the enormous
thickness of the excrement dried and accumulated upon them. It
required much effort and washing to free this bird of its filth, and it
would probably have died had it remained in its befouled condition.
It is not known what causes were responsible for this.

It has not been determined whether unattached birds—one may

call them widows, spinsters, or bachelors—ever enter into the care of
a Starling family, where one or both parents die. It seems probable
that this may occur, however, for on several occasions when the
legimitate parents of the family under observation were away, strange
starlings have occasionally appeared at the box entrance to peep in
curiously. These are quickly driven away when the parents arrive.
_ One of the most striking features of starling behavior is the rigid
observance of carrying out all introduced material, food or otherwise,
that has been dropped into the box. It would appear that all extrane-
ous matter not brought in by the parents themselves, even though
it may be acceptable food, is treated as if it belonged in the category
of excrementitious material.

Some biologists would explain all the niceties of adjustment and
accomplishment we have observed as an example of a train of mech-
anized integrated behaviors from which there was no escape. The
writer feels that we can not talk too dogmatically on this point. At
times there seems to be some element of experienced judgment in

46 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 1

operation as much as anything else. It is obvious that rarely has there
been any useless, accidental behavior shown, all behaviors operating
smoothly, rapidly and progressively to meet in a timely manner, all
the needs and adjustments demanded by a rapidly changing develop-
mental cycle.

The writer does not feel qualified at the present time to say what
moods in these behaviors are consciously reasoned and what are
purely mechanical, to be dogmatically referred to reflexes, tropisms,
or whatnot. Even we, in our highly civilized moods, at times show
many instances of mechanized, stereotyped behaviors that have no
longer logical meaning for the situation, as when we cling to archaic
procedures simply because they are sanctioned by custom or habit or
had become legal usage long ago.

A study of the starlings’ behavior and family life indicates a very
high order of bird intelligence throughout and a close attention to all
phases of the family welfare. It is evident that the intimate under-
standing and cooperation of the two parents in all the stages of
preparation for and the care of the family have helped to make the
starlings highly adaptive and successful birds wherever conditions
are favorable for their survival.

PROCEEDINGS OF THE ACADEMY AND
AFFILIATED SOCIETIES

THE ACADEMY

COMMITTEE ON CATALOGING SCIENTIFIC SOCIETIES OF WASHINGTON
AND DEFINING QUALIFICATIONS FOR AFFILIATION

As its meeting on November 10, 1939, the Board of Managers received
and accepted a report from a committee, consisting of C. P. CLausEn,
F. M. Deranporr, W. D. Lampert, J. E. McMurtrey, Jr., and W. T.
ScHALLER (chairman), dealing with the cataloging of the scientific societies
of Washington and the definition of the qualifications of a society for affila-
tion with the Academy.

To be eligible for affiliation the given scientific society, association, or club
must be concerned with natural science, with the social, economic, or his-
torical sciences, or with any phase of engineering; must hold its meetings
within an area that includes the District of Columbia and the territory lying
within 25 miles of it; must have a membership of whom the majority reside
within the area indicated; must have a regularly constituted organization
with elected officers; must concern itself primarily with the search for facts
and truths rather than with the popularization or commercial exploitation
of them; and must have at least an effective nucleus of members actively
engaged i in pure or applied scientific research who control the policies of the
society.

The list of scientific societies, associations, and clubs reported by the pres-
ent committee, which completed work begun by an earlier committee of the

JAN. 15, 1940 PROCEEDINGS: THE ACADEMY A7

Academy, includes (1) regularly organized bodies that hold their meetings
within the District of Columbia or the territory lying within 25 miles of it,
(2) local sections of national societies and associations, (3) national societies
that have their headquarters in the indicated area, and (4) scientific organ-
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ANTHROPOLOGY. —Prehistorie culture waves from £
‘s UIAMOND JENNESS (08 00. 5 ue

Botany.—Two new North A American species of

thay,

Botany.—Iwo new species: of M ublenbergia. :
DING. pleas oui dati Cn yee One nee ae

Fos our. —Studies on the ae habits of the fairy a
LAWRENCE AVERY. 2,00: 13 i st

ORNITHOLOGY. —The starling’ 8 tagaly life and behaviors
| 7.

Procerpines: Tue ACADEMY..................... : 4

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JOURNAL

OF THE
WASHINGTON ACADEMY OF SCIENCES
VoL. 30 Frsruary 15, 1940 No. 2

BOTAN Y.—Naming molds.1. CHARLES THom, U.S. Bureau of Plant
Industry. —

I owe an acknowledgment to this Society for rehabilitation as a
botanist. The office I am laying down tonight was my first elective
position in a botanical society. I gave up a graduate assistantship un-
der Atkinson at Cornell to turn dairyman in 1904, and that spelled
heresy to some. In that period, I once joined the crowd outside the
dining room door at the Annual Dinner for all Botanists. Some one
presented me to one of the elect who looked at me rather sharply and
said, “‘I don’t remember any Thom as a botanist.’’ I replied, ‘All
right, call me a dairyman.” He answered, ‘“‘Oh, then I do know you!”’
I never found out who he was and have always wished I had made
sure of his name. I should like to meet him right now.

When I left Cornell, I was assigned to the mycological phases of
producing certain varieties of cheese already recognized as ripened by
molds. My knowledge of molds was vague—I was superficially ac-
quainted with a few Mucorini, and with one or two bright-colored
Aspergilli; I was vaguely conscious of the general appearance of Penz-
cillum and a few more of the common genera. I knew nothing at all of
the technological task that I had acquired. Professor Atkinson had
written the recommendation. He was frank about it; he assured the
appointing power that he knew nothing about the project, neither did
Thom, but that he had more work already than he could do, whereas
Thom needed the job and had brains enough to fill it. No superlatives
appeared (I have read the letter). I thanked him and reported for
duty. You will readily understand that you are attending a kind of
confessional or experience meeting where the confessor has spent
some 35 years working with molds.

AN INDUSTRIAL MYCOLOGIST

Thus I became an industrial mycologist. I entered a field in which
existing mycological literature was mostly useless and in which the be-

1 Address of the retiring president of the Botanical Society of Washington, Decem-
ber 5, 1939. Received December 6, 1939.

49

50 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

ginner was left to feel his way among materials, factory processes,
ripening conditions, and biochemical aims, all unknown to him and
only vaguely known to his fellow workers who were dairymen, bac-
teriologists, and chemists. In some groups of botanists, my status as
apostate was quickly evident. It was several years before I dared to
face a botanical club, point to a distinguished ecologist in the room,
and say, “I am an ecologist—just as much as my friend over there.
The only difference is that I make the environment for my organism
while he goes out and hunts for his.’’ Academically, then, an indus-
trial mycologist must be an experimental ecologist, and the user of
ecological studies quickly learns that sound taxonomy is the essential
background of ecology. Failure in the correctness of identification of
the components of a plant formation in the field destroys the value of
a report.

Nevertheless, as a beginner I was about in the fix of the rookie
cavalryman who had never learned to ride but reported to the top-
sergeant for training in horsemanship. That hard-boiled individual
blurted out, ‘‘What? Never been on a horse before! Fine! Here is a
horse that’s never been ridden. You two may begin together.”’

Professor Atkinson was right—none of us had more than the va-
guest idea of the task before us. When we went to the literature it
paralleled closely a fellow-worker’s characterization of the German
literature about sauerkraut—“‘it was very extensive and not worth
reading.’’ Many cheese-ripening practices were described in countries
of origin as ‘‘rule-of-thumb”’ procedures in which climatic or other fac-
tors supplied conditions often not defined even in the worker’s
mind, but actually necessary to success. However inadequate the
available technological descriptions, the mycology was worse.

The molds present in the cheese industry were not difficult to iso-
late. Rigid preliminary survey did not leave many doubts as to which
were significant. Verification involved technical problems that re-
quired years of experiment after the organisms were recognized. But
when I started to find out what was already known about those molds
I was in trouble.

My connection with the taxonomy of saprophytic hyphomycetes
began at that point and has since led me in many directions. Some of
this experience will be discussed here. If at any point I may seem to
generalize, please remember that the title of this paper is just ““Nam-
ing Molds’—not ‘‘Systematic Mycology’’—however widely I might
be tempted to apply my ideas.

Why give Latin names to those wretched little molds? I can not

Frs. 15, 1940 THOM: NAMING MOLDS ol

answer, fundamentally, but I can make the trite observation that the
human mind works that way. The objects one meets get named. If
we are to understand each other then, we must be able to find out
what the names used by our fellows mean. Thus far the reasoning is
purely practical. For illustration, I once ran across a doctor of philoso-
phy to whom the word butter meant apple butter; if you meant the
stuff made from cow’s milk, it was necessary to call it cow’s butter to
make him understand. And he took a job in the dairy! A name then,
to be useful, must be an accepted designation for a very definite
thing.

Mold information, whatever its value, is indexed under the Latin
names of the molds themselves. And sometimes it is not very de-
pendable. For illustration, one very up-to-date practical and modern
journal in a perfectly proper technical article, published a new spe-
cles name and description a few years ago for a mold so well known
among culture men that nobody but the author and the editorial
committee could possibly have failed to have met it before. Since it
was new to that writer, he had burst into print with his one and only
new species. Judged by the tables of contents, that journal puts no
limitation upon the rotten practice of the “discovery” of an organism
new to the “one-project worker’ who thereupon prepares a descrip-
tion based upon his own lack of contact with the literature of the
group and sends it forth to plague all subsequent students with the
addition of another synonym, or, upon the equally obnoxious practice
of collecting all the miscellaneous organisms occurring in connection
with a special problem, labeling as new species all those that the
sender does not know and sending them to a specialist to identify,
always reserving the right to describe any organism that the specialist
verifies as new. Such men, not being mycologists, escape the ban.

Let us get back to our cheese molds. The technological writers had
copied Latin binomials from the books, with confidence, and added
the describer’s name in each case. Penicillium glawcum Link, P. candi-
dum Link, P. album Preuss, and several other molds were listed with-
out question marks. When I searched the cheese literature to find out
how they had sorted out names and fungi, I was driven to believe they
did it by Fisher’s method—‘‘at random.”

Then I tried the mycological literature to see what the names had
originally meant. I worked paper by paper back to Preuss (1851),
then back to Link (1809). I went on back to Micheli (1729). Many be-
lieve that Micheli intended Penzcillia by one of his figures, but it is
too doubtful to trouble us now.

52 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

PENICILLIUM

There is no question about Link’s generic idea of Penicillium, but
there is no evidence as to which actual species he had when he de- |
scribed either P. glaucum or P. candidum. One man’s guess is as good
as another’s—or as Link’s (yes, or Brefeld’s), for that matter. Link
did not know his P. glawcum from any one of 100 green molds. Fifteen
years later he put the whole green lot together and called them all P.
glaucum, which was designated ‘“‘the common green mold” in the
fourth edition of the Species plantarum (1824). And the idea crops up
yet, after more than 100 years!

Occasionally some one raised a doubt about a universally distrib-
uted green mold that grows upon and in everything, but the name
was convenient; it satisfied the pedantic requirement for a Latin bi-
nomial to be applied to material that people were not willing to study.
All local fungous floras report it. The popular writers accepted P.
glaucum as the green mold ; chemists took it up and tested “its” activ-
ity against every kind of substratum and reagent. With probably a
hundred green species to pick from, at random, each was able to ex-
pand the range of biochemical activity reported. Naturally with dif-
ferent agents, contradictions crept in and raised controversy between
individuals, but the popularity of P. glaucum was not abated—the
mistake was always charged against the worker. Penicillium glaucum
was sacred to the shades of Link and Brefeld.

A LIVE PROBLEM

I might go on and tell more of the story of nomenclature in Peni-
ciluum and Aspergillus, but all that has been published—and the in-
dexes are good. Suppose I shift to a problem of nomenclature that 1s
worrying a whole group of men today, and hang a general discussion
around it.

These men work with human mycotic diseases; they have had ex-
perience in the great laboratories of the world; they have excellent
instruments and refined technique; they have access to literature.
From a series of rather horrible lesions on perhaps 60 patients living
in widely separated places, they have isolated a number of strains of
mold with certain characters in common and differences that offer a
chance for individual judgment in classification. Between 1915 and
1939 these cultures have been assigned to about a dozen genera even
though it is doubtful if there are really more than three species. These
men disagree among themselves. I have been asked several times to
express an opinion as to which is the proper name, but I hardly expect

Fes. 15, 1940 THOM: NAMING MOLDS 53

any of them to accept my conclusion. The real question, however, is:
Why is there so much disagreement when the descriptive data are not
only readily established but in fact generally conceded?

A sketch of the nomenclatorial situation is necessary.

In 1915, a medical worker in Boston described a skin disease but without
very definitely naming it. From the lesions studied he isolated a fungus that

he decided belonged to an undescribed genus and species: Phialophora ver-
rucosa Medlar.

In 1920, A. Pedroso and J. M. Gomes, working in Sao Paulo, isolated a
similar organism from cases of Chromoblastomycosis. They accepted the
same name, P. verrucosa, which has since been commonly cited as originated
by Thaxter.

In 1921, Brumpt decided that the SAo0 Paulo fungus was not Medlar’s
organism but another species he called Trichosporium pedrosot Brumpt.
(Brumpt worked for a period in Brazil about that time.)

In 1922, Brumpt, having restudied the old literature, resurrected Bonor-
den’s generic name, Hormodendrum, and changed the name to H. pedrosot.

In 1922, Terra, Torres, da Fonseca, and Areo de Leao, in Rio de Janeiro,
called the same organism Acrotheca pedrosot (Brumpt) T. T. de F. and L.

In 1928, Ota distributed material under the name T'richosporium pedro-
stanum but later decided not to publish that species name.

In 1929, Langeron (Ota and Langeron collaborated about that time) again
assigned Ota’s mold to T'richosporium pedrosot (Brumpt, 1921).

1930, da Fonseca and Areo de Leao again published the name Acrotheca
pedrosot.

In 1935, Dodge resurrected the name Gomphinaria from Preuss’s 1851
paper and moved the species to that genus.

In 1936, Negroni proposed another new name, Fonsecaea, calling the
fungus F’. pedrosot (Brumpt) Negroni.

In 1937, Moore and Almeida, after collecting and comparing strains,
added three more generic names for the variations encountered, basing the
usage upon the presence and combinations of spore-bearing structures.
These names are Botrytoides, Hormodendroides, and Phialoconidiophora.

In 1939, L. Briceno-Irragorry proposed another generic name, Carrionia,
with C. pedroso: (Brumpt) as its type species, arguing that this genus should
include the organism of Chromoblastomycosis in South America.

CADOPHORA

In the field of forest pathology, Lagerberg, Lundberg, and Melin
(1928) found species with the sterigmatic cups, which characterize the
genus Phialophora, upon woody materials both in America and Swe-
den. They proposed the generic name Cadophora for those forms with-
out recognizing their essential identity with Phialophora. More re-
cently morphological and serological comparison of materials from
human and forest sources in culture (Conant, Martin) supports the

54 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

identity of these genera; hence Cadophora, 1928, gives way to Phialo-
phora, 1915. From the epidemiological point of view, however, Em-
mons (personal communication) finds grounds for belief that human
infections find their inoculation in spores or mycelium from the plant-
inhabiting fungi rather than from those upon fellow humans. As far
as cases have been studied, man-to-man communication seems to
have been excluded; hence he holds that we must search in the field
among fungi growing upon vegetation for the strains responsible for
cases of Chromoblastomycosis in man.

If further field study proves that these organisms are members of
species regularly found upon decomposing plant remains, occasional
infection of an individual human by spores or mycelia from such plant
material does not warrant the establishment of either a genus or a spe-’
cies for that organism as a parasite. Many of these strains have grown
well for me on sterilized plant material. This supports the view that
search for them by culture from field samples offers a hope for solving
some of these problems. Miscellaneous observation of great numbers
of colonies of this dematiaceous series in connection with soil and food
microbiology shows quite general growth at 37° to 38° C.—a condi-
tion usually regarded as a prerequisite to parasitism of warm-blooded
animals.

HORMODENDRUM OR CLADOSPORIUM

The identification of these fungi from human lesions as congeneric
at least with saprophytes or parasites of plant material turns our
quest for nomenclature back to such older names as Hormodendrum.
Bonorden (1851) distinguished his genus Hormodendrum from Clado-
sporium of Link (1816) by only one tangible character. Link had re-
ported the spores of Cladosporium as 2-celled. Bonorden described the
spores of his genus as l1-celled and transferred to it four species de-
scribed by Corda as Penicillia but without personally seeing any of
them, then added some ‘‘Penicillia’’ described by Fresenius, making
a few disparaging remarks about Fresenius (1851). Two years later
(1853) he clarified his ideas of Hormodendrum by actually describing
one from fresh material (H. atrum). He left little doubt as to the gen-
eral morphology of his mold.

Bonorden in his discussion clearly admitted that his material may,
at least in part, have already borne the name Cladospor1um. Among
mycologists familiar both with specimens from the field and with the
molds in culture, the 1-celled or 2-celled condition of the conidia is
found utterly unreliable; hence the identity of Hormodendrum with
Cladosporium has been quite generally conceded for 50 years at least.

Fes. 15, 1940 THOM: NAMING MOLDS DD

As a consequence, application of the rules of nomenclature relegates
Hormodendrum to synonymy.

Although Link’s description of Cladosporium is no more definite
than the other abbreviated Latin diagnoses of that time, the general
nature of his material appears to have been recognized and the infor-
mation handed down through continuous usage in the mycological
laboratories and collections of his own and succeeding generations of
workers. In this way his concept became more definite than Dematium
herbarum of Persoon, which he believed to be the same material.

HISTORY OF CLADOSPORIUM

Before 1816, Link had assigned such forms as Dematiwm herbarum
of Persoon to Acladiwm in 1809, after presumptively satisfying him-
self that they should be separated from the other species left in De-
matium. Later he must have looked at them more carefully and con-
cluded that they should be excluded from Acladium (branchless) and
put into a genus whose name pointed to the distinctive character,
branching (clados) spore chains. The code of nomenclature we use
today for these groups of fungi begins to apply the priority rule with
Fries’s Systema mycologicum (1921-1932), rather than the previous
publications of Link, Fries, or Persoon. Up to that time such arbi-
trary changes as we find in Link, Persoon, or even Fries merely furnish
background for understanding the conditions under which the usages
we have today were developing. Cladosporium is definitely recognized
by Fries; hence it is valid.

CLADOSPORIUM OR SOME SEGREGATE

If, then, our reasoning is correct, these fungi isolated from Chromo-
blastomycosis must be assigned to Cladosporium unless adequate
characters are available to separate some one or more of the series into
one or more other genera. Without repeating details, I have already
noted that the more careful workers agree that all the strains in ques-
tion are closely related, at least (Emmons and Carrion, 1936).

In seeking lines for separating the series, describers have empha-
sized three kinds of spore production. All agree on the observation of
some strains predominantly producing the Cladosporium or, to use
their term, Hormodendrum type of spore chains: i.e., a more or less
complex system of branching chains in which the newest cells or spores
are constantly developing on the tips of the branches. They equally
agree that other strains show progressive reduction of the branching
system toward the ultimate simplicity of clusters of primary spores

56 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

ageregated rather densely about the clavate ends of the fertile hyphae
(suggesting Botrytis to Moore and Almeida). Only an occasional in-
dividual cell in these groups develops a short series of buds of the
Cladosporium type—some of them call this the Acrotheca type of spore
formation. The fact of progressive reduction but never complete re-
duction, from the complex Cladosporium type of branching chains
toward the Acrotheca type, leaves the homologies definite and readily
recognizable. |

PHIALOPHORA MEDLAR?

In contrast, the third type of fruiting structure is that described
for Phialophora. Medlar figured hyphae with black or brown walls
upon which directly or on short branches, singly or clustered, basal
or sterigmatic cells develop. These cells termed by him phialids have
firm brown to black walls and contract abruptly near the tip into
spore-producing tubes, which then abruptly flare to form cups or
cupules also with heavy brown walls. Colorless, thin-walled spores
develop successively within the bases of the cups and tend to adhere
about the tips in more or less sticky masses or spore balls. My state-
ment is one of observation that they develop there but not how they
are formed.

This structure is known to many mycologists from its presence in
other series of imperfect fungi. In some species it is reported to be
functional in connection with producing the ascosporic phase of life
history. Outside of appearances in lesions, the life histories of the
fungi of Chromoblastomycosis are entirely unknown; hence these
cupules are for the present merely additional morphology, which can
be used in diagnosis. Some species or strains show them regularly,
some under limited conditions, while they are not known in other
strains.

In culture, Miss Margaret Church and I studied the ‘‘Cadophora’”’
type of structure from decaying plant materials at least 15 years ago.
I have examined an occasional culture from pathogenic sources.
Finally Dr. Emmons passed me 20 cultures from his collection, in-
cluding transfers of strains received from Dr. Morris Moore as rep-
resenting Phialoconidiophora, Hormodendrum, and Botrytoides. I have
kept them in petri-dish and test-tube cultures for at least two months.
It requires no imagination to find colony differences perhaps justify-
ing separation into species, but essential similarities are equally ap-

2 Not Thaxter! Even though Medlar acknowledged consulting Thaxter, he took en-

tire responsibility for the naming and description of his species. Citation of the species
as P. verrucosa Thaxter is common but unauthorized by any rule.

Frs. 15, 1940 THOM: NAMING MOLDS Oo”

parent. I can find no reason for separating them widely. I can not
agree that they should be placed in different genera.

Then how about selecting a generic name from the available dozen
and putting them into it? The objection is raised that particular
names are based upon the presence of particular types of spore forma-
tion, while in the cultures these do not always satisfactorily corre-
spond with any one of the descriptions. If the group of strains from
human sources appears to be too homogeneous to place in separate
genera, aS I think, the priority rule settles the question without
further debate. Phialophora verrucosa of Medlar was certainly one of
the series and was described first. A generic name once established
loses its etymological limitation and becomes the designation of an
aggregate rather than a unit or single species. Medlar’s organism
would be the type species. His generic description would need emen-
dation, but that is readily furnished. Such a separation appears to be
justified by the preponderance of observations to date and should be
broadly enough established to include some at least of the “‘Cado-
phora”’ series described by Lundberg, Lagerberg and Melin, and
others. The student of this ‘“‘Cadophora” series upon vegetation and in
culture is sufficiently impressed with its contrast in colony and spore
producing characters in comparison with the “herbarum” lot in
Cladosporium to be unwilling to assign them to Cladosporium; hence
he would choose the alternative of broadening the generic diagnosis of
Phialophora to cover the series showing these common characters un-
til further life history studies determine real relationships elsewhere,
if any. In other words, as I see it, the well-established saprophytic
series “‘Cadophora’”’ determines the placement of the vagrant mem-
bers of that series which are found here and there throughout the
Western Hemisphere as the cause of Chromoblastomycosis in indi-
vidual humans, each time apparently de novo.

EPIDEMIOLOGICAL ARGUMENT

Epidemiological isolation of these cases clearly disposes of any ne-
cessity to recognize the parasitization of the individual human as
justifying the separation of the causal organism, as isolated from the
patient, generically or specifically from the inoculum that produced
the lesion. Each organism isolated from such a lesion is to be con-
sidered merely a stray member of a species abundant in another en-
vironment. Such an occurrence is not essentially different from

growth in a petri dish, which frequently diverges in superficial char-
acters from the colony as seen in nature. In this series, then, the imag-

58 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

inary wall between fungi growing on vegetation and as fungous para-
sites of man has broken down. As in many other biological situations,
the idea of specificity which limited organisms to particular and nar-
row biochemical roles proves to be false.

BACK TO BIBLIOGRAPHIC HISTORY

Having illustrated my topic by proposing an answer to a specific
question, I must now go back and discuss the broad aspects of my
task—naming molds. The causes of such controversies as I have just
described lie back in the history of mycology covering the past 200
years. The original mycologists were essentially microscopists—lab-
oratory examiners of material, who applied rapidly evolving micro-
scopic methods to the description of specimens collected by them-
selves or others. I am sorry to express the conviction that there are
many today who go little farther in their examination than Link or
Persoon even though with better microscopes and more adequate
literature they can not avoid seeing more.

THE FIRST 100 YEARS

If we follow back the descriptions of molds to Micheli (1729) we
find that he separated the half dozen or so molds that he called Asper-
gillus (rough headed) as yellow, white, green, black, etc. There were 2
or 3 Asperigilli, probably a Penicillium or two, or some Mucors among
them. Color seemed to him all he needed for separation. By the be-
ginning of the nineteenth century, Persoon and Link, with a larger
series of molds to separate, had raised the requirement to 3 or 4 lines
of Latin. Their figures clearly indicate that their microscopes were low
in magnification, but their descriptions and figures seemed adequate
to men who had only a few forms to separate but did not help the next
fellow who had only a few, but a different few. Details of structure
and cell arrangement were not seen and were not thought necessary.

Corda (1830-1840) went a bit farther—he had a better microscope
and figured individual cells, but their origin and relationship were not
even considered among his recorded data. Corda could draw nice
pictures—the only trouble with them has been that no one else has
ever been able to find anything like his pictures of some of these eva-
nescent molds, however valuable his drawings may be in other groups.

By 1850 Montagne concluded that most of these old descriptions of
delicate fungi were entirely uninterpretable.

These men were busy naturalists—explorers of the new domain
opened up by the compound microscope. Their colleagues were clam-

Fes. 15, 1940 THOM: NAMING MOLDS 59

oring for information. Collectors in distant lands piled the tables of
Linnaeus, Persoon, Link, Fries, Berkeley, Cooke, and the rest with
unknown specimens. What wonder that each described in hasty terms
everything that came or matched it briefly against his predecessor’s
briefer description, then identified or separated it and passed it into
his collection. Unfortunately, most of these specimens of the more
delicate fungi kept even under the most careful management quickly
dried up, separated into powder, and disappeared. Verification from
type specimens is thus impossible.

CULTURE ENTERS THE CONTROVERSY

Culture did not appear in the mycological literature until the time
of deBary, about the 1850’s, and was not used seriously for descriptive
purposes before van Tieghem in the 1860’s and Brefeld in the 1870’s.

Brefeld did a prodigious amount of work, but he also was an artist
as well as a mycologist, so that one who studies his figures with a
hand lens finds that he established an interpretation in his mind, then
covered the paper with diagrammatic drawings that often tally with
the idea more closely than with the material under his microscope. I
am not undervaluing Brefeld as a pioneer. He did much, but as to de-
tails he left much undone, and we must not hesitate to correct mis-
takes incident to method and equipment.

There was a parallel development of mold culture in France in the
laboratories of Raulin, van Tieghem, Bainier, Gueguen, and others.
Both groups of pioneers tended to assume that every form found was
new, that the number of species was small; hence fragmentary de-
scriptions run from one line to ten in length, without enough compar-
ative work among groups of congeneric species to develop the dis-
crimination between fundamental and ephemeral characters.

EXPANDING THE DIAGNOSIS

Technological mycology, as far as molds are concerned, i.e., the
controlled utilization of particular and definitely known molds in ac-
complishing biochemical processes, began to appear in the literature
in the 1860’s. By the 1890’s, a number of such processes were fairly
well recognized. Sopp and Wehmer, both students of Brefeld, made
extensive studies of Penicillia and Aspergilli in connection with in-
dustrial problems. Again, the items included in an adequate descrip-
tion were greatly increased.

In this period (1860-1890) culture as a basis of description was
deemed satisfactory if a colony was obtained by any procedure. Data

60 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 2

from such a colony were regarded as information accessory to the
study and description of natural masses presumed to be typical but
were not included in diagnosis of the species. By 1890 some efforts
were made to insure the purity of the colony. In the succeeding decade
(1890 to 1900) species diagnoses based on colonies grown on labora-
tory media began to appear, limited to saprophytic molds. The sub-
stratum upon which the colony had been originally found continued
to hold the key position as “habitat,’? which might mean much or
nothing, according to the care with which the natural situation was
studied and described. Many still consider designation of the place
where the original specimen was obtained to be the only proper habi-
tat to be cited. Unfortunately the habitat-substratum very commonly
means only that the first gross inoculum was found there, not that
the organism isolated was specifically active upon that substratum.

DEMANDS OF “APPLIED MYCOLOGY”’

Thus far the methods and descriptive practices applicable to the
molds used in industrial work were not differentiated from those of
general mycology. The exacting demand for molds with specific
adaptability to important biochemical uses began to be felt in the
1890’s. Wehmer took out patents about 1893 for the manufacture of
citric acid, using Crtromyces as the fermenting agent. By 1905, he ad-
mitted that he thought at first that there were only two species,
whereas he had now found that there were not less than six and he did
not know which was which. Now we know there are many more.

Industrial use calls for exact information as to the biochemical pos-
sibilities of each mold. Culture media and conditions must be de-
scribed with such definiteness that experimental work can be re-
peated and checked by analysis. Species must be described in terms
sharp enough to insure identification.

Green Aspergilli are not adequately covered by the name Asper-
gillus glaucus nor all green Penicillia by P. glawcum. Means must be
found to make species more tangible than rough aggregates held to-
gether by one or a few vague adjectives. The number of genera and
species have increased beyond the wildest dreams of the early mycolo-
gists. In the effort to produce descriptions that will insure identifica-
tion, species diagnoses have become progressively detailed and com-
plicated. An extreme case may be cited: Strains of a great group were
collected for many years. In working them over the monographer
first developed punctilious notes as to the culture reactions of his
whole series upon about half a dozen selected media. While making

Fes. 15, 1940 THOM: NAMING MOLDS 61

these notes, he preserved colonies of each species in alcohol. After the
biochemical record was completed, a mount of each species was made
with extreme care and a plate of drawings representing that species
was prepared. When all the plates were finished, he prepared his
Latin diagnoses by describing the structures and variations depicted
in his plates! Then the accumulated mass of data was sorted out to
species, pieced together, and published. This is one illustration in con-
nection with a monograph, purely academic in character. In practical
fact, there are many unrecognizable species among those described in
detail within the last few years, and by all of us! WHY?

UNINTERPRETABLE DESCRIPTIONS

There are several reasons for failing to identify species from de-
scriptions. There is little agreement as to just what characters are
fundamental to genera and to species, and which are incidental varia-
tions representing direct response to environment. Great stress has
has been put upon numbers and measurements of parts or details of
branching systems. Large numbers of spores have been measured to
the fraction of the micron, then the totals averaged or statistically
analyzed to the fraction of a micron. As a result, emphasis upon un-
important details has often claimed the users’ attention while the
fundamental information escaped.

ESSENTIALS OF DIAGNOSIS

A safe description requires the exact identification of the culture
substratum, of the biochemical effects of the culture upon that sub-
stratum, and correspondingly a series of observations of the organism
itself upon that substratum.

Purity of culture is essential. The presence of bacteria, Actinomy-
cetes, or other molds often alters colony characters. Different con-
taminants produce different alterations; hence entire elimination of
other organisms is essential. Unfortunately, there are quite well-
known laboratories from which cultures consistently show mites, as
well as molds or bacteria. One is compelled to believe that some
workers have never yet seen mites, recognized their ravages in cul-
ture, or distinguished the characteristic smell that usually betrays
their presence. |

More important yet, actual relations and sequences between cells
are fundamental. New cells may be formed by fission: The older cell
is cut into equal halves; or, they may bud out at one of a dozen places

62 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

and in one of a dozen ways. Chains of spores may arise by budding
so that the newest cells are always farthest from the basal cell or sporo-
phore. Unbranched chains commonly arise from a basal cell—always
showing the distal cell as the oldest while the newest cell is attached
directly to the basal or sterigmatic cell. Again, this observation is
fundamental.

A description of a ripe spore mass as a mass, or, pulled to pieces and
the pieces described as found, may miss completely the significant
facts. Whole series of descriptions that disregard cell succession in
spore formation are simply uninterpretable, except in species in which
satisfactory material was preserved or which have been found iden-
tifiable from some other line of observation.

Finally, details of cell wall structure (as Jeffrey said about paleo-
botanical specimens) may furnish as many real clews to relationship
as the orthodox observation of sporogenous masses. Whole series of
Aspergilli and Penicillia show such consistent markings of the stalk
wall that examination of the sporophore wall with oil immersion ob-
jectives furnishes the most useful, most general, in fact, most easily
determined series character.

But the nomenclatorial sins are not all chargeable to describers.
The most striking lack among users of descriptive literature is in the
appreciation of the cell relations involved in whatever structures they
find. They fail to give proper consideration to cell contents, cell walls,
their structure, color, and markings, to cell-succession in the forma-
tion of the sporogenous tissue seen under the microscope, and to the
methods of aggregation of spores shown by heads, chains, or discharg-
ing mechanism. In other words, they fail to understand that identifi-
cation of a mold is not accomplished by a superficial examination with
a hand lens or with the low magnifications of the microscope. Exact-
ing observation of the detail indicated in descriptive keys is ordinarily
guided by those same keys. Failure to follow out definite instructions
is hardly justifiable.

In the more complicated groups, it is not just matching a culture
at a glance against a list of names, but the integration of all that one
can learn by painstaking examination against the literature and in-
vestigations of perhaps 100 years. Into that investigation must go a
first-hand knowledge of the life history of not one or two species but
whole groups of species; it must include many years of observation in
the field, controlled development in the culture room, and diligent
reading in the library.

Fes. 15, 1940 THOM: NAMING MOLDS 63

The need for care in establishing one’s right to use generic and spe-
cific names can not be brushed away with a wave of the hand.

I sat beside a well-known worker not long ago and heard him tell an
inquirer, ‘‘Why bother about the name—-select one of them, go ahead
and study your organism, and let future systematists decide where
your organism belongs!’ That dictum may suffice for some folks! But
it creates chaos when the industrial or technological scientist makes
the wrong selection and puts a series of industrial or biochemical pa-
pers about genus “‘X”’ and species “Y’’ into the literature when he
actually worked with genus ‘‘A’’ and species “B.”’ Indexed that way,
an error is often cited many times as a fact by workers who have no
means at hand to protect themselves. This is no hypothetical dilem-
ma. I can name instances. Men regularly ask me for cultures based on
such papers. Sometimes I can guess what they want; sometimes I
have no idea why they make the request. Again I am sure they se-
lected names at random.

In direct controversion of the dictum above, I believe it safe to say
that a critical cultural and microscopical study of saprophytic molds
will in the vast majority of cases throw together into homogeneous
groups the things that eventually prove to belong together even
though exact relations among them can not always be predicted from
the hyphomycete stage.

To the industrial mycologist who is confronted by a mold isolated
from where you please—important or merely questionable—the prob-
lem of what to call it is not theoretical, but practical. He should be
able to examine the thing before him and reach some diagnostic char-
acters that will lead to the correct literature of the species. In other
words, however unimportant naming may be, as an end in itself, the
descriptive and taxonomic problem must be solved before he can
reach what his predecessors and perhaps his colleagues have written

about the particular thing on the table.

If you are to do technical work with a particular mold, the funda-
mental dictum is: Know your organism by name and relationship,
know it morphologically and physiologically, macroscopically and mi-
croscopically—know it so well that if anything goes wrong, you will
detect the abnormality and correct it or make an adequate record.
That applies whether you are a mycologist, a pathologist, a chemist,
a physiologist, or any other brand of specialist; the man who fails to
know his organism thoroughly is helpless before contamination,
losses, or replacements, which often destroy the value of the results.

64 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

SUMMARY

I have tried to picture to you some of the problems of the ‘“‘applied
mycologist’? who works with the so-called “‘common molds.” As fac-
tors in human affairs, they spoil man’s food, mildew his clothes, pol-
lute his storerooms, and even attack his body. He gets some return by
eating a few. of them and using others in controlled fermentations of
many kinds. He can not escape from them—he must live with them.
The alternative, then, is to know them—individually, that he may use
or combat the single species; as groups, that he may so compare and
systematize his information that each item in it may contribute to
arranged and ordered systems of knowledge. In a recent memorial to
a great museum specialist, his services to mankind are listed as ‘‘(1)
Scientific research, (2) Nomenclatural,’’ etc. One makes bold to say
that there is fully as much reasoning exhibited in comparing, recon-
structing, and classifying a fossil as in digging it out of a hillside. The
one dictum that must not be forgotten is that no single item has per-
manent value unless it represents the closest approximation to truth
that can be reached by using all the means available. Any work, to
be worth while, must be a rigorous search for truth. No bypath can
be permitted to lure the worker aside for fancied results. If rigorous
good faith in method, in performance and in interpretation are main-
tained, usefulness from the results can not possibly detract from the
purity of the science. It is not the “‘pot of gold” that pollutes; it is the
method of getting it. It is true in applied mycology as everywhere else
that ‘‘He that entereth not by the door into the sheepfold, but climb-
eth up some other way, the same is a thief and a robber.” If the solu-
tion of problems in human service makes our work applied science,
then let us glory in the name.

Frs. 15, 1940 PHILLIPS: ANSELME PAYEN 65

CHEMISTRY.—Anselme Payen, distinguished French chemist and
pioneer investigator of the chemistry of lignin. Max PHILLIPS,
U. S. Bureau of Agricultural Chemistry and Engineering.

In these days when there is so much research activity on the chem-
istry and utilization of lignin and cellulose, it is fitting that we pause
and pay our respects to the memory of a man who was a pioneer in-
vestigator of the chemistry of these two substances and who is now
virtually forgotten. This man, Anselme Payen by name, published
the results of his investigations on lignin and cellulose about 100
years ago. Although the percentage elementary composition of wood
had previously been determined by Gay-Lussac and Thenard, Payen
was the first one to attempt a separation of wood into its component
parts. He treated wood with nitric acid and obtained a fibrous sub-
stance, which was relatively resistant to this reagent. This substance
Payen called “‘cellulose,” a term that he coined and introduced into
chemical literature (Compt. Rend. Acad. Sci. Paris 8:51. 1839). He
showed that cellulose had the same percentage composition as starch,
and was apparently isomeric with it. He found that irrespective of its
origin, cellulose had the same chemical composition, which could be
represented by the formula C,H,.0; and was apparently the identical
substance in all plants. Payen also found that in isolating the cellulose
he had to remove a substance or a group of substances, which had a
higher percentage of carbon than cellulose. These substances Payen
called “incrusting materials’ (‘‘Jes matieres encrustantes’’), and he
considered that the cellulose was mechanically incrusted or impreg-
nated by them. These incrusting materials were later (1857) designat-
ed by Schulze as lignin, a term previously used by the botanist and
plant physiologist de Candolle. The “‘incrustation hypothesis’’ of
Payen was in the main supported by Schulze, although other inves-
tigators, among them Erdmann, opposed Payen’s views. It is of in-
terest, however, to point out that Payen’s “‘incrustation hypothesis’’
has in more recent years been vigorously supported by Wislicenus and
by Freudenberg.

Anselme Payen was born in Paris, France, on January 6, 1795. He
was the son of Jean Baptiste Pierre Payen and Marie Frangoise Jean-
son de Courtenay. The elder Payen was educated at the Collége de
Navarre in Paris, where he distinguished himself especially in philoso-
phy and in the sciences. Through the insistence of his parents, how-

* Read before the Division of History of Chemistry at the 98th meeting of the Amer-

ican Chemical Society held in Boston, Mass., September 11 to 15, 1939. Received
December 11, 1939.

66 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 2

ever, he studied law and was for a time assistant to the procurator of
the King for the city of Paris. This legal work, however, did not prove
to his liking and in 1792 he established a factory at Grenelle, a suburb
of Paris, for the production of various chemicals, chiefly sal ammo-
niac. This enterprise proved to be quite successful, and we soon find
him engaged in the production of other chemical substances, such as
sulphuric acid, hydrochloric acid, borax, refined sulphur, soda, and
gelatin. He also established at Vaugirard a factory for the production
of sugar from sugar beets. This was then a new industry in France, and
during the continental blockade of the Napoleonic wars the cultiva-
tion of sugar beets and the extraction and refining of sugar therefrom
were greatly encouraged by the French Government.

Anselme Payen received his early education from his father, who
was a strict disciplinarian and very early inculeated in his son the
habit of systematic study, a characteristic he retained for the rest of
his life. However, in order to get a thorough grounding in the sciences
his father sent him to the Ecole Polytechnique in Paris where he
studied chemistry under Vaugeulin, physics under Fourcroy, and
mathematics under Trémery.

When young Payen was hardly 20 years of age his father placed
him in charge of a plant for the refining of borax. This was then a rela-
tively new industry in France for the Dutch virtually had a monopoly
of the production of refined borax in Europe. The crude borax con-
taining material was imported by the Dutch from the Orient and then
refined by them. Young Payen conceived the idea of preparing borax
synthetically from soda and boric acid, which was then obtained al-
most entirely from the hot springs and lakes of Tuscany. Payen’s
efforts were crowned with success, and in 1820 he placed on the
market synthetic borax at one-third of the then prevailing price and
thus succeeded in establishing a new industry in France.

In 1820 Payen’s father died, and at the age of 25 years the young
man had to assume full responsibility for the management of several
factories established by his father, including the factory for the pro-
duction of sugar from beets. For the clarification of the sugar solu-
tions vegetable charcoal, principally wood charcoal, was employed.
This peculiar property of wood charcoal was first brought to the at-
tention of chemists in 1785 by an apothecary by the name of Lowitz
of St. Petersburg, Russia, although according to E. O. von Lippmann
the decolorizing property of wood charcoal was known as early as the
fifteenth century to some of the German alum and saltpeter producers.
However, this observation was apparently entirely forgotten. As al-

Fes. 15, 1940 PHILLIPS: ANSELME PAYEN 67

ready mentioned, among the chemicals made by Payen in his factory
at Grenelle was sal ammoniac. This was prepared from the complex
mixture of volatile substances obtained by the destructive distillation
of bones and other products of animal origin. In the retort there re-
mained a charred mass known as animal charcoal or bone char, which
was then largely a useless by-product. Although Figuier of Montpel-
lier in 1811 called attention to the decolorizing property of animal
charcoal no application was made of this discovery until Payen got
interested in this problem. Payen was motivated in this study by the
double objective of improving the process of sugar refining, and of
utilizing the animal charcoal. In a paper published in 1822, entitled
“Theory of the Action of Animal Charcoal and Its Application to the
Refining of Sugar,’’ Payen gave results of a thorough study of the de-
colorizing properties of animal charcoal and pointed out that it owes its
activity to the peculiar shape and state of aggregation of the carbon
in this material. He also showed that it had the capacity of removing
certain salts from solution and thus facilitated the crystallization of
the sugar. He also developed an apparatus, called a “decolorimeter,”’
for the determination of the decolorizing ability of various lots of ani-
mal charcoal. This may be considered the first important scientific
paper published by Payen.

The credit for first using animal charcoal industrially for the refin-
ing of sugar clearly belongs to Payen. It was as a result of his zeal and
industry that the popular prejudice against the application of animal
charcoal to the refining of substances used for human consumption
such as sirup and sugar was overcome. In addition to thus improving
the process of sugar refining, Payen’s work resulted in establishing an
important new industry, namely, that of animal charcoal production.
We thus see that a product which had heretofore been entirely useless
became the basis of an important new industry. Few sugar technolo-
gists are today aware of the fact that when they decolorize their sugar
‘solutions by means of animal charcoal they are making use of a meth-
od of sugar refining introduced more than a century ago by the
French chemist Payen, now all but forgotten.

Because of his work on sugar beets, Payen very early in his life be-
came interested in agricultural chemistry, and in the application of
science in general to the improvement of agricultural practices, an in-
terest he retained throughout his long and useful life. In these days
when we hear so much about the industrial utilization of agricultural
products, it may be of interest to point out that Payen more than 100
years ago not only advocated this very thing, but also made impor-

68 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

tant contributions toward its realization. In a book he published in
1826 together with Chevalier, entitled ‘‘Traité de la Pomme de
Terre’’ (Treatise on the Potato), he described in considerable detail
not only the preparation of various foods and feeding stuffs for human
and animal consumption, including the production of sugar and sirup,
but also the methods for the preparation of starch and alcohol from
potatoes. Part of the results contained in this book were published in
the Proceedings of the Central Agricultural Society of France in the
year 1823 and in recognition of this work Payen was given a gold
medal by that society.

In the early part of the nineteenth century the need arose in France
for the development of more economical methods of disposal of the
carcasses of domestic animals that died of accident or of diseases of an
epidemic character. Accordingly, in 1825 the Central Agricultural
Society of France offered a first prize of 1000 francs for the most prac-
tical and economical method for the utilization of the carcasses of
these animals. In 1830 the first prize was awarded to Payen for his
132-page memoir, which was published in the proceedings of the so-
ciety for that year. There is a great deal of analytical data in this pa-
per, which in many cases were obtained by methods devised by Payen.
The method developed by him for the determination of nitrogen con-
sisted in heating the sample to red heat and collecting the gaseous
products in dilute sulphuric acid. While this did not, of course, give
him all the nitrogen in the sample in the form of ammonia, it is never-
theless of interest from the historical standpoint as it was the fore-
runner of the Will and Varrentrapp method, where it is recalled the
sample is heated with soda-lime and the ammonia collected. This was
later superseded by the now well-known Kjeldahl method.

The methods that Payen described for the utilization in industry
and in agriculture of various products of animal origin have proved to
be practical, and many of the processes now in use for the utilization
of the by-products of the meat-packing industry can be traced to
those described by Payen in his paper published more than 100 years
ago.

In 1835, at the age of 40, Payen gave up all active participation in
the various manufacturing enterprises that he was interested in and
accepted the position of professor of industrial and agricultural chem-
istry at the Ecole Centrale des Arts et Manufactures in order that he
might devote all his time to teaching and scientific research. In 1839
in addition to his duties at the Ecole Centrale he also accepted the
professorship of applied chemistry at the Conservatoire des Arts et.

Frs. 15, 1940 PHILLIPS: ANSELME PAYEN 69

Métiers. These two positions he held until the time of his death.
The next 36 years of Payen’s life were undoubtedly the most fruit-
ful of his entire career, judged from the many contributions of a fun-
damental character he made to chemistry. He published about 200
scientific papers in various scientific journals such as the Comptes
Rendus, Bulletin de la Société d’Encouragement pour |’ Industrie Na-
tional, Annales du Conservatoire des Arts et Métiers, Annals de Chi-
mie et de Physique, Annales d’Histoire Naturelle, Bulletin de la
Société Chimique, Journal de Chimie et Médecine, Mémoires de la
Société Centrale d’Agriculture de France, Annals des Mines, and
others. These papers covered a wide field of investigation and included
such subjects as starch, dextrin, sugar, lignin, cellulose, bitumen, and
various phases of agricultural chemistry including studies on plant
and animal nutrition, the latter studies being conducted in coopera-
tion with the famous agricultural chemist Boussingault. He also pub-
lished papers in the field of inorganic chemistry. It was in connection
with his studies on starch that Payen together with Persoz discovered
the enzyme diastase. The paper on starch that he published in 1836
is remarkable for its clearness and precision. He showed that starches
obtained from different sources differed as to size, shape, and state of
aggregation, but that they all had the same chemical composition.

Payen had a practical turn of mind, and in the midst of his work on
the purely scientific phases of chemistry he was always interested in
the application of chemistry to industry, agriculture, hygiene, and
medicine.

Payen was a prolific writer of books on industrial, agriculural, and
food chemistry, and many of his books became standard works of
reference and were translated into English and into several other
Kuropean languages. Special mention should be made in this con-
nection of his ‘“Traité de Chimie Industrielle,’ in which Payen de-
scribes in great detail the various processes then used in chemical in-
dustry. Even the superficial reader of this book can see at once that
its author was not a mere compiler, but rather one who has actually
had many years of practical experience in the operation of the several
processes described by him. In fact many of the processes there de-
scribed were developed and improved by Payen. From the titles of the
various books published by Payen one may get some idea of his ex-
tensive knowledge and interests.

The following is a list of his more important books (for a complete
list of Payen’s books the reader is referred to H. C. Bolton’s ‘‘A Select
Bibliography of Chemistry,’’ Washington, 1893):

70 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

(1) Traité de la pomme de terre; sa culture, ses divers emplois dans les
préparations alimentaires, les arts économiques, la fabrication du sirop,
de l’eau-de-vie, de la potasse, ete. Paris, 1826. (Published with Chevalier.)

(2) Traité de la fabrication et du raffinage des sucres. Paris, 1832.

(3) Des engrais. Théorie actuelle de leur action sur les plantes principaux,
moyens d’en obtenir le plus d’effet utile. Paris, 1839.

(4) Mémoire sur l’amidon, la dextrine et la diastase consideres sous les
points de vue anatomique, chimique, et physiologique. Paris, 18—.

(5) Précis de chimie industrielle 4 l’usage des écoles préparatoires, aux
professions industrielles, des fabricants et des agriculteurs. Paris, 1849.

(6) Précis d’agriculture théorique et pratique a l’usage des écoles d’agri-
culture, des propriétaires et des fermiers. Paris, 1851. (Published with A.
Richard.)

(7) Traité de la distillation des betteraves. Paris, 1854.

(8) Traité complet de la distillation des principales substances qui
peuvent fournir de l’alcool; vins, grains, betteraves, fecule, tiges, fruits,
racines, tubercules, bulbes, etc. Paris, 1857.

(9) Précis théorique et pratique des substances alimentaires et des moyens
de les améliorer, de les conserver et d’en reconnaitre les alterations, 4th ed.,
Paris, 1865.

According to Ostwald most great men can be divided into two
classes. Those who in their youth, that is to say between 20 and 35
years of age, do their most outstanding work belong to the romantic
type. Those who belong to the classical type are men who do out-
standing work during their entire life. Payen clearly belonged to the
latter class.

Payen was a member of many scientific societies. He was elected a
member of the Central Agricultural Society of France (La Société
Centrale d’Agriculture de France) in 1833 and was its permanent
secretary for 26 years. He was elected a member of the French Acad-
emy of Sciences in 1842. Among the other societies to which he be-
longed may be mentioned |’ Academie de Médecine, Société d’Encour- -
agement pour l’Industrie Nationale, Société d’Horticulture de la
Seine, and of the Council of Hygiene and Public Health.

Payen served the French Government in diverse capacities, particu-
larly as a member of various governmental commissions. He was made
a Knight of the Legion of Honor by Charles X in 1828. Louis Philippe
made him an officer of the Legion of Honor in 1847, and in 1863 Na-
poleon III elevated him to the rank of commander.

One of the characteristics of Payen was his intense patriotism, and
for 40 years he served as commander of the battalion of National
Guard of the town of Grenelle, where he made his home.

In 1821 Payen married Zelie Charlotte Mélanie Thomas, which
proved to be a very happy union. There were five children in the
family, four of which, however, died in childhood and only one child,
a daughter, survived him.

Fes. 15, 1940 SWALLEN: NEW SPECIES OF PARIANA 71

Payen’s last days were greatly saddened by the disasters that
France suffered as a result of the Franco-Prussian War. In spite of his
advanced age he refused to leave the city of Paris on the approach of
the Prussian army and displayed a great zeal in the study of all prob-
lems relating to the feeding of the besieged and famished city. On
May 9, 1871, while attending one of the sessions of the Academy of
Medicine he became ill and died three days later (May 12). Few of his
friends and comrades were able to pay their last respects to this dis-
tinguished savant, for those were the days preceding the establish-
ment of the Paris Commune and civil war raged on the streets of
Paris. He was buried in the cemetery at Grenelle.

The biographic material contained in this paper was obtained from the
following sources:

(1) Amer. Journ. Pharm. (ser. 4) 1: 432. 1871. (Anon. obituary notice.)

(2) Eloge biographique de M. Anselme Payen, by J. A. Barral, in Mé-
moires Publies par la Société Centrale d’Agriculture de France, pp. 67-87.
1873.

(3) Eloge de M. Payen, by Aimé Girard, in Annales du Conservatoire
Imperial des Arts et Métiers 9: 317-331. 1873.

A complete list of the scientific papers published by Payen can be found

in the Catalogue of Scientific Papers, compiled by the Royal Society of
London.

BOTANY.—Eight new species of Pariana. Jason R. SWALLEN,
U.S. Bureau of Plant Industry.!

Pariana is an anomalous genus of grasses that inhabits tropical
forests and forest borders from Panama to Brazil and Peru. Because
of the paucity of material in herbaria, the species have been con-
sidered as rare. The author, however, in 1933-34 observed these
grasses growing rather commonly in northern Brazil, in places even
completely covering the ground over rather large areas. The inflor-
escences are frequently entirely hidden by the foliage, which may
account for the earlier collectors overlooking them. Tutin? recently
revised the genus recognizing 23 species. Since then, one has been
described from Ecuador by Pilger.* The eight species herein described
bring the total number to 32.

Pariana nervata Swallen, sp. nov.

Perennis; culmi uniformes e rhizomatibus crassis erecti, 90-130 cm alti;
vaginae inferiores internodiis longiores vel breviores laminis obsoletis,
superiores internodiis multo longiores foliosae, auriculatae, sparse fimbriat-

1 Received October 26, 1939.
2 Revision of the genus Pariana. Journ. Linn. Soc. Bot. 50: 337-362. 1936,
§ Notizbl. Bot. Gart. Berlin 14; 323, 1939,

72 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

ae, glabrae vel sparse pilosae; ligula 3 mm longa, truncata; laminae 15-26
cm longae, 4-6 cm latae, acuminatae, infra scabrae; inflorescentia pendula,
20 cm longa, 1.3 cm lata; spiculae masculae 6.5-8 mm longae, pedicellis
occultis, 3 mm longis minute pubescentibus; glumae 5-6 mm longae, 1.5—
2.5 mm latae, 2—3-nerves, abrupte acutae, scaberulae, marginibus ciliatis;
lemma 6.5-8 mm longum, 3.5—4 mm latum, 5-nerve, subobtusum, scabrum ;
spiculae femineae uniseriatae; glumae 8 mm longae, 3-nerves, acutae, sca-
berulae, reticulatae; fructus 6 mm longus apice scaber.

Perennial, with leafy fertile culms; culms 90-130 em tall, erect from rather
short thick rhizomes; lower sheaths longer or shorter than the internodes,
the blades obsolete, the upper blade-bearing ones much longer than the
internodes, somewhat crowded, auriculate, sparsely fimbriate, glabrous or
sparsely pilose; ligule 3 mm long, truncate; blades 15-26 cm long, 4-6 cm
wide, acuminate, the lower surface scabrous, the upper glabrous; inflores-
cence drooping, 20 cm long, 1.3 cm thick; staminate spikelets 6.5-8 mm
long, the pedicels hidden, 3 mm long, minutely pubescent; glumes 5-6 mm
long, 1.5-2.5 mm wide, 2—3-nerved, abruptly acute, scaberulous, the margins
ciliate; lemma 6.5-8 mm long, 3.5-4 mm wide, 5-nerved, subobtuse, sca-
brous; pistillate spikelets in one row; glumes 8 mm long, 3-nerved, acute,
scaberulous, reticulate; fruit 6 mm long, scabrous at the apex.

Type in the U.S. National Herbarium, no. 1613658, collected at edge of
forest, Mosqueira, near Belem, Para, Brazil, June 15, 1934, by Jason R.
Swallen (no. 4873).

This species of Pariana differs from all the others in the conspicuously 5-
nerved lemma of the staminate spikelets.

Pariana ovalifolia Swallen, sp. nov.

Perennis; culmi 135 cm alti, basi decumbentes; vaginae inferiores elongat-
ae internodiis breviores laminis valde reductis, superiores internodiis
multo longiores, glabrae, reticulatae, in ore sparse fimbriatae; ligula trun-
cata 2-4 mm longa; laminae 17—24 cm longae, 6—7 cm latae, acuminatae,
infra pubescentes, supra glabrae, petiolis 5-8 mm longis glabris; inflores-
centia pendula, 16 cm longa; spiculae masculae pedicellis 3 mm longis,
pubescentibus; glumae 2 mm longae, 1—2-nerves, acutae, pubescentes,
marginibus ciliatis; lemma 5-6 mm longum, 2.5-3 mm latum, subobtusum,
dense pubescens; spiculae femineae 7 mm longae; glumae aequales, 3-
nerves, pubescentes; lemma 6 mm longum, acutum, glabrum apice pu-
bescens.

Perennial; culms about 135 ecm tall, erect from a decumbent base, the
fertile one leafy toward the summit, bladeless below, glabrous; lower sheaths
elongate, shorter than the internodes, the upper crowded, longer than the
internodes, glabrous, reticulate-veined, occasionally bearing a few fimbriae
at the mouth; ligule truncate, 2-4 mm long; blades 17-24 cm long, 6—7 cm
wide, the two uppermost reduced, narrowed toward the base, acuminate,
glaucous and pubescent on the lower surface, green and glabrous on the
upper, the petiole 5-8 mm long; inflorescence slender, drooping, 16 cm long,
the whorls of spikelets distant, not overlapping in the lower part, gradually
becoming more crowded toward the summit, but the staminate florets never
obscuring the pedicels of the staminate spikelets in the next higher whorl;
staminate spikelets 5-6 mm long, the pedicels 3 mm long, pubescent, es-
pecially at the base and on the margins; glumes 2 mm long, 1 mm broad at
the base, 1 or sometimes 2-nerved, acute, more or less pubescent, the mar-
gins ciliate; lemma 5-6 mm long, 2.5-3 mm wide, subobtuse, densely pu-.

Fre. 15, 1940 SWALLEN: NEW SPECIES OF PARIANA 13

bescent; fertile spikelet 7 mm long; glumes thin, equal, 3-nerved, pubescent;
lemma 6 mm long, acute, pubescent toward the tip, otherwise glabrous.

Type in the U. S. National Herbarium, no. 1614216, collected in forest
borders 35 kms north of Monte Alegre, Para, Brazil, January 28—Febru-
ary 1, 1934, by Jason R. Swallen (no. 3379).

Pariana ovalifolium belongs to the Foliosae group in which the fertile
culms are leafy. It is most closely related to P. lunata Nees, which is a much
smaller plant, about half as high, with smaller lanceolate rather than oval
blades, which are broad and rounded at the base.

Pariana aurita Swallen, sp. nov.

Perennis; culmi biformes, erecti vel basi decumbentes. Culmi steriles ad
80 cm alti in parte superiore foliosi, scabri; vaginae inferiores internodiis
multo breviores, superiores internodiis multo longiores, carinatae, auricu-
latae; ligula 8-15 mm longa; laminae 15-28 cm longae, 3.5—-5.8 cm latae,
acuminatae vel subattenuatae, glabrae vel scaberulae, petiolis crassis, 7-10
mm longis, pubescentibus. Culmi fertiles graciles, 50 cm alti, glabri; vaginae
internodiis longiores laminis obsoletis; ligula curta; inflorescentia 12 cm
longa, gracilis, pendula; spiculae masculae 4 mm longae, pedicellis 2-4 mm
longis, pubescentibus vel pilosis, marginibus ciliatis; glumae 2.5-3 mm
longae, angustae, acuminatae, 1—-2-nerves; lemma 4 mm longum, 1.5 mm
latum, reticulatum, scaberulum; antherae +12, lineares, 2.6 mm longae;.
spiculae femineae biseriatae 6 mm longae; glumae l-nerves, acutae, pu-
bescentes, marginibus ciliatis; lemma 5 mm longum, apice minute pu-
bescens. ;

Perennial; culms erect or somewhat decumbent at the base. Sterile culms
80 em tall, leafy in the upper fourth; lower sheaths bladeless, much shorter
than the internodes, rounded on the back, the upper ones crowded, much
longer than the internodes, distinctly keeled, glabrous; ligule 8-15 mm long,
joined at the base with the conspicuous auricles of the sheaths; blades 15-28
em long, long-acuminate, narrowed toward the base, glabrous or scaberulous;
fertile culms 50 cm tall, the bladeless sheaths mostly longer than the inter-
nodes, rounded on the back, glabrous; ligule very short or wanting; in-
florescence drooping, 12 cm long, the whorls of spikelets scarcely over-
lapping; staminate spikelets 4 mm long, the pedicels conspicuous, 2-4 mm
long, sometimes fused toward the base, pubescent or pilose, the margins
ciliate; glumes 2.5-3 mm long, usually 1 or sometimes 2-nerved, acuminate,
rather narrow at the base, glabrous or minutely pubescent, the margins
very shortly ciliate; lemma 4 mm long, 1.5 mm wide, finely reticulate-
veined, scaberulous; anthers +12, linear, 2.5 mm long; pistillate spikelet
6 mm long, the glumes acute, 1-nerved, pubescent, narrower than the ma-
ture fruit; fruit 5 mm long, the lemma minutely pubescent near the tip and
on the lateral nerves.

Type in the U. 8. National Herbarium, no. 1539376 collected in dense for-
est at the mouth of the Rio Santiago, above Pongo de Manseriche, Depart-
ment Loreto, Peru, November 17, 1931, by Ynes Mexia (no. 6116).

P. aurita is a very striking species, which is probably most closely related
to P. gracilis Doell, resembling it in the slender inflorescence of rather
distant whorls of spikelets. However, P. gracilis has fewer, much smaller
blades 10-15 cm long, and a ligule only 2-2.5 mm long.

Pariana distans Swallen, sp. nov.

Perennis; culmi difformes e rhizomatibus erecti, scabri. Culmi steriles 40
cm alti in parte superiore foliosi; vaginae inferiores laminis obsoletis inter-

74 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

nodiis breviores, superiores internodiis longiores, pubescentes; ligula trun-
cata, 2 mm longa, pubescens; laminae 10-15 cm longae, 1.8-2.5 em latae,
lanceolatae, acuminatae, infra scaberulae supra pubescentes. Culmi fertiles
40 cm alti, infra inflorescentiam dense pubescentes; vaginae internodiis
longiores, glabrae, laminis obsoletis; inflorescentia 9.5 cm longa, erecta,
verticillis spicularum remotis; spiculae masculae 7 mm longae pedicellis
3 mm longis, dense pubescentibus; glumae 4-5 mm longae, 3-nerves, sca-
brae, marginibus ciliatis; lemma 7 mm longum, 2.5 mm latum, pubescens;
spiculae femineae incognitae.

Perennial; culms numerous, erect from rhizomes, about 40 cm tall, sca-
brous. Sterile culms leafy toward the summit, the lower sheaths bladeless;
lowermost sheaths short, longer than the internodes, the one of the mid-
culm elongate, shorter than the internode, the upper bladebearing sheaths
again longer than the internodes, glabrous or minutely pubescent; ligule
truncate, about 2 mm long, pubescent; blades oblong-lanceolate, 10-15 em
long, 1.8-2.5 em wide, acuminate, shortly soft pubescent on the lower sur-
face, scaberulous above, the petiole 2 mm long, pubescent. Fertile culms
densely pubescent below the inflorescence; sheaths all longer than the in-
ternodes, bladeless, the uppermost one elongate, glabrous; inflorescence 9.4
em long, erect, the whorls of spikelets rather distant, the staminate florets
scarcely reaching the base of the pedicels of the staminate spikelets of the
whorl above; pedicels of the staminate spikelets 3 mm long, densely pu-
bescent, more or less fused together; glumes 4-5 mm long, 3-nerved, acute,
scabrous, the margins shortly ciliate; lemma 7 mm long, 2.5 mm wide.
pubescent; pistillate spikelet undeveloped.

Type in the U. 8S. National Herbarium, no. 1614187, collected in forest
borders, Santarem, Para, Brazil, January 19-26, 1934, by Jason R. Swallen
(no. 3286).

This species differs from all the others in the Graciles group in the 3-nerved
glumes of the staminate spikelets and the longer staminate lemma. In the
other species of this group the glumes of the staminate spikelets are only
l-nerved (rarely 2-nerved), and the lemmas are not more than 5.5-6 mm
long.

Pariana ligulata Swallen, sp. nov.

Perennis; culmi difformes caespitosi, erecti, basi decumbentes. Culmi
steriles 70-95 cm alti, in parte superiore foliosi; vaginae inferiores inter-
nodiis breviores laminis obsoletis, superiores internodiis multi longiores,
carinatae; ligula 6-10 mm longa, firma, obtusa vel truncata; laminae 15-32
cm longae, 3-4 cm latae, subattenuatae, reticulatae, petiolis crassis, 6-10
mm longis glabris vel minute pubescentibus. Culmi fertiles 50-60 cm alti,
laminis obsoletis; vaginae internodiis longiores; inflorescentia 12 cm longa,
pendula, parte inferiore in vagina suprema inclusa, verticillis spicularum
approximatis; spiculae masculae 5-6 mm longae, pedicellis 4-5 mm longis
sparse pubescentibus; glumae 1.5-2 mm longae, 1—2-nerves, triangulae,
acutae, scabrae; lemma 5-6 mm longum, 2—2.5 mm latum, 3-nerve, acutum,
reticulatum, scabrum; antherae lineares, 3 mm longae; spiculae femineae
9 mm longae; glumae 1—3-nerves acutae vel subacuminatae, scabrae; lemma
7mm longum, acutum, subapiculatum.

Perennial; culms in rather large spreading clumps, erect from a decum-
bent base. Sterile culms 70-95 cm tall, leafy in the upper half; sheaths
elongate, the lower ones a little shorter than the internodes, the upper ones

longer than the internodes but scarcely crowded, keeled toward the summit, |

, Ey

75

NEW SPECIES OF PARIANA

SWALLEN

Frs. 15, 1940

oN

SEN

I

, showing
Pe Curia
All drawings

5]

3

]
°

; 8, P. velutina

P. ovalifolia;
Weintraub.

’

2

, P. sociata; 7, P. modesta

2

C

1, P. nervata;

Figs. 1-8.—Front view of staminate spikelets of new species of Pariana
type specimens by Mrs. Frances

the pedicel, glumes, and lemma, X5
4, P. distans; 5, P. ligulata; 6

made from the

76 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 2

reticulate-veined; ligule 6-10 mm long, firm, obtuse or truncate; blades
15-82 em long, 3-4 cm wide, more or less plaited, the transverse veins con-
spicuous on the lower surface, acuminate, narrowly rounded at the base,
glabrous on both surfaces; petiole 6-10 cm long, glabrous or minutely
pubescent. Fertile culms 50-60 cm tall, nearly hidden under the foliage of the
sterile culms; sheaths somewhat inflated, all longer than the internodes,
bladeless or sometimes with blades 1—2.5 cm long; inflorescence drooping,
about 12 cm long, the whorls of spikelets distant, scarcely overlapping;
staminate spikelets 5-6 mm long, the pedicels 4-5 mm long, sparingly
pubescent; glumes 1.5—-2 mm long, triangular, acute, 1 or sometimes 2-
nerved, scabrous; lemma 5—6 mm long, 2.5 mm wide, strongly 3-nerved,
reticulate-veined above, scabrous; anthers about 3 mm long; fertile spikelet
9 mm long, the glumes l-nerved, scabrous or pubescent at the base; fruit 7
mm long, abruptly acute, almost apiculate.

Type in the U. 8. National Herbarium, no. 1613673, collected in forest at
Boa Vista, Rio Tapajos, Para, Brazil, January 7-13, 1934, by Jason R.
Swallen (no. 3177).

P. ligulata differs from all other species in the Graciles group in the large
pistillate spikelets and the long comparatively narrow plaited blades.

Pariana sociata Swallen, sp. nov.

Perennis. Culmi difformes e rhizomatibus erecti vel basi decumbentes.
Culmi steriles +40 cm alti, glabri, in parte superiore foliosi; vaginae glabrae
vel in parte superiore pubescentes, inferiores internodiis multo breviores,
superiores internodiis multo longiores in ore dense fimbriatae pilis 10-15 mm
longis; laminae 8.5—11 cm longae, 1.7—2.7 cm latae, acuminatae, infra minute
pubescentes, supra scaberulae, marginibus serratis; ligula truncata 1 mm
longa. Culmi fertiles 30 cm alti; vaginae internodiis longiores, inflatae, gla-
brae vel minute pubescentes, in ore dense fimbriatae, laminis obsoletis vel
ad 2 cm longis; inflorescentia 7-8 cm longa, 1—1.3 cm lata, erecta, verticillis
spicularum approximatis; spiculae masculae 5 mm longae, pedicellis 4 mm
longis dense pubescentibus; glumae 2-3 mm longae, 1—3-nerves, acuminatae,
pubescentes, marginibus ciliatis; lomma 5 mm longum, 2.5-3 mm latum,
acutum, dense pubescens; antherae +8, lineares, 2 mm longae; spiculae
femineae uniseriatae, 6 mm longae; glumae acutae, 3-nerves, aequales,
pubescentes; lemma 5mm longum, 3 mm latum, acutum, glabrum. }

Perennial; culms erect from a more or less decumbent rhizomatous base,
glabrous. Sterile culms leafy toward the summit, about 40 cm tall; lower
sheaths much shorter than the internodes, bladeless or the blades much
reduced, the upper ones crowded, keeled at least toward the summit, densely
fimbriate in the mouth, the hairs 10-15 mm long; ligule truncate, 0.5 mm
long; blades 8.5 em long, 1.7—2.7 em wide, rather abruptly acuminate,
rounded at the base, minutely pubescent on the lower surface, scaberulous
above, the margins finely but distinctly serrate, the petiole about 1 mm
long, pubescent. Fertile culms about 30 cm tall; sheaths inflated, all longer
than the internodes, bladeless or sometimes the blade as much as 2 cm
long, 1 cm wide, sparsely to rather densely fimbriate at the mouth; in-
florescence 7-8 cm long, 1—1.3 em wide, stiffly erect, the staminate spikelets
about half overlapping the pedicels of the staminate spikelets in the whorl
above; staminate spikelets 5 mm long, somewhat spreading; pedicels 4 mm
long, densely pubescent; glumes 2-3 mm long, 1-3-nerved, acuminate, pub-
escent, the margins ciliate; lemma 5 mm long, 2.5-3 mm wide, short but
densely pubescent; anthers 2 mm long, pistillate spikelet 6 mm long, the

Frs. 15, 1940 SWALLEN: NEW SPECIES OF PARIANA Oe

glumes acute, 3-nerved, pubescent; fruit 5 mm long, 3 mm wide, acute, gla-
brous.

Type in the U. 8S. National Herbarium, no. 1613672 collected in open
sandy forest between Caxias and Barra do Corda, Maranhéo, Brazil,
February 18-26, 1934, by Jason R. Swallen (no. 3599).

P. sociata is similar in appearance to P. zingiberina Doell, which differs in
having thicker, shorter blades, almost no fimbriae at the mouth of the
sheaths, and smaller glabrous staminate spikelets.

Pariana modesta Swallen, sp. nov.

Perennis; culmi difformes, graciles e rhizomatibus erecti. Culmi steriles
15-35 em alti, in parte superiore foliosi; vaginae glabrae, inferiores inter-
nodiis breviores, superiores internodiis multo longiores in ore fimbriatae;
laminae 3-7.6 cm longae, 5-14 mm latae, lanceolatae, minute pubescentes.
Culmi fertiles 20 cm alti; vaginae internodiis breviores inflatae, glabrae vel
minute pubescentes, laminis obsoletis; inflorescentia 3-4 cm longa, erecta,
verticillis spicularum approximatis; spiculae masculae 5 mm longae, pedi-
cellis 4 mm longis; glumae 3—4 mm longae, 3-nerves acutae vel subacumina-
tae basi latae, pubescentes, marginibus ciliatis; lemma 4.5-5 mm longum,
2mm latum, 3-nerve, acutum, scaberulum vel pubescens; palea acuta lemma
aequans; antherae +20, 1-1.5 mm longae; spiculae femineae 6 mm longae;
glumae aequales, 1—3-nerves, acuminatae, pubescentes; lemma 5 mm lon-
cum, acutum.

Perennial; culms slender, erect from rhizomes. Sterile culms 15-35 cm
tall, leafy toward the summit, glabrous; lower sheaths much shorter than
the internodes, the upper ones crowded, much longer than the internodes,
conspicuously fimbriate at the mouth; ligule very short, thick; blades 3-7.6
em long, 5-14 mm wide, thin, lanceolate, minutely pubescent on both sur-
faces, the pubescence sometimes obscure. Fertile culms 20 cm tall, glabrous
or minutely pubescent; sheaths all shorter than the internodes, inflated,
bladeless, never fimbriate; inflorescence 3-4 cm long, bearing 3 or 4 rather
distant whorls of spikelets, the whorls scarcely overlapping exposing the
pedicels of the staminate spikelets; staminate spikelets 5 mm long, the
pedicels 4 mm long, densely pubescent toward the base becoming nearly
glabrous at the summit; glumes 3-4 mm long, strongly 3-nerved, acute or
subacuminate, broad at the base, more or less pubescent, the margins ciliate;
lemma 4.5-—5 mm long, 2 mm wide, 3-nerved, abruptly acute, scaberulous
and more or less pubescent; palea acute, equaling or slightly exceeding the
lemma; anthers about 20, 1-1.5 mm long; pistillate spikelet 6 mm long, the
glumes equal, acuminate, 1—3-nerved, pubescent, the lemma 5 mm long,
acute, rather broad, almost entirely enclosing the palea.

Type in the U. 8S. National Herbarium, no. 1613666, collected in high
open forest between Caxias and Barra do Corda, Maranh&o, Brazil, Febru-
ary 18-26, 1934, by Jason R. Swallen (no. 3544).

In the region where the type was collected, this species was very abundant,
in some cases being the dominant herbaceous plant in moist open woods
called ‘‘Carasco.”’ The fertile culms were never plainly evident being ob-
scured under the foliage of the sterile culms, the young ones even hidden in
the ground cover of old leaves. Only a small proportion of the plants had
fertile culms at the time the specimens were collected.

DISTRIBUTION: Open forests, east-central Maranhado, Brazil. Between
Caxias and Barra do Corda, Swallen 3597, 3544 (type); Barra do Corda to
Grajahu, Swallen 3631; Carolina to San Antonio de Balsas, Swallen 4132.

78 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

This species was found growing with P. sociata and resembles it in several
ways, especially in having thin blades, numerous fimbriae, and a short erect
inflorescence. It differs, however, in being a much smaller plant with short
narrow blades, short slender inflorescences of distant whorls of smaller
glabrous staminate florets, the glumes of which are very broad at the base,
strongly 3—5-nerved.

Pariana velutina Swallen, sp. nov.

Perennis; culmi difformes basi adscendentes. Culmi steriles 45-50 cm
longi infra nodos dense pubescentes; vaginae inferiores obsoletae, mediae
elongatae internodiis breviores laminis obsoletis, superiores foliatae in-
ternodiis multo longiores, auriculatae, fimbriatae, carina dense pubescente;
laminae 12-16 cm longae, 2—4.6 cm latae, lanceolatae vel ovato-lanceolatae,
acuninatae, infra dense pubescentes, supra glabrae nervo medio dense hir-
tello. Culmi fertiles graciles, 25 cm longi, infra inflorescentiam dense pu-
bescentes; vaginae 2 vel 3, internodiis breviores, inflatae, glabrae; alaminis
obsoletis; inflorescentia erecta 6 cm longa; spiculae masculae 4.5 mm longae,
pedicellis occultis 2-2.5 mm longis pilosis basi villosis; glumae 2—2.5 mm
longae, 1 mm latae, 1—2-nerves, acutae vel acuminatae, scabrae; lemma
4.5 mm longum, 1.5-2 mm latum, 3-nerve, acutum, scabrum, basi pu-
bescens; antherae 2.5 mm longae; spiculae femineae uniseriatae; glumae 5
mm longae, 1—3-nerves, acutae vel subobtusae, pubescentes; fructus 5 mm
longus, 3 mm latus, glaber, superne pubescens.

Perennial; culms biform, erect from an ascending base. Sterile culms
45-50 cm long, densely pubescent below the nodes; lowest sheaths obsolete,
the median one elongate, shorter than the internodes, the blades obsolete,
the upper leaf-bearing ones crowded, much longer than the internodes,
auriculate, fimbriate, densely pubescent on the keel; blades 12-16 cm long,
2-4.6 em wide, lanceolate or ovate-lanceolate, acuminate, softly pubescent
on the lower surface, glabrous on the upper surface, the midnerve densely
hirtellous. Fertile culms slender, 2.5 cm long, densely pubescent below the
inflorescence; sheaths 2 or 3, shorter than the internodes, inflated, glabrous,
the blades obsolete; inflorescence erect, 6 em long; staminate spikelets 4.5
mm long, the pedicels hidden, 2—2.5 mm long, pilose, densely villous at the
base; glumes 2-2.5 mm long, 1 mm wide, 1—2-nerved, acute or acuminate,
scabrous; lemma 4.5 mm long, 1.5—2 mm wide, 3-nerved, acute, scabrous,
pubescent toward the base; anthers 2.5 mm long; pistillate spikelets in
one row; glumes 5 mm long, 1—3-nerved, acute or subobtuse, pubescent;
fruit 5mm long, 3 mm wide, glabrous, pubescent toward the summit.

Type in the U. S. National Herbarium, no. 1458908, collected at Gami-
tanacocha, Rio Mazan, Department Loreto, Peru, altitude 100-125 meters,
by Jose M. Schunke (no. 164).

Pariana trichosticha Tutin is the only other species that has the midnerve
of the blades hirtellous on the upper side but differs from P. velutina in
having glabrous blades, densely pubescent sheaths, and longer glumes on
the pistillate spikelet (7 mm long), which extend beyond the fruit.

Fes. 15, 1940 SWINGLE: ORANGE SUBFAMILY AURANTIOIDEAE 79

BOTANY .—WNew varieties and new combinations in the genera Clau-
sena, Oxanthera, and Triphasia of the orange subfamily, Auran-
tioideae.1 WALTER T. SwWINGLE, U.S. Bureau of Plant Industry.

In connection with the preparation of a synopsis of the orange
subfamily, soon to be published, I have brought to hight a number of
new genera, species, and varieties. I have already published a paper
on my new taxonomic arrangement of the subfamily.”

I have had the good fortune to be able to borrow from several of the
great herbaria of Europe, Asia, and the United States a large number
of specimens of the species of Clawsena and of other genera of the sub-
family Aurantioideae. Thanks to the use on a large scale of the modi-
fied Juel method? of restoring herbarium material of flowers, young
fruits, etc., and making serial microtome sections of them, I have been
able, with the skilled help of Dr. Albert H. Tillson, to throw new light
on some of the taxonomic problems that have arisen in separating the
numerous and often highly variable species of some of the genera of
this subfamily.

Genus CLAUSENA Burm.

Clausena luxurians (Kurz) Swingle, comb. nov.

Clausena Wallichit Oliv. var. luxurians Kurz, Journ. Asiat. Soc. Bengal
44 (2): 183. 1876.

Clausena Wallichii affinis sed differt (1) foliis multo grandioribus, (2) folio-
lis grandioribus et multo paucioribus, (3) floribus minoribus, (4) partes
florales sine cellulis residuiis tanniniferis repletis, (5) alis petiolis duplo lati-
oribus, (6) marginis foliorum integris vel leviter crenulatis nec denticulatis,
(7) foliolis in sicco pallido-viridis, nec fusco-viridis.

A small, meagre shrub, leaves large, 33-50 by 20-27 cm, 5-7- (often 6-)
foliolate, leaflets lanceolate or broadly oval, variable in size, lower lateral
leaflets 6.5-7.5 by 4-5 cm, upper lateral and terminal leaflets 15-21 by 6.5-
9.5 cm, larger leaflets acuminate or broad-acuminate at the apex, variably
cuneate at base and decurrent into the petiolules, lateral veins variable in
prominence, larger ones few, 7-9, arising at angles of 70—75° with the midrib,
the intermediate, fainter and shorter ones often arising at greater angles
(80-89°!), margins entire or slightly crenulate; rachis narrowly winged,
wings 0.8-1.5 mm wide on each side of the rachis; inflorescences terminal,
paniculate; flowers small, 4-5 mm in diameter, 4-(or 5-?) merous, borne on
short pedicels, 1-2 mm long with minute ciliate bracts at the base, calyx
lobes acute, with a large oil gland at the tip (not seen in Fig. 1 but visible in
other sections on the same slide), anthers with a small oil gland in the con-
nective, pistil (including gynophore) 2.5—2.8 mm long, ovary 1—1.1 mm long,

1 Received January 6, 1940.

2 SWINGLE, WALTER T. A new taxonomic arrangement of the orange subfamily,
Aurantioideae. Journ. Washington Acad. Sci. 28: 530-533. 1938.

3 See SwINGLE, WALTER T. New methods utilized in studying the taxonomy of the
orange subfamily. Journ. Washington Acad. Sci. 29: 270. 1939.

80 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

1—1.8 wide, with one medium-sized protuberant oil gland at the tip of each
locule, style 1.3 mm long, cylindrical, 0.5—0.6 mm in diameter, except at the
base, which is 0.3 mm in diameter, where it joins the ovary; stigma not de-
limited from the style; fruits (immature) globose, 5-6 mm in diameter.

Type specimen.—Burma, rare in eastern slopes of Pegu Yomah in tropical
forests, Sulpiz Kurz 1995, flowering branches, Herb. Roy. Bot. Gard. Cal-
cutta; photographs and serial sections, 8S. & T. Nos. 519 A, slides 1—4 (168
cross sections of a flower bud), 519 B 1, and 519 C 1 (95 longitudinal sections
of 2 pistils), Herb. National Arboretum.

Fig. 1.—Clausena luxurians: Serial longitudinal sections
of pistil of type specimen. X10

Cotype specomen.—Same locality, Kurz 1995, ‘‘Choungmerah Chg.,”’ Herb.
Roy. Bot. Gard. Calcutta; photographs and serial microtome sections, 8. &
T. Nos. 520 A, slides 1-4 (198 cross sections of 1 flower bud), 520 B 1, 520
C 1 (76 longitudinal sections of pedicel, calyx, and pistil), Herb. National
Arboretum.

Remarks.—The original description of Clausena Wallichit var. luxurians
by Kurz is very short; it reads, in full: “Clausena Wallichit Oliv. Var. 6
luxurians, rachis leafy-winged; leaflets only 4-2 pairs with an odd one, 4-8
in. long, remaining green in a dried state.”

The leaves of this species differ markedly from those of C. Wallichii in hav-
ing much larger, broader, and fewer leaflets (5—7 instead of 13-17!). On the
other hand, the flowers of C. Wallichit (which has much smaller leaflets) are
decidedly larger, the pistil being from one-fifth to one-quarter longer and
the style nearly two-thirds longer!

Another species, C. Guillauminit, native to French Indochina, resembles
C. luxurians and C. Wallichit in having leaves with narrowly winged rachis
and also has flower parts not greatly different from those of C. Wallichiz; itis,
however, strikingly different from both of them in having dimorphic oil
glands in the leaves, many very small ones, and a few very large ones, 4 to 3
mm in diameter, clear bright red by transmitted light. These three species
are doubtless descended from a common ancestral form but have diverged
so much in taxonomic characters that they are now good, easily distinguish-
able species.

One of the most striking characters of C. luxurians, in which it differs
markedly from all species at all closely related to it, is the almost complete
absence in the flower parts of tissue cells filled with brownish, nearly opaque
tannin residues that occur in abundance in all the other species of Clausena
except C. suffruticosa, a very aberrant species; C. dentata var. longipes has

Frs. 15, 1940 SWINGLE: ORANGE SUBFAMILY AURANTIOIDEAE 81

such cells but only a few of them; note the contrast between Figs. 1 and 2 in
this respect.

C. luxurians in its general aspect, flower morphology, and especially in its
leaf characters, somewhat resembles C. heptaphylla of British India, Burma,
Indochina, and Malaya, but the leaves of this latter species do not have a
winged or margined rachis and have floral organs with abundant cells filled
with tannin residues. The many-sided affinities of this newly recognized spe-
cies are characteristic of the situation found among the many closely related
species of this large genus.

Fig. 2.—Clausena Wallichii: Serial longitudinal sections of pistil. Burkzill
30288, Amherst Distr. Burma, Herb. Bot. Gard. Calcutta. X10.

Clausena dentata var. Henryi Swingle, var. nov.

A typo differt (1) fructu ovoideo nec sphaerico, negro, (2) foliis foliolisque
duplo majoribus hirsutioribus, (3) petalis hirsutis, (4) foliolis lateralibus su-
perioribus saepe grandioribus quam folioli terminales.

Type specimen.—China, Ichang, Henry 4122, flowering branch,- May
1888, Herb. Kew Gardens; photograph and serial microtome sections, 8. &
T. Nos. 528 A, slides 1-3; 528 B, slides 1-3 (381 cross sections of flower bud) ;
628 C 1 and D 1 (85 longitudinal sections of two pistils); Herb. National
Arboretum.

Cotype specimen 1.—China, Ichang, Henry 3127, February 1887, fruiting
branch, Herb. Kew Gardens; photograph Herb. National Arboretum.

Cotype specimen 2.—China, Ichang, Henry 13028, Herb. Kew Gardens;
photograph and serial microtome sections, 8. & T. No. 538 A, slides 1-3,
B 1-8. (836 cross sections from 2 flower buds); C 1, D 1 (105 longitudinal
sections from 2 flower buds); Herb. National Arboretum.

Serial microtome sections made from a flower bud of the type specimen
(Henry 4122) show the flowers to be tetramerous; the anthers show small oil
glands in the connective, obscured by dense tannin residues in the surround-
ing cells; total length of pistil from calyx to tip of stigma 4-43 mm, gyno-
phore hourglass-shaped, 0.6-0.7 mm high, ovary 1-1.2 mm high, 1.1-1.8 mm

|

82 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

wide, with a minute, slightly protuberant oil gland at the top of each locule,
style 1.2-1.8 mm long, subclavate, 0.5—-0.6 mm wide at base where it joins
the ovary, gradually expanding toward the tip where it is 0.7—-0.8 mm wide;
stigma not clearly delimited.
. Remarks.—This striking variety of C. dentata (Willd.) Roemer has been
identified wrongly as C. suffruticosa (Roxb.) Wight, which is a very different
plant which has been confused with Henry’s Clausena doubtless because of its
rather slender ovoid fruits. The fruits of C. dentata var. henry1, 9-12 by 6-8
mm, black when ripe and said to be edible, are very different from the
fruits of the typical C. dentata of British India, which when ripe are globose,
whitish green, and about 8-12 mm in diameter. Henry’s Clausena has the
northernmost range yet reported for any species of the genus. It grows near
Ichang, Hupeh Province, China, about latitude 31° N. Many varieties of C.
dentata have been described, but most of them are still inadequately known.
Another Chinese variety of this species, C. dentata var. robusta Tanaka, is a
vigorous plant having rather small globose fruits, 7-8 mm in diameter.

Still another Chinese variety very inadequately described and named by
Léveillé as a distinct species, C. Dunniana Lev., can better be considered a
variety of C. dentata.

Clausena dentata var. Dunniana (Lev.) Swingle, comb. nov.

Clausena Dunniana Léveillé, in Fedde’s Report. 10:67. 1912.

Type specimen.—Pin-fa, Kweichow Province, China, J. Cavalerie 1072,
Herbarium of Edinburgh University; photograph in Herb. National Arbo-
retum.

Cotype specumen 1.—Same locality, J. Cavalerie 1072, June 18, 1903, flow-
ering branch, Herb. Kew Gardens.

Cotype specimen 2.—Same locality, J. Cavalerie 1072, flowering branch,
Herb. Univ. Edinburgh; photograph and fragments ex Herb. Monseigneur
Léveillé, in Herb. Arnold Arboretum.

This plant is still very little known, as Léveillé’s description was very
short. It looks much like some specimens assigned to Clawsena dentata var.
robusta Tanaka, and more study is needed to separate these two poorly
characterized varieties. Some specimens of C. dentata var. Dunniana have pet-
iolules 4-6, rarely 8, mm long, longer than have been seen on any other
Chinese forms of C. dentata.

Clausena brevistyla var. halmaheirae (Miq.) Swingle, comb. nov.
Clausena halmahetrae Miquel, Ann. Mus. Bot. Lugd.-Bat. 1: 211. 1863.

This variety has diminutive pistils, only 1.4-1.5 mm long, with the gyno-
phore only 0.2—-0.3 mm long, the ovary 0.8 mm long and 0.8 mm wide, rough
with oil glands, the style 0.5-0.6 mm long and cylindrical. The inflorescences
are large, 25-35 cm long, with widely spreading primary branches and sturdy
terminal pedicels, 2-4 mm long.

The above measurements were taken from serial microtome sections made
from a topotype specimen from Halmaheira Island (Teysmann 5667) in
Herb. Roy. Bot. Garden Calcutta.

The styles are slightly longer than those of C. brevistyla but otherwise
similar.

The southernmost species of Clausena, C. brevistyla Oliver, native to
northeastern Australia, has, as its name indicates, a very short style, less
than one-half as long as the ovary. It now appears that a species described
from Halmaheira (or Gilolo) Island in the Moluccas, C. halmahetrae Miq..,

Fes. 15, 1940 SWINGLE: ORANGE SUBFAMILY AURANTIOIDEAE 83

also has a very short style and doubtless belongs close to C. brevistyla and
can best be made a variety of it.

Genus OxXANTHERA Montr.

Oxanthera is a genus of the Wenzelia group of the subtribe Triphasiinae,
tribe Citreae. It is limited to New Caledonia and with the species here trans-
ferred to the genus has four species with large orangelike flowers but with no
pulp-vesicles in the fruit. This excludes them not only from the genus Citrus
but from the whole subtribe Citrinae.

Oxanthera undulata (Guill.) Swingle, comb. nov.
Citrus undulata Guillaumin, Bull. Soc. Bot. France 85: 304. 1938.

This species has thin leaves unlike the other three, which have thick, more
or less coriaceous leaves. This seems to indicate they are more or less pro-
nounced xerophytes, whereas C. undulata is more likely a mesophyte.

All the species of this very curious genus are discussed in some detail in
my synopsis of Citrus and related genera of the orange subfamily, now in
press.

Genus TRIPHASIA Lour.

The genus T'riphasia is a small one that has been known to botanists for
several centuries and is the type of the subtribe Triphasiinae of the tribe
Citreae. The best-known species is Triphasia trifolia with trifoliate leaves,
small white flowers, and tiny red berries, often grown in hothouses and in
subtropical gardens as an ornamental plant.

Recently an autotetraploid form of this species was discovered in the
U.S. Bureau of Plant Industry citrus greenhouse.

Triphasia trifolia var. tetraploidea Swingle, var. nov.

A typo differt (1) chromosomatis duplo numerosioribus in omnibus cellulis
(autotetraploideis), (2) foliis paulo crassioribus, floribus paulo grandioribus.

Type specitmen.—Washington, D. C., citrus greenhouses, U. 8. Bureau of
Plant Industry, Swingle C.P.B. No. 2889, P.E.I. 27778, fruiting branch,
S. & T. No. 575, slide 1, numerous pollen mother cells showing tetraploid
chromosomes, made by Dr. A. E. Longley, March 3, 1937; Herb. National
Arboretum, No. 70839.

Remarks.—This variety differs from the species in having larger flower
parts and thicker leaves, and especially in having 36 chromosomes in all the
somatic cells instead of 18, the normal number for almost all the plants of
this orange subfamily so far studied.

This autotetraploid form of the common trifoliate limeberry looks much
like tetraploid forms of Citrus, having, like them, thicker leaves and larger
flower parts. Longley,* who has studied the chromosome numbers of Cztrus
and many related genera, first detected the tetraploid nature of this plant.
This variety doubtless originated as a mutation from the common diploid
form of the species. This tetraploid form of the common limeberry is a vigor-
ous plant that flowers and fruits freely in the greenhouse. It may prove to
exe value in ornamental plantings both in the hothouse and in subtropical
gardens.

*Lonetey, ALBERT E. Polycarpy, polyspory, and polyploidy in citrus and citrus
relatives. Journ. Washington Acad. Sci. 15: 347-351, 1 fig. 1925.

84 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 2

ZOOLOGY .—A new pycnogonid from Pescadero, Calif., and distribu-
tional notes on other species.| JonL W. HEDGPETH. (Communi-
cated by Waxpo L. ScHumiIrT.)

The pebble beach near Pescadero, Calif., is not a good collecting
ground for marine invertebrates, since the reef is somewhat like a sea
wall, but it is the only habitat so far known of a hitherto undescribed
species of Ammothea. A specimen of this pycnogonid was collected in
September 1930, and when the beach was revisited in August 1939
three more specimens were taken from the same small area of the reef
at the south side of the pebble beach cove. The recent collection was
made from worn eelgrass holdfasts just below the Littorina scutulata
horizon at a 2.8 tide (about the upper limit of zone 3 of the classifica-
tion of Ricketts and Calvin, 1939), which is a rather high distribution
for littoral pyecnogonids on the California coast and supports Giltay
(1934) in his contention that the genus Ammothea is composed of
littoral forms adapted to various biotopes and that a large number of
species are still to be discovered in regions not yet carefully investi-
gated for pyenogonids.

Ammothea euchelata, n. sp. Fig. 1

Holotype.—Male, with egg clusters. Pescadero, Calif., August 15, 1939.
U.S.N.M. no. 78411.

Paratypes.—One male, in U.S.N.M. collections; egg-bearing male, dis-
sected, in author’s collection.

Description.—Body circular, about 1.5 mm in diameter, lateral processes
contiguous but not fused, with low tubercles on the anterior and posterior
distal corners of the processes. Proboscis ovoid, directed ventrally, not
quite so long as trunk. Eye tubercle near anterior border of first trunk seg-
ment, low and without spines or projections; eyes large and distinct. Abdo-
men moderately long, about twice the length of the lateral processes of the
last pair of legs, with a few small spines near the apex. Chelifores nearly
as long as the proboscis although apparently a little over half as long; scape
stout, little more than half the length of the chelifore; chelae well developed,
subglobular, the fingers broadly curved so that only the tips oppose, with-
out teeth or setae. Palpus slightly longer than proboscis, first joint short,
about equal to third joint; second joint slightly longer than first and third
combined; fourth joint about equal to second; last four joints short, more or
less equal, together a little longer than the fourth; joints 5, 6, and 7 with
ventral processes, terminal joint oval, a few setae on the ventral processes
and on the distal portion of the terminal joint. Oviger of male about twice
as long as palpus, first joint short, stout; second joint three times as
long as wide; third joint slightly longer than second but half the diameter;
joints 4 and 5 equal, shorter than the third; joint 6 one-half as long as 5,
joints 5 and 6 with spines on outer margin directed backward; joints 7, 8,
and 9 nearly equal, 7 and 8 stout, 9 slender; terminal joint short and blunt.
The last four segments of the oviger bear a spine with teeth on its inner

1 Received September 20, 1939:

Frs. 15, 1940 HEDGPETH: A NEW PYCNOGONID 85

Fig. 1.—Ammothea euchelata, n. sp.: a, Front view of holotype, X30; b, dorsal view
of trunk of holotype, X80; c, ventral view of right chelifore of paratype, X45; d,
right palpus of paratype, X45; e, right oviger of paratype, X45; f, detail of terminal
joints of oviger, X120; g, denticulate spine from oviger, 1,000; h, right third leg of
paratype, X33. (All drawings made with the aid of a camera lucida; magnifications
approximate.)

86 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

margin at the ventral base of the joint. In the terminal segment this spine
is sickle-shaped, situated near the distal ventral margin, and on the slide
mount obscures a minutely lobed leaflike terminal spine. In Fig. 1, f, this
spine has been drawn away to reveal the structure of the terminal spine.
Eggs about 0.1 mm in diameter, in small scattered masses of 12—20 along the
oviger. Legs stout, comparatively short; first and second coxal joints armed
with spines laterally; genital protuberance of second coxa of male very
conspicuous; third coxa and remainder of leg with a few scattered spines.
Tarsus curved, terminal claw strong, nearly as long as tarsus, auxiliary
claws about half as long as terminal claw. Proportions of the joints in milli-
meters approximately as follows: 0.5, 0.5, 0.5, 1.0, 1.0, 0.8, 0.25, 0.5. Meas-
urements (approximate) as follows:

Holotype Paratype (from dissected parts)
Bodiystn ms sede io) mam 1.5mm
IPTODOSCIS!.. seer 1.25 1.0 |
Abdomen...... — 0.6
Chelifores.;.---) £40 1.0 (scape 0.6, chela 0.4)
Palpusiic. ch ee Ie)
Ovigerie meso. — 2.0
Third leg....... 2.5 (fromabove, unextended) 5.0 (sum of segments)

Relationships.—No other species of Ammothea known to me has such
large and well-developed chelae in the adult as A. euchelata, and this char-
acter separates it from the rest of the California species, in which the chelae
are rudimentary knobs in the adult. The ventral protuberances of the ter-
minal segments of the palpus are similar to those of Ammothea pribilofensis
Cole, which has a higher, pointed eye tubercle and a broader, blunter pro-
boscis than A. euchelata. Denticulate spines occur only on joint 10 of the
male oviger of A. pribilofensis. Ammothea euchelata is evidently related to
A. spinosa (Wilson), in which the chelifores are sometimes chelate. They
are much smaller, however, and the legs as a whole are spinier. The ter-
minal joints of the palpus of Ammothea spinosa do not have the ventral pro-
jections of A. euchelata, and the denticulate spines of the oviger are sym-
metrical and the proportions of the joints themselves different in A. spznosa.?

Distribution.—Although I have collected intensively for pyecnogonids at
various reefs along the central California coast from Marin: County to
Pacific Grove, I have so far collected this species only at Pescadero. On
September 30, 1930, one egg-bearing male was collected from Bugula along
with a specimen of Ammothea gracilipes, apparently the southern record for
the latter, which is most abundant just north of the Golden Gate.

DISTRIBUTION NOTES

Moss Beach, San Mateo County, Calif—Ammothella tuberculata Cole,
five specimens, from eelgrass holdfasts and among hydroids on pitted rocks,
2-11-34. Lecythorhynchus marginatus Cole, four specimens, 4-23-32; 12
specimens, including egg-bearing males, on Abietinaria, 2-10-34; six speci-
mens, 11-6-38. Tanystylum intermedium Cole, three specimens, 2—10—34;
three specimens, on Abzetinaria, 11-6-38. Phoxichilidium femoratum
(Rathke), 10 specimens, including egg-bearing males, 4-23-32; three speci-
mens, 2-10-34. Pycnogonum stearnsi Ives, one female, under rock, 3-26-82.

Muir Beach, Marin County, Calif—Ammothea gracilipes Cole, 12 speci-
mens, including egg-bearing males, from Bugula, Abietinaria, and pitted
rocks, 5-11-33. Ammothella tuberculata Cole, one specimen, 5-23-32; three

2 Ammothea acheloides Wilson (Pl. 5, Fig. 1) =A. spinosa, auctoritate Schimkewitsch.

Frs. 15, 1940 KENT: A NEW TREMATODE 87

specimens, 5-11-33; one specimen, April 1934. Lecythorhynchus marginatus
Cole, 12 specimens, including egg-bearing males, from Abzetinaria, 5-11-33;
12 specimens, including egg-bearing males, from Abzetinaria, April 1934.
Tanystylum intermedium Cole, 14 specimens, including egg-bearing males,
from Abietinaria, April 1934. Pycnogonum stearnsi Ives, three males from
Abietinaria, April 1934.

Jenner, Sonoma County, Calif. (near Bodega Bay).—Ammothea gracilipes
Cole, one female, and Pycnogonum stearnsi: Ives, one female, both from
Campanularia, collected by A. R. Grant, February 1934.

It will be noted in the Moss Beach collections that in 1982 Phoxichilidium
femoratum was common and Lecythorhynchus marginatus was scarce, but
that the situation was reversed in 1934. Unfortunately it was not possible
to make regular collections to determine if this is a case of periodicity or
simple coincidence. At Muir Beach, Lecythorhynchus marginatus and Tany-
stylum intermedium were collected from the same fronds of Abietenaria in
equal numbers. These two species and Pycnogonum stearnst occur in the
Abietinaria along the sheltered rock wall at the north border of the beach,
while Ammothea gracilipes occurs in hydroids and bryozoan colonies among
the maze of large boulders that marks the south border of the Muir Beach
cove. There is some variation in the length of the protuberance on the dorsal
part of the second coxal joint in this species, in some specimens being as
long or longer than the coxal segment. There was no variation observed in
the direction of increased body size and reduction of these protuberances
as described by Lozina-Lozinsky (1933) in his variety borealis of Ammothea
gracilipes from the Gulf of Kastri.

LITERATURE CITED

Coz, L. J. Pycnogonida of the west coast of North America. Harriman Alaska Ex-
pedition 10: 249-298, 16 pls. 1904.

Gittay, L. Remarques sur le genre Ammothea Leach et description d’une es péce nouvelle
dela mer Irlande. Bull. Mus. Roy. Hist. Nat. Belgique 10(18): 1-6, 3 figs. 1934.

Lozina-Lozinsxy, L. K. Pantopoda vostochnykh morei S. S. S. R. Gosudarstvennyi
Gidrologicheskii Institut Issledovaniia morei 8. S. S. R. 17: 438-74. 1938.

Ricketts, EH. F., and Catvin, Jack. Between Pacific tides, xxii+320 pp., 46 pls.
1939. Stanford University Press.

Witson, HE. B. Synopsis of the Pycnogonida of New England. Trans. Connecticut
Acad. Arts and Sci. 5: 1-24, 7 pls. 1882.

ZOOLOGY .—A new trematode from Siren lacertina: Diplostomulum
sirenis, n. sp.1. GEORGE C. Kent, Jr., Vanderbilt University.
(Communicated by Pau BARTSCH. )

A number of Siren lacertina (Linnaeus), collected from marshes
along the banks of the Cumberland River, near Nashville, Tenn., in
October 1938, were kept in an aquarium until May 1939, when the
animals were sacrificed for routine histological examination. Of four
specimens examined (all that were available), three (length approxi-
mately 25 cm) were found to be infected with a trematode, herein
described as a new species of the genus Diplostomulum. Over 100
specimens were obtained from one host, 50 from a second, 6 days

1 Received October 27, 1939.

88 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 2

later, and 30 from a third, 13 days after the first animal was sacri-
ficed. The fourth animal (14 em) did not appear to be infected. The
Diplostomula were not encysted but were free in the pericardial
cavity, where they were clumped in a white mass about the ventricle.
Inspection of the alimentary tract and of the great vessels revealed
no similar forms in these organs.

Materials and methods.—The parasites were collected with a pen
point previously dipped in black India ink on which background the
white specimens were easily visible, and with which instrument they
were more easily separated from the slimy exudate than would have
been possible with a pipette. They were transferred to a watch glass
containing physiological saline at room temperature. Specimens to be
permanently mounted were fixed with acetic-sublimate, or formalin-
alcohol-acetic acid. For staining, Harris’s hematoxylin, borax-car-
mine, and hematoxylin and eosin were employed in various dilutions,
and for periods of 10 minutes to 48 hours. When necessary, acid
alcohol was used for destaining. Undiluted Harris’s hematoxylin
applied 10 minutes gave the most satisfactory differentiation.

Diplostomulum sirenis, n. sp.

Description.—General form and anatomy typical. Forebody and hind-
body distinct. Forebody in living specimens thin, foliaceous, elongated,
slightly concave ventrally, which concavity is retained in many preserved,
unmounted specimens. Hindbody short, located posterodorsally on fore-
body. Oral sucker terminal, mouth ventroterminal. Acetabulum situated
anteriorly in posterior half of body, circular in outline in living specimens,
transversely elongated in stained mounts as result of longitudinal con-
traction of entire body.

Hold-fast organ prominent in stained specimens. Lateral suckers short,
broad prominences in living and preserved specimens.

Pharynx long, prepharynx short, esophagus longer than prepharynx.
Caeca slim, containing, in living, active forms, a dark, granular substance,
which is moved slowly backward and forward in the caeca as if by peristaltic
waves, and from the presence of which the dilation of the posterior termina-
tion of the caeca may be observed.

Fundament of reproductive system a darkly staining, single or irregularly
lobulated mass, situated in midline at posteriormost extremity of forebody
in preserved specimens.

Urinary bladder occupies greater part of hindbody, with pair of saclike
anterior diverticula terminating as separate lobes at a body level immedi-
ately posterior to hold-fast organ. In stained sections these appear as 1so-
lated vesicles occupying a position between the hold-fast organ and the
reproductive fundament. Two or three diverticula also extend laterad from
the urinary bladder. Bladders discharge through a short, common duct and
dorsoterminal pore, which contracts and relaxes rhythmically. Anteriorly,
the urinary bladder proper communicates on either side with the lateral
tubular portion of the reserve bladder. The remaining reserve bladder sys-
tem is essentially the same as described by Hughes (1929). Numerous short

Fes. 15, 1940 KENT: A NEW TREMATODE 89

branches of the common collecting trunks, and of their tributaries, end as
blind, ellipsoidal vesicles containing, in living and in certain preserved speci-
mens, an ellipsoidal, highly refractile, calcareous concretion. Flame cells
appear in pairs.

Living specimens average 0.825 mm extended, without pressure. Measure-
ments on 16 specimens mounted in toto are given in Table 1. The 16 speci-
mens were chosen at random from those subjected to one of each of the
different fixatives and stains employed.

TABLE 1.—MEASUREMENTS (IN MILLIMETERS) OF 16
MouNTED SPECIMENS OF Diplostomulum sirenis

Measurement Minimum Maximum Average

Body:

Mem ste: ie ee Se 0.370 O2507. 0.482

“VAGUE: Se GP erate ine ieee eee 0.192 0.281 0.243
Oral sucker

1 TERE] OY a le re 0.044 0.061 0.051

Milligan ei. et 0.037 0.047 0.042
Pharynx: .

BPC ee eo a 0.024 0.051 0.041

Wirteli ine ets tec ee Sods 0.013 0.029 0.021
Acetabulum

BSN oe St. LS si 0.034 0.062 0.048

VIVE 4 eee er - 0.047 0.078 0.065
Hold-fast organ

LCE ee Slee ea eee 0.068 0.119 0.092

Widtt 2... 25. ogee age Bale 0.047 0.081 0.070

Behavior.—In physiological saline, the animals move by attaching the
oral sucker to the substrate and contracting the body. The ventral sucker
next becomes attached, the oral sucker relaxes, and the anterior portion of
the body stretches forward. Progress is slow. When suspended in the liquid,
the movements differ in that the posterior portions of the body are al-
ternately flexed on the sides of the body, and extended, without progress
forward in space. Light from beneath results in writhing movements.

Host.—Siren lacertina (Linnaeus).

Habitat.—Pericardial cavity, in masses attached near ventricle.

Locality —Marshes of Cumberland River, near Nashville, Tenn.

Type specimen.—U. 8S. N. M. Helm. Coll. no. 9284, mounted in toto.

Discussion.—The method and duration of fixation and staining have far-
reaching effects on the size and appearance of the mounted specimen, es-
pecially in the relative size of individual structures. Lateral suckers may, in
the same species, be either inverted as cuplike depressions, or extended as
protuberances. The acetabulum becomes elongated transversely. The hind-
body especially becomes contracted, and the urinary bladder occupies an
abnormal position in stained specimens. Therefore, great care should be ex-
ercised in the setting up of new species on the basis of minor measurement
differentials, the condition of the lateral suckers, or of other superficial char-
acters, in stained mounts. Partially to offset these variations, the measure-

90 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 2

ments in Table 1 were made on specimens subjected to one of each of the
several different techniques employed in this study.

Fig. 1.—Diplostomulum sirenis, n.sp.

ac Acetabulum. ls Lateral sucker.

cc Calcareous corpuscle. oe Esophagus.

ce Caecum. os Oral sucker.

di Bladder diverticulum. ph Pharynx.

ep Excretory pore. pph Prepharynx.

gr Granular material in caecum. rf Reproductive fundament.
ho Hold-fast organ. ub Urinary bladder.

lc Lateral collecting vessel.

D. sirenis resembles five other forms found in North America [D. browni
Hughes (1929), D. gigas Hughes and Berkhout (1929), D. huronense (La
Rue), described by Hughes and Hall (1929), D. tritur7 Kelley (1934), and D.
ambystomae, described by Rankin and Hughes (1937)]. D. szrenzs differs from
D. brownt and D. huronense in being larger; from D. ambystomae in being
consistently smaller in the stained condition, and longer in living specimens,
body extended, without pressure. It most nearly approaches D. gigas in
relative size of structures, stained. D. szrenis differs from all these species in
the presence of paired, anterior diverticula of the urinary bladder, which lie
just posterior to the hold-fast organ. In this respect it resembles D. triturz
but differs from the latter in relative measurements, which, as pointed out
above, should not constitute the sole basis for introducing a new species. The
most notable difference in these two forms is the presence of an additional
series of conspicuous outpocketings of the reserve bladder in D. étriturt, re-

Fes. 15, 1940 KENT: A NEW TREMATODE 91

sulting in a condition of complexity of this apparatus. Two or three similar
outpocketings only are found in D. sirenis, and these are confined to the
urinary bladder proper. D. sirenis is, therefore, intermediate in complexity
of the reserve bladder system between D. gigas, which is without outpocket-
ings, and D. triturz, which has many.

Finally, it is probable that the occurrence of D. sirenis in the pericardial
cavity of Siren is a physiological character of specific importance, since T'r7-
turus viridescens spends four years strictly on land (Reinke and Chadwick,
1939), while Siren lacertina evidently remains, except for short excursions
in underground burrows, in water where its food supply exists, and in which
medium it may best respire.

Whether the hosts were infected before being placed in captivity is prob-
lematical. Three large Sirens, examined several months previously, did not
appear to be infected. Opportunities were offered for infection at intervals
during the latter two months of captivity, when snails and insect larvae were
transferred to the aquarium from marshes where Siren are known to exist.
Since the latter have not been observed to eat snails, infection may have
arisen from one of at least two possible sources: (1) From the insect larvae,
by ingestion; (2) from the snails, by swimming of the cercaria in the water,
and entrance into the host via the skin (which is very thick), or via the gills
or pharynx. Until D. sirenis may be isolated from the blood stream it seems
logical to assume that the metacercaria found in the pericardial cavity prob-
ably are transmitted to the succeeding host when Szren is eaten by another
vertebrate. Since the latter is often prey to snakes, the final host may there-
fore be a reptile, although the possibility of Siren being devoured by a bird
may also exist.

A study of the life history of each of these closely related forms is indi-
cated, to ascertain their exact taxonomic relationships.

LITERATURE CITED

Hugues, R. C. Studies on the trematode family Strigeidae (Holostomidae) No. XIV.
Two new species of Diplostomula, Occ. Pap. Mus. Zool. Univ. Michigan 202.
1929.

Hueues, R. C., and Berxuout, P. G. Studzes on the trematode family Strigeidae
(Holostomidae) No. XV. Diplostomulum gigas, sp. nov. Pap. Mich. Acad. Sci.,
Arts, and Lett. 9: 483-488. 1929.

Huaues, R. C., and Haut, L. J. Studies on the trematode family Strigeidae (Holo-
stomidae) No. XVI. Diplostomulum huronense (La Rue). Pap. Mich. Acad. Sci.,
Arts, and Lett. 9: 489-494. 1929.

Kewiey, R.S. The trematode parasites of Triturus viridescens viridescens (Rafinesque).
Univ. Pittsburgh Bull. 31(10): 201-210. 1934.

RanKIN, J. S., and Hueuss, R. C. Notes on Diplostomulum ambystomae, n. sp.
Trans. Amer. Micr. Soc. 56: 61-66. 1937.

Reinke, E. E., and Cuapwick, C. 8. Inducing land stage of Triturus viridescens to
assume water habitat by pituitary implantations. Proc. Soc. Expl. Biol. and Med.
40: 691-693. 1939.

Obituary

WALDEMAR LINDGREN, professor emeritus of geology at the Massachu-
setts Institute of Technology, died on November 3, 1939. He was born at
Kalmar, Sweden, on February 14, 1860. He became interested in geology
and mining at an early age and in 1882 was graduated from the Mining
Academy at Freiberg, Germany, with the degree of Mining Engineer. In
1884 he began work with the United States Geological Survey, with which
he maintained connection for 31 years. In 1905 he was made chief of the
Division of Mineral Resources of the Geological Survey, in 1908 was placed
in charge of the Section of Metalliferous Deposits, and in 1911 was ap-
pointed Chief Geologist of the Survey. In 1912 he was appointed William
Barton Rogers Professor of Economic Geology, and head of the Department
of Geology, at the Massachusetts Institute of Technology. His book
“Mineral Deposits,” the first edition of which appeared in 1913, became the
leading text and reference on this subject throughout the world. Other pub-
lications include about 185 titles. 7

While chairman of the Division of Geology and Geography of the National
Research Council (1927-28) Dr. Lindgren established the Annotated Bib-
liography of Economic Geology, to which he contributed (with a reading
knowledge of at least eight languages) about 2,500 abstracts in a 10-year
period. He was one of the founders of the Journal of Economic Geology (1905
and served as associate editor.

Dr. Lindgren received many honors in recognition of his achievements.
Princeton (1916) and Harvard (1935) conferred honorary degrees, and the
Society of Economic Geologists, l’Association des Ingénieurs de Liége, the
Geological Society of America, and the Geological Society of London award-
ed him their medals. The American Institute of Mining and Metallurgical
Engineers issued in tribute to him the Lindgren Volume on Ore Deposits of
the Western States (1933). He was chosen honorary chairman of the Six-
teenth International Geologic Congress held in the United States in 1933.

Dr. Lindgren held membership in the Washington Academy of Sciences,
the National Academy of Sciences, the American Philosophical Society, the
American Association for the Advancement of Science, the American In-
stitute of Mining and Metallurgical Engineers (honorary life member), the
Mining and Metallurgical Society of America (president, 1920), the Geo-
logical Society of America (president, 1924), the Society of Economic Geolo-
gists (president, 1922), the Geological Society of Washington, and the
Geological Society of Boston. He was a foreign member of the Royal Acad-
emy of Sciences of Sweden, the Academy of Engineering of Sweden, the Geo-
logical Society of London, and the Academy of Sciences of Leningrad. He
was a corresponding member of the Canadian Institute of Mining and Metal-
lurgy and the Geological Society of Stockholm and an honorary member of
the Geological Society of Belgium.

In 1886 Dr. Lindgren married Ottolina Allstrin of Gothenburg, Sweden,
who died in 1929. Lindgren’s stimulating and forceful influence on geologic
thought is tempered to all who knew him personally by his sympathetic con-
sideration to the viewpoints of others and his kindly and unemotional man-
ner in objections. He possessed an engaging personality and keen sense of
humor, and together with his great capacity for accomplishment in his vo-
cation he maintained broad interests in history, art, languages, and world
politics.

92

§
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B tok

CONTENTS |

Botany.—Naming molds. CHARLES THOM.....................05.

CHEMISTRY.—Anselme Payen, distinguished French chemist and
pioneer investigator of the chemistry of lignin. Max PHILuips.

Botany.—Ejight new species of Parzana. JASON R. SWALLEN.......

Botany.—New varieties and new combinations in the genera Clausena,
Oxanthera, and Triphasia of the orange subfamily Aurantioideae.
Wavren f) Swett CMY eee tae

ZooLocy.—A new pycnogonid from Pescadero, Calif., and distribu-
tional notes on other species. JonL W. HEDGPETH.............

ZooLtocy.—A new trematode from Siren lacertina: Diplostomulum
sirenis, n. sp. Grorer C. KENT, JR... 000. 2) 4) 0 ee

OBITUARY: WALDEMAR’ LINDGREN) (03. 00. 0 Oe a BE OTE &. 0.

This Journal is Indexed in the International Index to Periodicals

Page

87

92

Se ee ee eC

SS i Eta RAT pee eR age 8 - Ate cama & eee

bE AER ET Nae a a 5

ate

a manta Mi celtinn se TB te SPE PT

No 3

IENCES

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JOURNAL

OF THE
WASHINGTON ACADEMY OF SCIENCES

VoL. 30 Marcu 15, 1940 No. 3

GEOLOGY .—Placer gold in Alaska.! J. B. Mertis, Jr., U. 8. Geo-
logical Survey.

The study of metalliferous placers, if pursued in its many aspects,
is a very comprehensive one, because it leads directly into several
branches of geology and indirectly into the fields of mineralogy,
metallography, inorganic and physical chemistry and allied topics. A
lode of some sort must necessarily have existed before a placer could
be formed. Therefore the local bedrock geology is of primary interest,
and its investigation calls for petrographic work and possibly the
study of ore deposits. But usually neither the rocks nor the ores now
exposed at the surface are exactly like those that existed at the time
when the placers were formed, so that an understanding of surficial
geologic processes is also required not merely to interpret the pre-
existing bedrock geology, but more particularly to decipher the his-
tory and mode of accumulation of the placers. Yet physiography and
the related surficial earth sciences do not always suffice, because the
metals contained in many placers were liberated from their bedrock
sources long before the present surficial features originated. Hence
stratigraphy and paleontology not infrequently constitute another
part of the investigation. It is hardly necessary to add that structural
geology is likewise an essential topic, because the structure of bedrock
is an important factor in all geomorphic interpretations. And finally,
certain of the more specialized branches of geology and geophysics
may enter into the investigation.

By these remarks it is not intended to imply that the writer has
made any such comprehensive studies. Unfortunately many reasons
exist why the gold placers of Alaska have not received such treatment,
and not the least of these is that the time allocated to such work, both
in the field and in the office, has been inadequate for investigations of
such an extended scope. Nevertheless, even casual field examination
and limited laboratory work, when extended over a considerable pe-

_ | Address of the retiring president of the Geological Society of Washington, de-
livered December 13, 1939. Published by permission of the Director, Geological
Survey, United States Department of the Interior. Received January 12, 1940.

| 93
warn 15 1940

94 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 3

riod of years, lead to the formulation of more or less definite ideas;
and, though such ideas merely point the way toward future investiga-
tions, they may be worthy of statement.

PHYSICAL AND CHEMICAL PROPERTIES OF GOLD-SILVER ALLOYS

The physical and chemical data presented below are in no sense
intended to be a compendium of the mineralogy, physics, and chem-
istry of gold and silver, as an ordinary volume would not suffice to
cover such a vast field. Instead there will be given certain discon-
nected but significant data that seem to bear upon the origin and
accumulation of placer gold. ,

Pure gold has not been found in nature. Instead, it is always al-
loyed with more or less silver, together with a small proportion of base
metals, mainly copper and iron, which collectively are designated as
the dross. Gold has been found alloyed with some of the platinum
metals; and other specialized species, such as alloys of gold and bis-
muth, and natural amalgam have also been described. The purest free
gold that I have been able to find any record of is a sample of lode
gold from the Great Boulder Mine, in the Kalgoorlie district of West-
ern Australia, which had a fineness of 999.1 parts gold in a thousand;
but several samples from the Cripple Creek district are recorded as
having finenesses of 999. Most of the Australian gold is exceptionally
high grade, yet the quartz veins of the Wiluna area, of Western
Australia, have produced the lowest grade of natural gold on record,
ranging in fineness from 400 to 580 parts gold in a thousand, the re-
mainder being mainly silver. Native silver is relatively uncommon,

TABLE 1.—FINENESS OF PLACER GOLD IN ALASKA

Fj Fj
(parts wold pen Anercernat) Nurabeniof samples (parts geld pee chousad) MAE 90) SEES
975-965 4 795-785 2
965-955 13 785-775 28
955-945 33 775-765 Pa)
945-9385 6 765-755 iL?/
935-925 16 755-745 12
925-915 47 745-735 4
915-905 115 735-725 9
905-895 122 725-715 6
895-885 P27 715-705 if
885-875 104 705-695 2
875-865 119 695-685 0
865-855 166 685-675 1
855-845 125 675-665 1
845-835 101 665-655 0
835-825 143 655-645 0
825-815 79 645-635 1
815-805 30 635-625 1
805-795 39 625-615 1

Mar. 15, 1940 MERTIE: PLACER GOLD IN ALASKA 95

and is usually free of gold. The placer gold so far mined in Alaska
ranges in fineness from 620 to 973 parts gold in a thousand. Philip 8.
Smith? has recently compiled from the files of the U. S. Geological
Survey 1,528 assays of the placer gold alloys that have been mined
in Alaska. Seven additional assays, showing finenesses of less than
620, are also on record from the Territory, but these are considered
probably to represent bullion that included man-made impurities such
as lead shot, solder, and other such materials. This record is presented
in Table 1.

The above data are also presented in the form of a frequency curve
in Fig. 1. The graph was prepared by C. E. Van Orstrand and H. C.
Spicer, both of the U. 8. Geological Survey. The following statement,
prepared by Mr. Van Orstrand, gives the essential facts bearing upon
the choice of an interpolation formula, and a summary of the com-
putations.

nresequacion.yi— axe" (bo-- bit bet). 2 oe ew ee es (1)
is here used primarily as an interpolation formula. In this equation, x repre-
sents the series of integers, 1,2,3, .. . beginning with 1 at 970.

SURE GPT ONION hoe ik MOR OG Pie oe oo kan ce ele es (2)
has been used to represent production curves in the oilfields expressed as a
function of the time.* Calculation of the coefficients in (1) has been made
by first evaluating the constants in (2) and then multiplying the computed
values of (2) by (bo +bia+bex?) and solving the product relation equated
to the observed values of y for the values of bo, bi, and be. All the adjustments
were made by the method of least squares.

The results of the calculations show that the sum of the observed values
(Zyo) is 1,524; that of the computed values (Ly-.) is 1,493. The sum of the
squares of the residuals (2v?) is 8,423. The first two quantities can be brought
into closer agreement, and the last one can be reduced somewhat by making
a more precise determination of n. The modal frequency is x=11.11, or, in

terms of fineness, 868.9. The computations involved in obtaining this graph
were performed by Mr. Spicer.

The histogram shown in Fig. 1 may possibly suggest to the reader
that a less regular curve is justified by the numerical data. But the
1,528 assays, upon which this figure is based, are in reality only a
small part of the thousands of assays known to have been made upon
the placer golds of Alaska. Hence the irregularity of the histogram is
interpreted as a condition due entirely to imperfect sampling, and
not to any physical-chemical relationship that might be shown in an
equilibrium diagram. For this reason, a probability curve is believed
to be the best method of representing the available data.

2 SmituH, Puitie 8. Fineness of gold from Alaska placers. U.S. Geol. Surv. Bull.
910: in preparation.

3 VAN ORSTRAND, C. E. On the empirical representation of certain production curves.
Journ. Washington Acad. Sci. 15(2): 19-33. Jan. 19, 1925.

96 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

From this graph it appears probable that the placer gold alloys
most commonly mined in Alaska have a fineness of about 870 parts
gold in a thousand. As the dross seldom exceeds | percent, it follows
that the corresponding content of silver is about 120 parts per thou-
sand. One of the most significant features of these data, concerning
which more will later be said, is the absence in Alaska or elsewhere in
the world of placer gold alloys of a fineness much less than 600; and

160

140 y = axe? (6, 4b,x46, x7)
Jog a = — 0.092506
n= 2975559
b= 0057/85
bo= 0.0915 4/
b, = 0144579
b, =- Q005843

NUMBER OF SAMPLES
@
ce)

D
°o

40

20

Au +AQ +DROSS

Fig. 1.—Frequency curve, showing fineness of placer gold in Alaska.

also the world-wide absence of any gold-silver alloys having a fineness
of less than 400.

One of the most ill-defined terms applied to natural gold alloys is
the term electrum. According to Rose and Newman,‘ electrum means
pale yellow natural alloys of gold and silver containing 15 to 35 per-
cent of silver; but Dana® mentions two species of electrum containing
over 38 percent of silver; whereas Lindgren® restricts electrum to an
alloy containing 50 percent of silver. According to the first definition,

4 Ross, Sir Tuomas K., and Newman, W. A. C. The metallurgy of gold, ed. 7,
DiAGy Lose

5 Dana, JAMES W. The system of mineralogy, ed. 6, p. 15. 1914.

6 LINDGREN, WALDEMAR. Mineral deposits, ed. 2, p. 229. 1919.

Marz. 15, 1940 MERTIE: PLACER GOLD IN ALASKA 97

if 1 percent of the alloy is allocated to dross, all natural gold alloys
in the range of fineness between 840 and 640 parts gold in a thousand
should be classified as electrum; and therefore about one-third of the
natural gold alloys of Alaska are electrum. If the pale yellowish color
is an indispensable part of the definition, many of the Alaskan alloys,
in the range from 840 to 640 fineness, are not necessarily electrum. An
increasing percentage of silver does modify the color of a gold-silver
alloy, and the color of an artificial alloy containing 50 percent of
silver is actually a faint yellowish white. But I suspect that the nat-
ural alloys that the ancients designated as electrum owed their pale
color as much to elements of the dross as to their content of silver.
Therefore, although the term electrum is of historical interest, it does
not seem that it belongs in a handbook of mineralogy as a species or
variety of the natural gold alloys, unless it is given a more exact chem-
ical definition than it now has.

Many elements, usually in small quantities, have been found to
form a part of the natural gold-silver alloys. Such accessory metals
usually constitute from a half of 1 percent to 1 percent of the alloys.
Copper and iron are the principal metals of the dross, but the ratio
of these two elements to one another is extremely variable, such that
either of them may preponderate greatly over the other. Yet a sum-
mation of the chemical analyses of gold, given by Hintze,’ shows
that the average quantities of copper and iron in the natural gold-
silver alloys, is nearly equal. Small quantities of a number of other
elements have also been detected in placer gold by chemical analysis,
and doubtless traces of still others may be found by spectrographic
analysis. The principal accessory metals that have been recognized by
chemical analysis of the natural gold-silver alloys, are zinc, mercury,
tin, lead, antimony, bismuth, nickel, cobalt, rhodium, palladium,
iridium, and platinum; and of these all but antimony, bismuth, co-
balt, and rhodium have been identified chemically in the placer
golds of Alaska.

The physical properties of the natural alloys of gold and silver,
ordinarily designated as free gold, are given scant consideration in
books on mineralogy and economic geology; and some quite erroneous
statements have been published. Thus in a recently published hand-
book it is stated that ‘‘gold is a yellow malleable metal with a specific
gravity from 15.6 to 19.3, and a melting point at about 1,062° C.”
Pure gold does have the melting point stated, but its specific gravity

7 Hintze, Cart. Handbuch der Mineralogie 1: 316-320. 1904.

98 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

is not to any considerable degree variable, being approximately 19.3
to the nearest digit in the first decimal place. And if the author’s de-
scription referred to natural gold alloys of variable composition, then
their melting points were not 1,062° C. Moreover, natural alloys of
gold and silver are known that have specific gravities as low as 12.5
instead of 15.6. Hence the whole statement is erroneous.

The physical properties of gold and silver are well known and are
carefully tabulated in reference books such as the International Criti-
cal Tables. But in this paper it is the natural alloys of gold and silver,
rather than the pure metals, that are being discussed. Therefore, for
present purposes, the following short tabulation of the physical prop-
erties of pure gold and pure silver will suffice:

TABLE 2.—PuysIcAL PROPERTIES OF GOLD AND SILVER

Property Gold Silver
ACOMICH Wels Initia 7h: Sopa) ee eae erode oe 197.2 107.9
Specihcreravitye oo eke eon rae wees nan 19.3 10.5
Meltingépoints(G@) ea en ee 1063° 960°
Boling, point) (C@)tesaic. fae Lea en ea ae 2600° 1950°
Hiardness:(Mohysiscale)= a5. sees a 2 5 3.0
Electrical resistivity. Ohms per cméat 20°C. Dali ine oO <0)
Magnetic susceptibility (mass unit) at
IMSS Et © oi tae Mea rte Bex OPA re i, tiownen feat Oh eee Cate —0.15X107° —0.20 X10
Thermal coefficient of linear expansion at
DOS Or aes ob oe near nt mientn even. See eeeS te Wet Ae 2 lOme 18.9X10%
Crystallographye ese ier. oa ud ohne Face-centered cubic|/Face-centered cubic
space lattice. space lattice.
Length of side of cubic lattice (em)....... 4.07 X1078 4.08 X1078
ACOMIC a aGius* (Cim)eluss ele as elt ees oe 1.441078 1.44 x1078
Wiorkalbilit yarns cc sose Ann eee cree arceeas - Very malleable and Very malleable and
ductile. ductile, but less so
than gold.

Placer golds are well known to be natural alloys, principally of gold
and silver, but judging from published descriptions little is known of
their true nature. Much investigative work, to be sure, has been done
on artificial alloys of different metals, including gold and silver, but
even this information does not appear in our textbooks on mineralogy.
In dry melts, gold and silver are continuously miscible in all propor-
tions, like alcohol and water, and remain so after solidification, form-
ing a solid solution analogous to albite and anorthite. Raydt® has
studied the binary system of gold and silver, and has derived an
equilibrium diagram, relating composition to temperature, which is
shown in Fig. 2.

This diagram shows continuous solidus and liquidus curves, with-
out maximum or minimum points. It should also be emphasized that

8 Raypt, U. Uber Gold-Silberlegierungen. Zeitschr. anorg. Chemie 75: 58-62.
1912. |

Mar. 15, 1940 MERTIE: PLACER GOLD IN ALASKA 99

gold and silver, as well as all other metals that have solidified from
dry melts, exist as crystallites, usually in allotriomorphic aggregates.
These crystallites, in any one alloy, do not necessarily have the same
chemical composition; nor, in fact, is any one crystallite necessarily
chemically homogeneous.

Copper and iron, however, the principal metals of the dross, behave
differently, when melted with gold and silver. Copper, for example, is
miscible in all proportions in the liquid state, either with gold or with
silver, and forms a solid solution with gold in the solid state; but cop-
per and silver have limited ranges of miscibility in the solid state,
from 0 to 5 percent and from 954 to 100 percent copper. Iron, on the

oo

1050

TEMPERATURE
fo}
fo)
°.

950

Au
Au + Ag

Fig. 2.—Equilibrium diagram for artificial gold-silver alloys.

other hand, shows unlimited miscibility with gold in the liquid state,
but complete immiscibility with silver; and in the solid state, alpha
iron at 20° C shows solid solution with gold in the ranges from 0 to
20 percent, and 82 to 100 percent iron, whereas with silver iron is
completely immiscible in the solid state.

The phase relations shown by Raydt’s equilibrium diagram, and
additional phase relations known to exist between binary and ternary
systems of gold, silver, copper, iron, and other elements studied in
artificial melts, do not necessarily hold for natural alloys, because the
effects of still other components, such as quartz, water, base metals,
and the so-called mineralizers have not been evaluated. It seems
likely, however, that the relationship observed in dry melts will in
some measure, perhaps in large measure, apply to the natural alloys
of gold and silver; and for this reason placer gold should be considered

100 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

to be altogether or in large part a solid solution of crystalline gold and
silver, with small quantities of other metals, in unknown phase rela-
tionships. Yet certain significant differences between natural and
artificial alloys have been observed, which seem worthy of emphasis.

If any two metals were completely immiscible in the solid state, so
that they solidified as individual grains in a eutectic or mechanical
mixture, it should be possible from a chemical analysis to compute
those physical properties such as specific gravity, which are functions
of additive quantities like mass and volume. But gold and silver ecrys-
tallize from dry melts as solid solutions, or, if you will, as atomic mix-
tures; and if the natural alloys of gold and silver are likewise solid
solutions, it should not be possible from chemical data alone to com-
pute their specific gravities. This is found to be true. But it is a matter
of considerable interest to assume that the eutectic relationship holds,
and to compute, either from artificial mixtures of known weights of
gold and silver or from chemical analyses of the natural alloys of
these two metals, the specific gravities that should result. These the-
oretical values can then be used as standards, with which to compare
the respective alloys, in order to determine whether the latter show
contraction or expansion in volume, and the order of such changes,
as compared with mechanical mixtures of the two components. For
artificial alloys of gold and silver, this has been done by Matthiessen?
and by Hoitsema.'’° The specific gravities have also been computed
theoretically by McKeehan"™ from X-ray analysis.

The molecular composition indicated by Matthiessen and Mc-
Keehan were converted to percentages of gold and silver by weight,
to correspond with the form of the ratios given by Hoitsema. From
these ratios, and from the specific gravities of the pure gold and pure
silver used by these three men, the specific gravities and specific
volumes were then computed, which should exist if the two compo-
- nents were mechanically mixed. The differences between these specific
volumes, and those found by actual measurement or by X-ray analy-
sis, were then compared, giving a column of differences which show
whether volumetric contraction or expansion took place. All these
data are shown in Tables 3-5.

An adjusted graph of the specific gravities, utilizing the data of
Matthiessen, Hoitsema, and McKeehan has been published in the

® Marruiessen, A. On the specific gravity of alloys. Philos. Trans. Roy. Soc.
London 150: 188. 1859.

10 Horrsema, C. Die Dichte von Goldkupfer und Goldsilberlegierungen. Zeitschr.
anorg. Chemie 41: 66-67. 1904.

1 McKernan, L. W. The crystal structure of silver-palladium and silver-gold alloys.
Phys. Rev. (ser. 2) 20: 429. 1922.

Mar. 15, 1940.

MERTIE: PLACER GOLD IN ALASKA

101

TABLE 3.—SPECIFIC GRAVITIES AND SPECIFIC VOLUMES OF ARTIFICIAL ALLOYS

Composition!

Spec. Grav.? . Spec. Grav. Spec. Vol.
(true) (computed) (true)
19.265
18.041 17.999 0.05548
17.540 17.493 .05701
16.354 16.315 .06115
14.870 14.847 .06725
13.432 13.383 .07445
PA PAE. UPA PAS) .08159
11.760 Neeley .08503
10.468

Spec. Vol.
(computed)

0.05556
.05717
.06129
.06735
.07472
.08187
.08536

Spec. Vol.
(differences)

+0 .00013
.00016
.00014
.00010
.00027
.00028
.00033

++++4+4+

1 Atomic weights of 197 and 108 were used by Matthiessen in determining the percentages of gold and silver
in the specified alloys.
2 Specific gravity determinations by A. Matthiessen.
3 Owing to a typographic error, this reads AuAgs in the original paper.

TABLE 4.—SpEcIFIC GRAVITIES AND SPECIFIC VOLUMES OF ARTIFICIAL ALLOYS

Parts per thousand

Au Ag
1000 0
917 83
843 157
750 250
667 333
583 417
500 500
417 583
333 667
250 750
167 833
0 1000

Spec. Grav.!
(true)

Spec. Grav.?
(computed)

.000
.009
BOOM
.038
. 250
. 049
913
.328
OY)
.314

1 Specific gravity determinations by C. Hoitsema. ce : p
2 A number of corrections were made in the specific gravities and specific volumes computed by Hoitsema.

Spec. Vol.
(true)

.05531
.05896
.06238
.06636
.07023
.07353
.07692
.08070
.08489
.08861

Spec. Vol.
(computed)

.05556
.05879
.06287
.06650
.07018
.07381
.07744
.08112
.08475
.08839

Spec. Vol.
(differences)

I++ i++ |

TABLE 5.—SPECIFIC GRAVITIES AND SPECIFIC VOLUMES OF ARTIFICIAL ALLOYS

Composition!

Spec. Grav.? Spec. Grav. Spec. Vol.
(X-ray) (computed) (X-ray)
19. 24 —— ——
18.24 18.365 .05482
17.44 17.490 .05734
16.06 16.615 .06227
15.31 15.740 .06523
14.74 14.865 06784
13.50 13.990 .07407
11.93 12.240 .08382
10.49 Saaeee ——

Spec. Vol.
(computed)

.05445
.05718
.06019
.06353
.06727
.07148
.08170

Spec. Vol.
(differences)

—0.00037
.00016
.00208
.00179
.00057
.00259
.00212

[eel eleel elem

_ 1} Atomic weights of 197.2 and 107.88 were used by McKeehan in determining the percentages of gold and
silver in the specified alloys.

2 X-ray observations and derived specific gravities by L. W. McKeehan.

International Critical Tables.!? This, with some corrections, is shown

in Fig. 3.

590. 1927.

2 International critical tables of numerical data: Physics, chemistry and technology 2:

102 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

20

16 by e
°
15
>
2
>
q
o
14
°
=
(S)
wW
a
” 13
12 .
u
10
0 2 3 4 5 § 7 8 B-) 10
Au
Au +Ag

Fig. 3.—Adjusted curve, showing specific gravities of artificial gold-silver alloys
as obtained by Matthiessen©, HoitsemaA, and McKeehan@.

The computation of the specific gravity of a binary alloy from
its chemical composition, if the two components are considered to be
mechanically mixed, is simply done by means of the following formula:

ia Me Maa Ma Cn ei See

Vite Vo Mine elie MS. + MoS,

Si Se
where M, and M, are the weights, and S, and S; are the corresponding
specific gravities of the two components. To compute the specific
gravity of a n-ary alloy from its chemical composition, upon the
assumption that all the components are mechanically mixed, the
above formula has been expanded to read as follows:

ah QUE a ley Wy ae Se UE Sms 4 2 S)a)

CMS2SsS4- Sz)! + CMaSi8s84 > S;) 4- CUGSiS 25a ashe ieee

;
a

Mar. 15, 1940

(ob (My28iS2 - -

MERTIE: PLACER GOLD IN ALASKA

+ (M8183 -

Z Spesnasn) as (M ,,-1SiS2 oll

2 Sn—3Sn—2S fal)

103

‘Sn—3S n—29 n) =F

Now let us apply this formula to four complete chemical analyses of
placer gold alloys from Alaska, which have recently been made in the
laboratory of the U. 8S. Geological Survey. These are presented in

Table 6.
TABLE 6.—ANALYSES OF PLACER GOLD From ALASKA

Constituent 1 2 3
Sold veh. 83 .90 80.59 87.53
SUWeR, 30a 10.42 oA EL
Peraghmunims, <. es .01 . 20 28
Unchiinne se .02 .05
fgalleciuim.. 3... wee | trace
Lo Bitl , o oi gine .06 O07
MNeRCURY Se. ss sl) .05
LSI. loo) exO2 .08 .07
WRG. gn .007
C ODDGCs 6 sie .002 .03 sOik
DUNG. 6 Oe ee .04 .03
tio ob.06:6.0 trace trace
SiiGs..6 36 35 ee .43 .40
SiOz, Al.O3, and Cr.0; 5 432
Insoluble in aqua regia. 1.67
OIA. = 4s ae 97.461 99.76 99.61

100.26

trace
ath

1. Squirrel Creek, Goodnews Bay district, southwestern Alaska. E. T. Erickson,

analyst.

2. Seventymile River, Eagle district, east-central Alaska. R. E. Stevens, analyst.
3. Fourth of July Creek, Eagle district, east-central Alaska. R. E. Stevens, analyst.
4. Woodchopper Creek, Circle district, east-central Alaska. R. E. Stevens, analyst.

When the specific gravities of these four samples are computed,
upon the assumption and with the formula above given, some anoma-
lous results appear. These are given in Table 7.

TABLE 7.—SPECIFIC GRAVITIES AND SPECIFIC VOLUMES OF PLACER GOLD FROM ALASKA

Sample Spec. Grav.
Oo. (true)
1 LB,o4
ee, aes
3 ee 16.00
A dae 15.87

Le
16
17
18

Spec. Grav.
(computed)

.43
. 69
.61
dl

Spec. Vol.
(true)

0.07508
.06588
.06250
.06301

fr

Spec. Vol.
(computed)

0.05737
.05992
.05679
.05461

Spec. Vol.

(differences)

=O ONiaial
— .00596
= sWWS71
— .00840

The experimental work by Matthiessen shows that when pure gold
and pure silver are melted together in definite proportions, there re-
sults a contraction in volume of small magnitude, regardless of the
proportions of the two metals. Hoitsema’s results show both a con-

104 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 3

traction and an expansion of volume, for different compositions, but
the differences between the true and computed specific gravities are
at least of the same order of magnitude as those found by Matthies-
sen. The results obtained from McKeehan’s data are not strictly com-
parable with those obtained by Matthiessen and Hoitsema but are
given for the sake of comparison. When the same computations are
made from chemical analyses of placer golds, as shown above, there
results an expansion in volume, of large and variable magnitude.
Parenthetically it may be added that sample no. 1 is held by a strong
electromagnet, although both gold and silver are diamagnetic. The
reason for these differences between the artificial and natural alloys
of gold and silver is not known, but there seem to be ample grounds
for suspecting that complex phase relationships may exist. If this is
true, a careful study of placer gold by physical and chemical methods
is needed, in order to discover the present compositions of such alloys,
both in bulk and in crystallites or grains of microscopic dimensions.
A knowledge of the present physical and chemical conditions of these
alloys should also lead to a better understanding of the temperatures,
pressures and other conditions that existed in gold-quartz veins at
the time of their formation.

The solubilities of the natural alloys of gold and silver in inorganic |
acids also have a bearing upon their occurrence in nature, both in
their bedrock habitats and in their subsequent occurrence in placers.
Gold is not appreciably attacked by any single acid except selenic acid
(H.SeO,), but it dissolves readily in solutions generating the nascent
halides, and therefore in aqua regia. Silver, on the other hand, is dis-
solved by boiling concentrated sulphuric acid, by dilute nitric acid
and by hydriodic acid, and is attacked by all the halides, by warm
selenic acid, and to a very slight extent by hydrochloric acid in the
presence of oxidizing agents. Silver has also been shown to be slightly
soluble in distilled water, to the extent of 3.7 x10~ grams per liter.
But just as the physical properties of the alloys of gold and silver can
not be directly inferred from the physical properties of the two pure
components, so the solubilities of these alloys are not linear functions
of the solubilities of pure gold and silver.

One profitable line of investigation in connection with the study of
gold placers would be a study of the solubilities of artificial and
natural gold-silver alloys in the inorganic acids and other solvents
that might be expected to occur in surface and ground waters. At
present the most suggestive fact of this sort results from the well-
known process of quartation, or inquartation (as it is sometimes

Mar. 15, 1940 MERTIE: PLACER GOLD IN ALASKA 105

called), which has been used for over 600 years for parting silver from
gold-silver alloys. Natural gold alloys are not attacked either by
nitric or sulphuric acids, but if a sufficient amount of silver is added
to the alloy, practically all the silver can then be dissolved by nitric
acid at a temperature of 115° F. Essentially the same result is ob-
tained by the use of sulphuric acid. The really interesting feature,
however, is the fact of the limits of partial and complete solubility;
and these are shown in the accompanying diagram, taken from
Tammann’s® work. (See Fig. 4.)

100

80

60

40

PERCENTAGE SOLUBILITY OF SILVER

20

0.2 0.3 0.4 0.5 0.6 0.7

Au +Ag
Fig. 4.—Sketch showing amounts of silver dissolved by nitric acid
from artificial gold-silver alloys.

The fact particularly to be noted in this diagram is that no silver
can be dissolved by inorganic acids from an artificial gold-silver alloy
having the general composition of placer gold, because the high con-
tent of gold renders them practically insoluble. What would be the
probability, then, that any appreciable amount of silver could be dis-
solved from placer gold by cold surface waters? Further reference
will later be made to this topie.

GEOGRAPHIC DISTRIBUTION OF GOLD PLACERS IN ALASKA

It is not the purpose of this paper to tabulate and describe the gold
placers of Alaska, though such a compilation might indeed be a useful

13 TaAMMANN, Gustav. A textbook of metallography, ed. 3, p. 314. 1925. (Trans-
lated from the German by R. S. Dean and L. G. Swenson.)

106 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

piece of work. Instead, the writer aims principally to point out the
known facts that have a bearing upon the origin and accumulation of
such placers; and in order to accomplish this objective, geographic
references will be made only secondarily.

The principal fact to be noted regarding the geographic distribution
of gold placers in Alaska is that all the really important deposits occur
in the unglaciated part of the Territory. (See Fig. 5.) This unglaciated
placer-bearing region lies north of the Alaska Range, south of the
Brooks Range, and extends westward and southwestward to the
Arctic Ocean and Bering Sea. To be sure, several small placer camps
are located south of the Alaska Range, but these in toto produce no
considerable part of the placer gold mined in Alaska. Granitic rocks, —
which are considered to be the ultimate source of most of the placer
gold, are widely distributed in Alaska south of the crest of the Brooks
Range, and therefore no a priori reason exists why lode and placer de-
posits should not have coexisted north and south of the Alaska Range
before the beginning of the Ice Age, and doubtless this was true. But
glacial action in the Alaska Range and coastal ranges, and seaward
therefrom, was so severe that the country rock was largely denuded,
thus eroding and dissipating most of the pre-Quaternary alluvial de-
posits. In general, there has been insufficient time and erosive action
since the retreat of the ice to produce new gold placers from bedrock
sources. But at some localities in the glaciated regions, auriferous
glacial deposits have been reconcentrated by Recent streams, thus
producing workable placers; and, in fact, at one locality in the Cache
Creek district, a glacial deposit has constituted alow grade placer. At
other localities in southern Alaska there remain remnants of pre-
glacial placers, which were located in valleys that were transverse to
the principal flow lines of the ice, or were otherwise protected from
glacial scour. These explainable exceptions in no way vitiate the
general rule that the important placers are restricted to the unglaci-
ated part of Alaska. |

Most of the workable gold lodes, however, lie south of the Alaska
Range, or in southeastern Alaska southwest of the Coast Range,
though a number of small gold lodes have been discovered and are
being mined in the unglaciated region of interior Alaska. The mean-
ing of this distribution of the gold lodes is less apparent, but it is —
possible that the denudation of bedrock by glaciation has been an im-
portant factor favoring the discovery of lodes in southern and south-

144 Capps, STEPHEN R. Glaciation in Alaska. U.S. Geol. Surv. Prof. Paper 170-A, pl.
i L931

107

MERTIE: PLACER GOLD IN ALASKA

Mar. 15, 1940

NOILUNV1dX

‘BYSB[Y JO S¥ale poyVIov[duN UI S1e0B][d PjOS Jo UOT} eZI[vVIO0T—'g “BI

3 , ‘ : 2

108 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 3

eastern Alaska. On the other hand, the heavy cover of residual and
eluvial deposits in interior Alaska has certainly been a handicap to
lode prospecting, and this may be an important factor in their ap-
parent scarcity in that region.

CLASSIFICATION OF GOLD PLACERS

A complete classification of gold placers is beyond the scope of this
paper. There are probably types of placers with which the writer is
not familiar, and there are certainly some, such as aeolian placers,
that have not been recognized in Alaska. The following simple classi-
fication, therefore, aims mainly to tabulate the kinds of placers that
are known in Alaska:

I. Residual and eluvial placers.
II. Fluviatile placers.
A. Recent stream placers.
B. Ancient stream placers.
1. Bench stream placers.
2. Buried stream placers.
3. Compound stream placers.
C. Hard rock sedimentary reefs.
III. Glaciofluviatile placers.
IV. Beach placers.
A. Recent beach placers.
B. Ancient beach placers.
1. Elevated beach placers.
2. Buried beach placers.

Residual and eluvial placers are those that have been formed in or
almost in situ, as a result of deep residual alteration and disintegra-
tion of a gold-bearing bedrock. Placers of this type are geographically
uncommon and are of small economic significance in Alaska. No sharp
distinction can be made between residual and eluvial placers, and in
fact it is doubtful whether a truly residual deposit can be said to exist,
because this would imply an almost total absence of lateral movement
of the gold-bearing detritus, and a concentration largely by chemical
removal of rock constituents, and a downward segregation of gold to-
ward bedrock. For this reason these two types are grouped into a sin-
gle class. Perhaps the best example of combined residual and eluvial
placers in Alaska is in the Iditarod district, where a mass of monzonite
forms the country rock along a divide at the head of several streams,
notably Flat Creek. This monzonite is cut by many small seams and
stringers of gold-bearing quartz, some of which have been exposed by
mining operations. At this locality deep residual alteration of the
monzonite has taken place, and by the combined action of gravity,
frost-thrusting, and ground water a great volume of loosened and dis-

Mar. 15, 1940 MERTIE: PLACER GOLD IN ALASKA 109

integrated bedrock is slowly moving downhill into the adjoining val-
leys. Some of this material might possibly be classed as a residual
placer, but another part, which has clearly migrated down the slopes
of the ridge but has not yet been handled by running surface water,
is a good example of an eluvial placer.

Most of the commercial placers of Alaska are of the fluviatile type.
These have been divided upon the basis of their relative ages into
present and ancient stream placers, but in this connection a sharp
distinction must be made between the age of the present stream grav-
els, and the age of the alluvial gold contained in them. In speaking of
southern Alaska, it was stated that sufficient time had not elapsed
since the retreat of the ice for the present streams to erode from bed-
rock sources, and to concentrate workable gold placers. This is be-
lieved also to be true in interior Alaska, but postglacial and Recent
time in Alaska are not necessarily synonymous terms, and the term
Recent may connote a longer interval of time, at least at some locali-
ties, than the term postglacial. But a more important consideration is
that a large volume of already disintegrated debris, and also older
placers, have been continuously available to the Recent streams of
interior Alaska, throughout their life history. Hence Recent placers
do really exist in interior Alaska, though it is doubtful whether any
considerable part of the gold contained in them was eroded from bed-
rock in Recent time. Recent placers must therefore be defined as plac-
ers in which the component gravels have been moved to their present
sites during a single erosional cycle, controlled by a nearly static base
level of erosion, and within the Recent epoch. Many examples of
Recent stream placers could be cited, but their common occurrence
renders this unnecessary. Generally speaking, the Recent stream
placers are of less economic value than the older fluviatile placers.

The ancient stream placers consist of gold-bearing fluviatile gravels
that were deposited during one or more cycles of erosion preceding the
last one. Such placers are arbitrarily divided into bench and buried
placers, upon the basis of the altitude of their underlying bedrock, as
compared with the altitude of the bedrock lying below a body of ad-
joining stream gravels. Bench placers are commonly developed as the
result of a renewed cycle of erosion, in which the local base level of
erosion is lowered at a rate greater than the rate at which a stream can
lower by erosion its preexisting valley floor. If this occurs, there will
remain uneroded remnants of the old bedrock floor, covered by more
or less alluvial material at altitudes appreciably higher than the bed-
rock and alluvium of the new valley floor. Such uneroded remnants of

110 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

auriferous alluvium constitute true bench placers. If, however, the
rate of lowering of the local base level of erosion is equal to or less than
the erosional rate of the stream, the older bedrock surface will be con-
currently lowered and its overlying cover of alluvial material will be
continuously reworked, in such a manner that neither bedrock nor
alluvial terraces will be evolved. The same result more frequently ma-
terializes as a result of variable rates of the two functions, so that
terraces have been formed and subsequently destroyed, leaving little
or no trace of their former existence. In such cases the gold is really
ancient bench gold, but the gravels now containing it are Recent
placers. Similarly, however, the gold of the present bench placers may
have rested temporarily for considerable periods in still older placers.
Many splendid examples of true bench placers occur in the Fortymile
district. On the other hand, Mastodon Creek, in the Circle district, is
a good example of a valley in which the terraces were destroyed al-
most as fast as they were created.

Buried placers are those that resulted from an elevation of the local
base level of erosion, or from other causes such as aeolian action, so
that ancient auriferous gravels that formerly constituted a valley
floor were progressively buried by a general alluviation. In a certain
sense, all placers are buried placers because erosion in the headwater
part of a valley can take place contemporaneously with alluviation in
its lower stretches, even during an erosional cycle initiated by a lower-
ing of the local base level of erosion. But the term buried placers is
here used to designate gold-bearing alluvial deposits which were
formed in an erosional cycle that antedated the cycle during which
they were buried. Unlike bench placers, buried placers can not be
simultaneously created and destroyed, for theoretically in the sites
of buried placers, erosion of bedrock is nil, though it will continue to
exist in the headwater parts of a valley unless or until aggradation
reaches that far upstream. Some of the placers of the Fairbanks dis-
trict are examples of buried placers, but for the most part they are
more completely described as compound placers, because they have
been materially affected by one or more erosional cycles that post-
dated their burial.

Compound stream placers comprise many gold-bearing alluvial
deposits, which on geomorphic grounds could be divided into an al-
most endless number of species. The development of these subordi-
nate types is a function of the number and character of the variations
in the local base level of erosion, and also of the rate of change of such
variations; and these factors, in turn, are functions of simpler ones,

Mar. 15, 1940 MERTIE: PLACER GOLD IN ALASKA felt

which often are still too complex for simple exposition or analysis. The
placers of the Livengood district, to take a relatively simple example,
were at one stage in their development buried placers. As a result of
an elevation of the base level of erosion, accompanied by extensive ag-
gradation, Livengood Creek was superposed onto one side of its old
valley wall. By a subsequent lowering of the base level of erosion, a
new valley floor was carved contiguous to the old valley floor, but
separated therefrom by a bedrock reef. Finally the new valley floor
was so greatly eroded that the altitude of its bedrock floor became less
than that of the old bedrock floor, whereupon the old buried placer
came to have the general appearance of a bench placer. Similarly
bench placers can be rebuilt to simulate simple buried placers; bed-
rock terraces may be evolved which are younger than their contiguous
valley floors; and in the geologic interval ordinarily considered to
comprise the geomorphic record, these and other anomalous processes
may be several times repeated. Consequently compound stream plac-
ers are as difficult to classify as sedimentary rocks, and at present
their characteristics are much less well known. Many of the ancient
placers of interior Alaska are compound placers, but at few places
have their complete histories been deciphered.

Alaska contains no auriferous hard-rock reefs of economic value,

similar to the Witwatersrand deposits of South Africa. But Tertiary
sedimentary rocks carrying sparsely disseminated gold, occur south
of the Yukon River in a narrow belt extending from the international
boundary west-northwest for more than 100 miles. The conglomerates
that constitute a part of this sequence of rocks have clearly acted as a
proximate source of the gold now contained in the bench and stream
placers of this general district. This belt of Tertiary rocks is the best,
though not the only, example of auriferous hard-rock reefs known in
Alaska.
_ Glacial action tends to dissipate, rather than to concentrate, the
heavy metals and minerals, but few glacial deposits are altogether free
of the effects of running water. Some glaciofluviatile deposits, however,
are fairly well sorted, and except for their original heterogeneity are
not especially different from normal stream deposits. Hence if glacial
and more particularly glaciofluviatile deposits were derived from a
gold-bearing bedrock, and were not too far removed by glacial action
from their original sources, the glacial dissipation of the gold might
be small enough to render such deposits workable as placers. One of
the best examples of a glaciofluviatile placer conforming to these
conditions may be seen on Bird Creek, in the Cache Creek district.

112 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

If gold-bearing rocks occur close to a body of standing water large
enough to produce marked wave action along its bounding beaches,
gold placers may be formed by the sorting action of the waves. The
formation of such beach placers, however, is often aided by the prior
concentration of gold as fluviatile placers in valleys which discharged
to the sea. Beach placers are known in Alaska at a number of locali-
ties, but the richest and best known of these are at Nome, bordering
upon Norton Sound. Here are found Recent beach placers and ancient
ones, the latter including both the elevated and the buried types.

The conditions at Nome can best be understood by visualizing a
sloping foreland, with a width of more than 3 miles, which lies be-
tween low hills to the north and Norton Sound on the south. Within
this foreland, the bedrock is covered by alluvial deposits ranging in
thickness from 30 to 120 feet, and of these the lower few feet super-
jacent to bedrock are beach gravels, mainly of Pliocene age. The over-
lying deposits are both marine and terrigenous in origin, the terrige-
nous deposits lying for the most part closer to the hills. Starting 3 miles
from the sea, a bedrock profile drawn southward toward the sea shows
an initial altitude of about 70 feet above sea level, but passes below
sea level about three-fourths of a mile north of the present beach. At
the north end of this bedrock profile is a nearly vertical cliff, 12 to 14
feet high; and here there exists an alluvial cover about 85 feet thick,
of which the lower 3 to 5 feet are Pliocene beach gravels, whereas the
overlying material consists mainly of terrigenous deposits of Quater-
nary age. From the base of this cliff, bedrock slopes gently and gradu-
ally southward to and beyond the present strand line; and upon this
sloping surface are a number of sites at which pronounced concentra-
tions of gold were effected by an advancing Pliocene sea. Without cit-
ing the local names of these successive sites, it suffices to state that
there are five such beaches on bedrock, of which the two most south-
erly, which lie below the present level of the sea, are the oldest;
whereas the other three, which are above the present sea level, are
progressively younger to the north, the youngest being at the base of
the above-mentioned bedrock cliff. As the sea subsequently retreated,
another auriferous beach was formed about half a mile from the sea,
but not on bedrock; and finally, the present auriferous beach was
built. From the fossil record, the sixth (non-bedrock) beach is also of
Pliocene age, so that a long record of Pleistocene marine sedimenta-
tion appears not to be present in this area. This conforms with con-
ditions found elsewhere along Bering Sea, which point to the presence
of a strand line far seaward of the present strand line, during a large

Mar. 15, 1940 MERTIE: PLACER GOLD IN ALASKA 113

part of Pleistocene time. From these general conditions, it is easily
understood how and why there occur at Nome present, ancient ele-
vated, and ancient buried beach placers.

BEDROCK SOURCES

The natural gold-silver alloys of Alaska are associated, either di-
rectly or indirectly, with granitic rocks, or with their surficial equiv-
alents; and the general term granitic rocks is here used to include
granite, monzogranite, quartz monzonite, granodiorite, and quartz
diorite, together with their quartz-poor equivalents. Few if any quartz-
free granitic rocks appear to be genetically associated with gold, but
there are some, such as the monzonite at the head of Flat Creek, in
the Iditarod district, that are distinctly low in silica.

Granitic rocks of several ages have been identified in Alaska. The
oldest of these is a pre-Cambrian formation called the Pelly gneiss,
which occurs in east-central Alaska and consists dominantly of mas-
sive augen gneiss grading into feldspathic quartz-mica schist. Cock-
field,’ of the Canadian Geological Survey, implies though he does not
definitely state, that some part of the placer gold of the Sixtymile
district, of Yukon Territory, is genetically related to the Pelly gneiss,
but no evidence of such a relationship has so far been discovered in
Alaska. | |

The granitic rocks of the Coast Range and the Alexander Archi-
pelago of southeastern Alaska, of the coastal ranges of southern
Alaska, of parts of interior Alaska, and of the Brooks Range of north-
ern Alaska are considered to be mainly of Mesozoic age. It is believed,
however, that these granitic rocks range in age from Middle Jurassic
to Lower Cretaceous, being emplaced in more than one geologic epoch
and probably in severai stages. The granitic rocks of the Coast Range,
according to Buddington," consist principally of granodiorite, quartz
monzonite, and quartz diorite, in the order named; but to the south-
westward, in the Alexander Archipelago, the granitic rocks appear to
be less silicic and may be somewhat older than those of the Coast
Range proper. The granitic rocks of the coastal ranges of southern
Alaska are also believed to antedate the granitic rocks of the main
Coast Range, being probably of Middle Jurassic age. In interior
Alaska, the Mesozoic granitic rocks include mainly quartz diorite and
granite, with few intrusives of monzonitic character. They may be

16 CocKFIELD, W. HE. Szixtymile and Ladue Rivers Area, Yukon. Can. Geol. Surv.
Mem. 123: 49. 1921.

16 BuppiINnGTON, A. F., and CHapin, THEODORE. Geology and mineral deposits of
southeastern Alaska. U. S. Geol. Surv. Bull. 800: 177. 1929.

114 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

either of Jurassic or Cretaceous age, as the stratigraphic evidence of
their geologic age is lacking. On rather insufficient data the grano-
dioritic rocks of the Brooks Range are likewise considered to be either
of Jurassic or Cretaceous age. Workable gold-bearing quartz veins
and lodes of other types have been found in the vicinity of some of the
Mesozoic granitic rocks, and to a greater extent where placers are not
now present, than where they are. Such lodes appear to represent
mainly the mesothermal type of mineralization.

The Tertiary period also was characterized by the intrusion of
granitic rocks, in one or more stages of emplacement. Certain early
Tertiary granitic rocks appear to be normal granite, quartz monzo-
nite and quartz diorite, but another type believed to be of later Ter-
tiary age are nearly everywhere quartz monzonite or monzonite, rela-
tively low in free silica, and carrying pyroxene as the principal dark
mineral. Such rocks are widely distributed in southwestern Alaska,
and are rated as second in importance as producers of placer gold.
Much mineralization has been found in and near the mid-Tertiary
granitic rocks, in the vicinity of some of the derived placers, but few
important lodes of this age have been discovered. In general, the gold
occurs in small discontinuous quartz veins and stringers, which appear
to represent fillings in shattered epithermal zones. Cinnabar is almost
everywhere associated with the placer gold that has been derived from
the mid-Tertiary granitic rocks, and some mercury lodes of this age
have been successfully mined.

FORMATION OF PLACERS

Since most of the placers of Alaska are of fluviatile origin, and even
the beach placers were in some measure concentrated as preexisting
fluviatile deposits, the formation of placers deals mainly with the ero-
sion of gold from bedrock sources, and its transportation and con-
centration by the action of streams.

Many data indicate that most of the placer gold of Alaska was liber-
ated from its bedrock sources long before it was finally deposited
in the placers that are now being mined. In other words, it is believed
that much of this gold has been handled and rehandled by streams in
many successive geomorphic cycles. In east-central Alaska, for exam-
ple, the gold that originated in the Mesozoic granitic rocks south of
the Yukon River, began to be freed from its bedrock sources when
those rocks were first bared to erosion; and since a considerable part
of the gold was deposited, and some of it still remains, in the early
Tertiary conglomerates of this area, the long alluvial history of the -

Mar. 15, 1940 MERTIE: PLACER GOLD IN ALASKA 115

gold in this area is not open to question. At most localities, however,
this generalization is not directly provable, though it is strongly sug-
gested by the presence of large quantities of placer gold in certain
localities where few or no evidences of mineralized bedrock can be
found; by great differences in fineness between placer gold and the
gold of geographically contiguous lodes; and in fact by the mere pres-
ence of deeply truncated laccoliths and stocks of granitic rocks in the
drainage basins where workable placers occur. Naturally, since the
parent rocks range in age from Middle Jurassic to mid-Tertiary, no
generalized statement is possible regarding the ancient climatological
conditions under which the country rock was first disintegrated and
made available to stream action. But during some epochs, notably
just before the general epeirogenic uplift at the end of the Pliocene
epoch, a large volume of residual and eluvial material is believed to
have mantled much of Alaska. In interior Alaska, for example, there
are placer camps, as at Poorman, where practically all the gravels of
the placers are vein quartz and chert. In such localities, it seems cer-
tain that the concentration of siliceous rocks is due to the disintegra-
tion and destruction of the other rocks with which they were orig-
inally associated. Probably, therefore, a large part of these siliceous
gravels were derived from bedrock sources long before the Quaternary
period.

In streams having gradients of the same order as those obtaining
in the medial courses of the usual placer streams of interior Alaska,
the downstream vector of movement for gold appears to be small.
Splendid examples of this feature are apparent in Fourth of July,
Coal, and Woodchopper Creeks, in east-central Alaska, where impor-
tant placers have been concentrated from the above-mentioned Ter-
tiary bedrock in the present valleys during several erosional cycles.
The Tertiary rocks cross these three valleys as a belt several miles in
width. Upstream from these gold-bearing rocks no placers exist, and
downstream from them the workable placers terminate in a very short
distance. The same feature may also be noted in Hunter and Little
Minook Creeks, in the Rampart district, where the present placers
have been reconcentrated from a belt of unconsolidated auriferous
Pliocene gravels, that crosses the valleys of these two streams. Some
of the very fine gold, of course, travels many miles downstream, and
lodges in the large trunk valleys; but the amount appears to be very
small in comparison with that which is repeatedly handed by streams
in successive erosional cycles, and still remains in the original valleys
where it was first concentrated.

116 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

Since most of the placers of interior Alaska are classified as com-
pound types, no simple exposition of their general geomorphic history
can be attempted; but it is relevant to review the classical hypothesis
of their formation during a single cycle of erosion. (See Fig. 6.) In
most small streams much of the alluvial material is in course of pro-
gressive movement from the headwaters downstream. In the upper-
most stretches all this alluvium, from the surface to bedrock, at times
of flood is moved downstream, and redeposited. But in most small
streams there is a zone in the valley downstream from which the al-

(q
Or,
Le

ee
= — ——
LATEST Ol SS =

Fig. 6.—Successive longitudinal profiles of a valley, showing formation of a pay-
streak progressively upstream, with overlying alluvium omitted. (Vertical scale greatly
exaggerated.)

luvial material on or near bedrock will not be further disturbed, even
at the highest flood stages, unless the stream is rejuvenated by a
lowering of its base level of erosion. The position and length of this
critical zone varies with the strength of the current, the size and spe-
cific gravity of the alluvial materials, and with several other factors;
yet its existence is fairly well substantiated. If a gold lode occurs at or
near the head of a valley, the gold on being liberated by the process
of weathering migrates downstream with the other stream detritus,
gradually working its way toward bedrock. Somewhere in the critical
zone, however, most of this gold, and all the coarse gold, finally comes
to rest; and from this zone downstream the current of the stream is
slower, and the detritus becomes thicker, so that the stream can no
longer erode to bedrock. This critical zone, which lies between the
headwater stretch of intermittent movement of all debris and the
downstream stretch of no movement of the debris near bedrock,
marks the downstream terminus of the paystreak; but the gold in
process of downstream migration is also present upstream from the
critical zone, and such gold may or may not constitute a paystreak,
depending upon various factors. But stream erosion is a continuous
process, in the course of which the valley is either extended backward
into its divide; or, if another headwater stream is flowing in the op-
posite direction, the divide between the two streams will be lowered.

Mar. 15, 1940 MERTIE: PLACER GOLD IN ALASKA eT,

In either case, the net result is a change in the longitudinal profile of
the stream bed, such that the headwater gradient is diminished and
the critical zone of deposition migrates slowly upstream. Hence that
stretch of gold placers, no longer subject to downstream movement,
is lengthened, and a paystreak is deposited progressively upstream.
The concept thus results that the part of a placer farthest downstream
was deposited first and that the formation of the paystreak took
place progressively upstream, as a series of overlapping wedges. This
mode of paystreak formation is particularly applicable in areas where
the local base level of erosion has remained sensibly constant over a
long period of time; and such conditions have apparently obtained
during certain erosional cycles in parts of interior Alaska.

If a paystreak was derived from a lode located in the headwater
part of a valley, and if it was formed in the manner above outlined, it
should be expected that most of the placer gold will be formed near,
on, or in bedrock. This is actually true for most of the placer streams
of Alaska, the gold being found in the lowermost few feet of gravel,
on the surface of bedrock, and if the latter is greatly fractured to a
depth of as much as 6 feet in bedrock. Some of the early dredge opera-
tors in Alaska failed to recognize the depth to which gold can pene-
trate in bedrock; and as a result of this, and also of inadequate
washing in the trommel, some of the old dredging sites are now being
reworked at a good profit. The absence of this localization of the gold
becomes immediately a reason for searching a valley for uneroded lode
sources not in the headwaters, or for postulating a rapidly changing
local base-level of erosion during the deposition of the gold, or after
a part of it had been deposited. One of the most striking examples of
gold that is not concentrated near bedrock, is found in the placer
streams which derived their gold from the Tertiary sedimentary reefs,
above described.

It is obvious that this idealized mode of placer accumulation, which
stresses the lack of geologic simultaneity in the formation of a
paystreak, may be modified in many ways, not merely by a succession
of erosional cycles, but also by conditions and events that may exist
within a single cycle of erosion. The theory, as sketched, applies
particularly to placers that accumulate from lodes that are localized
in or near the headwater portion of a valley, and many examples of
such conditions in interior Alaska could actually be cited. But the
bedrock source of gold is not always thus localized, as for example
where the locus of a lode system is more or less coincident with a
valley, or where mineralized zones occur intermittently, crossing the

118 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

valley at different places. Likewise there may be present proximate
sources of gold, other than bedrock gold, such as auriferous bench
gravels of any origin, which are distributed along the sides of a valley,
or auriferous gravels or conglomerates which cut across valleys. All
such conditions, and combinations of them, tend either to modify or
to render entirely inapplicable the idealized concept; and many such
examples are known in interior Alaska where the general hypothesis
does not apply, even within a single erosional cycle.

Summarizing, it needs to be stressed that no general hypothesis of
placer accumulation can be presented. The volume of available de-
bris, derivable from bedrock, is affected by climatological conditions,
past and present; and such conditions also control the volume of water
formerly, and at the present time, existing in valleys. The character
of the alluvial deposits is likewise affected by climatological condi-
tions, as for example in interior Alaska where these deposits are
frozen to great depths. The velocities and erosive power of streams,
on the other hand, are functions that depend upon many variables,
among which are the volume of water, the transported load and the
valley gradients. And finally, the valley gradients, though locally
influenced by the character of bedrock and other factors, are in large
measure controlled by the duration, changes, and accelerations in
local base levels of erosion. Some of these data can be deduced or in-
ferred from geologic studies; others can not. Hence, the history and
mode of formation of placer deposits in a region constitute a series
of individual, yet related problems, which are seldom completely
solvable.

THE PROBLEM OF FINENESS

The fineness of lode and placer gold is an economic factor of con-
siderable significance. Gold that is 900 fine, for example, yields a
profit 20 percent greater than gold having a fineness of 750; and since
many examples could be cited of finenesses of this order, the illustra-
tion is by no means overdrawn. In general, therefore, the matter of
fineness has been approached from a purely economic rather than a
genetic point of view; and thousands of assays have been made of
Alaska gold, with few attempts to correlate and to understand these
significant data.

The genetic problem of fineness has several aspects, of which the
most general has to do with the range and limits of the ratios of gold
to silver in all the natural alloys of these metals. A second phase of
the problem is concerned with the recognition and explanation of
variations in the grades of lode and placer gold, and this veers into’

Mar. 15, 1940 MERTIE: PLACER GOLD IN ALASKA JAY)

and is really a specialized part of the theory of placer genesis. Still
another phase of the problem is what might be called a problem of
fineness of lower magnitude, dealing with the variations in fineness
within individual nuggets, grains, and crystallites of gold-silver alloys;
and this in turn leads to a physical-chemical study of free gold.

It has already been shown that artificial alloys of gold and silver
form a continuous series of solid solutions, with all possible composi-
tions from pure gold to pure silver. The natural alloys of gold and
silver, however, seldom contain more than 40 percent and never
more than 60 percent of silver; whereas at the other end of the system,
native silver is usually free of gold, and seldom contains more than
traces of it. Therefore, it is possible that a miscibility gap exists in the
solid state of the gold-silver system, as developed in nature. It has
also been shown that the specific volumes of the natural alloys of
gold and silver depart materially, and in a reverse manner, from the
specific volumes of the corresponding artificial alloys. Hence, it ap-
pears doubtful that the natural gold-silver alloys are altogether solid
solutions; or, if they are, the presence of a small percentage of base
metals in the dross has produced some remarkable atomic readjust-
ments. Moreover, since most geologists believe gold-quartz veins to
be of hydrothermal origin, what basis is there for assuming that gold
and silver will crystallize in the presence of silica, water, base metals,
and mineralizers to produce a solid phase identical with that which
solidifies from dry melts of gold and silver alone? And finally, is it not
possible that allotrophic modifications of gold and silver may crystal-
lize under the conditions of cooling formulated by geologists? These
questions will be answered only by laboratory work on specimens
of the natural gold-silver alloys; and such studies will probably have
to include the investigation of many physical properties of such alloys,
both on macroscopic and microscopic scales, in addition to chemical
and thermal analyses.

High-grade placers can be developed by the partial erosion of high-
grade lodes, thus making it possible to interpret preexisting bedrock
conditions in the light of the present bedrock; yet this condition
seldom obtains, because high-grade lodes are rare. On the other hand,
high-grade placers may also be produced from low-grade lodes, but
under such conditions the life history of the placers is so extended
that the original lodes may be largely or completely removed by
erosion. At some localities the roots or basal parts of the lodes may
still remain, but the contained ores may be quite different from the
medial and apical horizons that have been removed. Hence, it is

120 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

seldom that observations on bedrock lead to a complete understand-
ing of the character and genesis of the preexisting ore deposits that
served as the sources of the placers.

Few data have been collected in Alaska that bear upon the char-
acter of the eroded lodes, but one generalization has been adduced,
partly from theoretical reasons and partly from observed relation-
ships, that seems to bear upon the distribution of fineness in placers.
It is probably the usual, rather than the unusual, condition for the
fineness of gold to vary in different parts of a lode. Pertinent data on
this point are lacking, partly because the depth of many gold mines
is less than the thickness of rocks eroded to produce the present
placers, and partly because this topic has not received the con-
sideration it could have received in our deep mines. But since the
primary ores of the precious metals are known in some mines to give
place at depth to base ores, it is a reasonable hypothesis that free gold
itself, in the apical horizons of an original ore body, may be of higher
fineness than that which occurs at considerably greater depths, as a
result entirely of hypogene processes. The apical enrichment of lode
gold by supergene processes can hardly be questioned at some locali-
ties.

Gold lodes have been either wholly or partly eroded to produce
their derived placer deposits. Let it be assumed that the apical por-
tions of lodes contained gold of higher grade than the lower horizons.
Then if they have been wholly eroded, and if no enrichment of the
gold has occurred during or after the transformation of the lode gold
to placer gold, the average fineness of the placer gold should equal
approximately the average fineness of the preexisting lode gold. But
if the lode deposit has been only partly eroded, as is usually true, then
the average grade of the gold in any one paystreak should be higher
than that of the gold in the uneroded part of its antecedent lode.
Moreover, and regardless of the degree of erosion, it follows that the
average fineness of all the gold recovered from existing lodes should
be less than that of the average fineness of all placer gold; and this is
actually true. It should also follow from these considerations that
gold of lower grade could be found in existing lodes than in placers;
and this is likewise true, as little or no gold having a fineness of less
than 600 has been found in placers, whereas lode gold has been found
to have a fineness as low as 400.

In addition to variations in the primary fineness of gold within a
preexisting lode system, it is probable, at least in interior Alaska, that
a zone of oxidation and enrichment has existed continuously from the

Mar. 15, 1940 MERTIE: PLACER GOLD IN ALASKA 121

time that these lode systems were bared to erosion to the present day.
If surficial enrichment within alode system has operated to increase
the fineness of lode gold, then in a country free from glaciation, this
has been a continuous process to which all the lode gold was sub-
jected, though perhaps in varying degrees, before it was liberated
from its bedrock sources. This process of enrichment may, and in
some areas certainly has, operated to produce marked differences in
the grades of lode gold and placer gold derived therefrom; but unless
this process varies greatly in intensity from one geologic epoch to
another, it is improbable that it has been more than a minor factor
in producing variations in the grade of the gold within any one pay-
streak.

One of the best examples known in Alaska of a great divergence
between the lode gold and the placer gold derived from it, has been
described by the writer!’ in the Nixon Fork district. Here occurs a
quartz monzonite of Tertiary age, which has been the bedrock source
for gold placers in the streams draining from it; yet the roots of the
lodes are still preserved, and are being actively mined as gold lodes.
From the records of thousands of ounces of this lode gold, its average
fineness is known to be 735, with maximum and minimum values,
respectively, of 781 and 715; yet in Hidden Creek, which drains out
of the lode area, the maximum, minimum, and mean finenesses are, re-
spectively, 9612, 8924, and 928 parts gold in a thousand. It should be
stressed also that at this particular site the fineness does not increase
progressively downstream; and although most of the paystreak that
has been mined is underlain by a bedrock of limestone, nevertheless
this paystreak extends upstream into the zone of quartz monzonite
bedrock. Certainly no better example than this could possibly be
found to prove that the gold eroded from the apical part of this lode
system was of higher grade than that now being mined in the roots
of the lode; but such conditions indicate an enrichment of the alloy
in gold in the zone of weathering, before it was liberated from its
bedrock source. In other words, as no marked variation of fineness is
known within the paystreak, this locality may not be cited as an
example of primary differences in the grade of the lode gold.

The general accepted theory has been stated, which pictures the
paystreak as a series of overlapping wedges of gold-bearing alluvium,
which are progressively deposited upstream during a single erosional
cycle. According to this view the downstream end of a paystreak is

17 Mertis, J. B., Jr. Mineral deposits of the Ruby-Kuskokwim region, Alaska. U.S.
Geol. Surv. Bull. 864: 193-194, 229-242. 1934.

122 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

its oldest, and the upstream end is its youngest part, if the deposition
has taken place during a single cycle of erosion. A hypothesis has also
been stated, which assumes that the apical part of a gold lode contains
gold of higher grade than its lower horizons. Taken together, these
two concepts lead directly to the conclusion that the gold of highest
grade is likely to be found in the downstream end of a paystreak, and
that the grade of the gold may diminish progressively upstream. This
relationship actually exists in some placer paystreaks in Alaska, but
in others the fineness has been observed to change erratically, or not
at all.

The progressive increase in the fineness of placer gold, in going
downstream, has heretofore been explained as due to the removal of
silver from gold-silver alloys as a result of solution by cold surface
waters, during or after the formation of a paystreak. According to this
hypothesis, the gold farthest downstream has traveled a greater
distance from its parent lode than the gold farther upstream. There-
fore it has suffered the most handling by streams, has been to the
greatest degree comminuted, and for both these reasons has been the
most vulnerable to solution, Also, it has been longest separated from
a bedrock source, and for this reason, too, should have been most af-
fected by solution. This hypothesis is further supported by an experi-
ment performed by McConnell,'® on some of the placer gold of the
Klondike district, in Yukon Territory, Canada. He found that gold
shaved from the outer surface of a nugget assayed 60 to 70 parts per
thousand finer than gold from the inside of the nugget; and this has
been generally accepted as a proof that surface waters dissolve an ap-
preciable amount of silver from gold-silver alloys in a paystreak. Let
us look at the supporting data.

It has already been shown that the silver contained in artificial
alloys of gold and silver can be completely dissolved by strong in-
organic acids, only when the ratio of gold to silver is 1:3 or less; also
that if this ratio is greater than 1:1.5, corresponding to a fineness of
400 parts gold in a thousand, no silver will be dissolved. It has also
been shown that the solubility of silver in pure water is very slight.
But practically all placer gold is of higher grade than 600 fine, and
moreover we are dealing with ordinary cold surface waters, instead
of strong inorganic acids. Finally, it must be remembered that any
gold that has remained undisturbed in the placer paystreaks of in-
terior Alaska since the beginning of Pleistocene time has probably

18 McConngELL, R. G. Report on the gold values in the Klondike high-level gravels. |
p. 979. Geological Survey of Canada, 1907.

Mar. 15, 1940 MERTIE: PLACER GOLD IN ALASKA 123

been entirely untouched by water for the last million years, because
these alluvial deposits were then, as now, largely frozen.

Still other facts have to be considered. The assayer knows that even
strong inorganic acids do not readily remove all the silver from large
pieces of a gold-silver alloy; and for this reason, a sample of the alloy,
after quartation, is hammered flat on an anvil before treatment with
nitric acid, in order to present a large surface to solution. Hence, the
weak solvents postulated to exist in cold surface waters should affect
only the outer layers of grains of placer gold; and for this reason the
surficial volume of nuggets and coarse grains of gold, in which solution
of silver might occur, is a smaller proportion of the total volume than
in small grains of placer gold. It, therefore, follows, insofar as enrich-
ment by solution of silver is concerned, that nuggets and coarse grains
of placer gold should be of lower grade than the finely comminuted
grains of placer gold. Actually, in any one paystreak, and at any one
place in the paystreak, the reverse is usually true. Furthermore, when
any appreciable amount of silver is dissolved in the laboratory from
a gold-silver alloy, the sample becomes distinctly porous; and if much
of or all the silver is removed, the sample becomes very fragile and
may even crumble to a powder. Therefore, if solution of silver is
actually accomplished by cold surface waters to an extent sufficient
to change appreciably the fineness of placer alloys, this process should
be reflected in a marked surficial porosity. But this also has not been
observed.

As for McConnell’s experiment, nothing is proved except that the
outer surfaces of certain nuggets were of higher grade than the inner
parts. It does not at all follow that this relationship is due to solution
of silver by cold surface waters, for either it may have been an original
characteristic of the primary lode gold, or, more probably, it was
caused by surficial enrichment in the zone of oxidation, long before
the gold was liberated from its bedrock source.

As a result of these considerations, the writer is not disposed un-
reservedly to accept the idea of any progressive change of fineness in
a placer paystreak, as a result of solution of silver from gold-silver
alloys by the action of cold surface waters. But it must be admitted
that the alternate hypothesis is also unproved. In the first place it
rests upon another hypothesis regarding the vertical distribution of
different grades of gold in preexisting and present lodes. And secondly,
it rests upon physical-chemical data that assume an essential identity,
or at least the great similarity, of artificial and natural alloys of gold
and silver, in so far as their chemical reactions are concerned. Hence,

124 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

additional geologic, mining, and chemical data will be needed to prove
or to disprove the hypothesis above outlined. If, for example, the em-
placement of a granitic body long antedated the formation of its as-
sociated lode deposits, and if a period of diastrophism had intervened
between these two processes, the apex of the intrusive mass and the
apex of the mineralized zone would not necessarily correspond. And
if diastrophism had occurred after either or both of these processes
had occurred, the present cropping of an intrusive might correspond
to neither of these antecedent apices.

Mining and chemical data are required, most of all to learn the
three-dimensional variations in fineness that may exist in present gold
lodes. Assays, both of bulk samples and of individual grains, should
be made of uncontaminated samples of free gold taken from different
horizons in lodes; and for this purpose assays of run-of-mine bullion
may or may not suffice, depending upon the methods employed in re-
covering the gold. Assays, for example, of bullion recovered by cy-
anidization would certainly be useless. Complete chemical analyses
should also be made, in order to learn the character and quantity of
the metals in the dross; and for this purpose, even amalgamated free
gold would not be serviceable.

Finally, the physical and physical-chemical properties of natural
gold-silver alloys need to be studied, in order to learn how they differ
from those of the corresponding artificial alloys. One of the most im-
portant problems of this work should be a complete physical and
chemical examination of individual grains and nuggets of natural
gold. According to Raydt’s equilibrium diagram, it is possible, if the
solidification of gold took place rapidly enough, that individual crys-
tals of gold could be zonally grown, in the manner of the plagioclase
feldspars. If this occurs, and if his equilibrium diagram applies to
natural alloys, the outer zones of crystallites should be lower in gold
than the cores. Such a condition could hardly influence materially the
range of fineness in a placer paystreak, as it would be a microscopic
phenomenon, of a lower order of magnitude. But the investigation of
this and related phenomena is equally a part of the general problem
of fineness.

Mar. 15, 1940 EVERITT AND SULLIVAN: ACTION OF SULPHUR COMPOUNDS 125

CHEMISTRY.—The fungistatic and fungicidal action of certain or-
ganic sulphur compounds.! Epwarp L. Everitt, Georgetown
University School of Medicine, and M. X. SuLuivan, Chemo-
Medical Research Institute, Georgetown University.

Both bacteria and molds are in general useful to man, as for exam-
ple in decomposing plant and animal debris, which, if allowed to ac-
cumulate, would sooner or later be to man’s detriment. In both fields,
certain forms have become parasitic on man or on the fruits of the
field on which man depends directly or indirectly for food and cloth-
ing. The part that bacteria may play in the production of pathological
condition has become common knowledge and has long been under
study by the bacteriologist and the practitioner of medicine.

On the other hand, the relation of common molds to health and
disease has been given less attention and study since, unlike the bac-
teria, molds do not in general cause acute and killing diseases, in man
at least. However, Kuchenmeister (1857) lists 14 species of fungi that
invade the skin, hair, nails, lungs, or mucous membranes. In the list
are included Tricophyton tonsurans, which invades the hair follicles,
and several species of Aspergillus, which infect the auditory duct,
finger nails, toe nails, etc. Also ringworms caused by fungi of the
Tricophyton or Microsporum families are often met with in man.

In medical practice attention is often called to Moniliae, fungi that
invade the mucous membranes, especially of the mouth. Among the
diseases of man attributed to this class of fungi are thrush and sprue.
There is some suggestion also that fungi, at times at least, may be
involved in such allergic conditions as asthma and hay fever. Thus,
Prince, Selle, and Morrow (1935) report findings that indicate that
molds may play a causative role in some cases of asthma .and hay
fever, while Brown (1936) considers that hypersensitiveness to fungi
must take its place along with sensitization to pollens, animal epi-
dermis, food, and bacteria in the causation of bronchial asthma,
eczema, perennial hay fever, and other allergic conditions.

Since molds may destroy the necessary fruits of the field, causing
rots of various kinds, and may also invade the skin, the hair, and
lungs of man and are incriminated in certain forms of dermatitis, ring-
worms, actinomycosis of the lungs, loss of hair in some cases, and oc-
easionally in certain allergics, whatever can be found about the
metabolic processes and especially about means of preventing them

1 The data in this paper are taken from the dissertation presented by Edward L.

Everitt in partial fulfillment of the requirements for the degree of Doctor of Philosophy,
Georgetown University, 1937. Received January 20, 1940.

126 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

from getting a foothold on plants, animals, or man would be decidedly
worth while.

It is well known that molds and bacteria may be killed or inhibited
in their growth by drying, heat, light, pressure, and by various chem-
icals. Chemical substances that kill are known as fungicides or bac-
tericides, while those that simply inhibit growth are called fungistatic
or bacteriostatic. Some of the chemical compounds used in the pre-
vention of the growth of bacteria and molds are silver nitrate, mer-
curic chloride, iodine, phenol, cresol, benzoic acid, alcohol, and vari-
ous simple sulphur compounds. Among the sulphur compounds used
are carbon disulphide, hydrogen sulphide, sodium thiosulphate, flow-
ers of sulphur, and colloidal sulphur. Lime and sulphur mixtures have
been used for years to prevent the growth of fungi on plants and trees.

Our interest in the possibility of inhibiting the growth of molds
that invade man was aroused by results obtained in a study of the
changes brought about by certain wilt-producing organisms. In a
study of acid-base and oxidation-reduction phenomena, it was noted
that sodium thiosulphate added to the medium considerably reduced
the growth of the molds. This finding suggested a trial of various
organic sulphur compounds because elemental sulphur had long
been in use in medical practice in ointments for eezema and various
skin conditions and parasitic skin infections, and sulphur baths have
long been believed to have medicinal value. A direct bearing on the
relation of sulphur to fungi is the work of Lynch (1933), who reports
the successful use of a sulphur ointment in the treatment of an As-
pergillus infection in a scalp lesion caused by the bite of the red bug
(Leptus), and the review of Roark and Busbey (1935), who list a
number of organic sulphur compounds of high value as insecticides.

As pointed out by Roark and Busbey, sulphur in various forms and
combinations is one of the most valuable and widely used insecticides
and fungicides. In the form of elemental sulphur it is applied to fruit
trees and ornamental plants both as a dust and in suspension in water
for combating red spiders and fungous diseases.

We had on hand a large number of organic sulphur compounds
made or secured in a general study of the relation of sulphur and sul-
phur compounds to health and disease. Some of these we hoped would
be of value in medical and agricultural practice.

Accordingly, some 50 sulphur compounds were tested for their
fungicidal action in vitro. The molds used were Fusarium oxysporum
and F’. lycopersicum, which cause, respectively, wilt of potato plants
and tomato plants; Aspergillus fumigatus, which invades the ear of

Marz. 15, 1940 EVERITT AND SULLIVAN: ACTION OF SULPHUR COMPOUNDS 127

man and occasionally the lungs; Aspergillus niger, which spoils food-
stuffs and like fumigatus may invade man; and the Penicillium of
Fleming, which according to Reid (1935) generates a material toxic
to bacteria. The culture media used and the procedure employed for
testing the inhibitory action of the various compounds are detailed in
the following sections.

The culture medium—The synthetic culture medium used in this investi-
gation was the same as used by Anderson, Everitt, and Adams (1933) in
their study of the carbohydrate metabolism of Fusarzwm oxysporum, which
causes wilt of potatoes. It was first used by Tochinai (1920) to study the
carbohydrate metabolism of Fusarium lint. The composition of the medium
is as follows:

AUMIMONIUIN MbLAtCE yee asec as. oe 1.00 gm
Miagmesiumusulphatves aes an soi - 0.25 gm
Monopotassium phosphate.......... 0.50 gm
GUICOS CRE sect ice ecel icode an neare wants. che 20.00 gm
Wraterbosmakes:. 0s sees was 1,000 cc

In the present work, to 100-cc portions of this culture medium in 250-cc
Erlenmeyer flasks various quantities of the organic sulphur compounds were
added, and the flasks were plugged with cotton and sterilized at 15 pounds
pressure for 20 minutes. Control flasks containing the medium without addi-
tion of the sulphur compounds were sterilized in a similar manner.

Inoculation of the medium.—The stock medium for the development of
the molds was Sabouraud’s dextrose agar described in Difco Manual, ed. 5,
1935. Spores collected from the agar slants were suspended in sterile distilled
water, and 1 cc of the spore suspension was added to each culture flask by
means of a sterile pipette. The sulphur compounds tested are listed in Table
ile

Compounds (1)—(18) were obtained from Dr. H. L. Haller, Bureau of
Entomology and Plant Quarantine, U. S. Department of Agriculture. The
remaining compounds were at hand at Georgetown University.

Inasmuch as most of these compounds were ineffective, that is, did not
inhibit the growth of the molds, the formulas are not given here. Those
compounds that were effective will be discussed in detail and their formulas
will be given later.

With Fusarium oxysporum and F. lycopersicum used as test fungi, only a
few of the compounds listed in Table 1 inhibited or stopped growth. The in-
hibitors were Nos. (10), (11), (83), (84), (86), and (47). These compounds in-
hibited growth for a period varying from 4-15 days, after which time the
organism slowly developed. These compounds could be utilized as fungistatic
material and might have application in medical practice, in external applica-
tion. One compound, (24), the disulphide of ortho-thioaminophenol, abso-
lutely prevented growth of Fusarium oxysporum but allowed a slight and

128 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3
delayed growth of Fusarium lycopersicum with increase in growth after the
nineteenth day. Colloidal sulphur, (83) (sulphur diasporal), was labeled as
containing 10 mg of 8 in 2 ce. One ce was added to 100 ce of the culture me-
dium. Colloidal sulphur (sulisocol) was labeled as 2 cc containing 20 mg of
sulphur. One-half of 1 ce was added to 100 cc of culture media. Both types of
sulphur in a colloidal complex markedly inhibited the growth of the molds,
even at concentrations calculated to be 50 mg per liter. Since a question arose

TABLE 1.—SULPHUR COMPOUNDS STUDIED FOR FUNGICIDAL AND
BACTERICIDAL PROPERTIES.

(1) Phenothiazine-6-carboxylic acid | (26) Thiazolidine carboxylic acid
chloride (27) Formy!-dl-cystine

(2) 6-Acetyl-phenothiazine (28) S-carboxymethyl-cysteine

(3) Phenothioxine (29) Cystineamine hydrochloride

(4) Phenothiazine (30) Thiazolidine hydrochloride

(5) Tetrathiopentone (31) Thiobarbiturie acid

(6) Cuprous methylxanthate (32) Benzyl disulphide

(7) Cuprous isoamylxanthate (33) Colloidal sulphur (diasporal) 2 ce =

(8) Chlorbenzoketothiazine 10 mg

4-Chloro-2-nitrophenyl thioglycolic
acid

Colloidal sulphur (hyposols or suli-
socol) 2 ce=20 mg

(10) Phenyl thioarsenite (35) Sulphanilic acid

(11) 4-Chloro-2-nitrophenyl sulphur | (36) 1,2 Naphthoquinone-4-sodium sul-
amine phonate

(12) 4-Chloro-2-nitrophenyl sulphur | (37) 1,2 Naphthoquinone-4-6-scdium di-
bromide sulphonate

(13) Bis(2-nitrophenyl) disulphide (38) Sodium alizarine sulphonate

(14) Thioacetamide (39) Tropaolin 000

(15) Mercaptobenzothiazole (40) Congo red

(16) Phenylbenzothiazole (41) Bromcrescl green

(17) Sodium diethyldithiocarbamate (42) Methyl orange

(18) Diethyl thiourea (43) Cresol red

(19) Thiourea (44) Bromphenol blue

(20) Disulphide of thiotyrosine (45) Strychnine sulphate

(21) Dithiosalicylic acid (46) Thymol blue

(22) Phenylthioglycolic-ortho-carboxylic | (47) Prontylin (Winthrop)
acid (48) Trional

(23) Benzidine sulphonate (49) Sulphonal

(24) Disulphide of ortho-thioaminophe- | (50) Cystine

(51) Sulpharsphenamine

no
Cysteic acid

as to whether the inhibiting action was due to the sulphur as such or to the
changes in the reaction of the medium, the study of the inorganic sulphur in
the protective colloid solution was put aside for later development. Two
compounds, mercaptobenzothiazole and phenylbenzothiazole, allowed no
growth whatsoever of the two molds mentioned above when present in the
culture media at the concentration of 5-10 mg in 100 cc of solution. Because
a number of the organic sulphur compounds were found effective in inhibit-
ing or utterly preventing the growth of the two molds, the experiment was
extended to other molds as given in Table 2.

An interesting effect of chemical constitution on the growth of the molds
was exhibited by compounds (36) and (37), namely, 1, 2 naphthoquinone-4-
sodium sulphonate and 1, 2 naphthoquinone-4-6-sodium disulphonate, with
the formulae—

-Mar. 15, 1940 EVERITT AND SULLIVAN: ACTION OF SULPHUR COMPOUNDS 129

|
aon
| | and
SO;Na

|

Na0,S— |

SO,Na

The 1, 2 naphthoquinone-4-sodium sulphonate had marked power of in-
hibiting the growth of the molds, while the 1, 2 naphthoquinone-4-6-sodium
disulphonate was an excellent stimulator of growth. These findings are in
harmony with the early work of Ehrlich and Herter (1904), who found the
sodium salt of the 1, 2 naphthoquinone monosulphonic acid to be toxic.
Since these compounds are relatively expensive and since the work of Ehr-
lich and Herter, especially that of Herter (1905), indicated that the 1, 2
naphthoquinone-4-sodium sulphonate was toxic to animals, no further at-
tention was given to the compounds as fungistatic or fungicidal substances
for use in medical or agricultural practice.

Of the compounds studied in relation to the growth of the molds in vitro,

TABLE 2.—StTupDY OF THE Most PROMISING SULPHUR COMPOUNDS HAVING FUNGISTATIC

Com-
pound
No.

10

11

15

47

47

Amount

used

12.5

12.5

10.0

10.0

25.0

F. oxysporum | F. lycopersicum

First growth
in 15 days,

First growth
in 15 days

Growth in 8
days, becoming
heavy

No growth at |

any time

No growth

Slight growth
in 8 days, be-

| coming heavy
| No growth

| No growth

No growth in
11 days, some
spores in 15
days, slowly in-
creasing

Slight growth
in 4 days

No growth in 4
days, slight
growth in 12
days

|

| No growth in
| 25 days
slow thereafter |

First growth

in 15 days

Slight growth
in 8 days, be-
coming heavy
No growth at
any time

No growth
Slight growth

in 8 days, be-
coming heavy

No growth
No growth
No growth in

25 days

Slight growth
in 8 days

Slight growth
in 7 days

|

|

A. niger

No growth in
11 days, slight
in 25 days

No growth in
11 days, slight
in 15 days

Good growth in
8 days, very
few spores

Slight growth
in 8 days, no
spores

No growth
Heavy growth

in 8 days, with
spores

| Slight growth
/in8 days

No growth in

25 days
Good growth in

| 8 days

Slight growth
in 7 days

|

A. fumigatus

No growth in
15 days, slight
in 25 days

No growth in
25 days

No growth in
25 days

| No growth in |
| 25 days

|

No growth

Slight growth
in 11 days,
slowly increas-
ing

No growth in
25 days

No growth

No growth in
25 days

Slight growth
in 7 days

AND FUNGICIDAL PROPERTIES WHEN ADDED TO 100-cc CuLTURE MEDIUM.

Penicillium
of Fleming

Very few spores
in 15 days, slow
thereafter

A few spores in
15 days, slight
thereafter

Very slight
growth in 25
days

No growth in 15
days, a few
spores in 25 days

| No growth

Slight growth in
8 days

Slight growth in

days

| No growth in 25

days

No growth in 15
days, slight
growth and a few
spores in 25 days

130 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

number (15), mercaptobenzothiazole, and number (16), phenylbenzothia-
zole, were the most effective. Their chemical constitution, respectively, is—

Vie
een oN oan
Cane

Of these two, mercaptobenzothiazole was on hand in plentiful supply, so
some attention was paid to it from chemical and clinical viewpoints and to
its possible toxicity toward animals. As may be seen from its formula it is an
organic sulphur compound containing nitrogen and an (SH) group. This ~
compound, which is cheap and readily available, was first made by Hof-
mann (1887) and has been used in the rubber industry as an accelerator of
vulcanization for a number of years. Its use for such purposes seems to have
been first suggested by Bedford and Sebrell (1921). It is soluble in alcohol,
chloroform, and benzol but is not very soluble in water. It is sufficiently solu-
ble in water, however, to be used as a germicide. It is more soluble as a so-
dium salt and is rather soluble in sodium bicarbonate.

As judged by lack of growth for 50-60 days in the case of F’. oxysporum,
F. lycopersicum, and A. fumigatus, the compound has marked fungicidal
power. In the case of A. niger and Penicillium of Fleming, mercaptobenzo-
thiazole showed strong fungistatic activity, since no growth occurred until
after a period of 25 days. Without prejudice as to whether a compound can
be found that is fungistatic or fungicidal toward molds in general, it can be
said that with the molds studied by us mercaptobenzothiazole had marked
fungicidal or fungistatic activity.

Roark and Busbey (1935) state that mercaptobenzothiazole in concentra-
tion of 0.01 to 0.10 percent was effective in controlling a fungus living on wood
and that it has been used in controlling aphids and mosquito larvae. Davis
(1930) reported that this compound had little if any toxicity. He injected an
aqueous solution of it into guinea pigs and a total injection of 14.5 mg in 20
days did not produce any injurious effects on the animals. Medical examina-
tion of the men working with mercaptobenzothiazole over a period of years
in the Goodyear Tire & Rubber Co. did not show any toxic conditions or
dermatoses. In a recent personal communication, Dr. Davis (1939) reiterates
the conclusion that the compound has shown no toxic action on men working
with it in the vulcanization of rubber.

We have given 20-100 mg of the mercaptobenzothiazole by mouth to
guinea pigs weighing 400 grams with no gross evidence of toxicity and have
injected 20 mg in aqueous suspension intraperitoneally into a 200-gram
guinea pig with no effect on his activity, appetite, or general well-being. Dr.
William B. Wardrop, of Washington, D. C., found the mercaptobenzothia-
zole practically as effective toward ‘‘athlete’s foot”’ as salicylic acid. The var-

Mar. 15, 1940 EVERITT AND SULLIVAN: ACTION OF SULPHUR COMPOUNDS 151

ious sulphur compounds were also tested for their bactericidal and bacterio-
static properties. The findings will be detailed in a subsequent paper.

SUMMARY

About 50 organic sulphur compounds were tested for their fungi-
static and fungicidal action.

The molds used were the wilt-producing Fusarium oxysporum and
F.. lycopersicum, the pathogenic Aspergillus fumigatus, the common
Aspergillus niger, and the Penicillium of Fleming.

Fungistatic activity was manifested by phenylthioarsenite, 4-
chloro-2-nitrophenyl sulphur amine, 1, 2 naphthoquinone-4-sodium
sulphonate, and prontylin or sulphanilamide.

Fungicidal activity was manifested by mercaptobenzothiazole and
phenylbenzothiazole.

The most effective compound was mercaptobenzothiazole, which
inhibited the growth of the molds in concentration of 50 to 100 parts
per million. This compound is cheap and readily available.

The investigation deals only with in vitro tests, and no conclusion
can be drawn as yet as to the therapeutic application of the various
compounds. Preliminary toxicity tests on guinea pigs with both oral
and intraperitoneal application indicate that mercaptobenzothiazole,
the most effective fungicidal compound, has little if any toxicity.

LITERATURE CITED

ANDERSON, A. K., Everitt, E. L., and Apams, P. D. The carbon metabolism of
Fusarium oxysporum on glucose. Journ. Agr. Res. 46: 473. 1988.

BEpForD, C. W., and SEBRELL, L. B. Reactions of accelerators during vulcanization.
III. Carbo-sulfhydryl accelerators and the action of zinc oxide. Journ. Ind. Eng.
Chem. 13: 1034. 1921.

Brown, G. T. Hypersensitiveness to fungi. Journ. Allergy 7: 455. 1936.

Davis, P. A. Toxic substances in the rubber industry. Pt. XI: Mercaptobenzothiazole.
Rubber Age 27: 2491. 1930. |

Euruicy, P., and Herter, C. A. Uber einige Verwendungen der Naphthochinonsul-
fonsdure. Zeitschr. physiol. Chem. 41: 379. 1904.

Herter, C. A. The color reactions of naphthoquinone sodium monosulfonate and some
of their biological applications. Journ. Exp. Med. 7:79. 1905.

Hormann, A. W. Zur Kenntniss des o-Amidophenylmercaptans. Ber. deutschen
chem. Ges. 20: 1788. 1887.

KUCHENMEISTER, F. Animal and vegetable parasites of the human body, vol. 2. (Trans-
ned from the second German edition by E. Lankester.) Sydenham Society,

BG

Lyncu, K. N. Aspergillus in scalp lesions following red bug (Leptus) bites. Arch.
Dermat. and Syphil. 7: 599. 1923.

Prince, H. E., Sette, W. A., and Morrow, M.B. Molds in the etiology of asthma and
hay fever. Texas State Journ. Med. 30: 340. 1934-35.

Reip, R. D. Some properties of a bacterial-inhibitory substance produced by a mold.
Journ. Bact. 29: 215. 1935.

Roark, R. C., and Bussey, R. L. A list of organic sulphur compounds (exclusive of
mothproofing materials) used as insecticides. U. S. Dept. Agr., Bur. Ent. and Plant
Quar., Div. Insecticide Investigations. 1935.

Tocuinal, Y. Studies on the food relations of Fusarium lini. Ann. Phytopath. Soc.
Japan (3): 22. 1920.

132 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

PALEOBOTAN Y.—Additions to the Pensauken flora.! Epwarp W.
Berry, Johns Hopkins University.

_ In 1935 the writer in collaboration with Alfred C. Hawkins pub-
lished? an account of the plants found in the Pensauken formation in
Middlesex County, N. J. From time to time Dr. Hawkins has sent in
small amounts of additional material, which I have not had time to
study thoroughly until the past summer. The material is in the
same limonitic, coarse, micaceous sandstone as the original specimens,
and the preservation of the plants leaves much to be desired in the
way of details of venation.

The forms identified in the later collections are the following:
Cebatha carolina, Epigaea repens, Fagus americana, Nyssa sylvatica,
Salix humilis, and Viburnum alnifolcwm. Among these the Epigaea,
Fagus, and Nyssa are additions to the Pensauken flora. They add little
in the way of interpretive evidence of Pensauken time.

Epigaea repens Linnaeus

A single relatively small leaf, very coriaceous and with the margins in-
clined to be revolute. So far as I know the trailing arbutus has not heretofore
been found fossil. In modern times its range is from Newfoundland to
Florida in the Coastal Plain, usually on sandy soil. It occurs also beyond the
limits of this province.

Fagus americana Sweet

This species, represented by leaves or fruits, is exceedingly abundant in
the Pleistocene of southeastern North America, having been recorded from
11 States from Massachusetts on the north to Mississippi on the south,
chiefly in river terraces in the Coastal Plain province, although also recorded
from cave deposits in Pennsylvania and interglacial deposits in West Vir-
ginia.

Nyssa sylvatica March

A typical leaf of this species. Stones of this species have been recorded
from the Pleistocene of both Maryland and the District of Columbia. In the
modern flora it ranges, in rich, mostly swampy soil, from Maine and Ontario
to Florida and Texas.

There is some confusion among our recent species as well as among the
Pleistocene records. Two species, Nyssa caroliniana Poir and Nyssa unzflora
Walt., are recorded from the Bridgeton sandstone of southern New Jersey
and stones referred to Nyssa biflora have been recorded from the Pleistocene
of New Jersey (Fish House), Maryland, Virginia, North Carolina, and Ala-
bama. Probably some of these should really be referred to Nyssa sylvatica.

1 Received October 11, 1939.
2 BERRY, Epwarp W. , and AuFRED C. Hawkins, Flora of the Pensauken formation
in New Jersey. Bull. Geol. Soc. Amer. 46: 245-252. 1935.

Mar. 15, 1940 HOERNER: NOTE ON PSEUDOPERONOSPORA 133.

BOTANY.—A nomenclatorial note on Pseudoperonospora.! G. R.
HorERneER, U. 8. Bureau of Plant Industry. (Communicated
by JoHN A. STEVENSON.)

Miyabe and Takahashi,” in describing a new species reported for
the first time on hops, state:

This fungus with its dichotomously branched conidiophores and with its
conidia germinating by zoospores shares the characters of two genera
Peronospora and Plasmopara, and may form an intermediate genus together
with the downy mildew fungi of cucumber and Celtis. In 1901, Berlese’
created a new subgenus, Peronoplasmopara, in the genus Plasmopara and
placed under it Peronospora cubensis Berk. et Curt., and Peronospora
Celtidis Waite. Rostowzew‘ reported in 1903 a detailed account of his
study of different species of the Peronosporaceae, but especially of Perono-
spora cubensis B. et C., on which he founded a new genus Pseudoperonospora,
apparently without knowing the above mentioned work of Berlese.

In 1904, Clinton’ made a critical study of the cucumber mildew and
came to a similar conclusion in regard to the recognition of its generic
position. But regarding Berlese’s subgenus, Peronoplasmopara, because of
priority of publication, and also because it was given definite descriptive
characters as deserving preference over Rostowzew’s rather indefinite
Pseudoperonospora, Clinton elevated Peronoplasmopara to generic rank and
included under it two species, Peronoplasmopara cubensis (Berk. et Curt.)
Clint., and Peronoplasmopara Celtidis (Waite) Clint.

In the present paper, we have adopted Clinton’s view and name our
fungus Peronoplasmopara Humuli Myb. et Tak., thus adding another
species to this interesting genus.

However, on the basis of priority, if present generally accepted
rules of nomenclature are followed, it appears that Rostowzew’s name
for the genus is still valid. In view of the characteristics assigned to
the genus Pseudoperonospora and the validity of this genus name, it
seems necessary to assign this generic designation to three species
that have not hitherto been transferred to it although they clearly
belong there. The following new combinations are therefore proposed:

Pseudoperonospora cannabina (Otth), n. comb.

meee pore cannabina Otth, Mitt. Naturf. Ges. Bern, 1868, pp. 37-88.

Peronoplasmopara cannabina (Otth), Peg. Atti Accad. Naz. Lincei 114
isenro, 26): 620. 1917.

aa Technical Paper No. 327, Oregon Experiment Station. Received December 21,

oon TARE K., and TaKanasui, Y. Trans. Sapporo Nat. Hist. Soc. 1: 153. 1905-
’Buruuse, A. N. Riv. Pat. Veg. 9: 123-126. 1901.

* RostowzEw, 8S. J. Flora 92: 405-430. 1903.

* Cuinton, G. P. Rpt. Connecticut Agr. Exp. Sta. 1904: 329-362. 1905.

134 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 3

Pseudoperonospora elatostemae (Togashi and Onuma), n. comb.

Peronoplasmopara elatostemae 'Togashi and Onuma, Bull. Imp. Coll. Agr.
and For. Morioka 17: 2, fig. 1. 1934.

Pseudoperonospora portoricensis (Lamkey), n. comb.
Peronoplasmopara portoricensis Lamkey, 7n Stevens, Mycologia 12: 52.

PROCEEDINGS OF THE ACADEMY AND
AFFILIATED SOCIETIES

THE ACADEMY
358TH MEETING OF THE BOARD OF MANAGERS

The regular monthly meeting of the Board of Managers of the Academy
was held in the Holmes Room of the Cosmos Club, January 12, 1940. In
addition to the customary reports of the standing committees the Treasurer
presented and discussed an epitome of his annual report. He showed that
despite the publication of the ‘‘Red Book” the balance for the year’s opera-
tions is only slightly red.

The report of the special Committee on Clerical Assistance for the Board
of Editors, of which Joun A. STEVENSON was chairman and H. EK. Ewine
and HERBERT FRIEDMANN members, was read by Secretary N. R. Smirs in
the absence of the committee. The report recommended discontinuance of
the title ‘Editorial Assistant’? but expressed no opinion on the question as
to whether the Board of Editors should be granted or denied assistance.
Following a general discussion on the advisability of permitting the editors
to employ clerical assistance, the report was accepted.

The Custodian of Publications, W. W. Dinu, presented a request from
the Peiping National Library for a free subscription to the JoURNAL for the
current year. This institution pleaded that the continued hostilities in China
which had driven it to Yunnan made it impossible to renew its subscription.
The Board instructed the custodian to grant the request for the year 1940.

The subcommittees on awards of Certificates of Merit for scientific
achievement reported as follows:

E. A. GotpmaAn, chairman of the Committee for the Biological Sciences,
presented the decision of his committee, which selected HERBERT FRIED-
MANN to receive the Award for Achievement in the Biological Sciences for his
work on the parasitic cowbirds and cuckoos and for his studies of avian tax-
onomy and geographic distribution. Dr. Friedmann was born in New York
City on April 22, 1900. He received his B.S. degree from the College of the
City of New York in 1920 and his Ph.D. from Cornell University in 1923.
After a three-year National Research Council fellowship he taught first at
Brown University, then at Amherst College. In 1929 he succeeded Robert
Ridgway as curator of birds in the U. 8. National Museum, a position he
still holds.

H. L. Curtis presented the decision of the Committee on the Engineering
Sciences, of which he was chairman. For the award in this field PAUL ALBERT
SmitH was selected for his contributions to topographic surveys of the ocean
bottom along the eastern coast of the United States. Mr. Smith was born at
Rising Sun, Iowa, January 9, 1901. He received his B.S. degree at the Uni-
versity of Michigan in 1924. In August 1924 he joined the staff of the U.S.
Coast and Geodetic Survey and has remained with that organization.

Mar. 15, 1940 SCIENTIFIC NOTES AND NEWS 135

H. D. Miser presented the decision of the Committee for the Physical
Sciences, of which J. F. Coucu was chairman. For this award W1iLtmMot HypE
BRADLEY was selected for his investigations of varves in the Green River
formation of Wyoming and his contributions to the geology of abyssal sedi-
ments. Dr. Bradley was born at New Haven, Conn., April 4, 1899. He re-
ceived his Ph.B. degree at Yale University in 1920 and his Ph.D. in 1927.
During his summer vacations he worked as field assistant in the U. 8. Geo-
logical Survey, and at the completion of his academic work he joined the
staff of that organization, where he has remained.

After a discussion of the age limit, in which it was agreed that investiga-
tors eligible for honors are those who have not attained their forty-first birth-
day at the close of the year for which the awards are made, the Board voted
without division to award Certificates of Merit to the above three nominees.
Following a brief consideration as to how the Certificates evidencing this
honor accorded by the AcapEMy were to be presented—which was without
conclusion—the Board adjourned to resume its discussions informally over
a surprise buffet supper provided by the retiring President.

The attendance of Board members follows:

Present Absent
C. E. CHAMBLISS (Pres.) O. S. ADAMS (Rec. Sec.)
N. R. SmitH (Corr. Sec.) J. F. Coucu (Board Member)
H. G. AVERS (Treas. ) Neiut M. Jupp (Board Member)
F. G. BRICK WEDDE (Board Member) H. C. FULLER (Board Member)
J. H. HIBBEN - (Board Member) H. S. GRAVES (nonresident V.P.)
G. STEINER (Board Member) R. B. SosMAN (nonresident V.P.)
Henry B. Co.uuins, Jr. (An) W. B. BELL (B)
A. WETMORE (G) B. H. Nicotet (C)
H. D. MisER (Gl) AVE CLARK (En)
W. A. Dayton (F) FRED O. Cor (M)
H. L. Curtis (Ke) ALLEN C. CuarKk (H)
H. L. WittEMORE (Me) G. F. Gravatt (B)
KE. W. PRIcE (Hl) ALES HrpuicKa (An)
F. D. Rossini (Sr. Ed.) P. C. Wuitnry (5B)
W. W. Diets (Custodian) L. A. RoGERs (Ba)
Wo. BowliE (M1)

J. H. DELLINGER (R)

In addition to the above members there were present by invitation F. M.
SETZLER, chairman of the Meetings Committee; EK. A. GOLDMAN, chairman
‘of the Subcommittee on Awards for the Biological Sciences; and the editors,
C. L. Gazin and J. H. Kempton.

SCIENTIFIC NOTES AND NEWS

The Department of State has notified the AcapEMy that the Eighth
American Scientific Congress will be held in Washington, D.C., from May
10 to 18, 1940, under the auspices of the United States Government. The
Secretary of State has appointed an organizing committee composed of
‘government officials and distinguished scientists. Dr. ALEXANDER WETMORE,
Assistant Secretary of the Smithsonian Institution, is the Secretary General
of the Congress and the Secretary of the Organizing Committee. The
Congress will be divided into 11 sections covering the various disciplines.
The chairmen of these sections are as follows:

136 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 3

1. Anthropological Sciences—HERBERT J. SPINDEN, Brooklyn Museum.

2. Biological Sciences—Epwin G. Conxuin, Princeton University.

3. Geological Sciences.—T. WAYLAND VAUGHAN, Geological Society of America.

4, Agriculture and Conservation.—Huau M. Bennett, U. S. Soil Conservation
Service.

5. Public Health and Medicine.—Tuomas Parran, U.S. Public Health Service.

6. Physical and Chemical Sciences—LyMANn J. Briaes, National Bureau of Stand-
ards.

7. Statistics Stuart A. Ricu, Central Statistical Board.

8. History and Geography. CLARENCE H. Harina, Harvard University.

9. International Law, Public Law, and Jurisprudence.—JAMES BRown Scott, Car-
negie Endowment for International Peace.

10. Economics and Sociology—H aroup G. Mouton, Brookings Institution.

11. EHducatton.—Nicuotas Murray Butier, Columbia University.

Obituary

Harvey (WILuiIAMS) CUSHING, master-surgeon, neurologist, physician,
critical investigator, medical bibliophile, inspiring teacher, littérateur, artist,
died at New Haven, Conn., on October 7, 1939. Born in Cleveland, Ohio,
on April 8, 1869, the ninth child in his family, he became the fourth in a line
of doctors.

From school in Cleveland he went to Yale College (A.B., 1891) and then
to the Harvard Medical School (A.M. and M.D., cum laude, 1895). After a
surgical internship at the Massachusetts General Hospital, he went to Balti-
more in 1896 as junior assistant in the surgical service of Halsted at Johns
Hopkins Hospital. During 1900-1901 he studied abroad under Kocher and
Kronecker in Berne, Mosso in Turin, and Sherrington in Liverpool.

After his return to Baltimore Dr. Cushing became neurosurgeon and then
associate professor of surgery at Johns Hopkins. In 1912 he went to Boston
as Mosley professor of surgery at Harvard and surgeon-in-chief to the
Peter Bent Brigham Hospital. From 1915 to 1919 he saw service in France,
during which time he became senior consultant in neurosurgery of the
American Expeditionary Forces. Upon reaching the hospital retirement age
of 63 in 1932, Dr. Cushing gave up his work in Boston and accepted appoint-
ment as the Stirling professor of neurology at Yale, which chair he held from
1933 to 1937.

Dr. Cushing’s writings, as compiled at the time of his seventieth birthday,
number 318 items, largely in the fields of neurosurgery, neurophysiology,
endocrinology, and medical history. He was perhaps more widely known as
the author of the Pulitzer prize winner The Life of Sir William Osler (1925).

Dr. Cushing was a member of many scientific societies and served as presi-
dent of the American Society of Clinical Surgery (1921), the American
College of Surgeons (1922), the American Neurological Association (1923),
and the American Surgical Association (1927). Honors came to him in pro-
fusion, culminating in 1938 with the degree doctor of science, “honoris
causa, ” from Oxford.

With Dr. Cushing surgery of the brain became for the first time scientifi-
cally established. By nature he was a perfectionist, and his accomplishments
seemed to flow from his extraordinary capacity for sustained work. Of him
truly it may be said that he was in all that he attempted greatest of the
great.

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OBITUARY: Harvey WinraMs Cusuine. ,

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JOURNAL

OF THE
WASHINGTON ACADEMY OF SCIENCES
VoL. 30 Aprit 15, 1940 No. 4

PALEONTOLOGY .—Mammals and land bridges.1. GroRGE Gay-
LORD Simpson, American Museum of Natural History, New
York. (Communicated by C. Lewis Gazin.)

It was well known to the ancients that different regions of the earth
were characterized by different sorts of animal life. The Roman em-
perors seeking all manners of beasts for their diversions knew that
they must send to various countries each inhabited by characteristic
animals. Later, when European travelers began to penetrate the far
reaches of the earth, among the first questions asked them was what
peculiar creatures inhabited the deserts of Tartary or the jungles of
Ethiopia. Cartographers delighted in putting pictures of native ani-
mals on their maps, and their efforts to amaze and to embellish pro-
duced the first zoogeographic charts. Generations secure in the belief
in the creation of things as they are seldom sought any explanation of
the differences in fauna between one region and another, and few
men obscurely guessed that this might be the outcome of a shifting
history rather than the static result of divine command.

The rise of science in its modern form found here a whole series of
fascinating problems ready to hand. From a descriptive point of view
the main outlines of the present distribution of mammals were long
since correctly sketched, and now almost all the details are also
known. Confident that the processes of nature are orderly and can be
summarized by general theories and explained by general principles,
the students of the nineteenth century began the attempt to deduce
from the present faunal distribution the historical sequence that led
to it. In this new field of inference many blunders were made (and
we are surely still making some) because of the lack of historical
documents. On this basis alone, the history really can not be de-
ciphered, any more than one could reconstruct the political history of
Europe from the present boundaries of its nations if all actual records
of the past were destroyed. Here the paleontologist came to the res-
cue. His discoveries are the historical documents of animal dis-

1 Address delivered before the Washington Academy of Sciences, February 15,
1940. Received February 9, 1940.

137

138 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 4

tribution. They have solved many problems in this field and at the
same time they have revealed many others as yet unsolved.

Among the plainest inferences from the study of recent mammals
is the fact that some of them have been able to cross regions that are
now impassable to them. Aquatic animals have somehow traversed
areas now dry land, and land animals have gone from one area to
another now isolated by a barrier of water. The paleontologist was
called on to reveal how such movements were possible and when and
under what conditions they occurred.

Now geologists became vitally interested. Caring nothing about the
distribution of animals as such, they care a great deal about the past
distribution of land and sea, the evolution of climates, the rise and
fall of connections between the continents, and other problems that
are involved in or that depend on paleontological studies of distribu-
tion. Research in this field constantly assumes new aspects and
touches new fields of knowledge until from being a curiously special-
ized and abstruse detail it has become vital for work in several differ-
ent sciences and has acquired importance and meaning for anyone
who takes any intellectual interest in the world in which he lives.

When I undertook to discuss this subject, it was my first intention
to take up the various theoretical land bridges from one continent to
another and to summarize the evidence for and against each one in
order to produce a historical account of where and when such bridges
have existed. It soon became apparent that such an account, if it
were to have any value, would involve a mass of detail that would,
indeed, be of interest only to specialists in this field. It also became
evident that relatively few such specialists have risen above this mass
of detail to make a conscious survey of the general principles in-
volved and of the basic assumptions underlying their studies. Such a
general survey is, then, not only of wider interest but also fresher
and more needed in the present stage of study.

To review all the broad problems and principles in one paper is a
manifest impossibility, and attention will be directed to two aspects
on which it now seems possible and useful to make some suggestions.
The first is the broadest problem of all in this field, the general way
in which land mammals tend to become distributed and in which
their distribution tends to change in time. The second is more par-
ticular: the different types of migration routes between major land
areas, the way in which one type or another can be inferred from the
faunal evidence, and the effect that a given type has on the faunas
that use it. In order to lend reality to these abstractions and to point
out some further promising leads for research, one specific example of

Apr. 15, 1940 SIMPSON: MAMMALS AND LAND BRIDGES 139

MOVEMENT IN
ONE DIRECTION

MOVEMENT OUTWARD
FROM ACENTER

| |
CONTRACTION (HOLOGENESis)

MOVEMENT OUTWARD
FROM ACENTER
FOLLOWED BY
RESTRICTION (WITHOUT
NECESSARY MOVEMENT).

3
EXPANSION & CONTRACTION

Fig. 1—Diagram showing various theoretical explanations of the spread of a
group of mammals from one place to another. The given facts are that the group oc-
curs at both A and B and is known at B later than at A. The numbered circles represent
the limits of distribution of the group at successive times, from 1 to 5.

140 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

the rise of a migration route between continents is then taken and
what happened to the continental faunas as a consequence is briefly
considered.

TEMPORAL PATTERNS OF MAMMALIAN DISTRIBUTION

We commonly speak of changes in mammalian distribution as
being caused by migration and extinction. “Migration” suggests a
trek from one area into another or periodic movement back and forth
between two regions, both rare and unimportant phenomena in deal-
ing with the broader outlines of mammalian distribution. It would be
more accurate to substitute ‘““expansion and contraction”’ for ‘‘migra-
tion and extinction.’’? However the words be used, it is clear that
mammals do not as a rule acquire new territory simply by traveling
into it but by a less purposeful peripheral expansion in all possible
directions. Similarly, they do not usually lose territory simply by
traveling away from it, but by a complex sequence of attenuation and
local extinction that can be called contraction. (Fig. 1.)

Regarding the usual relationship of spatial distribution to time,
there are two extreme theories, that of ‘‘age and area,’’ expounded by
Willis (1922), and that of “‘hologenesis,’”’ advanced by Rosa (1931)
and supported in its zoogeographic implications by Fraipont and
Leclerq (1932). Willis is a botanist and bases his theory mainly on
plants but believes it probably also applicable to mammals. His basic
postulate is that new forms of life originate in definite, limited
regions from which as centers they expand slowly and steadily as
time goes on. Then, as a rule with exceptions, at any given point in
time, the area occupied by a form of life should be directly propor-
tional to the age of that form of life. The theory involves various
interesting corollaries, such as the belief that endemics or isolated
forms of life with narrow distribution are usually young forms that
originated where they are found and are just starting on their careers
of expansion.

Rosa’s theory of hologenesis, on the contrary, has the basic postu-
late that a new form of life appears simultaneously over a great area,
over the entire range occupied by an ancestral form or predecessor.
There is, then, no such thing as a center of distribution or a cradle of
any form of life. The distribution, as a rule with unimportant ex-
ceptions, is at the beginning as wide as it will ever be. Migration
(in any sense, or expansion), if it occurs at all, is so insignificant that

_ ? Although, since usage makes meaning, I am not prepared to grant that “‘migra-
tion’ can not mean what nine zoogeographers out of ten use it to mean.

Apr. 15, 1940 SIMPSON: MAMMALS AND LAND BRIDGES 141

the broad features of distribution are about what they would be if
migration never occurred. The area covered by a form of life tends
always to decrease, not to increase. Hence as a rule area is inversely,
not directly, proportional to age. The corollary regarding isolated
forms of narrow distribution is that they are necessarily the relicts
of old groups once more widely distributed.

To a more or less orthodox zoologist Rosa’s theory seems at first
sight so fantastic as hardly to warrant serious discussion. This is still
more true of some of the nongeographic aspects of the theory of
hologenesis not pertinent here. It seems so obvious that most of the
essential geographic implications of the theory are incorrect that I
shall not devote time to disproving them, but it is necessary to recog-
nize considerable merit in the work of Rosa, especially as supple-
mented by Fraipont and Leclerq, less on the theoretical side than in
the description and emphasis of real sequences of geographic events.
From this limited point of view both the age and area theory and that
of hologenesis give true but incomplete pictures. One theory reaches
an unsatisfactory conclusion, as far as mammals are concerned, and
the other departs from an unsatisfactory postulate, but the combi-
nation of the less disputable parts of the two gives a satisfactory
result.

One of the many moderate opinions intermediate between the
extreme views of Willis and of Rosa is that of Matthew (1915, 1939).
Matthew’s main thesis, now well known, is that groups tend to spread
from centers, that the marginal forms are generally conservative and
the central forms progressive, and that most of the main, primary
centers of such spreading have, for mammals at least, been in the
Northern Hemisphere, most southern mammals being relatively
primitive types pushed away from the north by peripheral expansion
about these centers. This thesis is not under discussion in the present
paper, but the general type of geographic history assumed by Mat-
thew to be typical for mammals is that here more explicitly supported.
Writing in 1915, before the recent denials of the existence of centers
of dispersal, Matthew took these as universally admitted. His work is
- full of examples of contracting phases in mammalian geographic his-
tory, and it was mainly on a consideration of these that he built
his theory.

As a concrete example of expansion and contraction, the distribu-
tion of the mastodonts is enlightening and was chosen by Fraipont
and Leclerg as one item of evidence for hologenesis. Their map shows
a Tertiary distribution essentially world-wide except for Australia

142 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

and a Quaternary distribution including all North America, a spot in
Ecuador, one in India, and one in Java. It is not fatal to their theory
that their facts are not straight. Mastodonts entered South America
only at the end of the Tertiary and were typically Quaternary all over
that continent. They died out in the Old World near the beginning
of the Quaternary and are typically Tertiary, only, in those continents.

The fatal flaw in the hologenetic presentation of mastodont history
is not factual. but in the method of generalization. Lumping the Ter-
tiary as if it were a single point in time, they make it appear that
mastodonts arose 7n situ everywhere, which is their thesis but which
is certainly contrary to fact. In the Oligocene mastodonts are known
only from northern Africa. Many great Oligocene faunas from other
continents are known, and it is inconceivable that mastodonts or any
possible ancestors of mastodonts would be (as they are) entirely un-
known in them if these then already had anything comparable to
their maximum distribution. Similarly it is as nearly certain as such
conclusions can ever be that mastodonts were present in Eurasia
(known in the Lower Miocene) earlier and thence spread to North
America (not known until Upper Miocene) and that they were in
North America long before they reached South America (not known
until the end of the Pliocene) and spread from North America to
South America. These facts are consistent with the age and area idea
of expansion from a center and are radically inconsistent with the
hologenetic idea of simultaneous appearance throughout the whole
range.

On the other hand, as mastodonts declined it is evident that their
area greatly diminished until only one or a few relicts were left in
relatively limited regions. This part of the history, if taken alone, is
consistent with hologenesis. It is not, in itself, inconsistent with age
and area, which admits the reality of such cases as exceptions, but it
becomes inconsistent if shown to be usual rather than exceptional,
and this can, I think, be shown.

The accompanying map (Fig. 2) epitomizes what is known of
mastodont distribution in space and time. I hold no brief for the ac-
curacy of this map in detail: there are great gaps in knowledge, and
later discoveries will necessitate changes in the distribution bound-
aries of the map, which are time contours or isochrones of mastodont
expansion and contraction. These isochrones are, however, consistent
with what is now known (which the map of Fraipont and Leclerq is
not), and I venture to predict that later changes of detail will not
much affect the general character of their pattern.

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Apr. 15, 1940 SIMPSON: MAMMALS AND LAND BRIDGES > 143

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This I believe to be the type of pattern that would be shown by
almost any form of life’ that had run its entire course from origin to
extinction. A form appears in some center or “‘cradle,’’ not an exact
spot that could be marked with a monument but, say, a single biotic
district or province. Thence it tends to spread steadily in all directions
until it encounters insuperable barriers. After a time it begins to
contract, possibly but not usually toward its center of origin and
often splitting into disjunctive spots as it contracts. Finally it
disappears. (Fig. 3.)

The expansion of a group of animals involves actual motion. Indi-
vidual animals must move from place to place, and some of them
must travel where their immediate ancestors had never been. The
population as a whole must move outward along its periphery. Con-
traction does not, or need not, involve any motion. It does not
necessarily mean and in reality very seldom means a contraction of
the population in the sense that there is predominant inward motion
along the periphery. It involves rather a process of disappearance or
extinction, commonly preceded by a general lowering or attenuation
of the population. A population may decrease greatly and actually
be well along in its contraction phase before it loses any significant
amount of its range by local complete extinction. This phase of con-
traction can not be simply represented by contours as in the ac-
companying diagrams, and this essential difference between expan-
sion and contraction must be understood if the diagrammatic
representation is not to be misleading.

An excellent descriptive analogy is provided by the expansion and
contraction of ice caps. In their expanding phase there is actual
movement outward from a center. They may begin to contract even
while the movement is still outward, but their definitive contraction
is accompanied by stagnation, with thinning of the ice (attenuation
of the animal population) before any considerable regression is ob-
vious. Commonly parts of the ice mass will be isolated and remain
an situ until they melt entirely, just as relicts of once widespread
animal groups may be isolated in one or in several separate regions
before they become extinct.

It is tempting to go into many of the details and corollaries of this
history, but I must limit myself to mention of only one or two. As

_ ° The expression ‘‘form of life’’ is intentionally vague fer the purpose of generaliza-
tion. Of course, it is not supposed that a single race or species goes through the whole
course of such a cycle unchanged. General racial evolution, modification in local en-
vironments, and many other factors greatly complicate the issue. It would be impossible
in limited space to attempt consideration of such modifications and it would merely
confuse the broader trends that are believed to be real despite these complications.

Apr. 15, 1940 SIMPSON: MAMMALS AND LAND BRIDGES 145

regards age and area, it is evident that this pattern is partly in agree-
ment with that theory, but the theory is unbalanced in tending to
stress the expansive phase as usual or normal and to consider the
diminishing phase, which seems really to be an inevitable, integral
part of the whole process, as unusual or abnormal. Whether a ma-
jority of animals at any given time were really distributed in accord-
ance with age and area would depend on whether more were then in

eos Nagas BARRIER

IMPERMEABLE BARRIER

Fig. 3— Diagram of a common type of mammalian expansion and contraction,
exemplified in varying detail by the mastodonts and other groups. The numbered lines
1 to 11, represent limits of distribution at various times. 1 to 5 represent the primary
expansion of the group on the land-mass where it originated, 5 to 8 contracting phases
here. At time 5 it crosses a barrier and from 5 to 8 expands on a second land-mass, con-
tracting there from 8 to 11. From 6 to 8 the group has discontinuous (disjunctive) dis-
tribution in two areas. After 8, it is extinct in its home-land but survives abroad. After
11, it is everywhere extinct.

the expanding or in the contracting phase and on the relative speed
of these phases. It seems probable that at the present time, including
the recent past, more mammals are actually in the contracting phase,
so that age and the area is a poor guide to the recent distribution of
this particular group of animals.

One other striking detail is that we can as yet seldom follow the
actual expansion of a group of mammals within its set of barriers.
Sometimes related mammals do really seem to appear all at once over
the whole of a great area inhabitable by them, and subsequent ex-

146 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

pansion, if it occurs, is not such in the simple age and area sense but
is by flooding through a broken barrier: this is in part true of the
spread of mastodonts into North America in the Miocene.‘ It is these
cases of sudden widespread appearance that are used to support holo-
genesis, but the support is spurious.’ One reason for the apparent
widespread simultaneous appearance is the imperfection of the record
and of our interpretation of it. For any one age we are lucky to get one
good fossil deposit on a continent and almost never have deposits
so placed all over a land mass that the expansion could be recorded.
And even if we did have such ideal data, our usual methods of cor-
relation would very seldom permit so precise a following of the real
sequence. We usually establish a theoretical sequence by assuming
(doubtless contrary to fact but as a workable approximation) that
given types of mammals did appear simultaneously over the whole
area—obviously it is then ridiculous to expect this sequence to show
that they did not. Only when different faunas cross barriers and im-
pinge on one another is it easy to show that expansion has occurred.
The other reason is that the expansive phase of mammals is normally
very rapid unless definite obstacles slow it down. Once a group of
mammals gains access to a land mass, it tends to spread over it in the
wink of an eye, geologically speaking. A century or a millennium may
suffice, and in most cases such periods are imperceptibly short to the
paleontologist. Only in dealing with recent mammals is one likely
really to see expansion taking place on a smaller scale.

Mammalian distribution as the paleontologist sees it is thus sel-
dom concerned with the spread of any group on a single land mass.
Relatively local differences are usually to be assigned to environ-
mental or facial causes, while differences between larger areas are
usually to be interpreted not primarily from the age and area view-
point of simple time elapsed but more from the point of view of the
rise, fall, and character of intervening barriers. The paleontologist’s

* But I do not doubt that they would have passed the barrier earlier if they had
reached it earlier, so that this is only a modification, not a contradiction, of the age and
area type of expansion.

> To mention only one of several cogent reasons, because adequate data always show
that new forms appear first only on one side and never on both sides of a barrier.

6 Willis foresaw that the great mobility of mammals might vitiate the application
to them of his age and area theory, which as a matter of practical observation demands
that spread should be very slow, as it commonly is among plants and some animals but
rarely among mammals. His thought that the theory might, after all, apply to mammals
was based largely on the fact that some of them gave a “hollow curve” for number of
genera plotted against number of species, but such a curve seems to me inevitable
either in the expanding “‘age and area” phase or in the contracting phase. Perhaps in an
eu eile relatively stable maximum phase it would not occur, but even this is
doubtful.

Apr. 15, 1940 SIMPSON: MAMMALS AND LAND BRIDGES 147

cardinal principles (open to exception) are (a) that strong differences
between approximately contemporaneous mammalian faunas of
similar facies imply an intervening barrier and (6) that strong
resemblances between such faunas denote an intervening connection.

Thus the paleontologist would seldom conclude that a given sort
of mammal occurred on one continent but not on another simply be-
cause it had not had time to reach the second, but because there was
no likely way for it to get there. If, then, this sort of animal did later
appear on the second continent, he would normally conclude that
something had happened to provide the means of getting there, and
not that the animal only then got around to using the means that
existed all along. These interpretive principles are widely accepted, so
much so that real exceptions to them have greatly confused zooge-
ographers.’ Generally true, they are the basis on which the paleon-
tologist and zoogeographer collaborate with the geologist in estab-
lishing the probable presence or absence of land connections between
the continents in past times.

- TYPES AND EFFECTS OF MIGRATION ROUTES
Corridors

If no barrier at all exists between two areas, it is to be expected
that their faunas will be very similar, or as far as genera or larger
groups are concerned practically identical. Such radical differences as
exist will be mainly or wholly caused by the survival or development
of local forms in some narrow environment, that is, will be facial and
not geographic in a broader sense.

As an example, a comparison of the living mammals of Florida and
New Mexico (Simpson, 1936) shows the degree of similarity attained
by areas in which there is no significant geographic barrier but where
the local climates and facies are almost completely different in the
two areas. For various reasons not pertinent here, the mammalian
fauna of Florida is relatively small, with only a quarter as many
species as in New Mexico, but of the orders of mammals present in
Florida, all occur in New Mexico, of the families over nine-tenths, of
the genera two-thirds, and of the species nearly one-fifth. If these
were fossil faunas resemblance this great (or, as is often the case,
greater) would warrant the conclusion that no barrier did exist be-
tween the two. This criterion can be applied in close parallel. It was
formerly sometimes supposed that when Florida first definitively

7 Cases of spread over “‘sweepstakes routes,’’ discussed on a later page, are the
most confusing of these real exceptions.

148 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 4

appeared as dry land in the mid-Tertiary it was not yet connected
with North America. Now we have from there Middle Miocene
mammalian faunas with ten genera surely and six others doubtfully
identified. Of these, all but one are common in contemporaneous beds
in western North America. The conclusion that there was no sea or
other notable barrier between Florida and these States is inescapable.
Such evidence suggests not merely that a bridge existed but that none
was needed; that the two areas were part of a single land mass.

Filter-Bridges

When two regions are separated by a strong barrier, they develop
quite different faunas, the differences being roughly proportional to
the lapse of time since the regions were connected. If now some means
of passing the barrier appears, the two faunas intermingle, but usu-
ally the result is not the production of a single fauna even in the sense
that Florida and New Mexico have one fauna. Several factors are
concerned in the usual fact that such regions tend indefinitely after
they are united still to have distinctive faunas, despite their sharing
of some faunal elements. From this point of view the fact that the
regions often are different environmentally exerts a profound effect,
but one not of primary importance in the phenomena here con-
sidered because the effect might have been analogous even if the
regions had always been united. A more important factor is that
biological pressure of immigrant forms may inhibit the expansion of
some groups in one region without being sufficient to cause rapid ex-
tinction, although in such eases extinction usually follows sooner or
later. Equilibrium does occur but is seldom or never permanent.

Another and for the present subject a more important reason for
the continued distinction of two faunas between which a barrier-
crossing has been established is the character (including the position)
of that crossing. Its approaches may be inaccessible for some animals,
and of course they can not use a bridge that they can not reach. From
the animals that do expand into a new land mass, it is sometimes
possible to infer where the bridge was. Thus when North America and
Asia had a great faunal interchange in the Pleistocene, no mammals
then confined to southern North America reached Asia and none then
confined to southern Asia reached North America. Obviously the
bridge was in the north and exclusively southern animals could not
reach it. It is also noteworthy that none of the mammals that had
come into North America from South America reached Asia. To
reach North America they had to come through the Tropics, and none

Apr. 15, 1940 SIMPSON: MAMMALS AND LAND BRIDGES 149

was sufficiently adaptive also to pass over a relatively cold bridge.

Here the character of the mammals themselves is a determining
factor. What is a barrier for one is not for another, and conversely
what is an open route for one is not for another. The Asia—North
America bridge opened the barrier for elephants (mammoths) but not
for gazelles. The North America—South America bridge opened the
barrier for horses but not for bison. This strongly selective action
depending on the position and character of the bridge and the con-
sequent environmental conditions of it and of its approaches is a rule
with few exceptions. Another way of putting this would be to say
that the true barrier in such cases was not the presence of a stretch
of sea but some less obvious environmental factor, such as climate or
vegetation, and that for these animals the apparent bridging of a
barrier had no meaning because the true barrier remained untouched.
(Fig. 4.)

In the inference of intercontinental land connections from faunal
relationships it is, therefore, wrong to demand that anything like a
complete faunal interchange be adduced as evidence of the existence
of the connection. A wide-open, nonselective connection, a corridor,
is the only sort that could approach such a result, and these are rare.*®
In the whole history of mammals there are exceedingly few cases
(e.g., Lower Eocene between Europe and North America) where the
evidence really warrants the inference of a wide-open corridor be-
tween two now distinct continental masses. The usual sort of con-
nection is selective, not acting as a corridor or open door but as a sort
of filter, permitting some things to pass but holding back others.
From the probable mechanism of such filtering of faunas, it follows
that these connections were usually of narrow environmental scope
and their continental abutments limited, drawing only on one faunal
zone of the continent, not on its fauna as a whole. In other words, the
usual evidence for such connections does not suggest “‘lost conti-
-nents’’ comprising parts of two or more as they exist today, or even
broad transoceanic pathways, but relatively restricted links. The
analogy of a bridge for such selective or filtering connections is fairly
good, and it is to them that the term “‘land bridge’ most properly
applies.

From the point of view of paleogeography, the sort of bridge that

8 Hurope and Asia are now connected by,a corridor, but zoogeographically they are
not distinct continents. One of the many arguments against the Wegener hypothesis,
at least in any application to mammals, is that the connections that it provides are

corridors, but the faunal relationships on which it depends for evidence would not be
produced by corridors.

JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 4

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best fits the zoological evidence in such cases of extensive but filtered
faunal interchange is an isthmian link in the sense of Bailey Willis
(1932). The broad land bridges of many paleogeographers should be
corridors from a faunal point of view, but isthmian links, more nearly
than any other geologically postulated connections, fill the require-
ments of a filter-bridge, which the faunal evidence shows to be the
usual type of intercontinental connection although, of course, by no
means the only type.

When it is recognized that a filter-bridge does not lead to an in-
tegral transfer of continental faunas, it is a practical problem to
determine what sort and degree of resemblance does indicate such a
bridge. There have been students who did not hesitate to build ex-
tensive individual bridges in all directions to account for peculiarities
of distribution in single forms of life. Thus, to mention only a few of
his many connections,’ Joleaud (1924 and elsewhere) has an indi-
vidual Late Oligocene route from Haiti to west-central Africa for
insectivores, one diagonally across this from Brazil to northwestern
Africa in the Late Eocene for certain rodents, one in the Early Mio-
cene straight across the Atlantic from the United States to Spain for a
genus of horses, Anchitherium, one at the same time parallel to but
south of this from northern Africa to Florida for the mastodonts, and
so on. Similarly, von Ihering built a special bridge across the Pacific
from South America to Asia for raccoons and bears,!° and examples
could be multiplied. Aside from geological considerations, which in
themselves are almost enough to exclude these particular bridges at
these places and times, and aside from what are now known to be
errors in the factual data adduced for them, such individual, self-
service bridges are supposed to have acted in a way in which no
surely established bridge is known to have acted, and I can not
believe in their reality.

One good criterion of the reality of a bridge is that it should have
acted in both directions. Provided that both areas had land faunas,
there seems to be no proved case in which a bridge has conducted
animals only from one to the other and not in both directions. This is
true even when one fauna was decidedly dominant and tended as a

® Postulated not necessarily as bridges but possibly as connections of similar effect
but a different sort by an ‘‘accordion”’ motion of the continents on Wegenerian lines.

10 Such a rapid summary is hardly fair either to Joleaud or to von Ihering, who
adduced considerable evidence for their views (although some of the evidence has since
been shown to be erroneous), but it is necessary to mention one or two instances as
briefly as possible in order to demonstrate that I have not set up a straw man. Citation
of the vagaries of less distinguished men would not warrant mentioning the point in so
general a review.

152 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 4

general rule to suppress the other or to inhibit its expansion. For
instance, the South American ground sloths were doomed to extine-
tion when they came in contact with the North American fauna, but
first they penetrated far into North America. The armadillos, also
archaic animals such as might be expected to contract in distribution,
have gained an even more enduring foothold in North America and
are now (for at least the second time) expanding there. One of the
best arguments against the disputed derivation of South American
marsupials by land bridge from Australia (direct or via Antarctica) is
that the evidence favors migration only from Australia to South
America, with none in the reverse direction even though the South
American mammals must have been at least as capable of expansion
as the Australian.!! This and other evidence regarding this particular
hypothetical migration route have been discussed elsewhere (Simp-
son, 1940). (Fig. 5.)

The second and perhaps the best criterion of the reality of a land
bridge is that even though it rarely transports whole faunas, it does
tend to transport integrated faunules. It does not transport all the
genera of a continent, but neither does it transport one genus all by
itself. For instance, it is improbable that only herbivores or only car-
nivores would cross such a bridge (although they need not both cross
in the same direction). Where herbivores go, carnivores can and will
accompany them, and carnivores can not go where there are no
herbivores. The postulation of land bridges on the basis of one or a
few mammals is thus very uncertain. Unless there is reasonable possi-
bility that their companions have not been discovered, a theoretical
bridge based on such evidence is probably unreal.

Sweepstakes Routes

There are, however, instances of migrations of single groups of
mammals or of unbalanced faunas that did occur but that do not
meet these criteria for filter-bridge connections and, of course, still
less those for corridors. Many insular faunas are of this type, as a
whole. Madagascar and the West Indies are classic examples. As
carnivores, Madagascar has only peculiar viverrids, relatives of the
civets, although nearby Africa is abundantly provided with cats

11 Tt is conceivable that a bridge might function in one direction by a sort of lock or
storm-door action, an otherwise uninhabited region receiving a fauna first from one
source, losing that connection, and only then being united with a second continent, so
that animals would be transported from the first to the second but not in the other
direction. There is, however, no good evidence that such a peculiar sequence of events

ever actually happened and it should hardly be postulated except in the absence of any
acceptable alternative hypothesis.

153

SIMPSON: MAMMALS AND LAND BRIDGES

Apr. 15, 1940

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154 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 4

large and small and various other carnivores. Madagascar’s insecti-
vores and rodents are also peculiar and each group is related to only
one of many African types. Madagascar has many primitive primates
and lemurs, but no apes or monkeys. These are all ancient forms
and constitute a very unbalanced fauna that must have entered
(whether together or separately) by the middle Tertiary at latest.
The only ungulates are a pigmy hippopotamus (now extinct) and a
bush-pig, both of which must have reached Madagascar much later
than its other mammals and which are, again, an example of migra-
tion that can not possibly be explained by an ordinary filter-bridge.
In the West Indies the Pleistocene land mammals included only
peculiar rodents, insectivores, and ground sloths, without any of the
ungulates, carnivores, and other groups abundant on all adjacent
continental areas. This fauna, too, is inexplicable as a result of normal
filtering on a land bridge such as is here envisioned. I am aware that.
some excellent authorities do maintain that these faunas arrived over
bridges (see general summary in Schuchert, 1935), but I can not feel
that they have clearly seen or considered the conditions that could
give such a result. (Fig. 6.)

There are also instances of the appearance of isolated immigrants
on continental masses. A curious and relatively neglected example,
among many that might be cited, is that of the sudden appearance
in South America of small relatives of the North American raccoon.
These procyonids appear as fossils in the Late Miocene or Early
Pliocene of Argentina definitely before any of the other carnivores or
any of the abundant North American ungulates reached there. Since
in this case a filter-bridge certainly existed at a later time, it is usual
to assume that the procyonids came on this bridge and that their
appearance dates the formation of the bridge as a practicable migra-
tion route or true and complete filter-bridge. If, however, we consider
only the time when the procyonids did appear, disregarding our
knowledge of what was destined to happen later, such a conclusion is
not warranted. If my previous remarks as to filter-bridges are true,
or are acceptable as a theory of general tendencies, then it is wrong
to conclude that a bridge can account for the appearance of this one
group of small carnivores and no other animals of similar geographic
origin at that time, unless the bridge was then so nearly impassable
as not to warrant the name in its usual accepted sense.

The late W. D. Matthew, who was probably the most distinguished
and best informed student of problems like this, concluded that insu-
lar and highly unbalanced faunas were probably to be accounted for

155

SIMPSON: MAMMALS AND LAND BRIDGES

Apr. 15, 1940

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156 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

by sporadic transportation of land animals on natural rafts, without
the existence of a dry-land route (Matthew, 1918, 1939). This opinion
has been severely criticized in some quarters. It has been claimed or
felt, even by some adherents of Matthew’s general thesis of ‘Climate
and Evolution,’’ that this sort of adventitious migration is dragged
in when necessary to explain away any facts that contradict the main
thesis. |

It has not been sufficiently emphasized even by Matthew that the
role of such a theory may be positive and primary, not merely nega-
tive and supplementary. Adventitious migration has indeed been
used and sometimes abused simply to get inconvenient facts out of
the way of a favored hypothesis, but there are instances in which
adventitious migration is itself the most probable hypothesis and the
most economical theory. In the cases of the faunas of Madagascar
and the West Indies, for instance, I strongly favor this explanation,
and I do so not at all in order to explain away data for a land bridge
where I do not want to believe in one—as Matthew has, quite in-
correctly, been accused of doing. It is to be favored because it does
explain, simply and completely, facts that the land-bridge theory
does not explain.

This sort of migration can be extended to include cases other than
those of transportation by natural rafts, although doubtless these
provide the most common instances. Any barrier, whether of water,
climate, biota, or other, may or will be involved in such migration if
its crossing at any one time is highly improbable but is not impos-
sible. The action is not merely like that of a relatively less permeable
filter but is different in kind as well as in intensity. A filter-bridge
permits some animals to pass and holds others back, but in general
those that can cross it do cross it and do so fairly soon after the bridge
becomes available to them. It is relatively deterministic as to the fact
of crossing, as to the animals that do or do not cross, and as to the
time of crossing. An adventitious route, which I call ‘‘a sweepstakes
route’ to emphasize this characteristic, is indeterministic. Its use
depends purely on chance and is therefore unpredictable and, except
in a broad way, can not be clearly correlated with other events in
time and space, as filter-migration can.

If a sweepstakes route exists, it depends on chance whether a given
type of animal that can cross it will really do so, which of two types
of animals will cross first, and when any particular types will cross
it. It is, for instance, my belief that such a sweepstakes route for land
mammals now exists between Asia and Australia, that it has existed

Apr. 15, 1940 SIMPSON: MAMMALS AND LAND BRIDGES 157

since toward the end of the Mesozoic, and that no more tangible
route, such as a filter-bridge, has existed there during that time. Cer-
tain Asiatic mammals can not follow such a route, and in this sense
it, too, has a filtering action. It is not really a route for such mammals:
they do not hold tickets in the sweepstakes. Other mammals, par-
ticularly small arboreal types, can. All these have tickets in the
sweepstakes, some types holding more tickets than others and so
having more chances, and any of them might win at any time, but any
one is unlikely to do so at any one time and the less likely can win be-
fore the more likely do. A given sort of mammal might have crossed
at once, might have crossed at any time from Cretaceous to Recent,
or might never have crossed. Whether it crossed and when it crossed
were matters of chance, in a sense almost exactly analogous to the
chance of throwing a given point with dice. (Fig. 6.)

This is, I think, the only theory yet advanced that really is capable
of explaining all the peculiarities of the Australian fauna and many
similar but less extreme peculiarities of land faunas in other parts of
the world. That such theories have not received much attention and
that they are uncongenial to many zoogeographers are perhaps a
reflection of the mechanistic scientific philosophy dominant in the
Victorian age, from which zoogeography has not fully emerged. Land-
bridge migration seems more mechanistic because it is often more
simply predictable. In fact, of course, it too depends on chance, but
here on the chances of a probable event, whereas sweepstakes migra-
tion depends on the chances of an improbable event.” The viewpoint
involved is, I believe, new, and it merits detailed consideration, but
this can not be given it here. Among other points, the physical nature
of such sweepstakes routes needs study. It is not to be supposed that
they are invariably island stepping-stones or that natural rafts are
the sole means of transport involved.

A FILTER-BRIDGE IN ACTION

As an example of what actually happens when two continents are
united by a filter-bridge, the case of North and South America is one
of the most interesting and the facts about it are now fairly well
known. These continents were separated (except, probably, for a
sweepstakes route) almost throughout the Tertiary. Toward the end
of the Pliocene they were united by an isthmian link antecedent and

12 Students of statistics will recognize a relationship with the binomial of probability
approaching forms like the normal distribution when chances are about equal and ap-

proaching forms like the very different Poisson distribution when chances are very
unequal.

158 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 4

similar to that now existing, the Isthmus of Panama. For the mam-
malian faunas this was and is a filter-bridge.

Just before the two continents were united, South America had
about 29 families of land mammals and North America about 27.
With two doubtful exceptions,!* they did not then have any families
in common. Shortly after the union of the continents, in the Pleisto-
cene, they had 22 families in common, 7 of South American origin,
14 North American, and | doubtful. Some extinction already having
taken place, South America then had 17 native families still confined
to it and North America 9.1° With further extinction and some
further migration, the Recent faunas of these continents have 14
families of land mammals in common and there are 15 families con-
fined to South America (not all native) and 9 confined to North
America. There was thus a great faunal interchange but one that
never produced even approximately identical faunas, involving many
mammals from each continent but never all or even the majority—
a typical picture of the action of a filter-bridge on the continental
faunas at each end of it.

In passing, there are various interesting facts involved in these
summary figures. The South American fauna is now about as rich as
it was before the interchange, but very different. North America has
a decidedly poorer fauna than before the interchange, but its general
composition has not changed so much as in South America. Both
faunas reached their maximum in variety soon after the interchange
and later declined.

The broad outlines of what actually happened can be seen by
summary of the histories of the various major groups of mammals
involved.

Certain groups expanded into the other continent and became per-
manently at home there, without losing much of their former range,
the ‘‘age and area” type of expansion. The groups of which this is true
were almost entirely of North American origin and include some ro-
dents, especially the cricetids, most of the carnivores, and, among
ungulates, the deer. Among South American mammals only the por-
cupine can unquestionably be placed in this category although there
are one or two other less clear or more complex cases, e.g. the peba
armadillo.

13 The exact figures depend on the classification used and are not important except
as they express relative values.

14 Didelphidae and Procyonidae, possible exceptions for different reasons not af-
fecting the basic situation here described.

15 Some of these, on each side, did manage to spread slightly beyond the isthmus,
but not to colonize the other continent widely.

Apr. 15, 1940 SIMPSON: MAMMALS AND LAND BRIDGES 159

Some groups seem to have been almost unaffected. It must be
supposed of these that they had not reached a contracting phase in
their history, that a barrier continued to exist for them (the filter-
bridge filtered them out of the flow of mammals), and that the con-
tact of new types of mammals was not lethal. This is true of a few
more South American than North American mammals, but the differ-
ence is not significant. In North America the moles, pocket gophers,
beavers, kangaroo-rats, prongbucks, bison, and a few others belong
here. In South America the (tree) sloths, anteaters, most of the
armadillos, the monkeys, and most of the native rodents (eight fami-
lies out of twelve, and lesser groups in the other families) may be
mentioned. Some of these managed to get onto the bridge (for in-
stance several sorts of monkeys), but none really succeeded in cross-
net.

I do not know of any single unified theory that would account well
for the fact that these animals did not cross the bridge and yet did
not markedly contract. The age and area theory demands that they
(or most of them) be new groups that have not yet had time for this
expansion, but this is clearly false. Most of these are ancient types of
animals in their own continent. All certainly have had ample time
to cross the bridge if they were going to do so. They are not incon-
sistent with Matthew’s ‘‘Climate and Evolution” theory, but neither
does it explain them; these data are outside that field of theory. The
reasons are probably too varied to be reduced to a formula more spe-
cific than that of general filter action. For many of these animals, such
as the monkeys, the absence of necessary environmental conditions
beyond the bridge is an evident reason for their stopping where they
did. Others, like the bison, were evidently kept by analogous environ-
mental barriers from reaching the bridge. In some cases, for instance
many of the rodents, it is hard to believe that the physical, climatic,
or floral environment can have sufficed to prevent their spread and
the most reasonable inference seems to be that these animals were
able to maintain their places in the shifting fauna around them, in the
region where they were well acclimated, but not quite able to invade
the same ecologic niches where these were already occupied under
somewhat different conditions, even though these conditions would
not have been deterrent if there were no competition. The explanation
is vague and not very satisfactory because it seems unlikely that so
delicate an equilibrium could long be maintained.

Some groups began to contract at or soon after the time of faunal
interchange. Doubtless some would have contracted anyway, but it

160 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

can not be coincidence that so many did so just at this time. In North
America there were few examples of this. Some North American
groups have contracted since the connection with South America, but
in these cases there is little doubt that the contraction had quite
different causes and would have occurred regardless of the rise of the
land bridge in question. In South America, however, the sharp con-
traction and eventual extinction of all the native carnivores (the
borhyaenid marsupials) and all the native ungulates (notoungulates
and litopterns) undoubtedly were related to this event and so, prob-
ably, was the contraction, with or without extinction as yet, of
various native rodents and of the caenolestid marsupials.

It is highly improbable that none of these animals could have
crossed the bridge successfully as far as most environmental factors
go. Some of the notoungulates and borhyaenids, for instanze, were
ecologically similar to animals that did cross the bridge and they lived
in environments abundantly available in North America. The only
probable explanation is that these animals were biologically inferior to
immigrants from North America. The impact of the latter not only
prevented the expansion of these South American groups but also
started or hastened their contraction. The contraction was slow in
some cases, occupying a million years or more, but it effectively pre-
vented acquisition of new territory and in most cases has now ended
in extinction. In this instance, and probably this is the rule for mam-
mals, expansion of groups that did expand was plainly more rapid
than the contraction of those that did contract.

A final category is provided by the various sorts of mammals that
expanded when the continents were united but that later contracted
again. This was true. of about as many North American as South
American mammals. It is a phenomenon still more complex than those
already mentioned and the land mammals so affected mie be placed
in three categories:

Hs Hees that expanded into the other continent and then became extinct
In both:
(a) Of South American origin: Glyptodonts, ground sloths (several
families).

(b) Of North American origin: Gomphotheres (bunodont masto-
donts),!° horses.
2. Those that expanded into the other continent and then contracted
(or in one case became extinct) there but were not much restricted in their
original home:

146 These were not ultimately of North American origin, but those involved in this
interchange were. Throughout this discussion North American origin means simply not
South American, only these two continents being considered.

Apr. 15, 1940 SIMPSON: MAMMALS AND LAND BRIDGES 161

(a) Of South American origin: .Capybaras, armadillos.
(b) Of North American origin: None.
3. Those that expanded into and survived in the other continent and
became restricted or extinct in their continent of origin:
(a) Of South American origin: None.
(b) Of North American origin: Tapirs, camels, peccaries, short-faced
bears."

The first of these three categories can be dismissed (although hardly
explained) as including groups that would have become extinct in any
case but that happened to share in this last expansive movement
before fatal restrictions set in. The last two are complementary and
show an interesting relationship. No North American groups be-
came extinct in South America and not in North America. If they
became extinct in South America they were, so to speak, slated
for extinction anyway and the new environment did not save them.
On the other hand several North American groups became ex-
tinct at home but not in South America.!® These were, then, contract-
ing groups, for which extinction was postponed by the change of
environment. South America was an asylum for them in their re-
treating phase and the preceding expansion was rather an incident
than an indication of potency against their old environment. Here
again both the age and area and the hologenetic theories are far
beside the point when confronted by the actual facts. On the other
hand, this particular class of facts is broadly consistent with Mat-
thew’s views, especially when details here omitted are considered.

South American groups that were contracting, or were destined
soon to contract, in that continent either were unable to reach or, in
rarer cases, did reach but could not survive in the northern con-
tinent. Even some animals that remained potent and at least did not
markedly contract in South America were unable to maintain them-
selves in North America after reaching there. Generally speaking, the
faunal interchange was far from equal. In the long run the two faunas
did not mingle as much as one invaded the other. The North Ameri-
can mammals were on the whole definitely more potent and more
expansive than the South American, both in their ability to migrate
and in their ability to survive, a generalization supported by the
following tabulations, in which the figures are numbers of families of
land mammals known to have existed in the two continents at about
the time when the bridge arose. (Doubtful cases are omitted.)

17 These have also contracted considerably in South America.

18 [ include peccaries in this group because they contracted greatly in North
America (also, but to far less extent, in South America) and in all probability would
have become extinct in the north if no asylum had been offered them, and may indeed
still become extinct first in the north.

162 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

TABLE 1.—ASSOCIATION OF MIGRATION AND SURVIVAL WITH GEOGRAPHIC ORIGIN

Of South Of North Ratio of ratios,
American American favoring North
origin origin America
Micrated to other continent... ..425.... U 14
Did not migrate to other continent...... 2. 11
Ration. Gis suc ee eas ara eae .3o Ik A 3.8
INOW SUTPVIVAN GE nv tate oe eee 7 21
INOW eX Lime taht. comet ay roads ME ene eran ae ii 4
FiatlOm ee eee de cc taal epee ie, Cae ae ee ame eae 1 5H) 5.25 3.4

These differences are statistically significant, the first surely, the
second probably.

A priori it would be expected that the ability to accomplish such a
migration, an indication of expansive power at the time, would be
related to ability to survive. It is possible that there is a relationship
here, but if so it is more complex and involves other factors. Simple
tabulation of the same families shows no such tendency:

TABLE 2.—ASSOCIATION OF MIGRATION WITH SURVIVAL

Migrated to other Did not migrate to
continent other continent
INOiweSUnvivamMemie soe aie Te 15 23
INO wee xtineteasy tee eee ee ee 6 9
art ea ie Seen ire ee aN ey ean a (ens 2.50 2.56

The difference is far from significant. As far as these figures show, a
family capable of spreading to the other continent was no more likely
to survive than one that did not spread.!® Thus in the final outcome
of the interchange, as far as yet reached, the ability of these faunas
to expand and their ability to survive are both associated with geo-
graphic origin, or biologically with the general character of the his-
torically northern, Holarctic, as opposed to the historically southern,
Neotropical, fauna. But ability to expand and ability to survive are
two different faunal characteristics in this instance with no apparent
relationship to each other.

Like so many phases of this great subject on which I have barely
been able to touch in passing, this unexpected conclusion has far-
reaching implications and merits much more detailed consideration
than can now be given it. An enormous amount of work has been done
to unearth the facts of faunal distribution in the past and present.

19 Use of smaller taxonomic units, such as genera, gives larger figures but obscures
the conclusion sought. Commonly the act of spreading from one continent to the other

was accompanied by evolution of generic rank. The use of actual phyla would be ideal
but is impractical because these are not sufficiently well known in many cases.

Apr. 15, 1940 AXELROD: ESMERALDA FLORA 163

Far less progress has yet been made in finding the broad interpretive
principles that may be revealed by these facts. Here an effort has been
made to indicate what a few of these principles may prove to be and,
more particularly, to suggest a few of the lines of attack that may
lead to clearer grasp of these and to the discovery of others.

LITERATURE CITED

Fraipont, C., and 8. Lecrera. L’évolution. Adaptations et mutations. Berceaux et
migrations. Actualités Scientifiques et Industrielles 47: 1-38. 19382.

IneRInG, H. von. Die Umwandlungen des amerikanische Kontinentes wahrend der
Tertidrzeit. Neues Jahrb. Min. Geol. Pal., Beil-Bd. 32: 134-176. 1911.

JoLeEAuD, L. L’histoire brogéographique de l Amérique et la théorie de Wegener. Journ.
Soe. Amér. Paris, n.s., 16: 325-360. 1924.

Mattruew, W. D. Climate and evolution. Ann. New York Acad. Sci. 24: 171-318.
1915. (Reprinted, with additions, in Matthew, 1939.)

—— Affinities and origin of the Antillean mammals. Bull. Geol. Soc. Amer. 29: 657—
666. 1918. (Reprinted in Matthew, 1939.)

— Climate and evolution, ed. 2, revised and enlarged. New York Acad. Sci., Special
Publ. 1: i-xiii, 1-223. 1939. (Also several other papers bearing on this subject.)

Rosa, D. L’ologénése. Nouvelle théorte de lévolution et de la distribution géographique,
pp. i-xii, 1-368. 19381. Félix Alcan, Paris. (Revised and translated from the
Italian edition, 1918.)

ScuucHERT, C. Historical geology of the Antillean-Caribbean region or the lands border-
ing the Gulf of Mexico and the Caribbean Sea. John Wiley & Sons, New York,
1935.

Simpson, G. G. Data on the relationships of local and continental mammalian faunas.
Journ. Pal. 10: 410-414. 1936.

— Antarctica as a faunal migration route. Proc. Pacific Congress of 1939, in press.
1940.

Wiuuis, B. Isthmian links. Bull. Geol. Soc. Amer. 43: 917-952. 1932.

Wiis, J. C. Age and area. A study of geographical distribution and origin of species,
pp. i-x, 1-259. University Press, Cambridge (England), 1922.

PALEOBOTANY.—The Pliocene Esmeralda flora of west-central
Nevadat. DanigL JI. AXELROD. (Communicated by Ro-
LAND W. Brown.)

One of the results of recent collections of later Tertiary floras over
_ the Great Basin province has been the discovery that the Esmeralda
flora described by Knowlton (1900) from the northern end of the
Silver Peak Range, Esmeralda County, Nev., is distinct from the
Coal Valley flora reported by Berry (1927), which lies in the drainage
of the East Walker River 75 miles northwest. A well-preserved flora
of approximately 50 plants has been collected at Coal Valley and will
form the basis of a subsequent paper. The present brief report adds
six species to the Esmeralda flora and includes an analysis of pre-
viously collected material now at the United States National Mu-
seum. Acknowledgement is made to Dr. Roland W. Brown for
assistance in examining the collections, to the Carnegie Institution
of Washington under whose auspices the collections were made, and

1 Received November 20 1939.

164 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

to the National Research Council for the grant of a fellowship to
study the later Tertiary floras of the Great Basin province.

The Esmeralda formation is in large part a fresh-water deposit,
made up of shales and coal seams in the lowest part, with sandstones,
conglomerates, and volcanics occurring higher in the section. Al-
though a vertebrate fauna has been collected from these beds (Stir-
ton, 1936), determinable plant remains are nowhere abundant, and
with the exception of an occasional specimen the flora is poorly
preserved. Under ordinary circumstances any revision of the Esmer-
alda would at best be largely tentative. However, by comparing it
with the Miocene Tehachapi flora 200 miles southwest and the Lower
Pliocene Truckee floras 110 and 150 miles north, it has been possible
to arrive at a more accurate evaluation of the Esmeralda species for
nearly all of them occur in those beds. The following 16 plants are
considered to represent the Esmeralda flora:

SYSTEMATIC LIST OF SPECIES
PTERIDOPHYTA:
POLYPODIACEAE:
Dryopteris obscura (Knowlton) Berry.

SPERMATOPHYTA:

GYMNOSPERMAE:
Cupressaceae:
Juniperus nevadensis, n. sp.
ANGIOSPERMAB:
Monocotyledonae:
Cyperaceae:
Cyperacites sp.
Dicotyledonae:
Salicaceae.
Populus alexanderi Dorf.
Populus nevadensis (Knowlton) Axelrod, n. comb.
Salix kernensis Axelrod.
Salix vaccinifolia Knowlton.
Fagaceae:
Quercus pollardiana (Knowlton) Axelrod, n. comb.
Quercus turnert Knowlton.
Ulmaceae
Celtis lacustris (Knowlton) Axelrod, n. comb.
Berberidaceae
Mahonia marginata (Lesquereux) Arnold.
Lauraceae:
Umbellularia salicifolia (Lesquereux) Axelrod.
Nymphaeaceae:
Nymphaeites nevadensis (Knowlton) Brown.
Rosaceae:
Cercocarpus cuneatus Dorf.
Prunus nevadensis (Knowlton) Axelrod, n. comb.
Ericaceae
Arctostaphylos preglauca Axelrod.

Apr. 15, 1940 AXELROD: ESMERALDA FLORA 165

The presumed relationships of the 13 woody plants to modern species are
indicated below. The rush (Cyperacites), water-lily (Nymphaeites), and fern
(Dryopteris) have not been included because they are represented by ma-
terial too lacking in diagnostic characters to warrant reference to any living
species.

Fossil Species Nearest Related
Modern Equivalents
Arctostaphylos preglauca A. glauca

Celtis lacustris C. reticulata

Cercocarpus cuneatus C’. betuloides

Juniperus nevadensis J. utahensis (californica)
Mahonia marginata M. fremontir

Populus alexanderi P. trichocarpa

Populus nevadensis P. tremuloides

Prunus nevadensis P. anderson

Quercus pollardiana Q. chrysolepis

Quercus turnerz Q. turneri

Salix kernensis S. bonplandiana var. towmeyi
Salix vaccinifolia S. exigua

Umbellularia salicifolia U. californica

Oaks dominated the fossil flora and together with the juniper comprised
a savanna-woodland community whose regular associates include the moun-
tain mahogany (Cercocarpus), barberry (Mahonia), manzanita (Arcto-
staphylos), and desert peach (Prunus). The remainder are either hygrophi-
lous, such as the water-lily (Nymphaeites) and rush (Cyperacites), or are
forms largely inhabiting stream banks in arid regions.

Although seven of these plants have survived down to the present in the
White Mountains 30 miles west of the fossil locality, and nine of them
commonly occur along the desert slopes of southeastern California, there are
two regions in particular that are significant, as shown in table 1.

TABLE 1.—DISTRIBUTION OF THE NEAREST RELATED MopDERN EQUIVALENT
SPECIES OF THE ESMERALDA FLORA

White Desert Slopes,

. Kern River Southern
1 c 5
Species Mountains Poutbesstern Canyon Arizona
Arctostaphylos glauca... x
Celtis reticulata........ a x Xe Xx
Cercocar pus betuloides. . x Ke x Xx
Juniperus utahensis.... x 5 x x
Mahoniafremoniu..... x x
Populus tremuloides.... x xX a2
Populus trichocarpa.... x x x
Prunus anderson... .. x x x
Quercus arizonica...... X
Quercus chrysolepis.... x x x
Salix bonplandiana.... x
ISGLUL CLIGUA LE coe sen: x X x x
Umbellularia californica x

ee 8) eee OE SS sss aa

MOtalen sy eels kw ih 9 9 9

166 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

The flora of the middle Kern River canyon area is characterized by oaks
and juniper scattered over the canyon slopes, with the willow, hackberry,
California laurel, and desert peach limited to the stream banks. Two of these
plants, the California laurel and manzanita, clearly form a climatically sen-
sitive unit in the flora, for they do not range with the remaining species onto
the desert slopes where conditions are more extreme. Nevertheless, it is to be
emphasized that the Arctostaphylos and Umbellularia are approaching
desert-border conditions in the Kern River canyon. This is shown by the
occurrence of Prunus anderson in the area, in addition to such typical
desert species as EHncelia frutescens, Ephedra nevadensis, Hymenoclea salsola,
Lycium andersonu, and Tetradymia spinosa (Coville, 1893, p. 8). It seems
clear that conditions now obtaining in areas of the southern and interior dis-
tribution of the manzanita and California laurel approximate the Esmeralda
climate. Although the aspen does not occur in the central Kern River can-
yon area, it grows under a wide range of conditions in western North Amer-
ica. The writer has observed it in eastern California and western Nevada
with many plants having equivalent species in the Esmeralda flora. It may
be recalled that the aspen has been recorded in the Pliocene Mount Eden
flora of southern California (Axelrod, 1937), where it is associated with five
plants that occur also in the Esmeralda. —

Nine of the modern equivalents of the flora may be found in southern Ari-
zona, where they also form part of the woodland community, but the Cali-
fornia laurel, black cottonwood, manzanita, and desert peach are absent.
However, the desert peach is replaced here by Prunus fasciculata, one of its
common associates in southeastern California, and both the cottonwood and
manzanita are represented in southern Arizona by species having somewhat
similar ecological requirements. Since the climate of southern Arizona dif-
fers from that of California in its biseasonal distribution of rainfall, the wil-
low (Salix bonplandiana) and oak (Quercus arizonica) limited to this area
clearly represent another climatically important group in the flora. It is
critical that both have fossil representatives in the Miocene Tehachapi flora
(Axelrod, 1939), and that they lived in California at a time when rainfall had
a distribution of the present southern Arizona type. Additional plants in
the Esmeralda flora occurring also in the Tehachapi include all species ex-
cept the juniper (Juniperus nevadensis), manzanita (Arctostaphylos preglau-
ca), and willow (Salix vaccinifolia), but all these would also form a consistent
element in that flora. The persistence of these plants into the later Tertiary
in the southern Great Basin shows clearly that they form a floristic group
which has been segregated into regions now isolated by desert in compara-
tively recent geologic time (Axelrod, 1939, p. 65).

Since the conditions in the Kern River canyon and in southern Arizona
are considered to most nearly approximate the climate of Esmeralda time,
it is desirable to compare briefly the climate of these areas with that now pre-
vailing at Esmeralda. Present conditions at the fossil locality, as judged from
data for Mina 25 miles north and at the same elevation, are as follows:

Apr. 15, 1940 AXELROD: ESMERALDA FLORA 167

ANTON ATENEO, Sslg ve cob ne Sales Neen eee ee 4 inches
mevetace annual vemperatuker yn 48s... 54°F.
mMverace July MmaximouUMteMmperazbumes. ...--..-...... 95°F.
Average January minimum temperature............. 19°F.
hishest temperature mecorded 5 92. 925 ik ee 109°F.
Kowest temperagunemrecordeds: . 2.4.20. ode 22°F.

Although precipitation at Esmeralda amounts to only 4 inches annually, it
ranges from 12 or 14 to as much as 18 inches in areas where vegetation re-
sembles the fossil flora. From these data it is clear that annual rainfall has
been lowered about 10 inches since Esmeralda time. Temperature data are
more difficult to evaluate, but the following points seem worthy of note: (a)
Average annual temperatures in areas showing relationship to the fossil
flora are about 5°F. higher than those now at Esmeralda; (b) average maxi-
mum summer temperatures are 4° or 5°F. lower, and average January mini-
mum temperatures are from 5° to 10°F. higher, in areas where the fossil
flora has the greatest proportion of its nearest related species; (c) extreme
summer temperatures are nearly alike in all of the areas; (d) extreme winter
temperatures are at least 30°F. higher in areas where the Esmeralda has its
greatest relationship to modern vegetation.

From this brief consideration of the floristic and climatic relationships of
the Esmeralda flora, it may be concluded that the arid basin was surrounded
by an oak-juniper community whose associates included species of barberry,
manzanita, mountain mahogany, and desert peach. Growing along the
streams that flowed into the lake, and about its margins, were species of
hackberry, California laurel, aspen, cottonwood, and willow. Rushes and
ferns formed dense communities along the borders of the lake, which also
supported colonies of water-lily. Although the occurrence of Prunus nev-
adensis in the flora suggests that certain members of the sagebrush for-
mation may have occupied drier sites over the area, the predominance of
grazing vertebrates in the formation indicates grassland was dominant.
Rainfall was in the neighborhood of 12 to 15 inches annually and was dis-
tributed as summer thundershowers and winter rains. This is shown by the
presence of the southern oak (Quercus turnerz), willow (Salix kernensis), and
barberry (Mahonia marginata) in the flora, and by the occurrence also in
relatively contemporaneous floras to the north and south, of fossil species
which may be referred to such characteristic southern plants as walnut
(Juglans rupestris), locust (Robinia neo-mexicana), oak (Quercus hypoleuca),
ash (Fraxinus velutina), and soapberry (Sapindus drummondiz). In addition,
it is to be pointed out that the early Pliocene floras of central California pro-
vide ample evidence of summer precipitation (Axelrod, 1939, p. 56; Axelrod,
1940). It is to be expected that rainfall was distributed biseasonally in west-
ern Nevada at this time, when the present Sierra Nevada barrier at the west
was a relatively low mountain range. Temperature conditions in the Es-
meralda area were more nearly like those now found on the western slopes
of the southern Sierra Nevada and in southern Arizona, which differ most

168 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

from the present Nevada climate in having considerably milder winters.

The lowering of rainfall 10 inches in post-Esmeralda time and the more
extreme winter temperatures at the fossil locality are reflected clearly in the
character of the modern flora, for all the area is Great Basin Sagebrush
(Clements, 1920). Although scattered junipers occur in the Silver Peak
Range several miles south, the nearest forest cover of any considerable ex-
tent is at higher elevations in the White Mountains along the California—Ne-
vada boundary 30 miles west. The major factor accounting for the restriction
of tree growth into the higher mountains has been the decrease in rainfall
due to the interception of rain-bearing winds by the rising Sierra Nevada
barrier at the west. The delimitation of the interior region resulted also in
greater extremes of winter temperature, and these have played an important
role in limiting such plants as the California laurel and manzanita to the
western slopes of the Sierra Nevada. Plants whose modern representatives
now occur in the southwestern United States and northern Mexico were
eliminated gradually as summer thundershowers disappeared over the west-
ern portion of the province. Although this floristic evolution over the area
has already been outlined (Axelrod, 1939a), it will be discussed in detail
after the floras over the region, which are distributed from southern Oregon
southward for 800 miles into the Mohave Desert, have been more thorough-
ly studied. :

There is no need of reviewing here the various ideas with respect to the
age of the Esmeralda formation. It is sufficient to point out that Stirton has
recently summarized the vertebrate evidence (1936), and places the forma-
tion in the Lower Pliocene (p. 183). Paleobotanical data point also to a Low-
er Pliocene age for the flora. The absence of the typical Miocene Asiatic and
broad-leaved deciduous genera in the flora definitely establishes its Pliocene
age (Chaney, 1936, 1936a, 1938). Whereas such Miocene genera as Cebatha,
Nelumbo, Trapa, and Zelkova occur in the Coal Valley flora 75 miles north-
west, none of them are known to appear in the Esmeralda flora. Their ab-
sence here indicates its post-Coal Valley age, a fact consistent with the
occurrence of vertebrates representing the Nannippus tehonensis zone (Up-
permost Miocene or basal Pliocene) at Coal Valley.? Although characteristic
Miocene genera and species are known to have survived into the Middle
Pliocene of west-central California under the influence of moderating coastal
conditions (Axelrod, 1940), are abundant in the early Pliocene along the
windward slopes of the Sierra Nevada 140 miles west (Condit), and have
been recorded from the basal Pliocene of the northern Great Basin (Dorf,
1936), they are known only to have persisted as occasional relicts into the
Pliocene of west-central Nevada. It is to be expected that the later Tertiary
floras of the western Great Basin, situated on the leeward side of the rising
Sierra Nevada barrier, would show a reduction in the regular members of the
northern Miocene elements earlier in the epoch than floras on the windward

2 R. A. Stirton, oral communication, 1939.

Apr. 15, 1940 AXELROD: ESMERALDA FLORA 169

slopes of the range or to northward. The replacement of the northern Mio-
cene elements by plants whose modern descendants now occur in the south-
western United States and northern Mexico, is a regular feature of the Lower
Pliocene floras of this area (Axelrod, 1938, 1939), and clearly indicates the
age of the Esmeralda flora.

The Esmeralda is intermediate in its composition and climatic indications
with respect to the Ricardo flora 175 miles south (Webber, 1933; Axelrod,
1939, pp. 78-84), and the Lower Pliocene Truckee flora at Fallon now under
investigation, which lies 110 miles northwest. The Ricardo is dominated by
oaks’ and these appear to have formed a savanna community. Fan-palms
and locust occurred along the streams in this area, and from higher eleva-
tions cypress and pinyon pine contributed occasional stem fragments to the
Ricardo deposit. The climate at Ricardo during Lower Pliocene time was
characterized by high ranges of temperature and a yearly rainfall of from 10
to 14 inches, distributed as summer thundershowers and winter rains. The
Truckee flora at Fallon is dominated also by an oak-juniper community, and
the hackberry, cottonwood, and willow occur here as well. A limited repre- ©
sentation of the montane forest (Pinus, Abies, Arbutus, Castanopsis, Amelan-
chier) at Fallon shows that the climate in this area, which lies 110 miles
northwest, received a greater amount of rainfall and had lower ranges of
temperature than the Esmeralda region.

From these relationships it is possible to conclude that the lowlands over
the area from Fallon southward into the Mohave Desert were characterized
by a woodland formation during Lower Pliocene (Axelrod, 1939a). North-
ward, where annual rainfall ranged from 14 to 17 inches in the lowlands,
members of the montane forest were limited to the adjacent hills. Precipita-
tion decreased southward, limiting members of the montane forest to higher
elevations, and in the western Mohave area, as well as along the borders of
the Colorado Desert (Axelrod, 1937), plants of desert-border character oc-
curred in the woodland region which had a rainfall of from 10 to 14 inches
yearly. .

Conclusions.—(1) Recent collections of later Tertiary vegetation over the
Great Basin have shown that the Esmeralda flora described by Knowlton is
distinct from the Coal Valley flora reported by Berry, which is located 75
miles northwest.

(2) The Esmeralda flora contains 16 species, including a fern, two aqua-
tics, and 13 trees and shrubs whose nearest related modern descendants now
form an oak-juniper community. The assemblage displays a close relation-
ship to modern vegetation now living in southern Arizona and along the
western slopes of the southern Sierra Nevada.

(3) In its climatic and floristic relationships, the Esmeralda is intermediate
with respect to the Ricardo flora at the south and the Fallon flora to the
north.

3 Continued excavations at the Ricardo Petrified Forest have shown oak to be the
dominant species.

170 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 4

(4) Whereas the Esmeralda is of Lower Pliocene age, the Coal Valley flora
contains members of the northern Miocene forest flora and is of Uppermost
Miocene age.

SYSTEMATIC DESCRIPTIONS*

Family PoLYPoODIACEAE
Dryopteris obscura (Knowlton) Berry

Dryopteris obscura (Knowlton) Berry, Proc. U. 8. Nat. Mus. 72(15):
1927.
Fragments of fern pinnae are exceedingly abundant in the shales associ-
ated with the coal workings at the base of the formation south of Coaldale.

Family CUPRESSACEAE
Juniperus nevadensis, n. sp.

A twig 3 cm long and 1 mm wide, with closely appressed leaves appar-
ently in whorls of three, is clearly that of j juniper.

On the basis of foliage alone it is not possible to indicate relationship to
any one modern species of the genus now in western North America. From
an ecological standpoint, however, this material seems referable to the
modern Juniperus californica Carriere of California or to J. utahensis
Lemmon of the Great Basin area. The montane J. occidentalis does not come
under consideration for it is regularly associated with plants whose fossil
equivalents have only a rare representation in this flora. Accordingly, the
Esmeralda material is recognized as distinct from Juniperus sabinoides
Ashlee of the Latah and Wieser floras because it is believed to represent a
different species. The northern juniper seems comparable to the western
J. occidentalis Hooker rather than to the more arid J. californica and J.
utahensis. Juniperus nevadensis is represented also in the Fallon flora 110
miles north and will be figured when this flora is completed.

Family CYPERACEAE
Cyperacites sp.

Remains of rushlike or reedlike plants, so lacking in diagnostic char-
acters that relationship to any one modern genus is questionable, are com-
mon in the shales throughout the section and are particularly abundant in
the lower part of the formation associated with the coal seams. These
dense tangled masses suggest a habitat somewhat similar to the present
sedge- and rush-covered borders of lakes and marshes throughout the arid
portions of the western United States.

Family SALICACHAE
Populus alexanderi Dorf

Populus alexanderi Dorf, Carnegie Inst. Washington Publ. 412: 75-77, pl. 6,
figs. 10, 11 (not fig. 9, which is Populus prefremontit Dorf); pl. 7, figs.
2,3 (not fig. 1, which is Populus prefremonti Dorf). 1930.

4 All species except the juniper, which is in the paleobotanical collections at the
University of California, are represented in the collections at the United States Na-
tional Museum. Although 4 new combinations are presented, it has seemed desirable
not to place in synonymy 3 species that may be shown eventually to belong here:
Populus nevadensis =P. pliotremuloides Axelrod, Quercus pollardiana=Q. hannibal
Dorf, Celtzs lacustris =C. kansana Chaney and Elias. This synonymy has been delayed
until a more complete and diagnostic Esmeralda collection can be obtained.

Apr. 15, 1940 AXELROD: ESMERALDA FLORA ECA

A well-preserved specimen among the undetermined Esmeralda material
at the United States National Museum represents this species, which is
closely similar to the living Populus trichocarpa Torrey and Gray of the
Western United States.

Populus nevadensis (Knowlton) Axelrod, n. comb.

Cercis nevadensis Knowlton, U.S. Geol. Surv. Ann. Rept. 21(2): 217, pl. 30,
fig. 23. 1900.
The leaf figured as Cercis is clearly that of an aspen, and another basal
fragment among the unidentified material of the Esmeralda flora is also
referred to the Pliocene species.

Salix kernensis Axelrod

Salix kernensis Axelrod, Carnegie Inst. Washington Publ. 415: 90-91, pl. 6,
fig. 7. 1939.

Salix vaccinifolia Knowlton (in part), U. 8. Geol. Surv. Ann. Rept. 21(2):
212, pl. 30, fig. 20 only (not fig. 8, which remains Salix vaccinifolia).
1900.

Salix angusta Al Braun, Knowlton, Idem: 212, pl. 30, fig. 22.

Salix sp. Knowlton, Idem: 213, pl. 30, fig. 13.

All these specimens are clearly of the same type and differ in no manner
from the Tehachapi species. Two of the Esmeralda species were figured
incorrectly. Salix vaccinifolia (fig. 20) should have long, thin and looping
secondaries like those of Salzx sp. in figure 13. The margins of the specimen
figured as Salix angusta are entire, not serrate, and the secondaries are
likewise long, looping, and thin.

This material compares rather closely with the entire-margined forms of
Salix bonplandiana Humboldt, Bonpland, and Kunth, of the southwestern
United States and northern Mexico, and particularly with the variety
toumey? of southern Arizona.

Salix vaccinifolia Knowlton

Salix vaccinifolia Knowlton, U.S. Geol. Surv. Ann. Rept. 21(2): 212, pl. 30,
fig. 8 only (not fig. 20, which is Salix kernensis Axelrod). 1900.
Knowlton originally pointed out the close resemblance between this speci-
men and the smaller leaves produced by the modern Salix exigua Nuttall
of the Western United States.

Family FaGacEAE
Quercus turneri Knowlton

Quercus turnert Knowlton, U. 8. Geol. Surv. Ann. Rept. 21(2): 214, pl. 30,
fig. 21. 1900.—Axelrod, Carnegie Inst. Washington Publ. 516: 100, pl.
7, figs. 10, 11, 14. 1939.
Quercus argentum Knowlton, U. 8. Geol. Surv. Ann. Rept. 21(2): 215, pl. 30,
fig. 12. 1900.
Cinchonidium? turneri Knowlton, Idem: 218, pl. 30, figs. 9-11.
All this material is one botanic species and forms the dominant of this
flora. Among modern plants there is a close relationship to Quercus arizonica
Sargent of the Southwestern United States.

Quercus pollardiana (Knowlton) Axelrod, n. comb.

Chrysobalanus pollardiana Knowlton, U. 8. Geol. Surv. Ann. Rept. 21(2):
216, pl. 30, fig. 19. 1900.

172 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

The Chrysobalanus may be matched by the large oval leaves of Quercus
chrysolepis Liebmann (see sheet nos. 1521956, 667634, 319924, 319884 in
the United States National Herbarium).

Family ULMACEAE
Celtis lacustris (Knowlton) Axelrod, n. comb.

Ficus lacustris Knowlton, U. 8. Geol. Surv. Ann. Rept. 21(2): 215, pl. 30,
fig. 26. 1900.

Although Berry has placed Ficus lacustris in the genus Populus (1927,
p. 9), Chaney and Elias (p. 22) have pointed out its resemblance to hack-
berry. Comparison of the type material with the leaves of the modern Celtis
reticulata reveals no differences between them. The specimen is entire-
margined, not serrate as figured.

Family BERBERIDACEAE
Mahonia marginata (Lesquereux) Arnold

Mahonia marginata (Lesquereux) Arnold, Contr. Mus. Pal. Univ. Michigan
5(4): 64-65. 1936. =

Odostemon marginata (Lesquereux) Knowlton, U.S. Geol. Surv. Prof. Paper
131: 189, pl. 43, figs. 7-10. 1923.

Although there appears to be a gradation between this species and M.
hakaefolia (Lesquereux) Arnold, they may be distinct. The typical leaflets
of M. hakaefolia are similar to those produced by the modern M. trifoliata
Fedde, which regularly has two or three lateral teeth and a slender shape.
The leaflets of M. marginata are more nearly like those of the modern
M. fremonti Torrey in that they are broader, the sinuses are shallower, and
the teeth are not so large and heavy. By this definition, the leaves figured by
Arnold as M. hakaefolia would belong more properly to M. marginata. It is
hoped that further collections from the Creede flora, where both of these
species are represented, will settle this problem.

Family LAURACEAE
Umbellularia salicifolia (Lesquereux) Axelrod

Umbellularia salicifolia (Lesquereux) Axelrod, Carnegie Inst. Washington
Publ. 516: 102-108, pl. 8, fig. 4. 1939. (See synonymy.)

Salix? sp. Knowlton, U. S. Geol. Surv. Ann. Rept. 21(2): 213-214, pl. 30,
fig. 14. 1900.

A relatively large leaf in the writer’s collection, in addition to the small
willowlike leaf figured by Knowlton, seems properly referable to Umbellu-
laria.

Family NYMPHAEACEAE
Nymphaeites nevadensis (Knowlton) Brown
Nymphaeites nevadensis (Knowlton) Brown, Journ. Washington Acad. Sci.
27: 509, pl. 1, fig. 10. 19388. (See synonymy. )

The water-lily is not only represented at the type Esmeralda locality, but
is exceedingly abundant in the Coal Valley flora where it forms a regular
associate of Nelumbo, Equisetum, Trapa, Typha, and other hygrophilous
plants.

Family Rosackar
Cercocarpus cuneatus Dorf

Cercocarpus cuneatus Dorf, Carnegie Inst. Washington Publ. 412: 98-99,
pl. 12, fig. 3. 1930.

Apr. 15, 1940 AXELROD: ESMERALDA FLORA 173

The upper portion of a leaf impression in the undetermined Esmeralda
material at the United States National Museum is clearly that of the moun-
tain mahogany.

Prunus nevadensis (Knowlton) Axelrod, n. comb.

Rhus? nevadensis Knowlton, U. 8. Geol. Surv. Ann. Rept. 21(2): 218, pl. 30,
fig. 15. 1900.

The leaf figured as Rhus nevadensis has been incorrectly illustrated be-
cause the-margin is serrate and the lower secondaries loop well into the upper
part of the blade. An additional unfigured specimen is clearly of the same
type, but is about two-thirds as large. Both of these specimens are indis-
tinguishable from the leaves produced by the modern Prunus andersont
Gray of the Great Basin area.

Family ERICACEAE
Arctostaphylos preglauca Axelrod

Arctostaphylos preglauca Axelrod, Carnegie Inst. Washington Publ. 476(3):
178, pl. 6, figs. 8, 9. 1937.

A single leaf among the undescribed Esmeralda material is clearly that
of manzanita.. Although it has been referred to this Mount Eden species,
which is related closely to the living Arctostaphylos glauca Lindley, it shows
affinity also to such species as A. pringlei Parry of southern Arizona and
A. glandulosa Eastwood of California. Although further collections may
show this Esmeralda manzanita to be a new species, it seems clear that it
has its relationships among species no longer occurring over the Great Basin
area.

LITERATURE CITED

AxELrop, D. I. A Pliocene flora from the Mount Eden beds, southern California.
Carnegie Inst. Washington Publ. 476: 125-183. 1937.

The stratigraphic significance of a southern element in later Tertiary floras of

western America. Journ. Washington Acad. Sci. 28: 313-322. 1938.

A Miocene flora from the western border of the Mohave Desert. Carnegie Inst.

Washington Publ. 516: 1-128. 19389.

Late Tertiary vegetation and climate of the Great Basin and border areas. Proc.

Geol. Soc. Amer. (1939a) 1940.

The Green Valley florule: its position in the Pliocene sequence east of the Berkeley
Hills, California. MS. submitted to Geol. Soc. Amer. 1940.

Berry, E. W. The flora of the Esmeralda formation in western Nevada. Proc. U.S.
Nat. Mus. 72(23). 1927.

CHANEY, R. W. The succession and distribution of Cenozoic floras around the northern
Pacific basin. Essays in Geobotany in Honor of William Albert Setchell, pp. 55-
85. 1936.

Plant distribution as a guide to age determination. Journ. Washington Acad.

Sci. 26: 313-324. 1936a.

Paleoecological interpretations of Cenozoic plants in western North America.
Bot. Rev. 4: 371-396. 1938.

CiemEnts, F. E. Plant indicators. Carnegie Inst. Washington Publ. 290. 1920.

Conpit, C. Late Tertiary floras from the central Sierra Nevada. MS. submitted to
Geol. Soc. Amer. 1940.

CovitueE, F. V. Botany of the Death Valley expedition. Contr. U. 8. Nat. Herb. 4.
1893.

Dorr, E. A late Tertiary flora from southwestern Idaho. Carnegie Inst. Washington
Publ. 476: 73-124. 1936.

-174 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

Know ton, F. H. Fossil plants of the Esmeralda formation. U.S. Geol. Survey Ann.
Rept. 21(2): 209-220. 1900.

StTirTON, R. A. Succession of North American continental Pliocene mammalian faunas.
Amer. Journ. Sci., ser. 5, 32: 161-206. 1936.

WEBBER, I. W. Woods from the Ricardo Pliocene of Last Chance Gulch, California.
Carnegie Inst. Washington Publ. 412: 1138-134. 1983.

PROCEEDINGS OF THE ACADEMY AND
AFFILIATED SOCIETIES

THE ACADEMY
42D ANNUAL MEETING

The 42d Annual Meeting of the Washington Academy of Sciences was
held in the Assembly Hall of the Cosmos Club, January 18, 1940, with 47
members present. President CHARLES I. CHAMBLIss called the meeting to
order at 9:45 p.m. The minutes jof the 41st Annual Meeting were presented
and approved as published in the JouRNAL. The Corresponding Secretary,
NatuaNn R. Smit, submitted the following report on the membership and
activities of the Academy:

During 1939, 52 were elected to resident, 9 to nonresident active member-
ship, and 6 to honorary membership. Of the 52 for resident membership, 34
had become members in good standing at the end of the year, 2 had de-
clined membership, 3 had not replied to our letters, and 13 had accepted
membership but had not paid their dues (most of these were elected in
December). Of the 9 elected to nonresident membership, 5 qualified, 1 de-
clined, 3 have not replied as yet. Of those who accepted membership in 1939,
14 were elected in recognition of their work in Chemistry; 6 in Botany; 5
each in Entomology, Bacteriology, and Medicine; 2 each in Physics, Biology,
Zoology, and Ichthyology; and 1 each in Geology, Soil Science, Helmin-
thology, Pharmacology, Genetics, Statistics, Education, Food Research, and
Ethnology. There were 4 resignations, 3 in the resident and 1 in the nonresi-
dent membership. Five resident and 3 nonresident members were dropped
for nonpayment of dues. Because of their retirement from professional work,
4 resident and 5 nonresident members were considered active members
without the payment of dues.

There were 9 deaths during the year, 4 resident and 5 nonresident
members, as follows:

GrEorRGE H. Girry, resident, on January 27, 1939.
EDMUND C. SHOREY, resident, on January 30, 1939.

W. D. BicELow, resident, on March 3, 1939.

J. G. LipMaANn, nonresident, on April 19, 1939.

JOSEPH GRINNELL, nonresident, on May 29, 1939.
WENDELL C. MANSFIELD, resident, on July 24, 1939.
Harvey W. CusHIna, nonresident, on October 7, 1939.
WALDEMAR LINDGREN, nonresident, on November 3, 1939.
E. E. FRE#, nonresident, on November 24, 1939.

To summarize, on January 1, 1940, the membership consisted of 17
honorary members, 3 patrons, and 572 active members, of which 2 are life
members. Of the active members, 43 (29 resident and 14 nonresident) do not
pay dues because of retirement. There were, therefore, 529 active members
who pay dues, 411 resident and 118 nonresident. Since the number of resi-

Apr. 15, 1940 PROCEEDINGS: THE ACADEMY 175

dent members is limited to 450 and the nonresident to 200, there were 39
vacancies in the resident and 82 in the nonresident.

The status of the resident active membership is satisfactory. The large
number of vacancies is due to the fact that the limitation was raised from
400 to 450 by vote of the membership in July. As was stated above, 13 duly
accepted, but there was not time for the collection of dues before January 1.
In addition, 15 have been recommended for membership by the Committee
on Membership and may be acted upon at the next meeting of the Board of
Managers. There are, therefore, potentially only 11 vacancies in the resident
membership.

The status of the nonresident membership is progressively unsatisfactory
owing to the small number elected.

The Board of Managers had a very busy year, holding 9 meetings with
an average attendance of 18. In addition to the routine business of the
Academy, the Board authorized the appointment of the following special
committees, all of which completed their work and reported to the Board:

On Editorial Assistant: Chairman, J. A. STEVENSON.

To Consider Publishing Red Book: Chairman, W. A. Dayton.

On Ways and Means for Selling Complete Sets of the Journal: Chair-
man, H. E. Ewine.

Memoirs and Awards: Chairman, J. F. Coucu.

To Finish Cataloging the Scientific Societies of Washington and to De-
fine Qualifications of a Society for Affiliation and to State the Ad-
vantages of Affiliation to the Society: Chairman, W. T. SCHALLER.

To Review the Bylaws and Make Recommendations: Chairman, F. C.
KRACEK.

To Consider the Revision of the Bylaws and to Make Definite Recom-
mendations to the Board of Managers: Chairman, W. A. Dayton.
On Awards: Subcommittee for Biological Sciences: Temporary Chair-

man, EK. A. GOLDMAN.
Subcommittee for Physical Sciences: Temporary Chairman, J. F.
CoucH
Subcommittee for Engineering Sciences: Temporary Chairman,
H. L. Curtis.
On Monographs: Temporary Chairman, H. B. Couuins, JR.
On Certificate of Award: Chairman, H. B. HumpHrey.

By authorization of the Board, the President appointed W. W. Dinu.
Custodian and Subscription Manager of Publications for 3 years.

On motion it was voted to accept the Corresponding Secretary’s report
_ and to place it on file.

The Recording Secretary, Oscar 8. ADams, presented the following re-
port:

The 42d year of the Academy began with the 290th meeting and ended
tonight with the 295th meeting. All the six meetings of the year were held
in the Assembly Hall of the Cosmos Club.

The 290th meeting was held on February 16, 1939. An illustrated address
was given by WALDO L. ScumiIrTT, curator, Division of Marine Invertebrates,
U. 8. National Museum, Smithsonian Institution, on the subject The
Galapagos Islands. Attendance about 200.

The 291st meeting was held on March 16, 1939. An illustrated address
was given by Enrico Fermi, Columbia University, New York City, on the
subject The Mesotron. Attendance about 200.

176 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 4

The 292d meeting was held on April 20, 1939. An illustrated address was
given by WitToN Marion KroGmMan, associate professor of anatomy and
physical anthropology, University of Chicago, on the subject The Skeleton
Teils Its Story. Attendance about 170.

The 293d meeting was held on November 16, 1939. The address was given
by Maj. Gen. Hanson E. Ey, former commandant of the Army War
College, on the subject Improvements in War Weapons since the World War
and in Methods of their Use. Attendance 103.

The 294th meeting occurred on December 21, 1939. The address was given
by Cart 8. MaRvEL, professor of organic chemistry, University of Illinois,
on the subject Plastic Glasshike Resins (Vinyl Polymers). Samples of many of
the Polymers were shown and their properties discussed. Attendance about
200.

The 295th meeting was held on January 18, 1940. The address was given
by the retiring President of the Academy, CHARLES E. CHAMBLISS of the
Bureau of Plant Industry, U. 8. Department of Agriculture, on the subject
The Botany and History of Zizania aquatica L. Attendance 1238.

This was the final meeting of the 42d year of the Academy.

On motion it was voted to accept the Recording Secretary’s report and to
place it on file.

The report of the Treasurer, H. G. AveRs, was read by Howarp 8.
RAPPLEYE:

CASH RECEIPTS AND DISBURSEMENTS

RECEIPTS:

Bromo baekedues:.2. 04 Wee: acs piers Sener ee ee $ 140.00
Bron dues fOr 1939 Gs 1s ee ee ee 2,450.00
From: ducsshOr a O40 eh Oa Ne nyo one 40.00
Krom subscriptions tor 1939) 223 ee. 432.30
Prom subscrptons tor lO4 0... oe se eee 236.10
Promesalessor J ounmalisis vi) sine ie ae 227 .26
Bromapaymentsoriteprints. sel) 2h en ee 450.94
Krom sales of 1937 Directory................ ib is
rom sales or 1939) Directory. 2). .). 2.2. 108.00
Krom interest.onidepositsy=.) 2) 95.82
Prom imferest on imvestments..\. 245085. .o0r 940.60

Motal receipusenpr ake as aay ee, oe $5,122.17
Cash Balanceranuany ti LOS We aes eee 1,514.94

To be accounted for yo. ei ee eee $6,637.11

DISBURSEMENTS:

Hor Secretaryes Omicer!93Ser an, eee ia 10)
Hor Secrevaryas:Otnce: 939. ne ee 227 .20
Hor Ereasurer’s Olicen 37. sl eee 188.15
Hor Journal lapramtine sl OS Srna er er 281.78
Hor: Jourmal moraine elo OF vn wate eee PARENT AD)
Por wournalare primus os Snr mre tae acne 70.88
Hor Jourmalreprimtshl 939). sce ee eee 558.18
Korillustrations, Qs Se.) ee ee 39.58
Ror-illustratioms NO30n 05 kee eae 402.61

Hor Journalt@ tices 1933)... nena 36.88

Apr. 15, 1940 PROCEEDINGS: THE ACADEMY > PLE

ier. ournal OmicelO39nt sry seh 202.42
For Meetings Committee, 1938.............. 25.00
For Meetings Committee, 19389.............. 238 .05
RigemVIFCCLOLY. 2 2 ee oe eee 702.59
For dues of retired members returned......... 10.00
For refund of overcharge on reprints......... 3.60
Bank debit memos, as follows:
Bs er es ns ee I od es $0.35
ISO SUbSCKIpPtIONS. ............. 2: .50
HOO SuwbSCrIptIONS 4.62 ie eS .08 .93
Deposited im savings account................. 95.82
Motaledisbursements y.s5 )-i5. 5 Be. $5,618.62
Cash balance December 31, 1939............. 1,018.49
TD Guba Ls 575 Roo een RGR art rec ow Doma ek yt cere Oe arenes $6,637.11
RECONCILIATION OF BANK BALANCE
alamee as per cash book, December 31, 1939............... $1,018.49
Bank balance, American Security & Trust Co. per’
statement of December 31, 1989.............. $1,158.48
PPEGenpesmMOb OeDOSILGU..... 006. we ee ee ce 76.00
$1,234.48
Checks outstanding, not cashed:
HSROMBINO Ross oats ent os ae $e oO
Niomeoualrere chen fo ea 9.12
OME ee es 6.00
UINGOMESHO ne ee or. cn ake. L337
TOO Mhe aee e ocld acee e We e ce 215.99
ReeHCNINC CR eke Na ees Oe a eee a amin Lt SE 2 $1,018.49

Check No. 170 listed as outstanding was issued in May 1934
but has not been cashed.

INVESTMENTS

409 shares stock of Washington Sanitary Improvement Co.,
patavaue- pl Oper share: COSt 28 sees. 6. oo ke $ 4,090.00
20 shares stock Potomac Electric Power Co., 6% Pref.; cost... 2,247.50

-1 bond of Interborough Rapid Transit Co., no 37020, interest
at 5%; due Jan. 1966; par value $1,000; cost............ 995.00

*1 bond of Chicago Railways Co., no. 1027; interest at 5%,
due 1927> par value $1,000 less $250; cost. .-.....2... 05. 713.87

1 real-estate note of T. Q. Donaldson (no. 6 of 12) dated June
PalOoqe tons) years; mberest ati o45 COSU.4. 5 s5ee: 5... 1,000.00

2 real-estate notes of Yetta Korman et al., dated Oct. 5, 1938,
for 3 years (no. 7 of 37 for $500, and no. 8 of 37 for $500);
HMERE SU AUTO 4. COSU.c ak Mend. el ro ee eee, Oo gale, 1,000.00

* The bond of the Chicago Railways Co. was not paid upon maturity due to the
expiration of franchise and failure of the Legislature to enact continuing legislation;
interest has been paid to date under authority of the courts, and $250 has been paid
on the principal since maturity.

178 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

3 real-estate notes of Ell & Kay Bldg. & Investment Co., dated
Oct. 15, 1938, for 3 years (no. 75 of 165 for $2,000, no. 83 of
165 for $1,000, and no. 101 of 165 for $1,000); interest at
DG COSG LST Wee ep. BO ap eee eee cn ee ee Oni eee 4,000.00
Butler notes—Property at 1707 LSt., NW., bought in by note
holders—Academy’s share 4/250ths; interest amounting to

$40:recervediin 1939; cost... : aie soa: ee) eee 2,000.00
$16,046.37
Deposited in savings account, American Security & Trust Co.. 8,409.94
A OAL: sed gg hE Ne ores a ee ae ce ee $24 456.31
Cash-book balance December 31, 1939................... 1,018.49
Total assets. (oct ee . $25,474.80
ALLOTMENTS
Allotted Receipts Expended
Secretary's Office....... $450 OR. bie ela ae ae $ 227.20 (1)
Treasurer’s Office..... 9, 225000) ass Be Oe ee 188.15
Journals eee. oe. 2,500.00-+- $1,444.20 = $3,944.20.. 3,580.46 (2)
Meetings‘Commuttee: 9 350.00 2.4... ee 238 .05 (3)
Membership Committee . 102000 oc. See ee -——
Executive Committee.... 10.000 = ce Sa eee -—
Directony Wie oe 700 .00-+$108.00 = $808.00..... 702.59
Bills outstanding, not paid, at end of year (estimated):
Gl) eSecretany: suOiicers 4 ele weer ea ee ee 6 30.00
(25) Jo iene ae ge ey Mak Stee oe Te ee mee ei ane ir 280.00
(S)e Meetings: Commmibtee we) 252s os ae ee 110.00

The Auditing Committee. H. G. Dorsny, Paut WHITNEY, and ELMER
Hiaains, reported:

Your committee appointed to audit the accounts of the Treasurer of the
Washington Academy of Sciences for the year 1939 submits the following
report:

The Treasurer’s records of receipts and expenditures as shown in his ac-
count books and included in his report have been examined and found cor-
rect. All vouchers have been examined and found to be correct and properly
approved.

The balance sheets submitted by the bank and the securities listed in the
Treasurer’s report have been examined. The statement of the assets of the
Academy was found correct.

No coupons not yet due were missing from any of the securities bearing
coupons.

The records of the Treasurer’s office have been carefully and systemati-
cally kept, thus greatly facilitating the work of the auditing committee.

The auditing committee congratulates the Treasurer on the manner in
which he has carefully conducted his office during his long faithful tenure.

It was moved to accept the reports of the Treasurer and the Auditing
Committee and to place them on file. By vote it was so ordered.

The Board of Editors, F. D. Rossini, C. L. Gazin, and J. H. Kempton,
submitted the following report:

Apr. 15, 1940 PROCEEDINGS: THE ACADEMY 179

I. NATURE AND AMOUNT OF THE MATERIAL PUBLISHED

Volume 29 consists of 560 pages, as compared with 556, 548, 554, 588,
576, 588, 572, and 552 pages, respectively, for the years from 1938 back to
1931.

The space in volume 29 was distributed as follows:

Number Number

Class Subject of papers of pages
Biology Anthropology if 5.60
. Biology if 14.00
Botany 8 49.22
Entomology 7 1.99
Evolution 1 20.00
Ichthyology 2 25.83
Malacology 2 4.15
Mammalogy 2 Shoo
Medical Entomology IL 5.90
Ornithology 3 1:89
Plant Physiology 1 15.00
Zoology 6 All
Geology Geology 24 18.50
Paleobotany 4 18.90
Paleontology 7 60.25
Chemistry Chemistry 2 14.15
Geochemistry 1 5.25
Physical Chemistry 1 25.30
Physics Biophysics 1 6.85
| Geophysics 1 30.00
Physics 2 29.15
Obituary 7 5.00
Scientific News and Notes if 0.70
Index Authors 4.00
Subjects 4.00
Proceedings Academy 10.40
Botanical Society Ho 0D
Chemical Society 13.00
Geological Society 5.65
Philosophical Society 14.25

Divided as above, the various classes of material occupied the following
_ percentages of the total space in the Journal:

Class No. of papers Percentage
Biology 30 51.0
Geology 13 17.6
Chemistry 4 Sell
Physics 4 11.9
Obituary 0.9
Scientific News and Notes 0.2
Proceedings 8.8
Index iS

Of the 56 original scientific papers, 36 were written by members of the
Academy and 20 were communicated.

180 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

II. AMOUNT AND DISTRIBUTION OF INCOME AND EXPENDITURES

Income
From the Academy $2,500.00
From subscriptions 740.10
From sales of back numbers 216.42
Total $3,456.52
Expenditures
Charged to Charged to
Cost authors or the

sponsors Academy

Printer’s office:
Printing, binding, mailing, etc. $2,618.59 $$ 8.72 $2,609.87
Engravings 409 .90 143.01 266.89
Reprints 568 .07 306 . 22 261.85

Editor’s office:

Postage and miscellaneous 43.14 — 43.14
Clerical assistance 180.00 — 180.00
Total $3,819.70 $457.95 $3,361.75
Unexpended balance $ 94.77

The amount actually expended by the Academy in maintaining the
Journal for the year 1939, which is the amount allotted to the Journal by the
Academy less the unexpended balance returned to the Academy by the
Journal, was $2,405.23.

The report of the Board of Editors was accepted and ordered placed on
file.

The following report was submitted by the Custodian and Subscription
Manager of Publications, W. W. DIEHL:

On December 1 a report was made to the Board of Managers recording the
result of an inventory of stocks made on November 13. This inventory
showed the surprising fact that there was then but one complete set of the
Journal available for sale in respect to volumes 1, 2, and 3; and in the case
of the next 12 volumes in no instance over nine sets to the volume. Personal
inquiry at the Waverly Press, Baltimore, where they are stored, revealed
that this very small number of old volumes was due to the elimination in
1937 of most of the old stocks save a very small number of which some have
since been sold. Because of this limited number of the older volumes of the
Journal some changes in policy seemed needful. The Board on December 1,
therefore, ordered the reservation from sale except as parts of complete sets
of all numbers in volumes 1-15. It is hoped that additional numbers of the
limited 15 volumes may be donated to the Academy in order to make more
complete sets available for sale in compliance with this Board action.

The Custodian and Subscription Manager has planned a course of action
in soliciting new subscriptions, but no action has yet been taken because it
is dependent upon the allotment of funds in the 1940 budget.

By vote the report of the Custodian and Subscription Manager of Publica-
tions was accepted and ordered placed on file.

The following report on awards and memoirs was submitted by President
CHAMBLISS:

Apr. 15, 1940 PROCEEDINGS: THE ACADEMY | 181

BYLAWS
Article VII—Recognition of Merit in Scientific Work

Section 1. The Academy may award medals and prizes or otherwise ex-
press its recognition and commendation of scientific work of high merit
and distinction.

Section 2. Awards shall be made only on approval by the Board of
Managers of a recommendation of a committee on awards.

The outgoing Board of Managers made this article operative on October
20, 1939, by the adoption of the Report of Committee on Awards and
Memoirs, appointed March 6, 1939.

By this action the Board of Managers established awards for noteworthy
discovery, accomplishments or publications in the biological, engineering,
and physical sciences, respectively.

The award will take the form of a suitably inscribed certificate. It will be
limited to persons 40 or less years of age, who are members of the Academy,
resident or nonresident, or to members of societies affiliated with the
Academy resident in the Washington area.

The Board of Managers on January 12, 1940, approved the reeommenda-
tions of the Subcommittees on Awards as follows:

Subcommittee on Awards representing the biological sciences recom-
mended that a certificate of award be granted HERBERT FRIEDMANN,
curator of birds, U. 8. National Museum, for noteworthy research and
publication in ornithology.

Subcommittee on Awards representing the engineering sciences recom-
mended that a certificate of award be conferred on Pau A. Smriru, hydro-
graphic and geodetic engineer, U. 8. Coast and Geodetic Survey, for his
contributions to our knowledge of the ocean bottom along the eastern coast
of the United States.

Subcommittee on Awards representing the physical sciences recommended
that a certificate of award be given to WitmoTt H. BRADLEY, senior geologist,
U. 8S. Geological Survey, for his contributions to our knowledge of the oil
shale of the Green River formation of Wyoming and of the Tertiary history
of the adjoining Uinta Mountains (Utah).

These certificates will be appropriately presented at an early date.

The action of the Board of Managers regarding awards was approved by
the Academy.

The tellers, CHARLOTTE ELLioTT, ANNA JENKINS, and DorotHy NICKER-
SON, reported the election of the following officers: President, EK. C. Crit-
TENDON; Nonresident Vice-presidents, P. G. AGNEW and GIFFoRD PINCHOT;
Corresponding Secretary, F. D. Rossin1; Recording Secretary, F. C.
- Kracex; Treasurer, H. 8. Rappueye; Board of Managers, J. F. Coucu and
J. E. GRAF.

All submitted amendments to the bylaws were approved.

In accordance with the report of the Tellers, the officers as submitted were
declared duly elected by the President.

The Corresponding Secretary read the list of nominations for Vice-
presidents submitted by the affiliated societies as follows:

Philosophical, R. E. Grsson
Anthropological, Frank M. SETZLER
Biological, W. B. BELL

Chemical, A. T. McPHERSON
Entomological, A. H. CLark

182 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 4

National Geographic, A. WETMORE
Geological, J. B. Mrrtin, Jr. (to be elected in March)
Medical, FRED O. Coz

Historical, ALLEN C. CLARK

Botanical, CHARLES THOM
Archaeological, ALES HRDLICKA
Foresters, W. A. Dayton

Washington Engineers, P. E. WHITNEY
Electrical Engineers, H. L. Curtis
Mechanical Engineers, WALTER RAMBERG
Helminthological, HE. W. Pricz
Bacteriological, R. R. SPENCER

Military Engineeers, C. L. GARNER
Radio Engineers, H. G. Dorsry

By vote of the Academy, the Recording Secretary was instructed to cast
one ballot for the list as read. This was done and the Vice-presidents were
declared duly elected.

As business from the floor, L. B. TucKERMAN presented a petition to the
Board of Managers for the revocation of the action doing away with the
nonresident Vice-presidents of the Academy. This petition was signed by
six members of the Academy and hence must be acted on by the Board of
Managers at a future meeting.

President CHAMBLISS appointed Past Presidents O. E. Mrrnzmr and W. J.
Humpureys to escort President-elect CRITTENDEN to the chair. After a short
address President CRITTENDEN declared the meeting adjourned at 10:30
p.m.

NEW MEMBERS ELECTED

The following have been elected to resident membership in the Academy:

Ipa A. BENGTSON, senior bacteriologist, National Institute of Health, in
recognition of her contributions to bacteriology, especially work on anaerobic
bacteria and rickettsia.

Miriam Lucite Bomuarp, associate forest ecologist, U. S. Forest Service,
in recognition of her contributions to botanical knowledge, especially in the
morphology and taxonomy of seeds of eastern plants, and to the ecology and
taxonomy of palms, particularly the genera Sabal, Butia, and Ceroxylon.

SELWYN DEWiIrTT CoLuins, principal statistician, National Institute of
Health, in recognition of his contributions to statistical analysis pertaining
to public-health problems.

Harotp RAYMOND CURRAN, associate bacteriologist, U. S. Bureau of
Dairy Industry, in recognition of his contributions to the science of milk
bacteriology.

Luioyp Derr FELTON, senior surgeon, U. S. Public Health Service, in
recognition of his studies on pneumococcus and pneumonia.

WititiaAM N. FrEnton, associate anthropologist, Bureau of American
Ethnology, Smithsonian Institution, in recognition of his contributions on
the ethnology of the Iroquois Indians.

EpWARD HARRISON GRAHAM, biologist, U. S. Soil Conservation Service,
in recognition of his contributions to botanical science, including flora of the
Kartabo region, British Guiana, and botanical studies in the Uinta Basin of
Utah and Colorado.

SAMUEL FREDERICK HILDEBRAND, senior ichthyologist, U. 8. Bureau of
Fisheries, in recognition of his demonstration of the utility of fishes in

Apr. 15, 1940 PROCEEDINGS: THE ACADEMY 183

mosquito control, his work on the life histories of marine fishes of the
Atlantic coast, and his contributions to the ichthyology of Central America.

ROBERT SULLIVAN HOLLINGSHEAD, acting chief, Food Research Division,
U.S. Bureau of Agricultural Chemistry and Engineering, in recognition of
his studies on the use of the data of food analysis in the improvement of the
technology of food handling and in the improvement of public health.

Fioyp Ervin Kurtz, associate chemist, U. 8. Bureau of Dairy Industry,
in recognition of his contributions to the science of milk chemistry.

Morris ABEL RaInzEs, professor of botany, Howard University, in recog-
nition of his researches in plant physiology.

SYLVESTER T’. SCHICKTANZ, senior chemist, U. 8. Bureau of Internal
Revenue, in recognition of his work on distillation and of methods for the
separation of azeotropic mixtures.

LEONARD PETER SCHULTZ, curator, Division of Fishes, U. 8. National
Museum, in recognition of his contributions to ichthyology, especially his
studies of the fishes of the north Pacific.

CLARENCE R. SHOEMAKER, assistant curator, Division of Marine Inverte-
brates, U. S. National Museum, in recognition of his special knowledge of
Crustacea, especially the Amphipoda.

EpGar, REYNOLDS SMITH, senior chemist, National Bureau of Standards,
in recognition of his contributions in the field of physical chemistry, es-
pecially for his work on “‘heavy water’ and ebulliometry.

FRANK E. A. THONE, staff member, Science Service, in recognition of his
contributions to plant ecology and his popularization of science.

RaupuH P. Tirtsuer, associate bacteriologist, U. 8. Bureau of Dairy In-
dustry, in recognition of his contributions to the science of bacteriology,
especially in connection with Escherichia colt and Escherichia-aerobacter.

Byron H. Wess, dairy manufacturing specialist, U. 8. Bureau of Dairy
Industry, in recognition of his contributions to the chemistry of dairy
products.

FREEMAN ALBERT WEISS, senior pathologist, U. S. Bureau of Plant In-
dustry, in recognition of his work in plant pathology, especially on diseases
of ornamentals, soil fungicides, and toxicity relations of fungi.

The following have been elected to nonresident membership in the
Academy:

ERNEST BROWN BaBcock, professor of genetics, University of California,
Berkeley, Calif., in recognition of his contributions to knowledge of genetics
and plant breeding, the nature and mechanism of germinal substance,
taxonomy, and plant evolution.

CHARLES Marius BARBEAU, anthropologist and folk-lorist of Canada,
Canadian National Museum, Ottawa, Ontario, Canada, in recognition of his
many fundamental researches in ethnology, folk-lore, and folk-music.

Oscar 8. Apams, Recording Secretary

Obituary

GEORGE CHARLTON Matson died in Tulsa, Okla., on January 3, 1940.
He was born at Strang, Nebr., on February 4, 1873. He attended Doane
College, where he obtained a B.S. degree in 1900, subsequently studied at
the University of Nebraska and Cornell University, and in 1920 was
awarded the degree of Doctory of Philosophy by the University of Chicago.
He was an assistant in geology at Cornell University, 1901-03, instructor in
geology at the University of Illinois, 1903-04, and fellow in geology at the
University of Chicago, 1904-06. During the 10-year period 1906-16, he was
an assistant geologist and a geologist with the United States Geological Sur-
vey. His scientific publications include reports on the ground-water re-
sources, clay, and phosphate deposits of Florida, the late Tertiary and Qua-
ternary formations of Mississippi, the oil and gas resources of Louisiana and
the Coastal Plain of Texas, and other geologic subjects. In 1916 he left the
Geological Survey and joined the geological staff of the Mexican Gulf Oil
Co. Subsequently he was connected with the Gulf Refining Co. of Louisiana,
the Gypsy Oil Co., and the South American Gulf Oil Co. From 1921 until
his death he was a consulting oil geologist and from 1922 until 1929 he was
also vice-president of the Schermerhorn Oil Co.

Dr. Matson was quiet and reserved in demeanor, an independent thinker,
industrious and thorough in his investigations, and successful in his business
undertakings. His published writings are authoritative, and some of them
were prophetic of important developments in oil and gas production in the
Gulf region. He held membership in numerous scientific and technical
societies, including the Geological Society of America, the American Asso-
ciation of Petroleum Geologists, the Geological Society of Washington, the
Washington Academy of Sciences, and the American Society of Mining and
Metallurgical Engineers. In 1921 he served as president of the American
Association of Petroleum Geologists.

184

CONTENTS

PALEONTOLOGY. —Mammals and land bridges. GEORGE GA’

SIMPSON) (oe ee eee

Dante I. AXELROD! 0 oo 3 ee ee

Procempincs: Toe ACADEMY..........4:..0.).(.2.7).08

OBITUARY: GEORGE CHARLTON MATSON.................-

May 15, 1940

No. 6

JOURNAL

OF THE

WASHINGTON ACADEMY

OF SCIENCES

C. Lewis GAZIn
U. 8. NATIONAL MUSEUM

BOARD OF EDITORS

James H. Kempton

U. 3. BUREAU OF PLANT INDUSTRY

ASSOCIATE EDITORS

RAYMOND J. SEEGER
GEORGE WASHINGTON UNIVERSITY

My
6 Lewis V. JuDSON Austin H. Cuark
PHILOSOPHICAL SOCIETY ENTOMOLOGICAL SOCIBTY
ip, H. H. T. Jackson GEORGE TUNELL
bs BIOLOGICAL SOCIETY GEOLOGICAL SOCIETY
oh Jason R. SwALLEN T. Date Stewart
ae BOTANICAL SOCIETY ANTHROPOLOGICAL SOCIETY
ae Horace S. IsBeiu
me CHEMICAL SOCIETY
By PUBLISHED MONTHLY
cn WASHINGTON ACADEMY OF SCIENCES =
iy 450 Aunaip Sr. a
ae at MenasnHa, WISCONSIN
ee
s : Entered as second class matter under the Act of August 24, 1912, at Menasha, Wis.
<i A@ceptance for mailing at a special rate of postage provided for in the Act of February 28, 1925.
a Authorized January 21, 1933
<a
. r
7
3

ee ee

aa

This JouRNAL, the official organ of the Wee aa f Sei
(1) short original papers, written or communicated by members of t. a,
proceedings and programs of meetings of the Academy and affilia d soci
notes of events connected with the scientific life of Washington. The Jour
monthly, on the fifteenth of each month. Volumes correspond to cal

Manuscripis may be sent to any naer of the Board of Editors. I urgently re
quested that contributors consult the latest numbers of the JourNau and conform th
manuscripts to the usage found there as regards arrangement of title, subh sy
onymies, footnotes, tables, bibliography, legends for illustrations, and other matt
Manuscripts should be typewritten, double-spaced, on good paper. Footnotes
be numbered serially in pencil and submitted on a separate sheet. The editor f
assume responsibility for the ideas expressed by the author, nor can ues und ak
correct other than obvious minor errors. BM

or

Illustrations in excess of the equivalent (in cost) of two ee line drawing ar
to be paid for by the author.

Proof.—In order to facilitate prompt publication one proof will scneeae be. en
to authors in or near Washington. It is urged that manuscript be submitted in final
form; the editors will exercise due care in seeing that copy is followed. Can

Author’s Reprints —Fifty reprints without covers will be furnished oe to aetiees
of scientific papers. Covers bearing the name of the author and title of the article, with —
inclusive pagination and date of issue, and reprints, will be furnished at cost a
ordered, in accordance with the aes schedule of prices. ;

Copies 4 pp. 8 pp. 12 pp. 16 pp. oa

50 ay eT ci tae oO
100 $ .75 $1.20 $1.50 $2.00 | 2.75 |
150 1.25 1.90 2.55 3.25. 3.50
200 1.75 2.60 3.60 4.50 4.25 |
250 2.00 3.30 4.65 6.75 6.00

Envelopes for mailing reprints with the author’s name and nddness printed i in the
corner may be obtained at the following prices: First 100, $4.00; additional 100, $1. 00.

Subscription Rates.—Per volume............. Sitettcereet certs sere (86, 00
To members of affiliated societies; per volume. eee e eee teens 2. 50
Single numbers.......... EG ik EU NP ere ener 8 said

Correspondence relating to new subscriptions or to the purchase of back numbers >
or volumes of the JouRNAL should be addressed to William W. Diehl, Custodian and
Subscription Manager of Publications, U.S. Bureau of Plant Industry, bee
D ae

Remitiances should be made payable to ‘‘Washington Academy of Sciences” ie
addressed to 450 Ahnaip Street, Menasha, Wis., or to the Treasurer, H. S. Reppleye,
U.S. Coast and Geodetic Survey, Washington, bc.

Exchanges.—The JourRNaAL does not exchange with other publications. 1 le

Missing Numbers will be replaced without charge provided that claim is made to
the Treasurer within thirty days after date of following issue.

OFFICERS OF THE ACADEMY

President: EUGENE C. CritTENDEN, National Bureau of Standards. i
Corresponding Secretary: FREDERICK D. Rossin1, National Bureau of Standards.
Recording Secretary: FRanx C. Kracex, Geophysical Laboratory.
Treasurer: Howarp 8. Raprpieye, U. S. Coast and Geodetic Survey.

JOURNAL

OF THE
WASHINGTON ACADEMY OF SCIENCES
Vou. 30 May 15, 1940 No. 5

BOTANY.—The botany and history of Zizania aquatica L. (‘wild
rice’).. CHARLES EK. CHamBtiiss, U. 8S. Bureau of Plant In-
dustry.

For this evening I have assembled some of my field notes on
Zizania aquatica, together with some historical data on the species,
which I trust will meet the requirements of this occasion. And this is
my story.

Life was easy for Wenibozho.? His indulgent grandmother, with
whom he lived, demanded no work of him, and in consequence he
passed through his early boyhood days without exhibiting any par-
ticular interest in those things that must be learned and thoroughly
understood by people who depend largely upon self for the neces-
saries of life. :

At last, the grandmother awoke to the fact that her grandson
lacked the initiative so essential to meet the requirements of their
race, and convinced that her solicitous care was responsible, the aging
woman urged the indifferent youth to prepare himself with a training
that would fit him to endure such hardships as hunger, thirst, and
cold. She told him that experiences of this kind would make him re-
sourceful and teach him how to care for himself and those who might
be dependent upon him. Probably somewhat irked by these plain
words, Wenibozho later said goodbye to his grandmother, who for
many years had provided him with food and shelter.

Equipped with only a bow and some arrows, he started on a long
_ journey through the forests. For meat, he had to depend upon the
flesh of small animals. Not because there was a scarcity of animals,
for they abounded in the woods, but because of his unskilled use of
the bow, his kills were few. Therefore, he had to subsist on seeds,
roots, and tubers. Without knowing the plants that could furnish
nourishing food, he naturally made mistakes.

One day when thoroughly exhausted from want of food he heard a
voice saying, ‘“Sometimes they eat us.’’ He heard this voice several

1 Address of the retiring president of the Washington Academy of Sciences, de-

livered on January 18, 1940. Received February 12, 1940.
2? GILMORE, MELVIN R. Prairie smoke, pp. 195-198. 1929.

185

186 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

times and finally asked, “‘To whom are you talking?’’ A small bush
replied that it had spoken. As no part of the bush above the ground
seemed edible, Wenibozho thought that the roots might be good to
eat. He uncovered the roots, tasted them, and liked the flavor of them
very much. Being hungry, he ate many of them and suffered from
overeating. For several days he was unable to travel, and when he
attempted to do so he found himself as hungry as before and quite
weak. As he passed along seeking food, many plants spoke to him.
Wenibozho gave no heed to their entreaties until he was attracted by
the beauty of a graceful grass growing in a small lake basking in the
sunshine of the open woodland. Some of these plants beckoned to him
and said, ‘‘Sometimes they eat us.’’ He was quite hungry now, and
observing that the upper part of the plants was loaded with long
seeds, he soon gathered some of them. Removing the hulls, he ate the
kernels and found the taste of them so pleasing and their effect upon
his hunger so gratifying that he exclaimed, ‘Oh, you are indeed
good! What are you called?’’ The plants replied, ‘‘We are called
manomin.”’

These adventures and discoveries of Wenibozho have served as a
foundation for many legends that have been handed down through
generations of Chippewa, Menomini, and related Indians. In their
childhood, this story excited their imagination, and as they grew
older they came to have veneration for this fruitful grass that provides
such palatable and nourishing food.

Manomin,* an Algonquian word meaning ‘‘good berry,’’ is sugges-
tive and descriptive. By most of the tribes of this linguistic stock,
this grass and its fruit are called manomin, and by the same name the
plant became known to the early white settlers of the upper Mis-
sissipp1 Valley. Many common names for it came into use when the
French and English population increased in this region. By the French
it was folle avoine, a name most frequently found in the earlier ac-
counts of that part of North America around the Great Lakes. The
English names for the plant are quite numerous. Such names as wild
rice, Indian rice, squaw rice, Canadian rice, black rice, Indian oats,
blackbird oats, wild oats, and water oats are found in the literature.
This plant, however, is not a species of rice or of an oat, though the
vernacular names so designate it. Some of these names are only
locally used. For example, this plant is known only by the name of
water oats to the older inhabitants living along the tidal streams in

the south Atlantic States.

3 Jenks, ALBERT Ernest. The wild rice gatherers of the Upper Lakes. Ann. Rept.
Bur. Amer. Ethnology, 1897-98, 19: 1024. 1900.

May 15, 1940 CHAMBLISS: BOTANY AND HISTORY OF ZIZANIA AQUATICA 187

Among the adventurers who flocked to North America shortly
after Columbus missed his way to India and discovered a new world,
there was a sprinkling of naturalists who came to gather seeds and
bulbs of plants that might be useful in the gardens, fields, and forests
of Europe. Probably the most noted among them was Peter Kalm.
Shortly after his return to Sweden he published in 1751 a small octavo
pamphlet of 48 pages,* containing ‘‘a comprehensive summary of his
observations on the habitat, use, and care of American plants which
he considered of sufficient economic importance to warrant experi-
mental introduction into Sweden.’’ In the text Kalm says: “I have
chosen this means to give an index and short account of some of the
useful plants [a total of 126 species] the seeds of which I brought
home with me from North America where I made a journey at the
command of the Royal Academy of Science.”’

In reference to wild rice, which is included in this collection, he
records: “In North America where the plant grows wild, it is used as
food by all the savage nations, who yearly collect quantities. Wild
ducks are particularly delicious when the rice is ripe, for at that time
they live on it almost entirely. If we could succeed in getting this
rice to grow and ripen here we would have gained a great deal, for
the wettest places would become as productive as fields if the plant
would stand our winters. Cattle are more than greedy for the leaves
and stalks. The greatest difficulty will be to find a method of sowing
seeds so they will germinate. We still know very little about nature’s
method of sowing the seeds of plants growing in water.”’

This attempt to introduce this wild plant into Europe was likely a
failure, for Lambert, in a paper presented before the Linnaean Society
of London in 1804, states that “the seed of [this species] Zzzania
aquatica in a vegetating state from America was long a desideratum
among the botanists of this country; for although seeds were received
here at different times, yet none of them grew. At last, Dr. Nooth
by the desire of Sir Joseph Banks sent them from the lakes of Canada
put up in jars of water. As soon as they arrived they were sown in a
proper situation, where they came up in a few days and the plants
ripened their seeds extremely well in the autumn.”’ This importation
of seed was made in 1791.

Resident collectors, among whom may be mentioned Bartram and
Clayton, also aided in this work and in addition supplied the botan-

4 LARSEN, EstHER Louise. Peter Kalm’s short account of the natural position, use,
and care of some plants, of which the seeds were recently brought home from North America

for the service of those who take pleasure in experimenting with the cultivation of the same
in our climate. Agr. Hist. 13: 34, 48-44. 1939.

188 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

Fig.1.—Typical mature plants of the broadleaf Zizania aquatica
growing in a Potomac River marsh.

ists of the continent with material that greatly enriched the herbaria
of many botanical centers of the Old World. In this way a specimen
of manomin, called wild rice by the white man, got to Europe, where
in 1753 it was described and named Zizania aquatica by the Swedish

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BOTANY AND HISTORY OE ZIZANIA AQUATICA 189

May 15, 1940 CHAMBLISS

190 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

botanist Linnaeus. Until then the ‘“‘good berry” plant was unknown
to botanists, although for centuries it had fed many tribes of wild
men and for nearly 200 years had supplied the food wants of many
European adventurers.

This grass, indigenous to North America, is found from Lake Win-
nipeg to the Gulf of Mexico and eastward from the Rocky
Mountains to the Atlantic coast. Throughout these latitudes, and in
this area where conditions are favorable, it is conspicuous among the
aquatic plants growing in shallow lakes and in slow-moving streams.
In the extreme northern and eastern limits of this region it often
covers several hundred acres. Within its natural range the species
often occupies the small bays of the large lakes, covers the mud flats
on tidal rivers of the Atlantic Coastal Plain, fills the lakelike ex-
pansions of rivers near their source, and grows luxuriantly in the quiet
bends of sluggish streams. It is seldom found in the inland lakes with
no outlets. It grows well on a variety of soils under fresh-water
streams and lakes. Its best growth, however, is made wherever the
plants can anchor themselves in a thick layer of mud, regardless of
the kind of soil. |

Zizania aquatica is an aquatic, annual, self-sowing grass having tall,
erect, cylindrical, and hollow stems, which bear the inflorescence and 4
to 6 long leaves with flat blades, conspicuously marked by a very
thick midrib (Fig. 1). The slender stems have a comparatively thin
wall, and when seen by transmitted light, thin, transverse partitions
are revealed, dividing the internodal space into compartments,
which give the stems a light banded appearance.

The stems vary in height from 5 to 10 feet and in diameter from
one-fourth to five-eighths of an inch. The taller plants are character-
istic of the tidal flats and the shorter plants of the northern lakes and
streams. Plants with stalks as thick as 2 inches near the crown are
not unusual in southern marshes. In thin stands and among isolated
plants a single plant may have many stems, some arising from the
base of the mother stem, though frequently as branches from the first
and second nodes.

The principal roots are slender, fibrous, and numerous and do not
penetrate deeply into the soil.

The first leaves to appear are long and narrow. In the later and per-
manent leaves the basal part known as the sheath is thick and spongy
in structure and completely wraps the stem, thereby adding much
to its rigidity. The sheaths vary in length from 9 to 25 inches.

The blade is the free end of the leaf. In the terminal leaves it may

May 15, 1940 CHAMBLISS: BOTANY AND HISTORY OF ZIZANIA AQUATICA 191

Fig. 3.—Panicles of Zizania aquatica. Left: Of the broadleaf form. Its natural
length is 26 inches. Right: Of the narrowleaf form. Its natural length is 163 inches.

be 2 to 4 feet long and from less than an inch to 1 inch or more wide.
The inflorescence (Fig. 3) is borne on the last node of the stem.
It consists of two parts—an upper, with slender straight branches
bearing the female or seed-producing flowers, and a lower, with
drooping branches bearing the male flowers.
The female flower is very simple in structure, consisting of lemma

192 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

Fig. 4.—Seeds and kernels of Zizania aquatica. (Natural size.) Narrowleaf form:
A, seeds; B, kernels; C, parched kernels. Broadleaf form: D, seeds; H#, kernels.

> ne eee eee eee

May 15, 1940 CHAMBLISS: BOTANY AND HISTORY OF ZIZANIA AQUATICA 193

and palea that enclose a much-branched stigma and a comparatively
small ovary. The lemma bears a long awn or beard. In the male flower,
there are also a lemma and palea, which are much shorter than in the
female flower. They enclose six bright yellow stamens.

The seed is long and slender and almost cylindrical (Fig. 4, A, D).
A thin brown hull made up of the lemma and palea encloses the kernel
and bears a long, stiff, straight awn that is covered with numerous
barbs or bristly hairs. The surface of the hull itself is covered with
similar hairs.

The kernel of the southern plants averages 20 mm in length and
1.4 mm in width (Fig. 4, £). In the northern plants the kernel has an
average length of 12 mm and a width of 2 mm (Fig. 4, B). There is
a shallow groove on the ventral surface of the kernel in which a long
embryo is concealed. When fully matured the kernel is purplish black
in color.

Zizania aquatica has two rather distinct forms, and within each
there are many variants. The form having broad leaves and long
needlelike kernels grows along the tidal rivers emptying into the
Atlantic Ocean and is also found locally in the interior as far north as
southern Minnesota and Wisconsin. The form with narrow leaves and
short thick kernels grows in the upper Mississippi Valley and east-
ward along the Canadian border.

In ascending a tidal stream such as the Potomac River, which is
salty or brackish almost to the head of tidal water, the halophytie
vegetation begins to disappear with the appearance of Scirpus
americanus, a fresh-water rush quite resistant to brackish conditions.
As the conditions become less saline, fresh-water plants begin to oc-
cupy the marshes, and principal among them is Zizania aquatica.
If the Zizanza marsh slopes gradually into a slow-running stream,
several distinct zones of plants will be present. The deeper water will
be inhabited by several species of Potamogeton, Vallisneria spiralis,
and such free-swimming plants as Lemna and other duckweeds.
In water less than 5 feet in depth, the yellow pond lily may occur.

The next zone is usually narrow and is populated with the pickerel
weed Pontederia cordata, which grows in approximately 2 feet of
water. Having a very large root system it is capable of holding its
place in a moderate current and serves as a protection to the adjoin-
ing zone containing Zzania aquatica, the plants of which are not
strongly anchored in the soil. Although such a marsh is inundated,
the depth of water covering it varies with the condition of the tides.

194 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

The Zizania zone is broad and usually parallels the stream for
some distance (Fig. 2). The luxuriant growth of Zizania aquatica
shades out most of its competitors, leaving it in possession of the
land except where the stand is thin on the margin of the stream.
Here straggling plants of Polygonum sagittatum and species of Sagit-
tarva and Bidens are sometimes present. On the land side of this zone
the dominant species is Typha latifolia, which under conditions of
less water and greater silt deposit may displace Zizania aquatica.
The Typha area itself is soon invaded by Peltandra virginica, Iris
versicolor, and several species of Hibiscus and Polygonum. As the
marsh becomes more elevated and drier, shrubs and finally trees be-
come dominant. |

In the lakes of Minnesota and Wisconsin, just beyond the zone con-
taining such submerged plants as the Potamogeton species and Val-
lisneria spiralis, the bottom is often thickly covered with plants
having lime-encrusted leaves and stems called stoneworts. The
Zizania area is usually bordered on the outside margin by several
species of Scirpus, never by Pontederia cordata as along the Potomac
River, and on the land side by Carez, though. Typha is often present.

During early April in the vicinity of Washington, D. C., the seed-
lings of Zizania aquatica have already emerged from the muck-
covered flats and are strong enough to stand erect when the tide is
out, like seedlings in any grainfield. Within a week or 10 days after
emergence the young plants have three leaves. Growth is slow at
first largely because of the low temperature of soil and water and in-
termittent sunshine. During these days they are strengthening their
grip upon Mother Earth, for good anchorage in the soil at this time
reduces the hazard of being washed away by tides, especially the ebb
flow. The protection of the marginal plants that they have in later
life is lacking now. The plants become more robust as spring ad-
vances, and by June, where the stand is good, the growth gives
the marsh the appearance of a low meadow. In another month the
stems that have been concealed by the enveloping leaf sheaths during
this vegetative growth begin to give some evidence of their existence.

The part of the stem embraced by the sheath of the last leaf is
growing rapidly now and being confined within a narrow space dis-
tends the sheath into a spindlelike appearance. Ten days after the
inception of this condition is perceptible, the panicle, which is the
terminal of the stem, begins to emerge. The panicle emerges slowly,
often requiring seven days to free itself from this cover. It carries on
its distal end the female flowers that bloom almost immediately upon

May 15, 1940 CHAMBLISS: BOTANY AND HISTORY OF ZIZANIA AQUATICA 195

emergence. They, at once, are receptive to wind-borne pollen from
nearby or distant plants, for the male flowers of the same plant are
still within the leaf sheath.

The male flowers on the lower part of the panicle hang like elon-
gated purple bells, which open a few days later, revealing six yellow
bodies like so many clappers, which are filled with pollen that is soon
discharged into the air to be carried by the wind to other plants.
This pollen takes no part in the fertilization of the female flowers of
the same flower cluster. The male flowers on long drooping branches
arranged in whorls add beauty and symmetry to this tall, slender,
stately plant attractively dressed with long, broad, hanging leaves.

The slender, flexible panicle, when fully protruded, may be 30 to
50 inches above the terminal leaf. On its topmost branches the seeds
are developing very irregularly. As they approach maturity, which
normally occurs within 15 days after fertilization, the seeds drop very
readily, passing quickly through the water to the mud bottom be-
low. They do not float and are soon anchored in a soft bed by the
many bristlelike hairs on their outer surface. These structures, by
their number and arrangement, serve to fasten the seeds more se-
curely in the mud. Here they lie until the following spring, when the
majority of them germinate.

Zizania seed can not be kept in dry storage like other seed. To
retain its viability it must be kept in a wet state and at a tempera-
ture that will prevent fermentation and control germination. There
is no harvesting of this seed in Eastern United States for human use.
Many birds, however, feast upon it and in so doing assist in the
natural sowing of enough seed to provide for next season’s crop. The
dense brown mat of fallen plants and crumpled foliage, which soon
covers the marsh, will again look green in spring when the young
plants, sprouting from this self-sown seed, push their way through
and above this organic debris.

The birds that feed upon this maturing grain are the bobolinks and
red-winged blackbirds. Late in August and early in September these
birds may be seen in large flocks settling on the plants of Zizania
aquatica at meal time, which continues throughout the day. The
wading birds, such as the sora, feed upon the fallen seeds that lie in
shallow water, or exposed on the ground, when the tide is out. Many
species of diving ducks feed upon the seed that has settled in the mud.

The center of the largest area of this uncultivated grain is in the
region of the adjacent sections of Minnesota, Wisconsin, Manitoba,
and Ontario, which is crowded with alluvial bottom lakes, serving as

196 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

sources of many rivers that for great distances meander through a
flat country and give the landscape the appearance of one immense
marsh. The Fox River, on which the earlier explorers traveled from
the Great Lakes region to the Mississippi River, may be taken as
typical of such streams, filled for the greater part of its length with
Zizania. The Indians?® tell us that this river was made by a mon-
strous serpent that spent the night in the marshes between Lake
Winnebago and the Wisconsin River. Having obtained during the day
enough food to satisfy its hunger, this creature at dusk crawled in
among the vegetation covering this low land to sleep off the lethargy
that accompanies a full meal. While it slept, the dew accumulated
upon its body. At sunrise it awoke and shook the moisture from its
back and in it wriggled toward the larger lake, leaving behind a chain
of small lakes that now are expansions of the river that became the
waterway to the great Northwest. In his account of ascending this
river in 1673, Marquette says, ‘“The way is so cut up by marshes and
little lakes (Fig. 5) that it is easy to go astray, especially as the rive)
is so covered with wild oats [Zizania aquatica] that you can hardly
discover the channel. Hence, we had good need of our two guides.”

In many places throughout this region such conditions exist today,
in normal seasons, from the middle of June until the first of October.
When the waters are free of ice, usually about the middle of May,
Zizania seed begins to germinate. Most of it was sown by nature
early in autumn of the preceding year and some of it through accident
by the Indians themselves in spite of their skill in harvesting the
crop.

The seedlings grow very slowly at first, too weak to stand erect
without the support of the water, which not only surrounds but covers
them, often deeply. During this early growth, with their narrow leaves
floating and reaching upward and outward, the plants appear, when
seen from a canoe floating over an old Zizania bed, like so many
hydrae seeking their prey. In less than a month the young plants
push their leaves to the water surface, spreading them upon it in long
streamers, which at a distance upon good light conditions give the
lake the appearance of having a low verdant island. In approaching
small beds of this plant, the emerging leaves could be taken, even at a
short distance, for a thick growth of duckweeds. In this stage of
growth, the plants are greatly exposed to wave action. By it they may
be detached from the soil and brought to the surface of lake or
stream, leaving only open water where a few days before the young

> THWAITES, REUBEN GoLp. Alistoric waterways, p. 153. 1888.

May 15, 1940 CHAMBLISS: BOTANY AND HISTORY OF ZIZANIA AQUATICA 197

Fig. 5.—Upper: A small lake in northern Minnesota filled with Zzzania aquatica
(“wild rice’). It has the appearance of a low meadow. August 1938. Lower: Young
Chippewa Indians, in boats, have just finished harvesting ‘‘wild rice’ in a Minnesota
lake. The plants are still erect and there is no open water to be seen. September 1938.

plants gave the area the appearance of a green field. Entire stands
covering a hundred acres or more are often destroyed in this way.
When water and weather conditions are favorable, the plants are
strong enough in July to push their stems upward. A few weeks later
the flower clusters begin to emerge. The stems are stronger now and
have become more erect. Continuous sunshine and a mild tempera-
ture ripen the seed in about three weeks.

Among the Indians about this time the topic of conversation is

198 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

“ricing,’’ a word coined by the white man and used only in reference
to harvesting this grain. The urge to assemble at their favorite lakes
begins now to grow upon these people. They drift in singly and in
family groups, settling usually in a wooded spot overlooking the rice-
beds. In a few days a camp of many families is established. The
Indians’ information on the crop is rather definite, for during the
season the rice-producing lakes are frequently scanned by them to
ascertain the stand, vigor of plants, and probable seed production.
Besides their interest in the crop as a source of food and revenue, this
harvest time is the great social event of the year. Within these camps
life presents a picture that is quite primitive even though here and
there are evidences of contact with the white man’s world.

When it becomes known after an inspection that about one-fourth
of the seeds appear ripe, the men and women and the older boys and
girls take to boats (Fig. 5).

The grain is harvested from canoes that may or may not be the
handiwork of Indians. Narrow flat-bottom boats, made of planks and
pointed at each end, are also used. These are made by the Indians
and are as expertly handled by them as the canoes. Either type of
craft is preferred to broad-bottom boats. The latter kind, because of
difficulty in handling, destroys many plants and shatters much grain
that would be gathered from a boat more easily handled. In using the
narrow boats the gatherers may return to the crop as the seed ripens
and this may be done two or three times. With the broad-bottom
boats, which are used by the white men, only one passage over the
ricebeds is ever made.

Our cultivated grain ripens rather uniformly, but not so with this
wild plant. The harvest in a certain locality may extend over a period
of two to three weeks when weather conditions are favorable and
when the crop is in the hands of Indians. The white man who gathers
this grain is not a conservationist. By heritage the Indian is.

Each craft is occupied by two persons, one who stands in the stern,
using a long forked pole to push and guide the boat slowly among the
plants, and the other usually a squaw, who gathers the grain, seated
near the middle of the boat, facing the bow. With two small pointed
sticks, about 30 inches long, one in each hand, the seated person runs
one of the sticks into the plant growth, bending a few plants over the
boat, and strikes the grain-bearing part of the panicles with the other
stick quickly and lightly. The grain, which is easily dislodged, drops
upon the covered bottom of the boat or canoe. This performance 1s
repeated on the other side of the boat and continued alternately |

May 15, 1940 CHAMBLISS: BOTANY AND HISTORY OF ZIZANIA AQUATICA 199

Fig. 6.—Upper: A Chippewa woman parching ‘“‘wild rice’’ in a typical Indian camp,

northern Minnesota. September 1923. Lower: A Chippewa Indian hulling (“‘jigging’’)
parched ‘“‘wild rice’ in a typical Indian camp, northern Minnesota. September 1923.

200 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 5

while the boat is moving until 75 to 100 pounds of seeds Hare been
gathered.

The harvested grain can not be kept perfectly dry while in. ‘the
boat. The added moisture is usually driven off by thoroughly airing
the grain, spread out in the sun on skins, birch bark, blankets, or
canvas. After the grain is dried in this manner for several dave,
though sometimes this step is omitted, it is put into a large iron kettle
or a galvanized iron tub, about 25 pounds at a time, and parched
slowly over a wood fire, being constantly stirred with a paddle to
prevent burning (Fig. 6). Parching requires from 15 to 25 minutes,
depending upon which of the above containers is used. The grain
parched in the iron kettle requires more time for this process and
usually is a better quality than that parched in a tub, probably owing
to the fact that a uniform heat is more easily maintained in the former
than in the latter.

The grain is now ready for the most primitive kind of a mill. This
equipment, which may be termed a mortar, is a hole in the ground
about 14 feet wide and 2 feet deep and lined with a skin. When ready
to operate, about 20 pounds of the parched grain is poured into this
receptacle. A buck Indian (Fig. 6), taking the part of a pestle, steps
upon this loose pile of grain and with a half jump on one foot and then
on the other, combined with a kind of shuffle, treads out the kernels.
While supplying this power the Indian supports himself by poles
driven into the ground near the hole. This process, usually called
“jigging,” detaches the hulls and completes the milling.

The mixture of kernels and hulls taken from the skin-lined hole is
now put into a birch-bark tray, about 30 inches long, 20 inches wide,
and 6 inches deep, to be separated by means of the wind. A windy
day is usually used for this purpose, yet in the hands of a skillful
squaw the tray without the aid of wind becomes a very efficient
fanning mill. The operator, while standing, holds the partially filled
tray even with her waist and slightly inclined. At regular intervals she
tosses the contents of the tray. After each toss, the kernels tend to fall
toward the lower side of the tray and the hulls toward the upper side.
After this partial separation the mixture is tossed higher into the air,
and at the same time with a quick movement of the wrists the tray
is turned forward, producing enough wind to throw much of the chaff
several feet away. This operation is repeated until the chaff is com-
pletely removed. If the grain was fully matured when gathered and
the hulls loosened and detached by parching and “‘jigging,’’ this
primitive method of cleaning leaves only the heavy kernels, which,

May 15, 1940 CHAMBLISS: BOTANY AND HISTORY OF ZIZANIA AQUATICA 201

after washing in several changes of water, are ready for cooking
(Fig. 4, C).

Some new methods of preparing the grain, brought about through
intercourse with the white man, are gradually being used by the
Indians. The primitive method of parching and hulling, however, al-
though it does take time and labor, produces a product superior in
quality to that so far obtained by the white man. The use of modern
machinery for cleaning in place of the birch-bark tray could be used
to advantage and is now being considered by dealers who are seeking
larger markets for this cereal.

The Indians known as wild-rice gatherers belong to the two great
linguistic stocks,* the Algonquian and the Siouan. The former includes
the Chippewa, Menomini, Cree, Fox, and other small tribes. Among
the latter the principal tribes are the Sioux, Winnebago, and Assini-
boin. Probably the Menomini and Winnebago were the first im-
portant tribes to enter the eastern border of the great wild-rice
country, for when found by Nicollet in 1634 they were well estab-
lished in the vicinity of Green Bay, an area now within the State of
Wisconsin. They had migrated from the Atlantic seaboard.

According to Indian tradition the Menomini tribe has been identi-
fied with wild rice for remote ages. Their name is usually translated to
mean “‘wild-rice people.” It has been their belief that ‘“whenever the
Menomini enter a region the wild rice spreads ahead, whenever they
leave it the wild rice passes.’’’ In their economy agriculture had a very
minor place. They lived almost exclusively on game and on plants
requiring no cultivation. They put a high valuation on wild rice and
considered it a gift from the spirit powers, and therefore it has always
been an essential basis for their ceremonial feasts and offerings.

In 1852 the Federal Government assigned to this tribe a large
timber tract on the upper Wolf River as a permanent reservation.
Here they are today, no longer ‘“‘wild rice people”’ but foresters en-
gaged in lumbering, having a tribally owned mill at Neopit that has
been in operation since 1908. Within their reservation there are a few
wild-rice patches, but they receive no attention because they are too
small to produce a worth-while crop. Although the Menomini do not
gather wild rice today, they still use it ceremonially.

The Winnebago Indians were less nomadic than the other Siouan
tribes and lived near the waterways in preference to a life on the
plains, which the Sioux enjoyed so much. For food they, like the

6 JENKS, ALBERT ERNEST. Op. cit., p. 1038.
’ Kexsine, Fevix M. The Menomini Indians of Wisconsin. Mem. Amer. Phil.
Soc. 10: x1+261. 1939.

202 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 5

Menomini, depended upon fish, small mammals, wild rice, maple
sugar, and berries.

When the Chippewa, one of the largest tribes north of Mexico, be-
gan to move westward, they were driven forward by the Iroquois,
who occupied land that was not overstocked with game but well
suited for cultivation. These newcomers, being hunters, were not wel-
come by the tillers of the soil and were forced by circumstances to
continue westward. As the Chippewa moved onward, they encoun-
tered the eastern bands of the Sioux tribe occupying the lake region
now a part of Wisconsin and Minnesota.

This part of the great central valley of this continent is filled with
innumerable shallow lakes and sluggish streams that at one time
contained an unfailing supply of food. To the red man centuries
ago this region was a hunter’s paradise. Besides wild rice and fishes and
other water-inhabiting animals, it contained for the aborigines an
inexhaustible supply of land animals too, the flesh and hides of which
provided food and clothing. These natural resources were considered
by the Sioux who first possessed them and by the Chippewa who de-
sired them as tribal property of the greatest value, and each fought
fiercely to get control of them. To these gifts of Nature should be
added the beautiful and useful birch, the bark of which the original
inhabitants and their descendants have used to cover their lodges,
wigwams, and canoes. The bark of the basswood, too, has con-
tributed much to the wants of these people.

Without a decisive engagement at any time, the struggle between
these tribes for the full possession of this country continued at in-
tervals for several centuries until 1862, when the Government re-
moved the Sioux.

The Chippewa never had an undisputed control of these lakes,
though for several hundred years they ventured upon them to gather
wild rice, often at the cost of a heavy loss of life. The Indian who
gathers wild rice in the United States today is of this tribe. His ap-
praisal of these lakes and woodlands made centuries ago has not
changed with the years. So, on many occasions, when new treaties
had to be made to gratify the greed of the white man, the Chippewa
Indians have asked our Government to give heed to their needs.

As late as 1863, Hole-in-the-Day, the leading chief of the Chip-
pewa, addressed a pathetic appeal to the Great Father at Washing-
ton, which, in part, is as follows :8

® Report of the Commissioner of Indian Affairs for the year 1863, pp. 328-331.

May 15, 1940 CHAMBLISS: BOTANY AND HISTORY OF ZIZANIA AQUATICA 203

My people are unhappy and dissatisfied. I want to see them happy and
contented. It is both to their interest and the interest of the white man
that they should be so, and they require but little to make them so. * * *

The present treaty gives us little but swamps or marshes, where locations
can be selected that combine all these elements of comfort and content to
our people, that is, good land, game, fish, rice and sugar. Here, we have
neither to any considerable extent. True, we may find a little rice and a
few fish, but not sufficient for my people, not enough to save them from
starvation. If a treaty were made with the Red Lake Indians, a tract of
country of the best character for my people might be secured without any
outlay of expense to the government: say that strip of land lying on the
Wild Rice River between the 47° and 48° north latitude and east of the
Red River. There is every advantage of good soil, game, fish, rice, sugar,
cranberries and a healthy climate. * * *

This late treaty never will, never can satisfy our people. A reservation
on the Wild Rice would satisfy them all, and they would leave their present
homes and go to their new ones happily and with a feeling that a better
future was before them. * * *

The sooner this is done the better, as it would have a tendency to quiet
the discontent now existing among our people generally, by holding out to
them a prospect of a good and pleasant home somewhere near or in the
valley of the Wild Rice. * * *

Believe me, then, my Father, to be what my people always have been
and what they and I now am, Your friend and the friend of the white man.

Wild rice has played an important part in providing subsistence
to explorers and trappers who penetrated this great continent two or
three centuries ago along the waterways that now separate the United
States from the Dominion of Canada.

A traveler among the North American Indians during the years
1652 to 1684, after referring to his reception by the natives, states,
“Our songs being finished we began our teeth to worke. We had there
a kinde of rice, much like oats. It growes in the watter in 3 or 4 foote
deepe.’’ After comments on God’s care over His creatures and a brief
description of how the grain is gathered, he continues: ‘“‘That is their
food for the most part of the winter and doe dresse it thus: ffor each
man a handfull of that they putt in the pott, that swells so much that
it can suffice a man.’’®

In his Travels and adventures in Canada and the Indian Territories,
Alexander Henry” tells us a hundred years later about obtaining from
Indian women by barter 100 bags of this grain and adds that ‘‘with-
out a large quantity of rice the voyage could not have been prose-
cuted to its completion.”’

9 Voyages of Peter Esprit Radisson. Transcribed from original manuscripts in the
Bodleian Library and the British Museum. The Prince Society, Boston, 1885

10 Henry, ALEXANDER. Travels and adventures in Canada and the Indian Terri-
tories, 1760-1776, new ed., p. 241. 1901.

204 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

An ample supply of wild rice was always included among the
winter provisions for the outposts of the fur companies trading in this
region. David Thompson" records in his Narrative of explorations
in western America that a superintendent of a fur company in north-
ern Minnesota and his men “passed the whole winter on wild rice and
maple sugar.’’ Under such circumstances he considered this grain
‘“‘a weak food”’ and says that “those who live for months on it enjoy
good health and are moderately active but very poor in flesh.”

The old journals of the earlier hunters, trappers, and priests seek-
ing adventure in this great wilderness of the north contain many
stories about the use of this grain in fighting hunger when other food
was hard to get. In the great outdoors, white men, like Indians, can
retain health, develop endurance, and enjoy life on a very simple diet.

The aborigines of this country were fond of soups, broths, and stews
thickened with wild rice.’ To this day, their descendants have not
lost that fondness. In addition to its simplicity, this dish has the char-
acter of elasticity, which appeals to the Indians. With them mealtime
is often visiting time. A meal started with three people may be in-
creased to ten or more before it is finished. The uninvited guests
never seem to cause embarrassment. To provide for them it is neces-
sary only to add water to increase the volume of the soup.

With a squaw as a cook a favorite dish in the camp of rice-gather-
ing Indians has been wild rice, corn, and fish boiled together, called
‘“’Tassimanonny.’’!® This combination of foods has also tickled the
palate of the white man, as may be judged by the enthusiasm of one
who spoke of it in his later years as being ‘‘an object of early love.’
The Indian likes sweets and often eats his boiled rice with maple
sugar.!* He may also flavor his boiled rice with cranberries and his
soup with blueberries. Boiling does not, as a rule, reduce the kernels
to a paste, but should this condition occur, the pastelike mass is used
by the Indians as a substitute for bread.

Some Indians parch wild rice until the kernels burst open as pop-
corn does when heated, and eat it in this condition when away from
camp. Because of its keeping qualities, the parched grain is recom-

11 THompson, Davin. Narrative of explorations in western America, 1784-1812.
The Champlain Society. Toronto 1916.

12 SmirH, Huron H. Ethnobotany of the Meskwaki Indians. Bull. Publ. Mus.
City of Milwaukee 4: 259. 1928.

13 BippLE, JAMES W. Recollections of Green Bay in 1816-1817. Appendix No. 4.
First Ann. Rept. and Collections of the State Hist. Soc. of Wisconsin for the year 1854,
1: 63.

144 DunBaAR, SIR GEORGE. Other men’s lives, p. 149. 1938.
15 STICKNEY, GARDNER P. Indian use of wild rice. Amer. Anthrop. 9: 115-121.

May 15, 1940 BLAKE: NEW FOSSIL FISH 205

mended to vacation campers for use in either the dry or boiled state.
In the land of ‘‘Ten Thousand Lakes” where the parched grain is
comparatively cheap and is usually a part of the daily meal, the
woodman has the advantage of the man of the city, who must pay
exorbitant prices for it to cover handling charges and profits. Under
these circumstances wild rice in the city home is seldom used except
on special occasions.

Knowing how some like the grain, we may assume that all would
be just as enthusiastic “‘ffor each man a handfull of that they putt
in the pott”’ and would exclaim with Wenibozho, ‘“‘Oh, you are indeed
good!’’

PALEONTOLOGY .—Paralbula, a new fossil fish based on dental
plates from the Eocene and Miocene of Maryland.’ 8. F. BLAKE,
Washington, D. C.

In the fall of 1935, my wife, Doris H. Blake, found a nearly perfect
specimen of the crushing dentition of some fish on the shore of the
Potomac River at Liverpool Point, Charles County, Md. The deposits
exposed in the bank at this place belong to the Piscataway member of
the Aquia formation (Lower Eocene), and the specimen, although not
found in situ, can be attributed to this period with as much confidence
as the teeth of Lamna obliqua and other sharks found in abundance
on the Potomac shore at the same locality. The specimen was given
to the U. 8. National Museum and has since been recorded and figured
by Dr. George 8. Myers.? Dr. Myers identified the specimen as one
of the Albulidae (ladyfishes), which he regarded as including among
living forms only two monotypic genera, but he refrained from giving
a name to the specimen or even assigning it to a genus because the
crushing tooth plates of the two existing genera are not distinguish-
able. |

The nearly perfect condition of this specimen and the fact that it
was the first fossil representative of the Albulidae to be recorded from
America, with the exception of a scale from the Cretaceous of Florida
described as Albula antiqua Cockerell,* seemed to make it desirable to
assign a specific name to it, if only to prevent it from being over-
looked. Examination of the pertinent literature and of alcoholic
specimens of Albula vulpes kindly made available by Dr. L. P. Schultz

soon showed that this dental plate could not be referred to either of

1 Received January 12, 1940.

* Myers, G.S. A third record of the albulid fish Dixonina nemoptera Fowler, with
notes on an albulid from the Eocene of Maryland. Copeia 1936(2): 83-85, fig. 1. 1936.

3 Copeia 1933: 226. 1933.

206 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

the recognized existing genera of the Albulidae. In Albula, the best-
known representative of the family, the lower dental plate is twice as
long as wide, or more, and bears only a single layer of teeth, which on
dropping out leave a circular rounded-out pit in the surface of the
supporting bone. This description applies to the fossil species, two of
which are described from the Eocene of England, and one from that
of Nigeria, as well as to the living one. The Maryland fossil is much
wider in proportion to its length, and its teeth are superposed in ir-
regular piles of 4 to 6, or probably more in the center of the plate, and
on breaking or wearing they do not leave a cavity but are replaced
from beneath, as in Phyllodus and some other genera of fossil fishes.
The teeth, moreover, as shown in the accompanying sketches, are
quite different in structure from those of Albula.

In superficial appearance the plate in question is very similar to the
original illustrations of Hgertonza, a genus described by Cocchi from
the Eocene of England. Errol I. White, of the British Museum, to
whom an enlarged photograph was sent for comparison with the type
of Egertonia, reported that the latter is very different, the teeth re-
sembling those of the related Phyllodus, being very thin, usually
only slightly convex, with very numerous successors in piles, while
both upper and lower plates are sigmoid in profile. Mr. White regards
the Maryland specimen as representing an apparently undescribed
genus, which may bear somewhat the same relationship to the albu-
lids that Labrodon does to the labroids and Diaphyodus to the sciae-
nids—that is, a fossil form with pharyngeal dental plates similar in
general to those of living forms, but having each tooth subtended by
a pile of successional teeth. Its assignment to the Albulidae, however,
must be regarded as only tentative.

Paralbula Blake, gen. nov.

Pharyngeal dentition similar to that of Albula, but each tooth with a pile
of subvertically arranged successors.
Type species, P. marylandica, n. sp.

Paralbula marylandica Blake, sp. nov.

A nearly oval lower pharyngeal dental plate measuring 35 mm in length,
26 mm in width, and 11 mm in depth, broadly rounded at anterior end,
subtruncate behind, convex-rounded on the oral surface, somewhat flat-
tened along midline especially posteriorly, slightly concave on the attached
surface. Triturating surface densely covered with jet-black, shining, sub-
circular, depressed-hemispherical teeth, about 141 in all (including bases of
broken-off teeth), not arranged in definite lines, the central ones 3.2-3.5 mm
in diameter, the lateral gradually decreasing in size, the outermost 2-2.5 mm
in diameter. The teeth are irregularly superposed in several layers separated
by bony tissue, not in direct vertical piles, and have a very smooth, rounded, |

May 15, 1940 BLAKE: NEW FOSSIL FISH 207

Fig. 1—A, Paralbula marylandica, tritoral surface; B, attached surface; C, lateral
view; D, Paralbula dorisiae, tritoral surface; E, lateral view; F, attached surface.
All X nearly 2.

208 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

depressed-hemispheric crown inflexed at base to form a horizontal, some-
what radially striate rim about 0.5 mm wide. On the attached surface, the
specimen is somewhat more deeply eroded at the anterior end, and is there
loosely and irregularly cellular, with occasional interspersed bases or in-
teriors of teeth. The posterior half is better preserved and shows the bone
cells arranged mostly in definite longitudinal rows, with on each side near
the margin of the bone a shallow more or less wedge-shaped depression,
bounded in front and on the inner side by a rather indefinite ridge, these
evidently being the scars marking the points of attachment to the support-
ing pharyngeal bones.

Type.—U.S.N.M. no. 138855, collected on shore of Potomac River at
Liverpool Point, Charles County, Md., October 13, 1935, by Doris H. Blake.
The deposits at this point, from which the specimen undoubtedly came,
belong to the Piscataway member of the Aquia formation, Lower Eocene.

In my own collection is a detached tooth with eroded base, measuring 4.5
mm in diameter, and evidently belonging to the same species, which I
picked up on the beach at the same locality in 1938.

As this paper was being prepared for publication, Mrs. Blake picked up on
the beach near the wharf at Plumpoint, Md., a much smaller and more im-

Fig. 2.—a-c, Paralbula marylandica: a, Cross section of tooth; b, view of tooth from
beneath, showing the inflexed rim; c, view of scar left on oral surface of plate by a de-
tached tooth. The outer ring, which is somewhat impressed and slants slightly toward
the center, represents the area occupied by the inflexed rim of the tooth; the inner ring,
which is elevated above the outer, is evidently a ring of denser bony tissue subtending
the tooth inside the inflexed rim, and fitting against the somewhat beveled inner edge
of the rim; the central circle is cellular bony tissue. d—e, Paralbula dorisiae; d, Typical
tooth from beneath; e, from the side. All <5.

perfect specimen that seems to represent a second species of the genus. The
teeth are higher in proportion to their width, the root is more slanting and
much more strongly grooved, and the opening in its center, which is scarcely
at all excavated, is much smaller. As no Eocene deposits are exposed within
miles of Plumpoint, it seems necessary to ascribe this specimen to the Mio-
cene, and probably to the Calvert formation, the one best developed at Plum-
point.

Paralbula dorisiae Blake, sp. nov.

An irregularly oval, much eroded dental plate 15.5 mm long, 12 mm wide,
and about 4 mm deep, obscurely convex in cross section on oral surface and
essentially plane in longitudinal section, flattish on attached surface. Tri-
turating surface densely covered with blackish brown to light brown, sub-
circular, depressed-subspherical teeth, about 40 in all (including bases of
broken-off teeth), not arranged in definite lines, 1.5-2.2 mm in diameter,
1.2-1.7 mm deep. Teeth irregularly superposed in about three layers, or
perhaps more in the center of the plate, separated by bony tissue, not 1n

May 15, 1940 SWALLEN: NEW AMERICAN GRASSES 209

direct vertical piles, with a smooth, almost hemispherical crown and
strongly striate, slanting root often almost equaling the height of the crown
and tapering to a subtruncate tip 0.8-1 mm wide. The cavity left in the
bone by detached teeth is deeper than that in P. marylandica, with more
strongly grooved sides, and the striated inner ring is not at all elevated;
the central area is perforate in all the cavities visible in the specimen.
Type.—U.S.N.M. no. 16134, collected on shore of Chesapeake Bay close
to and just south of the wharf at Plumpoint, Calvert County, Md., August
13, 1939, by Doris H. Blake. Probably from the Calvert formation of the
Miocene, the best developed formation in the vicinity of Plumpoint.

BOTAN Y.—Miscellaneous new American grasses.:. JASON R. SWAL-
LEN, U.S. Bureau of Plant Industry.

During the identification of various collections of grasses in the
past few years, the following species were found to be new, three of
which are from the United States, i.e., Poa fibrata, Stipa diegoensis,
and Digitaria albicoma.

Chusquea sulcata Swallen, sp. nov.

Culmi graciles ramis floriferis fasciculatis adscendentibus 10-25 cm longis;
vaginae internodiis longiores, glabrae, in ore hispidae marginibus ciliatis;
ligula 0.5 mm longa, truncata; laminae 6-12 cm longae, 4-6 mm latae,
acuminatae, glabrae marginibus sparse papilloso-hispidis; paniculae 4-10
em longae, 5-10 mm latae, densae, ramis brevibus appressis; axis pubescens;
glumae minutae obtusae enerves; lemmata sterilia obtusa vel subacuta,
obscure 1—5-nervia, 2.5-3 mm longa; lemma fertile 6-7 mm longum, sub-
acuminatum, obscure 7-nerve, exaristatum, glabrum; palea lemma aequans,
sulcata, carinis approximatis pubescentibus, marginibus latis; antherae 4
mm longae.

Culms slender, probably clambering, the flowering branches in rather
small ascending fascicles, 10-25 cm long; sheaths longer than the internodes,
glabrous on the back, hispid in the throat, the margins ciliate; ligule about
0.5 mm long, truncate; blades 6-12 cm long, 4-6 mm wide, acuminate, nar-
rowed toward the base, glabrous, sparsely papillose-hispid on the margins;
panicles 4-10 cm long, 5-10 mm wide, rather dense, the short branches ap-
pressed, floriferous to the base, the axis glabrous or obscurely scabrous on
_ the angles; glumes minute, obtuse, nerveless; sterile lemmas obtuse or sub-
acute, apiculate, 1—5-nerved, the lateral nerves often obscure, 4 to 4 as long
as the floret; lemma 6-7 mm long, subacuminate, obscurely 7-nerved, awn-
less, glabrous; palea as long as the lemma, sulcate, the keels close together,
cent, the broad glabrous margins enclosing the flower; anthers 4 mm
ong.

Type in the U. 8. National Herbarium, no. 1722000, collected on Mount
Orando, Chiapas, Mexico, April 1936, by E. Matuda (no. 321).

This species superficially closely resembles C. nelsoni Scribn., which has
shorter and broader blades, pubescent panicle axis, larger glumes, acuminate,
sterile lemmas, and strongly nerved, sparsely pilose, fertile lemma.

* Received October 26, 1939. All drawings by Mrs. Frances C. Weintraub.

210 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 5

Chusquea longifolia Swallen, sp. nov.

Rami floriferi dense fasciculati adscendentes 20-60 cm longi; laminae 12-
25 cm longae, 6-12 mm latae, marginibus scabris; paniculae 10-16 cm longae,
angustae ramis gracilibus appressis paucifloris scabris; spiculae 12-13 mm
longae appressae; glumae obtusae, enerves, prima 0.5 mm secunda 1 mm
longa; lemma sterile primum 7—9 mm longum, 9-nerve, acutum, apiculatum,
basi latum; lemma sterile secundum lemma fertile aequans, 9-nerve, acumi-
natum, superne scabrum; lemma fertile 11-12 mm longum, obscure nervo-
sum, puberulum, acuminatum, minute bifidum, aristatum, arista 2-4 mm
longa; palea lemmate fertili paulo longior, carinis approximatis, bidentata,
marginibus latis.

Main culm about 5 mm thick (in flowering part) the flowering branches
fascicled, ascending, 20-60 cm long; blades 12-25 cm long, 6-12 mm wide
with scabrous margins; panicles 10-16 cm long, narrow, the slender branches
appressed, scabrous, few-flowered; spikelets 12-13 mm long, appressed;
glumes obtuse, nerveless, the first 0.5 mm long, the second 1 cm long; first
sterile lemma 7—9 mm long, 9-nerved, acute, mucronate, very broad at the
base; second sterile lemma as long as the fertile one, 9-nerved, acuminate,
scabrous toward the tip; fertile lemma 11-12 mm long, obscurely nerved,
puberulent, acuminate, minutely bifid, with an awn 2—4 mm long; palea as
long as or a little longer than the lemma, bidentate, the keels approximate,
the margins very broad, incurved.

Type in the U.S. National Herbarium, no. 1724503, collected on Mount
Tacana, Chiapas, Mexico, altitude 2,000—4,088 meters, in August 1938,
by E. Matuda (no. 2373).

Poa fibrata Swallen, sp. nov.

Perennis, rhizomatosa; culmi rigidi, erecti vel basi adscendentes, 15-35
em alti, glabri vel scaberuli; vaginae inferiores tenues, lucidae, undulatae,
fibrosae, glabrae, superiores internodiis longiores, firmae, scabrae; ligula
1-1.5 mm longa acute vel obtusa; laminae 4—8 cm longae vel supreme re-
ductae firmissimae, conduplicatae, curvatae, scabrae, marginibus minute
serratis; panicula 4-10 cm longa, densa, ramis brevibus appressis ad basin
floriferis vel basi nudis, scabris; spiculae 3—4 florae, 5-6 mm longae; glumae
acutae vel subobtusae, 1—3-nerves, prima 2-3 mm longa, secunda 3 mm
longa; lemmata 2.5-3 mm longa, acuta vel subobtusa, glabra vel scabra vix
pubescentia; palea subacuta, lemma subaequans; antherae 2 mm longae.

Perennial with slender, creeping rhizomes; culms 15-385 ecm tall, rigid,
erect from an ascending base, glabrous or scaberulous; lower sheaths thin,
smooth and shining, undulate, becoming more or less fibrous with age, the
upper sheaths longer than the internodes, firm, scabrous; ligule 1-1.5 mm
long, acute or subobtuse; blades 4-8 em long, or the uppermost some-
times reduced, very firm, conduplicate, curved, pungent, scabrous, the
margins minutely serrate; panicle 4-10 cm long, dense, with short appressed
branches floriferous to the base, or the lower ones sometimes naked at the
base, scabrous; spikelets 3—4-flowered, 5-6 mm long; glumes acute or sub-
obtuse, 1—3-nerved, the first 2-3 mm long, the second 3 mm long; lemmas
2.5-3 mm long, acute or subobtuse, glabrous or scabrous, sometimes rather
obscurely pubescent near the base and on the margins; palea subacute,
about as long as the lemma or a little shorter.

Type in the U. S. National Herbarium, no. 1646953, collected in a dry
subsaline flat, 2 miles south of Grenada, Shasta Valley, Siskiyou County,
Calif., altitude 2,600 feet, June 30, 1935, by Louis C. Wheeler (no. 3629),

May 15, 1940 SWALLEN: NEW AMERICAN GRASSES 211

Poa mulleri Swallen, sp. nov.

Perennis; culmi caespitosi erecti 25-40 cm alti, compressi; foliae prope
basin congestae; vaginae carinatae glabrae, eae culmorum elongatae; ligula
0.5 mm longa, membranacea, truncata; laminae innovationum laxae, elonga-
tae, involutae, eae culmorum planae, 3-6 cm longae, 1-2 mm latae, glabrae;
paniculae 5-8 cm longae pyramidatae; rami solitarii vel bini, graciles, flex-
uosi, reflexi, ad 3 cm longi, basi nudi; spiculae biflorae, 4 mm longae; gluma
prima 3 mm longa, 3-nervis, acuta, navicularis; gluma secunda 3.5 mm
longa, 5-nervis, abrupte acuta; lemma, inferius 3.5 mm longum, 5-nerve,
acutum, carina et nervis marginalibus inferne pubescentibus; palea lemmate
paulo brevior; antherae 2 mm longae.

Perennial; culms loosely cespitose, 25—40 cm tall, erect from an ascending
base, conspicuously flattened; leaves mostly crowded toward the base of the
culms; sheaths compressed keeled, glabrous, those of the culm elongate;
ligule 0.5 mm long, truncate; blades of the innovations lax, elongate, in-
volute, those of the culm flat, 3-6 cm long, 1—2 mm wide, glabrous; panicles
5-8 em long, pyramidal, the branches solitary or in pairs, slender, flexuous,
reflexed, the lower ones as much as 3 cm long, naked in the lower half;
spikelets clustered on the upper half of the branches, appressed, short
pediceled, 2-flowered; glumes navicular, the first 3 mm long, 3-nerved, the
second 3.5 mm long, 5-nerved, usually rather abruptly acute; lower lemma
3.5 mm long, 5-nerved, acute, sparingly pubescent on the keel and marginal
nerves toward the base; palea a little shorter than the lemma; anthers 2
mm long.

Type in the U. 8. National Herbarium, no. 1645320, collected in pine
woods on the Peak of Cerro Potosi, Municipio de Galeana, Nuevo Leon,
Mexico, July 21, 1935, by C. H. Muller (no. 2251).

The very flat culms with short reflexed panicle branches are characteristic.

Stipa linearis Swallen, sp. nov.

Perennis; culmi graciles, dense caespitosi, erecti, 50-75 cm alti, 2-nodes;
foliae ad basin dense congestae; vaginae culmorum elongatae internodiis
breviores, subplanae, glabrae, eae innovationum breves, angustae; ligula

Fig. 1.—Stipa linearis, mature floret, <1, and the body of the lemma, X5.

brevissima vel ad 3 mm longa; laminae culmorum 3-10 em longae, planae
vel involutae, eae innovationum ad 35 cm longae, 1—-1.5 mm latae, planae vel
subinvolutae, infra glabrae supra scaberulae, marginibus scabris; panicula
ad 13 cm longa, angusta, ramis gracilibus, appressis, paucifloris, 1-3.5 cm
longis; glumae aequales 5 mm longae, acutae, prima 3—5-nervis, secunda
5-nervis; lemma 4-4.5 mm longum, fuscum, sparse pilosum; callum 0.5 mm
longum, hebes, dense barbatum; arista 2 cm longa, bigeniculata.

Perennial; culms slender, densely cespitose, erect, 2-noded, 50-75 cm tall,

212 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

the numerous leaves of the innovations crowded toward the base in a dense
cluster; culm sheaths elongate, shorter than the internodes, becoming almost
flat, glabrous, those of the innovations short, narrow; ligule very short in
the innovations, 1.5-3 mm long on the culm leaves; culm leaves 3-10 em
long, flat or involute, those of the innovations as much as 35 cm long, 1—1.5
mm wide, flat or subinvolute, glabrous below, scaberulous above, the mar-
gins scabrous; panicle up to 13 cm long, narrow, the slender appressed, few-
flowered branches 1-3.5 em long; glumes equal, 5 mm long, acute, green,
hyaline toward the tip, the first 3-5-nerved, the second a little broader,
always 5-nerved; lemma 4—4.5 mm long, abruptly narrowed to the asym-
metrical summit, dark brown, sparsely pilose with pale or brownish hairs;
callus 0.5 mm long, blunt, densely barbate; awn 2 cm long, somewhat ob-
scurely twice geniculate.

Type in the U. S. National Herbarium, no. 1610598, collected in Diente
Canyon, 12 miles south of Monterrey, Nuevo Leon, Mexico, July 13, 19383,
by C. H. Muller (no. 483).

This species is closely related to Stzpa augustifolia Hitche., which is a
smaller plant, 10-30 cm tall, with short involute blades and longer glumes
and lemma.

Stipa acuta Swallen, sp. nov.

Perennis; culmi graciles caespitosi, erecti, ad 95 cm alti; vaginae inter-
nodiis breviores, glabrae; ligula 1.5-2 mm longa vel eae innovationum
brevissimae; laminae involutae ad 23 cm longae, firmae, glabrae, marginibus
scabris; paniculae longe exsertae, 15-20 cm longae, laxae, ramis gracilibus,
adscendentibus vel divergentibus, basi nudis, paucifloris, ad 6 cm longis;
spiculae appressae; glumae acutae, hyalinae, reticulatae, basi purpurascentes,
prima 11-12 mm longa, 5-nervis, secunda 9-10 mm longa, 5—7-nervis; lemma
7mm longum, fusiforme, fuscum, pilosum pilis albis appressis; callum 1 mm
longum, pungens, dense pilosum; arista 4—4.5 mm longa, bigeniculata.

Perennial; culms slender, tufted, erect, about 95 cm tall; sheaths shorter
than the internodes, glabrous; ligule 1.5 to 2 mm long, or very short on the
innovations; blades involute as much as 23 cm long, firm, glabrous with
scabrous margins; panicles long exserted, 15-20 cm long, lax, the slender
ascending or spreading, somewhat flexuous glabrous branches naked at the
base—usually in the lower half, few-flowered, as much as 6 cm long; spike-
lets appressed, the pedicels 2-5 mm long; glumes acute, hyaline, reticulate
veined, tinged with purple at least toward the base, the first 11-12 mm long,
5-nerved, the second 9-10 mm long, 5—7-nerved; lemma 7 mm long, fusiform,
very dark brown, rather densely appressed-pilose with white hairs; callus
1 mm long, sharp pointed, densely appressed pilose; awn 4—5 cm long, twice
geniculate, scabrous, the terminal segment slender, somewhat flexuous,
3-3.5 cm long.

Type in the U.S. National Herbarium, no. 1760238, collected on rocky soil
on Carneras Pass, 21 miles south of Saltillo, Coahuila, Mexico, September 1,
1938, by Forrest Shreve (no. 8545).

Stipa acuta is most closely related to S. eminens Cav., which has nar-
rower, 3-nerved, acuminate glumes, a pale lemma 5-6 mm long, and a some-
what shorter awn.

Stipa diegoensis Swallen, sp. nov.
Perennis; culmi densi caespitosi, erecti, 70-100 cm alti, scaberuli, infra

nodos densi pubescentes; vaginae inferiores internodiis longiores, superiores
internodiis breviores, elongatae, glabrae vel scaberulae; ligula 1-2 mm longa,

May 15, 1940 SWALLEN: NEW AMERICAN GRASSES 213

obtusa vel truncata, ciliata, pubescens; laminae 15-40 cm longae, 2-4 mm
latae, planae vel involutae, attenuatae, infra scaberulae, supra pubescentes;
panicula 15-30 cm longa, densa, angusta, ramis appressis ad 10 cm longis;
gluma prima 9-10 mm longa, 1-nervis, acuminata; gluma secunda 8-9 mm
longa, 3-nervis, acuminata; lemma 6.5-7.5 mm longum, pilosum, pilis su-
premis 1-2 mm longis; callum 0.5 mm longum, curvatum, pungens, pilosum;
arista 2-3.3 cm longa, bigeniculata, scabra; palea 4 mm longa, acuta, pilosa;
antherae 4 mm longae, purpurascentes.

Perennial; culms densely tufted, erect, 70-100 cm tall, scaberulous,
densely pubescent below the nodes; lower sheaths longer than the inter-
nodes, the upper ones shorter than the internodes, elongate, glabrous or
scaberulous; ligule 1-2 mm long, obtuse or truncate, ciliate, pubescent;
blades 15-40 em long, 2-4 mm wide, flat or involute, scabrous on the lower
surface, pubescent on the upper; panicle 15-30 cm long, narrow, dense, the

Fig. 2.—Stipa diegoensis, mature floret, X1, and the body of the lemma, X65.

branches appressed as much as 10 em long; first glume 9-10 mm long, 1-
nerved, acuminate; second glume 8-9 mm long, 3-nerved, acuminate;
lemma 6.5-7.5 mm long, pilose, the hairs at the top of the lemma, 1-2 mm
long; callus 0.5 mm long, curved, sharp-pointed, appressed pilose; awn 2-3.3
cm long, bigeniculate, scabrous; palea 4 mm long, acute, pilose; anthers 4
mm long, purple.

Type in the U. 8. National Herbarium, no. 1761177, collected along
vernal stream in chaparral, Proctor Valley near Jamul, San Diego County,
Calif., May 23, 1938, by Frank F. Gander (no. 5778).

This species is apparently related to S. editorwm Fourn., which is a much
more slender plant, glabrous below the nodes and has thinner glumes, the
second 1-nerved or obscurely 3-nerved, a shorter lemma 5-6 mm long, and a
more slender obscurely geniculate awn.

Stipa bracteata Swallen, sp. nov.

Perennis; culmi ad 125 cm alti, caespitosi, erecti, 3-nodes, infra nodos
dense retrorse pubescentes, infra paniculam bracteatae; vaginae internodiis
breviores inferne pubescentes, in ore sparse pilosae; ligula 1.5-3 mm longa;
laminae 25-40 cm longae, 1-3 mm latae, planae vel involutae, infra glabrae,
supra pubescentes; panicula 35-42 cm longa, laxa, fasciculis ramorum re-
motis; rami graciles 4-10 em longi adscendentes vel divergentes, basi nudi;
spiculae appressae; glumae acuminatae, hyalinae, 3-nerves, prima 10-11
mm longa, secunda 8-9 mm longa; lemma 5.5-6.5 mm longum, fusiforme,

214 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

fuscum, pilosum pilis albis appressis; callum 0.5 mm longum, curvatum,
dense barbatum; arista 20-24 mm longa, bigeniculata, glabra.

Perennial; culms as much as 125 cm tall, erect, tufted, 3-noded, densely
retrorsely pubescent below the nodes, with a conspicuous ciliate bract 4-6
mm long below the panicle; sheaths shorter than the internodes, pubescent
near the base, sparsely pilose at the throat, those of the innovations densely
pubescent on the collar; blades 25-40 cm long, 1-3 mm wide, flat or involute,
glabrous on the lower surface, pubescent on the upper; panicle 35-42 cm
long, lax, the branches in small remote fascicles, 4-10 cm long, some long
and some short in the same fascicle, slender, ascending or spreading, naked
at the base, the longer ones usually spikelet bearing only near the ends;
spikelets appressed; glumes acuminate, hyaline, 3-nerved, the first 10-11
mm long, the second 8-9 mm long; lemma 5.5—6.5 mm long, fusiform, brown,
pilose with white appressed hairs; callus 0.5 mm long, somewhat curved,
rather blunt, densely barbate; awn 20-24 mm long, bigeniculate, glabrous
or nearly so, the two lower segments twisted, the upper straight.

Type in the U. 8. National Herbarium, no. 1721797, collected on grassy
flats 25 miles north of Ensenada, Baja California, April 4, 1931, by Ira L.
Wiggins (no. 5153). |

Stipa bracteata is related to S. editorum and S. diegoensis but is at once
distinguished by the conspicuous bract below the panicle.

Digitaria albicoma Swallen, sp. nov.

Perennis; culmi caespitosi, erecti, 65-71 cm alti, simplices vel basi ramosi;
vaginae inferiores internodiis multo longiores, dense villosissimae, superiores
elongatae internodiis breviores, glabrae vel ad basin papilloso-pilosae;

Fig. 3.— Digitaria albicoma, spikelet and fruit, X5.

ligula 1.5-—2 mm longa, truncata; laminae planae, 10-80 cm longae, 3-5 mm
latae, pilosae vel papilloso-pilosae, marginibus scabris; panicula longe ex-
serta; racemi 5-9, graciles, adscendentes vel divergentes, 8-12 cm longi
basi nudi; spiculae 2.5-mm longae, solitariae vel binae, inferior subsessilis,
superior pedicello scabro 2 mm longo; gluma prima obsoleta; gluma secunda
angusta 3-nervis, glabra, marginibus hyalinis; lemma sterile lemma fertile
aequans, subobtusum, 5-—7-nerve, glabrum, marginibus hyalinis; fructus
2.5-2.8 mm longus, acutus, plumbeus.

Perennial; culms cespitose, erect, 65-75 cm tall, simple, or branched at
the base; lower sheaths much shorter than the internodes, densely villous,
the upper ones elongate but shorter than the internodes, glabrous or papil-
lose-pilose toward the base; ligule 1.5-2 mm long, truncate; blades flat,
10-30 cm long, 3-5 mm wide, pilose or papillose-pilose, the margins scabrous;
panicle long exerted; racemes 5—9 slender, ascending or spreading, 8-12 cm
long, naked at the base; spikelets 2.5 mm long, solitary or paired, the lower

May 15, 1940 SWALLEN: NEW AMERICAN GRASSES 215

one subsessile, the upper pedicellate, the pedicel 2 mm long, scabrous; first
glume obsolete or nearly so; second glume narrow, 3-nerved, glabrous, the
margins hyaline; sterile lemma as long as the fruit, 5—-7-nerved, glabrous,
the margins hyaline; fruit 2.5—-2.8 mm long, acute, lead-colored.

Type in the U. S. National Herbarium, no. 1761179, collected in low
open sandy woods, Chinsegut Hill Sanctuary, Brooksville, Hernando
County, Fla., November 18, 1938, by Jason R. Swallen (no. 5644).

Digitaria albicoma is closely related to D. subcalva Hitche., which has
many noded culms decumbent at the base and sometimes rooting at the
lower nodes, less conspicuously villous sheaths, and fewer appressed racemes.

Mesosetum comatum Swallen, sp. nov.

Perenne; culmi graciles, erecti, 50 em alti; vaginae internodiis longiores,
glabrae, in ore pilosae, marginibus minute ciliatis; laminae 4-6 cm longae,
2—2.5 mm latae, superiores et eae innovationum reductae, firmae, erectae,
pungentes, infra glabrae, supra glabrae vel hispidae, marginibus sparse
hispidis; gluma prima 3.5 mm longa, obtusa, basi hirsuta pilis ad 4 mm
longis, in parte superiore scabra; gluma secunda 3 mm longa, 3-nervis, basi
pilosa, superne glabra, marginibus breviter ciliatis; lemma sterile 4 mm
longum, acutum, marginibus ciliatis pilis 2-2.5 mm longis, quam pilis glumae
secundae crassioribus; lemma fertile 3.8-4 mm longum, minute striatum.

Perennial; culms slender, erect, about 50 em tall; sheaths as long as or
longer than the internodes with a small tuft of hairs in the throat and a
pubescent line on the collar, the back glabrous, the margins finely ciliate;
ligule about 0.2 mm long, ciliate; blades 4-6 cm long, 2-2.5 mm wide, the
uppermost culm blades and those on the innovations much reduced, firm,
erect, pungent, involute toward the tip, the lower surface glabrous, becoming
hispid toward the tip, the upper surface glabrous or hispid, the margins
sparsely papillose-hispid; raceme 7 cm long, erect; spikelets appressed; first
glume 3.5 mm long, obtuse, obscurely keeled, not greatly narrowed to-
ward the base, hirsute across the lower part, some of the hairs as much as
4 mm long, sparsely hirsute on the midnerve to the middle, the upper dorsal
portion scabrous; second glume 3 mm long, 3-nerved, the tip rounded, gla-
brous on the back, pilose toward the base, the margins evenly appressed
ciliate nearly to the summit; sterile lemma 4 mm long, acute, the margins
densely ciliate, the hairs 2-2.5 mm long, coarser than those on the second
glume; fruit 3.8 mm long, minutely striate.

Type in the U. 8. National Herbarium, no. 1721241, collected at San
Ignacio, Cerro Piedras, Province of Misiones, Argentina, March 10, 1914,
by Rodriguez (no. 31). This specimen was communicated by L. R. Parodi,
Buenos Aires, Argentina.

M esosetum comatum resembles M. loliiforme (Hochst.) Chase, which is a
stoloniferous plant with smaller appressed pilose spikelets, the sterile lemma
not conspicuously fringed.

Panicum abscissum Swallen, sp. nov.

Perenne; culmi densi caespitosi, 55 cm alti, compressi; vaginae inferiores
carinatae, 3-4 mm latae (e carina ad marginem), truncatae, ad basin cul-
morum congestae, superiores longiores, internodiis breviores, auriculatae;
ligula curtissima; laminae 15-25 cm longae, 1-2 mm latae, conduplicatae,
firmae, curvatae vel flexuosae, infra glabrae supra scaberulae; paniculae
terminales et axillares, 7-15 cm longae, ramis adscendentibus vel appressis,
paucifloris, ad 10 cm longis; pedicelli 1-4 mm longi appressi; spiculae 2.8-3

216 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

mm longae; gluma prima 1.6—2 mm longa, 3-nervis, acuta, carina scabra;
gluma secunda et lemma sterile aequalia, 5-nervia, subobtusa, glabra; fruc-
tus 2.2-2.3 mm longus, 0.5 mm latus, acutus, pallidus, nitens.

Perennial; culms densely cespitose, 50-70 cm tall, compressed; lower
sheaths keeled, somewhat crowded, 3-4 mm wide from keel to margin, one
side of the sheath sometimes 5-8 mm longer than the other, truncate or
extended at the summit into short, very broad, obtuse auricles, the upper
sheaths much longer than the lower but much shorter than the internodes,
the auricles not so prominent; ligule membranaceous, very short; blades
15-25 cm long, 1-2 mm wide, conduplicate, firm, curved or flexuous, gla-
brous on the lower surface, scabrous on the upper; panicles terminal and
axillary from all the nodes, long exserted, 7-15 cm long, the branches ascend-
ing or appressed; spikelets 2.8-3 mm long; first glume 1.6—2 mm long, 3-
nerved, acute, scabrous on the keel; second glume and sterile lemma, equal,

Fig. 4.—Panicum abscissum, spikelet and mouth of sheath, X5.

5-nerved, subobtuse, glabrous; fruit 2.2-2.3 mm long, 0.5 mm wide, acute,
pale, smooth and shining.

Type in the U. 8. National Herbarium, no. 1259952, collected in dry sand
at tourist camp near lake, Sebring, Fla., October 3, 1925, by Paul Weather-
wax.

Panicum abscissum belongs to the Agrostoidea group of Panicum. It is
distinguished from all the other species of the group by the broad sheaths
truncate at the summit, the firm flexuous blades, and the axillary panicles
from all the nodes.

DisTRIBUTION: Sandy or swampy ground, central Florida. De Soto
County: Sebring, Weatherwaz in 1925 (type); Polk County: Winter Haven,
O’ Neill 5207, 5208; Florinda, Pzper in 1917.

Panicum irregulare Swallen, sp. nov.

Perenne; culmi 80 em longi, prostrati, ramosi, radicantes, glabri; vaginae
internodiis breviores vel superiores internodiis longiores, glabrae, in ore
pubescentes; laminae 4.5-6.5 cm longae, 1-1.5 cm latae, ovato-lanceolatae,
acuminatae, glabrae, marginibus albis; paniculae 16-17 cm longae, racemis
15 remotis adscendentibus 1-2.5 cm longis; spiculae biflorae, binae, 2—2.2
mm longae, prima subsessilis, secunda pedicellata; gluma prima 0.7-—1 mm
longa, l-nervis, acuta; gluma secunda et lemma flosculi primi subaequalia, -

May 15, 1940 HERMANN, KEARNEY, AND PEEBLES: ARIZONA PLANTS 217

acuta, 5-nervia, glabra; palea flosculi primi 1.5 mm longa, hyalina; caryopsis
libera; fructus 1.5 mm longus, acutus, lucidus.

Perennial; culms 80 cm long, prostrate spreading, ascending at the ends,
branching, rooting at the lower nodes; sheaths shorter than the internodes
or the upper ones a little longer than the internodes, glabrous, pubescent at
the mouth; blades short pedicellate, 4.5-6.5 cm long, 1—-1.5 em wide, ovate-
lanceolate, acuminate, with white cartilaginous margins; panicles 16-17 cm
long with 15 rather remote ascending racemes 1-2.5 cm long; spikelets
rather densely arranged on one side of the rachis, 2-flowered, paired, 2—2.2
mm long, spreading, the first subsessile, the second pedicellate; first glume
0.7-1 mm long, 1l-nerved, acute; second glume and lemma of the first floret
similar, subequal, acute, 5-nerved; palea of first floret 1.5 mm long, hyaline;
earyopsis free; fruit 1.5 mm long, acute, smooth and shining.

Type in the U. S. National Herbarium, no. 1644874, collected on stony
river bank in the vicinity of El General, Province of San Jose, Costa Rica,
altitude 760 meters, February 1939, by Alexander F. Skutch (no. 4115).

Panicum irregulare is related to P. frondescens Meyer, which has a denser,
shorter panicle of more numerous racemes and larger spikelets 2.6—2.8 mm
long with the lower floret sterile.

BOTAN Y.—Arizona plants: New species, varieties, and combinations.'
F. J. Hermann, T. H. Kearney, and R. H. Persuss, U. 8. Bu-
reau of Plant Industry.

A paper on this subject appeared in this JouRNAL on November 15,
1939 (29: 474-492). A few additional novelties have since come to
light, and they are published here in order to avoid publication of new
names in a flora of Arizona, now in preparation.

Juncus interior Wieg. var. arizonicus (Wieg.) Hermann, comb. nov.
Juncus arizonicus Wieg., Bull. Torrey Bot. Club 27: 517. 1900.

Juncus interior Wieg. var. neomexicanus (Wieg.) Hermann, comb. nov.

Juncus neomexicanus Wieg., Bull. Torrey Bot. Club 30: 447. 1903.

Both J. arizzonicus and J. neomexicanus possess the acuminate to aristate
bracteoles of J. interior, which set this species off from its nearest allies.
Intermediates are frequent between the two varieties here proposed and
typical J. interior, but most material falls rather readily into one of the
three forms and may be distinguished by the following key:

Perianth equaling the capsule, 3-4 mm long................ J. interior

Perianth exceeding the capsule, 4-5 mm long.

Bracteoles lanceolate, acuminate; perianth-segments erect, rigid,
lanceolate, their hyaline margins relatively narrow and opaque....
eR EM ne rs ne te nen al J. interior var. arizonicus

Bracteoles broadly ovate, acute to abruptly aristate ; perianth-segments
spreading, not rigid, broadly ovate, with broad transparent scarious
margins and brown lateral bands bordering the green center... .
5-8 SUS RSs he ee ee No ea ce J. interior var. neomexicanus

1 Received February 1, 1940.

218 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

Juncus longistylis Torr. var. scabratus Hermann, var. nov.

A varietate typica recedit foliis pedicellisque scabratis.

Of the 20 collections of J. longistylis seen from Arizona, 9 represent the
typical form of the species and, except for one specimen from the Huachuca
Mountains (Cochise County), all are from the northern half of the State.
The others have the vegetative parts scabrate much as in the eastern J.
caesariensis Coville, and the auricles show a tendency to be prolonged, free,
and acute. The scabrosity is most conspicuous upon the pedicels and the
terminal portions and margins of the leaves. Intergradations with the typical
form are found in various degrees, however. The following collections in the
U.S. National Herbarium are characteristic of var. scabratus: Near Prescott,
alt. 5,350 feet, Yavapai County, Peebles, Harrison, & Kearney 2712, August
8, 1926 (TYPE): Willow Springs, Apache, County E. Palmer 556 (in part)
and 624 Gn part), July 1890.

Juncus tracyi Rydb. f. utahensis (Martin) Hermann, comb. nov.

J. utahensis Martin, Rhodora 40: 69-71. 19388.

Juncus tracyi manifests two forms, a few-headed, many-flowered phase,
characterized by the type specimen of Rydberg’s species, and a many-
headed, few-flowered phase, represented by the type specimen of J. utah-
ensis. The extremes of these forms are strikingly different in appearance
yet intermediates are plentiful and there is no apparent geographic segrega-
tion of the extremes. Juncus ensifolius Wikstr., J. phaeocephalus Engelm..,
J. saximontanus A. Nels., J. canadensis J. Gay, and J. acwminatus Michx.
are other species that exhibit a parallel series of forms.

Juncus saximontanus A. Nels. f. brunnescens (Rydb.) Hermann,
comb. nov.

Juncus brunnescens Rydb., Bull. Torrey Bot. Club 31: 400. 1904.

This form shows the same instability as does J. tracyz f. utahensis and
like it seems to have no geographic significance. The extremes may be dis-
tinguished by the following key:

Inflorescence composed of few (seldom more than 10) heads, which aver-

age 7-10 mm in diameter and are many (15 to 25)-flowered........
PSR estas. ae 2, A Nik Seah cut arecanimee oe te, T SeaN Re BO a ee J. saximontanus
Inflorescence composed of numerous (usually more than 10) heads, which
average 5-6 mm in diameter and are few (5 to 12)-flowered........
ee ER MRE tie OR Sal te) EO cee J. saximontanus f. brunnescens

Astragalus cobrensis Gray var. maguirei Kearney, var. nov.

Ab A. cobrensi f. typica pilis longioribus patentibus, foliolis supra parce
pubescentibus, calycis dentibus quam tubo longioribus, recedit.

Type from White Tail Canyon, Chiricahua Mountains, Cochise County,
Ariz., in sandy soil under oak and walnut, Bassett Maguire et al. 11079,
May 2, 1935 (U. S. National Herbarium no. 1768940).

The variety here described has rather long spreading hairs on the herbage
and pods, leaflets sparsely pubescent above, and calyx with teeth longer
than the tube, whereas in typical A. cobrensis the hairs are shorter and
appressed, the leaflets glabrous above, and the calyx-teeth about as long
as the tube. The flowering stems in var. maguiret (probably also in A. co-
brensis) arise from long, slender, creeping rootstocks.

May 15, 1940 THORNE: HAIRWORM AS PARASITE OF MORMON CRICKET 219

Astragalus cobrensis, known only from southwestern New Mexico, is
sparingly represented in herbaria, and var. maguirez is known only by the
type collection. Study of additional material of these forms may prove
maguirei to be a distinct species.

Echinocereus robustus Peebles, sp. nov.

Echinocereus rectispinus Peebles var. robustus Peebles, Amer. Journ. Bot.
252015. 1938.

ZOOLOGY .—The hairworm, Gordius robustus Lezdy, as a parasite of
the Mormon cricket, Anabrus simplex Haldeman.1 GERALD
THORNE, U.S. Bureau of Plant Industry.

INTRODUCTION

Parasites effective against the Mormon cricket, Anabrus simplex
Haldeman, apparently are very rare. It was, therefore, of considera-
ble interest when there were reports of a heavy parasitism by Gordius
robustus Leidy in the vicinity of Arrowrock Dam, Idaho (1). Visits to
that locality were made by the writer on July 26 and October 9, 1935,
and on May 16 and August 24, 1936. The unusual populations oc-
curring there presented excellent opportunities for studying the para-
site from the standpoint of life history, habits, host relationship,
ecology, and economic importance. Specimens for more detailed ex-
amination and experiments were taken to the Salt Lake City, Utah,
laboratory.

This observation of Gordius robustus coincided with the unprece-
dented populations of Anabrus simplex which first appeared at Fort
Hall, Idaho, in May, 1932, and by August, 1936, had infested almost
2,000,000 acres in 24 counties of Idaho (11). Similar outbreaks of the
cricket occurred in other western States (Fig. 1).

HISTORICAL

Gordius robustus Leidy, 1851, belongs to the rather common group of or-
ganisms known as “hairworms,” the adults of which are usually found in-
habiting warm shallow pools or streams. In its immature stages it is known
as a parasite of certain insects, principally Orthoptera.

The first record of Gordius robustus in Anabrus simplex (synonym A. pur-

’ The writer is indebted to Claude Wakeland and R. W. Haegele, who gave valued
preliminary information on the distribution of Gordius robustus in the Arrowrock sec-
tion and joined in visiting the area on two occasions. J. Perey Moore, of the University
of Pennsylvania, kindly lent the Leidy collection of G. robustus from Milk River, Mont.,
and Dr. Henry B. Ward forwarded the May collection from Urbana, IIL., for com-
parison with the Idaho and western Montana specimens collected in 1935. G. Steiner
and Edna M. Buhrer have given valuable suggestions on the manuscript and bibliog-
raphy. The map used in Fig. 1, giving the 1937 distribution of Anabrus simplez, is taken
from a U. S. Department of Agriculture Press Service release, March 26, 1937. Re-
ceived November 18, 1939.

220 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

purascens’) is given by Riley, Packard, and Thomas (9, footnote 83, p. 327),
who mention “Gordius robustus, Leidy.—A o& from 9 Anabrus purpurascens
Uhler; in Mr. Riley’s cabinet.’’ No records remain as to the locality in which
this specimen was collected. Riley, Packard, and Thomas (10, p. 64) quote
Bruner’s remarks on parasites of the Mormon crickets observed along the
Portneuf River, Idaho: ‘‘Hair-worms (Gordius), too, are frequently found
wound about their intestines.”

MMMME HEAVY INFESTATIONS
T1110 \\GHT INFESTATIONS

Fig. 1.—Range of Anabrus simplex in 1937 and known distribution of Gordius
robustus: 1, Cutbank, Mont.; 2, Augusta, Mont.; 3, Arrowrock Reservoir, Idaho.

Leidy (3) reported Gordius robustus from New Jersey as a parasite of a
grasshopper, later (5) identified as Conocephalus fasciatus fasciatus (DeGeer)
(synonym Orchelimum gracile Harris), and (4) of the carabid beetle Ptero-
stichus (Gastrellarius) honestus Say (synonym Pterostichus fastiditus De}.).
Thomas (9) found a female in Stenopalmatus fuscus Haldeman (synonym S.
fasciatus Thomas) at Wasatch, Utah. May (6) reports it in the vicinity of
Urbana, Ill., as a frequent parasite of Orchelimum vulgare Harris, O. nigripes
Scudder, Conocephalus memoralis (Scudder) (synonym Xiphidium nemorale
Scudder), and occasionally Phaneroptera furcata furcata (Brunner) (syno-
nym Scudderia furcata Brunner).

2 Synonyms given are the host names of the original records.

May 15, 1940 THORNE: HAIRWORM AS PARASITE OF MORMON CRICKET 221

THE IDENTITY OF GORDIUS ROBUSTUS LEIDY, 1851

The type specimen of Gordius robustus from Concocephalus fasciatus fas-
ciatus has not been preserved, but a collection from Milk River, Mont.,
made in 1876 and identified by Leidy, is still available. This consists of four
males and six females varying from 31 to 46 cm in length. The hosts of these
specimens are unknown, but they were collected within the range of Anabrus
simplex and, judged from their size, it appears quite probable that they grew
in these large insects. These 10 specimens are identical to those recently col-
lected in Idaho and Montana except for their lighter color, which is due to
bleaching during their long preservation in alcohol.

Leidy’s original designation of the name Gordius robustus was given with
a very meager description (3). Later (4, p. 57) he published a more detailed
description but listed the species as a synonym of G. aquaticus? Gmelin
(which is G. aquaticus Linn., 1758). He followed the name with a question
mark, indicating that his specific designation was questionable. Montgomery
(7) placed the species as a variety, G. aquaticus robustus; but later (8) made
it a synonym of G. villoti Rosa, 1882. May (6) considered G. robustus as a val-
id species and prepared an excellent, detailed morphological description.

After examining the specimens from Milk River, Mont., the writer con-
siders that May’s action was fully justified. To further establish the fact,
specimens from the Arrowrock section were sent to Kurt Heinze, a leading
European authority on Gordioidea, who pronounced them as being neither
Gordius aquaticus nor G. vllotz.2 Caballero (2) records both G. robustus and
G. aquaticus from Mexico, thus supplying additional evidence that the two
species are distinct.

Since Leidy determined the specimens from Milk River, Mont., as being
the same as his type specimen of Gordizus robustus from New Jersey and also
in view of the fact that May found the species prevalent in Illinois, the writer
has no basis for assigning the form to any other species.

DISTRIBUTION OF GORDIUS ROBUSTUS AS A PARASITE OF ANABRUS SIMPLEX

A survey of the Arrowrock section on July 26, 1935 (Fig. 2), revealed that
there was a great difference in the number of Gordius present in the various
localities. They were most numerous on Elk Creek where tangled masses
(Fig. 3) were present in almost every pool, and many more were scattered
along the stream or buried in the loose gravelly soil or under rubbish. At
Grouse Creek and near Long Gulch Ranger Station they were almost as
numerous, while small numbers were present in Slide Gulch, Cottonwood,
Highland Valley, and other points along the route.

After the first reports of the parasitism of Mormon crickets by a Gordius,
workers in the various cricket infested areas of the Western States were on
the alert for additional localities in which the hairworm might be present.

3 Personal communication dated February 20, 1937.

VOL. 30, No. 5

JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

222

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May 15, 1940 THORNE: HAIRWORM AS PARASITE OF MORMON CRICKET 223

However, the only places added were in Montana, where Frank Cowan and
Horace J. Shipman, of the U.S. Bureau of Entomology and Plant Quaran-
tine, collected a considerable number near Cutbank and a single female near
Augusta (Fig. 1).

ECONOMIC IMPORTANCE OF GORDIUS ROBUSTUS

Reliable reports indicate that between 25 and 50 percent of the Mormon
crickets invading the Grouse Creek section early in July 1935, were para-

Rs

Fig. 3.—Entangled mass of Gordius robustus collected
on upper Elk Creek.

sitized by Gordius robustus, and this appeared to be corroborated by the
great numbers present in the stream and buried in wet areas. At that time on
Arrowrock Reservoir the crickets entered the water in great numbers, until
patches 2 or 3 square rods in extent sometimes were floating about and from
them large numbers of Gordii emerged and congregated along the shore. By
July 26 it was evident that the larger portion had left their hosts, and it was
difficult to find crickets in which Gordii still remained; between Arrowrock
Dam and Grouse Creek only five parasitized crickets were found among
1,000 examined, and in other localities they were just as rare. However, the

224 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 5

Fig. 4.—Anabrus simplex parasitized by Gordius robustus. Note in the top
figure how thoroughly the body contents of the insect have been absorbed, leav-
ing only the alimentary canal. Natural size.
crickets are migratory, remaining in one locality only a short time, and it is
probable that those observed by the writer were from different breeding areas
than those in the earlier invasions.
Commenting on the parasitized crickets collected on September 26, 1935, |

May 15, 1940 THORNE: HAIRWORM AS PARASITE OF MORMON CRICKET 225

near Cutbank, Mont., Horace J. Shipman wrote, ‘The parasitized crickets
were found near a pond, which was the only one in the vicinity. Crickets
taken within 100 yards of the pond were approximately 99 percent para-
sitized, and only one of the crickets contained eggs. One mile north of this
pond the crickets were about 50 percent parasitized.’’4

On July 20, 1936, Claude Wakeland' reported that about one-fourth of the
crickets in the Slide Gulch and Rattlesnake localities near Arrowrock Reser-
voir were parasitized.

None of the parasitized crickets collected by the writer contained eggs.
The body cavity was invariably found to be completely filled with the para-
site, and only the alimentary canal remained (Fig. 4). The parasitized female
containing a few eggs reported above by Shipman may have been a case of
very late parasitism after the ovaries had been developed.

It is evident that female crickets parasitized by Gordius robustus rarely
produce eggs; therefore, in regions of heavy parasitism the cricket popula-
tion must be considerably reduced. Since only a minor portion of the breed-
ing grounds of the Mormon cricket lies near streams, ponds, and swamps
suitable for the propagation of Gordius, the effective control exercised over
the total cricket population during extensive outbreaks probably is negligi-
ble. The migratory habits of the cricket bring great numbers to streams
where they may become parasitized but at the same time a large portion of
these individuals wander far from water and die, the parasites being lost be-
cause water is essential to completion of their life cycle.

During the many years between extensive cricket outbreaks only small
bands occur, and these generally are confined to the higher elevations where
there is more abundant moisture. In those regions where Gordius is abundant
the water-frequenting habits of the crickets no doubt bring practically all of
them in contact with Gordius, and during such periods it seems quite prob-
able that economically more effective parasitism occurs.

MORPHOLOGY OF GORDIUS ROBUSTUS

Adults.—But little can be added to May’s description of the adults (6).
The specimens observed all correspond closely with his observations except
in minor points, which apparently are due to individual variation. Very
light-colored specimens, especially females, apparently lack the dark dorsal
and ventral bands and light-colored areas of the cuticle that May men-
tions, but they show distinctly on all the darker-colored females and males.
These markings were also absent on the bleached, alcohol-preserved speci-
mens from the Leidy collection. The branching hairs of the cuticle men-
tioned by Montgomery (7) could not be found, and since May also failed to
observe them this statement must have been an error. Considerable varia-
tion in the male postcloacal ridge occurred among the specimens collected,

4 Personal communication dated December 9, 1935.
> Personal communication dated June 20, 1936.

226 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 5

the most common form being slightly angular (Fig. 5, A), while a more an-
gular type (Fig. 5, B) was comparatively rare.

The oral opening is degenerated to a minute scar (Fig. 5, C), which is visi-
ble only in a face view. In no instance was it observed that the larval spear
remained within the oral aperture.

A brownish ring generally surrounds the cloacal opening of the female, but
on occasional specimens it is entirely lacking. When present it varies con-
siderably in its distinctness and width.

Fig. 5.—A, B, Male tails showing variations in the form of the cloacal ridge, «25.
C, Rudimentary oral opening, X750. D, Larva, X750. E, Head of larva with pro-
truded proboscis, X750. FF, Posterior portion of larva with blunt terminus, x 750.
G, Sperms, 1500.

Specimens of Gordius robustus from Anabrus stmplex frequently are very
large. Wakeland and Shull (11) state that they reach a length of 48 inches
(121.9 em), and the writer collected a female 120.6 cm long. A random collec-
tion of 200 specimens contained 63 females varying in length from 13 cm to
120 cm and 137 males from 18 cm to 91 cm.

Eggs.—The slightly ovate eggs average about 38u wide by 51ly long and
are deposited in slender broken cords held together by a gelatinous mucus,

May 15, 1940 THORNE: HAIRWORM AS PARASITE OF MORMON CRICKET 227

which swells on contact with water until the egg cord is about 1 mm in di-
ameter (Fig.6, A). The ovary occupies almost the entire body cavity, and the
number of eggs produced by a female is enormous, those from a 106-cm fe-
male being estimated at 28,902,500. There is a short uterus, 2 to 3 mm long,
in which huge numbers of minute spermatozoa (Fig. 4, G) can be found in
fertilized females. The body of the spermatozoon appears to be somewhat
flattened and concave, probably giving it a rotary motion as it is propelled
by its tail. Fertilization of the eggs occurs as they pass through the uterus.

Fig. 6.—A, Egg cord after deposition, X1.
B, Female hibernating among roots, X1.

Eggs are regularly deposited in spring, but rare females occasionally were
observed depositing in fall. Segmentation and larval development may occur
immediately or be delayed for some months.

LIFE CYCLE OF GORDIUS ROBUSTUS IN THE ARROWROCK SECTION

Overwintering females emerge from their hibernating quarters in the
loose gravelly soil or under sticks, stones, and rubbish (Fig. 6, B) during
April and May and enter the shallow water of swamps, pools, or streams
where the eggs are deposited. Many of the eggs hatch soon after deposition,

228 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 5

while others may remain for several months without hatching. Some of these
taken to Salt Lake City contained living larvae until September, and it is
doubtful if they would have hatched that fall.

The minute larvae move slowly about in the water, frequently lifting them-
selves up on the tips of their tails and swinging about in circles. Here they
apparently are picked up by the young crickets when drinking, or when eat-
ing aquatic plants, and pass to the intestine. Presumably they force their
way through the intestinal wall into the body cavity with the toothed pro-
boscis while they cling to the intestinal tissues by means of the three rows of
cephalic hooks.

After entering the body cavity growth takes place so rapidly that by late
in June many are full grown and ready to emerge from the crickets. Appar-
ently the time of leaving the host depends upon some sense of proximity to
water, for during the heat of the day, when crickets congregate in the shade
along streams or moist areas, the mature Gordius break through the abdom-
inal wall and emerge, moving into the water or burying themselves in the
moist soil.

At the time of emergence from the host the color of the females varies
from a milky white to light brown, and that of the males from light to dark
brown. After exposure to light the color deepens, especially in the males,
which are usually darker than the females, sometimes almost black.

Copulation takes place in the water, perhaps also in the soil, soon after
emergence. Females collected July 26 usually contained sperms in the uterus.

Larvae.—The larvae (Fig. 5, D) range in length from 0.12 to 0.16 mm and
average about one-ninth as wide as long. The flexible cuticle is marked by
annules that vary in distinctness and width as the individual moves about.
The broad anterior end bears three circlets of cephalic hooks, the second row
is arranged directly behind the first, while the third row alternates, the hooks
being back of the interspaces of the first two rows. When at rest these hooks
are retracted into the oral opening until the last row points almost straight
forward. As the proboscis is extruded the hooks roll out and back (Fig. 5, E).

The slightly clavate proboscis bears two or three very minute projections
on its distal end. Back of these, on the anterior conoid portion, are about six
rows of fine corrugations. The proboscis joins the tubular esophagus, their
combined length varying from one-eighth to two-fifths that of the entire
body.

The larvae generally have a pointed tail, but occasional specimens had
blunt, rounded tails (Fig. 5, F).

The larvae are soft bodied and unable to withstand drying. Exposure to
air on a bamboo splinter for one minute was fatal in every case.

The manner in which the larvae of Gordius robustus enter Anabrus semplex
was determined by the following method: A supply of crickets was secured
near Tooele, Utah, where G. robustus is not known to occur. Lettuce was fed
to the crickets, and as soon as one was busily feeding a drop of water con-

May 15, 1940 THORNE: HAIRWORM AS PARASITE OF MORMON CRICKET 229

taining several larvae of G. robustus was placed just in front of it. Generally
the cricket continued feeding and swallowed the larvae along with the food.

Thirty crickets were fed in this manner on June 15 to 18 and placed in a
cage. A few crickets were removed each week thereafter, three to five at a
time, and dissected. On July 1, one female contained a male Gordius 260 mm
long and on July 9 another female was found with a male Gordius 245 mm
long. Of the 30 crickets, 18 died and 12 were dissected. Only the two men-
tioned were parasitized.

At the same time 23 crickets were selected, and a drop of water containing
several larval Gordius was placed in the folds between the sclerites of the
abdomen. Of these 23 crickets, 9 died and the remainder were dissected,
none being found infested. Because of their sensitiveness to drying it would
appear impossible for the larvae to enter the body of the host in any manner
other than through the mouth.

Unfortunately the season was so far advanced that very young crickets
could not be secured, and those used in the above experiments were all in the
second and third instars. In these large crickets the larval Gordius apparently
are carried through the intestinal tract before they have an opportunity to
penetrate the intestinal wall. This appeared to be proved by the fact that
among 50 adult crickets confined in wet swampy pens, in which thousands
of Gordius eggs and larvae were kept in excellent condition, not one became
parasitized. An abundance of small aquatic plants was present and these,
together with the water consumed, certainly would appear to have given
every cricket ample opportunity to become parasitized.

ECOLOGICAL FACTORS

In the postparasitic stages Gordius robustus thrives best in warm shallow
pools with loose gravelly banks well covered with vegetation. These condi-
tions enable them to congregate during the mating season and later work
their way into the loose soil, under stones and sticks, or into masses of roots
or leaves, where they hibernate during winter. Seep areas where warm
ground water rises during winter are especially favorable, and in the Arrow-
rock Dam section these areas were generally inhabited by large numbers.
In the years preceding these investigations such conditions had prevailed

in the Arrowrock section. There had been no unusual runoff and the stream
beds were littered with sticks, pieces of logs, and loose stones, making ideal
breeding conditions for Gordius robustus (Fig. 7, A).

The months of January and February, 1936, were unusually cold, each
with average mean temperatures of 23.4° F., 1.7° and 7.4° respectively below
the normal at the Arrowrock Reservoir station. This temperature was ac-
companied by a heavy snowfall with a water content of 10.14 inches, 4.87
inches above the mean for these two months. As a result the loosely packed
snow melted rapidly during March and especially during April. Streams
flooded over their banks, channels were widened and deepened, and all

230 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

Fig. 7.—A, Upper Elk Creek, Idaho, July 26, 1935.
B, Same location, May 16, 1936.

debris was removed (Fig. 7, B). Most of the hairworms that had hibernated
in the stream channels and adjacent banks were carried away by these
floods. Scores of them could be found hanging high on limbs and roots
where they had been left as the waters receded. As a result the Gordzus
population in the Elk Creek and neighboring basins was estimated to be

May 15, 1940 THORNE: HAIRWORM AS PARASITE OF MORMON CRICKET 231

only 5 to 10 percent as great as in the preceding year. Many years will be
required to repair the erosion damage to these stream beds and return them
to their former conditions, which were so favorable as breeding grounds for
Gordius.

SUMMARY

The hairworm Gordius robustus was first observed as an important para-
site of the Mormon cricket, Anabrus simplex, in 1935 in the vicinity of Arrow-
rock Reservoir, Idaho. In certain localities of this area there are favorable
breeding grounds for the hairworm along small mountain streams and in
pools or swamps. In these localities they may be of economic importance in
the control of ordinary populations of Mormon crickets. However, during
heavy outbreaks of crickets, such as occurred in 1935 to 1937, the percentage
of parasitism was too small to be of economic importance, although in some
instances it reached 25 to 50 percent of the crickets. Similar parasitism was
also observed near Augusta and Cutbank, Mont., where in certain small
areas 50 to 99 percent of the crickets contained Gordii.

Parasitized crickets rarely produce eggs. They live normally until the
mature Gordius breaks through the abdominal wall and enters water or wet
soil. Here the Gordii mate and later hibernate in moist places under sticks
and stones or among roots and rubbish. Some eggs are deposited in fall but
more generally in spring, when the females emerge from hibernation and
deposit them along the shores of streams, ponds, or swampy areas. Here the
minute larvae hatch and probably are picked up by the young crickets when
drinking water or eating aquatic vegetation. Upon reaching the intestine of
the cricket they apparently bore through the wall and enter the body cavity,
remaining there until maturity.

Under present erosion conditions in breeding areas of the Gordius most
of them are carried away by floods.

LITERATURE CITED

1. Anonymous. Long worm parasites bring death to crickets, scientists discover. Salt
Lake Tribune 131(84-B2). July 7, 1935.
2. CaBALLERO, E. Contribucion al conocimiento de los Gordius de Mexico. An. Inst.
Biol. Univ. Nac. Mexico 7: 477-488, illus. (Cover page dated 1937, title page
dated 1936.)
Leipy, J. On the Gordiaceae. Proc. Acad. Nat. Sci. Philadelphia 5: 275. 1851.
. A synopsis of Entozoa and some of their ecto-congeners observed by the author.
Proc. Acad. Nat. Sci. Philadelphia 8: 42-58. 1856.
. The Gordius, or hair-worm. Amer. Ent. and Bot. 2: 193-197. 1870.
May, H.G. Contributions to the life histories of Gordius robustus Letdy and Para-
gordius varius (/ ezdy). Illinois Bicl. Monog. 5(2): 120-240, illus. 1919.
Monteomery, T. H., Jr. The Gordiacea of certain American collections with par-
ticular reference to the North American fauna I. Bull. Mus. Comp. Zool. 32:
23-59, illus. 1898.
. The distribution of the North American Gordiacea, with description of a new
species. Proc. Acad. Nat. Sci. Philadelphia 59: 270-272. 1907.

9. Ritey, C..W., Packarp, A.5S., Jk., and THomas, C. Hair-worms. U.S. Ent.
Comm. Ann. Rept. (1877) 1: 326-334, illus. 1878.

10. . The western cricket. U.S. Ent. Comm. Rept. 3: 61-64. 1883.

11. WaxkELAND, C., and SHuutt, W. E. The Mormon cricket with suggestions for its
control. Univ. Idaho Agr. Coll. Ext. Bull. 100, 30 pp., illus. 1936.

Oo wow

232 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 5

SCIENTIFIC NOTES AND NEWS
CONFERENCE ON A NEW FLORA OF THE DISTRICT OF COLUMBIA

A conference on a new flora of the District of Columbia (to replace the
one published in 1919) was held at George Washington University, Febru-
ary 20, 1940.

An Executive Committee was appointed to administer the venture. The
chairman of this committee is E. H. Waker, of the U. 8S. National Her-
barium, and the other members are C. R. Batu, Extension Service, U. S.
Department of Agriculture; AGNErs CHaAsE, Smithsonian Institution;
Rosert F. Griaes, George Washington University; Nei Horcuxiss,
U.S. Biological Survey; and Huexu O’ Nei, Catholic University.

The conference discussed the area to be included in the new flora, and
though no final decision was taken it was the consensus that the limits be
extended beyond those established for the current flora.. It will be several
years before the new flora goes to press, and the intervening period will be
utilized in collecting. The George Washington University has provided
space, with tables and storage cases, in which several botanists have offered
to deposit their local collection.

All those desirous of participating in this enterprise should communicate
with EK. H. WALKER.

Obituary

FREDERICK GALE TRYON, economist, National Bituminous Coal Com-
mission, died on February 15, 1940. Mr. Tryon was born in Minneapolis,
Minn., in 1892. He was graduated from the University of Minnesota with an
A.B. in 1914, and in 1916 he received his A.M. degree from that institution.
Later he did additional graduate work at Johns Hopkins University. In 1916
and 1917 he was employed as an assistant geologist with the Empire Gas
& Fuel Co. During the Great War he was commissioned as a captain and
assigned to the statistics branch of the General Staff of the Army in Wash-
ington and with G.H.Q. in France, and later was American secretary for the
raw materials section, Supreme Economic Council, American Peace Com-
mission, Paris, 1919. In 1920 to 1925 he served as a mineral geographer in
charge of coal and coke statistics for the U. 8. Geological Survey. During a
part of this time, from 1922 to 1923, he further acted as statistical adviser
to the U. 8S. Coal Commission. From 1925 to 1937 he was connected with the
U. 8. Bureau of Mines in fuel industries, during 1936 and 1937 as principal
economist. In 1937 he was appointed economist in the Bureau of Research
and Statistics, National Bituminous Coal Commission.

Mr. Tryon served on the staff of various institutions including the Uni-
versity of Pennsylvania, Brookings Institution, American University, and
the Institute of Economics (Washington, D. C.). He was appointed on the
President’s Research Committee to study social trends in 1932-1933, and
in 1934-1935 to study population redistribution.

In addition to the Washington Academy of Sciences Mr. Tryon was a
member of the Economics Association, the American Institute of Mining and
Metallurgical Engineers, the Mining and Metallurgical Society, the Geologi-
cal Society of Washington, and the American Association for the Advance-
ment of Sciences and was a fellow of the Statistical Association.

CONTENTS

Botany.—The botany and history of Zizania aquatica L. (‘‘wild
rice’). CHARLES HE. CHAMBLISS... Oe. a

Pagnonconcus —Paralbula, a new fossil fish based on dental plates
from the Eocene and Miocene of Maryland. 8. F. BuakE.......

Botany.—Miscellaneous new American grasses.. Jason R. SWALLEN —

Botany.—Arizona plants: New species, varieties, and combinations.
F. J. Hermann, T. H. Kearnny, and R. H. PHEBLES........... ‘

ZooLocy.—The hairworm, Gordzus robustus Leidy, as a parasite of the
Mormon cricket, Anabrus simplex Haldeman. GERALD THORNE

Screnriric NovEs AND-NEWS. 2). 4600200 0

OBITUARY: FREDERICK GALE. TRYON. 200.0 2. 45 eee

This Journal is Indexed in the International Index to Periodicals

Vou. 30 | JUNE 15, 1940 No. 6

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JOURNAL

OF THE
WASHINGTON ACADEMY OF SCIENCES
Vou. 30 JUNE 15, 1940 No. 6

CHEMISTRY.—A study of the association of magnesia with silica in a
pure magnesium clay.! P. G. Nuttrine, U. 8. Geological Sur-
vey.

A white clay occurring near Hector, Calif., described by Foshag
and Woodford (1) was shown to be a nearly pure magnesium silicate
of the montmorillonite type. It swells largely and disperses readily in
water like the familiar Wyoming bentonites. Although composed al-
most entirely of a crystalline mineral, it is remarkable in that its
magnesium may be completely removed (leaving pure silica) by elec-
trodialysis. In view of this ready separation by electrical means, it
seemed desirable to make solution analyses of this clay by the method
previously described by the author. This method consists essentially
of bringing the clay to equilibrium with various hot dilute solutions of
acid or alkali and analyzing the resulting solution (2, 3). To clarify
the interpretation of these analyses, pH determinations were made
on each solution. Some rather remarkable results appear worth put-
ting on record.

The magnesium is not removable by electrodialysis from a pure
montmorillonite to any observable extent. From the Wyoming ben-
tonites only the alkalies are thus removable. The swelling property of
the Hector clay is doubtless due to its being saturated with alkali to
base-exchange capacity, a property that, as shown by Mattson (4),
is true for clays and soils in general. Alkali in the Hector clay amounts
to 4.19 percent, of which 3.00 percent is Na,O and 1.12 percent is
Li.O (1). Magnesium can not be exchanged for lithium or the lithium
for magnesium, so that the exchangeable base is the 3.00 percent of
Na,O. The 1H,0 of water of crystallization is very firmly held, being
released only above 650° C. (nearly 100° higher than in pure mont-
morillonite) and is completely removed only by heating to 1,000° C.
Boiling with ammonium oxalate separates magnesium from silica as
do of course the mineral acids. Sodium carbonate solutions remove
the silica from the magnesium.

_1 Published by permission of the Director of the U. S. Geological Survey. Re-
ceived January 14, 1940.

233

234 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

The white Hector clay forms a permanent hydrogel in water, but
the light-brown clay associated with it does not and may be readily
separated by settling and decantation. The stock suspension used
contained 8 grams per liter. To 100-ce portions of this stock in pyrex
flasks was added hydrochloric acid to make concentrations ranging
from 0.01 to 4.00 percent, as shown in Table 1. The flasks were kept
at 90°-92° C. for 24 hours, the clay filtered out with Whatman 42
filter paper, the filtrate evaporated, dried at 160° C., and weighed in
pyrex beakers. The residue consisted partly of water-soluble chlorides.
These were removed by boiling in water. The water-insoluble residue,
after drying and weighing, was treated for 20 minutes with concen-
trated hydrochloric acid to separate bases and silica, and these were
determined. Calcium, iron, and aluminum were negligibly small.
Magnesium was determined as pyrophosphate. The pH determina-
tions were made with a glass electrode apparatus on portions of the
filtrate carefully brought to room temperature (29°+1°). The inter-
mediate range of concentrations (0.063 to 0.63 percent) were repeated
six times since in this range equilibrium is extremely sensitive to op-
erating conditions.

The more significant results are given in the following table and are
graphed in the figure.

TABLE 1.—SoOLUBILITY OF HECTOR CLAy IN HCl SoLutTIons AND pH oF
VARIOUS CLAY SOLUTIONS

In solution: Filtrate: pH ,
Initia Goaias Donner Initial acid: m.e.q. HCl
peat MgO | MgO me . 2) cea
ercen SiO» Teseinbist Soluble! Hector Paria |Wyoming
0.0 0.082 | 0.00 0.00 | (0.01) 2.70 8.65 OY 4.10 0.274
0.016 0.010 | 0.00 0.00 2.49 Coes} 4.52 3.39 0.436
0.025 0.048 | 0.032 0.00 2.28 7.00 3.40 Phe 1D 0.690
0.040 0.121 | 0.099 0.00 2.09 6.20 76 ll Da PAGE Wolo)
0.063 O82 250 OlG2Z (0) 5 EA 1.88 5.40 2.08 1.90 We 0
0.100 0.276 | 0.190 0.32 1.70 5.60 1.80 1.63 24
OR59 0.246 | 0.168 0.62 1.49 4.48 1.39 1.40 4.36
(0) PABA 0.182 | 0.140 0.98 1.30 4.38 1.10 ileal 6.90
0.40 0.4381 | 0.295 1.39 5 Bo PAPA 0.97 1.04 | 10.96
0.63 0.495 | 0.302 as 0.95 1.39 0.88 O2L9o2 Tw eivers
1.00 0.508 | 0.282 1.93 0.78 1.01 0.80 OPEL B74
oo 0.496 | 0.252 1.98 Fresh 9.90 9.52 9.55 | (In H2O
72, O74 0.473 | 0.216 2.00 Cooked — 9.10 — only)
4.00 0.439 | 0.175 2.00 a= = = = =

1 After heating at 160° C.

The solubility of the Hector clay is of the same general character as
that previously found (3) for the familiar Wyoming swelling ben-
tonite. The reversal in slope in the concentration range of 0.1 to 0.25 —

JUNE 15, 1940 NUTTING: STUDY OF A MAGNESIUM CLAY 239

Si02 and MgO in Filtrate g/|

H-Hector clay

P-Paria swelling clay
W-wyoming swelling clay
V-Polkville Vicksburg
D-N.Dakota acid clay

pH of Filtrate

Fig. 1.—Solubilities and pH values for Hector clay.

percent acid occurs also in the curves for Wyoming bentonite in the
range of 0.4 to 1.0 percent—four times the concentration. This re-
versal has been found in the nonswelling adsorbent clays low in so-
dium (2), but it can not be fully explained.

In pure water silica appears to dissolve as sodium silicate and the

236 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

solubility (0.082 gm/liter) is higher than in dilute acid. It is nearly or
- quite insoluble in 0.01 percent acid but is rising rapidly at 0.04 per-
cent reaching a first maximum at 0.1 percent. Magnesium, in a form
insoluble in water (after evaporation to 160° C.), appears with about
0.016 percent acid and closely parallels the silica curve, the ratio of
MgO to SiO, remaining about 0.7 throughout the entire middle range.
In the clay that ratio is 0.5. Mg as water-soluble chloride appears in
the filtrate residue at about 0.04 percent acid and steadily increases
to a constant amount (0.2 gm/liter) above 1 percent acid, owing to
the limited amount of clay present.

The pH curve of the equilibrium clay-acid filtrates descends from
the initial value 9.9 for clay in pure water in a linear relation from 0.01
to 0.1 percent acid, then remains nearly constant from 0.1 to 0.4 per-
cent, drops suddenly to the normal value, and continues just above
the curve for pure acid (dashed line). The slope on the 0.1 to 0.4 per-
cent shoulder is less than that for the acid solution alone; trebling the
acid content lowers the pH only from 4.6 to 4.4. Such an effect might
be produced by adsorption, as pH is a measure of the hydrogen ion
activity, but the total magnesium going into solution is steadily in-
creasing in that range. Living bodies maintain a similar constant pH
regardless of the intake, possibly for a similar reason.

The pH curve for the Wyoming swelling bentonite lies very much
lower than that for the Hector clay and exhibits a barely perceptible
shoulder in the critical region. Three other swelling bentonites from
Ontario, Oreg., Como, Colo., and Paria Valley, Utah, were inves-
tigated. The Paria clay runs higher than the Wyoming, the others are
lower. For comparison pH runs were made on a Vicksburg bentonite,
a very pure montmorillonite. The curve for this was almost identical
with that for the acid solution, despite the fact that it is attacked by
acid as freely as the Hector clay (2). Florida fuller’s earth gave a pH
curve almost identical with that for the Wyoming bentonite. An acid
clay from northeastern North Dakota and a field soil gave curves
slightly lower than that for straight acid. Not one of these clays, ex-
cept the Hector, showed more than a slight irregularity in its pH
curve where the Hector clay has a pronounced shoulder. At higher
acid concentrations, the Paria clay curve dips well under the curve
for pure acid as though an excess of silicic acid were released.

Summary.—The white swelling Hector clay breaks down in hot
acid solutions like the swelling Wyoming bentonite but at markedly
lower acid concentrations. The pH curve of the filtrates from these
suspensions not only runs much higher than that of any of the seven —

JUNE 15, 1940 MORAN, BRIESE, COUCH: NEW CYANOGENETIC PLANTS 237

other clays tested but exhibits a pronounced shoulder in an inter-
mediate acid range not shown by the others. This shoulder indicates
that hydrogen ions are inactivated either by physical adsoprtion or
by chemical reaction, or that hydroxy] ions are released largely over

a particular range.
LITERATURE CITED

1. Fosuac, W. F., and Wooprorp, A. O. Bentonitic magnesian clay mineral from

California. Amer. Min. 21: 238-244. 1930.

. Nurrine, P. G. A study of clay sclubshty. Franklin Inst. Journ. 224: 339-362.

1937.

The action of scme aqueous solutions on clays of the montmorillonite group.

U.S. Geol. Surv. Prof. Paper. (Not yet published.)

. Mattson, Sante. The law of soil colloidal behavior: XIII. Forms and functions of
water. Soil Science 33: 301-822. 1932.

PHYTOCHEMISTRY.—Some new cyanogenetic plants! E. A.
Moran, R. R. Briss, and J. F. Coucn, U. 8. Bureau of Animal
Industry.

This paper reports the detection of hydrocyanic acid or of com-
pounds capable of developing it in six species of plants not hitherto
known to be cyanogenetic together with some figures for the HCN
content of Triglochin palustris.

Emplectocladus fasciculatus Torr.—California desert almond, one
of the Amygdalaceae, is found in California, southern Nevada, south-
ern Utah, and Arizona, according to Tidestrom.’ It 1s a small spines-
cent shrub closely related to the well-known cyanogenetic genus
Prunus. It has not hitherto been reported as being cyanogenetic.
Material was collected in 1936 in Wah Wah Valley, 30 miles west of
Milford, Utah. This was dried and shipped to Washington. Analysis
of the leaves showed a high HCN content of 172 mg per 100 g. Leaves
collected on June 19, 1937, at 2 p.m. in the same locality were pre-
served in 1 percent mercuric chloride solution. Analysis of these
samples at 4, 8, and 12 weeks after collection gave 132, 135, and 134
mg HCN per 100 g green. Neither acetone nor benzaldehyde could be
detected in the distillates from these samples, and no clue was ob-
tained as to the nature of the cyanogenetic glucoside.

Linum king 8. Wats.—A perennial yellow-flowered flax found in
Wyoming and Utah, according to Tidestrom, belongs to a recognized
cyanogenetic genus. The common flax, Linum usitatissimum, is a
well-known cyanogenetic species,’ and several cases of poisoning of
livestock have been traced to it. Fresh material was collected on

1 Received January 3, 1940.
ee 2 pe Ivar. Flora of Utah and Nevada. Contr. U. S. Nat. Herbarium
: oe
8 JORISSEN and Harrs. Bull. Acad. Roy. Belg. (3) 21: 529-540. 1891.

238 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 6

July 13, 1938, from the mountain sides near the Salina (Utah) Experi-
ment Station of the U. 8. Bureau of Animal Industry and at an alti-
tude of 7,800 feet. The plants were then beginning to blossom. The
fresh material was preserved in 1 percent mercuric chloride solution
and shipped to Washington. On analysis after one month the sample
gave 0.98 mg HCN per 100 g of plant. A sample preserved in water
alone gave 1.07 mg under the same circumstances. This species is,
therefore, only shghtly cyanogenetic in the flowering stage.

Linum lewisi Pursh.—Prairie flax. A perennial blue-flowered spe-
cies found, according to Tidestrom, from Manitoba to Alaska, south-
ward to Texas, California, and Mexico. Fresh material of this species
was collected on June 24, 1938, from a locality 10 miles southeast of
Salina, Utah, at about 6,000 feet altitude. The fresh plant preserved
in 2 percent mercuric chloride was shipped to Washington. The
plants were in late-blossom and immature fruit stage of growth. Anal-
ysis after 2 months showed 69 mg HCN per 100 g or considerably
more than L. kingzt contained. If grazed by cattle or sheep, L. lewisu
is likely to produce fatal poisoning. |

Glyceria septentrionalis Hitchc.—The cyanogenetic properties of
eastern mannagrass were discovered by George B. Reynard, of the
U. S. Bureau of Plant Industry, who collected samples at Oakland,
Md., on September 4, 1937. Mr. Reynard stated that the fresh grass
gave a strong Guignard test for HCN. A partly dried sample that he
submitted to the authors was identified by Mrs. Agnes Chase. On
analysis it gave 30.9 mg of HCN per 100 g. Several other species of
Glyceria have previously been reported to be cyanogenetic.* At the
level of HCN found in the specimen G. septentrionalis should be
dangerous to ruminants if eaten in quantity.

Centaurea americana Nutt.—Basket flower, American star thistle.
Seeds of this plant were submitted by Robert Merz, Jr., of College
Station, Tex. Hydrocyanic acid has been reported from the following
species: C. montana and C. solstitialis by Couperot® and from C.
crocodylium by Mirande.® The seed of C. americana gave 5.8 mg. of
HCN per 100 g.

Stillingia dentata (Torr.) Britton and Brown. Dried plants col-
lected by O. G. Babcock in the vicinity of Garden City, Tex., were
supplied by Dr. H. L. Haller, of the U. S. Bureau of Entomology and
Plant Quarantine. Mr. Babcock stated that the plant was very poi-
sonous to sheep and that the circumstances of the poisoning indicated

4Cf. WeumerR, C. Die Pflanzenstoffe, Jena, 1929, Erginzungsband, 1935; and

ALSBERG, C., and Buacx, O. F. Journ. Biol. Chem. 21: 601-609. 1915.
5 WEHMER, C. Loc. cit.

JUNE 15, 1940 HEWETT: NEW FORMATION NAMES, KINGSTON RANGE 239

the possibility of cyanogenetic glucosides and alkaloid being present.
Analysis of the dry leaves and stems showed that 77.7 mg. of HCN
per 100 g could be obtained after 24 hours of maceration in water.
This method is known to give somewhat low results, and it may be
concluded that the actual quantity potential in the dried plant is 10
to 20 percent higher than the figure stated. The quantity of material
at hand was insufficient to permit further investigation of this point.
At this level of HCN it can be said that this plant is very dangerous
to cattle and sheep.

Triglochin palustris L.—Arrowgrass has been reported cyanoge-
netic’ but much less is known about this species than the more im-
portant 7. maritima, and no figures for its HCN content have been
published. Material of T. palustris collected by J. H. Ehlers at Che-
boygan, Mich., in August 1938 and dried was analyzed September 20.
It gave 13.5 mg HCN per 100 g. The plant had flowered but was not
yet in seed. The quantity of HCN found is considerably smaller than
is customarily given by 7’. maritima dried samples of which commonly
yield more than 100 mg of HCN per 100 g, and specimens running
from 200 to 300 mg are not uncommon. One specimen growing under
water contained only 13.4 mg calculated on the basis of fresh plant.

GEOLOGY .—New formation names to be used in the Kingston Range,
Ivanpah quadrangle, Californa.! D. F. Hewerrt, U. 8. Geo-
logical Survey.

A thick series of sedimentary rocks of late pre-Cambrian age that
occur in the Kingston Range have not heretofore been divided into
mappable units. They have been referred to in adjacent regions by pre-
vious writers? as Algonkian strata. The series comprises about 6,500
feet of sandstone, conglomerate, quartzite, shale, and dolomite that
crop out on the north and east slopes of the Kingston Range. They are
somewhat similar to a series of rocks that occur in the Grand Canyon
_ region of Arizona. This assemblage of sedimentary rocks in the King-
ston Range is named the Pahrump series, from a nearby valley lying
north and northeast of the Kingston Range. It rests unconformably
on pre-Cambrian granite gneiss and is overlain unconformably by
sedimentary rocks that probably represent the basal formation of the

6 GresHoFF, M. Pharm. Weekbl. 45: 1165-1169. 1908; BuanKsma, ibid. 50:
1295-1302. 1913.

1 Published by permission of the Director, Geological Survey, U. 8S. Department of
the Interior. Received March 23, 1940.

2 Nose, Levi F. Rock formations of Death Valley, Calif. Science, n.s., 80 (2069):
Mees). 1934. HazzarvD, JOHN C. Paleozoic section, Nopah and Resting Springs
Mountains, Inyo County, Calif. Calif. Journ. Mines and Geology, State Mineralogist’s
Rept. 33(4): 288-339. 1938.

240 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 6

Paleozoic rocks, Lower Cambrian in age. The formations to be de-
scribed are as follows:

Kingston Peak formation.
Pahrump series; Beck Spring dolomite.
Crystal Spring formation.

Crystal Spring formation.—This formation crops out in a belt that extends
for 5 miles around the north slope of the Kingston Range and in two areas
on the eastern slope of the range. The formation comprises about 2,000 feet
of quartzite, arkose, shale, and limestone, intruded by sills of diorite and
syenite, with beds of dolomite and chert at the top and thick coarse con-
glomerate at the base. The base of the formation has been observed at
only one locality, on the west end of a ridge 2 miles east of Horse Spring.
Here conglomeratic quartzite rests on pre-Cambrian gneissic granite. No
fossils were found in the formation. The name of the formation is derived
from Crystal Spring on the north slope of Kingston Range.

Beck Spring dolomite-—The Beck Spring dolomite conformably overlies
the Crystal Spring formation and crops out conspicuously in a belt 6 miles
long on the north slope of Kingston Range. Two smaller blocks of the
dolomite, each about 1 mile long, crop out in the hills northeast and east of
the range. The formation is made up largely of beds of light bluish-gray
dolomite, 2 to 4 feet thick, separated by layers of shaly material, and is
about 1,000 feet thick. No clearly identifiable fossils have been found in it,
but layers of roundish concretions about half an inch in diameter, found
at one place, may be of algal origin. Beds of oolite several feet thick are also
numerous. The name of the formation is derived from Beck Spring on the
north slope of the range.

Kingston Peak formation.—The Kingston Peak formation is an unusual
assemblage of sediments that conformably overlie the Beck Spring dolomite.
It crops out in a belt 20 miles long around the north and east slopes of
Kingston Range but is interrupted by several faults. The formation is
about 1,900 feet thick, along the western part of the belt, but at the east end
it appears to be much greater. The formation consists of sandstone and lime-
stone with a thick coarse conglomerate in the middle. The conglomerate is
made up largely of subangular cobbles of limestone up to 12 inches in di-
ameter with quartzite and granite cobble predominating near the base. This
uncommon assemblage of sediments resembles the debris that forms fans on
the border of the present valleys, called fanglomerate. The name of the
formation is derived from Kingston Peak, the highest point in the range.

BOTAN Y.—Three fungi destructive to free-living terricolous nema-
todes.1| CHARLES DREcHSLER, U. S. Bureau of Plant Industry.

In several earlier papers (5, 6, 8, 9, 10) comparative treatment was
accorded to 24 fungi that had been observed to subsist on free-living
nematodes infesting old agar cultures started from diseased rootlets
or from other decaying vegetable materials. As the agar media em-
ployed were of a concentration sufficient to insure a rather firm con-
sistency together with relative freedom from liquid water, the cul-

1 Received December 6, 1939.

JUNE 15, 1940 DRECHSLER: FUNGI DESTRUCTIVE TO NEMATODES 241

tures provided approximately terrestrial rather than aquatic condi-
tions, and therefore not only encouraged the multiplication of eel-
worms mainly terrestrial with respect to source and adaptation, but
also permitted development of the similarly terrestrial fungi habitu-
ally destructive to them under natural conditions. The 24 fungi
hitherto treated include representatives of two distinct groups—21
species, mainly predaceous in mode of attack, being referable to an
interrelated series of hyphomycetes wherein are embraced members
of the genera Arthrobotrys, Trichothecium, Dactylella, Dactylaria, Tripo-
sporina, and Tridentaria; the remaining 3 species being referable to
the Zoopagaceae, a distinctive family of conidial phycomycetes most-
ly destructive to terricolous rhizopods.

Apart from recognizable members of the predaceous mucedinaceous
series and of the Zoopagaceae, about a score of additional fungi have
been observed to destroy free-living nematodes in agar cultures.
Nearly all these forms are parasitic in their mode of attack, infecting
their hosts by germination of adhering conidia. Only one among them,
the widely distributed and very destructive Harposporium anguillulae
Lohde, can with any certainty be referred to a species previously re-
ported as parasitic on nematodes. This meager agreement with earlier
findings is not surprising when it is considered that in large part the
literature on fungus parasites of nematodes records observations made
on animals kept under conditions widely different from those obtain-
ing in agar cultures. The relevant contributions by Braun (3), Soro-
kine (23), Sadebeck (19), Dangeard (4), Lindner (13), Maupas (16),
Micoletzky (17), and Sparrow (24, 25) dealt presumably with infec-
tions of actively motile eelworms that came about under conditions
apparently more nearly aquatic than terrestrial. Baunacke (1), Gof-
fart (11), Korab (12), and Rozsypal (18), on the other hand, were
concerned with parasites destructive to the eggs and the almost
equally inert larvae in cysts of Heterodera schachtii Schmidt. Now,
manifestly, infection of active eelworms by fungus spores under
aquatic conditions, and infection of inert stages under terrestrial con-
ditions, are not attended by the difficulties usual in infection of ac-
tively motile eelworms under terrestrial conditions. For in their brisk
movement through soil or through decaying organic materials, terric-
olous nematodes constantly tend to scrape away the spores adhering
to them externally, much as cattle rid themselves of the larger sting-
ing flies in walking through dense shrubbery. Small size, slender or
flattened shape, and strong adhesiveness are among the physical at-
tributes whereby the spores of successful parasites are enabled to

242 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

withstand the adverse mechanical action to which they are exposed.
In the two phycomycetous parasites herein described some of these
attributes are derived from developmental modifications that appear
to be of taxonomic significance.

HAPTOGLOSSA HETEROSPORA

A fungus evidently referable to one or another of the several zoospore-
producing groups in the Phycomycetes has frequently been observed de-
stroying enormous numbers of nematodes in agar cultures prepared from
media of rather soft consistency and planted with decaying portions of roots
or stems originating in greenhouse or field. It has also been found destroying
nematodes in agar cultures that after being permeated with Pythium myce-
lium had been further planted with pinches of leaf mold collected in decidu-
ous woods in Virginia, Maryland, or Wisconsin. Most of. the species of
eelworms commonly infesting soft agar cultures started from vegetable
materials that have undergone partial decomposition in contact with moist
eround, appear susceptible to attack by the fungus, though differences in the
rapidity of their destruction are often noticeable. The appearance of the
fungus in a petri-dish culture leads usually within a few days to the extermi-
nation of all active individuals of the susceptible species present. In its
spectacular destructiveness and widespread distribution the fungus invites
comparison with Harposporium anguillulae and with the larger of the
retiary predaceous hyphomycetes, particularly Arthrobotrys oligospora Fres.,
A. conoides Drechsl., A. musiformis Drechsl., and Dactylaria thaumasia
Drechsl.

During the earlier stages of its development within a nematode the fungus
is relatively inconspicuous. At about the time the infected animal becomes
incapable of further locomotion, the parasite may be discerned imbedded in
the granular and globulose material resulting from degeneration of muscula-
ture and organs (Fig. 1, A). With further appropriation of the degenerating
material it improves markedly in visibility and increases somewhat in
volume (Fig. 1, B), so that when the last remnant of digestible substance
has been absorbed, it is revealed with much distinctness inside the otherwise
empty host integument (Fig. 1, C-G).

In an individual host animal the parasite may be represented by a single
thallus (Fig. 1, F) or by many thalli (Fig. 1, A-E, G). Instances of infection
by a single thallus or by two or three thalli are especially frequent early in
the course of an epizootic. As the epizootic continues the number of thalli
in the individual infected eelworm increases rapidly, with the result that in
the end instances of infection by more than 50 (Fig. 1, C) or even more than
100 thalli are not infrequent. This increase in number, naturally, is accom-
panied by proportional decrease in size, and, somewhat secondarily, by some
change in shape; the larger vegetative bodies being generally of cylindrical
conformation, with bluntly rounded ends, whereas the smaller specimens are
mostly of prolate ellipsoidal conformation. Irregular modifications of out-
ward form, such as median constrictions and narrowed terminal prolonga-
tions, occur here and there. The larger vegetative bodies are frequently
somewhat curved or crooked, either because of crowding by their fellows
(Fig. 1, B, G), or because of constraint from the posture taken by the host
animal in its death (Fig. 1, F).

Following exhaustion of their food supply the thalli of the fungus are
transformed into sporangia. The smaller specimens (Fig. 1, I, J, K) each put

JUNE 15, 1940 DRECHSLER: FUNGI DESTRUCTIVE TO NEMATODES 243

forth laterally a single broad protrusion that presses the animal’s integument
outward in a perceptible bulge. More or less simultaneously cleavage lines
appear in the granular protoplasmic contents, dividing them into units of
approximately equal volume. Suddenly the broadly rounded tip of the pro-
trusion yields, and the delimited protoplasmic lumps are in large part
forcibly expelled through the opening. Transformation of the larger thalli
usually entails development of several protuberances at some distance from
one another. In instances where five or six beaks are formed, two or three of
them may participate in dehiscence. Regardless of the number of openings
present, a considerable proportion of the protoplasmic lumps usually fail to
be discharged (Fig. 1, H, b, c; L, O), and must await their liberation by the
disintegration of host integument and sporangial membrane.

After being propelled some distance into the adjacent medium, the dis-
charged protoplasts round up individually to form irregularly globose im-
motile spores (Fig. 1, H, c; L). Each of the encysted bodies soon gives rise
to a bud, which then grows out (Fig. 1, Q) to form eventually a flattened
structure with an elongated lateral lobe that is usually curled at its tip in a
tonguelike manner (Fig. 1, M). Often the axial or proximal portion of the
flattened structure, like the envelope of the immotile spore to which it
remains attached, is found evacuated of contents, all of the protoplasmic
material having been collected in the glossoid lobe. Before long, hundreds of
the curiously constituted units thus resulting from asexual reproduction
surround the integumentary remains of each host animal like a dense cloud.

When a healthy nematode passes through such a cloudlike array, some of
the reproductive bodies become fixed to it; attachment being accomplished
apparently through adhesion of the concave surface of the glossoid lobe to
the animal’s integument. Upon perforation of the integument by a minute
opening the protoplasmic contents of the reproductive body migrate inward
to form a small saclike structure. This structure then proceeds to develop
autonomously, thereby again giving rise to a cylindrical or ellipsoidal thallus.

During the later and more readily discernible stages in their development
some of the thalli can be seen attached to the animal’s integument by a
conical connection devoid of protoplasmic contents (Fig. 1, G). In this
conical part is to be recognized unquestionably the saccate body by which
invasion was initiated. The larger number of thalli, however, show no evident
connection to the integument, a circumstance that might imply either that
thalli often multiply by division, or that connections originally present often
are destroyed as a result of movement by the animal. An approximate end-
to-end arrangement of thalli (Fig. 1, A, B) occasionally to be observed argues
somewhat in favor of the former alternative, while the latter is supported
_more especially by the frequent absence of any observable connection be-
tween integument and fungus in animals parasitized by only a single thallus
(Fig. 1, F). Since such single thalli, when occurring in large animals, attain
impressively large dimensions without dividing into units of smaller size, it
may be presumed that vegetative division is at least not a constant feature
in the development of the fungus. The presence of very numerous thalli in
infected animals during the later stages of an epizootic in an agar culture is
very obviously attributable to the enormous numbers of germinated spores
then distributed through the medium. Often in such later stages the incipient
infections from individual spores are so numerous and close together that
the intruding protoplasts, operating virtually as a mass, will extensively
separate integument from naked musculature. In the head region, especially,
such wholly pathological separation—not to be confused with hastened

244

JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 6

foes

ee z

O CLIO,
Geeeeeveey

C.Drechster del. as

Fig. 1.—(See opposite page for explanation)

JUNE 15, 1940 DRECHSLER: FUNGI DESTRUCTIVE TO NEMATODES 245

molting—is often associated with conspicuous deformation of fleshy struc-
tures.

With respect to size the immotile spores from each individual sporangium
show only moderate variation. Most frequently, perhaps, they range in
diameter between 4.5 and 6y (Fig. 1, H, 6, c; L; M). Many sporangia, how-
ever, give rise to spores ranging in diameter between 8 and 10u (Fig. 1,
N; O: P). It is not apparent that spores of intermediate dimensions are pro-
duced except as unusually oversized specimens in clusters referable to one
category, or as undersized specimens in clusters referable to the other cate-
gory. As sporangia producing the larger spores frequently occur in the same
host animal with sporangia producing the smaller spores, the difference in
dimensions can not readily be held traceable to differences in environmental
conditions. Unstained living material reveals no divergence in manner of
development or in mode of infection associated with the inequality of size.

Presumably the production of flattened lobate or glossoid infective struc-
tures by the discharged immotile sporangiospores represents a develop-
mental process equivalent to the emission of motile zoospores from encysted
sporangiospores familiar in the saprolegniaceous genera Achlya and Apha-
nomyces. Certainly, the type of asexual reproduction concerned here shows
less similarity to the Pythiwm-like reproduction that is found associated
with more or less swollen endoparasitic thalli in the genera Lagena, Lagendt-
dium, and Myzocytium, than to the Achlya-like reproduction associated with
swollen endoparasitic thalli in Ectrogella and Aphanomycopsis, as these
genera were understood by Scherffel (21). With Ectrogella and A phanomycop-
sis, moreover, the fungus under consideration offers further morphological
parallelism in the pronouncedly grandular consistency of the protoplasm in
its thalli—a consistency which Scherffel held with some justification to dis-
tinguish the Saprolegniaceae and Peronosporales from the Ancylistales and
Chytridiales.

In its curious emission of immotile infective structures from immotile
sporangiospores the fungus would seem to provide a plausible transition be-
tween the Achlya-like diplanetism of Ectrogella and A phanomycopsis, on the
one hand, and the direct discharge of immotile infective sporangiospores in
Protascus subiliformis Dangeard (4), a parasite destructive to nematodes,
on the other. The existence of such a transitional type of asexual reproduc-
tion lends support to Maupas (16) and Maire (14) in their interpretation of
the conical sporangiospores of P. subiliformis as bodies homologous to motile

Fig. 1.—Haptoglossa heterospora, drawn to a uniform magnification with the aid of
a camera lucida; X 500 throughout. A, B, C, D, Nematodes probably referable to
Rhabditis dolichura Schneider, containing respectively 16, 5, 54, and 10 thalli of the
- fungus. E, Nematode, referable to Cephalobus sp., containing 10 thalli of the fungus.
F, G, Specimens of Acrobeloides biitschlii (De Man) Thorne occupied respectively by
1 and 18 thalli of the fungus; owing to lack of space F is shown in three parts, which
connect at the points y and z. H, Three neighboring thalli: a, in an early stage of trans-
formation into a sporangium, showing two papillae of dehiscence; b, a sporangium,
partly evacuated, within which the numerous retained spores have encysted; C, & Spo-
rangium showing in profile a papillar orifice through which many zoospores have been
discharged, though a large number, too, have encysted within. I, J, K, Three small
sporangia, each showing a papilla of dehiscence some little time previous to discharge of
spores. L, Sporangium after partial emission of spores. M, Discharged spores of the
usual smaller size, each of which has given rise to a lobate infective body. N, Spo-
rangium with two ‘papillae of dehiscence; its contents showing demarcation into spores
of the larger size. O, Sporangium within which three spores of the larger size have
germinated to form infective bodies. P, Spores of the larger size, each with an infec-
tive body attached to it. Q, Spores of the smaller size, showing different stages in the
emission of infective bodies.

246 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

zoospores of the aquatic phycomycetes. Indeed, the transitional type of
sporangiospore development herein set forth may have been known to these
authors, as Maupas took occasion to allude to “les spores, également im-
mobiles, de mon Ectrogella? sp.”’ I have not found any published account of
the form to which this allusion applied, yet as it is recorded by his obituarist
(22) that Maupas worked rather extensively on fungi parasitic on nematodes,
the doubtful Ectrogella species may have been encountered as a parasite on
eelworms, and may therefore have been a fungus related to, or perhaps even
identical with, the one under discussion.

Achlyogeton entophytum Schenk and A. rostratum Sor., both reported by
Sorokine (23) as being parasitic on nematodes, differ from my fungus in that
their thalli become partitioned into somewhat distended segments, which,
though functioning individually as sporangia, remain connected with one
another. In their development the zoospores of A. entophytum reveal an
Achlya-like diplanetism, except for the difference, if Schenk’s account (20)
is to be trusted, that the motile stage is uniciliate rather than biciliate. While
Schenk’s representations relative to the number of cilia are so much at
variance with expectations that their correctness has been questioned, the
observations of Martin (15) and Tokunaga (26) have corroborated the exist-
ence of a secondary motile stage; and it seems certain, at least, that in A.
entophytum the zoospore protoplast does not remain attached to the empty
cyst envelope as an immotile infective body.

Erection of a new genus, under a name compounded of two words meaning
“to fasten’ and ‘‘tongue,” is made necessary by the curious development of
the sporangiospores in the present fungus.

Haptoglossa gen. nov.

Thalli singuli vel complures in viva animalia crescentes, elongato-cylind-
rati vel elliptici, maturitate protoplasmatis dense granulosi repleti, omnino
sporangia evadentes; sporangio uno vel aliquot tubis papillisve mox in-
structo, sporas inertes expellente; sporis inertibus a vesicula non circum-
datis, quisque corpus planum inertem inficiens proferente.

Vegetative thalli elongate-cylindrical or ellipsoid in shape, developing
singly or in variable numbers within living animals, at maturity filled with
densely granular protoplasm and in their entirety becoming converted into
sporangia; the individual sporangium provided with one or several evacua-
tion tubes or evacuation papillae, through which ready-fashioned spores are
forcibly expelled without formation of any enveloping vesicle; spores im-
motile and without cilia, each putting forth an immotile infective body, often
of flattened lobate shape.

Haptoglossa heterospora sp. nov.

Thalli hyalini, elliptici vel elongato-cylindrati, 7-350u longi, 4-18 crassi;
sporangiis 1-6 humilibus papillis praeditis, per 1-4 ora 4—7y lata dehiscenti-
bus, aliis sporas 4.5—6u crassas aliis sporas 8-10u crassas eicientibus; sporis
hyalinis, globosis vel rotundo-angulosis; corporibus inficientibus planis,
5-10u longis, a latere lobo lingulato curvato 3—-8u longo apice attenuato
instructis.

Vermiculos nematoideos multarum specierum enecans habitat in terra
et humo silvestri et materiis plantarum putrescentibus in Maryland et
Virginia et Wisconsin.

Vegetative thalli hyaline, ellipsoidal or elongate cylindrical, 7 to 350u
long, 4 to 18u in transverse diameter; sporangia developing 1 to 6 short |

=

JUNE 15, 1940 DRECHSLER: FUNGI DESTRUCTIVE TO NEMATODES 247

broad papillae of which 1 to 4 may be functional in providing apertures of
dehiscence mostly 4 to 7u wide. Spores hyaline, globose or with rounded
angles, those produced in some sporangia 4.5 to 6u in diameter, those pro-
duced in other sporangia mostly 8 to 10u in diameter. Infective bodies flat-
tened, consisting individually of an axial portion 5 to 10u long, together with
a broad-based glossoid lobe mostly 3 to 8u long, more or less upcurved at its
tapering end.

Destroying nematodes of many species, it occurs in soil, leaf mold, and
decaying vegetable materials in Maryland, Virginia, and Wisconsin.

MERISTACRUM ASTEROSPERMUM

A conidial phycomycete manifestly related to the entomogenous Ento-
mophthoraceae was observed destroying nematodes in some agar plate
cultures to which had been added pinches of leaf mold taken from deciduous
woods near Butternut, Wis., in September, 1938. Among the eelworms sub-
ject to destruction several species belonging evidently to the genera Rhab-
ditis, Aphelenchoides, and Plectus could be distinguished.

Infection is accomplished by the fungus in question through the germina-
tion of a conidium or of several conidia attached to the integument of a
susceptible animal, most often toward its anterior end (Fig. 2, A, B, C).
Usually the means of attachment is clearly visible in a sizable deposit of
adhesive material between spore and integument. The proximal portion of
the germ tube—the portion most directly concerned in penetration of the
integument—is commonly set off from its distal continuation within the
fleshy interior by yellow coloration and somewhat irregular thickening of its
wall. When the invading element has attained a length of approximately
10y, it widens abruptly to form a terminal swollen body, in which the proto-
plasmic contents of the conidium and infection tube then migrate. The
swollen body thereupon continues its development by extending itself
lengthwise through the animal as a stout assimilative hypha or thallus.
Progressive destruction of musculature leads to increasingly pronounced
disablement of the eelworm, and finally to its death. Vegetative growth of
the assimilative hypha comes to an end only when the digestible contents of
the host have been completely appropriated.

Early in the course of an epizootic within a petri-dish culture, infection
of nematodes by single conidia is more frequent than infection by plural
conidia. Later, as conidia of the fungus become more abundant, plural in-
fection, with resultant development of two (Fig. 2, B, C, D), three, or four
thalli, tends to predominate. On conclusion of their vegetative growth the
assimilative hyphae, with the exception of occasional small specimens,
_ undergo division into a number of segments, which appear rather obviously
homologous to the “‘hyphal bodies”’ familiar in the entomogenous genera
Empusa and Entomophthora. Division in each instance is accomplished by
deposition of a cross wall (Fig. 2, D, b, c) and subsequent disarticulation of
the delimited parts. The thallodic segments may be elongate-ellipsoidal
(Fig. 2, D, a, e), or somewhat curved-cylindrical (Fig. 2, D, 6, c, g); or, again,
may have a more irregular shape with bulging expansions (Fig. 2, D, f) or
short diverticulate lobes (Fig. 2, D, d).

Asexual reproduction may be considered to begin when the individual
thallodic segment puts forth a hypha, about 3.5 to 5u wide, that after
emerging from the enveloping host integument grows through the overlying
medium (Fig. 2, EK, F) out into the air. If the host animal has succumbed
deep in the medium, the outgrowing hypha often needs to make its way

JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

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JUNE 15, 1940 DRECHSLER: FUNGI DESTRUCTIVE TO NEMATODES 249

through solid material for a distance of 1 to 3 mm; whereas if the animal
happened to die on the surface the hypha may extend itself into the air im-
mediately. The aerial hypha or aerial prolongation, usually less than 0.5 mm
in length, widens out perceptibly in its distal portion, and rather commonly
is disposed here in a helicoid spiral of two, three, or four turns (Fig. 3, A).
Into this somewhat modified terminal portion of hypha is soon transferred
all the protoplasm from the entire thallodic or reproductive unit. Transverse
septa are now laid down in basipetal succession, dividing the densely filled
hyphal termination into cells of approximately equal protoplasmic content.
After a bud has been put forth terminally from the apical cell, a similar bud
is put forth laterally in basipetal succession from each of the lower cells
(Fig. 2, G), each of the buds receiving the entire protoplasmic contents of
the parent cell (Fig. 2, H, I, J), and then being delimited by a basal septum
as an obovoid conidium (Fig. 2, K-Q). Detachment of the conidia appears
to ensue spontaneously soon after they attain full development. Once they
lie on the surface of the medium they are ready to adhere to any susceptible
nematode that may come in contact with them; such adhesion then in-
augurating another sequence of endoparasitic and reproductive develop-
ment. However, when opportunity for infection is not presented within a
certain period of time, the detached conidium gives rise to a secondary
conidium closely resembling the primary one except for its somewhat smaller
size (Fig. 2, R, 8, T). The secondary spore is regularly borne aloft a few
microns above the surface of the medium by means of a delicate filament
often more or less prostrate in its proximal portion, but ascending or erect
in its distal portion. Through development similar to that whereby it origi-
nated a secondary conidium may in turn give rise to a tertiary one (Fig.
2s (UD)

: Frequently a thallodic segment, instead of putting forth a conidiophorous
filament, forms a resting spore. By continued enlargement of a vacuole at
each end of the segment (Fig. 2, D, 6) the granular materials are brought
together into a globose mass, which then lays down two somewhat convex
end walls to form a subspherical cell (Fig. 2, V). The densely granular con-
tents of this cell now are converted into an endogenous spore that is provided
with a separate wall of its own. This wall appears prominently scalloped in
profile, and only rather loosely surrounds the coarsely granular subspherical

Fig. 2.— Meristacrum asterospermum, drawn to a uniform magnification with the
aid of a camera lucida; X 1000 throughout. A, Specimen of Plectus communis Biitschli
showing early stage of infection by a germ-tube from an adhering conidium. B, An-
terior portion of a specimen of Rhabditis dolichura with two adhering conidia, one of
which has given rise to a growing vegetative thallus in the animal. C, Parasitized
nematode, possibly referable to Aphelenchoides sp., the contents of which have been
almost completely appropriated by two thallii. D, Portion of a specimen of R. do-
lichura containing seven thallodic segments of the fungus, a—g; one of these segments, b,
which is not yet separated from its neighbor c, shows an early stage in the development
of an azygospore; the other segments, a, c-g, subsequently put forth conidiophorous
filaments. E, Thallodic segment from which a conidophorous filament is being put forth.
I’, Thallodic segment (with envelope of empty spore attached) likewise with an elongat-
ing conidiophorous filament. G, Sporiferous termination of conidiophorous filament,
showing basipetal deposition of septa, and early stage in the budding forth of conidia
from the delimited cells. H, I, J, Later stages in formation of conidia. K—Q, Disarticu-
lated primary conidia. R, Primary conidium in process of producing a secondary co-
nidium. U, Empty envelopes of a primary anda secondary conidium shown attached to
a tertiary conidium. V, Same thallodic segment as D, b, but at a later stage, showing de-
limitation of the sporangial cell within which an azygosvore is tc be produced. W—Z
Mature azygospores, each within its sporangial wall.

’

250 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

protoplast, in which, on advancing maturity, a homogeneous central reserve
globule may be distinguished (Fig. 2, W—Z).

With respect to the disposition of the membrane and contents of its resting
spores the fungus shows striking similarity especially to the members of the
genus Hndocochlus (7), in the Zoopagaceae. Although in the few known
species of Hndocochlus the resting spores are always formed as a result of
sexual union between portions of separate thalli, development of azygospores
in a manner inviting comparison with the fungus under discussion is present
in at least one member of the Zoopagaceae, namely in Zoopage cladosperma
Drechsler (7). Resemblance to the genus Endocochlus is expressed also in the
production of conidia by individual segments delimited through deposition
of cross walls in aerial hyphae. This parallelism in asexual reproduction may
well be of taxonomic significance despite some divergence in outward
morphology apparent not only in the widening of the sporiferous termina-
tion, but also in its helicoid modification. The fungus under discussion shows
further similarity to many endoparasitic Zoopagaceae in the frequently
rather extended migration of its protoplasm from deeply immersed thallodie
segments to the aerial terminations of its conidiiferous hyphae.

However, if the granular consistency and ready mobility of its protoplasm
are considered together with the division and disarticulation of its vegetative
hyphae into disconnected segments, the fungus appears more intimately
akin to the Entomophthoraceae than to the Zoopagaceae. The indeterminate
elongation of its conidiophorous filaments finds a close parallel in the inde-
terminate extension of external hyphae from the intramatrical mycelial
segments in species of Ancylistes, a genus recently shown by Berdan (2)
to be properly referable to the Entomophthoraceae. In their obovoid shape
the conidia of the fungus reveal a very obvious similarity to those of many
species of Empusa and Entomophthora. Often, besides, a fully developed
conidium was seen to be attached to a dome-shaped protrusion of the
parent cell; and the denuded empty conidiiferous cells were nearly always
found badly collapsed, much like conidiophores of Conidzobolus villosus
Martin some time after their energetic discharge. Although these details
indicate a parallelism with the genera in the Entomophthoraceae known for
their forcible projection of conidia, I did not succeed in demonstrating that
the fungus under discussion likewise discharges its asexual spores forcibly.
It seems probable, however, that the primary conidia are actively propelled,
though with much less violence than in species of Empusa, Entomophthora,
and Conidiobolus. The secondary and tertiary conidia are almost certainly
never shot away, but are produced and held in slightly elevated positions,
where opportunity for contact with nematodes given to surface locomotion
appears especially favorable.

In the Entomophthoraceae the fungus is distinguished from those hitherto
made known by its production of plural conidia, for the most part laterally,
on a multiseptate conidiophore. It is accordingly described as the type of a
new genus of that family. A term compounded of two words meaning
“divided” and ‘‘tip” respectively, is deemed an appropriate name for the
genus. The sculpturing of its azygospore wall suggests for the fungus a
specific epithet made up from two words meaning “‘star’” and “‘seed,”’ re-
spectively. :

Meristacrum gen. nov.

Hyphae assumentes intra animalia viva crescentes, simplices vel non-
nihil ramosae, primo continuae, postea—animali emortuo et carne ejus
consumpto—in aliquot partes se diffindentes; his partibus disjunctis aut

JUNE 15, 1940 DRECHSLER: FUNGI DESTRUCTIVE TO NEMATODES 251

hyphas fertiles emittentibus aut sporas perdurantes (zygosporas vel azygo-
sporas) gignentibus; hyphis fertilibus in aere se porrigentibus, sursum proto-
plasmatis repletis, ibi primo continuis, deinde transverse multiseptatis, ex
quaque cellula conidium proferentibus. _

Vegetative hyphae developing within living animals, simple or somewhat
branched, at first continuous, later (after death of the animal and exhaustion
of its fleshy contents) often dividing into disjointed segments, which either
put forth conidiophorous hyphae or give rise to resting spores (zygospores or
azygospores) ; the individual conidiophorous hypha extending some distance
into the air, in its densely filled terminal portion becoming divided by trans-
verse septa into a number of cells, each of which then produces a single
conidium.

Meristacrum asterospermum sp. nov.

Hyphae assumentes hyalinae, simplices vel parvulum ramosae, saepius
50—500u longae, 5-12 crassae, in articulos disjunctos plerumque 25-60u
longos se diffindentes; quoque articulo mox utrimque rotundato, aut unam
hypham fertilem emittente aut unam azygosporam gignente; hyphis fertili-
bus saepe 0.5-3 mm longis, 200—500u in aere se porrigentibus, parte propiore
saepe immersa 3—5y crassa mox vacua et sparsum septata, parte ulteriore
aeria saepius in spiram levem contorta, 6-9» crassa, primo protoplasmatis
repleta, continua, deinde in 5-15 cellulas 4—20u longas divisa quae singula
conidia proferunt. Conidiis hyalinis, obovoideis, plerumque 8—11y longis,
6—-Su crassis, modo singulas hyphas germinationis circa 10u longas, 1.5u
crassas in vermiculum nematoideum intrudentibus, modo quoque unum
conidium ordinis secundi ex hypha germinationis assurgente 5-10 longa,
basi circa 2u et apice 0.8u crassa ferente. Azygosporiis hyalinis vel flavi-
dulis, globosis, circa 15-20 crassis, maturitate membrana late verrucosa
vel undulata cellulam viventem sphaeroideam 11—14u crassam laxe circum-
dante.

Vermiculos nematoideos enecans habitat in humo silvestri prope Butter-
nut, Wisconsin.

Vegetative hyphae hyaline, simple or slightly branched, mostly 50 to 500u
long, 5 to 12u wide, when well developed dividing into segments 25 to 60u
long; the thallodic segments becoming rounded at their ends, each putting
forth a single conidiophorous filament or giving rise to a single azygospore.
Conidiophorous filaments mostly 0.5 to 8 mm long, usually extending 200
to 500u into the air; the proximal part usually 3 to 5u wide, soon becoming
empty and septate at long intervals; the distal prolongation, often 6 to 9u
wide and noticeably disposed in a helicoid spiral, becoming divided through
basipetal deposition of transverse cross walls into 5 to 15 cells, mostly 4 to
20u long, each of which by budding gives rise to a single sessile, hyaline,
obovoid conidium, 8 to 11y long and 6 to 8u wide. Conidium after disarticu-
lation often adhering to a nematode, then thrusting into it an infective germ
tube, about 10u long and 1.5u wide, which develops into a vegetative hypha;
or, failing to adhere to a nematode, giving rise on an ascending sporophore,
5 to 10u long, 2u wide at the base, and 0.8u wide at the tip, to a somewhat
smaller secondary conidium. Azygospore loosely contained within the wall
of its parent cell, colorless or slightly yellowish, subspherical, 15 to 20u in
diameter, its pronouncedly undulated wall loosely enveloping a subspherical
protoplast 11 to 14y in diameter.

Parasitic on nematodes it occurs in leaf mold in deciduous woods near
Butternut, Wis.

252

JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

C.Drechsler del.

Fig. 3.—(See opposite page for explanation)

JUNE 15, 1940 DRECHSLER: FUNGI DESTRUCTIVE TO NEMATODES 253

CEPHALOSPORIUM sp.

A fungus subsisting on eelworms, yet hardly deserving to be considered
either predaceous or vigorously parasitic on these animals, was observed in
a few nematode-infested agar cultures to which had been added some pinches
of partly decayed tobacco leaves (Nicotiana tabacum L.) from a greenhouse
near Beltsville, Md. In these cultures it apparently obtained its nourishment
exclusively from specimens of Plectus communis Biitschli and P. parvus
Bastian that had become disabled as a result of extensive infection by a
protozoan endoparasite producing minute subspherical spores. When a dis-
abled animal was halted near mycelial hyphae of the fungus, these hyphae
would send out one or several branches, which soon perforated its integument
and intruded a corresponding number of haustorial elements (Fig. 3, B, C,
D). In most instances of plural penetration only one or two of the haustorial
elements attained extensive development, apparently because the fleshy
contents of the eelworm had already in large part been appropriated by the
protozoan. Similar restriction of hyphal development within nematodes
disabled and partly expropriated of contents by a protozoan parasite was
noted earlier in my account (10) of the feebly predaceous Tridentaria
doedycoides Drechsl.

Although its general habit recalls that of the more delicate species in the
predaceous series of hyphomycetes, the fungus at present can neither be
definitely assigned to nor excluded from that series. Its mycelial hyphae vary
from 1 to 3u in width. In addition to branches of indeterminate length, they
bear here and there short spurlike branches that are often more or less
appressed to the parent filament (Fig. 3, B, a). Ordinary cross walls occur
in the mycelial hyphae at moderate intervals, and are supplemented by
curiously oblique partial septa, which sometimes appear alone (Fig. 3, D, b),
and at other times occur united to partial cross walls (Fig. 3, D, a, c). In-
fective branches are often noticeably stouter than the filaments from which
they arise. The haustorial hyphae are often wider in their proximal portions
than in their distal prolongations.

Hyaline ellipsoidal spores, commonly 2.5 to 3u long and 1.5 to 2u wide,
are produced by the fungus on tapering branches, mostly 1.5 to 4u long, and
1 to 2u wide at the base. Owing, perhaps, to repeated disturbance by nema-
todes, these spores, when developed on the surface of a culture, show only a
very promiscuous arrangement (Fig. 3, E); whereas in submerged and con-
sequently better protected positions they are found in capitate clusters
numbering ten to twenty individuals (Fig. 3, G, F). They have never been
observed to infect nematodes, yet their small size would seem to make them
incapable of giving rise to mycelia of such extent that disabled eelworms

Fig. 3—Drawn to a uniform magnification with the aid of a camera lucida; X 1000
throughout. A, Sporiferous termination of a conidiophorous filament of Meristacrum
asterospermum, showing its distal disposition in an extended helicoid spiral. B, Speci-
men of Plectus communis which, after its disablement by a protozoan parasite, is being
invaded besides by two hyphae of Cephalosporium sp.; a, a short spurlike hyphal
branch. C, Another disabled specimen of P. communis, penetrated by five hyphal
branches of Cephalosporium sp. D, Specimen of P. parvus Bastian, disabled by proto-
zoan parasite and then further invaded by a branching filament of Cephalosporium sp.;
a—c, three oblique partial septa. E, Superficial hypha of Cephalosporium sp. with sev-
eral spore-producing sterigmata. F, G, Submerged filaments showing clusters of spores
at the tips of lateral sterigmata. (Owing to lack of space, A is shown in two parts, which
are connected in a. The small sketch, in which the lower portions of the successive
turns are stippled, is intended to show more clearly the direction of rotation in the
helicoid termination.)

254 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 6

might become a likely source of nourishment. These circumstances suggest
that the Cephalosporrum stage present here may constitute an accessory
reproductive phase in the development of the fungus, rather than its main
conidial phase.

LITERATURE CITED

1. BaunackE, W. Untersuchungen zur Biologie und Bekimpfung des Riibennematoden
Heterodera schachtii Schmidt. Arb. Biol. Reichsanst. Land- und Forstw. Berlin-
Dahlem 11: 185-288. 1922.

2. BerRpDAN, H. B. Revision of the genus Ancylistes. Mycologia 30: 396-415. 1938.

3. Braun, A. Uber einige neue Arten der Gattung Chytridium und die damit ver-
wandte Gattung Rhizidium. Monatsb. Ko6nigl. Preuss. Akad. Wiss. Berlin
1856: 587-592. 1856.

4. DanGEARD, P. A. Les ancétres des champignons supérieurs. Le Botaniste 9:157—
303. 1908.

5. Drecuster, C. A new species of conidial phycomycete preying on nematodes.
Mycologia 27: 206-215. 1935.

6. ———. A new species of Stylopage preying on nematodes. Mycologia 28: 241—
246. 19386.

7. ———. New conidial phycomycetes destructive to terricolous Amoebae. Mycologia
28: 363-389. 1936.

8. — . Some hyphomycetes that prey on free-living terricolous nematodes. Myco-
logia 29: 447-552. = 1937.

9. . Five new Zoopagaceae destructive to rhizopods and nematodes. Mycologia

31: 388-415. 1939.

10. ————. Three new hyphomycetes preying on free-living terricolous nematodes.
Mycologia 32. 1940.

11. Gorrart, H. Untersuchungen am Hafernematoden Heterodera schachtii Schm.
Art. Biol. Reichsanst. Land- und Forstw. Berlin-Dahlem 20: 1-26. 1932.

12. Koran, I. I. [Results of a study of the beet nematode Heterodera schachtii Schm.,
according to investigations carried out at the nematode laboratory at Belozerkovsk.
Ukrainian Research Institute for the sugar industry, Kiev] 8(16): 29-67. 1939.
[In Russian with English summary.]

13. Linpner, P. Hin neuer Alchenpilz Rachisia spiralis n.g. n. sp. Deutsche Essig-
industrie 17: 465-468. 1913.

14. Marin, R. Remarques sur le Protascus subiliformis a propos de la communication
de M. E. Maupas. Bull. Soc. Hist. Nat. Afrique du Nord 7: 50-51. 1915.

15. Martin, G. W. Two unusual water molds belonging to the family Lagenidiaceae.
Mycologia 19: 188-190. 1927.

16. Maupas, E. Sur un champignon parasite des Rhabditis. Bull. Soc. Hist. Nat.

l Afrique du Nord 7: 34-49. 1915.

17. Micoterzky, H. Dre frecbelenden Stisswasser- und Moornematoden Ddnemarks
nebst Anhang viber Amébosporidien und andere Parasiten bei freilebenden Nema-
toden. Danske Vid. Selsk. Skrifter Naturw. Math. (ser. 8) 10: 57-310. 1925.

18. Rozsypau, J. Houby na had’ atku fepném Heterodera schachtii Schmidt v morav-
skich piddach (Pilze in Cysten von Heterodera schachtii Schmidt aus mahrischen
Riibenbéden). Véstn. Ceskosl. Akad. Zeméd. (Bull. Acad. Tchécosl. Agr.) 10:
413-422. 1934.

19. Sapepeck, R. Pythium anguillulae aceti nov. spec. Bot. Centralbl. 29: 318-319.
1887.

20. ScumNK, A. Achlyogeton, erne neue Gattung der Mycophyceae. Bot. Zeitung 17:
398-400. 1859.

21. ScuerFret, A. Endophytische Phycomyceten-Parasiten der Bacillariaceen und
einige Monadinen. Arch. Protistenk. 52: 1-141. 1925.

22. Seurat, L. G. Emile Maupas. Bull. Soc. Hist. l’Afrique du Nord 8: 318-323.
1916.

23. SoroKINE, N. Note sur les végétaux parasites des anguillulae. Ann. Sci. Nat. Bot.
(ser. 6) 4: 62-71. 1876.

24. Sparrow, F. K. Inoperculate chytridiaceous organisms collected in the vicinity of
Ithaca, N. Y., with notes on other aquatic fungi. Mycologia 25: 513-535. 1988.

—. A contribution to our knowledge of the aquatic Phycomycetes of Great Britain.
Journ. Linn. Soc. London 50: 417—478. 1986.

26. Tokunaga, T. Notes on the Lagenidiaceae in Japan. Trans. Sapporo Nat. Hist.
Soc. 13: 227-232. 1934.

JUNE 15, 1940 LOHMAN AND CASH: ATROPELLIS FROM PINE CANKERS 200

BOTAN Y.—Atropellis species from pine cankers in the United States.!
M. L. Lonman, Indiana University, and Epiry K. Casu, U. 5.
Bureau of Plant Industry. (Communicated by JOHN A. STEVEN-
son.)

INTRODUCTION

In various areas of the United States certain dermateaceous or
tryblidiaceous fungi occur in association with twig or stem cankers of
one or more species of Pinus, most frequently of species of hard or
pitch pines (subgenus Diploxylon Koehne). These diseases, recently
summarized by Boyce (1), have attracted the attention of pathologists
only during the past 25 years.

The first was reported in 1921 by Weir (11), who described Cenan-
gium piniphilum as a new species associated with stem cankers of
Pinus contorta Loud. and P. ponderosa Lawson in Idaho and Mon-
tana. At that time Overholts (8) had observed cankered twigs of P.
rigida Mill. in Pennsylvania, and later in the same State similar can-
kering of P. pungens Lamb., and considered the fungus associated to
be a species of Crumenula, one possibly identical with C. pinicola
(Rebent.) Karst., which has been reported (5, 7) as pathogenic to
pines in Europe. Subsequently, Zeller and Goodding (13) described
Atropellis pinicola as a new species in association with branch and
stem cankers of several species of pine in the Pacific Northwest. They
allied the fungus with Scleroderris treleasei Sace. of Alaska in a new
genus, Atropellis.

In 1932 specimens of Atropellzs canker collected on Pinus sylvestris
L. near Portsmouth, Ohio, and near Mount Alto, Pa., were re-
ceived by the Division of Forest Pathology, U. 8S. Bureau of Plant
Industry.” The following year an investigation of the disease was un-
dertaken by the Civilian Conservation Corps, in cooperation with the
Forest Service and the Division of Forest Pathology, and from the
field surveys made approximately 500 collections of canker fungi ac-
crued. The writers have had the opportunity to classify these speci-
mens, which are largely from the Eastern and Southeastern States
(2, 3). The present paper is concerned with the taxonomy of speci-
mens representative of the various susceptible hosts in these areas,
studied in comparison with authentic specimens of the canker fungi
in the Western States.

1 Received December 13, 19389.

2 Collections by Dr. Curtis May and Dr. L. W. R. Jackson, respectively; also re-
ported to the Division of Forest Pathology by Prof. J. S. Boyce as occurring on P.
sylvestris and P. resinosa Soland. in Massachusetts and New Hampshire.

256 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

TAXONOMY OF SPECIES ON PINES

With respect to the previously mentioned so-called Cenangium,
Crumenula, and Atropellis cankers in the United States, which are
characterized by a grayish-green to blue-black coloration of the wood
(Fig. 1, c) beneath each lesion (1), the causative fungi seem to be suf-
ficiently closely related to be properly considered within a single
genus. In some features, particularly the roughened exciple and the
elongate, septate spores, the species here referred to Atropellis re-
semble those of Crumenula described as occurring on pines in Europe.
The latter, however, are characterized by Rehm (9) and Lagerberg
(7) as having the apothecium with circular opening, entire margin,
pale hymenium, and long, fine excipular hairs. The American fungi
differ from Crumenula in the furfuraceous exciple, stellate or irregu-
larly lacerate aperture, and blue-black epithecium, as well as in the
type of canker produced and the characteristic discoloration of the
host tissue by the mycelium. In their clavate to elongate-fusoid as-
cospores the species are suggestive of Scleroderris as used by Karsten
(6) and Rehm (9) but are dissimilar with respect to paraphyses, aper-
ture, and exciple. It has been suggested that a possible relationship
exists between these species and S. bacillifera (Karst.) Sacec., as men-
tioned by Zeller and Goodding (13). This fungus, however, as repre-
sented by Karsten’s Tympanis bacillifera (Fung. Fenn. 756, Cent. 8,
1868, in the Farlow Herbarium, Harvard University) on bark of
Picea (‘‘granbark’’) has smooth, patellate apothecia, which conform in
gross and microscopic features to the description by Karsten (6, efr.
Sacecardo, Sylloge Fung. 8: 595) and, except for arrangement of spores
to that by Rehm (9). In the writers’ preparation of F. Fenn. 756
ascospores are 0- to 5-septate, 22-86 by 3-8.5u, subspirally overlap-
ping, with three or four in the apex of the ascus.

It seems advisable, therefore, to recognize Atropellis as a distinct
genus for the species associated with pine cankers in the United
States, with certain revisions of the genus and species as described by
Zeller and Goodding (13, p. 561) for pines.

Atropellis Zeller and Goodding, emend.

Apothecia erumpent, externally furfuraceous and brownish to black,
fleshy coriaceous, single or cespitose, sessile to substipitate, cupulate to
patellate, laciniately dehiscent, hymenium concolorous, bluish black or
lighter; asci clavate, 8-spored; spores fusoid to narrow cylindrical-clavate,
hyaline, continuous, then 1- to 3- (rarely 5-) septate; paraphyses hyaline,
septate, simple or branched, exceeding the asci, end cells equal or slightly
swollen, forming a colored epithecium, agglutinated below and granular

JUNE 15, 1940 LOHMAN AND CASH: ATROPELLIS FROM PINE CANKERS 207

encrusted above by yellowish-brown, pinkish, or violaceous excretion;
hypothecium typically thick, subtended by a hyaline medullary tissue; ectal
layer dense, dark, furfuraceous.

Conidial stromata, where known, of similar habit, texture, and color;
fertile cavities variable, usually coalescing, the pustules dehiscent by one or
more papillulae; conidiophores simple or subverticillately branched; conidia
minute, acrogenous, bacillar, continuous, hyaline.

In each of the following species the pinkish or violaceous material in and
above the hymenial tissue of both conidial and apothecial fructifications is
readily dissolved by sodium or potassium hydroxide solutions, which are im-
mediately colored greenish or bluish. If a hydroxide solution is used as a
swelling agent in microscopic preparations, the cytoplasm of young asci and
ascospores appears greenish rather than hyaline. Overholts (8) describes
this reaction in his account of the fungus associated with the canker of
Pinus pungens, presumably the second of the following species.

Atropellis pinicola Zeller and Goodding Fig. 2, A-B

Zeller and Goodding (13) record A. pinicola on Pinus contorta Loud. and
P. lambertiana Dougl. in Oregon, and on P. monticola D. Don. in Oregon,
Washington, Idaho, and British Columbia. Zeller (12) also reports the
species on P. lambertiana in California and on P. strobus L. in Oregon.

Atropellis pinicola differs from the three following species primarily in its
longer and narrower spores, which Zeller and Goodding, upon the examina-
tion of numerous specimens, record as ‘‘382—-63 X 1.5-3.5u (average 40 X 2u),”
and in its more frequent occurrence on soft pines. Specimens examined by
the writers have spores typical in size and shape, although not uniformly
1-celled as originally described, but frequently 2- to 4-celled, and occasion-
ally 6-celled.? The paraphyses are septate, simple or forked, with the tip
cells equal, or swollen and up to 4u in diameter. The conidial fructifications
and conidia observed conform to the descriptions under the succeeding
species. (Cf. 12, p. 464.)

Atropellis tingens sp. nov. Fig. 1, B, C, F—H; Fig. 2, C—-L

Apotheciis sessilibus vel substipitatibus, ex cortice erumpentibus, sub-
globosis dein patelliformibus, coriaceis, 2-3 mm diam., furfuraceis, atris,
margine laciniato, hymenio atro-coeruleo; ascis clavatis, longe pedicellatis,
octosporis, (70) 90-110 (150) X8-10 (14); ascosporis 2-8-seriatis, anguste
fusiformibus, rectis, curvatis vel sigmoideis, (20) 24-40 (46) x (2) 2.5-8.5
(4)u; paraphysibus simplicibus ramosisve, apice incrustatis, epithecium
atro-coeruleum formantibus; hypothecio brunneo, strato interiore hyalino,
cortice denso, atro, rugoso; fructibus conidicis pustulatis, atris, furfuraceis,
plicatis; conidiophoris ramosis, 35X1.5u; conidiis bacillaribus, continuis,
hyalinis, 4-8 X0.5—0.7u. Hab. in cortice ramorum vivorum Pini.

Apothecia sessile to substipitate, single or gregarious, erumpent through
the bark of cankered areas, often concentrically arranged, subglobose when
young, expanded to cup-shaped or patellate, contorted or compressed when
crowded, opening by irregular splitting, coriaceous, 2-3 mm diam., exterior
black, furfuraceous, margin laciniate, incurved, when dry often hysteroid

3 Specimens examined: Oregon Agr. Coll. Herbarium no. 4877 (courtesy of S. M.
Zeller); nos. 1304, 1635 (ex type), 1679, 1968, and 1969 in the herbarium of J. 8. Boyce,
Yale University; Forest Pathology nos. 86401-86405; Mycological Collections, U.S.
Bureau of Plant Industry, nos. 66565-66567, 71100-71104.

i}
U

opposite page for explanation)

JUNE 15, 1940 LOHMAN AND CASH: ATROPELLIS FROM PINE CANKERS) 259

in form, hymenium blue-black, pruinose. Asci clavate, long-pedicellate,
apex rounded, 8-spored, (70) 90-110 (150) X8—-10 (14)y; spores irregularly
2-3 seriate in the upper part of the ascus, slender fusiform, broadest near
the apex and tapering below, straight, curved or sigmoid, hyaline, continuous
then 1-3 (rarely 4-5) septate, (20) 24-40 (46) X (2) 2.5-3.5 (4.0)u; paraph-
yses sometimes simple, more frequently branched and slightly thickened
at the tips, septate, subhyaline, granular-encrusted, forming a dense
purplish- or bluish-black epithecium. Hypothecial layer 20-25 thick,
brown; underlying tissue of fine, loosely interwoven, hyaline or subhyaline
hyphae, forming a layer 70— 110y thick; cortex 30- 100. thick, very dense,
black, roughened on the outer surface by clumps of thick- walled, brown,
closely septate hyphae. Conidial fructifications pustulate, erumpent, black,
furfuraceous as in the apothecia, irregularly chambered or folded, the folds
lined with branched conidiophores 35 X1.5u; conidia borne at the tips of the
branches, bacillar, continuous, hyaline, 4-8 X0.5-0.7u.

On cankered twigs, branches and small stems of Pinus banksiana Lamb.,
P. caribaea Morelet, P. clausa (Engelm.) Sarg., P. densiflora Sieb. & Zucc.,
P. echinata Mill., P. nigra Arn., P. pinaster Ait., P. pungens Lamb., P. re-
sinosa Soland., P. rigida Mill., P. rzgida var. serotina (Michx.) Loud., P. stro-
bus L. (infrequent), P. taeda L. and P. virginiana Mill., from New Hampshire
to Florida, west to Ohio and Arkansas (2, 3). Type specimen: Dzller 24 on
Pinus rigida, Camp Roosevelt, George Washington National Forest, Va.,
June 12, 1933.4 All of the specimens on Pinus strobus were obtained near
Mount Solon, Va.

The ascospores of this species are regularly shorter and broader than those
of Atropellis pinicola, averaging approximately 30 X3y, but of the same de-
gree of variation. With respect to a single apothecium, spores in general are
less variable in width than in length. When relatively broad ascospores are
encountered, they are of the shape characteristic of the species and average
considerably longer than those of the next succeeding species.

The specimens examined show that apothecia with mature spores may be
found throughout the year, as Zeller and Goodding note for Atropellis
pinicola. Yet much sterile material is encountered among samples from the
surveys and among the writers’ casual collections made during all seasons of
the year. Possible perennial sporulation on living or weakening twigs and
stems is suggested by the arrangement of the apothecia and degrees of de-
velopment on cankers of long standing.

While this species is suggestive of Atropellis pinicola in the shape and ar-
rangement of the ascospores and of the following species in its general oc-
currence on hard, rather than soft, pines, it is clearly intermediate to the
_ two with regard to spore size.

4 The type and other respresentative specimens are deposited in the Mycological
Collections, U.S. Bureau of Plant Industry; portions of the type and several other col-
lections are also in the Farlow Herbarium of Harvard University and the herbaria of
the New York Botanical Garden and the University of Michigan.

Fig. 1.—A—B, Seven-weeks-old cultures of Atropellis on malt agar medium, kept at
room temperature: A, Atropellis arizonica from Pinus ponderosa; B, A. tingens re-
isolated 21 months after Pinus echinata was inoculated with a culture from P. caribaea.
C-H, Atropellis on various pines: C, A. tingens on P. pungens, North Carolina; D, A.
piniphila on P. ponderosa, Idaho; E, A. arizonica on P. ponderosa, Arizona; F, A. tin-
gens on P. sylvestris, Ohio; G, A. tingens on P. taeda, Virginia; H, A. tingens on P. pin-
oe North Carolina. Figures A—-H, X2; all photographie negatives bye OME. ie By:

oubert.

260 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

The species grows very slowly in cultures held at room temperature, pro-
ducing uneven, elevated, black, compacted or loosely stromatic surface mats
of irregular outline and with scattered and marginal, fine grayish to viola-
ceous tomentum on potato-dextrose and malt agar media, and less erumpent
colonies with black subsurface mycelioid margins on corn meal agar. Identi-
cal colonies were obtained from plantings of infected wood of various species
of pines. The conidial stage developed in irregular, convoluted areas, was
observed in isolations from Pinus banksiana Lamb., P. caribaea Morelet,
P. echinata Mill., P. pinaster Ait., and P. pungens Lamb..,

Atropellis piniphila (Weir) comb. nov. Fig. 1, D; Fig. 2, M

As mentioned above Weir (11) reports this fungus as Cenangium pini-
philum on Pinus contorta and P. ponderosa in Montana and Idaho. More

F
D Fs G

iy) OUR

Fig. 2.—Ascospores of four species of Atropellis; 500. A, A. pinicola on Pinus
monticola, Oregon; B, same on P. contorta, Oregon; C—L, A. tingens on various pines in
the Eastern States (Pinus virginiana, taeda, pungens, caribaea, sylvestris, banksiana,
echinata, rigida, resinosa, and strobus, respectively); M, A. piniphila on Pinus contorta
(type specimen); N, A. arzzonica on Pinus ponderosa.

J K L

recent accounts (1, 12) indicate its further occurrence in the Pacific Coast
States on these hosts and on P. albicaulis Engelm., and (4) in Arizona and
New Mexico on P. ponderosa. Specimens examined in the present study add
other hosts and extend the range of the species to New Mexico and stations
in the Southeast.> The species occurs on both soft and hard pines but in-
frequently on the former.

The apothecia are as characterized for the genus, with the hymenium
brownish to black, averaging somewhat larger than those of the two preced-
ing species. The asci are longer and broader, measuring (100) 120-135
(170) X 10-15, and the tendency for the spores to be clumped in the upper
end also obtains in this species, with the upper two to six spores biseriate
overlapping. Paraphyses are septate, simple or branched, with the tip cells

5 Specimens examined: Weir 2631 (locality unknown: probably Dakotas), on Pinus
banksiana Lamb.; Weir 2624 and 2625 (Idaho), 2626 (Montana), 2627 (Washington),
and Forest Pathology 14973 (type) and 15532 (Idaho), all on P. contorta Loud; Weir
2633 (Oregon), P. jeffreyi ‘‘Oreg. Com.”’; Weir 2628 (Idaho), F. P. 89367 Gill & Ellis
(New Mexico), P. ponderosa Lawson; Weir 2630 (Alabama), P. taeda L.—all in Myco-
logical Collections, U.S. Bureau of Plant Industry; Boyce 766 (ex-type), and 1659
(Idaho), P. monticola D. Don.—Herbarium of J. 8S. Boyce; Diller 311 (Tennessee),
P. virginiana Mill.—courtesy of J. D. Diller.

———

= 7 =

JUNE 15, 1940 LOHMAN AND CASH: ATROPELLIS FROM PINE CANKERS 261

equal or swollen. The hymenium in section shows the purplish tints char-
acteristic of species of this group. Ascospores are elliptical-fusoid, with ends
rather acute, straight or subsigmoid, and measure (14) 16-22 (24) X (3.5)
4-5 (5.5), averaging 18 X 5u in the type specimen. While Weir describes and
illustrates the spores as 1-celled, 1- and 2-celled spores occur in half of the
specimens listed herein, including the type. The conidial stage observed in
association with apothecia (Boyce number 1659) is morphologically identical
with that of A. tengens.

Sydow and Petrak (10) determine this fungus as Cenangium farinaceum
(Pers.) Rehm, which occurs on branches of Pinus sylvestris in Europe,
making particular note of the coloration in the hymenium (“‘Paraphysen’’)
and epithecium. Rehm (9) notes that in this respect C. farznaceum differs
from other species of Cenangium. The accounts, however, do not indicate
that the European fungus is pathogenic or that it discolors the wood. On the
basis of Rehm’s description C. farinacewm has smaller asci and shorter and
broader, 1-celled spores, which have obtuse rather than acute ends. What-
ever the relationship between these fungi may be, the American specimens
can not be identified as Peziza pinicola beta caespitosa of Fries (Syst. Mye.
2: 113; Peziza farinacea Pers., Syn. Fung., p. 672), which in the specimen
distributed by Fries (‘‘294. Peziza farinacea b. Syst. Mye.”’ in the Farlow
Herbarium, Harvard University), shows numerous flattened, smooth, black
and shining apothecia up to 0.6 mm broad when dry, occurring singly or
clustered on a large sheet of bark sufficient to indicate that fruiting was not
associated with a canker condition. Therefore, the synonymy that Sydow
and Petrak suggest is considered untenable.

Atropellis arizonica sp. nov. Fig. 1, A, E; Fig. 2, N

Apotheciis substipitatibus, ex cortice erumpentibus, subglobosis dein
expansis, coriaceis, atris, 1.2-2.5 mm diam., furfuraceis, margine undulato,
lacinato, incurvato, hymenio pruinoso, atro-coeruleo; ascis cylindrico-
clavatis, apice obtusis (90) 100-120 X9-14u; ascosporis 2-3-seriatis, el-
liptico-fusoideis, rectis vel subsigmoideis, continuis vel 1-3-septatis, (20)
24-28 (32)xX(4) 4.5-6 (7)u; paraphysibus filamentosis, septatis, ramosis;
hypothecio brunneo; strato interiore hyalino, cortice crasso, atro, rugoso;
fructibus conidicis pustulatis, nigris; conidiophoris ramosis, 12-20 X2u;
conidiis bacillaribus, hyalinis, continuis, 4-6 X<1-1.5u. Hab. in cortice
ramorum vivorum Pini.

Apothecia substipitate, erumpent singly or in groups of two to three, scat-
tered over bark of cankered areas of the host, subglobose when young, then
expanded and patellate, sometimes irregularly contorted and compressed
when crowded, leathery, black, 1.2-2.5 mm diam., exterior furfuraceous,
margin undulate, laciniate, incurved when dry, hymenium pruinose, blue-
black. Asci cylindrical-clavate, obtuse at the apex, gradually attenuated
toward the base, 8-spored, (90) 100-120 X9-—14y; spores irregularly 2-3-
seriate in the upper part of the ascus, 1-2-seriate below, elliptic-fusoid,
straight or subsigmoid, tapering at both ends, continuous to 1-3-septate,
equal or slightly constricted at the central septum, (20) 24-28 (82) x (4)
4.5-6 (7)u; paraphyses filamentous, septate, branched above, the tip cells
equal or swollen, dilute pinkish in mass, covered by a dense blue-black
epithecium. Hypothecium brown, 45—65u thick; inner layer of tissue 100-—
150u thick, of loosely interwoven, subhyaline hyphae; cortex 100—250u
thick, black, dense, exterior roughened and covered with loose clumps of

262 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

dark, closely septate hyphae. Conidial fruits pustulate, black, furfuraceous
as in the apothecia; conidiophores simple or branched, pale violaceous in
mass, 12-20 X2u; conidia bacillar, hyaline, continuous, 4—6 X 1—1.5n.

On stems of Pinus ponderosa Lawson, Hospital Flat, Safford, Ariz.,
September 22, 1934, D. J. Stouffer.

This species is known only from the locality of the type specimen, which
was obtained at an elevation of 9,000 feet. It is closely allied with A.
piniphila but has longer and broader spores, in their form suggestive of cer-
tain species of Crumenula on pines in Hurope. In cultural aspects the species
resembles A. tingens. The conidial stage was obtained in cultures from iso-
lated groups of ascospores discharged upon plates of nutrient agar.

SUMMARY

The genus Atropellis is revised to include along with A. pinicola
Zeller and Goodding, of the Pacific Northwest, two new species,
namely, A. tingens and A. arizonica, and Cenangium piniphilum Weir,
all of which cause characteristic cankers of pines. Of these A. tengens
is of widest occurrence, on various species of native and introduced
pines in the eastern half of the United States, and A. arizonica is
most limited in distribution, being known only from the type local-
ity, Safford, Arizona. Each of the species produces a localized, dark
stain of the wood in cankered twigs or stems.

LITERATURE CITED

. Boycs, J. S. Forest pathology, 600 pp., illus. New York, 1938.

. Dinter, J. D. The Atropellis canker of eastern pines. Service Letter, Pennsyl-

vania Dept. Forests & Waters, ser. 5, no. 46, pp. [2-3]. 1934.

The Atropellis canker of eastern pines. Plant Dis. Report. 19: 17. 1935.

ELLIS, D. E. Conifer diseases hitherto unreported from the Southwest. Plant Dis.

Report. DSioak. 1939)

. JARSTAD, I. Innberetning fra statsmykolog Ivar J¢rstad om co DpSUMaeTN ne: pa
skogtraerne 1 drene 1926-1930. Beretning om det Norske Skogwesen for 1930,
Skogdirektgren, Oslo, pp. 78-96. 1931. Rev. Appl. Myc. 11: 1386-137. _ 1932.

6. Karsten, P. A. Mycologia Fennica. Pars prima, Discomycetes. Bidr. Finl. Nat.

Folk 19: 1-264. 1871.

7. LaGERBERG, T. Studier 6fver den norrlindska tallens sjukdomer, sdrskildt med

hdnsyn till des foryngring. Medd. Stat. Skogsférséksanstalt 9: 135-170, illus.

1912

oo RO Ne

8. OvERHOLTS, L. O. Mycological notes for 1925. Mycologia 18: 179-184, illus.
1926.

9. Reum, H. Ascomyceten: Hysteriaceen und Discomyceten in Rabenhorst’s Krypto-
gamen-Fl., ed. 2, Bd. 1, Abt. 3, 1270 pp., illus. 1896.

10. Sypow, H. and PETRAK, 'F. Ein Beitrag zur Kenntnis der Pilzflora Nordamertkas,
insbesondere der nordwestlichen Staaten. Ann. Myc. 20: 178-218. 1922.

11. Weir, J. R. Cenangium piniphilum n. sp., an undescribed canker-forming fungus
on Pinus ponderosa and P. contorta. Phytopath. 11: 294-296, illus. 1921.

12. Zevurr, 8S. M. Some miscellaneous fungi of the Pacific Northwest. Mycologia 27:
449-466, illus. 19385.

13. — and Gooppina, L. N. Some species of Atropellis and Scleroderris on cont-
fers in the Pacific Northwest. Phytopath. 20: 555-567, illus. 1930.

JUNE 15, 1940 CHAPIN: NEW LADY-BEETLES 263

ENTOMOLOGY .—WNew genera and species of lady-beetles related to
Serangium Blackburn (Coleoptera: Coccinellidae).1 Epwarp A.
CHAPIN, U.S. National Museum.

This paper has been prepared primarily to supply names for cer-
tain species of coccinellid beetles of economic importance. In the
course of the work dissections of representatives of a great many
genera were made, and it became evident that in many places the ac-
cepted classification of the family is faulty. More attention must be
paid to the structures of the head and abdomen before a system can
be outlined that will be justified by the morphology of the species in-
volved. The division of the family into its major groups will, I believe,
be based mainly on the structure of the mandible. Such a system
was suggested in 1843 by Redtenbacher.?

Thanks are due to Miss Hazel Bowen, who has prepared the figures
illustrating the essential characters of the genera discussed.

The genera and species considered in this paper form a very. com-
pact group within the family Coccinellidae. This group is the Oeneini
of Casey, 1899, minus the genus Oenezs Mulsant, 1850. As the species
placed in Oenezs must be associated with Cryptognatha Mulsant, 1850,
and not with Delphastus Casey, 1899, and as the name Oeneis is not
available for use in the Coccinellidae because of its prior use in Lepi-
doptera by Hubner, the tribal name Oeneini must be abandoned. It
seems undesirable at present to propose a new name in view of our
lack of exact knowledge of the structure of most of the small Coc-
cinellidae.

It is advisable here to discuss certain generic segregates that have
been presumed to be related to this group:

Lioscymnus Champion, 1913, is determined, after an examination of a
paratype of the type species, to be a synonym of Delphastus.

Delphastobia Casey, 1924, is determined, after examination of the type

species and specimen, to be a chrysomelid related to or belonging to the
genus Lamprosoma.

Delphastopsis Casey, 1924, is determined, after examination of the type
species and specimen, to be close to and possibly a member of the genus
Scymnillus.

Semichnoodes Weise, 1892, is, according to notes by Sicard, 1909, not re-
lated to Serangium.

Serangium monticola Sicard, 1909, is, according to the original description
and figure, not related to Serangium.

The species belonging to this group have the following characters in com-

1 Published by permission of the Secretary of the Smithsonian Institution. Re-
ceived January 15, 1940.

> Tentamen dispositionis generum et speciorum coleopterorum..., 32 pp., Vido-
bonae, 1843. Reprinted in Germar’s Zeitsch. Ent. 5: 113-132. 1844

264 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

mon: Form very compact; head convex in front; antennae with the terminal
segment spatulate and very long in comparison with other segments;
pronotum finely margined throughout; prosternum strongly developed an-
teriorly as a plate protecting the mouth parts when head is in position of
repose, notched on each side near lateral margins for the reception of the
antennae; elytral epipleura moderately broad, with cavities for the reception
of the tips of the middle and posterior femora; legs received in deep cavities
on under side of body, those for the front legs being on the prosternum, for
the middle legs on the suture separating the meso- and metasternum and for
the hind legs mostly on the first abdominal sternite; femora broad and flat,
completely concealing the tibiae when legs are retracted; abdomen with five
visible sternites, the second, third, and fourth being quite short in compari-
son with the first and fifth.

After the elimination of the genera mentioned above, there remain two
described genera having the characteristics listed in the preceding paragraph
and two as yet unnamed. These four may be separated in the following
manner:

1. Antennae 8-segmented; size large (length 2—2.5 mm); middle and hind
tibiae not angulate externally; Oriental region..... Catana, new genus
Antennae 9-segmented; size small to medium (upto2mm)............ 2

2. Middle and hind tibiae slender, not angulate externally; size medium
(1.5-2 mm); Oriental, Ethiopian, and Australian regions...........

PUP aU AR Mat ale Wicca als (OREO ae Arie eC MRL FU Ne era Serangium Blackburn
Middle and hind tibiae angulate externally... 2 ee 3

3. Third segment of antennae obtriangular, twice as wide at apex as fourth
segment; tibiae without teeth at apex of angulation; size small (about
ia) Ceylon i,k Steere fees chee eee aah ee Serangiella, new genus
Third segment of antennae cylindrical, not much wider at any point
than fourth; tibiae with usually two denticles at angulation of tibiae;

size small to medium (0.8-1.5 mm); New World... .Delphastus Casey

Delphastus Casey, 1899

While it is not possible at present for me to give a key to all the species, it
is desirable to give names to two species that are frequently submitted for
identification. The essential characters of the type species of the genus, Del-
phastus pusillus (LeConte), are illustrated in Figs. 1-4.

Delphastus nebulosus, n. sp.

Length 10 mm, width 0.8 mm, altitude 0.5 mm. Pale yellow-brown, legs
whitish, elytra each with a single basal castaneous spot of indefinite extent.

Integuments shining. Head not visibly punctured except for a very few minute

setigerous punctures. Scutellum very small, not visibly punctured. Elytra
without visible punctures, but with pellucid dots grouped along the suture
and lateral margin and on apical portion. Two or three setae occur on each
elytron in the region of the humeral callus, and afew very short setae along
the lateral margin toward the apex; otherwise the elytra are glabrous. The
dark spot extends along base from near scutellum to beyond humeral callus
and backward to just beyond basal third. The margin of the spot is not
sharp but the area covered is fairly constant in the series of specimens
examined. Under parts not visibly punctured except for the apical abdominal

Figs. 1-4.—Delphastus pusillus (LeConte): 1, Mandible; 2, maxillary palpus;
3, hind leg; 4, antenna. Figs. 5-8.—Catana clauseni, n. sp. Figs. 9-12.—Catana
parcesetosa (Sicard). Figs. 18-16.—Catana spilota (Weise). Figs. 17—20.—Serangium
maculigerum Blackburn. Figs. 21—24.—WSerangiella flavescens (Motschulsky).

I!

266 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 6

sternite, which is finely and densely punctured and conspicuously pubescent:
Femora shining, impunctate. Middle and hind tibiae with two small denticu-
lations on outer margin at apex of angulation.

Type and 17 paratypes.—U.S.N.M. no. 53941.

Type locality—Puerto Rico. Type and seven paratypes from Villalba,
November 15, 1932, R. G. Oakley (San Juan no. 3195); six paratypes from
Rio Piedros, March 19, 1913, T. H. Jones (P.R.S.G.A.Acc. 274-13); one
paratype from same locality, January 25, 1914 (P.R.S.G.A.Ace. 16-14); one
paratype from Barceloneta, March 22, 1932 (San Juan no. 2069); one from
Santurce, April 1, 1932 (San Juan no. 2121); and one from Ponce, March 13,
1933, R. G. Oakley (San Juan no. 3796).

Remarks.—D. nebulosus is most closely related to D. pallidus (LeConte)
but is distinguished from that species by the spot on the elytron and by the
smooth and polished side pieces of the metathorax (in D. pallidus sclerites are
finely alutaceous). While D. pallidus is known from Cuba as well as from
Florida, it is not known from Puerto Rico, and D. nebulosus is known from
Puerto Rico only.

Delphastus collaris, n. sp.

Length 1.3 mm, width 1.0 mm, altitude 0.5 mm. Deep piceous; male with
head, entire prothorax, and legs pale yellowish brown, female with head,
under part of prothorax, and legs somewhat paler. Integuments shining.
Head not very convex, sparsely and irregularly set with a mixture of very
fine and moderately coarse punctures, some of which are setigerous. Pronotum
slightly more coarsely punctured, very sparsely hairy, anterior angles bent
down outside the fine marginal line, which cuts across the angle and which
is very fine anteriorly. Scutellum with only two or three punctures. Elytra
distinctly and rather densely though not coarsely punctured, sparsely set
with erect hairs. Metasternum finely and rather sparsely punctured, the
side pieces finely alutaceous. First abdominal sternite punctured as meta-
sternum, second sternite with a complete, third and fourth sternites with an
incomplete, row of punctures, fifth sternite finely and sparsely punctured.
Femora not visibly punctured. Denticles on tibiae elongated into crests as in
D. pusillus (LeConte).

Type and 37 paratypes.—U.S.N.M. no. 53940.

Type locality —Paraiso, Canal Zone. Type and 28 paratypes taken Janu-
ary to March 1911 by E. A. Schwarz; nine paratypes taken March to April
1911 by A. H. Jennings. In addition to the type material I have referred to
this species two specimens from Porto Bello, Panama, February 15-24, 1911,
EK. A. Schwarz; one specimen from Panama City, April 30, 1911, A. H. Jen-
nings; and five specimens from El Cermeno, April-May 1938, J. Zetek
(Zetek no. 4401).

Remarks.—tThis species is easily separated from all species of Delphastus
known to me by the strong punctures on the second to fourth sternites and
in the male by the entirely pale prothorax.

Catana, n. gen.

Serangium auct. nec Blackburn, 1889, p. 187.

Coccinellidae of nearly hemispherical form, always slightly longer than
greatest width, altitude greater than half the transverse diameter (the three
dimensions approaching the ratio 8:7:4). Head convex, eyes relatively
coarsely faceted, very slightly emarginate adjacent to antennal sockets,
without canthus; distance between inner margins of antennal sockets equal

JUNE 15, 1940 CHAPIN: NEW LADY-BEETLES 267

to three-fourths distance between eyes; antennal sockets moderately large,
with beaded margin, the marginal bead continuous around the clypeus but
finer across anterior margin of clypeus; clypeus transverse anteriorly with
rounded angles; antenna 8-segmented, first segment stout, second globular,
third elongate, subcylindrical, as long or slightly longer than fourth to
seventh combined, fourth longer than fifth, sometimes showing the beginning
of a division and therefore morphologically equal to fourth and fifth com-
bined, fifth to seventh short and subequal, eighth long, broad and thin,
setigerous, longer than second to seventh combined; mandible with slender
apical portion somewhat curved, prostheca conspicuous; maxillary palpus
with terminal segment not strongly inflated. Pronotum short, strongly
transverse, anterior angles acute and prominent, posterior angles acute,
marginal line complete, basal margin slightly sinuate near scutellum. Pro-
sternum strongly developed and subtruncate anteriorly, lateral portion
deeply notched for reception of antennae, the median portion strongly
bicarinate, the carinae strongly divergent anteriorly. Posterolateral portions
deeply excavate for the reception of the front legs. Mesosternum very
short. Metasternum long. All coxae widely separated, the hind coxae es-
pecially so. Femora broad, tibiae rather slender, tarsi with four distinct seg-
ments, the third about one-third length of fourth. Claws broadly toothed at
base. Scutellum small, triangular. Elytra with broad, somewhat concave
epipleura, which are impressed deeply for reception of apices of femora.
Abdomen of five visible sternites, first and fifth long, second to fourth very
short. See Figs. 5-16.

Genotype.—Catana clauseni, n. sp.

In addition to the type species, I refer the following previously de-
scribed species to Catana: Serangium parcesetosum Sicard, 1929, from India,
and Serangium spilotum Weise, 19138, from Luzon, Philippines.

These three species may be distinguished in the following manner:

meUmitonm pale yellow-brown..:...0:..........0 4.2: parcesetosa (Sicard)
Upper parts deep piceous, elytron with a large castaneous spot....... 2

2. Pronotum rather densely hairy at sides; spot on elytron elliptical......
5 oo. pach enna ee ame Re a ee sae es AST LOE (WEISE)
Pronotum sparsely but evenly hairy; spot on elytron reniform........
PM ee VeA IT een PANE eile CRVOMEES ott ine tek shhh. clausent, N. sp.

Catana clauseni, n. sp. Figs. 5-8

Cryptognatha sp. Clausen and Berry, 1932, pp. 36—40.

Length 2.3 mm; width 2 mm; altitude 1.14 mm. Head, legs, under parts,
and a large reniform spot on each elytron bright castaneous, pronotum,
scutellum, and rest of elytron deep piceous. Integuments shining. Front of
head minutely and sparsely punctulate, vertex slightly more densely so, a
fine hair arising from each puncture. Pronotum more coarsely punctured
than head, the punctures sparsely and irregularly set, each furnished with
a fine and moderately long hair. Scutellum with a very few fine punctures.
Elytra very minutely and sparsely punctulate on disks, slightly more
coarsely so on apical third and each with a regular row of fine punctures
that is parallel to and near lateral margin, each puncture of the row bearing
a short seta. The castaneous reniform spot on each elytron is large, com-
mencing near scutellum and reaching to about apical fourth, its concave
margin toward the suture. The spots are variable in size, but in no specimen
do they coalesce either anteriorly or posteriorly. Under parts shining and
minutely and sparsely punctulate except for the side pieces of the metaster-

i
i

268 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

num, which are strongly alutaceous, and for the apical abdominal sternite,
which is rather densely punctured and furnished with rather long hairs.
Femora finely and rather densely punctulate, hairy, strongly margined on
basal half of posterior border. Tibiae slender.

Type and 23 paratypes.—U.S.N.M. no. 53942.

Type localityw—Medan, Sumatra. Other localities: Kuala Lampur,
Federated Malay States, and Cuba in the West Indies where the species has
been introduced.

Remarks.—The material before me consists of 24 specimens from Medan,
seven from Kuala Lampur, both lots collected by C. P. Clausen, and 27
specimens from Cuba, five from Santiago de las Vegas, August 20, 1934,
A. R. Otero (H.E.A. Cuba, Ent. no. 10350), and 22 from Camito, Province
Habana, May 17, 1938, P. A. Berry. As the species was introduced into
Cuba in 1930, the 1938 collection would indicate that it is firmly established
on the island. Recently the species has been planted at Nassau, Bahamas,
but no recoveries have so far been made from that locality.

Catana spilota (Weise) Figs. 13-16

Serangium spilotum Weise, 1913, p. 241.

The national collection contains a series of 23 specimens collected in May
1911 on Negros Island, Philippines, by C. V. Piper. Although the type lo-
cality of this species is Luzon, I can see no reason to doubt the specific
identity of the Negros specimens.

Catana parcesetosa (Sicard) Figs. 9-12

Serangium parcesetosum Sicard, 1929, p. 184.

Sixty specimens of this species from India are in the collection. Most of
these are labeled ‘‘Panjab and U Provinces, VI-X, India, R. 8. Woglum”’;
some carry the additional data “‘predaceous on Aleyrodidae, Saharanpur,
India, R. S. Woglum,”’ and two were taken at Mangalore by J. C. Bridwell.
Except for a slight variation in size, the specimens are remarkably uniform.

Serangium Blackburn, 1889

Several species of this genus were studied in connection with the species
now placed in Catana and those that appear to be new are described in the
following pages. For convenience, a key to the new ones, together with two
well-known Australian species, is given. For illustration of the salient char-
acters, Serangium maculigerum Blackburn was selected, and Figs. 17—20
refer to this species.

Grandi (1914) made a very careful study of an African species, Serangizuwm
giffard: Grandi, and gave a detailed description with excellent figures of the
adult, larva, and pupa. With this evidence there can be no doubt concerning
the occurrence of the genus in Africa. It is unfortunate that Grandi was not
permitted to dissect a specimen of Semichnoodes kunowt Weise and so dem-
onstrate the true affinities of that genus.

1. Elytra densely hairy; front femur alutaceous; side pieces of metasternum
roughly sculptured; abdominal sternites strongly punctured.........

A Gece ae Coes. ona Pee ee ee eer oonorosiie Jsllackklourrin
Elytra very sparsely hairy, usually with a few setae at base and along
TATE UNS i. 36) Diet Stan « WOM ee eae a eee aca 2

JUNE 15, 1940 CHAPIN: NEW LADY-BEETLES 269

BAO Se nmunmralUibaceCOUSe .os2) ty) ne eed yet Pet ths Pd be eae ee 3
EOsrermMimmrs mnine. HdistimMeE tye pUMetUTeds ie.) 22 ae ee... 4

3. Second to fourth abdominal sternites finely strigillose ; metasternum with-
out a triangular alutaceous area at side...... maculigerum Blackburn
Second to fourth sternites smooth and polished; metasternum with lateral
MAMeUAr AlUvaACeOUS AFA. . 0. .5 ey eo metasternalis, n. sp.

4. Outer face of front femur shining, not evidently alutaceous, pronotum
STMEMIPSI [OR EG RE goa ae tart ed) oa eee em See ae comperet, Ni. Sp.
Outer face of front femur strongly alutaceous; pronotum at most with
MC MMLONCRAMN CICS iat ee ena ee eee eae cis eke ekg 5

5. Abdominal sternites highly polished, without punctures except along
ROCA mana Came Cte tt Mr wey Nee CRM. 3 yh Viele poets. bakeri, n. sp.
Abdominal sternites very sparsely but distinctly punctured except fifth,
which is rather densely punctured at middle..................... 6

6. Elytra noticeably and rather strongly punctured..... japonicum, Nn. sp.
Elytral punctures visible only with high magnification...............

Serangium japonicum, n. sp.

Length 2 mm; width 1.5 mm; altitude 0.9 mm. Deep piceous, head, flanks
of pronotum narrowly, and legs pale castaneous. Integuments shining. Head
with a few distinct punctures more or less definitely grouped in two patches,
one on each side of the median line, sparsely hairy. Pronotum finely, sparsely
and very irregularly punctured. The punctures near the median line, which
is narrowly impunctate, somewhat coarser than those near the lateral mar-
gin, surface sparsely hairy. Scutellum occasionally with one or two punctures,
usually impunctate. Elytra moderately coarsely, densely, and evenly punc-
tured, except that the punctures near the apical angles are somewhat coarser
than the others. Elytra without vestiture except for a submarginal row of
short setae. Epipleura finely punctured and sparsely hairy. Prosternum
somewhat rough, meso- and metasternum polished with a few fine punctures,
side pieces of metasternum alutaceous. Abdominal sternites sparsely punc-
tured except last, which is densely punctured and finely pubescent. Front
femora strongly alutaceous, middle and hind femora very sparsely punctured
and finely hairy.

Type and 15 paratypes.—U.S.N.M. no. 53943.

Type locality —Japan. Type and seven paratypes from Yokohama, where
they were found feeding on Ceroplastes rubens by S. I. Kuwana; three
paratypes labeled “Japan. Koebele’’ (Koebele Note no. 12638); and five
paratypes from Nagasaki, taken May 20, 1922, feeding on aleurodid eggs
ly 22 Ishi.

In addition to the type material, I ascribe to this species a single specimen
ao Nanking, Kiangsu Province, China, taken August 15, 1919, by H. F.

oomis.

Serangium comperei, n. sp.

Length 1.7 mm; width 1.3 mm; altitude 1.1 mm. Pale yellow-brown,
pronotum sometimes infuscate at base (female?), elytra piceous except for
narrow basal, humeral, and apical margins, which are yellow-brown. In-
teguments shining. Head finely and very sparsely punctured, the punctures
bearing fine, moderately strong setae. Pronotum sparsely and irregularly
punctured, the punctures of two sizes, the larger ones bearing fine setae.
Scutellum not visibly punctured. Elytra without visible punctures except

270 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

for a very few in the scutellar region, and one or two setigerous punctures
near humeral callus. Epipleura finely and sparsely hairy. Prosternum finely
and moderately densely punctured and hairy, meso- and metasternum and
abdominal sternites one to four shining, without visible punctures, fifth
(apical) sternite shining and impunctate laterally, finely and densely punc-
tured and hairy at middle. Femora shining and not visibly punctured.

Type and four paratypes.—U.S.N.M. no. 53944.

Type locality.—Fiji Islands. A series of five specimens was taken at the
type locality by George Compere in October or November 1899.

Serangium bakeri, n. sp.

Length 1.4 mm; width 1.1 mm; altitude 0.6 mm. Under parts, head, and
sometimes portion of pronotum castaneous, legs slightly paler than sternites,
upper parts generally deep piceous. Head very finely and sparsely punctured.
Pronotum with a few setigerous punctures, mostly on disk. Scutellum not
visibly punctured. Elytra without visible punctures, with a few long setae
near scutellum, humeral callus, along basal half of suture, and on lateral
margin near apex. E/pipleura almost impunctate, apparently not setose.
Prosternum very finely and sparsely punctured, mesosternum finely punc-
tate and rather densely hairy, metasternum with setigerous punctures ad-
jacent to mesosternum at middle and on the median portion, side pieces of
metasternum together with the narrow side margins of the metasternum
proper strongly alutaceous, abdominal sternites highly polished, without
punctures except for a few along apical margin. Front femora alutaceous,
middle and hind femora finely punctured and sparsely hairy.

Type and paratype.-—U.S.N.M. no. 53945.

Type locality—Luzon, Philippine Islands. The type and paratype were
taken on Mount Makiling by C. F. Baker. A third specimen that appears
to belong to this species was taken at Zamboanga, Mindanao, P. I., by
Baker.

Serangium luzonicum, n. sp.

Length 2.2 mm; width 1.9 mm; altitude 0.6 mm. Dark castaneous, head
and under parts somewhat paler. Head finely, irregularly, and rather
sparsely punctured, punctures poorly defined, mostly setigerous. Pronotum
with similar though slightly coarser punctures, which are largely wanting
along a narrow median longitudinal line, setae sparse and not long, rather
more densely set in anterior angles. Scutellum without visible punctures.
Elytra distinctly and rather evenly punctured, the punctures very poorly
defined, setigerous punctures present across base and in a row parallel to
the lateral margin. Epipleura sparsely set with short setae. Prosternum shin-
ing, moderately densely punctured and hairy. Mesosternum very short,
shining, without visible punctures. Metasternum proper shining, very
sparsely set with very fine punctures, which bear short setae, side pieces
strongly alutaceous, sparsely hary. Abdominal sternites shining, first four
very sparsely set with setigerous punctures, fifth rather densely punctured
and hairy. Front femora strongly alutaceous, other femora shining, the hind
pair moderately densely hairy.

Type and four paratypes.—U.8.N.M. no. 53946.

Type locality —Luzon, Philippine Islands. Type and one paratype col-
lected at Manila by W. Schultze, two paratypes taken at the same place by
George Compere, and one paratype from Bacoor (9 miles south of Manila)
taken by P. L. Stangl.

JUNE 15, 1940 CHAPIN: NEW LADY-BEETLES PA AL

Serangium metasternalis, n. sp.

Length 1.7 mm; width 1.4 mm; altitude 0.6 mm. Dark castaneous to
piceous, head and legs paler. Head very finely punctured, the punctures more
densely placed toward the clypeal region, where the vestiture of fine hair is
rather dense. Pronotum with the punctures coarser than those on head and
gathered into two moderately dense patches on each side of the narrow
smooth median line. Vestiture sparse and fine. Scutellum not visibly punc-
tured. Elytra sparsely punctured, the punctures visible only under high
magnification, without setae except near humeral callus and along lateral
margin. Epipleura not visibly punctured. Prosternum evenly and rather
strongly alutaceous, very sparsely hairy. Mesosternum smooth at middle,
with a small patch of setae at side. Metasternum proper smooth at middle
and sparsely and finely punctured; laterally there is a triangular area ad-
jacent to the side piece that, like the side piece itself, is strongly alutaceous.
Abdominal sternites one to four finely and sparsely punctured, fifth sternite
much more densely punctured and moderately densely hairy. Front femora
strongly alutaceous, middle and hind femora smooth and sparsely hairy.

Type and paratype.—U.S.N.M. no. 53947.

Type locality.—Luzon, Philippine Islands. The type and paratype are
from Manila, collected by George Compere. With these I have associated a
third specimen from Biliran Island, collected by C. F. Baker.

Serangiella, n. gen.

Oeneis Motschulsky, 1866, nec Mulsant, 1850.
Cryptognatha Crotch, 1874, nec Mulsant, 1850.

Coccinellidae similar in form to Serangium. Head convex, eyes relatively
coarsely faceted, shghtly emarginate adjacent to antennal sockets, without
canthus; distance between inner margins of antennal sockets equal to one-
half distance between eyes; antennae sockets strongly transverse, with
finely beaded margin, which is apparently not continuous across anterior
margin of clypeus; clypeus very short and strongly transverse: antennae
9-segmented, first segment stout, second stout and but little more than half
length of first, third strongly obtriangular, a little longer than second and
not so long as fourth to eighth combined, these segments becoming succes-
sively wider until the eighth, which is almost as wide as the third, ninth
segment almost as long as third to eighth combined, width equal to more
than half its length; mandible very small, roughly quadrate, the inner or
cutting margin feebly curved, the outer margin acutely angulate just before
middle of length, prostheca apparently not present; terminal segment of
maxillary palpus rather strongly inflated, subtruncate apically. Pronotum
short, strongly transverse, anterior angles slightly prominent and strongly
rounded, posterior angles acute, marginal line complete though very fine
anteriorly. Prosternum strongly developed, truncate anteriorly, deeply
notched laterally for reception of antennae, median portion strongly bi-
carinate, the carinae strongly divergent anteriorly. Mesosternum extremely
short. Metasternum long and broad, all coxae widely separated, the hind
coxae especially so. Femora broad, middle and hind tibia conspicuously
angulate externally beyond middle, tarsi with four distinct segments, the
third half as long as but of same diameter as fourth. Claws thickened at base
but without distinct tooth. Scutellum small, triangular. Elytra with rather
narrow somewhat concave epipleura, impressed for reception of apices of
femora. Abdomen of five visible sternites, as in Catana.

Genotype.—Oeneis flavescens Motschulsky, 1866.

a

272 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 6

Although I have been able to examine only the one species, I feel reason-
ably certain that the other two Ceylonese species described by Motschulsky
(1866, p. 423), Oenezs laterale and O. nigritulum, also should be referred to
this genus. In fact, it is not evident from the original description that the
three species are truly distinct. Figs. 21-24.

LITERATURE CITED

BuacKBURN, T. Further notes on Australian Coleoptera, with descriptions of new species.
Trans. Royal Soc. South Australia 11: 187, 209. 1889.

CasEy, T. L. A revision of the American Coccinellidae. Journ. New York Ent. Soc.
7: 71-169. 1899.

Addition to the known Coleoptera of North America. Memoirs Coleopt. 11: 155—
176. 1924.

Cuampion, G. C. Notes on various Central American Coleoptera, with description of new
genera and species. Trans. Ent. Soc. London, 1918, pp. 120-128. 1913.

CLAUSEN, C. P., and Berry, P. A. The citrus blackfly in Asia, and the importation of
its natural enemies into tropical America. U.S. Dept. Agr. Techn. Bull. 320:
36-42. 19382.

Crotcu, G. R. A revision of the coleopterous family Coccinellidae, pp. 206-208.
London, 1874.

GRANDI, G. Descrizione di un nuovo Coccinellide africano. Boll. Lab. Zool. Portici
8: 165-178, figs. 1-8. 1914.

MotscHutsky, V. DE. Essai d’un catalogue des insectes de Vile Ceylan. Bull. Soc.
Hist. Nat. Moscow 39: 422-426. 1866.

Mutsant, E. Species des coléoptéres triméres sécurtpalpes, pp. 496-501. Paris and
Lyons, 1850.

StcarD, A. Revision des coccinellides de la faune malgache. Ann. Soc. Ent. France 78:
150-155. 1909.

Descriptions de quelques especes nouvelles de coccinellides de la faune Indo-
Malaise. Ann. Mag. Nat. Hist. (10) 3: 179-184. 1929.

Weisk, J. Kleine Bettrdge zur Coccinelliden. Fauna Ost-Afrika’s. Deutsche ent.
Zeitsch., 1892, pp. 15-16. 1892.

———Uber Chrysomeliden und Coccinelliden der Philippinen. II Teil (Coleoptera.)
Philippine Journ. Sci. 8D: 241-242. 19138.

ENTOMOLOGY—Siphonaptera from Western United States.! InvING
Fox, Iowa State College, Ames, Iowa. (Communicated by
C. F. W. MvuESEBECK.)

The following descriptions, involving three new species and a new
subgenus, are based upon material in the United States National
Museum and in the author’s private collection. Type material is in
the United States National Museum, to whose authorities the author
expresses his appreciation for the privilege of studying the collections
in their charge. Particular thanks are due to Dr. H. E. Ewing, of
the Bureau of Entomology and Plant Quarantine, for his kindness
and assistance.

Family HYSTRICHOPSYLLIDAE

Corypsylloides Hubbard
Corypsylloides Hubbard, Pacific Univ. Bull. 37: 7. 1940.
Front reduced; frontal tubercle and notch absent. Gena much enlarged;

genal ectenidium obliquely vertical consisting of six spines, the lowermost
reduced in width. Eyes absent. Labial palpus 5-segmented, the last segment

1 Received February 2, 1940.

JUNE 15, 1940 FOX: SIPHONAPTERA FROM WESTERN UNITED STATES 273

tapering distally. Pronotum rounded cephalad, well separated from the
head, bearing on the posterior margin a ctenidium consisting of spines of
various sizes. Mesopleuron not divided by a vertical rodlike sclerotization
into a mesoepisternum and a mesepimeron. Line of division between mete-
pisterna and metepimeron not distinct. Abdominal tergites armed with
apical spines but no heavy transverse incrassations. Each abdominal tergite
with one row of four bristles, two situated laterally and two dorsally. In
male the posterior arm of sternite 1X absent; sternite VIII not modified;
manubrium laterad to rest of the clasper rather than ventrad to it. Coxa III
with a patch of spinelets on the inside. Last segment of each tarsus with
four pairs of lateral plantar bristles.

This genus is most closely allied to Corypsylla C. Fox from which it may
be readily separated by the absence of heavy transverse incrassations on
the abdominal tergites and by the character of the genal ctenidium. It is
easily distinguished from Nearctopsylla Rothchild, which it superficially
resembles, by the structure of the thorax.

Corypsylloides spinata, n. sp. Figs. 1, 2, 4

Male.—Preantennal region of head armed with a frontal row of four
bristles and a small bristle at the base of first, third, fourth, and fifth genal
spines; several other small bristles located along dorsal margin of head.
Spines of genal ctenidium arranged as in Fig. 1. Postantennal region armed
with three bristles in addition to a marginal row of about five. Labial palpus
barely reaching to apex of fore coxa. Pronotum with a medial row of but
three long bristles and a ctenidium of about 15 spines on a side. Mesopleuron
with four bristles, two in center and two at posterior margin; metepisternum
with two or three bristles, of which one is long and one or two are minute;
metepimeron with four bristles. One antepygidial bristle present on a side.

Modified segments—Process of clasper and movable finger as in Fig. 2.
Spring short, not completing a single turn. Total length, 1.8 mm. Greatest
depth of abdomen, 0.5 mm.

Female.—A large bristle located on gena near third antennal segment;
otherwise, chaetotaxy of head as in male. Pronotal ctenidium consisting of
about 13 spines on a side. Two antepygidial bristles present on a side.
Receptaculum seminis and sternite VII as in Fig. 4. Total length, 1.9 mm.
Greatest depth of abdomen, 0.6 mm.

Type host and type locality— Meadow mouse, Microtus townsendii, at
Portland, Oreg.

Type slide—U.S.N.M. no. 54011.

Type materral—Male holotype and female allotype collected by 8S. G.
Jewett from Microtus townsendii, December 24, 1931, at Portland, Oreg.;
in the United States National Museum. Four male and two female para-
types bearing the same data in the author’s private collection.

Family DOLICHOPSYLLIDAE
Amphipsylla neotomae, n. sp. Fig. 3

Male.—Frontal tubercle indistinct. Preantennal region of head armed
with an ocular row of three bristles, which lies below two bristles located
near antennal groove. Postantennal region with a single bristle in addition
to a marginal row of about seven. Eye vestigial, the vestiges not pigmented.
Labial palpus extending to about three-fourths the length of fore coxa.

) 6

Fig. 1.—Corypsylloides spinata, n. sp., head of male. Fig. 2.—idem, process of
clasper and movable finger. Fig. 3.—Amphipsylla neotomae, n. sp., process of clasper
and movable finger. Fig. 4.—Corypsylloides spinata, n. sp., receptaculum seminis
and sternite VII. Fig. 5.—Dactylopsylla (Spicata) rara, n. sp., process of clasper and
movable finger. Fig. 6.—zdem, sternite VIII of male.

JUNE 15, 1940 FOX: SIPHONAPTERA FROM WESTERN UNITED STATES 275

Pronotal ctenidium with about 13 spines on a side. One long antepygidial

bristle between two minute ones present on a side.

Modified segments.—Process of clasper and movable finger as in Fig. 3;
the latter armed on posterior margin with three prominent spiniform bristles
in addition to several others. Spring short, not completing a single turn.
Total length, 3.4 mm. Greatest depth of abdomen, 1.1 mm.

Type host and type locality—Wood rat, Neotoma desertorum, at Dos
Palos, Calif.

Type slide.—U.S.N.M. no. 54012.

_ Type material—Male holotype collected by H. 8. Gentry from Neotoma
desertorum, March 1934, at Dos Palos, Calif.;in U. 8. National Museum.
But two members of this genus have heretofore been reported from North

America, A. sibirica pollionis Rothschild from Alberta and A. ewingz I. Fox

from Alaska. The above described new species may be separated from both

of these by the structure of the male genitalia. It represents the first record
of the occurrence of the genus Amphipsylla within the United States.

Genus DAcTYLOPSYLLA Jordan
Spicata, n. subgen.

Frontal tubercle prominent, not acuminate. Eye vestigial, the vestiges
indistinct. Pronotum armed with a ctenidium of long slender spines. In
male sternite VIII broad and prominent, armed on the posterior margin
with a number of bristles of which some are robust and heavily pigmented
(Fig. 6). Anterior arm of sternite IX much narrower than in Dactylopsylla
(Dactylopsylla). Manubrium not short and triangular, tapering and curved
distally.

Type species.—Dactylopsylla (Spicata) rara, n. sp.

This new subgenus may be readily distinguished from Dactylopsylla
(Dactylopsylla) and from Foxella Wagner by the structure of the male geni-
talia. In the type subgenus sternite VIII is large but not armed with heavy
pigmented bristles. In Fowella this sternite is small and bears but a single
long bristle near the apex. The manubrium of the new subgenus, being nar-
row and tapering distally, is markedly different from both Forella and
Dactylopsylla (Dactylopsylla) where this structure is short, broad, and
triangular.

Dactylopsylla (Spicata) rara, n. sp. Figs. 5, 6

Male.—Preantennal region of head armed with two rows of bristles; the
upper row consisting of about five, the lower row of four. Eye vestigial, the
vestiges exceedingly small and inconspicuous. Postantennal region of head
armed with a marginal row of six or seven bristles. Labial palpus extending
beyond basal half of fore coxa. Pronotal ctenidium consisting of about 14
spines on a side.

Modified segments—Movable finger and process as in Fig. 5; the former
differing in structure on the two sides. In one case the movable finger is
armed with one bristle on the posterior margin (Fig. 5), while in the other
it is armed with three bristles and there are slight differences in the shape
of the structure. Posterior arm of sternite LX with a number of small bristles
distally. Sternite VIII broad, wider distally and armed on posterior margin
with five or six heavily pigmented bristles and about a dozen much more
slender ones. Total length, 4.56 mm. Greatest depth of abdomen, 1.5 mm.

276 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 6

Type host and type locality.x—Pocket gopher, Thomomys fossor, Jackson
County, Colo.

Type sliide.—U.S.N.M. no. 54013.

Type material.—Male holotype collected by S. C. McCampbell from
Thomomys fossor in Jackson County, Colo., July 13, 1926; in the United
States National Museum.

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pure magnesium clay, 2. G. Novena. ee

PHYTOCHEMISTRY.—Some new cyanogenetic anil —&
R. R. Brigse, and J. F. COUCH... 6... nes

GroLocy.—New formation names to be used i in the Kings
Ivanpah aes California. D. F. Hewert.. ;

M. L. LonMtan and Eprra K. Chane ee

Seance Blackburn Fane Coceinllidae).
Cae a ee

ENTOMOLOGY. Fie ie a from Westen United: States
FOR ee

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JOURNAL

OF THE
WASHINGTON ACADEMY OF SCIENCES
Vou. 30 Juty 15, 1940 INOS 7

CHEMISTRY.—The separation of isotopes by chemical means.'
Haroup C. Urry, Columbia University. (Communicated by
Frep L. MouuEr.)

Ten years ago no successful separation or even extensive concentra-
tion of the isotopes of any of the elements had been made. Methods
for the separation of isotopes that had been suggested and investi-
gated depended upon differences in the physical properties of isotopes
and isotopic compounds which were readily predictable from simple
kinetic theory, as, for example, separation by diffusion through porous
tubes and by unidirectional evaporation. Today several methods are
available for the concentration of isotopes, and the isotopes of a few
elements have been separated quite completely. It is my purpose here
to review the present status of this problem, and particularly to em-
phasize the part of the problem with which my colleagues and I have
been most closely associated.

In 1931 two successful attempts at the partial separation of iso-
topes were made. The first of these, by Keesom and van Dijk,’ dealt
with the separation of the isotopes of neon by the fractional distilla-
tion of liquid neon. The second of these was the concentration of the
rare isotope of hydrogen by Brickwedde, Murphy, and myself,’ in
connection with our experiments leading to the discovery of this iso-
tope. These distillation methods could be predicted from current
theories in regard to the solid state and were independently predicted
by these two groups of authors. Recent work by Keesom, van Dijk,
and Haantjes‘ has produced rather high concentrations of the neon
isotopes in substantial amounts, and Scott and Brickwedde*® have
prepared hydrogen deuteride, HD, in nearly a pure form by the frac-
tional distillation of a mixture of hydrogen, deuterium, and hydrogen
deuteride.

* The Tenth Joseph Henry Lecture of the Philosophical Society of Washington,
delivered on March 16, 1940. Received April 19, 1940.

> Kexsom, W. H., and van Disx, H. Proc. Acad. Sci. Amsterdam 34: 42. 1931.

° Urey, H. C., Brickweppz, F. G., and Murpuy, G. M. Phys. Rev. 40:1. 1932.

* KeEsom, W. H., van Dix, H., and Haanrsus, J. Proc. Acad. Sci. Amsterdam
37: 615. 1934; Physica 1:1109. 1934.

* Scott, R. B., and Brrckweppz, F.G. Phys. Rev. 48: 483. 1935.

277

JUL 16 19%

3S) SS Sip

278 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

Differences in vapor pressures of deuterium and protium oxides, .
the O' and O'8 waters, and the N!* and N’ ammonias, as well as sim-
ilar differences in the cases of many of the isotopic compounds of
hydrogen, are now known and have been used to concentrate the iso-
topes of oxygen and hydrogen by fractional distillation methods.
These differences in vapor pressures are small, and effective separation
can be made only by very efficient fractionation columns.

The first complete separation of isotopes was accomplished by means
of the electrolytic separation of the hydrogen isotopes, discovered by
Washburn.® This method of separation is so effective and so easily
carried out, and the demand for the heavy isotope of hydrogen so gen-
eral, that deuterium and its compounds are now produced commer-
cially by this method.

In 1932 Hertz’ developed his cascade diffusion method, making use
of a series of diffusers arranged in a cascade, so that the light fractions
diffusing through the porous tubes, or through a moving stream of
mercury vapor, were fed to a preceding unit of the cascade, while the
heavy fractions were fed to the following unit of the cascade. In this
way Hertz prepared nearly pure neon isotopes and also succeeded in
preparing very pure samples of deuterium. Others have used this
method to concentrate other isotopic substances.

In 1935, Urey and Greiff* presented a method for the separation of
isotopes by chemical means, depending upon the slight differences in
the chemical properties of isotopic compounds. This method has since
been developed and has led to the separation of nitrogen and carbon
isotopes in substantial amounts.

Recently Clusius and Dickel® discovered the thermal diffusion
method, which is applicable to many gaseous substances. The method
has many interesting features and undoubtedly can be used widely
for the separation of small amounts of rarer isotopes.

Recently Beams! has made some orienting experiments on the cen-
trifugal fractionation column method. These have not gone so far as
to produce separated isotopes, but they have demonstrated the feasi-
bility of the method.

Smith and Scott" have investigated the possibility of using oscillat-
ing electric fields acting upon streams of ions as a means of separating
isotopes. The difficulty of mass spectrographic methods and these al-

6 WASHBURN, E. W., and Urry, H.C. Proc. Nat. Acad. Sci. 18: 496. 1932.
7 Hertz, G. Zeit. Phys. 79: 108, 700. 19382.

8 Urny, H. C., and Greirr, L. J. Journ. Amer. Chem. Soc. 57: 321. 1935.

®° Cuusius, K., and DicxeL, G. Naturwissenschaften 26: 546. 1938.

10 Beams, J. W., and Skarstrom, C. Phys. Rev. 56: 266. 1939.

4 SmiruH, L. P., and Scort, G. W., Jr. Phys. Rey. 55: 946. 1939.

Juty 15, 1940 UREY: SEPARATION OF ISOTOPES 279

- ternating field methods resides in the low intensity of beams of ions.
Owing to the repulsion between the ions, the beams spread rapidly,
and sharp separation can not be secured if large amounts of the
separated isotopes are desired. Smith has found that it is possible to
arrange an apparatus so that electrons are present in the beam with
the ions, thus preventing the spread of the beam.

With all these methods available, or their development actively in
progress, it appears now that the separation of the isotopes of the ele-
ments in amounts necessary for adequate physical, chemical, and bio-
chemical research is only a matter of time. The past 10 years have
brought the problem from a position where no separation had been
secured to a point where the entire discussion is now based upon the
best methods. The methods that my colleagues and I have developed,
namely, the chemical exchange methods and distillation methods, ap-
pear to be those most suitable for large production of separated iso-
topes in the limited number of cases where they are applicable. It is
the purpose of this lecture to discuss principally these chemical meth-
ods of separation for the elements carbon, nitrogen, and sulphur, and
to compare this method with the others mentioned above.

THEORY OF THE SEPARATION OF ISOTOPES BY
FRACTIONATION COLUMNS

The most common method for the separation and purification of
substances used in chemistry is that of distillation, making use of
fractionation columns. As mentioned before, this method was used
for the concentration of the hydrogen and neon isotopes where the
differences in vapor pressures could be readily predicted by the use
of the Debye theory of the solid state. Early in the investigation of the
properties of hydrogen and deuterium compounds, Lewis and Cor-
nish” and Washburn, Smith, and Frandsen™ observed that there were
differences in the vapor pressures of hydrogen oxide and deuterium
oxide, and it was also observed that there were slight differences in the
vapor pressures of the O'* and O18 waters.

The theory of distillation apparatus has been discussed in standard
textbooks for many years, and little new can be added. However,
there are certain points to be emphasized when one comes to the frac-
tional distillation of substances with only slight differences in vapor
pressures. The simple process fractionation factor may be defined as
the ratio of the two constituents in the vapor divided by this ratio in

12 Lewis, G. N., and Cornisu, R. E. Journ. Amer. Chem. Soc. 55: 2616. 1933.
13 WASHBURN, HE. W., Smitu, E. R., and Franpsen, M. Bur. Standards Journ.
Res. 11: 453. 1988.

280 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

the liquid phase. Letting N and n represent the mole fraction of one
constituent in the liquid and the vapor, respectively, this fractiona-
tion factor a may be defined by the equation

n N
—/ =e" Cy

The fractionation secured by a column is then given, according to rea-
sonable theory (see below) as well as experiment, by the formula

N; Ny
petals v7 piles = One (2)
1—WN, 1— Nz

where N, and N, are the mole fractions of one constituent at the top
and bottom, respectively, a is the simple process fractionation factor
defined above, z is the length of the column and k is a constant. The
quantity kz is referred to as the number of theoretical plates. In the
case of the plate columns, common for many distillation processes in
the chemical industry, the plates are actual physical plates, although
each may not change the ratio of the two constituents by the simple
process factor. In the case of packed columns, which have been de-
veloped in recent years in considerable variety, no physical plates are
present in the column but the fractionation follows this exponential
form, as determined by experiment. It is of considerable interest to de-
rive expressions by which the time required for equilibrium to be es-
tablished can be calculated at least approximately. Such a calculation
was made by Huffman and Urey™ for the case of total reflux with a
large reservoir at one end of the column. The formula secured by them
is

1 NG) ENA = ee
a

Nios Sa a

a alle ie H
where H is the holdup of the column per unit length, NV and N> are the
mole fractions of one constituent at the reflux end at time t, and in the
natural substance, respectively, and L is the rate of flow through the
column. This formula shows that in order to secure a rapid equilibri-
um, a small holdup, large natural abundance, rapid rate of flow, and
a fractionation factor that deviates markedlyfrom unity, are required.
In the distillation of isotopic substances and their separation by
chemical exchange methods, however, the value of the simple process
factor is very nearly unity, and the time required to reach a steady
state may be very long indeed unless a column of proper design is

144 HUrrMAN, J. R., and Urny, H.C, Ind. Bung: Chem. 29:53) 19378

JuLy 15, 1940 UREY: SEPARATION OF ISOTOPES 281

secured. This means that columns with small holdup, rapid rate of
flow, and a large number of theoretical plates per unit length are re-
quired. In most industrial processes, fractionation columns that have
from 10 to 50 theoretical plates are quite adequate. However, if ade-
quate separation is to be secured for isotopic molecules, a very large
number of theoretical plates, e.g., from 300 to 500, is required. This
large number is necessary because the ratios of vapor pressures of
isotopic substances deviate only slightly from unity.

The transport of such apparatus is, of course, equal to the differ-
ence in the flow of one isotopic substance in the two directions. If the
mole fraction of the isotope to be concentrated is small, the transport
at the beginning of the operation of such a column, using total reflux,
is given by the equation

ft == Ibi g(@ = 1) (4)

where L is the total flow in either direction and N, the mole fraction
of the rare isotope in the mixture. This formula is approximate and
applies only when the fractionation column is not near the steady
State. :

A more exact theory of the operation of such columns can be given
if one assumes that back diffusion is not an important process in the
column, and that the rate of transport of material from the one phase
to the other depends upon the concentrations in the two phases in the
way required for the equilibrium process. Two differential equations
can be secured for the rate of change of concentration in the two
phases. These are:

aN aN

H — — L— — k,N(l — n) + kiintl — N) =0 (5)
ot 02
On On

h—-+1 —4+k,N(l — n) — k_in(l — N) = 0 (6)
ot Oz

where L, H, N andJh,n are the rate of flow, the holdup, and the mole
fraction in the liquid and gas phase, respectively, and ki and k_1 are
the velocity constants for the transport of this isotope from one phase
to the other. The solutions of these two simultaneous equations ina
form suitable for convenient calculation have not been made. How-
ever, if we are interested only in the final steady state, so that the
derivatives with respect to the time are zero, the solution of the equa-
tions is simple. The first integral is

EN — In = no(L — 1) (7)

282 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

This equation states that the net flow in the column is equal to the
amount of material withdrawn at one end of the column. The second
integral leads to the following relation between the forward flow and
the composition of the material produced:

1—2N,+ 4
y +0 = C+ Hy) tanh } — ofl + Hy) + tan (<=) (8)
, Late OU

where == Ay, ) = » and o = 3lnF).

angie 9.4
In these equations 7 is the mole fraction of the isotope in the for-
ward flow and /) is the overall fractionation factor when the forward
flow is zero. By making use of these equations it is possible to show
how the effective fractionation factor for the column changes with the
rate at which material is withdrawn. Our apparatus seems to give bet-
ter results than the simple theory would indicate.

In the case of the fractional distillation of the hydrogen isotopes,
the values of a are in the neighborhood of from 3 to 10, depending
upon the temperature and whether one is working with H, and HD or
H, and D,». The distillation of the hydrogens is therefore a problem
like that of the separation of ordinary substances, except that the
fractionation must take place at a very low temperature. In the case
of the fractionation of the hydrogen and deuterium oxides, however,
or of the O' and O!° waters, the ratios of the vapor pressures are near-

TABLE 1.—RaATIO OF VAPOR PRESSURES OF WATERS

Le p(H:201*) /(pH2018) p(H20)/p(D20)
Bute} 1.188
125 1.0138

20.0 1.148
237.0 1.0088

35.6 1.008?

40.0 Weiltes
46.35 1.0077

80.0 1.070

100.0 1.003 (extrapolated) 1.052

ly equal to unity, as shown in Table 1. In this case it may require long
periods of time for the fractionation column to come to a steady state.

CONCENTRATION OF OXYGEN ISOTOPES

Huffman and Urey“ made a substantial concentration of the O18
isotope by means of distillation methods. Water was pumped in at the
top of a fractionation column at a fixed rate, evaporated completely

Juuy 15, 1940 UREY: SEPARATION OF ISOTOPES 283

at the bottom, and the vapor returned upward through the column.
At the top, the vapor was discarded since it was not convenient to
condense the water for re-use. In this way water containing 0.85 per
cent O!8 was produced, i.e., with somewhat over four times the nat-
ural concentration of O18. Also, considerable amounts of water con-
taining half the normal abundance of O!* were produced by boiling a
large container of water at the bottom of the column and totally re-
fluxing at the top. The operation of the column in this case was not
steady and this probably accounts for the smaller change in the ratio
secured.

It does not seem probable that distillation methods can be very
useful in other cases for the separation of isotopic substances. The
N"™ and N® ammonias have but slight differences in their vapor pres-
sures, as is shown by Table 2. The great difficulty in carrying out a
distillation at the temperature of boiling ammonia, using extensive

TABLE 2.—VAPOR PRESSURES OF THE AMMONIAS

‘ pN“H, pN”H, A 4
r*k (em of Hg) (em of Hg) p(N“Hs)/p(N™Hs)
15 5.488 5.459 1.0053
204 .4 9.060 9.018 1.0046
PAT all 10.904 10.858 1.0042
208.8 12.593 12.541 1.0041
PAT 7 21.570 21.488 1.0038
228.2 41.030 40.899 1.0032
228.4 41.550 41.418 1.0032
234.9 59.560 59.399 1.0027
239.4 75.340 75.153 1.0025

fractionation columns, makes this an unsatisfactory way for the frac-
tionation of isotopes. A study of the vapor pressures of the hydrogen
and deuterium oxides shows that there is a marked difference of vapor
pressure even at the critical temperature of water, i.e., as the tempera-
ture increases, differences in vapor pressures do not necessarily at
once become small. The same may be true in the case of the ammo-
nias, and the question deserves further study to determine whether a
fractionation column at ordinary temperatures, operating under pres-
sure, might not produce a considerable fractionation of the nitrogen
isotopes. A similar behavior might be observed in other cases, but as
yet our knowledge of this subject is incomplete.

CHEMICAL EXCHANGE REACTIONS

The discovery of the isotope of hydrogen and the very extensive
studies that have been made of the differences in chemical properties

284 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

of hydrogen, deuterium, and their compounds show that the chemical
properties of isotopic compounds are not identical, and that slight but
definite differences may be expected. The first calculation of such dif-
ferences in chemical properties, made by Urey and Rittenberg” on the
hydrogen-hydrogen iodide exchange reaction, showed again that
marked differences could be expected. Subsequent calculations and
experiments by many workers show that even larger differences in the
chemical properties are present in many other cases, as, for example, in
the exchange reaction between hydrogen and water. This work nat-
urally led to a calculation of the equilibrium constants for exchange
reactions involving other isotopes. The results of such calculations
are given in Table 3. These calculations are from Urey and Greiff? ©

TABLE 3.—EQUILIBRIUM CoNnsTANTS OF GASEOUS ExcHANGE REACTIONS

4 Equilibrium constant (a)
Reaction “998.1 °
a5Oo'® + HoO!? = FSOs!®-- MeO oa ee ee 1.028
+CO,!6 + H,O!}8 = +CO,}8 +H,O16 Ono 'o G0 00 0060.0 04 6-0 O00 1 6 054
40,!6 +H,0!18 => 40,18 +H,01'6 eoonoeoD OOO DOG DOD Oo OOOO 1 . 020
CEOz CLO = CZO CEO an a note ee eee 1.086
PON ope lal GR KO) Ge SIU CIOS oe, eee A ei eels bb oo 1.003
PU DIO CO ale sieeve lab eyes sbi bs bidle 5 lgie ule o Ble 1.0004
FING CN Os ING ee NG Oh ete are ee nee eae een 1.015
Uy 10s ee ig a hs Fs pera eA MR Ra Mie, hlbont 1.025
ClO, + 4C],37 = Cl370, + 4C]l,35 do Goo HbRODD ODDO DG ROO 0 1.023
ClO, +HCl7 = C70, +HCl3 canodooooop oD oOo e oo OOO ] 5 027

with the exception of the last two equilibria recorded, which were
made by Dwayne T. Vier. These fractionation factors, a, deviate from
unity by as much as 9 per cent in one case. The table shows that some-
what larger deviations from unity are obtained in the case of elements
of low atomic weight than in the case of those of high atomic weight,
but that even for elements as heavy as chlorine, a considerable frac-
tionation may be expected. The calculations given in these tables are
made by the use of well-established statistical mechanical theory, us-
ing experimental molecular spectra data for abundant molecules. The
necessary vibration frequencies and moments of inertia for the rarer
molecules are calculated by the use of the theory of small vibrations
apphed to molecules of this kind.

Exchange reactions involving two phases must be considered if
countercurrent methods similar to distillation, using a fractionation
column, are to be used. In the case of the liquid-gas phase equilibria
the examples listed in Table 4 may be considered. The calculated val-

15 Urey, H. C., and Ritrensere, D. Journ. Chem. Phys. 1: 137. 1933; 2: 106.
1934.

JuLy 15, 1940 UREY: SEPARATION OF ISOTOPES 285

TABLE 4.—EQUILIBRIUM CONSTANTS OF Two PHASE EXCHANGE REACTIONS

Equilibrium Equilibrium
Reaction constant constant
calculated observed
N&H;3(g) +N"“H,*(aq) = N“H3(g) + N!°5H.* (aq). 1.033 1023
HC®N (g) +C8N-(aq) =HCN(g) + C2N-(aq). 1.026 1.013
HCN#4(g) +CN (aq) =HCN#*(g) +CN" (aq). 1.003 Slightly less than 1
S#4O0.(g¢) +HS?0O; (aq) =S*2O2(g) +HS*O;3 (aq). — Sil

ues of the equilibrium constants, which are also the values of the frac-
tionation factors, a, have been secured by assuming that the distri-
bution functions for the ions in solution are the same as those to be
expected for a gas. Such calculations can not be exact, but experiment
does show rather good agreement with the calculated values recorded
in the second column of the table. The observed values depend upon
our experiments made on the separation of the isotopes of carbon,
nitrogen, and sulphur, and depend upon the rate of production of the
heavy isotopes, assuming that equation (4) applies. Since our separa-
tion apparatus is run under conditions where this formula may give
too low a value for the fractionation factor, and since experiments of
this kind never run perfectly, the observed constants as given deviate
less from unity than actually is the case. This is due to the fact that
the transport is proportional to the deviation from unity of this con-
stant, and any interruption of the process is likely to decrease the
yield, and can not increase the yield under any circumstances. The ta-
ble shows that the values calculated in this way are at least a good
approximation to the experimental values.

Exchange reactions involving two liquid phases have been used by
Lewis and MacDonald" to secure a partial separation of the lithium
isotopes. The exchange reaction involved is given as the first reaction
in Table 5. Exchange reactions between solid and liquid phase sys-
tems are illustrated by the last three equations of Table 5. In these
cases it is impossible to make satisfactory calculations of the equilib-
rium constants, and only the constant observed by Taylor and Urey"
in one case, namely, the lithium ion-lithium zeolite exchange reaction
is recorded. In the potassium ion-potassium zeolite and the ammo-
nium ion-ammonium zeolite reactions, only the direction of the devia-
tion of the constant from unity can be given, and it seems difficult to
deduce from the results of Lewis and MacDonald any value for the
equilibrium constant for their exchange reaction.

aan Lewis, G. N., and MacDonaup, R. T. Journ. Amer. Chem. Soc. 58: 2519.

17 Tayior, T. 1., and Urgy, H.C. Journ. Chem. Phys. 6: 429. 1988.

286 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

TABLE 5.—EQUILIBRIUM CONSTANTS OF ZEOLITE EXCHANGE REACTIONS

Reaction! Equilibrium constant

observed
Li? (amalgam) +Li6t =Li® (amalgam) +Li7*............ Sl
Bsc SI Grou Wh ele Cn Ae sneer le pe NIE Cn ke tas RE ak Oe 1022
TRS SERS KGL RS Za tidy eens cue ante arias oat re eaad a <1
NYE => NYE = NPE EIN es et ee : el

1 The zeolite radical is indicated by the symbol Z.

Such two phase exchange reactions can be used for the separation
of isotopes, and the effect of the simple process fractionation in the
case of gas-liquid equilibria can be multiplied many times by the use
of countercurrent apparatus of the distillation column type. In the
case of liquid-liquid equilibria, countercurrent liquid flow can be used
in a similar way, but in the case of the liquid-solid equilibria, a coun-
tercurrent flow of this kind would be very difficult to maintain. In this
latter case, procedures similar to those of chromatographic analysis
can be used effectively.

The apparatus which has been used by my colleagues and myself for
the fractionation of the nitrogen, carbon, and sulphur isotopes consists
of the packed column type of fractionation column. The time which
is required for such apparatus to come to a steady state may be very
long indeed, and in the experiments which we have conducted years
would have been required in some cases if special types of operation —
had not been employed. The transport of the column is given approxi-
mately by equation (4), and this shows that the transport is propor-
tional to the total flow in the column and to the concentration of the
less abundant material, provided that the mole fraction of this less
abundant material is small. Thus if one fractionation column will in-
crease the concentration by some factor, F, the total flow could be
decreased in the next column by the same amount and still the trans-
port would remain the same. Therefore a second column is made to
operate on the product of the first, the second column having a smaller
total flow and a smaller hold up, and hence the final steady state is
reached in a much shorter time than if this additional fractionation
had been carried out with the column of the same size as the first one.
In this way the steady state is reached much more quickly, with the
same transport. A third still smaller column can now be used to oper-
ate on the product of the second in a similar way.

Fig. 1 shows diagrammatically the arrangement of the apparatus
which we have used for the concentration of the nitrogen isotopes."®
In this case the heavy isotope concentrates in the liquid phase. An

18 THopE, H. G., and Urny, H.C. Journ. Chem. Phys. 7:34. 1989.

Juuy 15, 1940 UREY: SEPARATION OF ISOTOPES 287

ammonium-nitrate solution containing 60 per cent by weight of the
salt was pumped into the top of the first unit. This unit increased the
concentration of N by a factor of about 7. Six-sevenths of the ma-
terial was withdrawn at the bottom, sodium hydroxide added, and
_ the ammonia removed from the solution by boiling and passed back
upward through this unit. One-seventh of the material flowed to Unit
2, which changed the concentration of the nitrogen isotopes by an ad-
ditional factor of 9. About eight-ninths of the material was removed

Fig. 1.—Arrangement of fractionation units (HZ, E’, E’’) in nitrogen apparatus.
Stripping units are indicated by S, S’ and 8’, liquid flow by solid lines (F), and gas
flow by broken lines (V).
at the bottom of this column, sodium hydroxide added, and the am-
monia sent upward through the unit, while one-ninth went to Unit 3,
which further changed the ratio of the nitrogen isotopes by a factor of
11. The ammonia was finally liberated at the bottom, passed upward
through all the units, as indicated by the arrows, and finally absorbed
in water at the top of Unit 1. The arrangement is similar to a distilla-
tion apparatus except that a large reservoir of material is maintained
at the top of the first unit and total reflux at the bottom. The solu-
tion takes the place of the liquid in a distillation apparatus, and the
ammonia gas replaces the vapor in distillation. Rapid exchange takes

288 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

place between the ammonium ion and ammonia just as equilibrium is
maintained between the liquid and the gas in a distillation.

The arrangement used in the carbon case’? is illustrated in Fig. 2.
Only two units were used since we make use of hydrogen cyanide and
not too large quantities could be used in a laboratory building. In this
case, C® concentrates in the gas phase and therefore the general ar-

a ee
NaOH
Absorption bas

:
. i
Capillary

Manormeter
1, iguid Flow

NaOH ‘| Gas Flow

————— Electrical
Connection
Unit |

NacN
dng er
u ;
q Ta Droin

Fig. 2.—Arrangement of fractionation units in the carbon apparatus.

rangements must be inverted as compared with the nitrogen separa-
tion. Solutions of sodium cyanide and sulphuric acid are pumped into
the bottom of Unit 1 and the hydrogen cyanide removed from the
solution by a stripping column. Hydrogen cyanide gas rises through
Unit 1. At the top, sufficient sodium hydroxide (free of iron) is added
to absorb about four-fifths of the hydrogen cyanide. This unit changes
the ratio of the carbon isotopes by a factor of about 5. The remainder

19 Hutcuison, C. A., Stewart, D. W., and Urry, H. C. Journ. Chem. Phys.
In press. )

Juty 15, 1940 UREY: SEPARATION OF ISOTOPES 289

of the hydrogen cyanide gas passes to the bottom of Unit 2, and at the
top of Unit 2 sufficient sodium hydroxide is added to absorb it. This
second unit changes the ratio of the carbon isotopes by approximately
a factor of 6.

Small amounts of air are introduced into the apparatus with the
solutions, and if these are not removed from the top of the second
unit the air will accumulate there and prevent the hydrogen cyanide
from reaching the top of this unit. As time went on the apparatus
would completely fill with air, and no fractionation would be obtained
since all absorption of the hydrogen cyanide would occur at the bot-
tom of the first unit. To avoid this it is necessary to remove the inert
gas at the top of the second unit. This is done by connecting an addi-
tional absorbing unit with the top of Unit 2 by a capillary tube. The
sodium hydroxide is pumped first to the absorbing unit and from the
bottom of this to the top of Unit 2. Part of the HCN is neutralized in
the absorbing unit and the remainder at the top of Unit 2. As the inert
gas accumulates at the top of the absorbing unit, the flow of hydrogen
eyanide through the capillary decreases. A manometer arranged with
an electrical contact is attached to the ends of the capillary so that
the pressure difference measures the rate of flow of HCN. When the
pressure difference falls to zero, a pump is started which removes inert
gas from the top of the absorbing unit. This causes rapid absorption
of HCN in the absorbing unit again, and the pressure drop across the
capillary is restored. In this way inert gas is removed automatically
without difficulty.

The apparatus is controlled by fixing the rates with which alkali
enters the tops of the two units. The rate of supply of hydrogen cya-
nide at the bottom is regulated by maintaining a constant pressure
at this point through a manometer with an electrical contact which
starts and stops the feed pump.

The actual form of the apparatus is illustrated in Fig. 3 for the case
of the nitrogen separation. Only the two smaller units are used for the
carbon separation. A detailed description of this apparatus has been
published elsewhere.!®

Taylor and Urey’ have made use of the liquid-solid equilibria to ef-
fect a change in the ratio of the lithium, potassium, and nitrogen iso-
topes. In these cases a 30-foot tube was filled with a zeolite, say, for
example, a sodium zeolite and covered with a solution of sodium chlo-
ride. Lithium chloride solution was then fed into the top of the tube
and the solution withdrawn at the bottom. Since lithium 6 is concen-
trated in the zeolite rather than in the lithium chloride solution, it is

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JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

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retarded relative to lithium 7 in its movement down the column. After
some time the solution leaving the bottom of the column will contain
lithium ion and this will be enriched in lithium 7; that is, the leading
sample contains the heavier lithium. One can then wash out the lith-
Fig. 3.—Complete apparatus as used in the separation of the nitrogen isotopes.
ium chloride from the zeolite and in this case the trailing sample is
enriched in lithium 6. Fig. 4 represents results secured in such experi-
ments by Taylor. A rather marked change in the relative abundance
of lithium is secured by this simple method. Similar experiments using
potassium and ammonium salts showed that changes in the relative

=)
for)
N

JuLy 15, 1940 UREY: SEPARATION OF ISOTOPES 291

i seal ec I ll (a

40 60 80
PERCENT LITHIUM THROUGH COLUMN

Fig. 4.—Distribution of lithium isotopes coming through a zeolite column.

abundances of the isotopes of potassium and nitrogen result from such
experiments, but that the heavier isotopes are concentrated in the
trailing samples in these cases.

CRITIQUE OF SEPARATION METHODS

In considering the relative advantages of various methods for the
separation of isotopes, several factors must be kept in mind. In the
first place there is the question of the amounts of isotopic materials
that are required for experiments. For effective work most chemical
experiments require amounts of the order of grams of these separated
isotopes. In the second place, the cost of production is an important
item. If one is separating only small amounts as a laboratory curiosity
or for a very few special problems, then the cost becomes relatively
unimportant. However, if one looks upon separated isotopes as a new
tool for chemical research in all its various aspects, then the matter
of production cost on a commercial basis is important. The value of
such stable isotopes as tracers, as compared with the value of radio-
active isotopes, depends upon the amount by which the material can
be diluted before the tracer can no longer be detected, upon the size of
sample that is required for analysis, and upon the convenience of the
analytical method. It is very difficult for me to speak authoritatively
in regard to other methods of separation than these chemical ones,
but in the case of the chemical method it is of interest to record some
facts in regard to the cost, and the usefulness of the separated isotopes
particularly for tracer use.

The rate of production by chemical methods, using apparatus of the
type I have described, is considerably larger than that of any other
type of separation that has been used, so far as I am aware. In the
case of our nitrogen experiments, it was possible to transport 0.75

292 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

gram of N® per 24-hour period, while on the carbon experiment (us-
ing only the two smaller units) we have been able to transport 0.15
gram of C per 24-hour period. There seems to be no reason why these
methods can not be increased almost indefinitely on a commercial
scale. It should be possible to increase the production by at least a fac-
tor of 10 on the basis of present knowledge in both cases. The cost of
production depends greatly upon the scale on which one works. In our .
experiments on nitrogen, using the laboratory size apparatus, we
were able to produce N* for a cost of about $50 per gram of N®, in-
cluding the cost of chemicals and the salaries of the men operating the
apparatus. There seems to be no reason why this cost can not be low-
ered to something in the neighborhood of $15 per gram of N®, or less.
In the case of carbon, our cost for salaries and chemicals amounts to
about $200 per gram, although there appears to be no reason why this
cost can not also be brought to the neighborhood of $15 or $20 per
gram if the production is maintained on a sufficiently large scale.

These figures, of course, can not be regarded as total costs for the
reason that in any commercial venture it is necessary to charge over-
head, which is an important part of the cost of production, but it does
seem probable that these isotopes can be produced and sold for some-
thing in the neighborhood of $50 per gram.

The dilution factor of the material we are now producing, 25 per-
cent C, is approximately 2500 to 1, if we assume that the precision of
a mass spectrometer is limited to 1 percent of the natural abundance
of C®. If mass spectrometers can be devised which are able to detect
0.1 percent of the natural abundance of C™, and this does not seem
impossible, the dilution factor for our present material is 25,000.

The sample required for satisfactory analysis with a mass spectrom-
eter is in the neighborhood of 1 ce of gas at atmospheric pressure,
and if necessary this could quite easily be reduced to one-fifth of this,
or approximately 0.1 milligram of C. The method of analysis requires
a rather extensive and somewhat expensive piece of apparatus, and
one that at the present time is often difficult to keep in satisfactory
adjustment. There appears to be no reason, however, why such an
apparatus can not be made commercially in a much more convenient
form than those now in use in our laboratories, and at a cost com-
parable to medium sized spectrographs.

Distillation, chemical exchange reactions, thermal diffusion, and
centrifugal evaporation all depend upon the possibility of using vola-
tile compounds. The relative advantages of the thermal diffusion and
the chemical methods can be rather well evaluated at the present

Juty 15, 1940 UREY: SEPARATION OF ISOTOPES 293

time. The chemical methods rapidly become ineffective as atomic
weights increase. For example, it would appear that fractionation fac-
tors in the neighborhood of 1.0002 might be expected in the case of
elements of atomic weight 200 or more, instead of 1.02 as in the case
of carbon and nitrogen. It is thus evident that it will probably be im-
possible to effect any separation of isotopes on an effective scale by
chemical methods for elements above 40 in atomic weight. On the
other hand, the thermal diffusion coefficient can be expected to be
about 0.1 as great for elements of atomic weight 200 or more, as for
carbon. Thus we see that its use over a wider range of the periodic
system is possible, although the speed of the method per unit appara-
tus is only about 5 or 10 milligrams of C® per 24-hour period. To at-
tain a production of 1 gram of C® per 24 hours would require at least
100 thermal diffusion tubes, while the chemical separation can be
effected on the same scale with a single apparatus of somewhat more
extensive character than those already used. While the thermal dif-
fusion method can be easily operated without attention, chemical
methods require constant attendance. The comparison for a case such
as carbon comes to a matter of cost of production. Electrical current is
particularly expensive for the thermal diffusion method, probably
running in the neighborhood of $100 per gram of C® transported.
Other methods of heating can hardly be expected to have the uni-
formity and the steady character required for the efficient operation
of the process. It seems quite certain that the construction and opera-
tion of 100 thermal diffusion tubes can not be expected to be as eco-
nomical as the operation of a single chemical separation apparatus,
even though the chemical method must have constant attention. More-
over, if larger amounts are needed the chemical method can be ex-
tended to larger scale apparatus by methods that are well understood
in the chemical industry.

Turning to the possible separation of the elements of high atomic
weight, we see that the centrifugal fractionation column is the most
feasible method. It should have a transport per unit comparable to
chemical exchange units of approximately the same size. Moreover,
the effectiveness of separation should decrease very slowly with in-
creased atomic weight, the decrease being due to slower rates of dif-
fusion in the case of high atomic weight substances, and not due to a
decrease in the effective fractionation factor.

It is always interesting to speculate on where the next important
development of a subject will take place. At the present time the
separation of the uranium isotopes would probably lead to most in-

i

294 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

teresting progress in the study of the fission process of the uranium
isotope of mass 235. Thermal diffusion seems to be a possible method
for accomplishing this, although no very great speed can be expected.
It would seem to me that the centrifugal fractionation column or
some modification of this would be a most likely method for securing
results on this most fascinating problem, and in my opinion this is the
most important direction for research on the concentration of iso-
topes at the present time.

I have been greatly aided in the work that I have reported in this
lecture by my colleagues who have worked with me for various peri-
ods of time since this work was started about five years ago. Prof.
John R. Huffman, now at New York University, aided with the dis-
tillation experiments on the O' and O18 waters and with the early
experiments on the separation of the nitrogen isotopes. Prof. Harry
G. Thode, now at McMasters University, did much of the work on
the final separation of the nitrogen isotopes and aided with the first
experiments on C. Prof. Clyde A. Hutchinson, now at the Univer-
sity of Buffalo, Dr. Irving Roberts, of Weiss & Downs, Inc., and Dr.
David W. Stewart, of Columbia University, are in a large measure re-
sponsible for the success of the separation of carbon. Dr. Karl Cohen
has helped with the more recent experiments and has carried through
the calculations on the fractionation process reported in this paper.
From time to time other men have aided us with the actual mechanical
work of carrying on these rather arduous experiments. My thanks
and appreciation are due to all these men, for the program could not
have been performed without the loyal help of my co-workers.

BOTAN Y.—Aublet the botanist, a pioneer against slavery, with a me-
morial genus of palms... O. F. Coox, Lanham, Md.

Darwin’s reaction against slavery, during his visit to Brazil in 1832,
has figured as an example of humanitarian sentiments combined with
scientific pursuits, to show that science is not so aloof from human in-
terests as often supposed. Cuvier affirmed ‘“‘the power of feeling to
exalt the intelligence,” but curiosity affects the mysterious and re-
mote. Darwin noted in Galton’s questionnaire on heredity that more
originality is implied in “‘discoveries with regard to common objects,’
and Aublet saw significance in simple racial reactions that continued
to be disregarded through the slavery controversy. From a scientific
standpoint it must be expected that the racial and social problems,

1 Received December 22, 1939.

JuLy 15, 1940 COOK: AUBLET THE BOTANIST 295

since they lie in the field of biology, eventually will be studied by
consecutive scientific methods, and that each discovery or original
approach to a significant fact will be recognized, appreciated, and
commemorated.

Aublet not only was much earlier than Darwin but was well in ad-
vance of most of the philanthropists who gained distinction through
their efforts for the suppression of the slave trade and the abolition
of slavery, as Sharp, Wilberforce, Buxton, Garrison, and Phillips.
Abbé Gregoire did not organize his ‘‘Friends of the Blacks’”’ till 1788,
the same year when Brissot de Warville traveled in the United States
with a zealous interest in the ‘“‘wretched Africans.’’ Clarkson’s survey
of his “forerunners,” in The history of the abolition of the slave trade,
1808, does not refer to Aublet, nor does Saco’s Historia de la esclavitud,
1879-1903.

Jean Baptiste Christophe Fusee Aublet, a contemporary of the
Jussieus, was born in Provence in 1720, and is chiefly known for his
Histoire des plantes de la Guiane Francoise, 1775, in four octavo vol-
umes, the foundation of forest botany in tropical America. Nearly
400 new plants were described and figured, including many new gen-
era of trees. Piso, Plumier, Browne, Sloane, and Jacquin had explored
the plant world of the West Indies and the shores of South America,
but Aublet entered the continental forest and began the study of its
seemingly inexhaustible flora, continued by Ruiz and Pavon, Martius,
Spruce, Pittier, and many others. Aublet was bent on botany even as
a boy, ran away, to Spain in search of plants, and worked a year for
an apothecary at Granada before he was traced and taken home.

The Guiana collection was made in two years, 1762-1764, but
Aublet had spent 9 years, 1753-1761, in establishing a garden of drug
plants in Mauritius, then a French colony, ‘‘l’Isle-de-France.”’ In
returning from Guiana he spent several months in ‘‘Saint-Domingue,”’
at Mole Saint Nichols, Bombardopolis, and other points in the north-
ern peninsula of Haiti. Thus Aublet had active experience in three of
the tropical colonies as a basis of his chapter on ‘‘Observations sur les
Negres Esclaves,”’ published in the supplement of the Histozre.

Slavery had been challenged on grounds of religious sensibility from
the time of Las Casas in the sixteenth century, more frequently after
George Fox visited Barbados in 1671 and urged a gradual emancipa-
tion of the slaves, as did John Woolman in Maryland and Virginia
before the Revolutionary War. Aublet was a secular voice, perhaps
the first to be raised, objecting to the system of slavery as injurious to
the European settlers. He saw that the French colonists, from being

296 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

kindly, good-natured people, became haughty, hard, and inhuman in
dealing with another race, but saw no object in this severity, the Ne-
groes being, in his view, not turbulent savages, but naturally quiet,
peaceable people, not entirely irresponsible or unwilling to work for
what they might need.

Thus Aublet reached the conclusion that holding the Negroes in
slavery was not necessary for maintaining the colonies, a racial judg-
ment that received a practical demonstration in hundreds of separate
communities on tropical coasts and islands when the system of forced
labor eventually was abandoned. Much of the calamitous history of
Haiti and of the United States might have been avoided if Aublet’s
factual view of the Negro character had prevailed before the explosive
tensions were generated. It generally was believed that continuous
repression was required and that the Negroes would revolt at the first
opportunity. John Brown had that idea, and many other abolition-
ists.

In the light of subsequent history it will not be questioned that the
effect of slavery on the ruling class was a serious factor of the racial
problem of the West Indies. From Bolingbroke’s Voyage to the De-
merary, 1807, it appears that many of the islands were going back-
ward long before the slaves were liberated. Emancipation did not
change the essential conditions, and the decline continued. ‘‘A ruling
class always rules itself out.’’? Some of the finest islands have been en-
tirely abandoned to the Negroes, and now this ‘‘New Africa”’ is re-
ported in serious distress. Macmillan’s Warning from the West Indies,
1936, urges an extension of the policy of trusteeship, ‘‘development
in the interest of the weaker classes,” as projected for the African
colonies. The notion of ruling another race takes another form,
benevolent, of course, but requiring ‘‘a firm exercise of authority in
their behalf.’’ In South Africa it is being recognized that the exploitive
relations are detrimental to both races, and measures of segregation
are being applied.

A few years after Aublet an English botanist, Henry Smeathman,
outlined a plan for settling colonies of civilized Negroes on the coast
of Africa, in order to end the slave trade and eventually to return the
slave population from America. Smeathman went to Sierra Leone in
1771 to collect plants for a scientific group of English Quakers de-
scribed by R. Hingston Fox in Dr. John Fothergill and his friends,
1919. An African tree of the passion-flower family was named Smeath-
mania by Robert Brown, in Tuckey’s Congo Narrative, 1818. Two of
Smeathman’s letters, dated 1783 and 1786, relating to a proposed

JuLy 15, 1940 COOK: AUBLET THE BOTANIST 297

settlement at Sierra Leone were included in Wadstrom’s Essay on
colonization, 1794. Smeathman’s account of the social economy of the
African termites, published by the Royal Society in 1781, is praised
in Maeterlinck’s Life of the white ant as an “inexhaustible treasure’”’
of basic observations.

The policy of colonization had the approval of Washington and
Jefferson, and settlements in Liberia were developed by Ashmun
(1822-1828) to the point of showing that the project was feasible,
though later mismanaged and discredited, but still finding a few sup-
porters in each generation. Senator Bilbo of Mississippi recently has
proposed in Congress a “‘Greater Liberia Act,” for resuming and ex-
tending ‘“‘the voluntary resettlement of American Negroes in Africa.”’
A century of peaceable history in Liberia confirms Aublet’s assay of
the racial temperament.

A GUIANA PALM GENUS TO COMMEMORATE AUBLET

The palm called ‘‘maripa” in French Guiana, figured on the frontispiece
of Aublet’s Hzstozre and described in the Supplement, is distinct as a genus
from the South American palms with which it has been associated and may
receive the name Ethnora, to symbolize the interest of Aublet in the racial
problems. The name Attalea maripa was given to Aublet’s palm by Martius,
and Drude called it Maximiliana maripa. The type species would appear as
Kihnora maripa (Martius). The ample duplex perianth, uniform, crassate
endocarp, and superficial submedial foramina, are diagnostic characters.
The pulp of the fruit yields an edible oil. The related genera were reviewed
in the National Horticultural Magazine for October 1939, in a paper entitled
“Bornoa, an Endemic Palm of Haiti.”

Other distinctive features of Hthnora are the low, robust trunk, rather
short leaves, and the pedunculate inflorescence, less compact than in Bornoa.
The large orbicular-apiculate petals are half as long as the fruit, twice as
long as the broadly cordate-auriculate sepals, and about three times as long
as the tomentose laciniate staminal cup. The stigma is persistent, the ros-
trum indurated, the exocarp firm, the mesocarp fleshy and fibrous with a
compact inner layer of longitudinal mesocarp fibers adhering closely to the
endocarp but not fused, the cross section showing only the few fibers that
outline the sutural sclerites. One or two carpels are fertile, the endosperm
narrowly striate on the outer face, and the slender curved embryo protected
by a papilliform process of the indurated testa. Fruits from Dutch Guiana
collected by Harold F. Loomis for the Allison V. Armour Expedition of 1932
have been examined.

The endocarp of Ethnora is remarkable, the walls very thick and extremely
hard, of uniform texture, with no peripheral fibers, but the sutural sclerites
distinctly outlined in the cross section of the nut by parallel series of fibers,
rather regularly spaced. In Bornoa the sutural bands appear as segments of
the outer wall separated by a few fibers in radial or oblique rows, but in
Ethnora it is plain that the bands project as radial flanges from the center
of the nut and are structurally distinct from the thickened walls of the
carpels, which doubtless must be considered as the endocarp proper. The
central structure is hardly to be viewed as a receptacle or a placenta but

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may represent an intermediate metamere, between the stamens and the
pistils. The nut of Temenia (Maximiliana) shows the same structure as
Ethnora, but the walls are much thinner, with the pits that mark the sutural
bands more numerous, closer to the surface of the chambers, and distinctly
flattened.

The personal name Aubletia is not available, Gaertner in 1788 and four
other botanists within a few years having dedicated new genera to Aublet.
Only the first genus could be recognized, and that proved invalid.

A reduced photograph of Aublet’s frontispiece is reproduced (Fig. 1), with
an enlargement of the details of the maripa palm. The four other palms
“comon,” “‘avoira,”’ “‘bache,” and ‘“‘zaguenete”’ are identified respectively as
Oenocarpus, Astrocaryum, Mauritia, and Manicaria. No other palms were
illustrated in Aublet’s Histoire, though several were briefly described, with-
out botanical names. The leaf above the medallion may represent the
“comon.”’

The palm of Dutch Guiana may be a different species, since it develops
a tall, clean, cylindrical trunk. A single fruit of this palm is 7 cm by nearly
4 em, the petals attaining 4 cm and the sepals 2 cm. An inflorescence branch
with 6 fruits is 18 cm long, the basal joint nearly 4 cm the other joints about
2 cm, and the slender male section 4 cm.

The name “‘cocorite” or ‘‘kokerite” also is applied to palms of this group
in Guiana and Trinidad, usually classified under Maximiliana.

In British Guiana, according to Im Thurn, the cocorite palms are short
or even stemless in the coast districts, but in the interior grow very tall,
“the grandest column-like stem that palm ever had.”’ The name “‘mareepa”’
is given by Im Thurn as relating to the seeds of the cocorite palm.

From Aublet’s descriptions a genus Avoira, with six formally named
species, was recognized by Giseke in 1792, though generally overlooked in
taxonomic literature. Giseke’s genus antedates Astrocaryum Mayer, 1818,
but may prove distinct. The type species, Avozra vulgaris, is represented in
Aublet’s drawing by a branch of the inflorescence with several fruits, while
a branch of Astrocaryum aculeatum Mayer appears from the original descrip-
tion to have only one female flower, at the base of a slender “‘stipe’”’ equal
in length to the thickened male spike.

BOTANY.—A_ second note on Georgia Discomycetes... Epitu K.
CasH, U. S. Bureau of Plant Industry. (Communicated by
JOHN A. STEVENSON.) |

During the past few years the writer has had the privilege of study-
ing numerous specimens of Discomycetes collected in the vicinity of
Athens, Ga., by J. H. Miller and G. E. Thompson, of the University
of Georgia. Several of these fungi have been discussed previously (2);
notes are given here on five additional species, four of which are de-
scribed as new. Specimens are in the herbarium of the University of
Georgia and in the Mycological Collections of the U. 8. Bureau of
Plant Industry, and type material has also been deposited in the
Farlow Herbarium of Harvard University and the herbaria of the
New York Botanical Garden and the University of Michigan.

1 Received January 2, 1940.

300 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

Sphaeropezia arundinariae, n. sp. Fig. 1

Apothecia irregularly scattered over the stems of the host plant, ellipsoid
or elongate, 0.7—1 by 0.3-0.5 mm, deeply embedded in the tissue, at first
completely covered by a lid concrete with the darkened and convexly
arched epidermis, which later splits longitudinally to expose the sunken
hymenium; hymenium pale flesh color to vinaceous-buff (Ridgway),
pruinose; asci cylindrical, rather abruptly narrowed to the apical pore,
gradually attenuated toward the base, pore blue with iodine, 8-spored,
75-85 by 6-8u; spores irregularly biseriate, hyaline, cylindrical-clavate,
3-septate, not constricted, rounded at the ends and narrowed at the lower
end, 16-18 by 3—4u; paraphyses hyaline, filiform, simple or branched near
the tips, often twisted or circinate; hypothecium hyaline or subhyaline, of
small-celled plectenchyma; exciple prosenchymatous, brown, enclosing the
hymenial layer at the base and sides and the cover above it in a continuous
layer; cover concrete with the epidermis, the outer layer prosenchymatous,
brown, with a thick inner layer of hyaline, gelatinous longitudinal cells,
opening by a lengthwise slit, the sections extending perpendicularly or more
frequently breaking away at maturity and leaving only a few fragments
around the margin of the hymenium.

Apotheciis conspersis, ellipsoideis vel elongatis, 0.7—1X0.3—-0.5 mm, im-
mersis, clypeo epidermide adnato et longitudinaliter fisso et hymenium pal-
lide-roseum pulverulentum patefacienti; ascis cylindricis, octosporis, 75-85
X 6-8; ascosporis biseriatis, hyalinis, cylindrico-clavatis, 3-septatis, base
attenuatis, 16-18 3-4u; paraphysibus hyalinis, filiformibus, simplicibus
vel apice ramosis, saepe circinatis, hypothecio subhyalino, plectenchymatico;
excipulo prosenchymatico, brunneo.

On the small branches of Arundinaria tecta, Bobbin Mill, Athens, Ga.,
April 27, 1938, J. H. Miller, and April 5, 1939, G. E. Thompson and J. H.
Miller (type).

This fungus was at first tentatively placed in the genus Phragmonaevia,
which it resembles in its sunken apothecia and hyaline, septate spores. The
inadequate descriptions of many species of this genus make their real char-
acters and affinities questionable. Details of the structure and development,
particularly the manner of opening and the nature of the covering layer, if
present, are usually not noted. Although von Hoehnel in several instances
(4, 5, 6) notes the resemblance of some species of Naevia to Phacidium, at one
time classifying the genus Naevia in the Phacidiaceae, he consistently re-
garded Phragmonaevia as belonging to the Stictidaceae, as have Saccardo
and Rehm, among other writers. The fungus on Arundinarza, however, can
not be considered as stictidaceous, being plainly most closely allied to the
Phacidiaceae. The structure of the apothecium is strikingly similar to that
of Phacidium lacerum Fr., the type of the genus Phacidium, as illustrated
and described by von Hoehnel (4, p. 317, fig. 15). Von Hoehnel’s figure shows
a similar covering lid, open and extending vertically at right angles to the
hymenium, with an inner layer of mucilaginous cells, which he interprets as
an opening mechanism.

2 Color terminology follows Ridgway, Color standards and color nomenclature,
Washington, D. C., 1912.

JuLy 15, 1940 CASH: GEORGIA DISCOMYCETES : 301

Several species of Phragmonaevia growing on grasses are transferred by
von Hoehnel and Nannfeldt to Hysteropezizella v. Hoehn., a genus charac-
terized by elongate ascomata, a covering lid concrete with the epidermis and
opening at one side, simple spores, and acute lanceolate paraphyses. S.
arundinariae differs from Hysteropezizella in the filiform, flexuous paraphyses,
septate spores, and most especially in the prosenchymatous exciple. No
pseudoparenchymatous basal tissue, such as that illustrated and described
by von Hoehnel and Nannfeldt, could be found in this fungus.

The original description of Sphaeropezia Sace. (10, p. 253) is brief, and the
type species, S. alpina Sacc., appears not to have been available for study
by later workers, so that the position of the genus is not clear. First de-
scribed as a genus of the Patellariaceae by Saccardo, it was referred by Rehm
(9, p. 72-73) to the Phacidiaceae, a classification that was adopted by Sac-
cardo in the Sylloge Fungorum, volumes 8-24, and followed by von Hoehnel
(5). Nannfeldt (8, p. 214), on the other hand, on the basis of his examination
of S. empetri (Fckl.) Rehm, considers Sphaeropezia a member of the Pseudo-
sphaeriales, removing it from the Discomycetes completely, but not assign-
ing it definitely to any family. In the absence of more adequate information,
the writer has followed the usage of Rehm and von Hoehnel in describing
this fungus as a Sphaeropezia.

Lachnum arundinariae, n. sp. Fig. 2

Apothecia sessile, scattered, amphigenous, mostly hypophyllous, super-
ficial, minute, 0.1—0.2 mm in diameter, translucent when moist, white-pilose,
- subglobose then patellate, soft-fleshy, hymenium hyaline to sea-shell pink
(R), margin fimbriate; asci cylindrical, short-pedicellate, rounded at the
apex, 8-spored, 27-33 by 3-3.5u; spores narrow-clavate to acicular, biseriate,
unicellular, hyaline, straight, 5-6.5 by 0.7—1u; paraphyses lanceolate, 45-55
by 3-4y; exciple hyaline, prosenchymatous; hairs hyaline, septate, finely
echinulate, not swollen at the apex, 50—70 by 3un.

Apotheciis sessilibus, amphigenis, 0.1—0.2 mm, molle carnosis, albo-pilosis,
margine fimbriato, hymenio carneo; ascis cylindricis, breve pedicellatis,
octosporis, 27-33 X3-3.54; ascosporis anguste clavatis vel acicularibus,
unicellularibus, hyalinis, 5-6.5X0.7-1u; paraphysibus lanceolatis, 45-55
<3-4u; excipulo hyalino, prosenchymatico; pilis hyalinis, septatis, echinu-
latis, 50-70 X 3p.

On Arundinaria tecta, Bobbin Mill, Athens, Ga., April 27, 1938, J. H.
Miller, and May 12, 1989, G. E. Thompson and J. H. Miller (type).

Judged from the description, Pezzza caulicola Fr. (8, p. 94), reported on
Arundinaria, is close to Phialea cyathoidea. Both the former species and
Peziza arundinariae Berk. (1, p. 155) differ from Lachnum arundinariae in
color and in the presence of a stipe. Among species of Lachnum that occur
on grasses, L. carneolum (Sacc.) Rehm f. hyalinum (Rehm, Ascomyceten no.
1931) is more densely pilose, with thicker, longer, crystal-bearing hairs,
larger asci and spores, and shorter paraphyses; L. nardi Rehm is a darker,
stipitate species; and European specimens of L. controversum (Cke.) Rehm
show larger, stouter apothecia, with clavate-tipped hairs and spores equally
acute at both ends.

302 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 7

Pyrenopeziza minuta, n. sp. Fig. 3

Apothecia hypophyllous, subcuticular, becoming superficial with the dis-
organization of the host tissue, closely aggregated in pale spots of withering
leaves, patellate, pale brown when moist with subhyaline hymenium, entire
fungus black when dry, very inconspicuous, 150-175 in diameter, soft
fleshy to membranous, margin uneven; asci clavate to fusoid, rather abruptly
narrowed at the apex and base, 8-spored, 28-33 by 4—5y; spores biseriate or
irregularly 3-seriate in the upper part of the ascus, acicular clavate, 4-5 by
0.7—1u; paraphyses filiform, hyaline, swollen at the tips to 2-3y in diameter
and conglutinate in a yellowish mazaedium; hypothecium thin, subhyaline,
outer layer dark at the base, of elongate, yellow-brown hyphae forming a
dentate margin.

Apotheciis in maculis hypophyllis, subcuticularibus dein superficialibus,
patellatis, pallide brunneis, 150-175u diam., molle-carnosis vel membra-
nosis; ascis clavati-fusoideis, octosporis, 28-33 X4—5yu; ascosporis 2-3-
seriatis, aciculari-clavatis, 4-5 X0.7—1.2u; paraphysibus filiformibus, hya-
linis, apice inflatis et conglutinatis; hypothecio tenui, subhyalino; excipulo
brunneo.

In withered leaves of Tilia heterophylla var. michauxi, Princeton, Ga.,
April 26, 1938, and March 25, 1939 (type), J. H. Miller and G. KE. Thompson.

This species differs from Naevia tiliae (Kleb.) Nannf. (8, p. 190) in smaller
asci and spores, longer paraphyses, and in the thinner, darker basal tissue.

Pyrenopeziza prinicola (EH. & E.), comb. n.

Synonym.—Mollisia prinicola EK. & E., Journ. Myc. 4: 99. 1888.

Apothecia erumpent, then superficial, scattered rather thickly and evenly
over the lower surface of the leaf, minute, 150—275y in diameter, subturbi-
nate then patellate, membranous to fleshy, hymenium alutaceous to dark
brown, the entire fungus black when dry, margin slightly crenulate; asci
clavate, abruptly narrowed at the base and apex, 8-spored, 33-40 by 6-8y;
spores irregularly 2-3-seriate, hyaline, acicular-clavate, 1-celled, guttulate,
becoming pseudoseptate near the middle, the lower end acute and nar-
rowed, straight, 10-15 by 1—1.5u; paraphyses filiform, hyaline, simple or
branched near the tips; hypothecial layer hyaline, plectenchymatic, 15—20u
thick, gradually changing to subhyaline pseudoparenchyma at the base;
exciple subhyaline at the base, dark toward the margin, covered with brown
hyphae 2—3y in diameter adnate with remnants of the host cuticle.

On dead leaves on Quercus stellata, Bobbin Mill, Athens, Ga., April 27,
1938, J. H. Miller; on Quercus rubra, same locality, May 12, 1939, G. E.
Thompson and J. H. Miller. Type on Q. prinus, Louisiana, 1888, A. B.
Langlois 1329.

Comparison with the type specimen of Mollisia prinicola EK. & E. in the
Mycological Collections of the Bureau of Plant Industry shows that the
Georgia material constitutes recent collections of the same species, which is,
however, more properly a Pyrenopeziza than a Mollisia.

So far as can be ascertained from descriptions and available specimens this
American fungus is not identical with similar European species occurring
on oak leaves. Mollisia rabenhorstit (Auersw.) Rehm (9, p. 537) has smaller
asci and spores, apothecia free from the host tissue at the border instead of

Juty 15, 1940 CASH: GEORGIA DISCOMYCETES 303

adnate with it, and a darker, large-celled pseudoparenchymatous exciple
with conspicuous, broad, clavate ends of hyphae projecting at the margin.
Rabh. F. Eur. 2312, issued as Pyrenopeziza foliicola Fckl. and cited by Rehm |
as M. rabenhorsti1, agrees with Rehm’s description of the latter species; |
Sace. Myc. Ven. 975, labeled Pyrenopeziza foliicola (Karst.) Sace. f. quercus
pedunculata, and Peziza nervicola Desm. in Desm. Pl. Crypt. Fr. 1067 also
appear to be the same fungus, differing from P. prinicola in the characters
mentioned.

—

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|
'

Fig. 1.—Sphaeropezia arundinariae on Arundinaria tecta (X 5). Fig. 2—Lachnum
arundinariae on Arundinaria tecta (X 20). Fig. 3.—Pyrenopeziza tiliae on Tilia
heterophylla var. michauxii (X 18). Fig. 4—Phaeangium magnisporum on Betula
nigra (X 5). Photographs by M. L. F. Foubert.

304 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

Phaeangium magnisporum, n. sp. Fig. 4

Apothecia erumpent singly or in groups of two to three, closely sur-
rounded by fragments of broken bark, 1-1.5 mm in diameter, fleshy, sub-
turbinate or obconic, smooth, black, often compressed and contorted, readily
falling out, leaving cavities in the bark, hymenium fuscous-black (R), shin-
ing, rough; asci broad-cylindrical, abruptly narrowed at the base, rounded
at the apex, 4-spored, 175-200 by 25-30y; spores uniseriate, broad-ellipsoid,
dark brown, 1-celled, with one large guttule evident when young, smooth,
surrounded by a thick hyaline gelatinous envelope 2—2.5y thick, 40-50 by
19-—22u (exclusive of enveloping sheath); paraphyses numerous, filiform,
with granular contents, branched and interwoven at the tips, forming a
subgelatinous mazaedium; hypothecium subhyaline, of very loosely formed
plectenchyma, becoming darker and more closely interwoven toward the
base; outer layer of cells breaking up irregularly and forming a furfuraceous
cortex. :

Apotheciis erumpentibus, carnosis, subturbinatis vel obconicis, 1-1.5 mm
diam., nigris, hymenio fusco, nitenti, rugoso; ascis late-cylindricis, 4-sporis,
175-220 X 25-304; ascosporis uniseriatis, late-ellipsoideis, nigro-brunneis,
unicellularibus, 40-50 X 18—22y, tunica gelatinosa, 2—2.5u4 crassa involutis;
paraphysibus numerosis, filiformibus, apice ramosis et intertextis, mazae-
dium gelatinosum formantibus; hypothecio subhyalino, plectenchymatico,
cortice furfuraceo.

In bark of Betula nigra, Agr. Campus, Athens, Ga., March 2, 1939, G. E.
Thompson and J. H. Miller.

P. magnisporum is related to P. tetrasporum (Ell.) Sacc. & Syd., described
by Ellis on Quercus coccinea from New Jersey as Dermatea tetraspora. In the
specimen of the latter species examined (Ellis, N. Am. Fungi no. 70), the
spores are subspherical, 22—28.5 x 18-20u, none reaching even the minimum
length of those in P. magnisporum. P. peckianum Sace. (11, p. 190) on Acer
differs in more caespitose apothecia and much smaller asci and spores.

Nannfeldt has pointed out (7, p. 202) (8, p. 92) that several fungi de-
scribed as Phaeangium are in reality species of Dermatea, Pezicula, and Velu-
taria, the brownish spore coloring being apparently due to age and poor con-
dition of the specimens. This does not seem to be the case, however, in P.
tetrasporum and P. magnisporum, in which the spores are brown in a very
early stage, becoming when mature an intense dark brown, similar to the
color of Xylaria spores. The name Phaeangium is therefore retained for the
present for these two species, in order to avoid the necessity of naming a new
genus.

LITERATURE CITED

. BERKELEY, M.J. Notices of North American fungi. Grevillea 3: 145-160. 1875.
. Casu, E. K. Some Georgia Discomycetes. Journ. Washington Acad. Sci. 29:
47-51, illus. 1939.

. Fries, E. Systema mycologicum, vol. 2, 620 pp. 1822-1823.
HoEHNEL, F. von. Fragmente zur Mykologie no. 1011. Sitzungsb. Akad. Wiss.
Wien, math.-nat. Klasse 126: 314-322, illus. 1917.

Mycologische Fragmente no. 120-190. Ann. Myc. 15: 293-383. 1917.
System der Phacidiales v. H. Ber. deutsch. bot. Ges. 35: 416-422. 1917.
. NANNFELDT, J. A. Type specimens of British inoperculate Discomycetes (First part,
notes 1-50). ‘Trans. Brit. Myc. Soc. 20: 191-206. 1936.

No Boo Ne

JuLy 15, 1940 SAYLOR: NEW WEST INDIAN BEETLES 305

: Studien iiber die Morphologie und Systematik der nichtlichenisterten ino percu-
laien Discomyceten. Nova Acta Soc. Sci. Upsal. (ser. 4) 8(2): 368 pp., illus. 1932.
9. Reum, H. Ascomyceten: Hysteriaceen und Discomyceten. Rabh. Kryptogamenfl.,
eae? yol. 1, Abt. 3, 1270 p., illus. 1896.
10. Saccarpo, P. A. Conspectus generum Discomycetum hucusque cognitorum. Bot.
Centralbl. 18: 213-220, 247-256. 1884.
Notae mycologicae series XX. Nuovo Giorn. Bot. Ital. 23: 187-234. 1916.

ENTOMOLOGY.—Ten new West Indian scarab beetles of the genus
Phyllophaga, with two new names.1. LAWRENCE W. SAYLor,
U. 8S. Bureau of Biological Survey. (Communicated by E. A.
CHAPIN.)

The species described as new in this paper further enlarge our
knowledge of the relatively numerous species of June beetles known
from the West Indies. For the privilege of studying the majority of
these new species I am indebted to Dr. E. A. Chapin, of the United
States National Museum.

Phyllophaga (Phyllophaga) bimammifrons, n. sp. Fig. 2

Male.—Elongate, shining, glabrous above; head piceous, thorax and legs
rufotestaceous, otherwise testaceous above. Clypeus with moderately fine
punctures, these dense at center of disk but nearly absent along apical
margin of clypeus; clypeal margin slightly reflexed, narrowly and shallowly
incised at middle, the angles very broadly rounded; underside of outer
apical margin with a single row of punctures, each bearing a long hair, which
curves up and back over the clypeal apex. Front with two moderately sepa-
rated mammilliform tubercles; apical half of disk of the front with fine,
sparse, and irregularly placed punctures, basal half of disk and vertex with
very dense and extremely fine punctures and several larger intercalated
punctures. Antenna 9-segmented, club not quite equal in length to the four
preceding segments. Thorax with sides entire, with a few cilia near the
apical angles, both hind and fore angles rounded, base and apex with strong,
complete margins, punctures of disk of moderate size, rather regularly
placed and separated by one to two times their diameters, the center of disk
with a small irregular, impunctate area. Elytron with sutural stria obsolete
at base and apex, other striae absent; disk subrugose, punctures slightly
smaller than those of thorax and less densely placed. Pygidium convex,
subrugose, with fine, regularly placed punctures, separated by about two
times their diameters and each bearing a short, erect hair. Abdomen convex,
polished, fifth sternite sparsely punctured; sixth sternite three-fifths as wide
as the preceding, slightly transversely impressed and with fine, moderately
dense, setigerous punctures. Claws with a strong median tooth, claw base
obtusely dilated.

Female.—Antennal club equal to the three preceding segments; pygidial
puncturation somewhat coarser than in male, and disk somewhat tumid be-
fore apex; abdomen with the sixth sternite flat. Otherwise as in the male.

Length, 21-23 mm. Width, 11-12 mm.

The type series consists of 12 males and 8 females, 10 from ‘“‘Sta. Fe, I. de
Pines, Cuba, A. R. Otero collector, March 1939 (E.E.A. Cuba Ento. No.

1 Received February 9, 1940.

306 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 7

11013)’ and 10 from the same locality collected March 21, 1939. The
holotype (U.S.N.M. no. 54014), allotype, and designated paratypes are in the
United States National Museum; paratypes also have been returned to

S. C. Bruner, who kindly sent the series for study, and others have been
deposited in the Saylor Collection.

Db

Fig. 1.—Phyllophaga adjuntas, n. sp. Fig. 2.—Phyllophaga bimammaifrons, n. sp.
Fig. 3.—Phyllophaga cneda, n. sp. Fig. 4.—Phyllophaga blackweldert, n. sp. Fig. 5.—
Phyllophaga bahama, n. sp. a, and b, Lateral view of male genitalia. c, Tarsal claw.

P. bimammifrons is most closely related to P. tuberculifrons (Chev.), with
which species it has in common the bimammillate front, but it may be
readily separated by the more median tooth of the tarsal claw and the very
different male genitalia. In this new species only the females have the smooth

JuLy 15, 1940 SAYLOR: NEW WEST INDIAN BEETLES 307

reniform impression adjacent to each eye that Chapin describes for tuberculi-
frons; in the males the area is punctate to the eye margin.
Phyllophaga (Phyllophaga) wolcotti, n. sp. Fig. 10

Male.—Elongate, testaceo-ochreous to testaceopiceous, head and greater
part of the thoracic disk piceous, legs rufopiceous; upper surface pruinose,
dull, glabrous. Head with clypeus polished, very sparsely punctate, apex
narrowly and shallowly emarginate, angles very broadly rounded. Front
pruinose, with sparse, small, irregularly placed, umbilicate punctures, vertex
and occiput polished and impunctate. Antenna 9-segmented, club oval,
equal to or slightly longer than the preceding three segments. Thorax with
small, irregularly placed, umbilicate punctures, separated by one to two or
more times their diameters; lateral margins entire, angles well marked but
obtuse, base entirely margined. Elytron with sutural stria obsolete basally
and apically; disk subrugose, with punctures of same size and density as
those of the thorax. Pygidium convex, polished, glabrous, with small,
moderately dense umbilicate punctures. Abdomen slightly convex, with very
fine punctures, glabrous or nearly so at middle, fifth sternite long and with
sparse transverse punctures; sixth sternite one-third the length of the pre-
ceding, with a deep transverse sulcus, surface hardly punctate. Claws with
a strong median tooth, which is long and as strong as the apical one; claw
base angularly dilated. Metasternal hairs of moderate thickness. Propygid-
ium strongly polished but with several very minute punctures visible.

Female.—All characters as in the male except that the sixth ventral ab-
dominal sternite is much less strongly impressed and the antennal club is a
little shorter. |

Length, 23—24.5 mm. Width, 11—-12.5 mm.

The holotype (U.S.N.M. no. 54015), allotype, and 2 paratypes are from
“El Yunque, Porto Rico,” collected by G. N. Wolcott on April 6, 1939. An
additional pair of paratypes are from ‘‘Mts. N. of Yauco, P. Rico, [V-20-36,
coll. by R. G. Oakley.”’ The holotype, allotype, and a paratype are in the
United States National Museum; a pair of paratypes is in the collection of
Wolcott, and one paratype has been deposited in the Saylor Collection.

The species is named in honor of G. N. Wolcott, who has contributed
much to our knowledge of Puerto Rican melolonthids and who transmitted
part of the specimens for description. P. wolcotti is most closely related to
P. yunqueana Chapin, but besides the quite different male genitalia, it may
be separated by the darker color, less densely punctate head, nontumid
elypeus, and shorter antennal club.

Phyllophaga (Phyllophaga) spinifemora, n. sp. Fig. 6
-Male.—Oblong oval, shining, glabrous above; pale testaceous, the head
and legs varying to rufotestaceous. Clypeus short, densely punctured, apex
widely and moderately deeply emarginate, almost bilobed, angles well
rounded, margin hardly reflexed. Front with very dense punctures of moder-
ate size, with or without an indication of a slightly impressed, longitudinal
sulcus; vertex polished, impunctate. Eyes rather large. Antenna testaceous,
apparently 8-segmented, the fourth segment as long as the second and third
combined and widened at the middle; fifth segment one-third as long as the
preceding; club equal to funicle in length. Thorax with regularly placed
punctures of moderate size, separated by one and one-half to two times their
diameters, somewhat closer along the front margin and near sides; sides
rounded, angles blunt, lateral margins entire, with very short cilia; hind
marginal line absent. Elytron with sutural stria ill-defined, hardly raised,

308 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30. NO. 7

and punctate; disk subrugose, with moderately dense punctures. Pygidium
convex, polished, either glabrous or with a few short and erect hairs, and
with regularly placed moderate to rather large punctures, the punctures
separated by one to two times their diameters. Abdomen polished, slightly
convex; fifth sternite slightly transversely impressed, the apical margin at
middle prolonged into a slight though evident lobe; sixth sternite one-third
as long as fifth, transversely impressed and punctate. Claws short, with a
stuby triangular tooth at the middle. Upper tooth of front tibia well defined.
First two segments of hind tarsus equal in length. Hind femur with a single

=

Fig. 6.—Phyllophaga spinifemora, n. sp. Fig. 7.—Phyllophaga manchesterea, n. sp.
Fig. 8.—Phyllophaga pseudocalcaris, n. sp. Fig. 9.—Phyllophaga mandevillea, n. sp.

Fig. 10.—Phyllophaga wolcotti, n. sp. a, Lateral view of male genitalia. b, En face
view of male genitalia. c, Front tibia. d, Dorsal view of front tarsus.

row of 6 to 10 backwardly projecting, short stubby spines well inside of, but
parallel to, the hind margin. Spurs free, the longest longer than the first
tarsal segment. Metasternum nearly glabrous at center, sides coarsely punc-
tate and the hair very short and scarcely obvious.

Length, 10-10.5 mm. Width, 5—5.5 mm.

The holotype (U.S.N.M. no. 54016) and 6 paratypes, all males, were col-
lected by E. A. Chapin and R. E. Blackwelder at Santa Cruz, Jamaica,
February 24, 1937, ‘flying at dusk.” The type material, except for a pair
in the Saylor Collection, is in the National Museum collection. This species
is quite distinct from all West Indian species known to me and is not closely
related to any of the known Jamaican species.

JuLy 15, 1940 SAYLOR: NEW WEST INDIAN BEETLES 309

Phyllophaga (Phyllophaga) pseudocalcaris, n. sp. Fig. 8

Male.—Oblong-oval, shining; head and elytra with sparse and scattered
short hairs, otherwise glabrous above; color testaceous to testaceopiceous,
with dark castaneous cloudings on greater part of the head, thoracic disk,
and on the elytral apices—entire dorsal surface in some lights with a faint
metallic sheen. Clypeus short, almost rounded, but very transverse, apex
not reflexed and hardly emarginate, disk densely and somewhat coarsely
punctate. Front with moderately coarse, irregularly placed, moderately
dense, and variolate punctures. Antenna 9-segmented, testaceous, club
ovate and subequal to the preceding four segments in length. Thorax with
moderately coarse, somewhat regularly placed variolate punctures, sepa-
rated by one to three times their diameters; sides roundly dilated, margins
entire, with sparse, short cilia; angles blunt, base margined only at sides.
Elytron with moderately dense and regularly placed punctures of moderate
size, surface smooth, some of the punctures with a very short scarcely
obvious hair; an oblique, strialike, impunctate but not raised line on the
disk, which is otherwise not striate. Pygidium convex, rugose, with moder-
ately dense punctures, disk apparently glabrous, apex ciliate and sub-
rounded. Abdomen convex, polished; fifth sternite depressed laterally,
center of base with a small, ill-defined, and transversely tumid area; sixth
deeply and transversely sulcate, disk rugosely punctate. Claws almost like
those of Phytalus but too widely cleft for that subgenus, the upper claw
subapical and as long as, but thicker than, the apical claw; claw base ob-
tusely dilated. Hind spursfree and slender; lower tibial margin immediately
adjacent to the shorter spur extended into a short but very obvious pseudo-
spur. First two hind tarsal segments subequal in length. Metasternum nearly
glabrous and impunctate at middle, sides moderately densely punctured, the
hair short and but little evident. Hind tarsus very slightly shorter than its
tibia. Upper tooth of front tibia somewhat distant from the apical two.

Length, 10.5 mm. Width, 6 mm.

The unique male type (U.S.N.M. no. 54017) is from Rio San Juan, Domini-
can Republic, collected by G. S. Miller in March 1938. This species is abun-
dantly distinct from all West Indian species known to me both in the external
characters and in the male genitalia.

Phyllophaga (Phyllophaga) blackwelderi, n. sp. Fig. 4

Male.—Oblong. oval, polished; testaceous, with the head, thorax, and legs
rufotestaceous to piceotestaceous; pygidium, thorax, and elytra apparently
glabrous. Clypeus with dense, coarse punctures; apex slightly emarginate,
but little reflexed, the angles very broadly rounded. Front with coarse punc-
tures and a few scattered short hairs, the punctures dense in part toward
vertex but much sparser in the front half bordering the clypeal suture—
vertex impunctate. Eyes large. Antenna 9-segmented, unicolorous testaceous
club not quite equal to the funicle. Thorax with strong and entire basal
margin; sides roundly dilated, coarsely crenate and with a few cilia in apical
half, and entire in basal half; disk with coarse, regularly placed, moderately
sparse punctures, the latter separated by one and one-half to three times
their diameters; in some specimens the disk hasa small and irregular median
smooth area; angles obtuse and not well indicated. Elytron subrugose, punc-
tures slightly more dense than on thorax; striae, other than sutural, scarcely
indicated, the sutural stria later becoming obsolete before reaching the
elytral apex. Pygidium convex, polished, with coarse, regularly placed,
moderately dense punctures, the apex rounded and ciliate. Abdomen flat-

310 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

tened, polished, fifth sternite with coarse, setigerous punctures; sixth sternite
three-fifths the length of fifth, base and apex carinate and the disk trans-
versely flattened and punctate. Metasternum coarsely, densely punctate,
the hair short and not very conspicuous. Front tibia tridentate, the teeth
approximately equidistant. Hind spurs free and slender, the longest one and
one-half times the length of the first tarsal segment, the latter being slightly
shorter than the second. Claws widely cleft, approaching closely the
Phytalus-type; the short but sharp triangular tooth is situated very slightly
beyond the middle; claw base roundly dilated. Hind tarsus subequal to the
tibia.

Female.—Antennal club equal to the preceding four segments; hind tibial
spurs shorter and broader than in male; last segment of abdomen flattened
as in male but a little longer, the basal carina broader and less obvious; first
segment of hind tarsus a little shorter than in the male and the hind tarsus
in some individuals a little shorter than the tibia. Otherwise as in the male.

Length, 9-12 mm. Width, 4.5-7 mm.

The holotype male (U.S.N.M. no. 54018) and 4 paratypes are from station
234, Santa Lucia, British West Indies, collected May 15, 1936, by Dr. R. E.
Blackwelder. The allotype female and 19 additional paratypes have the same
locality and collector but were taken at station 205 on March 27, 1936.
The types are in the National Museum and designated paratypes are in the
Saylor Collection.

The two sexes of this species are surprisingly alike, though they can be
separated by giving attention to the details of the antennae, hind spurs, and
abdomen. The species is not closely related to any described West Indian
species. The only other species described from Santa Lucia is brevipes of
Froelich (1792), which may not be a Phyllophaga, or even a rhizotrogid. In
any event, blackwelderi differs from it in the femora being elongate and not
“ovate-oblong,’’ in the moderately tridentate not “profoundly dentate”
front tibiae, and in the marginal base of the thorax, as well as in other
details.

Phyllophaga (Phyllophaga) cneda, n. sp. Fig. 3

Male.—Oblong-oval, testaceocastaneous; thorax slightly dull, the beetle
otherwise shining above; pygidium elytra, and thorax glabrous. Clypeus
short, transverse, with fine and dense punctures; apex moderately reflexed,
scarcely emarginate at middle, the whole clypeal margin appearing very
nearly semicircular. Front faintly impressed at middle, punctures coarser
than those of clypeus, irregularly and rather sparsely placed. Antenna 9-seg-
mented, club very slightly longer than the funicle. Thorax with complete
basal marginal line, sides entire and ciliate, angles obtuse and fairly well
marked; disk with moderate-sized, regularly placed punctures, separated
by one and one-half to three times their diameters. Elytron with sutural
stria obsolete apically and nearly so basally, otherwise unstriate; disk sub-
rugose, with moderately dense, regularly placed punctures. Pygidium convex
and polished, regularly, moderately densely, but not coarsely punctured.
Abdomen slightly convex, polished; fifth sternite plane, sixth three-fourths
the length of the fifth, and transversely impressed. Eyes large. Claw tooth
situated very nearly at the middle, claw base hardly dilated. Second seg-
ment of hind tarsus not longer than the first. Metasternal hairs very short
and relatively sparse.

Length, 9.7 mm. Width, 5 mm.

Juty 15, 1940 SAYLOR: NEW WEST INDIAN BEETLES 311

The male holotype (U.S.N.M. no. 54019) from Montserrat Island, West
Indies, collected by H. G. Hubbard, is in the National Museum. A paratype
male, from “Basse Terre, St. Kitts Island, March 26,” remains in the Saylor
Collection. This species appears to be most closely related to blackwelderz
Saylor and is considerably smaller than P. montserratensis Arrow, the only
other known species from the island. From blackwelderi the present species
may be most easily separated by the strong medium (not subapical) tarsal
tooth and also by the slightly different male genitalia.

Phyllophaga (Phyllophaga) bahama, n. sp. Fig. 5

Male.—Subelongate, highly polished; color testaceous, the head, thorax,
and legs rufotestaceous to rufopiceous; glabrous above. Clypeus with
moderately dense and regularly placed punctures; apex widely and shallowly
emarginate, angles broadly rounded, margin slightly reflexed. Front a little
more sparsely punctate than the clypeus, the variolate punctures of moder-
ate size. Antenna 9-segmented, club oval and equal to the preceding 4
segments. Thorax regularly, somewhat sparsely punctate, the umbilicate
punctures separated by one to three times their diameters; base not mar-
gined; lateral margins evenly rounded, with a few cilia, not crenulate; hind
angles broadly rounded and indicated only by a very slight point, front
angles nearly rectangular. Elytron with sutural stria obsolete basally and
apically, otherwise unstriate; disk subrugose, punctures of same size and a
little more dense than those of the thorax. Pygidium convex, rugose, the
moderately sized punctures dense and regularly placed, surface apparently
glabrous. Abdomen convex, polished, nearly glabrous and very finely punc-
tate at middle; fifth sternite with moderately dense punctures, the surface
gradually declivous in apical half, which is flattened; sixth sternite three-
fifths the length of the fifth, transversely impressed, and the surface very
densely and not coarsely punctate. Claw with a short, stubby, median tooth,
claw base not obviously dilated. First segment of the hind tarsus varying
from as long as, to very slightly longer than, the second. Metasternal hairs
moderately dense, but quite short.

Female.—Antennal club equal to either the 3 or 4 preceding segments;
pygidium more flattened than in the male, and with or without very short hair;
abdomen less convex, with the fifth sternite flat and the sixth slightly con-
vex, the latter finely and moderately densely punctured in the apical three-
fifths. Otherwise as in the male.

Length, 14-16 mm. Width, 8-9 mm.

The holotype male (U.S.N.M. no. 54020) and allotype female, from ‘‘Nas-
sau, Bahama Island, West Indies,” are in the National Museum. A second
pair from the same locality are in the Saylor Collection. Four additional
paratypes from ‘‘Mangrove Cay, Andros Island, Bahamas, coll. May-June
1917 by Wm. Mann,” agree with the types in all respects; one pair of these
paratypes is deposited in the collection of the American Museum of Natural
History, from whence they came, and the other is in the Saylor Collection.

P. bahama is most closely related to P. youngi Cartwright from Florida,
but may be separated by the smaller size and the different male genitalia.
The close relationship of these two species and the fact that all their char-
acters are closely similar to those of the West Indian type, rather than the
mainland type of Phyllophaga, give added force to the opinions of both
Cartwright and myself that P. young? is not a very old Floridian species but
rather a recent immigrant.

312 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

Phyllophaga (Phyllophaga) adjuntas, n. sp. Fig. 1

Male.—Subelongate, wider behind; testaceous to rufotestaceous, the
thorax and head more rufous; surface shining, head, thorax, and pygidium
glabrous. Clypeus with sparse, regularly placed punctures; apex widely and
shallowly emarginate, hardly reflexed, angles broadly rounded. Front with
moderately dense, irregularly placed punctures, those at the center of apex
slightly smaller, vertex impunctate. Antenna 9-segmented, club subequal to
funicle, segments 5 and 6 of funicle transverse. Thorax with strong basal
margin and wide apical margin; hind angles subrounded, front angles sub-
angulate; disk with moderately fine and umbilicate punctures, separated by
one to three times their diameters. Elytron subrugose, punctured like thorax
but more densely so, the punctures densest near base. Pygidium convex,
polished, fifth sternite with several coarse setigerous punctures at center.
Sixth abdominal sternite a little shorter than the fifth, the basal half being
a smooth but nearly flat process, widest at the middle where it is suddenly
interrupted, thus forming two very blunt and dentate lobes; surface between
apex, and basal and apical processes, transversely impressed and with fine
punctures and sparse erect hairs, the entire sternite with a faint suggestion
of a median longitudinal sulcus. Claw tooth a little stronger than apical one
and situated a little beyond the middle, claw base obtusely dilated. Meta-
sternal hair of moderate length and density. First two segments of hind
tarsus subequal in length.

Female.—Antennal club equal to or slightly longer than the preceding 4
segments; sixth abdominal segment slightly convex, with fine and not dense
setigerous punctures; first segment of hind tarsus shorter than the second.
Otherwise as in the male. :

Length, 13-15 mm. Width, 6-7 mm.

The holotype male from Isolina, Puerto Rico, collected on April 16, 1901,
is in the United States National Museum (U.S.N.M. no. 54021). The
allotype female and a paratype male, both from Adjuntas, Puerto Rico,
collected May 2, 1917, at the ““Fed. Exp. Station, Acc. no. 49, 1917,” are
in the American Museum of Natural History, whence they came as a loan
for study.

P. adjuntas does not resemble closely any known West Indian species in
the type of male genitalia; externally it resembles P. citri Smyth but differs
from that species in the nonpruinose dorsal surface, the much longer male
antennal club, and the quite different male sixth abdominal segment.

Phyllophaga (Phyllophaga) mandevillea, n. sp. Fig. 9

Male.—Rufocasteneous; polished and glabrous above. Head with front
convex and with a suggestion of a median longitudinal sulcus, and with
rather coarse, variolate, and somewhat sparse punctures. Clypeus very

’ short and transverse, the apical margin narrowly and very strongly emargi-

nate, the emargination in connection with the strongly rounded angles
giving the clypeus a distinctly bilobed appearance; disk coarsely and not
densely punctate. Eyes large. Antennae 8-segmented, the club very slightly
longer than the funicle; fourth segment twice the length of third, fifth seg-
ment transverse and about one-fourth as long as fourth. Thorax with the
disk variolately punctate, the punctures moderately coarse but sparse, and
separated by two or three times their diameters on the center of the disk,
but closer at sides; base of thorax not margined at middle, front margin
very weakly indicated but entire, sides roundly dilated and entire, with
sparse and very short cilia; front and hind angles obtusely subangulate.

Juty 15, 1940 SAYLOR: NEW WEST INDIAN BEETLES 313

Scutellum impunctate. Elytron with coarse, variolate punctures, surface
unstriate and subrugose; sutural stria not well indicated at middle and
entirely obsolete basally and apically. Pygidium strongly convex, pruinose,
with moderately dense, fine punctures, and short erect hair; apex sub-
truncate and ciliate. Abdomen polished, flattened at middle, coarsely punc-
tured, and with sparse short hairs; fifth sternite slightly longer than fourth,
middle of the apical margin with two sharp but short teeth separated from
each other by a shallow emargination; sixth sternite as long as fifth, surface
rugose, flattened, with coarse punctures and slightly carinate basal and
apical marginal lines, each bearing a single row of rather long hairs. All claws
short with the apical tooth rather strongly curved; middle tooth small,
situated slightly basad from the middle and strongly inclined toward the
base; basal dilation obtusely rounded. All tarsal segments with moderately
dense hairy soles; first segment of each tarsus longer than the second. Front
tibia strongly tridentate, the upper tooth slightly beyond the middle. Meta-
sternum strongly, very densely, and umbilicately punctate, with very short
procumbent hair; somewhat more sparsely punctate at middle. All tarsi
longer than their tibiae. Apex of middle tibia with a strongly serrate margin;
each lateral angle of the apex is somewhat prolonged into a slightly larger
tooth.

Female.—Antennal club ovate, hardly equal to segments 3 to 5 combined;
eyes smaller; clypeus larger than in male; pygidium more narrowed apically
and the hairs longer; abdomen semiconvex and very coarsely punctate,
with the apical margin of the fifth sternite plane; sixth abdominal segment
shightly convex; hind tarsus distinctly shorter than the tibia; all other
characters including the middle tibia, exactly as in the male.

Length, 11.5 to 13.5 mm. Width, 5.5 to 6.5 mm.

The holotype male, the allotype female, and a pair of paratypes are from
Mandeville, Manchester, Jamaica, collected between November 16 and
December 19, 1919, at altitudes between 2,131 and 2,250 feet. The holotype
and allotype are deposited in the American Museum of Natural History
and the paratypes in the Saylor Collection.

This species is most closely related to a species (from exactly the same
locality in Jamaica) that is now in manuscript by M. W. Sanderson. From
this species of Sanderson’s the new species differs especially in having the
third tarsal segment two and one-half times longer than wide (rather than
only one-half longer than wide), the elytral punctures finer and evenly
spaced (not confusedly punctate), the hairy ‘‘sole’’ of the tarsus much less
dense, and the slightly different male genitalia. I am indebted to Dr. Sander-
son for comparing my specimens with the type of his new species.

Phyllophaga (Phyllophaga) manchesterea, n. sp. Des 7

Male.—Castaneous, shining, glabrous above. Head with front very coarse-
ly, densely and variolately punctured, with a strong median, impressed, longi-
tudinal sulcus. Clypeus short, the apex reflexed, narrowly and deeply
emarginate and the angles rounded, the entire clypeus appearing bilobed;
disk coarsely punctate. Antenna 8-segmented, club subequal to funicle;
segments 3 and 4 moderately long, the fourth slightly longer than the pre-
ceding; fifth transverse. Thorax with disk variolately punctate, the punc-
tures moderately coarse and separated by one to two times their diameters
on the disk, much closer at sides and nearly contiguous at center of apex;
base not margined at middle; front with strong marginal lines; sides roundly
dilated, entire; angles obtrusely subangulate. Scutellum with several coarse

314 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

punctures. Elytron rugose, nonstriate on disk, the punctures coarse, vario-
late, and moderately dense, sutural stria obsolete basally and apically.
Pygidium convex, slightly pruinose, with moderately dense and very coarse
punctures, and apparently glabrous except for the row of hairs at apex.
Abdomen slightly flattened, coarsely punctate, with scattered short hairs;
fifth sternite coarsely punctate, apical margin evenly rounded; sixth sternite
half as long as fifth, and transversely impressed. All claws of moderate
length, each with a very small median tooth. Hind tibial spurs free, the
longest longer than the first tarsal segment. First segment of hind tarsus
shorter than the second; in the front and mid tarsi the first segment is equal
in length to the second. Metasternum coarsely and very densely punctate
at sides and very sparsely so at center, with short and hardly obvious hair. ©
Front tibia tridentate, the upper tooth slightly beyond the middle in pos-
tion; the middle tooth is distinctly closer to the apical than to the basal
tooth.

Length, 9.6 mm. Width, 4 mm.

The unique male type is from Mandeville, Manchester, Jamaica, collected
November 30 to December 4, 1919, at an altitude of 2,131 feet, and is de-
posited in the collection of the American Museum of Natural History, from
where it was lent for study by Dr. R. EK. Blackwelder.

P. manchesterea is nearest P. mandevillea in general relationships but
differs especially in the smaller size, tarsal claws, much less pilose tarsi,
pygidial and abdominal characters, and the coarser and denser puncturation.
From Sanderson’s new species it differs especially in having a median tarsal
claw tooth that the former lacks.

NEW NAMES

Whereas the name Listrochelus can not be maintained as a genus by any
set of characters yet discovered! and must be considered as a subgenus of
Phyllophaga, several specific names become homonyms, and new names are
needed. Two are here proposed:

Phyllophaga (Phyllophaga) temaxa, new name

Lachnosterna parilis Bates, Biol. Cent. Amer. 2(2): 404. 1889. (Non
Listrochelus parilis Bates, 1888, l.c., p. 172.)

Phyllophaga (Listrochelus) yaqui, new name

Listrochelus debilis Arrow, Ann. Mag. Nat. Hist. (ser. 10) 11: 145. 1933.
(Non Gynnis debilis LeConte, 1856, non Lachnosterna debilis Horn, 1885.)

1 Saytor, L. W. Necessary changes in important rhizotrogid genera. Rev. Ent.
7: 318-322. 1987.

2 Savior, L. W. Revision of the beetles of the melolonthine subgenus Phytalus of
the United States. Proc. U. S. Nat. Mus. 86 (8048): 157-167. 1989.

JuuLy 15, 1940 REHDER: A NEW MOLLUSK FROM MEXICO ills

MALACOLOGY.—A new urocoptid mollusk from Mexico... HARALD
A. REHDER, U.S. National Museum.

Among some mollusks from the State of Nuevo Leon, Mexico, sent
to the United States National Museum by E. J. Koestner, was an
interesting new urocoptid, which was determined to be related to
Propilsbrya Bartsch.

This group was created by Dr. Paul Bartsch in 1906? as a subgenus
of Epirobia to include a single species Epirobia (Propilsbrya)
nelsont Bartsch, from the Sierra Guadalupe, Coahuila, Mexico.
It differs radically from Epirobia in having a broader axis, an
axial lamella, and a denticulate parietal lamella; moreover, it
is far out of the range of the more southerly Epzrobia. In the same
publication, on page 151, Bartsch defined a new subgenus of Holospira
under the name Stalactella, for the species Holospira (Stalactella) rosei
Bartsch, from Tehuacan, Puebla, Mexico. This species has the same
type of lamellae as found in Propilsbrya and has the same general
form, but is decollate. In this group, however, the presence or absence
of the early whorls is apparently not a differentiating character, as the
first four or five whorls seem to be readily detachable. Among the 11
specimens of the present new species that I have been able to examine,
two had the apical whorls still adherent, and in one of these the first
4 or 5 whorls dropped off during the examination of the specimen.

We therefore may unite Propilsbrya and Stalactella, retaining the
first name as the generic designation for the group in question and
placing it near Holospira, from which it differs in being slenderer and
decollate and in possessing denticulate lamellae.

The shell submitted by Mr. Koestner is related to Propilsbrya (Sta-
lactella) roset, differing from it in the internal armature. For this spe-
cies I propose the following new subgenus:

Pectinistemma, n. subgen.

Shell of moderate size, axially ribbed, with the nuclear whorls usually
deciduous. An axial lamella is present, originating anteriorly as a cord and
gradually becoming broader and more lamellar, furnished in the third and
fourth whorls from the last with long, flattened, rather crowded, forward-
curving teeth. A strong parietal lamella is also present, originating in the
fifth whorl from the last, and in the next two whorls scalloped, ‘or bearing
crowded, short, broad, flattened, forward-curving teeth.

Type.—Propilsbrya (Pectinistemma) koestner, new species.

This subgenus differs from typical Propilsbrya in the axial lamella being
thin and bearing denticles and in the teeth of the parietal lamella being

* Published by permission of the Secretary of the Smithsonian Institution. Re-

ceived March 9, 1940.
2 BartscH, Pauu. Proc. U. S. Nat. Mus. 31(1483): 121. 1906.

316 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

broader and more crowded, forming almost a thin, pendant, platelike
lamella.
Propilsbrya (Pectinistemma) koestneri, n. sp. Figs. 1-3

Shell of moderate size, slender, cylindrical, gradually tapering at the
apex, vinaceous-brown to bluish gray in color. The nuclear whorls about 3,
rather smooth, somewhat bulbous, the following 184 whorls rather flat-
tened, strongly impressed at the suture, rather closely axially ribbed. The
first 5 whorls are easily detachable and generally lost, so that the shell is
decollate, with the minute round perforation of the hollow axis visible from

1

e

3

Figs. 1-3.—Propilsbrya (Pectinistemma) koestneri, n. sp. Fig. 1.—Holotype, <3.
Fig. 2.—Paratype, immature, showing nuclear whorls, X3. Fig. 3.—Internal view,
showing lamellae, <7.

above. The base is evenly rounded, the axial ribs continuing into the minute
umbilical perforation. The aperture is oval, the posterior portion somewhat
flattened, and the lip entire, slightly reflexed. The lamellae are adequately
described in the subgeneric diagnosis above. Both lamellae terminate just
behind the aperture and hence are not visible from the exterior, the colu- |
mella showing only a slight axial swelling.

The type, U.S.N.M. no. 535762, was collected by R. A. Schneider and E. J.
Koestner on the Cerro Potosi at Galeana, State of Nuevo Leon, Mexico.
Here it was hanging on mosses under rock outcroppings at an elevation of
12,000 feet. It, a decollate specimen, measures: Height, 17.4 mm; width,
5.0 mm.

Seven other specimens, U.S.N.M. no. 535763, are from the same lot.

Three further specimens, U.S.N.M. no. 467392, were received some years
ago from Dr. 8S. 8. Berry. They were collected by 8. Dickin on the north
slope of Cerro Potosi at an elevation between 10,000 and 11,000 feet.

From Propilsbrya roset Bartsch, described from Tehuacan, Puebla, this
species differs externally in color and in the axial riblets being stronger and
more crowded. It is, moreover, a larger, stouter shell, and the apical whorls
are slenderer and attenuated. From Propilsbrya nelsoni Bartsch it differs in
being a stouter shell with more flattened whorls. :

JuLY 15, 1940 PROCEEDINGS: THE ACADEMY Sly,

PROCEEDINGS OF THE ACADEMY AND
AFFILIATED SOCIETIES

THE ACADEMY
360TH MEETING OF THE BOARD OF MANAGERS
The 360th meeting of the Board of Managers was held in the Board Room

of the Cosmos Club on Friday, April 19, 1940. President CriTTENDEN called
the meeting to order at 8:02 p.m. There were 19 present, as follows:

E. C. CrRiTTENDEN Me Ee Curtis

F. D. Rossini W. RAMBERG

F. C. KRacEK E. W. PRIcE

H. S. RappLEyYE C. L. GARNER

G. STEINER H. G. DorsEy

F. M. SETZLER C. L. GaziIn

W. B. BEL W. W. DIEHL

A. T. McPHERSON and by invitation:
C. THom J. H. Kempton
W. A. Dayton R. J. SEEGER

The customary reports of the standing committees were read and accepted.
H. L. Curtis, chairman of the special committee, charged with reeommend-
ing to the Board the appropriate action to be taken with respect to amending
the bylaws on the subject of the offices of nonresident vice-presidents, re-
ported his committee in disagreement but with the possibility of accord by
the next meeting of the Board.

M. W. StiRLING was elected by the Board to fill the vacancy caused by the
resignation of J. F. Coucn who had been elected manager at the last elec-
tion.

President CRITTENDEN was authorized to appoint delegates from the
Academy to the Fiftieth Anniversary celebration of the Ohio Academy of
Sciences at Columbus, Ohio, May 9-11, and to the Eighth American Scien-
tific Congress at Washington, D. C., May 10-18. He was also authorized to
appoint two committees—one to consider Societies eligible for affiliation
with the Academy and the other to consider the question of a meeting place
for the Academy when the Cosmos Club leaves its present quarters.

The Board adjourned at 10:02 p.m.

ANTHROPOLOGICAL SOCIETY

The Anthropological Society of Washington at its annual meeting held
January 16, 1940, elected the following officers for the ensuing year: Presi-
dent, Frank M. Srerzuer; Vice-president, JuLian H. Srewarp; Secretary,
REGINA FLANNERY; Treasurer, T. DALE StewaRt; members of the Board of
Managers, GrorGcEeS. Duncan, Hersert W. Krizcer, JULIAN H. STEWARD,
RutH UNDERHILL, WaLpo R. WEDEL.

A report of the membership and activities of the Society since the last
annual meeting follows.

Membership:
aie smnemlbenshe ue ne 2
PNEUIVEe MENDES. ue. 6: 4. 4 a 39
Associate members. 4.12) 4..8. Jal
‘LUG ie Ro a Si tee a ey 2,

318 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30,‘NO. 7

The members elected during the year were: Witi1am N. FEnTon,
ANTONIO SAnTA Cruz, and HELEN CULLINANE, active members; WALTER
Bart GREENWoop and JosnPH R. SINGER, associate members.

The Treasurer’s report is as follows:

Funds invested in Perpetual Building Association

(Interest for 1939 not included) $1,453.42
21 shares Washington Sanitary Improvement Co.
par value $10 per share 210.00
2 shares Washington Sanitary Housing Co.
par value $100 per share 200.00
Cash in bank 576.62
$2,440.04
Bills outstanding: |
To American Anthropological Association $80.00
To printer 3.75
To Treasurer 1.93
$85.68 85.68
Total $2 , 354.36
Total as of 1/16/39 ZAVALA GO
Increase $ 182.60

Papers presented before the regular meetings of the Society were as
follows:

January 17, 1939, 683d meeting, Joonn M. Cooper, Are the simplest living
cultures representative of early prehistoric cultures?

February 21, 1939, 684th meeting held jointly with the Smithsonian In-
stitution on the occasion of the Eighth Arthur Lecture, HERBERT J. SPINDEN,
Sun worship.

March 21, 1939, 685th meeting, WitiiAM E. Capirux, The Blackfoot
Indians of Montana.

April 20, 1939, 686th meeting, held jointly with the Washington Academy
of Sciences, W. M. Kroeman, The skeleton tells its story.

October 17, 1939, 687th meeting, W. W. Hin, Economic history of the
Navaho Indians.

November 21, 1939, 688th meeting, Davin C. GraHam, The customs and
myths of the Ch’uan Miao of southwest China.

December 19, 1939, 689th meeting, Joun R. Swanton, De Soto’s route
through the Southeast.

REGINA FLANNERY, Secretary

BOTANICAL SOCIETY

293D MEETING

The 293d regular meeting was held in the assembly hall of the Cosmos
Club, December 6, 1938, President GravaTT presiding; attendance 70. —
RocErs McVaveu, BRITTAIN B. Rosinson, Marcus M. RHOADES,
DANIEL SULLIVAN, and CLAUDE Hops were elected to membership.

Juty 15, 1940 PROCEEDINGS: THE ACADEMY 319

Notes and reviews.—M. B. Warte called attention to the scarlet oak,
which still showed a brilliant red coloring while the usual autumnal coloring
was about October 10. He also displayed a plant of Mammoth Maryland
tobacco in flower which had survived two heavy frosts. The tobacco crop
had been harvested about the first of September. CHARLES THOM exhibited
a new book, Introduction to industrial mycology, by George Smith.

Program.—PERKINS CovILLE: Prairie shelter belt work of the U.S. Forest
Service.

38TH ANNUAL MEETING

The 38th annual meeting was held immediately following adjournment
of the 293d regular meeting, G. F. GRAvaTT presiding.

James F. Couch read an obituary for V. K. CuyEsnut; John A. Stevenson
for B. T. GatLtoway; Charles E. Chambliss for FRanK LAMSON-SCRIBNER;
H. P. Barss for Freep C. Mermr; C. E. Leighty for Guy N. Cotuins. Emma
F. SrRRINE was elected to honorary membership.

The following officers were elected for 1939: President, CHARLES THOM;
Vice-President, CHARLOTTE ELuLioTT; Recording Secretary, ALice M.
ANDERSEN; Corresponding Secretary, NELLIE W. Nanczu; Treasurer, KEN-
NETH B. Raper. Nominated for Vice-President of the Washington
Academy of Sciences, G. F. GRAVATT.

294TH MEETING

The 294th regular meeting was held in the assembly hall of the Cosmos
Club, January 3, 1939, President CHARLES THom presiding; attendance 90.
Harry HUMFELD was elected to membership.

Notes and reviews.—M. B. Waits called attention to the fact that many
tree seeds require a resting period before they will germinate, but that he had
observed that the acorn of the rock chestnut oak does not require a resting
period and germinates immediately upon falling to the ground. In November
the roots of the seedlings were 6 inches long.

Program.—W. M. Lunn, D. E. Brown, J. E. McMurtrey, and W. W.
GARNER: Tobacco and weeds.—It was soon observed in colonial days that
virgin soil produced good yields of tobacco and that the leaf had a finer tex-
ture and lighter body, better suited to the market demands of the period,
than that grown on the older cultivated areas. The same holds true at the
present time. Consequently, in order consistently to produce the desired
type of leaf the early settlers and their successors continued to clear the
forested areas until in time most of the land suited to tobacco in the older
tobacco-producing regions had been brought into cultivation. Subsequently,
crop rotation and the use of manures and fertilizers have been tried in an ef-
fort to maintain yields and quality on old land but these practices have not
proved satisfactory on all soils and with all crop combinations.

It is clearly evident from the results presented that tobacco that is fer-
tilized intelligently and grown after natural weed fallow of sufficient duration
possesses in a large measure those characteristics which were early observed
in the crop grown on virgin land. The crop grown after bare fallow has shown
a rapid decline in yield and gross value, demonstrating that the cover of
spontaneous vegetation is the keystone of the system rather than simply al-
lowing the land to remain idle.

The tests conducted with individual weed and crop plant species have
consistently shown that certain species are much more desirable than others.
Ragweed and horseweed as the preceding cover crop have produced marked

320 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 7

increases in yields and especially in values:of the tobacco crop over results
obtained from bare fallow. On the other hand, following lamb’s-quarters the
tobacco crop has actually shown reductions in yields and value as compared
with bare fallow. In these tests annual lespedeza has shown no advantage
as a cover crop to precede tobacco, sweet clover, rabbit’s-foot clover, and
wild pea have not always shown a decided advantage, while partridge pea
has produced some increase in yield. While the natural weeds occurring in
these tests consisted principally of those species in pure stands which pro-
duced high quality leaf, it is possible that the ones found to be objectionable
might predominate under some conditions, with a resulting harmful effect
on the succeeding tobacco crop. It is hardly to be expected that a given weed
species would have the same effect on tobacco on all soils or under all condi-
tions.

The general beneficial effect of the weed fallow was the promotion of a
quick start and very rapid and uniform growth of the tobacco plants from
transplanting time to maturity. Within normal limits this result is, in turn,
associated with a uniformly high market value per acre and average price
per pound which demonstrates that the resulting product meets current de-
mands for most manufacturing purposes.

J. H. Martin: Cleaning 20 million bushels of seed grain (lantern).—Aver-
age grain from the spring-wheat States contains more than 2 percent by
weight of weed seeds, chiefly wild oats, mustard, and pigeon grass. Nearly
20 million bushels of Government-owned wheat, oats, barley, and flax were
cleaned for seeding in drought-stricken areas in 1934-35. Seed separation by
mechanical means is possible when the seeds differ appreciably in specific
gravity, length, or thickness. The most complete separations are accom-
plished by means of pocketed discs or indented cylinders when seeds differ in
length. Suction separates better than air blasts. Mustard seeds roll over
screen perforations and are difficult to separate completely from grain.
Quackgrass seeds could not be separated completely from oats or flaxseed.
Mechanical cleaning increased the test weight of wheat slightly, barley con-
siderably, and oats remarkably, but reduced that of flaxseed by roughening
the seed coats and preventing close packing.

AuicE M. ANDERSEN, Recording Secretary.

Curmistry.—The separa ation of isotopes by.
Haroun CO URey ie ee

Borany. —Aublet the botanist, a pioneer —
memorial genus of palms. ©! F. Coox.. ee

ho

- Botany.—A second note on Georgia Diseomycetes, oh So

Maxacotocy.—A new urocoptid mollusk from Mexico
REHDER. a ts SO hee ease Res Ce ey ie

PROCEEDINGS: THe ACADEMY./.. n= (77

PROCEEDINGS: ANTHROPOLOGICAL SOCIETY............

PRocEEDINGS: BoranicaL SOCIETY.

RayYMonpD J. SEEGER
GRORGH WASHINGTON UNIVDRSITY

keane Hy Gaiey

- ENTOMOLOGICAL SOCIETY

GEORGE TUNELL
_ GEOLOGICAL SOCIETY

\

T. DALE STEWART

_ ANTHROPOLOGICAL SOCINTY

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OFFICERS OF THE ACADEMY

President: KucENE C. CrittENDEN, National Bureau of Standards. |
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JOURNAL

OF THE
WASHINGTON ACADEMY OF SCIENCES
Vou. 30 Aveust 15, 1940 No. 8

PALEONTOLOGY .—Seven new genera of Carboniferous Crinoidea
Inadunata.1. Epwin Kirk, U. 8. Geological Survey.

A considerable number of inadunate crinoids with turbinate dorsal
cups, three anal plates in the cup, and with varying arm structures
have been described from the Carboniferous. In earlier years these
species were, as a rule, described as Poterzocrinus or Scaphiocrinus.
Latterly many of them have been referred to Pachylocrinus or Scytalo-
crinus. The assignment has been based primarily on whether the arms
divide above the first dichotom. A study extending over a number of
years of practically all the described species and a considerable series
of undescribed species has shown that such a casual grouping obscures
both the biologic and stratigraphic relationships of the crinoids. In
some cases, owing to the poor quality of the types and uncertainty of
accurate placement of well-preserved material, description of the
genera will have to wait upon description of adequate specific material.

In forming these inadunate genera and others to be established,
consideration has been given to all known species and large numbers of
undescribed species. It may seem at times that some of the genera are
too closely circumscribed. However, one must either use the inclusive
genera of the past with stratigraphic ranges in some cases from the
Silurian to the Carboniferous, or attempt to delimit phyletic groups
that have real stratigraphic and genetic significance. Hundreds of
complete crowns are available for study in the museums. These give
pertinent information as to ontogenetic development and permissible
variation in structures. In the case of most genera they also furnish
adequate phylogenetic series.

Blothrocrinus, n. gen.

Genotype.—Poteriocrinus jesupt Whitfield.
Generic diagnosis.—
Crown. Very high, widening gradually distad, then gradually contracting.
Dorsal cup. Turbinate; plates smooth.
IBB. High, forming an appreciable amount of the cup wall.
BB. Large.

1 Published by permission of the Director, Geological Survey, U. 8S. Department
of the Interior. Received March 19, 1940.

o21

Aue 13 8

322 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 8

RR. Large; articulating facet full width of R, shghtly curved, suture not
gaping.

IBr. Two in all rays except anterior (irregularly one IBr in early species) ;
in ant R, [Br range in number from 3 (one specimen) to 14 as seen, the
average being about 10. 3

Arms. The arms are very long, rounded, and with slightly cuneate
brachials. In earlier species and in some of the later ones the arms are
endotomous, with two or three divisions above the primaxil. In some
species the first admedian ramus divides, giving approximately iso-
tomous arms but leading toward para-endotomy.

Post IR. Three anal plates in cup; RA large, penetrating between the post
B and r post B to about one-half their height; X large, meeting post B
on wide, horizontal face, about one-half the plate rising above the level
of the RR. RT large, extending well above level of RR.

Ventral sac. The ventral sac is imperfectly known. A portion of ventral
sac with fragments of arms was identified by Springer (1900) as
Blothrocrinus swallowi. According to this, the ventral sac was long and
slender and made up of fragile plates, plentifully pierced with pores
along their margins. Another specimen, a partial set of arms, shows a
portion of the sac similar to that figured by Springer. The sac extended
nearly to the tips of the arms.

Column. Circular in section with pentagonal lumen.

Characteristic species of the genus.—

Blothrocrinus cultidactylus (Hall), n. comb.

Poteriocrinus cultidactylus Hall, p. 62, 1860. ‘‘Base of the Burlington lime-
stone, near Hamburgh, Illinois.”—Worthen, p. 301, pl. 30, fig. 1, 1883.

Poteriocrinus (Scaphiocrinus) cultidactylus Wachsmuth and Springer, p. 114
(337), 1880.

Scaphiocrinus cultidactylus Wachsmuth and Springer, p. 237 (161), 1886.

Blothrocrinus jesupi (Whitfield), n. comb.

Poteriocrinus jesupi Whitfield, p. 7, pls. 1, 2, 1881. “‘Burlington limestone,
Burlington, Iowa.’’ (Upper Burlington.)
Pachylocrinus jesupi Springer, p. 71, 1926.
Scaphiocrinus swallowi (Meek and Worthen) (pars). (Cited as synonym of.)
Wachsmuth and Springer, p. 235 (159), 1886.

Blothrocrinus spartarius (Miller and Gurley), n. comb.

Poteriocrinus spartarius Miller and Gurley, p. 37, pl. 7, fig. 1, 1890a. ‘“Kinder-
hook, Le Grand, Iowa.’ (Hampton formation. )

Pachylocrinus spartarius Springer, p. 71, 1926.—Laudon and Beane, p. 254,
De Oe hee Se solut Seamer 2 Own

Potertocrinus scopae Miller and Gurley, p. 38, pl. 7, fig. 2, 1890a.

Poteriocrinus genista Miller and Gurley, p. 38, pl. 7, fig. 3, 1890a.

Blothrocrinus swallowi (Meek and Worthen), n. comb.

Poteriocrinus swallovt Meek and Worthen, p. 397, 1860.—Meek and Worthen,
p. 183, pl. 16, figs. 4a, b, 1866. “Burlington limestone, Burlington, lowa.”’
(Upper Burlington.)

Poteriocrinus (Scaphiocrinus) swallovi Wachsmuth and Springer, p. 114
(337), 1880.

Avuaust 15, 1940 KIRK: NEW CARBONIFEROUS CRINOIDEA 323

Geologic and geographic distribution.—Blothrocrinus as known ranges from
the Kinderhook to the Burlington-Keokuk ‘‘transition beds’”’ of the Missis-
sippi Valley.

Relationships.—In its arm pattern Blothrocrinus resembles Pachylocrinus,
the only described genus with which it can be compared. The presence of
numerous [Br in the anterior ray and the long, slender rami of Blothrocrinus
as against the consistent two IBr and short rami of Pachylocrinus are the
more obvious differences in arm structure. The turbinate dorsal cup of
Blothrocrinus is in marked contrast to the depressed bowl-shaped cup of
Pachylocrinus. The ventral sac of Blothrocrinus is long and slender, while
that of Pachylocrinus is relatively short and stout.

Remarks.—It has been the custom to identify all large Blothrocrinus from
the upper Burlington as swallow: Meek and Worthen. Blothrocrinus jesupt,
here chosen as genotype, may or may not be synonymous with swallow.
The original description and the poor figure of the badly preserved type of
swallowit make it impossible to identify the species with certainty. It has
seemed wise to choose the splendid specimens used as types of jesupz.

Cydrocrinus, n. gen.

Genotype.—Potertocrinus coxanus Worthen.
Generic diagnosis.—

Crown. Of medium height, compact, expanding gradually to about three-
fourths its height, then contracting.

Dorsal cup. Broadly turbinate, cup plates relatively thin, unornamented.

IBB. Large, making up an appreciable amount of the cup.

BB. Large. |

RR. Articulating facet full width of the R, slightly crescentic. Suture not
gaping.

IBr. Two in all rays (variation possible in ant R, as apparently in the type
specimen of C. concinnus, the only case known).

Arms. Para-endotomous, that is, each half-ray is endotomous. Two to
three divisions in each half of the half-ray, giving a great number of
closely crowded rami. Br wedge-shaped to cuneate. Pinnules long,
moderately stout.

Post IR. RA large, pentagonal, not entering deeply between the post and
r post BB. X large, extending well above the level of the R. RT smaller.

Ventral sac. Unusually broad and stout, composed of large plates with
radiating ridges, nodose, but not spinous. The sac has a height of about
three-fourths or three-fifths the height of the crown.

Column. Circular in section, lumen pentalobate.

Characteristic species of the genus.—

Cydrocrinus concinnus (Meek and Worthen), n. comb.

Potertocrinites (Zeacrinus?) concinnus Meek and Worthen, p. 26, 1870.—
Meek and Worthen, p. 490, pl. 14, fig. 3, 1873. (Called Zeacrinus con-
cinnus in explanation of plate.) ‘Keokuk, Crawfordsville, Indiana.”’
(Upper Borden.)

Poteriocrinus (Pachylocrinus) concinnus Wachsmuth and Springer, p. 116
(339), 1880.

324 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 8

Cydrocrinus coxanus (Worthen), n. comb.
Potertocrinus coxanus Worthen, p. 4, 1882. ‘‘Upper part of Keokuk lime-
stone, Hamilton, Illinois.’’,—Worthen, p. 269, pl. 27, fig. 1, 1883.
Scaphiocrinus coxanus Wachsmuth and Springer, p. 237 (161), 1886.
Pachylocrinus coxanus Springer, p. 71, 1926.

Cydrocrinus subramulosus (Worthen), n. comb.
Poteriocrinus subramulosus Worthen, p. 14, 1882.—Worthen, p. 284, pl.
27, fig. 6, 1883. ‘Keokuk limestone, Keokuk, Iowa.”
Poteriocrinus (Scaphiocrinus) swallowt Meek and Worthen (pars). (Cited as
synonym of.) Wachsmuth and Springer, p. 237 (161), 1886.

Geologic and geographic distribution.—The genus as known ranges from
the upper Burlington of Iowa (undescribed species) to the upper Borden of
Indiana and Keokuk of Iowa. ?

Relationships.—Superficially Cydrocrinus most nearly resembles Blothro-—
crinus. The relatively short, compact crown of Cydrocrinus, with its numer-
ous rami, as against the very long, slender crown of Blothrocrinus is a most
obvious difference. In Cydrocrinus all rays normally have two IBr against
the numerous IBr in the ant R of Blothrocrinus. The relatively short, very
stout ventral sac of Cydrocrinus, composed of large heavy plates, isin marked
contrast to the long, slender, fragile sac of Blothrocrinus. The turbinate cup
and para-endotomous arms of Cydrocrinus are in marked contrast to the de-
pressed cup and endotomous arms of Pachylocrinus.

Ascetocrinus, n. gen.

Genotype.—Scaphiocrinus rusticellus White.
Generic diagnosis.—

Crown. Tall, slender, compact, expanding slowly to about three-fourths
its height, then contracting.

Dorsal cup. Subturbinate, with flattened base. Pits at angles of plates.
Very fine linear ornamentation.

IBB. Small, practically concealed by column, but showing in side view. |

BB. Medium size. |

RR. Large; articulating facet extending full width of R, linear to slightly
erescentic; suture slightly gaping.

IBr. One in all rays, high, deeply constricted medially.

Arms. Long, slender, endotomous. Two or three divisions above the prim-
axil. In one specimen in one ray the first admedian ramus divides. Br
high, with marked lateral shoulders for support of pinnules, giving the
ramus a Decadocrinus-like zigzag appearance. Pinnules long, slender.

Post IR. RA large, pentagonal, seated well down between post and r post
BB. X large, extending upward well above the radial facet. RT large,
lying mostly above the level of the radial facet.

Ventral sac. Slender, composed of about eight vertical series of plates on
the posterior side, with pores at angles. Sac about one-half the height
of the arms, recurved.

Column. Stellate in section in proximal position; pentagonal, with concave
faces, changing to circular in section going distad. Nodals prominent.
Lumen pentagonal.

Characteristic species of the genus.—

Aveust 15, 1940 KIRK: NEW CARBONIFEROUS CRINOIDEA 329

Ascetocrinus rusticellus (White), n. comb.
Scaphiocrinus rusticellus White, p. 505, 1863. ‘‘Lower division of the Burling-
ton limestone, Burlington, lowa.’’—Keyes, p. 212, pl. 26, fig. 1, 1894.
Poteriocrinus (Scaphiocrinus) rusticellus Wachsmuth and Springer, p. 113
(338), 1880. |
Abrotocrinus rusticellus Springer, p. 72, pl. 16, fig. 11, 1926.

Ascetocrinus whitei (Hall), n. comb.
Scaphiocrinus whiter Hall, p. 306, 1861.—Hall, p. 7, 186la. ‘Burlington
limestone, Burlington, Iowa.’’ (Lower Burlington.)
Poteriocrinus whitec Wachsmuth and Springer, p. 112 (837), 1880. ‘‘Lower
Burlington limestone.”’

Geologic and geographic distribution.— Known at present only in the lower
and upper Burlington limestones of Iowa.

Relationships.—Springer (1926, p. 72) referred Ascetocrinus rusticellus to
Abrotocrinus Miller and Gurley. Abrotocrinus he defined as a Pachylocrinus
with a pentagonal column, this character being ‘‘not a very reliable one.”’
As a matter of fact, Abrotocrinus is a well-characterized genus. Ascetocrinus
differs from Abrotocrinus in many respects. The compact, narrow crown of
Ascetocrinus is in marked contrast to the irregular, expanded crown of
Abrotocrinus. In Abrotocrinus there are many [Br in the ant Ras against the
single IBr in Ascetocrinus. The dorsal cup of Ascetocrinus is much less de-
pressed than that of Abrotocrinus. The ventral sac in Abrotocrinus is stout,
expanded in its apical portion, and usually spinous. The sac of Ascetocrinus
is very slender, short, and delicate.

Hypselocrinus, n. gen.

Genotype.—Poteriocrinus hoveyz Worthen.
Generic diagnosis.—

Crown. Very high, slender.

Dorsal cup. High, narrowly turbinate.

IBB. High, making up an appreciable part of the dorsal cup.

BB. lLarge;r post B supporting RA on its upper sloping shoulder; post B
supporting X and RA.

RR. Large; articulating facet full width of the R, linear.

IBr. One in all rays, except anterior. Ant ray one [Br or atomous. IBr
long, constricted medially in earlier forms, slightly constricted, broad
and ne in later species (two IBr exceptionally found in one or two
rays).

Arms. Very long, slender, typically not dividing above main dichotom (ir-
regular divisions of rami rarely noted in upper Borden). Ant ray
typically atomous, in some species dichotomous. Br cuneate. An adult
specimen of Hypselocrinus hoveyi with a cup 1.7 em in height has arms
25 cm long. Pinnules long, slender.

Post IR. Three anals in cup; RA high on upper shoulders of post and r
post BB; X large, extending well above level of r post R. RT approxi-
mately one-half within the cup.

326 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 8

Ventral sac. Long, slender, composed of 10 or more vertical series of hexa-
gonal plates, wider than high. Sac six to seven times the height of the
cup. A ventral sac 9.8 cm in length has a width of but 1.2 em.

Column. Circular in section.

Characteristic species of the genus.—

Hypselocrinus arcanus (Miller and Gurley), n. comb.

Poteriocrinus arcanus Miller and Gurley, p. 29, pl. 5, fig. 4, 1890. “Keokuk
Group, Washington County, Indiana.”’

Hypselocrinus boonvillensis (S. A. Miller), n. comb.

Poteriocrinus boonvillensis 8. A. Miller, p. 42, pl. 8, figs. 3, 4, 1891. “‘Keokuk
Group, Booneville, Missouri.”

Hypselocrinus calyculus (Hall), n. comb.

Poteriocrinus calyculus Hall, p. 553, pl. 9, figs. 6a—c, 1858. ‘‘Burlington lime-
stone, Burlington, lowa.’’—Hall, pl. 2, fig. 11, 1860.

Scaphiocrinus calyculus Hall, pl. 6, fig. 1, 1872—Wachsmuth and Springer,
p. 120 (345), 1880.

Hypselocrinus douglassi (Miller and Gurley), n. comb.

Potervocrinus douglasst Miller and Gurley, p. 83, pl. 5, figs. 16, 17, 1896a.
(Madison limestone.)
Decadocrinus douglass: Laudon, p. 68, pl. 7, fig. 7, 1933.

Hypselocrinus hoveyi (Worthen), n. comb.

Poteriocrinus hoveyt Worthen (in Worthen and Meek), p. 516, pl. 29, fie 6,
1875. “Keokuk, Crawfordsville, Indiana.” (Upper Borden.)

Scytalocrinus hoveyt ‘Springer, 10s 145, pl. 16, fig. 8, 1900.

Poteriocrinus (Scytalocrinus) robustus (Hall) (pars)—Wachsmuth and
Springer, p. 118 (343), 1880.

Hypselocrinus maccabei (Miller and Gurley), n. comb.

Poteriocrinus maccabei Miller and Gurley, p. 34, pl. 3, figs. 3-6, 1894.
‘Kinderhook Group, Le Grand, Iowa.’’ (Hampton formation. )

Scytalocrinus maccabei Laudon and Beane, p. 258, pl. 17, fig. 13; pl. 18, figs.
6, 7; pl. 19, fig. 6, 1937.

Poteriocrinus maccabet var. decrepitus Miller and Gurley, p. 36, pl. 3, figs.
9-12, 1894.

Poteriocrinus hammondi Miller and Gurley, p. 35, pl. 3, figs. 7, 8, 1894.

Hypselocrinus macrodactylus (Meek and Worthen), n. comb.

Poteriocrinites (Scaphiocrinus) macrodactylus Meek and Worthen, p. 140,
1869. ‘Lower Burlington, Burlington, lowa.’’—Meek and Worthen, p.
ANS, ley 2, Ol S73)

Poteriocrinus (Scytalocrinus) macrodactylus Wachsmuth and Springer, p.
117 (842), 1880.

Hypselocrinus neglectus (Miller and Gurley), n. comb.

Poteriocrinus neglectus Miller and Gurley, p. 31, pl. 4, figs. 3-5, 1896.
“Keokuk Group, Booneville, Missouri.’’ (Warsaw. )

Avucust 15, 1940 KIRK: NEW CARBONIFEROUS CRINOIDEA 327

Hypselocrinus pleias (Hall), n. comb.

Poteriocrinus plevas Hall, p. 8, 1863.—Hall, p. 57, 1864.—Hall and Whitfield,
p. 173, pl. 12, fig. 8, 1875. ‘‘Waverly group, Richfield, Summit county,
Ohio.”’

Poteriocrinus (Decadocrinus) plecas Wachsmuth and Springer, p. 119 (342),
1880.

Hypselocrinus tethys (Meek and Worthen), n. comb.

Poteriocrinites (Scaphiocrinus) tethys Meek and Worthen, p. 1438, 1869.
“Upper Burlington, Burlington, lowa.’—Meek and Worthen, p. 419,
_pl. 2, fig. 13, 1873.—Wachsmuth and Springer, p. 114 (839), 1880.

Geologic and geographic distribution.—As known, the genus ranges from
the upper Burlington to the upper Borden.

Relationships.—H ypselocrinus among described genera most nearly re-
sembles Scytalocrinus. Scytalocrinus has relatively short, stout arms as
against the long, relatively slender rami of Hypselocrinus. In Scytalocrinus
the IBB lie in a depression in the species with bowl-shaped dorsal cups and
are barely visible in side view in the campanulate species. In Hypselocrinus
the IBB are prominent and high. They form an appreciable part of the cup.
The ventral sac of Hypselocrinus is very long and slender. In Scytalocrinus
the sac is short and relatively stout.

Remarks.—It does not seem desirable to separate those species with two
rami in the anterior radius from the typical form with a single ramus.
Atomous and dichotomous anterior radii are both found in such compact
genera as Scytalocrinus and Decadocrinus, while in Graphiocrinus arm reduc-
tion is carried outside the anterior radius, and one specimen has but seven
rami. In Phanocrinus of the Chester one or more species show but five rami
in a usual 10-rami genus, and in other species an occasional ray, usually the
anterior, is atomous. In this case there could be no valid excuse for separat-
ing the forms generically.

Sladen (1878, p. 245) proposed the new genus Dactylocrinus pro D. loreus
n. sp. = Poteriocrinus tenuis Austin non Miller. The generic name had been
preoccupied by Quenstedt (1876). The genotype is obviously a very young
individual and its affinities uncertain. Such characters as are shown in the
poor illustrations are essentially those of Hypselocrinus. The adult crinoid
may have been quite different, however.

Poteriocrinus fusiformis Hall is referable to this genus. The name was pre-
occupied by Roemer, but no new name should be proposed. The species was
based on a dorsal cup and [Br only and may well be identical with Hypselo-
crinus macrodactylus (Meek and Worthen).

Histocrinus, n. gen.

Genotype.—Poteriocrinus (Scytalocrinus) grandis Wachsmuth and Springer
Generic diagnosis.—
Crown. Compact, high.
Dorsal cup. Subturbinate, plates thin, smooth, or with faint radial plica-
tions.

328 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 8

IBB. Well exposed in side view.

BB. Medium size.

RR. Articulating facet slightly crescentic, extending nearly the full width
of the R. Suture not gaping.

IBr. Twoin all rays except anterior. Ant ray two [Br or atomous.

Arms. Moderately stout, composed of cuneate brachials. Arms dichotomous
except in anterior ray, which may be dichotomous or atomous (typical. )
The Burlington species alone of the known species is dichotomous in the
anterior ray. The pinnules are moderately stout and stand out stiffly
from the ramus.

Post IR. RA is large, resting subequally on post and r post BB. X is of
about the same size as RA and extends well above the level of the RR.
RT is smaller and lies almost entirely above the level of the RR.

Ventral sac. The ventral sac is subcylindrical, expanding slightly distad.
The tube is reflexed, carrying the tip of the sac and the anal opening
well down on the anterior side. On its posterior side the sac shows five
vertical series of plates. This is one of the genera in which Springer
(1900, p. 144) demonstrated the presence of pores in the ventral sac. The
plates at the distal recurved portion of the sac are tumid to subspinous.

Column. The column is circular in section and made up of prominent
series of nodals and internodals.

Characteristic species of the genus.—

Histocrinus grandis (Wachsmuth and Springer), n. comb.

Poteriocrinus (Scytalocrinus) grandis nom. nov. pro Poteriocrinus corey
Worthen 1875, non Poteriocrinites (Scaphiocrinus) coreyi Meek and
Worthen 1869.—Wachsmuth and Springer, p. 118, 1880.

Decadocrinus grandis Wachsmuth and Springer, pl. 7, figs. 4, 5, 1897. (The
figures are, however, Decadocrinus tumidulus (Miller and Gurley).)
Poteriocrinus coreyt Worthen Gin Worthen and Meek), p. 516, pl. 29, figs.

2, 3a, b, 1875.

Histocrinus graphicus (Miller and Gurley), n. comb.

Scaphiocrinus graphicus Miller and Gurley, p. 50, pl. 10, fig. 4, 1890. ‘“Keo-
kuk Group, Crawfordsville, Indiana.” (Upper Borden. )

Histocrinus juvenis (Meek and Worthen), n. comb.

Poteriocrinites (Scaphiocrinus) jguvenis Meek and Worthen, p. 146, 1869.
“Lower Burlington, Burlington, lowa.”—Meek and Worthen, p. 417,
pl. 2, fig. 8, 1873.

Poteriocrinus (Decadocrinus) juvenis Wachsmuth and Springer, p. 119 (842)
1880.

Geologic and geographic distribution.—The genus as known is found in the
Burlington and Keokuk of the Mississippi Valley and the Borden of Indiana.

Relationships.—Histocrinus combines characters diagnostic of the un-
related genera Decadocrinus and Hypselocrinus. The erect, stout arms with
their heavy comblike pinnules are similar to those of Decadocrinus. The
subturbinate cup, the relatively thin, smooth cup plates, and the large
prominent IBB could fall under Hypselocrinus. The ventral sac is struc-
turally like Decadocrinus but is relatively shorter. The presence of two IBr,

Aveust 15, 1940 KIRK: NEW CARBONIFEROUS CRINOIDEA 329

the character of the arms, and the more turbinate cup distinguish H7zsto-
crinus at once from Scytalocrinus.

One must consider the possibility of intergeneric crosses among the
crinoids. This is by no means the only instance of combined generic char-
acters suggesting such a cross.

Phacelocrinus, n. gen.

Genotype.—Poteriocrinus wetherbyz 8. A. Miller.
Generic diagnosis.—

Crown. High, subcylindrical or spreading distad.

Dorsal cup. Subturbinate to campanulate.

IBB. High, making up an appreciable part of the dorsal cup.

BB. Medium size; r post B supporting RA; post B supporting X and RA.

RR. Large; articulating facet full width of R, slightly crescentic, suture
gaping.

IBr. ‘Two or fusing irregularly to one compound brachial, though suture
can often be seen. In both cases the compound brachial is very high
and deeply constricted medially.

Arms. ‘Two undivided rami to the ray is typical. In some of the Chester
specimens irregular branching occurs. Br cuneate, with long, slender
pinnules. The maximum length of arms seems to be about six or seven
times the height of the cup.

Post IR. Three anal plates in cup. RA penetrates well down between post
and r post BB. X extends above level of RR, sometimes separated from
post B in the Chester forms. RT mostly out of the cup.

Ventral sac. Cylindrical, with a maximum height of about four times that
of the cup. Composed of vertical series of hexagonal plates of nearly the
same height and breadth.

Column. Pentagonal in section.

Characteristic species of the genus.—

Phacelocrinus bisselli (Worthen), n. comb.

Poteriocrinites bisselli Worthen (in Meek and Worthen), p. 546, pl. 21, fig.
4, 1873. “Chester, Chester, Illinois.”

Poteriocrinus (Scytalocrinus) bisselli Wachsmuth and Springer, p. 117 (340),
1880.

Phacelocrinus columbiensis (Worthen), n. comb.

Poteriocrinus columbiensis Worthen, p. 22, 1882. ‘‘Chester limestone, near
Columbia, Monroe county, Ill.’’? (Renault.)—Worthen, p. 298, pl. 29,
fig. 6, 1883. 3

Decadocrinus columbiensis Wachsmuth and Springer, p. 239 (163), 1886.

Phacelocrinus dactyliformis (Hall), n. comb.

Scaphiocrinus dactyliformis Hall, p. 670, pl. 17, fig. 6, text fig. 105, 1858.
“St. Louis limestone: St. Louis, Missouri.”’

Poteriocrinus (Scaphiocrinus) dactyliformis Wachsmuth and Springer, p. 112
(335), 1880.

300 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 8

Phacelocrinus decabrachiatus (Hall), n. comb.

Scaphiocrinus decabrachiatus Hall, p. 679, pl. 25, fig. 1, text fig. 106, 1858.
‘Kaskaskia limestone: Kaskaskia, Illinois.”’ (St. Louis limestone fide
Worthen, 1883.)

Poteriocrinus (Scytalocrinus) decabrachiatus Wachsmuth and Springer, p. 117
(340), 1880.

Phacelocrinus gracilis (Troost), n. comb.

Agassizocrinites gracilis Troost, p. 420, 1849. (Nom. nud.)—Troost, p. 62,
1850. (Nom. nud.)—Troost, p. 88, pl. 11, fig. 9, 1909.

Scytalocrinus? gracilis Wood (in Troost), p. 88, 1909. ‘‘St. Louis limestone.
Huntsville, Alabama.”’ (‘“‘Ste. Genevieve.’’)

Phacelocrinus internodius (Hall), n. comb.

Scaphiocrinus internodius Hall, p. 679, pl. 25, fig. 2, text fig. 107, 1858.
‘“‘Kaskaskia limestone: Chester, Illinois.”” (St. Louis limestone fide
Worthen, 1883. Possibly Ste. Genevieve.) Wachsmuth and Springer, p.
113 (836), 1880.

Phacelocrinus longidactylus (McChesney), n. comb.

Scaphiocrinus longidactylus McChesney, p. 7, text fig., 1860. ““Kaskaskia
division of the Carboniferous limestone, Kaskaskia, Illinois.’’-—Mce-
Chesney, pl. 4, fig. 4, 1865.—McChesney, p. 4, pl. 4, fig. 4, text fig.,
1868. ‘‘Chester division of the sub-carboniferous limestone, Kaskaskia,
Illinois.” 3

Poteriocrinus (Scytalocrinus) decabrachiatus Hall. (Cited as synonym of.)
Wachsmuth and Springer, p. 117 (840), 1880.

Phacelocrinus vanhornei (Worthen), n. comb.

Poteriocrinus vanhornet Worthen (in Worthen and Meek), p. 517, pl. 31, figs.
2, 3, 1875. “Upper division of St. Louis group, Alton, Illinois.”

Poteriocrinus (Scytalocrinus) vanhornet Wachsmuth and Springer, p. 118
(343), 1880.

Scytalocrinus vanhorner Keyes, p. 2138, pl. 26, fig. 3, 1894.—Springer, p.
145, pl. 16, figs. 18, 14, 1900.

Poteriocrinus arrectarius Miller and Gurley, p. 33, pl. 4, fig. 6, 1896.

Phacelocrinus wachsmuthi (Wetherby), n. comb.

Scytalocrinus wachsmutht Wetherby, p. 155, pl. 5, fig. 4, 1880. ‘““Kaskaskia
(Chester) Group, Pulaski county, Kentucky.”’ (Glen Dean.)—Wachs-
muth and Springer, p. 238 (162), 1886.

Phacelocrinus wetherbyi (S. A. Miller), n. comb.

Poteriocrinus wetherbyi S. A. Miller, p. 36 (6), pl. 8, figs. 1, la, b, 1879.
‘Kaskaskia, Pulaski county, Kentucky.”’ (Glen Dean.)

Poteriocrinus (Scytalocrinus) wetherbyz Wachsmuth and Springer, p. 118
(343), 1880.

Geologic and geographic distribution.—Phacelocrinus ranges as known from
the St. Louis to the upper middle Chester (Glen Dean) of the Mississippi
Valley, Alabama and Kentucky. An undescribed species is known in the
Greenbrier of Maryland.

Aveust 15, 1940 KIRK: NEW CARBONIFEROUS CRINOIDEA dol

Relationships.—Phacelocrinus in its simplicity of form resembles Hypselo-
crinus. No one, after handling specimens of the two genera, would have
difficulty separating them, even in the case of dorsal cups. The arms of
Phacelocrinus are relatively shorter and stouter than in Hypselocrinus in
comparing similar stages in phylogeny. In Phacelocrinus the [Br are either
clearly two in number or have fused into a high, compound brachial, deeply
constricted medially. This lack of fusion of the two plates is very apparent
in the St. Louis species, though the degree of fusion varies among the rays
of the same specimen. Even in the Chester, however, either through incom-
plete fusion or fracture on the suture, the compound nature of the brachial
is often seen. In Hypselocrinus there is but a single primibrach, and when
occasionally a variation showing two primibrachs is found in one or two
rays, the second is obviously an additional brachial. The gaping articulating
sutures of Phacelocrinus are in marked contrast to the close union in
Hypselocrinus as usually seen. The column in Phacelocrinus is pentagonal in
section, as against the round column of Hypselocrinus. The ventral sac of
Phacelocrinus is relatively shorter and composed of larger plates than in
Hypselocrinus.

- Pegocrinus, n. gen.

Genotype.—Poteriocrinus bijugus Trautschold.
Generic diagnosis.—

Crown. High, compact, arms attaining a length 15 times or more that of
the dorsal cup.

Dorsal cup. Broadly turbinate. Plates thick.

IBB. Small, partially anchylosed into a firm disk, clearly visible in side
view. Sutures well shown.

BB. Variable in size and shape, barely meeting laterally or on very short
faces. The post B is high and narrow, supporting RA on its long, upper
right shoulder. Above it barely meets anal X. R post B is large, sup-
porting RA on a narrow face.

RR. Large. Articulating facet the full width of the R. Suture not gaping.
The upper surface of the R is very deep. There is a prominent fulcral
ridge and a deep ligament pit. There is a large, well-defined pair of
muscular fossae. The apposing edges of the R and IBr along the suture
are strongly crenulate.

IBr. Variable in number without respect to the rays. Two or three in
number, usually two.

Arms. Long and massive, two undivided rami to each ray. Br low, quad-
rangular or with slightly sloping faces, each bearing along, stout pinnule.
Food groove shallow, no axial canal. Margins of Br crenulate.

Post IR. RA small, elongate, narrow. Anal X small, high, narrow. RT
small, in some cases barely contacting RA.

Ventral sac. Poorly known. One specimen shows the partially dissociated
plates in approximately their original position. According to this, the
sac is short, erect, and fairly stout in relation to the dorsal cup. The
plates are small and thin, forming a fragile structure.

Column. Circular in section, with stellate lumen.

Characteristic species of the genus.—

302 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 8

Pegocrinus bijugus (Trautschold), n. comb.

Poteriocrinus bijugus Trautschold, p. 14, pl. 4, figs. 1-8, text fig., 1867.
Junger Bergkalk (Moscovian), Miatschkowa, Russia.—Trautschold,
text fig., 1879.—Jaekel, p. 64, fig. 54a, 1918.

Poteriocrinus (Scytalocrinus) bigugus Wachsmuth and Springer, p. 118 (348),
1880.

Geologic and geographic distribution.—The genus is known at present only
in the Moscovian (Pennsylvanian) of Russia.

Relationships.—There seems to be no described genus with which Pego-
crinus may be confused. Two poteriocrinoid genera associated with Pego-
crinus are readily distinguished, although one of them was apparently con-
fused with it by Trautschold. Both Moscovicrinus and Ophiurocrinus have
high, rather narrowly turbinate dorsal cups as against the broadly turbinate
cup of Pegocrinus. The IBB of Pegocrinus, meeting on very narrow faces and
practically anchylosing into a solid disk, are in marked contrast to the high
IBB of the other two genera. The patterns of the plates of the posterior IR
are unlike. Moscovicrinus has branching arms and a remarkable circinate
ventral sac, both wanting in Pegocrinus.

Of the original specimens figured by Trautschold (1867, pl. De specimen
figure 2 was sent by Trautschold to Wachsmuth and is now in the Springer
collection in the United States National Museum. Jaekel (1918, p. 64, fig.
54a) states that his figure is from the “‘holotype” in Breslau. It could not be
either specimen figures 2 or 3. Owing to the great number of inaccuracies in
this work of Jaekel, it is doubtful that it really represents any one of the
three specimens of Trautschold. At any rate, Jaekel does not indicate which
specimen he chose as type. Owing to the uncertainties of the matter, I have
chosen specimen figure 2, plate 4, of Trautschold (1867) as holotype.

The specimen figured by Trautschold (1879, pl. 14, fig. 3) as Poterzocrinus
bajugus is certainly not that species. It probably is referable to Potertocrinus
originarius, for which Jaekel erected the genus Ophiurocrinus.

LITERATURE CITED

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(1859) 1860.

Haun, James. Descriptions of new species of Crinoidea from the Carboniferous rocks of
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Hau, JAMES. Descriptions of new speceis of Crinoidea; from investigations of the Iowa
Geological Survey. Preliminary notice (privately issued, Albany, N. Y.), pp. 1-12,
February 14, 1861. Complete paper, pp. 1-18. February 25, 13861a.

Hau, James. Preliminary notice of some spectes of Crinoidea from the Waverly sand-
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of New York. Pp. 1-11. ‘‘Printed in advance, from the 17th Report on the New
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Hau, JAMES. Preliminary notice of some species of Crinoidea from the Waverly sand-
stone series of Summit County, Ohio, supposed to be of the age of the Chemung group
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Aveust 15, 1940 KIRK: NEW CARBONIFEROUS CRINOIDEA 300

Haut, JAMES. ‘Photographic plates.’ Plates1—7. Privately issued. Plates distributed
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JAEKEL, O. Phylogenie und System der Pelmatozoen. Paleont. Zeitschr., Band 3,
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Keyes, C. R. Paleontology of Missouri. Part 1. Missouri Geol. Survey, vol. 4, pp.
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Laupon, L. R. The stratigraphy and paleontology of the Gilmore City formation of Iowa.
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Laupon, L. R., and Bransz, B. H. The crinoid fauna of the Hampton formation at Le
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McCuesney, J. H. Descriptions of new species of fossils, from the Paleozoic rocks of
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McCuesney, J. H. Plates illustrating in part the new species of fossils, from the Paleo-
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McCuesney, J. H. Descriptions of fossils from the Paleozoic rocks of the Western States,
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Meek, F. B., and WortTHEN, A. H. Descriptions of invertebrates from the Carboniferous
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1866.

Merk, F. B., and WorTHEN, A. H. Descriptions of new Carboniferous fossils from the
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Miuier, 8. A. A description of some Lower Carboniferous crinoids from Missourt.
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Miter, 8. A., and Guruey, W. F. E. Description of some new genera and species of
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Miter, 8. A., and GurutgEy, W. F. E. New species of crinoids from Illinois and other
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Mier, 8. A., and GurLEy, W. F. E. New species of Echinodermata and a new crus-
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SHumMARD, B. F. A catalogue of the Palaeozoic fossils of North America. Trans. Acad.
Sci. St. Louis, vol. 2, no. 2, pp. 334-407. 1866.

SLADEN, W. P. On the genus Poteriocrinus and allied forms. Proc. West Riding
Geol. aS Polytech. Soc., new ser., vol. 1, pp. 242—253, pl. 10. 1878. (Reprint
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SPRINGER, FRANK. On the presence of pores in the ventral sac in fistulate crinoids.
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334 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 8

SPRINGER, FRANK. Unusual forms of fossil crinoids. Proc. U. 8. Nat. Mus., vol. 67,
art. 9, pp. 1-137, pls. 1-26. February 15, 1926.

TRAUTSCHOLD, HERMANN. LEHunige Crinoideen und andere Thierreste des jungeren Berg-
kalks 1m pa ae Moskau. Soc. Imp. Nat. Moscou Bull. 3, pp. 1-49, pls.
1-5. 1867.

TRAUTSCHOLD, HERMANN. Dive Kalkbriiche von Miatschkowa. Eine Monographie des
oberen Bergkalks. Soc. Nat. Moscou Nouv. Mém., vol. 14, pp. 101-180, pls.
12-18. 1879.

Troost, GERARD. Communication. Amer. Journ. Sci. and Arts, ser. 2, vol. 8, no. 24,
pp. 419-420. November 1849.

Troost, GERARD. A list of the fossil crinoids of Tennessee. Proc. Amer. Assoc. Adv.
Sci., 1849, pp. 59-64. 1850.

Troost, GERARD. A critical summary of Troost’s unpublished manuscript on the cri-
noids of Tennessee. U.S. Nat. Mus. Bull. 64, pp. i-xi, 1-50, pls. 1-15. 1909.
(Edited by Elvira Wood.)

W ACHSMUTH, CHARLES, and SPRINGER, FRANK. Revision of the Paleocrinoidea. Pt. 1,
pp. 1-153, pls. 1-3 (15-17). Proc. Acad. Nat. Sci. Philadelphia, 1879, pp. 226—
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WACHSMUTH, CHARLES, and SPRINGER, FRANK. Revision of the Paleocrinoidea. Pt.
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1886.

WacHSMUTH, CHARLES, and SPRINGER, FraNK. The North American Crinoidea
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Weruersy, A. G. Remarks on the Trenton limestone of Kentucky, with descriptions of
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Wuitsr, C. A. Observations on the summit structure of Pentremites, the structure and
arrangement of certain parts of crinoids, and descriptions of new species from the
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Nat. Hist.), vol. 7, pp. 481-506. 1868.

WHITFIELD, R. P. Description of a new species of crinoid from the Burlington limestone,
at Burlington, Iowa. Bull. Amer. Mus. Nat. Hist., art. 1, pp. 7-9, pls. 1-2. 1881.

Woop, Exvrra, in Troost, Grrarp. A critical summary of Troost’s unpublished
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WortTHEN, A. H., in Merk, F. B., and WortHen, A. H. Descriptions of invertebrates
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pp. 265-338, pls. 27-30. 1883.

Aveust 15, 1940 ALLARD & ALLARD: CHROMOSOME NUMBER OF BEAN 3309

BOTANY.—The wild bean Phaseolus polystachyus (L.) B.S.P.: Its
chromosome number.! H. A. ALLARD and H. F. ALuLarp, U. 8.
Department of Agriculture.

Eleven species of beans of the genus Phaseolus are known to be
more or less generally cultivated in various parts of the world, all of
tropical, subtropical, or warm-temperature origin and sensitive to
frost. Considerable work has been done by various investigators to
determine the chromosome number of the cultivated species, as re-
ported by J. Kawakami in his paper, Chromosome numbers in Legumi-
nosae (Bot. Mag. Tokyo 44: 319-329. 1930). It appears that in all
our well-known species of beans of the genus Phaseolus, 22 chromo-
somes is the somatic or 2n number. Among those reported are the
species P. acutifolius, angularis, aureus, coccineus, lunatus, and
mungo, as well as varieties of some of these. Dolichos lablab also has
22=(2n) chromosomes, as reported by Karpetschenko (1925) and
Kachidse (1925).

In the Eastern United States from Connecticut southward to the
Gulf, a native wild bean (Phaseolus polystachyus) occurs, which is of
considerable interest in some of its behaviors. Although stated other-
wise in Gray’s Manual of botany and other botanical works, a char-
acteristic of its germination is the hypogean life of its cotyledons,
whereas the cotyledons of most of our beans have an epigean history.
The Searlet Runner bean (P. multiflorus), however, is an exception to
this, like the wild bean.

The senior writer has found this bean to be an occasional constit-
uent of the flora of the lower mountain areas of Virginia, within the
Upper Piedmont physiographic province, and he has studied its ecol-
ogy and length-of-day behaviors in some detail.

It is eminently unsuited to tropical lengths of day of 12 hours or
less and to the winter conditions of daylight in the greenhouses of our
northern latitudes. Under such conditions its growth is so slow that it
becomes a much dwarfed and practically a dormant plant, devoid of
the typical twining habit, and producing few or no flowers.

The rootstock of this bean is perennial, the aerial stems dying down
to the ground, where, under favorable conditions of protection, winter
buds are formed that originate the new stems of the next summer.
The laying down of dormant buds and the gradual dying of the aerial
parts at the close of the summer season are nicely adjusted behaviors
dependent upon shortening days and lowering mean temperatures.

1 Received February 1, 1940.

306 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 8

The suppression of vegetative elongation together with normal flower
production when the days are sufficiently shortened artificially repre-
sents extremes of this late-summer behavior. As a matter of fact,
when germination takes place in the greenhouse in winter time, the
seedling plants remain very small and almost dormant, tending to
produce thickened rootstocks with the downward translocation of
material. It would appear that there can be no utilization in upward
elongation and flowering as in the summer season of long days and
high mean temperatures.

In nature the wild bean in the upper Piedmont area is ecologically a
plant of deciduous woodlands and warm well-drained rocky slopes.
It will grow readily in the open in rich garden soils, however, and re-
main in a flourishing condition until autumn arrives, but the roots
readily freeze and rot in wet, unprotected garden soils.

Cold, wet, exposed soils and the aggressions of vigorous competi-
tors do not favor the survival of this bean in open grassland fields and
pastures. Even in the woodland environment of the Piedmont up-
lands, where it is driven, its scarcity would indicate precarious con-
ditions of survival.

A study of the chromosome number of this bean was made by the
junior author, and the n number of the pollen mother cells was found
to be 11. The somatic (2n) number, then, is 22, which, as previously
stated, is that of all our common garden varieties reported upon by
Kawakami. The chromosomes of the wild bean are of small size.

On the basis of the constant chromosome number of the species of
Phaseolus, so far as such determinations have been made, this genus
appears to be a relatively stable one. It might also be assumed that
it is of more recent evolution, without marked specialization in this
direction as yet, owing in part perhaps to uniformity of habitat.

There is reason to believe that the genus is definitely of warm-cli-
mate origin, since the center of distribution is within the Tropics. Those
beans that have been investigated have been shown to grow and to
flower readily in response to lengths of day of 12 hours or less, thus
adapting them to tropical and subtropical conditions of daylight.
Some species and varieties, in addition, are able to flower in response
to the much longer days of high latitudes. In this respect these beans
are either indeterminate in their day-length requirements or are short-
day plants with very high critical length-of-day limitations that
favor flowering far north of the Tropics.

Other species or varieties of beans are more rigidly confined to the

Avuaust 15, 1940 ALLARD & ALLARD: CHROMOSOME NUMBER OF BEAN 337

Tropics or to low latitudes, owing to the narrow limits of their length-
of-day adaptations.

Several tropical lima beans (Phaseolus lunatus) from Peru (F.S.P.I.
no. 46153) and Ecuador (F.S.P.I. no. 46339) reported upon in 1920
by Garner and Allard? were found to be unable to flower in the lati-
tude of Washington, D. C., owing to the fact that these were short-
day plants with a rather narrow range of lengths of day favorable to
flowering. In other words, these beans were adapted only to tropical
lengths of day or to those lengths of day of low latitudes, so far as
sexual reproduction was concerned.

A bean of the species Phaseolus multiflorus, also from South Amer-
ica, reported upon by Garner and Allard? in 1923, showed a similar
tropical short-day behavior, since it could not flower at Washington,
D. C., when long days were experienced, but was induced to flower
late in summer with 10 hours of daylight afforded each day. This
variety, also, when afforded only 10 hours of daylight, showed a
marked reduction in aerial stem growth and the roots became strik-
ingly tuberiferous (loc. cit., pl. 6). With 12 hours of light there was
less tuberization, and with exposure to the full length of the summer
day, and likewise to artificially lengthened winter days, by the use
of electric light, there was none.

The native wild bean of the eastern North American territory can
be definitely excluded from the tropical category. This bean, with its
present constitutional inability to grow and to fruit on lengths of day
around 12 hours or less, could not well have originated in the Tropics
or in equatorial regions. There is reason, then, to believe that it is a
somewhat isolated outlying offshoot of the great tropical group, since
it not only must have the greatly increased lengths of day of higher
latitudes to persist, but its greater resistance to winter cold than most
beans can withstand is an additional adaptation for the habitat of a
temperate climate.

2 Garner, W. W., and Atuarp. H. A. Effect of the relative length of day and night
and other factors of the environment on growth and reproduction in plants. Journ. Agr.
Res. 18(11): 600. 1920.

3’ GARNER, W. W., and AtuarpD. H. A. Further studies in photoperiodism, the re-

soe of the plant to relative length of day and night. Journ. Agr. Res. 23(11): 893.
1923.

338 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 8

MYCOLOGY.—A ringworm disease of muskrats transferable to man.
Vera K. Cuarzes, U.S. Bureau of Plant Industry.

In the fall of 1936 a culture of a fungus apparently pathogenic to
man was received from Dr. Paul L. Errington, of Iowa State College
of Agriculture and Mechanic Arts. Dr. Errington had been working
with muskrats (Ondatra zibethica) near Ruthven, Iowa, which were
affected with a serious skin disease. The culture was made from his
arm, which had become infected with a type of ringworm, and as the
infected arm was the one he had habitually used in examining nests
of muskrats and handling the young, it occurred to him that he might
have contracted the disease affecting the muskrats. The culture sub-
mitted was somewhat old and contained only one type of spores,
which were small and 1-celled, suggesting the microconidia of a ring-
worm fungus. The culture was transferred to Sabouraud and corn-
meal agar, and in the course of 3-5 weeks developed other types of
organs and spores characteristic of the genus T'richophyton.

APPEARANCE ON MAN

The appearance of ringworm was first observed by an assistant
who spent July 4, 1935, on the marsh helping to capture for tagging
the more active sizes of young animals. In this case definite “rings”’
appeared on her legs. While this would seem a more unlikely place
than exposed arms, the work of helping capture the animals for
tagging often necessitated her wading in the water about the lodges,
and in this manner infection could have easily taken place.

Dr. Errington describes the history of his own case as follows:

On June 24, 1936, I noticed a glazed-appearing area about 20 mm in
diameter on the back of my left hand; by the next day this was almost com-
pletely surrounded by an inflamed ring. A similar area was to be seen higher
up on my arm and a smaller one near the elbow. My left arm was usually
more or less scratched, as it was the one I used in digging into and feeling
around in the muskrat lodges in connection with my research. I had handled
badly diseased animals on several occasions during the preceding two weeks
and had been in the habit of washing thoroughly in soap and water and
touching recognized scratches with iodine each evening after coming in
from the field.

The evening of June 25, I applied a salve extensively used for the treat-
ment of burns, etc., but by morning the two eruptions were enlarged and
itching. I went to a local physician, who gave me three different ointments
to try, none of which proved more than partially efficacious. These were,
I think, preparations containing salicylic acid, but the thick greasy base
evidently prevented really effective penetration. The result was that salves
strong enough to have even a superficial effect blistered my skin severely

1 Received February 5, 1940.

Auaust 15, 1940 CHARLES: A RINGWORM DISEASE OF MUSKRATS 309

and yet did not seem to reach the organisms intrenched at the base of the
hairs; at any rate, whenever I would try to give my skin a rest, fungus
activity would start up again.

I had occasion to make a trip to Ames about July 8 and while there called
on two other physicians. By this time, I was in misery, with 32 square inches
of my left arm involved either by the fungus or dermatitis from the salve.
One of the physicians directed me to pull out the hairs on the raised fungus
infected areas and to keep the arm wet-packed with an aqueous solution of
allantoin and also to apply 2 percent allantoin and 1-10,000 phenyl mercuric
nitrate in a greaseless base as an ointment. The almost intolerable itching
was relieved, and, in a few days, the troublesome areas consisted only of the
raised foci of the fungus infection. These foci, however, remained unyielding
to treatment prescribed so far and were four in number, namely, two raised
rings, 35 mm in diameter on back of hand and lower arm, a solid raised area
20 mm in diameter near the elbow, and a fourth area of about 8 mm at the
base of the little finger. All were firm or hard to the touch and dark red, the
larger ones being purplish red and spotted by pus pockets.

It was Dr. Errington’s experience that there was an apparent dif-
ference in the effectiveness of a penetrating solution and a thick
salve even when salicyclic acid was the active chemical used in both.
He stated in a subsequent letter that on occasions of suspected ex-
posure he had washed his hands in 1:500 aqueous solution of mer-
curic iodide, which appeared to give protection.

APPEARANCE OF ANIMALS

Dr. Errington reported that during the midsummer of 1935 a
considerable number of young muskrats under observation on Round
Lake near Ruthven were found to be suffering heavy mortality from
a skin disease:

Advanced cases were marked by underweight, by watery swollen pustules
on ventral surface or by hairlessness about legs, base of tail and under-parts.
Incipient cases in individuals of apparently normal size and health may at
times be detected by a slight baldness or dandruff-like scurf on head or
back or possibly by one or a few watery blisters on the end of the tail (4).
[Later he wrote] A skin disease (I think it is only one but there may be
more) affecting young muskrats has been found in about 10 percent of the
litters examined. When the disease is represented in a litter, usually a third
to a half of the members are affected, though sometimes the whole litter.
Mortality seems to be complete if the muskrats contract the disease while
less than ten days old; larger young may recover, little the worse for the
experience except perhaps for a slightly bobbed tail.

In very young animals, the disease seems characterized by dermatitis
pustules, usually on the underparts. Young thus affected typically cease
growing and soon disappear. In animals of around two weeks, pustules are
seldom seen except on the bare scaly tails, but either a dry, dandruff-lke
scurf may be noted or a localized loss of hair, especially about the extremi-
ties, underparts, and base of tail. This sort of case is commonly attended
by stunted growth and ultimate death. When fairly large young have only
a blister or two on the tail the case is not so likely to be serious, but some

340 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 8

animals with only this slight degree of visible infection may become stunted
and die also.

It is believed that only one organism was involved in the skin dis-
ease of the muskrats because only one organism was consistently
found in the material submitted for examination, and in the cultures
isolated from the skin and hair of diseased animals. All the other
fungi that appeared in culture or were present on the fresh material
were clearly secondary. A species of Fusarium was observed in one
culture, but it is not thought that a species of this genus would pro-
duce the clinical symptoms presented in this case.

ISOLATION OF THE FUNGUS

The first specimen of muskrat material was received by the writer
in January 19387 and consisted of a dry skin of a muskrat that had
been affected by a ringworm fungus but had recovered from the
attack. Cultures were made from this skin, but no pathogenic fungus
was recovered. As the season for work with very young muskrats had
passed, the mycological studies had to wait until the following year.
In May 1938, Dr. Errington reported that in his study of muskrats
in the area near Ruthven, Iowa, he had found no evidence of muskrat
skin disease, but that he had expectations of finding infected animals
late in June or in July, when past experience indicated that the
trouble would be more widespread and conspicuous. Anticipating this
reappearance of the disease, tubes of culture media were forwarded to
Dr. Errington in order to enable him to make cultures direct from
the animals in the field. This supposition proved to be correct, and
in the later part of June 1938 cultures made from skins of diseased
muskrats from Round Lake, Iowa, were sent to the author. Six cul-
tures were received at this time, but no pathogenic organism was
found, although the cultures were made from the scurf. However, this
may have been due to the fact that the work in the field presented
many opportunities for contamination by vigorous but nonpatho-
genic fungi. Still believing that a species of Microsporum or a related
fungus was the cause of the trouble, the writer requested more cul-
tures from diseased animals, and on July 14 and 16 twelve additional
cultures were received. One culture made from hairs from the bare
foot of a muskrat, removed by heat-sterilized tweezer-tips, was sug-
gestive of Trichophyton, and after being cultured it proved to belong
to this genus. When this culture was received it contained only micro-
conidia, but transfers made to other culture media developed a
striking polymorphism. The original culture was found to be ex-

Aveust 15, 1940 CHARLES: A RINGWORM DISEASE OF MUSKRATS 341

ceedingly vigorous and grew luxuriantly on various culture media.
Cultures were also made from a small dried piece of skin from a dis-
eased muskrat. Pure, vigorous growths of the fungus were obtained
from these isolations.

TERMINOLOGY OF CERTAIN STRUCTURES IN
THE DERMATOPHYTES

The polymorphic character of many human and animal fungus
pathogens has led to a more or less specialized terminology of the
different organs. In order to avoid any confusion the terms employed
in this paper may be defined as follows:

Thyrse, a contracted paniclelike spore bearing structure.

Microconidia (Aleurospores) may be acrogenous or pleurogenous and are
borne in thyrses. They are small, 1-celled, and spherical to oval or clavate
in shape. The term aleurospore has been applied to these spores and is em-
ployed in most papers on medical mycology. They are generally produced in
large numbers and when they predominate impart a powdery appearance
to the surface of the culture.

Macroconidia are referred to as closterospores or fuseaux. They are
elongate, septate, and thin or thick walled. The ends are blunt, and the
base is broad and encircled by a collar, which marks the point of attachment
to the conidiophore. Constrictions are sometimes evident at the septa.

Chlamydospores, so-called, may be formed in the hyphae or they may be
terminal. They do not possess the thick wall characteristic of true chlamydo-
spores and when intercalary resemble the vesiculose cells of species of the
genus Fusarium.

Arthrospores are rows of undifferentiated hyphal cells, which function in
the distribution of the fungus and in carrying it over periods of unfavorable
conditions.

Nodular organs may resemble haustoria in shape or may appear as a
tangle of hyphae resulting from a number of short branches near the tip
of the hyphae.

Spirals are produced from vegetative hyphae and may consist of loosely
or tightly coiled hyphae with smooth thin walls. It has been suggested that
these structures may be connected with an ascogenous stage, but this has
not been demonstrated. Spirals occur in several species of Dermatophytes
and are not regarded as characters useful in the taxonomic distinction of
species.

CULTURAL CHARACTERS

The organism from muskrats was grown in culture for over a year
and a half. On solid media the early cultures produced a powdery
growth at first white, later becoming cream. On liquid media the
growth was more cottony, although microconidia were developed
rather abundantly. Subsequent cultures during the course of some 15
months showed less tendency to form macroconidia and other organs
but continued to produce microconidia in abundance. An exception

042

JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

VOL. 30, No. 8

to this rule was observed in the case of cultures grown on egg-meat
broth. The original culture was made on Sabouraud and transferred
to egg-meat on September 24, 1938, on which it produced a white,
fluffy growth at the surface of the medium, an abundance of micro-
conidia but no coils. Transferred to Sabouraud on November 10,
1939, the growth was rapid, vigorous, and characteristically poly-
morphic, producing microconidia, macroconidia, and short coils. It is
interesting to note that after this length of time and after transfer to
semiliquid media and return to solid media (Sabouraud) the macro-
conidia were noticeably longer and more septate than in the original
cultures or in the cultures grown continuously on solid media. These
cultures had been kept in an ice-box at a temperature of about 40°F.
for a period of a year.

TABLE 1.—TyrE oF GROWTH ON NINE MEDIA

Medium Type of growth Microconidia Macroconidia | Spirals * Color of medium

Blood serum....| Growth felty | Abundant on | Present, 3-4 Absent. No change.
when later pow- | sides of walls in | septate.
dery. young cultures.

Corn-meal agar.| Growth scanty; | Abundant. Absent. Present but few | Not colored.
chlamydospores and not highly
well developed. developed, about

3-4 coils.

Sabouraud....... Cottony at first | Abundantslight- | Present; in | Weak develop- | Reverse of
later powdery; | lylargerandless | young cultures | ment. medium wine
chlamydospores | uniformly spher- | 2-3 celled. red when old.
present. ical than in

growth on other
media.

IRGCehas Se secbncen Growth rapid at | Abundant. Few, 2-3 celled. | Numerous. Back of old
first, pure white, cultures tan.
finally cream;
aerial mycelium
in cultures car-
ried over one
year remained
white.

Potato dextrose.| Growth white, | Abundant. Fair develop- | Absent. Little change,
cottony, finally ment of blunt back of old
powdery at top 4-septate co- cultures dark.
of culture; lux- nidia.
urlant develop-
ment of chlamy-
dospores.

iBrainiviedl aera: Mycelium luxu- | Numerous. Blunt or slight- | Absent. No change.
riant; chlamydo- ly pointed.
spores abundant.

Egg-meat.......| Growth at top | Numerous (ae- | Absent. Absent. No change.
of culture fluffy, | rial spores
white. smaller).

Rosenau Dext. | Growth mostly | Numerous. Few, mostly | Few, short.

brain broth..... at surface of me- 5-septate.
dium, white
then cream;
chlamydospores
few.

Phenol-red tar- | Growth white. Abundant. Fair, develop- | Absent. Little change.

trate agar...... ment, mostly

3-septate.

Auaust 15, 1940 CHARLES: A RINGWORM DISEASE OF MUSKRATS 343

IDENTITY OF THE FUNGUS

As mentioned in the introduction the cultures isolated by Dr.
Errington from his arm and from the muskrat after numerous culture
studies were determined as belonging to the genus Trichophyton ac-
cording to Emmon’s classification. This author (8) recognizes three
groups of Dermatophytes, which are closely related but may be
separated by the distinctive types of macroconidia that they form
in culture. These three types are represented by the following genera:

Trichophyton (Malmsten, 1845) (6).
Epidermophyton (Sabouraud, 1907) (8).
Microsporum (Gruby, 1848) (5).

In the genus Trichophyton the mycelium is generally hyaline though
it may be yellow, violet, or even brown. The reproduction is princi-
pally by small conidia, the so-called microconidia. The macroconidia
are clavate and thin-walled but are not always formed and are gener-
ally absent in old cultures or those that have been repeatedly trans-
ferred.

The genus Epidermophyton is characterized by the oval to egg-
shaped, smooth, thick-walled macroconidia. The mycelium is usually
yellow.

Species of the genus Microsporum are easily determined by the
mostly numerous, spindle-shaped, thick-walled macroconidia and the
clavate conidia. In certain species of this genus, however, the macro-
conidia may be few or abortive. The mycelium is generally hyaline,
or according to certain authors it may range from hyaline to brown.

The organism isolated from the muskrat and grown over a period
of 14 years in pure culture conformed to the generic characters of the
genus Trichophyton and was identified as T. mentagrophytes (Robin)
(7) Blanchard (1). 3

The identity of the two organisms, the one isolated from man and
the other from muskrats, was proved by comparative cultural studies.
All probable sources of the case of human infection other than the
muskrats were eliminated. It was observed that the strain obtained
from the muskrat produced a much more rapid and vigorous growth
than the one isolated from man. The longevity of the former organ-
ism was also much more marked.

A very similar case of the occurrence of this fungus on an animal
host has recently been described by DeLamater (2) on common gray
squirrels living on or near the Johns Hopkins University Campus at
Baltimore. The author described the virulence of the fungus on squir-

344 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 8

rels and stated that cats and rabbits were shown to be susceptible to
the strain and in two cases accidental infection of human subjects
occurred. There was no mention of any mortality of the squirrels.

As far as our information goes this is the first report of T. menta-
grophytes on muskrats. This occurrence is not only of interest because
of the economic importance of the host, but because it presents an-
other record of the transference of an animal parasite to man.

LITERATURE CITED

1. BLANCHARD, RAPHAEL. Parasites végétaux a lexclusion des bactéries. In Bov-
CHARD, CHARLES, Traité de pathologie général 2: 811-926. 1896.

DeLamater, HE. D. The squirrel as a new host to a ringworm fungus. Mycologia
31: 519-526. 1939.

Emmons, C. W. Dermatophytes. Natural grouping based on the form of the spores and
accessory organs. Arch. Derm. and Syph. 30: 337-362. 1934.

ERRINGTON, Paut L., and CaroLtyn Storm Errineton. Experimental tagging of
young muskrats for purposes of study. Journ. Wildlife Management 1: 49-61.
1937.

5. GruBy, Davin. Recherches sur la nature, la siege et le développement du Porrigo

decalvans ou Phyto-alopecte. Compt. Rend. Acad. Sci. 17: 301-303. 1843.

6. MaumstEn, P. H. Trichophyton tonsurans harskérande Mégel. Stockholm, 1845.
Translated by F. C. H. Crepiin: Arch. Anat. Physiol. Wiss. Med. (J. MULLER)
1848, pp. 1-19, 1 pl.

7. Rosin, CHAruES. Histoire naturelle des végétaux parasites. Paris. 1853. 430 pp.

8. SapouraAup, R. Sur lVeczeema marginatum de hebra, trichophytie inguinale et son
parasite econ inguinale Sab.). Arch. Med. Exp. Anat. Path. 19: 565—
586. 1907.

Het Co cel,

PALEOBOTANY.—WNew species and changes of name in some Ameri-
can fossil floras... Rotanp W. Brown, U.S. Geological Survey.

Information uncovered by the writer during the past few years
involves a number of new species that seem worthy of immediate re-
port, new occurrences of described species that shed further light on
the floras in which they occur, and changes of name demanded by un-
equivocal evidence. In the study of the material assembled here the
writer has been aided in part by the generous cooperation of W. R.
Maxon, of the National Herbarium; H. A. Gleason and colleagues, of
the New York Botanical Garden; J. B. Reeside, Jr., of the Geological
Survey; F. M. Carpenter, of Harvard University; and D. I. Axelrod,
National Research Council Fellow.

OSMUNDACEAE
Osmunda occidentale (Berry) Brown, n. comb. Fig. 1

Asplenium occidentale Berry, 4, p. 236, pl. 49, figs. 3, 4.
Fern fragment. Knowlton, 25, p. 24, pl. 9, fig. 10.
Pteris sp. Berry, 4, p. 237.—Idem, 6, p. 103.
When compared with the pinnules of the royal fern, Osmunda regalis

1 Published with the permission of the Director, Geological Survey, United States
Department of the Interior. Received February 28, 1940.

Auveust 15, 1940 BROWN: SOME AMERICAN FOSSIL FLORAS 045

Linnaeus, these specimens show such close resemblance that little doubt
as to their being Osmunda can be entertained. This identification harmonizes
with the general ecological setting of these floras.

Occurrence.—Latah formation, Washington and Idaho. Also lake beds in
the Lemhi Valley, Baker, Idaho (Fig. 1).

POLYPODIACEAE
Adiantum anastomosum Brown, n. sp. Fig. 2

Frond or pinna 10 cm or more in diameter, strongly cordate at the base;
deeply lobed. Lobes apparently 8, long, narrow, of unequal length, lygodioid
in form. Venation without a midrib, reticulate, anastomosing freely. Sori on
the margins of the lobes narrow, in discontinuous patches. Margin entire.

Except for the more cordate base, this frond is almost identical with that
described by Gardner and Ettingshausen (19, p. 42, pl. 11, fig. 1) as Hew-
ardia regia, from the middle Eocene Middle Bagshot beds at Bournemouth,
England. Hollick (21, p. 41, pl. 6, figs. 1-5) has described what seems to be
a species of this genus as Anthrophyopsis hamiltonensis, but it has ligulate
or lanceolate foliage. However, as no sori are present, there is no positive
assurance that Hollick’s specimens are related to Adzantum.

Botanists regard Hewardia as a section of Adiantum. The fossil species
may be compared with a number of living species, but there is no complete
agreement with any. Adiantum cordatum Maxon, from Panama, has cordate
pinnae with entire margins, no tendency to lobing, and with a distinct
midrib. Adiantum wilsoni Hooker, from Jamaica, is similar to cordatum but
has smaller, more numerous pinnae, with serrate margins. Adiantum adi-
antoides (J. Smith) Christensen, from Guiana, is similar to wilsonz, but the
pinnae are not serrate. Adiantum olivaceum Baker, from Guiana, has disjunct
sori, giving the margins of the pinnae a toothed appearance.

The close correspondence between Adiantum anastomosum and Hewardia
regia suggests a comparison of the two fossil floras of which these ferns are
members. The Bagshot beds at Bournemouth, according to a citation by
Reid and Chandler (29, p. 84) of the work so far done by Dr. Helen Ban-
dulska on the cuticles of leaves from these beds, carry, besides the ferns—
Chrysodium, Asplenites, Glossochlamys, Podoloma, Meniphyllum, Osmunda,
Pteris, Phegopteris, Gleichenia, Lygodium, Hewardia, Aneimia, Adiantum—
and Nzpa, described by Gardner and Ettingshausen, species of the following
genera: Araucarites, Sequoia, Taxodium, Aniba, Neolitsea, Litsea, Lindera,
Cinnamomum, Nothofagus, Rhodomyrtus, and Tristania.

Compared with this list, the flora from Steel’s Crossing, Wash., not yet
completely identified, has the following: Adiantum, Woodwardia, Equisetum,
~ Sequoia, Ulmus or Zelkova, Cercidiphyllum, Ptelea, Dillenites, Acer, and
Platanus.

Occurrence.—In light-colored, grayish to buff shales at Steel’s Crossing,
3 miles west of Renton, Wash. Upper Eocene.

Pteris idahoensis (Knowlton) Brown, n. comb.

Dryopteris idahoensis Knowlton, 22, p. 721, pl. 99, figs. 1, 2.
Pteris calabazensis Dorf. Smith, 33, p. 467, pl. 12, fig. 2.
The venation and habit displayed by these specimens are those of Pieris,
a species tentatively regarded as different from P. szlvicola Hall (Chaney,
13, p. 100, pl. 8, figs. 2, 4-7) from Crooked River, Oreg.
Occurrence.—Marsh, Idaho. Payette formation.

ite page for legend)

-18 (See oppos

1

igs.

F

Aveust 15, 1940 BROWN: SOME AMERICAN FOSSIL FLORAS 347

PINACHAE
Abies chaneyi Mason

Abies chaneyi Mason, 28, p. 149, pl. 4, fig. 2 [not other figures].
Abies magnifica var. shastensis Lemmon. Mason, 28, p. 150, pl. 4, fig. 6.

Although Mason compared the cone scale of Abzes chaneyz with that of
the living venusta, he noted that the bract of this scale lacked the basal
lateral appendages of venusta. The scale itself is twice as wide as it is long,
whereas those of venusta are only a little wider than they are long. Further-
more, if a venusta scale with bract were to occur as a fossil the bract would
scarcely display the wide, expanded base seen in the fossil figured by Mason;
it would appear more like those called Abzes longirostris Knowlton from
Creede, Colo., although here also reservations should be made when com-
paring with venusta. The second scale figured by Mason and compared
with shastensis appears to the writer to belong to the same species as the
first but has had the long point of the bract broken off. The shape of these
scales does not compare favorably with that of shastensis, but the writer
has no other suggestion to offer, except that this fossil species may not have
been in the direct line of either venusta or shastensis but may have ended
without further issue.

Occurrence.—Mascall formation, John Day basin, Oreg.

Abies concoloroides Brown, n. sp.

Abies chaneyt Mason. Brown, 10, p. 167, pl. 45, figs. 238, 24.
Abies chaneyi Mason, 28, p. 149, pl. 4, figs. 1, 7, foliage [not other figures].
Abies magnifica var. shastensis Lemmon. Mason, 28, p. 150, fig. 5.
Mason compared the foliage of Abzes chaneyz with that of the living
venusta but pointed out differences, notably that the needles are slenderer
than those of venusta. Another significant difference is the strong curvature,
which venusta does not display. The needles of the living concolor, however,
match this foliage very well. The seed, here synonymized, can also be dupli-
cated by seeds of concolor. That several species of fir are represented in the
fossils from the Mascall and related formations is quite likely.
Occurrence.—Mascall formation, John Day basin, and Blue Mountains,
Oreg.
Callitris potlatchensis Brown Fig. 6

Callitris potlatchensis Brown, 9, p. 575, pl. 67, fig. 16.

This species was first reported from the Latah formation on Potlatch
Creek, Idaho. An unremitting search for foliage to confirm the identification
of these seeds has so far failed to produce anything satisfactorily comparable
with the foliage of the living species.

Occurrence.—Gray ranch, on Crooked River, Oreg. Specimen collected
by Richard Mote, Corvallis, Oreg.

Fig. 1—Osmunda occidentale (Berry) Brown, n. comb. Fig. 2.—Adiantum ana-
stomosum Brown, n. sp. Fig. 3.—EHucommia montana Brown, n. sp. Fig. 4.—
Eucommia ulmoides Oliver, a living species from central China. Fig. 5.—Frazxinus
flexifolia (Lesquereux) Brown. Fig. 6.—Callitris potlatchensis Brown. Figs. 7, 8.—
Acer knowltont (Berry) Brown, n. comb. Fig. 9.—Tetrapteris simsont Brown, n. sp.
Fig. 10.— Engelhardtia olsonit Brown, n. sp. Fig. 11.—Banksites ineatus Lesquereux.
Fig. 12.—Porana speirit Lesquereux. Fig. 13.—Porana tenuis Lesquereux.—Figs.
14, 15.—Potamogeton heterophylloides Berry. Fig. 16.—Potamogeton parva Brown,
n. sp. Fig. 17.—Dipteronia insignis (Lesquereux) Brown. Fig. 18.—Umbellularia
eocenica Brown, n. sp.

348 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 8

Pseudolarix americana Brown, n. sp.

Piceasp. Brown, 10, p. 167, pl. 45, fig. 8.

At the time this cone scale was tentatively reported as that from a species
of Picea, its resemblance to the scales of the golden larch, Pseudolarixz
kaempfer: Gordon, of eastern China, was noted. Although foliage to confirm
the identification has not yet been collected, or at least not recognized, it
seems to the writer now that this scale should be definitely referred to
Pseudolariaz.

Occurrence.—Republic, Wash. Probably lower Miocene.

NAIADACEAE
Potamogeton heterophylloides Berry Figs. 14, 15

Potamogeton sp. Knowlton, 25, p. 29, pl. 10, figs. 5, 6.
Potamogeton heterophylloides Berry, 4, p. 240 [not pl. 50, figs. 1-8, which are
Keteleeria heterophylloides (Berry) Brown].

Fig. 14 shows a linear, somewhat spatulate, floating leaf and Fig. 15 a
linear, submerged leaf of a species of pondweed. Margins entire. Venation
of the floating leaf 6-8, thin, parallel veins, with delicate cross-veinlets;
that of the submerged leaf is an intramarginal vein with reticulate connec-
tions to the midrib. In. Knowlton’s Fig. 6, which is a two times enlargement
of Fig. 5, the venation is retouched and shown as a series of parallel veins.
‘The venation, however, is like that shown here in Fig. 15, and indicates
that Knowlton’s leaf was a submerged, not a surface leaf. These leaves can
be matched readily by those from a number of living species.

Occurrence.—Latah formation, Spokane, Wash. Both specimens were
collected by Kentworth Buxton, of Spokane.

Potamogeton parva Brown, n. sp. Fig. 16

A small, linear-elliptic, floating leaf of a pondweed. Principal veins
parallel, delicate, and at their basal origin, closely appressed to the midrib.
Cross-veinlets scattering, indistinct.

Occurrence.—Gray Ranch, Crooked River, Oreg. Oligocene. Specimen
collected by Charles B. Read, U. 8. Geological Survey.

FAGACEAE
Castanopsis perplexa (Knowlton) Brown, n. comb.

Castanopsis convexa (Lesquereux) Brooks, 7, p. 288, pl. 6, fig. 5; pl. 10, figs.
1, 3; pl. 12, figs. 1-6; pl. 13, figs. 4-6; pl. 18, figs. 3-5; pl. 20, fig. 4;
pl. 21, fig. 1b. Omit first three items of the synonymy and fig. 7 of the
fourth item.—Brown, 10; p. 171, pl. 49) figs. S—11— Dori aoe 2,
pl 1, figs? 7, 3:.— Smith; 32) p. 102) pl 2. fen 7 plea, noma

Quercus browni Brooks, 7, p. 291, pl. 14, figs. 5, 6 [not figs. 3, 4, 7, 8].—
Smith a2 pal 2 sole 2 tess

Quercus treleasit Berry. Smith, 32, p. 113, pl. 6, fig. 2.

Rhododendron cdahoensis Smith, 33, p. 180, pl. 13, figs. 6, 10.

Axelrod (2, p. 95) identifies as Quercus the types upon which Brooks
based her combination, and considers the Sucker Creek, Oreg., and allied
Castanopsis material as different from Castanopsis chrysophylloides Lesque-
reux, from Chalk Bluffs, Calif. A new name is therefore required for the
Sucker Creek specimens, and those from other localities subsequently identi-

Avaust 15, 1940 BROWN: SOME AMERICAN FOSSIL FLORAS 349

fied as the same species. The new name is based on the types figured by
Knowlton (23, p. 31, pl. 2, figs. 5, 6, 8 [not. fig. 7]) as Salix perplexa, and
cited by Brooks as the fourth item of her synonymy.

Smith questioned the assignment of Berry’s specimens of Quercus treleasi
to Sophora spokanensis Knowlton on the ground that their specific identity
is not evident from the published figures and descriptions. It is true that at
least the petioles in the figures are not clear; but the specimens were cleaned
several years ago before the transfer to Sophora was made, and the petioles
as well as the form and venation were found to be definitely leguminous and
identical with the corresponding characters of the specimens previously
described as Sophora spokanensis.

Occurrence.—Mascall formation, John Day basin; and Sucker Creek,
Oreg.

JUGLANDACEAE

Engelhardtia olsoni Brown, n. sp. Fig. 10

This is a small, 3-lobed bract with a conical nutlet at the base. The lobes
are entire. The venation of each of the lobes consists of a midrib from which
arched veinlets connect with intramarginal veins parallel to the midrib for
a short distance. In the apex of the lobes the secondary veins form a series
of loops. 3

Although this specimen is smaller, it is comparable in other respects to
those called Carpinus grandis Unger (Chaney, 18, pl. 9, fig. 7-9) from
Bridge Creek and Crooked River, Oreg. Although living species of Carpinus
only rarely produce fruit bracts with entire margins, it is possible that
ancient species may have done so. Thus far no leaves of Carpinus have been
reported from the Latah formation.

Occurrence.—Latah formation, 19 miles up Orofino Creek, Idaho. Col-
lected by Boyd H. Olson, Orofino, Idaho.

EUCOMMIACEHAE

Eucommia eocenica (Berry) Brown, n. comb.

Simarubites eocenicus Berry, 5, p. 94, pl. 44, figs. 15, 16.
Carpolithus banisteroides Berry, 5, p. 134, pl. 33, figs. 5, 6.

Although somewhat smaller and with more attenuate bases than the
samaras of Hucommia ulmordes Oliver (Fig. 4), a tree of central China, the
seeds figured by Berry agree very well in all other details. It is not yet clear
to the writer what foliage among the abundant remains in the Wilcox group
belongs with these seeds, but at least one leaf species 1s under suspicion.

The fact that Cercidiphyllum, a genus now restricted to eastern Asia, has
a representative in the Wilcox flora (Brown, 12, pp. 486, 492) makes it less
surprising to find Eucommia there also.

Occurrence.— Wilcox group, near La Grange and Somerville, Tenn.

Eucommia montana Brown, n. sp. Fig. 3

Samara as in that of Eucommia ulmoides Oliver (Fig. 4) of central China,
except somewhat wider in proportion to length, which is about half that of
ulmoides. Tip similarly cleft, margins winged. The pedunculate base is
slightly more attenuate. Venation pattern over the seed a reticulate, forking
meshwork.

This species is almost identical with that described above as Hucomma
eocenica from the Wilcox group of Tennessee; but it differs in being a little

—

000 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES’ VOL. 30, NO. 8

smaller in size and somewhat wider in the apical half. No leaves that could
be definitely assigned to this species were found at this locality.
Occurrence.—One mile west of Grant, Mont., in deposits probably of
upper Oligocene age. |
LAURACEAE

Sassafras bendirei (Knowlton) Brown, n. comb.

Cinnamomum bendirer Knowlton, 23, p. 59, pl. 10, fig. 4.
Philadelphus bendire: (Knowlton) Chaney, 13, p. 118, pl. 17, figs. 1, 2 [not
other figs. |].

These specimens can be matched readily with leaves from the living
Sassafras varufolium. (Salisbury) Kuntze. This fossil species may prove to
be the same as Sassafras hesperia Berry (Brown, 10, p. 174, pl. 53, figs.
7-10), but the writer is unwilling to synonymize them at the present time.

Occurrence.—Bridge Creek and Crooked River basins, Oreg.

Umbellularia eocenica Brown, n. sp. Fig. 18

Leaf oblanceolate to elliptic with acute apex and petiole 1 cm long. The
venation is characterized by the presence of a pair of thin, lateral primaries
arising from the top of the petiole. The secondaries, departing from the
midrib at wide angles, form loops with those above. Margin entire.

This specimen can be matched closely, though not exactly, with leaves
from the California laurel, Umbellularia californica Nuttall, which is found
on mountain slopes and stream bottoms in the Coast Range and Sierra
Nevada region of California, and in southern Oregon.

Occurrence.—In shales of the Green River formation, 3 miles north of
Watson, Utah.

LEGUMINOSAE
Robinia oklahomensis (Berry) Brown, n. comb.

Diospyros pretexana Chaney and Elias, 14, p. 44, pl. 7, figs. 6-8. See syn-
onymy.

Salix coalingensis Dorf. Chaney and Elias, 14, p. 37, pl. 4, figs. 1, 3 [not
Fig. 6, which is Sapindus oklahomensis Berry].

Bumelia oklahomensis Berry, 3, p. 634, pl. 94, fig. 1.

A comparison of the published figures here cited with the specimens
themselves supplies an example of the occasional, unfortunate deceptiveness
of photographs and drawings, and suggests a warning that if retouching is
necessary it be done as faithfully as a satisfactory magnifying glass and a
steady hand can make it. Berry’s figure of Bumelia oklahomensis, for ex-
ample, has far too many secondary veins. Chaney and Elias have shown
Fig. 1 of Salix coalingensis and Fig. 8 of Diospyros pretexana with no sug-
gestion of intermediary secondaries. Their Fig. 7 of Diospyros pretexana
shows the petiole directed toward the lower right, whereas in the specimen
it is directed toward the lower left, a faint hint of which may be had from
the illustration. In all these specimens the petioles are typically leguminous,
that is, broad and cross-wrinkled, indicating their glandular nature. In form,
venation, and petioles these leaflets can be matched by the leaflets of
Robinia, particularly R. pseudacacia Linnaeus, the black locust of the region
east of the Rocky Mountains. The smaller leaflets may also be compared
with those of Amorpha fruticosa Linnaeus.

Occurrence.—Beaver County, Okla.

Avuaust 15, 1940 BROWN: SOME AMERICAN FOSSIL FLORAS Jol

SIMARUBACEAE
Ailanthus indiana (MacGinitie) Brown, n. comb.

Apocynum indiana MacGinitie, 27, p. 66, pl. 12, fig. 1—Smith, 32, p. 117,
lenorenee lvl. 7, fie. 2.

Ailanthus lesquereuxt Cockerell. Arnold, 1, p. 95, pl. 8, figs. 3, 4.

Ailanthus (?) americana Cockerell. Oliver, Carnegie Inst. Wash. Publ.
455(1): 23. 1934.

Cedrela oregoniana (Lesquereux) Brown [part]. Brown, 10, p. 513.

For a long time the writer has been looking for leaf remains that would
confirm the identification of the undoubted seeds of Azlanthus found in the
Green River formation in Wyoming and Colorado; in the lake beds at
Florissant, Colo.; and in the strata on Trout Creek and Sucker Creek,
Oreg. It now appears that the specimen reported by MacGinitie from Trout
Creek, supplemented by those reported recently by Smith from Sucker
Creek, as A pocynum indiana, may be the required foliage. In form, venation,
and length of petiole they compare favorably with the leaflets of the living
Asiatic species, Azlanthus altissima (Miller) Swingle. The published figures
do not show clearly the large basal marginal teeth with glands characteristic
of the living species, but Smith’s Fig. 1 appears to have them.

Seeds called Azlanthus americana Cockerell, but no Azlanthus leaflets,
have been reported from the Green River formation and from Florissant.
It is very likely that when and if the leaflets are found they will be sufh-
ciently different to justify the new specific name for the Trout Creek and
Sucker Creek material.

Occurrence.—Trout Creek, Sucker Creek, and Tipton, Oreg.

ACERACEAE
Acer knowltoni (Berry) Brown, n. comb. Figs. 7, 8

Phanerophlebites knowltoni Berry, 5 p. 48, pl. 7, fig. 6.
Negundo knoultont Berry, 5, p. 98, pl. 21, fig. 7; pl. 42, fig. 7.

Additional material collected by the writer in Tennessee demonstrates
that the specimen called Phanerophlebites knowltoni is a maple samara of
the negundo type, and undoubtedly belongs with the leaves called Negundo
knowltoni, these being the only maple seeds and leaves from the same
localities and the Wilcox group. The samaras have unusually long wings,
and narrow heads which are seldom well enough preserved to show the
attachment scar. Acer negundoides MacGinitie is the well-known relative
in the later Tertiary of the western United States.

Occurrence.—One mile north of Somerville, Tenn. Wilcox group (Eocene).

Dipteronia insignis (Lesquereux) Brown Bitora h7,

Dipteronia insignis (Lesquereux) Brown, 10, p. 181, pl. 59, figs. 10-12. See
synonymy.

The leaflet figured here, although somewhat fragmentary, has a well-
preserved apex and several large rounded, apical teeth, together with the
characteristic venation, sufficient to identify the species. The rounded,
winged fruits have not yet been reported from the same locality, but they
may be looked for with confidence.

Occurrence.—In shales on Bridge Creek, northwest of Mitchell, Oreg.

352 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 8

MALPIGHIACHAE
Tetrapteris simsoni Brown, n. sp. Fig. 9

A 4-winged samara resembling a propeller, one of the wings of which is
shorter and narrower than the other three. Venation of wings subparallel
to flaring near the apex.

This fruit resembles those of a number of living species of Tetrapteris,
particularly 7. schiedeana, found from Mexico to Brazil. The species is
named for A. G. Simson, U.S. Forest Service, who guided me to this locality.

Occurrence.—Four miles west of Mitchell, Oreg., in strata probably in
the lower part of the Clarno formation.

OLEACEAE
Fraxinus flexifolia (Lesquereux) Brown Fig. 5

Fraxinus flexifolia (Lesquereux) Brown, 8, p. 64.

This is the first seed to be reported in confirmation of the previous
identification of ash foliage in the Green River formation. Although the
wing is narrow, it may originally have been wider near the apex.

Occurrence.—Green River formation (middle Eocene), on Piceance Creek,
west of Rio Blanco P. O., Colo.

MENYANTHACEAE
Limnanthemum circularis (Chaney) Brown, n. comb.

Asarum circularis Chaney, 13, p. 115, pl. 18, fig. 11.

The resemblances in form and venation between leaves of Asarum and
Limnanthemum are very striking. Nevertheless, there are distinctive differ-
ences, so that when the fossil leaf ascribed to Asarum by Chaney is sub-
jected to close scrutiny, its assignment to Asarum becomes untenable. In
Asarum there is a midrib flanked by two strong lateral primary veins, from
which basal as well as higher branches arise. The base of the blade, together
with the midrib and primary veins, make a broad, flaring transition into
the petiole. In Limnanthemum the leaves are peltate or nearly so with a
sharp transition of petiole to blade. As should be expected in such leaves,
the primary venation is radiate, although one pair of primaries, depending
upon the length of the leaf, may appear stronger than the rest. This is the
condition in the fossil specimen.

Asarum is a genus of low herbaceous land plants. The possibility of such
remains getting into the fossil record is perhaps not unlikely, but the proba-
bility is remote. Limnanthemum, on the other hand, is a genus of some 20
species of aquatics widely distributed in temperate and tropical regions and
is very likely to get into the fossil record, especially in sediments in quiet
waters, such as those in the Bridge Creek and Crooked River basins of
Oregon must have been.

Occurrence.—Gray Ranch on Crooked River, Oreg.

OF UNCERTAIN AFFINITY
Banksites lineatus Lesquereux Fig. 11

Banksites lineatus Lesquereux, 26, p. 165, pl. 32, fig. 21.
Banksites lineatulus Cockerell, 16, p. 8, pl. 2, fig. 3.

Originally described from the Florissant lake beds and subsequently from
the Green River formation, this species has since been found in strata at

Aveust 15, 1940 BROWN: SOME AMERICAN FOSSIL FLORAS 3090

Gray Ranch on Crooked River, Oreg., and in beds probably in the lower
part of the Clarno formation, 4 miles west of Mitchell, Oreg. The present
occurrences are reported to indicate further relationship between a number
of western Tertiary floras.

Occurrence.—Gray Ranch, Crooked River, Oreg. Collected by Charles B.
Read.

Porana speirii Lesquereux rex a2,

Porana speirit Lesquereux. Brown, 9, p. 583, pl. 69, figs. 1-8. See syn-
onymy and discussion.—10, p. 185, pl. 61, fig. 14.
Viburnum palmatum Chaney and Sanborn, 15, p. 97, pl. 40, fig. 4 [not fig. 3].
These calyces, although readily identifiable, continue to be mysterious in
regard to their botanic affinity. The specimen figured here is the largest the
writer has seen, the figure of that described by Chaney and Sanborn as
Viburnum palmatum being twice natural size. Smaller, but otherwise similar
specimens were found in 1938, 4 miles west of Mitchell, Oreg., in beds pre-
sumably in the lower part of the Clarno formation.
Occurrence.—Gray Ranch, Crooked River, Oreg. Collected by Charles B.
Read.

Porana tenuis Lesquereux Fig. 13

Porana tenuis Lesquereux, 26, p. 173.—Knowlton, 1916, p. 286, pl. 27,
figs. 4-6.

Philadelphus bendirei (Knowlton) Chaney, 13, p. 118, pl. 17, fig. 5.

Cornus ovalis Lesquereux. Chaney, 13, p. 131, pl. 19, fig. 4.

These specimens, like Porana speirii Lesquereux, also remain a mystery
as to their correct botanic affinity. Specimens similar to that figured here
were also found in 1938 in the Green River formation exposed along Piceance
Creek, west of Rio Blanco P. O., Colo.

Occurrence.—One mile west of Grant, Mont., in strata of probable upper
Oligocene age.

ADDITIONAL CHANGES

Abies chaneyt Mason (28, p. 149, pl. 4, figs. 1, 7, seeds) = Cedrela oregoniana
(Lesquereux) Brown.

Acer bolanderit Lesquereux. Smith (31, p. 562, pl. 28, fig. 5; pl. 29, fig. 8;32,
p. 116, pl. 3, fig. 3) = Acer osmonti Knowlton. These are small leaves
of the silver maple type. j

Acer negundoides MacGinitie. Chaney and Elias (14, p. 42, pl. 7, fig. 10) =
Gymnocladus casei Berry.

Acer osmontt Knowlton. Smith (32, p. 116, pl. 4, fig. 1) = Platanus dissecta

Lesquereux.

Azolla berry Brown (8, p. 52, pl. 8, fig. 2) = Azolla berryi Brown. The omission
of the ‘‘i’’ at the time this species was published was deliberate. Al-
though the writer still believes that the principle is sound, it would
seem that in this and in Pinus ball Brown the application was un-
fortunate and confusing because of the connotations of the specific
names.

Betula ? dayana Knowlton (23, p. 41, pl. 4, fig. 4)=Ulmus speciosa New-
berry. This leaf has doubly serrate teeth and is evidently a small leaf
of the speciosa type.

Betula lacustris MacGinitie (27, p. 50, pl. 4, fig. 2)= Alnus carpinoides
Lesquereux.

304 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 8

Cedrela oregoniana (Lesquereux) Brown. Smith (82, p. 115, pl. 3, fig. 7) =
Castanopsis perplexa (Knowlton) Brown.

Fagus sp. Hollick (20, p. 401, pl. 30, fig. 4) = Fagus washoensis LaMotte.

Fagus sanctieugeniensis Hollick (20, p. 399, pl. 30, fig. 3; pl. 31, fig. 3; pl. 32,
figs. 2, 3) =Fagus washoensis LaMotte. Although sanctieugeniensis has
priority, the writer urges that an exception to the rules be made here
on the ground that the term is lengthy and cumbersome—a coinage
contrary to the recommendations in the codes.

Glyptostrobus europaeus Lesquereux. Smith (30, p. aS pl. 1, fig. 14) =
Glyptostrobus oregonensis Brown.

Oreopanax dissecta (Lesquereux) Smith (GWA; 105 ILIG, jole Ike pilin Ay eS, IO)
Oreopanax precoccinea (Brooks) Arnold. If Smith’s specimens are differ-
ent from those called O. precoccinea, they certainly are not identical
with the Florissant species called Aralia dissecta.

Phylloteuthis subovata Meek and Hayden (1876, U. 8S. Geol. and Geog.
Survey Terr. Rept. 9: 505, pl. 33, fig. 3) = Nzlssonza gibbsiz (Newberry)
Hollick. The illustration of Phylloteuthis subovata is in part recon-
structed to show an alleged resemblance of the specimen to the ‘‘pen’”’
of a squid. The fact that this specimen, from Fox Hills strata (Upper
Cretaceous) on the Moreau River, 8. Dak., is embedded in a block
carrying marine mollusks, was further cause for believing it to be the
remains of a marine animal. The form and venation, however, identify
it as part of a cycad leaf. (See Brown, 11, p. 245, pl. 49, figs. 5, 6;
pl. 52, fig. 6b.) This specimen was called to my attention by John B.
Reeside, Jr.

Pinus ball Brown (8, p. 53, pl. 9, figs. 6-10) = Pznus balla Brown. See com-
ment on Azolla berry Brown.

Platanus regularis Knowlton. Smith (32, p. 115, pl. 4, fig. 2) = Platanus
dissecta Lesquereux. Given a large enough suite of Sucker Creek ma-
terial, this specimen can be duplicated readily as a variant of P. dissecta.
What does Mrs. Smith mean by the paradox: ‘‘The specimens are
easily separated from P. dissecta, with which they have much in
common’’?

Populus balsamoides Goeppert. Elias (18, pl. 367, fig. 6)=Celtis kansana
Chaney and Elias.

Ptelea enervosa Smith (31, p. 560, pl. 29, fig. 6)=Ptelea miocenica Berry.
The lack of venation in this specimen can be explained by the fact that
the Sucker Creek matrix is notoriously poor for preserving fine details.
The small size of this fruit does not eliminate it from falling within the
limits of variation of P. miocenica, if the latter be compared with living
species of Ptelea.

Ptiloteuthis foliatus Gabb (1869, Paleont. California 2: 128, pl. 19, fig. 4) =
Sagenopteris elliptica Fontaine. Originally described from the Shasta
group (Cretaceous) of California, by Gabb as the ‘‘pen”’ of a squid,
and recently redescribed (Rehn, John W. H., 1939, Acad. Nat. Sci.
Philadelphia Notulae Naturae no. 9, pp. 1, 2) as a cockroach wing, this
specimen represents a species of fern common in the flora of the Shasta
group. This specimen was called to my attention by F. M. Carpenter.

Quercus malheurensis Smith (31, p. 560, pl. 29, fig. 1) = Acer bendzrez (Lesque-
reux) Brown. The venation of this leaf, together with the form of its
lobes, would, if the leaf were completely reconstructed, show clearly
that it is a leaf of the macrophyllum type.

Aveust 15, 1940 BROWN: SOME AMERICAN FOSSIL FLORAS 300

Rhus oregonensis Smith (31, p. 561, pl. 29, fig. 9) = Acer glabroides Brown.
This is a deeply lobed leaf. Instead of having rounded teeth as in Rhus
trilobata Nuttall, with which it was compared, the teeth are sharp and
in the lower left lobe are represented as being double, which they
never are in R. trilobata.

Salix coalingensis Dorf. Chaney and Elias (14, p. 37, pl. 4, fig. 6)=
Sapindus oklahomensis Berry.

Salix perplexa Knowlton (23, p. 31, pl. 2, fig. 7) = Pterocarya mixta (Know]-
ton) Brown. Fig. 7 depicts an entire-margined specimen, which, in fact,
has serrate teeth.

Sciadopitys americana Smith (30, p. 225, pl. 1, fig. 1) = Pznus sp. This is a
needle of a species of pine, of which three are described from the same
formation and region.

LITERATURE CITED

1. ARNoLD, C. A. Observations on the fossil flora of eastern and southeastern Oregon.
Part 1. Univ. Michigan Mus. Pal. Contr. 5(8): 79-102. 1937.
2. AxELrRop, D. I. A Muocene flora from the western border of the Mohave Desert.
Carnegie Inst. Washington Publ. 516. 1939.
3. Berry, E. W. Fossil plants from the late Tertiary of Oklahoma. Proc. U.S. Nat.
Mus. 54: 627-636. 1918.
4. —— A revision of the flora of the Latah formation. U.S. Geol. Surv. Prof. Paper
154: 225-265. 1929.
5. Revision of the lower Eocene Wilcox flora of the southeastern States. U.S. Geol.
Surv. Prof. Paper. 156. 1930.
6. Miocene plants from Idaho. U. 8S. Geol. Surv. Prof. Paper 185: 97-125.
1934.
7. Brooks, Betty W. Fossil plants from Sucker Creek, Idaho. Annals Carnegie
Mus. 24: 275-336. 1935.
8. Brown, R. W. The recognizable species of the Green River flora. U.S. Geol. Surv.
Prof. Paper 185: 45-77. 1934.
9: Miocene leaves, fruits, and seeds from Idaho, Oregon, and Washington. Journ.
Pal. 9: 572-587. 1935.
10. Additions to some fossil floras of the western United States. U.S. Geol. Surv.
Prof. Paper 186: 163-206. 1937.
ae Fossil plants from the Colgate member of the Fox Hills sandstone and adjacent
strata. U.S. Geol. Surv. Prof. Paper 189: 239-275. 1939.
Ps ae leaves, fruits, and seeds of Cercidiphyllum. Journ. Pal. 13: 485-499.
1939.
13. CHANEY, R. W. Geology and paleontology of the Crooked River basin, with speczal
enone to the Bridge Creek flora. Carnegie Inst. Washington Publ. 346: pt. 4.
192
14. CHaney, R. W., and Exias, M. J. Late Tertiary floras from the High Plains. Car-
negie Inst. Washington Publ. 346: te ole OSG=
15. CHANEY, R. W., and SANBORN, ETHEL I. The Goshen flora of west central Oregon.
Carnegie Inst. Washington Publ. 439. 1933.
16. CocKERELL, T. D. A. Plant and insect fossils from the Green River Eocene of
Colorado. Proc. U. S. Nat. Mus. 66(19): 1-138. 1925.
17. Dorr, Eruine. A late Tertiary flora from southwestern Idaho. Carnegie Inst.
Washington Publ. 476(2). 1938.
18. Enras, M. K. Grasses and other plants from the Tertiary rocks of Kansas and
Colorado. Univ. Kansas Bull. 33: 333-367. 1932.
19. GARDNER, J. S., and ETTINGSHAUSEN, C. <A monograph of the British Eocene flora,
1. Filices. . 1879-1882.
20. Houiick, Artuur. The flora of the Saint Eugene Silts, Kootenay Valley, British
Columbia. New York Bot. Gard. Mem. 7: 389-464. 1927.
ile The Tertiary floras of Alaska. U.S. Geol. Surv. Prof. Paper 182. 1936.
22. KNowuton, F. H. The fossil plants of the Payette formation. U.S. Geol. Surv.
Ann. Rept. 18 (3): 721-744. 1898.
ao. Fossil flora of the John Day basin, Oregon. U.S. Geol. Surv. Bull. 204.
1902.
24. —— A review of the fossil plants in the United States National Museum from the

306 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 8

Florissant lake beds at Florissant, Colorado. Proc. U.S. Nat. Mus. 51: 241-297.
1916.

25. —— Flora of the Latah formation of Spokane, Washington, and Coeur d’ Alene,
Idaho. U.S. Geol. Surv. Prof. Paper 140: 17-81. 1926.

26. LesquEerREuX, Lro. The Cretaceous and Tertiary floras. U.S. Geol. Surv. Terr.
Rept. 8. 1883.

27. MacGinitiz, H. D. The Trout Creek flora of southeastern Oregon. Carnegie Inst.
Washington Publ. 416 (2). 1983.

28. Mason, H. L. Fossil records of some west American conifers. - Carnegie Inst.
Washington Publ. 346 (5). 1927.

29. Reip, ELeanor M., and CHAaNnpLER, MArRsoriz, E. J. The London Clay flora.
British Mus. Nat. Hist., pp. 1-561. 1933. |

30. SmirH, HELEN V. Notes on fossil plants from Hog Creek in southwestern Idaho.
Trans. Michigan Acad. Sci., Arts and Letters 23: 223-230. 1988.

31. Some new and interesting late Tertiary plants from Sucker Creek, Idaho-Oregon
boundary. Bull. Torrey Bot. Club 65: 557-564. 1938.
By Additions to the fossil flora of Sucker Creek, Oregon. Trans. Michigan Acad.

Sci., Arts and Letters 24: 107-120. 1939.
33. —— A flora of eastern American aspect in the Miocene of Idaho. Bull. Torrey Bot.
Club 66: 465-481. 1939.

ANTHROPOLOGY .—The finding of an Indian ossuary on the York
Raver in Virginia. T. D. Stewart, U.S. National Museum.

As recently as 1935, when Davidson wrote his paper on the burial
customs in the Delmarva Peninsula, there were no published descrip-
tions of Indian burials from the Algonkian territory south of Chesa-
peake Bay. Since that time, however, detailed records have been made
for at least 17 ossuaries—the prevailing mode of burial—on the tide-
water section of the Potomac River (Graham, 1935; Ferguson, 1937a
and 1937b; Stewart and Wedel, 1937; Stewart, 1939 and 1940a). The
present report adds to this record the only carefully investigated « case
from the tidewater region south of the Potomac.

HISTORY OF THE REGION

The earliest description of the York River is that of Capt. John
Smith in 1608:

This Riuer of Pamaunke [York] is not past twelue mile from that we dwell .
on [James], his course northwest and westerly as the other. Weraocomoco
is vpon salt water in bredth two myles, and so [the river] keepeth his course
without any tarrying some twenty miles; where at the parting of the fresh
water and the salt, it diuideth it selfe into two partes, the one part...
[Pamunkey], as broad as Thames, and nauigable with a Boate threescore or
fourescore miles. . .

The other branch [Mattaponi] a little lesse in breadth, yet extendeth
not neare so farre.... (Arber, 1884, pp. 20-21)

Repeating essentially the same description in 1612, Smith adds:

Where this river is divided, the Country is called Pamavnke [West Point],
and nourisheth neere 300 able men. About 25 miles lower on the North
side of this river is Werawocomoco, where their great King inhabited when

1 Published by permission of the Secretary of the Smithsonian Institution. Re-
ceived March 1, 1940.

Aveust 15, 1940 STEWART: INDIAN OSSUARY ON YORK RIVER 300

Captain Smith was deliuered him prisoner; yet there are not past 40 able
men. (Arber, 1884, p. 51.)

In addition, Smith’s map of 1612 shows three small Indian villages
on the north side of the river between the fork and Werawocomoco:
Pasaughtacock, Poruptanck, and Mattacock. The name of the second

/ MILE

Fig. 1.—Map showing the location of the ossuary in relation to the York River
and the highway system passing through the town of West Point. (Based on the
Geological Survey maps of Virginia: New Kent, Urbana, Toano, and Williamsburg
quadrangles, 1906-1919.)
village in the course of time may have been extended to the creek that
forms part of the boundary between King and Queen and Gloucester
Counties and now known as Poropotank Creek.

Beginning about 1642 (Nugent, 1934, p. 131 et seq), large tracts of
land on the north side of the York River (then known as the Charles)
were patented. The boundaries of these patents are usually indefinite
unless physiographical landmarks are mentioned. In this connection

358 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 8

it is noteworthy that “Poropotanck”’ Creek figures prominently in the
land records.

These are the essential historical data pertaining to the site of the

Fig. 2.— Appearance of the north shore of the York River in the vicinity of the
ossuary. Looking north. Human bones from the ossuary may be seen on the beach and
protruding from the bank.
ossuary, which, it will be seen from the accompanying map (Fig. 1) is
located on the north bank of the river between 4 and 5 miles below the

fork, or where the present town of West Point is situated.

Auecust 15, 1940 STEWART: INDIAN OSSUARY ON YORK RIVER 309

DESCRIPTION OF THE SITE

The circumstances of the find are as follows: Several years ago fol-
lowing a spring flood, B. R. Anderson, the present owner of Belleview
farm, saw for the first time some bones exposed along the river bank
about 850 feet upriver from his house. He reburied them. From time
to time since then, however, bones have been seen along the beach,
and finally in the fall of 19389 L. R. Treat, of West Point, obtained
some and brought them to the attention of Dr. Malcolm H. Harris,
also of West Point. Being familiar with the history of the colonial pe-
riod, Dr. Harris at once sensed the significance of this find and im-
parted the information to the Smithsonian Institution, suggesting at
the same time a scientific examination of the site. As a result, I was
detailed from October 26 to 28, 1939, inclusive, to make this investi-
gation. 7

At the time of my visit, bones could be seen protruding from the
river bank (Fig. 2) for a distance of 10 feet. Also, loose bones were
seen along the beach, both nearby and at a considerable distance.
When the sand was dug away from the base of the bank it appeared
that the waves had undercut the bone layer and caused blocks of it to
drop down, as is typical of this form of erosion. The soil in this area
apparently contains much clay, for the vertical surface of the bank
had been baked by the sun so that it resisted the point of a pick de-
livered with considerable force. Under these circumstances, and since
the protruding bones were badly damaged, there was nothing to do
but remove the outer layer and thus reach the soft soil. When this
was done it became possible to collect whole bones and to observe
their relationship.

It was determined that the bone layer (8 inches to 1 foot thick) ex-
tended into the bank only about 4 feet in the middle and less at either
end. It seems likely, therefore, that the original shape of the pit was
ovoid—perhaps 8 by 10 feet. If this is true, then about half of the os-
suary had been washed away before it could be examined.

It was observed, also, that the remains found around the periphery
of the pit—that is, farthest within the bank—were at a higher level
(3 feet from the top); in other words, the sides of the pit were not ver-
tical, but sloping, as is the usual finding. Moreover, the limits of the
pit were easily distinguishable, because the fill was softer and of a
darker color than the undisturbed soil. The fill surrounding the hu-
man bones yielded many large oyster shells, six potsherds, parts of a
dog’s skull, a turkey humerus, and the calcaneus of a deer.

The human bones within the ossuary were distributed in definite

360 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 8

bundles and frequently with related parts in positions indicating that
they were still articulated when interred. At the point on the pe-
riphery of the pit occurring deepest within the bank a quantity of
charred bone fragments was found scattered about, but seemingly as-
sociated with a well-preserved large male skull. It was obvious, how-
ever, that these bones had been burned prior to burial.

As the ossuary was explored a selection of bones was made for
further study, consisting of whole skulls and long bones and such
other parts as past study (Steward and Wedel, 1937) has shown to be
of most value in this type of investigation. The examination of this
material is reported below.

CULTURAL MATERIAL

Potsherds constitute the only cultural material from the ossuary
that requires further description. The six sherds recovered (U.S.N.M.
no. 380715) suggest two types of ware: (1) A thick (7-9 mm) coarsely
shell-tempered variety with irregular fracture lines; and (2) a thin
(8-5 mm) lightly shell-tempered variety with straight fracture lines
suggesting a coiled structure. Fairly large vessels are suggested by the
curvature of the bigger sherds especially those of the coarser variety.
Color ranges from black through variable gray to orange-buff. Leach-
ing of the shell has left the surfaces pitted, particularly in the coarser
variety. Otherwise the surfaces show mostly the striations due to a
smoothing implement with an occasional faint suggestion of net im-
pression or a cord-wrapped paddle.

Only one rim sherd was recovered; it belongs to the thin variety of
ware and has a crudely everted lip. The surface of this sherd has
flaked off somewhat but still shows a diagonal striation that may be
an attempt at decoration.

During the time spent at this site I took the opportunity to examine
the neighboring field and river bank. Local residents asserted that un-
til recent years the shore at this point had been a part of the swamp
that now ends a short distance upriver (see figure). The field here is
quite flat, and although it had not been plowed recently, except back
about 100 feet from the river, it was possible to see some of the sur-
face. Pottery was not in evidence; indeed, I found only one artifact, a
small triangular quartz point.

Downriver from the old colonial farmhouse, the fields, unlike those
above, are full of oyster shells. Along this section of the river bank it
was possible frequently to find small‘potsherds just below the sod. A
small collection from this place (U.S.N.M. no. 380716) differs from

Avucust 15, 1940 STEWART: INDIAN OSSUARY ON YORK RIVER 361

the ossuary sample chiefly in including thick grit-tempered sherds
with exterior surfaces probably net marked. Because of the smallness
of the samples, it is impossible to say whether these differences are
significant. In general, however, these wares are typical of the area.

SKELETAL MATERIAL

Number.—Since part of this ossuary had been washed away, we
can never know the accurate number of individuals buried here. Even
an approximation, however, is of value for comparative purposes. For
this reason the skeletal parts recovered (U.S.N.M. nos. 379062-379-
088) are listed in Table 1. According to this listing more skulls are
represented than other skeletal parts. In round numbers the maxi-
mum is 25. Assuming that half of the ossuary was washed away, the
number originally buried here was approximately 50. As compared
with the Potomac ossuaries of which only 5 out of 17 contained 50 or
fewer bodies, this one from the York River thus ranks with the mi-
nority as small. ;

Sex.—Exact determination of sex is difficult in fragmentary mate-
rial and has not been attempted here. Both sexes, however, are repre-
sented and not disproportionately.

Age.—Most of the skeletal remains are those of adults, only 7 sub-
adults being noted. As judged by the teeth of the lower jaw, these 7
subadults fall into the following age periods: 2-6 years, 2; 6-12 years,
3; over 12 years, 2.

Physical type —Of the recovered skulls, 8 (adults: 4 males, 4 fe-
males) were sufficiently complete to permit measurement. These have
cranial indices ranging from low dolichocrany to low brachycrany
70.7, 72.0, 72.7, 74.0, 74.1, 75.3, 78.3, 81.8 (asymmetrical). The mean
height indices of these skulls, with two exceptions (81.9 each), indi-
cate high vaults (83.9-88.5). These figures are typical of the general
Algonkian range (Hrdli¢ka, 1927). Further skull measurements will
not be given here because a detailed study of the accumulating skele-
tal remains from this area is planned for the future.

TABLE 1.—NuUMBER OF BONES RECOVERED FROM THE OSSUARY

Bone Right Left
BNCTINPOnAE hee ab oy Saks Bea date a's 23 19
HO OWE Tea We si A oes cles sl ade 20
Ecmerus (distalend)....05...... IL 7 62,
emai @oroximeal end). ...45. 5. 16 18

1 With symphysis; possibly 6 others represented.

062 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 8

The remaining skeletal parts present among other things two anom-
alies, the frequency of which, even in such a small sample, suggests
relationship with other Algonkian groups: (1) The external auditory
meatus is generally free from exostoses, only traces being noted in
7.1 per cent of the ears (42). This compares with 10 per cent for the
two Anacostia ossuaries (Stewart and Wedel, 1937). (2) Of 26 humeri
about 54 per cent show septal apertures. Although this is somewhat
higher than the 43.2 per cent recorded for the Anacostia ossuaries, it
is nevertheless fairly typical of the Northeastern Indians (Hrdli¢ka,
1932).

Stature in such a small sample is perhaps best gauged by the maxi-
mum encountered. Thus, the longest humerus is 36.3 em and the
longest femur 51.5 em. According to Manouvrier’s tables (Hrdlicka,
1939) these lengths correspond to a statute of nearly 6 feet.

Pathology.—Pathological bones include 5 right and 8 left tibia with
lesions such as are generally attributed to syphilis. One frontal bone
shows scars perhaps caused by the same disease.

As usual among the Indian remains from this region the teeth are
frequently decayed. Caries appear to be chiefly of the developmental
type; that is, beginning in the fissures on the occlusal surface.

Burnt bones.—This lot of fragments—all that were found in the os-
suary—includes parts of most of the major bones of the body and ap-
parently of one individual. The lower jaw indicates that this individu-
al was an adult. Unfortunately, however, the sex characters are not
positive; it was either a small male or a female.

DISCUSSION

The conclusion reached in this study is that the York River ossuary
has the same general features as those found along the Potomac River.
In view of the proximity of the two areas, and their history, this is to
be expected. Among the features possessed in common by the major-
ity of these ossuaries are the following:

. Association with a recognizable habitation site.

. Rounded pit, 3-4 feet deep, with sloping sides.

. Bones arranged in bundles and indicating at least a partial articula-
tion.

. Poverty of accompanying cultural remains.

Presence in the pit of burnt human bones.

. Ages and ratio of sexes reflecting normal death rate.

. Physical type of the Northeastern Indians.

. Presence of dental caries and (probably) syphilis.

The finding of the burnt bones in the York River ossuary is of in-
terest in view of the following statement by Stewart and Wedel (1937,

Avuaust 15, 1940 STEWART: INDIAN OSSUARY ON YORK RIVER 063

pp. 218-219): “The finding of calcined bone in the second pit at Ana-
costia is something new for this area and cannot be clearly explained.’’

Of the ossuaries investigated since 1937 by Mrs. Ferguson, the late
Judge Graham, and me, almost all have contained burnt bones. Mrs.
Ferguson” informs me that each of her 5 major ossuaries contained
deposits of burnt bones, varying in number up to 7 and estimated to
have included at least 38 individuals. In no ease did it appear that the
firing had occurred in the pit, for adjacent bones were normal in ap-
pearance. Moreover, whole skulls were sometimes found to contain
fragments of charred bone.

The records left by the late Judge Graham are not so complete but
indicate essentially the same thing. Of the 4 ossuaries that he exca-
vated, 3 contained deposits of burnt bone, ranging up to 4 in number
and possibly including 10 individuals. In only one instance do I find
an opinion expressed in his notes regarding the place of burning. This
is to the effect that they could not have been burned 7n sztu. In two
cases Judge Graham noted that the burnt bones belonged to infants.

Following up Judge Graham’s work at Potomac, I succeeded in lo-
cating and partly excavating a fifth ossuary (Stewart, 1940a). Among
the first human bones encountered on the periphery of this pit was a
mass of burnt fragments representing remains of more than one in-
dividual.’? The position of this deposit—on the sloping side of the pit—
and the close proximity of normal bones, make it seem improbable
that the firing occurred in the pit.

In addition, both Mrs. Ferguson and Judge Graham, but especially
the former, have found individual secondary burials in which the
bones were burned.

The uniformity of these findings suggests that the burning of one or
more individuals was commonly a part of the burial ceremony in this
area.* Whether this ceremony was simply a cremation or a sacrifice
naturally can not be ascertained from the bones. However, Mrs.
Ferguson has called my attention to the following statement in Henry
Spelman’s Relation of Virginea (Arber, 1884, pp. ev—cvi):

In y® Patomecks cuntry they haue an other god whom they call Quio-
quascacke, and unto ther Images they offer Beades and Copper if at any
time they want Rayne or haue to much, and though they obserue no day to
worshipe ther god: but uppon necessitye, yet oncfe] in the yeare, ther

preests whch are ther coniurers with y* [people] men, weomen, and children
doe goe into the woods, wher ther preests makes a great cirkell of fier in y°

> Personal communication. February 26, 1940.

3 Material not yet studied.

* In 1936 I examined the remains of an ossuary exposed in a gravel pit near Cam-
bridge, Md. (Dorchester County). Here, too, a charred adult bone was noted.

364 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 8

which after many obseruanses in ther coniurations they make offer of 2
or 3 children to be giuen to ther god if he will apeare unto them and shew
his mind whome he [will haue] desier. Vppon which offringe they heare a
noyse out of y® Cirkel Nominatinge such as he will haue, whome presently
they take bindinge them hand and footte and cast them into y° circle of the
fier, for be it the Kinges sonne he must be giuen if oncle] named by ther god,
After y® bodies which are offered are consumed in the fier and ther ceri-
monees performed the men depart merily, the weomen weaping.

It is possible, therefore, even though burned children’s bones are
uncommon in the ossuaries, that all such remains represent some re-
ligious rite other than a burial ceremony.

Finally, a word may be said about the age of these ossuaries. Of the
17 ossuaries to which reference has been made, 6 contained European
articles—glass, copper, iron, or silver—in considerable abundance;
the remainder had only native artifacts—shell beads, clay pipes, etc.
This finding represents, I believe, a clear distinction in time: before
and after trade goods became plentiful, or roughly before and after
1608.° Since this dividing date is more than 100 years after the dis-
covery of America, it seems unlikely that few or any of the ossuaries
lacking European objects are pre-Columbian in age. This view is sup-
ported by the presence of syphilitic bones in all of the ossuaries thus
far examined. I have summarized elsewhere (Stewart, 1940b) the evi-
dence for believing that this disease appeared in North America in
relatively recent, possibly even post-Columbian, times.

LITERATURE CITED

ARBER, Epwarp. Capt. John Smith’s works, 1608-1631. The English Scholar’s
Library, No. 16. Birmingham, 1884.

Davipson, D. 8. Burial customs in the Delmarva Peninsula and the question of their
chronology. Amer. Antiq. 1(2): 84-97. 1935.

Fereuson, Avice L. L. Burial area in Moyaone. Journ. Washington Acad. Sci.

27(6): 261-267. 19387a.

Moyaone and the Piscataway Indians. (Privately printed.) Washington, 1937b.

GRAHAM, Wm. J. The Indians of Port Tobacco river, Maryland, and their burial places.
(Privately printed.) Washington, 1935.

—— Notes on excavations at Potomac. Copy filed under U. 8. National Museum ac-
cession no. 147346.

HrpuritéKa, A. Catalogue of human crania in the United States National Museum col-
lections. The Algonkins and related Iroquois, etc. Proc. U. 8. Nat. Mus. 69(5).
1927.

—— The humerus; septal apertures. Anthrop., Prague, 10: 31-96. 1932.

Practical anthropometry. Philadelphia, 1939.

Nugent, Nett Marion. Cavaliers and pioneers. Abstracts of Virginia land patents
and grants 1623-1800. I. Richmond, 1984.

Stewart, T. D. Excavating the Indian village of Patowomeke (Potomac). Expl.
Field-Work Smithsonian Inst. 1938, pp. 87-90. Washington, 1939.

—— Further excavations at the Indian village site of Patawomeke (Potomac). Expl.
Field-work Smithsonian Inst. 1939, pp. 79-82. Washington, 1940a.

—— Some historical implications of physical anthropology in North America. In Smith-
sonian Misc. Coll. 100. 1940b.

Stewart, T. D., and WrepEL, W. R. The finding of two ossuaries on the site of the
Indian village of Nacotchtanke (Anacostia). Journ. Washington Acad. Sci. 27(5):
213-219. 1937.

> Probably 10-20 years later in some places.

CONTENTS

PALEONTOLOGY.—Seven new ice of Carboniferous Crinoidea i
Inadunata. ie win RK. so fae Te See ees

Botany.—The wild bean Phaseolus polystachyus (L.) B.S. Pp: tte
chromosome number. H. A. Atuarp and H. F. Anuarp. sf

Mycotocy.—A ringworm disease of muskrats transferable to man. ee
VerA K. CHARLES..... Ai Ce ian ee ee ee ree ahs OG ea

PaLEOBOTANY.—New species and changes of name in some American —
fossil floras. RoLAND W. BRowN......... ene te ho. :

ANTHROPOLOGY.—The finding of an Indian ossuary on the York River
in Virginia. T. D: STEWART. .0..0 622 7.) 4 ae

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JOURNAL

OF THE
WASHINGTON ACADEMY OF SCIENCES
Vou. 30 SEPTEMBER 15, 1940 No. 9

PALEON TOLOGY.—Fossil pearls from the Colorado group of western
Kansas... Routanp W. Brown, U. 8. Geological Survey.

Although pearls taken from living mollusks are familiar to everyone
as lustrous ornaments, often of great beauty, fossil pearls are seldom
seen. As arule they are not spectacular, for, with few exceptions, they
have lost their original luster; and, unless still embedded in or asso-
ciated with an identifiable shell, they have also lost some of their sci-
entific significance. Whether they are as rare as the 45 or 50 records
would indicate is not definitely certain, because it is altogether likely
that by many collectors they may hitherto have been unsought or
may have been mistaken for other objects that simulate them, such as
ordinary rounded pebbles, concretions formed by inorganic processes,
cave ‘‘pearls,”’ oolites, pisolites, otoliths, fish teeth like those of Sphae-
rodus, algal nodules, and even seeds or fruits of plants.

The literature about ornamental pearls runs the whole gamut of
fact and fancy, as one may learn from the extensive bibliography in
The book of the pearl by Kunz and Stevenson (18). Of fossil pearls,
however, the literature embraces only about 30 titles. These are cited
in part in the recent papers by Russell (24), Berry (2), Zilch (33), and
Frenguelli (9). All these students have also published tables showing
the distribution of fossil pearls known to the date of publication. Zilch
(33), in particular, has reviewed every occurrence known to him and
has made two changes that should be noted by those who compare
these tables, namely, Perna oblonga Seeley is P. seeleyi Zilch, and Per-
na sandbergeri, cited erroneously at first by Zilch (32),is P. oblonga
Rohmer and Biichner (33). In the latest table, that by Frenguelli,
Perna oblonga should read P. seeleyi; Perna sandbergeri should read P.
oblonga and the occurrence 5 should be changed from Pliocene to Oli-
gocene; Nautilus sp. should read Pleuronautilus pseudoplanilaterus;
also the occurrence cited as 30 in the Eocene should be changed to Oli-
gocene. All the tables cited have omitted J. Marwick’s record of pearls
from Melina zealandica in the Pliocene of New Zealand (20). Further

1 Published by permission of the Director, Geological Survey, U. S. Department
of the Interior. Received April 29, 1940.

365

Sep 24 1940

366 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

Figs. 1-22.—(See opposite page for explanation.)

Sup. 15, 1940 BROWN: FOSSIL PEARLS 367

records not appearing in previous tables are those by Dorn (7) and
Kutassy (19). Here too may be added a note that Dr. 8. F. Blake, of
the U. 8. Bureau of Plant Industry, has found subsequent to Berry’s
record (2) a large blister pearl also in a Panopea americana shell from
the Miocene at Jones Wharf, Md. Blake’s pearl is slightly larger than
Berry’s but has the same outline and occurs in exactly the same posi-
tion near the hinge line. The exterior of the shell does not now show
evidence of having been perforated, but irregularities in the amount
and contour of shell substance indicate that some pathologic condition
prevailed. The instances to be recorded here are worthy of being
added to the list of occurrences of fossil pearls, because the pearls are
not only firm, unusually large, well formed, and characteristic, but the
records are from strata that, although carrying marine faunas chiefly
of Foraminifera and Mollusca, have not yet been cited as yielding
pearls. |

Five of the new specimens (Figs. 1, 3, 4, 5, 12) were collected by
George F. Sternberg, of the Fort Hays Kansas State College, at Hays,
Kans., from scattered outcrops of the Niobrara formation (Upper
Cretaceous) along the Smoky Hill River valley west and southwest
of Hays and extending into Gove and Logan Counties toward the
head of that valley. Several of these specimens were taken from re-
mains of Inoceramus, but a few were found free. Two specimens (Figs.
7, 10) were collected by A. H. Schutte, of Ellis, Kans., from the
Benton shale, which underlies the Niobrara limestone, along a tribu-
tary of Smoky Hill River, 16 miles south of Hays. These 7 specimens,
together with 36 more collected by a friend of Mr. Sternberg from
the Benton shale 18 miles east of Hays, were sent to the writer by
Mr. Sternberg for confirmation of the tentative identification as
pearls.

All the Niobrara pearls are now lusterless and are dull, yellowish
gray to dark gray in color. Four are of discoid or oblate spheroid
form, the largest (Figs. 5, 6) having a long diameter of 2 cm and a

Figs. 1-6, 12-16, 19, 20.—Fossil pearls from the Niobrara limestone. Figs. 1, 2.—
Different views showing same pearl partially embedded in the prismatic layer of an
Inoceramus shell. Fig. 6.—Side view of 5. Fig. 13.—Side view of 12, showing equa-
torial line where cut was made for section 14. Fig. 15.—Section at right angles to the
long diameter showing irregularities in deposition of the laminae. The dark spots are
limonite. X4. Fig. 16.—Same as 14, x4. Fig. 19.—A sector of 14, showing radial
prismatic calcite, X40. Fig. 20.—Polygonal surface pattern of 5. X16. Figs. 7-11,
21, 22.— Fossil pearls from the Benton shale. Figs. 7-9.—Top, side, and bottom views
of a hemispheric pearl. Figs. 10, 11.—Top and bottom views of a baroque. Figs. 17,
18.—Bottom and side views of a modern hemispheric pearl from the common clam,
Venus mercenaria, for comparison with 8 and 9. Fig. 21.—A hemispheric pearl resting
on the prismatic layer of an Inoceramus shell. Fig. 22.—Group of spherical pearls.
All figures natural size except as indicated.

368 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 9 ~

short diameter of 1.3 em. The smaller specimens (Figs. 3, 4) are some-
what irregular and appear to have suffered slight fracturing and dis-
tortion by mechanical causes probably subsequent to entombment in
the Niobrara sediments. Portions of the smooth outer laminae of
some of the pearls have been spalled off. This outer surface, where
well preserved, is seen under a 14-power hand lens to be a pattern of
minute, closely packed polygons (Fig. 20), each convexly domed out-
ward. These polygons represent the slightly eroded ends of radial
prisms. The smooth surface of the prismatic layer exposed on the
interior of an Jnoceramus shell displays the same pattern.

Figs. 1 and 2 illustrate an almost spherical specimen, similar in
color and texture to the others but partially embedded in a piece of
the prismatic portion of the host Inoceramus shell. Nothing appar-
ently now intervenes between the pearl and the shell, and none of the
nacre of the shell remains, but whether this means that the growth of
the prismatic layer was gradually enveloping the pearl is a matter of
conjecture, though parallels to this condition are not rare in modern
mollusks. The writer has been unwilling to risk destruction of the
pearl by attempting to detach it from the shell merely to satisfy
curiosity on this point.

One specimen (Figs. 12-16) was cut in half through the equator by
using a thin copper wire and carborundum powder. The sections re-
veal numerous light and dark, concentric, closely-spaced rings or
laminae around a small, indefinite, rusty-stained nucleus. Darker
parts of some rings are transitional cireumferentially into lighter por-
tions of the same rings. Under the microscope the rings show some
slight variations in smoothness of curvature and in width both in
themselves and as compared with one another (Fig. 19). They are
composed for the most part of large, radially disposed crystals of
calcite. No conchiolin appears to be present now. The section does not
display a black cross under crossed nicols, and in this respect is similar
to the prismatic portion of Russell’s second pearl (24, p. 422). Narrow
radial zones of distortion along which secondary calcite has been
sporadically deposited may be seen in the right half of Fig. 16. Fig. 15
is a section cut through the short diameter at right angles to the long
diameter. It differs from the previous section chiefly in showing
greater irregularity in deposition of the concentric laminae. The dark
spots are flecks of secondary limonite, probably after pyrite. These
sections compare well with those reproduced by Russell (24, figs. 4-7),
Newton (22, pl. 5, figs. 3-6), Kunz and Stevenson (18, plate facing
p. 53), and others.

SeprT. 15, 1940 BROWN: FOSSIL PEARLS 369

It has already been intimated that the Niobrara pearls were pro-
duced by a species of Inoceramus. The large size of the pearls predi-
cates a large mollusk as host, and this suggests that Inoceramus
[Haploscapha] grandis may have been the species, but certainty con-
cerning this can not be established until a pearl is found in a shell or
portion of shell which can be specifically identified.

The Benton pearls are of various forms: spherical (Fig. 22), oblate,
hemispheric (Figs. 7, 21), oblong, and irregular (Fig. 10). These pearls
are, in general, darker in color than the Niobrara specimens. The
largest is an irregularly rounded baroque, 3.5 cm in diameter, with a
corrugated surface. The surfaces of these pearls display the distinctive
polygonal pattern noted in the Niobrara pearls (Fig. 20), and there is
no doubt that the Benton pearls were also produced by one or several
large species of Jnoceramus, for a number of specimens are still at-
tached to portions of shell showing the characteristic hinge line.

Figs. 7, 8, and 9 represent a specimen, hemispherical in general
form, its top (Fig. 7) slightly flattened with a depression in the center.
The under side (Fig. 9) displays a series of concentric growth rings,
but at the outer edge can be seen the now ragged margin of the smooth
coating of the upper surface lapping over on the under surface. This
condition can be matched by the structures observable on the under
sides of modern hemispheric pearls (Figs. 17, 18) from the common
clam, Venus mercenaria. The dimple on the top side also finds its
counterpart in some modern pearls. The development of hemispheric
pearls is explained by Kunz and Stevenson (18, p. 57) as follows:

When a growing pearl is very near to the nacreous lining of the shell, the
pressure between the two hard substances results in a rupture of the pearl-
forming sac and the epithelial layer of the shell, and the pearl comes in
actual contact with the nacre. The pearl gradually becomes attached to the
shell, and the under portion is prevented from growing further; the upper
or exposed surface receives other layers, resulting in the formation of a
bouton.

Figs. 10 and 11 represent the top and under sides of an irregular,
flattened pearl. In structure and appearance this specimen seems un-
related to those already described. On the under side, however, ec-
centric lines of growth around a distorted nucleus can be detected,
and the smooth patch on the upper side under a 14-power hand lens
displays the same polygonal pattern as the surface of the other pearls.
Because this pearl is thin, flat, and irregularly but smoothly lobed or
corrugated it would seem to compare best with a modern baroque or
perhaps a blister pearl. The latter are said to be formed when foreign

370 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 9

TABLE 1.—DISTRIBUTION OF FossiL PEARLS. (Numbers refer to citations. —
B, N =Benton and Niobrara pearls described in this paper.)

Binet Nal Tri-| Juras-| Cre- Bocene | Olusoae Nias Plio- | Pleisto-
assic| sic | taceous cene | cene | cene cene
Ceratites compressus .| 16
Exogyra texana...... 1, 24
Gryphaea dilatata.... 14, 21
Ean purites Spears 17
Inoceramus expansus. 22
goldfussianus.. . 10
Labtatusinn- ae 22
SQUCRSUS re 27
subundatus..... 24
spp. 14,21, 22,

Emarscabna. oe 14
Megalodus amplus...| 19
Melina zealandica... 20
Mytilus edulis...... 14
LLIGOMUS . ae 9
Ostrea cymbula...... 6
eduliformis..... a
CUULIS ie ona 14

CONOR Gigs hoes ee 14

SPoaees uaa ae 6, 29, 30
Panopea americana. . ; 2
Rerna heberti.. 52. 33
TOP PONKTHITS 5 5 5 ot 12
ObDIORGG ee 32, 33
sandbergert..... BA, Be
SCCVCT A cin alan ausee Dale, PAO, B38
JERI: OPUDOBE aoe 6 13, 24
Pleuronautilus pseudo-
DIGNTHALERUS so: 15
PLCThUG? Sh ae ee 24
Volsella modiolus.... 23
is Ane Ri tue 3 | 4, 33

bodies such as sand grains become lodged between the mantle and the
shell, or when a mollusk attempts to repair injuries to the shell.

Many of the fossil pearls so far reported were derived from species
of Inoceramus of the Cretaceous period, a circumstance that would
seem to label these species as the pearl ‘‘oysters’’ par excellence of
that time, 100,000,000 years ago. The only other Cretaceous records
are from species of Perna, Exogyra, and Hippurites.

No definite pearls have been recorded from the Paleozoic, but
J. Smith? reports from the Upper Silurian:

Pearls (?).—In the shale at Sedgley, Woolhope, Dormington, Lincoln Hill,
Benthall Edge, and Gleendon Hill, a number of minute spheroidal bodies

occurred. Some are silvery white, some yellowish, and others of a dark-brown
color when viewed by reflected light. All of them have a peculiar pearly

2 Smitu, J. Notes on a collection of bivalved Entomostraca and other Microzoa from
the Upper Silurian strata of the Shropshire District. Geol. Mag. n. ser., dec. 2, 3: 72.
1881.

SEPT. 15, 1940 BROWN: FOSSIL PEARLS 371

lustre. One that had been split through the middle shows a concentric lami-
nated structure. On washing some of the rotten limestone from the Much-
Wenlock quarries, I found these pearls (?) very abundant.

All the Mesozoic and Cenozoic periods are represented by examples.
In Table 1 showing the distribution of fossil pearls as now known the
numbers refer to citations which, in several instances, include more
than one occurrence. |

It will be observed that all the mollusks in the list, with the excep-
tion of Ceratites and Pleuronautilus, are bivalves (pelecypods). Theo-
retically, however, any mollusk that builds a shell by means of a
mantle that, besides depositing the horny periostracum or outer layer
and the calcareous prismatic middle layer, also secretes the calcareous
nacre or mother-of-pearl] of the inner layer, should be able to produce
pearls. Today most commercial pearls come from species of bivalves:
pearl “‘oyster’’ (variously assigned to Pinctada, Meleagrina, or Mar-
garitifera), sea mussel (Mytilus), fresh water mussels (Unio, Elliptio,
Anodonta, Dipsas); but species of many other genera also yield pearls.
Among univalves (gastropods or snails) the following are known to
produce pearls: common conch (Strombus gigas), abalone (Haliotis),
and chank (Turbinella). The pearly nautilus (Nautilus pompilius),
a cephalopod, sometimes produces pearls of a yellowish color.

The inevitable question always asked concerning pearls is, How do
they originate? Considerable investigation has been conducted in
attempts to penetrate the mollusks’ secret, with the result that some
specific conclusions now seem sound. To understand the causes and
methods of pearl formation it is necessary first to consider what the
end product, a pearl, is. The usual spherical pearl is a calcareous con-
cretion composed of a nucleus and many more or less concentric layers
or laminae of minutely platy aragonite (mother-of-pearl, nacre,
CaCO;), separated by thin layers of organic conchiolin (C3>H1sN 2Ou).
Spherical pearls, however, are only one of the many forms pearls may
take. These forms depend largely on the position of the pearls in the
mollusk, whether attached to the shell or free in the mantle or muscle.
Pearls originally free may eventually be buried by enveloping shell
growth. The free pear! is formed within a sac lined by epithelium, the
layer of the mantle that secretes the different parts of the molluscan
shell. The methods by which epithelial sacs are formed and how they
enclose instigating nuclei are discussed and illustrated by Dakin (5,
pp. 91-116), and Boutan (4, p. 294).

Inasmuch as different areas of the mantle surface deposit different
parts of the molluscan shell, how does it happen that epithelium de-

372 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 9

rived from one spot in the epithelium of the mantle and now lining
an isolated pearl sac can produce a pearl that duplicates the com-
position and structure of all the substances of the host shell? Dakin
(5, pp. 24, 25) answers:

If a mollusc shell is broken (or a piece of the shell removed) some distance
away from the edge, a most extraordinary thing happens, which is of the
greatest importance in regard to theories of pearl formation. The shell is
repaired, but the repair substance is not, as was once thought, only nacre.
The mantle cells, which normally would be secreting nacre, now secrete at —
the area of breakage, first periostracum, then prismatic layer, and finally
mother-of-pearl. Thus, although certain parts of the mantle surface secrete
normally certain definite parts of the shell, the cells which secrete the nacre
are capable of secreting all the other layers if necessary.

It follows that the segregated epithelium surrounding the nucleus
of a pearl may deposit layers of both calcareous and organic sub-
stances. Part of the epithelium may even be secreting aragonite, while
simultaneously another part is depositing conchiolin. Next to an in-
citing nucleus, therefore, epithelial cells play the essential role in the
process of pearl formation.

The importance of the epithelium was demonstrated by the experi-
ments of Alverdes and also by the practical experience of Mikimoto
with culture pearls. In the modern culture pearl industry as practiced
by the Japanese, a pellet of nacre is first placed on the epithelium of
the mantle of one mollusk. After enclosing the pellet and suturing, this
part is excised and transplanted to the mantle of another mollusk,
which is then returned to the water. Upon removal, after a period of
seven years, about 60 percent of the mollusks yield pearls. Of these,
©) percent are said to be marketable (8, p. 423).

Although the age-long empirical methods of the Chinese in arti-
ficially stimulating mollusks to produce pearly growths by introduc-
ing foreign objects, such as small images of Buddha, between the shell
und mantle, suggest that some natural pearls may originate from the
accidental intrusion of grains of sand, particles of mud, fragments of
shell, ete., nevertheless evidence is accumulating to the effect that
many, and perhaps most, natural pearls have their inception from
particles originating within the epithelium of the mantle, or from
parasitic larvae of worms that become encysted within the epithe-
lium. In the latter case, if the mollusk is not devoured by that one of
its predators which is the next required host in the predestined life
cycle of the worm, a death sentence automatically passes upon the
larva, and it is forthwith entombed in a pearly mausoleum. To quote
Kunz and Stevenson (18, p. 45): ‘‘Some truth exists in the statement
that the most beautiful pearl is only the brilliant sarcophagus of a

Sept. 15, 1940 BROWN: FOSSIL PEARLS 31d

worm.’’ Some of the Benton and Niobrara pearls, therefore, may be
mute testimonials to the Cretaceous existence of a cestode or trema-
tode of which we may never know anything more. Some may also
indicate shell-boring predators.

The writer is indebted to many of his valleapues for help in this
study, but particularly to Dr. W. F. Foshag, curator of mineralogy
at the U. 8. National Museum. Specimens (Figs. 1, 5, 7, 14, 21, 22)
are deposited in the U. 8. National Museum.

REFERENCES

1. Apxins, W.S., and Winton, W. M. Paleontological correlation of the Fredericks-
burg and Washita formations in north Texas. Univ. Texas Bull. 1945: 64. 1919.

2. Berry, CHartes T. A Miocene pearl. Amer. Midl. Nat. 17: 464-470. 1936.

3. BoOTrTGER, Oskar. Ueber die nachweisbaren Spuren des Lebens der Tier und
Pflanzenwelt in der Vorzert. (Full citation not obtained, but see Kirchner 16,
p. 149.) 1867.

» Boutan, L:., La perle, p. 67... Paris, 1925.

Dakin, W. J. Pearls. New York, 1913.

DartTEVELLE, EK. Les perles fossiles. Journ. Conchyl. 78(3): 169-195. 1934.

Dorn, Pauu. Fossile Perlen in Ostreen des Dogger Delta Schwabens nebst palao-

geographischen Bemerkungen. Zentr. Miner. Abt. B (7): 295-304. 1937.

. ENCYCLOPEDIA BRITANNICA, ed. 14, 17: 420-423. 1929.

. FRENGUELLI, JOAQUIN. Sobre una perla fosil del Aonikense de Punta Norte en la
peninsula Valdez (Chubut). La Plata Univ. Nac. Inst. Mus. Notas 2, Paleon-
tologia 11:155-162. 1937.

10. Goupruss, A. Petrefacta germaniae 2: 116, pl. 112, fig. 4d. 1886.

11. Haas, F. Fossile Perlen. Nat. u. Mus. 61(3):120. 1931.—Bau und Bildung der

Perlen. Leipzig, 1931.

12. Hornes, M. Die fossilen Mollusken des Tertiar-beckens von Wien, Bd. 2. Abh. -
geol. Reichsanst. 4. 1867.

13. Jackson, J. F. Fossil pearls. Proc. Isle of Wight Nat. Hist. Soc. 1: 466. 1926.

14. Jackson, J. W. On some fossil pearl growths. Proc. Malac. Soc. London 8: 318,
pl. 14, figs. A-E. 1909.

15. Krestineer, Atois. Die Nautiliden der Mittleren und Oberen Trias von Timor.
Jaarboek van het Mijnwezen en Nederlandsch Oost-Indie, s’ Gravenhage.
Verhandlungen 1. 1922.—Untersuchungen an triadischen Nautiloideen. Pa-
laeont. Zeitschr. Berlin 7: 101-122, fig. 7. 1925.

16. Kircuner, H. Perlbildung bei einem Cerititen. Centralbl. fiir Min., Geol. und
Palaont. 4: 148-152. 1927.

17. Kuinenart, Franz. (Full citation not obtained, but see Kirchner 16, p. 149.)
1922.

18. Kunz, GrorGeE F., and STEVENSON, CuHarutes H. The book of the pearl. .548 pp.,
with bibliography. New York, 1908.

19. Kurassy, ENpre. Dve dlteste fossile Perle und Verletzungsspuren an einem tra-
adischen Megalodus. Math. und Naturw. Anz. (Ung. Akad. Wiss.) 55(3):
1005-1023. 1937.

O00 NOR

20. Marwick, J. Fossil pearls in New Zealand. New Zeal. Journ. Sci. 5: 202. 1922.

21. Morris, J. Paleontological notes. Ann. Mag. Nat. Hist. 8: 85-90, pl. 4. 1851.

22, Newton, R. B. Fossil pearl growths. Proc. Malac. Soc. London 8: 128-139.
1908.

23. Ropertson, D. On the post-Tertiary beds of Garvel Park, Greenock. Trans. Geol.
Soc. Glasgow Mel) PA Sie ASS oe

24. Russet, Ricuarp D. Fossil pearls from the Chico formation of Shasta County,
California. Amer. Journ. Sci. 18: 416-428. 1929.

25. Rutot, A. Nouvelles decowvertes faites dans le Tongrien inferreur du Limbourg, par
M. le Comte Georges de Looz-Corswaren. Ann. Soc. Malac. Belgigue 14. Bull.
des Seances, p. 77. 1879.

26. SpELEY, H.G. Notes on Cambridge paleontology. Ann. Mag. Nat. Hist. 7: 1-21,
pl. 6, ‘fig. Grp glSok

27. WHITFIELD, R. P. Brachiopoda and Lamellibranchiata of the Raritan clays and
Greensand marls of New Jersey. U.S. Geol. Survey Mon. 9: 79, pl. 14, fig. 16.
1885.

374 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

28. Woopwarp, J. Essay toward a natural history of the earth 3:24. 1723.

29. WrietEy, A. Thelower Eocene Mollusca of Abbey Wood and High Halstow (Kent).
British Mus. Nat. Hist. 1931.

. A Lutetian fauna of Southampton docks. Proc. Geol. Assoc. 45(1): 1-16.

1934.

31. YoxoyamMa, M. Versternerungen aus den japanischen Kreide. Palaeontogr. 36:
ial: 18, fig.6. 1890.

OZ: ZILCH, Apotr. Eine Perle aus der Meereszeit der Wetterau. Nat. und Volk 64:
93-95. 1934.

. Unsre Kenntnis von fossilen Perlen. Archiv. fiir Molluskenk. 68(6): 238-

252, pl. 10. 1936.

PALEONTOLOGY.—Two new corals from the Avenal formation
(Hocene) of California: JoHN W. WELLS, Ohio State University.
(Communicated by Joun B. Rexsipe, JR.)

The two new species of solitary corals described below were col-
lected by Dr. Ralph Stewart, of the United States Geological Survey,
who kindly turned them over to the writer for description. They are
particularly interesting because of their close relationships with EKo-
cene corals of the Gulf Coastal Plain and the Antillean regions.

Family CALAMOPHYLLIIDAE
Genus ANTILLOSERIS Vaughan, 1905
Antilloseris? vaughani, n. sp. Figs. 1, 2

Corallum solitary, turbinate, flaring rapidly from a subcylindrical base.
Calice circular, shallow. Corallite wall synapticulothecal, thickened to a
septotheca basally, irregularly perforate near calicular margin, nonepithe-
cate, costate. Costae corresponding to all septa, equal, crispate or finely
granulose laterally and on margins. Septa very numerous, in six systems,
probably with six complete cycles and part of the seventh (about 250 septa
in holotype). Septa thin, mostly imperforate and laminar, except those of
the first two cycles which are relatively thick and laterally and basally sub-
splnose, with inner ends free and thickened. Synapticulae rare except in
mural region. Endotheca and exotheca absent. Columella wholly absent.

Dimensions of holotype.-—Diameter of calice, 20 mm; height of corallum,
7 mm; diameter of corallum 7 mm below calice, 7 mm.

Holotype.—U.S.N.M. no. 498697.

Type locality.—U. 8. Geological Survey locality 14482; California, Kettle-
man Plain quadrangle, T. 23 8., R. 17 E., sec. 35, 400 feet south of north
boundary and 2,110 feet east of west boundary of section, near Little Tar
Spring. Avenal formation, near Domengine horizon.

Remarks.—The agaricid nature of this coral is shown by the thin, slightly
and irregularly perforated septa and by the synapticulothecal wall. The ab-
sence of any dissepiments, even basally, and of any trace of columella further
indicates a generic assignment in or very close to Antilloseris, a genus con-
fined to the Eocene and heretofore found only in the Antillean region. In all
the described species? of Antzlloserzs, however, the corallum is more or less

1 Received April 10, 1940.
2 Duncan, P.M. Quart. Journ. Geol. Soc. London 29: 558-560, pls. 21, 22. 1878.
VauGcHAN, T. W. Bull. Mus. Comp. Zool. 34: 245-246, pl. 40. 1899. In both these
papers the species included by Vaughan in Antilloseris in 1905 are described as Tur-
binoseris.

Sept. 15, 1940 WELLS: TWO NEW CORALS 375

compressed and tapers basally to a very small point of attachment, and
since the present specimen is not compressed and apparently possessed a
broad base of attachment, the reference to this genus is not positive.

Family FUNGIIDAE
Genus Discotrocuus Edwards and Haime, 1848
Discrotrochus californicus, n. sp. Figs. 3-5

Corallum solitary, depressed-cupoloid in shape, with flat, circular base
with faint scar of very early fixation. Corallite wall septothecal, thick, solid,
horizontal, nonepithecate, costate. Costae corresponding to all septa, alter-
nating regularly in size, obsolescent towards center of base. Those cor-
responding to septa of lower cycles larger, subacute with a few stout granu-
lations on edges. Septa 36 in number, in six systems, imperforate, highly
exsert, outer margins vertical, inner ones gently convex to the circular cen-
tral fossette, laterally ridged or granulated perpendicularly to margins
which are peripherally crossed by transverse denticulations where opposing
ridges meet over margins. Septa of the fourth cycle not developed in ‘‘dorsal’’

I 3 5

Fig. 1.—Antilloseris? vaughani, n. sp., holotype; oblique basal view, showing polished
section of base (note absence of columella and dissepiments), X2. Fig. 2.—Same,
lateral view, X1. Fig. 3.—Discotrochus californicus, n. sp., holotype, view of calice,
x5. Fig. 4.—Same, lateral view, X5. Fig. 5.—Same, basal view, X65.

half of each of the six systems. Columella trabecular, small, extending up-
ward about half the thickness of the corallum.

Dimensions of holotype-—Diameter of corallum at base, 6 mm; height of
corallum, 2.5 mm.

Dimensions of paratypes.— Qo ey re (4) (5)
Diameteron corallum.. >... :- OD Oo O00 0.8. Deh worry,
herent or corallum..-... 0, AOe Aa) A) A, o a0

Paratype 2 1s much worn.

Holotype.—U.S.N.M. no. 498698.

Paratypes.—U.S.N.M. no. 498699, five specimens.

Type locality—Same as for Antilloseris? vaughant.

Paratype locality —U. 8. Geological Survey locality 14482a. Same horizon
as type locality but on next ridge to east of type locality.

Remarks.—This species is based upon a holotype (the first collected) and
five paratypes from a nearby locality. All are in a fair state of preservation

“Kinks 2

sa ee

376 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 9

and show the characters of Dzscotrochus, a genus associated by most authors
with the Caryophyllinae. Recent study of the type species, D. orbignianus
Edwards and Haime,’ by Dr. T. W. Vaughan and the writer, however, indi-
cates that Dzscotrochus is scarcely distinguishable from the fungid Cycloseris
except for the rarity of synapticulae, smaller number of septa, and thicker
basal wall in Discotrochus. The present species is distinguished from the
genotype, and only other American species, D. orbignianus, of the lower
Claibornian (Kocene) of the Gulf Coastal Plain, by the different proportions
of the corallum. The ratio of height to diameter of D. orbignianus is 1:4,
in D. californicus, 1:2.5. The latter is therefore relatively much higher than
D. orbignianus. Further, D. orbignianus usually possesses four complete
cycles of septa, whereas all the specimens of D. californicus examined show
the fourth cycle only half developed in each of the six systems.

PALEONTOLOGY .—A new Gisortia.1. WiLL1am M. INGRaM, Cornell
University. (Communicated by Juntia A. GARDNER.)

The Gisortia described here, from the Capay stage, Llajas forma-
tion, lower zone, Simi Valley, Ventura County, Calif., is the only spe-
cies belonging to this genus thus far reported from North America.
It is one of the species of mollusks used by Clark and Vokes (1936) for
intercontinental correlation of west-coast Eocene and European Eo-
cene faunas. These workers have compared it to Grsortia tuberculosa
(Duclos) from the Ypresian stage, Paris Basin, France. They state,
“The California specimen |[Gisortia clarki Ingram] is somewhat more
globose and the outer lip is not as prominently reflected posteriorly as
in the Paris Basin form [Gisortia tuberculosa (Duclos)]. The evidence
indicates that in both species the spire was submerged beneath a cal-
lus wash.’’?

Gisortia clarki, n. sp.

Shell globose, heavy; posterior canal prominently produced, and covered
dorsally by a shelf 8.5 mm broad on the outer side; dorsally on the columellar
side this shelf narrows to a width of approximately 3 mm and slopes abruptly
toward the ventral shell surface; posterior canal produced 11 mm; spire
almost totally submerged beneath outer enamel, the spire peak projecting
3.5 mm from the body of the shell; a flattened surface persists to the left
of the posterior canal; maximum width of posterior canal is 14 mm; esti-
mated maximum width of anterior canal about 16 mm, a shelf with a maxi;
mum width of 5 mm occupies the columellar side of the anterior canal-
this shelf is angled dorsally from the shell base; the anterior canal is ap-
parently compressed dorsoventrally at its outermost extremity; outer and
columellar lips of aperture broadly rounded; aperture curves to the left
anteriorly and posteriorly.

3 VauGHAN, T. W. U.S. Geol. Surv. Mon. 39: 78-80, pl. 5, figs. 13-19b. 1900.

1 Received February 2, 1940.

2 CLARK, B. L., and Voxss, H. E. Summary of marine Eocene sequence of western
North America. Bull. Geol. Soc. Amer. 47: 851-878. 1936. |

Sept. 15, 1940 INGRAM: A NEW GISORTIA Ot7

Dimensions—Length 121 mm, breadth 94 mm, height 64 mm.

Holotype.— Univ. California, Mus. Pal., invertebrate collection, no. 14844.

Type locality—Locality no. 4052, Museum of Paleontology, invertebrate
collection; Capay stage, Llajas formation, lower zone, Simi Valley, Ventura
County, Calif,

The holotype specimen is moderately eroded on the dorsal surface. The
greater part of the dorsal shell convexity is represented by an intact internal

Fig. 1.—Gisortia clarki, n. sp. Dorsal view. Approximately natural size.

mold. The shell substance is well preserved on the base, the extreme pos-
terior region, columellar lateral shell boundary, and on the posterior three-
fourths of the outer lip. The aperture is filled with a matrix, which prohibits
a description of the internal surfaces of the columellar and outer lips.

This species is named in honor of Dr. Bruce L. Clark, of the Department
of Paleontology of the University of California, Berkeley, Calif. The photo-
graph used here was obtained through his courtesy. The holotype specimen
was also sent to the writer by Dr. Clark.

378 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

BOTANY.—Spore formation in Matula.1 G. W. Martin, State
University of Lowa.

The genus Matula was established by Massee (Journ. Royal Mier.
Soc. 1888: 173) to accommodate M. poroniaeformis, a fungus first de-
scribed from Ceylon by Berkeley and Broome (Journ. Linn. Soc. Bot.
14: 73. 1873) as Artocreas poroniaeforme. Berkeley believed that the
species so named was congeneric with Michenera Artocreas Berk. and
Curt., but by an error he transposed the generic and specific names
of the latter species, writing it, in a note appended to the description
of A. poroniaeforme, Artocreas Michenert. Shortly afterward (Journ.
Linn. Soc. Bot. 15: 83. 1876) he corrected the error, without, how-
ever, actually publishing the combination Michenera poroniaeformis,
which he had evidently meant to use in the first place but which seems
to have been first printed in the Sylloge Fungorum 6: 653. 1888. Lloyd
(Myc. Writ. 3: 443. 1909) examined the specimens at Kew and satis-
fied himself that Berkeley had really transposed the names, and Petch
in connection with his careful study of the Ceylon species (Trans.
Brit. Mye. Soc. 11: 67-81. 1926) checked the situation independently
and came to the same conclusion.

Massee restudied Berkeley’s material and, believing that he saw
basidia, made Matula the type and sole representative of a new “‘or-
der’’ of Gasteromycetes, the Matuleae, which he regarded as inter-
mediate between the Nidulariaceae and the Hymenogastraceae.
Many years later Rick (Broteria 5: 10. 1906), referring to what he
regarded as a distinct species, Michenera Rompela Rick, from Brazil,
said: “Es gehort natiirlich unter die Hymenogastrineae als Hemihy-
menogastria.”’ Lloyd (Mye. Writ. 2: 391. 1908) transferred Rick’s
species to Matula, and said, referring to both species: ‘““The relations
appear to be entirely with the Nidulariaceae.’’ Petch, however, pre-
sents convincing evidence that the Ceylon species is an imperfect
stage of Peniophora habgallae Cke.

Although Massee’s supposed discovery of basidia seems clearly to
be based on misinterpretation of what he saw under the microscope,
his generic name has been very generally adopted. Petch, however,
uses the generic name Artocreas on the ground that the species de-
scribed by Berkeley as A. poroniaeforme is generically distinct from
Michenera and was technically validly published. With this interpre-
tation I can not agree. Berkeley believed his A. poroniaeforme be-
longed in the same genus as Michenera Artocreas, and his publication

1 Received March 21, 1940.

Sept. 15, 1940 MARTIN: SPORE FORMATION IN MATULA BYA)

of Artocreas as a genus was no more than the result of a trick of mem-
ory, as he himself pointed out. Such publication certainly does not
comply with the conditions clearly stated in Article 37 of the Inter-
national Rules. Massee was the first to recognize the generic distinct-
ness of Matula, and, however erroneous certain of his conclusions
may have been, his description does bring out the characteristics by
which the genus may readily be recognized. So long as it is regarded as.
distinct from Michenera, his name should be applied.

Berkeley described the spores of A. poroniaeforme as 0.0009 inch in
diameter (24-25u). Michenera Rompelii Rick (Ann. Myce. 2: 243.
1904) was based largely on the supposedly smaller spore size, less than
20u. Lloyd reported the spores of both species to be 18—20u in diam-
eter and concluded they were probably the same. Petch, however,
noted certain differences between them, particularly a continual basal
layer beneath the cup in Rompelzz, which he finds lacking in poroniae-
forms, and concludes that it is doubtful whether they are identical.

A species of Matula was fairly common in western Panamdé in
July 1935, being represented by five collections. Four of these are
from the valley of the upper Rio Chiriqui Viejo, above 1,600 m., and
the other is from the forest south of the Llanos del Volcan between
1,100 and 1,200 m. I have also a single collection from the Sierra
Nevada de Santa Marta, Colombia, between 1,250 and 1,500 m.
gathered in August of the same year. In addition, I have examined
one of Rick’s collections from Brazil, now in the Lloyd Herbarium at
Washington (no. 22541) and a collection of G. H. Cunningham’s from
New Zealand, now in the herbarium of the Missouri Botanical Gar-
den. All, I am convinced, represent the same species. Some fructifica-
tions appear to have the continuous basal layer beneath the cup
referred to by Petch as characteristic of the American forms, but
others lack it, appearing in section, except for the absence of bark
fragments imbedded in the fructifications, essentially like the forms
illustrated by Petch in figures 1—5 of his plate 3. As in the Old World
specimens, the fructification is at first subglobose and closed, opening
above very early and assuming the characteristic cupulate form. The
spores are borne in peridiole-like chambers, originating from the walls
of these chambers. In the older portions near the surface the walls
disappear, at which time this part has become a solid mass of the
large, thick-walled spores, held together by their gelatinous walls
(Fig. 2). The fructification is tough-gelatinous when fresh, drying
somewhat horny, but readily regaining its original texture and ap-
pearance when soaked (Fig. 1). The temptation is strong to regard

380 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 9

the New World and New Zealand specimens, on the one hand, and
the Ceylon specimens studied by Berkeley, Massee, and Petch, on the
other, as referable to the same species. The manner of spore develop-
ment in the former group, as I interpret it, differs decidedly, however,
from Petch’s account for the Ceylon form, and while a careful reading
of Petch’s paper makes it seem not entirely impossible that his is
merely a different interpretation of the same structures, in the ab-
sence of opportunity to examine Ceylon collections, it seems proper
to refer the specimens here discussed to Matula Rompelii (Rick) Lloyd.

Fig. 1.—Matula Rompelii (Rick) Lloyd: Group of four fructifications, soaked, X3.
Fig. 2.—Matula Rompelit (Rick) Lloyd: Longitudinal section through fructifica-
ee center, showing peridium, basal chambers, and solid mass of spores at surface,

According to Petch, the spores of poroniaeforme are borne on coni-
diophores that produce first a terminal spore, additional spores being
borne basipetally. The lumen of the young spores stains deeply but the
wall surrounding it remains hyaline and gelatinous, fusing with ad-
joining hyphae so that the outline is indistinct. Finally, the central
protoplasmic mass develops an independent wall of its own, and the
wall of the parent cell deliquesces, freeing the spore. We have, then, a
conidium formed within a sac, surely a curious phenomenon. Petch
argues that such a spore can not be a chlamydospore and regards the
spore of Matula poroniaeformis as homologous with that of Michenera
Artocreas, the essential difference being that in the latter species the
wall of the mother cell persists as an apical appendage and a basal
pedicel, while in the former species the wall gelatinizes and disap-
pears. Petch notes that the hyphae of Matula bear clamp connections,
but he makes no mention of any relationship between such structures
and the spores.

Sep. 15, 1940 MARTIN: SPORE FORMATION IN MATULA aol

In the American forms studied, including Rick’s Brazilian collec-
tion, and also in the New Zealand material, clamp connections are
abundant on the hyphae at the bases and sides of the spore sacs,
where the spores are being formed (Fig. 3, a). but are only rarely and
obscurely visible elsewhere. I have never seen any suggestion of
conidiophores, as described by Petch. The spores seem always to
originate as intercalary structures on the hyphae and always at the
apical side of a clamp connection. This is uniformly apparent in the
young spores but progressively less so as the spores mature. The con-
tents of the apical cell are retracted toward the septum, causing a
pronounced swelling (Fig. 3, b, c, d). The lumen stains deeply, while

5 Ae
Fig. 3.—Matula Rompeliz (Rick) Lloyd: a, A clamp connection from region of
spore formation; b, c, d, early stages in spore formation, showing accumulation of
protoplasm on distal side of clamp connection; e, f, g, h, progressive stages in develop-
ment of spore; 7, spore freed from basal hypha, projections arising from lumen; 7, ma-
ture spore with double wall. All 1,000.
the outline of the thick, gelatinous wall is very faint. Sooner or later
this wall ruptures or is dissolved, and then the spore appears to be
terminal (Fig. 3, d, e, h). Finally it breaks loose from the stalk and
gradually becomes globose, apparently developing, in its later stages,
at the expense of the gelatinous matrix in which it is imbedded. At
complete maturity (Fig. 3, 7) the lumen occupies approximately two-
thirds of the total diameter of the spore. The wall is composed of two
layers, an outer highly refractive portion, at complete maturity 0.6-
0.8u thick, and an inner clear layer 3—4y thick or somewhat more. Oc-
casionally the inner layer is divided by faint, concentric rings into
two or more portions. Six or eight conical projections arise from the
lumen and reach nearly or quite to the outer spore coat. Two or three
of these are usually visible when the lens is focused on the optical
center of the spore. It is not improbable that these represent germ-

382 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 9

pores. The outer wall above them appears, however, to be perfectly
homogeneous. It seems clear that spores formed in the manner de-
scribed can not be referred to as conidia, and the term chlamydospore
comes nearer than any other to describing their true nature.

The spores of the Brazilian specimen are more variable in size than
those of the Panama and Colombia material, some of them reaching
28u in diameter, and the peglike protrusions from the lumen, while
often present, are usually fewer in number and sometimes not evident.
I doubt whether such differences are of taxonomic significance.

No suggestion of possible connection with a perfect stage is to be
found in any of the material at hand. Lacking information to the
contrary it may be assumed that any perfect stage will prove to be a
member of the Thelephoraceae.

ZOOLOGY.—Boceardia proboscidea, a new species of spionid worm
from California. OLGA Hartman. (Communicated by WaLpo
L. ScHMITT. )

The commonest representative of the family Spionidae, of the genus
Boccardia, occurring in the intertidal zones of California, remains un-
named and undescribed. This species is of interest not only because it
occurs in abundance in the littoral zones where it is readily available
but also because of several biological features, which may warrant
more thorough investigation.

My attention was first called to this species several years ago, when
I was a member of Prof. 8. F. Light’s courses in invertebrate zoology
at the University of California. At that time its designation was ques-
tionable, and after a perusal of the literature I (Univ. California Publ.
Zool. 41: 48. 1936) referred it to Boccardia natrix (Séderstrém)
without, however, consulting Sdoderstrém’s type materials. Since then
the species has been noted many times along the coast of California.
At the Scripps Institution of Oceanography, through the courtesy of
Dr. Martin W. Johnson, it was possible to observe it also as an ele-
ment in the plankton and to maintain adults and larvae under labora-
tory conditions.

This study has been supported through a grant from the American
Association of University Women. I wish to express my thanks not
only for the privileges already mentioned, but also to Dr. Waldo L.
Schmitt, of the United States National Museum, for further cour-
tesies.

1 Received March 30, 1940.

Snpt. 15, 1940 HARTMAN: A NEW SPIONID WORM 383

Boccardia proboscidea, n. sp. Fig. 1

?Polydora californica Treadwell, Univ. California Publ. Zool. 13: 203-204,
pl. 12, figs. 23-29. 1914 (not Spzo californica Fewkes, 1889, which is a
Polydora, sensu latiore).

Boccardia natrix Hartman, Univ. California Publ. Zool. 41: 48. 1936.
Journ. Washington Acad. Sci. 26: 32. 1936 (not Polydora natrix Séder-
strom, 1920; not Berkeley, 1936).

The general form of the body is long, depressed, widest in the region of
the eighth to sixteenth segments, and tapers gradually posteriorly. Number
of segments is about 125 to 150; total length 30 to 35 mm but capable of
much greater extension in life; greatest width (preserved) is about 1.5 mm.
Color in life is deep yellow to orange, with bright-red branchial filaments,
and considerable dusky pigment along the prostomial ridges and palpal
grooves; in preserved individuals all color fades except the sooty markings
along the prostomium and some along the palpal length.

The prostomium is an elongate, entire lobe, without median groove such
as characterizes most species of this genus; a weakly developed emargination
is visible only on the ventral side (Fig. 1, a). The lobe is snoutlike (for which
the specific name is proposed), it extends posteriorly between the palpal
bases where it forms a low crown visible in lateral view (Fig. 1, d) and passes
more posteriorly as a low, broad ridge to the posterior margin of the third
setigerous segment (Fig. 1, b). In the region just anterior to the insertion of
the palpal bases there are two or three pairs of dark eye spots, clearly visible
when the palpi are pushed somewhat to the side. In immature specimens
there may be several additional dark spots in these areas, but at least two
pairs persist even in the largest individuals. The prostomial lobe is sharply
set off from the peristomium by grooves at the sides of the snoutlike eleva-
tion. The lateral and ventral sides of the oral opening are bounded by the
apodous, achaetous peristomial ring (or first segment). Except for some
longitudinal and transverse furrows and some wrinkles of contraction, it is
quite smooth (Fig. 1, a, b, d).

The second segment (herein designated the first setigerous) is proximal to
the peristomium; it bears a dorsal and a ventral fascicle of pointed setae,
each provided with a small postsetal lobe (Fig. 1, d). These parapodial
structures are comparatively weakly developed, less than half as large as
those of the following segment, the setae both fewer in number and shorter.
The second setigerous segment resembles the next two. There are well-
developed setigerous fascicles, the dorsal and ventral postsetal lobes are
auricular, and there is a long, filamentous branchia (Fig. 1, d). The fifth
(modified) setigerous segment is nearly twice as long as the preceding; it is
provided with a dorsal fascicle of stout hooks and an inconspicuous ventral
fascicle of capillary setae (Fig. 1, 7); there are no postsetal lobes. The stout
hooks are of two kinds—a longer, falcate, smooth hook, accompanied by an
equally heavy, though shorter, bushy-topped seta. In unworn condition
(Fig. 1, h) the falcate spine terminates distally in a tapering fang; the other
has a characteristic subterminal constriction, then widens suddenly and
ends in a spinous cap that is strongly asymmetrical (Fig. 1, h). In worn con-
dition both of these spines lose these characters (Fig. 1, 7), but the cuplike
base of the latter is still notable.

The next (sixth setigerous) segment is provided with pointed setae in
both dorsal and ventral fascicles; its structures are similar to those of the
second to fourth segments, but the postsetal lobes are progressively larger.
From the seventh the neuropodium is provided with about eight hooded

\ a
0.5mm 0.1m ™ 0.1 Tm ™ 0.03 ™ ™m

Fig. 1.—Boccardia proboscidea, new species: a, Anterior end, in ventral view, show-
ing the rounded margin of the prostomial lobe and the peristomial ring bounding the
oral aperture, followed by the first two setigerous segments; b, the same, in dorsal view,
with only the palpal bases shown, the prostomial caruncle continued posteriorly; the
eye spots are largely hidden because of their position at the sides of the prostomial lobe;
c, anal disk in posterior view, showing the 4-lobed arrangement; d, anterior end from
the left side, the left palpus removed so that the eyes may be seen, setal fascicles in-
dicated; e, posterior end from the left side, showing the prolongation of the ventral
lobes of the disk; f, parapodial gland from the ninth segment seen from the front, show-
ing its characteristic outline; g, hooded crotchet from tenth neuropodium; h, com-
panion stout setae from the fifth setiger in unworn condition; 7, pointed seta from in-
ferior fascicle of the modified fifth setigerous segment; 7, worn companion setae from
the fifth setigerous segment.

Sept. 15, 1940 HARTMAN: A NEW SPIONID WORM 3890

crotchets, disposed in a single vertical series, and a smaller, inferior fascicle
of about five or six slender, pointed setae. The latter persist through about
four segments, but from the eleventh there are only hooded crotchets. Noto-
podia are provided with only pointed setae throughout. Hooded crotchets
of anterior and posterior segments resemble one another. The distal end is
bifid, the lateral fang nearly at right angles to the main stem, the distal
tooth smaller (Fig. 1, g).

Branchiae are distributed on setigerous segments 2 to 4, and from the
sixth to near the posterior end; the last few (two to four) segments are
abranchiate. Setigerous glands, characteristic of members of this family,
are best developed posterior to the modified segment; they form a thick,
sacklike, slightly L-shaped, pouch (Fig. 1, f), terminating under the body
_ wall adjacent to the hooded hooks. The last setiger is followed by a broad,
flaring anal disk (Fig. 1, c, e) unequally 4-lobed, the dorsal lobes the smaller.
The anal aperture lies in the depression near the center of the disk (Fig. 1, c).

B. proboscidea is thus characterized in having branchiae distributed on
setigerous segments 2 to 4, and from segment 6 nearly to the posterior end;
the first setigerous segment has weakly developed nodopodial and neuro-
podial fascicles; hooded crotchets are present from the seventh, the first few
accompanied by an inferior fascicle of pointed setae, the rest forming a
single series of hooks only; the prostomial lobe is long, entire, snoutlike,
without median emargination; it is continued posteriorly as a ridge to the
posterior margin of the third setigerous segment; the modified segment is
provided with smooth falcate setae and bushy-topped setae in which the
spinous area is asymmetrical. Its affinities are with B. polybranchia (Has-
well), from which it differs in its entire prostomial lobe and in having setal
fascicles both dorsally and ventrally in the first setigerous segment. Another
nearly related species is B. natrix (Séderstr6m), but this has well-developed
parapodial lobes on the first setigerous segment and the prostomial lobe is
bifid; also the bushy-topped setae have paired bosses subdistally.

Holotype.—U.S.N.M. no. 20217. Caspar, Calif.

Distribution.—B. proboscidea inhabits a great variety of niches in the
intertidal zones of California, south at least to San Diego and north to
Mendocino, but its range may extend north to Puget Sound, Wash. It has
never been encountered in dredged collections from deeper water. Careful
investigations in other areas, however, may extend these ranges.

Biology.—B. proboscidea is conspicuously abundant in shale and limestone
reefs, penetrating the softer rocks, boring usually at right angles to the sur-
face, and sometimes present in such numbers as to cause the rock to break
~ away. It is also an interesting dweller in high tide pools, where hollows are
formed in sedimentary rocks. In such places the minute burrows of the
spionid may be observed at the apertures of which the two long, waving
palpi of the worm reach far out, after particles of food in the vicinity.
A common association in such pools is with a small red harpacticoid copepod,
of the genus T7zgriopus, on which the spionid feeds. Prof. 8. F. Light, of the
University of California, has often called attention to this association, as
well as to the high incidence of the Boccardia in these areas. An abundance
of the latter is usually correlated with small numbers of Tzgriopus. Its pres-
ence in such habitats is also indicative of great tolerance for variations in

386 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

salinity and temperature. Another favorite habitat is in narrow crevices in
intertidal zones. Under such conditions a loosely constructed tube surrounds
the individual.

B. proboscidea is a voracious predator, feeding not only on algal particles,
Bryozoa, Hydrozoa, and other attached organisms but actually capturing
free-swimming animals. The greatly extensile, prehensile palpi are the chief
organs of food getting; the tip senses out favorable objects for ingestion and
is firmly wrapped about any desirable object that is nipped off, taken up in
the ciliary groove, moved orally, and swept into the mouth.

Only fragmentary information is known of the life history. The eggs are
deposited in ovoid capsules, 50 or more eggs in each, and five or more cap-
sules in a tube. Aeration, produced by rhythmic movement of the adult, is
continued while the young develop. The rate of development in any one
capsule may be about equal for the various individuals, or very irregular.
Development is fairly rapid and can be conveniently observed because of
the ease with which they develop under laboratory conditions.

Capsules collected from the tube of a single adult differ among them-
selves. Some will have numerous larvae, all in a similar stage of develop-
ment, with as many as 40 or more rapidly moving young. When such a cap-
sule is split open there is a great scurry to escape. If given nourishment, they
flourish as plankton organisms and after some weeks metamorphose into
settling young. In other capsules there may be only one or two much larger
larvae, cannibalistic on the other contents in the capsule, and escaping only
when all food has been used. Whether the other eggs fail to develop because
of injury or some enzymatic influence has not been ascertained. At any rate,
when such young emerge they are ready to settle immediately. Incidence of
the larvae in the plankton is readily observed because of a unique pigmenta-
tion pattern; already in a 7- to 9-segmented stage the modified segment is
differentiated.

Systematic discussion.—Several species of Polydora (sensu latiore) have
been described from the west coast of the Americas, but the status of some
of them is stillin doubt. Unless type specimens are extant, some of the names
may need to be dropped. The first spionid to be described from California
was Spio californica Fewkes (Bull. Essex Inst. 21: 37-38. 1889) from Santa
Barbara. It is difficult (perhaps impossible) to know what this is, but it must
be considered in this discussion because of the statement, ‘“‘On the second,
third and fourth body segments, counting from the head, we find a dorsal
and ventral bundle of setae, and a dorsal and a ventral cirrus. The ventral
cirrus [postsetal lobe?] is smaller than the dorsal. In the fifth body segment
[modified segment?] there is a fan-shaped, deeply embedded bundle of large
spines in addition to the dorsal and ventral clusters.’’ There is, however, no ~
mention of the distribution of branchiae, nor is it clear that the author dis-
tinguished between branchiae and postsetal lobes, nor what is meant by
dorsal and ventral cirrus. It is not possible to distinguish it as either a Boc-

Supt. 15, 1940 HARTMAN: A NEW SPIONID WORM 387

cardia or Polydora (sensu stricto). There is another statement regarding its
tubes, which contradicts its inclusion in the family Spionidae: ‘‘The tubes
... resemble those of Sabellarza but differ from them in color, size, and form
of the openings. The edges of the orifices are sharper and the tubes them-
selves are more compact.” I know of no spionid which constructs such tube
masses. However, the head is said to have ‘‘two long tentacles’ (as in
Spionidae) but they are described as papillated (Magelonidae??). Other
features exclude it from the latter. I am unable to regard this as anything
more than a Polydora in the broad sense.

Later Polydora californica Treadwell (Univ. California Publ. Zool. 13:
203-204, pl. 12, figs. 23-29. 1914) was described, its locality given as un-
known. I have seen the single incomplete specimen, deposited in the collec-
tions of the University of California. It may be the same as the species
herein described, but since the name turns out to be a questionable homo-
nym, it should be replaced by another. A few years ago, while going over the
collections at Berkeley, I reported this as B. natrix (Univ. California Publ.
Zool. 41: 48. 1936), and identified Treadwell’s name with it. Since then,
I have seen Séderstr6m’s type of B. natrix and found it to be something
quite different. Through the courtesy of Prof. Sixten Bock, of the Swedish
State Museum at Stockholm, these materials were made available. The re-
sults are being reported on in a separate study. It is sufficient here to say
that B. natriz is an inhabitant of a sponge, from a depth of 135-150 meters,
off southern South America, that it is clearly separable morphologically from
the California species, and that B. natrix can not be applied to the latter.
B. natrix has a prostomial lobe that is clearly bifid at its anterior margin,
the groove continued for some distance along the dorsal surface; there are
eye spots but differently disposed; the first setigerous segment has unusually
well developed fascicles and postsetal lobes, nearly or quite as large as those
of the following segments, and the modified hooks of the fifth segment are
provided with paired lateral bosses. Polydora (Boccardia) natrix was later
reported by Berkeley (Ann. Mag. Nat. Hist. (10) 18: 472. 1936) from the
Nanaimo district. These specimens are partly described, showing therein

differences from Sdderstr6m’s species, but differing also from B. proboscidea.
Dr. Berkeley recorded the absence of eyes and described the prostomium
as bifurcate.

388 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 9

ZOOLOGY.—WNotes on the amphipod Gammarus minus Say and de-
scription of a new variety, Gammarus minus var. tenuipes.
CLARENCE R. SHOEMAKER, U.S. National Museum.!

Thomas Say (1818, p. 376) described Gammarus minus as follows:

Eyes reniform; superior antennae longer than the inferior ones, terminal
joint with about twelve articulations. Inhabits rivulets and small fresh-
water streams. Body whitish, with a few very pale fulvous lateral spots; eyes
blackish, placed at the exterior base of the superior antenna; superior an-
tennae obviously longer than the inferior ones, seta short, attaining the tip
of the second articulation of the terminal joint, terminal joint with about
twelve articulations. Length three-twentieths of an inch. Found in brooks
under stones, and may be readily discovered by taking a stone out of the
water, and inspecting its inferior surface.

His description would be quite inadequate for the recognition of
the species, except for his mentioning that the first antennae are ob-
viously longer than the second and that it inhabits rivulets and small
fresh-water streams. It is, as he states, an inhabitant of the small
streams, and, though he does not say streams of the east coast, he
undoubtedly means such, as all the other amphipods he described
were from the east coast. It is a very common and widely distributed
species of the cold streams of the eastern section of the United States,
in contradistinction to Gammarus fasciatus Say, which inhabits the
warmer waters of our eastern rivers.

Though Gammarus minus is so common and widely distributed,
there have been comparatively few references to it in literature since
its discovery in 1818. H. Milne Edwards (1840, p. 46) says that the
species described by Say under the name of Gammarus minus does not
seem to differ in any important character from G. fasciatus and it will
perhaps be necessary to unite them. A. A. Gould (1841, p. 334) says,
“Found [in Mass.] in ditches and sluggish fresh water, adhering to
sticks.’”’ Since the habitat he cites does not seem suitable for G. minus,
it is doubtful whether his reference applies to this species. J. E.
DeKay (1844, p. 37) says, ‘“This species is common [in New York]
in most of our fresh-water streams, and may often be detected under
stones and pieces of wood. It is extremely active, and is popularly
known under the name of Fresh-water Shrimp.” He gives a short
description, but fails to add any useful characters, and he also gives
a very small inaccurate figure. Adam White (1847, p. 88), in his List
of the specimens of Crustacea in the collection of the British Museum,

1 Published by permission of the Secretary of the Smithsonian Institution. Re-
ceived April 11, 1940.

Sept. 15, 1940 SHOEMAKER: NOTES ON GAMMARUS MINUS 309

uses the name Gammarus minimus Say and gives the locality as
United States; specimens presented by Thomas Say. Spence Bate
(1862, p. 221), in Catalogue of the specimens of amphipodous crustaceans
in the collection of the British Museum, repeats Say’s description and
habitat.

Sidney I. Smith (1874, p. 654) says, “‘I have not yet been able to
rediscover this species, which is very likely not a true Gammarus.”’
He believes that the specimens referred to by White and Bate were
the ones that Bate described as Allorchestes knickerbockeri (1862, p.
36). Lucien M. Underwood (1886, p. 357) mentions the species as
occurring doubtfully in New York. Thomas R. R. Stebbing (1906,
p. 513) puts G. minus in the list ‘“Gammari nominatim, reapse in-
certae sedis.’’ Henry W. Fowler (1912, p. 194) in The Crustacea of New
Jersey, lists Gammarus minus Say and questionably Lepleurus rivu-
laris Rafinesque as synonyms of Gammarus fasciatus Say. From the
localities he cites for G. fasciatus it is quite evident that he is dealing
with both G. fasciatus and G. minus.

As the references to Gammarus minus in literature add nothing to
Say’s inadequate diagnosis, I here redescribe and figure the species
from specimens collected by John W. Price from a small stream at
Gable’s Woods, Lancaster, Pa., on March 6, 1936. As Say did not
designate any locality, a male from this lot has been selected as the
neotype, U.S.N.M. no. 79152. I have selected this locality for the
neotype because Say collected G. fasciatus at Philadelphia, and it is
probable that he also collected G. minus in the brooks of the surround-
ing country, which would be only about 60 miles from Lancaster.

Gammarus minus is placed by Dr. A. Schellenberg (1937, p. 270) in
the subgenus Rivulogammarus, which he established for those almost
exclusively fresh-water species of the cold and temperate regions of
the northern half of the earth. He characterized this subgenus as
_having the side lobes of the head truncate or rounded off without the
sharp upper corner. Eyes mostly small, oval or kidney-shaped. First
and second urosome segments without or with only weakly developed
central humps and with only one pair of central spines. Third urosome
segment mostly without central spines. A part of the urosome spines
sometimes replaced by bristles. Accessory flagellum 1- to 6-jointed.
Lower margin of the coxal plates mostly unbristled. Metacarpus of
the first gnathopod of male with peglike spines on rear margin. Rami
of the third uropod varied in length, but the inner ramus never re-
duced to a scalelike joint.

Ree f ss

390 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

Gammarus (Rivulogammarus) minus Say Figs. 1, 2

Gammarus minus Say, Journ. Acad. Nat. Sci. Philadelphia 1(2): 376. 1818.
G. propinquus Hay, Proc. U. 8S. Nat. Mus. 25(1285): 224. 1902.
G. purpurascens Hay, Proc. U. 8. Nat. Mus. 25(1292): 433, fig. 7. 1902.

Male.—Head with lateral lobes squarely truncate, upper and lower angles
of lobes rounding. Eyes large, black, reniform, and slightly broader below.
Antenna 1 about two-thirds the length of the body and considerably longer
than antenna 2; first joint of peduncle a little longer than second, which is
about twice as long as third; flagellum of about 30 joints; accessory flagellum
of three or four long joints and a very minute terminal joint. Antenna 2,
gland cone of second joint prominent; third joint prominent; fourth and
fifth joints about equal in length; flagellum nearly as long as fourth and
fifth joints combined and consisting of about 14 joints, the first five or six
of which bear calceolli.

Right mandible with cutting edge rather narrow and armed with four
teeth; accessory plate with double cutting edge; five spines in spine row;
molar strong and bearing two short plumose setae on front margin and a
long seta on inner margin; second joint of palp slightly longer than the third.
Inner plate of maxilla 1 with about 16 plumose setae; outer plate with 11
spine teeth, some of which are blunt and some pectinate; palp of right
maxilla broad, armed distally with four or five stout teeth, one slender
pectinate spine tooth, and one slender plumose seta; palp of left maxilla
with about seven slender spine teeth and four slender setae. Maxilla 2
normal and bearing the usual oblique row of plumose setae on inner plate.
Maxillipeds with inner plate narrower, but nearly as long as outer plate,
bearing distally three spine teeth and numerous slender curved plumose
spines, inner margin bearing long, slender, plumose setae and one short
spine near the upper angle; outer plate armed with a row of spine teeth on
distal inner margin, a row of curved plumose setae on distal margin and
numerous submarginal setae near the inner margin; palp rather short and
stout, third joint with a low distal lobe, the inner edge of which extends
obliquely across the inner face of the joint bearing a row of serate spinules;
fourth joint rather slender and armed distally with a fine nail at the base of
which are several closely set setules. Lower lip with broad blunt side lobes.

Coxal plates 1 to 3 bearing a few spinules at the rounding anterior and
posterior corners. Coxal plate 4 bearing a few setules at anterior corner and
a row on the lower posterior margin: Sixth joint of gnathopod 1 with the
very oblique palm bearing a stout central spine and a stout defining spine,
hind margin and inner surface of joint bearing several short blunt spines.
Sixth joint of gnathopod 2 with front and hind margins about parallel, the
oblique palm bearing a stout central spine and two stout spines at the
broadly rounding defining angle, inner surface of joint bearing two small
submarginal spines at defining angle. Peraeopods 1 and 2 very similar in
structure, but 1 the longer and stouter and bearing longer and more numer-
ous bristles. Second joint of peraeopod 3 broad, with the lower hind margin
forming a prominent angle. Peraeopods 4 and 5 about equal in length,
second joint with distinct lower hind angle.

Metasome segments 2 and 3 with lower hind corners sharply produced,
lower margins bearing spinules and hind margins setae. Urosome segments
1 and 2 somewhat raised dorsally, and each bearing the two median dorsal
spines characteristic of the subgenus Rivulogammarus. Urosome segment 3
may have either one or two median dorsal spines, or it may be without any,

SEPT. 15, 1940 SHOEMAKER: NOTES ON GAMMARUS MINUS o9l

but there are always median setae. Urosome segments 1 to 3 may bear either
one or two lateral spines on either side. All urosome spines are accompanied
by setae or bristles.

The number of spines on uropods | and 2 varies with the size of the speci-
men, but the number is always small. Uropod 3 with inner ramus about two-
thirds the length of the outer, and both rami furnished on their margins
with fascicles of long setae. Outer ramus without plumose setae on the
second joint or apex of first joint; outer margin of first joint with a plumose
seta in each fascicle except those which contain spines; inner margin with a
plumose seta in each fascicle. Inner ramus without plumose setae on apex,
but outer and inner margins with a plumose seta in each fascicle.

The telson varies somewhat in different individuals; the general arrange-
ment being a single lateral spine accompanied by several setae, three apical
spines with long setae on each lobe, and a group of lateral setae near the
apex of each lobe. Occasionally there is an extra lateral spine either above
or below the regular lateral spine. Length of the largest males about 16 mm.

Female.—The gnathopods are smaller than in the male, and the fifth and
sixth joints of gnathopod 2 are much narrower and are about equal in length.
Antenna 2 bears no calceoli. Uropod 3 shorter than in the male.

Remarks.—There are in the National Museum females of Gammarus
minus from the District of Columbia carrying eggs each month from January
to August. There are hatched young in the marsupium in January, April,
May, June, and August. Free-swimming young are found in January, April,
July, and August collections. There are no specimens taken in the District
of Columbia from September to December, but specimens from Virginia and
Pennsylvania taken in November and December carry eggs. It appears,
therefore, from these records that the species breeds throughout the year.

Specimens of G. minus from a cave at Speed Creek, about 8 miles north-
east of Bedford, Lawrence County, Ind., and those from a cave on the
Mullindore property, Rohrersville, Washington County, Md., differ very
slightly from the normal surface water form.

In the National Museum there are collections of G. minus from Penn-
sylvania, Maryland, District of Columbia, Virginia, West Virginia, Indiana,
Tennessee, Kentucky, and Georgia.

I have examined the type material of Gammarus propinquus Hay from a
spring about 2 miles north of Mammoth Cave, Ky., and of G. purpurascens
Hay taken at the mouth of Nickajack Cave, Shellmound, Tenn., and I find
that they do not differ specifically from G. minus Say. Hay does not give
the measurements of either species, but the largest specimens of G. pro-
pinquus are about 10.5 mm from the front of the head to the tip of the third
uropods, and those of G. purpurascens are about 10 mm. The specimens
collected by Hay in spring at Rossville, Ga. (referred to in Proc. U.S. Nat.
Mus. 25: 434. 1902) and identified as G. purpurascens, are in the National
Museum. They are all of medium size, the largest about 9 mm and are un-
doubtedly G. minus Say.

The Lepleurus rivularis of Rafinesque (1820, p. 7) said to occur in the
brooks of the mountains of Pennsylvania and at Shannon Run and Bedford

392 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 9

Springs, Va., is probably the Gammarus minus of Say, but Rafinesque’s
description is so imperfect that nothing can be learned from it.

Gammarus minus var. tenuipes, new variety

Specimens from Organ Cave, Greenbrier County, W. Va., found in a
stream a quarter of a mile in from the entrance, appear to be a subterranean
variety of Gammarus minus. These specimens have the eye greatly reduced
and occasionally almost entirely lacking. The lateral lobes of the head have
the corners very rounding. The gnathopods and peraeopods are slender,
especially noticeable in the narrowing of the second joint of the last three
peraeopods. In some of the specimens there is a reduction in the number of
spines on the urosome. The inner ramus of the third uropod is proportion-
ately longer than in the normal form. The whole animal has a weaker and
more delicate appearance than the surface form. Length about 14 mm.

The type, a male, U.S.N.M. no. 79195, was taken in Organ Cave, Green-
brier County, W. Va., on October 21, 1939, by T. Kenneth Ellis.

The National Museum has also a specimen of this variety from McClung’s
Cave, West Virginia, and several from Higgenbotham’s Cave, Frankford,
Greenbrier County, W. Va., collected by J. M. Valentine.

LITERATURE CITED

Batsz, C. SpeENcE. Catalogue of the specimens of amphipodous Crustacea in the collection
of the British Museum. Pp. 1-399, pls. 1-58. London, 1862.

DeKay, J. E. Natural history of New York, or the New York fauna. Part 6: Crustacea.
Pp. 1-70, pls. 1-13. 1844.

Fowiter H. W. Crustacea of New Jersey. Ann. Rep. New Jersey State Mus. for
1911: 35-461, pls. 1-150. 1912.

Goutp, A. A. Report on the Invertebrata of Massachusetts. Crustacea. Pp. 11-23;
321-341. 1841.

Hay W. P. Observations on the crustacean fauna of the region about Mammoth Cave
Kentucky. Proc. U.S. Nat. Mus. 25 (1285): 223-236. 1902.

— Observations on the crustacean fauna of Nickajack Cave, Tennessee, and vicinity.
Proc. U.S. Nat. Mus. 25 (1292): 417-439 figs. 1-8. 1902.

MiILtNE-Epwarps H. Histoire naturelle des crustacés, 3: 1-638. 1840.

RAFINESQUE C. 8. Annals of nature or Annual synopsis of new genera and species of
animals, plants, etc., discovered in North America. First Annual Number, for
1820: 1-16. 1820.

Say, Tuomas. An account of the Crustacea of the United States. Journ. Acad. Nat.
Sci. Philadelphia 1: 374-457. 1818.

ScHELLENBERG, A. Schliissel und Diagnosen der dem Stisswasser-Gammarus naheste-
henden Einheiten ausschliesslich der Arten des Baikalsees und Australiens. Zool.
Anz. 117: 267-280. 1937.

Smitu, 8. I. The Crustacea of the fresh waters of the United States. Report of the
United States Commissioner of Fisheries for 1872 and 1873, pt. 2: 637—709, pls.
1-3. 1874. :

STEBBING, T. R. R. Amphipoda I Gammaridea. Das Tierreich, pp. i-xxxix; 1—806.
1906.

UnpvEerwoop, L. M. List of the described species of fresh water Crustacea from America,
north of Mexico, Article V. Bull. Illinois State Lab. Nat. Hist. 2: 323-386. 1886.

Wuirtsr, ApAMm. List of the specimens of Crustacea in the collection of the British Museum.
Pp. i-viii; 1-143. 1847.

Supt. 15, 1940 SHOEMAKER: NOTES ON GAMMARUS MINUS 393

Fig. 1.—Gammarus minus Say, male: a, Anterior end of animal; 6, right mandible;
c, maxilla 2; d, right maxilliped; e, inner and outer plates of maxilliped on larger
scale; f, lower lip; g, gnathopod 2; h, inner surface of metacarpus of gnathopod 2;
1, peraeopod 2; j, urosome; k, uropod 1; 1, uropod 2.

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394 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 9

EZ

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tit7-tj,
Sy

Fig. 2.—Gammarus minus Say, male: a, Right maxilla 1; b, palp of maxilla 1 on
larger scale; c, palp of another male showing five spine teeth; d, left palp showing
terminal spines; e, distal end of outer plate of maxilla 1 showing the 11 spine teeth;
f, peraeopod 1; g, peraeopod 3; h, peraeopod 4; 7, peraeopod 5; 7, metasome; k, uropod
3; 1, m, apex of outer and inner rami of uropod 3 on larger scale; n, telson; 0, telson of
another male. The figures of telson are on a slightly larger scale than the uropods.

Sept. 15, 1940 COKER AND MORGAN: A NEW COPEPOD 395

ZOOLOGY.—A new harpacticoid copepod from North Carolina.
R. E. Coker and Juanita Moraan, University of North Caro-
lina. (Communicated by Waxpo L. ScHMITT.)

Harpacticoid copepods collected by a graduate student, Paul Mc-
Kee, from Baker Lake (Bladen County), Swamp Pond (Craven
County), Odum’s Millpond (Cumberland County), and other waters
of eastern and southeastern North Carolina were found to resemble
in many respects both Canthocamptus staphylinoides Pearse (1905)
and the subspecies that the senior author has described as sinwus
(1934). There are, however, such notable points of difference from one
or both of these forms that it merits at least subspecifie designation.
We have named it:

Canthocamptus staphylinoides vagus, new subspecies

Holotype.-—Female from Swamp Pond, Craven County, N. C., U.S.N.M.
no. 79249. Collected by Paul McKee, January 16, 1938. Males have not
been found. (McKee and Coker, 1940.)

Diagnosis.—Second foot conforming to the type; vestigial seta of fifth
foot rather long—about one-sixth the length of outermost seta; relative
lengths of spines on that segment about as in C. staphylinoides; anal spines
small, 10 to 12 in number; many small spinules just lateral to the straight
mesial borders of furcal rami on ventral side. (Now known only from waters
of the lower Coastal Plain in North Carolina.)

Description.—Second foot: The endopod has only one seta on the inner
border of the third segment (Fig. 1, c) where sinuus has two (Fig. 1, 7);
apparently it is the distal seta that is lacking. Pearse did not describe or
figure this foot, but copepods we have from Massachusetts seem to agree in
every other particular with the description of C. staphylinoides and are found
to have only one seta in the place mentioned.

Fifth foot: Examples from Baker Lake have a distal joint of the fifth foot
virtually identical with that of C. stnuus, but the short seta between the
bases of the outermost setae of the proximal segment is distinctly longer
than the corresponding seta of sinuus or of staphylinoides from Massachu-
setts (compare Figs. 1, e, and 1, a, with Figs. 1, k, and 1, g). The third well-
developed seta, counting from the outer border of this segment, is distinctly
shorter than the fourth, whereas in C. sinuwus (Fig. 1, k) the two are ap-
proximately equal; in this respect the copepods in question are like C.
staphylinoides (Fig. 1, g). The ecpepods from Swamp Pond (Fig. 1, a) have
all spines, except perhaps the two outermost setae and the innermost seta
of the distal joint, decidedly shorter, blunter, and stronger. The short seta
of the proximal segment is intermediate between that of Baker Lake and
C. sinuus, but its length, relative to that of the longest spine, conforms
roughly with the corresponding relation in the Baker Lake copepods. This is
true also of the third well-developed spine.

Caudal furea: The form of the furea (Figs. 1, b, f) is quite unlike that
characteristic of C. sinuwus (Fig. 1, 2) but like that of C. staphylinoides (Figs.
1, /, h). On each branch of the furca there is a submarginal row of fine spinules
or setae on the ventral surface a little lateral to the inner margin (Figs.
1, b, f); these spinules are not well seen until the furca is turned at an angle,

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396 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 9

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ULL SSN

Fig. 1.—Canthocamptus staphylinoides vagus, holotype, female, from Swamp Pond:
a, Fifth foot; b, furea—ventral aspect, slightly turned. Canthocamptus staphylinoides
vagus from Baker Lake: c, Endopod of second foot, female; d, endopod of third foot,
female; e, fifth foot; f, furca. Canthocamptus staphylinoides (s. str.) from Massa-
chusetts: g, Fifth foot; h, furea—dorsal. Canthocamptus staphylinoides (after Pearse,
1905): 74, Furca. Canthocamptus sinwus (from Coker, 1934): 7, Second foot, female;
k, Fifth foot; U, furca.

Sept. 15, 1940 COKER AND MORGAN: A NEW COPEPOD 397

as shown in Fig. 1, 6. The furca of the Massachusetts copepod (Fig. 1, 1)
seems to be typical of C. staphylinoides as described and illustrated by
Pearse (Fig. 1, h), with a few rather strong setae showing on the inner mar-
gin or ventrally.

Anal plate: The spinules of the anal plate (Figs. 1, b, f) are relatively
small, there being 10 on the examples from Baker Lake, 12 on those from
Swamp Pond. The number in C. senuwus (Fig. 1, 7) and C. staphylinoides
(Fig. 1, h) is about 6, and the spinules are distinctly stouter.

The differences between the copepods from Baker Lake and those from
Swamp Pond are insignificant. In respect to the fifth foot, the differences
between these copepods on the one hand and either C. staphylinoides or
C. sinuus are also inconsiderable.

Remarks.—Comparison of the new copepod with the previously described
forms is given in the following table, where the mark ‘‘X”’ is used when the
character is common to two, and the mark ‘‘—” where a distinctive char-
acter is found.

Rudi-
Second S| Sebati ne Anal Furea | Arma-
Species Roos |S foot spines form ture
seta
Cesiapnylenoides.. 0... ... xX XS xX x xX —
CASvagusmN. SubDSp......,. xX = xX — xX —
Ls SUUUTS =e — x — x — —

1 Relative lengths of seta on first segment.

Since C. sinuus may now be presumed to differ from C. staphylinoides, not
only in the form and armature of the furea, but also, and significantly, in
the armature of the second foot, it should be regarded as a distinct species.
The furca is unmistakable at a glance.

For direct comparison with the foregoing diagnosis of C. s. vagus we pre-
sent here diagnoses of C. staphylinoides Pearse and C. sinwus Coker (here
raised to the status of a species; described by Coker, 1934), the only other
members of the restricted genus Canthocamptus, as defined by Chappuis and
followed by the senior author in Contribution to knowledge of North American
freshwater harpacticoid copepod Crustacea, 1934, now known to occur in the
United States.

C. staphylinoides Pearse: Characters of the genus (both rami of second to
fourth swimming feet 3-jointed, a vestigial seta on mesial expansion of fifth
foot, etc.); second foot with one seta on inner border of third segment of
endopod; short seta between bases of two outermost setae of proximal seg-
ment of P5 very short—about one-twelfth as long as the adjacent outermost
seta or shorter; third well-developed seta of that segment, counting from the
outermost border, shorter than fourth; anal spines large, 6 to 8 in number;
a few prominent spinules on the straight mesial side of each fureal rami.
(Now known only from Northern United States and Canada.)

C. sinuus Coker: Second foot with two setae on inner border of third seg-
ment of endopod; vestigial seta of P5 about as in C. staphylinozdes; third
well-developed seta on that segment about equal to fourth; anal spines 6 to
8 in number; mesial profile of furcal rami sznwous and unarmed. (Known
from Connecticut, New Jersey, and North Carolina.)

398 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

Our new copepod is nearer to C. staphylinoides than to C. sinuus in the
second foot, the fifth foot, and both the form and armature of the furca,
although it differs from the former in the rudimentary seta of the fifth foot,
the anal spines, and the armature of the furca. Since, however, the differ-
ences between our copepod and C. staphylinoides are quantitative and may
be presumed to be subject to considerable variation, I would propose only
a new subspecific name vagus, referring to its apparent habit of wandering
out into open water. Harpacticoid copepods are characteristically littoral in
the extreme and it is worthy of note that this copepod was taken repeatedly
in plankton hauls during a survey covering 45 waters. The only other har-
pacticoid encountered was C. sinwus in one collection from a shallow swamp
lake.

Some of the older lists of American harpacticoid copepods include C.
staphylinus Jurine, but it is very probable that the records of the European
species, which is clearly distinguishable by the conspicuous blunt, spinelike
projection on the posterolateral margins of the fifth abdominal segment, as
well as by other characters, are attributable to errors in identification.
We know of one such case.

LITERATURE CITED

CoxErR, R. E. Contribution to knowledge of North American freshwater harpacticoid
copepod Crustacea. Journ. Elisha Mitchell Sci. Soc. 50: 75-141, 15 pls. 1934.

McKes, Pav, and Coxrr, R. E. Notes on plankton Entomostraca of the Carolinas.
Journ. Elisha Mitchell Sci. Soc. 56: 177-187. 1940.

Prarseg, A.S. Contributions to the copepod fauna of Nebraska and other States. Trans.
Amer. Micr. Soc. 26: 145-160, pls. 13-17. 1905.

Fresh-water Copepoda of Massachusetts. Amer. Nat. 40: 241-251, 9 figs. 1906.

PROCEEDINGS OF THE ACADEMY AND
AFFILIATED SOCIETIES

PHILOSOPHICAL SOCIETY

1155TH MEETING

The 1155th meeting was held in the Cosmos Club Auditorium, Saturday,
October 14, 1939, President BrIcKWEDDE presiding.

Program: T. F. W. Bartu, Geophysical Laboratory: Geysers and other
hotsprings in Iceland.—The thermal activity in Iceland, which is but one
phase of the recent voleanicity, is not equaled in intensity or extent by any
other region. Pioneers of our modern science, men like Bunsen, Descloizeaux,
and others, visited Iceland to study these phenomena, and through their
work the local name of the most famous spouting fountain in Iceland at that
time, viz., ““Geysir’”’ (meaning ‘‘spouter’’) was accepted as a general term
for hotsprings of this type.

The acid springs are intimately associated with recent volcanic activity,
the alkaline springs are more independent in this respect, but are clearly
most plentiful in areas possessing a superior water supply. Both acid and
alkaline springs may show geyser action. Some of the geysers form deep wells,
but measurements show that, although the temperature increases with
depth, the boiling point is not reached at any depth—this is also true for
the Great Geysir, which is 27 m deep.

SEPT. 15, 1940 PROCEEDINGS: PHILOSOPHICAL SOCIETY 399

The biggest boiling stream in the world is a copious spring in western
Iceland the discharge of which is nearly 300 liters per second. The thermal
energy it carries with it is approximately 140,000 horsepower. (Author’s ab-
stract. )

A. Naval, Westinghouse Electric and Manufacturing Co.: Formation of
surfaces of slip under plastic states of stress —A characteristic phenomenon
accompanying the plastic deformations of natural rock materials or of the
ductile metals is the formation of very regular markings on the surface of
test specimens. They are the traces of deformations localized in thin layers
of the material which become plastic under the given system of forces. The
laws of the formation of these slip layers are not yet fully understood and are
rather intricate. Several important factors are required to be satisfied for
the localized yielding of materials, which were discussed. Formation of regu-
lar spirals around a punch mark was shown for mild steel. Similar lines seem
to mark the location of secondary cones on certain voleanoes in Japan. The
formation of faults in geology seems to follow similar laws than the one de-
scribing shp in metallic materials. A motion picture was shown on the forma-
tion of gliding lines in mild steel bars. (A uthor’s abstract.)

The first paper was discussed by Messrs. WRIGHT and GisH; the second
one by Messrs. HAWKESWORTH, GISH, ROLLER, MAXWELL, and Brick-
WEDDE.

1156TH MEETING

The 1156th meeting was held in the Cosmos Club Auditorium, Saturday,
October 28, 1939, President BrickKWEDDE presiding.

Program: K. H. Bris: Model testing at the National Hydraulic Laboratory.—
The National Hydraulic Laboratory is authorized to conduct general re-
search in hydraulics on problems of its own choice and to work on specific
engineering problems at the request of other Government organizations. For
obvious reasons ship-model work is excluded, and for practical reasons pump
and turbine testing is not feasible at the present time. The general study of
the theoretical bases underlying hydraulic model studies is an important
field of research that has not received sufficient attention. It appears likely
that the laboratory will enter this field in the near future. Model tests for
specific purposes have been conducted for and with the cooperation of
several Government departments. The purposes of such model tests are
(1) to check calculations of design, (2) to suggest improvements either to
assure better performance or to lower costs without material sacrifice in
performance, and (8) to discover and correct undesirable or even dangerous
conditions which could not have been foreseen in the design. Lantern slides
were shown illustrating representative model tests, and some of the problems
encountered in making these studies were discussed. These problems include
on the one hand, the construction, operation, and instrumentation of models,
and on the other hand, the difficulties and uncertainties arising from the
impossibility of obtaining strict similarity of model to prototype. (Author’s
abstract.)

H. E. Saunpers, U.S. Navy: The David W. Taylor Model Basin at Carde-
rock, Md.—The completion of the buildings for the new ship model testing
establishment of the United States Navy at Carderock, Md., just northwest
of Washington, has already received considerable publicity in the press.
These buildings have a function much different from that of the multitude
of new public buildings erected throughout the country in the last decade.
Pictorially they are of great interest to the public because of the simplicity

400 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

and dignity of their architecture and the use of precast exposed aggregate
concrete for the exterior finish. The large building housing the model basins,
with its arch roof, is a quarter of a mile long and is the only one of its type
in the United States.

The new establishment has been named the David W. Taylor Model
Basin in honor of Rear Admiral David W. Taylor, Construction Corps,
U. S. Navy (Retired), whose researches in ship propulsion and whose ex-
periments with ship models have made his name renowned throughout the
world.

The model testing plant at Carderock includes four separate model basins,
each of which is designed to undertake and prosecute to the best advantage
a particular kind of ship model research. These four basins will, in addition,
provide a plant of vastly increased capacity for undertaking a steadily in-
creasing volume of work. The old model basin at the Washington Navy
Yard, built in 1899, is now running sixteen hours a day in an endeavor to
keep pace with the requirements of the current naval and merchant ship
program. The ship-model basins at Washington are the only ones in America
equipped to undertake tests of ship models when they are driven by their
own propellers. They perform towing tests and self-propelled tests of this
kind for private shipbuilders and naval architects of the United States as
well as for the Navy Department and all other Government departments.

To enable the staff to make correct predictions of ship performances to
within a few percent, as modern designs require, the new basins at Carderock,
embody a number of distinctly novel features. To obtain the desired accu-
racy for model tests, the carriages that tow the ship models through the basins
must run at constant speed on tracks that are straight and level within a
few thousandths of an inch.

The walls of the basins upon which these tracks rest are of massive con-
crete construction, built directly upon bed rock. The track structure is of a
new design, using a rail heavier than that employed on any railroad in the
world and a rail supporting structure which is much more rigid than that in
any other model basin track. As it is expected that at least a year will be
required to lay the carriage tracks to the requisite degree of accuracy, it is
intended by this means to provide a permanent track structure which will
require no further attention for many years to come.

The work of laying the tracks alongside the new model basins is now being
undertaken by the staff of the David W. Taylor Model Basin and they hope
that by the summer of 1940 the work of ship model testing in the new plant
will have begun. (Author’s abstract.)

The second paper was discussed by Messrs. LirERocK and PAWLING.

1157TH MEETING

The 1157th meeting was held in the Cosmos Club Auditorium, Saturday,
November 11, 1939, President BRIcKWEDDE presiding.

Program: L. V. BERKNER: Fluctuations in the earth’s outer atmosphere dur-
ing a great magnetic storm.—The great magnetic disturbance that occurred
on April 16, 1938, was one of the largest in recent years. Records of iono-
spheric changes on the magnetic equator at Huancayo, Peru, are complete
during this disturbance. Actual distribution of ion-density at successive
quarter-hourly intervals can be represented by parabolic distributions which
would produce the observed records using a method recently described by
Booker and Seaton.

SEPT. 15, 1940 PROCEEDINGS: PHILOSOPHICAL SOCIETY 401

With advent of the storm, just after midnight, maximum electron density
of the F-region (above 250 km) fell in the first hour by three orders of mag-
nitude and essentially to zero, with the disturbance progressing downward
through the ionosphere. Then appeared an intensely ionized absorbing
region at 85 km (near where maximum conductivity per electron is experi-
enced) which is shown to have covered a substantial portion of the earth’s
surface. The disturbance then progressed rapidly upward with the height of
F-region apparently rising at about 8 km per second first with increasing and
then decreasing ion-density until there remained a maximum density of

only 5,000 electrons per cc in the neighborhood of 1,000 km. Recovery came -

with daylight.

A number of interesting problems are raised in view of the restrictive
influence of the earth’s magnetic field on motions of charged particles at the
magnetic equator. Possibly the ionosphere above Huancayo might be imag-
ined to explode toward the polar regions during the disturbance. Mechanism
of entrance of the disturbing influence from the sun which produced the
effect offers room for speculation. (A uthor’s abstract.)

Evuis A. JoHNSON: Scattering of light from a modulated searchlight beam
at high altitudes.—The measurement of the light scattered by the atmosphere
from an intense searchlight beam up to heights of 34 km was described. The
method makes it possible to determine the temperature and pressure with
height above the earth’s surface and to make determinations of some of the
atmospheric constituents. The experimental procedure consists in directing
an intense searchlight beam nearly vertical and modulating it at 10 cycles
per second with Venetian blind shutters. A receiver, consisting of a large
mirror with a photoelectric cell at its focus, is placed about 6 km away. The
beam can be scanned by tilting the receiver and the height at any particular
angle calculated from the geometry. The sensitivity is limited by the back-
ground statistical fluctuation of shot-effect due to photocell current caused
by light from the night sky. Measurements above a low haze bank agreed
with the atmosphere given by Humphreys up to 34 km. With 60-inch mirrors
at the receiver and transmitter, it is calculated that heights of 80 km could
be reached, and problems such as the distribution of ozone, water vapor,
turbulence, winds, dust, fluorescence, etc., could be studied. (Author’s ab-
stract. )

The first paper was discussed by Messrs. SkEGER, GOLDBERG, BITTENGER,
Mouuerr, BRickKwEppDs, 8. 8. Huruny, Hows, MeNisu, and SMITH ; the
second paper by Mr. BRICKWEDDE.

J. PAWLING presented an informal communication, which was a report of
progress on The new general catalog of Albany.

1158TH MEETING

The 1158th meeting was held in the Cosmos Club Auditorium, Saturday,
November 25, 1939, President BRicKWEDDE presiding.

Program: J. C. Hupsparp, Johns Hopkins University: Ultrasonics and the
ratio of specific heats of gases —Among the indirect methods of studying the
specific heats of gases, those involving measurements of sound velocity have
been considered most ideal. However, until the advent of ultrasonics, values
obtained by acoustic methods have been disappointing, in spite of a vast
amount of empirical and theoretical work on corrections entailed by the
confinement of sound waves within a chamber necessarily of dimensions of
the same order of magnitude as the wavelength. Ultrasonics provides wave
lengths measured in millimeters or less, and experiments with them are com-

——— i.

402 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

parable to experiments with ordinary sound waves in chambers so large as

to eliminate the boundary corrections. The way is thus opened to a study of
various gases; moreover these need be used only in small amounts. The re-
sults are of a highly satisfactory precision and yield satisfactory values of
specific heats except insofar as they are influenced by the dynamic character
of sound waves. In many cases it is found that there is a lag in the adjustment
of certain energy states of the molecule. This effect was responsible for ultra-
sonic dispersion found in COs, in 1925 by G. W. Pierce and accounted for
theoretically in 1928 by Herzfeld and Rice. Ultrasonic methods have now
been developed so that precise acoustic data on specific heats may be secured
over a very great range of temperatures and pressures. The character of the
results permits the detection of those regions of pressure or frequency in
which energy lags are present, and are thus useful in determining lifetimes
of energy states. Examples were shown. (Author’s abstract.)

Francis E. Fox, Catholic University of America: Supersonics in liquids.—
For the production of ultrasonic waves in liquids the source is usually a
magnetostrictive rod or piezo-electric quartz plate with its vibration elec-
trically maintained and coupled by direct contact with the liquid. The in-
vestigation of the waves, and through them of acoustic properties of the
liquid, may be carried out by electrical, mechanical, or optical effects pro-
duced by the waves.

(1) The reaction upon the driving electrical circuits produced by waves.

reflected back to the source, as in the acoustic interferometer, may be ex-
pressed as functions of the reflector distance and the half-wavelength in the
medium, the absorption factor, and the coefficient of reflection. (2) The
steady radiation pressure exerted cn an obstacle in the sound field offers a
direct means of investigating the intensity. (8) The variations in the index
of optical refraction that are produced by the sound wave have been the
basis of many optical methods. (a) in the Sears-Debye method one uses the
sound beam as an optical grating. Progressive or stationary waves may be
used. (b) With Schlieren metheds, of which there are many variations one
renders the stationary sound fields visible with constant illumination, or
employs stroboscopic light to investigate progressive waves. (c) The dif-
fraction-interference methods enable one to ‘see’ the sound field in the liquid.

The choice of a method depends of course on the problem to be investi-

gated. Methods that permit accurate determination of wavelength may yield

little information about intensity or absorption. It may be possible to extend
the range of frequency in which some one method is applicable by combining
that method with a different one, e.g., the acoustic interferometer with an
optical indicator or resonance for very short wavelengths.

The demonstration included the acoustic interferometer, radiation pres-
sure, and several optical effects. (A uthor’s abstract.)

The first paper was discussed by Messrs. H. L. Curtis, Herzrevp, and
BRICKWEDDE; the second one by Messrs. BRICKWEDDE, LEwIs, and others.

1159TH MEETING

The 1159th meeting, constituting the 69th annual meeting, was held in
the Cosmos Club Auditorium, Saturday, December 9, 1939, President
BRICKWEDDE presiding.

The treasurer reported that the income from dues, interest on investments,
and sale of reprints was $1,392.94 and that the expenditures exclusive of
investments was $1,342.46, leaving a net surplus of $50.48 on ordinary ex-
penses. The ordinary expenditures were at the rate of $4.18 per member.

a!
el

Sept. 15, 1940 PROCEEDINGS: PHILOSOPHICAL SOCIETY 403

The secretaries’ joint report showed an active membership as of December
1, 1939, of 321, of whom the following were elected during the year: JoHN
Berek, JR., Pau Brapt, Brick A. Brooks, L. O. CoLBERT, RicHarp M.
Fraps, Cuas. K. GREEN, RaupH F. Haupt, ALBERT J. HOSKINSON, HENRI
JORDAN, J. W. Joycr, THoBuRN C. Lyon, Harotp W. Murray, F. W.
REICHELDERFER, C.—G. Rosssy, W. H. Seaquist, Douauas R. Tats, W. A.
WILDHACK, and WALTER J. Youna. The following were elected in 1938 and
qualified in 1939: L. Lron SHERESHEFSKY and H. M. O’Bryan.

The following death was reported: JAMES H. Gore.

The following officers were declared elected for the year 1940:

President, R. E. Gipson; Vice-Presidents, H. E. McComs and W. G. Brom-
BACHER; Corresponding Secretary, H. F. Stimson; Treasurer, W. E. DEMING;
Members-at-Large of the General Committee, L. V. BERKNER and R. W.
CURTIS.

Program: G. GaMow: Uncertainties in atomic and common life.—In classical
physics, the notions of the trajectory, position, and velocity of a moving
object were always considered as given a priorz and self-evident. However,
the discovery of the quantum-phenomena leads to a conclusion that these
seemingly fundamental notions represent actually only very good approxima-
tions to quite different and rather involved basic principles of a correct sys-
tem of mechanics.

Analyzing the method by which an idea of trajectory, for example, could be
constructed, we find that this cannot be done if the physical action can be
delivered only in the form of certain finite portions or quanta. This criticism
leads to fundamental uncertainty in the description of physical phenomena,
expressed in the so-called Heisenberg’s uncertainty relations between the
coordinates and mechanical momenta of moving particles. It can be shown
that the product of the uncertainties of these two quantities is equal to the
basic quantum constant, so that if the position is known very accurately,
the momentum is not, and vice versa.

Analogous relation exists between the energy and time, which, according to
the theory of relativity, represents the fourth pair of data describing the
event in the four-dimensional time-space. The possible methods for simul-
taneous measurement of energy and time have been discussed by N. Bohr
who has shown that such measurements necessitate taking into account the
effects of general relativity. Making use of Einstein’s “red-shift-formula,”’
Bohr proved that the uncertainty relation between energy and oe must
hold in any such measurement.

It is usually believed that owing to extremely small value of the quanti:
constant, uncertainty phenomena can be of importance only in the atomic
world. It is, however, not so. If, for example, we fix a steel ball 1 inch in
diameter rigidly on the table and drop another ball of the same size from the
point several feet above the fixed ball, classical mechanics will tell us that
our ball will jump up and down until the energy will be dissipated by fric-
tion. According to quantum-mechanics, however, the existing uncertainty
of the position and momentum of the ball at the moment it is dropped, will
cause a quite appreciable deviation and will make the ball drop aside after
only a few jumps. Unfortunately no technically produced steel is homogene-
ous enough to permit such experimentation.

We can imagine, however, some crazy worlds in which the quantum-
constant is so large that the uncertainty relations could be observed micro-
scopically. One fellow, called Mr. Tompkins, used to dream about such worlds
and had a lot of very unusual experiences in them. Not only was he unable

404 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

to play a good billiard game with ‘‘spreading-out”’ quantum balls, but also
his car ran out of the garage leaving through the wall just as an @ particle
from the atomic nucleus. Going to hunt tigers, he was quite scared to see a
tremendous number of tigers jumping on his elephant simultaneously from
all sides. He explained, however, that there was only one tiger in the first
quantum state around the elephant, and the tiger was soon shot by a quan-
tum bullet. Mr. Tompkins awoke after that event, but forever after remem-
bered what the uncertainty-relations actually are. (Author’s abstract.)
This paper was discussed by Messrs. O’ BRYAN and BRICKWEDDE.

1160TH MEETING

The 1160th meeting was held in the Cosmos Club Auditorium, Saturday,
January 6, 1940, President GIBSON presiding.

The Retiring President, F. G. BRickwEppDE, gave an address entitled
Some complexities of the simple element hydrogen. It is expected that this
address will be published in this JouRNAL.

1161ST MEETING

The 1161st meeting was held in the Cosmos Club Auditorium, Saturday,
January 20, 1940, President Gipson presiding.

Program: F. 8. Brackett: Spectroscopic methods and their importance in
biological research.—When radiation produces a measurable change, be it
physical, chemical, or biological, the plot of wavelength dependence of the
measured change may be termed an ‘“‘action spectrum.” In biophysics action
spectra are fully as important as emission and absorption spectra. Familiar
examples: In physics, the wavelength dependence of photocell sensitivity;
in chemistry, the photochemical yield as a function of wavelength; and in
biology, the visibility curve of the eye.

The photochemical problem is simpler than the biophysical, since the
absorption spectrum of the absorbing molecules may be measured and the
concentrations of the components of the system varied in order to determine
the kinetics of the system. Here the purpose of the action spectrum is to
establish the photochemical threshold and the quantum efficiency. In bio-
physics, even the absorbing molecule may be unknown. The purpose of the
action spectrum may therefore be to determine the effective absorption
spectrum and the efficiency or number of quanta required to produce the
effect. A common form of biophysical action spectrum is the wavelength plot
of the reciprocal of the incident energy required to produce an equal bio-
logical change. Typical example—the lethal action of radiation on micro-
organisms. Assuming Beer’s law a general expression 1s deduced which leads
to simple interpretation in two important cases. One, where the population
of microorganisms in suspension is small, that is, less than ten percent of
the beam absorbed. Here it is shown that the action spectrum yields the
wavelength dependence of the product of efeiency and absorption coefficient
per organism. The second case is where the number of organisms in suspen-
sion is large, so that practically all energy is absorbed. Here the action spec-
trum yields directly the efficiency or the reciprocal of the number of quanta
required to kill the organism.

The requirements on spectrographic equipment for obtaining action spec-
tra are fundamentally different from those to obtain emission or absorption
spectra. In general an output of great radiant power is necessary in order to
produce a measurable change in a reasonable length of time, but high in-
tensity is not necessary. High purity, that is, freedom from scattered radia-

Spr. 15, 1940 PROCEEDINGS: PHILOSOPHICAL SOCIETY 405

tion is required though high resolving power is not. This leads to instruments
with a large linear aperture but not necessarily large relative aperture;
high dispersion but low resolving power. For this purpose, large quartz prisms
are ideal in the ultraviolet, the number of prisms being as great as is com-
patible with maintenance of high transmission. Gratings are preferable in
the visible. Slit widths may be wide, but a double monochromator to elimi-
nate scattered radiation is often necessary. (Author’s abstract.)

P. A. Couz, National Institute of Health: Determination of cell structure by
means of the ultraviolet microscope.—A brief summary of the historical develop-
ment of the ultraviolet microscope was given. An eventual goal of all previous
workers has seemed to be to attain ultraviolet absorption spectra of micro-
scopic objects. The difficulties encountered in attaining this goal were dis-
cussed. Preliminary photographs were shown taken at various ultraviolet
wavelengths and a method was described by which the absorption spectrum
of a portion of a microscopic organism could be determined. (Author’s ab-
stract. )

The first paper was discussed by Mr. O’Bryan; the second one by Messrs.
Gisson, Brackett, Briacs, TELLER, Humpureys, H. L. Curtis, F. Con,
and others.

1162D MEETING

The 1162d meeting was held in the Cosmos Club Auditorium, Saturday,
February 3, 1940, President G1iBson presiding.

Program: P. A. Smitu: Exploring the continental shelves and slopes—Know]l-
edge of the topography of the sea floor has grown rapidly during the past
decade, largely through the development of echo-sounding. Owing to the
accompanying need for more complete nautical charts, several of the mari-
time nations have extended hydrographic surveys seaward from their shores;
and due to this seaward extension of the surveys a whole new physiographic
realm is being opened to students of earth science. The perfection and use of
offshore position finding methods and echo sounding by the U.S. Coast and
Geodetic Survey has added many thousands of square miles of surveys of the
continental shelves and slopes of United States and waters under their juris-
diction. The discovery of the stream-erosion type of topography on the sea-
ward faces of the continental shelves under hundreds of fathoms of water
has raised questions that seriously affect a number of cherished geological
theories. While topographic characteristics of numerous submarine canyons
are clearly and unmistakably those commonly observed in valleys. eroded
by running water above the sea, many geologists have felt that the simple
explanation of a lowered sea or rising land in the order of 10,000 feet and
back in late Tertiary or Pleistocene time is untenable in the face of appar-
ently well-established theories. A number of hypotheses have been advanced
in the attempt to explain these features by some process that may harmonize
with other geologic evidence, but the origin of submarine canyons 1s still
controversial.

Through a number of contributors, including Dr. H. S. Stetson of the
Woods Hole Oceanographic Institution, Dr. Maurice Ewing of Lehigh Uni-
versity, and collaborators, and Dr. C. 8. Piggot of the Geophysical Labora-
tory, Carnegie Institution, our knowledge of the materials and underlying
strata of the sea floor is being expanded. In all this work the Geological
Society of America has been especially interested, and has contributed
greatly to the growth of knowledge in this field of science through grants to
various investigators.

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406 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

Automatically recorded profiles of submarine topography of the Atlantic
Continental Slope show decidedly V-shaped cross sections, in the extreme
heads of the canyons as well as in depths of 6,000 feet and over. Due to cer-
tain limitations of echo sounding it is sometimes impossible to record the
bottom of steep, narrow ravines in deep water, or to record true slopes in
very rugged topography. Graphic records of echo sounding always give a
more complete picture of the profile than visual indicators and in many in-
stances reveal phenomena that might otherwise be missed, such as the
great sand.waves found south of Nantucket Island during the summer of
1939 by the Coast and Geodetic Survey Ship Oceanographer.

The abrupt change from the topography of the shelf which is plainly ma-
rine made in character, to that of the slope which shows no indications of
marine erosion, but is plainly that commonly attributed to stream erosion
is one of the most remarkable of the many interesting facts demonstrated
by the new surveys. The cooperation of the Geological Society of America
through the late Dr. A. C. Veatch and the Coast and Geodetic Survey has
resulted in the publication of these new surveys in the form of special charts
for use by various investigators. (Author’s abstract.)

T. C. Lyon, U. 8. Coast and Geodetic Survey: Methods of air navigation
and aeronautical charts.—Air transportation is still making rapid strides, and
the distribution of aeronautical charts by the Coast and Geodetic Survey
during the past year exceeded the distribution of nautical charts.

The basic methods of marine navigation are also the basic methods of air
navigation, although new instruments and stream-lined technique are re-
quired for the higher speeds of aircraft.

Aeronautical charts are comparable to nautical charts. They differ from
topographic maps in their emphasis upon landmarks, in the degree of accu-
racy required, and in the necessity for frequent revisions.

Three standard series of aeronautical charts are now being published by
the Survey: 87 sectional charts, scale 1:500,000; 17 regional charts at
1:1,000,000; and 6 direction finding charts at 1:2,000,000—all on the Lam-
bert conformal conic projection. These are designed to meet the needs of air-
craft of widely different speeds and methods of navigation.

Aeranautical charts are compiled from the best available data, and checked
from the air before distribution. For the future, air navigation will be at new
altitudes, with new methods, new instruments, and possibly new charts, in
prospect.

Specimens of the standard charts, and also of several experimental charts,
were on display. (Author’s abstract.)

The first paper was discussed by Messrs. Eaton, Paw ine, and Hum-
PHREYS.

1163D MEETING

The 1163d meeting was held in the Cosmos Club Auditorium, Saturday,
February 17, 1940, President GiBson presiding.

Program: H. R. Moraan, U. 8. Naval Observatory: The motions of the
earth.—Observations of the sun, moon, and planets in the past 300 years
show rather large and irregular deviations from their gravitational orbits.
These are attributed to fluctuations in the rate of rotation of the earth giving
a variable time amounting to as much as 60 seconds of time in a century as
compared with a perfectly uniform time.

To determine the motions of the planets over long periods of time it is
necessary to have a uniform time unit. The new tables of the moon are so

SEPT. 15, 1940 PROCEEDINGS: PHILOSOPHICAL SOCIETY 407

accurate they may be used for this. In an investigation of the motion of the
planet Mercury, now being carried on at the Naval Observatory, the ob-
served deviations of the moon from its gravitational orbit are used as the
corrections to variable earth time to reduce it to uniform or Newtonian
time. (Author’s abstract.)

C. B. Warts: A traveling-wire micrometer with photographic registration.—
The 6-inch transit circle of the U. 8S. Naval Observatory has been equipped
with a photographic register for use with the traveling-wire micrometer. The
usual method of recording on a chronograph the signals originated by a con-
tact device on the micrometer has been discarded. It is replaced by a system
in which a portion of the graduated head of the screw is photographed
periodically on recording paper while a transit is being observed. The ex-
posures are initiated by the clock and no chronograph is required. The result
is equivalent to that yielded by the older method; there are, however, several
advantages in the new system. One of these is the fact that the position of
the traveling wire is recorded at the same time intervals for all declinations.
These intervals are sufficiently great to insure the independence of succes-
‘sive recordings; closely spaced signals for an equatorial object and exces-
sively long intervals for a circumpolar star are both avoided. The new de-
vice also serves as an efficient recorder of pointings made on the collimators,
meridian marks, and reflected nadir images. When used for this purpose the
exposures are initiated by the observer. (Author’s abstract.)

The first paper was discussed by Messrs. Mracurrs and Warts; the sec-
ond by H. L. Curtis.

F. G. BRicKWEDDE gave an informal communication on a new tempera-
ture scale that has been proposed. He showed by a quotation from a paper
by J. P. Joule and William Thomson entitled On the thermal effect of fluids
in motion, which appeared in Phil. Trans. 144: 32 (1854), that the idea was
original with them. This was discussed by Messrs. GIBson and TUCKERMAN.

1164TH MEETING

The 1164th meeting was held in the Cosmos Club Auditorium, Saturday,
March 2, 1940, President GiBson presiding.

Program: A. H. Prunp, Johns Hopkins University: A partial analysis of
gaseous mixtures without the use of a spectroscope.—This procedure is based
on the fact that certain gases and vapors such as COz, H2O, CHa, ete. have
absorption bands in characteristic wave-length regions in the infrared spec-
trum. The apparatus consists of an incandescent solid whose radiations pass
through an initially evacuated absorption-cell and then enter a detector con-
taining a gas such as CO». Some of this radiation will be absorbed and trans-
formed into heat, thus raising the temperature of the gas and affecting ther-
mo-junctions not in the direct path of the radiation. Upon admitting air
containing CO, into the absorption-cell, this COs prematurely removes or
weakens those radiations which, previously, had heated the COs, in the de-
tector. As a result, the temperature of the gas in the detector drops.

Applications of this procedure to ventilation, fermentation, and respiration
were discussed. (A uthor’s abstract.)

A. G. McNisu: The geomagnetic field and its variations.—The general mag-
netic field of the earth may be represented to within the reliability of the
observations by a dipole near the center (moment 8 X10” egs units), giving
rise to about 80 percent of the field, and 14 secondary radially directed
dipoles (average moment 0.1 X10? egs units) located at specified positions

408 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

midway between the surface of the earth and its center, giving rise to the
remainder or residual field.

Secular change can be represented by the yearly addition of 13 dipoles
of equal strength (moment 1.410” cgs units) at the same depth as the
dipoles of the residual field. Continuance of secular change at the present
rate for one hundred years would thus build up a new residual field.

Interpretation of this model leads to the beliefs that (1) at least a consid-
erable portion of the earth’s magnetism (the residual field) originates at a
lesser depth than the central core revealed by seismological evidence, and
(2) secular change involves this residual field and therefore is due to changes
taking place between the surface of the earth and the surface of the central
core. (Author’s abstract.)

The first paper was discussed by Messrs. Guise MAXWELL, Brick-
WEDDE, Karrer, MuniuErR, McNisuH, HumMpuHrReys, and Astin; the second
one by Messrs. GIBSON, SEEGER, TELLER, McComp, SmirH, BRICKWEDDE,
and Howe.

1165TH MEETING

The 1165th meeting was held at the Cosmos Club Auditorium, Saturday,
March 16, 1940, President Gi1BSoN presiding.

The Tenth Joseph Henry Lecture entitled Differences in physical proper-
ties of isotopes was delivered by Prof. Haroutp C. Urny, of Columbia Uni-
versity. This lecture has been published in this JouRNaL 30: 277-294,
1940.

1166TH MEETING

The 1166th meeting was held in the Cosmos Club Auditorium, Saturday,
March 30, 1940, President GiBson presiding.

Program: R. M. Zane, Hygrade Sylvania Corporation: Fundamentals
of fluorescence.—Since Edison’s invention of the incandescent lamp in 1879,
incandescent lamp efficiencies have increased from slightly over 1 lumen per
watt to 14 lumens per watt for a standard 60 watt tungsten lamp. The in-
troduction of the fluorescent lamp has increased this efficiency to approxi-
mately 42 lumens per watt (including lamp wattage and auxiliary loss).

The visible light in a fluorescent lamp is generated by the passage of a
discharge through a mixture of argon gas and mercury vapor, the discharge
being operated under such conditions that as high a percentage as possible
of the radiation generated is in the resonance line of mercury (2537 A). This
ultraviolet radiation is absorbed by the fluorescent material on the inside of
the lamp and re-radiated as visible light of a wavelength depending upon
the type of fluorescent material used.

The effect of small quantities of foreign materials upon the fluorescent and
phosphorescent characteristics of a silicate or tungstate, such as is most com-
monly used in fluorescent lamps, is very marked. For example, the introduc-
tion of 1 percent manganese in zine silicate greatly increases the fluorescent
output, and moves the maximum of the output curve from approximately
4,150 A to 5,230 A. On the other hand, the introduction of iron, copper, or
nickel greatly decreases the fluorescent output. This effect is presumably
due to the introduction of additional energy levels in the fluorescent material
due to the replacement of the normal metallic constituents of the crystal by
the foreign metallic elements. In some cases these additional energy levels
are properly placed to radiate light in the visible region. In other cases, they
may be so placed as to re-radiate the light outside the visible, generally in
the infrared. This, of course, wastes the energy.

Sept. 15, 1940 PROCEEDINGS: PHILOSOPHICAL SOCIETY 409

Phosphorescence differs from fluorescence in that a time lag exists before
the hght output of the phosphor builds up to a maximum and also a lag exists
in the decline of the hight output to zero after the starting or stopping of
the exciting radiation. Many fluorescent materials exhibit both fluorescence
and phosphorescence. Phosphorescence in the type of materials under con-
sideration is ascribed to the presence of potential traps which hold appreci-
able quantities of electrons which escape slowly through the barrier of the
trap before returning to their normal place in the crystal structure and
emitting the radiation normally coincident with this energy interchange.

Various fluorescent materials are mixed to obtain the color desired in a
finished fluorescent lamp.

The quantum efficiency of the best fluorescent material found to date is
approximately 70 percent. (Author’s abstract.)

W. P. Lowe tu, Jr., Hygrade Sylvania Corporation: Practical application
of fluorescent lighting —To use fluorescence in practical lighting it is neces-
sary to combine a source of ultraviolet and the proper phosphor in a con-
venient form, and to devise means of operating this lamp on the 115-volt
60-cycle circuits generally available in this country. The lamps which have
been developed are long tubes having electrodes at both ends, powdered
phosphor on the inner surface of the tube, and a filling of mercury vapor and
rare gas at low pressure.

The electrical control circuit consists of inductive ballast in series with
the lamp, to control the current, and an automatic switch to aid in starting
the lamp. This switch provides for preheating of the cathodes, then applies
sufficient voltage across the tube to insure starting the arc. Several types of
starting switches were demonstrated, including Manual, Magnetic, Thermal,
“Glow Tube,” and High Resistance Thermal. |

The power factor of this lamp and inductive ballast being inherently low
(and lagging) it is our practice to ballast alternate lamps with a combination
of capacitance and inductance, thus operating one lamp with leading cur-
rent, and the resultant power factor of each pair of lamps very near to unity.

Practical lighting units, each one complete with lamps, ballasts, starters
and all necessary reflectors and glass diffusers were demonstrated, including
the illumination of one-half the lecture hall with two of the “Miralume”
units. Lantern slides were shown of many actual installations of fluorescent
lighting.

In conclusion, the effect of Daylight Fluorescent color was demonstrated
on flowers and many other objects. The ordinary tungsten filament in-
candescent lamp is, of course, far more yellow and red than natural daylight,
while the Daylight Fluorescent Lamp is very close to actual average day-
light (color temperature 6,500° K.). Blue flowers and blue cloth are particu-
larly striking when their true color under Daylight Fluorescent is compared
to identical samples under an incandescent source. (Author’s abstract.)

1167TH MEETING

The 1167th meeting was held in the Cosmos Club Auditorium, Saturday,
April 13, 1940, Vice-President McComps presiding.

Program: L. Marton, R. C. A. Manufacturing Co.: Electron microscopy.—
The discovery of the optical behavior of electron beams led to the develop-
ment of a new branch of science: ‘‘Electron microscopy.” It is based on the
analogous use of light optical elements in a compound light microscope and
electron optical elements in an electron microscope; and on the highly in-
creased resolving power due to the much shorter wavelength of electrons.

410 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 9

On considerations of wavelength alone, an enormous increase in resolving
power should be obtainable. However, this is limited by a number of factors
given, chief among them being the optical aberrations, which were discussed
in detail and numerically evaluated. It was concluded that at the present
state of development of electron microscopy, resolving pewers of the order
-of 10 A should be obtainable.

After a discussion of constructional details of electron microscopes, a se-
ries of micrographs taken by the electron microscope was shown, and the re-
sults discussed. It was pointed out that the main applications of the electron
microscope at the present time are in bacteriology, virus research, and in
the wide field of colloidal research, including not only industrial colloids,
but also all the building stones of living matter. (A uthor’s abstract.)

The paper was discussed by Messrs. Smita, HuMPHREYS, ROLLER, Cox,
Mauan, READING, BEEK, Farno, and others.

1168TH MEETING

The 1168th meeting was held in the Cosmos Club Auditorium, Saturday,
May 11, 1940, President GiBson presiding.

Program: J. H. Swartz, U. 8. Geological Survey: Resistivity studies of
some geological problems.—In El Paso, Texas, salt water was found to be
encroaching upon a well from which a considerable portion of the city’s
water supply was at that time being drawn. Resistivity measurements were
successful in determining the area underlain by salt water and in delineating
the boundary between fresh and saline water areas. Salt water predicted
for a city well at Clint, Texas, was encountered as predicted when the well
was drilled despite the adverse prediction.

In the Hawaiian Islands large amounts of water are developed from thin
basal lenses of fresh water floating on the salt water permeating the various
islands. It was found possible by resistivity measurements to tell the thick-
ness of such basal fresh water lenses and to determine quite accurately the
elevation of the basal water table. Five test checks so far obtained at shafts,
drill holes, and wells, have shown an average difference of 0.56 feet between
predicted and observed water table elevations.

Studies of perched water horizons are now being made on the island of
Maui. While still in an early stage due to the research character of the prob-
lem and the stratigraphic complexity involved, encouraging results have
been obtained. (A uthor’s abstract.)

F. W. Lez, U.S. Geological Survey: Some problems in geophysics.—As the
science of geophysics has grown, especially in its application to prospecting,
there has been a marked evolution in design and use of geophysical appara-
tus. During this evolution many traditional features of instruments have
been improved whereas others have become obsolete. In the light of this
progress it is very interesting to examine the many stages of development
that scientific prospecting has undergone and is undergoing. Old dousing
paraphernalia have been replaced by the magnetic dip needle, which in turn
has been largely replaced by the more accurate and much more sensitive
magnetic variometer. Magnetic interpretations have made a decided ad-
vance by the introduction of magnetic models and magnetic scales. Marked
advances have also been made (by the sister methods) in electrical, seismic,
and gravitational fields, and scientific prospecting is entering into a new
era in which its application will, doubtless, be greatly broadened. In all of
these branches of geophysical prospecting there are three distinct divisions
which must be closely intercoordinated for securing the optimum results.

SEPT. 15, 1940 PROCEEDINGS: PHILOSOPHICAL SOCIETY 411

First, there is the instrumentation or the design and construction of precision
field instruments. Experience in the oil industry shows how much advance
can be made in instrument technique if the scientific results are coupled
with the prospects of high economic returns. Second, there is the broad
research associated with the many types of geologic bodies which commonly
differ from territory to territory and also have many modifying controls and
factors. Third, there is the theoretical principle involving the application of
the magnetic, electric, gravitational, and seismic deformation fields to geo-
logic and engineering problems. The analyses of physical field measurements
in these fields often involve very complicated procedure. (Author’s abstract.)

1169TH MEETING

The 1169th meeting was held in the Cosmos Club Auditorium, Saturday,
May 25, 1940, Vice-President BROMBACHER presiding.

Program: Joun P. Hacen, Naval Research Laboratory: Beam power tubes
as ultra-high radio frequency generators.—The increasing use of the lower
radio frequencies by the existing commercial services has forced new services
into the ultra-high frequency range. There are many features in the behavior
of ultra-high frequency circuits and waves which invite their use even
though a condition of saturation had not been reached in the lower frequen-
cles.

The advent of tubes of the beam type, such as the Inductive Output Tube,
the Klystron and the Velocity Modulated Tube, has made the use of the
ultra-high frequencies practicable. The essential features in the action of
these tubes are: the production of a cylindrical beam of electrons; the ac-
celeration of the beam by means of a high voltage anode which collects little
current; the amplitude modulation of the beam by conventional grid action
as in the Inductive Output Tube, or the velocity modulation of the beam
as in the latter two types; the subsequent induction of the power from the
beam to a plate tank circuit which collects little or no direct current. This
tank circuit is usually some form of resonant transmission line; and the
eventual collection of the beam, which has given up its radio frequency
power, by means of a collecting anode at lower direct current potential.
(Author’s abstract.)

Oscar NorGorDEN: Propagation characteristics of ultra-high radio frequen-
cves.—One of the important links in a radio communication system is the
propagation of the radio waves between the transmitting and receiving an-
tennas. The electric field strength is proportional to the square root of the
power and must exceed some definite value if to be useful for communication
purposes.

At the ultra-high frequencies, frequencies above 100 megacycles, the trans-
mitting and receiving antennas are elevated several wavelengths above
the surface of the earth. For distances less than the optical range or line
of sight the field strength # at the receiving antenna is

EH=EH,+E£,

where #; and £» are the electric fields at the receiving antenna due to the
direct and reflected rays respectively. Thus an interference pattern should
be observed for the variation of the field strength with distance. Measure-
ments show that this interference pattern does exist and is in agreement with
the theory. For distances greater than the optical range the field strength
decreases exponentially with the distance. Thus the maximum useful com-
munication distance for the ultra-high frequencies is nearly independent of

412 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 9

the power and is approximately equal to the optical range. (Author’s ab-
stract.)

The first paper was discussed by Messrs. RamBerc, SmiTtH, MAxwELL,
and Buarr; the second by Messrs. MaxXwELL, BROMBACHER, BRICKWEDDE,
SEEGER, Buair, and MoHLER.

RayYMOND J. SEEGER, Recording Secretary.

@Obituary

Cyrus ADLER, president of Dropsie College for Hebrew and Cognate
Learning, died at his home in Philadelphia on April 7, 1940. He was born
in Van Buren, Ark., on September 13, 1863. He received the A.B. degree
from the University of Pennsylvania in 1883, the A.M. in 1886. From
Johns Hopkins University he received his Ph.D. in 1887. Hebrew Union
College gave him his L.H.D. in 1925; University of Pennsylvania, his
Litt.D. in 1930. He was in turn Fellow, Instructor, and Associate in Semitic
languages at Johns Hopkins from 1884 to 1893. At the Smithsonian Institu-
tion in Washington he served successively as librarian, 1892-1905; as as-
sistant secretary, 1905-1908; and as assistant curator and curator of historic
archeology and historic religions, 1888 to 1908. He was elected president
of Dropsie College in 1908; acting president of Jewish Theological Seminary
of America in 1916, president in 1924. Throughout his life he was a tireless
worker in the field of education and in recent years served as a member of
the board of education in Philadelphia; as president of the board of trustees
Philadelphia Free Library; and as a member of the board of Gratz College.
He was a delegate to the conference on an International Catalogue of Sci-
entific Literature in 1908 and since served as member of the international
committee. Widely known as an educator, writer, and philosopher, Dr.
Adler was also one of the recognized leaders of the Jews in America.

The bibliography appearing in “Lectures, Selected Papers, Addresses”’
by Cyrus Adler, published by his friends and colleagues on the occasion of
his 70th birthday, includes nearly 600 titles. In addition, there is a list of
17 publications ‘‘edited by Dr. Adler, or issued under his supervision.” He
was editor of the Jefferson Bible; of the American Jewish Year Book, 1899-
1906; of the Jewish Quarterly, since 1910; and one of the editors of the
Jewish Encyclopedia. His writings included Oriental, archeological, and
philological subjects; he was an authority on comparative religion and
American Jewish history. Among other organizations, Dr. Adler was a
member of the Washington Academy of Sciences, a member and past
officer of the American Philosophical Society; the American Jewish His-
torical Society; and the American Oriental Society. He was a member of
the Cosmos Club in Washington and of the Oriental and University Clubs
in Philadelphia.

ee

PALMowtonoay, —Fossil pearls from ae Colorado. g
Kansas. RoLaNnD W. oe ERE aa ae

“of California. JOHN W. Was. te

PALEONTOLOGY.—A new Gisortia. “Wii ‘Mz. Inera

abi seat

Borany. ae, formation in-M atula.

s

California, OLGA HARTMaN. eS

ZooL.ocy.—Notes on the aiplipee Conia: minus Sa
tion of a new variety, Gammarus minus var. we
R. SHOEMAKER.......... a

OBITUARY? CYRUS ADLER.) F900

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JOURNAL :

OF THE
WASHINGTON ACADEMY OF SCIENCES
Vou. 30 OcToBER 15, 1940 No. 10

CHEMISTRY.—Cyanogenesis and enzyme activity in sorghum varie-
ties.1 JAMES F. CoucH and REINHOLD R. Brissz, U.S. Bureau
of Animal Industry.

Our knowledge of the decomposition of cyanogenetic glucosides
by their appropriate enzymes is based to a large extent upon the
interaction of emulsin and amygdalin. This system has been subjected
to much careful study (1, 3, 6, 9). Less is known about the interaction
of other cyanogenetic systems such as prunasin-prunase and lina-
marin-phaseolunatase and very little about the highly important
system dhurrin-dhurrinase which occurs in the sorghums.

Since the enzyme component regulates the rate at which HCN may
be developed in a cyanogenetic system, the quantity or activity of
that component in any given specimen of cyanogenetic plant is an
important factor in the toxicity of the plant. Evidence accumulated
in this laboratory during the past three years indicates that the active
proportion of enzyme in cyanogenetic plants may vary considerably
at different periods and under different conditions not only in the
sorghums but also in species of wild cherry. The greater number of
samples of sorghum appear to be deficient in dhurrinase, but oc-
casional specimens occur in which there is ample enzyme to liberate
the greater portion of the potential HCN in 3 hours. The wild-cherry
species usually contain an abundance of enzyme, liberating all or
nearly all the potential HCN in 2 to 3 hours, but rare samples occur
in which the enzyme is deficient and cannot develop the entire quan-
tity of potential HCN in the plant.

Variation in enzyme activity may explain the failure of some sam-
ples to produce poisoning in animals when chemical analysis indicates
a toxic level of HCN. The lack of cyanogenetic enzyme in Hremophila
maculata has been advanced by Finnemore and Cox (5) to explain the
nontoxic nature of that plant when fed alone and its ability to produce
_ HCN poisoning when some other material containing the enzyme is
fed at the same time.

1 Received May 9, 1940.
413

OCT 26 1949

es ae

414 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

It is of interest to know whether the enzyme activity of plants like ©
the sorghums bears any constant relationship to the quantity of po-
tential HCN in the plant or whether these two factors may vary in-
dependently so that the rate at which the plant develops HCN may
vary at different times. In the case of dried plants, such as hay, it is
known that there is some loss of HCN during the curing process and
the question of the effect upon the enzyme naturally arises. Some
dried samples of sorghum and Sudan grass that have been kept for
several years appear to have lost the greater part of theirenzyme
activity, although they retain considerable potential HCN. It is ob-
vious that such hay would not be dangerous to livestock if the enzyme
is insufficient to develop a toxic quantity of HCN when the hay is
eaten. It is evident, therefore, that in some cases the quantity of
enzyme present in the plant may be the factor that determines
whether poisoning occurs. In view of these considerations it becomes
highly desirable to work out some method whereby the activity or
quantity of the enzyme can be evaluated. Until this has been accom-
plished it will be difficult to study the factors that influence the forma-
tion and storage of the enzyme in the living plant.

In attacking the problem of working out some method by which a
numerical index could be assigned to the enzyme activity of a cyano-
genetic plant, it was thought that some indication might be obtained
by observing the percentage of the total potential HCN that might be
obtained after various periods of maceration. A number of data were
obtained in this way which, while they clearly indicated variation in
the enzyme activity at different times, were not adapted to mathe-
matical treatment and did not lead to any quantitative knowledge of
the enzyme. Samples of these same plants to which an enzyme had
been added and others to which a solution of the glucoside was added
were run at the same time as the untreated samples. The purpose was
to supply enzyme if that factor was deficient or in case the enzyme
should be in excess to supply enough glucoside to get an index of the
excess. In both cases the results, while informative, were not of such a
nature as to yield a numerical index of the enzyme activity. The
failure in these attempts may be due to the fact that in the decomposi-
tion of dhurrin or of prunasin, two enzymatic processes are occurring,
the hydrolysis of the ether linkage with splitting off of the sugar mole-
cule and the hydrolysis of the nitrile to produce HCN. Whether both
of these reactions are catalyzed by the same enzyme or whether two
enzymes are concerned is not known, nor have we any knowledge that
one reaction takes precedence over the other or that the two proceed

Oct. 15, 1940 COUCH AND BRIESE: CYANOGENESIS A15

concurrently. If one enzyme only is concerned and if one reaction pro-
ceeds more rapidly than the other, it is possible that the enzyme might
be quite inhibited in partially hydrolyzing a large quantity of gluco-
side and thus produce very little of the completely hydrolyzed prod-
ucts. We have some evidence that in the case of dhurrin glucose is
split off the molecule more rapidly than HCN. Should some of these
possibilities be true, the problem of accurately evaluating the quan-
tity of enzyme may be quite complex.

A study of the rate at which the enzyme hydrolyzes a definite
quantity of glucoside should yield figures from which the approximate
quantity of enzyme present could be inferred. This is especially true
in the earlier stages of the process where, as shown by Auld (1) the
reaction follows the law for monomolecular reactions. O’Sullivan and
Tomson (8) used the time required for the performance of a certain
amount of work as an index of the activity of invertase and the idea
was applied to emulsin by Helferich (6) and by Willstatter and
Csanyi (9). Helferich determined this “‘Zeitwertquotient”’ as the time
required to hydrolyze 50 percent of substrate. The application of this
method to plants is limited by the fact that it is not possible arbi-
trarily to alter the relative proportions of glucoside and enzyme pres-
ent but one must accept the mixture as it naturally occurs. The
addition of enzyme or glucoside to macerating plant mixtures is only
partially satisfactory since much of the natural enzymolysis takes
place within the plant tissues and there is considerable doubt that
much of the plant enzyme goes into solution in the external fluid
where the added glucoside is. On the other hand, there is little doubt
that much of the added enzyme does not penetrate the plant tissues
and come in contact with the plant glucoside.

EXPERIMENTAL

Twelve varieties of sorghum grown at the Bureau of Plant Industry Ex-
perimental Farm, Arlington, Va., were made available for this season
through the courtesy of Dr. J. H. Martin, of the Bureau of Plant Industry,
who helped in the selection of the varieties and arranged for the planting
and cultivation of the crop. The 12 varieties studied were Dwarf Yellow milo,
Ajax, Spur feterita, Sharon kafir, Grohoma, Sagrain, hegari, Leoti sorgo,
Sumac sorgo, Kansas Orange sorgo, Rex sorgo, and Atlas sorgo.

It was planned to determine in each plant collection the HCN liberated
without preliminary maceration and after water maceration for 1, 2, 3, 4,
24, and 48 hours, and after mercuric chloride preservation for 4 weeks, 8
weeks, and 6 months. When there was a shortage of material, certain of the
least essential samples in this series were omitted. Samples were collected

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VoL. 30, No. 10

JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

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‘O LG LY WaLV MA NISGOIAd LINAYAAdIG, YOU NOILVUYROV IY HALAV SHILGIYVA WOHDYOS WOU GAHHAOOAY GIOYV OINVAOOUGAT{—'T ATEAV EE

Oct. 15, 1940 COUCH AND BRIESE: CYANOGENESIS 417

daily throughout the season at 8:30 a.m. and taken at once to the laboratory,
where they were immediately sliced fine or ground for analysis. The whole
plants were sliced so long as no pith was present. As soon as pith began to
appear, that part of the stalk containing the pith was discarded. When the
plants became fibrous and tough the leaves were stripped off and ground ina
food chopper, the stalks and heads being discarded. Fifty-gram samples
were used whenever sufficient material was on hand and the plant was
weighed out (before grinding or slicing) in slight excess for the number of
analyses planned. The sliced or ground plant was mixed thoroughly and
weighed at once into flasks and jars assembled in advance. The samples for
water maceration were placed in 5-liter round-bottom flasks and stoppered
with a rubber stopper. Those for mercuric chloride preservation were placed
in pint fruit jars. As soon as the last sample had been weighed, the contents of
the jarswere flooded with water and 4 gram (1 percent) of mercuric chloride in
solution added to each. The Jars were then filled to the shoulder with water,
covered, shaken well and stored at 25° C. until the time of analysis. In the
meantime a flask of water had been brought to boiling. For the time series it
was necessary to stop all enzyme action promptly at the beginning of the
analysis. This was accomplished quite effectively with boiling water and
mercuric chloride. To the sample marked zero hours, 1,500 ce of boiling
water containing | gram (2 percent) of mercuric chloride was added. The
flask was connected to a condenser and the burner lighted at once. As soon
as vapor began to condense in the delivery tube above the stopper, 2 grams
of stannous chloride in water was added, the connections quickly reinserted,
and without removing the flame the mixture was well shaken while holding
the stoppers firmly in place. The HCN was then distilled off and caught in a
500-ce Erlenmeyer flask containing 10 ce of 5 percent potassium hydroxide.
Two distillates of 400 cc each were taken for each sample. For the remaining
samples a quantity of warm tap water sufficient for 500 cc per sample was
run into a large container and adjusted to 37° C. To each flask was added
500 ce as rapidly as it could be measured in a graduate and the rubber stop-
pers reinserted. The flasks were placed at once in the incubator at 37° C.
The whole operation of adding the water required from 3 to 6 minutes, de-
pending on the number of samples in the series. When the period of macera-
tion had elapsed for each sample, it was diluted to 1,500 ce with 1,000 cc of
boiling water containing 1 gram (2 percent) of mercuric chloride. The flask
was then connected, heated, stannous chloride added, and the contents dis-
tilled just as in the case of the zero hour sample. The HCN content of the
distillates was determined as previously described (2).

The amount of HCN liberated each hour for the first four hours was de-
termined in these experiments because an animal that is to become poisoned
from sorghum will ordinarily show the effect within four hours. The animal
will very likely not become poisoned if no effect is produced within four
hours after the plants are eaten.

i _—_ ss,

418 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

DISCUSSION OF RESULTS

The data obtained in the experiments are tabulated in Tables 1 and 2.
The data in Table 1 are expressed in milligrams of HCN per 100 grams of
plant calculated to the dry basis. Columns 6 to 12 give the figures for the
amount of HCN liberated by water maceration during intervals of 0, 1, 2,
3, 4, 24, and 48 hours. The total amount of potential HCN in the plant as
determined by 4 and 8 weeks of maceration in mercuric chloride is shown in
column 13, the highest value thus obtained being given and the values simi-
larly expressed in mg of HCN per 100 grams of dry plant. Column 14 gives

TABLE 2.—SorRGHUM VARIETIES LIBERATING THE HIGHEST AND LOWEST PERCENT OF
THE PoTENTIAL HypRocYANIC ACID FOR DIFFERENT PERIODS OF MACERATION OF
THE SAMPLES IN WATER AND IN MERcURIC CHLORIDE

ae oh Highest Lowest
maceration
In water Variety Coll. no. ese Variety | Coll. no fee
hrs. 1939 % 1939 %
0 Sharongkahir an 5 ee 25 17.6 HMegartc eer ieee Bil 4.0
1 Hegari (frosted)...... 57 36.8 SUMAC Aa see 16 4.5
2 AGIASISOL2On ose aoe 30 Sato Fle anit. cin eee 14 ey?
3 7-NI Vp: Geran RO na Cee a eal 43 69.0 Hegart.: 25. een 42 14.1
4 NAS CECE Baty nae een ste Eerste 43 80.5 iegaria sch ones ee 31 14.2
24 Spur feterita......... Pit 97.5 legart ..... <2 ee 55 41.2
48 Spur feterita......... 26 93.8 Leoti sorgo.......... 17 54.6
Ina @lo i Aviaxens crease eee 43 487 mg. Megaris ss i. 4 54 73 mg.

the time in hours at which 50 percent of the potential HCN had been liber-
ated. These values were obtained from graphs of the figures for the number
of milligrams of HCN liberated against the time intervals 0, 1, 2,3, 4, and 24
hours corresponding to these figures. Column 15 gives the rate index for 50
percent hydrolysis. This figure indicates the number of milligrams of HCN
obtained per hour up to the time at which 50 percent of the potential HCN
had been liberated and was calculated from the data in columns 13 and 14.
The data in Table 1 are arranged according to height of plants. The number
of milligrams of HCN liberated at the stated time for hegari are given in the
first four sections of the table. Similar figures are given in sections 5 and 6 for
11 varieties of first-growth plants and 4 varieties (5 samples) of second-
growth plants all less than 15 inches in height. The last section gives the
figures for plants 2 feet or more in height for all varieties excluding hegari.
Spur feterita, 20-22 inches high, was placed in this section although slightly
less than 2 feet high. This sample is the exception mentioned in the next
paragraph.

The data in Table 1 reveal the outstanding fact that young sorghum liber-
ates HCN at a much higher rate than older plants. The rate index expresses
this fact numerically. All the high values for the rate index are seen to be
associated with the sorghums less than 15 inches in height, sections 1, 5, and
6, with one exception, namely, Spur feterita, 20-22 inches high in section 7,
which has a rate index of 31 although it is nearly 2 feet in height. Another
exception is Sharon kafir, section 5, young first growth which has a rate index

Oct. 15, 1940 COUCH AND BRIESE: CYANOGENESIS 419

of 13.8 while the next lowest value in this section is 20.9. No explanation for
these exceptions can as yet be given.

A comparison of first and second growth sorghum brings out the fact that
some samples of second growth have an exceptionally high rate index. For
example second growth Ajax with a potential HCN value of 487 milligrams
has a rate index of 100.5 and Spur feterita has corresponding values of 240
and 60. The highest value for young first growth was that of Spur feterita
with a potential HCN value of 292 milligrams and a rate index of 56.1. This
comparison between first and second growth is well brought out by the re-

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HOURS

Fig. 1—Average percent of the potential hydrocyanic acid liberated from sor-
ghums after maceration in water for different periods of time.

spective averages calculated from Table 1, as given in Fig. 1, in which the
percentages of the potential HCN liberated are plotted for the various time
intervals. The curves show that 50 percent hydrolysis is reached earlier in
the case of second growth plants so far as can be determined by a small
number of samples.

The activity of the enzyme as indicated by the rate index is a factor de-
termining the rate of evolution of HCN and hence the toxicity of sorghum.
The high rate indices associated with young plants, including second growth,
first growth and suckers, indicate that sorghum in its early stages of growth
is a dangerous food for livestock. From the present state of our knowledge it
is as yet impossible to say whether plants with low rate indices are toxic to
animals when eaten in large quantities and if so what determines the toxic
level for such plants.

The rate index is independent of the quantity of potential HCN in the

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420 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES’ VOL. 30, NO. 10

plant. A comparison of the amount of potential HCN in sorghum with the
corresponding rate indices showed that there was no correlation between the
two. Samples with a potential HCN content of 200 and 206 milligrams
showed rate indices of 32.3 and 4.3, respectively. Similar discrepancies were
numerous throughout the list.

It is of interest to note which varieties of sorghum liberated the lowest
percent of the potential HCN and which the highest for the different periods
of maceration in water and in mercuric chloride. Data for such a comparison
are given in Table 2. Hegari is shown to be the lowest in 6 out of 8 instances
and Sumac sorgo and Leoti sorgo each in one instance. Among the highest
are Atlas sorgo, Ajax, and Spur feterita.

In Fig. 1 the curve or the averages of all first growth plants 2-6 feet in
height shows a slowing up of the rate of evolution of HCN between the 2-
and 3-hour points followed by an increase. All samples from plants of this
size were ground in a food chopper for analysis. Hence a certain amount of
glucoside and enzyme from the crushed sample were in intimate contact
and therefore the hydrolysis proceeded more rapidly from the beginning than
in the case of sliced first growth plants. The grinding, however, was not suf-
ficiently fine to expose all the glucoside and enzyme and a certain time
would be required for the penetration of the solvent and subsequent dif-
fusion of the reactants and reaction products. The result is a slowing up of
the reaction rate as indicated by the curve after the 2-hour point in the
averages. Young plants, which contain more water and are less fibrous, per-
mit a rapid diffusion of glucoside and enzyme and do not show this phe-
nomenon. No analyses were made this season for periods between 4 and 24 |
hours. It is conceivable that this slowing of the reaction could occur later
than at the 4-hour period in a sample with a slow reaction rate and if at the
same time the curve in this area was rather flat a dip could occur due to the
rate of destruction of HCN for a time exceeding the rate of its formation.
Such a dip has been discussed in a previous publication (2).

It is of interest to calculate how much difference in toxicity the variation
in rate of evolution of HCN would be expected to make. Taking the m.].d.
of HCN for cattle as 2 mg per kg and for sheep as 2.3 mg per kg, then on the
basis of the average of actual HCN evolved from hegari in 4 hours, the
8-14 inch plants would furnish one m.1.d. for a 800-kg cow in 281 g of plant;
the 24—26 inch plants in 1,215 g and the 6-foot plants in 1,378 g. For a 50-kg
sheep the figures are: 53.8 g, 233 g, and 264 g. respectively. The potential
HCN contents (average) of the plants are as 100: 71.7:32 percent.

SUMMARY

The activity of the enzyme determines the rate at which a cyano-
genetic plant evolves HCN and so is a determining factor in the tox-
icity of the plant. Unless the enzyme is sufficiently active to evolve a
toxic quantity of HCN quickly, poisoning is not likely to occur.

Oct. 15, 1940 COUCH AND BRIESE: CYANOGENESIS 421

Young second-growth and first-growth plants including suckers have
a much higher rate index than leaves of well-developed sorghum
varieties and consequently are to that extent more likely to cause
poisoning. Whether plants with a low rate index are toxic when larger
quantities are eaten and if so what determines the toxic level for such
plants are questions requiring further investigation.

The enzyme activity of a cyanogenetic plant may be numerically
defined as the number of milligrams of HCN evolved per hour until
50 percent of the potential amount has been set free under certain
conditions of temperature and dilution. The figure is referred to as the
“rate index.”

The rate index was determined for a number of varieties of sor-
ghum collected at Arlington Farm during the summer of 1939. The
rate index is high for young plants of all varieties less than 15 inches
high and low for plants of all varieties above 2 feet high. The rate
index is not correlated with the quantity of potential HCN and
varies independently of it. Consequently the activity or quantity of
the enzyme in the plants is not a function of the quantity of dhurrin.

LITERATURE CITED

1 Autp, 8. J. M. The hydrolysis of amygdalin by emulsin. Part I. Journ. Chem. Soe. T.
92: 1251-1275. 1908.

. BRIESE, R. R., and Coucu, J. F. Preservation of cyanogenetic plants for chemical

analysis. Journ. Agr. Res. 57: 81-108. 1938.

. CALDWELL, R. J., and Coutrauvtp, 8. L. The hydrolysis of amygdalin by acids.

Journ. Chem. Soc. T. 91: 666-671. 1907.

. Coucnu, J. F., and Brizsz, R. R. Hydrogen ton concentration and cyanogenesis in

sorghum. Amer. Journ. Pharm. 111: 55-64, 151-160, 193-201. 1939.

. FINNEMORE, H., and Cox, C. B. Cyanogenetic glucosides in Australian plants, Part II.

Eremophila maculata. Journ. Roy. Soc. New South Wales 63: 172-178. 1929.

. Hewurericn, B. Uber Emulsin. Zeitschr. physiol. Chem. 117: 159-171. 1921.

Martin, J. H., Coucn, J. F., and Brizssz, R. R. Hydrocyanic acid content of different

parts of the sorghum plant. Journ. Amer. Soc. Agron. 30: 725-734. 1938.

. O’Suuuivan, C., and Tomson, F. W. Invertase; A contribution te the history of an

enzyme or unorganized ferment. Journ. Chem. Soc. T. 57: 834-931. 1890.

. WILLSTATTER, R., and Csany1, W. Zur Kenninis des Emulsins. Zeitschr. physiol.
Chem. 117: 172-200, 1921.

co ONO oO - WwW WN

422 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

PALEOBOTANY.—A bracket fungus from the late Tertiary of south-
western Idaho... Rotanp W. Brown, U. 8. Geological Survey.

The specimen to be described here was collected in 1939 by James
L. Morris, registered engineer of Boise, Idaho, at a locality about
15 miles east of a point 5 miles south on the highway from Bruneau,
Idaho, to Owyhee, Nev. Associated with this specimen on the surface
at the same locality is much fossil wood, which includes species of fir
(Abies), alder (Alnus), poplar (Populus), oak (Quercus), and hickory
(Hicoria). From a sketch and photograph submitted by Mr. Morris,
it would appear that the fossils occur in a stratum lying just above a
white bed of volcanic glass sand. These strata may be part of the
Idaho formation, which in the course of its history has been made to
include a rather heterogeneous series of lake and basin deposits lying
above the Columbia River basalt. The presence of undoubted hickory
among the fossil woods suggests a much moister climate than that
now prevailing in the area, and it also suggests that the age of that
part of the formation is probably not later than early Pliocene.

Analysis of the specimen shows that it is composed principally of
calcium carbonate with minute amounts of other substances, some
of which may have come from clay or other extraneous material lodged
in the pores subsequent to the calcification of the specimen. The fact
that this fungus and the woods are calcified, with preservation of the
original cellular structure as perfect as the best examples of silicified
specimens of the same kind, raises some questions that the writer
can not answer at this time, such for example as: What was the prob-
able immediate source of the calcium carbonate? Why did calcifica-
tion take place instead of silicification? A thorough examination of
the geological setting would doubtless throw light on these matters,
but the writer has not had an opportunity to visit the locality. Ac-
cording to the sketch there are now hot springs 6 miles east of this
locality, but whether these or similar springs that may once have
existed closer by had anything to do with this fossilization is only
conjectural. It would seem that definite answers to these questions
might perhaps precipitate the long-sought solution to the problem as
to how calcification and silicification take place.

POLYPORACEAE
Fomes idahoensis Brown, n. sp. Figs. 1-4
This specimen is the sporophore or spore-bearing portion of a bracket
fungus. It is 13.5 em long and 4.5 cm thick. The upper surface (Fig. 1) is or

1 Published by permission of the Director, Geological Survey, U. S. Department
of the Interior. Received May 29, 1940.

Oct. 15, 1940 BROWN: TERTIARY BRACKET FUNGUS 423

\
r
i

Wem) ) poms

Figs. 1-4.—Fomes idahoensis Brown, n. sp.: 1, View of top; 2, pores, 4; 3, side
view showing context above and pore area below; 4, view of bottom. Figs. 1, 4, <é.

S33

bh;

424 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

was fairly smooth and displays a series of convex-rounded ‘‘rings”’ of growth.
The under surface (Fig 4): flat and porous, the pores numbering about 750
per sq. cm. These are shown in Fig. 2, magnified four times. Fig. 3 is a cross
section through the right hand end of Fig. 4 and illustrates the vesicular
corky context separated from the pore area by an irregular dark zone.
Microscopic examination reveals no spores or other significant cellular struc-
tures.

The specimen resembles the living bracket fungus, Fomes pinicola
(Swartz) Cooke, so closely that no doubt exists as to its affinities. The only
difference appears to be in the size of the pores, which are slightly larger
in the fossil, thus aggregating only about 750 per sq. cm as compared with
about 1,000 in F. pinicola. The species of the genus Fomes attack many
kinds of trees. Fomes pinicola is found on conifers generally and is not con-
fined to Pinus specifically as the name might suggest. The fossil may have
lived on one of the numerous species of conifers recorded from the late
Tertiary of the western United States.

Mason? records Fomes applanatus (Persoon) Gillet from the Pleistocene
of the Tomales Bay region, Calif. This appears to be the only other au-
thentic American fossil polypore fungus. Those described by Wieland and
Brown in 1934 and 1936, respectively, have since been shown by Brown?
to have been misidentified. Seward‘ in 1898 accepted as authentic several
records of Polyporaceae from Europe, the earliest being an Oligocene
species of Polyporus described by Conwentz from the Baltic amber deposits.
Meschinelli,®> however, records a number of additional species.

The writer expresses appreciation for help in this study to John A. Steven-
son, U. 8. Bureau of Plant Industry, and Kiguma J. Murata, U. 8. Geo-
logical Survey.

2 Mason, H. L. Pleistocene flora of the Tomales formation. Carnegie Inst. Wash-
ington Publ. 415(4): 148, pl. 5, figs. 1, 4. 1934.

3 Brown, Routanp W. Two fossils misidentified as shelf-fungi. Journ. Washing-
ton Acad. Sci. 28: 130-131. 1938.
4Sewarp, A.C. Fossil plants. 1: 207-222. 1898.

5 Mescuinetu, A. Fungorum fossilium omnium hucusque cognitorum icono-
graphia. 1898.

Oct. 15, 1940 KENG: A NEW SPECIES OF BAMBOO 425

BOTAN Y.—Oxytenanthera felix, a new species of bamboo from Yun-
nan, China. Y.L. Kena, National Central University, Chung-
king, China. (Communicated by AGNES CHASE.)

The genus Oxytenanthera Munro comprises about 20 species, from
tropical Asia, New Guinea, and Africa. The species here described is
the first known from China.

Oxytenanthera felix Keng, sp. nov.

Culmi teretes, 9 m alti, ramis foliiferis (in specimine) usque 90 cm longis,
5 mm crassis, basi squamis glabris stramineis cinctis; vaginae sese arcte
involventes, infima 12 cm longa, glabrae vel superiores pilis adpressis 0.5—1
mm longis strigosae; ligula firma, 3-4 mm longa, saepe bipartita; laminae
saepissime 30-45 cm longae, 4.5-6 cm latae, acutissimae, glabrae, basi in
petiolum 4-7 mm longum attenuatae, margine scaberulae vel veteres fere
laeves; rami floriferi simplices, aphylli, subherbacei, usque 2.5 m longi,
internodiis inferioribus cire. 15 cm longis, 9mm crassis; spiculae dense fascic-
ulatae vel plurimae in capita magna usque 5 cm crassa dispositae, pleraeque
4-florae, 16-20 mm longae, pallide virides vel ad maturitatem stramineae;
glumae 1-3, late ovatae, 3-9 mm longae; leommata ovato-lanceolata, 11-17
mm longa, multinervia, superiora convoluta, pungenti-mucronata; palea
multo angustior, 11-15 mm longa, suprema convoluta, inferior bicarinata,
carinis marginibusque superne longe ciliata; lodiculae nullae; stamina 6,
filis inferne connatis, hyalinis, usque 15 mm longis, antheris 4-7 mm longis,
apice minute hispidulo (interdum fere glabro) apiculatis; ovarius linearis,
stipitatus, in stylum hispidulum cire. 1 cm longum attenuatus, stigmato uno,
plumoso, circ. 9 mm longo.

Culms terete, 9 meters tall, the leaf-bearing branches (in specimen) arising
singly from the nodes of the main culm or primary branches, about 6 mm
thick and medullated with whitish-flocculose pith, covered at base with gla-
brous stramineous scales, about 90 cm long, the lower 4 internodes 6-16 cm
long, glabrous; sheaths 10 on the upper part of the leafy branch, tightly
overlapping, the lowermost 12 em long, keeled or the midnerve prominent
above, glabrous or the upper strigose with appressed hairs 0.5-1 mm long;
ligule firm, prominent, 3—4 mm long, frequently 2-partite or sometimes con-
cave at apex; blades mostly 30-45 cm long, 4.5-6 em wide, with a strong
midnerve and 10-13 secondary nerves on either side, cross-veined, sometimes
obscurely so, acuminate-pointed, cuneate at base with a petiole 4-7 mm long,
glabrous or the pale lower surface very sparsely strigose with appressed
deciduous hairs, the margins scaberulous or in age nearly smooth; floriferous
branches simple, aphyllous, up to 2.5 meters long, the lower greenish sub-
herbaceous internodes about 15 cm long, 9 mm thick; spikelets densely
clustered or numerous in large heads up to 5 em thick, usually 4-flowered,
16-20 mm long, pale green or stramineous at maturity; rachilla joints very
short or the nodes nearly approximate; glumes 1-38, broadly ovate, acute or
obtusish, successively longer, 3-9 mm long (when 3, the first sometimes
keeled, the keel ciliate, the second sometimes paleate), multinerved, cross-
veined, glabrous or the margins ciliolate; lemmas ovate-lanceolate, 11-17
mm long (the lowermost sometimes only 7 mm long, broadly ovate, stami-
nate), multinerved, the upper ones usually convolute, pungent-mucronate,

1 Received Apri! 30, 1940.

*
_. “..~

, apo?

426 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES’ VOL. 30, No. 10

glabrous or the margins ciliolate above; palea very narrow (about 1.5 mm
wide between the keels), 11-15 mm long, the uppermost convolute, the lower
ones 2-keeled the keels and the inflexed margins long-ciliate above; lodicules
none; stamens 6, the filaments connate into a tube around the pistil, hyaline,
up to 15 mm long, the anthers 4-7 mm long, apiculate with minutely his-
pidulous tip (or sometimes nearly glabrous); ovary linear or somewhat en-
larged at base when ripe, stipitate, hispidulous, attenuate into a hispidulous
style about 1 cm long, then continuous with a plumose stigma about 9 mm
long.

Type in the U.S. National Herbarium, nos. 1214319 (flowering specimen)
and 1214320 (leaf-bearing specimen), collected on plain of the Nam Ha,
between Muang Hai and Keng Hung, Yunnan Province, altitude 1,260-
1,350 meters, February 15-17, 1922, by J. F. Rock (no. 2462).

BOTANY.—Tazonomic relationships in the genus Gossypium.!
S. C. Haruanp, Sociedad Nacional Agraria, Lima, Peru. (Com-
municated by T. H. KEaRNgEy.)

There have been three recent papers dealing with the classification
of the genus Gossypium: Hutchinson and Ghose (1937), Hutchinson
(19388), and Hutchinson (1939). The third of these is a condensed
and modified version of the first two. The discussion of Hutchinson
and Ghose (1937) on the taxonomy and relationships of the Asiatic
cottons calls for no comment. It seems to the writer an accurate and
painstaking attempt to straighten out a confused and complicated
situation. The treatment of the other groups is less satisfactory. First,
a slight error is made in the statement that G. davidsoni and G.
klotzschianum do not cross with cultivated New World cottons. Both
in fact do cross with Sea Island cotton (G. barbadense L.) but produce
seedlings that, although germinating vigorously, die when young
through a progressive necrosis of the cotyledons.” Exception must
also be taken to the statement that the two Australian species G.
sturtaz and G. robinsonii are probably best classified with the New
World diploid cottons. About G. robinson we know nothing except
from herbarium specimens, so that it may be removed from the dis-
cussion. Information on the relationship of G. sturtiz to other diploid
species is derivable only from cytological sources (Webber, 1935, 1936,
and Skovsted, 1937).

1 Received June 14, 1940. The writer is indebted to Dr. T. H. Kearney and to
Dr. H. J. Webber, who have read this manuscript and furnished valuable suggestions.

2 Dr. J. M. Webber informs the writer (April 24, 1940) that he has obtained one
plant from the cross barbadense X davidsonit which grew normally and flowered pro-
fusely, exhibiting the most typical Drosera scheme meiotic chromosome behavior
(131 +13) of any Gossypium hybrid yet examined.

Oct. 15, 1940 HARLAND: TAXONOMY OF GOSSYPIUM 427

Webber (1936) reported the following results in crosses between
sturta and New World diploids:

Cross TYPE OF CONJUGATION
sturtii X amourianum Oi +26; (most) to 61 141
sturtit < harknessiz Or +26; (most) to 411 18;
davidsoni X sturta Or +261 (most) to 5m 167
thurbert X sturti 261 (all)

In a later communication Webber (1939) gave further data as fol-
lows:

Cross UNIVALENTS BIVALENTS
G. sturtia XG. armourianum 24.08 0.96
G. sturtit XG. harknessii 24.56 0.72
G. davidsoni XG. sturtit 23.94 A038
G. thurberi XG. sturti 24.52 0.74

Skovsted (1937) presented cytological data on crosses between sturtii
and (a) davidsoni, (b) armourianum, and (c) Asiatics. He gave the
following results:

Cross UNIVALENTS BIVALENTS TRIVALENTS QUADRIVA-
LENTS
davidsonit X sturtit 14.95 5.32 0.10 0.02
sturtui Xarmourianum 8.45 8.20 0.25 0.10
Asiatic X sturti 2.0-3.7 9.5-10.2 0.5-0.8 0.3-0.45

Here it is evident that sturtzz is cytologically closer to the Asiatic cul-
tivated cottons than to the New World diploids. Skovsted states that
pollen mother cells with 13 bivalent chromosomes are found in the
hybrids between Asiatic cottons and G. sturtiz and between Asiatic
cottons and G. anomalum, and again, ‘‘G. anomalum and G. sturtii
separated in different directions and to different continents. They yet
retain sufficient cytological similarity with Asiatic cottons to permit
of fairly normal conjugation in their hybrids.’’ If therefore sturtii is
to be classed with any other group, it should, agreeing with Skovsted,
go with the Asiatic and not with the American diploids. Webber, how-
ever (1939), obtained in the cross herbacewm Xsturtii 21.56 univalents
and 2.22 bivalents. Since cytological criteria disagree and genetical
criteria are entirely lacking, owing to complete sterility of all recorded
crosses, it seems best pending further evidence to put sturtii in a group
by itself, as has been done by the writer (1939).

Some comments may now be offered on certain changes that Hutch-
inson and Ghose (1937) and Hutchinson (1937-88) propose to make
in the taxonomic status of some species in the tetraploid New World
cottons and that differ from the schemes put forward by the writer
(1932, 1939).

=)

~

428 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

The specific rank assigned darwini, tomentosum, barbadense, and
tartense is not in dispute, but regarding the remaining species, two
species and one subspecies were recognized in the writer’s first scheme
(1932): (1) G. hirsutum L. (the American Upland of commerce); (2) G.
purpurascens Poir. (the Bourbon group), including G. punctatum Sch.
and Thon. (the punctatum group). Later, the writer (1939) tentatively
and hesitantly gave specific rank to punctatum. Hutchinson and
Ghose (1937) followed this scheme but changed the name G. purpuras-
cens to G. religiosum. In a later communication Hutchinson made a
further modification and proposed that purpurascens and punctatum
should now be considered as varieties of hirsutum under the names
var. purpurascens and var. religiosum.

Now since the writer disagrees with the first tax ononnte change in
so far as purpurascens is termed religiosum, and with the second
change in so far as purpurascens and punctatum are made varieties of
hirsutum, it seems to be necessary to discuss the interrelationships of
hirsutum, purpurascens, and punctatum rather fully. Let us first de-
fine what we mean by the terms. By G. hirsutum L. we mean the huge
and variable assemblage of forms comprised under the name Ameri-
can Upland cottons. The type of the species is in the Sloane Her-
barium, B. M., vol. 294, p. 45 (Miller’s specimen named by himself).
It is important to note that the type herbarium specimen consists of
a small branch with seven or eight leaves and two flowers (or buds).
To use it as an accurate means of discriminating between the modern
hirsutum and related species can be done only by a process of ration-
alization and might well be merely taxonomic pedantry.

A whole series of minor details of form which characterize hirsutum
are not visible in the specimen. It can not be said, for example, whether
the specimen is derived from a monopodial or sympodial plant, or
whether the flower is really characteristically hirsutum. It is true that
the very special leaf shape of hirsutum is present, and this delimits it
from barbadense, a species in which this leaf shape is unknown. The
process we have to go through, then, is this: We identify (rightly or
wrongly) the type specimen of G. hirsutum L. with the modern hir-
sutum. We then compile a taxonomic description of hirsutwm from
modern material and in the process discover what are the characters
of taxonomic significance, which probably are not visible in the origi-
nal specimen and which really do characterize hirsutum and mark it
off as a good species. We go through the same process with other spe-
cies thought to resemble an original type specimen, and then, by using
a combination of methods in which anatomy, morphology, ecology,

Oct. 15, 1940 HARLAND: TAXONOMY OF GOSSYPIUM 429

geographical distribution, and genetics all play a part, though in vary-
ing order of importance, we decide upon species nomenclature.

The question then is whether a detailed examination from the
above points of view justifies the merging of purpurascens and punc-
tatum into the species hirsutum. Such an examination must make use
of the criterion proposed but not employed by Hutchinson and Ghose
(1937) when they say: “In devising a satisfactory classification for
the species determined by genetic study, it is necessary to use only
such characters as are indicative of fundamental differences in genetic
composition.”’

About the punctatum group little will be said in this article, since
more genetical information is required before discussing its taxonomic
position. It is obviously related to both Uplands (hirsutum) and Bour-
bons (purpurascens), and Lewton (1912) thought it worth while to
describe a characteristic member of this group as a separate species,
G. hopi Lewton.

The use of the specific name G. purpurascens Poir. to denote the
Bourbon group was adopted by the writer (1932) from Watt (1907).

The drawing of G. purpurascens Poir. (No. 44 opposite p. 250) so
closely resembled the typical Bourbons in the writer’s collection, par-
ticularly in the calyx shape, number of bract teeth, narrow, cup-
shaped flower, and boll shape, that it seemed the most convenient
name to adopt, more especially since the general description of the
species given by Watt also conformed. It was recognized that the
Bourbon group was related to the Upland group, but how distantly
was not known.

Subsequently, descriptions were made of as many morphological

TABLE 1.—GENETICAL DIFFERENCES BETWEEN PURPURASCENS
AND HIRSUTUM

purpurascens — hirsutum
Gene Character Gene Character

Notkiown. ~....... 1 RS bk ite ta tee, aie Red leaf.
eemOne tS fe io. a Red leaf. INotekn owns aac
rere eh) creas Petal Spot (muta- || Not known.......

ble on hirsutum

background).
NoteknMowml........:. Spee came bee Petal spot.
Not known. 2... 6... GUE BO cM eee Green lint.
ChE 3 ae Corky (comple- || Not known.......

mentary).
oe a Naked seed (reces- || Not known.......

cessive).
Not KmOwin sc i. k)s0- Dy pene ee Eee Naked seed (domi-

nant).

430 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES’ VOL. 30, No. 10

characters as possible, and it was found that the means of many of
them were intermediate between barbadense and hirsutum. In other
characters, more especially the time of bursting of the anthers and
the features of the filaments, the Bourbons differed from both hirsu-
tum and barbadense. Genetical studies of crosses between Bourbon and
Upland have also been conducted by the writer since 1926 and have
revealed that the genetical differences between these two groups are
very considerable. The main ones are given in Table 1.

MODIFIER COMPLEX DIFFERENCES BETWEEN
PURPURASCENS AND HIRSUTUM

1. The modifier complexes affecting ‘‘crinkled’’ are completely dif-
ferent in hirsutum and purpurascens. The gene for ‘‘crinkled’’ of
barbadense was transferred to a typical purpurascens (morrilli Cook,
and Hubbard) by several repeated backcrosses, and then isolated by
selfing. When crossed with barbadense crinkled and hirsutum crinkled,
respectively, it gave the following results in F,:

(a) purpurascens crinkled X barbadense crinkled.

Complicated series of forms from supercrinkled to pseudonormal.
(b) purpurascens crinkled X hirsutum crinkled.

Complicated series of forms from crinkled to pseudonormal.

The variation in grade of crinkled in the purpurascens-hirsutum
cross, although very considerable, was rather less than in the
barbadense-hirsutum or purpurascens-barbadense series. This set of re-
sults is probably the most important of those yet available in de-
ciding on the taxonomic position of purpurascens, and indicates such
a specificity in the crinkled modifier complex as to confirm pur-
purascens in its position as a good taxonomic species, perhaps occupy-
ing a position intermediate between barbadense and hirsutum but
nearer to the latter.

2. The fact that the petal spot gene S’ of purpurascens becomes
reduced in intensity and becomes mutable, also indicates differing
modifier complexes in hirsutum and purpurascens.

STERILITY IN CROSSES OF PURPURASCENS AND HIRSUTUM

Some sterility has been noted in the F, of purpurascens—hirsutum
crosses. Such sterility is not to be expected between varieties assigned
to the same species.

CHARACTER DIFFERENCES

Some character differences between purpurascens and hirsutum are
worthy of note.

Oct. 15, 1940 HARLAND: TAXONOMY OF GOSSYPIUM 431

CHARACTER PURPURASCENS HIRSUTUM
Flower shape Narrow cuplike Widely expanded
Anthers—time of bursting Late Early
Number of bract teeth Few Many
Leaf shape Mean L/S* of 2.5 L/S about 1.6-1.8
Kidney seed Known but rare Not known

* L/S = Length of leaf +length to sinus.

LEAF SHAPE

There are several allels for leaf-shape in purpurascens. The char-
acteristic broad leaf of hzrsutwm is extremely rare in purpurascens, be-
ing present in one type only of 20 standard purpurascens types se-
lected from widely differing localities.

BRACT TEETH

The mean number of bract teeth in purpurascens varies from 4.0
to 9.5 with a mean of 6.0 in the 20 types. This character appears to be
of quite definite significance in purpurascens. :

Table 2 shows the correlation diagram between leaf shape (L/S-

TABLE 2.—CORRELATION: LEAFSHAPE AND NUMBER OF Bract TEETH
20 Representative Bourbons

s[slejele| ai Mets

Leaf Shape L/S

—_
>
ez)
iw)
j=)
ee)

ice

|
|

| 92

5.0) 5.5) 6.0) 6.5) 7.0) 7.5) 8.0) 8.5) 9.0} 9.5)10.0/10.5/11.0/11.5)12.0

ese ers ie St hie a

Mean number of bract teeth

The numbers in the squares refer to the type number in the World Collection.
© =Mean of Bourbon Group.
@ =Approximate position of Upland.

432 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

length +length to sinus), and number of bract teeth in a representa-
tive group of Bourbons. The position of a representative hirsutum is
marked, from which it will be seen that the distribution of all the
Bourbons is clearly demarcated from hirsutum.

SUMMARY

Enough has been said to make it evident that the Bourbon group
is distinct from the Upland group on both morphological and genet-
ical grounds. In the writer’s view, the assemblage of Bourbons is a
good taxonomic species and is as distinct from Upland as, say, arbor-
eum is from herbaceum. Bourbon should continue to be known as G.
purpurascens Poir. pending a better name.

LITERATURE CITED

Haruann, 8. C. The genetics of Gossypium. Bibl. Genet. 9: 107-182. 1932.

. The genetics of cotton. 193 pp. London, 1939.

Hurcuinson, J. B. The distribution of Gossypium and evolution of the commercial cot-
tons. Conf. Sci. Res. Workers on Cotton in India. Bombay, 1938.

. The classification and evolution of cotton. Trop. Agr. 16: 82-83. 1939.

, and Guoss, R. L. The classification of the cottons of Asia and Africa. Indian
Journ. Agr. Sci. 7: 233. 1937.

LewtTon, F. L. The cotton of the Hopi Indians: A new species of Gossypium. Smith-
sonian Misc. Coll. 60 (6): 1-10. 1912.

SxovstepD, A. Cytological studies in cotton. IV : Chromosome conjugation in interspecific
hybrids. Journ. Genet. 34: 97-134. 1937.

Watt, G. The wild and cultivated cotton plants of the world. London, 1907.

Wesser, J. M. Interspecific hybridization in Gossypium and the meiotic behavior of
F, plants. Journ. Agr. Res. 51: 1047-1070. 1935.

. Cytogenetic notes on cotton and cotton relatives, IJ. Science 84: 378. 1936.

. Relationships in the genus Gossypium as indicated by cytological data. Journ.

Agr. Res. 58: 237-261. 1939.

ENTOMOLOGY.—Some new species of Syrphidae (Diptera).!
FRANK M. Huu, University of Mississippi. (Communicated
by E. A. CHAPIN.)

This paper describes some neotropical Syrphidae, some of which
were collected by the author at Barro Colorado Island and others ac-
cumulated. Types are in the author’s collection.

Mesogramma lyrata n. sp.

Male.—Length 6 mm. Head: Front and face pale yellow, cheeks black,
antennae wholly pale orange, arista black on apical two-thirds. Vertex violet
immediately behind the ocelli, brassy brown posteriorly. Thorax: With a
median, bluish-gray, linear vitta outside of which it is very broadly brassy
brown. Viewed from the rear there is sublaterally, above the complete yel-
low lateral margin, a light brown and then a dark brown vitta. Scutellum
brownish yellow, diffusely darker brown upon the disk, its discal pile pale,
its marginal pile black, the posterior part of mesopleurae and the upper
part of sternopleurae pale yellow. Abdomen: Brownish orange, marked with

1 Received June 6, 1940.

Oct. 15, 1940 HULL: NEW SPECIES OF SYRPHIDAE 433

dark brown. First segment yellow, with a pair of posterior, slender, black
fascia separated in the middle, not reaching the sides. Second segment with
a posterior, marginally evanescent, complete brown fascia, a quite evanes-
cent incomplete subbasal fascia which shows traces of connection with the
posterior one. Third segment with a slender subapical brown fascia curving
up for a short distance along the posterior margin, also reaching the posterior
corner and near the midline turning sharply upward to form a pair of out-
wardly turned, comma-like brown spots. Fourth segment with similar pat-
tern but only the comma-like brown spots apparent. Fifth segment and
hypopygium brownish orange. Legs: Pale yellow, the anterior tarsi orange,
the posterior tarsi brownish, the hind femora and tibiae with subapical and
subbasal blackish annulae respectively. Wings: Clear hyaline, stigma pale
yellow.

Holotype.—Male. August 30, 1938, Barro Colorado Island (F. M. Hull
collector).

Mesogramma basilaris Wd. var. flavocuneus n. var.

Female.—Length 6 mm. Head: Face yellow, a small obscure blackish spot
beneath vertex violet; the front flat, smooth, metallic. The sides sharply
marked with pale yellow almost to the ocelli. Cheeks black, antennae gray-
ish brown, orange below. Thorax: With a wide, metallic gray, median
vitta and a similar sublateral one. Between these vittae are others which are
dark, golden-brown or brassy, and in some lights themselves appear to be
divided. Scutellum metallic black, the posterior part of the rim narrowly
yellow. Humeri, the posterior part of mesopleurae, and the upper part of
sternopleurae yellow. Abdomen: Black marked with yellow, the first seg-
ment largely black, the anterior corners and a narrow anterior margin yel-
low, second segment black with a pair of small oval yellow spots, rather
widely separated in the middle of the segment and a quite narrow lateral
margin reaching nearly to the posterior corners also yellow. Third segment
with a pair of widely separated yellow basal spots, which on their median
ends are widely produced in a posterior direction to just past the middle of
the segment and at the same time slightly curving toward the midline.
These posterior ends are rounded and the lateral, basal portion of the spot
reaches to the lateral margin of the segment where it is just a little expanded
posteriorly. Fourth segment with almost identical pattern. Fifth segment
somewhat similar, the lateral portions wider, the median portion about the
same but the two spots more closely approximated. Legs: Pale yellow, the
anterior tarsi pale brown, the hind tarsi and a prominent subapical annulus
on the hind femora black. Hind tibiae with faint brownish black annulae on
either side of the middle. Wings: Hyaline, stigma pale brown.

Types.—Holotype and three paratypes: Females. Salada River, near
Ceiba, Spanish Honduras, September 2, 1938 (F. M. Hull collector). One
additional paratype, female, from Tuxtepec, Mexico, in the U.S. National
Museum.

Baccha flata n. sp.

Male.—Length 14 mm. Head: Face, cheeks, and front pale yellow, the
front with a rhomboid, blackish spot of some size in the center, confluent
with a smaller black spot just before the antennae and on each side of which
are brownish triangles touching the eye margins. Pile of front and face erect,
thick, and blackish, somewhat paler below. Antennae orange-brown, darker
upon the upper part of the third joint, antennal pile black. Thorax: With
four, dully shining, brassy black vittae upon a brownish-yellow background;

434 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

the sublateral vittae widest, interrupted at the suture; the submedian vittae
connected at a spot opposite the suture and giving the impression of a nar-
row H. Between the several vittae the thorax is golden-pollinose. Thoracic
pile thick, erect and dark brown, becoming reddish or yellow along the
lateral margins of the mesonotum. Humeri and practically the whole of the
pleurae, except for a tiny spot below the base of the wing, light yellow.
Scutellum light yellow with a subtranslucent, transverse fascia. Abdomen:
Chiefly brownish orange, flattened and spatulae, the first segment pale yel-
low except for a narrow posterior margin, the second segment with a narrow
yellow posterior margin and a pair of prominent, hyaline, rectangular win-
dows in the anterior corners. Third segment wholly orange except for a
narrow subapical, narrowly attenuated black fascia, interrupted in the
middle and not reaching the sides. Fourth segment similarly colored. Fifth
almost.wholly orange. Legs: Pale brownish orange, more yellow upon the
anterior pair. Hind femora with thick, very long, shaggy, reddish pile and
some black pile at its base and on hind trochanters and coxae. Wings:
Hyaline, with a sharply marked brown band along the anterior margin.
Holotype.—Male. Petropolis, Brazil, March 1, 1914.

Salpingogaster halcyon n. sp.

Male.—Length 17 mm. Head: Cheeks and face and upper part of front
pale yellow. Greater anterior portion of front is black, expanded triangularly
to the sides and to the eye margin. Antennae dark brown, first joint nearly
black. Thorax: Obscurely shining black with three obscure, dark, reddish-
brown vittae on the anterior two-thirds, and viewed from behind a more
or less square area covered with pale yellowish-gray pubescence that is
rather prominent. Pleurae black. The humeri, propleurae, notopleurae, a
wide conspicuous oblique cuneiform stripe upon the posterior part of the
mesopleurae, upper sternopleurae, anterior pteropleurae, and a second spot
upon the anterior part of the metapleurae all pale yellow. Scutellum pale
yellow with a transverse brown fascia not reaching the sides in the middle.
Abdomen: Slender, with a prominent median black vittae from base of
second segment to apex. First segment pale yellow with a diffuse ight brown
posterior margin. The anterior corners of the second segment and the narrow
sides, as well as the somewhat wider sides of the third segment are pale
yellow but diffuse. The apical portion of the sides of the third segment as
well as the fourth segment reddish brown. Legs: Light yellow, the posterior
tarsi, apical two-thirds of posterior tibiae, base and apical annulus of hind
femora obscurely blackish or dark brown. Pile of anterior legs yellow, except
on fore tarsi; black on the entire hind legs. Wings: Pale brown, strongly
yellow on the basal half anteriorly, the stigmal cell very dark brown, the
marginal cell somewhat lighter brown.

Holotype-—Male, Barro Colorado Island, August 28, 1938 (F. M. Hull
collector).

Oct. 15, 1940 CREASER: A NEW SPECIES OF PHYLLOPOD 435

ZOOLOGY.—A new species of phyllopod crustacean from Stone Moun-
tain, Georgia.1 Epwin P. Creasmr, U. S. Fish and Wildlife Ser-
vice. (Communicated by Waupo L. ScumiIrTT.)

On June 8, 1939, I was informed by Don E. Eyles that certain fairy
shrimps were occurring in rain pools on the summit of Stone Moun-
tain in DeKalb County, Ga. Mr. Eyles had been requested to be on
the watch for these crustaceans, as he was studying these pools and
a few specimens new to science had previously been found at this lo-
cation. Accordingly, on June 9, 1939, I climbed to the summit of the
mountain, amid a rain and hail storm, and obtained a fine series of
these phyllopods.

These crustaceans inhabited shallow rock pools 3 to 6 feet in di-
ameter with fine silt bottoms, occurring in water not exceeding 4
inches in depth. They were found in company with a bloodworm (chi-
ronomid larva?) and a species of Hulimnadia (Phyllopoda: Concho-
straca), which is possibly also undescribed and which is being studied
by Prof. J. G. Mackin. These phyllopods prove to belong to the genus
Chirocephalus, another species of which is widespread throughout
Europe. The genus has not previously been recorded from North
America. The species, Chirocephalus lithacus,? is here described for the
first time.

Dr. A. 5S. Pearse very kindly arranged the loan of some microscopic
equipment needed during the course of this study.

BRANCHIPODA
PHYLLOPODA: ANOSTRACA
Family CHIROCEPHALIDAE
Chirocephalus lithacus, new species

Description.—Male: Total body length of adult, 8.3 to 10.5 mm. Frontal
appendage rising from middle of head as a single organ, then dividing into
two similar branches, each branch in turn terminating in two lateral
branches, one of which is club-shaped and armed at apex with a stout spine
and studded over entire surface with papillae of probable sensory nature;
the other branch laminate, with fingerlike weak spines along margins. Each
branch of frontal appendage about 3 mm long. Clasping antenna sickle-
shaped; basal article half length of terminal article. Inner margin of terminal
article with fine oblique grooves. First antennae shorter than terminal
article of clasping antennae. Eyes stalked, with greatest diameter less than
0.5 mm. Abdominal segments consisting of genital-bearing segment, 7 post-
genital segments, and cercopods. Cercopods uniformly tapering, with setae
of approximately one-fourth total length of cercopod. Swimming appendages
with one branchial lamina.

1 Received June 5, 1940.

2 The name lithacus was suggested for this animal by Dr. Carl L. Hubbs. Translated
from the Greek, it means of a stony place.

436

Fig. 3.—Male cercopods. Fig. 4.—Front view of head of male. Fig. 5.—Front view
of head of female. Fig. 6.—Papillae of frontal appendage of male.

JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

1mm. 5 No mm. &

_ SS ee

Fig. 1.—Frontal appendage. Fig. 2.—Clasping or second antenna of male.

Oct. 15, 1940 ROTHSCHILD: A NEW TREMATODE 437

Female: Total body length of adult, 10.5 to 11.1 mm. Frontal appendage
absent. First antennae about as long as clasping antennae. Clasping an-
tennae flattened, pointed at apex, uniformly rounded on margins. Eyes
stalked, with greatest diameter less than 0.5 mm. Ovisac one-half as long
as abdomen with cercopods excluded. Ovisae gourd-shaped, not uniformly
tapering, with apex turned abruptly toward the abdomen. Cercopods uni-
formly tapering, with setae of approximately one-fourth total length of
cercopod.

Remarks.—This species may be distinguished from other North American
phyllopods by the shape and structure of the bilaminate male frontal ap-
pendage and by the sickle-shaped appearance of the clasping antennae. _

Types.—I collected the types of this new species in temporary pools in
the granite rock on the summit of Stone Mountain, DeKalb County, Ga.,
within 100 yards of the airway beacon, June 9, 1939. They have been de-
posited in the United States National Museum: Holotype male, no. 79294;
allotype female, no. 79295.

LITERATURE CITED

Creaser, E. P. Division of Phyllopoda, in Pratt, H. 8., Manual of the common in-
vertebrate animals, pp. 373-381. Philadelphia, 1935.

Dapay, E. Monographie systematique des phyllopodes anostraces. Ann. Sci. Nat.
(9) 11:213. 1910.

ZOOLOGY .—Cercaria pricei, a new trematode, with remarks on the
specific characters of the ““Prima’’ group of Xziphidiocercariae.}
Miriam RotHscuHiItp, London, England. (Communicated by
E. W. PRIcE.)

It is not an unusual phenomenon to discover several very closely
related species of larval trematodes parasitizing the same host. Cer-
caria pricei n. sp., described in this paper, is the third species of the
“Prima”’ subgroup of Cercariae ornatae (Lithe, 1909) to be found in
the snail Pseudosuccinea columella Say. Although this type of cer-
caria was originally recorded from Europe, almost all the important
experimental work on their life histories has been carried out in bril-
liant fashion by Krull (1931, 1933) and other workers in the United
States.

The individual author’s spelling of host names is used herein.
Furthermore the generic and specific names of the adult fluke used by
the author in his description of its cercaria and life history are re-
tained to avoid confusion.

I should like to express my gratitude to Dr. E. W. Price for his
kindness in affording me every facility for work during my unexpected
visit to Washington at the outbreak of the European war, to the Spen-

1 Received June 3, 1940.

438 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

cer Lens Company for lending me superlative microscopical appara-
tus, and to Dr. J. P. E. Morrison and Countess Mary Roberti for as-
sisting in the collection of specimens.

CHARACTERS OF THE ‘‘PRIMA’’ SUBGROUP
OF CERCARIAE ORNATAE

Sewell (1922) recognized that certain species of “‘Ornatae’’ Xiphi-
diocercariae formed a compact natural group of which Cercaria prima
Sinitzin, 1905, was selected as the ‘“‘type.” Sewell’s definition can be
emended on several unimportant points. He regarded only the stem
of the Y-shaped excretory bladder as the bladder proper and de-
scribed the crura as dilatations of the excretory tubes. The subse-
quent development of the bladder in the metacercaria shows that this
is erroneous. Two species with six pairs of penetration glands have
also been discovered since 1922.

McMullen (1937) has shown that the presence or absence of a tail
fin-fold may not be a character of great importance when forming a
natural classification within a large group like the Plagiorchioidea.
With this opinion I am in entire agreement. In the matter of tail
structure and modification or loss of fins there is repeated evidence of
parallel evolution in distantly related genera within one superfamily.”
McMullen writes:

In the cercariae of the frog lung flukes and related forms a similar situa-
tion arises. The cercariae of these trematodes... all have a fin-fold on the
tail except the cercaria of Haplometra cylindracea. Here the absence of the
fin-fold would, in the present classification, eliminate this form from the
Ornatae group of Xiphidiocercariae and thus separate it from closely re-
lated forms. It seems probable, therefore, that the fin-fold of the tail and
other such larval modifications are of little more than specific value in the
Xiphidiocercariae.

However, the adult forms of ‘‘Prima”’ cercariae have all proved to be
frog lung flukes of the subfamily Haematoloecinae Freitas and Lent,
1939 (=Pneumonoecesinae Mehra, 1937). Haplometra cylindracea
Zeder is not in my opinion closely related to this subfamily, and in the
opinion of Mehra (1937) it pertains to another family altogether. The
absence of a tail membrane in this cercaria is therefore not considered
to have any special bearing on the subgroup now under consideration,
and the presence of a ‘“‘Prima’’ type of fin-fold is for the present re-
tained as a diagnostic character.

2 Another type of parallel evolution to be found in the tail structure of cercariae is
worthy of note. This may be typified by the giant tails found in unrelated cercariae
which are swallowed as free swimming organisms by fish, for example, Petasiger nitidus
Linton, 1928 (Echinostomidae) and Proterometra macrostoma (Faust, 1918) (Azygiidae).

Oct. 15, 1940 ROTHSCHILD: A NEW TREMATODE 439

4

100 u

40

mY :

Fig. 1.— Diagram of Cercaria pricet from ventral aspect with tail contracted (pene-
tration glands shown slightly larger than in reality. Fig. 2.—Fin-fold under pressure
of a coverslip. Fig. 3—Stylet. Fig. 4—A-—F, Diagrams showing appearance of ex-
cretory bladder during different stages of contraction.

440 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10 —

THE “‘PRIMA’’ SUBGROUP SEWELL, 1922 (EMENDED)

(1) Distome Xiphidiocercariae of medium size in which the acetabulum
is situated behind the middle of the body length.

(2) The tail is shorter than the body and is furnished with a dorsoventral
fin-fold in its distal portion; the ventral portion of the fin extends farther
forward than the dorsal portion.

(3) The alimentary canal possesses a prepharynx and a pharynx, and the
intestinal caeca reach back to a point between the margin of the acetabulum
and the posterior end of the body.

(4) Four to six pairs of penetration gland cells are present.

(5) The excretory bladder is Y-shaped and does not extend beyond the
lateral margins of the ventral sucker. The main excretory tubes enter the
bladder at the extremity of the arms of the Y. They divide into anterolateral
and posterolateral branches at the level of the ventral sucker. The excretory
pattern is 2[(1-+1+1)+(1+1-+1)]=12 or 2[(8+8+3)+ (84+3-+3)] =36.

(6) Development occurs in small unbranched sporocysts.

(7) The first intermediate hosts are fresh-water gastropod mollusks, and
the second intermediate hosts are arthropods.

Thirteen species may with certainty be included in this group:

Cercaria prima Sinitzin, 1905.

Cercariae indicae XXIV Sewell, 1922.

Cercariae indicae X XVIII Sewell, 1922.

Cercaria longistyla McCoy, 1929.

Cercaria prima Dubois, 1929 (nec Sinitzin, 1905).

Cercaria of Pneumonoeces medioplexus (Stafford, 1902) Krull, 1931.

Cercaria of Pneumobites pariiplexus (Irwin, 1929) Krull, 1931.

Cercaria prima Wesenberg-Lund, 1934 (nec Sinitzin, 1905, nec Dubois,
1929).

Cercaria of Haematoloechus complexus (Seely, 1906) Krull, 1933.

Cercaria of Ostiolum oxyorchis (Ingles, 1932) Ingles, 1933.

Cercaria herbert McMullen, 1938.

Cercaria merchants Rankin, 1939.

Cercaria pricet n. sp.

Of these species C. prima Wesenberg-Lund was examined from a
crushed snail only, a fact that undoubtedly accounts for the unde-
veloped state of the intestinal caeca. From time to time various spe-
cies such as Cercaria pela Porter, 1938, C. elemensis Porter, 1938, and
C’. laticauda Wesenberg-Lund, 1934, have been erroneously assigned
to this subgroup. These larvae generally display features that place
them in Lihe’s Cercariae ornatae but that exclude them from the
circumscribed ‘‘Prima”’ subgroup. There are certain other species,
such as Cercaria ornata Lutta, 1934 (nec La Valette, 1855), Cercaria
pratensis Skwortzoff, 1924, and Cercaria prima Ruszkowski, 1925
(nec Sinitzin, 1905), whose exact systematic position must remain
doubtful until more adequate descriptions are available.

441

A NEW TREMATODE

ROTHSCHILD

Oct. 15, 1940

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442 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

SPECIFIC CHARACTERS OF THE ‘‘PRIMA’’ SUBGROUP

Judged from the descriptions by such excellent observers as Krull
and Sewell it is only a little less difficult to distinguish between these
larvae than certain heterophyid, notocotylid, and microphallid cer-
cariae. Up to a point the same difficulties are encountered as those
indicated for Opisthorchioidea cercariae (Rothschild, 1938). Meas-
urements of soft parts can only be regarded as supplementary data,
although in some cases the discrepancy in size is so great that it can
not be due to difference in the technique adopted by individual ob-
servers. Measurement of the stylet is, however, of much importance.
As in other groups, the relative degree of development and transpar-
ency of certain organs, such as sensory setae, cuticular spines, intes-
tinal caeca, reproductive anlage, etc., are specific differences that do
not lend themselves well to written descriptions. Perhaps one of the
most tantalizing characters of this type is the relative thickness or
toughness of the cuticle. The cercaria of Pneumobites parviplexus, for
example, fragments after a few seconds under a coverslip, whereas
C’. pricet n. sp. remains intact for many hours. This is no doubt an ex-
treme case, but in comparing cercariae almost every worker comes
across subtle differences of this type that, although very obvious, can
not be accurately described and at times even defy analysis.

The number and arrangement of penetration glands can vary spe-
cifically, but as the range is small this may not be very helpful (Table
1). The ducts apparently can open at the base or the tip of the stylet
(C. indicae XXVIII, C. indicae XXIV). The shape and color of the
parthenitae, together with the number of cercariae maturing simul-
taneously within each sporocyst, also vary specifically, but the age of
an individual infection and the temperature of the water (Rothschild,
1935) appear to exert some influence upon these characters. Two dif-
ferent excretory patterns are met with. In one the full complement of
flame cells found in the adult fluke is already developed, and in the
other each group is represented by one flame cell only. No more than
three patterns have been completely worked out. The distribution of
sensory setae may be of much importance, but apparently in many
cases these structures are not visible to the ordinary observer. Sinit-
zin undoubtedly possessed superior eyesight and described them in
wonderful detail for nearly all his cercariae. The accuracy of his ob-
servations is verified from time to time when a species is found in
which they are more obvious than in others.

aA)
SH
sH

A NEW TREMATODE

ROTHSCHILD

Ocr. 15, 1940

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444 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES’ VOL. 30, No. 10

Specific descriptions of ‘‘Prima”’ cercariae can be based on the fol-
lowing characteristics:
(1) Size:
(a) Relative length of body and tail.
(b) Relative size and position of suckers, pharynx,? prepharynx, and
esophagus.
(2) Shape and dimensions of stylet (see Table 1).
(3) Precise extent and shape of caudal fin-fold.
(4) Behavior (see Table 2):
(a) Time of emergence from the snail host.
(b) Duration of free swimming life.4
(c) Tropisms.
(d) Choice of second intermediate host (including negative data).
(5) Cyst:
(a) Shape (see Dubois, 1929, p. 56).
(b) Size.
(c) Location.

Supplementary characters:

(6) Excretory pattern.
(7) Number of penetration gland cells.
(8) Distribution of sensory setae.
(9) Presence of sensory papillae.
(10) Sporocysts:
(a) Shape.
(b) Color.
(c) Number of cercariae maturing simultaneously within each
sporocyst.

It is probable that these characters are valid for differentiating be-
tween other subgroups of Cercariae ornatae such as the ‘‘Hemilo-
phura’’ subgroup. These differ essentially from ‘‘Prima”’ cercariae in
the possession of an I-shaped excretory bladder, a fin-fold extending
along the whole or two-thirds of the ventral side of the tail but not
along the dorsal side, and elongated ‘“‘worm’’-shaped sporocysts. Vari-
ation in behavior has also been noted. The cercaria of Macroderoides
typicus emerged “‘in the mid-morning hours... and died within a
few hours’? (McMullen, 1935), whereas the cercariae of Haplometrana
utahensis, ‘“‘are shed throughout the day and night with the greatest
numbers appearing towards the evening... and lived as long as 40
hours’”’ (Olsen, 1937).

3’ The conspicuously large pharynx of the cercaria of Ostiolum oxyorchis is also a
feature of the adult fluke.

4The term “duration of free swimming life’? I understand to mean the period
during which the cercaria is capable of swimming in the water after natural emission
from the snail host (Rothschild, 1938), and not the period during which the cercaria
manifests movement or “signs of life.’”” In most instances the latter character does not
lend itself so well to accurate and clear cut comparison (Dubois, 1929), although it is
useful as supplementary data (Wheeler, 1939).

Oct. 15, 1940 ROTHSCHILD: A NEW TREMATODE 445

Cercaria pricei,® n. sp. Figs. 1-4

Description.—Xiphidiocercaria of the ‘‘Prima”’ type, with the characters
of the group. Measurements in microns (under light pressure of coverslip):
Body length 375 to 257, mean 327; width 149 to 107, mean 135. Tail length
232 to 166, mean 200; width 49 to 41, mean 45. Fin 41 long ventrally, 22
dorsally; maximum width 17. Oral sucker 68 by 68 to 51 by 51, ventral
sucker 44 by 51 to 31 by 41. Pharynx 25 by 22, prepharynx length 11.9 to
15.3; esophagus length 25. Stylet 30 to 40 by 4.2 to 5.1. Sporocyst (maxi-
mum) 249 by 124.

Body regularly oval in outline. Tail when extended two-thirds length of
body, when contracted one-half length of body. Ventral sucker slightly
more than half diameter of oral sucker, situated somewhat less than two-
thirds of body length from anterior extremity. Esophagus short, barely
longer than pharynx. Intestinal caeca well developed. Five pairs of penetra-
tion gland cells of which the posterior three pairs are coarsely granular,
slightly larger, and dorsal to the two anterior pairs. Ducts of glands arranged
in two groups opening near anterior extremity of stylet. Crura of bladder
not reaching posterior border of ventral sucker. Cuticle faintly rugose, with-
out spines or sensory tubercles. Two rows of long sensory setae along
lateral edges of body, arranged as in C. prima. Parthenitae stumpy, sausage-
shaped, pale yellow; one to two cercariae maturing simultaneously within
each sporocyst.

Behavior.—The cercariae emerge throughout the day, the length of free
swimming life being nine hours in tap water at room temperature.

No phototropic responses.

First intermediate host.—Pseudosuccinea columella Say.

Locality and date of collection Shaw Lily Gardens, Washington, D. C.,
September-October, 1939.

Remarks.—Cercaria pricet most closely resembles the Cercaria of
Haematoloechus complexus described by Krull and Cercaria merchants de-
scribed by Rankin, both from the same host. It can be separated from the
former by its much greater size, number and arrangement of sensory setae,
the relatively shorter prepharynx, and greater distance between the fork
of the intestine and the ventral sucker. In C. merchanti the forks of the
Y-shaped bladder extend to the lateral borders of the acetabulum; the
long sensory setae are absent but sensory tubercles are present around the
oral sucker; the stylet is narrower in comparison with its length; and there
are also differences in behavior (see Table 2).

- One of the most striking features of C’. pricez is the periodic contractions
of the bladder. First the cavity of the stem is obliterated and then the
branches of the Y. This curious phenomenon is not mentioned by either
Krull or Rankin, but Ingles (1933) has noted it for the Cercaria of Ostzolum
oxyorchis, and Miller (1935) and Dubois (1929) record it for ‘‘Polyadena”’
cercariae. Krull’s drawing of the bladder of the Cercaria of H. complexus
does not entirely agree with the description, and in fact the bladder figured
for Cercaria tricystica Miller, 1935, C. helvetica V Dubois, etc., more closely

5 Named for Dr. E. W. Price, of the U. S. Bureau of Animal Industry, in honor of
his many contributions to our Knowledge of the Trematoda.

446 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

resembles that of C. price: than several of those figured for ‘‘Prima’”’
cercariae!

It has already been mentioned that the ducts of the penetration glands
are arranged in bundles. They are so closely approximated that anterior to
the esophagus there appear to be only two ducts and two apertures on each
side. It sometimes happens that one of the posterior pairs of glands, which
are situated dorsally, presents the finely granular appearance characteristic
of the anterior ventrally situated pairs (see also Sewell, 1922, p. 222). A
large spherical nucleus and prominent nucleolus are conspicuous in each
penetration gland cell. |

Unlike C. merchanti, C. pricet does not collect on the lighted side of the
vessel and in fact shows no phototactic responses whatsoever. At times it
manifests a definite tendency to accumulate near the surface. At other times,
however, the cerceriae remain distributed at all levels of the water or segre-
gate in the lower levels. It seems probable that the nature of their geotropic
response is correlated with temperature changes as suggested by Krull
(1931) for cercariae of P. medioplexus and P. parviplexus. C. pricet swims
continuously from the time of emergence, with occasional momentary pauses
when the body is suddenly extended horizontally. There is also a slight tend-
ency to swarm around small objects in the water, particularly the snail
host, but this again seems to depend on certain unknown factors, as it is
manifested on some days and not on others.

One very peculiar habit is typical of this cercaria. A fairly large number
were kept in glass tumblers, and while swimming they quite frequently came
into contact with the sides of the vessel. They then attached themselves
and crawled about on the glass by the usual ‘‘leeching’’? movements. At
death they did not drop off and fall to the bottom of the container but be-
came fully extended and remained flattened against the glass like a series of
permanent preparations. Water beetles, dragonfly nymphs, and other insect
larvae were fished from the same pond and introduced into the jars with the
cercariae, but penetration was not observed.

EFFECT ON THE SNAIL HOST

The effect of trematode parasites on the ultimate size and growth
rate of the molluscan host and the shape of its shell is probably far
more universal than is generally recognized. Kelly (1899) apparently
first drew attention to this phenomenon in the United States when
he recorded change of shell shape in Unionidae due to castration by
Bucephalus cercariae. He writes:

From the shape of the shells eight of these thirty-one individuals (Gnfected
with cercariae) were pronounced males and two females. Others, also belong-
ing to species in which the shells of the two sexes are normally characteristic,

had shells of such shape as to render the sex problematical and to suggest
that infestation by Bucephalus or other cercariae, when early acquired and

Ocr. 15, 1940 ROTHSCHILD: A NEW TREMATODE 447

long continued, may so alter the form of the shell of the female as to cause
it to resemble that of the male or, if acquired later, may produce an inter-
mediate form.

Rankin (1939) found slight abnormalities of the shell of Pseudo-
succinea columella infected with echinostome cercariae, but “‘little or
no effect on the snail host from infection with Xiphidiocercariae was
observed.”

The snails harboring C. pricez n. sp. were collected from a tiny iso-
lated pond about 4 by 3 feet, the surface of which was entirely cov-
ered with vegetation. A certain number of relatively large specimens
of P. columella were found resting on the upper side of waterlily
leaves, exposed to direct sunshine even during the hottest period of
the day (100° F. in the shade). All these proved to be infected with
Xiphidiocercariae. No uninfected snails were found in this unpro-
tected situation; there was thus a marked difference in the behavior
of parasitized snails.

It is believed that all the mollusks® present in this pond were col-
lected and examined. Only those that could have passed through a
fine-mesh sieve might have escaped notice. The mean height of un-
infected P. columella was 3 mm, the maximum size being 4 mm. In-
fected specimens reached 10 mm in height, with a mean of 9.75 mm.
It is not suggested that parasitism is responsible for this great con-
trast in size, but it might well prove a contributing factor. Without a
fairly prolonged study of the snail population of any given area it
would be impossible to come to any definite conclusion on this point.

BIBLIOGRAPHY

Dusotis, G. Les cercaires de la région de Neuchatel. Bull. Soc. Neuchatel Sci. Nat.
5354-177. 1929.

Freitas, J. F.T., and Lent, H. Consideracoes sobre algu as especies americanas don
genero Haematoloechus Looss, 1899 (Trematoda: Plagiorchoidea). Livro Homena-
gem Profs. A. e M. Ozorio de Almeida, pp. 246-256. 1939.

IneiEs, L. G. Studies on the structure and life-history of Ostiolum oxyorchis (Ingles)
from the California red-legged frog Rana aurora draytoni. Univ. California Publ.
Zool. 39: 135-161. 1933.

Kewiy, H. M. A statistical study of the parasites of the Unionidae. Bull. Illinois Lab.
Nat. Hist. 5: 399-418. 1899.

Kruuu, W.H. Life-history studies on two frog lung flukes, Pheumonoeces medioplexus .-
and Pneumobites parviplexus. Trans. Amer. Micr. Soc. 50: 215-277. 1981.

. Studies on the life-history of a frog lung fluke, Haematoloechus complexus

(Seely 1906) Krull, n. comb. Zeitschr. Parasitenk. 6: 192-206. 1933.

. Some additional notes on the life-history of a frog lung fluke Haematoloechus
complexus (Seely, 1906) Krull. Trans. Amer. Micr. Soc. 53: 196-199. 1934.

Line,M. Parasitische Plattwiirmer:I.Trematoda. Siisswasserfauna Deutschlands 17:
1-217. 1909.

Lutrta, A. Die Fauna der parthenogenetischen Trematoden-generationen in den Siiss-
wassermollusken Peterhofs. Trav. Soc. Nat. Leningrad (Sect. Zool.) 63: 261-309.
1934.

6 Four species, all of which were identified by Dr. J. P. E. Morrison, to whom I
tender my best thanks.

WN

448 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

eres O. R. Notes on cercariae from Missourt. Journ. Parasit. 15: 199-208.

] :

McMoutten, D. B. The life histories and classification of two allocreadiid-like plagi-
orchids from fish, Macroderoides typicus (Winfield) and Alloglossidium corti
(Lamont). Journ. Parasit. 21: 369-3880. 1935.

. A discussion of the taxonomy of the family Plagiorchiidae Liihe, 1901, and re-

lated trematodes. Journ. Parasit. 23: 244-258. 1937.

. Notes on the morphology and life cycles of four North American cercariae.
Livro Jubilar L. Travassos, Rio de Janeiro, pp. 299-306. 19388.

Meura, H. R. Certain new and already known distomes of the family Lepodermatidae
Odhner (Trematoda), with a discussion of the classification of the family. Zeitschr.
Parasitenk. 9: 429-469. 1937.

Miuuter, E. L. Studies on North American cercariae. Univ. Illinois Bull. 33(35)
[Illinois Biol. Monogr. 14(21)]: 1-125. 1936.

OusEeNn, O. W. Description and life history of the trematode Haplometrana utahensis
sp. nov. (Plagiorchidae) from Rana pretiosa. Journ. Parasit. 23: 13-28. 1937.

Porter, A. The larval Trematoda found in certain South African Mollusca, with
special reference to Schistosomiasis (Bilharziasis). South African Inst. Publ.
Med. Res. 8(42): 1-492. 1938.

RANKIN, J. 8. Ecological studies on larval trematodes from western Massachusetts.
Journ. Parasit. 25: 809-328. 1939.

RorTuscHitp, M. The trematode parasites of Turritella communis Lmk. from Plymouth
and Naples. Parasitology 27: 152-170. 1935.

. The excretory system of Cercaria coronanda n. sp. together with notes on tts life-
history and the classification of cercariae of the superfamily Ophisthorchioidea Vogel
1934 (Trematoda). Novit. Zool. Tring. 41: 148-163. 1938.

Ruszkowskl, J. 8S. Materialy do fauny helmintologicznej Polski. Spraw. Komis.
Fisjograf. Polsk. Akad. Krakow 60: 173-185. 1925.

SEWELL, R. B.S. Cercariae indicae. Indian Journ. Med. Res. 10(suppl. no.): 1-370.
1922.

Sinitzin, D. F. Distomes of fishes and frogs in the vicinity of Warsaw. Materials for
natural history of trematodes, pp. 1-210. Varshava, 1905.

Skwortzorr, A. H. Materialien zur Fauna der Larvenformen von in den Mollusken
der Wolga und Wetluga lebenden Trematoden. Arb. Biol. Wolga-Station Saratow
7(4—5): 201-211. 1924.

WESENBERG-LUND, C. Contributions to the development of the Trematoda Digenea.
Part II. The biology of the freshwater Cercariae in Danish freshwaters. Danske
Vidensk. Selsk. Skr. (ser. 9) 5: 1-228. 1934.

WHEELER, N. C. A comparative study on the behavior of four species of pleurolopho-
cercous cercariae. Journ. Parasit. 25: 343-353. 1939.

PROCEEDINGS OF THE ACADEMY AND
AFFILIATED SOCIETIES

THE ACADEMY

361ST MEETING OF THE BOARD OF MANAGERS

The 361st meeting of the Board of Managers was held at the Cosmos Club
on Friday, May 24, 1940. There were 20 present as follows:

E. C. CritTENDEN A. T. McPHERSON EK. W. Prict
F. D. Rossini A. WETMORE R. R. SPENCER
F. C. Kracex C. THom C. L. GARNER
H. 8S. RaPpplEYE W. A. Dayton C. L. Gazin

J. H. HispBEN P. C. WHITNEY W. W. Dieei
G. STEINER H. L. Curtis and by invitation
F. M. SETZLER W. RAMBERG J. H. Kempton

Acting upon instructions from the Board, President Crittenden appointed
the following committees:

(1) To consider societies qualified for affiliation with the Academy:
F. M. Serzumr, chairman; R. E. Grsson and R. R. SPENcER, members.

Oct. 15, 1940 PROCEEDINGS: THE ACADEMY 449

(2) On awards for scientific achievement: A. H. Cuark, general chair-
man; Physical Sciences, O. R. Wuur, chairman; Engineering Sciences,
F. M. Drranporr, chairman; Biological Sciences, A. H. CuarxK, chairman.
The other members of these committees are to be appointed later.

(3) To conserve the archives of the Academy: N. R. Smitu, chairman;
F. D. Ross1n1, member.

The following persons, whose names were presented to the Board by the
Committee on Membership, were elected to membership:

Resident Members

OLAF SVERRE AAMODT, principal agronomist, in charge of the Division of
Forage Crops and Diseases, U. 8. Bureau of Plant Industry, in recognition
of his work on cereal pathology, particularly with respect to breeding for
disease resistance, and also of his studies on forage grasses and legumes.

JAMES KENNETH ABLEITER, senior soil technologist, Division of Soil Sur-
vey, U.S. Bureau of Plant Industry, in recognition of his work in land-use
planning and in productivity ratings of mineral soils.

CHARLES WALTER Bacon, physiologist, Division of Tobacco and Plant
Nutrition, U.S. Bureau of Plant Industry, in recognition of his contributions
to various aspects of the physiology of tobacco plants, nutrition, etc.

Mark BauLpwin, senior soil scientist, Division of Soil Survey, U. S.
Bureau of Plant Industry, in recognition of his outstanding work on the
genesis and classification of soils and on the coordination of federal and state
soil surveys.

EpwArRD EASTMAN CLAYTON, senior physiologist, Division of Tobacco
and Plant Nutrition, U. S. Bureau of Plant Industry, in recognition of his
contributions on the diseases of tobacco and other plants.

PAUL SYDNEY CONGER, research associate, Carnegie Institution of Wash-
ington, and custodian of diatoms, U. 8. National Museum, in recognition
of his studies on diatoms and their economic use and his discovery of im-
portant commercial deposits of diatoms in Wisconsin.

Henry TRENDLEY DEAN, dental surgeon, Division of Infectious Diseases,
National Institute of Health, in recognition of his work on the relationship
of fluorine to dental caries.

SAMUEL LEONARD EMSWELLER, principal horticulturist, in charge of Flori-
culture and Ornamental Horticulture Plant Investigations, Division of Fruit
and Vegetable Crops and Diseases, U. S. Bureau of Plant Industry, in
recognition of his contributions to horticulture and the cytology and genetics
of ornamental plants.

EARL STEINFORD JOHNSTON, assistant director, Division of Radiation and
Organisms, Smithsonian Institution, in recognition of his studies on radiation
in connection with plant life.

CHARLES EDWIN KELLOGG, principal soil scientist and chief of the Di-
vision of Soil Survey, U. 8S. Bureau of Plant Industry, in recognition of his
work on the genesis and classification of soils and on rural land-use plan-
ning.

ConsTANTIN NIKIFOROFF, soil scientist, Division of Soil Survey, U. S.
Bureau of Plant Industry, in recognition of his work on dynamic processes
in soils, on frozen soils of Siberia, on desert types of soil formation, and on
the great soil zones of western North America.

GLENN LANE Parker, chief hydraulic engineer and chief of the Water
Resources Branch, U. 8. Geological Survey, in recognition of his work in
hydrology, particularly on water power and snow surveys in the Cascades
and on water resources as related to placer mining.

450 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

EUGENE ALFRED SIEGLER, pathologist, Division of Fruit and Vegetable
Crops and Diseases, U. 8. Bureau of Plant Industry, in recognition of his
coutalbareu to phytopathology, particularly his researches on crown
gall.

JAMES THORP, soil scientist, Division of Soil Survey, U. S. Bureau of
Plant Industry, in recognition of his work on the study and classification of
the soils of China.

JACOB OSBORN WARE, senior agronomist, Division of Cotton and other
Fiber Crops and Diseases, U. 8. Bureau of Plant Industry, in recognition
of his work in cotton breeding and genetics and his contributions to agron-
omy.

Nonresident Members

EKUBANKS CARSNER, senior pathologist, Division of Sugar Plant Investiga-
tions, U. 8. Bureau of Plant Industry, Riverside, Calif., in recognition of his
work in the field of virus diseases in plants.

JoHN NATHANIEL Coucu, professor of botany, University of North
Carolina, Chapel Hill, N. C., in recognition of his contributions to mycology,
particularly on the genus Septobasidium.

GorDOoN Enocu Gates, professor of biology and head of the Department
of Biology, Judson College, Rangoon, Burma, India, in recognition of his
special researches on terrestrial oligochaetes.

Moses Ben Naputaui LEVINE, pathologist, Cereal Disease Investiga-
tions, U. S. Bureau of Plant Industry, University Farm, St. Paul, Minn.,
in recognition of his work on cereal rusts and smuts, on physiologic special-
ization and generic nature of rust pathogenes, and on environment and pre-
disposition of grain crops.

MavricE Buoop LINForD, pathologist, Experiment Station, Pineapple
Producers Cooperative Association, Honolulu, Hawaii, in recognition of his
work in plant pathology, especially his studies on diseases of peas and pine-
apples, on plant-parasitic nematodes, on nematode biology and populations,
and on natural enemies of nematodes.

ARTHUR SPERRY PrEARSE, professor of zoology, Duke University, Durham,
N. C., in recognition of his researches in zoology and ecology.

LOWELL Fitz RANDOLPH, chief in research, New York Experiment Sta-
tion, Geneva, N. Y., in recognition of his work in cytology of chlorophyll
types in maize, morphology of developing kernel in maize, chromosomes in
maize and related genera, and induced polyploidy in maize and other plants.

WILLIAM JAcOoB Rossrns, director, New York Botanic Garden, and pro-
fessor of botany, Columbia University, New York, N. Y., in recognition
of his work on plant nutrition, on enzyme secretion by fungi, on tissue cul-
ture with higher plants, and on growth substances.

ELVIN CHARLES STAKMAN, professor of plant pathology, University of
Minnesota, and agent of the U. 8. Department of Agriculture, University
Farm, St. Paul, Minn., in recognition of his contributions to phytopathology
particularly on rust diseases of cereals.

JAMES ZETEK, associate entomologist, U. S. Bureau of Entomology and
Plant Quarantine, and director, Barro Colorado Island Biological Labora-
tory, Institute for Research in Tropical America, Balboa, Canal Zone, in
recognition of his researches in entomology, his services as a consultant in
entomology to certain Latin-American republics, and his management of the
Barro Colorado Island Laboratory.

Percy WHITE ZIMMERMAN, plant physiologist, Boyce Thompson Insti-
tute for Plant Research, Yonkers, N. Y., in recognition of his work in plant

physiology.

Oct. 15, 1940 PROCEEDINGS: BOTANICAL SOCIETY 451

The Committee on Revision of the Bylaws, H. L. Curtis, chairman,
C. THom and P. C. WHITNEY, members, recommended the following state-
ment on the ballot to be issued on the restoration of the offices of nonresident
vice-presidents:

In December 1939 the Academy adopted certain amendments to its bylaws, one
group of which had for its purpose the elimination of the two nonresident vice-presi-
dents of the Academy. The section of the bylaws dealing with amendments provides
that any three members of the Academy can propose an amendment and that the
Board of Managers shall submit this proposal to the members before the next annual
meeting. The Board received a petition on January 17, 1940, signed by six members,
proposing changes to the newly adopted bylaws, the purpose of which is to restore the
two non-resident vice-presidents. Petitioners claim that the provision of two non-
resident vice-presidents as members of the Board of Managers is a proper representa-
tion of the nonresident members of the Academy. As required by the bylaws, the
Board of Managers submits the following to the Academy for action:

Amend Article 2, Section 1, by inserting in the fourth line, “‘and two vice-presidents
chosen from nonresident members.” Article 2, Section 3, insert “‘resident’”’ before vice-
president in the fifth line. Article 5, Section 1, insert in the fifth line after the word
president “‘two persons for the nonresident vice-presidents.”

In favor of amendment

Opposed to amendment
To be valid, all ballots must be received by the Secretary within two weeks from
date at which they were mailed.

It was moved and carried that this proposed amendment, with the pre-
ceding statement of explanation, be submitted for balloting by letter to the
members of the Academy at the time of the distribution of the notices for
the next regular meeting of the Academy.

The Corresponding Secretary announced the following deaths:

Cyrus ADLER
Tuomas A. GROOVER

The Board adjourned at 9:42 P.M.

BOTANICAL SOCIETY

295TH MEETING

The 295th regular meeting was held in the assembly hall of the Cosmos
Club, February 7, 1939, President CHARLES THom presiding; attendance 95.
-K. D. Doak, Hata DeRMEN, and DanrieL ReEapy were elected to member-
ship.

Notes and reviews.—M. B. Waits called attention to the winter injury
that had occurred on the bamboos growing near the Henderson castle and
also on Japanese honeysuckle. He believed this was due to the weather at
Thanksgiving time when, after a heavy rain, the temperature fell to 17° fol-
lowed by snow and sleet. He also displayed a very old book, Twaite’s Early
western travels, vol. 3, 1748-1846, which contains ‘‘The Travels of André
Michaux.”

Program.—G. F. Gravatt: Effect of epidemic tree diseases on the food sup-
plies of animals (lantern).—Chestnut blight, Phytophthora root rot of Cas-
tanea species, Cephalosporium wilt of persimmon, a nectria disease of beech,
and willow diseases, which threaten American forests and woodlands, are

452 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

examples of rapidly spreading, supposedly introduced diseases that reduce
materially food supplies of wildlife. Diseases reducing winter foods are most
serious. One minor objective of the chestnut breeding and selection program
is a strain combining small nuts, productiveness, and resistance to blight,
which has destroyed nearly all commercial chestnut growth. Species that re-
place those killed may supply considerably less wildlife food. Even though
white-pine seeds are not of outstanding value as wildlife food, the introduced
white-pine blister rust has changed the food habits of some wildlife. When
food is scarce some animals eat the young blister rust cankers; and other
wildlife probably find species replacing Azbes eradicated to protect white
pines no less valuable as sources of summer food. Diseases that do not kill
whole trees may also decrease the food supply of wildlife. Probably our for-
ests have so far been exposed to relatively few of the tree-disease organisms
of other continents.

Father ARTHENE DU TILLY: Botanical exploration in the Canadian Arctics
(lantern).—Father pu TiLuy paid a tribute of admiration and gratitude to
his predecessors in the exploration and conquest of the Arctic. Touching
rapidly on the habitat of the Eskimos, he gave more consideration to the
Arctic flora, showing its supreme importance in the life of men and animals,
since both live mainly on plants. He compared this vegetation to that of the
deserts and explained their physiognomic differences by conditions incident
to sunlight, water, wind and snow. He told of the edible roots and berries,
the medicinal properties of the greens, the different lichens used as fuel or
food and fodder for animals; also of plants which by their variety and rich-
ness of coloring adorn the Boreal world. In concluding, Father pu TILLy ex-
pressed the wish that more scientists may continue in the footsteps of those
pioneers in the discovery of the treasures of the polar regions.

296TH MEETING

The 296th regular meeting was held in the assembly hall of the Cosmos
Club, March 7, 1939, President CHarLES THom presiding; attendance 90.
Frep A. ABrcG, Cari O. Grassi, R.A. STEINGBERG, and JOSEPHINE
Ross were elected to membership.

Notes and reviews.—M. B. WaitTE showed some specimens of a rare moss,
Buxbaumia aphylla, which he and A. V. Smirx collected on his farm near the
Patuxent River in Maryland.

Program—DorotHy BuaisDELL: Variability studies in the tomato Fu-
sarium wilt organism; cultural studies.—Thirty geographical strains of Fu-
sarium bulbigenum var. lycopersicz were collectedfrom widely separate regions
of the United States. These were grown on many different types of solid,
liquid and tissue media to determine range of variability, and constancy of
cultural characters. The largest number of isolates were divided into two
main cultural types: the white, raised sclerotoid forms (RS), and the vina-
ceous purple, slimy, appressed form (A). Types intervening between these
occurred: a group with dark undermats covered with coarse water-soaked
mycelium of intermediate raised appearance (IR), and an intermediate ap-
pressed group with dark undermats having light fibrilous aerial growth (1A).
A raised type was also found (R) similar to RS but without sclerotia and
of more stable character. The raised and appressed strains could easily be
distinguished on all types of media.

Sectors often appeared in the RS forms. Transfers from these yielded the
appressed, slimy forms or intermediate types. Single sporing of the raised
sclerotoid forms also yielded both appressed and raised types.

Ocr. 15, 1940 PROCEEDINGS: BOTANICAL SOCIETY 453

F. L. Weviman: Variability studies in the tomato Fusarium wilt organism;
pathogenicity studies (lantern).—Studies of 30 tomato wilt Fusarium isolates
obtained from diverse regions of the United States showed all, except one,
to be Fusarium bulbigenum var. lycopersici. These cultures were readily
separable on the basis of growth appearances into five types, which were
found to be correlated with pathogenicity variations observed on tomato
varieties susceptible, tolerant, and resistant to Fusarium wilt.

Cultures with raised (R) light-colored mycelium were most virulent;
raised types with sclerotial-like bodies (RS) were erratic but next in patho-
genicity; intermediate raised (IR) types were somewhat lower in patho-
genicity; intermediate appressed (IA) were next to the least pathogenic
type; and the dark colored completely appressed (A) cultures with no aerial
mycelium were the least effective pathogenically. Data frominfection studies
on the three tomato varieties showed marked divergences in intensity of
pathogenic effects due to differences in relative disease resistance.

C. O. ERutanson: The first South American Meeting of Botanists, October
12-19, 1988. ;

Ernst J. SCHREINER: Forest tree breeding (lantern).

297TH MEETING

The annual banquet and 297th meeting of the Botanical Society was held
in the Italian Garden of the Mayflower Hotel, April 4, 1939; attendance 125.

Program.—Ira N. GaBRIELSon: Western American alpine plants (lantern).
—A discussion of personal experiences in seeking out and photographing
alpine plants in many western mountain areas. The Olympics, Siskiyous,
Cascades, Blue Mountains, and several desert ranges in southeastern Ore-
gon were the principal areas visited. It is necessary to make many seasonal
trips into each area in order to find the flowers at their best. General floras
at or near timberline are strikingly similar, but in many ranges distinct and
often beautiful forms and species have been developed. The Siskiyous and
Olympics are particularly rich in localized species.

298TH MEETING

The 298th regular meeting was held in the assembly hall of the Cosmos
Club, May 2, 1939, President CHarLEs THom presiding; attendance 110.

Notes and reviews —JouN A. STEVENSON called attention to his observa-
tion of squirrels cutting off elm leaves and twigs at the rate of 4a minute.
They were also cutting off seed pods.

M. B. Waire displayed a moss protonema that he had found growing in
abundance on ground in which potatoes had grown. The ground had been
bare in November and now was covered with this moss protonema.

Program.—W. A. ArcHER: Indian medicinal plants in Nevada.

H. L. Cranu: Tung-oil industry in the South.

JOINT MEETING

A joint meeting of the Botanical Society and the Biological Society was
held in the assembly hall of the Cosmos Club, April 14, 1939, President
CHARLES THOM presiding; attendance 200.

Program.—Lots JotTrEeR: A botanical trip down the Colorado River by boat
from Greenriver, Utah, through the Grand Canyon, to Boulder City, Nevada,
August 1938.

454 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

SPECIAL MEETING

A special meeting was held in the assembly hall of the Cosmos Club,
September 26, 1939, President CHarLES THom presiding; attendance 110.

Program.—A. H. Recinautp BuuuEr: The sexual process in the rust fungi
(lantern and models).—In long-cycled heteroecious heterothallic rust fungi,
e.g., Puccinia graminis, the sexual process may be initiated in two different
ways: (1) In compound pustules where a (+) mycelium derived from a (+)
basidiospore has fused with a (—) mycelium derived from a (—) basidio-
spore, by an exchange of nuclei, (+) nuclei traveling through the (—)
mycelium to the (—) proto-aecidia and (—) nuclei traveling through the
(+) mycelium to the (+) proto-aecidia; and (2) where (+) and (—) pustules
are widely separated from one another so that compound pustules cannot be
formed, by insects conveying (+) pyenidiospores from (+) pustules to
(—) pustules and (—) pyenidiospores from (—) pustules to (+) pustules,
then by (+) pyenidiospores fusing with (—) flexuous hyphae and (—)
pycnidiospores fusing with (+) flexuous hyphae and, finally, by the nuclei
of the pycnidiospores traveling down flexuous hyphae to the proto-aecidia.
Flexuous hyphae are special hyphae projecting beyond the periphyses into
the drop of nectar that is exuded from the ostiole of every active pycnidium.

299TH MEETING

The 299th regular meeting was held in the assembly hall of the Cosmos
Club, October 3, 1939, President CHARLES THoM presiding; attendance 100.
OuAF 8S. AaomptT was elected to membership.

Notes and reviews.—M. B. Watts exhibited a moss, Webera sessilis, which
he found growing near Patuxent, Md. It does not have any stem and is
about one-fifth the size of a kernel of wheat and is commonly called “powder
gun.” He also displayed a plant, Oldlandia uniflora, which he found growing
at the Horticultural Station at Beltsville, Md. It is related to the common
spring bluet. He also called attention to a Bartram’s oak (Quercus hetero-
filia), which is growing at Twelfth and C Streets, SW., Washington, D. C.
J. B. S. Norton stated there was a Bartram’s oak on the Boulevard near
Riverdale, Md.

Program.—s. F. Buaxe: Early American botanists, John Bannister (1650—
1692), John Clayton (1686-1773), Humphrey Marshall (1722-1801).—John
Bannister, born in England, visited the West Indies and settled in Virginia
in 1678. He collected plants and animals and published several papers on
these subjects. His most important botanical work is his list of about 147
Virginia plants, published in Ray’s Historia plantarum in 1698. John Clay-
ton, born in England came to Virginia in 1705 and was assistant clerk and
clerk of Gloster County for over 50 years. His extensive botanical collection
from Tidewater Virginia were the foundation of Gronovius’s Flora Virginica
(1739-1743), the most important single basis for North American species
in Linnaeus’s Species plantarum. Humphrey Marshall, a cousin of John
Bartram, was born and lived all his life in eastern Pennsylvania. He estab-
lished a private botanical garden, and published his Arbustrum Americanum
in 1785. This was the first work on United States botany published in the
United States.

M. Tuomas Bartram: An early American botanist, John Bartram 1699-
1777.—John Bartram, grandson of a colonist under William Penn, can best
be described as a plain farmer possessing only a country school education
supplemented by wide reading in physics and medicine. His interest 1n
botany was acquired while still in his twenties, and his knowledge of the

Oct. 15, 1940 OBITUARY 455

subject was self-taught through extensive correspondence with American
and European botanists, and the reading of the contributions of Linnaeus
and others. His garden of native plants and shrubs, started in 1731 and in
existence today, is recorded as the earliest in America and served as a source
for the shipment of all types of botanical specimens to subscribers in Europe.
The revenue derived from this activity supported extensive explorations
from ‘“‘New England to Florida; and from the sea coast to Lake Ontario and
branches of the Ohio.” In 1765 he was appointed ‘‘Botanist to the King”
and commissioned to explore Florida and the St. John River.

RosBertT F. Martin: An early American botanist, Thomas Walter 1740?-
GSO.

JO0OTH MEETING

The 300th regular meeting preceded by an informal dinner at Hogate’s
Restaurant, was held at the restaurant, 8:30 p.m., November 7, 1939;
attendance 150.

Notes and reviews.—M. B. WatTE as master of ceremonies briefly remi-
nisced about the early botanists of the society as their pictures were shown
on the screen.

Program.—C. Stuart GAGER: How botany advances.—This address stressed
the importance for the advancement of botany, and of science in general,
of strict adherence to the scientific method, and illustrated the errors and
the absurdities into which students of plant life have been led by failure to
employ what is now generally recognized as ‘‘the method of science.’’ These
illustrations were drawn from the writings of Aristotle (384-322 B. c.),
Caesalpino (1583), Oken, the ‘“‘physio-philosopher”’ (1809), Nicolaus of
Cusa (1514), Van Helmont (1648), Gerard (1597), Parkinson (1640), and
others.

In addition to the more commonly recognized steps of scientific method
(observation, experiment, the formulation and testing of hypotheses), the
the speaker emphasized the importance of another factor less often stressed,
namely, what has been called the ‘‘divination” or ‘‘revelation”’ of the solu-
tion of scientific problems, which Lord Kelvin described as characterizing
his own process. The contrast was also noted between Darwin’s statement of
his procedure, from facts to concept (natural selection), and de Vries’s state-
ment of his procedure, namely, from concept (intracellular pangenesis) to
observation of facts and finally the formulation of the generalization of
mutation. AnicE M. ANDERSEN, Recording Secretary.

Obituary

_ Tuomas ALLEN GROOVER, physician and radiologist, died on April 20,
1940, from x-ray injuries dating from the early days of his investigations.
Dr. Groover was born at Pideock in southern Georgia on May 1, 1877, a
son of Thomas Alfred and Sarah Jane (Joiner) Groover. His father, a farmer,
was a Confederate soldier during the Civil War and had lost an arm in
action. Dr. Groover received his early education in the public schools of
Brooks County, Ga. Because of financial considerations it did not seem
possible for him to continue his education, but through the good offices of
the Hon. Hoke Smith, of Georgia, Secretary of the Interior in President
Cleveland’s Cabinet, he was appointed assistant messenger in the Depart-
ment of the Interior. He came to Washington in 1893 and resided here the
remainder of his life. In 1894 he entered the Medical Department of Colum-
bian University, later George Washington University, and graduated in

456 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 10

1898. In 1926 his alma mater conferred upon him the degree of doctor of
science. After internship at the Garfield Memorial Hospital he was appointed
physician with the Isthmian Canal Commission and in that capacity spent
the next year in Nicaragua.

He returned to Washington in 1900 and entered the general practice of
medicine. At the same time he began x-ray work at the Central Dispensary
and Emergency Hospital. Although he did not completely specialize in
radiology until 1912, he was actively engaged in that field from 1900 up to
within three months of his death. It was in those early days from 1900 to
about 1904 or 1905 that he suffered the injuries to his hands that resulted
in amputation through the left forearm in 1926 and to final involvement of
the right axilla and lung, which cost him his life.

Dr. Groover was closely associated with the scientific progress and the
organizational development of radiology in this country. He made numerous
contributions to the literature and was always interested and quietly helpful
in everything that affected the welfare of the specialty he loved. He was
insistent, however, that the radiologist is primarily a physician, and as such
he always established a personal relationship with every patient who came
under his care.

Dr. Groover’s work as a practicing radiologist was characterized by the
utmost painstaking care and attention to detail. He not only required the
best possible technical work but spared no time or personal effort to arrive
at a correct interpretation and final diagnosis. He had developed in himself
to a very high degree those qualities that are indispensable in the good
physician, accuracy in observation and a keen sense of relative values. He
was a diagnostician of unusual ability. In manner Dr. Groover was quiet and .
reserved. Being of a very studious habit, he made it a rule to spend at least
one hour a day in reading medical literature. He had great qualities both of
mind and heart that were unusual and outstanding. His ability as an or-
ganizer and executive was recognized. His careful, methodical attention to
all the details of bookkeeping, records, and management laid the foundation
for his professional work. He always looked upon business arrangements,
however, as a means to more important ends. He was first and last a phy-
sician. Even radiology, in every branch of which he was proficient, he in-
sisted must always be looked upon as an integral part of general medical
practice. His outstanding mental characteristics were patience and a great

capacity for methodical, painstaking care in every diagnostic and thera- _

peutic procedure. His dominant moral characteristic was a keen sense of
justice, right, and fair dealing. He had an unfailing patience in compromising
differences of opinion among his associates, a sane judgment in arriving at
important decisions, and above all, a kindly, tolerant charity toward all
with whom he could not agree. He will be greatly missed by his close associ-
ates who were accustomed to call upon him almost daily for his helpful
counsel, and his loss will be keenly felt in many organizations in which his
advice was highly valued. His death adds another martyr to the long roll of
those who have sacrificed their lives in the interests of science and humanity.

Dr. Groover was a member of the following organizations: The Washing-
ton Academy of Sciences (vice-president, 1925); the Medical Society of the
District of Columbia (president, 1925); the American Medical Association
(fellow); the American College of Physicians (fellow); the Southern Medical
Association (vice-president, 1924); the American Roentgen Ray Society
(president, 1925); the Radiological Society of North America; and the Ameri-
can College of Radiology (fellow), in which he served both as president and
as chancellor.

CONTENTS

CHEMISTR Y.—Cyanogenesis and enzyme activity in sorghum varieties.
James F. Coucn and Reinnoup R. BRIESE..................

4

PaLEOBOTANY.—A bracket fungus from the late Tertiary of south-
western Idaho. RouaAnp W. BROWN: >... 2... (236.

Botany.—Ozytenanthera felix, a new species of bamboo from Yunnan,
China, #Y.. ib ISENG SO Bp eS

Botany.—Taxonomic relationships in the genus Gossypium. 8S. C.
HARLAND. 203. ale ey a sais Nyse te a koh ues GE OM a ear ame

ENTOMOLOGY.—Some new species of Syrphidae (Diptera). Frank M.

ZooLocy.—A new species of phyllopod crustacean from Stone Moun-
tain, Georgia:.. Hpwin P. CREASER 22... 7.0% 6,0 3

ZOOLOGY.—Cercaria pricei, a new trematode, with remarks on the
specific characters of the ‘‘Prima’’ group of Xiphidiocercariae. —

MintAM ROTHSCHILD. 63 o. 4 ee eae oon + sie ot tie de oe
Progeepwes: Toe ACADEMY...» 0
PROCEEDINGS: BOTANICAL SOCIETY. .........0...0c2 ces ceeue ee.
OBITUARY: THomMAS ALLEN GROOVER............... Re be Poon.

This Journal is Indexed in the International Index to Periodicals

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JOURNAL

OF THE
WASHINGTON ACADEMY OF SCIENCES
Vou. 30 NovemBeER 15, 1940 IN@s 1

BIOCHEMISTRY.—The bacteriostatic and the bactericidal action of
certain organic sulphur compounds.! Epwarp L. Evreritr and
M. X. SULLIVAN, Georgetown University.

For a number of years in the field of chemotherapy investigations
have been carried on relative to the treatment of various infections
of man by certain chemicals. The importance of the use of chemical
substances lies in the possibility of checking or destroying the invad-
ing organisms, or in offsetting the products of the activity of the or-
ganisms in question. Since many compounds having fungicidal or
bactericidal properties are harmful to the body tissues, very few
chemical compounds have been found useful for internal application,
such as injection into muscles or directly into the blood stream.

Within the past few years, however, several compounds have been
discovered that are of value in the treatment of pneumonia and of
certain streptococcic infections. Thus, Rosenthal (1934) investigated
the action of sodium formaldehyde sulphoxylate on virulent pneu-
mococci and found it effective against one strain of the organism
among the 32 strains studied with mice. The sulphoxylate, as the
complex is called, was injected into mice inoculated with the particu-
lar strain of pneumococci under investigation. In contrast to its
effectiveness in mice was the fact that in vitro tests showed that the
compound had little if any bactericidal or bacteriostatic action. Do-
magk (1935) in Germany then announced a discovery that has
revived chemotherapeutic research. He found that the addition of a
sulphamide group into certain azo dyes would protect or cure mice
infected with fatal doses of hemolytic streptococci and that this effect
was not correlated with in vitro experiments, in which the compounds
had little effect on the organisms.

The compounds found most effective by Domagk were: 4-sulphon-
amide-2, 4-diamino-azobenzene, and 4-sulphonamide-phenyl-2-azo-
7-acetylamino-1-hydroxynaphthalene-3, 6-sodium disulphonate, which

1 The data in this paper are taken in part from the dissertation presented by

Edward L. Everitt in partial fulfillment of the requirements for the degree of doctor
of philosophy, Georgetown University, 1937. Received July 9, 1940.

457

s Tt. 1940
Nov 25

458 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES. VOL. 30, No. 11

were named respectively prontosil and prontosil soluble. Domagk’s
work was soon followed by that of Trefouel, Nitti, and Bovet (1935),
who described a sulphonamide product that did not contain the azo-
group but that still had chemotherapeutic activity. This compound,

p-aminobenzene sulphonamide, HNC »SO.N H2, was first known

as prontylin and has been extensively used in this country and abroad
under the name sulphanilamide. It has been found very valuable
therapeutically against several infectious diseases, streptococci infec-
tions in particular. An excellent review with 162 references has been
made by Holman and Duff (1938).

In work with various sulphur compounds, started long before the —
publication of Domagk’s work and subsequent work of others on
sulphanilamide and its main derivatives, we found that a number of
organic sulphur compounds investigated in vitro had a fungistatic
or a fungicidal action on various molds (Everitt and Sullivan, 1940).
The compounds made use of in the mold studies were then tested as
to their bacteriostatic and bactericidal actions, with especial atten-
tion to the streptococci and staphylococci.

The culture media employed in the study of the growth of the
bacteria were those commonly used in bacteriological work and were
made up as follows, (A) and (B):

(A) BROTH MEDIUM

Neopeptome Moan cola ac et eG eee 10 grams
Sodium: chioride.. 66 oe se. ie. a ee oe 5 grams
Water tovmakeé 8. s5 os fies cl es Bee 1,000 ce

This culture medium was adjusted to pH 7.6, and 50-ce portions
were placed in 125-cc Erlenmeyer flasks. The sulphur compounds
were added in specified amounts, and the flasks were plugged with
cotton and sterilized at 15 pounds pressure for 30 minutes. Certain
flasks without the particular compounds served as controls.

(B) BLoop AGAR SLANTS

The viability of the microorganisms on medium (A) with and with-
out the added sulphur compounds was further determined by making
transfers to blood agar slants. These slants were made by adding 0.5
ce of sterile blood to tubes of agar made up as follows:

Beef extract: fe: cascode os eo ee 3 grams
Peptone soe. PAs eee RE ek a 10 grams
Sodium chlorides ee ee AS eco 2S 5 grams
PA CATA OAT iis le emte te tae Meena Me ee ge Abe ore et 15-18 grams

Water tommake:. teyn edb eee een ae cas tee ee 1,000 cc

Nov. 15, 1940 EVERITT AND SULLIVAN: SULPHUR COMPOUNDS 459

The agar solution was then tubed, plugged with cotton, and steri-
lized at 15 pounds pressure for 30 minutes. The sterile blood was
added by means of sterile pipettes to these sterilized tubes of agar
after they had been cooled in a slanting position.

In the study, the bacterial culture medium (broth medium) was
inoculated by the addition to the various flasks of two drops of an
18- to 24-hour peptone medium growth of the particular organism.

The inoculation of the blood agar slants was made by transferring
a 6-mm loopful of the culture media to the surface of the slants.

In each case, whether in broth or agar slants, the bacterial cultures
were incubated at 37° C.

A series of preliminary tests with a number of sulphur compounds
to determine their bacteriostatic and bactericidal properties was
undertaken with the use of Staphylococcus aureus as the test organism.
As this organism was found very resistant, its study was deferred
until other possibly more potent compounds could be made, and at-
tention was paid to the streptococci, considered of more importance
from the standpoint of human infections. The streptococci employed
were (a) Streptococcus hemolyticus (oyler strain) isolated from a pa-
tient with an infection on the dorsal surface of the hand and found
capable of producing beta hemolysis; (b) Streptococcus hemolyticus
epidemicus Group A, obtained from Dr. Alice Evans, National In-
stitute of Health. It belonged to Lancefield’s group A and was viru-
lent to white mice; (c) Streptococcus viridans No. 2, obtained from
Dr. Cecil in 1935. It produced alpha hemolysis. All the streptococci
used were provided by the Department of Bacteriology, Georgetown
University Medical School.

In preliminary work about 50 chemicals previously tested in mold
growth and listed in the reference to the fungicidal and fungistatic
action of various sulphur compounds by Everitt and Sullivan (1940)
were tested for their effect on the streptococci. Only a few seemed
promising from a bacteriostatic or bactericidal standpoint. These
possibly useful compounds are as follows:

(3) Phenothioxine

(10) Phenyl thioarsenite

(11) 4-chloro-2-nitrophenyl sulphur amine
(14) Thioacetamide

(15) Mercaptobenzothiazole

(16) Phenylbenzothiazole

(17) Sodium diethyldithiocarbamate

(18) Diethylthiourea

460 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

(31) Thiobarbituric acid

(32) Benzyl disulphide

(33) Colloidal sulphur (sulphur diasporal)

(34) Colloidal sulphur (sulisocol)

(36) 1,2 naphthoquinone-4-sodium sulphonate

The effect of the various compounds was determined by incubating
the broth culture containing the test material for 24 to 96 hours and
at each twenty-fourth hour transferring a loopful to blood agar slants
for a further judgment as to growth. The effect of the compounds on
the bacteria is given in Tables 1, 2, and 3.

TABLE 1.—Stupy or THE Most PRomMISING SULPHUR COMPOUNDS FOR ACTION
ON STREPTOCOCCUS HEMOLYTICUS (OYLER STRAIN)

Time interval of incubation in hours on medium (A)

eae ql pout before transplanting to blood agar!
No. 100 ce 24 48 72 96
mg
Control... —— + + +
(3) 20 a 0 0 0
15 + + 1 colony 0
CEA eee 15 15 colonies + + 0
CUD). 10 0 0 0 0
Gor: 10 0 FP == ap
Glia 15 0 0 -0 0
(tS). 3 15 + 0 0 0
(@lD) . 15 1 colony “- 0 0
(2) ie 20 0 0 0 0
(33) 20 0 0 == ote
(34) . 20 “+ 0 0 0
(36). 15 0 0 0 0

1 The sign (+) signifies growth on the blood agar, (0) no growth.

TABLE 2.—StTupy oF SULPHUR COMPOUNDS ON GROWTH OF STREPTOCOCCUS
HEMOLYTICUS EPIDEMICUS

Time interval of incubation in hours on medium (A)

Sulphur Amount before transfer to blood agar!

compound used in
ND. 100 ee 24 48 72 96 120
mg
Control... — + + + + 0
(Galas 10 0 0 0) 0 0
ClO) Se 10 0 0 0 0 0
(GED ees 15 1 colony OF 0 0 0
(Ob i ee 10 0 0 0 0 0
(iS) ee 10 0 0 0 0 0
(UG s 10 + 0 + 0 0
CLS) eee 15 6 colonies 0 0 0 0
(BilWag & 15 a + + +
G2) eee 15 aa + + +
G3)ine 20 0 0 0 0
(4) ee 20 0) 0 0 0
(GIS) 10 + 0 0 0 0)
5 + 0 0 0 0

La-=e Dp

1 The sign (+) signifies growth on the blood agar, (0) no growth.

Nov. 15, 1940 EVERITT AND SULLIVAN: SULPHUR COMPOUNDS 461

TABLE 3.—ACTION OF CERTAIN OF THE SULPHUR COMPOUNDS ON STREPTOCOCCUS
VIRIDANS INCUBATED IN 24-HourR PERIODS ON MeEpivum (A) AND A LOOPFUL
oF BrotH TRANSFERRED TO BLoop AGAR SLANTS TO DETERMINE GROWTH

Sulphur Amount Time in hours
compound used in. |—---
No. 50 ce 24 48 72 96 120
mg
(3) ie 5.0 ar =e ls le le
Tf fs “te aks 0 0 0
GIES) 0) 15 colonies — — -- 0
(16). 5.0 = =r =e a
GIy): C5 ae + =F + 3 colonies
(18). ao qr ar ae aE
GIO). 5.0 1 colony — 2 colonies + 0
GIS)? les a =F ts

A study of Table 1 will show that a number of compounds have a
retarding action in vitro on the growth of Streptococcus hemolyticus
(oyler strain). These compounds are 3, 15, 17, 18, 32, 33, 34, and 36,
and of these 15, 17, and 36 were the most effective. Similarly, as
shown in Table 2, a number of compounds inhibited the growth of
Streptococcus hemolyticus epidemicus. These compounds are 3, 10, 11,
14, 15, 16, 18, 33, 34, and 36.

Of the compounds tested and found inhibiting on both strains of
streptococci there may be listed nos. 3, 15, 18, 33, 34, and 36. Com-
pounds 33 and 34, colloidal sulphur preparations, may be effective
because they are alkaline in reaction and may change the hydrogen-
ion concentration of the culture medium sufficiently to inhibit the
growth of the organisms. Since this question called for a detailed
study, these compounds were set aside for further consideration. Since
compound 36 (1, 2 naphthoquinone-4-sodium sulphonate) was early
shown by Herter (1905) to have more or less toxicity toward animals,
the use in offsetting the action of streptococci in vivo might be ques-
tionable. :

Finding that various compounds checked the growth of Strepto-
coccus hemolyticus and Streptococcus hemolyticus epidemicus, we de-
cided to study their effect on the highly virulent microorganism
Streptococcus viridans, the checking of which has long been a desired
need in medical practice. The results with Streptococcus viridans are
shown in Table 3.

It is obvious from Table 3 that Streptococcus viridans is more re-
sistant to the action of the various compounds than are Streptococcus
hemolyticus and Streptococcus hemolyticus epidemicus. Streptococcus
viridans infection in human beings usually has a very unfavorable
outcome and is one of the organisms for which some effective com-

ee ee ee a

462 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

pound should be found. Of the compounds tested, only three seem
| to have some inhibiting action on Streptococcus viridans. These com-
y pounds are nos. 3, phenothioxine,

| x Or .

10, phenyl thioarsenite, As—-S< >
Sea
if n

\
and 15, mercaptobenzothiazole, C-SH.

A
a
t Of these compounds, mercaptobenzothiazole was on hand in plenti-
' ful supply, and some attention was paid to it from a chemical and a
f clinical viewpoint. As may be seen from its formula, it is an organic
, sulphur compound containing nitrogen and in addition a -SH group.
v This compound, which was first made by Hofmann (1887), has been
used for a number of years in the rubber industry as an accelerator of
: vulcanization. Its use for this purpose seems to have been first sug-
3 gested by Bedford and Sebrell (1921). It may be purchased from the
Eastman Kodak Co. It is moderately soluble in alcohol, ether, chloro-
A, form, and benzol, alkalies, and alkali carbonate solutions but not
| very soluble in water.
fl Roark and Busbey (1935) state that mercaptobenzothiazole in
; concentrations of 0.01 to 0.10 percent was effective in controlling a
fungus living on wood and that it has been used in controlling aphids
iia; and mosquito larvae. Davis (1930) reported on the toxicity of the
compound. He injected an aqueous solution of it into guinea pigs
NR and a total injection of 14.5 mg in 20 days did not produce any in-
NB jurious effects on the animals. Also, medical examinations of the men
| working with mercaptobenzothiazole over a period of years in the
a

Goodyear Tire & Rubber Co. did not show any toxic condition or
dermatoses. We have given 20-100 mg of the compound by mouth to
guinea pigs weighing 400 grams with no gross evidence of toxicity.
The disadvantages of using mercaptobenzothiazole are its slight
solubility and its tendency to chemical change when in an alkaline
solution.
In preliminary work, phenothioxine in a slightly alkaline medium
\ and at a level of 20 mg per 100 cc has shown good bacteriostatic action
on the streptococci, even on Streptococcus viridans. Phenothioxine

>) ~~ _

Nov. 15, 1940 ©®vVERITT AND SULLIVAN: SULPHUR COMPOUNDS 463

however, is rather difficultly soluble in the culture media, but it can
be converted readily to a more soluble sulphonate which will be in-
vestigated further in vitro and in vivo.

SUMMARY

In testing about 50 organic sulphur compounds as regards their
effect on the growth of various streptococci, a number of compounds
were found to have more or less bacteriostatic or bactericidal power
at a level of 100 parts per million of peptone broth. The most promis-
ing compounds are phenothioxine, which showed marked bacterio-
static action on Streptococcus hemolyticus (oyler strain) and Strepto-
coccus hemolyticus epidemicus and considerable inhibition of Strep-
tococcus viridans; and mercaptobenzothiazole, which showed bacteri-
cidal action on Streptococcus hemolyticus (oyler strain) and on
Streptococcus hemolyticus epidemicus, but not on Streptococcus viridans.
The criterion of biological action of the various compounds was the
degree of growth on blood agar slants after a period in peptone broth
culture with and without the compound under study.

LITERATURE CITED

Beprorp, C. W., and SEBRELL, L. B. Reactions of accelerators during vulcanization
III. Carbosulfhydryl accelerators and the action of zinc oxide. Journ. Ind. Eng.
Chem. 13: 1034. 1921.

Davis, P. A. Toxic substances 1n the rubber industry. Pt. XI: Mercaptobenzothiazole.
Rubber Age 27: 2491. 1930.

Domack, G. Ein Beitrag zur Chemotherapie der bacteriellen Infecktionen. Deutsch.
Med. Wochschr. 61: 250. 1935.

Everitt, E. L., and Sutuivan, M. X. The fungistatic and fungicidal action of certain
organic sulphur compounds. Journ. Washington Acad. Sci. 30: 125. 1940.

Herter, C. A. The color reactions of naphthoquinone sodium monosulfonate and some
of their biological applications. Journ. Exper. Med. 7: 79. 1905.

Hormann, A. W. Zur Kenntniss des 0o-Amidophenylmercaptans. Ber. 20: 1788. 1887.

Houtman, W. L., and Durr, G. L. Sulfanilamide and similar compounds in chemo-
therapy. Amer. Journ. Med. Sci. 195: 379. 19388.

Roark, R. C., and Bussey, R. L. A list of organic sulphur compounds (exclusive of
mothproofing materials) used as insecticides. U. S. Dept. Agr., Bur. Ent. and Plant
Quar., Div. Insecticide Investigations. 1935.

RosENTHAL, 8S. M. Studies in chemotherapy. I. The action of sodium formaldehydesulf-
oxylate in bacterial infections. Publ. Health Rept. U. 8. Public Health Service,
49: 908. 1934.

TREFOUEL, J.. TREFOUEL, Mme. J., Nitti, F., and Bovet, D. Actwité du p-amino-
phenylsulfamide sur les infections streptococciques expérimentales de la souris et du
lapin. Compt. Rend. Soc. Biol. 120: 756. 1935.

Fn et emnine + adie
Te. B=) sy) ss a

—— a + oa Se

464 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

PALEOBOTANY.—A Cusparia from the Pliocene of trans-Andean
Bolwia.t Evwarp W. Berry, Johns Hopkins University.

The specimens on which the following note is based were collected
many years ago by Dr. Edwin Kirk, to whom I am indebted for them
as well as the information regarding their geological setting.

They were collected at Agua Salada (lat. 21° 50’ 20” S., long. 63°
30’ W.), which is about 25 kilometers northeast of Yacuiba, Bolivia,
from the surface of an argillaceous deposit, some thousands of feet
above the base of what Bonarelli? called the “‘Terciario subandino.”
This is more or less the equivalent of what Madgwick called the
Upper Sacarigua beds; Heald and Mather the San Ysidro forma-

tion; and Mather the Tatarenda formation. Similar deposits in this

general region were earlier christened the Jujuy formation by Stein-
mann. Beds of this character extend in the sub-Andean zone through
many degrees of latitude and appear to be lithologically and geneti-
cally similar. They appear to be poorly fossiliferous everywhere,
although doubtless detailed searching of outcrops in this superficially
explored region might greatly modify this statement. Bonarelli and
others have reported obscure molluscan remains and ostracods;
Harrington collected insects, which were described by Cockerell (1925) ;
Kraglievie records a sloth cranium (Scelidodon); and Kirk collected
a mammalian jaw which Gidley regarded as of Pliocene age. Fossil
leaves of Cassia chrysocarpoides Engelhardt were collected at Caran-
daiti,? which is about 150 miles north of Agua Salada.

This last is a common Potosi form and a Corbicula mentioned by
Bonarelli also occurs in the inter-Andean Pliocene.

In previous discussions of the date of the final uplift of the Andes men-
tion has been made of the thick series of erosion products that accom-
panied this uplift, remnants of which skirt the mountains from eastern
Colombia southward to western Argentina and from which a consid-
erable number of fossil plants have been described, as at the Rio
Aguaytia locality in eastern Peru, or molluscan faunas as near Pebas,
Peru. Some of the latter as well as many of the plants are identical
with inter-Andean forms from the Cuenca and Loja basins in Ecuador,
from the headwaters of the Orinoco system in Colombia, or the head-
waters of the Amazon system in eastern Ecuador and Peru, the Terri-
tory of Acre in western Brazil, and in eastern Bolivia.

It appears to me that they all clearly represent a part of the same

1 Received July 25, 1940.

2 BONARELLI, G. "Anal. Min. /fag, hse) (U1), GPa
3 Berry, E. W. Johns Hopkins Univ. Stud. Geol. No. 13:51. 1939.

Nov. 15, 1940 BERRY: A CUSPARIA FROM BOLIVIA 465

geological epsiode and are of about the same age, although it obvi-
ously would be inaccurate to assume exact correlation of named litho-
logic units through this vast distance, especially with the realization
of the difficulty of the country and the reconnaissance nature of its
exploration. The fossil fruits from Agua Salada are tentatively iden-
tified as belonging to the rutaceous genus Cusparia and may be de-
scribed as follows:

Cusparia yacuibana Berry, n. sp. Figs. 1—7

Carpels unsymmetrically egg-shaped ranging from irregularly obovate to
subrhomboidal in lateral profile, somewhat compressed, rounded above and

Figs. 1-7.—Cusparia yacuibana Berry: 1, Two specimens assembled to depict a
fruit consisting of two capsules; 2—4, lateral views of valves; 5, 6, internal views of
valves showing outline of the single seed, placenta, and dorsal dehiscence canal; 7,
ventral view of partly opened capsule. All X2.

obtusely pointed proximad, a slight sinus on the ventral border. Substance
crustaceous or bony. Surface with a characteristic sculpture, heavily rugose.
Ribs comprising a broad, rounded, lunate rib originating near the middle of
the ventral margin curving outward and downward subparallel with the dor-
sal margin. This central rib divides the surface into two regions—above it
there are at least two major and one or two minor ribs, all subparallel; below
it there are three or four similar ribs more parallel with the axis of the fruit
and gradually becoming obsolete at its base. All the ribs are subject to con-
siderable variation in both extent and magnitude. Dimensions: Length,
ranging from 7 to 10; maximum width, 5 to 7; thickness, 4 to 5 mm.
Carpel is tardily dehiscent into two equal valves, well shown in Figs. 1-4.
At least one specimen preserves the single reniform seed, which is hard and
smooth and exactly fills the cavity. By voluntary shifting the focus of the
_ eyes the observer of Figs. 5 and 6 can get the exact picture of the valve with
the seed preserved except that the seed is smooth and rounded dorsally
and does not extend into the canal through the upper dorsal wall of the cap-
sule. This canal has been puzzling. It can not be detected in all the speci-
mens, nor is any trace of it shown from the outside in the unopened speci-

466 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

mens. It is present in the majority of dehisced valves and appears to make
the dorsal limit of dehiscence. The wall must have differed naturally in
structure along this line although nothing can be made out microscopically.
In Fig. 3 a considerable sinus on the upper dorsal margin is shown at this point.

The seed is interpreted as pointed at the base, although no trace of a stylar
canal is preserved in the capsule wall. The hilum is supposed to be at the
concavity on the ventral side, and the swollen wall around which it fits is
supposed to represent the placenta. No trace of a funicular canal can be de-
tected in any of the specimens.

These carpels, seemingly so individualistic and well marked, have baffled
both identification and interpretation for many years. I have submitted
them to systematists at some of our larger herbaria as well as to experts in
the study of fossil carpology, such as Mrs. Reed, the well-known British
authority, to all of whom I am grateful. One insuperable difficulty as usual
is to find comparable recent material since most botanical species are founded
on herbarium material of flowers and leaves.

For a long time I was convinced that the specimens represented the stones
of a drupe or berry. Arguments for such an interpretation are the traces of a
dark organic substance in the grooves between the ribs; the stony character;
the lack of traces of attachment; the fact that 10 specimens were unopened
and there were but 28 half specimens. Arguments for a capsular nature was
the dehiscence of a majority of the specimens; the prevalence of the capsule
habit in this region of the Rutaceae and the somewhat similar ornamentation
in many of the Recent genera some of which have bony capsules. The con-
sensus of the authorities consulted was that the fossils were closely related
to the genus Cusparia, and since I can not believe that an extinct genus was
in existence as late as the Pliocene in tropical South America, I have ventured
to describe them as an extinct species of that genus, and I am optimistic
enough to expect that when the botanical exploration of that vast region shall
be more complete than it now is this extinct species may be found to be a
still existing one. Decision as to habit and orientation has also been difficult.
At first I assumed that the more pointed end to which the ribs converged was
the apex, but a review of the literature and such recent material as was avail-
able convinces me that the widest part of the capsule is distad and the nar-
rower end from which the ribs diverge is the proximal part. All the material
that I have seen has the ornamentation disposed in this pattern.

There are many similar, but not identical, Recent forms in the Cusparieae
especially in the genera Cusparia, Pilocarpus, and Raputia. In Cusparia, to
which the fossils conform more closely than to the others, the mature cap-
sules are one to three and rarely four in number. I am giving a restoration of
the fossil in which two capsules are shown as having matured.

The subfamily Cusparieae comprises 16 genera with upward of 100 known
species, all American, in the existing flora. Engler, who prepared the account
of the Rutaceae in Martius’s Flora Brasiliensis and for Das nattirlichen
Pflanzenfamilien, discussed the geographical distribution in a special pa-

- Nov. 15, 1940 BLAKE: ARIZONA ASTERACEAE 467

per? in 1816, since when there has been considerable botanical exploration,
and it now seems probable that most of the genera of the Cusparieae, certain-
ly all the larger genera, may be expected to be found anywhere in the neo-
tropical region of South America centering in the Orinoco and Amazon
basins, extending to the Greater Antilles (Cuba, Haiti, and Puerto Rico) in
the genus Ravenia, to Central America and southern Mexico in the genus
Erythrochiton, to Paraguay and Uruguay in the genus Pilocarpus, and to the
eastern Andean slopes of Peru and Bolivia in the genera Erythrochiton and
Cusparza, and probably in the majority of other genera of this subfamily.

At least two species of Cusparia (ramiflora and toxicaria) are recorded
from the upper Amazonian drainage basin, and one (pilocarpoidea), although
with capsules not close to the fossil, is found in eastern Bolivia.

Cusparia, so far as I know, has not previously been found fossil, and the
same statement is true of 12 of the other genera comprising this subfamily.
The three genera, in addition to the present occurrence of Cusparia, which
have furnished fossils are (1) Pzlocarpus, species of which are recorded from
the lower Miocene of Coronel, Chile, the Pliocene of Psillypampa, Bolivia,
and the Pliocene of Aramary and Marahii in eastern Brazil; (2) Ticorea,
which is recorded from the lower Miocene of Chile; and (3) Evrythrochiton
with a species associated with T7zcorea and a second from the Pliocene of
eastern Brazil (Aramary in the State of Bahia and Fonseca in the State of
Minas Geraes).

BOTAN Y.—New species and new names among Arizona Asteraceae.'
5. F. Buaxs, U. 8. Bureau of Plant Industry.

The following new species, new varieties, and new combinations are
published here in order that they may be available for the Flowering
plants and ferns of Arizona, by T. H. Kearney and R. H. Peebles, now
in course of publication by the U. 8. Department of Agriculture.

Kuhnia rosmarinifolia Vent. var. chlorolepis (Woot. and Standl.). Blake
Kuhnia chlorolepis Woot. and Standl. Contr. U.S. Nat. Herb. 16: 177. 1913.

Aplopappus croceus Gray var. genuflexus (Greene) Blake

Pyrrocoma genuflexa Greene, Pittonia 3: 348. 1898.
Haplopappus croceus subsp. genuflecus Hall, Genus Haplopappus 99. 1928.

Chrysothamnus pulchellus (Gray) Greene var. baileyi
(Woot. and Standl.) Blake

Chrysothamnus baileyi Woot. and Standl. Contr. U. 8S. Nat. Herb. 16: 181.
1923.

Chrysothamnus pulchellus subsp. baileyz Hall and Clements, Phylogen. Meth.
Taxon. 194. 1928.

4ENeter, A. Uber die geographische Verbreitung der Rutaceae im Verhdltnis zu
threr systematischen Ghiederung. Abs. Preuss. Akad. Wiss. 1896: 1-27, pls. 1-3. 1896.
1 Received October 8, 1940.

Bate

468 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

Chrysothamnus viscidiflorus (Hook.) Nutt. var. elegans (Greene) Blake

Chrysothamnus elegans Greene, Erythea 3:94. 1895.
Chrysothamnus viscidiflorus subsp. elegans Hall and Clements, Phylogen.
Meth. Taxon. 188. 1923.

Chrysothamnus viscidiflorus var. molestus Blake, var. nov.

Suffrutex 8-12 cm altus pluricaulis; caules albo-corticati foliosi normaliter
infra inflorescentiam simplices ut folia subdense hirtelli pilis brevibus pa-
tentibus albis conicis saepe glanduloso-capitatis; folia linearia vel infima
anguste lineari-spathulata calloso-apiculata sessilia plana v. concava 1-nervia
pallide viridia 7-15 mm longa 0.8-1.5 mm lata; cymae terminales folia su-
perantia pauci- v. multicapitatae 1.5-4 cm latae ut caules hirtellae; capitula
anguste cylindrica 5-flora fructu 1.2-1.5 cm alta (pappo incluso) ;involucri
valde gradati 4-seriati 9.5-11 mm alti phyllaria in ordinibus 5 perpendiculatis
ordinata linearia v. extima lineari-oblonga obtusa v. acutiuscula non valde
carinata 0.8-1.2 mm lata dorso glabra vel extima parce hirtella substraminea
scarioso-marginata margine prope apicem eroso-ciliata exteriora apice viridia ;
flores maturi involucro multo longiores; corollae tenues prope basin faucis
parce puberulae ut videtur pallide flavae (6.3) 7.8-8.2 mm longae (tubo
1.8—2.3 mm, fauce 4.5-4.7 mm, dentibus 1-1.5 mm longis); achenia linearia
4.2-6 mm longa 6-8-nervia infra glabra supra dense glandulosa glandulis
brevissime stipitatis; pappus vix copiosus stramineus corollam maturam
aequans 8-9 mm longus, setis subequalibus hispidulis prope apicem longius
hispidulis; antherarum appendices anguste lanceolatae 0.5—0.8 mm longae;
styli rami 3-4 mm longi, lineis stigmaticis 1.83-3 mm appendicibus anguste
subulatis hispidulis (1) 1.3-1.7 mm longis.

ARIZONA: On limestone slope in edge of pifion belt, easterly base of San
Francisco Mountain, 45 miles from Flagstaff, Sept. 26, 1920, H. M. Hall
11184 (Herb. Univ. Calif. no. 205824) ; between Williams and Grand Canyon,
west of San Francisco Mountain, on plains, Sept. 27, 1920, Hall 11199 (type
no. 1288511, U. S. Nat. Herb.; duplicate, Herb. Univ. Calif. no. 205819);
open rocky park, Tusayan Forest, T. 23 N., R. 3 E., altitude 2,135 m, Sept.
9, 1912, A. D. Read 165 (U.S. For. Serv. Herb. 5439).

This little shrub, known from three collections from the general region of
San Francisco Peak in northern Arizona (Coconino County), is of consider-
able interest. Hall’s specimens were identified by him as Chrysothamnus de-
pressus Nutt., and the measurements of floral details (not made by Hall
himself, and in part not agreeing with my own measurements from the same
specimens) attached to the sheets in the University of California Herbarium
are given in his table of measurements of that species. The sheet in the U. 5.
Forest Service was originally identified by Dr. E. L. Greene as C. pulchellus
(Gray) Greene, and later by the writer as C. vaseyi (Gray) Greene. It is
rather surprising that the specimens should have been identified as C. de-
pressus and C.. pulchellus, since they completely lack the outstanding charac-
teristic of these two species, namely strongly acuminate or cuspidate and
strongly keeled phyllaries arranged in five very distinct vertical ranks. Their
separation from C. vaseyi, with which as well as the two species just named
they share the character of a merely glandular achene (not densely pubes-

Nov. 15, 1940 BLAKE: ARIZONA ASTERACEAE 469

cent, as in the forms of C. viscidiflorus hitherto known), is much less readily
made. Chrysothamnus vaseyt, which is not known from Arizona, is closely
similar in habit to the new plant; its branches, although described by Gray,
Rydberg, and Hall and Clements as glabrous, are often puberulous, but much
less coarsely so than in the new plant, and never with glandular-capitate
hairs developed; its leaves (sometimes finely puberulous) are normally longer
and wider, with a weaker costa, usually indicated on the upper leaf surface,
where the midrib in the new form is most prominently developed, only by an
impressed line; its involucre is shorter (5.5-7 mm high), of broader and
blunter phyllaries with less evident keel; its corollas are shorter (5.5-6.5 mm
long), with relatively longer teeth (1.5—2 mm long); and its achenes are 9—10-
nerved.

In achenial characters var. molestus is more like C. vaseyz, in which the
achene is 9-10-ribbed, glabrous or glandular or rarely sparsely pubescent
above, than the forms of C. viscidiflorus, in which the achene is about
5-nerved and almost always more or less densely pubescent but not glandu-
lar. The only previous record of the occurrence of nearly glabrous achenes
among the forms of C. viscidiflorus seems to be in C. marianus Rydb.,
synonymized by Hall and Clements with C. viscidiflorus subsp. puberulus,
apparently correctly. In this the achenes were originally described as only
slightly strigose, and in the type specimen (Rydberg & Carleton 6993) are said
by Hall and Clements to be sparsely strigose or nearly glabrous.

The degree of variation in the proportions of the parts of the style in var.
molestus is considerable and suggests that the emphasis placed on this feature
by Hall and Clements in their monograph of Chrysothamnus may have been
exaggerated. In the three collections now at hand, coming from a restricted
region and extremely similar in all other characters, the style branches in
normally developed flowers vary from 3 to 4 mm in length, the stigmatic lines
from 1.3 to 3 mm, and the appendages from 1 to 1.7 mm, the appendages
being sometimes distinctly longer than the stigmatic lines, sometimes slightly
less than half as long.

Var. molestus is distinguished from all the forms of C. viscidiflorus hitherto
_known by its coarser pubescence consisting in part of gland-tipped hairs, its
longer involucre, and its glandular achenes. In many of its characters it is
more or less intermediate between C. depressus and C. viscidiflorus var. ele-
gans, and might almost as well be considered a variant of C. depressus with
blunt phyllaries. The varietal name refers to the difficulties its recognition
introduces into the preparation of a key to the species of the genus. Hall
states that the specimens collected under his no. 11184 were much browsed
back by grazing animals.

Chrysothamnus greenei (Gray) Greene var. filifolius (Rydb.) Blake

Chrysothamnus filifolius Rydb. Bull. Torrey Club 28: 503. 1901.
Chrysothamnus greenet subsp. filzfolius Hall and Clements, Phylogen. Meth.
Taxon. 191. 1923.

\
\
a

470 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

Chrysothamnus nauseosus (Pall.) Britton var. turbinatus (Jones) Blake

Bigelovia turbinata Jones, Proc. Calif. Acad. II. 5: 691. 1895.
Chrysothamnus turbinatus Rydb. Fl. Rocky Mts. 859. 1917.

Chrysothamnus nauseosus subsp. turbinatus Hall and Clements, Phylogen.
Meth. Taxon. 217. 1923.

Achaetogeron chihuahuensis Larsen, sp. nov.’

Herba annua (?) 10-30 cm alta pluricaulis; caules simplices v. parum
ramosi adscendentes v. decumbentes 1—2-capitati saepe purpurascentes foli-
osi subdense pilosi v. hirsuti pilis infra patentibus supra adscendentibus v.
subappressis; folia infima anguste spathulata v. oblanceolata longe petiolata
utrinque pilosa v. hirsuta integra v. utroque latere 1-dentata obtusa v.
acutiuscula 2-4 em longa (petiolo incluso) 3-4 mm lata, caulina anguste
oblanceolata v. linearia sessilia v. inferiora petiolata sursum sensim reducta;
pedunculus 1.5—6 em longus nudus v. subnudus adscendenti- v. subappresso-
pilosus; capitula solitaria 1.5—2 em lata; involucri 2-seriati subaequalis 4 mm
alti phyllaria lanceolata acuminata patenti-pilosa et minute glandulari-
puberula; radii numerosi pauciseriati sicco albidi v. lavandulaceo-tincti;
achenia paullum compressa oblonga 2-nervia albida minute hirsutula 0.8 mm
longa; pappus minimus coroniformis multisetulosus basi concretus 0.1 mm.
longus.

Arizona: White Mountain Apache Reservation, Apache County, June 25,
1907, Mrs. Zuck; semiwaste places, Thompson’s Ranch, Black River, White
Mountains, July 12, 1910, L. N. Goodding 539.

CHIHUAHUA: Vicinity of Madera, alt. 2,250 m, May 27—June 3, 1908,
E. Palmer 301 (type in Mo. Bot. Gard.; dupl. in U. 8. Nat. Herb.), 302438;
Colonia Garcia, in Sierra Madre, alt. 2,218 m, June 12, 1899, Townsend &
Barber 32.

Of interest as the first species of this Mexican genus to be reported from
the United States. In general appearance it resembles species of Hrzgeron
such as H. divergens Torr. and Gray, but it is readily distinguished by the
minute coroniform pappus. All the specimens cited are in the U. S. National
Herbarium except Townsend & Barber 32, which may be present there but
has not been located.

Aster commutatus (Torr. and Gray) Gray var. crassulus (Rydb.) Blake

Aster crassulus Rydb. Bull. Torrey Club 28: 504. 1901.
Aster commutatus crassulus Blake in Tidestrom, Contr. U. 8. Nat. Herb. 25:
560. 1925.

Aster commutatus var. polycephalus (Rydb.) Blake

Aster scoparius DC. Prodr. 5: 242. 1836. Not A. scoparius Nees, 1818.

Aster polycephalus Rydb. Bull. Torrey Club 33: 153. 1906.

Aster commutatus polycephalus Blake in Tidestrom, Contr. U. 8S. Nat. Herb.
25: 560. 1925.

2 Description contributed from her unpublished revision of this genus by Esther L.
Larsen, now Mrs. Kenneth D. Doak.

Nov. 15, 1940 BLAKE: ARIZONA ASTERACEAE 471

Aster arenosus (Heller) Blake

Aster ericaefolius var. tenuis Gray, Syn. Fl. 17: 198. 1884.
Leucelene arenosa Heller, Cat. N. Amer. Pl. ed. 1. 8. 1898.

Aster adenolepis Blake, nom. nov.

Machaeranthera mucronata Greene, Pittonia 4: 72. 1899. Not Aster mu-
cronatus Sheldon, 1903.

Aster aquifolius (Greene) Blake

Machaeranthera aquifolia Greene ex Woot. and Standl. Contr. U. 8. Nat.
Herb. 16: 188. 1913.

Erigeron perglaber Blake, sp. nov.

Perennis ut videtur suffrutescens 20 em altus pluricaulis sparse et obscure
stipitato-glandulosus primo visu glaberrimus; caules erecti tenues simplices
v. ramo unico erecto donati infra foliosi superne nudi; folia anguste spathu-
lata longe petiolata integerrima obtusa v. acutiuscula coriacea pallide viridia
1-nervia; capitula solitaria longiuscule pedunculata ca. 2 cm lata radiata;
involucre ca. 3-seriati paullum gradati 5 mm alti phyllaria anguste linearia
acuminata 0.3-0.5 mm lata appressa subchartacea costa glandulari-
incrassata minute sed dense stipitato-glandulosa apice sparse ciliata; radii
numerosi angusti sicco laete rosei; achenia compressa 2-nervia subappresse
hirsuta; pappus duplex, exterior e squamellis lineari-oblongis 0.5 mm longis,
interior e setis ca. 12 fragilibus ca. 3 mm longis compositus.

Basal leaves 4-5 cm long including petiole, the blade about 10-14 mm long,
2—2.5 mm wide, gradually decurrent on the petiole, this at base widened and
1-nerved; stems leafy to above the middle, the upper leaves similar to the
lower but smaller; petiole bases of old leaves persistent at base of stem;
peduncle (naked tip of stem) about 3.5-5.5 cm long, very slender, striate like
the stem; disk (as pressed) 6-8 mm high, 10-12 mm thick; rays 40 and more,
the tube 2 mm long, puberulous above, the lamina linear, 3-denticulate,
4-nerved, 10 mm long, 1—1.3 mm wide; disk corollas yellow, glabrous on tube
and teeth, densely puberulous on lower half of throat, 3.4 mm long (tube 0.6
mm, throat cylindric, 2.3 mm, teeth 0.5 mm long); achenes oblong, 1.5-1.8
mm long; inner pappus of about 12-14 fragile hispidulous bristles, the outer
of about as many linear or linear-oblong paleae, sometimes acute and entire,
sometimes obtusish and few-dentate, all free; style appendages deltoid, ob-
tusish, hispidulous.

Arizona: Without definite locality, 1869, H. Palmer (type no. 45116, U.S.
Nat. Herb.).

The data for Edward Palmer’s ‘“‘Arizona”’ specimens of 1869 are so uncer-
tain that this plant is ascribed to the State with some hesitation. However, it
appears to represent a species so distinct and unique as to merit description
despite some doubt about its habitat. In most of its characters it resembles
E. concinnus (Hook. and Arn.) Torr. and Gray so strongly that it might
almost be considered an essentially glabrous form of that species. The mate-
rial consists of two specimens, each with a decidedly woody, erect, once- or
twice-forked base about 3 cm high, possibly representing the upper part of a
much-branched caudex, clothed above with the marcescent bases of the peti-
oles of fallen leaves, and emitting several erect flowering stems. The glands

Bee cog

papvuie

= & ~

472 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

of stem and leaves are so minute that even under a lens the plant appears
glabrous or essentially so except on the involucre and upper part of the
peduncles, and even there the glands are very small; the leaves for the most
part seem to be completely glabrous.

Erigeron brandeget Gray, still known definitely only from the single scanty
specimen collected by T. 8. Brandegee on adobe plains, southwest Colorado,
in 1875, is intermediate between EF. concinnus and E. perglaber. The extreme
base of the specimen (a single stem with abortive terminal head and four
1-headed branches) is not present; the whole plant is finely and sparsely sub-
glandular-puberulous and more densely hirsute with mostly spreading hairs,
especially on the lower half of stem, leaves, and apex of peduncles; the in-
volucre is finely subglandular-puberulous, and sparsely spreading-hirsute on
the outer phyllaries; and the phyllaries are broader (0.50.8 mm) than in
E. perglaber. The pappus squamellae are long (often 1-2 mm), and irregu-
larly united with each other and with the inner bristles in a manner that
certainly indicates an abnormal condition, as Gray suggested.

Galinsoga semicalva var. percalva Blake, var. nov.

Achenia radii et disci glabra epapposa.

ARIZONA: Santa Rita Mountains, Pima County, David Griffiths & J. J.
Thornber 162 (type no. 497226, U. S. Nat. Herb.); Santa Rita Mountains,
1912, E. O. Wooton (U.S. Nat. Herb.).

With the narrow leaves and other characters of Galinsoga semicalva (Gray)
St. John and White, but with achenes of both ray and disk glabrous and
completely without pappus. In the typical form the disk achenes are hispidu-
lous and bear a well-developed pappus, those of the ray glabrous or usually
hispidulous above on the inner face and provided with a reduced pappus or
none.

Actinea quinquesquamata (Rydb.) Blake

Hymenoxys coopert subsp. grayz Cockerell, Bull. Torrey Club 31: 495. 1904
Hymenoxys quinquesquamata Rydb. N. Amer. Fl. 34: 114. 1915.

Gaillardia arizonica Gray var. pringlei (Rydb.) Blake

Gaillardia pringlec Rydb. N. Amer. Fl. 34: 139. 1915.
Gaillardia arizonica pringlei Blake in Tidestrom, Contr. U.S. Nat. Herb. 25:
598. 1925.

Artemisia carruthii Wood var. wrightii (Gray) Blake

Artemisia wrightit Gray, Proc. Amer. Acad. 19: 48. 1883.
Artemisia vulgaris subsp. wrightit Hall and Clements, Phylogen. Meth.
Taxon. 80. 1923.

Artemisia dracunculoides Pursh var. dracunculina (S. Wats). Blake

Artemisia dracunculina 8. Wats. Proc. Amer. Acad. 23: 279. 1888.
Artemisia dracunculus subsp. dracunculina Hall and Clements, Phylogen.
Meth. Taxon. 116. 1923.

Nov. 15, 1940 PEEBLES AND WHEELER: ARIZONA PLANTS 473

BOTAN Y.—Arizona plants: A new variety and new names and com-
binations.1 RoBpERT H. PrEsues, U.S. Bureau of Plant Industry,
and Louis C. WHEELER, University of Missouri. (Communi-
cated by T. H. KEARNEY.)

In recent numbers of this JoURNAL, several novelties were pub-
lished by the writers and their collaborators, in the course of prepara-
tion of a flora of Arizona. The need has arisen since for publication of
a few additional names.

Ephedra cutleri Peebles, nom. nov.

Ephedra coryt var. viscida Cutler, Ann. Missouri Bot. Gard. 26: 413. 1939.

In view of the ease with which E. cutleri can be recognized, its great abun-
dance, dominating habit, comparatively wide distribution, and doubtful
affinity with EL. coryz Reed, it seems proper to regard it as a specific entity.
According to Cutler (zbzd.), this Ephedra occurs in southwestern Colorado,
southeastern Utah, northwestern New Mexico, and northeastern Arizona,
thus being geographically distinct from EH. coryz, which has been reported
only from western Texas. Morphologically, it differs from E. coryz in having
the aerial stems more or less woody, frequently branched, often viscid, and
the bracts of the mature fruiting cones not fleshy. E. cutlerz seems to be most
closely related to E. viridis Coville, although differing from that species in
having shorter, often viscid stems, and long-peduncled ovulate cones. The
geographical distribution of E. cutler2 and E. viridis are mainly distinct, al-
though both species occur in Coconino County, Ariz.

Dalea amoena Wats. var. pubescens (Parish) Peebles, comb. nov.

Parosela johnsoni var. pubescens Parish, Bot. Gaz. 55: 308. 1913.
Psorodendron pubescens Rydb. N. Amer. FI. 24: 44. 1919.

Euphorbia palmeri 8. Wats. var. subpubens (Engelm.) Wheeler, comb. nov.
Euphorbia subpubens Engelm. in 8. Wats. Bot. Calif. 2: 76. 1880.

Euphorbia incisa Engelm. var. mollis (Norton) Wheeler, comb. nov.

Euphorbia schizoloba Engelm. var. mollis Norton, N. Amer. Euphorbia sect.
Tithymalus. 43. 1899 (preprint from Ann. Rep. Missouri Bot. Gard.
11: 127. 1900).

Opuntia whipplei Engelm. & Bigel. var. enodis Peebles, var. nov.

A forma typica tuberculis baccae latis humilibus distinguitur.

Type: North end of Hualpai Mountain, Mohave County, Ariz., altitude
4,200 feet, Peebles SF 883, April 19, 1935.

Although the fruit of Opuntia whipplez is described as not being very prom-
inently tuberculate, Engelmann and Bigelow evidently used the term in a
comparative sense. The illustration accompanying the original description
shows the fruit to be rather strongly tuberuclate, and it was thus interpreted
by Britton and Rose in their monograph of the Cactaceae. The usual form of
this common Arizona species bears prominent, high, laterally compressed
tubercles. In the variety enodis the tubercles are broad, low, and not prom-
inent. The variety is known only from the type collection.

Sterile fruits occur in typical O. whipplei and in var. enodis, but the pres-
ence or absence of seeds does not materially affect the appearance of the berry.

1 Received August 6, 1940.

474 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

ZOOLOGY.—The nematode genus Raillietnema Travassos, 1927.1
A. C. WauLTon, Knox College. (Communicated by E. W. Prics.)

The genus Razllietnema was established by Travassos in 1927 for
those species of cosmocercoid nematodes that show simple basically
amphidelphic uteri, a small number of relatively large ova, a vulva
opening posterior to the midregion of the body, a much simplified
male reproductive structure, subequal spicules, and a very small
accessory piece. Oxysomatium simples Travassos, 1925—from Hyla
faber, Brazil—was made the type species. In 1931 Travassos added
Aplectana loveridget Sandground, 1928—from Scolecomorphus vittatus,
Africa—to the genus, and the present paper adds a third species.
Semenow (1929) added Oxyuris praeputialis Skrjabin, 1916, to the
genus, but this form lacks an accessory piece and has complex ovarian
coils ending in uteri containing many eggs of relatively small size.
As was pointed out earlier (Walton, 1933) this species must be ex-
cluded from the genus Razllietnema as it is now constituted.

Restudy of material believed to be Razllietnema simples (from Hyla
faber—Brazil) and R. loveridger (from MScolecomorphus unicolor—
Tanganyika Territory, Africa) supports the grouping of the two spe-
cies into a single genus, and the finding of a third species with similar
characteristics in Scolecomorphus uluguruensis (from Lyingwa, Tan-
ganyika Territory, Africa) seems to justify the establishment of
Raillietnema as a valid genus of the family Cosmocercidae.

Raillietnema simples (Travassos, 1925) Travassos, 1927 Fig. 1

Examination of both male and female material from the type host, Hyla
faber (from Brazil), adds little to the original description except to point
out that the male possesses very narrow caudal alae, which extend from just
in front of the cloaca to a point midway along the length of the tail, and that
these alae are supported by 4 of the 10 pairs of papillae characteristic of the
species. Narrow lateral ridges are present in both sexes, extending from the
anterior esophageal to the postanal region. This character is present in all
three species of the genus. The excretory vesicle is heavily cuticularized, as
it is in the other two species, but is characteristically cylindrical, not spheri-
cal, in form.

Male.—Length, 1.6-1.75 mm; greatest width, 0.076—0.082 mm; pharynx
length, 0.029—0.032 mm; esophagus length, 0.37—0.405 mm; esophageal bulb
measures 0.052—0.057 mm by 0.052—0.057 mm, with a neck 0.047—0.05 mm
in length; head-nerve ring distance, 0.195-0.21 mm; head-excretory pore
distance, 0.3885-0.405 mm; cloaca-tail distance, 0.165—0.171 mm; spicule
length, 0.218—0.221 mm; accessory piece length, 0.0159—0.0163 mm; caudal
papillae arrangement, 2 pairs precloacal and 8 pairs postcloacal, with 1 pair
of precloacals and 3 pairs of postcloacals supporting the caudal alae.

1 Received June 20, 1940. Contribution from the Biological Laboratories of Knox
College, no. 68.

Nov. 15, 1940 WALTON: THE NEMATODE GENUS RAILLIETNEMA 475

One median unpaired precloacal papillus is present. Narrow lateral ridges
extend almost the entire length of the body.

Female.—Length, 2.35—2.72 mm; widthat vulva, 0.105—0.141 mm; pharynx
length, 0.034—0.038 mm; esophagus length, 0.49-0.51 mm; esophageal bulb
measures 0.07—0.072 mm by 0.07—0.072 mm, with a neck 0.05—0.053 mm in
length; head-nerve ring distance, 0.245—0.251 mm; head-excretory pore dis-
tance, 0.48—0.51 mm; anus-tail distance, 0.22—0.245 mm; vulva-tail distance,
1.19-1.38 mm; larvated eggs measure 0.09-0.12 mm by 0.22—0.24 mm; 2-4
eggs in each uterus. Narrow lateral ridges along almost the entire length of
the body.

Host.—Hyla faber—Brazil.

i Habitat.—Large intestine and the lower end of the small intestine of the
ost. |

Raillietnema loveridgei (Sandground, 1928) Travassos, 1931 Fig. 2

Study of both male and female material collected from three specimens
of Scolecomorphus unicolor (taken July 1939, at Magrotto Mountain, Tan-
ganyika Territory, Africa) adds but little to the original description. Both
sexes have narrow lateral ridges extending almost the entire length of the
body, and the excretory vesicle of each is subspherical in shape. One pair of
precloacal and one pair of postcloacal papillae support the caudal alae in the
male; the third pair of papillae is postalar in position. The number of ova in
each uterus is somewhat greater than in R. szmples, but is still relatively low.

Male.—Length, 1.81—2.01 mm; greatest width, 0.109—0.12 mm; pharynx
length, 0.021—0.032 mm; esophagus length, 0.363—0.4 mm; esophageal bulb
measures 0.055—0.08 mm by 0.055-0.08 mm, with a neck 0.06—0.07 mm in
length; head-nerve ring distance, 0.145—0.15 mm; head-excretory pore dis-
tance, 0.326—0.363 mm; cloaca-tail distance, 0.07—0.08 mm; spicule length,
0.175—0.2 mm; accessory piece length, 0.035-0.039 mm; caudal papillae
arrangement, one pair precloacal and two pairs postcloacal, with the last
pair caudad to the alae. One median unpaired precloacal papillus is present.
Narrow lateral ridges extend almost the entire length of the body.

Female.—-Length, 2.141-3.025 mm; width at vulva, 0.127-0.15 mm;
pharynx length, 0.024—-0.03 mm; esophagus length, 0.34—0.42 mm; esopha-
geal bulb measures 0.075—-0.09 mm by 0.075—0.09 mm, with a neck 0.07-
0.085 mm in length; head-nerve ring distance, 0.152—0.175 mm; head-excre-
tory pore distance, 0.35-0.375 mm; anus-tail distance, 0.09-0.115 mm;
vulva-tail distance, 0.6—0.95 mm; larvated. eggs measure 0.076-0.1 mm by
0.14-0.16 mm; 6-7 eggs in each of the uteri. Narrow lateral ridges extend
almost the entire length of the body.

Hosts.—Scolecomorphus unicolor, S. vittatus—Africa (Tanganyika Terri-
tory).

Habitat.—Large intestine of the host.

Raillietnema multipapillata, n. sp. Fig. 3

Scolecomorphus uluguruensis (taken at an altitude of 8,000 feet near Ly-
ingwa, Tanganyika Territory, Africa, in October, 1926) is the host for a
third species of Raillietnema. Both male and female specimens are available
for study.

Male.—Length, 1.82-1.83 mm; greatest width, 0.09-0.1 mm; pharynx
length, 0.027—0.03 mm; esophagus length, 0.33—0.345 mm; esophageal bulb
measures 0.07—0.08 mm by 0.07—0.08 mm, with a neck 0.07—-0.075 mm in

476 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 11

| Fig. 1.—Raillietnema simples: a, Esophageo-intestinal junction; 6b, lateral view
hy of tail of male. Fig. 2. Razllietnema loveridget: a, Esophageo-intestinal junction;
b, lateral view of tail of male. Fig. 3. Raillietnema multipapillata: a, Esophageo-
intestinal junction; b, lateral view of tail of male. (All drawn to the same scale.)

length; head-nerve ring distance, 0.17—0.178 mm; head-excretory pore dis-
tance, 0.28-0.3 mm; cloaca-tail distance, 0.12—0.13 mm; spicule length,
0.28-0.291 mm; accessory piece length, 0.036—0.041 mm; caudal papillae
arrangement, 10 pairs precloacal and 4 pairs postcloacal, with 7 pairs of
precloacals and 1 pair of postcloacals supporting the narrow caudal alae.
One median unpaired precloacal papillus present. Narrow lateral ridges
| extend practically the entire length of the body.

if

Nov. 15, 1940 WaLTON: THE NEMATODE GENUS RAILLIETNEMA 477

Female.—Length, 2.582—2.702 mm; width at vulva, 0.232—0.261 mm;
pharynx length, 0.033—0.037 mm; esophagus length, 0.36—-0.4 mm; esopha-
geal bulb measures 0.1—0.125 mm by 0.1-0.125 mm, with a neck 0.1—0.13
mm in length; head-nerve ring distance, 0.175-0.185 mm; head-excretory
pore distance, 0.36—0.371 mm; anus-tail distance, 0.161—0.169 mm; vulva-
tail distance, 1.075—-1.092 mm; nonlarvated eggs measure 0.109—-0.113 mm
by 0.125—0.13 mm ; 6-8 eggs in each of the uteri. Narrow lateral ridges extend
almost the entire length of the body.

Host.—Scolecomorphus uluguruensis—Tanganyika Territory, Africa.

Habitat.—Large intestine of host.

Type specomens.—Cotypes are deposited in the collections of the United
States National Museum (no. 42090).

Discussion.— Railletnema multipapillata may be separated from the other
two members of the genus on the bases of: (1) A larger total number of cau-
dal papilla, (2) by the greater number of papillae supporting the caudal alae,
(3) by the larger number of smaller ova, (4) by the longer spicules, and (5)
by the distinctive form of the heavily cuticularized excretory vesicle. R.
loveridger has the least number of caudal papillae and the shortest caudal
alae, the shortest spicules, and the most complex accessory piece of the three
species. The number of ova is intermediate and the excretory vesicle is sub-
spherical in form. In R. simples the number of caudal papillae and the size
of the caudal alae increase, the spicules are longer, the accessory piece is
much reduced, the number of ova is low, and the excretory vesicle is a short
cylindrical structure. R. magnipapillata has the greatest number of caudal
papillae, the longest caudal alae, the longest spicules, a large accessory piece
of simple structure, a larger number of smaller ova, and an excretory vesicle
shaped like a collapsed sphere with very heavy walls.

The genus Raillietnema at present consists of three species, two from Afri-
can examples of the Apoda and one from a Brazilian tree frog. It seems to
act as a connecting link between the cosmopolitan Oxysomatium-A plectana
group of quite complex species, on the one hand, and the extremely simpli-
fied, secondarily evolved, and geographically localized Schrankza species, on
the other. The species of this latter genus are reported only from one species
of Leptodactylus (L. pentadactylus) from Brazil. Only one or two eggs seem
to mature at any one time in the members of the genus, and the number of
spermatozoa is likewise greatly reduced. Oxysomatium appears to be nearer
the original stem of the cosmocercoids, and from this base has evolved
Raillietnema and then Schrankia. The reduction in the complexity of the
reproductive systems is regarded as evidence of secondary, not primary,
simplicity of the genera.

LITERATURE CITED

SANDGROUND, J. H. Some new cestode and nematode parasites from Tanganyika Ter-
ritory. Proc. Bost. Soc. Nat. Hist. 39(4): 131-150, 6 pls. 1928.

SEMENOW, W.D. Beitrag zur Charakteristik des Nematoden Raillietnema praeputiale
(Skrjabin, 1914). Zool. Anz. 86(5/8): 149-158, 5 pls.

2 a

478 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

SKRJABIN, K. J. Parasitic trematodes and nematodes collected by the expedition of Prof.
V. Dogiel and I. Sokolow in British East Africa. Sci. Res. Zool. Exp. Br. E. Afr.
and Uganda. V. Dogiel and I. Sokolow, 1914 (English translation). 1: 99-157,
10 pls. 1916.

Travassos, L. Contribuigdes para 0 conhecimento da fauna helmintolojica dos Batraquios
do Brazil. Trab. Inst. Oswaldo Cruz, Serv. San. Rur. Est. Rev., pp. 1-17, 10 figs.
1925.

. Schrankia brasili. Oxyuroidea de Batrachio. Bol. Biol. 9: 147-152, 4 figs. 1927.

. Pesquizas helminthologicas realisadas em Hamburgo. IX. Ensaio monographico
da familia Cosmocercidae Trav. 1925 (Nematoda). Mem. Inst. Oswaldo Cruz.
25(3): 2387-298, 43 pls. 1931.

Watton, A. C. The Nematoda as parasites of Amphibia. I. Journ. Parasitol. 20(1):
1-32, 1 pl. 1933.

ENTOMOLOGY.—The North American empoascan leafhoppers of
the alboneura group (Homoptera-Cicadellidae).1 Nancy H.
WHEELER,’ U.S. Bureau of Entomology and Plant Quarantine.
(Communicated by J. 8. WADE.)

In 1898, Gillette,? using external characters only, described the
widely distributed American leafhopper Empoasca alboneura. At the
same time he also described two other species of Empoasca, mexicana
and tumida, having certain external characters similar to those of
alboneura. In the revision of the genus Empoasca, DeLong‘ rede-
scribed the external characters of alboneura and included a description
and illustrations of the internal male genitalia of this species, pointing
out that twmida was a synonym and placing alboneura next to aspersa
in the subgenus Hebata. Later, DeLong and Davidson’ described and
illustrated confusa, which externally closely resembles alboneura but
internally is quite distinct.

The most distinctive external characters of alboneura and closely
related species are the more or less bluntly rounded crown and the
pale nervures of the forewings, the latter, by contrast, giving to most
of the species of this group a more or less striped appearance. The
species are all rather robust and range from 2.5 to 3.3 mm in length.
In size and shape they are somewhat similar to the species of the
aspersa group,® but are of a duller green color and lack the mosaic
pattern of the forewings that is characteristic of the aspersa group.

With a view to bringing together the various species of this rather
homogeneous group, of which alboneura is considered typical, a care-

1 Received June 19, 1940.

2 The writer expresses her appreciation to P. W. Oman, of the Division of Insect
Identification, Bureau of Entomology and Plant Quarantine, for his cooperation and
for furnishing specimens of species recognized by him as undescribed, and to Dr. R. H.
Davidson and Dr. D. M. DeLong for the loan of valuable type material.

$ Proc. U. 8S. Nat. Mus. 20: 748-744. 1898.

4U.S8S. Dept. Agr. Tech. Bull. 231: 35. 1931.

5’ Ohio Journ. Sci. 36: 229. 1936.

6 Proc. Ent. Soc. Washington 40: 133-147. 1938.

Nov. 15, 1940 WHEELER: LEAFHOPPERS OF THE ALBONEURA GROUP 479

ful study has been made of the specimens standing under the name
alboneura in the United States National Museum collection. As a
result, the type series has been found to be mixed, containing both
alboneura and confusa. From a further examination of available col-
lections for specimens resembling alboneura, two apparently unde-
scribed species have been segregated. The group is now known to con-
tain five different species, three of which have been previously de-
scribed and illustrated.’ For convenience, however, illustrations and
descriptions of all the species of the alboneura group now known to
occur in North America are included in this paper, together with some
synonymical changes. A key to the males of the group is also offered
as an aid in separating the various species.

KEY TO THE SPECIES OF THE ALBONEURA GROUP

ie Lateral processes converging distally....... 4. confusa DeL. and D.
Waverale processes divercine distally. - 05 0....0.. 025-27... oe 2

2 (1). Dorsal spine vestigial. Lateral process very broad basally, in lateral
view tapering abruptly and strongly curved dorsad.............

ER ee ae Sate OU Ra ee Le Wee 1. sonorana, Nn. sp.

Dorsal spine prominent. Lateral process relatively slender basally, in
lateral view more or less sinuate, with sides almost parallel to near

SE OCTESO| DIOR. tee erecta aa eet ict ol Aire erg cee 3

3 (2). Aedeagus with two pairs of spinelike processes projecting dorsocau-
dad. Apex of dorsal spine short, bluntly pointed. .5. mexicana Gill.
Aedeagus without spinelike processes. Apex of dorsal spine attenu-
BCOMOMSMALD IY OMIM =) ih 2 Ay oe ce ko oka e eee ee eee 4

4 (3). Dorsal spine sickle-shaped, apex attenuated. Lateral process, in lat-
eral view, with both proximal and distal portions definitely sinuate;

distal portion convexly curved on dorsal margin. .3. alboneura Gill.

Dorsal spine not sickle-shaped, broad basally, abruptly narrowed
medially, and terminating in a slender, pointed, toelike projection.

Lateral process, in lateral view, with proximal and distal portions

only slightly curved; distal portion concavely curved on dorsal

ODEN ROR ia cas 2 so 3 ue io eae I 2. stylata, n. sp.

Empoasca sonorana, n. sp. Fig. 1

Resembling alboneura in size and general structure, but lacking the
strongly contrasting veins of forewing; lateral process very broad basally
and dorsal spine vestigial. Length, 3.25 mm.

External characters—General color pale sordid green; head, thorax, and
scutellum sometimes with irregular, paler areas. Posterior margin of seventh
sternite of female faintly incised medially and faintly sinuate each side of
median incision.

Male internal structures.—Lateral process very broad basally, tapering

7 Subsequent to the submission of this paper for publication it was found that
Empoasca knulli, described by Davidson and DeLong (Ohio Journ. Sci. 39: 111. 1939)
is a member of the alboneura group. The original description, in which the species was
compared with similis and bicuspida (members of the aspersa and fabae groups, re-
spectively), appeared without accompanying illustrations. Empoasca knulli may be
differentiated from the other members of the alboneura group, here treated, by the
bifurcate dorsal spine.

480 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

rather abruptly but evenly, in lateral view to sharply pointed apex curved
dorsad, in ventral view to bluntly pointed apex directed caudolaterad. Style
long, relatively slender, gradually tapering to sharply pointed apex directed
caudolaterad. Dorsal spine vestigial. Aedeagus slender, extending upward
from base, then bent directly caudad and broadly curved dorsad. Sternal
apodemes rudimentary.

Holotype male, allotype female, and 3 male and 20 female paratypes from
Cajene, Sonora, Mexico, June 1, 1927, A. W. Morrill, collector.

Type, U.S.N.M. no. 54190.

Empoasca stylata, n. sp. Fig. 2

Related to alboneura, but larger, with venation of wings less conspicuous,
dorsal spine stouter, and lateral process not broadened distally in ventral
view. Length, 3.3 mm.

External characters.—General ground color pale sordid green tinged with
fuscous; venation of wings obscure except apically. Crown with ivory spots
and an ivory median line. Pronotum marked with three irregular ivory areas
anteriorly.

Male internal structures.—Lateral process long, slender, with sides almost
parallel to near pointed apex, with distal portion in lateral view concavely
curved on dorsal margin and directed dorsocaudad, in ventral view curved
caudolaterad. Style long, relatively broad at base, with tip bent laterad.
Dorsal spine strongly curved first caudad, then ventrad and cephalad, broad
basally, narrowed medially, and terminating in a slender, pointed, toelike
projection extending cephalomesad from ventrocephalic margin. Aedeagus
stout, process for muscular attachment extending at nearly right angles to
shaft. Sternal apodemes fuscous, large, sacklike, and rounded apically.

Holotype male and one male paratype from Cajon Pass, Calif., June 6,
1935, collected by P. W. Oman, and one male paratype from Los Angeles,
County, Calif., collected by D. W. Coquillett. The last-mentioned specimen
was standing under the name alboneura in the Museum collection.

Type, U.S. N. M. no. 54191.

Empoasca alboneura Gillette Fig. 3

Empoasca alboneura Gillette, Proc. U. 8S. Nat. Mus. 20: 743-744. 1898.

A relatively robust, dull-green species characterized by the pale nervures
of the forewings, giving a conspicuously striped appearance. Lateral
processes definitely sinuate and diverging distally; spine stout and sickle-
shaped. Length, 2.5 to 3.5 mm.

External characters.—General color varying from yellowish green tinged
with orange to dull green tinged with brown, veins pale and conspicuous.
Crown relatively broad, bluntly rounded and slightly produced, with a
light median stripe and an oblique dash on each side near the eye. Posterior
margin of seventh sternite of female produced and rounded medially, with
a very shallow and broad lateral emargination.

Male internal structures.—Lateral process in lateral view long, relatively
slender and sinuate, with distal portion convexly curved on dorsal margin
and tapered to pointed apex directed dorsocaudad; in ventral view, curving
mesad to near middle, then laterad, with distal portion slightly broadened,
then narrowed to pointed apex directed caudolaterad. Style long, with
basal portion relatively broad, sinuate and abruptly narrowed near middle,
with apical portion broadly curved and gradually tapered to bluntly pointed
apex directed caudolaterad. Dorsal spine relatively stout and sickle-shaped,

Nov. 15, 1940 WHEELER: LEAFHOPPERS OF THE ALBONEURA GROUP 481

ooorernes

oes
- .

a ne Td

Yq SONORANA

Wi

\
Mh Vf XN
i if Vy e
ae Hh

'
,
,
'
' D
1 ‘ ae:
1
. 2cocre”
# --reee
.

oe ‘STYLATA

3 ALBONEURA
ae oo es er

.
.
‘
‘

one
perl mene

wo ole ®

5 are ceawh

Figs. 1—5.—Lateral and ventral (A) views of male genital capsule and ventral view
(B) of sternal apodemes of (1) Empoasca sonorana, n. sp.; (2) HE. stylata, n. sp.; (3)
E. alboneura Gill.; (4) EF. confusa DeL. and D.; and (5) E. mexicana Gill. Xca. 60.

a

—

482 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

broadly curved and evenly tapered to attenuated apex directed ventro-
cephalad. Aedeagus broad distally. Sternal apodemes rather small, narrowed
on lateral margins to bluntly rounded and slightly fuscous apices.

Originally described by Gillette from 15 females and 9 males from Mis-
sissippi Agricultural College, 2 specimens from Virginia, and 7 specimens
from Colorado; specimens were also recorded from ‘‘D. C.,’”’ and from the
U. 8. National Museum marked ‘‘Va.,” ““Los Angeles, Calif., Coquillett,”’
“Nevada Co., Cal., Sept.,” “Horace, Ks., July 28,” and ‘‘Neb.,” the type
being designated ‘‘No. 3437 U.S. N. M.”

The type series now extant in the U.S. National Museum consists of two
females with the Type no. 3437, one labeled Horace, Kans., and the other
Lakin (?), Kans. Therefore, Gillette’s cotype female in the U. 8. National
Museum (Type no. 3437) from Horace, Kans., is here designated lectotype.

More recently this species has been identified in miscellaneous collections
from Alabama, California, Colorado, Connecticut, District of Columbia,
Kansas, Kentucky, Maryland, Mexico, Missouri, Montana, Nevada, New
Mexico, New York, North Carolina, Ohio, Oregon, Pennsylvania, Tennes-
see, Texas, Utah, Virginia, and Wyoming.

In the original description Gillette reported alboneura from ‘“‘native
plants” and from plum in Colorado. Later, DeLong‘ recorded this species
as “distributed almost throughout the United States on herbaceous plants.”
Other records have been procured, from collections in various localities, of
the occurrence of alboneura on alfalfa, Anthemis cotula, Artemisia, arti-
chokes, aster, Chrysothamnus, giant ragweed, and white pine. This species
has also been taken in trap lights at Arlington, Va., Knoxville, Tenn., and
Richfield, Utah.

The accompanying illustrations and descriptions of this species have been
made from a study and comparison of a number of males and females from
series collected both in the East and in the West. There seems to be some
variation in this species, both externally (in size and coloration) and in-
ternally Gn the breadth and sinuation of the apical portion of the lateral
process), but no greater differences have been noted between eastern and
western specimens than between specimens of a series from the same locality
and date of collection.

Empoasca confusa DeLong and Davidson Fig. 4

Empoasca confusa DeLong and Davidson, Ohio Journ. Sci. 36: 229. 1936.

Superficially resembling alboneura but less conspicuously striped and less
robust, with crown narrower and only slightly produced; lateral processes
sinuate but converging distally. Length, 3 mm.

External characters—General color fuscous-green, with venation of fore-
wing pale. Posterior margin of seventh sternite of female produced and
broadly rounded medially, with a small notch, on each side, laterally.

Male internal structures.—Lateral process long and sinuate; in ventral
view broad, with distal portion narrowed to long slender point directed
caudomesad; in lateral view more slender and tapered gradually to pointed
apex directed caudad. Style relatively long, broad at base, with distal por-
tion abruptly narrowed to long, slender apex directed caudolaterad in
ventral view. Dorsal spine short, broad, concave on anterior margin, with
a bluntly pointed, toelike projection extending cephalad from ventro-
cephalic margin. Aedeagus simple and relatively slender. Sternal apodemes
rudimentary.

Originally described from a single male collected at Burns, Oreg., holo-

Nov. 15, 1940 WHEELER: LEAFHOPPERS OF THE ALBONEURA GROUP 483

type male in the DeLong collection. The male and female herein treated,
the former redescribed and reillustrated, are from a series of 18 males and
48 females collected at Mount Shasta, Calif., by R. H. Beamer. Other
males and females at hand are from the following localities: CALIFORNIA:
Bray (Oman). ConLorapo: Durango, Livermore (Oman); Mesa Verde
(Beamer); Poudre River Canyon (Sanderson). NEvapa: Reno (Beamer).
OrEGON: Bend (Oman). WasHINGTON: Cliffdell (Beamer); Mount Rainier
(Oman). Four females of this species from Nevada County, Calif., were
found under alboneura type no. 3437 in the U. 8. National Museum
collection.

Empoasca mexicana Gillette Fig. 5

Empoasca mexicana Gillette, Proc. U. 8. Nat. Mus. 20: 737-738. 1898.
Empoasca tumida Gillette, Proc. U. S. Nat. Mus. 20: 744. 1898.
Empoasca bitubera DeLong, Ohio Journ. Sci. 32: 395. 1932.

One of the smallest species of the alboneura group, with crown rounded,

but shghtly more produced than in alboneura. Aedeagus with two pairs of -

spinelike processes. Length, 2.5 to 2.75 mm.

External characters.—Color yellowish to fuscous-green, with veins of fore-
wing pale but not so distinct as in alboneura, except apically. Posterior
margin of seventh sternite of female slightly produced and rounded medially,
with slight lateral emargination.

Male internal structures.—Lateral process relatively long, broad, and
tapered apically; in lateral view extending dorsocaudad, with distal portion
narrowed on dorsal margin and curved upward; in ventral view sinuate,
curved mesad about middle, with distal portion broadened slightly, then
tapered to sharply pointed apex curved laterad. Dorsal spine broad at base,
broadly curved caudoventrad, with bluntly pointed apex directed cephalo-
mesad. Aedeagus unusual, having a pair of long, narrow, spinelike processes
projecting dorsocaudad from near base and another, shorter pair arising
similarly but more dorsally. Sternal apodemes medium sized, slightly
fuscous, and rounded apically.

This species was originally described by Gillette from five females col-
lected near Veracruz, Mexico, and was given U. 8. N. M. type no. 3430.
Later, DeLong redescribed mexicana, in the absence of Gillette’s type ma-
terial, from specimens in the collection of the U. 8. National Museum,
labeled Marfa, Tex., June 5, 1908, Mitchell and Cushman, collectors. As
previously indicated by the writer,’ from an examination of this series now
extant, the external markings and internal structures of the male were found
to be identical with those described and figured later by DeLong for bitubera,
and bitubera, therefore, was placed as a synonym of mexicana.

An examination of the type of tumida, a female in the U. S. National
Museum, labeled ‘‘Colo. 1658” (type no. 3488), has shown this form to be
specifically identical with females from Marfa, Tex., identified as mexicana
by DeLong. Therefore, Empoasca tumida is also placed as a synonym of
E. mexicana.

For the sake of uniformity, mexicana is herein redescribed and reillus-
trated from the series collected at Marfa, Tex., June 5, 1908, now consisting
of 1 male (dissected), 3 females, and 2 specimens with abdomens missing.
Other material has been examined from various localities in Arizona,
California, Colorado, Nevada, New Mexico, Texas, and Utah.

8 Journ. Washington Acad. Sci. 29: 300. 1939.

484 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

PROCEEDINGS OF THE ACADEMY AND AFFILIATED
SOCIETIES

THE ACADEMY
362D MEETING OF THE BOARD OF MANAGERS

The 362d meeting of the Board of Managers was held at the Cosmos Club
on Friday, June 28, 1940. President Crittenden called the meeting to order
at 8:04 p.m. There were 11 present as follows:

E. C. CrittENDEN E. W. Prick

F. D. Rossint1 C. L. GARNER

F. C. KrRacek W. W. DirHu

H. 8S. RaAPPLEYE and by invitation
A. H. CLark F. G. BRICK WEDDE
P. C. WHITNEY N. R. Smit

President CRITTENDEN announced the appointment of the following com-
mittees:

COMMITTEES ON AWARDS FOR SCIENTIFIC ACHIEVEMENT FOR
THE CALENDAR YEAR 1941 ;

For the Biological Sciences

A. H. Cuark, chairman Il, 40, Jalan

F. O. Cor C. F. W. MugseBeck
J. M. CoorER H. W. ScHOENING

H. A. Epson G. STEINER

E. A. GOLDMAN A. WETMORE

For the Physical Sciences

O. H. Gisu, chairman F. L. MouLER
A. K. Baus W. T. ScHALLER
F. 8S. BRACKETT J. H. Taytor
W. E. DEMING O. R. WuLF

H. E. McComs E. G. Zins

For the Engineering Sciences

F. M. Dreranporr, chairman H. N. Eaton
H

C. H. BirDSEYE A. C. FIELDNER
H. L. Curtis H. C. Havers
H. G. DorsEy G. W. MusGRAVE

W.N. SpaARHAWK

COMMITTEE ON MEETINGS TO SERVE UNTIL JUNE 1941

C. L. GARNER, chairman W. C. LowDERMILK
R. E. Gipson P. A. SmitH
L. V. JuDson CHARLES THOM

Chairman N. R. Smrru, of the Committee on the Records of the Academy,
presented the following recommendations, which were accepted by the
Board with instructions to the committee to prepare the appropriate amend-
ments to the bylaws for submission to the members:

1. That there be created the office of Archivist, whose duties would be: First, to
go over the present accumulation of material, sort out the important records, and

Nov. 15, 1940 PROCEEDINGS: GEOLOGICAL SOCIETY 485

arrange and keep them in permanent order; second, accept and care for the records
turned over to him by the Corresponding Secretary each year.

2. That the present duties of the Corresponding Secretary as regards the main-
tenance of the Academy’s permanent records be transferred to the Archivist.

3. That the office of Recording Secretary be abolished, his purely secretarial
duties transferred to the Corresponding Secretary, and such permanent records as
are now kept by him transferred to the Archivist.

4. That the name of the office of Corresponding Secretary be changed to Secretary.

Under the proposed plan, the Archivist would have charge of the following records
which are now in the hands of the Corresponding Secretary:

. A set of the Proceedings of the AcADEMyY.

. A set of the bound Journals.

. A set of the Directories (incomplete, 1900 to 1910).
Permanent Minutes of the Board of Managers.

. A record of Academy officers and organization.
Nominations for membership.

. Important correspondence.

. Material of permanent interest, medals, invitations, ete.

DONA MAP WN He

Your committee further recommends that the Archivist be appointed by the
President for a period of three years and that the person selected for the position be
one who is interested in this type of work and who has had appropriate experience in
the affairs of the ACADEMY.

On the recommendation of the Committee, of which H. L. Curtis was
chairman and CHARLES THoM and P. C. WHITNEY members, the Board
approved for presentation to the AcADEMY a revision of the bylaws entitled
‘‘Amendment to modify the procedure for amending the bylaws’’:

ArtIcLE IX. Delete lines 1 to 10 and insert the following:

‘“‘Amendments to these bylaws shall be proposed by the Board of Managers and
adopted by a letter ballot of the members of the AcapEmy, with a two-thirds majority
of those voting being required for adoption. When a proposed amendment is sent to
the members, there shall also be submitted a statement of the reasons for the proposed
amendment. The ballots shall be counted one month after mailing to the members.

‘“‘Any affiliated society or any group of ten or more members may request, in
writing, the Board of Managers to propose an amendment. The action of the Board in
accepting or rejecting this request shall be by a verbal (yea and nay) vote on a roll
call, and the complete roll call shall be entered in the minutes of the meeting.”

The Corresponding Secretary presented for the Committee on Member-
ship the nominations of 17 persons; one resident and 16 nonresident.
_ After fixing the next meeting for September or October, the Board ad-
journed at 9:50 P.M.

GEOLOGICAL SOCIETY

569TH MEETING

The 569th meeting of the Society was held at the Cosmos Club, January
11, 1939, President J. B. Mrrtin, Jr., presiding.

Informal communications.—T. 8. Kester described some unusual rock
structures in northern Georgia.

Program.—C. WytTHE Cooke: Neptune’s racetracks.

EK. CauuaGHAN: Recent fault scarps in the western part of the Great Basin.

C. B. Hunt: Pediments around the Henry Mountains, Utah.

486 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

070TH MEETING

The 570th meeting of the Society was held at the Cosmos Club, January
25, 1939, President J. B. Mertts, JR., presiding.

Program.—F. P. SHeparp: Nondepositional environments off the California
coast.—Bottom sampling by P. D. Trask and reports of bottom conditions
coming from the work of the U. 8. Coast and Geodetic Survey gave the
impression that there were considerable areas off the California coast where
deposition is either very slow or nonexistent. Recent work on the E. W.
Scripps, involving dredging and coring operations over a wide area off this
coast, has shown that these nondepositional surfaces are widespread, that
they are associated with various topographic environments, and that all
depths up to 3,500 meters are involved. The environments having either rock
bottom or sediment that appears to be too coarse to be introduced at the
present time include: Banks or submarine mountains rising above any part
of the sea floor; steep submarine slopes; portions of the continental shelves;
and the walls and floors of submarine canyons. With the possible exception
of some banks, none of these nondepositional areas were entirely free from
recent sediments, but coring and dredging operations gave the indication
that the recent sediments in these places form only a thin discontinuous
cover over rock or sediments of an older generation.

While basins and troughs off the southern California coast appear to be
areas where deposition is important, several of these features were found to
have relatively clean sand deposits underneath thin covers of mud. Since
silt and clay are known to be the chief sediments contributed at the present
time to the basins and troughs, the existence of this sand leads to the sus-
picion that even in some of these deep depressions deposition of fine grained
sediment may have been inhibited in relatively recent time.

The nondepositional surfaces are found for the most part at depths too
great for effective wave agitation. However, investigation of the bottom cur-
rents by R. Revelle and the speaker in all the principal environments has
shown that large eddy movements are taking place with sufficient current
velocity to move fine-grained sediment. These currents show no sign of de-
creasing velocity with increased depth. Furthermore, the discovery of rela-
tively rapid increase in depth from time to time at the heads of submarine
canyons and that these depth changes may occur during calm sea conditions
suggests that mud flows or other types of submarine landslides are carrying
sediments seaward. It seems probable that bottom currents and mass gravity
movements are responsible for the nondepositional surfaces.

ErRuING Dorr: The stratigraphic significance of the flora of the type Lance
formation.—The Lance formation at its type locality in Niobrara County,
Wyo., has previously yielded a dinosaur fauna of the Triceratops stage, a
fresh-water invertebrate fauna, and a small flora. The flora has never been
described or fully studied. During the past two summers a large collection
of fossil plants was obtained in this area by the speaker, under joint auspices
of Carnegie Institution of Washington and Princeton University.

At present this flora comprises about 75 species, of which 8 are apparently
new and 6 are not specifically determinable. More than 75 percent of the
species have previously been reported elsewhere. Over 80 percent of these
species occur only in formations whose Upper Cretaceous age is not in doubt:
the Laramie of Colorado, the Medicine Bow of Wyoming, and the Hell
Creek of Montana and the Dakotas.

Only about 10 percent of the species are known from the widespread and
well-known Fort Union flora of Paleocene age. Equally distinct from the

Nov. 15, 1940 PROCEEDINGS: GEOLOGICAL SOCIETY 487

type Lance flora is the flora of the non-dinosaur-bearing beds (Tullock and
Ludlow), which overlie the dinosaur-bearing beds (Hell Creek) in Montana
and western Dakotas. These have previously been considered the upper part
of the Lance formation. The known flora of over 50 species of the Tullock
and Ludlow is clearly of Paleocene aspect, with about 85 percent of the
species recorded only from the overlying Fort Union.

It is apparent that published statements that point out similarity between
the Lance and Fort Union floras are erroneous and are based on inclusion in
the Lance flora of species known now to have come from horizons in the
Tullock or Ludlow of post-Lance age, or from beds that are actually basal
Fort Union. The paleobotanical evidence indicates that the Mesozoic-
Cenozoic boundary should be placed between the dinosaur-bearing beds
(type Lance or Hell Creek) and the non-dinosaur-bearing beds (Tullock
or Ludlow). This is in harmony with the known vertebrate evidence and
does not appear to be seriously contradicted by the marine invertebrate
fauna (Cannonball) associated with the Ludlow of the western Dakotas.

F. E. Matruess: Post-Pliocene origin of the glaciers of the Sierra Nevada.

5718ST MEETING

The 571st meeting of the Society was held in the Cosmos Club, February
8, 1939, President J. B. Merrie, JR., presiding. The following resolution
presented by J. S. WILLIAMS, seconded by J. B. Rersip#, JR., was unani-
mously approved by the Society:

“Mr. President: Dr. GEoRGE HERBERT GIRTY, a member of this Society
since January 1897, died of heart failure at his home in this city on Friday,
January 27. He had recently passed his 69th birthday.

““Dr. Girty was well known to most of the members of this Society either
personally or professionally as a specialist on Mississippian, Pennsylvanian,
and Permian fossils. In that capacity he has cooperated, both in the labora-
tory and in the field, with many members in the solution of geologic problems
in many States and in Alaska. The demands of his position made him spend
considerable time on stratigraphic paleontology, but his greatest interest
was in the field of descriptive paleontology, and he used to sit by the hour
often with the bright summer sunshine streaming in through an open
window, dictating descriptions of species to his stenographer. Never a show-
man in his professional work, many observations of great importance are
rather apologetically made and obscurely placed within the texts of his de-
scriptions of species. Many of these have been given prominence by later
writers. Other observations of equal importance have never been written,
because Dr. Girty belonged to the school of geological thought that believes
that principles, correlations, and classifications should be based on evidence
that is well in hand before they are enunciated. This fact, together with the
fact that so much of his time was employed in examining and writing reports
on field collections, accounts in part for the relatively small proportion of
philosophical papers he published.

“Dr. Girty was author or part author of 9 professional papers and bulletins
of the U. 8. Geological Survey and of about 50 articles published in various
journals and State survey reports. His interests were wide, and at the time
of his death he had but recently completed or was working on papers de-
scribing fossils from the Mississippian, Pennsylvanian, or Permian rocks
from many scattered areas extending from Pennsylvania and Ohio on the
east to the Pacific Coast States on the west; and from Texas on the south to
Alaska on the north. Perhaps his work best known the world over was his

488 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

Guadalupian report, which gave the first extensive description of the Per-
mian faunas of the Southwest. Other papers, however, are known almost as
widely. He has contributed through descriptive papers or reports to the
knowledge of the Carboniferous geology of almost every State and territory
in the United States in which Carboniferous deposits occur and also of
Canada, Mexico, and countries on other continents.

‘““A short biographical sketch written by Dr. Girty a few years ago at the
invitation of a British Journal gives many personal details of his life. In it
he tells that he was born in Cleveland, Ohio, on December 30, 1869. His
parents appear to have been extremely well to do, and from the beginning
he lived a rather sheltered life. A great shock, which he says profoundly in-
fluenced his life, came when he was 12 years old, when his father died. He
was educated in the Cleveland public schools and in Yale University, re-
ceiving his Ph.D. degree there in 1894. He came to the Geological Survey
in 1895. He was very fond of music and was an accomplshed pianist. In his
early youth he considered selecting music as a career, but he gave it up
because he felt that his talents were inadequate—‘‘not much above medioc-
rity,’ he says. He wrote several compositions for his own pleasure—some
of them during his last illness, but he published none of them. He was also
fond of poetry and himself wrote poetry. When he started in his college
career, he had, as he put it, ‘‘an idea’’ that he would like to teach Latin and
Greek, both of which, especially Greek, made a strong appeal to him. His
proficiency in these subjects later made him a frequent consultant of Survey
members. In his senior year in college he became interested in paleontology,
and this interest, as we all know, continued to be his most ardent interest
until his death.

“With Dr. Girty’s passing, the Society and the science of paleontology
have lost an extremely able contributor, and many members of the Society
have lost a sincere friend who will be sadly missed. Therefore, I move that
these words of appreciation be incorporated in the minutes of this meeting
and the Secretary of the Society be instructed to convey to Mrs. Girty the
sincere sympathy of the membership of the Society.”

Informal communications.—F. E. Mattues described some artificial
geysers near Mono Lake in California, resulting from wells.

Program.—Harry H. Hess: Gravity anomalies and structure of the West
Indian Island arcs.

J.C. Resp: Mineralized faults in the Chichagof mining district, Alaska.

0/2D MEETING

The 572d meeting of the Society was held at the Cosmos Club, February
22, 1939, Vice-president, JoHN B. REeEsIDE, Jr., presiding.

Informal communications.—RoBERT P. Bryson described some crater-
shaped depressions in reservoirs, whose origin seems to be associated with
gas bubbles that rise through the sediments.

Program.—k. L. StePHENSON, The results of magnetometer surveys on lac-
coliths in the Highwood Mountains, Mont.—Magnetometer measurements
were made on and near the Shonkin Sag and Cowboy Creek laccoliths in
the Highwood Mountains in Montana, to trace the buried extension of a
horizontal pipelike body and to locate, if possible, the feeders of the lac-
coliths. Approximately 2,500 readings were made with a standard Askania
vertical variometer.

By means of magnetic anomalies the horizontal pipelike body was traced,
with reasonable certainty, for a distance of 9,000 feet under a Tertiary gravel
bench to a dikelike body exposed at the opposite side of the bench near the

Nov. 15, 1940 PROCEEDINGS: GEOLOGICAL SOCIETY 489

Cowboy Creek laccolith. The measurements show no connection between the
laccolith and these two bodies, which, therefore, are not feeders of the
Cowboy Creek laccolith.

Areas of great magnetic disturbance occur over parts of each of the two
laccoliths. The general form of the anomalies is that commonly associated
with relatively narrow bodies of considerable vertical extent. These anoma-
lies may indicate the presence of feeders of the laccoliths, and no other
explanation seems more probable at present; however, because other condi-
tions might produce similar anomalies, only tentative conclusions are
drawn pending the completion of additional measurements.

JAMES L. Dyson: Structure and motion of cirque glaciers.— Detailed study
of two cirque glaciers, Sperry and Grinnell, in Glacier National Park has
shown that a small cirque glacier in an extreme state of decadence exhibits
its structure and other glacial phenomena very clearly. Such glaciers have
motion, deposit morainic material, and perform other typical glacier func-
tions. Sperry and Grinnell glaciers are both about the same size, each have
a length of approximately one mile.

Stratification due to the annual accumulation of snow is well exhibited on
these glaciers. The ice strata, each of which represents one season’s snowfall,
range in thickness from 2 to 6 feet and are readily apparent because their
upper surfaces are covered with dust and small rock particles which make
visible contact planes between each stratum. In the narrow firn zone, which
lies at the base of the cirque headwall and comprises usually about 8 percent
of the surface of the glacier, layers of snow are wedge- or shingle-shaped and
have a steep initial tilt toward the glacier front. Weight of the overlying
layers causes these wedge-shaped layers of snow to move downward and to
rotate so that after traveling a relatively short distance they assume a back-
ward dip, which gradually increases until about midway between the firn-ice
boundary and the glacier front a maximum of 45° may be attained. Near the
firn-ice boundary, ice strata usually are nearly horizontal or have rotated
sufficiently to possess a gentle dip toward the cirque headwall. The thin
part of each wedge is removed by melting, and only ice which was originally
next to the headwall reaches the glacier front. On both Sperry and Grinnell
glaciers the edges of ice strata are distinct enough so that between the firn
area and the glacier front their number may be ascertained. Thus the ap-
proximate length of time necessary for ice at the glacier front to have moved
from the cirque headwall may be determined.

In spite of the small size of these glaciers they are actively eroding the
surfaces over which they move. This activity is made manifest by con-
spicuous amounts of rock flour in their melt waters and by abundant striae
on rock surfaces from which the ice has recently retreated. The nature of
many of these striae indicates that in places the ice in a very thin zone at the
base of the glacier is quite plastic. Because of obstructions in the cirque
floor many of the striae are extremely sinuous. Boulders were found that,
after being transported in the basal portion of the glacier, were forced into
crevices in the cirque floor from which the ice was unable to extricate them.
Subsequently the ice was deflected around these boulders as indicated by
striae that approach a boulder, curve around it, and then resume a course
in direct line with their original trend.

It is suggested also that plasticity of the ice and to a lesser extent its ero-
sive power, depend not so much upon the thickness of the ice as upon the
type of internal motion it exhibits.

C. F. BrockMan: Glacial recession in Mount Rainier National Park.

on ot oe eee ee

490 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

573D MEETING

The 573d meeting of the Society was held at the Cosmos Club, March 8,
1939, President J. B. Murrtin, Jr., presiding.

Informal commumncations—W. W. Rusry described the work of an ad-
visory committee of naturalists in preserving and making accessible features
of interest in the new Government park being developed along the old
Chesapeake and Ohio Canal and asked members to help by drawing atten-
tion to features that they considered should be pointed out.

R. E. StEvEns presented the results of an analytical study of intergrown
albite and mica from a California pegmatite, in which although the two
minerals had apparently crystallized simultaneously, the mica contained 10
percent potash and only 1 percent soda, a feature that has a bearing on the
existence of the soda mica paragonite.

Program.—MarGaret D. Foster: Geochemical relations of ground waters
in the Houston-Galveston area, Tex.—Ground-water supplies in the Houston-
Galveston area are obtained principally from permeable sands in the
Willis (?), Lissie, and Beaumont formations. The shallow waters in the dif-
ferent formations differ characteristically in their content of calcium bi-
carbonate and in their total mineral content. These differences may be
correlated directly with the amount and character of soluble material in the
sands and clays from which the waters are derived and with the organic
content of the soils through which the waters passed in entering the forma-
tion.

As they pass downward, the waters in all the formations show a progres-
sive alteration in character, changing from calcium bicarbonate to sodium
bicarbonate—a result, apparently, of reaction between the waters and base-
exchange minerals in the sands. In waters in the Willis (?) sand, this altera-
tion in type is accomplished by a continuous increase in total mineral con-
tent and in sodium bicarbonate; in the waters in the Lissie and Beaumont
formations there is a gradual decrease in calcium and magnesium, accom-
panied by a gradual increase in sodium, the total mineral content remaining
fairly constant. |

The depth at which the change in character begins to take place differs
in the different formations; it is greatest in the Willis (?), least in the Beau-
mont. Calcium chloride waters found in the vicinity of some of the salt domes
may be attributed to the reverse of this softening process; deep salt water, —
moving upward through faults and fissures and invading the surface strata,
reacts with exhausted shallow base-exchange minerals, exchanging its
sodium for their calcium and magnesium. Distortion of the strata is indi-
cated by the chemical character of some of the waters in the vicinity of
Blue Ridge and Pierce Junction domes.

J.S. Vuay: Structural features on the north side of the Beartooth Mountains
near Nye, Mont.—From a study of the geology of the Nye #2 quadrangle on
the north side of the Beartooth Mountains, Mont., it is concluded (a) that
relief to compression during the Laramide orogeny was localized by major
zones of weakness in the basement rocks to form two zones of complex struc-
tures, and (b) that porphyry bodies along the north zone were intruded
previous to the period of major deformation.

The Beartooth Mountains in the southern part of the area studied consist
mainly of pre-Cambrian rocks. Along the approximately east-west trending
front are steeply dipping Paleozoic, Triassic, and Jurassic formations, simi-
lar to those found elsewhere in southern Montana. In the western part of
the area the mountains swing to the north and are made up for the most part —

Nov. 15, 1940 PROCEEDINGS: GEOLOGICAL SOCIETY 49]

of Paleozoic rocks intruded by numerous porphyry masses. The central and
eastern part of the area consists of the rather broad valley bottoms of the
Stillwater River and its tributaries and intervening terraces and rolling,
moderately dissected pediments; it is underlain by Upper Cretaceous
formations. To the northeast is an upland of considerably greater relief
developed on the pyroclastic Livingston formation.

Regional structural studies have shown that the Beartooth block over-
thrusts in an easterly direction east of the Nye area, whereas low-angle
underthrusting toward the northeast occurs west of this area.

Within the Nye area two zones of complex structures are found. The
south zone coincides approximately with the mountain front but passes back
into the pre-Cambrian to the west. It trends about N. 80° W. and consists
of nearly parallel east-west faults arranged en echelon and dipping steeply
south; movement on the faults has been mostly eastward and probably some-
what upward on the south side. The Paleozoic rocks within the fault zone
stand at high angles and in places are overturned and cut by subsidiary
faults; at the mouth of the Stillwater Canyon they are piled up in a schup-
penlike structure north of the main fault.

The northern structural zone is on the line of the Nye-Bowler lineament.
In the east half of the area it is characterized by a rather sharp anticlinal
structure broken by many small en echelon faults, believed to have resulted
from a dominantly horizontal movement along the zone, the south side
moving eastward. Farther west along this zone are a number of porphyry
intrusives, generally laccolithic in structure. Limestone Butte shows a ring
fault offset by radial faults, within which the Paleozoic formations have
been pushed up asymmetrically, so that the greatest displacement is on the
south side, where Ordovician is against Jurassic, and porphyry masses occur
along and on both sides of the fault. Round Mountain is a complex laccolith
intruding a syncline, and the elongated dome to the north shows a partly
concordant porphyry mass in the nose and northeast flank. Farther west,
in Meyers Canyon and its north fork, compound porphyry intrusions have
domed up the Lower Paleozoic rocks. The synclines around these domes as-
sociated with the porphyry are tightly compressed and in places overthrust
by the domes.

Between these two zones in the central and eastern part of the area there is
a Shallow syncline having sharply upturned edges. Farther west the rocks
dip rather uniformly northeast, except near the edge of the quadrangle
where they are folded asymmetrically, the axial planes dipping northeast;
the folds are cut by a few thrust faults dipping in the same direction.

It is believed that two major zones of weakness, trending about N. 80° W.,
existed in the basement rock previous to the Laramide deformation. In an
early stage of the orogeny porphyritic magmas intruded the sediments along
the north zone, and perhaps the eastern part of the Beartooth block was
bowed up relative to the western part and to the surrounding plains area.
During the period of major compression the Beartooth block was pushed
toward the ENE. relative to the plains area to the north, so that transverse
movement took place on the east-west en echelon faults of the south zone
where the Beartooth block stood high, and low angle underthrusting toward
the northeast occurred farther west where it stood relatively lower and was
opposite the area intruded by porphyry. On the north zone deepseated move-
ment was mainly horizontal, and the individual porphyry masses com-
pressed and overthrust the surrounding synclines.

T. A. Henpricks: Structural interpretation of recent gravity observations in
southeastern Oklahoma.

’
Ni
\

492 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

574TH MEETING

The 574th meeting was held at the Cosmos Club, March 22, 1939, Presi-
dent JouHn B. MertTIbx, Jr., presiding.

Informal communications —W. H. Monrok described a cave in sandstone
that had been formed by animals licking away the rock to obtain the small
amount of salt in the rock.

Program.—J. F. Bruu: The mechanical deformation of crystals and its
relation to structural geology.

W.H. Bravery: Fossil fish of the Green River formation.

S. R. Capps: Some Pleistocene placers in Idaho.

575TH MEETING

The 575th meeting of the Society was held at the Cosmos Club, April 12,
1939, President J. B. Mrrtiz, Jr., presiding.

Program.—W. F. Fosuac: Problems in the study of meteorites.

Louis 8. GARDNER: Displacement along the Hurricane fault in Utah and
Arizona.—The Hurricane fault extends north and south for at least 160
miles in southwestern Utah and northwestern Arizona. At some places it is
a single normal fault, but elsewhere it consists of a zone of several closely
packed faults showing three periods of movement in Tertiary and Quater-
nary times. The faults occur at the base of a high, almost unbroken, west-
facing escarpment, whichis a fault scarp for 30 miles in Utah, but isa fault-line
scarp near Mount Trumbull, Ariz. The stratigraphic displacement increases
northward from 1,500 feet at the Colorado River to about 10,000 feet near
the Pine Valley Mountains in Utah.

At many places 500 to 3,000 feet of the displacement is due to strata of the
dropped block bending downward toward the fault. In a distance of half a
mile or more the easterly dip increases progressively from less than 5° to
more than 25° at the fault. In the raised block to the east the strata are
again essentially horizontal. Farther to the east, such faults as the Sevier
and West Kaibab also show this feature.

Dutton and others interpreted the structures as representing monoclines
that were later cut by great normal faults. This explanation is unsuited to
the Hurricane fault zone because the bending, first, is not like that in ordi-
nary monoclines; second, is strictly confined to one side of the fault; third,
is closely associated with the fault and apparently dependent upon it; and
fourth, it has resulted from repeated movements, the latest of which affected
a land surface that is still in existence. Explanations such as drag along the
fault, or simple rotation of the dropped block, are also inconsistent with
field evidence.

The bending seems to have resulted from a mobile western block sagging
irregularly downward away from a stable eastern block. Evidences for this
are: (1) The physiographic contrast between the structural valley of Ash
Creek immediately west of the fault and the complexly eroded valley of La
Verkin Creek nearby to the east; (2) recent headward erosion of Ash Creek
tributaries into Great Basin alluvium; and (3) the anomolous arrangement
of ridges and valleys that would exist if the land surface deformed by the
last down-flexing were restored to its original condition.

J. B. Mertin, Jr.: Stratigraphic measurements in parallel folds—Two
topics are treated in this paper. The first is concerned with the development
of methods for analyzing the form and curvature of parallel folds, utilizing
actual field data. The second is a presentation of empirical methods for the
measurement of stratigraphic dimensions in parallel folds.

Nov. 15, 1940 PROCEEDINGS: GEOLOGICAL SOCIETY 493

The greatest desideratum of structural geologists is a better understand-
ing of the mechanics of folding. For this objective two principal sets of data
are available; first, the strength of the materials composing rocks, and
second, their observed deformation. While assumptions must still be made
regarding the behavior of deeply buried rocks when subjected to stress, it is
nevertheless true that greater progress might be made in the study of folded
rocks if all the available data could be treated quantitatively. The methods
given in the first part of this paper should be useful in analyzing the geo-
metrical forms of parallel folds.

Folded rocks having bedding surfaces which are approximately parallel
are said to lie in parallel folds. Utilizing the geometrical idea of an evolute
and its involutes, the author offers a more precise two-dimensional definition
of parallel folds, and at the same time points out inconsistencies in earlier
concepts. Suggestions are also made regarding a three-dimensional classifica-
tion of parallel folds, and particular attention is directed to the similarity
between Appalachian folding and certain three-dimensional graphs of the
Bessel function J,(x), of the real variables p and z.

It is shown that a family of involutes may be regarded as the traces of a
set of stratigraphic surfaces, in a plane normal to the axis of an elongate
parallel fold; and that comparable planes and involutes may exist for qua-
quaversal folds. Methods are presented for deriving the differential equation
of the normals to a family of involutes, and for obtaining therefrom the dif-
ferential equation of the involutes. The derivation of the equation of the
corresponding evolute is also given. All these equations are obtained from
the relationship 6=f(#), where x represents the varying distances along a
geological traverse normal to the strike of the rocks, and 6 represents the dip
of the rocks corresponding to various values of x. The relationship 6=kz,
which is often assumed by geologists in the measurement of stratigraphic
thickness, is utilized to exemplify the principle involved in this analysis; and
the type of fold which must exist under this assumption is deduced. Geo-
metric methods are also given for accomplishing the same results that are
obtained by mathematical analysis.

The second part of the paper deals with the application of mean trigono-
metric functions in the determination of stratigraphic thickness and other
stratigraphic dimensions. In such applications the most general case is as-
sumed, namely, that both the strike and dip of the rocks change from point
to point. Where it is possible to obtain a number of observations lying along
a straight line of traverse, it is shown that the best possible results will be
obtained by deriving the mean trigonometric functions corresponding to
these strikes and dips, and by substituting such mean values in homoclinal
formulae. This is essentially the method earlier proposed by E. L. Ickes.
Where sets of observations lying upon a straight line cannot be obtained,
or where for other reasons it is desirable to consider observations in pairs,
different methods should be used. One such method is based upon the
assumption that 6=kx. Another method, proposed by the author, is de-
veloped under the empirical assumption that the traces of parallel strati-
graphic surfaces in a plane, normal to such surfaces, may be considered, over
short distances, to have the curvature of concentric circular ares. Under this
assumption it is found that 6=are tan (kx+g) where k and g are constants,
depending upon the initial and terminal values of the strike and dip. The
corresponding mean trigonometric formations are then found to be the fol-
lowing:

ee

}

— Ps

= te

\ a _F

494 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

ey 1 k
tan 6 = Jc + ode =— 49
0

: 1
simon — fain [arc tan (kx +g)] = i (sec d2 — sec 61)
0

eae , 1
cos 6 = Joos [are tan (ka + g)| = — log

= do + sec =|
7 i

tan 6; + sec 6,

For purposes of rapid computation, the logarithms of sin 6, cos 6, and tan 6
have been computed, with an increment of five degrees for the variable,
which may be either the strike or the dip. These logarithms are utilized in
substituting such mean trigonometric functions in homoclinal formulae.

576TH MEETING

The 576th meeting of the Society was held at the Cosmos Club, April
26, 1939, President J. B. Mrrrtn, Jr., presiding.

i nformal communications—J. J. Fawry described peculiar crystals of
dolomite embedded in gypsum, which have an octahedral form that simu-
lates the pattern of the isometric octahedron, but the angle between the face
and base is 75.5° instead of 70.5° in the octahedron, The crystals actually are
the steep-sided rhombohedron, 4041.

H. D. Misrr described two thin dolomite layers in the Whitehorse forma-
tion in southwest Oklahoma, which, though less than half an inch thick,
can be traced over an area of several hundred square miles and form good
horizon markers for correlation.

Program.—Davip Griees: Convection currents and mountain-building.—
A review of the known forces available for deformation of the earth’s crust
shows that only those of thermal contraction and subcrustal convection are
sufficient in magnitude. Thermal contraction seems unable to supply the
amount of contraction required by geologic analysis of the Tertiary moun-
tain system. Mathematical analysis of convection by Vening Meinesz,
Pekeris, and Hales indicates that subcrustal convection is probable, and
would supply force of the proper magnitude. This paper presents an analysis
of the convective movements which suggests that such currents would be
periodic, and discusses the implications of this in mountain-building.

A convection-current cycle is suggested which may be correlated with the
mountain-building cycle. To study the action of the currents on the crust,
a model has been designed to be dimensionally similar to the conditions in
the earth’s outer shells, and to reproduce in one minute a process which takes
a million years in the earth. Moving pictures of this model in action suggest
a similarity between the structures so produced and those that are observed
in mountains.

C. K. Lreiru: Present state of some international mineral problems.

577TH MEETING

The 577th meeting of the Society was held at the Cosmos Club, November
8, 1939, President J. B. Mrrtin, Jr., presiding.

The following resolution presented by Miss JuLt1A GARDNER was un-
animously adopted by the Society:

“WENDELL CLAY MANSFIELD was born on June 9, 1874, in Charlotte
Center in western New York, a little south of Lake Erie and east of Lake

Nov. 15, 1940 PROCEEDINGS: GEOLOGICAL SOCIETY 495

Chautauqua. His A.B. degree was received from Syracuse University in
1908. For the two years following his graduation he taught science in the

secondary schools. On June 20, 1910, he entered the Federal service as pre- |

parator to Dr. William Healey Dall, replacing the veteran collector Frank
Burns. Mr. Mansfield advanced through the usual stages to the rank of
geologist and in the meantime had in 1913 received a M.S. degree from
George Washington University and in 1927 his doctorate. He was a member
of the Geological and Biological Societies of Washington, the Paleontological
Society, the Washington Academy of Sciences, and the Geological Society
of America. His wife Katherine Gibson Mansfield died several years ago
after a long illness, and there were no children. He accepted without bitter-
ness, but with full realization, the comparative loneliness of his later life.
He himself was in wretched health for a number of years, but there was no
word of complaint or of morbid reflection. He was spared, however, a long
dependence that would have been harder than physical pain, for the end
came suddenly on July 24, his first day of absence from his office desk. Two
sisters, a brother, and two nephews, whom he regarded with pride and af-
fection, survive him.

‘Trained under the aegis of Dr. Dall, his natural regard and respect for
books and museum collections was deepened and strengthened, and the
Tertiary molluscan collections at the United States National Museum bear
the imprint of his care. His faunal and stratigraphic studies were focused on
the middle and later Tertiary and Pleistocene of the Atlantic seaboard from
Virginia to Florida. To those familiar with the deliberation with which he
worked, the sum total of his published writings is surprisingly large, and
most of his papers include a distinct stratigraphic contribution. He was the
first to extend the Yorktown formation inland to Petersburg, Va. The de-
scription of the upper Miocene faunas of Florida—those of the Choctaw-
hatchee formation—and their careful zoning are perhaps his most compre-
hensive works. In this he established the upper faunal and stratigraphic limit
of the Miocene, while his work upon the Tampa had greatly aided in fixing the
lower limit of the Miocene. His study of the controversial faunal and field
relations of the Chickasawhay, on which he was engaged for some months
before his death, was sufficiently far advanced so that it can be published
in part.

‘Possibly the foremost quality of the man and of his work may be indi-
cated by the outmoded word integrity. Too little self-assertive and too little
self-confident to be sure of the rightness of his own opinions, he: was yet
tenacious of a belief that he had well considered and made his own. He was
never too busy to be approached and gave generously of all that was his,
whether material or immaterial. He asked little of life during the later years,
and those of us accustomed to the simple tenor of his ways were surprised at
the diversity of interests and organizations represented in the chapel filled
by those who gathered to pay the final tribute.

‘Mr. President, I move that these words of appreciation be incorporated
in the minutes of the Society and a copy of this statement be sent to his
sister in New Jersey.”

The following resolution, presented by W. C. ALDEN, was unanimously
adopted by the Society:

‘““ARTHUR JAMES COLLIER, a member of the Geological Society of Wash-
ington, died at Coronado, Calif., on October 13, 1939. Mr. Collier was a
member of the United States Geological Survey from 1896 until the time of
his retirement in 1935. He was born in Wheaton, IIl., on January 5, 1866.

496 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

After moving to the West he attended the University of Oregon, and from
this school he received the degrees A.B. and A.M. In 1894, Harvard Uni-
versity conferred on him the degree of Bachelor of Science in consequence
of studies there. During the years 1910 to 1914 he served as professor of
geology at the University of Oregon.

“Wield studies for the Federal Geological Survey resulted in the publica-
tion of several papers, by 1908, on gold, tin, and coal deposits of Alaska.
Following these came field mapping and published papers on coal and ore
deposits in Oregon, Washington, and Idaho. Several years were devoted to
mapping and study of the areal geology and structure of the rock formations
in several districts in Montana, including coalfields, the Kevin-Sunburst
oilfield, the Little Rocky Mountains, Bowdoin dome, and other possible
reservoirs of oil and gas. Several papers were issued by the Geological Survey
based in these studies, including also oilfields in Wyoming and the Nesson
anticline in North Dakota.

“Mr. President, I move that a copy of this statement be incorporated in
the minutes and that appropriate words of sympathy be sent to Mrs. Col-
lier.’”’

Program.—K. J. Murata: Volcanic ash as a source of silica for the silicifi-
cation of wood.—The chemical and physical properties of voleanic ash make
it a rich source of readily available silica for geochemical processes. In recent
years more and more sedimentary formations are being recognized as con-
taining volcanic ash or its decomposition products. The common associa-
tion of silicified wood with volcanic ash is noted in a list of occurrences of
silicified wood in the United States, and this association is viewed as the
result of a genetic relationship arising from the alterability of the ash.
Silicified wood may serve as an indicator for volcanic material in sedimen-
tary deposits.

W. D. Couuins: Water analyses —At one time the main use of water
analyses was as a help in judging whether waters were safe to drink. There
are still a good many who have no other thought about water analysis.
The water analyses made by the Geological Survey have very little value in
connection with questions relating to health. The analyses generally show
the quantities of a larger or smaller number of dissolved mineral con-
stituents, depending on the use that is to be made of the results. Analyses
may be faulty, valueless, or misleading because the samples were not col-
lected properly, because they changed in composition between the time of |
collection and the time of analysis, or because of errors in the analysis.

It is not easy to obtain samples that are truly representative of the sources
from which they are taken. Samples collected in the course of drilling a
well may represent water added in drilling rather than water present in the
formations reached by the drill. Most surface waters vary so much in com-
position with the seasons that care must be taken in using analyses of single
samples of surface waters. It is possible to take a sample from a river that
will represent the water in a particular part of the stream and fail to repre-
sent the main body of water in the stream.

There have been occasions when a sample of water has changed apprecia-
bly in composition during storage because of solution of the bottle, because
of the solution of insoluble matter that was in suspension in the water when
collected, or because of the deposition of material that was originally in
solution. This deposited material is generally calcium carbonate or hydrated
iron oxide.

The special methods and precautions necessary to obtain reliable analyses

Nov. 15, 1940 PROCEEDINGS: GEOLOGICAL SOCIETY 497

of the mineral content of natural waters are not fully appreciated by some
analysts who are expert in other lines of analytical work. Not all are familiar
with the various checks that can be applied to test the reliability of an
analysis. The inaccuracy of many analyses can be detected by inspection,
but others equally in error may appear, by every test, to be entirely reason-
able and reliable.

To obtain a useful and dependable water analysis it is necessary that the
sample be so collected as to be truly representative; that it be submitted for
analysis unchanged, and that it be analyzed with due attention to details,
with careful scrutiny and checking of the results.

Tom F. W. Barta: Thermal activity in Iceland.—A broad belt of geologi-
cally Recent formations traverses Iceland from southwest to northeast. It is
the so-called Palagonite formation, which consists of tuffs and breccias as-
sociated with basic intrusions, moraines, boulder-clays, and glacial sedi-
ments. To this belt is confined the modern volcanicity (volcanoes and
fissures such as Trolladyngja, Hekla, Katla, Laki, Vatnajokull, Askja,
Krafla, etc.), as well as the acid hot springs, which in Iceland invariably are
intimately associated with active volcanism. Indeed, acid springs seem to
depend on it for their existence; they are given birth by active volcanism,
they are changed by volcanic events, and they die when in any local region
the voleanism abates. But this does not mean that the acid thermal activity
is a modern phenomenon in Iceland. It has been inferred from certain meta-
morphic clay deposits that the pre-glacial voleanicity also was accompanied
by hot-spring action.

The alkaline hot springs are geographically independent of the modern
voleanism. They are found in the palagonite belt, as well as on either side of
this belt, in the formation of Tertiary basalts of both East Iceland and West
Iceland.

Other characteristic differences between acid springs and alkaline springs
are: (1) Their relation to the surface topography: acid springs are found
along tectonic lines and usually on high, dry ridges, but alkaline springs are
typical for areas possessing a superior ground-water supply, and break out at
the foot of long hill slopes, or in depressions in the land surface, often in
river beds or in the bottom of lakes; (2) the alkaline springs are much more
copious than the acid ones; (3) the temperature curve is typically different
for the two kinds of springs.

Both types of springs are fed by volcanic contributions; they receive their
thermal energy as well as certain chemical constituents from volcanic sources.
It can be shown that the farther the volcanic emanations have to travel
underground before they find egress to the surface and break out as hot
springs, the more alkaline they become and the more do they become diluted
by ordinary ground water (thus becoming increasingly copious).

This evolution from what in a sense may be called ‘‘primary”’ acid springs
of insignificant water discharge to “‘secondary”’ alkaline springs of con-
siderable discharge, is in line with, and corroborates the studies conducted
by A. L. Day and E. T. Allen on the thermal activity in California and in
Yellowstone National Park.

578TH MEETING

The 578th meeting of the Society was held at the Cosmos Club, November
22, 1939, President Joun B. Mertin, Jr., presiding.

The following resolution presented by G. F. LouGHLIN was unanimously
adopted by the Society:

498 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

‘““We were very sorry, though not surprised, to learn that Dr. WALDEMAR
LINDGREN died on November 4, 1939, at the age of 79, after a long illness.
Though not a charter member of the Geological Survey, he was identified
with it and with American geology from its pioneer days. Born in Sweden
and educated there and at Freiberg, Germany, he came to America in 1883,
with a letter of introduction to Raphael Pumpelly, who was conducting the
Northern Transcontinental Survey established by the Northern Pacific
Railway. Pumpelly assigned him to assist W. M. Davis, who for years after-
ward used to refer to his young assistant who mounted a horse on the wrong
side. This work ended in 1884 and Lindgren worked for a short time as an
assayer at Helena and draftsman at Anaconda, Mont., before being ap-
pointed assistant geologist in Becker’s party, which was studying quick-
silver deposits in California for the U. S. Geological Survey. From then
until 1912 he remained with the Survey. His ability as a mining geologist,
reflected in his Survey reports and scientific papers, soon gained deserved
recognition and he became generally regarded as the world’s outstanding
authority on the science of ore deposits.

“His broad view of the subject is reflected not only in his writings on min-
ing districts and on processes of mineral deposition, but in the time and
energy that he spent in placing on a sound basis the Survey’s annual statisti-
cal reports, which from 1903 on have furnished the basis for a quantitative
approach to mining geology. In 1905 he organized the small group that
established the journal Economic Geology, which promptly became the
leader in its field. Lindgren continued for several years in charge of the
Metals Section of the Division of Mineral Resources, and in 1908 succeeded
S. F. Emmons as chief of the Section of Metalliferous Deposits. He held
both positions until 1911, when he was made Chief Geologist, succeeding
C. W. Hayes.

“In 1912, with the desire to give more time to original investigations than
his administrative duties on the Survey permitted, he resigned to become
William Barton Rogers professor of economic geology at the Massachusetts
Institute of Technology, where he had been a visiting lecturer for the three
preceding years. Years later, when he received the Penrose medal in recog-
nition of his leadership, he said that he never felt cheaper than when he
handed his resignation to George O. Smith, who was then director. He had
grown with the Survey, and it had been such a great part of his life for so
many years that it was hard to reconcile himself to the separation. In 1913
the first edition of his book Mineral Deposits, an outgrowth of his lectures,
was published. This book also promptly became the leader in its field. He
remained at Tech until his retirement about nine years ago and continued
to be an inspiration to many graduate students, and an outstanding con-
sulting geologist.

“Tn 1925 he served as chairman of the Division of Geology and Geography
of National Research Council and successfully laid the groundwork for the
publication of the Annotated bibliography of economic geology, the first volume
of which appeared in 1928. He supervised the first 10 volumes, his many
contributions to which made interesting and spicy reading, besides showing
in few words the relative value of many articles on the subject.

“The science of mineral deposits is by no means a one-man science, but,
when we reflect on its growth, we find that no one so much as Lindgren has
contributed to and guided its advance.

“Geology and the world has suffered an irreparable loss in his passing,
but we are most grateful that he was able to round out his work so well

Nov. 15, 1940 OBITUARY 499

during his long career. We regret that his period of comfortable relaxation
after retirement could not have been longer and we extend our sincere
sympathy to his surviving relatives and to the Massachusetts Institute of
Technology where he spent the last 27 years of his life.

“Mr. President, I move that the Geological Society of Washington prepare
an appropriate resolution of sympathy and send it to Dr. Lindgren’s nearest
relatives, particularly his niece, Helen Lindgren, who has been his close com-
panion for many years, and to the Massachusetts Institute of Technology.”

Informal commumnications.—J. C. REED and R. R. Coates reported on
the thermal gradient in mines on Chichagof Island, Alaska, which they found
to be about 1.55° F. per hundred feet. The communication was presented
by Mr. CoatEs.

Program.—P. B. Kina: Persistent structural trends in Guadalupe Mountain
region, Tex.

FraNK NEUMANN: Seismograph evidence of deep focus earthquakes.

W. D. Urry: Measurement of geologic time.

579TH MEETING

The 579th meeting of the Society was held at the Cosmos Club, December
13, 1939, President J. B. Mrertin, Jr., presiding.

Program.—Vice-president J. T. ParprEr took the chair while President
Menrtiz delivered his presidential address, Placer gold in Alaska, which was
published in this JouRNAL, Vol. 30: 93-124, Mar. 15, 1940.

AT7TTH ANNUAL MEETING

The 47th annual meeting of the Society was held immediately following
the 579th regular meeting. The reports of the secretaries, auditing commit-
tee, and treasurer were read and approved.

The Society elected the following officers for the year 1940:

President: J. T. PARDEE. !

Vice-presidents: C. WytHE Cooke and L. H. Apams.

Treasurer: ALICE S. ALLEN.

Secretary: R. W. Brown.

Council: D. A. ANDREWs, EUGENE CaLLaGcHan, C. B. Hunt, C. B. Reap,
R. E. STEVENS.

The society appointed J. B. Mrertin, Jr., as its nominee for vice-president
of the Washington Academy of Sciences. :
ParRKER D. Trask, Secretary

Obituary

David Myers Mortrtier, professor emeritus of botany at Indiana Uni-
versity, died at the home of his daughter in Indianapolis on March 25, 1940,
after a brief illness, at the age of 75 years. Born September 4, 1864, his early
years were spent near his birthplace in a Swiss settlement at Patriot, in
southeastern Indiana. Following his attendance at the public schools near
his home, he taught for several years in the elementary schools before enter-
ing Indiana University as a student. He received the A.B. degree from Indi-
ana in 1891 and the A.M. in 1892.

500 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 11

In his student days at Indiana and the few years immediately following,
while he served as instructor in botany, Mottier caught the enthusiasm of
such of his instructors and colleagues as David Starr Jordan, John Merle
Coulter, and Douglas H. Campbell and, as the next natural step for that
time, turned to Germany for further study. His formal graduate work with
Strasburger at Bonn resulted in his receiving the Ph.D. degree in 1897. He
later spent some time in research at the Biological Station at Naples and at
the University of Leipzig.

From the time of his first appointment at Indiana University in 1891 until
his retirement in 1937 he served continuously—as instructor, associate pro-
fessor, professor, and departmental head—except for two leaves for study in
Europe.

The pattern of Dr. Mottier’s life work took form in that bright decade
between 1890 and 1900 when so much was done to clarify the cytological de-
tails of the life histories of the higher plants and animals and lay the founda-
tion for the experimental work of the present century. At that time he was
acquainted, personally or by correspondence and exchange, with many of the
most active workers in his field in both Europe and America. More than two
years before the announcement by others of double fertilization in angio-
sperms, he had made excellent microscopic preparations showing a sperm in
contact with the polar nuclei, but he heeded the vigorously administered ad-
vice of Strasburger and ignored it as of no consequence. As a symbol of his
excellent cytological technique, one of his slides showing this phenomenon
was in good condition 30 years later, although it had many times been sub-
jected to projection with an arc light.

His most important early work was on various problems of the cytology
and embryogeny of vascular plants. Later he made significant contributions
to what was known of the origin and development of chloroplasts and mito-
chondria. His last work dealt chiefly with the prolonged culture of the
gametophytes of ferns under conditions preventing fertilization.

A keen mind, a thoroughness of application, and a master of detailed per-
fection made him an artist in the techniques of cytological preparation and
illustration. These qualities, with his seriousness of purpose in all aspects of
teaching, research, and study, and his gentlemanly old-school personal quali-
ties, are reflected in the respect of numerous students who felt his influence.

Besides numerous research papers, he prepared in 1902, under the title
Fecundation in plants, a résumé of what was then known of fertilization in all
groups. He was also the author of an elementary text in botany and a labora-
tory manual.

Dr. Mottier was a fellow of the American Association for the Advancement
of Science, a life member of the Botanical Society of America, a charter mem-
ber, fellow, and expresident of the Indiana Academy of Science, and a mem-
ber of the American Society of Naturalists, the Washington Academy of
Sciences, and, at times, of several other learned and honorary societies.

y

tain organic salphur ne “EDWARD ae “EVERIPD and
ME Xs SULLIVAN ge eee ee coe a eee

PALEOBOTANY. on -Cusparia from the Plioeene of =
Bolivia. _Epwarp W. BERRY So ois: |

-Botany.—Arizona plants: A new variety and aed)
eae Roce H. PEEBLES and Louts ¢ ve

_ ENTOMOLOGY. aS North American on leafhop oppers of
alboneura group (Homoptera-Cicadellidae). N. ANCY i. Wasi I

Procuspines: THE AcapEmy Dy 4 ete a

cag

PRocEEDINGS: GEOLOGICAL SocrEty.

;

OBITUARY: alee gees :

4 DrcemBer 15, 1940 No. 12

JOURNAL

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| OF SCIENCES

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JOURNAL

OF THE
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VoL. 30 D&EcEMBER 15, 1940 No. 12

BOTAN Y.—Two new genera of Acanthaceae from Guatemala. E. C.
LEonarp, U.S. National Museum. (Communicated by WILLIAM
R. Maxon.)

In the large collection of plants procured by Paul C. Standley on
the 1939 Sewell Avery Expedition of the Field Museum of Natural
History to Guatemala there were a number of interesting specimens
of the family Acanthaceae. Represented in this material are two new
genera, which are described herewith. One of these includes a species
previously placed in the genus Drejerella.

Averia Leonard, gen. nov.

Sect. Odontonemineae. Calyx 5-partitus, tubo brevi, segmentis angustis;
corolla subregularis, tubo gracili, lobis subaequalibus, obtusis vel rotunda-

a b 5 d e

Fig. 1.—Averia serrata Leonard: a, Por-
tion of plant, natural size; b, bract; c, bract-
let; d, calyx; e, stamen. (6, c, d, e, twice
natural size.)

tis; stamina 2; antherae biloculares muticae, loculis parallelis, subaequaliter
affixis; capsula stipitata, parva, 2- vel 4-sperma. Herbae; spicae terminales,
densae; bracteae ovatae, imbricatae.

Type species, Drejerella longipes Standley.

This genus is closely related to Drejerella Lindau, but differs in its muti-
cous, subequal, parallel anther cells. The anther cells of Drejerella are sub-

1 Published by permission of the Secretary of the Smithsonian Institution. Received
June 28, 1940.

501

pec 31

502 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12 |

superposed, the lowermost strongly calcarate. In both genera the pollen

erains are of the “Spangenpollen” type, with a series of longitudinal grooves
and an equatorial band of pores (three).

Fig. 2.— Megalostoma viridescens Leonard: a, Portion of
branch; b, calyx (both natural size).

The name Averza is given in honor of Dr. Sewell Avery, director of the
recent Field Museum expedition to Guatemala above mentioned.

Bracts serrate es et ee a AS ee A. serrata
Bracts entire ss. c either os A eee ee A. longipes

Averia serrata Leonard, sp. nov.

Herba, caulibus erectis vel adscendentibus, subteretibus, glanduloso-hir-
tellis; lamina foliorum ovata, apivte subobtusa, apiculata, basi truncata, ser-

Dec. 15, 1940 LEONARD: NEW ACANTHACEAE 503

rata, glanduloso-hirtella; petioli tenues; spicae multae; bracteae acutae vel
obtusae, apiculatae, serratae, glanduloso-hirtellae; bracteolae lineari-
lanceolatae, glanduloso-pilosae; calycis segmenta lanceolata, glanduloso-
hirtella; corolla glabra, ochroleuca; capsula glabra; semina plana, muricata.

Branched herbs, suffrutescent at base; stems erect or ascending, up to 20
em long or more, subterete, glandular-hirtellous or the older parts glabrate;
leaf blades ovate, 8 mm long, 6 mm wide, obtusish and minutely apiculate
at apex, truncate at base, serrate, glandular-hirtellous; petioles slender,
about 5 mm long, glandular-pilose; spikes numerous, up to 2.5 cm long and
1 cm in diameter; bracts closely imbricate, rhombic, up to 5 mm long, 3 to
4.5mm wide, acute or obtuse at apex, minutely apiculate, narrowed at base
to a short petiole (2 mm long, 1 mm wide), serrate, with one to three teeth
on each side, 3-nerved, glandular-hirtellous; bractlets linear-lanceolate, 6
mm long, 1 mm wide, densely glandular-pilose; calyx 5-parted, the seg-
ments lanceolate, 3.5 mm long, 0.5 mm wide, glandular-hirtellous; corolla
about 8 mm long, glabrous, cream-colored, the tube-3 mm long, slender, the
segments (4) obovate, subequal, 3 to 3.5 mm wide toward apex, obtuse;
stamens 6 mm long, the filaments slender, glabrous, the anthers purplish;
capsules clavate, 3 to 4 mm long, 2 mm broad, glabrous, 4-seeded, the seeds
flat, muricate.

Type in the herbarium of the Field Museum of Natural History, no.
981916, collected at Champerico, Department Retalhuleu, Guatemala, in a
dry thicket at sea level, February 26, 1939, by Paul C. Standley (no. 66612).
An isotype is in the U.S. National Herbarium (no. 1790340). _

In general appearance A. serrata is similar to A. longipes, but is readily dis-
tinguished by its serrate bracts and its smaller and more glandular leaves.

Averia longipes (Standl.) Leonard, comb. nov.

Drejerella longipes Standley, Field Mus. Publ. Bot. 8: 47. 1930.

The type, collected at Chichen Itza, Yucatan, February 27-28, 1899, by
C. F. Millspaugh (no. 1621), is at the Field Museum (no. 437703), as also
another specimen, collected at Buena Vista by G. F. Gaumer in 1899.

Megalostoma Leonard, gen. nov.

Sect. Louteridieae. Calyx 5-partitus, irregularis, coriaceus, segmentis
lateralibus falcatis, anterioribus angustissimis; corolla bilabiata, labiis late
apertis, labio superiore angusto, erecto, inferiore cochleariformi; stamina 2;
antherae sagittatae. Frutex; paniculae laxae terminales.

Type species, M. viridescens Leonard.

The genus Megalostoma has apparently no close affinities, although its
irregular coriaceous calyx and tubercular pollen grains suggest a possible
relationship to Louteridium Wats. The name Megalostoma alludes to the
widely open corollas.

Megalostoma viridescens Leonard, sp. nov.

Frutex, ramis glabris, subquadrangularibus, angulis anguste alatis; lamina
folhiorum oblongo-elliptica, acuminata, basi angustata, glabra; petioli tenues;
paniculae laxae, secundae; calyx glaber, segmento posteriore elliptico, ob-
tuso, lateralibus lanceolatis, subacutis, anterioribus angustissimis; corolla
viridescens, glanduloso-pubescens; ovarium glabrum.

Shrub, up to 2.5 meters high; branches subquadrangular, narrowly
winged, glabrous, the cystoliths subpunctiform; leaf blades oblong-elliptic,
up to 12 cm long and 5 cm wide, slenderly acuminate at apex (the tip often

504 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

curved), narrowed at base, glabrous, blackish in dried specimens, the cysto-
liths obscure; petioles slender, up to 5 mm long; flowers relatively few,
secund, borne in loose terminal panicles; bracts leaflike, soon deciduous;
calyx segments 1.5 cm long, the posterior segment elliptic, 7 mm wide, ob-
tuse, the lateral pair lanceolate, 5 mm wide, acutish, faleate, the anterior
pair narrowly lanceolate, 1.5 mm wide, all glabrous, coriaceous, blackish in
dried specimens; corolla greenish white, glandular-pubescent, the lips about
2.5 cm long, entire, the upper lip linear-oblong, about 5 mm wide, obtuse,
erect, the lower about 16 mm wide, obtuse, cochleariform; stamens equaling
the corolla lips, the anthers 7 mm long, acute at the base, the sacs parallel;
pollen grains ellipsoidal, tubercular, with an equatorial row of pores (“‘Stach-
elpollen’’); style as long as the stamens, reclining in the lower lip of the co-
rolla; ovary glabrous; capsule not seen.

Type in the herbarium of the Field Museum of Natural History, no.
990983, collected at Escoba, across the bay (west) from Puerto Barrios,
Department Izabal, Guatemala, in a wet forest near sea level, May 3, 1939,
by Paul C. Standley (no. 72948). Nos. 72896 and 73025 of Mr. Standley’s
collection, with identical locality data, are the same.

Remarkable for its widely divergent corolla lobes, spreading at a right
angle, which bear a fanciful resemblance to the yawning mouth of a pelican.

ENTOMOLOGY .—The oviposition habits of the Eucharidae (Hymen-
optera).1 Curtis P. Cuausen, U. 8. Bureau of Entomology and
Plant Quarantine.

The Eucharidae are a family of small wasps that are parasitic upon
the mature larvae and the pupae of ants. This limitation in host pref-
erences to a single family or superfamily, whichever status is given to
the group, is in sharp contrast to the wide host range of related fami-
lies of the Chalcidoidea. The family is world-wide in distribution but
is encountered most frequently in the Tropics. The adults are black,
metallic blue, or green, often with the thorax distinctively sculptured,
and many species have the scutellum variously modified, in may spec-
ies it being produced into a conspicuous bifurcate process that may
extend beyond the tip of the abdomen. Not only are they conspicuous
and often weird in form, but the habits and relationships of these in-
sects to their hosts are so unusual and varied as to be outstanding,
even in an order in which the most diverse adaptations to the parasitic
mode of life are found.

The first studies on the biology and habits of the Eucharidae were
by Wheeler (1907), who found several species of Orasema associated
with ants of the genera Pheidole and Solenopsis in Texas and Colorado.
He reared several successive ‘‘broods” of O. viridis Ashm.? upon
Pheidole instabilis Emery and described and figured the larval instars

1 Received August 10, 1940. ;
2 According to A. B. Gahan, the species here referred to is probably Orasema wheelert
Wheeler and not the true viridis Ashm. (See Proc. U. S. Nat. Mus. 88: 459. 1940.)

_ Dec. 15, 1940 CLAUSEN: OVIPOSITION OF THE EUCHARIDAE 505

and the pupa. Oviposition was not observed nor were the eggs found,
yet so certain was he that these must be placed directly upon the ant
pupae that he described in some detail the manner in which the female
was presumed to accomplish this act. The conclusions reached were
entirely logical on the basis of what was then known regarding the
habits of parasitic insects.

A few years later H. S. Smith (1912) presented a detailed account of
the biology of Perilampus hyalinus Say, of the family Perilampidae,
which is closely allied taxonomically to the Eucharidae. Here again
the act of oviposition and the egg were not seen, but the first instar
larva proved to be identical in general characters with that of Ora-
sema. These planidia (dimunitive wanderers) differed from other
known parasitic hymenopterous larvae in the possession of a fusiform
body and a highly sclerotized and darkened integument. Because of
the form and habits of the planidium of Perzlampus, Smith was con-
vinced that oviposition did not take place directly in or on the host,
and he believed that the eggs were more probably deposited upon the
food plant in the vicinity of a colony of hosts. In discussing Wheeler’s
observations on Orasema he pointed out the similarity in form and
habits of the larvae with those of Perrlampus and suggested the proba-
bility that oviposition takes place entirely outside the ant nest. Smith
was able later (1917) to verify the leaf-ovipositing habit in P. chryso-
pae Cwfd., and this discovery served to stimulate interest in the habits
of the two families.

The first discovery of the oviposition habits of a species of EKuchari-
dae was purely accidental. During the course of a study of the insect
fauna of wild cotton in Arizona, Pierce and Morrill (1914) chanced
to observe two females of Chalcura arizonensis Cwfd. with their ovi-
positors inserted in apparently healthy blossom squares of this plant.
Upon later examination these squares were found to contain masses of
minute eggs immediately beneath the points of insertion of the ovi-
positors. This record was entirely overlooked by later workers, and it
was not until the writer’s account of the habits of (Schizaspidia) Stil-
bula tenuicornis (Ashm.), published in 1923, that the plant-oviposition
habit became generally known. Since that time observations have
been made upon the oviposition habits of 7 additional genera and 12
Species occurring in various parts of the world,*® and several other in-

3’ The writer is indebted to A. B. Gahan, of the U. S. Bureau of Entomology and
Plant Quarantine, for the determination of species and for the descriptions of those that
have proved to be new (Proc. U. S. Nat. Mus. 88: 425-458. 1940). In a forthcoming
publication (Hntomophagous insects, New York) a short account is given of the biology
and habits of the family. At the time of submission of the manuscript the names of

~
\

506 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

vestigators have added to our knowledge of this subject, so that it is
now possible to present an account of several of the principal oviposi-
tion habits of the family. In every instance it has been found that the
eggs are deposited entirely apart from the host, that the association
with plants for this purpose is obligatory, and that a wide range exists
with respect to the manner of deposition of the eggs and the part of
the plant on or in which they are placed.

As an aid to others who may be interested in studying the habits of
the family, it may be mentioned that the oviposition habits can be
readily and quickly determined by observations on females in the
field. They oviposit almost immediately after leaving the ant nest and,
because of their limited and relatively slow flight, can be followed
until they alight upon the plant that is to receive their eggs. The
elapsed time from emergence to oviposition is usually less than 1 hour.
Botanical gardens provide exceptional opportunites for the study of
the Eucharidae, as the variety of plants grown there insures that some
will be present that are suitable for oviposition by such species as may
occur in that locality. Also, the ant population is usually relatively
high and of many species. The Botanical Garden at Peradeniya, Cey-
lon, was especially fruitful and yielded three species in abundance
during a short visit there in February 1930.

OVIPOSITION PLANTS

In Table 1 are given the species of which the habits are known and
the plant or plants with which each one is associated for oviposition.
The records are based on the author’s observations unless otherwise
indicated.

PART OF PLANT UTILIZED FOR OVIPOSITION

The part of the plant utilized for oviposition varies with the species
and may be the overwintering buds, opening flower buds, stems of
blossom clusters, seed pods, or leaves. The variations in habit, in rela-
tion to the part of the plant that serves to receive the eggs, are here
discussed under the following headings:

1. In overwintering buds. The single species that is definitely known
to pass the winter in the egg stage is Stdlbula tenuicornis of Japan and
Chosen, which places its eggs in the overwintering buds of mulberry
(Fig. 1) and, to a lesser extent, in those of chestnut, oak, birch, and

Gahan’s new species were not available, and several were referred to under the generic
name only. These are now assigned as follows: Psilogaster sp. from Malaya=P. anten-
natus Gahan; Parapsilogaster sp. from Ceylon =P. laeviceps Gahan; Eucharis sp. from
Chosen = £. scutellaris Gahan; and Schizaspidia sp. from Malaya =S. antennata Gahan.

507

OVIPOSITION OF THE EUCHARIDAE

CLAUSEN

Dec. 15, 1940

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008 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

Cladrastis amurensis var. floribunda (Clausen, 1923). The entire
quota of eggs of the female, numbering approximately 1,000, is depos-
ited within the interior of the bud at one insertion of the ovipositor.

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Fig. 1.—A mulberry flower bud cut away to show two egg masses of Stilbula tenut-
cornis (Ashm.). Fig. 2.—A seed, with fully expanded plumes, of Picris hieracioides var.
spinulosa bearing a mass of eggs (H.M.) of Stilbula cyniformis Rossi (from Parker,
1937). Fig. 3.—A portion of a leaf showing, on its under side, the paired rows of ovipo-
sition scars of Losbanos uichancoi Ishii (from Ishii, 1932). Fig. 4.—A group of eggs of
Psilogaster antennatus Gahan upon the leaf surface surrounding a freshly deposited egg of
Selenothrips rubrocinctus. Fig. 5.—A fleshy leaf showing the serpentine lines of oviposi-
tion scars of Schizaspidia antennata Gahan. Fig. 6.—A diagrammatic section through
a fleshy leaf showing (a) a group of three eggs of Parapsilogaster laeviceps Gahan and
(6) a group of four eggs of Schizaspidia antennata Gahan in their characteristic position
resulting from the insertion of the curved ovipositor at a sharp angle to the leaf surface.

Dec. 15, 1940 CLAUSEN: OVIPOSITION OF THE EUCHARIDAE 509

These eggs remain therein until the following spring, when the buds
expand and the great majority of egg masses fall to the ground with
the bud scales. A portion of the buds die, however, and the scales
draw apart somewhat, thus permitting the escape of the larvae, which
hatch from the eggs during July and early in August.

2. In expanding leaf and flower buds. A number of species, repre-
senting several genera, utilize the expanding leaf and flower buds of
various plants for oviposition. Hucharis scutellaris of Chosen, which
oviposits in those of a trailing vine of the genus Cebatha, may even de-
posit her eggs in fully opened blossoms. Less than 50 eggs are placed
in the center of each one, and consequently each female visits a con-
siderable number of blossoms before oviposition is complete. Kapala
furcata oviposits similarly in the flower buds of another vine, of the
genus Mikania, and of an undetermined species of Amaranthaceae.
A colony of this parasite was found inhabiting the small plot of open
eround immediately in front of the main laboratory building on Barro
Colorado Island, Panama Canal Zone.

Chalcura arizonensis deposits its eggs in the blossoms of wild cotton
in Arizona, while C. deprivata of Ceylon does so in the leaf buds of jak
fruit, though some individuals were seen to utilize the expanding
flower buds of croton and Cordia. Stilbula manipurensis of Assam
places its eggs in masses beneath one of the outer scales of the large,
loosely formed buds of Flamingia (Clausen, 1928). Schizaspidia con-
vergens deposits them in clusters of a few hundred beneath the outer
scales of leaf buds, and occasionally in the flower buds also, of jak
fruit in Ceylon, and usually high up in the tree. Buds containing egg
masses can frequently be recognized by the presence of a curled ribbon
or thread of white congealed sap, several millimeters in length, at the
point penetrated by the ovipositor. In all these species the incubation
period is relatively short, covering not more than two weeks, but be-
fore hatching takes place the bud scales and petals fall to the ground
and carry the great majority of eggs with them.

3. In seed receptacles. Parker (1987) describes the oviposition of
Stilbula cyniformis in the seed heads of a small composite plant of the
genus Picris in southern France. The eggs are deposited en masse
among the bracts or adhering to the outer layer of seeds. As the seed
head opens the plumes of each seed expand and the seed, with its
burden of eggs (Fig. 2), is blown away. Inasmuch as oviposition takes
place only during August, it is possible that this species, like S.
tenuicornis, passes the winter in the egg stage. After completion of
Oviposition the female often dies with her ovipositor still inserted in
the seed receptacle.

510 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

4. In incisions in leaf tussue. The females of Schizaspidia antennata,
the commonest of the eucharid species observed at Kuala Lumpur,
Federated Malay States, usually deposit their eggs in pairs, but at
times in groups of three or four, in incisions in the under sides of the
fleshy leaves of Hugenia, Medinella, etc. These punctures are evenly
spaced about | millimeter apart and are made in serpentine rows over ~
the leaf (Figs. 5, 6, b). When the trees are in bloom oviposition is fre-
quently in the fleshy stems of the blossom clusters rather than in the
leaves. Parapsilogaster laeviceps shows a pronounced preference for the
half-grown leaves of jak fruit, the eggs are laid singly in incisions on
the under side, usually near the leaf margin, and the incisions are
made somewhat at random rather than in rows.

In July 1932 J. C. Bridwell showed the writer a colon of Orasema
coloradensis at Barcroft, Va., that was restricted to a very limited area
along a railroad right-of-way and bordering a wooded area. The fe-
males were seen to be ovipositing in the younger leaves of the small-
leaf pencil-flower, Stylosanthes biflora, and also occasionally beneath
the bud scales of the large-leaf Jersey tea, Ceanothus americanus
(determinations by Dr. 8. F. Blake). In the pencil-flower the eggs
are placed singly or in pairs in incisions in the under sides of the
younger leaves. between the parallel veins. While there may be several
punctures in a row, yet because of the small size of the leaf any distinct
linear or serpentine arrangement of the punctures is prevented. An
elongate area of leaf-tissue about the puncture becomes discolored and
dies. In the limited area inhabited by this colony practically every leaf
of the pencil-flower contained one or more eggs, and the numerous
areas of dead tissue gave the plants the appearance of being diseased.
O. smithi oviposits in the same way, but this species is distinctive in
that it oviposits only in the upper sides of the leaves and limits itself
to those within 2 feet of the ground. Kapala terminalis likewise ovi-
posits in the upper rather than the lower sides of the leaves.

Ishii (1932) describes the somewhat similar leaf-ovipositing habits
of two species from the Philippine Islands. The females of Kapala
foveatella place one to four eggs in each incision in the lower sides of
the leaves of Gliricidia and Leucaena. The oviposition punctures made
by Losbanos uichancoi on the under sides of the leaves of Celtis and
Leucaena occur in two short parallel rows, each row comprising five to
ten punctures. This oviposition in a double rather than in a single row,
as is the habit of other species, is an interesting variation, but unfor-
tunately the author does not describe the manner in which it 1s accom-
plished. His illustration (Fig. 3) indicates that the two rows of eggs are

Dec. 15, 1940 CLAUSEN: OVIPOSITION OF THE EUCHARIDAE 511

deposited simultaneously, the ovipositor apparently being inserted
alternately right and left as the female moves forward.

5. At random on leaf surface. This oviposition habit was first ob-
served by Ishii in the case of Parapsilogaster montanus at Los Banos,
Philippine Islands. The eggs are deposited horizontally on the under
sides of the leaves of Sandricum and Premna, and, when abundant, they
give a white, powdery appearance to the leaf surface. A species of Ka-
pala found in Cuba, and not distinguishable in the adult stage from K.
terminalis, oviposits in the same way upon the leaves of T’ragza volubi-
lis, A female normally deposits her entire quota of eggs upon a single
leaf, and each of these may bear many thousands of eggs. She walks
about very slowly over the leaf, tapping its surface rapidly with the
tip of the ovipositor, and one or two eggs are extruded each time.
Leaves bearing fully incubated eggs appear as if covered with a fun-
gous growth, the slender egg stalks resembling hyphae and the deep
amber-colored egg bodies the conidia. These species, and Pszlogaster
antennatus, are among the very few Chalcidoidea that deposit eggs of
the stalked type in such a position that they are completely exposed.

6. Upon leaf surface associated with thrips eggs. One of the most
striking and highly specialized adaptations in oviposition in the
Kucharidae was observed in the Malayan Pszlogaster antennatus,
which was collected in some numbers in January 1930 near Kuala Lam-
pur. The eggs are placed vertically, regularly spaced and in numbers
up to 100, in the immediate vicinity of a freshly deposited thrips egg
(Fig. 4). Under cage conditions oviposition could not be secured in
the absence of these eggs, and the relationship appears to be obliga-
tory. The thrips species concerned, Selenothrips rubrocinctus (Giard),
was found commonly only on mango and Erythrina foliage. The
thrips female partially inserts the egg in an incision in the under side
of the leaf and covers it with a mass of excrement. The female para-
site is attracted to the egg itself rather than to excrementous covering.
The parasite’s association with the foliage of the two trees mentioned
is believed to be incidental and it very probably will be found to
frequent any type of plant which bears an infestation of Selenothrips.
The presumed ant host of Psilogaster is not known, so it is impossible
to give any convincing explanation of this association with thrips.
Several species of the genus have been reared from Myrmecia and
Pheidole, so there is little basis for believing in any radical departure
in host preferences. The association with thrips undoubtedly relates
to phoresy, as the eggs hatch simultaneously with those of the carrier
and the planidia attach themselves to the young thrips larva as soon

512 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

as it emerges from the egg and are carried about until the first molt of
the latter. Were the thrips attended by ants, or carried into their
nests, the relationship would serve a definite and obvious purpose, but
such is not the case.

The number of species for which information is available regarding
their oviposition habits is still too small to warrant any generaliza-
tions, and in all probability other and perhaps radical departures from
those discussed will be found. In examining the data given in the pre-
ceding paragraphs it is seen that there is little uniformity in habit even
among species of a genus. Parapsilogaster laeviceps places its eggs in
incisions in leaf tissue, whereas P. montanus deposits them at random
on the leaf surface. Kapala foveatella and K. terminalis oviposit in leaf
tissue, K. furcata in the expanding blossom buds of a vine, and an
undetermined species of Kapala places its eggs loosely upon the leaf
surface. Stilbula tenuicornis oviposits in overwintering leaf and flower
buds, S. manipurensis in expanding flower buds, and S. cyniformis
in seed receptacles. The three species of Orasema are, however, con-
sistent in placing their eggs in incisions in leaves.

MANNER OF OVIPOSITION

The species of Stzlbula, Kapala, Eucharis, Chalcura, and Schiza-
spidia that deposit their eggs in buds or seed receptacles have a uni-
form manner of oviposition. The female penetrates the scales or cov-
ering by a downward thrust of the ovipositor, and full penetration of
a bud with heavy scales may require 5 minutes or more. In no instance
has a female been seen to insert the ovipositor between the scales.

Two distinct methods of oviposition are found among the species
that oviposit in leaf tissue. Parapsilogaster laeviceps (Fig. 6, a) and
Orasema coloradensis merely puncture the leaf tissue by a perpendicu-
lar thrust of the straight ovipositor, and consequently the perforation
in the epidermis is found at the center of an area of dead tissue, which,
within a few days, becomes about 1 millimeter in diameter. Schiza-
spidia antennata, however, has a much heavier ovipositor, which is
distinctly curved downward, so that when it is lowered preparatory
to oviposition the tip is directed forward. Insertion into the leaf is
consequently effected by a forward pull rather than by a backward or
downward thrust. After being inserted to the proper depth in the
fleshy leaf tissue, it is swung from side to side through an are of about
45°, with the base held stationary. This results in the formation of a
fan-shaped recess within the leaf, in which two to four eggs are placed
(Fig. 6, b). On leaves in which eggs have been deposited several days

Dec. 15, 1940 CLAUSEN: OVIPOSITION OF THE EUCHARIDAE 513

previously the area of dead tissue is almost entirely posterior to the
surface incision.

It has been mentioned that Schizaspidia antennata oviposits also in
the heavy fleshy stems of the flower clusters of certain trees. At these
times it often happens that the eggs are deposited externally rather
than embedded in the stems. The tissues of the stem are very soft and
tender and offer very little resistance to the ovipositor. It is inserted
transversely to the axis of the stem but, because of its curvature, the
tip breaks through the surface farther on, thus making a double per-
foration of the epidermis such as is done with a curved surgeon’s
needle. The eggs are then extruded from the ovipositor and only the
slender anterior stalks remain embedded in the puncture.

Orasema smithi inserts the ovipositor in the same way as Schiza-
spidia antennata, and its incisions are readily recognized by the tri-
angular form of the surface puncture. The single egg deposited at each
insertion is found at the anterior end of the cavity rather than at the
center.

The female of Pszlogaster antennatus makes a very minute puncture
in the epidermis of the mango leaf and embeds the pointed posterior
tip of the egg therein so that it is firmly held in an erect position, with
the anterior stalk turgid and straight.

The form of the incision and the cavity produced in the plant tissue
are thus seen to differ among species and they permit of provisional
recognition in the field. An examination of the ovipositors of the fe-
males often gives a clue to the manner and place of oviposition. A
heavy curved ovipositor points to oviposition in succulent leaves and
stems whereas one which is long, straight, and slender indicates that
the eggs are probably placed in buds or seed pods.

SUITABILITY OF DIFFERENT PLANTS FOR OVIPOSITION

The suitability or otherwise of a plant for oviposition by Eucharidae
appears to be governed mainly by the physical qualities of the part
in which the eggs are placed, rather than by definite attraction.
Stilbula tenwicornis, a bud-ovipositing species, favors mulberry in
northern Japan, but it also oviposits to a lesser extent in chestnut,
birch, and Cladrastis, while in Chosen it was seen to do so in the buds
of oak. These buds have certain qualities in common. The scales, while
firm, are not too hard for penetration by the ovipositor, the interior
of the bud has at least a small amount of free space in which the eggs
may be placed, and the sap is nonresinous. The adults of this species
are present in the field for only about three weeks each year, and they

514 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

consequently are limited in oviposition to such trees as have their buds
fully formed at that time. The same requirements are encountered
among other species, such as S. cyniformis, which oviposit in seed
receptacles.

Among the leaf-ovipositing species, those that insert the eggs in
incisions in the tissue restrict themselves to plants having leaves with
certain physical qualities. When one surface is punctured the wound
must remain permanently open to permit of the later escape of the
larvae. This rules out the plants that exude appreciable amounts of
‘sap from leaf wounds. Plants with rather fleshy and smooth leaves,
such as Artocarpus, Citrus, and Codiaeum, are most frequently chosen.
The two leaf-ovipositing species observed in Ceylon, representing the
genera Parapsilogaster and Chalcura, both deposit the great bulk of
their eggs in Artocarpus. Orasema smithi and Kapala terminalis, which
were found in the same habitat at Hoyo Colorado, Cuba, both ovi-
posit in the leaves of Casearia. O. coloradensis, however, chooses the
small and very delicate leaves of Stylosanthes. The collection notes of
C. F. Baker, given by Gahan in his discussion of O. wheeler: Wheeler,
show three collections on separate dates on Eriogonum at Fort Collins,
Colorado. It is quite possible that the females were ovipositing in
the leaves of that plant.

In the temperate regions, where the various species apparently have
only a single generation each year and the adults are present for only
a very short period, the number of plant species that serve for oviposi-
tion is at a minimum and a single one may suffice. That chosen in one
locality may differ from the one favored in another. Some of the
tropical Eucharidae, most of which are assumed to have overlapping
generations throughout the year, may change their oviposition plants
with the seasons. This is of most probable occurrence among those
that oviposit in buds, particularly flower buds, and seed receptacles.

The above generalization regarding the factors influencing the
choice of plants for oviposition apparently does not hold true with
species that deposit their eggs in expanding flower buds, and some,
at least, exhibit a strong response to what is apparently an odor stimu-
lus. The most striking example of such a reaction was observed in
Eucharis scutellaris, which places its eggs in the flower buds of Cebatha
orbiculata. A sprig of this vine held near an ant nest from which
Eucharis is emerging will attract all females in the vicinity within a
few minutes. If shaken off they immediately return to the buds and
cling to them tenaciously. The males, however, are not attracted to
these buds.

Dec. 15, 1940 CLAUSEN: OVIPOSITION OF THE EUCHARIDAE 515

Only two species are known to deposit their eggs at random on the
leaf surface, but in these instances the plants favored for oviposition
have leaves with a hairless and glossy under surface.

RATE OF OVIPOSITION

The total egg capacity of the females of the Eucharidae ranges from
a minimum of about 1,000 to a maximum of 10,000 or more. The eggs
are very minute, seldom exceeding 0.2 mm in length even in the larger
species, and those of the smaller species may not exceed 0.1 mm. The
smaller total given above is for Stilbula tenuicornis, which deposits
the entire lot en masse in a bud during an elapsed time averaging 20
minutes. This is at the rate of one egg a second, and consequently
they must flow from the ovipositor in virtually an unbroken stream
during this period. Where the parasite population is high, as in one
locality near Koiwai, Japan, several of these masses are deposited in
each bud, and one rather sparse mulberry bush 7 feet in height was
estimated to contain 4,320,000 eggs. One bud was found to contain
24 egg masses. Because of the limited space available in the bud a
high pressure must be exerted by the later females in forcing their
eggs into the bud, and instances were seen where previously deposited
eggs had been forced out through the older ovipositor puncture holes
in the bud scales. The most striking illustration of such pressure was
observed in S. manipurensis. One Flamingia bud showed a “‘ribbon”’ of
eggs 2.5 mm in length and 1.0 mm in width that had been forced out
from beneath the margin of a bud scale at a distance of 2 mm from
the point of insertion of the ovipositor.

An undetermined species of Kapala from Cuba, which deposits its
eggs at random on the leaf surface, has an exceedingly high egg capac-
ity. One female deposited a total of approximately 10,000 eggs during
a period of six hours. While the rapidity of deposition does not equal
that of Stelbula, yet the total is much greater. Parker (1937) mentions
the possibility of the production of 10,000 to 15,000 eggs by each
female of Stilbula cyniformis.

In general it is the habit of the species that deposit their eggs en
masse to complete oviposition the day of emergence from the host
nest, and this appears to be true also of those that place them at ran-
dom on the surface of leaves. Those that insert them singly or in small
groups in leaf tissue are more deliberate in their oviposition activities
and this may extend over one week or more. The number of eggs de-
posited each day is consequently only a few hundred, though a female
of Schizaspidia antennata was seen to make 10 to 12 insertions of the

516 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

ovipositor a minute, which represents a deposition of about 30 eggs
during that period. In other species, however, the interval between
insertions 1s much longer.

LITERATURE CITED

CLAUSEN, Curtis P. The biology of Schizaspidia tenuicornis Ashm., a eucharid para-
site of Camponotus. Ann. Ent. Soc. Amer. 16: 195—[219], illus. 1923.

. The manner of oviposition and the planidium of Schizaspidia manipurensis n. sp.
(Hymen., Eucharidae). Proc. Ent. Soe. Washington 30: 80-86, illus. 1928.

GAHAN, A. B. A contribution to the knowledge of the Eucharidae (Hymenoptera: Chalci-
doidea). Proc. U. 8S. Nat. Mus. 88: 425-458. 1940.

Iso, Ter. Some Philippine eucharids with notes on their oviposition habits. Imp.
Agr. Exp. Sta. Japan [Nishigahara, Tokyo] Bull. 3: 203-212, illus. 1982.

ParKER, H. L. On the oviposition habits of Stilbula cynipiformis Rossi (Hymen..,
Eucharidae). Proc. Ent. Soc. Washington 39: 1-3, illus. 1937.

Pierce, W. D., and Morritu, A. W. Notes on the entomology of the Arizona wild cot-
ton. Proc. Ent. Soc. Washington 16: 14-23. 1914.

SmitH, H.S. The chalcidoid genus Perilampus and its relation to the problem of parasite
introduction. U.S. Bur. Ent. Tech. Ser. 19: 33-69, illus. 1912.

. The habit of leaf-oviposition among the parasitic Hymenoptera. Psyche 24: 63-—
68, illus. 1917.

WHEELER, WILLIAM Morton. The polymorphism of ants, with an account of some singu-
lar abnormalities due to parasitism. Bull. Amer. Mus. Nat. Hist. 23: 1-98, illus.
1907.

ENTOMOLOGY.—A genus of histerid beetles new to the United
States! Rupert L. WENZEL, Field Museum of Natural History.
(Communicated by Epwarp A. CHAPIN.)

During the summer of 1938, I had occasion to visit the U. 8. Na-
tional Museum and to examine briefly the Histeridae of that institu-
tion’s collection. A specimen collected in Texas and determined by
Herbert 8S. Barber as Reninus salvini Lewis was called to my attention.
Reference to the original description of that species revealed a dis-
crepancy between the specimen and the description; however, on the
basis of notes made by Dr. Gilbert J. Arrow, of the British Museum,
who kindly examined the type, there can be no doubt that the Texas
example is to be referred to that name. Since the genus Renznus has
not heretofore been known from America north of Mexico, a generic
and species description are given here. Acknowledgment is due Dr.
Edward A. Chapin and H. 8. Barber for their kind cooperation in
making the specimen available for study.

Genus Reninus (s. str.) Lewis

Reninus Lewis, 1889, p. 275.
Renia Lewis, 1885, iO. 467,

Form ablone: oval, moderately convex. Head with a marginal carina on
each side of the epistoma, the carina continuous with the supraorbital stria.
Labrum transverse, its anterior margin straight or with a slight production
inferiorly. Antennae with nine articles, the club consisting of a ‘“‘single”’
article, which is strongly sclerotized at base and densely pubescent at apex;

1 Received July 25, 1940.

Dec. 15, 1940 WENZEL: HISTERID BEETLES U7)

flagellum articulated laterally to the antennal scape, which is angulately
swollen distally. Pronotum transverse, anterior angles obliquely truncated;
marginal pronotal stria present, lateral pronotal striae absent. Elytral striae
earinate. Propygidium transverse, hexagonal, nearly twice as broad as long.
Pygidium subcircular, nearly vertical. Prosternal keel striate, basal margin
deeply, angulately incised; prosternal lobe broad. Antennal cavities deep,
visible from beneath. Mesosternum very short, consisting of little more than
a strongly angulate process, which fits into the incised prosternal base.
Tibiae rather strongly expanded; all the tarsal grooves distinct, straight, only
their inner margins well defined. Outer margin of anterior tibiae broadly,
evenly arcuate, multidenticulate, the denticles fine, short, spinelike. Outer
margin of middle and posterior tibiae more or less angulate, denticulate as
in the anterior tibiae. All the tarsi of five articles, ultimate article bearing
two claws.

Genotype.—Reninus meticulosus Lewis, 1885, p. 467.

According to Reichensperger (1935, p. 26), the genus Brachylister Bick-
hardt (1917, p. 234) should be placed under Reninus and is not to be ac-
corded any higher ranking than that of a subgenus. The arguments in favor
of this arrangement are well founded, and it will probably be most satis-
factory to follow it.

Fig. 1.—Reninus salvint Lewis: View showing gross structure of
under side of prothorax.

My diagnosis of Renznus would undoubtedly be of more value if it were
based on all the species known; however, since many of the forms are un-
available, the description is based on characters known to me, and it will
serve to separate A. salvini from the species of other North American histerid
genera.
~ Reninus belongs to the tribe Hetaeriomorphini of the subfamily Hetaeri-
inae and may be separated from the other North American genera of the
tribe as follows:

fet lwarrawiumecorsalestmaes. i222 eh a! So a es eh bee ee ee 2
Elytra rather densely punctate, without dorsal striae................ 4

2. Anterior margin of mesosternum deeply emarginate to receive the pro-
sternal base; anterior tibiae subcircular; a lateral pronotal stria present

6S AROS Bs eS Ol Ee eee ree ee eae Yarmister Wenzel (1939, p. 391)
Anterior margin of mesosternum produced (feebly in Ulkeus), the meso-
sternal process received in the emarginate prosternal base; anterior
tibiae not subcircular, though they may be strongly expanded; pro-
notum without a lateral stria in addition to the marginal stria...... 3

518 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

3. Pronotum without a lateral gibbous area; anterior margin of mesosternum
strongly, angulately produced, the process received in the strongly
emarginate prosternal base; elytra without rows of trichomes along the
Strigeyek.. tO Oe Soe ae Re ee Reninus Lewis

Pronotum with a longitudinal gibbous area on each side; anterior margin
of mesosternum feebly, obtusely produced (nearly truncate); elytra
with rows of trichomes along the striae... Ulkeus Horn (1885, p. 143)

4. Elytra without dorsal striae, rather densely punctate; pronotum gibbous
at the sides, the gibbous area divided into distinct cushionlike lobes;
anterior margin of mesosternum produced and received in the angu-
lately emarginate prosternal base.....Terapus Marseul (1862, p. 680)

The tribe Hetaeriini (composed in North America of the genera Echinodes
and Hetaerius) may be separated from the Hetaeriomorphini by the condi-
tion of the antennal club, which is oval in the latter tribe; in the Hetaeriini
the club is cylindrical and truncated at the tip, the truncated tip being
densely pubescent.

Reninus salvini (Lewis) JPiiazs I

Renia salvini Lewis, 1888, p. 220, pl. 8, figs. 6, 7.
Reninus salvint Lewis, 1905, p. 52.

Form oblong-oval, moderately convex. Color deep reddish brown, shining.
Head, labrum, and mandibles with fine, sparse, setigerous punctures, those
of the mandibles aciculate. Mandibles and vertex of head with a few fine
rugae. Supraorbital stria distinct but somewhat broken up, continuous on
each side with the cariniform marginal stria which commences within the
eyes and extends to the anterior margin of the epistoma.

Pronotum a little less than twice as wide as long, rather strongly impressed
on each side near the anterior angles, less strongly impressed on each side
near base, sharply, sparsely punctulate throughout; sides nearly straight,
moderately converging to the truncated anterior angles. Marginal pronotal
stria extremely fine, cariniform, complete laterally, interrupted behind the
head.

Elytra sharply, sparsely punctulate throughout. Epipleura finely strigose,
with two fine cariniform striae, of which the outer is short and apical, the
inner (marginal elytral stria) is very close to the epipleural margin basally
but extends dorsally to transverse the epipleural fossette on apical two-thirds,
and from thence extends half way across the elytral apical margin. External
subhumeral stria complete, cariniform, strongly sinuous, and “‘fused’’ with
the internal subhumeral stria for a short distance near the middle; internal
subhumeral stria complete, nearly straight, costiform basally, finely carini-
form and close to the first dorsal stria near apex, forming the margin between
the elytral dise and the epipleuron. First dorsal stria complete, cariniform
basally, costiform apically ; second dorsal stria finely cariniform, very shghtly
abbreviated at apex; third and fourth dorsal striae finely cariniform, extend-
ing to about apical third, the fourth stria arching broadly at base and re-
curving briefly along the suture; sutural stria absent; internal subhumeral
stria and the dorsal striae strongly, inwardly, transversely hooked at base.
Propygidium and pygidium sparsely punctulate.

Prosternum moderately broad, the keel completely margined, the carinal
striae finely cariniform, joined in an arch anteriorly and continuous at base
along the incised margin; lateral prosternal striae cariniform, divergent and
ascending, extending to the prosternal lobe; on each side between the carinal

Dec. 15, 1940 WENZEL: HISTERID BEETLES 519

and lateral striae are two fine abbreviated striae: prosternum (except the
enclosed keel) and prosternal lobe microscopically, densely strigose and
sparsely, finely punctate, the punctures elongate, linear. Prosternal lobe with
the anterior margin truncate, margined, the marginal stria deep and termi-
nating on each side in deep, elongate foveae.

Mesosternum very short, strongly, angulately produced at middle; meso-
metasternal stria slightly anterior to the meso-metasternal suture, and
trisinuate, the middle sinuation strongest. Metasternum sparsely, remotely
punctulate, with four apically diverging striae on each side, these striae
joined medial to the middle coxal cavity; the innermost stria is straight and
extends posteriorly one-half the length of the metasternum; the next stria
is complete and extends to the anterior margin of the posterior coxal cavity;
the two outer stria are arcuate and extend laterally on the elevated meta-
sternal sides. Meso- and metathoracic pleurites with a number of strioliform
punctures. Anterior margin of first abdominal sternite with a complete
transverse, crenate, marginal stria; within the coxae on each side are three
eariniform, longitudinal striae of varying length.

Length: 3.5—4.2 mm. Width: 2.7 mm.

Remarks.—The specimen upon which this description is based was col-
lected by E. V. Walters in a nest of Atta texana (Buckley) at a depth of
between 6 and 10 feet at San Antonio, Tex., January 17, 1935.

Lewis’s type of this species was collected in an ant nest beneath a stone at
Atlisea, Puebla, Mexico, but the host was not recorded. Later (1907, p. 105)
Lewis recorded Atta cephalotes as a host species, and Bickhardt (1917, p. 241)
gives Atta fervens Say (sic). Other Reninus are known to be attaphiles, and one
species (R. meticulosus Lewis) has been recorded from Atta nests (A. sexdens,
fide Reichensperger, loc. cit.) and from termite galleries (Lewis). The atta-
philous Histeridae, believed to be chiefly synechthrans, are few in number,
the only other species apparently known from the United States being
Acritus attaphilus Wenzel (loc. cit., p. 384).

LITERATURE CITED

BICKHARDT, HEINRICH. Histeridae, in Wytsman’s ‘“‘Genera Insectorum,”’ fasc. oe b,
SUZ Rp ps Loyple ~ 196-17.

Horn, GrorGE. Contributions to the coleopterology of the United States (No. 4). Trans.
Amer. Ent. Soc. 12: 128-162. 1885.

Lewis, GrorGE. New species of Histeridae, with synonymical notes. Ann. Mag. Nat.
Hist. (5) 15: 456-473. 1885.

———. Histeridae. Biol. Centr.-Amer. Insecta, 2 (1): 182-244. 188%.

On a new genus of Coleoptera [has appended note on substitution of Rendon for

Renia]. Ann. Mag. Nat. Hist. (6) 14: 273-275. 1889.

. A systematic cataloque of Histerrdae. 81 pp. London, 1905.

. On new species of Histeridae and notices of others. Ann. Mag. Nat. Hist. (7)

. 20: 95-107. 1907.

Marseut, M.S. A. pg. Supplément a la monographie des Histerides. Ann. Soc. Ent. Fr.
(4) 2: 669-702. 1862.

REICHENSPERGER, August. Beitrag zur Kenntnis attaphiler Histeriden aus Brasilien.
Rev. Ent. 5: 25-32. 1935.

WENZEL, Rupert L. A new genus and several new species of North American Histeridae.
Ann. Ent. Soc. Amer. 32: 384-394. 1939.

520 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

ZOOLOGY .—Austrobdella anoculata, a new species of fish leech from
Greenland! J. Percy Moors, University of Pennsylvania.
(Communicated by Waupo L. ScHMITT.)

In the course of his many expeditions to the Arctic, Capt. Robert
A. Bartlett has brought back extensive collections of animal and plant
life from that region. Among the material collected on opposite sides
of Greenland, seven years apart, he obtained two contracted examples
of a small marine leech, of which the host is unknown, one from north-
west Greenland between Capes Alexander and Chalon, and the other
from northeast Greenland.

I have been unable to harmonize them with the description of any
species hitherto reported from our northern seas. Except for minor
differences, they agree with the type of Badham’s genus Austrobdella.
Of the two, the paratype is somewhat less contracted and distorted
and for that reason was sectioned for study of the internal anatomy.
The drawings and description are based upon both specimens.

Austrobdella anoculata, new species Figs. 1-4

Diagnosis.—Similar to A. translucens Badham,? but distinguished by ab-
sence of eyes, abdomen less abruptly shouldered in adult, somites typically
tri- (sex-) annulate; gonopores separated by two annuli, ovisacs short, with-
out prolonged posterior lobes; last pair of gastric caeca with about one-fifth
of their caudal ends disunited.

Description.—Body divided into two regions, “neck” and “‘abdomen,”’ the
former short and subcylindrical, the latter about three times as long, ab-
ruptly wider and moderately depressed. Measurements in millimeters of
type: Length 4.6, to & pore 1.0; widths, cephalic sucker (contracted) about
0.4, at & pore 0.65, maximum (XIX—XX) 1.8, anus 0.7; caudal sucker 0.9;
depths not measured but in neck slightly less than widths, in abdomen about
three-fourths widths. Paratype at same points 6.8, 1, 4.7, 0.8, 2.1, 0.75, 1.0;
maximum depth about 1.4. Cephalic sucker small, about one-half diameter
of caudal sucker, normally cup-shaped, but so contracted in both specimens
that ventrally it appears as a thickened annular rim surrounding a deep cen-
tral depression; in dorsal aspect hemispherical, not definitely wider than the
first nuchal somites, without obvious markings, only a few very faint traces
of annuli on caudal part and a few scattered, very minute, sensory papillae.
Eyes absent, in sections a few pigment granules near middle of head, but no
pigment cups or visual cells. Mouth seen only in sections as a minute pore
on the cephalic slope of a slight papilla at the center of the ventral face of the
sucker. Neck sharply differentiated from abdomen (most so in paratype), sub-
cylindrical, slightly depressed, short, about one-sixth or one-seventh length
of abdomen, its maximum width about one-seventh that of the widest part
of the abdomen, increasing slightly in width cephalocaudad but again slight-
ly contracted at the clitellum; preclitellar annuli about 13, but irregular,
some of them double; intermetameric furrows, including the nuchal groove,

1 Received August 6, 1940.
2 Quart. Journ. Micr. Sci. (new ser.) 62:1. 1916.

1
fico sls ==

1. Annulation in dorsal aspect as worked out from a co
In most respects they agree. Where they differ the

clearest is usually represented, or in other

the left side only. a, Anus; at, atrium; de, ductu
vc, caecate intestine; zs, saccate intestine; m,
tum; s, stomach or crop; ¢ 1-5, testes of left side; v

522 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

generally deeper than the interannular furrows and the only visible external
metameric character; most somites 3-annulate. Clitellum ill-defined, somite
X not modified, XI and XII somewhat narrower, about as wide as the first
yuchal annulus, XII embraced at its caudal end by the anterior fold of the
wider first abdominal annulus, into which it is slightly recessed; somites XI
and XII triannulate dorsally, biannulate ventrally, the second annulus of
XI enlarged in the medial ventral field to encroach upon the first annulus of
XII, and bearing the & gonopore; on venter all clitellar annuli longer than
those of the preceding somites; gonopores (Fig. 2) separated by two annuli,
the o& on the caudal margin of XI a3, the 2 at XII a2/a3, both small and
obscure, especially the female, which on the type is concealed beneath the
edge of projecting rim of somite XIII but on the paratype is exposed. Ab-
domen much wider than neck throughout, outline elliptical, the greatest
width near middle at somites XVITI-XX (relatively much wider in the
type) moderately depressed; cephalic end truncate but not expanded shoul-
derlike, embracing the last clitellar annulus, but prepucal fold only slightly
developed; caudal end tapered to the sucker, for which it forms a definite
peduncle; no lateral pulsatile vesicles and no other external metameric struc-
tures (nephropores, sensillae) discernible, except that intersegmental fur-
rows are usually deeper than the others. Abdominal somites typically 3(6)-
annulate but some of them definitely 6-annulate with the primary triannula-
tion obscure, furrows often irregular or incomplete. Anus a well marked pore
on the caudal peduncle followed by two incomplete annuli. Caudal sucker
about twice size of cephalic in all dimensions, deeply cupped, regular, di-
rected caudad, with about 48 minute marginal crenulations; dorsal face
areolated and divided by faint furrows into three or four obscure concentric
rings, each of which bears a circle of very minute papillae which are most
distinct on the smooth ventral face. No natural color remains, but the type is
stained a uniform green, which may be due to preservation in a copper tank.

Annulation (Fig. 1).—Often irregular and in places, especially at clitellum
and caudal end of abdomen, difficult to interpret. On most somites primary
and secondary furrows are distinguishable by their relative depth but in
places this distinction is lost. I-V, cephalic sucker (head), no annuli distin-
guishable except some faint traces of one or two at caudalend. VI 2-annulate,
first annulus (al a2) larger and very distinct, separated from sucker by a deep
nuchal furrow, the second (a3) very small and not separated ventrally. VII
2 or 3-annulate, al separated as a very short annulus on paratype, not dis-
tinct on type; furrows irregular and incomplete on both. VIII 3-annulate,
similarly irregular, with split and spiral primary annuli. [X and X 3-annulate
both dorsally and ventrally, a3 largest on X with a faint secondary furrow.
Normally X is first clitellar, but on these specimens apparently not closely
united with XI externally. XI 2- or 3-annulate, definitely clitellar, (a1 a2) >
a3 with a faint al/a2 furrow on the dorsum, which disappears on the venter
where a8 is enlarged medially and produced caudad into XII as a small lobe
bearing the o gonopore on its caudal margin. XII 2- or 3-annulate, 3rd
clitellar, similar to XI but shorter, more crowded, and partly concealed by
XIII, ai less developed than on XI and a3 possibly slightly subdivided, 9
gonopore very minute at a2/a3, beneath prepucal fold on type. XIII 3-an-
nulate, sharply defined by deep furrows from both XII and XIV and ab-
ruptly larger than former, with a moderately developed prepucal fold, which
partly encloses it. XIV and XV 3-annulate, with bd and b6 indicated on both
paratype and type by a slightly developed 65/06 furrow and bi and 62 on the
former; owing to contraction the annuli of both crowded together and piled

Dec. 15, 1940 MOORE: A NEW FISH LEECH FROM GREENLAND 523

up, making the region very opaque. X VI-X XII normally 3(6)-annulate. The
normal condition is that the three primary annuli are approximately equal
and each divided by shallower furrows into two secondary annuli, but be-
cause of different degrees of contraction of the two specimens and of greater
or less engorgement of different gastric caeca of the same specimen there is
much irregularity. On the type specimen the triannulate condition dominates
but the shallower secondary furrows are usually present, especially on the
more contracted side, where annuli stand out more prominently, but a3 is
rarely and a2 more frequently undivided. On the paratype XVI to XIX are
much distended by the large blood-filled caeca, which stretch the integument
so that it is thin and transparent and the distinction between primary and
secondary furrows is nearly lost (Fig. 1). This shows best on XVI and XVII.
XVITI-X XII more typical, all annuli may be equal or a2 smaller and a3
larger. XXIII-XXV 3-annulate, but progressively reduced in size and the
annulation very irregular, especially on the type. XX VI and X XVII 2-an-
nulate, but irregular, the latter much smaller and bearing the anus on the
anterior margin of the first annulus.

Fig. 4.—Austrobdella anoculata: Dorsal and ventral photo-
graphic views of type. X 33.

Anatomy (Fig. 3).—Some of the internal organs can be seen through the
integuments sufficiently to determine their position and roughly their form,
but the details were worked out from sections of the paratype in which
some of the structures were obscured by the engorged gastric caeca. Anat-
omy closely patterned after that of the genotype, A. translucens. Ventral
nerve ganglia of complete somites in annuli a2, chiefly in b4. Pharynx with
medial ventral and paired dorsolateral muscular ridges; salivary glands
diffuse, of very large single cells in somites VII-X. Gastric caeca seven pairs
in XIII to XIX, the first six reaching nearly to the lateral body walls and
lobed at the ends, the last pair coalesced completely except at the caudal end,
where the two caeca remain distinct for a longer distance than in A. trans-
lucens; intestine with two or three pairs of small simple caeca at the anterior
end, following which it expands into a wide, simple sac reaching nearly to the
end of the united gastric caeca and giving rise from its dorsal face a short
distance anterior to its caudal end to a narrow, tubular rectum which ends
at the anus. Reproductive organs similar to those of A. translucens except
that the median chamber of the atrium is relatively smaller and the duct
shorter; cornu large and sperm ducts with several loose coils at the atrial
end; testes five pairs at XIV/XV to XVIII/XIX, alternating with gastric
caeca. Vagina a simple vertical duct with only a slight enlargement and not
expanded into a sac or bulb but encased in a mass of glands; it divides be-
neath the nerve cord into the paired ovarian sacs lacking the narrow duct
and anterior lobe shown by Badham, both ovisacs very short, the one reach-
ing to XIII only, the other to the middle of XIV, both containing developing
ova in early stages.

524 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

Material examined.—Two specimens, one of which, the holotype, U.S.N.M.
no. 20573, was obtained in 25 to 40 fathoms between Capes Alexander and
Chalon, northwest Greenland, by Capt. Robert A. Bartlett, August 2, 1937
(station 27, seine haul). The other specimen, the paratype, comes from
northeast Greenland, where it was collected by Captain Bartlett in 1930
(No. 14).

Remarks.—Owing to the limited material and its indifferent preservation,
it has not been possible to arrive at such unequivocal conclusions and to pre-
pare as satisfactory a description as could be wished. Many nominal species
of fish leeches have been described from Arctic and sub-Arctic waters. Some
of the early descriptions are so brief and ambiguous that the species to which
they refer have never been certainly determined. Synonymies are still con-
fused. It is possible that the species here described may belong to one of
these, but none were found to agree. Concerning the generic reference there
is little doubt. On a preliminary study it was thought that these leeches
might belong to Abranchus or Ottonia, but externally the form and annula-
tion differ from those genera and internally the coalescence of the last pair of
gastric caeca is sharply differential.

On the other hand, the resemblance to Austrobdella, both externally and
internally is very close. So far as it could be worked out, the annulation
agrees closely with that of A. translucens Badham. The external form and
proportions of parts are very similar. Neither of the two specimens is so
strongly shouldered at the junction of the neck and abdomen as in Bad-
ham’s most mature individuals. The anatomy of the alimentary canal and re-
productive organs, so far as worked out, is very close in the two species, the
principal differences being that in the type species the last pair of gastric
caeca are more completely united at the caudal end than in A. anoculata, and
that the ovisacs of the latter are not prolonged caudally and have the vaginal
duct short and simple. These ovarian differences may be due to immaturity.
Owing to the state of the material, a study of the distribution of the sinuses
was not attempted.

ICHTHYOLOGY.—Hadropterus palmaris, a new darter from the
Alabama River System.) Reeve M. Battey. (Communicated
by Lronarp P. SCHULTZ.)

In a collection of fishes from the Etowah River in northern Georgia,
six specimens of a hitherto undescribed species of Hadropterus were
taken. Twenty-seven additional specimens from two localities in
Alabama have been placed at my disposal through the generosity of
Dr. Carl L. Hubbs.? This handsomely colored species is herein de-
scribed under the name Hadropterus palmaris.*

1 Received August 24, 1940. Journal Paper no. J791 of the Iowa Agricultural Ex-
periment Station, Ames, Iowa. Project no. 651.

2 T am indebted to Dr. Hubbs for helpful suggestions in preparing this paper.

3 Palmarts =a prize.

Dec. 15, 1940 BAILEY: A NEW DARTER 525

Hadropterus palmaris, new species Fig. 1

BRONZE DARTER

Holotype—An adult male (Univ. Michigan Mus. Zool. no. 126179),
65.5 mm in standard length, collected in the Etowah River, known locally
as the High Tower River (tributary to the Coosa River, tributary to the
Alabama River), just above the bridge at U. S. Highway 19, 4 miles south-
west of Dahlonega, Lumpkin County, Ga., on August 25, 1939, by Reeve M.
and Marian K. Bailey (field no. B39:59).

Paratypes.—F ive specimens were taken with the holotype: U.M.M.Z.
no. 126180 (2), U. 8. National Museum no. 117881 (2), Iowa State College
no.1 (1). U.M.M.Z. nos. 111232 and 111133 (23) collected in Saugahatchee
Creek (tributary to the Tallapoosa River), Ala., October 24, 1930, by F. E.
Guyton. U.M.M.Z. no. 111234 (4) collected 3 miles east of Tuskegee,
Macon County, Ala., June 3, 1931, by F. E. Guyton.

Diagnosis.—A highly colored species of Hadropterus with moderate sized
scales, 59 to 73 (usually 69 or fewer) along lateral line; nape and cheeks fully
scaled; opercles imperfectly scaled (especially in adults); and midventral
scales greatly enlarged in males. Dorsal XI to XIV-10 to 13, the soft portion
high; anal II, 7 to 10; and pectorals 13 to 15. Preopercle entire. Gill mem-
branes separated; the distance from their union to insertion of pelvic fins
0.75 to 0.95 in distance from tip of mandible to union of gill membranes.
Snout short, 3.6 to 4.4 in head length. No subocular dark bar; dorsolateral
surface punctated with dark; 2 large light spots at base of caudal.

Description.—The various body proportions (Table 1), fin-ray and scale-
row counts (Table 2), and the photograph (Fig. 1) indicate clearly many of the
characteristics of the species. The juveniles are seen to differ from the adults
in their larger eyes, shorter snouts, and the somewhat greater distance be-
tween the union of the gill membranes and the tip of the mandible, but
variations in body proportions within the species appear to be slight. The
apparently broader interorbital width in the Alabama specimens seems to
be due to preservation. In most of the fin-ray and sceale-row counts (Table 2)
the Alabama and Georgia specimens agree rather closely; but in lateral-line
scale count the range of variation in the Alabama specimens is greater (59
to 73) than in those from Georgia (59 to 63). The increased number of pec-
toral rays in the Georgia specimens (average 14.8) as contrasted with those
from Alabama (average 13.7) is notable.

The preopercle is entire, the horizontal arm scarcely longer than the verti-
cal arm. The premaxillary frenum is slightly narrower than the thick, termi-
nal, upper lip, and about two-thirds the diameter of the pupil. The moderately
heavy mandibles are well separated, diverging but little for about three-
fourths of their length from the symphysis, behind which they flare outward
more sharply. The mandibular frenum is somewhat wider than the premaxil-
lary frenum. The snout is bluntly decurved; the upper edge of the upper lips
is on the level of the bottom of the eye, and the lower Jaw is slightly included.
In L.8.C. 1 the gill-rakers number 3+11, of which 1 above the angle and 4
below are very short. The longest gill-raker when depressed scarcely extends
to the base of the second raker below. In the same specimen the vertebral
count is 18+23=41 (hypural included); in the first caudal vertebra no
haemal spine is developed but the haemal processes are united. The vomer
bears a strong cluster of teeth, and the palatines are strongly toothed along
their length. In H. nigrofasciatus the palatine teeth are few and restricted to
the anterior portion of the bones.

526 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

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Dec. 15, 1940 BAILEY: A NEW DARTER 527

TABLE 2.— FREQUENCY DISTRIBUTIONS OF FIN-RAY AND SCALE Counts! on
HADROPTERUS PALMARIS (the counts of the holotype are indicated
by asterisks).

Dorsal Dorsal Anal anaes
ines soft rays? soft rays? ean
Locality ae rays?
WL |) ee |) iss) ez) TI) |) atk ie | 13 a | 8 | 9 1K) |) gs} |) aleb yas
Geortiaeerern. cis ok cs hoes ces — 1 5*| — | — 2 4*| — | — 1 5*, — | — 2 4%
JAE) OOTND 6.6 5.6 Oa ee aL |} 20) |} tks 1 els 6 2 A |) U2 |} I) 3 1 | 19 a
PIR Gall ee fey 3 scanesesev'e@ svevavaldians Wi} io yf BO 1 IL || 2X0) |) Wl@) 2 Py |) Wes Als) 3 iL }) AL |} dual
Pectoral rays Bcale ows
Locality (both sides) Above lateral line Below lateral line
Ig Nw 7 | 8 9 1) ah) TB ewe i Te
(GOORHE, 5 66.4 os OGnbe ae — 3% Q* 1 5* — 3 2* —- 1 _
PAN AD AIA ey ee aiess cls lo one 6 16 36 2, 1 20 6 3 2 8 12 2
Total. eee 16 39 11 2 | 25 6 6 4 8 13 2
Scale rows
Locality Along lateral line Around caudal
peduncle
59| 60] 61] 62] 63] 64] 65] 66] 67| 68] 69] 70) 71] 72] 73} 20) 21 22 23) 24) 25
(GXOTUE Son. G eRe eeee 2*/—| 1] —)| 3} —] — — — — — —| —| —| — —) 2} 1) 2% 1 —
JANE STTE Con GUO oOb OF TH BS} ap TN BA SB Tu aN a a SE HP RE 1B) AL
INO IGES 6. SOEs 3) fanaa | <>) | (| A re) | fue | => | > | mV | ——— | PT | ce ce) em Ce) He mT

1 The scales above the lateral line were counted downward and backward from the origin of the second
dorsal fin; those below the lateral line were counted upward and forward from the origin of the anal fin; those
along the lateral line were counted to the base of the caudal rays; those around the caudal peduncle represent a
minimum enumeration.

2 The last ray of the soft dorsal and anal fins was counted as double at the base.

3 Seventeen principal caudal rays were counted in all specimens.

Five pores are given off by the lateral canal of the head, the anterior from a
short downward projecting tube and the next three at the tips of long slender
tubes, which project downward and backward. The supratemporal canal is
complete, giving off a median pore from a short backward projecting tube,
and one lateral pore on each side from longer and slenderer tubes that project
backward and slightly downward. A single postorbital pore opens from a
short backward projecting tube from the supraorbital canal on each side, and
the interorbital pore of each side opens from a short tube that projects out-
ward or outward and backward. The posterior nasal pore opens from a short
tube just above the anterior end of the slitlike, posterior nostril. The an-
terior nasal pore lies anterodorsal to the tubular, anterior nostril. The
coronal pore opens at the end of a long tube which may be curved slightly
to the left or right. The infraorbital canal is complete with 8 pores; of which
the anteriormost lies below and behind the anterior nostril, the second and
third open from slender tubes near the edge of the maxillary groove, the

528 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 12

fourth emerges from a short upward projecting tube, and the posterior
four open from long slender tubes that project downward and backward.
Of the ten operculomandibular pores the six on the preopercle open from
short, side tubes. The complete lateral line is very slightly arched anteriorly.
(Nomenclature of canals and pores from Hubbs and Cannon, p. 10, pl. 2.4)

The genital papilla of the adult male is a depressed and rounded, conical
projection, which is smooth except for a short, longitudinal groove near its
tip; that of the female is broadly semicircular, with several deep grooves
radiating forward and outward from the orifice.

The pelvic fins are separated by a space equal to about four-fifths of the
pelvic base. Between the pelvics there are two or three enlarged median
scales, which may or may not be followed posteriorly by a naked strip for
two-fifths of the length of the pelvic fins. In males the remainder of the mid-
line of the belly is provided with a series of greatly enlarged scales with long
and very strong ctenil; in females these scales are scarcely or not at all en-
larged and the ctenii are little developed. There are one or two slightly to
moderately enlarged scales at the union of the pelvic bones near the middle

Fig. 1.—Paratype of Hadropterus palmaris, U.S.N.M. 117881, an adult male 67.5
mm in standard length, from the Etowah River, Ga. The photograph, of the right side
of the fish but reversed in printing, was taken by Max E. Davis.

of the breast, and the subtriangular area between the pelvic bones is provided
with small imbedded scales; otherwise the breast is entirely naked. The nape
and cheeks are covered with well-developed imbricating scales; those of the
cheek are in seven or eight rows as counted from the eye to the postero-
ventral angle of the preopercle. The opercles are provided with deciduous
scales, varying from a complete absence of scales in some of the larger adults
to an almost complete covering in juveniles. The holotype has five small,
nonimbricate, imbedded scales on the left opercle and nine on the right.

The spinous dorsal is long and rather low and is narrowly separated from
the soft dorsal. The soft dorsal and anal are large. The first of the two anal
spines is much stronger but slightly shorter than the second. The margin of
the caudal is slightly concave. The pectorals are broadest near their middle
and extend as far back as the pelvics.

Coloration.—With the exception of evides, palmaris is the most colorful
and handsome species of Hadropterus thus far described. In life the bodies of
adult males were deep yellowish brown above, shading to dull olivaceous on
the breast. On each side were eight to ten dark greenish-black vertical bars

4 The darters of the genera Hololepis and Villora. Misc. Publ. Mus. Zool. Univ.
Michigan, 30: 1—98, pls. 1-3. 1935.

Dec. 15, 1940 BAILEY: A NEW DARTER 529

(which retained their color in preservation), each considerably wider than
the interspaces. These bars are usually continuous with their fellows across
the back, especially anteriorly; and each narrows and fades ventrally, failing
to cross the midventral line; those on the caudal peduncle extend somewhat
lower than those on the body. In females the lateral blotches are longer than
high and fail to ascend high on the sides. The bands are scarcely in contact
with one another along the lateral line in some places; in others the contact
is equal to the eye’s breadth. The cheeks and lower portions of the opercles
were (in life) dull yellowish green. The throats were iridescent with bluish-
purple reflections; in preservation they are faintly dusky but less so than
the mandibles. Unlike most species of the genus Hadropterus there is no trace
of a subocular dark bar. A narrow dark band extends forward from the
middle of the eye through the nostril, usually joining with that of the op-
posite side across the frenum. A dark bar extends backward from the eye
across the upper part of the cheek to the anterodorsal corner of the opercle.
A few dark spots are scattered on the dorsolateral surfaces of the back of the
head, and somewhat larger dark spots are present between the lateral bars
on the dorsolateral surfaces of the body.

In life the spinous dorsal was light near the base, with a rich chocolate-
brown spot on each interradial membrane. This light area was succeeded
distally by a dusky band, which shaded gradually into a rich reddish-orange
band occupying most of the distal half of the fin, which was sharply and
narrowly edged with clear white. In preservation the white edging and the
dusky band are evident in the otherwise uniformly faint pigmentation. The
soft dorsal was dull orange-yellow, with a narrow light border; a few scat-
tered dark radial spots tend to form rows about one-third and two-thirds of
the distance from the base, but in some specimens they are faint or obsolete.
At the base of the caudal there are two large light spots (each about size of
eye), which were pale yellowish orange in life. Three wavy vertical bars on
the caudal are formed by pigment confined to the rays. The caudal, anal,
pectorals, and pelvics were suffused with dull yellow in life, and all had light
distal edges. In preservation the basal half of the anal is faintly dusky; the
pelvics have a few median dusky spots; and each pectoral has a narrow
curved bar just proximad of the middle.

The color description given above was drawn from the Georgia specimens.
Juveniles from Saugahatchee Creek, Ala., vary greatly in body pattern.
Some differ but little from the adults, whereas others are heavily and ir-
regularly mottled with dark on the sides and the lateral series of blotches are
scarcely or not at all developed.

Habitat—The Etowah River is an upland stream (elevation at site of
collection about 1,300 feet), which at the time of the collection was clear,

colorless, cool (67° F.), and with a moderate to swift current. It varied from
about 35 to 60 feet in width, and the flow, estimated at 250 cubic feet a
second, was somewhat greater than normal owing to recent rains. The bot-
tom was composed largely of sand, with coarse gravel and slab-rubble riffle
areas. Vegetation was limited to slight algal growth on the stones.

All the specimens of H. palmaris were taken on a swift riffle, which was
about 6 feet long and extended the width of the stream. It varied from a few
inches to over a foot in depth. Common associates of H. palmaris on the riffie
were Poecilichthys jordani and Cottus zopherus (probably this species although

530 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

the lateral line is nearly or quite complete). Other species taken in the col-
lection were Moxostoma sp., Hypentelium etowanum, Notropis callistius,
N. x. xaenocephalus, N. trichroistius, and Micropterus coosae. Local residents
reported that rainbow trout occurred in the stream, but none were collected.
No habitat data are available for the Alabama collections.

Relationships.—H. palmaris differs trenchantly from all other known
species in the genus in coloration and color pattern; especially distinctive are
the pattern of the spinous dorsal, the uniform absence of a subocular bar, and
the presence of dark spots on the anterior back and of two round light spots
at the base of the caudal. It differs from macrocephalus, phoxocephalus, and
oxyrhynchus especially in the blunter snout, the shorter distance from the tip
of the snout to the union of the gill membranes (see Hubbs and Raney, loc.
cit.), and in the larger scales. In its entire preopercle, palmaris contrasts with
H. scierus. The Atlantic drainage species, peltatus and roanoka, differ from
palmaris, among other respects, in the larger scales and in the absence of
scales on the nape. H. maculatus and H. palmaris are quite similar in form,
although in maculatus the soft dorsal and anal are much lower, but maculatus
typically lacks scales on the predorsal area, and has more perfectly scaled
opercles than palmaris. On the basis of color pattern H. palmaris is separable
at a glance from H. nigrofasciatus, the only other described species of the
genus from the Alabama basin. It also differs from that species in the im-
perfect squamation of the opercles, the much larger and more specialized
midventral scales of the males, and in the stronger fin spines, especially the
first anal spine. Although palmaris lacks the characteristic subocular dark
bar of. evides, the two species share certain features of coloration. Both have
paired light spots at the base of the caudal, lateral blotches which are similar
in shape and color and agree in continuing upward to cross the back, and
each is highly colored in life. H. evides is a heavier and more robust species,
and in it the spinous dorsal is conspicuously higher and more rounded, and
the soft dorsal lower than in palmaris. The absence of cheek scales in evzdes
and the imperfect squamation of the opercles in palmaris are clear-cut dif-
ferentiating features.

H. palmaris seems to share more similarities with evzdes than with other
species, the two apparently standing somewhat apart from the remaining
species in the genus.

Dec. 15, 1940 SMITH: THE REMORA PHTHEIRICHTHYS LINEATUS 531

ICHTH YOLOGY.—The remora Phtheirichthys lineatus and the first
specimen from United States waters: Hucu M. Situ, U.S. Na-
tional Museum.

In the year 1791 there appeared two descriptions of a new remora
called Echeneis lineatus by Archibald Menzies in Transactions of the
Linnean Society, London (vol. 1, p. 187, pl. 17, fig. 1), and Echeneis
tropica by Bengt Anders Euphrasén in Kongliga Vetenskaps Acade-
miens Nya Handlingar, Stockholm (vol. 12, p. 317). The habitat of
E. lineata was stated to be the Pacific Ocean between the tropics, and
the single specimen 5 inches long on which the description was based
was found adhering to a turtle. For EL. tropica the locality given was
latitude 2° 9’ N., longitude 20° 49’ W. from Paris, a place in the At-
lantic Ocean about 600 miles southwest of that part of Africa now
known as Liberia, and the single specimen mentioned was discovered
attached to the nautical log of a ship when the log was pulled aboard.

These two names have generally been regarded as applying to the
same species, and there is certainly nothing in the original descriptions
to indicate specific distinctness. The outstanding character is the
possession of only 10 laminae in the cephalic disk, combined with fea-
tures that are shared with the common remora Hcheneis naucrates,
such as the very elongate body, ventral fins narrowly adnate to the
abdomen, angulate pectoral fins, middle caudal rays produced in the
young, and strongly projecting lower jaw with a cutaneous symphys-
eal flap. In 1862 Gill (Proceedings of the Academy of Natural
Sciencies of Philadelphia) created the genus Phthetrichthys and desig-
nated Echeneis lineata Menzies as the type.

With two specific names for the same fish coming into use in the
same year, there arises the question of the proper one to adopt. While
the question may not be definitely settled by the internal evidence
afforded by the respective publications, there is ground for a reason-
ably satisfactory decision affecting both priority and convenience.

The Nya Handlingar in which Euphrasén’s Echeneis tropica was
described (in a paper entitled ‘‘“Scomber atun och Echeneis tropica
beskrisne’’) was issued in quarterly parts with consecutive pagina-
tion, the paper appearing in the part for October, November, and
December 1791. It is improbable that the part was printed and dis-
tributed before the last quarter of 1791; it is possible that it was not
issued until the last month of that quarter or even the beginning of
the next quarter. |

_1 Published by permission of the Secretary of the Smithsonian Institution. Re-
ceived September 25, 1940.

532 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, NO. 12

Menzies’s paper, ‘‘Description of Three New Animals Found in the
Pacific Ocean,” was read before the Linnean Society on April 6, 1790,
and was followed by papers by other authors read in October, Novem-
ber, and December, 1790, and March, 1791, and was preceded by
papers read in 1788, 1789, and 1790, with an addendum to one of the
papers bearing date of February 6, 1791. The plate accompanying
the paper contains, in addition to a figure of the remora, figures of a
worm and a leech described by Menzies and four figures of insects re-
ferred to in a paper by another author. The make-up and general ap-
pearance of the volume strongly indicate that it was published as a
whole, and there is nothing to suggest that any of the papers were is-
sued as separates; indeed, such issuance was practically precluded by
the fact that figures illustrating papers by two different authors ap-
peared on the same plate. The title page of the volume bears date of
1791, and it may be safely inferred that the volume was issued in the
first half of that year.

But even if it can not be conclusively shown from the evidence at
hand that the name Echeneis lineatus has priority over Echeneis tro-
pica, it may be proper, and it is certainly desirable, to render a deci-
sion in favor of the former because the description was accompanied
by an illustration and because lineatus was designated as the type of
a new genus. |

The first mention of this fish as an inhabitant of the coastal waters
of the United States was by Jordan and Gilbert (Proc. U. 8. Nat.
Mus. 5:591. 1882), who recorded a specimen 4 inches long taken by
Gilbert at Charleston, 8. C., in July-August, 1882. This specimen, ap-
parently no longer in existence, was fully described and was the basis
for the statement in Jordan and Gilbert’s Synopsis of the fishes of
North America (1883) that the species ranges in ‘‘tropical seas, north
to South Carolina.’”’ In Jordan and Evermann’s Fishes of North and
Middle America (part 3, 1898) the species is reported to range also to
Pensacola, although in Evermann and Kendall’s Check-list of the
Fishes of Florida (1900), which included ‘‘all salt-water species known
from Florida within the depth limit of 1,000 fathoms,” this remora is
not listed. In the Check List of Fishes and Fishlike Vertebrates of North
and Middle America (1930), by Jordan, Evermann, and Clark, the
species appears under the name Phtheirichthys tropicus, without men-
tion of P. lineatus as a synonym.

It now remains to present a record of what appears to have been |
the first specimen of Phthetrichthys lineatus taken in United States
waters. A short time ago Dr. Leonard P. Schultz, curator of fishes in

Dec. 15, 1940 PROCEEDINGS: THE ACADEMY 533

the United States National Museum, found in the National collection
a fish labeled Rhombochirus osteochir, which proved to be a Phthezrich-
thys indistinguishable from the form described and figured by Men-
zies. The specimen came from Woods Hole, Mass., was obtained in
1871 by Vinal N. Edwards, the indefatigable collector of the United
States Fish Commission, and was catalogued in 1876, bearing the
number 20331 in the Museum register. In view of my special interest
in the fish fauna of the Woods Hole region, Dr. Schultz has kindly per-
mitted me to make known the existence of this specimen, which is
apparently the first and only one reported from our northeastern
coast. The specimen, in an excellent state of preservation, is 15.6 cm
long, with discal laminae numbering 10, dorsal rays 31, and anal rays
30, and, although nearly 60 years in alcohol, it still shows the two
whitish longitudinal stripes along the side as represented in Menzies’s
plate and embodied in his specific name.

PROCEEDINGS OF THE ACADEMY AND
AFFILIATED SOCIETIES
THE ACADEMY
363D MEETING OF THE BOARD OF MANAGERS

The 363d meeting of the Board of Managers was held in the Board Room
of the Cosmos Club on Friday, October 11, 1940. President CritrENDEN
called the meeting to order at 8:03 p.m. There were 15 present, as follows:

EK. C. CRITTENDEN - J. H. HIBBEN W. RAMBERG

F. D. Rossini G. STEINER EK. W. PrRIicE

F. C. Kracex F. M. Setzer C. L. GARNER

H. 8S. RapPPLEYE A. T. McPHERSON and by invitation

Ae Cah uLiLeR ACwae C1 ARK J. H. Kemrton
© Grom

President CRITTENDEN appointed the following Committee of Tellers
to count the ballots on the three proposed amendments to the bylaws, which
were submitted to vote of the membership on October 7: H. N. Eaton,
chairman; R. 8. Jessup; B. J. Marr; Coarutes Moon, alternate.

Chairman GARNER reported for the Committee on Meetings that the
following program of meetings has been arranged for the Academy for the
season October to April, inclusive:

October 17, 1940. Dr. W. F. G. Swann, director of the Bartel Research Foundation
of the Franklin Institute, Swarthmore, Pa., on ‘“‘The Laws of Nature.”’

November 28, 1940. Dr. VeERGiL REED, assistant director of the U. S. Bureau of Census,
on ‘The Human Side of the Census.”’

December 19, 1940. Dr. W. C. LowpERMILK, chief of the Division of Research of the
U.S. Forest Service. Title not yet announced.

January 16, 1941. Annual meeting and presidential address.

534 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

February 20, 1941. Dr. P. W. Bripeman, professor of physics at Harvard University,
Cambridge, Mass. Title not yet announced.

March 20, 1941. Awards for scientific achievement.

April 17, 1941. Tentatively, Commander F. W. REICHELDERFER, chief of the U. S.
Weather Bureau.

The date for the November meeting was changed to the 28th because the
21st is Thanksgiving Day. The October and November meetings are sched-
uled to be held in the Assembly Hall of the Cosmos Club, as in the past.
Commitments for the following months will be made later.

On recommendation by the Committee on Membership, who submitted
the names at the last meeting of the Board, 17 persons were considered indi-
vidually and duly elected to membership.

Mr. SrTzLER presented the report of the Special Committee, of which he
is chairman with R. E. Gipson and R. R. SpeNcER members, on Society
Affiliation with the Academy. The committee considered the list, published
in Vol. 30, No. 1, Jan. 15, 1940, of the JourNAL, of societies not affiliated with
the Academy and recommended action on organizations representing dis-
ciplines having a small active stable membership within the District of Co-
lumbia. The report was accepted by the Board, and the committee was in-
structed to report in detail on the Washington Section of the American
Institute of Mining and Metallurgical Engineers; the Washington Chapter
of the American Institute of Metals, and the Washington-Baltimore Psycho-
analytic Association.

The secretary reported the following status of Academy membership:

Resident Nonresident T otal
NCIC A. steeey See eweae unre 420 WAZ 547
TO GURE Ca tea agate Rone nne ee hus 30 13 43
HOnOratye oe eee ce os 3 14 17
Patronee soak fg 5 hove Sete g: —— 7 DA
MOG a et ee eee 453 156 609

Cognizance being taken of present commitments in the form of nomina-
tions, acceptances, etc., there is a minimum of 22 vacancies in the resident
membership and of 5 in the nonresident membership.

The Board adjourned at 9:18 P.M.

ELECTIONS TO MEMBERSHIP

The following were recently elected to resident membership:

Ernest ApNA Back, principal entomologist, U. 8. Bureau of Entomology
and Plant Quarantine, in recognition of his contributions relating to the con-
trol of attacks of insects on stored products and to the biology and control of
citrus white flies.

NatHAN Lincotn DRAKE, professor of organic chemistry, University of
Maryland, in recognition of his contributions to organic chemistry, in par-
ticular his researches on the constitution of the polyterpenoids.

LYNDON FREDERICK SMALL, head chemist, National Institute of Health, in
recognition of his studies on morphine derivatives.

R. Cuirrorp Hatt, principal forester, U. 8. Forest Service, in recognition
of his researches and investigations on forest policy and forest tax problems.

WaLTER J. Hamer, associate chemist, National Bureau of Standards, in
recognition of his work on hydrogen ion concentration, in particular his in- .

Dec. 15, 1940 PROCEEDINGS: CHEMICAL SOCIETY 539

vestigations on the thermodynamics of sulphuric-acid solutions and the
development of a pH method using galvanic cells without liquid junctions.

VERNE LesTEeER Harpmr, senior silviculturist, U. 8. Forest Service, in
recognition of his contributions to oleoresin production in pines, in particular
the effect of weather and soil factors on gum flow.

JAY P KINNEY, supervisor of work program, Indian Service, U. 8. Office of
Indian Affairs, in recognition of his contributions to forest law and history
and to the improvement of forest management on Indian lands.

JacoB Korz, clinical professor of obstetrics and gynecology, George
Washington University, in recognition of his contributions to endocrinology.

JOSEPH THOMAS PARDEE, senior geologist, U. 8. Geological Survey, in
recognition of his contributions to the geology of metalliferous deposits
(including manganese) in the Northwestern States.

ELizABETH ParRKER Bartscu, physician, clinical instructor in obstetrics
and gynecology, George Washington University, in recognition of her re-
searches in female endocrinology.

RAYMOND PRIc#, senior forest ecologist, U. S. Forest Service, in recogni-
tion of his researches relating to forest and range lands, particularly in con-
nection with range management, artificial revegetation, and plant nutrition.

FREDERIC G. RENNER, chief of the division of range conservation, U. 8.
Soil Conservation Service, in recognition of his researches on erosion control
and watershed protection on forest ranges and other range lands.

Oscar WILLIAM TORRESON, observer, Department of Terrestrial Magne-
tism, Carnegie Institution of Washington, in recognition of his investigations
in atmospheric electricity.

RicHMOND TUCKER ZocH, associate meterologist in charge of library, U.S.
Weather Bureau, in recognition of his contributions to mathematical
hydrology.

The following was recently elected to nonresident membership:

GrorGE Burton Riae, professor of botany, University of Washington,
Seattle, Wash., in recognition of his contributions to plant physiology, in
particular his researches on the physiology and ecology of sphagnum-bog
vegetation.

CHEMICAL SOChn IY
512TH MEETING

The 512th meeting was held in the School of Government Building of
George Washington University on Thursday, October 12, 1939, President
KRACEK presiding. After the completion of routine business the following
communications were presented in three sections:

Inorganic and Analytical Chemistry, C. R. NAESER presiding
F. W. Scowas and E. Wicuers: The Preparation of Benzoic Acid of High
Purity.
A. K. Brewer: A Thermal Method for the Separation of Isotopes.

FP. W. Guaze and A. N. Finn: A Modification of the Partition Method for
Determining Boron in Glass.

Organic and Biological Chemistry, C. M. MacKAtt presiding
L. D. GoopuutE and H. L. J. Hatumr: Recent Progress in the Chemistry of
Dervis.

M. B. Matuack: The More Important Aspects of Our Work on Lipase and
Esterase.

0386 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

L. A. SHinn and B. H. Nicouet: The Action of Periodic Acid on Amino
Alcohols and Amino Acids.

Physical Chemistry, M. H. VAN Horn presiding

S. BRUNAUER and P. H. Emmerr: Chemisorptions of Gases on Iron Syn-
thetic Ammonia Catalysts.

A. BRENNER: The Electrodeposition of Copper-Bismuth Alloys from a Per-
chlorate Bath.

C. K. Horner and D. Burk: The Catalytic Function of Molybdenum in
Nitrogen Fixation by Azotobacter.

513TH MEETING

The 513th meeting was held in the auditorium of the East Building of the
National Bureau of Standards on October 26, 1939, President KracEK
presiding. Following the completion of routine business, the Society was ad-
dressed by H. Mark, formerly of the University of Vienna, on The Elastic
Behavior of High Polymeric Substances.

514TH MEETING

The 514th meeting was held in the auditorium of the Cosmos Club on
November 9, 1939, President KraceEx presiding. The Society was addressed
by Earu P. STEVENSON, president of Arthur D. Little Co., of Cambridge,
Mass., on the subject Research in Industry. Elections were held for the offi-
cers for the Society for the year 1940. The following were elected: President,
R. M. Hann; Secretary, NorMAN BEKKEDAHL; Treasurer, E. R. SMITH;
Councilors, F. G. BrickwEppE, N. L. Draxn, R. Gitcurist, H. T. HER-
Rick, G. E. Hitsert, B. H. Nicouet, B. D. Van Evpra; Managers, M. M.
Harina, 8. B. Henpricks, J. T. Scanian, 8. T. ScuicktTanz, L. A. SHINN,
H. P. Warp.

515TH MEETING

The 515th meeting was held in the auditorium of the Cosmos Club on
November 30, 1939, President Kracx presiding. Following the completion
of routine business, the Society was addressed by Roser? EK. Lutz, of the
University of Virginia, on Some Recent Work on Furans and Hydroxy-
furanones.

516TH MEETING

The 516th meeting was held in the auditorium of the Cosmos Club on
December 14, 1939, President Krackx presiding. Following the completion
of routine business, the Society was addressed by W. FRANKENBURG on
Relations Between Photochemical and Catalytic Processes.

517TH MEETING

The 517th meeting was held jointly with the Washington Academy of
Sciences in the auditorium of the Cosmos Club on December 21, 1939. The
President of the Academy, CHARLES E. CHAMBLIss, introduced B. F. Nico-
LET, Resident Vice-President of the Academy, who presented the speaker of
the evening, C. 8. MarvEt, professor of organic chemistry of the University
of Illinois. Dr. Marvel addressed the meeting on Plastic Glasslike Resins
(Vinyl Polymers).

518TH MEETING

The 518th meeting (56th annual meeting) was held in the auditorium of

Dec. 15, 1940 PROCEEDINGS: CHEMICAL SOCIETY 537

the Cosmos Club on January 11, 1940, President HANN presiding. After the
reading of reports of the officers and committees for 1939 and the completion
of other routine business, the Society was addressed by the retiring Presi-
dent, Frank C. Kracex, who spoke on Some Chemical Aspects of Order-
Disorder Phenomena.

519TH MEETING

The 519th meeting was held in the auditorium of the Cosmos Club on
February 8, 1940, President Hann presiding. Following the completion of
routine business, the Society was addressed by L. Micuagnis, of the Rocke-
feller Institute for Medical Research, New York City, on Reversible Oxida-
tion-Reduction in Organic Compounds and Semiquinone Radicals.

520TH MEETING

The 520th meeting was held in the auditorium of the Cosmos Club on
March 14, 1940. The occasion was the annual banquet of the Society, and
119 members and guests gathered together to celebrate the awarding of the
Hillebrand Prize to R. E. Gipson, of the Geophysical Laboratory, Carnegie
Institution of Washington, for this excellent work, The Behavior of Solutions
Under High Pressures. President HANnN acted as Toastmaster. He first in-
troduced L. H. Apams, the director of the Geophysical Laboratory, who
spoke on Some Significant Aspects of High-Pressure Research. Following this,
Past-President KRracrExk introduced the recipient of the award, Dr. GiBson,
who was then presented with the certificate by Dr. Hann. Dr. Grsson then
addressed the Society, giving a résumé of his work in the field of the be-
havior of solutions under high pressures.

521sT MEETING

The 521st meeting was held in the auditorium of the Cosmos Club on
March 28, 1940, President Hann presiding. Following the completion of
routine business, the Society was addressed by PuHitit1p ABELSON, of the
Department of Terrestrial Magnetism, Carnegie Institution of Washington,
who spoke on Indentification of the Products of Uranium Fission.

522D MEETING

The 522d meeting was held in the auditorium of the Cosmos Club on
April 11, 1940, 8. B. Henpricks presiding in the absence of President Hann.
The Society was addressed by two members of the National Institute of
Health:

_ SANFORD M. RosENnTHAL: Hxperimental Studies upon Sulphanilamide and
Related Compounds.

Huco Baurr: Chemical Structure of Some Arsenic and Phosphorous Com-

pounds Active against Bacterial Infections.

523D MEETING

The 523d meeting was held in the auditorium of Engineering Building,
University of Maryland, College Park, Md., President Hann presiding.
H.S. Ispeiu and C. E. Waits were elected to the Board of Managers to fill
vacancies arising from the resignations of J. T. SCANLAN and 8. T. Scuick-
TANZ. After the completion of other routine business, the following com-
munications were presented in three sections:

538 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

Physical Chemistry, M. M. Hartna presiding

J. STEINHARDT and M. Harris: Role of Anions and of Undissociated Acids
in the Combination of Acids with Proteins.

D. V. SickmMaAN and W. P. Raturorp: Induced Chains and the Mechanism
of the Thermal Decomposition of Acetaldehyde.

K. R. Smirx: Limitations of the Polarographic Method of Analysis.

Organic and Biological Chemistry, N. L. Drake presiding

W. H. Sesrevu and F. 8. Darr: Vitamin Bz, Complex Deficiencies.
J. J. Stusss, L. B. Lockwoonp, E. T. Ros, B. TaBEenKIN and G. EH. Warp:
Production of Ketogluconic Acids from Glucose by Acetic-Acid Bacteria.
- S. me WRENN and C. I. Pops: Reaction of Acetals with Anhydrous Hydrogen
romide.

Analytical and Inorganic Chemistry, W. J. SvinBELY presiding

C. E. Wuite: Fluorescence Analysis of Aluminum and Thorium.
W. D. MocermMan and J. A. ScHERRER: Determination of Tin in Non-
ferrous Alloys by the Distillation-Cupferron Method.
W. O. Roprnson: The Occurrence of Rare Earths in the Leaves of Hickory
and a Few Other Plants.
NorMAN BEKKEDAHL, Secretary

BOTANICAL SOCIETY

3018ST MEETING

The 301st regular meeting was held in the assembly hall of the Cosmos
Club, December 5, 1939, Vice-president CHARLOTTE ELLioTT presiding;
attendance 100.

Notes and reviews.—M. B. Warts called attention to the fact that he had
found a butternut tree (Juglans cinerea) growing on the Coastal Plains,
near Walshs Grove, 1 mile from Priests bridge on the Patuxent River. He
also displayed a puffball, Calastoma cinnabarinum, which he had collected
on his farm growing among sphagnum and ferns and a rare climbing fern,
Lygodium palmatum, which was growing in his apple orchard among dead
golden-rods. It is related to Schizaea.

Program.—Puiuuir BrieRuty: Mosaic of lulies.—Two viruses can be iso-
lated from Easter lilies (Lilzuwm longiflorum Thunb.) affected with necrotic
fleck mosaic: A strain of cucumber virus I and a virus similar to Mc-
Whorter’s latent virus of lily. Tulip viruses such as Mc Whorter’s latent type
fail to infect tobacco, and cucumber-virus strains are not recovered when
necrotic fleck is passed through tulip. Tulips inoculated with cucumber virus
develop leaf and flower symptoms distinct from the usual breaking of tulips.
Easter-lily seedlings show mild symptoms or none following inoculation with
latent virus only, and none with cucumber virus only; under some conditions
they develop fleck when cucumber virus is superimposed on latent virus.
The coarse mottle mosaic of Easter lily yields a tulip virus but no cucumber
virus. Lilium formosanum Stapf expresses symptoms on inoculation with
tulip viruses including that which is latent in Easter lily and other species.
This species is a convenient substitute for tulip in indexing lilies for the
presence of tulip virus. With this species and tobacco used as test plants,
cucumber and tulip viruses have been shown generally distributed both
separately and in double infections in many lily species in gardens.

CuarLes THom: Address of the retiring president, Naming molds, pub- |
lished in full in this JouRNAL, vol. 30, no. 2, February 15, 1940.

AuicE M. ANDERSEN, Recording Secretary

Obituary

FREDERICK EKUGENE FOWLE was born in Arlington, Mass., on November
20, 1869, the son of Frederick E. and Mary Proctor Fowle. He died at
Washington, D. C., on April 23, 1940. A precocious youth, he early undertook
the study of music and languages, becoming proficient in both. His formal
education was completed at the Massachusetts Institute of Technology,
from which he was graduated in 1894.

As a young man he was for several years assistant at the Harvard College
Observatory, but after graduation from the Massachusetts Institute of
Technology he accepted a position at the Smithsonian Institution, where he
spent more than 40 years in the measurement and study of the radiation of
the sun. He was coauthor in Volumes | to 5 of the Annals of the Astrophysical
Observatory, and he devised methods of measuring the total atmospheric
contents of water vapor and of ozone. He made valuable researches on the
influence of these vapors on the income and outgo of radiation through the
atmosphere which will long be referred to. Not less valuable was his assembly
and editorship of the Smithsonian Physical Tables, a reference work that he
made invaluable in these days of rapid discovery of the physics of the atom
and the extension of the universe. Nearly 20,000 copies of this book went
into the hands of research men during his editorship, and received universal
praise for broadness and accuracy.

By his first wife, Susie Gates, of Arlington, Mass., Mr. Fowle had two
children, who survive him. In 1930 he married Juliana Thomasson, of North
Dakota, who also survives him.

FRANCIS RANDALL HaGNeER died suddenly on July 7, 1940, at his home,
1824 Nineteenth Street, NW., Washington, D. C. Dr. Hagner was born in
Washington in 1873, the son of Dr. Charles E. and Mrs. Isabella Davis
Hagner. His family had long been prominent in the affairs of Maryland and
the District of Columbia. He received his preliminary education at the
Emerson Institute. In 1894 he was graduated from the School of Medicine
of the Columbian College, now George Washington University. After gradu-
ation he went to Johns Hopkins Hospital, becoming assistant resident in
surgery. In 1896 he returned to Washington to practice medicine. For a
time he carried on a general pr actice, but after a few years he began to spe-
cialize in urology.

He was appointed professor of genitourinary surgery at the Gates Wash-
ington University School of Medicine in 1905, which position he held until
his retirement in 1939 when he became professor emeritus. Dr. Hagner will
be remembered as a successful teacher and leader in his field. Many of his
students through his influence, have specialized in the field of urology.

Many honors came to him in the field of medicine, and he served as presi-
dent of the American Association of Genitourinary Surgeons, the Clinical
Society of Genitourinary Surgeons, the Clinico-Pathological Society of the
District of Columbia, and the Medical Society of the District of Columbia.
He was a life member of the American College of Surgeons, a fellow of the
American Medical Association, and a member of the American Urological
Association and the Southern Surgical Association.

Dr. Hagner invented numerous devices and made marked advances in
procedures involved in his specialty and wrote many medical articles relative
to it.

Among Dr. Hagner’s many interests one of the outstanding was his pro-
longed and thorough study of the life of John Wilkes Booth and of the cir-
cumstances surrounding the assassination of President Lincoln. He was

> Ps

540 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

deeply interested in the collection of etchings and early Western Americana.
He was a member of the Washington Academy of Sciences, the Biological
Society of Washington, and the Columbia Historical Society.

He was possessed of a charming personality which won him hosts of friends
and admirers both in Washington and wherever he was known. He was one of
the most beloved as well as one of the most distinguished physicians of our
time.

Dr. Hagner married Miss Elizabeth Allemong of Charleston, W. Va., in
1898. She and a sister, Mrs. Norman James of Baltimore, survive him.

INDEX TO VOLUME 30

An asterisk (*) denotes the abstract of a paper presented before the Academy or an affiliated society.

PROCEEDINGS OF THE ACADEMY AND AFFILIATED SOCIETIES

Anthropological Society of Washington.
318, 538.
535.

Botanical Society of Washington.
Chemical Society of Washington.
Geological Society of Washington.

Philosophical Society of Washington.
46, 134, 174, 276, 317, 448, 484, 533.

Washington Academy of Sciences.

317.

485.
398.

AUTHOR INDEX

ALLARD, H. A. The starling’s family life
and behaviors. 34.

and H. F. Auuarp. The wild
bean Phaseolus polystachyus (L.)
B. 8. P.: Its chromosome number.
335.

ALLARD, H. F. See H. A. AtuarpD. 335.

AVERY, JOHN LAWRENCE. Studies on
the egg-laying habits of the fairy
shrimp. 31.

AXELROD, DanieEL I. The Pliocene Es-
meralda flora of west-central Nevada.
163.

BAILEY, REEVE M. JHadropterus palm-
aris, a new darter from the Alabama
River System. 6524.

BartH, Tom F. W. *Geysers and other
hotsprings in Iceland. 398.

*Thermal activity in Iceland.

497.

Bartram, M. Tuomas. *An early Amer-
ican botanist, John Bartram 1699-
Wie 454:

Beis, K. H. *Model testing at the Na-
tional Hydraulic Laboratory. 399.

BERKNER, L. V. *Fluctuations in the
earth’s outer atmosphere during a
great magnetic storm. 400.

BERRY, Epwarp W. A Cusparia from
the Pliocene of trans-Andean Bolivia.
464.

Additions
florasnela2:

BLAISDELL, Dorotuy. *Variability stud-
ies in the tomato Fusarium wilt
organism; cultural studies. 452.

BuakeE, 8. F. *Early American bota-
nists, John Bannister (1650-1692),
John Clayton (1686-1773), Hum-
phrey Marshall (1722-1801). 454.

to the Pensauken

New species and new names

among Arizona Asteraceae. 467.

Paralbula, a new fossil fish based
on dental plates from the Eocene and
Miocene of Maryland. 205.

Brackett, F. §. *Spectroscopic meth-
ods and their importance in biologi-
cal research. 404.

BrRIERLY, Pointuie. *Mosaic of lilies. 538.
BrRIiESE, R. R. See James F. Covucu.
413. See E. A. Moran. 237.
Brown, D. E. See W. M. Lunn. 319.
Brown, Rotanp W. A bracket fungus

from the late Tertiary of southwest-
ern Idaho. 422.
Fossil pearls from the Colorado
group of western Kansas. 365.
New species and changes of
name in some American fossil floras.
344.

BuuuserR, A. H. ReGinaLp. *The sexual
process in the rust fungi. 454.
CARLGREN, OSKAR. Actiniaria from
Alaska and Arctic waters. 21.

CasH, Epita K. A _ second note on
Georgia Discomycetes. 299.
See M. L. Lonman. 255.

CHAMBLISS, CHARLES E. The botany
and history of Zizania aquatica L.
(‘wild rice’). 185.

CHAPIN, Epwarp A. New genera and
species of lady-beetles related to
Serangium Blackburn (Coleoptera:
Coccinellidae). 263.

CHARLES, VERA K. A ringworm disease
of muskrats transferable to man.
338.

CLausEN, Curtis P. The oviposition
habits of the Eucharidae (Hymen-
optera). 504.

541

——————

542 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

Coker, R. E., and Juanira More@an. A
new harpacticoid copepod from North
Carolina, 395.

Cott, P. A. *Determination of cell
structure by means of the ultra-
violet microscope. 405.

Coutuins, W. D. *Water analyses. 496.

Coox, O. F. Aublet the botanist, a
pioneer against slavery, with a
memorial genus of palms. 294.

Coucn, James F., and Reinyotp R.
BRIESE. Cyanogenesis and enzyme
activity in sorghum varieties. 413.

See E. A. Morgan. 287.

CrEASER, Epwin P. A new species of
phyllopod crustacean from Stone
Mountain, Georgia. 435.

Dorr, Eruine. *The stratigraphic sig-
nificance of the flora of the type
Lance formation. 486.

DRECHSLER, CHARLES. Three fungi de-
structive to free-living terricolous
nematodes. 240.

pu Titty, ARTHENE. *Botanical ex-
ploration in the Canadian Arctics.452.

Dyson, James L. *Structure and mo-
tion of cirque glaciers. 489.

Everitt, Epwarp L., and M. X. SuL-
LIVAN. The bacteriostatic and the
bactericidal action of certain organic
sulphur compounds. 457.

——— The fungistatic and fungicidal
action of certain organic sulphur com-
pounds. 125.

FENNELL, J. L. Two new North Amer-
ican species of Vitis. 15.

FERGUSON, FREDERICK F. See Way-
LAND J. HAyYEs, Jr. 27.

Foster, Marcaret D. *Geochemical
relations of ground waters in the
Houston-Galveston area, Tex. 490.

Fox, Frances E. *Supersonics in liq-
uids. 402.

Fox, Irvine. Siphonaptera from West-
ern United States. 272.

GABRIELSON, IRA N. *Western Ameri-
can alpine plants. 453.

GaGER, C. Stuart. *How botany ad-
vances. 455.

Gamow, G. *Uncertainties in atomic
and common life. 403.

GaRDNER, LouisS. *Displacement along
the Hurricane fault in Utah and
Arizona. 492.

GARNER, W.W. See W.M. Lunn. 319.

GoopDING, CHARLOTTE O. Two new
species of Muhlenbergia. 19.

Gravatt, G. F. *Effect of epidemic tree
diseases on the food supplies of ani-
mals. 4651.

Grices, Davin. *Convection currents
and mountain-building. 494.

HAGEN, JoHN P. *Beam power tubes as
ultra-high radio frequency genera-
tors. 411.

HARLAND, S. C. Taxonomic relation-
ships in the genus Gossypium. 426.

HARTMAN, Ote@a. Boccardia proboscidea,
a new species of spionid worm from
California. 382.

Hayes, WayLanp J., Jr., and FREDERICK
F. Fereuson. Notes on the mor-
phology of Macrostomum ruebushi
var. schmittt. 27.

HeEepGpeETH, JoEL W. A new pycnogonid
from Pescadero, Calif., and distribu-
tional notes on other species. 84.

Hermann, F. J., T. H. Kearney, and
R. H. Prresurs. Arizona plants:
New species, varieties, and combina-
tions. 217.

Hewett, D. F. New formation names
to be used in the Kingston Range,
Ivanpah quadrangle, California.
239.

HorErNErR, G. R. A nomenclatorial note
on Pseudoperonospora. i133.

Huspparp, J. C. *Ultrasonics and the
ratio of specific heats of gases. 401.

Huut, Frank M. Some new species of
Syrphidae (Diptera). 432.

IncrAaM, WitLiAM M. A new Gisoriza.
376.

JENNEss, D1amMoNpD. Prehistoric culture
waves from Asia to America. 1.
JoHNSON, Exuis A. *Scattering of light
from a modulated searchlight beam

at high altitudes. 401.

Kearney, T. H. See F. J. HERMANN.
217.

Kene, Y. L. Oxytenanthera felix, a new
species of bamboo from Yunnan,
China. 425.

Dec. 15, 1940

Kent, GEorRGE C., Jr. A new trematode
from Siren lacertina: Diplostomulum
strenis, n. sp. 87.

Kirk, Epwin. Seven new genera of
Carboniferous Crinoidea Inadunata.

aA
Ler, F. W. *Some problems in geo-
physics. 410.

LronarpD, E. C. Two new genera of
Acanthaceae from Guatemala. 4501.

_Louman, M. L. Atropellis species from
pine cankers in the United States.
255.

LowELL, W. P., Jr. *Practical applica-
tion of fluorescent lighting. 409.
Lunn, W. M., D. E. Brown, J. E. Mc-

Mourtrey, and W. W. GARNER.
*Tobacco and weeds. 319.
Lyon, T. C. *Methods of air navigation
and aeronautical charts. 406.
Martin, G. W. Spore formation in
Matula. 378.
Martin, J. H.
bushels of seed grain.

*Cleaning 20 million
320.

Marton, L. *Electron microscopy. 409.

McMortrey, J. E. See W. M. LunNN.
319.

MecNisu, A. G. *The geomagnetic field

and its variations. 407.
MertIis£, J. B., Jr. Placer gold in Alaska.
93.

*Stratigraphic measurements in
parallel folds. 492.

Moore, J. Percy. Austrobdella ano-
culata, a new species of fish leech
from Greenland. 520.

Moran, E. A., R. R. Brigsg, and J. F.

CoucnH. Some new cyanogenetic
plants. 237.

Morean, H. R. *The motions of the
earth. 406.

MorRGAN, JUANITA. See R. E. Coxer.
395.

Morata, K. J. *Volcanic ash as a
source of silica for the silicification of
wood. 496.

Napal, A. *Formation of surfaces of slip

under plastic states of stress. 399.

NoRGORDEN, Oscar. *Propagation char-
acteristics of ultra-high radio fre-
quencies. 411.

INDEX

-PHILLIPS,

543

Nuttine, P. G. A study of the associa-
tion of magnesia with silica in a pure

magnesium clay. 233.
PEEBLES, Ropert H., and Lovis C.
WHEELER. Arizona plants: A new

variety and new names and combina-

tions. 473.

See F. J. HERMANN. 217.

Prunp, A. H. *A partial analysis of
gaseous mixtures without the use of
a spectroscope. 407.

Max. Anselme Payen, dis-
tinguished French chemist and pio-
neer investigator of the chemistry of
lignin. 65.

REHDER, Harautp A. A new urocoptid
mollusk from Mexico. 315.

ROTHSCHILD, MirtaM. Cercaria pricei, a
new trematode, with remarks on the
specific characters of the ‘‘Prima”’
group of Xiphidiocercariae. 437.

SaunpDERS, H. E. *The David W. Tay-
lor Model Basin at Carderock, Md.
399.

SayLtor, LAwRENcE W. Ten new West
Indian scarab beetles of the genus
Phyllophaga, with two new names.
305.

SHEPARD, F. P. *Nondepositional en-
vironments off the California coast.
486.

SHOEMAKER, CLARENCE R. Notes on the
amphipod Gammarus minor Say and
description of a new variety, Gam-

marus minus var. tenuipes. 388.
Simpson, GEorGE GayLorp. Mammals

and land bridges. 137.

Smitu, HueuH M. The remora Phtheir-
ichthys lineatus and the first speci-
men from United States waters. 531.

Smitu, P. A. *Exploring the continental
shelves and slopes. 405.
STEPHENSON, E. L. *The results of

magnetometer surveys on laccoliths
in the Highwood Mountains, Mont.
488.

Stewart, T. D. The finding of an In-
dian ossuary on the York River in

Virginia. 356.
Suuttivan, M. xX. See Epwarp L.
EveRItT. 125, 457.

544 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

SWALLEN, JASON R. Eight new species
Oi IOC, Tale

Miscellaneous new American
grasses. 209.
Swartz, J. H. *Resistivity studies of
some geological problems. 410.
SWINGLE, WALTER T. New varieties and
new combinations in the genera
Clausena, Oxanthera, and Triphasia
of the orange subfamily, Auranti-
oideae. 79.

TuHom, Cuartes. Naming molds. 49.

THORNE, GERALD. The hairworm, Gor-
dius robustus Leidy, as a parasite of
the Mormon cricket, Anabrus simplex
Haldeman. 219.

Urry, Haroup C. The separation of
isotopes by chemical means. 277.

Vuay, J. S. *Structural features on the
north side of the Beartooth Moun-
tains near Nye, Mont. 490.

Wauton, A. C. The nematode genus
Raillietnema Travassos, 1927. 474.

Watts, C. B. *A traveling-wire microm-
eter with photographic registration.
407.

WELIMAN, F. L. *Variability studies in
the tomato Fusarium wilt organism;
pathogenicity studies. 453.

WELLS, JOHN W. Two new corals from
the Avenal formation (Eocene) of
California. 374.

WENZEL, Rurert L. A genus of histerid
beetles new to the United States. 516.

WHEELER, Lovis C. See Ropert H.
PEEBLES. 473..

WHEELER, Nancy H. The North Amer-
ican empoascan leafhoppers of the
alboneura group (Homoptera-Cicadel-
lidae). 478.

ZABEL, R. M. *Fundamentals of fluor-
escence. 408.

SUBJECT INDEX

Aeronautics. *Methods of air navigation
and aeronautical charts. T. C.
Lyon. 406.

Agriculture. *Cleaning 20 million bushels
of seed grain. J. H. Martin.
320.

*Tobacco and weeds. W. M. Lunn,
D. E. Brown, J. E. McMurtrey,
and W. W. GARNER. 319.

Anthropology. Prehistoric culture waves
from Asia to America. DIAMOND
JENNESS. 1.

The finding of an Indian ossuary on
the York River in Virginia. T. D.
STEWART. 356.

Biochemistry. The bacteriostatic and the
bactericidal action of certain or-
ganic sulphur compounds. Ep-
wARD L. Everitt and M. X.
SuLnIvan. 457.

The fungistatic and fungicidal action
of certain organic sulphur com-
pounds. Epwarp L. Everitt and
M. X. SULLIVAN. 125.

Biophysics. *Determination of cell struc-
ture by means of the ultraviolet
microscope. P. A. Cour. 405.

*Spectroscopic methods and their im-
portance in biological research.
F.S. Brackett. 404.

Botany. *An early American botanist,
John Bartram 1699-1777. M.
Tuomas Bartram. 454.

Arizona plants: A new variety and
new names and combinations.
Rosert H. PEEBLES and Louis C.
WHEELER: 473.

Arizona plants: New species, vari-
eties, and combinations. F. J.
HERMANN, T. H. Kearney, and
lai, JEL, 1Piobousienis) “HAIL (

A second note on Georgia Discomy-
cetes. EpitH K. Casu. 299.

Aitropellis species from pine cankers
in the United States. M. L.
Lowman and Epira K. Casu.
255.

Aublet the botanist, a pioneer against
slavery, with a memorial genus of
palms. O. F. Coox. 294.

545

*Botanical exploration in the Ca-
nadian Arctics. ARTHENE DU
Titty. 452.

*Harly American botanists, John
Bannister (1650-1692), John Clay-
ton (1686-1773), Humphrey Mar-
shall (1722-1801). S. F. Buaxg.
454.

Hight new species of Pariana. JASON
R. SWALLEN. 71;

*How botany advances. C. STUART
GaGcER. 455.

Miscellaneous new American grasses.
Jason R. SWALLEN. 209.

*Mosaic of lilies. PHILuirp BRIERLY.
538.

Naming molds. CHartEs THOM.
49,

New species and new names among
Arizona Asteraceae. S. F. BLAKE.
467.

New varieties and new combinations
in the genera Clausena, Oxanthera,
and Triphasia of the orange sub-
family, Aurantioideae. WALTER
T. SWINGLE. 79.

Nomenclatorial note on Pseudoperon-
ospora. G. R. HopRNER. 133.
Oxytenanthera felix, a new species of
bamboo from Yunnan, China.

Y. L. Kena. 425.

Spore formation in Matula. G. W.
Martin. 378.

Taxonomic relationships in the genus
Gossypium. S.C. HARLAND. 426.

The botany and history of Zizania
aquatica L. (“wild rice’).
CHARLES HE. CHAMBLIss. 185.

*The sexual process in the rust fungi.
A. H. REGINALD BULLER. 4654.

The wild bean Phaseolus polystachyus
(L.) B.S. P.: Its chromosome num-
bers) Hs VAe ATTARD (and: HE.
ALLARD. 335.

Three fungi destructive to free-living
terricolous nematodes. CHARLES
DRECHSLER. 240.

Two new genera of Acanthaceae from
Guatemala. E. C. Lnonarp. 501.

546 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

Two new North American species of
Vitis. J. L. FENNELL. 15.

Two new species of Muhlenbergia.
CHARLOTTE O. GooppiNnG. 19.
*Variability studies in the tomato
Fusarium wilt organism; cultural
studies. DorotTuHy BLAISDELL.

452.

*Variability studies in the tomato
Fusarium wilt organism; patho-
genicity studies. F. L. WELLMAN.
453.

*Western American alpine plants.
Ira N. GABRIELSON. 453.

Chemistry. Anselme Payen, distinguished
French chemist and pioneer inves-
tigator of the chemistry of lignin.
Max PHILuirs. 65.

A study of the association of mag-
nesia with silica in a pure mag-
nesium clay. P. G. NuvuttTINa.
233.

Cyanogenesis and enzyme activity
in sorghum varieties. JAMES F.
Coucu and REINHOLD R. BRIESE.
413.

Some new cyanogenetic plants.
E. A. Moran, R. R. Briese, and
J. F. Coucnw. 287.

The separation of isotopes by chem-
ical means. Haroutp C. UREY.
7 Te

Ecology. *Effect of epidemic tree dis-
eases on the food supplies of wild-
life. G. F. Gravatt. 4651.

Entomology. A genus of histerid beetles
new to the United States. Rv-
PERT L. WENZEL. 516.

New genera and species of lady-
beetles related to Serangium Black-
burn (Coleoptera: Coccinellidae).
EpWwarpD A. CHAPIN. 2683.

Siphonaptera from Western United
States. Irvine Fox. 272.

Some new species of Syrphidae
(Diptera). Frank M. HULL.
432.

Ten new West Indian scarab beetles
of the genus Phyllophaga, with two
new names. LAWRENCE W. Say-
LoR. 305.

The North American empoascan
leafhoppers of the alboneura group

(Homoptera-Cicadellidae). Nancy
H. WHEELER. 478.

The oviposition habits of the Eu-
charidae (Hymenoptera). CuRTIS
P. CLAUSEN. 504.

Geochemistry. *Geochemical relations of
ground waters in the Houston-
Galveston area, Tex. MARGARET
D. Foster. 490.

*Volcanic ash as a source of silica for
the silicification of wood. K. J.
Murata. 496.

*Water analyses. W. D. Co.uins.
496.

Geology. *Convection currents and
mountain-building. Davip Griaes.
494,

*Displacement along the Hurricane
fault in Utah and Arizona. Lovis
S. GARDNER. 492.

New formation names to be used in
the Kingston Range, Ivanpah
quadrangle, California. D. F.
Hewett. 239.

*Nondepositional environments off
the California coast. F. P. SHEp-
ARD. 486.

Placer gold in Alaska. J. B. MrEr-
TIE, JR. 93:

*Stratigraphic measurements in par-
allel folds. J. B. Mertiz, Jr.
492.

*Structure and motion of cirque
glaciers. James L. Dyson. 489.

*Structural features on the north side
of the Beartooth Mountains near
Nye, Mont. J. 8S. Vuay. 490.

*The results of magnetometer sur-
veys on laccoliths in the Highwood
Mountains, Mont. E. L. SrEPHEN-
SON. 488.

Geophysics. *Exploring the continental
shelves and slopes. P. A. SMITH.
405.

*Geysers and other hotsprings in
Iceland. T. F. W. Bartu. 398.

*Resistivity studies of some geologi-
cal problems. J. H. Swartz.
410.

*Some problems in geophysics.
F. W. Les. 410.

*The geomagnetic field and its varia-
tions. A. G. McNisxH. 407.

Dec. 15, 1940

*The motions of the earth. H. R.

MorGan. 406.
*Thermal activity in Iceland. Tom
F. W. BartH. 497.
Ichthyology. Hadropterus palmaris, a new

darter from the Alabama River
System. RerveE M. _ BalILey.
524.
The remora Phtheirichthys lineatus

and the first specimen from United
States waters. Hueco M. Smiru.
531.

Malacology. A new urocoptid mollusk
from Mexico. Haraup A. REu-
DER. 315.

Mycology. A ringworm disease of musk-
rats transferable to man. VERA
K. CHARLES. 338.

Obituary. ADLER, Cyrus. 412.
CusHING, HARVEY WILLIAMS.
FowLse, FREDERICK EUGENE.
GROOVER, THOMAS ALLEN.
HAGNER, FRANCIS RANDALL.
LINDGREN, WALDEMAR. 92.
Matson, GEORGE CHARLTON.
Mortiser, Davip Myers. 499.
TRYON, FREDERICK GALE. 232.

Ornithology. The starling’s family life
and behaviors. H. A. ALLARD.
34.

Paleobotany. A Cusparia from the Plio-
cene of trans-Andean Bolivia.
Epwarp W. Berry. 464.

Additions to the Pensauken flora.
EpWARD W. Berry. 132.

A bracket fungus from the late Terti-
ary of southwestern Idaho. Ro-
LAND W. Brown. 422.

New species and changes of name
in some American fossil floras.
Rotanp W. Brown. 344.

The Pliocene Esmeralda flora of
west-central Nevada. Danie. I.
AXELROD. 163.

*The stratigraphic significance of the
flora of the type Lance formation.

136.
539.
455.
539.

184.

ERLING Dorr. 486.
Paleontology. A new Gisortia. WiILLIAM

M. INcram. 376.

Fossil pearls from the Colorado

group of western Kansas. Ro-

LAND W. Brown. 365.

INDEX

547
Mammals and = land bridges.
GEORGE GAYLORD SIMPSON. 137

Paralbula, a new fossil fish based on
dental plates from the Eocene and
Miocene of Maryland. S. F.
BLAKE. 205.

Seven new genera of Carboniferous
Crinoidea Inadunata. Epwin
KoRKew ozs

Two new corals from the Avenal
formation (Eocene) of California.
JOHN W. WEuts. 374.

Physical chemistry. *A partial analysis of
gaseous mixtures without the use

of a spectroscope. A. H. Prunp.
407.
Physics. *A traveling-wire micrometer

with photographic
C. B. Watts. 407.
*Beam power tubes as_ ultra-high

registration.

radio frequency generators. JOHN
P. Hagen. 411.

*Electron microscopy. L. Marton.
409.

*Fluctuations of the earth’s outer
atmosphere during a great mag-
netic storm. L. V. BERKNER.
400.

*Formation of surfaces of slip under

plastic states of stress. A. NADAt.
399.

*Fundamentals of fluorescence.
R. M. Zaseu. 408.

*Model testing at the National Hy-
draulic Laboratory. K. H. Bets.
399.

*Practical application of fluorescent
lighting, W. P. Lowe, Jr.
409.

*Propagation characteristics of ultra-
high radio frequencies. Oscar
NoRGORDEN. 411.

*Scattering of light from a modulated
searchlight beam at high altitudes.
Exuis A. JOHNSON. 401.

*Supersonics in liquids. Francis E.
Fox. 402.

*The David W. Taylor Model Basin
at Carderock, Md. H. E. Saun-
DERS. 399.

*Ultrasonics and the ratio of specific
heat of gases. J. C. HusBarp.
401.

—“-—~-

548 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES VOL. 30, No. 12

*Uncertainties in atomic and com-

mon life. G. Gamow. 408.
Scientific notes and news. 135, 232.
Zoology. Actiniaria from Alaska and

Arctic waters. OSKAR CARLGREN.

Dale

A new harpacticoid copepod from
North Carolina. R. KE. Coxerr
and JuANITA Moraan. 395.

A new pycnogonid from Pescadero,
Calif., and distributional notes on
other species. JozrL W. HeEpc-
PETH. 84.

Austrobdella anoculata, a new species
of fish leech from Greenland. J.
Percy Moore. 520.

A new species of phyllopod crusta-
cean from Stone Mountain, Geor-
gia. Epwin P. Creaser. 435.

A new trematode from Szren lacer-
tina: Diplostomulum sirenis, n. sp.
GEoRGE C. KENT, JR. 87.

Boccardia proboscidea, a new species
of spionid worm from California.
Ouca HARTMAN. 382.

Cercaria pricet, a new trematode,
with remarks on the specific char-
acters of the “Prima” group of
Xiphidiocercariae. Miriam Rotus-
CHILD. 437.

Notes on the amphipod Gammarus
minus Say and description of a new
variety, Gammarus minus vat.
tenuipes. CLARENCE R. SHOE-
MAKER. 388.

Notes on the morphology of Macro-
stomum ruebusht var. schmitt.
WayYLANp J. Hayes, JrR., and
FREDERICK F. FErRGuson. 27.

Studies on the egg-laying habits of
the fairy shrimp. JoHn Law-
RENCE AVERY. 31.

The hairworm, Gordius robustus
Leidy, as a parasite of the Mor-
mon cricket, Anabrus~ simplex
Haldeman. GERALD THORNE. 219.

The nematode genus Razllietnema
Travassos, 1927. A. C. Wauton.
ATA.

CONTENTS

Borany.—T wo new genera of Acanthaceae from Guatemala. E. C. . Bi AN
LHONARD i iG ce i oe

Enromo.Logy.—The oviposition habits of the Bucharidae (Hymenop- ee
tera). Curtis P. CLAUSEN. er ee SLi ae a i eee ine) DOL.

EnromoLogy.—A genus of histerid beetles new to the United States i 3 é : ss
Rurert b. WENZD. 02. ee ie Oe ee ee rie

ZOOLOGY. hue once anoculata, a new species of fish oo from

Greenland, J. Peucy. MOORE... 0.0.0.4 cane 2 eA 520 ie
IcutHYoLoGy.—Hadropierus palmeris, a new darter from the Alabama :

River System. Reeve M. Batney............. ee sa. 524
IcutHyoLocy.—The remora Phtheirichthys lineatus and the first Peers ge

men from United States waters. Hucu M. SmirH............ 631
PROCEEDINGS: THE Acapaur 2 cee 21 ay oe
PROCEEDINGS: CHEMICAL SOCIETY........0+-..-005- eee oo Obs
PROCEEDINGS: BOTANICAL SOCIETY......... oii revs «Coenen a:

Ozrruary: Frederick Eugene Fowle, Francis Randall Hagner...... 539

INDEX TO VOLUME 300.0... 052. DES. i che os Or ee
This Journal is Indexed in the International Index to Periodicals

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