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JOURNAL

WASHINGTON ACADEMY

OF SCIENCES

VOLUME 44, 1954

BOARD OF EDITORS

JOHN C. EwERSs R. K. Coox FENNER A. CHACE, JR.

U.S. NATIONAL MUSEUM NATIONAL BUREAU U.S. NATIONAL MUSEUM

OF STANDARDS

ASSOCIATE EDITORS

J. I. HOFFMAN BERNICE SCHUBERT

CHEMISTRY BOTANY

DEAN B. CowleE Puitie DRUCKER

PHYSICS ANTHROPOLOGY

ALAN STONE y Davin H. DuNKLE

ENTOMOLOGY GEOLOGY

PUBLISHED MONTHLY

BY THE

WASHINGTON ACADEMY OF SCIENCES

Mount Royat & GUILFORD AVES.

BALTIMORE, MARYLAND

gL OE TEEN EY

LES eae Up. =

AN

=f

a

Cy

ACTUAL DATES OF PUBLICATION, VOLUME 44

No. 1, pp. 1-82, January 20, 1954

No. 2, pp. 33-64, February 22, 1954

No. 3, pp. 65-96, March 24, 1954

No. 4, pp. 97-1382, May 3, 1954

No. 5, pp. 1383-164, May 27, 1954

No. 6, pp. 165-200, June 16, 1954

No. 7, pp. 201-2382, July 28, 1954

0. 8, pp. 233-264, August 13, 1954

0. 9, pp. 265-296, September 28, 1954

o. 10, pp. 297-832, October 22, 1954

o. 11, pp: 333-3876, November 22, 1954

0. 12, pp. 377-408, December 22, 1954

A De

Vou. 44 JANUARY 1954 No. 1

\ FEBI1 1954 )

JOURNAL i LinnaRy pf

a rad

ee ee

OF THE

WASHINGTON ACADEMY

OF SCIENCES

BOARD OF EDITORS

J. P. E. Morrison JoHN C. EwrErs R. K. Coox

U.S. NATIONAL MUSEUM U.8. NATIONAL MUSEUM NATIONAL BUREAU

OF STANDARDS

ASSOCIATE EDITORS

F. A. Cuacsz, JR. EvBert L. LITT es, JR.

ZOOLOGY BOTANY

J. I. HorrMANn Puitie DRUCKER

CHEMISTRY ANTHROPOLOGY

Dean B. CowiE Davip H. DuUNKLE

PHYSICS : @EOLOGY

ALAN STONE

ENTOMOLOGY

PUBLISHED MONTHLY

BY THE

WASHINGTON ACADEMY OF SCIENCES |

Mount Rorau & GurILForD AVES.

BALTIMORE, MARYLAND

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Acceptance for mailing at a special rate of postage provided for in the Act of February 28, 1925

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ces aay AS

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2A Sat

JOURNAL

OF THE

WASHINGTON ACADEMY OF SCIENCES

Vou. 44

January 1954

No. 1

PHYSICS.—On research and education—in fluid dynamics.! RAYMOND J. SEEGER,

National Science Foundation.

“Many of the physical sciences seem to

have abandoned the laboratory for stu-

dents.”’ This observation was made by

G. E. Erikson of Harvard University in

I. B. Cohen’s and F. G. Watson’s General

education in science (1952). Another in-

teresting remark is one by 8. J. French of

Colgate University that ‘“‘general education

is only an improved version of liberal edu-

cation.’”’ Both these statements take us back

to the Academy which Plato conducted for

41 years in the Athenian grove honoring

the Greek hero Akademos. The disciples

who gathered about him there outside the

city were eager to learn, not special informa-

tion peculiar to observed phenomena, but

rather general principles underlying uni-

versal knowledge.” Two academic emphases

have had an influence upon education long

after the closing of the Academy by Jus-

tinian nine centuries later. Inasmuch as

Plato was not interested in the imperfect

material triangles that one observes, wooden

triangles, metal triangles, etc., but rather

in the idea of a perfect triangle, his followers

were intent primarily upon contemplating

this thoughtful world. The Greeks had a

word for such an attitude, viz., theory

(from the same root as the Greek word for

theatre—meaning, to view). Aristotle later

contrasted this point of view with the

practical (meaning, to do) business of doing.

Consequently, there has been a persistent

trend in traditional humanistic education

to keep it divorced from progressive scien-

tific research about things.

1 After-dinner address, Fluid Dynamics Divi-

sion, American Physical Society, July 2, 1953,

Pennsylvania State College.

2 Sarton, G. History of science. Cambridge,

1952.

On the other hand, the enterprise of scien-

tific research was initially disassociated from

education. The classic document along this

line was Francis Bacon’s New Allantis,

published incomplete in the year of his

death, 1626.4 You may recall that in his

Timaeus Plato mentions a lost island of

Atlantis somewhere west of Gibraltar. Re-

garding the discovery of America as vir-

tually that of Atlantis, Bacon visualizes a

new Atlantis some place in the Pacific west

of America. In this book, he describes an

ideal research institute, Salomon’s House,

which had the following objective: ‘The

end of our foundation is the knowledge of

causes and secret motions of things.’’ His

proposal in many respects is quite novel,

even modern.

Salomon’s House contained specialized

facilities for various types of investigations.

There were deep caves for exploring phe-

nomena beneath the earth, as well as high

towers on mountains for observing meteoro-

logical phenomena. There was a _ special

laboratory for high-temperature investiga-

tions, one for optics, one for acoustics. A

special room was provided for the artificial

production of rain. One room contained

primarily engines, including some for ord-

nance. Finally, there was a mathematical

room filled with appropriate instruments.

It was recognized that the proper utilization

of these facilities would requisition the entire

time of the workers.

Novices and apprentices were to be differ-

entiated from what we- nowadays all

' professional personnel, 36 fellows in all,

grouped as follows: 12 merchants of light

who roamed about the earth in search of

3 Famous new deals of history. New York, 1935.

$6m% 5 @ ines

2, JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

observed data; 3 deprepators who sought

such data in books; 3 individuals who col-

lected data from current experiments; 3

pioneers who collected data from new experi-

ments, and 3 compilers who classified and

tabulated all these data. In addition, there

were 3 benefactors whose primary function

was to determine how to use all the results;

3 planners of new experiments, 3 others who

performed these experiments and reported

on them, and finally 3 who interpreted all

the discoveries thus made.

It is to Bacon’s credit that he sensed the

importance of a cooperative enterprise of

specialists having available unusual equip-

ment and books. Although Bacon himself

was not a scientist and failed even to recog-

nize the outstanding scientists of his day,

such as Galileo, Gilbert, and Harvey, never-

theless, he successfully sowed and propa-

gated the concept of a research organization.

The New Atlantis went through ten editions

before 1670. Its influence may be seen in

that the frontispiece of the first (1667)

History of the Royal Society by Thomas

Sprat has as a central figure the bust of

Charles II, on one side the President of the

Royal Society, and on the other side, Francis

Bacon himselt.*

Another description of a utopian learned

society, this time with satirical intent, ap-

peared exactly 100 years later (1726) in

Jonathan Swift’s Gulliver’s Travels. In A

Voyage to Laputa (Part 3), attention is

called to the flying island Balnibarbi with

its Academy of Projectors in Lagado, the

metropolis. The academy had more than

500 rooms, each with more than one pro-

jector (in charge of a project). A comment

by the guide, a former projector, is perti-

nent: ‘“The only inconvenience is that none

of these projects are yet brought to perfec-

tion and in the meantime the whole country

lies miserably waste, the houses in ruins,

and the people without food and clothes.”

To cite a celebrated instance, for eight years

one man had been ‘‘extracting sunbeams out

of cucumbers, which were to be put into

vials hermetically sealed and let out to

warm the air in raw inclement summers.”’

An old established custom was the begging

4 Stimson, D. Scientists and amateurs—a history

of the Royal Society. New York, 1949.

vou. 44,.No. 1

by the projectors for funds to carry on their

work. On the other side of the academy,

there were ‘‘projectors in speculative learn-

ing,’’ such as a universal artist who had been

for 30 years employing his thoughts for the

improvement of life.

Meanwhile, the serious development of

scientific interests began to necessitate

special funds for apparatus. Up to the middle

of the seventeenth century, support for

scientific investigations had been largely

the personal concern of individuals. We

recall that Archimedes had been supported

by Kang Hieron of Syracuse. Galileo later

spent much effort to obtain his final appoint-

ment at the court of the Grand Duke of

Tuscany. He was succeeded in his position

there by Evangelista Torricelli. Occasion-

ally, the church was found supporting un-

usual priests, such as Edmé Marriotte. It is

noteworthy that in all these instances the

individuals apparently had adequate leisure

and freedom to carry on their own in-

vestigations in addition to fulfilling certain

obligations of a more applied or engineering

character. Now and then a person like Rob-

ert Boyle had sufficient wealth in his own

right to carry out his researches. Otto von

Guericke, the mayor of Magdeburg, was

apparently able to use $20,000 of available

funds for his experiment with the famous

hemispheres.

The increased interest in science in the

early 17th century made the acquisition

of funds of paramount importance. In view

of the Pope’s condemnation of Galileo’s

opinions, it 1s not surprising that scientific

research was little supported by the uni-

versities of that day. Academies of science,

therefore, were organized primarily for this

purpose.® An early society was the Acca-

demia dei Lincei in Rome from 1600 to

1630; it included Galileo in its membership.

The more active Accademia del Cimento of

Florence (1657-1667) was the first organ-

ized scientific academy; it was made up

largely of members of universities in the

vicinity. They pursued a careful program of

measurements guided by the motto, ‘‘Pro-

bando e Reprobando,” while cautiously

avoiding any theoretical interpretations in

5 ORNSTEIN, M. The role of scientific societies

in the seventeenth century, ed. 3. Chicago, 1938.

JANUARY 1954

view of Galileo’s indiscretions. In 1662, the

second year of the Stuart Restoration,

Charles II founded the Royal Society in

England, essentially a group of amateurs of

science. The French Académie des Sciences

was established in 1666 by Louis XIV. In-

eluding laboratory work, it was generally

supported and regulated by the government.

Each academician was asked to submit a

program of his own work. Following Bacon,

Christian Huygens organized cooperative

research for compiling and amassing facts.

For example, there were two scientific ex-

peditions: one to Uranienburg and one to

Cayenne, as well as two well-known co-

operative projects, a history of animals and

a history of plants. The Berlin Academy

was founded in 1700 through Gottfried W.

Leibniz by Frederick the Great.

It is significant that even when outstand-

ing scientific research emanated from a

university little stimulation might be given

a man like Isaac Newton by his colleagues

there. Indeed, the Cartesian physics which

Newton himself had disproved was. still

taught at his own University of Cambridge

in 1718 long after he had left. It is note-

worthy also that the Royal Society, whose

charter originally permitted the forming of

a college, definitely rejected such an institu-

tion as not being conducive to research in

view of its primary obligation to teaching.

(Abraham Cowley actually proposed the

support of such a college from the profits

out of resulting scientific inventions.)

The scientific academies did not pursue

their objectives without popular ridicule.

In “The Clouds” Aristophanes had long

ago poked fun at Socrates in his phronis-

terlum (thinking house).° There Socrates is

found suspended in a basket. In reply to

a question as to what he was doing Socrates

said, “I walk on air and contemplate the

sun.”’® Charles II himself is credited with

haying slyly proposed to the members of

the Royal Society at their charter banquet

a question as to why a pail of water weighs -

the same when two live fish are put into it.

After considerable discussion on this sub-

ject by the members he admitted knowing

that such a pail actually weighs more. He

® Coopmr, L. Fifteen Greek plays. Oxford, 1950.

SEEGER: FLUID DYNAMICS 3

is said to have laughed also at the professors

at Gresham College who weighed air. In

1673, the poet laureate, Thomas Shadwell,

culled data from the publications of the

Royal Society and wrote a play called “The

Virtuoso.”’ The leading character was Sir

Nicholas Gimerack, who suspended him-

self on a rope from the ceiling and imitated

the motions of a frantic frog swimming in

a nearby bowl of water. When asked why

he did not learn swimming by immersing

himself in water, Sir Nicholas replied, ‘I

content myself on the speculative part of

swimming....I care not for the practice.”

Later critics were Samuel Butler, Joseph

Addison, and Richard Steele. The private

views of scientists will always be open to

public criticism by non-scientists.

Thus we see that as traditional education

had developed independently of research,

so cooperative research originated inde-

pendently of education. It is not surprising

that these two merged later into research

and education.

The first such program, strangely enough,

was organized by Aristotle.’ After studying

with Plato, he started a school outside

Athens on the side of town opposite the

Academy in a grove sacred to Apollo Ly-

ceius. For 13 years, he was the head of this

peripatetic Lyceum, which combined formal

lectures with organized research, both

personal and collective. Those of us who are

physicists are prone to pass over Aristotle’s

contributions to research because of certain

flagrant errors in his physics which retarded

its growth when scholasticism later popu-

larized Aristotelianism. One of the promi-

nent errors was based on a false interpreta-

tion of observations in fluid dynamics. For

example, looking at matter in motion

Aristotle had perceived that the greater

the force applied to a body at rest the greater

the speed the body acquires and that the

greater the resistance of the medium tra-

versed the less the speed the body develops.

One might say that speed in this instance is

roughly proportional to the ratio of the

initiating force and the resisting force. On

this basis, a vacuum, which has zero resist-

ance, would produce an infinite’ speed.

7 Sarton, G. Op. cit.

4 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

Believing that so great a speed is impossible,

Aristotle rejected the underlying assump-

tion, namely, the concept of a vacuum. The

phrase, ‘‘Nature abhors a vacuum,”’ became

proverbial. Galileo, on the other hand, re-

viving this rejected idea of a vacuum, con-

cluded correctly that speed in this case is

proportional to time (Aristotle had _ erro-

neously decided that speed is proportional

to distance). Aristotle, however, was suc-

cessful in other observations of phenomena

such as those associated with meteorology.

Isolated cases of combined research and

education are found in the 17th and 18th

centuries. For example, Galileo achieved

his most outstanding scientific successes

while he was at Venice’s University of

Padua, prior to 1610, when he moved to

Florence. It was in the earlier period that he

conducted fundamental investigations on

his two new sciences (motion, including that

of projectiles, and the strength of materials,

including the power of a vacuum). It was

in these years, too, that he invented a

thermoscope, as well as the telescope which

he used in discovering the moons of Jupiter

early in 1610. Daniel Bernoulli participated

in the St. Petersburg Academy for 8 years,

but he published his famous book on ‘“‘Hy-

drodynamica”’’ during his later tenure as

professor of physics at the University of

Basle. An interesting development was the

Collegium Curiosum sive Experimentale at

Altdorf University. Christopher Sturm, the

professor of physics and mathematics there,

set up a laboratory with instruments for

teaching his university students in his home.

He is said to have trained many skilled

experimenters. Two volumes of experiments

were published and subsequently served as

a textbook of experimental physics.

The culmination of these isolated in-

stances and the initiation of a new era oc-

curred in the establishment of the Hcole

Polytechnique in 1797, a byproduct of the

French Revolution. The first professor in

Mathematics was J. L. Lagrange who had

spent most of his life engaged in research

at the Berlin Academy although earlier in

his life he had attempted to establish a

research institute in connection with his

teaching at the Turin Royal Artillery School.

It is worth noting that Lagrange had been

vot. 44, No. 1

recommended by Laplace who had once

examined a pupil by the name of Napoleon

in the Ecole Militaire. The mid-nineteenth

century became replete with university

posts filled by men who combined research

and education interests. We naturally think

of pioneers in fluid dynamics such as G. G.

Stokes at Cambridge, W. Thomson, Lord

Kelvin, at Glasgow, H. von Helmholtz at

Bonn and Heidelberg, and L. Prandtl at

Gottingen. Nowadays such activity is not

unusual.

In conclusion, I should like to say a few

words about some of the motivating princi-

ples underlying the National Science Foun-

dation’s program in fluid dynamics. It is

rather significant that the 1950 Act. which

established the Foundation stresses ‘‘the

promotion of basic research and education

in the sciences.’’ Not education per se, not

research per se, but rather research and

education. Some of us have begun to feel

that the most important word here is the

connective, ‘‘and’’.

We find our minds meditating on the

concept of the great scientist-scholar. You

know, and I know, individuals who have

been great teachers despite their lack of

competence in research. We have known

also great research people who have been

impossible as teachers. The average teacher,

however, would undoubtedly be a better

teacher because of some research and the

average researcher would probably be better

in his research because of some teaching.

In any event, the great scientist-scholar

must be motivated by superior intellectual

curiosity whether he engages primarily in

research at a university or engages mostly

in teaching at a college. I am reminded of

a statement made by Prof. A. M. Tyndall

of the University of Bristol when he was

president of Section A of the British Asso-

ciation for the Advancement of Science in

1952. He quoted from Prof. C. F. Powell’s

Nobel Prize citation: ‘“‘His special claim to

consideration is, In my view, the fact that

he has shown that discoveries of funda-

mental importance can still be made with

the simplest apparatus.’’ Powell had used a

photographic emulsion to record the tracks

of charged particles. Sometimes in our

awareness of the increasing importance of

JANUARY 1954 CAMPAIGNE: NUMBER OF HYPERGROUPS IN A GIVEN ORDER 5

large-scale facilities in pushing back the

frontiers of science, we forget that behind

the facilities there must be an individual,

and that behind the individual there must

be an idea. The Foundation is interested

in supporting such individuals with ideas.

The Foundation has no preconceived notion

as to whether these individuals will be

found primarily in the north or the south,

in the east or the west; in university or

college; whether they will belong predomi-

nantly to any one race or creed, to any one

nation or class.

Another fertile concept is that of the

great university or college. One readily

recognizes three areas of influence of any

modern educational institution. Initially

there is the fundamental concept of

teaching. Developing from this central moti-

vation is the sensitizing region of basic re-

search, while spreading from this stimu-

lating field is the outgrowth of public

MATHEMATICS.—A lower limit on the

Howarp H. CAMPAIGNE.

An outstanding problem in groups is to

determine the number of distinct groups of

a given finite order n. It is of some interest

to study this problem with the group axioms

relaxed. In this paper a lower bound for the

number of hypergroups of order n is shown

to be 8 X 11”-*. An upper bound can readily

be found.

A hypergroup is a set H of elements

satisfying three postulates.

1. If a and b are any two elements of H,

then there is associated with this ordered

pair a subset of H, not necessarily proper,

called the product ab.

If A and B are subsets of H, then their

product AB is defined to be the union of

all ab, for ain A and 6 in B. If O is the null

sen tnen Ob =O = BO. |

2. If a, b, and c are elements of H, then

a(bc) = (ab)ec.

3. If a and b are any elements of H, then

there are elements x and y, not necessarily

unique, such that b is in each of the sets

ax and ya.

Although the product ab is not neces-

sarily proper it cannot be void.

service, including applied research and

development. It does seem, however, that

the third function has often overbalanced

the other two which are the proper domain

of an educational institution (except pos-

sibly in times of national emergency).

Assuming that a college will be primarily

concerned with teaching, that a university

will be primarily concerned with research,

and that both have some responsibility for

public service, we believe the urgent need

today to be the establishing of more research

in colleges and the shifting of emphasis from

applied research to basic research in uni-

versities, at least insofar as fluid dynamics is

concerned.

In pursuit of these ideals of the American

scientist-scholar in the American college

and university, the National Science Foun-

dation is genuinely desirous of assisting

individuals interested in research and educa-

tion in fluid dynamics.

number of hypergroups of a given order.

In a hypergroup of order n there are

2” — 1 subsets which might appear as

products. There are n? products to be

defined. Therefore there cannot be more

than (2” — 1)’ hypergroups of order n,

and a crude upper limit is established.

In order to establish a lower bound we

will give rules for constructing a number of

distinct hypergroups. This will be done by

induction, that is, from each hypergroup

of order n will be constructed 11 of order

n + 1, which will be shown to be distinct.

The complete proof is tedious with special

eases, so only an outline will be given here.

Our problem then is to adjoin an ele-

ment a to a hypergroup H in such a way

as to form a new hypergroup G. Let hi,

hy, °-- be elements of H, and h;h; mean a

product in H, while h,h; means a product

in G. First define hih; = h;h;. Then for a

‘general hypergroup HH exhaustive trials

show that there are only six ways of defining

the products ah;, a’, and h,a so that G satis-

fies the three postulates. These are listed

below in tabular form.

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

Tyre 1 |d(xh)}. Furthermore any symmetric func-_

rule ah; a hia tion of one of these sets is an invariant. By

1 s H oO means of these invariants we will show the

2 a G a : : : :

3 GC i GC various extensions of H to be isomorphically

4 G G G distinct.

5 a G G Lemma |. Given any two nonisomorphic

6 G G a

There is another possibility for h,h;, that

is to have the element a adjoined to it, that

is, define h;h; = h;h;ua. Again there are

six ways of defining the remaining products.

Typr 2

rule ah; a? hia

1 G a G

8 jee a H

9 G H G

10 G G G

il H a G

12 G a H

There is a third type of rule conceivable

for which some products h;h; contain a and

others do not. If there is such a rule depend-

ing in no way on the hypergroup H then it

must apply to groups as well as to hyper-

eroups. Examining the application of the

rule to a group it appears that if zy =

xy Ua, Where xy = w, then a cannot be

adjoined to any product r-s ~ w. Further-

more w and a are inseparable, for if ts = w,

then ts = w Ua is a necessary consequence.

Now turn our attention to the hypergroup

T which has for every pair of elements ¢;t; =

T. If a is always associated with some ele-

ment then it is in every product ¢,t;. For T

then the rule is indistinguishable from a type

2 rule, and therefore useless to us. Thus only

the twelve rules tabulated above are pos-

sible.

In order to show that these extensions

are distinct we will need certain auxiliary

theory, which we outline here. Let d(h,h;)

be the number of distinct elements in the

product h;h;. Then the unordered set of n?

numbers D(H) = j{d(h:h;)}, where h; and

h; run through the hypergroup H, is an in-

variant under isomorphism. A necessary

and sufficient condition that H be a group

is’ that DG1)* consist. of all 1’s: The-set

S(H) = {d(h?)} of n numbers is also an in-

variant. The set J, = {d(ha)} where x ranges

through the n values of H, is also an in-

variant. By convention d(h?) is always the

last number in this set. Similarly i, =

hypergroups H and K of order n, with H

extended by a to hypergroup G, and K ex-

tended by b to hypergroup J. Then G and

J are not isomorphic provided rules 9 and

10 are not both used.

Outline of the proof. Suppose that the

rule used on K has a rule number not less

than that used on H. This is convenient

and quite general.

Now suppose the contrary of the lemma,

that G is isomorphic to J under a mapping

f(G) = J. Then f(a) ¥* 6b, for otherwise

f(H) = K, contrary to hypothesis.

Observe that d(h,h;) <,nif Gis formed by

a rule of type 1 and d(h,;h;) > 2 if the rule

is of type 2. A complete proof would require

the consideration of each rule applied to

each of the hypergroups H and K. In order

to shorten the exposition we will do only

a few rules on H.

Rules 7, 8, 11, and 12. In G we have

d(a?) = n + 1, which implies that J, if it

is isomorphic to G, was not formed by a rule

of type 2. Therefore these cases are impos-

sible.

Rule 9. Hypergroup K can have rules 9,

10, 11, or 12 applied to it. If rules 11 or 12

then d(b?) = 1 in J, and G cannot be iso-

morphic to J. If rule 9 is applhed to K to

form J, and if Gis isomorphic to J, f(G@) = J,

then there is an automorphism ¢ of J defined

as follows. Let é(6) = f(a), andi) — 3a

Ii f(a) # k # b then define t(k) = k. The

mapping tf(G) = J is a second isomorphism

of Gto J withif(a) = b. Thereforetf(H) = Kk,

contradicting the hypothesis. But J can be

formed from K by rule 10. In fact, if a hyper-

group A is extended by rules 9 and 10 to

give hypergroups B and C, respectively, and

if these are extended by rules 10 and 9 to

give B’ and C’ respectively, then B’ and C’

will be isomorphic, but B and C will not be.

By similar arguments all pairs of rules

can be excluded except 9 and 10, and this

will establish lemma 1.

There remains the possibility that the

same hypergroup H could be extended by

two different rules to isomorphic results.

JANUARY 1954 MAMAY:

Lemma 2. If H of order greater than 1 is

extended by two different rules to hyper-

groups F and G, then F and G are not iso-

morphic.

Outline of proof: Suppose F isomorphic

to G under the mapping m. We have F =

mua andG = Hub, andF = m(G). Evi-

dently a # m(b) by the definitions of the

various rules. Let p = p(#H) be the sum of all

the numbers in D = {d(h;h;)} over H. Then

p(G) is given for the 12 rules by the follow-

ing table.

rule p(G)

1 p + 3n

2 p+e3n-+ 1

3 p + 2n? + 2n + 1

4 p+ 2n? + 3n+ 1

5 and 6 p+ n* + 3n--+ 1

i p+ 3n?+ 2n+ 1

8 p + 3n? + |

9 p jon —-- on

10 p -- dn? —- on -+-1

11 and 12 p+3n?t+tn+1

It is evident that in general no two of these

are equal. There may be specific values of

n for which equality occurs; we will take

them up later. :

By rule 5 we have f, = {n+ 1,n+4 1,

-,n + 1}. By rule 6 there is no element

having this property. By rule 11 we have

d(a?) = 1, and by rule 12 there is no element

g except 6 for which d(g?) = 1.

The lemma has now been proved except

for those values of n mentioned above.

These exceptional values are n = 2 and 3.

For n = 2 rules 3 and 8 might conceivably

give isomorphic results. But if s(H) is de-

PERMIAN

DISCINITES CONE ré

fined to be the sum of all d(h?), we see that

s(F) — s(G@) = n = 2. Thus the results are

not isomorphic. For n = 2 rules 4 and 11

or 12 might give isomorphic results. But

by rule 11 we have J,°= {2, 2, 1}, and by

rule 4 every J; contains a 3. For rules 4 and

12 we have J, and J, similarly inconsistent.

For n = 38 rules 4 and 8 might give iso-

morphic results. By rule 8 we have J, =

{3, 3, 3, 1}, and by rule 4 there is a 4 in Jj.

This completes the proof of the lemma.

For n = 1 we can write out all the exten-

sions. There are exactly eight hypergroups

of order 2, and these are all given by our

twelve rules applied to the identity group.

Theorem: The number of distinct hyper-

groups of order n + 1 is at least 11 times

the number of order n, for n > 2.

Because rules 9 and 10 are exceptional

we discard one of them. Then the theorem

follows.

Theorem: The number of hypergroups of

order n > 3 is greater than 8 X 11”-.

For this many could be constructed by

starting with the hypergroups of order 2

and iterating the constructions. The groups

of order n > 3 are not among these, and

they bring the total above 8 x 11".

Other ways of extending a hypergroup

can be invented, such as adjoining one hyper-

group to another, or adjoining several ele-

ments simultaneously. But to prove the re-

sults isomorphically distinct by the methods

used here is too formidable to undertake

lightly.

PALEOBOTANY.—A Permian Discinites cone.! Serctus H. Mamay, U. 8. Geo-

logical Survey. (Communicated by James Stark Williams.)

The Paleozoic plant collections in the

United States National Museum include

an abundance of interesting Permian mate-

rial from the southwestern United States.

Aside from a few small lots, these collections

were made by the late David White and

C. B. Read, both of the U. 8S. Geological

Survey. With the exception of White’s de-

scription of Gigantopteris americana (1912)

and Darrah’s paper on Tingia (1938), ac-

1 Publication authorized by the Director, U.S.

Geological Survey.

counts of these floras have not been pub-

lished.

Recently the writer’s curatorial efforts

have primarily concerned these collections.

During the course of this preliminary survey,

conducted with the purpose of undertaking

-a long-range investigation of the Permian

floras of the American Southwest, several

unusual floristic features have been noted;

one of these is the presence of Dzscinites,

a genus not previously known to occur in

strata as young as the lower Permian. Four

fragmentary specimens (U.S.N.M. numbers

8 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

40626-40629) were found, representing two

localities in north-central Texas.

Specimen no. 40626 bears the U.S.GS.

locality number 8959. This collection was

made by C. B. Read and H. W. Ervin in

October 1940. Their collecting label reads:

“Upper part of Belle Plains formation in e.

central part of Emily Irish Grant, south side

of Salt Fork of Brazos River, Baylor County,

Texas. 16 to 1614 miles scaled due southeast

of Seymour, Texas.”’

Specimens 40627-40629 bear the U.S.G.S.

locality number 8877; these were among

collections made by David White in 1910.

According to White (1912, p. 495), this

locality is at: ‘‘...the bank of the stream

at the crossing of the old road, one-fourth

mile south of the ford of Little Wichita

River, 4 miles southeast of Fulda, a station

in Baylor County.” Material from this lo-

eality was the source of White’s paper

(1912) dealing with Gigantopteris americana;

this exposure, as well as that from which

the C. B. Read collection was made, occurs

in beds of the Wichita group, which is re-

garded as lower Permian (Moore, 1949).

Discinites is a genus of Paleozoic cones

that differ from other contemporaneous

strobiloid fructifications in that they possess

the following unique combination of struc-

tural features: |

1. The cones are verticillate, with a disc

of laterally fused sporophylls arising from

each node.

2. The upper surface of each disc is cov-

ered by numerous, closely set, apparently

sessile sporangia.

3. The cones are heterosporous, both

mega- and microsporangia being produced

in the same whorl. ;

Several species of Dziscinites are known,

all of pre-Permian age. The genotypic

species, D. bohemicus, was reported from

the Westphalian C of Czechoslovakia by

K. Feistmantel, founder of the genus, in

1879. Subsequently several additions to

the genus were made from the Westphalian

of Czechoslovakia. These are: D. major

Nemejc (1937), D. raconensis Nemeje (1941),

D. Hlizae Nemejc (1941), and three addi-

tional entities simply referred to by Nemejc

941) as Do spp. 7) 2) and:

The Westphalian of Holland has produced

vou. 44, no. |

one additional species, D. Jongmansii Hir-

mir (1940), and the known geographic

distribution of the genus has been extended

to the United States by Arnold’s (1949)

report of D. delectus (Arnold) Arnold and

D. Jongmansu among the Pennsylvanian

plants described in his flora of the Michigan

coal basin.

DESCRIPTION OF THE SPECIMENS

The transverse aspect of the cone is shown by

specimen 40626, which consists of the compres-

sion and counterpart of a single sporophyllar

disc, compressed parallel to its upper surface.

This dise, with its attached sporangia, is shown

in figures 1 and 2. The specimen is in a matrix

of blue-gray, fine-grained shale; there is little

carbonaceous material left in the specimen, parts

of its surface having been replaced by a powdery

white calcite deposit. However, such features as

the margins of the disc, position of the cone axis,

and the size and distribution of the sporangia

may be made out reasonably well.

The circular sporophyllar dise is about 2.5 em

in diameter, exclusive of the marginal teeth; at

one side of the specimen the margin is partly

buried in the matrix (broken line at the right

of Fig. 2). At the center of the disc there is a

slight elevation, which represents the cone axis

(Fig. 2, a). This is a rather stout axis, measuring

about 4 mm in diameter.

Part of the margin is completely flattened; in

this part several separate marginal teeth may

be seen (Fig. 2, mt). The teeth are 4 to 6 mm

long and about 1.5 mm wide at their bases, taper-

ing gently to blunt tips. Judging from their size

in relation to the diameter of the disc, there were

probably 40 or 50 teeth to the margin of the disc.

Aside from the central portion that represents

the cone axis, the surface of the disc is covered

by a pattern of closely fitting diamond-shaped

marks that doubtless represent the sporangia

(fig. 2, sp). They are slightly elongate in the

radial direction, reaching 3 mm in that dimen-

sion. There is no evidence to indicate the method

of sporangial attachment to the disc; nor can

the differences between mega- and microspo-

rangia be made out, if both kinds were present

in this whorl]. The sporangia cannot be accurately

counted, but, judging from the areas where they

can be counted and the sizes of these areas in

relation to the total area of the disc, probably

100 to 150 sporangia were attached to this disc.

JANUARY 1954

The size of this specimen and its attached

sporangia compares closely with that of the

specimen designated as Discinites sp. 1 by

Nemeje (1941, fig 1). In the shape of the spo-

rangia, however, it compares more closely with

D. bohemicus (cf. Nemejec, 1937, figure 2), and

in the toothed margin of its sporophyllar disc,

the specimen recalls D. Jonymansii (Hirmer,

1940, figs. 1-10).

The remainder of the material consists of

MAMAY: PERMIAN

DISCINITES CONE 9

three fragmentary specimens (40627-40629) that

clearly present the longitudinal aspect of Dis-

cinites (figs. 3-5). These are all from lo ality

no. 8877 and are preserved in a rather coarse-

grained, light-brown, sandy shale.

The largest fragment (specimen 40629) is

about 4.5 em long and 2 em wide (fig. 5). It

contains parts of 11 consecutive discs, each of

whose closely arranged sporangia may be seen.

The dises are about 2 mm apart.

Figs. 1-5.—Discinites sp.: 1, 2, Upper surface of a sporophylar disc (specimen 40626), Fig. 2 retouched

with white ink to show more clearly the marginal teeth (mt), the sporangia (sp), and the cone axis (a) ;

3, Fragment of a longitudinally compressed specimen (no. 40627) showing parts of nine successive whorls;

4, Fragment. of a longitudinally compressed specimen (no. 40628) showing parts of 13 successive whorls;

5, Fragment of a longitudinally compressed specimen (no. 40629) showing parts of 11 successive whorls.

Faint impressions of several of the upturned marginal teeth of the sporophyllar discs may be seen at

the right. (All figures 1.5.)

10 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

The sporangia appear to be in their natural

erect positions, apparently having undergone

little compaction during the process of preserva-

tion. They average about 2 mm in height and

1 mm in width as seen on their tangential surfaces

(Figs. 3-5). There is little organic matter left

in these specimens, and it was therefore im-

possible to isolate spores for comparison with

those of previously described Dviscinites speci-

mens. As in specimen 40626, there is no evidence

concerning the relative distribution of micro-

and megasporangia or the mode of attachment

to the discs.

At one side of specimen 40629, several up-

turned marginal teeth of the discs may be seen

(Fig. 5). As far as can be determined, these are

approximately the same size as those seen in

specimen 40626 (Figs. 1, 2), and they appear to

demonstrate the natural positions of the teeth.

The specimens shown in Figs. 3, 4, and 5 compare

favorably with D. delectus (ef. Arnold, 1949,

pl. 28, fig. 6). However, there is no suggestion of

pairing of the sporangia, as originally described

by Arnold (1944) when he first assigned his

specimens to the genus Bowmanites.

Although the generic characters of Discinites

are clearly demonstrated by the Texas speci-

mens, a specific designation for them is not

offered here, because of their imperfect preserva-

tion. There are close resemblances, pointed out

in previous paragraphs, between the Permian

specimens and certain of the older ones, but the

differences in both geographic and stratigraphic

occurrences present a strong likelihood that

additional material may eventually warrant the

erection of a new species for the Texas specimens.

DISCUSSION

The vexing problem of natural affinities

of detached reproductive organs of fossil

plants presents itself again in connection

with this occurrence of Dzscinites. While

there is some agreement that Dzscinites

is most probably referable to the Noeggera-

thiales, the broader relationships of the

Noeggerathiales in themselves are contro-

versial. The whorled construction of Dzs-

cinites, Tingiostachya, and other included

genera suggests a sphenopsid alliance for

this group, and although this feature alone

is insufficient evidence for an incontestable

assignment of the Noeggerathiales to the

Sphenopsida, such an interpretation seems

VOL. 44, No. 1

preferable to Hirmer’s (1940) inclusion of

the Noeggerathiineae within the Filicales.

A full discussion of the systematic position

of this group of plants is quite beyond the

scope of this paper; however, it should be

pointed out in passing that the reference of

these plants—with their strobiloid fructifica-

tions—to the Filicales represents a radical

and, in the writer’s opinion, unwarranted

departure from the established tenets of

fern morphology and systematics.

The more immediate problem lies in the

identity of the vegetative parts of the plant

that bore Discinites. On the basis of associa-

tion alone, Nemeje (1937) and Hirmer

(1940) have suggested the probability that

Discinites is the fructification of the same

plant that bore Palaeopteridium foliage;

this association has also been noted by

Arnold (1949) in the flora of the Michigan

coal basin. This Permian occurrence of

Discinites throws doubt on such a proposed

relationship, however, for as far as presently

known, Palaeopteridium is restricted to

horizons of pre-Permian age. It is, then, of

interest to examine briefly the floristic ele-

ments associated with the Texas Discinztes,

with the purpose of attempting to shed some

light on this problem.

White (1912), in his discussion of Gigan-

topterts americana, published a list of the

plant genera associated with Gigantopteris

(locality 8877). The following genera were

noted: Annularia, Araucarites, Aspidiopsis,

Cordaites, Diplothmema, Gomphostrobus,

Neuropteris, Odontopteris, Pecopteris, Poacor-

daites, Sigillaria, Sphenophyllum, Taeniop-

teris, and Walchia.

The writer’s survey of this collection has

added the following genera to this list:

Aphlebia, Callipteris, Daubreeia, Lebachia,

Lepidophyllum, and Odontopteris. A few

undetermined strobiloid fructifications are

also present in the flora.

The following genera have been tenta-

tively identified in the C. B. Read collection

(locality 8959): Annularia, Aphlebia, Arau-

carites, Callipteris, Gigantopteris, Lebachia,

Odontopteris, Pecopteris,. Sphenophyllum,

Taeniopteris, and Tingia. This flora is essen-

tially the same as that of White’s collection |

in generic composition; the most outstand-

ing difference between the two lies in the

JANUARY 1954

apparent absence of 72ngia from the White

collection; this genus is very abundant in the

Read collection.?

Thus far, then, 77ngza is the only vegeta-

tive element of supposedly noeggerathialean

affinity known in these two floras; if one

were to attempt to establish a fructification-

foliage alliance for Discinites on the basis

of association alone, Jingia might well

‘appear to be the logical foliage genus in

these assemblages with which the Dzscinites

specimens should be considered allied. How-

ever, the fructification of Tingia (Tingio-

stachya), as described by Kon’no (1929),

differs strikingly from Discinites in some

important morphological features. J ingio-

stachya bore unfused sporophylls in whorls

of four, each sporophyll producing a single

synangial group of four sporangia on its

upper surface; it seems unlikely that two

types of cones as morphologically dissimilar

as Tingiostachya and Discinites could belong

to one and the same natural genus of plants.

All the present evidence considered, then,

Discinites still retains the status of a genus

of detached fructifications, the identity of

whose vegetative parts is not established,

even though they themselves may be well

known as detached foliar organs under a

different generic name. This occurrence of

Discinites is of interest, however, in that

it provides not only a link between the

floras of the Pennsylvanian and the Permian,

2 Darrah (1938) described two species of Tingia

(T. taeniata and T. kempiae) from Texas; accord-

ing to his locality data, the source of his material

is within 1 or 2 miles of the C. B. Read locality

(locality 8959), in rocks of the same stratigraphic

horizon.

HUI-LIN LI: TRAPELLACEAE LL

but also an additional link between the

floras of Europe and North America. It has

also served the writer as one of many illus-

trations of the poorly understood nature of

American Permian floras, the alleviation of

which condition constitutes one of the out-

standing needs of American paleobotany.

REFERENCES

ARNOLD, C. A. A heterosporous species of Bow-

manites from the Michigan coal basin. Amer.

Journ. Bot. 31: 466-469. 1944.

———. Fossil flora of the Michigan coal basin. Con-

trib. Mus. Paleontol. Univ. Michigan 7: 131-

269. 1949.

Darra, W. C. The occurrence of the genus Tingia

in Texas. Harvard Univ. Bot. Mus. Leafl. 5:

173-188. 1938.

FEISTMANTEL, K. Hine neue Pflanzengattung aus

bohmischen Steinkohlenschichten. Sitzungsb.

Bohm. Ges. Wiss. 1879: 298-303.

HirMerR, M. Noeggerathiineae. In: Hirmer, M..,

and Guthorl, P. Die Karbon-Flora des Saar-

gebeites. Abt. 3: Filicales und Verwandte.

Palaeontographica, Suppl. Bd. 9: 3-60. 1940.

Kon’no, E. On genera Tingia and Tingiostachya

from the Lower Permian and the Permo-Triassic

beds in northern Korea. Trans. and Absts. Jap.

Journ. Geol. Geog. 6: 113-147, 1929.

Moore, R. C. Rocks of Permian (?) age in the

Colorado River Valley, north-central Texas.

U.S.G.S. Oil and Gas Inv. Prelim. Map 80.

1949.

Nemesc, F. On Discinites K. Feistm. Bull. Intern.

Acad. Sci. Boheme 38: 3-10, 1937.

———. Weitere Fructifikationem vom Typus Disci-

nites nebst einigen Bemerkungen wiber die

Archaeopteriden der Mittelbohmischen Kohlen-

becken. Mitteil. Tschech. Akad. Wiss. 1941:

1-13.

Wuitk, D. The characters of the fossil plant Gi-

gantopteris Schenk and its occurrence in North

America. Proc. U. 8. Nat. Mus. 41: 493-516.

1912.

BOTANY —Trapellaceae, a familial segregate from the Asiatic flora. Hu1-Lin Lt,

Morris Arboretum, University of Pennsylvania.

The genus Trapella, like many other

genera of plants of temperate eastern Asia,

is apparently of a relic nature. It is treated

as an anomalous member in the nearest

largest family Pedaliaceae, as a result of the

earlier prevalent view against unigeneric

families. However, the more favored view

among taxonomists and morphologists at

present is to recognize phylogenetically iso-

lated genera as representing distinct families

irrespective of their sizes. It seems that a

phylogenetic system of classification can be

more readily elucidated by recognizing more

‘families with clearcut circumscription rather

than fewer, larger, more inclusive but also

more ambiguous ones. The tendency now

is to segregate more and more such isolated

genera into independent families of their

own. The case of Trapella is summarized

below.

12 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

Trapellaceae, fam. nov.

Pedaliaceae tribe Trapelleae Oliver in Ann.

Bot. 2: 110. 1888; Stapf in Engler and

Prantl, Natur. Pflanzenf. IV. 3b: 260, 265.

1895.

Herba natans, foliis oppositis petiolatis, in-

ferioribus lineari-oblongis denticulatis superiori-

bus deltoideo-rotundatis vel cordiformibus cre-

nato-denticulatis.

Flores axillares solitarii pedunculati. Calyx

tubo ovario adnato, limbo libero 5-fido. Corolla

perigyna tubuloso-infundibuliforma, limbo pa-

tente bilabiato, labio superiore breviter bifido,

-labio inferiore trifido; tubo basi abrupte an-

gustato, aestivatione imbricata, labio superiore

exteriore. Stamina pollinifera 2 epipetala inclusa,

antheris bilocularibus, connectivo peltato carno-

sulo insidentibus; filamentis filiformibus glabris;

staminodia antica 2 elongata; stamen posticum

nullum. Ovarium inferum apice tantum lberum

biloculare, loculo antico rudimentario, loculo

postico biovulato; stylo gracile elongato, stigmate

bilabiato; ovula 2, anatropa, pendula, superiore

sessili, inferiore breviter funiculato deinde abor-

tivo. Fructus angustus elongatus monospermus

indehiscens, apice appendicibus 5 coronatus

elongatis arrectis apice uncinatim incurvis, 2

brevioribus spinosis subulatis rectis patentibus;

pericarpio tenuiter charteceo-lignoso. Semen

pendulum, cylindraceum, endospermio tenul,

embryonis recti radicula supra, cotyledonibus

lineari-oblongis radicula brevioribus.

One genus, Trapella, with two species, widely

distributed in eastern Asia.

Trapella is one of the most interesting plants

discovered in China. It is an aquatic plant with

the appearance of Trapa but is very different in

its characters. At first one species was known for

the genus, 7. sinensis Oliver (1887), first de-

scribed from central China but later known to

occur also in eastern, northern, and northeastern

China as well as Japan. Gltick (1939) later dis-

covered that the genus is actually composed of

two species, one, 7’. sinensis Oliver, in Kyushu

of Japan, maritime Siberia, and northeastern

as well as northern, eastern, and central China,

and another 7. antennifera (Léveillé) Gliick, in

Honshu of Japan, northeastern China, and mari-

time Siberia only.

Trapella, since its discovery, has remained in

the Pedaliaceae assigned by the original author.

When Oliver first described this plant, he was

VOL. 44, NO. 1

very much impressed by its uniqueness. His as-

signment of the genus in the ‘‘Pedalineae”’ is,

however, tentative, as he noted that exceptions

might surely be taken for this action. He did so

not because of the agreement of the genus in all

essential characters with the Pedaliaceae, but

mainly to avoid the alternative of making this

the type of a distinct ‘‘order.”’

Oliver noted that Trapella is intermediate in

its characters between the Pedaliaceae and Myo-

poraceae and that it can be considered as an aber-

rant member, taken to aquatic habits, of the

Pedaliaceae, which have all other species terres-

trial. The genus is distinct from other genera of

the family not only in the aquatic habit but also

in the geographical distribution. All other genera

of the Pedaliaceae are shore and desert plants in

tropical Africa, Madagascar, the Indo-Malayan

region, and tropical Australia. The tropical and

subtropical American genera formerly included

in the Pedaliaceae are now separated as a distinct

family Martyniaceae.

Trapella was treated as representing a distinct

tribe Trapelleae in the Pedaliaceae by F. W.

Oliver (1888-89) and this was followed by Stapf

(1895), who recognized the other genera as form-

ing the tribe Pedalieae.

A very thorough study on the morphology of

the vegetative as well as reproductive structures

of this very anomalous genus was made very

shortly after its discovery by F. W. Oliver (1888-

89), son of the original author. Young Oliver is

of the opinion that Trapella is related more

closely to Myoporaceae and Pedaliaceae than

other families of the “bilabiate monopetalae.”

He regards the wide discrepancies between Pedal-

iaceae and T'rapella as due to the change of habit

in the latter, and he refrains from making the

genus a new family because he considers the

Pedaliaceae, Myoporaceae, and other related

families as mostly ill-defined. A very detailed de-

scription of the genus is given in his paper, basing

on the original description of the genus with

modifications and additions resulting from his

intensive studies. The diagnosis of the family as

given above is adopted from his detailed descrip-

tion. An intensive taxonomical and morphological

study of the genus was made also by Glick

(1941).

In morphological characters, the chief distinc-

tion of Trapella from the Pedaliaceae are the in-

ferior ovary and the presence of only two fertile

JANUARY 1954

stamens. Its relationship with the Pedaliaceae is

indicated with Pedaliwm, in the bilocular ovary

with two pendulous ovules. The appendaged

fruit forms a strongly analogous character with

the Pedaliaceae. In the form and arrangement of

the seed it is near the Myoporaceae but it differs

in the fruit and the opposite leaves. Trapella thus

ean be considered as representing the type of a

distinct family somewhat linking the Pedaliaceae

and Myoporaceae.

CLARK: PRESCRIPTION AS APPLIED TO TAXONOMY 13

LITERATURE CITED

Guiick, H. ‘Uber eine neue Trapella des dstlichen

Asiens. Bot. Jahrb. 70: 149-152. 1939.

Die Gattung Trapella. Bot Jahrb. 71:

267-336. 1941.

OuivER, D. Trapella sinensis, Oliv. Hook. Icon.

Pr 16 2 pl. 1595. Uss7.

OuIvER, F. W. On the structure, development, and

affinities of Trapella Oliv., a new genus of

Pedalineae. Ann. Bot. 2: 75-115. 1888-89.

Starr, O. Pedaliaceae. In Engler and Prantl,

Natur. Pflanzenf. IV. 3b: 260-265. 1895.

TAXONOMY .—For and against the doctrine of prescription as applied to taxonomy:

A historical retrospect. AUSTIN H. Cuarx, U.S. National Museum.

In these days of renewed interest in

taxonomy and in revisions of and emenda-

tions to the International Code of Nomen-

clature it is perhaps of interest to put on

record the sentiment expressed by the

leading zoologists and paleontologists of

nearly 50 years ago.

In the early years of the present century

systematic zoology, including paleontology,

was in high favor. Previously unknown

species were being discovered in large num-

bers by expeditions to various little-known

parts of the world and through the intensive

exploration of the deep sea, and material

that had been collected earlier was being

intensively studied. This work, especially

the revisionary studies connected with it,

naturally focused attention on taxonomy.

Many names in general use were found

to be untenable according to the strict

application of the Rule of Priority. In ac-

cordance with the rule of priority an Inter-

national Code of Nomenclature had been

drawn up, based chiefly on previous codes

concerned mainly with terrestrial verte-

brates. This code proved of great value in

stabilizing zoological nomenclature, but its

strict application in certain cases led to the

suppression of many well-established names

in general use (as for example Holothuria)

and also to interminable controversies re-

garding species inadequately described by

early authors of which the type specimens

had disappeared. Furthermore, the fossils

did not come under the binomial principle

elaborated by Linnaeus; they were made

exceptions through ignorance of their true

nature.

The binomial system was slow in becom-

ing established in purely fossil groups, and

there was, and still is, much controversy

in regard to the binomial status, or other-

wise, of many of the works of the earlier

authors.

Dissatisfaction with the inflexibility of

the Code and with its arbitrary interpreta-

tion and application at that time was

becoming marked, and so in 1909 the Hon.

Frank Springer of the Territory of New

Mexico, the well-known authority on the

fossil crinoids, and leader of the New Mexi-

ean bar, and I decided to test the prevailing

attitude toward it on the part of our col-

leagues.

The genera Hnerinus, Pentacrinus, Iso-

crinus, and Mullericrinus, among the best

known and most firmly established of all

the genera of the Crinoidea, are all either

untenable or of doubtful availability ac-

cording to the strict application of the

current code of zoological nomenclature

adopted by the International Zoological

Congress. In a circular dated Burlington,

Iowa, May 1, 1909, Mr. Springer gave a

summary of the involved history of the

genus Hncrinus and showed that great

confusion would result if the genotype were

determined according to the strict applica-

tion of the rules.

Encrinus liliiformis is the best known of

all the stalked crinoids. It has been figured

and described under that name in countless

works, and specimens are found under that

name in all the important cabinets and

museums of the world. Encrinus Blumen-

bach, 1779, is generally accepted, with the

14. JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

genotype LHncrinus liliwformis Lamarck,

1801. But according to a strict application

of the code Hncrinus should date either

from Andreaé, 1763, with the genotype

E.. coralloides, which is supposed to be the

terminal stem branches or roots of a species

of Muillericrinus, one of which has been

referred to M. echinatus by de Loriol; or

from Blumenbach, 1779, with the genotype

Isis asteria Linné (= the Recent West

Indian Cenocrinus asteria); liliformis was

not included in Enerinus by Blumenbach

(see A. H. Clark, Ann. Mag. Nat. Hist. (8)

3 (15): 308-310. March 1909). Mr. Springer

wrote:

The results of either of these applications of

the name Encrinus, one or other of which would

be rendered necessary by a strict adherence to the

International Code, would be to throw the study

of the fossil crinoids into intolerable confusion,

not so much for the workers in the group, like

myself, who can take immediate cognizance of

any change, but for the much larger number of

students interested in stratigraphy, paleontology,

general geology, and allied subjects. The shifting

of the names, commonly accepted and heretofore

unquestioned, of the commonest genera—genera

predominantly characteristic of certain horizons—

would impose an intolerable burden upon every

one who ever had any occasion to refer to the

crinoids in any way. The change, even if it could

finally be brought about, would take years to

accomplish, and the burden would fall heaviest on

those to whom crinoids were only of incidental

interest, though of the greatest indirect import-

ance, in the identification of strata, or in the

instruction of students. The result would be

hopeless confusion, would benefit nobody, and

could not fail to bring ridicule upon the taxonomic

methods now in vogue.

Moreover, it must be borne in mind that the

fossils did not come under the binomial principal

as elaborated by Linnaeus; they were made ex-

ceptions through ignorance of their true nature.

Thus we find that the binomial system was slow in

becoming established in purely fossil groups, and

there will always be controversy in regard to many

of the works of the earlier authors.

The underlying principle of the rule of priority

is said, and properly said, to be fixity. Yet by

insisting upon its absolute and unbending applica-

tion in all cases, without regard to circumstances,

we may destroy the very fixity for which we

contend. There is no law more deeply rooted in the

foundations of civil government, or more essential

to the welfare and stability of society, than that

of the fixity of the titles to real estate based on

priority. But just as that law in actual adminis-

tration is subject to exceptions founded upon

principles of natural justice and the dictates of

public policy, so I think we may find reasonable

voL. 44, No. 1

basis for an exception to the rule of priority in

nomenclature which will meet such cases as this.

This would be that such cases, irrespective of

the actual state of the record as to their dates,

should be protected under an exception to the

rule, simply on the ground of long use, on the

doctrine of prescription, which is a principle well

known in law, recognized in continental Europe

as coming down from the civil law of Rome, and

now embodied in statutes in all English-speaking

countries. It is that the right of property will be

upheld by the courts in favor of one who can show

a long, continuous, and undisputed possession of

it, under a claim of right, however defective, not-

withstanding he has no paper title, and even

though the records may show the prior title to be

in some one else. This rule of law rests upon the

idea that it 1s for the public interest that there

be an end of controversy, and that there shall be

some reasonable time after which titles may be

held safe from attack on any ground. And this end

was attained in the beginning, not by denying or

abrogating the law governing the conveyance of

property by deeds, but by invoking a simple

- presumption, founded on the known as usual

conduct of men with regard to their interests,

that where such long and undisputed possession

existed there must have been a good title, the

evidence of which is lost. .

This principle of jurisprudence is now recog-

nized throughout the civilized world as one of the

most salutary and beneficial provisions for pre-

venting injustice and insuring that repose of titles

which the peace and order of society demand.

By virtue of its operation a title by lapse of time

merely, if properly proven under all the safeguards

which are prescribed in practice to prevent the

abuse of it, is as good in the actual possessor as a

paper title showing priority by an unbroken chain

of recorded deeds. If this be true with regard to

matters of such vital importance as the titles to

our landed property, why may not the same

principle be invoked in favor of repose and stabil-

ity of names in our scientific literature? It is not a

question of ‘‘doing justice’ to any particular

ancient author. The proposition is one of far

broader significance, and involves the paramount

interest of the scientific public.

In this way, by analogy to the practice which

prevails in courts of justice touching the most

solemn rights of property, a presumably just

conclusion can be reached independent of the rule

of priority, and without impairing its force in

cases to which no such considerations of public

policy apply.

In view of the above, I am in favor of making an

exception to the rigid application of the law of

priority in regard to the exclusively fossil genus

Encrinus, accepting it from. Schulze, 1760,1 by

1 Mr. Springer wrote that Schulze’s work ‘‘was

mainly a compilation from former authors, as

Linck, Lluyd, Seba, Capelier, and Ellis, and he

uses their names in the same manner as they did,

with but small pretense to binomial application.

JANUARY 1954

which means all the above mentioned genera of

erinoids [Encrinus, Pentacrinus, Isocrinus, and

Millericrinus| would be retained in the same

significance in which they are used today, and

have been used for the better part of a century.

Mr. Springer concluded “I am asking you

to indicate on the enclosed card whether,

in view of the above considerations, you

are in favor of making an exception to the

International Code in favor of Hncrinus,

or whether, in your judgment, the best

course would be to adhere strictly to it in

this case, notwithstanding the deplorable

confusion which would result.”

The enclosed postal card, which was ad-

dressed to me, had two alternatives, each

followed by a line for a signature. The

alternatives were:

(1) I am in favor of accepting Encrinus

from Schulze, 1760, and of retaining it and

the other crinoid genera affected in the same

sense as understood today.

(2) I am in favor of a strict adherence to

the International Code, regardless of the

effect on the present nomenclature.

This circular was sent to 1,000 zoologists

and paleontologists.”

We received 376 replies to the circular,

from Algeria, Brazil, Canada, Ceylon,

Denmark, Egypt, Eire, England, Finland,

France, Germany, Italy, Japan, Nether-

lands, New South Wales, New Zealand,

Norway, Philippines, Portugal, Queensland,

Russia, Scotland, South Australia, Sweden,

Tasmania, Trinidad, B.W.I., United States,

Victoria, and Western Australia.

Of these replies 297 (nearly 80 per cent)

favored retaining Hncrinus, and 62 (about

20 per cent) favored strict adherence to

the Code, a number of them with reluctance.

He did not propose Encrinus to represent a genus,

but only mentioned, by way of recital, the fact

that certain petrifactions resembling a lily have

been called the lily-stone, Encrinus (‘Man findet

eine gewisse Versteinerung, die, in Ansehung ihrer

Gestalt, einige Gleichheit mit einer Lilie zu haben

scheinet, daher man dieselbe enfanglich fur die

Versteinerung dieser Blume gehalten, und sie den

Lilienstein, ENCRINUM, genennen hat.’). On plate

IV is a figure of a complete crown of the fossil to

which he refers, and in the long description which

follows he mentions it four times by the name

‘Lilienstein,’ but never again as ‘Encrinus.’

The figured specimen is EL. liliiformis Lamarck.”

2See Austin H. Clark, The strict application

of the law of priority to generic names, Science,

n.s., 31 (787): 145-146. January 28, 1910.

CLARK: PRESCRIPTION AS APPLIED TO TAXONOMY in

To show the broad general interest taken

in this matter at that time it may be men-

tioned that among those who replied were

Alexander Agassiz, Count Arrigoni Degli

Oddi, Lord Avebury, E. G. Conklin, Theo-

dore Gill, Sir Sydney Harmer, Ernst

Hartert, John B. Henderson, Jr., Edgard

Hérouard, John C. Merriam, Edward S.

Morse, Adam Sedgwick, Sir D’Arcy Thomp-

son, A. E. Verrill, Charles D. Walcott, and

Alfred Russel Wallace.

Following is an analysis of the replies.

FOR RETAINING ENCRINUS

Of those who returned the cards or wrote letters

243 favored the first alternative, the acceptance

of Encrinus from Schulze, 1760, without comment.

Among these were:

George Abbott, Charles C. Adams, Nicolai

Adelung, Alexander Agassiz, M. J. Ahern, A.

Aleock, Edward Phelps Allis, Jr., Richard John

Anderson, A. W. Anthony, Prof. Dr. Appelléf,

Count Arrigoni Delgi Oddi, E. A. N. Archer, Chr.

Aurivillius, Lord Avebury [formerly Sir John

Lubbock], G. E. H. Barrett-Hamilton, Walter B.

Barrows, R.S. Bassler, F. E. L. Beal, Tarleton H.

Bean, C. William Beebe, F. Jeffrey Bell, Charles

P. Berkey, S. W. Berger, A. Bibbins, W. B. Ben-

ham, M. A. Bigelow, H. P. Blackmore, J. E. V.

Boas, L. A. Borradaile, Adam Béving, H. Bolton,

Aug. Brinkmann, Hjalmar Brock, Arthur Erwin

Brown, J. Buttikofer, W. T. Calman, Oskar

Carlgren, J. W. Carr, W. D. Carr, Thomas ‘L.

Casey, George H. Chadwick, Robert E. Coker,

Leon J. Cole, E. G. Conklin, H. Coutiére, William

A. Cunningham, Ulric Dahlgren, W. Boyd Daw-

kins, R. Etheridge, W. L. W. Field, G. H. French,

John H. Gerould; O: C. Glaser, E. L. Golds-

borough, Seitaro Goto, L. C. Graton, Laurence E.

Grifin, R. J. Lechmere Guppy, Robert Gurney,

H. J. Hansen, Chas. W. Hargitt, Clemens Hart-

laub, Sir Sydney F. Harmer, John B. Henderson,

Jr., Junius Henderson, P. P. C. Hoek, S. J.

Holmes, A. D. Hopkins, Walter Howchin, I.

Ijima, Hartley H. T. Jackson, Robert T. Jackson,

Otto Jaekel, O. A. Johansen, Lynds Jones, Chaun-

cey Juday, Hector F. E. Jungersen, W. C. Ken-

dall, John T. Kemp, J. Graham Kerr, H. Kirk-

patrick, K. Kishinouye, N. Knipowitsch, K.

Kraepelin, B. W. Kunkel, H. H. Lane, Torsten

Lagerberg, J. M. R. Levinsen, Edwin Linton, F.

A. Lucas, Wiliam Lundbeck, Henry H. Lyman,

Richard C. McGregor, A. Gibb Maitland, B.

. Pickman Mann, E. L. Mark, Geo. W. Martin, K.

Martin, O. Maas, S. E. Meek, E. A. Minchin,

John Mitchell, Henry Montgomery, Roy L.

Moodie, Carlos Moreira, Edward 8S. Morse, Th.

Mortensen, Henry F. Nachtrieb, John Treadwell

Nichols, A. M. Norman, Hj. Ostergren, Paul

Pallary, Raymond Pearl, G. Pfeffer, A. L. Quaint-

ance, Wilhelm Ramsey, Herbert W. Rand, Paul

M. Rea, C. Tate Regan, Jacob Reighard, Robert

16 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

Ridgway, Alice Robertson, Rudolf Ruedemann,

G. O. Sars, R. 8. Scharff, W. L. Selater, E. A.

Schwarz, H. H. Scott, Adam Sedgwick, H. W.

Shimer, C. Ph. Sluiter, Frank Smith, Grant

Smith, John B. Smith, Sanderson Smith, T.

Southwell, J. W. Spengel, E. C. Stirling, F. B.

Sumner, W. M. Tattersall, Hjalmar Théel, Sir

D’Arcy W. Thompson, Charles D. Walcott, P. R.

Uhler, HO. Ulrich, A: BE. Vermll; Gens A> W.

Vogdes, U.S.A., B. E. Walker, Henry B. Ward,

Stuart Weller, W. M. Wheeler, R. P. Whitfield,

C. O. Whitman, W. H. Wickes, 8S. R. Williams, 8.

W. Williston, Chas. B. Wilson, Herluf Winge,

Lorande Loss Woodruff, H. Woods, Horace B.

Woodward, J. B. Woodworth, and Dean C. Wor-

cester.

In addition to these, 54 zoologists and

paleontologists were in favor of retaining

Encrinus as suggested, but added comments.

These may be classified as follows.

1. Questions such as this should be pre-

sented to the International Commission to

be adjudicated by the Commission itself or

by a committee or committees appointed

by the Commission. Among those taking

this view were:

Glover M. Allen, E. P. Bailey, F. A. Bather,

James E. Benedict, R. P. Bigelow, A. J. Jukes

Browne, Charles B. Davenport, Hubert Lyman

Clark, Walter L. Hahn, Reginald Heber Howe,

Jr., Charles A. Kofoid, F. B. Loomis, Alfred G.

Mayer, Herbert Osborn, Raymond C. Osburne,

A.S. Pearse, Charles Schuchert, Hugh M. Smith,

T. Wayland Vaughan, and L. B. Walton. |

2. There were 15 replies that favored

retaining old established names. These were

from:

Robert Anderson, Prof. Apstein, J. W. Beede,

Lyman Belding, Wesley R. Coe, L. Cuénot, W. R.

Dudley, Charles L. Edwards, J. Stanley Gardner,

Francis H. Herrick, J.S. Kingsley, Alfred C. Lane,

John C. Merriam, Charles 8. Prosser, and Alfred

Russel Wallace. Of these, three suggested time

limits of general use—20-25 years (Wallace), 30

years or more (Hoek), 75 or 100 years (Beede), and

100 or even 50 years (Belding).

3. Among the replies 11 regarded strict

adherence to the Code as leading to con-

fusion or to absurd results, as “perfectly

idiotic,” or as “rank nonsense.” These

were from:

T. B. Bonney, Charles A. Chilton, Ludwig

Doderlein, J. H. Fleming, R. Fourtau, Robert H.

Gordon, L. P. Gratacap, R. Koehler, W. P. Py-

craft, Thomas Scott, and W. L. Tower.

vou. 44, No. |

There were also scattering comments.

Oldfield Thomas supported Encrinus on the

ground that it is technically valid. William

Sorensen said, ‘‘(1) I cannot see that thereby

the rules are broken, and (2) one must have

a motive to do a thing, but not to do noth-

ing.’ G. W. Kirkaldy wrote, “I would

have to examine the various papers myself

before giving an opinion, but I would point

out that Sherborn considers Schulze bi-

nomial.’’ David Starr Jordan said, ‘‘Under

the Code is not encrinus L. necessarily the

type of Hncrinus Blumenbach? I am on the

fence at present. The considerations are

strong on both sides. I think that if I were a

paleontologist I should wait before changing

these names. I am not sure that under the

Code Encrinus Schulze is not tenable. In

any case, this is a very difficult problem.

Did Andreaé have Schulze’s work in mind?

Is Pentaceros Schulze tenable?” W. L. Mc-

Atee wrote ‘Stability of nomenclature at-

tained by means however arbitrary is

preferable to the continual changing which

our best intentioned rules seem powerless to

prevent.”’? Warren D. Smith favored the

retention of Encrinus, but also favored a

strict adherence to the Code in future work.

Theodore D. A. Cockerell analyzed in detail

the status of Encrinus.

In addition to returning the cards, the

following wrote at length regarding their

views:

Thomas L. Casey T. D. A. Cockerell, Leon J.

Cole, H. Coutiére, L. Déderlein, J. Graham Kerr,

F. A. Lucas, Richard C. McGregor, Th. Mor-

tensen, Rudolf Ruedemann, Henry B. Ward, and

S. W. Williston.

FOR THE STRICT APPLICATION OF THE CODE

Of those who returned the cards, 42 fa-

vored the second alternative, the strict

application of the Code, without comment.

Among these were:

Paul Bartsch, Wilhelm Blasius, Sergius A.

Buturlin, C. Callaway, Morton L. Church, Robert

Collett, J. A. Cushman, A. A. Doolittle, C. H.

Eigenmann, Barton W. Evermann, W. K. Fisher,

C. H. Gilbert, Theodore Gill, O: PP.» Hay. W. ke:

Hay, Harold Heath, H. W. Henshaw, Charles W.

Johnson, Frederick Knab, F. H. Knowlton, G. W.

Lee, M. W. Lyon, Jr., Gerrit 8. Miller, Jr., E. W.

Nelson, Harry C. Oberholser, J. Douglas Ogilby,

A. E. Ortmann, Henry A. Pilsbry, Franz Poche,

JANUARY 1954

Julius Pohlman, Edward A. Preble, Mary J.

Rathbun, James A. G. Rehn, Harriet Richardson,

Charles W. Richmond, J. H. Riley, R. W. Sharpe,

Witmer Stone, W. E. Clyde Todd, George Wagner,

and W. M. Winton.

In addition to these, 20 zoologists and

paleontologists added comments. J. J.

Buckman said, ‘“‘I am in favor of a strict

adherence to the Law of Priority properly

interpreted when I hope it will be proved

that it will not have any effect on the

present nomenclature. August Busck wrote

that “‘similar cases just as tempting to make

exceptions of occur in Lepidoptera, on which

there is far more literature than on Encrinus.

One exception justifies another and gives

chance for differences of opinion.” A. N.

Caudell noted that ‘“To do otherwise would

set a bad example. Many genera in other

lines have the same claim on exception.”

Frederic Chapman said, “‘It is a bad surgical

ease, but I fear there is no way out if we

accept the rules.”” M. L. Fuller noted that

“This doubtless leads to confusion at times,

but on the whole it seems a good policy.

The way to uphold and establish it so that

in the end the greatest good will result is to

avoid making exceptions.’ Ernst Hartert

said, “I am in favour of a strict adherence

to the International Code, regardless of the

effect on the present nomenclature.’ Her-

mann von Ihering wrote, ‘‘We have already

proceeded against our desires in applying

the international rules In many cases and

shall do so also in the present....” F. A.

Jentink remarked, ‘“‘I am in favor of a strict

adherence to the International Code regard-

less of the effect on present nomenclature.” -

E. L. Morris said that ‘‘priority is the only

stable basis for all time.”’ R. I. Pocock was

in favor of strict adherence to the Inter-

national Code “‘because the anticipated ill

effects are always greatly exaggerated, and

because if one exception be made a thousand

will have to follow, each worker having pet

names he would like to preserve in statu quo

ante.”

However, 11 of those who voted for the

strict adherence to the Code were not en-

tirely satisfied with it. Louis B. Bishop said,

“T was not in favor of any of the recent

changes in ornithological names, believing

it far better for all to agree to stick to what

CLARK: PRESCRIPTION AS APPLIED TO TAXONOMY ay

we had that were thoroughly accepted

regardless of priority, but it is now too

late.” John M. Clarke wrote, ‘The prin-

ciples of judicial procedure if applied to

authors would ignore the element of equity

which is essentially safeguarded by the

International Code. The disturbance of

conventional use is a temporary incon-

venience to which science will eventually

adjust itself. Jonathan Dwight, Jr., said,

“At present there is no alternative except

to play the game according to the rules.

Nobody has a right to make exceptions

because there is no court of appeal to them,

and the whole discussion of Hncrinus is a

plea for preference instead of rule.’”’ William

H. Dall was in favor of accepting Hncrinus

‘af this can be authorized by the vote of

the International Zoological Congress.”’

George H. Girty thought that ‘dropping

Encrinus from the literature as not recog-

nizable [Encrinus of Andreaé]... might be

thought a pity but would not lead, I

would think, to much confusion.’’ Edgard

Hérouard upheld strict adherence to the

Code, but believed it would be useful to

modify the Code in this case. Wilfred H.

Osgood favored strict adherence to the

Code, but said, “I would favor modification

of the Code, even very drastically, in order

that such names might be retained.” H. E.

Summers said that any exceptions should be

made by the International Congress. Henry

L. Ward wrote, ‘“Taken by itself it would

seem advisable to retain Hncrinus, but there

are others, and some rule must be enforced,

and as we have no courts of law the rules

must be self-enforcing.’”? David White was

in favor of strict adherence to priority in the

binomial usage, thatis, for Encrinus Andreaé.

Letters giving their views in detail were

received from J. J. Buckman, W. H. Dall,

Jonathan Dwight, Jr., C. H. Gilbert, George

H. Girty, G. Douglas Ogilby, and Franz

Poche.

NO OPINION EXPRESSED

J. A. Allen, A. J. Bigney, Arthur M.

Edwards, and Arthur H. E. Mattingley

sent in cards signed without comment in

both places. One correspondent said he was

not competent to express an opinion, “But

in general I deplore the discarding of old

18

and long accepted familiar names in any

branch of natural history, resulting as it

does in constant confusion and discourage-

ment, especially to the uninitiated.”

Letters without an expression of opinion

were received from Henry B. Bigelow, A. J.

Jukes Browne, William E. Hoyle, and

Charles Wardell Stiles.

Finally, one card from Dorchester, Eng-

land, read “I am in favor of accepting Hn-

crinus Schulze, 1760... but I should prefer

a postal order for five shillings to be sent to

ZOOLOGY —On Polyclinum indicum, a

India. V. O. SEBASTIAN,! University

India. (Communicated by Fenner A.

While engaged in a study of the ascidian

fauna of Madras coast, I was able to identify

a new species of Polyclinum, the structure

of the zooid, larva, and postlarval stages of

which forms the substance of the present

paper. Herdman (1891) has described P. con-

stellatum and P. iscacum from the Indian

Ocean, and a doubtful species (1906),

P. nigrum, from the gulf of Manaar. Sebas-

tian (1942) has published an account of

the anatomy and larval organization of

Polyclinum sp. obtained from a dredge col-

lection off the coast of Madras and later

(1952) described as P. madrasensis Sebastian.

The present form, Polyclinum indicum, n.

sp., 1s the commonest synascidian found

along the rocky shores of the Madras coast.

The colonies are found attached to the under

surface of stones and boulders, at the level

of the tides, on the Royapuram coast, north

of Madras harbor. The places where they

grow are always subjected to the action of

violent waves.

Exiernal appearance-—The colonies vary in

shape, younger ones being oval or pear-shaped

with narrow bases of attachment and round up-

per exposed surfaces (Figs. 1, 2). The full-grown

colony has an umbrella-shaped upper exposed

surface, the base of attachment being broader,

but narrower than the diameter of the upper

region (Fig. 3). The mature colony has a diame-

1 | wish to express my thanks to Dr. C. P. Gna-

namuthu, director of the University Zoology Re-

search Laboratory, for his helpful suggestions,

and to the authorities of the Madras University

for varied assistance rendered.

JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

vou. 44, No. l

Timothy Scroggins, Warwick Gaol, and to

be given me when I have finished my

time.”

Since this poll was taken the principle of

prescription has been adopted by the Inter-

national Commission on Nomenclature, and

a list of nomina conservanda has been estab-

lished.

All the cards and the letters referred to

above have been mounted in a scrapbook,

which is filed in the Library of the Smith-

sonian Institution.

new ascidian from the Madras coast of

Zoology Research Laboratory, Madras,

Chace, Jr.)

ter of 2 to 21% inches, and a height of 1144 to 2

inches. The common cloacal openings are found

scattered on the exposed surface, raised on coni-

eal projections of the outer test. The surface is

encrusted with a thin layer of sand. The color is

hght brown or pale red in the majority of cases.

Rarely the color is dull green, but except for this

the anatomical features are the same. Several

colonies, large and small, could be seen closely

“applied to one another, the different-colored

colonies occurring in the same group.

The zooids are arranged in systems of about

20 to 40, three or more such systems found around

one common cloacal opening forming a pattern

(Fig. 4). The branchial openings have a whitish

color on their margins. A cross section of the

colony (Fig. 5) shows the disposition of the zooids

inside the test. They are arranged toward the

outer periphery, their long ampullae running

throughout the length of the test in various direc-

tions. The test is transparent, having a tinge of

either red or green according to the color of the

colony.

Structure of zooids—The length of the zooid

(Fig. 6) from the branchial siphon to the tip of

the postabdomen is about 2.3 to 3.2 mm. The

abdomen is about one-half and the postabdomen

three-fourths the size of the thoracic region. The

shape and proportion of the various regions of the

zooids may vary slightly according to the manner

in which each zooid is pressed into the group

forming a system. The branchial siphon is 6-lobed.

The atrial siphon is a wide space exposing a part

of the branchial sac, which is a characteristic

feature. At the anterior edge of the atrial siphon

there is an atrial languet, which is leaflike, but

JANUARY 1954 SEBASTIAN: A NEW

the distal extremity is blunt with a few protu-

berances. The size and shape of the languets vary

according to the distance of the zooid from the

common cloacal opening, those farthest being

longer and more pointed, those nearest, shorter

and blunt. The languet appears to be the en-

larged anterior lobe of the opening, the other

lobes having failed to develop to their full size.

Rarely vestiges of the other lobes are also seen

(Fig. 7). The muscle bands are six pairs on the

mantle wall, extending only up to about the

middle of the thoracic region.

The thorax is cylindrical. There are about 13

to 15 rows of stigmata, with 14 to 16 elliptical

stigmata in each row. In smaller zooids the num-

ber may vary from 12 to 14. Each transverse

vessel bears a series of minute papillae on its

inner free margin. The dorsal lamina is in the

shape of languets. The tentacles are simple vary-

ing from 12 to 16, their free ends being slightly

swollen.

The abdomen when viewed as a whole is kid-

pu

Tf)

f= AX)

Be |

=

=

=

SI

=

!

iD

=) = EaI=!|

iS!)

ASCIDIAN FROM INDIA 19

ney-shaped. There is a funnel-shaped esophagus,

followed by a midgut and rectum, the whole sys-

tem bent and twisted into a U with unequal

limbs. The rectum opens at the posterior edge of

the atrial siphon, the anal opening situated be-

tween two lobes.

The postabdomen is connected to the abdomen

by a short narrow stalk which gradually widens

to hold the gonads and heart. The testis consists

of follicles in the shape of bunches of grapes all

connected together by narrow ducts. The ovary

is found to occupy the middle portion among the

mass of sperm bunches, and shows eggs of various

sizes. The sperm duct and oviduct run together

and open near the level of the rectal opening.

Posterior to the gonads and slanting in disposi-

tion is the pericardium enclosing the heart. A

thin-walled epicardium runs along the side of the

gonads (Figs. 8, 9): its posterior extremity comes

to the middle bend of the heart.

Sexual reproduction—Eggs undergo develop-

ment inside the atrial chamber, and tadpole lar-

Fics. 1-9.—Polyclinum indicum, n.sp.: 1, 2, Young colonies; 3, mature colony; 4, surface view show-

ing the grouping of zooid systems; 5, a thin solid section of a colony; 6, mature zooid; 7, atrial siphon

with vestiges of lobes; 8, postabdomen; 9, cross section of postabdomen.

20 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

vae when mature are liberated in swarms. Breed-

ing occurs throughout the year. Colonies col-

lected and kept in troughs of sea water liberate

larvae at any time of the day. The egg measures

0.25 mm and has a yellowish tinge due to the

presence of yolk. A few stages of developing em-

bryos are shown in Figs. 10-12.

Tadpole larva.—The tadpole (Fig. 13) measures

1.66 mm from the adhesive papillae to the tip of

the tail fin, the trunk measuring 0.32 mm and

the tail 1.34 mm. The tunic covers the whole

body and tail of the larva. In the region of the

tail, the tunic is expanded into the tail fin, which

is horizontal in position owing to the rotation of

the tail (Fig. 14). Test vesicles (‘‘tunic vesicles,”

Scott ’46) are of two sets, one directed anteriorly

and the other posteriorly. The anterior ones are

club-shaped and arranged in a ring of eight in

two rows of four each, arising from the anterior

ectodermal margin of the body of the larva, and

ZLLL DCE say

ASE:

3

~ ;

RNID Ren

VOL. 44, No. 1

spreading out with a slant toward the dorsal side.

The middle two pairs are longer, measuring 63.2,

and the lateral ones smaller, measuring 47.4y.

The posterior test vesicles are in the nature of

bunches of grapes, one set dorsal and one ven-

tral in position, with long narrow tubular stalks

from which arise pinnately arranged branches

ending in round hollow vesicles containing mesen-

chyme cells. They are of ectodermal origin, the

stalks arising from the anterior margin of the

trunk, from their respective dorsal and ventral

sides, at the level of the origin of the anterior test

vesicles. Adhesive papillae are three in number

and arise from the anterior ectodermal margin

of the trunk between the ampullae. Hach one

of them has a long narrow tubular stalk, the dis-

tal extremity of which swells into a goblet con-

taining a central mass of columnar secretory

cells, all converging to a point in the central open-

ing (Fig. 19). Grave (1921) has stated that the

Fias. 10-15.—Polyclinum indicum, n.sp.: 10-12, Developing embryos; 13, tadpole larva, side view;

14, tadpole larva, dorsal view; 15, Transverse section of tail of larva.

JANUARY 1954 SEBASTIAN: A NEW

secretory cells of the adhesive papillae in Amarou-

cium constellatum are modified mesenchyme cells.

Sebastian (1942) has found them to be of ecto-

dermal origin jn Polyclinwm sp. (P. madrasensis),

formed by the intucking and elongation of the

ectodermal cells of the papillae, similar to the

condition found in the cement glands of certain

fishes and amphibian larvae (Asheton, 1896;

Jones, 1937; Bhaduri, 1935). Scott (1946), study-

ing the larval organization of A. constellatum, has

also found that these cells are of ectodermal ori-

gin. The same kind of ectodermal origin of the

secretory cells of the adhesive papillae in P. indt-

cum is illustrated in Figs. 16-19. The mantle or

ectodermal covering of the body and tail is made

up of one layer of cells with distinct nuclei, con-

taining a large number of yolk granules. The

layer covering the trunk has cubical cells which

gradually become thinner and flattened in the

00602

sieteanst

ASCIDIAN FROM INDIA el

tail region. In the region of the branchial and

atrial siphons these cells are columnar.

The nervous system consists of the sensory vesi-

cle with the contained ocellus and otolith, visceral

ganglion with the visceral nerve, and nerve cord,

of the larval action system, and the permanent

ganglion and hypophysial duct of the adult system

(Figs. 20, 21). The sensory vesicle is situated be-

tween the branchial and atrial siphons to the

right side. The ocellus consists of three lens cells,

pigmented optic cup and associated retinal cells.

The otolith is single-celled with a_ perfectly

spherical pigmented mass at its distal end.

The digestive tract (Fig. 13) is a bent tube in-

cluding the pharynx, esophagus, intestine, and a

short rectum, which ends blindly near the level

of the esophagus. There is a middle mass of yolky

cells, conical in shape, the narrow portion being

connected with the endodermal layer by a short

S50

one

Fies. 16-24.—Polyclinum indicum, n.sp.: 16-18, Stages in the formation of the secretory cells of ad-

hesive papillae; 19, goblet magnified; 20, sagittal section through the region of the sensory vesicle; 21,

section passing through the visceral ganglion; 22, Transverse section of yolky cell mass; 23, magnified

view of the posterior extremity of tail of larva; 24, secretory cells of the goblet being shot out.

Zp JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

stalk. In transverse sections. it is found to be

formed of two squarish portions (Fig. 22), each

having a narrow central cavity. The endostyle is

placed on the anterior margin of the pharynx,

one edge of it touching the anterior edge of the

yolky mass. The pharynx has two rows of stig-

mata on each side, each row having about eight

stigmata. On the ventral side of the yolky mass

toward the anterior side is situated the peri-

cardvum and heart.

The notochord forms the central core of the

tail (Fig. 15); it has 40 cells placed one behind

another in a row. In the full-grown larva the

boundaries of notochordal cells are not clearly

seen. Owing to the twist of the tail at an angle of

90° to the left the nerve cord is found on the left

and the endodermal strand on the right side of

the notochord. The muscle bands situated dor

sally and ventrally do not extend up to the pos-

terior extremity of the notochord (Fig. 23). Each

band is formed of three rows of muscle cells, each

row of seven cells, placed one behind the other.

There are only two rows of cells on the portion of

the notochord. which projects into the body. The

muscle cell has a darkly staining cortex, and an

inner vacuolated core with cytoplasmic strands

and nucleus. The disposition of the striations is

oblique, as described by Grave (1921), Conklin

28A

VoL. 44, No. 1

(1931), Scott (1946), and Berrill (1947). The

ectoderm of the tail forms the outer covering of

the muscles.

Metamorphosis of the larva.—The free-swim-

ming period lasts 5 to 8 hours. At the time of

fixation the secretory cells of the adhesive pa-

pillae shoot out of the goblet, exuding their

secretory product (Figs. 24, 25). The tail gets

resorbed, and the anterior ampullae elongate and

spread out in an irregular way (Figs. 25-27), at-

taining their maximum length of 0.24 mm in 2

hours after the tadpole has become fixed. The

heart begins to beat after 3 hours, the siphons

contract after the fifth hour, and the intestine

functions 10 hours after fixation. Complete re-

sorption of the tail does not take place in most

of the cases during the changes described above.

At least a quarter of the original length, and in

some cases even half, of the tail remains un-

changed for about 12 hours, and in certain cases

for more than 24 hours. The posterior ampullae

do not disappear immediately after metamorpho-

sis. They enlarge in size and remain all over the

surface within the tunic.as pyramidal projections

(Fig. 28a, b); their walls are made up of a very

thin unicellular layer. The ducts that connect

them are not found during this time. These vesi-

cles are clearly visible for a week or more, later

Fras. 25-28.—Polyclinum indicum, n. sp.: 25-27, Stages in the metamorphosis of tadpole larva; 28a, a

functional oozooid, 48 hours old; 28b, magnified view of a single posterior test vesicle.

JANUARY 1954 SEBASTIAN: A NEW

disappearing by bursting and releasing the mesen-

chyme cells lodged inside them. Under labora-

tory conditions it has not been possible to keep

alive the metamorphosed stages for more than 10

to 13 days; the size attained in 4 to 5 days re-

mains without any change for the rest of the

days. The abdomen descends partially (Fig. 28a),

but the descent of the postabdomen has not been

observed.

Validity of the Species——Zooids as well as lar-

vae of Policlinum are alike, differentiating char-

~ acters being minor. Berrill (1950) remarks that

‘tn zooid structure, though less so in form of

colony, species of Policlinwm are very much alike,

and even the tadpoles and larvae appear to differ

only in size.”’ Van Name (1945) says that ‘‘the

true Polyclinum are all very closely related to

each other, their zooids apparently having nearly

the same structure, so that we must depend

chiefly on the gross characters of the colonies for

distinguishing them. A supposed difference in the

number of branchial tentacles (whether in multi-

ples of 4 or 6) appears to be of questionable value

as a specific character, multiples of 6 being prob-

ably normal, though subject to irregularities.”

The present species closely resembles P. planum

in shape and number and rows of stigmata, but

it differs in the shape of the atrial siphon, the

atrial languet and the slanting disposition of the

heart. On account of these differences the present

form is given the status of a new species, P. in-

dicum.

LITERATURE CITED

ASSHETON, R. Notes on the ciliation of ectoderm of

the amphibian embryo. Quart. Journ. Micr. Sci.

38: 465-484. 1896.

BerRrRiLL, N. J. Metamorphosis in ascidians. Journ.

Morph. 81: 1947.

. The Tunicata, 354 pp. Ray Society, London

1950.

Buavpurt, J. L. The anatomy of the adhesive appara-

tus in the tadpoles of Rana afghana Ginther,

ASCIDIAN FROM INDIA 23

with special reference to the adaptive modifica-

tions. Trans. Roy. Soc. Edinburgh, 58: 339-349.

1935.

GRAVE, C. Amaroucium constellatum (Verrill): LJ,

The structure and organization of the tadpole

larvae. Journ. Morph. 36: 71-91. 1921.

HERDMAN, W. A. A revised classification of the

Tunicata, with definitions of the orders, sub-

orders, families, subfamilies, and genera, and

analytical keys to the species. Journ. Linn. Soc.

London, Zool., 28: 558-652. 1891.

. Report on the Tunicata collected by Prof.

Herdman in 1902. Rep. Pearl Oyster Fisheries

5: 295-348, 9 pls. 1906.

JONES, S. On the origin and development of the

cement glands in Etroplus maculatus. Proc.

Indian Acad. Sci., Sec. B 6 (4): 251-261.

1937.

SEBASTIAN, V. O. On the anatomy and larval or-

ganisation of Polyclinum sp. Journ. Madras

University 14 (2): 251-278. 1942.

. A new species of synascidian from Madras.

Current Science 21: 316-317. 1952.

Scorr, F. M. The development history of Amarou-

cium constellatum. JJ. Organogenesis of the

larval action system. Biol. Bull. 91: 66-80.

1946.

KEY TO LETTERING ON FIGURES

AB, abdomen; ADP, adhesive papilla; AL, atrial

languet; AMP, ampulla; AN, anal opening; AS,

atrial siphon; ATV, anterior test vesicle; BS,

branchial siphon; BRS, branchial sac; CCL, com-

mon cloacal opening; ECT, ectoderm; EN, endo-

style; EN.ST, endodermal strand; EP, epicar-

dium; HT, heart; LC, lens cell; MB, muscle

band; MES, mesenchyme cells; MG, midgut;

MUS, tail muscle; NC, nerve cord; NHC, neuro-

hypophysial canal; NT, notochord; O, ovary;

OES, oesophagus; OT, otolith; PAB, postabdo-

men; PC, pigmented optic cup; PGN, permanent

ganglion; PH, pharynx; PHW, pharyngeal wall;

PTV, posterior test vesicle; R, rectum; RC,

retinal cell; SC, secretory cells; SP, sperm duct;

ST stomach; SV, sensory vesicle; T, testis; TF,

tail fin; TN, tentacle; TU, tunic; VGN, visceral

ganglion; VN, visceral nerve; YC, yolky cells

connected with the growth of the postabdomen;

YK, yolk granules.

24 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

vou. 44, No. 1

HELMINTHOLOGY .—Psilocollaris indicus, n.gen., n.sp. (Psilostomidae Odhner,

1911: Trematoda) from an Indian stork, Dissoura e. episcopus. KUNWAR SURESH

SincH, Department of Zoology, Lucknow University. (Communicated by —

Robert Rausch.)

Seven specimens of an undescribed trema-

tode were recovered from the intestine of a

stork, Dissoura e. episcopus (Boddaert), shot

near Lucknow, India, in August 1950. The

trematodes were pink in color when alive

and were capable of active movement.

Seven genera have been described in the

family Psilostomidae: Psilostomuwm Looss,

1899; Psilochasmus Liihe, 1909; Apophar-

ynx Lithe, 1909; Sphaeridiotrema Odhner,

1913; Psilotrema Odhner, 1913; Lyperorchis

Travassos, 1921; Pszlorchis Thapar and Lal,

1935. The present form differs from all these

genera in the possession of a definite collar

at the anterior end of the body. It is most

closely related to the genera Lyperorchis and

Psilorchs.

For Lyperorchis, described from the cloaca

of Aramus scolopaceus, a slightly developed

collar was described by Travassos: ‘‘Ventosa

oral ladeada por duas saliencias papillifor-

mis.” In the present form, however, the

collar is well developed, and clearly of the

Echinostome type. Indeed if spines were

present, these forms would be regarded as

typical Echinostomes.

Further, the testes in Lyperorchis are

sinuous and much elongated whereas in the

present form they are typical of the family

(i.e., bean-shaped). The oral sucker and the

pharynx in the present form are poorly de-

veloped as compared to those of Lyperorchis

and other genera. The arms of the Y-shaped

excretory bladder are very much elongated

and are longer than the stem, extending to

the anterior region of the anterior testis. In

the present forms the excretory bladder is

Y-shaped but the arms are comparatively

very small. In Pszlorchis the collar is entirely

lacking, the oral sucker and the pharynx are

comparatively well developed and large in

size, and the stem of the excretory bladder

is very small and the arms comparatively

very long.

Accordingly a new genus is created for

these forms, with the following generic

diagnosis:

Psilocollaris, n. gen.

Psilostomidae: Body much elongated; oral —

sucker and pharynx weakly developed, pre-

pharynx small and oesophagus very long, in-

testinal ceca extend to posterior end of body;

slightly developed unarmed collar present at the

anterior end; ventral sucker well developed and

present in the anterior bodyhalf; testes bean-

shaped or oval, and situated one behind the

other in the posterior region; cirrus pouch small

and partly covered by ventral sucker; genital

pore just posterior to intestinal bifurcation;

small, rounded ovary in posterior half of body;

vitellaria consist of many follicles laterally dis-

tributed from ovary into posterior end, though

a clear midarea is usually present but the vitel-

laria may be confluent in regions; uterus with

ascending limb only containing many eggs.

Parasites of birds.

Type spectes.—Psilocollaris indicus, n. sp.

Psilocollaris indicus, n. sp.

Figs. 1-3

Diagnosis.—Body measures 5.3-16.4 mm in

length and 0.384—0.69 mm in maximum breadth.

Anterior end is provided with slightly developed

collar, not much unlike the collar of an Echino-

stome, but spines are absent. When viewed from

the side the collar appears as a flaplike process

(Fig. 2). Collar measures 0.26-0.3 mm across

and is not muscular. Oral sucker and the pharynx

are comparatively very small and poorly de-

veloped. The oral sucker measures 0.506 x

0.024 mm and the mouth opening is very small,

sub-terminal, and leads into a small prepharynx.

The pharynx, which is larger than the oral

sucker, measures 0.069-0.1 x_ 0.040.046 mm.

The oesophagus is very long, measuring 0.88-

2.0 mm in length. It divides into two intestinal

ceca which run laterally to the posterior end of

the body, but their extreme terminations could

not be observed because of the presence of large

number of vitelline follicles. Ventral sucker is

comparatively very large and muscular and is

situated posterior to the intestinal bifurcation in

the anterior fourth of body. The sucker is trans-

versely elongated, and measures 0.27-0.4 x

0.26-0.4 mm. The body is without spines.

JANUARY 1954

The excretory bladder is Y-shaped and the

arms are rather wide (Fig. 3). The stem of the

bladder extends medially to the posterior end

of the posterior testis and the arms are short and

do not extend beyond the base of the posterior

testis. The excretory pore is situated at the pos-

terior end and is terminal.

The two testes are present in posterior fourth

region of body, one behind the other. The

anterior testis, usually slightly smaller than

posterior testis, measures 0.43-0.72 x 0.19-0.34

mm. The posterior testis situated about 0.2

mm. posterior to anterior testis, is 0.43-0.83 x

0.21-0.32 mm. Both testes are beanshaped.

The cirrus pouch is comparatively small and

lies in between the ventral sucker and the

intestinal bifurcation. It is somewhat oval and

measures 0.16-0.26 x 0.069-0.14 mm. External

vesicula seminalis is absent, but a large internal

vesicula seminalis is present. Numerous prostate

gland cells surround the ejaculatory duct. The

genital pore is just posterior to the intestinal

bifurcation. Ovary is small and rounded and

situated in posterior half of body, anterior to the

anterior testis. It measures 0.12-0.184 x 0.099-

0.16 mm. Mehlis’s gland is slightly smaller

than ovary and is present just posterior to it.

It is almost rounded and measures 0.13-0.18

x 0.18-0.19 mm. The vitellaria consist of nu-

merous elliptical follicles distributed mainly

0.2 mm !

a2

A ee Bt Is aE

eae &

~

Sicy =P

se

“

bothek tee

eS Seige @ wie

2a rasta Roa = ae

SESsSan tee sae

Ne ey MINS ESAS GR Baten

Petals Daa rien ety

ae? ELS FS oe

SINGH: PSILOCOLLARIS INDICUS

erate, 3 A exe

sess ca

2S

‘

~_

5)

laterally from level of the ovary to the posterior

end of the body. In most specimens, the vitellaria

leave a clear space in the middle, but sometimes

they are confluent anterior to the anterior testis.

Two transverse vitelline ducts meet in the

region of Mebhlis’s gland and form a_ small

vitelline reservoir. Uterus consists of an ascending

limb only and after several folds immediately

anterior to the ovary it proceeds more or less

as a straight tube, opening at the genital pore

posterior to the intestinal bifurcation. The eggs

are numerous and measure 0.092-0.104 x 0.058

mm,

Host.—Dissoura e. episcopus (Boddaert).

Locality.—Lucknow, India.

Habitat.—Intestine.

Remarks.—The genus Psilorchis contains three

species, all described from India: P. indicus

Thapar and Lal, 1935; P. ajgainis Lal, 1938,

and P. thapari Baugh, 1949. In P. indicus the

cirrus pouch is present in front of the ventral

sucker but in P. ajgainis the cirrus pouch is

described as adhering to the ventral sucker and

this character has been used to distinguish the

two species (Lal, 1938; Lal, 1939). Consequently,

the diagnosis of the genus Psilorchis should be

emended and should read ‘‘Vesicula seminalis

and cirrus pouch retort-shaped, partially or

totally in front of the ventral sucker...”

Further, the genital pore in P. ajgainis is de-

FG

iS Paes 6

Ba eet 2%

ROPES

pee g Mase RISES.

SE OR Spee

a eh

Fics. 1-3.—Psilocollaris indicus, n.g.,n. sp.: 1, Entire worm; 2, collar, lateral view; 3, posterior end,

showing excretory bladder.

26 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

scribed as ‘‘in front of the oral sucker, between

it and the intestinal bifureation.” (Lal, 1938, p.

260) An obviously typographical error occurs

here.

The excretory system in the family Psilo-

stomidae has been described as ‘‘comprising a

subcutaneous net-work of vessels and two main

lateral canals which unite in front of the ventral

sucker to form a long median stem” (Dawes,

1946, p. 99). Unfortunately the details of the

excretory system of many of the genera are not

known, but in genera like Pstlostomum, Ly-

perorchis, Psilorchis, Apopharynx and the present

form, the excretory bladder is Y-shaped, the

comparative length of the stem and the main

branches varying from genus to genus, but in all

cases the median stem extends to the base of the

posterior testis only. It certainly does not extend

anterior to the ventral sucker, as stated by

Dawes (1946). Indeed, in the two species of

Psilorchis, the Y-shaped excretory system is

very small. Beaver (1939), who investigated the

life history of Psilostomum ondatrae_ Price,

1931, found that the excretory bladder in the

cercaria is not elongated and is confined to the

posterior end, posterior to the genital anlagen

and the ventral sucker. It retains the same

comparative position in the adult, though the

median stem becomes elongated and the ex-

cretory system looks typically Y-shaped. Hence,

in the opinion of the author the excretory ‘system

in the family Psilostomidae should be described

as Y-shaped, without limiting the length of the

median stem.

The families Psilostomidae and Echinostomi-

dae are closely related and there seems little

vou. 44, No. l

difference between the two except for the presence

of a collar with spines in the family Echino-

stomidae. The resemblance is not confined to

the adult characters only, since Beaver (1939)

found that the miracidia, redia and cercaria of

Psilostomes closely resemble those of Echino-

stomes and the pattern of the life history is the

same. In view of such evidence, the presence of a

genus with a collar, but without spines, is

especially interesting as it joins the two families

and may be considered an intermediate form in

the evolution of the families Psilostomidae and

Echinostomidae.

REFERENCES

Bauau, 8. C. On a new avian trematode, Psilorchis

thapari (Fam. Psilostomidae), with a record

of Psilochasmus oxyurus (Crep.) from India.

Ind. Journ. Helminth. 1: 79-84. 1949.

Beaver, P. C. The morphology and life history of

Psilostomum ondatrae Price, 1931 (Trema-

toda: Psilostomidae). Journ. Parasit. 25:

383-393. 1939.

Dawes, B. The Trematoda. Cambridge Univ.

Press, 644 pp. 1946.

Lau, M. B. On a new species of Psilorchis from the

intestine of the common teal, Nettion crecca.

Livro Jub. Prof. Travassos : 259-262. 1938.

. Studies in helminthology. Trematode para-

sites of birds. Proc. Ind. Acad. Sci., Sec. B.,

10: 111-200. 1989.

Lune, M. Die Stisswasser-Fauna Deutschlands 17:

1-217. 1909.

TuHapar, G.S., and Lat, M. B: On the morphology

of a new genus of trematode parasite from the

kingfisher from Lucknow. Proc. Ind. Acad.

Sci., Sec. B., 2: 88-94. 1935.

Travassos, L. Fauna helminthologica de Matto

Grosso. Mem. Inst. Oswaldo Cruz 21: 309-341.

1928.

MALACOLOGY.—Hydrobia totteni, new name for Turbo minuta Totten, 1834

(Gastropoda: Hydrobiidae). J. P. E. Morrison, U. 8. National Museum.

The species name still in use for the com-

monest New England salt-marsh-inhabiting

hydrobiud snail (Turbo minuta Totten, Amer.

Journ. Sci. 26 (2): 369, fig. 6. July 1834)

was originally thrice preoccupied, by 7’. mi-

nuta Brown, 1818; JT. minuta Michaud,

1828; and T. minuta Woodward, 1833. This

species has been so ‘‘well-known”’ that no

one has given any alternative name, even

after Sherborn (Index Animalium 1801-

1850, p. 4101. 1928) listed the homonyms.

Since there is no available name known to

me, I hereby rename the species Hydrobia

tottent in honor of its first describer.

Examination of the animals has shown

this species to be a true Hydrobia, congeneric

mm sensu stricto with the genotype of Hy-

drobhia Hartmann, 1821, namely Turbo

stagnalis Baster, 1765.

JANUARY 1954 NICOL:

PELECYPOD CLASSIFICATION 27

MALACOLOGY .—Trends and problems in pelecypod classification (the super-

generic categories). Davip Nicou, U. 8. National Museum.

Little has been written to guide workers

in the various animal phyla in problems of

classification in the supergeneric categories.

Simpson (1945, pp. 20-24) and Mayr, Lins-

ley, and Usinger (1953, pp. 46-59) have

contributed valuable suggestions; but gen-

eral principles of standardization that could

be applied to the higher categories of all

phyla are still lacking. The reason is that

many of the classification problems of the

entomologist or mammalogist, for example,

are entirely different from those of the

malacologist. From the standpoint of size

alone, if not on a morphologic basis, a pele-

eypod family or other supergeneric category

may not be comparable with a family of

insects or mammals; this in itself would tend

to create different problems in classification

in the Insecta, the Pelecypoda, and the

Mammalia. Other differences in higher cate-

gories of different classes of animals arise

from the percentage of described species

and the phylogenetic knowledge in the

various groups. Hence, standardization at

the level of the phylum would be virtually

impossible; in fact, about all that can be

hoped for in a large phylum is reasonable

standardization within a class.

For the purposes of brevity and continuity

in this paper, I have decided to treat the

Pelecypoda as a class. This has been the

most common treatment in the past, al-

though I realize that at least some of our

problems might be solved if the Pelecypoda

were given the rank of, let us say, a sub-

phylum...

It appears to me that the phylogenetic

relationships of the Pelecypoda are not

adequately or correctly shown by the pres-

ent classifications; but, with the excellent

fossil record of the pelecypods, these rela-

tionships could be shown and thus the use-

fulness of the classification could be im-

proved.

The ideas and problems presented here

are mainly those of a malacologist; however,

many valuable suggestions and criticisms

were given by Dr. R. E. Blackwelder,

entomologist at the U.S. National Museum,

and for these I am very grateful.

The purpose of this paper is to examine

some recent examples of classification of

pelecypods used by paleontologists and

neontologists, to discuss present trends, and

to offer possible solutions to some of the

problems. A list of some important works on

pelecypod classification is included.

THE SUBCLASS, SUPERORDER,

SUBORDER

ORDER,

The first example of classification is

taken from the paleontology textbook Jn-

vertebrate fossils by Moore, Lalicker, and

Fischer (1952). Moore, who wrote the chap-

ter on the pelecypods, used a modified

version of Dall’s classification based on the

hinge teeth. He divides the pelecypods

(pp. 409-412) into two subclasses—Prio-

nodesmacea and Teleodesmacea. In the

first group Moore includes five orders and

twelve suborders; in the second, in which

he combines Dall’s orders Anomalodesmacea

and Teleodesmacea, he includes three orders

and twelve suborders. According to Moore,

the Paleoconcha are an order of the subclass

Prionodesmacea; Dall, however, did not

include them in any of his three orders in

1895 (p. 513) but set them aside as incertae

sedis, although he later arbitrarily placed

them in the Prionodesmacea.

Moore discusses the structure of the

pelecypod ctenidia at considerable length

but gives the ctenidia little importance in

his classification—i.e., in the Prionodes-

macea, Moore includes pelecypods with

protobranch, filibranch, and eulamelli-

branch ctenidia. Furthermore, Moore places

the anomiids and the spondylids together

in the order Isodonta and places the nucu-

lids and arcids together in the order Taxo-

donta; but on the basis of phylogeny and

- morphology it is difficult to see these rela-

tionships. In the order Dysodonta, Moore

includes the mytilids, pectinids, pinnids,

ostreids, limids, and dreissensuds—truly

a heterogeneous assemblage.

28 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

The classification used by Shrock and

Twenhofel in their book (1953, pp. 386-393)

is that of Thiele, 1934, who divided the

Pelecypoda into three orders, Taxodonta,

Anisomyaria, and Hulamellibranechia, pri-

marily on the basis of ctenidia, hinge teeth,

and adductor muscles. In the order Taxo-

donta are included the nuculids and the

arcids. However, in a footnote (p. 389)

Shrock and Twenhofel make the following

statement:

The Areazea are now regarded by some in-

vestigators (MacNeil, 1937; Nicol, 1950) as more

closely related to certain groups of the Aniso-

myaria because the hinge structure is of a later

type and may possibly be the result of converg-

ence. It may well be, therefore, that this super-

family should be transferred to the order Aniso-

myaria or used as the basis for a new order.

Shrock and Twenhofel imply that the lack

of close relationship between the arcids and

the nuculids is a new idea, but if the authors —

had examined the writings of Pelseneer and

Douvillé they would have found that the

idea was presented much earher than 1937.

Furthermore, if my paper of 1950 had been

carefully read (p. 89), the following state-

ment would have been noted:

Pelseneer, Douvillé, and others have pointed

out the fact that the prionodonts are not closely

related to the true taxodonts such as Nucula,

Nuculana, and Yoldia. .

This is but one example of the lack of

knowledge of the basic works on pelecypod

systematics.

A recent and much more comprehensive

work is the T'razté de paléontologie edited by

Jean Piveteau, 1952. The major portion

of the chapter on Pelecypoda was written

by Colette Dechaseaux. The main divisions

Dechaseaux uses are based on hinge char-

acteristics, and she divides the pelecypods

into four orders—Taxodonta (with three

suborders), Dysodonta, Preheterodonta, and

Heterodonta. Only the taxodonts are di-

vided into suborders, one of which is the

Paleoconcha. By far the largest order is the

Heterodonta, which is comprised of more

than half of the pelecypod families.

Dechaseaux’s classification bears little

resemblance to her schematic table on the

evolution of the pelecypods (page 229). This

vou. 44, No. 1

is Just one example of workers who, although

cognizant of phylogenetic evidence, do not

base their classification on phylogeny but,

instead, follow the line of least resistance by

using the outmoded classification of a prede-

cessor. As long as this attitude occurs, little

progress can be made in the classification

of the pelecypods.

Another work worth analyzing is that of

T. Habe (1951-1953), who uses Dall’s classi-

fication with few modifications. Habe di-

vides the Pelecypoda into three subclasses—

Prionodesmacea, Teleodesmacea, and Anom-

alodesmacea. The first subclass is di-

vided into four orders, and the Teleodes-

macea is divided into two orders. Habe

does not group the Anomalodesmacea: above

the level of superfamily. The categories

superorder and suborder he does not use at

all in his classification of the pelecypods.

Habe places the nuculids and arcids in

the order Taxodonta, although, as has been

pointed out before, they bear only a super-

ficial resemblance to each other. Further-

more, although most of the Cenozoic arcids

and nuculids do have similar hinges, the

hinge teeth of the Mesozoic and Paleozoic

arcids and their allies are generally quite

unlike those of the nuculids. Thus, from

the practical standpoint of classification,

disregarding phylogeny, this grouping is not

workable.

Little attempt is made by Habe to show

relationships in the order Heterodonta, to

which he assigns 14 superfamilies and 34

families. :

The comprehensive treatment of the

classification of the Pelecypoda by Cotton

and Godfrey (The molluscs of South Aus-

tralia, 1938) is also noteworthy. These

authors subdivide the class into the three

orders proposed by Dall. The order Priono-

desmacea is subdivided into five suborders—

Palaeoconcha, Taxodonta, Schizodonta, Iso-

donta, and Dysodonta—which are in turn

divided into superfamilies and _ families.

The order Anomalodesmacea is not grouped

above the level of superfamily; however,

the authors use the category ‘‘section”’ be-

tween the superfamily and family. Cotton

and Godfrey subdivide the order Teleodes-

macea into five suborders—Pantodonta,

Diogenodonta, Cyclodonta, Teleodonta,

JANUARY 1954

and Asthenodonta—and these suborders

are further divided into numerous super-

families.

Once again we find the unlike nuculids

and arcids grouped together in the suborder

Taxodonta. Cotton and Godfrey place the

pteriids, ostreids, unionids, and _ trigoniids

in the suborder Schizodonta; certainly the

ostreids do not belong with such primitive

nacreous groups. The grouping of the ven-

erids, tellinids, solenids, and mactrids in

the suborder Teleodonta seems arbitrary

and appears to be based on little or no

phylogenetic and morphologic evidence.

All the foregoing examples show certain

common characteristics which are important

as well as interesting. They are as follows:

1. In none of the treatments reviewed of

the classification of the class Pelecypoda are

all the common categories used, 1.e., sub-

class, superorder, order, suborder, super-

family, family, and subfamily.

2. None of the classifications is basically

new. With one exception, each author fol-

lows one authority almost exclusively with

perhaps minor modifications; the one ex-

ception uses a combination of basic char-

acters and classifications.

The fundamental concepts for classifying

the Pelecypoda were mainly promulgated

between the years 1889 and 1912. It was

during this period that the morphologists,

embryologists, and evolutionists were most

intensively working on natural, or phylo-

genetic, classifications of the Pelecypoda.

Since that time only a few details of classi-

fication have been added. Even thorough

review and synthesis of the classifications

have received little interest lately. This

basic pattern of the development of classi-

fication may have counterparts in other

groups of animals.

Indifference to the classification of the

Pelecypoda began in 1913 and has con-

tinued for 40 years since. As a result, lack of

knowledge of the basic works on the subject

is continually being exhibited. One solution

to our present state of stagnation is to re-

examine the ‘‘classics’”’ on pelecypod classi-

fication—papers by Neumayr, Dall, Pelse-

neer, Bernard, Jackson, and Douvillé. Each

classification and set of facts on morphology,

NICOL: PELECYPOD CLASSIFICATION 29

embryology (including growth stages of the

Shell), and paleontology should be thor-

oughly studied and the evidence evaluated.

(Douvillé’s classification might have been

more widely accepted if he had assigned

definite categories for his three-fold division

of the Pelecypoda.) Incorrect data and

conclusions should be deleted.

3. Little or no attention is paid to phy-

logeny in classification even when the evi-

dence is clear and the author is aware of it.

This has led to serious errors in classifica-

tion from the standpoint of practical mor-

phology as well as phylogeny. For the past

40 years work on phylogeny has been con-

sidered relatively unimportant and unre-

warding; however, at the ordinal level of

pelecypod classification a careful analysis

and synthesis of the classical work on

pelecypods is our first need. Further work is

needed on pelecypod morphology and shell

structure, including more anatomical work

on the soft parts. Further studies on the

nepionic and later stages of the shell are

also needed. The greatest lack, and probably

the most fruitful line of investigation, is

careful work on Triassic and Paleozoic

pelecypods, for this work would lead to a

better understanding of the relations of the

varlous major groups of pelecypods.

THE SUPERFAMILY, FAMILY, SUBFAMILY

The superfamilies, families, and sub-

families have been undergoing some changes

in number and scope within the past quarter

of a century. The changes have been brought

about slowly by the great increase in num-

ber of proposed genera and subgenera. The

result has been for malacologists to group

genera into new subfamilies, families, and

superfamilies by redefining and restricting

them. Two examples of this occurrence

should show the involved problems.

Frizzell (1936) raised the family Veneridae

to the rank of a superfamily and excluded

the petricolaceans and glaucomyaceans from

the Veneracea. The superfamily was then

subdivided into nine families, and two of the

families were further subdivided into sub-

families. Since 1936 other workers have

erected more genera and subgenera of

veneraceans, and the total 1s now about 200.

30 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

Recently Keen (1951) downgraded the

Veneracea to the rank of a family, in which

eleven subfamilies were included. In a more

recent paper Tremlett (1953) followed

Kkeen’s classification and made the following

comments (p. 1):

D. L. Frizzell (1936), in one of the most recent

works on this group, has suggested that they

should be regarded as a superfamily Veneracea,

with the same limits approximately as the family

Veneridae as the term was used by Dall (1903),

Jukes-Browne (1914), Palmer (1927), and others.

I cannot see the advantage in raising the status of

the group which is thereby separated from the

closely related Petricolidae, and also from the

Oncophoridae which are probably related to it;

furthermore it unnecessarily increases the number

of superfamilies. Even though the Veneridae are

one of the largest families of pelecypods, the char-

acters defining it are of about the same importance

as those defining other families. Frizzell’s families

obviously have close similarities, and I prefer to

regard them as subfamilies and retain the term

Veneridae in its old sense.

The 11 subfamilies included by Keen in

the Veneridae do not all have the same

morphologic distinctness. How can _ these

subfamilies be grouped to show the relation-

ships? One solution that has been adopted

is the one taken by Frizzell. Certainly the

commonly used categories are available,

and to raise the rank of several of the larger

pelecypod families to the rank of' super-

family would not create chaos in the classi-

fication. This course of action would prob-

ably be most acceptable to the malacologists

and paleontologists. However, another solu-

tion is possible, if, as Tremlett claims, the

morphologic characters defining the Vener-

idae are equal to, or of the same importance

as, the morphologic characters defining other

pelecypod families. This solution is to insert

additional categories between the subfamily

and the genus—for examples, the categories

tribe and subtribe. The entomologists have

vou. 44, No. 1

done this for classifying many of the large

families of insects.

Three ways of classifying a part of the

veneraceans are shown on Table 1.

To add to the difficulties of an already

large family, there are undoubtedly some

aberrant groups which are venerids or

veneraceans. I have considered the genus

Euloxa a veneracean; but in order to fit it

into the classification, I used Frizzell’s

arrangement, considered the Chionidae as —

a family, and subdivided the Chionidae into

two subfamilies—the Chioninae and Eu- —

loxinae (Nicol, 1953, p. 60). This type of

problem was also encountered in the genus

Pliocardia. Once again I (1953a) used Friz-

zell’s classification in order to show the

systematic position of the genus. In each

of these cases the only other reasonable

solution would have been to create a cate-

gory, such as tribe, between the subfamily

and the genus.

It is true that some of the pelecypod

genera and families have been split un-

reasonably (e.g., the genus Jnoceramus);

but the veneraceans do not appear to have

received such disproportionate treatment,

at least at the generic level, and Keen dis-

agrees not with the number of groups desig-

nated by Frizzell, but with the rank to

which he assigns them.

Furthermore, one of Tremlett’s main ob-

jections to Frizzell’s classification—namely

that the morphologic characters defining the

Veneridae are of about the same importance

as those defining other families—apparently

overlooks the fact that much of our classifi-

fication of the pelecypods is based on the

size of the group in question rather than on

morphologic differentiation. For example,

on the basis of morphologic characters the

Cretaceous genera Pseudocucullaea and

Lopatinia are quite distinct from all other

prionodont genera; but, as they have few

TABLE 1.—THREE WAYS OF CLASSIFYING A PART OF THE VENERACEANS.

Frizzell, 1936

Keen, 1951

Another proposed solution

Superfamily Veneracea

Family Meretrecidae

Subfamily Meretrecinae

Subfamily Pitarinae

Family Veneridae

Subfamily Meretrecinae

Subfamily Pitarinae

Family Veneridae

Subfamily Meretrecinae

Tribe Meretrecini

Tribe Pitarini

JANUARY 1954

species, they have not been placed in a

separate subfamily or family. The Glycy-

meridae, on the other hand, although no

more distinct morphologically than Pseudo-

cucullaea or Lopatinia, have approximately

700 described species, ranging from the

Cretaceous to the Recent, and have there-

fore been classified as a family. Although I

do not assert that rank should be based

upon size, it is nevertheless true that in

many cases size has apparently been the

decisive factor, and Tremlett’s attitude is

not realistic.

However, my objection to the ideas of

Keen and Tremlett is not primarily that

the rank should be Veneracea rather than

Veneridae, but that their classification does

not allow for enough categories to show

adequately the relationships among the

200 genera and subgenera of the group.

A comparable situation is present in the

arcaceans. The latest classification (Frizzell,

1946, p. 41) raises the rank of the family

Arcidae to a superfamily, in which two

families are included, the Arcidae and the

Noetiidae. Of these, the first is subdivided

into three subfamilies, and the second into

two subfamilies. If Frizzell’s arrangement is

compared with the conservative arrange-

ment of Reinhart (1935, pp. 11-12), one is

astounded. Reinhart divides the Arcidae into

three subfamilies—Arcinae, Anadarinae, and

Noetiinae. The subfamily Litharcinae of

Frizzell is relegated to the rank of a sub-

genus of Arca by Reinhart. Although I have

found no published objection to Frizzell’s

arrangement of the Arcacea, objections

similar to those of his classification of the

Veneracea could, and probably will, be

raised in the future. My preference for

Frizzell’s treatment of both the veneraceans

and the arcaceans over more conservative

classifications is that it has more categories

in which to show more morphologic and

phylogenetic relationships. Whether all the

relationships as shown by Frizzell’s classi-

fications are correct or not is a matter to be

investigated further.

Blackwelder (personal communication)

has suggested to.me the most objective and

probably only satisfactory way of solving

the type of problem exemplified by the

NICOL: PELECYPOD CLASSIFICATION 3]

Veneridae versus the Veneracea and the

Arcidae versus the Arcacea. The genera of

the family or superfamily being studied

should be examined for morphologic simi-

larities and inferred phylogenies. ‘These

genera can then be grouped, and the groups

can likewise be grouped in a series of ascend-

ing categories. The number of categories

necessary in order to show the relationships

can then be ascertained. What category

should be used for the group as a whole

should be based primarily on what has been

used in related groups; and when the rank of

the group studied is decided, then the vari-

ous subdivisions of the group should fall

into place. In the case of the pelecypods this

will not be easy because the entire classifica-

tion at the familial levels is nebulous. How-

ever, much progress could be made if these

problems were approached in as objective

a manner as possible. Such studies would

undoubtedly result in many major changes

in the classification of the pelecypods above

the generic level.

There has been a tendency to redefine,

restrict, and propose more subfamilies,

families, and superfamilies, apparently as a

result of the rapid increase in the number of

proposed genera and subgenera of pelecy-

pods. This tendency has met with some

opposition, but some of the objections to

creating more families or raising the various

groups to higher categories seem to be ill-

founded. As MacNeil (1938, p. 1) stated:

With our increasing knowledge of the structure

and phylogeny of the Pelecypoda it becomes more

and more obvious that their supergeneric classi-

fication is short of satisfaction, the principal

defect being that not enough groups of high

ordinal rank have been recognized.

Recognition of more groups of high ordinal

rank would undoubtedly alleviate many of

our present problems of pelecypod classifica-

tion. Another solution might be to create

categories for groups between the generic

and subfamily levels as the. entomologists,

for example, have done.

To improve the classification more work

is needed at the genus and family levels,

and it should include a careful analysis of

all morphologic, embryologic, chronogenetic,

and geographic data. Many of the basic

32 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

data are in the literature—but careful

analysis and synthesis of the data are needed

to ascertain phylogenetic relationships.

SOME BASIC PELECYPOD LITERATURE

A thorough understanding of the following

references is necessary as a starting point for

a classification of the Pelecypoda. This list

is not intended to be complete, but it should

form a good basis for the student who is

interested in this group of mollusks.

BERNARD, Fruix. Sur le développement et la

morphologie de la coquille des les lamelli-

branches. Bull. Soc. Géol. France, sér. 3 (23):

104-154; (24): 54-82, 412-449; (25): 559-566.

1895-1897.

. Recherches ontogéniques et morphologiques

sur la coquille des lamellibranches. Ann. Sci.

Nat., Zool. et Pal., sér. 8 (8): 1-208. 1898.

Datu, W. H. Contributions to the Tertiary fauna of

Florida, etc.: Part III. A new classification of

the Pelecypoda. Trans. Wagner Free Inst. Sci.

Philadelphia 3 (3) : 483-570. 1895.

DovuvitLe, H. Classification des lamellibranches.

Bull. Soc. Géol. France, sér. 4 (12): 419-467,

69 figs. 1912.

Jackson, R. T. Phylogeny of the Pelecypoda, the

Aviculidae and their allies. Mem. Boston Soe.

Nat. Hist. (4) : 277-400, pls. 23-30, 53 figs. 1890.

Neumayr, M. Bevttrdége zu einer morphologischen

Eintheilung der Bivalven. Besonders Ab-

gedruckt aus dem 58 Bande der Denkschriften

der Mathematisch-Naturwissenschaftlichen

Classe der Kaiserlichen Akademie der Wissen-

schaften: 101 pp. Wien, 1891.

PELSENEER, Pauu. Sur la classification phylo-

génétique des pélécypodes (communication

préliminaire). Bull. Sei. France, Belgique:

27-52, 4 figs. Paris, 1889.

. A treatise on zoology (edited by E. Ray

Lankester), Part V, Mollusca: 355 pp., 301

figs. London, 1906.

Les lamellibranch2s de lVExpédition du

Siboga, partie anatomique. Monograph 53a:

125 pp., 26 pls. Leiden, 1911.

RipEwoop, W. G. On the structure of the gills of

the Lamellibranchia. Philos. Trans. Royal Soc.

London, ser. B (195): 147-284, 61 figs. 1903.

vot. 44, No. 1@

REFERENCES

Cotton, B. C., and GoprreEy, F. K. The molluscs

of South Australia, Part I. The Pelecypoda:

314 pp., 340 figs. Adelaide, 1988.

FRIZZELL, Don L. Preliminary reclassification of

veneracean pelecypods. Bull. Mus. Roy. Hist.

Nat. Belgique 12 (34): 84 pp., 1 fig. 1936.

. A study of two arcid pelecypod species from

western South America. Journ. Pal. 20 (1):

38-51, pl. 10, 13 figs. 1946.

Hass, T. Genera of Japanese shells: 4 vols., 326 pp.,

770 figs. Tokyo, 1951-1953.

Keen, A. M. Outline of a proposed classification of

the pelecypod family Veneridae. Minutes of the

Conchological Club of Southern California.

Minutes 113: 2-11. September, 1951.

MacNeIL, F. 8. The systematic position of the

pelecypod genus Trinacria. Journ. Washington

Acad. Sci. 27 (11) : 452-458, 1 fig. 1987..

. Species and genera of Tertiary Noetinae.

U.S. Geol. Survey Prof. Paper 189-A: 49 pp.,

6 pls., 2 figs. 1988.

Mayr, E., Linstey, E. G., and Usinemr, R. L.

Methods and principles of systematic zoology-

328 pp., 45 figs. New York, 1953.

Moorg, R. C., Laticker, C. G., and FIscHER,

A. G. Invertebrate fossils: 766 pp. New York,

1952.

Nicou, D. Origin of the pelecypod family Glycy-

meridae. Journ. Pal. 24 (1): 89-98, pls. 20-22,

2 figs. 1950.

. Systematic position of the pelecypod EKuloxa.

Journ. Pal. 27 (1): 56-61, 8 figs. 1953.

. Systematic position of the pelecypod Plio-

cardia. Journ. Pal. 27 (5) : 703-705, 7 figs. 1953a.

PIVETEAU, JEAN. Traité de paléontologie (2): 785

pp. Paris, 1952.

REINHART, P. W. Classification of the pelecypod

family Arcidae. Bull. Mus. Roy. Hist. Nat.

Belgique 11 (13): 68 pp., 5 pls. 1935.

Surock, R. R., and TwENHOFEL, W. H. Principles

of invertebrate paleontology: 816 pp. New York,

LO53:

Simpson, G. G. The principles of classification and

a classification of mammals. Bull. Amer. Mus.

Nat. Hist. 85: 350 pp. 1945.

THIELE, J. Handbuch der systematischen W evcntier-

kunde 3: 779-1022, figs. 784-893. Jena, 1934.

TremuettT, W. E. English Eocene and Oligocene

Veneridae. Proc. Malac. Soc. London 30 (1-2):

1-21, pls. 1-4, 1 fig. 1953.

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mepeescuinipe On, Counc Of A A. Al Qs.) cence as cock veate ok amiss esa Watson Davis

Committee of Auditors....... Loutst M. Russewu (chairman), R. 8. Dri, J. B. REESIDE

Committee of Tellers...... C. L. Garner (chairman), L. G. HenBrest, Myrna F. Jones

CONTENTS

Puysics.—On research and education—in fluid dynamics. RAYMOND

Js SEEGER: 286 Si cian sy eds ae eye ea Oe ames ole ee

MatueEmatics.—A lower limit on the number of hypergroups of a given

order. Howarp H. CAMPAIGNE...). (0.02.05... ...2. 2.5) ae

PALEOBOTANY.—A Permian Discinites cone. SERGIuS H. MAMay......

Botany.—Trapellaceae, a familial segregate from the Asiatic flora. Hut-

Jane Ta os as ae

Taxonomy.—For and against the doctrine of prescription as applied to

taxonomy: A historical retrospect. AusTIN H. CLARK..........

ZooLoey.—On Polyclinum indicum, a new ascidian from the Madras coast

of India... VO. SHBASTIAN 2.) c0. si one oer Ae

HELMINTHOLOGY.—Psilocollaris indicus, n. gen., n. sp. (Psilostomidae

Odhner, 1911: Trematoda) from an Indian stork, Dissoura e. epi-

scopus.’ KUNWAR SURESH SINGH. ........... 64-60: 5-2

MauacoLocy.—Hydrobia tottent, new name for Turbo minutia Totten,

1834 (Gastropoda: Hydrobidae). J.P. EH. MorrIson............

MatacoLtoGy.—Trends and problems in pelecypod classification (the

supergeneric categories). .Davip Nicol............:2.heeeeeee

This Journal] is Indexed in the International Index to Periodicals.

Page

VoL. 44 FrBruary 1954

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JOURNAL

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WASHINGTON ACADEMY OF SCIENCES

VoL. 44

MATHEMATICS—Inequalities restricting the form

February 1954

No. 2

of the stress-deformation

relations for isotropic elastic solids and Reiner-Rivlin fluids. M. Baker and

J. L. Ericksen, Naval Research Laboratory.

M. Trent.)

According to the natural state theory of

elasticity, the principal values t,, ts, ts; of

the stress tensor for an isotropic, perfectly

elastic material are given in terms of the

principal extensions 6; , 62 , 63 by”

20 OD OD OD

eee = rt —— |) + (1 +8,)

po ( al .) ar |

OD 1

ae ue

olf (1 + 6;)?

if the material is compressible, and by

oD if

—— +2 (1 +5) - 2 2

BBN Ss Parra + 6” 2

if the material is incompressible. Here, >

the strain energy, is a function of he

(1 + 6)" sgl te 2) + (1 + 53)’, LS

aes 1) (1 + 69)” ay (1 + 63)°(1 + 63)" ae

ie be): eee 6). and I/IT = (1 + 61)

(1 + 6.)°(1 + 63), p is an arbitrary hydro-

static pressure, p is the density in the

deformed state, and p) the density in the

undeformed state. The condition for incom-

pressibility is 77 = 1 so that 2 = ZU, IJ)

in (2).

Truesdell” has given a physical argument

which shows that, for an incompressible

material, the inequalities

6] O>

(+0)? = +>

all ee

1 See, e.g., TRUESDELL, C. A., The mechanical

foundations of elasticity and fluid dynamics, Journ.

Ratl. Mech. and Anal. 1: 173-182. 1952.

2 TRUESDELL, C. A., op. cit.: 181-182.

33

(Communicated by Horace

should always be satisfied. Rivlin’ derived

a special case of this inequality and used it

to obtain qualitative information about the

Poynting effect. These inequalities also

gave information concerning the propaga-

tion of waves in such materials.’

By considering general isotropic func-

tions, we shall deduce certain inequalities

which follow from conditions imposed on

the eigenvalues of the tensors involved. We

shall then discuss the significance of these

conditions as applied to elastic solids and

viscous fluids, and determine the form which

these inequalities assume in the different

physical situations. In particular, we shall

see that inequality (3) should hold for both

incompressible and compressible materials.

Suppose that the 3 X 3 real symmetric

matrix A is an isotropic analytic function

of the 3 X 3 real symmetric matrix B. We

then have’

Sta oe Bl Be (4)

where the scalars f; are functions of the three

principal invariants of B.

Now, if B’ exists, we can also write A in

the form

A = gol +g4iB™ + giB, (5)

where the scalars g; are also functions of the

three principal invariants of B. Equation (4)

3 Rivuin, R. S., and SaunpgErs, D. W., Large

elastic deformations of isotropic materials VII.

Experiments on the deformation of rubber, Phil.

Trans. Roy. Soc. London (A) 248: 251 288. 1951.

4 ERICKSEN, J. L., On the propagation of waves

in isotropic, perfectly elastic materials, Journ. Ratl.

Mech. and Anal. 2: 329-837. 1953.

5’ TRUESDELL, C. A., op. cit.: 131-132.

34 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

implies that the eigenvalues a, , d2, a; of A

are related to the eigenvalues 6, , bs, bs of

B by

a; = fo + fib; + fob, (6)

while (5) yields

a; =g +91/bi + mdi. (7)

We want to determine necessary and suff-

cient conditions that the inequality a; > a;

hold whenever b; > b; . Let us first consider

(6), which yields

Oi = Oy = (b; ray b [fi + foll: =e b;)|.

Thus a necessary and sufficient condition

that b; > b; imply a; > a; is that

fi + fo(b; + 6;) > 0, (8)

whenever 6; > b;. Since (8) is unaltered if

b; and 6; are interchanged, this inequality

must hold unless 6; = 6; (¢ ¥ 7), in which

case (8) is replaced by the weaker inequality

hh Ste fe (b; aie b;) aa Hal fe 2fob; 22) (peed Davie

latter condition follows from the fact that

A is a continuous function of B. Similarly,

from (7), we obtain, as an alternative neces-

sary and sufficient condition,

gi > g-r/b.b; , (9)

unless: b;:— -b,.,, and. ¢).. @=1/ 0j0 7, —s0 4/6;

Oe — Oral e

Let T denote the stress tensor in a con-

tinuous medium. At any point P, each of

the three perpendicular planes whose nor-

mal vectors are eigenvectors of T has the

property that the stress vector acting on it

at P is normal to it. These normal forces

(per unit area) are the eigenvalues 4 , fo, ts

of T. A positive eigenvalue corresponds to

a tension, a negative eigenvalue to a pres-

sure.. We assume the eigenvalues ordered

sO that ty a to 2 ts :

In an isotropic, perfectly elastic body, the

principal directions of T coincide with the

directions of principal extension. We re-

strict our attention to these directions. For

such a material, it is reasonable to suppose

that the greatest (least) tension occurs in

the direction corresponding to the greatest

VoL. 44, NO. 2

(least) extension. Expressing these condi-

tions analytically, we obtain

t; > t; whenever 6; > 6;. (10)

It may be possible to get further conditions

by comparing forces and extensions in other

directions. One might, for example, compare

normal stresses and extensions normal to an

arbitrary pair of perpendicular planes. How-

ever, because of the fact that a pure shear-

ing stress acting on a plane may give rise to

extensions normal to this plane, the validity

of results obtained from such a comparison

is questionable. Comparing (1) and (7),

we see that (10) implies (9) with b; =

(1 + 6,)°, 91 = 2(p/p) 02/0L, g-1 = —2(0/po)

III dz/dll. Since p/p > 0, we have for

0 Ze i,

az TOE a> |

ap his ea ta

(1+ 6)2(1 +6207] i

(11)

z al a>

7 ae

ea Geeyesc j

for a compressible material. Since //J =

(1 +8) +5) = 6.)*, the imequalidtes

(11) become

eae = 20

on ae

ax ‘

= + (1+ 5)# = > 0 |

if 8; ¥ & (i, j,k ), |

+ (2)

a> babe

|

|

if 6; = 6, (7,7, k ¥).

Comparing (7) with (2), we see that (10)

implies (9) with b; = (1 + 6,)’, g: = 202 /al,

g1 = —20z/odlI. Using the incompressi-

bility condition //7J = 1, one can show that

the resulting inequality reduces to (12),

which is in agreement with Truesdell’s result

(3). Results of a number of experiments on

rubber,” which is virtually incompressible,

show that d2/ol > 0, @2/oll > 0 fora

wide range of values of J and JJ so that (12)

certainly holds in this case.

Now, in the classical linear theory of

elasticity, the stress tensor T is given in

6 Rivuin, R.S., and SaunpERs, D. W., op. cit.

FEBRUARY 1954

terms of the infinitesimal strain tensor E by

T = dX 1 + 2uE, where 2d and wp are the

Lamé constants. Application of the above

analysis to this expression for T leads im-

mediately to the condition u > 0, which is

certainly true for all materials which are

adequately described by this theory.

According to the theory of compressible,

highly viscous fluids proposed by Reiner,’

the stress T is an analytic isotropic function

of the rate of deformation D, so that

T=fol + fiD + f2D’, (13)

where the scalars f; are functions of the three

principal invariants of D. In terms of the

eigenvalues d,, d., and d3 of D, these in-

variants are d; + dz + d3, didz + dods +

d3d,, and d,d.d;. Rivlin’s theory® differs

from this only in that D is assumed to satisfy

the condition of incompressibility, d; + d2 +

d; = 0, and —f is replaced by an arbitrary

hydrostatic pressure p. That 1s,

Beer 1 fiD + f2D?. (14)

At a given point P at a given time, the

rate of increase of distance between the

material particle at P and particles lying

on a sphere of fixed radius r about P will

take on stationary values for certain par-

ticles on this sphere. The directions de-

termined by drawing the radius vector from

P to these particles approach the principal

directions of D as r tends to zero. In first

approximation, the stationary values men-

tioned above are obtained by multiplying

the appropriate eigenvalue of D by r. Let us

restrict our attention to the principal direc-

tions of D which, in a Reiner-Rivlin fluid,

are simultaneously principal directions of T.

It seems reasonable to suppose that, at a

point P in such a material, the greatest

(least) tension will be exerted across a plane

whose normal is in the direction in which

the rate of increase of distance between the

7 REINER, M., A mathematical theory of dt-

latancy, Amer. Journ. Math. 65: 350-362. 1945.

8 Rivuin, R. 8., Hydrodynamics of non-New-

tonian fluids, Nature 160: 611-613. 1947.

BAKER AND ERICKSEN: STRESS-DEFORMATION RELATIONS 35

particle at P and particles equidistant from

P is greatest (least). That is, we should have

t; > t; whenever d; >d;. | (15)

It then follows from (6), (8), and (13) that,

for a compressible material,

fi + fo(di + d;) > 0 |

(16

fitfld: +d;)20 ifd; =d;( #)).

From (6), (8), and (14) it follows that (16)

must also hold for incompressible materials.

In this case one can, using the incompressi-

if dj + hae

bility condition d; + d, + d; = 0, write

(16) as

fi —fed; > 0 I a; CS Oe Gist, =),

(17)

fi —fod; 2 Ty e— Ned CO Iles

For a plane motion of an incompressible

fluid, d. = O and d; = —d,, so that (17)

becomes

fe 0 te fo ed at ay = 0

fiz0

(18)

if di = 0,

If d, = 0, then d, = d3; = 0, so the motion is

instantaneously rigid. From (18),

ay

fod

except perhaps when d; = OQ, in which case

the ratio on the left may be indeterminate.

It has been shown? that, in such a motion,

the equation f; + fod: cos 2¢ = O determines

two characteristic directions for the equa-

tions (14). According to (19), there exists

no real angle ¢ for which this equation is

satisfied. |

In the classical theory of isotropic viscous

fluids, fo = O and f; = 2u, where uw is the

coefficient of viscosity. Since uw is always

positive, (16) holds for materials to which

this theory applies.

Silt (19)

9 HRIcKSEN, J. L., Characteristic surfaces of the

equations of motion for non-Newtonian fluids,

Zeitschr. fiir Angew. Math. u. Physik 4: 260-267.

1953.

36 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

VoL. 44, No. 2

PALEONTOLOGY .—The development of the-hingée of Veniella conradi (Morton)

and some conclusions based on its study.

University.

While studying Cox’s excellent report on

Cretaceous and Eocene fossils from the Gold

Coast (1952) the writer’s attention was

struck by a statement (p. 19) made during

his comparison of the hinge of Venzella

conradi (Morton), the type species of

Veniella Stoliczka (1870, p. 181), with that

of ‘“V.”’ undata (Conrad), the presently ac-

cepted name for the type species of Roudairia

Munier-Chalmas (1881, p. 74). This state-

ment may be paraphrased as follows: If

good figures of the interior of the shell of

Veniella conradi ‘“‘are compared with those

of the hinge of V. wndata . . . it may be seen

that hinge-structure of the right valve is

essentially the same in the two species. In

the left valve of V. undata the anterior

lateral consists of two narrow, divergent

prongs united at the top, whereas in the

single satisfactory illustration of the left

valve of V. conradi (that given by Stephen-

son [1923, p. 66, fig. 4]) the same tooth ap-

pears to be thicker and its bifid character

is less obvious. The fine transverse crenula-

tion of the posterior laterals of V. conradz,

shown in Stephenson’s figures, has not, been

observed in V. undata, but this may be due

to imperfect preservation.”’

This suggestion of a difference in the left

valves of the two species without a corre-

sponding divergence in the right led me to

examine the series of specimens of V. conrad

in our collections, and especially those in

the Wade collection from Coon Creek, Tenn.

where the species is common and well-pre-

served. It was found that while the condi-

tion of the left anterior lateral in all the

adult left valves agreed with that figured by

Stephenson, in other respects the specimen

figured was in a somewhat immature stage

of development. On the other hand, certain

‘“adolescent”’ individuals show some trace of

bifidity of the anterior lateral, though never

is this as pronounced as in the illustrations

of “V.” undata given by Quaas (1902, pl.

34, fig. 22: as Roudairia Drui) and by Per-

vinquiére (1912, pl. 15, fig. 11b). Further-

more, a comparison of right valves with

those figured by Pervinquiére (1912, pl. 15,

H. E. Voxrs, The Johns Hopkins

fig. 12a), and by Rennie (1930, pl. 21, fig. 9;

as Veniella druz), shows that there are dif-

ferences in the anterior area of the hinge

that correspond with those of the left valve.

Another fact that became apparent dur-

ing the examination of these specimens was

that there was a pronounced change in the

nature and orientation of the anterior ele-

ments of the hinge structure during the

development of the individual. An examina-

tion of the published literature indicates

that this fact has not been noted by previous

students, and since it may have some sig-

nificance in elucidating the ancestry of the

veniellid type, it seems desirable to record

the observed facts at this time.

The hinge of the adult right valve has been

well figured by Wade (1926, pl. 24, fig. 16),

while illustrations and descriptions of some-

what less mature examples of this hinge have

been given by Gardner (1916, pp. 643-5, pl.

38, fig. 4) and Stephenson (1923, pp. 257-62,

pl. 66, fig. 5; 1941, pp. 168-70). Miss Gard-

ner (1916, pl. 38, fig. 3) also figures, but

without comment, the hinge of an immature

right valve of somewhat unusual outline.

Meek (1876, pp. 147-8, text figs. 9-11) re-

figured Morton’s type specimen, a right

valve, but misinterpreted the nature of the

hinge. Douvillé (1921, p. 121, text fig. 16)

gives a very diagrammatic drawing of a

right valve from ‘‘la Craie superieure du

Tennessee,’ presumably the Coon Creek

locality.

The only adequate figure of a hinge of the

left valve is that of the somewhat immature

specimen furnished by Stephenson (19238, pl.

66. fig. 4).

DEVELOPMENT OF THE HINGE OF THE RIGHT VALVE

The smallest complete right valve available

for study has a length of 7.6 mm and a height

of 6.6 mm. There are two well-developed con-

centric lamellae on the exterior of the shell.

A very similar specimen has a length of 8 mm

and a height of 7 mm. From these minima a

rather complete series of specimens up to adult

valves has been studied. The largest valves are

61.5 mm long and 50.5 mm high; and 57 mm

FEBRUARY 1954

long and 52.5 mm high, respectively. Fragments

of valves smaller than the smallest mentioned

above give suggestions as to earlier stages of

the hinge development but are too incomplete

to permit certain conclusions.

In the smallest complete specimens the hinge

of the right valve (Fig. 1) consists of cardinals

3a and 3b. Tooth 3a is long, relatively thin,

and sub-parallel with the anterior dorsal margin

of the valve. Its anterior end terminates at a

point approximately midway across the narrow

hinge plate. The posterior cardinal, 3b, is oblique

in position, subtriangular in shape, moderately

heavy and weakly grooved; in adult specimens

it is weakly bifid. The anterior end of the tooth

is, in the smallest specimen, definitely continu-

ous with the posterior end of 3a, although

the lamina from which these two teeth have been

derived is greatly thinned and the connection is

quite tenuous.

In addition to the two cardinals the hinge

carries long, deep anterior and posterior lateral

sockets, bordered on their ventral sides by rela-

tively strong laminae, AI and PI, and with low,

but sharp ridges on their dorsal sides marking

the position of laminae AIII and PIII. The sides

of both anterior and posterior sockets are finely

transversely grooved. The most striking feature

of this assemblage is the presence of a relatively

strong, elongate and moderately high tubercle

on the posterior half of AI. The tubercle, which

represents 1 in the Bernard system of numbering,

is here clearly lateral in position, being located

parallel with the ventral margin of the hinge-

plate, and having its posterior end distinctly

in front of and slightly ventrad of the anterior

end of 3a. 4

The socket between 3a and 3b is an elongate

triangle, whose apex lies approximately at the

posterior two-thirds of its total length. Tooth

3a margins the anterodorsal side of this socket,

but its anterior termination is furnished by the

posterior end of the tubercle 1 on AI.

A comparison of this hinge (Fig. 1) with the

adult hinge (Fig. 4) reveals that the principle

modifications have occurred in the anterior part

of the structure, particularly in the shape and

position of teeth 3a and 1, and in the total reduc-

tion of the remaining part of the anterior lateral

lamina AI. On a purely mechanical interpreta-

tion, the changes might be ascribed to the pos-

terior migration of the tubercle 1, from its anterior

lateral position to one that is almost wholly

VOKES: HINGE OF VENIELLA CONRADI od

subumbonal. This migration has brought it into

approximate contact with 3a and has forced

that tooth to change its shape from a narrow

lamina to that of an inverted triangle that has

its apex on the ventrad side where it is in juxta-

position with 1. Coincidental with this change in

shape there is a sharp change in the orienta-

tion of the tooth. Initially it is subparallel with

the anterodorsal margin so that its long axis

virtually intersects the anterior extremity of

the valve (Fig. 1); in its maximum distortion,

in what must on other characters, be considered

gerontic individuals, the axis of the tooth has

been shifted to a position where, if projected, it

would intersect the posteroventral margin of

the valve (Fig. 4).

The change in shape and the migration of the

axis of 8a markedly alters the shape and propor-

tions of the socket separating that tooth from 3b.

Initially the socket is elongately triangular with

its anterior termination well in advance of the

umbo; in the gerontic individuals, it has become

narrow, with subparallel sides, and has its long

axis trending posteroventrally.

The migration of 1, and its conflict with 3a

changes the shape and position of the former,

from its initial structure as an elongate tubercle

parallel with the ventral margin of the hinge

plate, to a heavy, triangular structure, whose

apex is directed dorsally toward the anterior

end of the base of the inverted triangle that is

3a. In the gerontic stage of development, 3a and

1 come to represent what is essentially a unit

transverse to the hinge plate that is formed of

two triangles in juxtaposition, the posterodorsal

side of 1 being parallel to and separated only by a

very narrow but deep groove from the. antero-

ventral side of 3a.

It is to be emphasized, as shown in Fig. 12,

that the change in shape and migration of teeth

1 and 3a is a dynamic process that does not appear

to have become stabilized at any time during

the development of the individual.

DEVELOPMENT OF THE HINGE OF THE LEFT VALVE

The changes in the hinge of the left valve are,

as would be expected, essentially similar to those

that mark the right. The smallest complete

specimen of this valve available for study has a

length of 10.56 mm and a height of 10.2 mm.

There are, however, smaller though incomplete

individuals that probably did not exceed 6 mm

in length, in which the hinge is adequately pre-

38 JOURNAL OF THE WASHINGTON ACADEMY OF'SCIENCES

served for the purposes of this study. The largest

specimens in the collection have lengths of 60.7

and 61.8 mm, and heights of 58.0 and 52.0 mm,

respectively.

The immature hinge of the left valve (Fig. 5)

consists of a long, thin anterior lamella that is

parallel to the: ventral margin of the hinge plate.

This lamella bears two elevations on its outer

face that are separated by a somewhat lower

area which is, however, distinctly raised above

the general level of the hinge plate itself. The

anterior of these elevations clearly represents

AII; the posterior, which is somewhat swollen

toward its obliquely terminated posterior end,

represents 2, but there is no feature visible that

would distinguish it as either 2a or 2b, or as a

union of both.

In addition to these elements there is a thin,

obliquely transverse 4b, and an elongate, strong,

PII that is markedly transversely striated. In

the smallest specimens available, the AII is

smooth, but it too, soon becomes transversely

striate, the striae being well developed on a

specimen 10.5 mm. in length.

The subsequent modification of this hinge is,

like that of the right, mechanically a factor of

the migration and enlargement of 1. In the earli-

est stages there is a slight flattening of a portion

of the ventral side of AII (Fig. 6) marking the

position of tubercle 1, which is still located on the

lamella AI. This becomes enlarged, particularly

in the low area between AII and 2, until the

lamella of which these two are a part is disrupted

with the posterior end of AII being deflected

dorsally. At this stage (Fig. 7), tooth 2 is elon-

gately triangular and relatively quite heavy. The

apex of.the triangle so formed lies immediately

below the umbo, the posterior dorsal slope is

short, almost vertical, while the anterior dorsal

slope is at least two and a half times as long as

the posterior and trends obliquely forward.

As the progressive posterior movement of 1

continues and involves the 3a, these two teeth

encroach into the space between AII and 2,

completely separating them and resulting in a

progressive modification of their shape. Cardinal

2 becomes more and more equilaterally trigonal,

and tends to develop a broad ‘‘V”’ shaped groove

on its ventral side; AII tends to maintain the

position of its anterior part at the ventral margin

of the hinge-plate, while its posterior part is

more and more deflected dorsally, with the result

that the anterior part becomes elongately trigonal

vou. 44, No. 2

in- Shape, with.a relatively tenuous “‘tail’’ pro-—

jecting posteriorly from the apex of the triangle

formed of the posterior part of the original

lamella! (Fig. 8). A continuation of this “tail”

across the interspace between AII aind the antero-

ventral edge of 2 forms the low, acute ridge —

that separates the interspace into sockets for the —

reception of 1 and 3a of the right valve. This

stage of development of the left hinge might be

termed an adolescent one and is the stage that

was figured by Stephenson (1923, pl. 66, fig. 4).

Continued modification eventually results

(Fig. 9) in a 2 that is broadly heavy, rather than

trigonal in shape, and which trends obliquely

posteriorly, so that a line drawn through the

axis of the tooth and prolonged to the shell

margin would approximately cut the middle of

the posterior end of the valve. At the same time

AII loses its ‘‘tail’” and becomes more and more

strongly triangular in shape eventually forming a

strong, almost equilaterally triangular tooth.

It is probably significant, however, that the ridge

crossing the ‘interspace’? between AII and 2

separating it into sockets for the reception of 1

and 3a continues to connect the posterior side

of the apex of AII with the anteroventral edge

of 2.

ON THE DENOMINATION OF TOOTH 2

Douvillé (1913, 1921) has shown that in the

development of the arcticid (ecyprinid) hinge the

left anterior lateral AII becomes extended to

the center of the hinge where it abuts against

3b, and is bent down at its posterior end to form

a chevron with its apex just beneath the umbo.

This downbent portion develops as 2b. He then

considered that the umbonal end of the remain-

ing portion of AII later differentiated to form

2a, leaving AII of the later forms as only the

anterior end of the original lamellar tooth.

Casey (1952, pp. 123-5, text fig. 1) postulates

a lucinoid ancestor for the cyprinoid group. In a

series of drawings he shows the lucinoid ancestor

as having a bifid 2 and a distinct AII. In the

next stage, which he labels the “‘cyprinoid”’

stage, the two limbs of the bifid lucinoid “2” are

designated as 2b; and 2bs, and a distinct 2a is

forming on the anterior end of AII. In his “ad-

vanced cyprinoid” and ‘early cyrenoid” stages

1 Since this is clearly part of the original lamella

it cannot be called a vinculum (Casey, 1952, p. 124),

since that term by definition is applied to second-

ary shelly matter only.

FEBRUARY 1954 VOKES: HINGE OF VENIELLA CONRADI 39

2a becomes strengthened and begins to separate AIT and assumed a position identical with that

from the posterior end of AII as it migrates pos- — originally held by prong 2a).

teriorly, displacing the anterior prong 2b, of the Since there is never more than one element of

lucinoid tooth. In the final stage, ‘“cyrenoid” the tooth 2 complex visible in the Veniellid

stage 2a has become completely separate from hinges available for study it is not clear just

Fics. 1-11.—Veniella conradi (Morton): 1-4, Development of the hinge of the right valve: (1)X 5;

Wren (e+) KL. 5-9, Development of the hinge of the left valve, (5) X 6, (6) X 5; (7) X 2; @,

Or. 10-11, Exterior of left and right valves: (10), same specimen as Fig. 8, X 1.3; (11) same

specimen as Fig. 1, X 3. Figs. 2, 7, specimens from Ripley formation, Owl Creek, Miss.; others from

Coon Creek tongue, Ripley formation, Coon Creek, Tenn.

40 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

which part is represented. In the earliest stages

(fig. 5) there is a suggestion of a down-bending

of the posterior end of the AII-2 lamella that 1s

very similar to the figures given by Douvillé

(1921, p. 118, fig. 6) for the hinge of Hotrapezvum

germart (Dunker), the types species of Hotra-

pezium Douvillé, considered by Douvillé as

representing the primitive stage in the develop-

ment of the cyprinid hinge. It soon loses this

characteristic, however, becoming first a broadly

triangular structure and later a heavy simple

tooth that comes to trend obliquely posteriorly.

It seems best, therefore, to designate it as

tooth 2, recognizing that, since it is certainly

continuous with AII in its earlier stages, it prob-

ably was formed as a complete fusion of ele-

ments 2a and 2b. It cannot be demonstrated,

however, whether or not the 2b element arose as

a result of the downward bending of AII, as

postulated by Douvillé, or as a separate element

that was joined by 2a, as postulated by Casey.

GENERAL OBSERVATIONS |

Relationships of Veniella and Roudairia.—

Available illustrations of the hinge of Roudairia

undata (Conrad), the presently accepted name

for the genotype species of Roudairia Munier-

Chalmas 1881 are not wholly satisfactory for

comparison with those of Veniella. Perhaps the

best are those given by Pervinquiére (1912,

p. 230, pl. 15, figs. 9-138, especially figs. 11b and

12a). By a particularly good fortune these are

re-illustrations of Munier-Chalmas’ types of his

Roudaria drut |=Opis undata Conrad] and

hence represent the specimens upon which he

based his genus. The most striking difference

between the hinges there figured and those of

Veniella conradi is the persistence, in Roudairia,

of the right anterior lateral AI, which is lost in

Veniella, and in the effect this tooth has on the

left anterior lateral AII. This tooth instead of

being broadly triangular and heavy, as in Veniella

is split into an inverted ‘‘V’’-shaped unit whose

two narrow prongs margin AI anterodorsally

and posteriorly. This condition is particularly

well-shown by Quaas (1902, pl 24, fig. 22). These

hinge differences, in conjunction with the differ-

ences in weight of the shell and in external orna-

mentation, mentioned by Olsson (1934, p. 39)

justify the retention of Roudairia as a separate

genus, rather than relegating it to the synonymy

of Veniella, as has been done by Newton (1909,

p. 69), Wade (1926, p. 77), Rennie (1929, pp.

voL. 44, No. 2

26-8; 1930, p. 191), Cox Q952. pp) ie i1oe

and others.

Roudairia and Veniella were, so far as can

be determined from published records, contem-

poraneous genera. Both ranged from the Turon-

ian? through the Maestrichtian, and both seem

to have been most abundantly represented in the

upper Campanian and the Maestrichtian.? It is

to be assumed therefore that the two genera

represent separate essentially parallel evolution-

ary developments within the very plastic arcticid

stock. The specimens from Pondoland and Zulu-

land identified by Rennie (1930, pp. 192, 244,

pl. 29, figs. 1-5) as ‘“Veniella forbesiana

(Stoliczka)”’ are now considered as being of

upper Campanian or Maestrichtian age and

can not, therefore, be considered as having an

intermediate position between the two genera,

despite the fact that they show a greatly reduced,

essentially tubercle-like, AI with a corresponding

strengthening of AII, called the left anterior

cardinal by Rennie. These, together with the

strong transverse striations developed on the

posterior laterals closely approach the conditions

observed in VY. conradi. The illustrations of

Stoliczka (1870, pl. 9, figs. 2-8) are too diagram-

matic to permit certain comparisons. There

seems, however, to be differences in the posterior

laterals and possibly also in the shape of 3a,

that would justify at least a question as to the

identity of Rennie’s species. The Albian species

2'The specimens of Veniella mortoni Meek

figured by Stanton (1898, pl. 23, figs. 6-9) were

collected on the Arkansas River, 18 miles west of

Pueblo, Colo., associated with a fauna including

such characteristic Carlile species as Collignoni-

ceras hyattt (Stanton), Inoceramus fragilis Hall

and Meek, and Ostrea lugubris Conrad.

Stoliczka’s ‘“‘Cyprina’’ forbesiana (1870, p.

192, pl. 9, figs. 2-8) was described from the Tri-

chinopoly Group of India, now generally consid-

ered as being of Turonian age. The figured hinge

of the left valve shows the characteristic split

AII of Roudairia, while that of the right valve

seems to lack an anterior lateral, thus suggesting

Veniella. Since, however, the species clearly pos-

sesses the strong carinate post-umbonal ridge of

Roudairia, its reference to that genus seems more

correct.

3 It is recognized that the application of Euro-

pean stage names to the geologic range of Veniella

applies a precision of intercontinental correlation

that is not wholly in accord with the present state

of our knowledge. Nevertheless, since these corre-

lations are based mainly upon the distribution of

ammonite genera who seem to have had a more

rapid rate of migration than the heavy-shelled

pelecypods, it is believed that the correlations im-

plied above are sufficiently close to have real

significance when applied to the genera under

consideration.

FEBRUARY 1954

“Veniella’” etheridgei Newton, as figured by

Rennie (1931, p. 242, pl. 31, figs. 1-3) differs

entirely in the nature of the posterior lateral and

in the bifid condition of tooth 2 of the left valve.

The drawing of the hinge is such as to make it

uncertain just what condition exists with respect

to AIT; it might be that the tubercle-like struc-

ture shown at the posterior end of the structure

represents 2a, in which case the bifid tooth would

be 2b and the species would be entirely separate

from the Veniella-Roudairia complex. In general,

the hinge suggests a closer relationship to Venzl-

cardia than to Veniella.

Relationship of Cicatrea Stoliczka.—Stoliczka

(1870, p. 191) described the subgenus Cicatrea

with Cyprina (Circatrea) cordialis Stoliczka

(1870, p. 199, pl. 10, figs. 1, a, b, c, 2) as the type

species. Among the important generic characters

mentioned were: (1) ‘‘a rather short but deeply

bifureate groove in which the ligament is lodged.”’

(2) “The posterior cardinal teeth are rather

narrow in both valves...the two anterior

cardinals in the left valve are very large, the

same superimposed teeth in the right valve,

however, very small.” (3) ‘‘The anterior muscu-

lar impression is anteriorly margined by a sharp

ridge.”

Douville (1904, p. 216), on the basis of a speci-

men from Madagascar, which he unfortunately

did not figure, concluded that Cicatrea cordialis

was actually a Roudmria and that Cicatrea

Stoliezka was therefore a prior name for Munier-

Chalmas’s genus. He refused, however, to adopt

the Stoliczka name on the grounds that it had

not been accurately defined. Rennie (1929, p. 27)

discusses the matter at some length and points

out that the difference between the nymph struc-

ture described by Stoliczka and that observed

in Roudairia is probably due to imperfect prepa-

ration of the Indian specimens. It should be

pointed out, however, that the heavy, triangular

“left anterior cardinal” described and figured by

Stoliezka (pl. 10, fig. 2) is totally unlike that to

be seen in illustrations of Roudairia undata,

but is entirely like the condition found in adult

specimens of Veniella conradi. In neither genus,

however, is there any suggestion of the ‘‘sharp

ridge” in front of the anterior muscle scar, de-

seribed by Stoliczka, and no structure has been

observed that corresponds to the peculiar feature

shown in this area in his illustration. If it is not

actually present in the species and represents

extraneous material impressed into the shell,

VOKES: HINGE OF VENIELLA CONRADI 4]

then it seems more likely that Cicatrea may be

regarded as a synonym of Veniella rather than

Roudairia. Since Veniella Stoliczka is a new

name proposed as a substitute name for Venilia

Morton, 1834, and also has page priority in

Stoliezka’s work, Cicatrea could safely be dis-

carded, without endangering Roudairia.

Petalocardia Vincent.—In 1924 E. Vincent

(1924, pp. 59-62, text figs. 1, 2) proposed Petalo-

cardia as a subgenus of Venvella, with “Venus ?”

pectinifera Sowerby, from the Upper Eocene,

Bartonian, of England, Belgium and France as

the type species. This is a small form that has in

common with Veniella external lamellar flanges

and a moderately sharp posterior umbonal ridge.

In the right valve 3a trends diagonally forward

across the hinge-plate coming into apposition

with the upturned posterior end of AI, which,

in Vincent’s figure seems to possess a small tu-

bercle possibly representing an incipient cardinal

1; 3b is strong and broadly grooved, while PI

(labelled LP2, by Vincent) is strong and smooth,

with a broad, deep socket above it for the recep-

tion of PII. In addition, however, there is a well-

developed AIII. The left hinge has a triangular

2 (labelled 2a by Vincent), and a lamellar 4b,

both of which are similar to those in immature

Fig. 12.—Relative position of the trend of the

ventral, later posterior, margin of 3a during the

development of the individual. The lines indicate

the relative trend of the margin with respect to

the outline of the valve in specimens whose length

is indicated, in millimeters, adjacent to the trend

line. The specimen of 46.7 mm length represents

an unusually obese gerontic individual from

Brightseat, Maryland. Specimens of 24.9 and 28.8

mm length are from the Ripley formation at Owl

Creek, Miss. All others from the Coon Creek

tongue of the Ripley formation, Coon Creek,

Tenn.

42 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

specimens of Venvella conradi (see fig. 7); AII is

transversely elongate and shows no evidence of

former connection with 2, while PII (called

LP1, by Vincent) is short, with a peculiar ventral

projection from the hinge plate to accommodate

the socket for PI. The inner margin of both

valves is strongly crenulate. Externally, as

figured by Glibert (1933, p. 156, pl. 9, fig. 9)

and Cossmann (1883, p. 169, pl. 6, fig. 7, 7a),

surface of the valves between the flanges shows

a well-developed radial ornamentation.

The external radial ornamentation, denticulate

inner margins, and, in the right valve, the pres-

ence of AIII and the smooth PII, together with,

in the left valve, the distinct separation of AII

from 2, and the short discreet PII with its un-

usual ventral socket arrangement, all serve to

distinguish Petalocardia from Veniella. These

factors, together with the long time interval

between the last known representatives of

Venella and the appearance of Petalocardia,

justify the divorcement of the latter from

Veniella, and its recognition as a distinct genus.

Comparison with Arctica Schumacher s.s.—

A comparison of the figure of a specimen of

“Cyprina” islandica (Linné) with a length of

but 6 mm, given by Bernard (1895, p. 129,

text fig. 14) with specimens and figures, such

as those given by Nicol (1951, p. 1038, text fig.

103) or Theile (1934, p. 856, text fig. 819),

seems clearly to indicate that the development

of the hinge here described for Veniella conradi

is also essentially that which occurs in Arctica‘

islandica. The principal differences between the

two so far as the anterior portion of the hinges

are concerned are: in the immature specimens,

in tooth 2 which is distinctly divided into two

limbs, so that elements 2a and 2b are easily

recognized at this stage of development, even

though later they apparently unite to a single

element; and, in the more adult valves, in the

great variability of the strength of the anterior

elements, 1 in the right valve and AII in the

+The name Arctica Moehring 1752, as repub-

lished in the 1758 translation by Nozeman and

Vosmaer under the title of Geslachten der Vogel,

was clearly not validated by that translation,

being analogous with the names Grus and Coturniz,

also used in that translation, that were discussed

by Hemming in the re-issue of Opinion 5 of the

International Commission on Zoological Nomen-

clature. (see ‘‘Opinions and Declarations. . . ete.,

vol. 1, pt. 14, esp. pp. 120-122, 1944). Arctica

Moehring, therefore, does not invalidate Arctica

Schumacher, 1817, and the latter is the valid name

for the present genus (see also Nicol, 1951, p. 102).

vou. 44, No. 2

left. In many specimens of Arctica these are

reduced to deeply corrugated remnants, or, as

in the diagrammatic drawings given by Cox

(1947, p. 144, text fig. 8a, b) to minute tubercles.

In other specimens, however, notably in that

figured by Nicol, both of these teeth are strongly

and heavily developed, being very similar, in~

this respect to the condition that is maintained

in Veniella conradi.

Developmental stages in the growth of the Veni-

ella hinge —It has already been pointed out that

Douvillé considered the earliest stage in the

evolution of the arcticid hinge to be that in

which AII becomes extended to the center of

the hinge line and is bent down at its posterior

end to form a chevron with its apex just below

the umbo. This he designated the “‘Hotrapezium

stage.”’ The smallest left valves present in our

collection show this condition. No equally minute

right valves are available for study. However,

there is a suggestion of a small socket dorsal to

the AII-2 combination in the lefts that suggests

the presence in the right valves of a small 3a;

if present it would indicate that even at this

small size, the species had passed beyond the

Eotrapezium stage, since a 3a is not present in

that genus.°®

Cox (1947, p. 142) in a review of the British

Jurassic species, recognizes three groups of

species within the family, stating: ‘‘In the right

valve, at a slightly more advanced stage than

that of Eotrapezvum, a small excrescence appears

at the posterior end of AI and is the origin of

tooth 1. In one group of forms, which includes

the Recent Cyprina, this remains merely a

tubercle, and an anterior cardinal 3a develops

close to the lunular margin with its apex touch-

ing that of 3b below the umbo. In other forms,

such as Pronoella and Eocallista of the Jurassic

and Pygocardia of the Miocene, 1 develops

into a strong triangular tooth with its apex

close to the umbo; it thus becomes a median

cardinal tooth inserted between 3b and 3a,

although when it is very prominent the develop-

ment of 38a may be impeded....In the poste-

riorly carinate Pseudotrapezium and the globose

Rollierella 1 is halfway between a tubercle and a

triangular tooth and 3a is quite well developed.”

’ Casey (1952, pp. 134, 136) who provisionally

accepts the genus Hotrapeziwm on other grounds,

considers that the development of 3a is so variable

in the arcticids as to have no taxonomie signifi-

cance. Chavan (1952, p. 83, fft. 1) does not agree

with this conclusion.

FEBRUARY 1954

It seems to the writer that the most important

factor in this grouping is the relative position

of teeth 3a and 1 at the time of their early develop-

ment. If 1 first appears ventral, or posteroven-

tral, to the anterior end of 3a then it may migrate

posteriorly and assume a median position without

coming into conflict with the developing 3a; if it

first appears anterior to the anterior end of 3a

its posterior migration seems to be impeded by

the development of 3a, unless the latter tooth

remains as a thin lamella adjacent to the dorsal

margin of the valve.

An examination of the many illustrations,

often unfortunately diagrammatic, available in

the literature for the old world Jurassic and

Cretaceous -arcticidae, especially the excellent

figures of Cox (1947) and Casey (1952), suggests

that the immediate post-Eotrapezium stage in

the development of the hinge of Veniella would

resemble the hinge of the Middle Jurassic,

Great Oolite species, Anisocardia (Antiquicy-

prina) loweana (Morris and Lycett) (see Cox,

1947, pl. 9, figs. 75, 76; Casey, 1952, text figs.

49a, b, 50) the type species of the subgenus

Antiquicyprina Casey (1952, p. 153). This hinge

is almost identical with that observed in the

smallest right valve of Veniella in our collection,

except that 1 is somewhat stouter in the Creta-

ceous specimen.

The available figures of the Upper Jurassic,

Kimmeridgian species Anisocardia (Anisocardia)

elegans Munier-Chalmas are too diagrammatic

to be of certain help here (see Douvillé, 1921,

text fig. 6; Cox 1947, text fig. 5). They do, how-

ever, suggest that it is within this group of spe-

cies that the continuation of the development

of the Veniellid type of hinge is to be found.

On the other hand, the figures of the Middle

Jurassic Antsocardia (Anisocardia) — truncata

(Morris) given by Casey (1952, text fig. 48a, b)

and said to agree “‘in all features,” in the right

valve at least, with that of A. elegans is clearly

not in an ancestral position. Tooth 1 is very large

and heavy, completely anterior to 3a which has

assumed a position where it crosses the entire

hinge plate in an anteriorly oblique position,

and seems to be displacing 2a of the left valve

in such a way that the latter could not unite

with 2b, as it clearly has at this stage in Veniella.

In the latter genus, 1 impinges against, deflects,

and later almost completely obliterates the

anterior end of AII, not 2a.

VOKES: HINGE OF VENIELLA CONRADI 43

Certain species of the genus Venilicardia

represent well the expectable Lower Cretaceous

stage in the development of the Roudaria and

Veniella-type of hinge. The figures of the Upper

Greensand species Venilicardia lineolata (Sow-

erby) given by Woods (1907, pl. 22, figs. 5a, b,

6a, b, 7, 8) depict a form that in all characteris-

tics save the nature of the left posterior lateral,

PII, agrees with the condition found in “adoles-

cent”? specimens of Veniella conradi similar to

those figured by Stephenson (1923, pl. 66, figs.

4, 5).

If, therefore, we may accept Douvillé’s

postulation of an Hotrapeziwm stage in the an-

cestry of the later Arcticidae, we would have

such a stage represented in the, at present little

known, representatives of Veniella that were less

than 6 mm. in length. At approximately that

length the hinge shows an Antiquicyprina-like

arrangement, that seems to pass through a modi-

fled Anisocardia-like type to enter a Venilicardia-

form arrangement at lengths of 30 to 40 mm.

The fully developed, characteristic Veniella

hinge does not appear until late in the life of

the individual specimen.

It is to be emphasized that this is a series of

stages in the ontogeny of the individual hinge

and that no suggestion of actual evolutionary

ancestry for Veniella is implied or is to be read

into the generic hinge types with which the

growth stages of the Veniellid hinge are here

compared. The genus Veniella is a North Amer-

ican Cretaceous group so far as our present

knowledge of its distribution is concerned,® and

it is quite probable that the ancestral types

were North American in their distribution also.

Our present knowledge of the Jurassic and Lower

Cretaceous pelecypod faunas of this continent

is so very inadequate that it is not possible at

this time to even consider possible ancestral

types for Veniella. Whether the types mentioned

were significant in the ancestry of Rouwdairia

can only be decided by an examination of a

series of immature specimens of species referable

to that genus.

6 The specimens from the Cameroon discussed

and figured by Riedel (1932, p. 54, pl. 1, la, 2, 2a,

2b) as Veniella mortont Meek do not represent

that species, the left hinge of which has been

figured by Stanton (1893, pl. 23, fig. 9), nor do they

seem to be correctly identified generically. Two

genera may be represented, Riedel’s fig. la show-

ing a bifid cardinal 2, while fig. 2b has an entire 2.

44

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VOL. 44, NO. 2

region. Bull. Amer. Paleont. 20 (69): 104 pp.,

11 pls., 2 text figs. 1934.

PERVINQUIERE, L. Etudes de paléontologie tunit-

sienne. II. Gastropodes et lamellibranches des

terraines Crétacés: xiv + 352 pp., 23 pls. Paris ©

1912;

Quaas, A. Beitrag zur Kenntniss der Fauna der

obersten Kreidebildungen in der libyschen

Wiiste. Palaeontographica 30 (2) : 153-336, pls.

20-33. 1902.

RENNIE, J. V. L. Cretaceous fossils from Angola

(Lamellibranchia and Gastropoda). Ann.

South African Mus. 28: 1-54, pls. 1-4, 2 text

figs. 1929.

RENNIE, J. V. L. New Lamellibranchia and Gas-

tropoda from the Upper Cretaceous of Pondo-

land (with an Appendix on some species from

the Cretaceous of Zululand). Ann. South

African Mus. 28: 159-260, pls. 16-31, 3 text

figs. 1930.

RIEDEL, L. Die Oberkreide vom Mungofluss in

Kamerun und thre Fauna. Beit. geol. Erforsch.

deut. Schutzgebiete 16: 154 pp., 33 pls., 47

text figs. 1932.

Stanton, T. W. The Colorado formation and its

Seneniebr aie fauna. U. 8S. Geol. Surv. Bull.

106: 288 pp., 45 pls. 1893.

STEPHENSON, L. W. The Cretaceous formations of

North Carolina. I. The invertebrate fossils of

the Upper Cretaceous formations. North Caro-

lina Geol. and Econ. Surv. 5: xi + ye pp.,

102 pls., 6 text figs. 1923.

SroticzKa, F. The Cretaceous fawna of Soukeen

India. hive The Pelecypoda, with a review of all

known genera of this class. Mem. Geol. Surv.

India, Pal. Indica: xvii + 5388 pp., 50 pls.

1870-71.

THEILE, J., Handbuch der systematischen Weich-

teerkunde 2 (3): 779-1022, text figs. 784-893.

Jena, 1934.

VINCENT, E. Observations sur la place systématique

de Venus pectinifera Sow. Ann. Soc. Roy.

Zool. Belg. 55: 59-62, 2 text figs. 1925.

Wave, B. The fauna of the Ripley formation on

Coon Creek, Tennessee. U. 8. Geol. Surv. Prof.

Pap. 137: 272 pp., 72 pls. 1926.

Woops, H. A monograph of the Cretaceous Lamel-

libranchia of England. Paleontolographical

Soe. 2: 133-180, pl. 20-27. London, 1907.

MYCOLOGY .—Some Discomycetes new to Alaska. Epirn K. Casu, U. 8. Bureau

of Plant Industry, Soils, and Agricultural Engineering.

The extensive collections of fungi made

in Alaska by Dr. Roderick Sprague during

the summer of 1952 included a large number

of Discomycetes which were referred to the

writer for examination. Four collections

were made of an apparently undescribed

Peziza, which is therefore named here as

new. Several species hitherto unreported

from Alaska are proposed as new combina-

tions, and twenty additional species of

Discomycetes are also briefly listed for

which Dr. Sprague’s collections constitute

the first reports from Alaska.

1. Peziza alaskana, n. sp.

Apothecia dispersa, carnea, cupulata, margine

leniter undulato, ex parte in terram arenosam

sepulta, extus fusco-nigra, furfuracea, hymenio

Fepruary 1954

glabro, purpureo-atro vel atro, 5-12 mm in diam.,

3-8 mm alta; asci teretes, apice obtusi et leniter

jodi ope azurescentes, gradatim basim versus

attenuati, 275-3800 x 15-18 uw; ascosporae oblique

uniseriatae, hyalinae, ellipsoideae, utrinque an-

gustatae, subtiliter echinulatae, hyalinae vel

pallide brunneolae; paraphyses numerosi, api-

cibus brunneis et usque 6-8 yu inflatis, in mazae-

dium brunneum agglutinati; textura excipularis

hyphis pallide brunneis, laxe intertextis extus

obseurioribus et furfuraceis composita.

Hab. ad terram arenosam, Alaska.

Apothecia scattered, fleshy, partially buried

in sand, deep cup-shaped, margin slightly un-

dulate, exterior fuscous-black,! furfuraceous,

hymenium smooth, dull purplish black, 5-12

mm in diameter, 3-8 mm deep; asci terete, obtuse

at the apex, faintly blue with iodine, gradually

attenuated toward the base, 275-300 x 15-18 yu;

ascospores obliquely uniseriate, hyaline, ellipsoid,

narrowed at the ends, minutely echinulate,

hyaline to pale brownish, 22-24 x 9-10 u; para-

physes numerous, brown and swollen to 6-8 yu

at the tips, becoming agglutinated into a dark

brown mazaedium; exciple of pale brown, loosely

interwoven, rather thin-walled hyphae, the outer

layer darker and roughened by loose ends or

clumps of hyphae. |

Auaska: Mendenhall area, July 9, 1952, R.

Sprague 3?; Crocker Station no. 1, Mendenhall

Glacier area, July 11, 1952, 20, type; Herbert

Glacier area, July 19, 1952, 100; base of Red

Mountain, Glacier Bay National Monument,

August 12, 1952, 259, and Bear Track Cove,

Glacier Bay National Monument, August 23,

1952, 469. ;

This small black Peziza resembles Peziza

brunneo-atra Desm. in some respects, but the

spores are narrower and more pointed and finely

echinulate rather than verrucose. The apothecia

in P. alaskana also remain deep cup-shaped,

never becoming applanate as in P. brunneo-atra,

and the hymenium is purplish-black, not tinged

with green. The pale brown spores suggest

Aleurina but they are evenly and finely echinu-

late, not reticulate.

1 Color readings are from Ripeway, R., Color

standards and color nomenclature. Washington,

1912.

2 Collection numbers throughout are those of

Roderick Sprague.

CASH: DISCOMYCETES NEW TO

ALASKA 45

comb.

Helvella arctica Nannf. Svensk Bot. Tidskr. 31:

60, illus. 1937.

2. Paxina arctica (Nannf.), n.

ALASKA: Glacier Bay National Monument:

Anchorage Cove area, August 9, 1952, 232, and

Forest Creek area, August 15, 1952, 276 and 282.

Helvella arctica was reported from arctic and

subarctic regions of Sweden and Spitzbergen by

Nannfeldt. The Alaskan collections agree with

the original description and illustrations, and

with type material issued in Lundell & Nannf.

F. Exsice. Suec. 369. The species may be readily

recognized by the furfuraceous white margin

surrounding the black hymenium. If Pazina is

recognized as distinct from Helvella, the species

would belong to the former genus.

comb.

Trichopeziza stipae Fckl. Symb. Mye. p. 297. 1869.

Helotium stigmaion Rehm Hedw. 21: 99. 1882.

Helotium stigmaion Rehm var. minusculum Rehm

Ascom. no. 767. 1883; Hedw. 24: 13. 1885.

Phialea stipae (Fckl.) Rehm Kryptogamenfl. Bd.

1, Abt. 3, p. 734. 1893.

3. Helotium stipae (Fckl.), n.

ALASKA: on Phleum alpinum, base of Red

Mountain, Glacier Bay National Monument,

August 12, 1952, 613; Poa alpina, lake shore,

Mendenhall Glacier area, July 9, 1953, 52; Poa

arctica, Mount Gastineau, July 18, 1952, 264;

Poa compressa, Glacier Bay National Monument,

August 18, 1952, 743; Poa trivialis, Mendenhall

Glacier area, July 10, 1952, 62.

This inconspicuous species, apparently con-

fined to grasses, is reported by Rehm on Stipa

and Phleum. The Alaskan specimens agree with

Thuemen Mycotheca univ. no. 2020 on Phleum

pratense and Krieger F. Saxon. 1835 on an un-

determined grass. No record has been found of its

occurrence in North America.

4. Dasyscypha aspidii (Lib.), n. comb.

Peziza aspidi Lib. Pl. Crypt. Ard. no. 226. 1832.

Trichopeziza aspidit (Lib.) Fekl. Symb. Myce. p.

297. 1869.

Lachnum aspidit (Lib.) Karst.

Faun. Fl. Fenn. 16: 27. 1888.

Meddel. Soc.

ALASKA: On Dryopteris sp., Sebree Island,

Glacier Bay National Monument, August 19,

(1952, 369.

As pointed out by Dennis, the use of the generic

name Lachnum sensu Rehm for species of Dasy-

scypha with lanceolate paraphyses has no justifi-

cation; the species is therefore referred to Dasy-

scypha.

46 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

In addition to the fungi listed above, Dr.

Sprague’s 1952 collections include the following

Discomycetes not previously reported from

Alaska: Ascobolus glaber Pers. ex Fr. on grizzaly-

bear dung, Belonioscypha campanula (Fr.) Rehm

on Hordeum, Dasyscypha calyculiformis (Schum.

ex Fr.) Saec. on Salix, D. lewcophaea (Pers. ex

Weinm.) Mass. on Lupinus nootkatensis, D.

virginea (Batsch ex Fr.) Fekl. on Populus tricho-

carpa and Salix alaxensis, Heloteum caudatum

(Karst.) Vel. on Alnus, H. cyathoideum (Bull.

ex Fr.) Karst. on Equisetum virgatum, Epilobium

latifolium, and Bosnickia, H. leucellum Karst. on

Alnus, H. scutula (Pers. ex Fr.) Karst. on Dryas

voL. 44, No. 2

drummondu and Epilobium, H. virgultorum (Vahl

ex Fr.) Fr. on Alnus and Sambucus, Humaria

hemisphaerica (Wigg. ex Fr.) Fekl., H. wmbrorum

(Fr.) Fekl., Lamprospora amethystina (Quél.)

Seaver, L. constellatio (Berk. & Br.) Seaver, and

Mollisia uda (Pers. ex Fr.) Gill. on Alnus,

(Cke.) Rehm, Pyrenopeziza ~

karstenit Sace. on Agropyron trachycaulum and

Poa, Rutstroemia nervisequia (Schroet.) W. L.

White on Alnus, Stamnaria persooni (Moug.

ap. Pers. ex Fr.) Fckl. on Equisetum, and Tapesia

fusca (Pers. ex Fr.) Fekl. on Alnus, Salix, and

Shepherdia.

Otidea auricula

ZOOLOGY .—Description of Eocyzicus concavus (Mackin) with a review of other

North American species of the genus (Crustacea: Conchostraca). N. T. Marrox,

University of Southern California.1 (Communicated by F. A. Chace, Jr.)

In a key to the phyllopods of Oklahoma

and neighboring states, Mackin (1939)

listed a previously undescribed species under

the name Estheria concava. As a result of

personal communications Dr. Mackin in-

formed me that the original four specimens,

on which the key characters were based,

had been lost. However, another collection

from the same locality contained eight speci-

mens which Dr. Mackin kindly presented to

me for study. Careful examination of these

specimens resulted in the unquestionable

decision that the species should be assigned

to the genus Hocyzicus Daday, 1915. I was

then asked by Dr. Mackin to make a com-

plete description of this unusual and in-

teresting conchostracan.

Meanwhile there appeared in the key to

the North American phyllopods by Pennak

(1953) a listing of a species, presumably the

species here under consideration, indicated

as Hocyzicus concava Mattox. The original

designation by Mackin must be recognized

even though it was based on the following

incomplete diagnosis: ‘‘Rostrum shaped like

a hatchet blade; with a row of large smooth

spines along the mid-dorsal line, one spine

for each trunk segment; hand of the male

deeply incised at the base of the thumb;

shell sway-backed.’’ The diagnosis given

by Pennak was: ‘‘Rostrum like hatchet blade;

1 Department of Zoology, Allan Hancock Foun-

dation. Allan Hancock Foundation Contribution

no. 1243 :

with large, smooth spine on the middorsal

line of most trunk segments; rare, poorly

known; Okla.”’ The specific name must be

that of Mackin even though the generic

designation is invalid. EHstherza Ruppell

1837 as used for the Conchostraca is a

homonym, as the name Hstheria was first

used for a genus of Diptera by Robineau-

Desvoidy in 1830. The name LHstheria, for

conchostracans, is replaced by Cyzicus

Audouin, Hocyzicus Daday, Caenestheria

Daday, Caenestheriella Daday, Leptestheria

Sars, Holeptestheria Daday, Leptestheriella

Daday and Cyclestheria Sars. The original

trivial name of the species under considera-

tion, must be changed to agree with that

of the genus, hence the name Locyzicus

concavus (Mackin, 1939) is here given. Since

the species has not previously been com-

pletely described a description is here pre-

sented and a neotype is designated. These

animals were collected on August 12, 1928,

in a temporary pool near Summerfield, Tex.

Eocyzicus concavus (Mackin)

Description.—Male: The shell is elliptical with

a straight dorsal hinge line extending two-thirds

the shell length, and with a rounded ventral

margin (Fig. 1, a). Posterior to the hinge the

dorsal edge is straight extending ventrally at

approximately a 20° angle. The anterior shell

margin is rounded, extending ventrally very

abruptly; the posterior portion is more attenu-

ated. The greatest height of the shell is slightly

Fepruary 1954 MATTOX:

anterior to the middle. The umbone is very con-

spicuous extending dorsally above the hinge line

and located approximately one-fifth the shell

length from the anterior edge. The anterior slope

of the umbone extends abruptly ventrad, the

posterior slope is more gradual. The form of the

umbone gives a ‘“‘sway-backed” appearance to

EOCYZICUS CONCAVUS 47

the shell. The lines of growth on the six male

shells in the collection varied in number from 18

to 22. The average shell size was 6.9 by 4.2 mm,

a shell width-length ratio of 1:1.6.

The head of the male possesses the characters

typical of the genus as established by Daday

(1915). Those characters are the roundly ex-

Wy

SA &

G

S as

LA WSN ea.

cere new =

ae oe

a

Ene: 1.—Bocyzicus concavus (Mackin): a, Shell of male; 6, first gnathopod of male; c, second

enathopod of male (4, 5, 6—fourth, fifth, sixth endite); d, lateral view of dorsal portion of trunk

and the telson; e, lateral view of head of male; f, lateral view of head of female; g, male third ap-

pendage; h, female ninth appendage with eggs attached to epipodite. Seales all equal 0.5 mm.

48 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

tended and shallowly notched occipital region,

also the broadly spatulate rostrum when viewed

in profile (Fig. 1, e). The dorsal surface of the

front is slightly concave. The anterior flagellum

of the second antennae is composed of 12 seg-

ments, the posterior flagellum of 14 segments.

The first antennae are elongate, dorsally papil-

lose, up to 20 papillae, and extend to the seventh

segment of the second antennae.

The body, or trunk, is composed of 19 seg-

ments each bearing a pair of appendages. The

posterior 11 segments bear a middorsal, smooth

spine; the anterior 8 segments bear no spines

(Fig. 1, d). The first two pairs of appendages are

developed into gnathopods, typical of the group.

The palpiform “thumb” of the fourth endite of

the first pair of gnathopods is notched at its base

(Fig. 1, b). The digitiform fifth endite is shorter

than the sickel-shaped sixth endite. On the second

gnathopod the notch at the base of the thumb is

not as pronounced as in the first (Fig. 1, c). The

fifth endite is approximately one-third longer than

the sixth. The third pair of appendages is folia-

ceous, as are all the others. The fifth endite of

the third appendages possesses a digitiform ex-

tension approximately twice the length of the

sixth endite (Fig. 1, g).

The telson is truncate, shortened, with a pair

of ventral, curved cercopods extending pos-

teriorly beyond the dorsal telson spines (Fig. 1,

d). The telson dorsal ridges possess 16 or 17

spines of variable length. The dorsal forked fila-

ment arises between the third and fourth telson

spines.

Female: The shell of the female is similar in

form to that of the male; no marked dimorphism

VoL. 44, No. 2 ©

is evident. On the two female shells in the collec-

tion each contained 22 growth lines.

The head of the female has the form charac-

teristic of the genus. The rostrum, in profile, is

accuminate; the occipital area is rounded and

with a shallow notch (Fig. 1, f). The front is

straight, not concave as in the male. The second

antennae are as in the male, the flagellae are 12

and 14 segmented. The first antennae are shorter

than in the male extending only to the fourth

segment of the second antennae and possess 15

to 18 dorsal sensory papillae.

The body appendages, 19 pairs, are all similar

in form, the typical fohaceous swimming legs.

The ninth and tenth pairs have the epipodite

elongated for the ovigerous function. The epipo-

dite of pair nine is approximately one third longer

than that of pair ten (Fig. 1, h).

The last 9 segments of the body possess a mid-

dorsal spine, the anterior 10 segments are smooth.

The telson is similar to that of the male with 16

pairs of dorsal spines.

Type locality —Summerfield, Texas.

Type.—Deposited in the U. 8. National Mu-

seum. Neotype, male, U.S.N.M. no. 95731. As

the original specimens of Mackin were lost it is

necessary to designate the type as a neotype.

One male and one female U.S.N.M. no. 95732,

also deposited.

Remarks.—Eocyzicus concavus represents the

third species of the genus to be described from

this continent. H. diguett (Richards, 1895) was

described from Purissima, Calif., and EH. van-

hoffent Daday, 1915, was described from Mexico,

the exact locality being uncertain. In the absence

of specimens for direct examination the descrip-

E. vanhoffent |

E. digueti E. concavus

SHELL 8.5 X 3.5 mm. 8.6 X 4.8 mm. 6.9 X 4.2 mm.

1:2.4 shell size ratio 1:1.8 ratio 1:1.6 ratio

14-16 growth lines 26 growth lines 18-22 growth lines

Male and female similar Male shell more rounded dorsally Similar in two sexes

HeEapD Female rostrum elongate and | Female rostrum short, roundly ac- | Female rostrum regularly accumi-

sharply accuminate cuminate nate

Occipital notch of female deeper | Occipital notch of male deeper than | Occipital notch of two sexes equal

than in male in female

Second antennae flagella 12-13 seg- | Second antennae flagella 14-16 seg- | Second antennae flagella 12-14 seg-

ments ments ments

First antennae with 13-18 papillae First antennae with 14 papillae First antennae with 15-20 papillae

TRUNK 16 segments 16 segments 19 segments

14 posterior segments with dorsal | 14 posterior segments with dorsal | 9-11 posterior segments with dorsal

spine spine spine

Telson with 12-15 pairs of spines 15 pairs of telson spines 16-17 pairs of telson spines

APPENDAGES Male “thumb” of gnathopod | Male ‘‘thumb’’ with deep basal cleft | Male ‘‘thumb’’ with shallow cleft

cleft deeply

Tenth epipodite of female twice | Ninth epipodite of female twice | Ninth epipodite of female one-third

length of ninth length of tenth longer than tenth

Fespruary 1954

tion given by Daday for vanhéffeni has been used

as a basis of comparison. The comparison with

digueti has been facilitated by a recent acquisition

of a collection of 3 males and 1 female from near

Reno, Nev. This represents a new locality record

for this species as it had been previously known

from only the type locality. In addition to the

characters indicated in the foregoing table,

concavus differs from digueti in a number of other

features. The hinge line is proportionately much

shorter, the umbones are less prominent in diguett,

and it does not have the “sway-back” appear-

ance of concavus. In digueti the first antennae of

the male extend only to the fifth or sixth segment

HOFFMAN: AMERICAN MILLIPEDS

OF FAMILY EURYURIDAE 49

of the second antennae; the telson is more trun-

cate, and the cercopods are proportionately

shorter than in concavus. Also, the “thumb” of

the male gnathopods of digueti are much more

deeply cleft at the base than in concavus. A tabu-

lated comparison of the three species is given

on the opposite page.

REFERENCES

Dabay DE Drss, E. Monographie systematique des

phyllopodes conchostraces. Ann. Sci. Nat. Zool.,

ser. 9, 20: 39-330. 1915.

Mackin, J. G. Key to the species of Phyllopoda of

Oklahoma and neighboring States. Proc. Okla-

homa Acad. Sci. 19: 45-47. 1939.

PEnNAK, R. W. Fresh-water invertebrates of the

United States: 769 pp. New York, 1953.

ZOOLOGY .—Further studies on American millipeds of the family Euryuridae

(Polydesmida). Ricuarp L. Horrman, Clifton Forge, Va.

My previous paper dealing with the

American euryurids (1951) endeavored to

provide‘a summary of the genera recognized

by me at the time of its writing in early

1950. Since that time I have accumulated

additional pertinent information and have

come to realize that my reliance upon

Attems’s treatment of the group in Das

Tierreich (1938) was in many instances ill-

advised. In all, so many changes are neces-

sary in the arrangement of the genera of

this family that a second paper becomes

advisable. While aware of the limitations

imposed by the acute lack of critical study

material, I am nonetheless convinced that

even preliminary attempts at synthesis are

badly needed at present. Half a loaf is better

than none at all. é

In the light of the preceding comments

it may seem improper to refer to the works

of the late Carl Attems in any vein other

than one of utmost respect. Attems was the

only recent worker with the industry and

ability to produce manuals of the scope of

his 3-volume ‘“Polydesmoidea,’’ yet while

this magnificent compilation stands as a

memorial to its gifted author, its minor

imperfections will long be the despair of

the uncritical user. Outstanding are Attems’s

disregard of the works of some of his col-

leagues (notably Cook and Silvestri), and

a most remarkable indifference to the

principles of type fixation. Some of these

idiosyncrasies will be noted further on in

the text. |

Aside from information gathered from

the literature, I have based this paper to a

considerable extent upon Central American

specimens preserved in my personal col-

lection and that of the United States

National Museum. This study material is

very uneven as regards the genera repre-

sented. In the case of Pseudamplinus, I

have adequate material to justify the prepa-

ration of a generic revision, which is now in

progress. There seems to be no advantage

in delaying the descriptions of the various

new forms in other genera, however; these

are given herewith, with at least a modicum

of attention to their relationships to es-

tablished species. In general the present

paper is concerned with changes and addi-

tions on a generic level. It is assumed that

the reader has access to the earlier paper

mentioned above, which lists the species in

the various genera not dealt with here.

It is with pleasure that I must again

mention my increasing indebtedness to

Dr. E. A. Chapin, for access to the collec-

tions of the National Museum and for much

advice and information pertinent to the

completion of the present study. I am also

grateful to Dr. W. J. Gertsch, through whose

cooperation I was able to examine material

in the collection of the American Museum

of Natural History.

Family EuRyvRIDAE Pocock

Trachelorhacidae Silvestri, Boll. Mus. Torino

13 (324): 5. 1898 (based upon T'rachelorhacis

Silvestri, a preoccupied name).

50 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

Euryurinae Pocock, Biologia Centrali-Americana,

Chilopoda and Diplopoda: 1138, 147. 1909.

Euryurini Brolemann, Ann. Soc. Ent. France

84: 584. 1915.

Euryuridae Chamberlin, Bull. Mus. Comp. Zool.

62: 249. 1918.— Hoffman, Proc. U. 8. Nat. Mus.

102: 235. 1951.

Genera.—18, of which 13 are found in the

Western Hemisphere.

Range—North Carolina, western Pennsyl-

vania, and Minnesota south through Central

America to Ecuador, northeastern Peru, and

northern Brazil. Celebes, Halmaheira, Ternate,

Timor, and the Solomon Islands.

Definition —A small family of the suborder

Eurydesmidea (= Leptodesmidi of Brolemann,

1915), resembling the Platyrhacidae in the pres-

ence of (1) closely-set antennae each subtended

by an oval swelling, (2) small sternites with the

legs basally approximate, (3) a broad distally

truneate terminal segment (telson), and (4)

small, rather simple gonopods in a proportionately

small, somewhat diamond-shaped aperture. The

gonopods are similar in lacking a definite pre-

femoral process. From the Platyrhacidae, the

members of this family differ chiefly in that the

repugnatorial pores are located in the sides of

definite marginal swellings of the keels. The pores

in species of the Platyrhacidae are more or less

remote from the edges of the keels, and are en-

circled by a flat polished area; the keels them-

selves have no marginal ridges or thickenings.

In the absence of any intermediate condition,

it seems best to follow the example of my prede-

cessors, who have found it expedient to set the

euryurid genera apart in a separate family.

Attems, however, lumps the two groups in his

recent (1938) survey.

With the increasing significance that is being

attached to the taxonomic value of the male

genitalia, it appears possible to further divide

the euryurids into several groups of genera, each

rather discrete and readily defined. Since the

recognition of such ensembles is a substantial aid

to an understanding of the affinities of the genera,

I think it advisable to designate them by the use

of subfamily names, proposed towards the con-

clusion of the discussion.

A comment on the use of the family name may

be of interest. In 1938 Attems proposed the name

Eutheatus as a substitute for Huryurus of Koch,

1847, as the latter name had already been pub-

lished in 1815 by C. 8. Rafinesque for a group

of polychaete annelids. I have examined a copy

of the Rafinesque paper, the Analyse de la nature

voL. 44, No. 2

ou Tableau de lVunivers et des corps organises

(Palerme, 1815), and find that the name Euryurus

is given merely in a list of generic names. There

is no description, no reference to a description,

and no specific name cited, and the name may be

regarded a nomen nudum. According to the Inter-

national Rules of Zoological Nomenclature,

generic names of this nature may be, if properly

proposed, used again for other forms by later

workers. Hutheatus in consequence was proposed

without justification, it becomes a junior synonym

of Huryurus Koch, 1847.

Genus Amplinus Attems

Amphinus (sic) Attems, Denk. Akad. Wien 68:

281, 396. 1899 (as subgenus of Pachyurus).

Polylepiscus (in part) Attems, Das Tierreich 69:

300. 1938 (not Polylepiscus in the sense of

Pocock 1909).

Phinotropis Chamberlin, Bull. Amer. Mus. Nat.

Hist. 78: 499. 1941—Hoffman, Proc. U. 8. Nat.

Mus. 102: 239. 1951 (type, Phinotropis tidus

Chamberlin).

Type.—Pachyurus (Amplinus) kalonotus At-

tems, by designation of Pocock, 1909.

Diagnosis —A euryurid genus characterized

as follows: head with prominent subantennal

swellings, collum as wide as second segment,

tergites with prominent quadrate areas, telson

broadly quadrate in shape, preanal scale sub-

acuminate or rounded and with very small lateral

setiferous tubercules.

Male gonopods with elongate trachial rods,

coxae somewhat rounded-elongate, prefemur and

femur fused into a short, straight trunk, a long

slender solenomerite and an elongage laminate

tibiotarsus are set off from the femur by a per-

ceptible joint.

Range-——Upper Amazonian basin, in western

Brazil, eastern Ecuador, and northeastern Peru.

Species.—Five, listed in my 1951 paper under

Phinotropis.

Remarks.—As originally proposed, Amplinus

included six species: Pachyurus kalonotus Attems

and P. acuticollis Attems, Polydesmus klugiw

Brandt, P. erichsont Brandt, P. abstrusus Karsch,

and P. ater Peters. No type species was desig-

nated. In 1909 Pocock selected kalonotus as type,

and added six more species from Central America

to the genus. But within the genus as understood

by Pocock there were representatives of two

different groups. In one of them (including

kalonotus, acuticollis, and presumably ater) the

preanal seale is of the usual polydesmoid form—

subtriangular to semicircular. In the other group,

FEBRUARY 1954 HOFFMAN: AMERICAN

however, it is trapeziform in shape, that is to say,

distally truncate instead of acute or rounded,

with the caudal margin parallel to the basal edge

and with the lateral setiferous tubercules con-

siderably enlarged. This difference has come to

be unanimously regarded by students of diplopods

as one of generic value. It is very unfortunate

that the name Amplinus has been misapplied,

by all workers subsequent to Pocock, to the group

having the truncate or concave preanal scale.

In his most recent treatment of the poly-

desmoids (Das Tierreich, Lief. 68-70) Attems

erroneously cites Polydesmus klugiw of Brandt

as type of Amplinus, and, moreover, places

kalonotus and its relatives in Pocock’s genus

Polylepiscus. That this association is untenable

taxonomically as well as nomenclatorially is evi-

denced by differences in the gonopods of the

various species involved. The Guatemalan species

included by Pocock in Polylepiscus have, in addi-

tion to the spiculiform solenomerite, a somewhat

similar branch from the base of the tibiotarsal

blade. The South American species under con-

sideration lack this additional process, and the

tibiotarsus of their gonopods is also longer and

more sinuate. Realization that they could not

be properly placed in Polylepiscus induced me to

group them under Chamberlin’s name Phino-

tropis, based upon P. tidws—a Peruvian species

which is obviously congeneric with kalonotus.

So the discovery that kalonotus is the true type

species of Amplinus requires relegation of Phino-

tropis to the status of a junior synonym, and

proposal of a new generic name for the Central

American species heretofore called Amplinus.

In reference to its mistaken identity, the group

may be called

4

Pseudamplinus, n. gen.

Type-—Amplinus orphinus Chamberlin 1922,

by present designation.

Diagnosis.—A euryurid genus characterized

as follows: head with prominent subantennal

swellings, collum as wide as second tergite,

tergites strongly tesselated in most species, telson

broadly truncate distally and quadrate in ap-

pearance, preanal scale trapeziform in shape

with the lateral setiferous tubercules very large

and the margin between them straight or concave.

Male gonopods with very long slender trachial

rods; coxae rather slender; prefemora and femora

fused into a straight trunk; a slender blade-like

solenomerite and a thin flattened tibiotarsal

branch, both directed at nearly a right angle to

MILLIPEDS

_

OF FAMILY EURYURIDAE 51

the femoral portion, are set off from this by a per-

ceptible joint or suture, both of these terminal

elements are directed away from the coxal joint.

This is the only genus of the family having

the truncate or distally concave preanal scale

which is so characteristic of the Platyrhacidae,

and may be regarded as a sort of intermediate

group. In the form of the keels, however, the

relationship is clearly with the other euryurids.

Range-—Middle America, from southern

Mexico (Guerrero and Vera Cruz) south to

Costa Rica, and also northwestern Venezuela.

Most of the species occur in Guatemala and in

Vera Cruz.

Species.—23, as follows: abstrusus (Karsch),

areatus (Pocock), armatus (Pocock), beebei

(Chamberlin), converus (Carl), crenus (Chamber-

lin), eutypus (Chamberlin), erichsont (Brandt),

flavicornis (Pocock), klugw (Brandt), leon (Cham-

berlin), manni (Chamberlin), nitews (Chamberlin),

nitidus (Brolemann), orphinus (Chamberlin),

palicaudatus (Attems), pococki (Cook), schmidti

(Chamberlin), tajywmulco (Chamberlin), tapa-

chulae (Chamberlin), triramus (Pocock), xelitus

(Chamberlin), zuniulus (sic) (Chamberlin).

Remarks.—Inasmuch as no members of this

genus have yet been collected in Panama, the

presence of two rather similar forms in north-

western Venezuela is of some interest. These

species, abstrusus and beeber, agree with each

other and differ from other members of the genus

in having a rather shortened solenomerite.

Karsch’s type came from Puerto Bello, Chamber-

lin’s from Rancho Grande, about 35 miles to the

south. The similarities between the two suggest

synonymy, or at best a subspecific relationship.

Redescription of several poorly known species,

based upon material in my possession, is planned

for inclusion in my forthcoming revision of this

genus.

Although Pocock clearly disposed of the

mystery surrounding the matter of the type

species of Orthomorpha—designating Polydesmus

beaumonti Le Guillou following the elimination

of all other species from the genus Paradesmus

by Saussure, a remarkable effort was made by

Cook in 1911 to restrict the name Orthomorpha

to members of the present genus! But Cook’s

proposal was based upon a highly subjective line

of reasoning having absolutely no basis in fact,

and no subsequent writer has ever discussed or

even referred to it. The matter, in brief, may be

summarized as follows: in 1859 the genus Para-

desmus was erected by Saussure for five species

52 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

distributed in three groups. The first contained

only Polydesmus carolinensis, the second P.

klugu, P. erichsoni, and P. pictett, and the third

only P. beaumonti. As Pocock pointed out, one

of these five species must be the type, the addi-

tion of coarctatus in 1860 having no bearing on

the question. In 1869 Humbert and Saussure

provided the name Euryurus of Koch for caro-

linensis, and Pachyurus for the three species of

the second group, leaving beawmonti as the only

species in Paradesmus. Cook observed that since

Saussure had implied in 1859 that beaumont

was not a typical member of the genus Para-

desmus (in the sense that it differed considerably

from the other species), that generic name, and

its replacement Orthomorpha) should be properly

applied to the members of the second. group—

klugit, erichsoni, and pictete.

On the other hand, Cook also pointed out that

the species described and illustrated as klugit

by Pocock in the Biologia differed in several .

respects from the type specimen of the species,

which he had seen in the Berlin Museum, and

suggested the name pococki for the form treated

in the Biologia as klugiit. Cook’s perception of

specific differences was a good one, and it may be

reasonably admitted that he was correct in this

particular. I therefore list pococki in the roster

of species given above. It is to be hoped that ma-

terial from Alvarado, Vera Cruz (the type locality

of klugit) will soon be forthcoming to permit a

final settlement of the confusion which surrounds

application of the name.

Genus Polylepiscus Pocock

Polylepiscus Pocock, Biologia Centrali-Amer-

icana, Chilopoda and Diplopoda: 154. 1909.

Euplinus Chamberlin, Ann. Ent. Soc. Amer. 45:

578. 1952 (type: EH. volcanicola Chamberlin, by

original designation).

Polylepiscus appears to be a genus more or

less endemic to the Guatemalan plateau, and

individuals seem to be scarce, to judge from the

few preserved in museum collections. It is note-

worthy that no additions to the genus have been

made since Pocock’s original description and

account in the Biologia'. It is to that reference,

1 Kuplinus volcanicola has recently been de-

scribed as a new genus and species and was con-

trasted only with Amplinus in the generic diag-

nosis. One can never understand why Polylepiscus,

beautifully illustrated and thoroughly described

in the Biologica, was not taken into consideration.

Not only is Huplinus a junior generic synonym,

but furthermore I can not see how volcanicola

differs in any respect from P. furcifer Pocock. —

VOL. 44, No. 2

therefore, that appeal must be made for informa-

tion pertinent to the allocation of the new forms.

Pocock found that the genus was divisible

into two groups on the basis of several correlated

structural features. The following key is based

upon that presented by him, with a few changes.

KEY TO THE KNOWN FORMS OF POLYLEPISCUS

1. Pores of 19th segment completely lateral;

metatergites smooth or nearly so, polygonal

areas well-defined or obsolete but not ob-

scured by granulation of dorsum........ 2

Pores of 19th segment on dorsal side of keels;

dorsum rugulose or granular, the polygonal

areas obseured!\.s.; aus ot 4

2. Dorsum entirely smooth, no trace of areas ex-

cept a few vaguely defined on keels; a mid-

dorsal row of large oval spots present

trimaculatus, n. sp.

Dorsum with polygonal areas well defined;

no middorsal row of spots................. 3

3. Polygonal areas smooth, not tubercular, ex-

cept obscurely so on the keels

stoll1 Pocock

Polygonal areas manifestly tubercular

furcifer Pocock

4. Keels of posterior half of body with anterior

border basally produced, posterior border

from 13th to 18th distinctly shouldered at

base, 19th tergite granular

actaeon Pocock

Keels of posterior half of body with anterior

border less produced; posterior border

strongly concave, not shouldered, 19th ter-

gite granular only posteriorly or smooth. 5

5. Tibiotarsus of male gonopod less curved, not

crossing behind tip of tibiotarsal process;

19th tergite granular over posterior half

h. heterosculptus Carl

Tibiotarsus of male gonopod more acutely

bent, in mesial aspect crossing behind tip of

tibiotarsal process; 19th segment almost

entirely smooth...... h. pococki, n. subsp.

It is to be especially noted that, with the excep-

tion of stolli (from Cholhuitz, Guatemala), the

previously known species have not been recorded

from any definite locality. A considerable amount

of work thus remains to be done in the way of

defining the ranges of the different forms. Ma-

terial of trimaculatus, on the other hand, has

been obtained at several localities; and hetero-

sculptus pococki is here described from north-

eastern Chiapas, extending the known range of

the genus slightly outside the political limits of

Guatemala.

Polylepiscus furcifer Pocock

Polylepiscus furcifer Pocock, Biologia Centrali-

Americana, Chilopoda and Diplopoda: 156, pl.

12, figs. 1-1h. 1909. -

FEBRUARY 1954. HOFFMAN:

Euplinus volcanicola Chamberlin, Ann. Ent. Soc.

Amer. 45: 578, fig. 48. 1952.

The type locality of volcanicola, Volein Taju-

muleo, Guatemala, provides the first definite

locality from which furcifer has been taken.

Pocock’s type specimen was without any locality

data.

Polylepiscus trimaculatus, n. sp.

Fig. 2

Type specimen.—Male holotype, U. 8. Nat.

Mus. no. 2098, from Sepaciute, Guatemala, col-

lected in March 1902 by O. F. Cook. Allotype

a female with the same collection data.

Diagnosis—Readily separable from the other

members of this genus by the smooth dorsum,

trimaculate color pattern, and configuration of

the male gonopod. The lateral position of the

pores allies this form with stolli and furcifer. The

prefemerofemoral portion of the gonopod is pro-

portionately smaller in relation to the tibiotarsus

than in the other species of which males are

known.

Description.—The general body form coincides

closely with the descriptions given by Pocock.

The following notes were made from the type

specimens:

Length of male, 62, width, 11 mm.; length of

female, 60, width, 9.5 mm. Dorsum entirely

smooth except for faint indications of areas on

the keels; metazonites considerably raised above

level of prozonites. Keels of segments 3-19 pro-

duced caudad, their caudal margins concave and

finely serrate; pores all completely lateral. Telson

broadly rounded. Preanal scale subtriangular

but distally rounded. Pleurites coarsely granular,

with a series of acute tubercules along the caudal

margins, becoming larger on the posterior seg-

ments, frequently a small cluster of spines just

below the projection of the keels.

Sternites and basal segments of legs smooth

and glabrous. The raised area between the leg-

pairs of each segment impressed by a longitudinal

and a-transverse furrow, creating a small tumid

area at the base of each leg. Distal segments of

legs sparingly hirsute. Bases of last pair of legs

almost in contact.

Gonopod aperture of male rather small and

ovoid, with raised margins. Gonopods with long

slender trachial rods; the coxae small and cylindri-

cal. Basal half of telopodite densely setose. Distal

half set off by a distinct groove or suture. Solenom-

erite long and slender, unmodified, very slightly

sinuous. Tibiotarsus somewhat broader, laminate,

AMERICAN MILLIPEDS OF FAMILY

EURYURIDAE 53.

its distal end bent at a right angle; tibiotarsal

process slender, unmodified, gently curved distad

toward the tibiotarsus.

Color of the preserved specimens as follows:

dorsum chocolate brown to blackish; underparts

light brown to tan. All of the upper surface of

the keels and a large ovate median spot on each

segment lighter. Collum with an _ hourglass-

shaped mark. A label with the specimens reads

“Spots, carinae, legs, and antennae pale grayish,

nearly white’, this presumably referring to the

condition in life.

Remarks.—Additional material of trimaculatus

has been seen from the following localities:

GUATEMALA.—Pancajche, several taken in May

1905 by G. P. Goll; Trece Aguas, April 21, 1906

and June 1907 by O. F. Cook (all U.S. N. M.).

Polylepiscus heterosculptus pococki, n. subsp.

Fig. 1

Type specumen.—Adult male holotype, U. S.

Natl. Mus. no. 2099, from Tumbala, State of

Chiapas, Mexico, collected on June 20, 1906,

by O. F. Cook.

Diagnosis.—Very similar to P. h. heterosculptus,

from which it may be distinguished by the con-

trasting features given in the following couplet:

Size large, length 80-90 mm, width, 13-15 mm;

dorsum with conspicuous transverse rows of

tubercules; caudolateral corners of keels of

segments 5-16 produced caudally; keels, legs,

and antennae reddish brown to blackish brown;

19th tergite granular posteriorly; tibiotarsus

of male gonopod slender, distally recurved

away from the solenomerite

h. heterosculptus Carl

Size smaller, length 60-65 mm, width, 9-11 mm,

dorsal rows of tubercules less pronounced;

caudolateral corners of keels of segments 4-16

produced caudad; legs, antennae, and keels

yellowish; 19th tergite entirely smooth; tib-

iotarsus of gonopod heavier, crossing behind

tip of solenomerite....h. pococki, n. subsp.

With the foregoing exceptions, pococki agrees

so well with the excellent description of hetero-

sculptus given by Carl that a description of my

type series seems unnecessary. The differences

between the two forms, while not impressive

singly, are sufficient when taken in combination

to warrant separation of a new subspecies. It is

to be regretted that we are ignorant of the prov-

enance of Carl’s material.

This form is named in recognition of R. I.

Pocock’s outstanding contribution to our knowl-

edge of the diplopod fauna of Central America.

o4 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

Genus Colomborus Chamberlin

Colomborus Chamberlin, Ann. Ent. Soc. Amer. 45:

587. 1952.

Type.—Colomborus martanus Chamberlin (=

Pycnotropis colombiensis Chamberlin, 1928).

Range.—Colombia.

Species.—1.

Remarks.—There is no doubt that the type of

this genus is worthy of systematic recognition.

It is interesting to note, however, that Colomborus

was proposed as a monotypic genus based upon

a new species, despite the fact that this species

had obviously already been described. One must

presume that Chamberlin, in working up the

material for his 1951 paper, did not even take

into consideration his own earlier (1923) con-

tribution on the millipeds of Colombia in which

the species was first recognized under the name

Pycnotropis colombiensis. I am unable to discern

any specific differences in the illustrations of the

male genitalia (19238: fig. 112; 1952: fig. 42).

Genus Protaphelidesmus Brolemann

Protaphelidesmus Brolemann, Ann. Soc. Ent.

France 84: 559. 1915—Attems, Das Tierreich

69: 313. 1988—Hoffman, Proc. U. 8. Nat. Mus.

102: 240. 1951.

Ptyxogon Chamberlin, Bull. Amer. Mus. Nat.

Hist. 78: 500. 1941.—Hoffman, Proc. U. 8.

Nat. Mus. 102: 240. 1951 (type: P. incus Cham-

berlin, by original designation).

Type.—Platyrrhacus ligula Brolemann.

Ranye.—Venezuela; northeastern Peru.

Species.—3: ligulus Brolemann, incus Cham-

berlin, levigatus Attems.

Remarks.—Since completing my first paper on

this group, in which Ptyxogon was held distinct

on the basis of smooth tergites, I have carefully

reconsidered the drawings and descriptions of all

three of the species involved, as well as the type

of P. incus, and no longer believe that two genera

can be recognized. Jeekel, writing on East Indian

strongylosomoids, has recently emphasized the

intrageneric variability of body form, and re-

iterated the primary importance of the male

gonopods. I have reached the same conclusions

from recently acquired familiarity with several

groups of tropical diplopods. The sculpture of

the tergites, without substantiating genital

features, can scarcely be considered of generic

value,

Genus Seminellogon Chamberlin

Seminellogon Chamberlin, Pan-Pacific

(1): 18. 1933.

Ent. 9

voL. 44, No: 2

Type.—sS. chitarianus Chamberlin, by original

designation.

Range.—Costa Rica; Panama.

Species.—2: chitarianus Chamberlin,

azulensis, Ni. sp.

Remarks.—This genus was overlooked in the

preparation of my previous paper, primarily be-

cause it was described as being closely related to

Aphelidesmus and thus listed by Attems in 1937

amongst the “unsichere Gatturgen’’ of the

Strongylosomidae. Seminellogon is of course a

euryurid, and very close to Amplinus, into which

it and several others may have to be withdrawn.

A possible generic character, of unproven value,

is the location of the pores on the under side of

the keels of the 18th and 19th segments. In this

respect it differs from the Central American

genera, but I have not been able to study the

South American forms in this respect.

Seminellogon heretofore has been monotypical.

A new species is here described from the high

mountains of northwestern Panama, and based

upon fairly large and homogeneous series. Con-

sidering the quality level of the differences, how-

ever, it would not be surprising if the new form

is later shown to be a subspecies of chitarianus.

cerro-

Seminellogon cerroazulensis, n. sp.

, Fig. 3

Type specimens.——Male holotype, U. S. Nat.

Mus. no. 2100, from Cerro Azul, Province of

Chiriqui, Panama, collected on March 26, 1911,

by E. A. Goldman. Male and female paratypes

from Boquete, Chiriqui Province, Panama,

January 1940, W. C. Wood (Amer. Mus. Nat.

Hist. A-7175) and from El Volcan, Chiriqui

Province, Panama, February 28, 19386, W. J.

Gertsch (Amer. Mus. Nat. Hist.).

Diagnosis —Differing from S. chitarianus pri-

marily in the characters of the male gonopod,

such as the longer and less curved solenomerite;

the much more slender tibiotarsus with a promi-

nent broadening near its base; and the distinctly

shorter basal, setose, portion of the telopodite.

Description.—Agreeing in most respects with

the description of chitartanus given by Chamber-

lin. The following specific notes were made from

the type specimen.

Length, 44, width, 6 mm. Dorsum, sides, and

ventral surfaces all entirely smooth and shining,

no evidence of granulation or other roughening.

Front edge of keels not shouldered, but forming

an even are back to the caudolateral corner.

FrBRuUARY 1954

Latter very little produced except on the last

few segments. Caudal margin of keels nearly

straight, not concave or shouldered basally.

Pores lateral on all keels except the 18th and

19th, where they are distinctly inferior, on the

under side. Telson almost square, with rounded

corners, but slightly longer than broad. Sternites

not distinctly impressed, nor lobed at the bases

of the legs.

Male gonopod (Fig. 3) with a short slender

tracheal rod; coxa rather globose and large in

proportion to the rest of the gonopod; prefemur-

femur short, heavily setose, its front margin

produced into a flange overlapping most of the

HOFFMAN: AMERICAN MILLIPEDS OF FAMILY

On

Or

EURYURIDAE

course of the seminal channel. Tibiotarsus set.

off by a conspicuous suture, its basal half broad

and laminate with a conspicuous lobe on one side,

distad of which it becomes abruptly more slender

and sigmoidally bent, tapering gradually to its

tip. Solenomerite very long, almost length of

tibiotarsus, gently curved and gradually tapering

distad, without branches or other modifications.

Dorsum light brown with the keels and a large

trapeziform area in the middle of each meta-

tergite yellow, the latter giving the effect of a

continuous, broadly serrate median dorsal band.

Prozonites entirely yellow. Color of legs, sternites,

and antennae probably also yellow in life.

Fries. 1-4+—1, Polylepiscus heterosculptus pococki, n. subsp., left gonopod of paratype, Tumbala,

Chiapas; 2, P. trimaculatus, n. sp., left gonopod of holotype, Sepaciute, Guatemala; 3, Sem-

nellogon cerroazulensis, n. sp., left gonopod of holotype, Cerro Azul, Panama; 4, Varyomus confluens

(Chamberlin), left gonopod of male holotype, Rancho Grande, Venezuela. All figures to same scale,

showing gonopod in mesial aspect.

56 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

Genus Aphelidesmus Brolemann?

Aphelidesmus Brolemann, Ann. Soc. Ent. France

67: 322. 1898; Ann. Soc. Ent. France 84: 584.

1915—Carl, Mem. Soc. Neuchat. Sci. Nat. 5:

936. 1914—Attems, Rev. Zool. Bot. Afr. 17:

280. 1929; Das Tierreich 68: 128. 1937.

Type.—A. hermaphroditus Brolemann, by orig-

inal designation.

Diagnosis—A genus of the Euryuridae in

which the tibiotarsus of the male gonopod is

generally elongated and considerably expanded-

laminate, forming a shield-like or spathe-like

arrangement which partially or entirely encloses

the long slender solenomerite. Femur of gonopod

distally rotated almost 360°, causing the seminal

groove to completely encircle the base of the

tibiotarsus before entering upon the solenomerite.

Range.—From the State of Para, Brazil, north

through the Guianas and Venezuela to Colombia

and Ecuador, thence north along the Atlantic

coast of Central America to extreme southern

Texas. Most of the species occur in Colombia.

Species.—29, as follows: albocarinatus (Peters),

ambiguus Carl, areatus (Peters), asper Attems,

aterromus Attems, atratus (Pocock), bellus At-

tems, converus Jeekel, dealbatus (Gervais), dit-

vergens (Chamberlin), elongatus (Brolemann),

frangens Chamberlin, fumigatus (Peters), glaphy-

ros (Attems), goudotr (Gervais), gwianiensis

Chamberlin, hermaphroditus Brolemann, hybridus

(Peters), atermedius Chamberlin, major Cham-

berlin, octocentrus (Brolemann), panamanicus

Chamberlin, rivicola (Silvestri), roulini (Gervais),

semicinctus (Peters), surinamensis Jeekel, tertius

(Chamberlin), tripunctatus (Peters), wncinatus

(Peters).

Remarks—Many of the species referred to

Aphelidesmus were originally described under

the generic name Huryurus (including two named

by Brolemann six years after he had proposed

Aphelidesmus!); and most subsequent writers

such as Pocock, Carl, and Chamberlin have as-

sociated the genus with other euryurid groups.

Despite this body of precedence, however, the

genus was removed from the Euryuridae and

relocated in the family Strongylosomidae by

Attems in his 1929 summary of the group. It

was out of respect of his authority that I omitted

Aphelidesmus from my 1951 paper. Later con-

sideration has convinced me that Attems was

2 Trachelorhacis Silvestri 1898, which has three

months priority over Aphelidesmus as a name for

this genus, is a junior primary homonym of T’rache-

lorhacis Agassiz, 1846.

voL. 44, No. 2

in error, an opinion which is shared by my col-

league C. A. W. Jeekel, who has worked upon

both Aphelidesmus and many strongylosomoid

genera from Asia. Mr. Jeekel suspects that Attems

may have been influenced primarily by the elon-

gated and partly concealed solenomerite of

Aphelidesmus species. In other respects, includ-

ing the two cited as diagnostic of the Strongyloso-

midae by Attems (i.e., median constriction of

the gonopod aperture and independence of the

gonopod coxae) these species do not qualify as

stronglylosomid.

This is the largest genus in the Euryuridae,

there being some 29 species (many of which are

known only from female type specimens); the

range extends from Texas to northern Brasil,

with most of the forms concentrated in the

Colombia-Venezuela region. It seems not un-

likely that the group as here comprised may be

divided into two or more genera. At least one

species described in the genus is not congeneric

with A. hermaphroditus, and a new name is pro-

posed for its accomodation.

Varyomus, n. gen.?

Type.—Aphelidesmus confluens Chamberlin

1950, by present designation. —

Diagnosis——A euryurid genus characterized

as follows: similar in most respects to Aphelides-

mus but differing in that the distal half of the

gonopod telopodite ‘is not twisted on its axis, and

the seminal canal proceeds directly to the solenom-

erite without first encircling the gonopod. The

solenomerite itself is long and slender and distally

protected by several laminate expansions of the

tibiotarsus, but even these are different in ap-

pearance from the analogous structures in Aph-

elidesmus.

Range.—Northern Venezuela.

Species.—One.

Remarks.—Through the kindness of Dr. W. J.

Gertsch, I was able to re-examine the type speci-

men of Aphelidesmus confluens in the American

Museum collection, and provide herewith an

illustration of the left gonopod as seen in mesial

aspect. It will be apparent that the appendage is

considerably different from those of typical

Aphelidesmids, and this situation is additional

evidence of the futility of drawing gonopods from

whatever position happens to be convenient.

3 Named for Dr. Ralph Vary Chamberlin, the

describer of perhaps the majority of American

milliped species.

FEBRUARY 1954 HOFFMAN: AMERICAN

SUMMARY

It now appears that three groups can be

discerned amongst the American genera of

the Euryuridae (perhaps the Oriental

genera, of which I have not seen material,

belong to still another) as reflected by vari-

ous trenchant differences in the male

genitalia. Two of these have already been

recognized by Brolemann, as long ago as

1915, who set Aphelidesmus and_ Pro-

taphelidesmus apart in a separate subfamily

which he called Aphelidesminae. His ar-

rangement of the platyrhacoid diplopods

was as follows:

Family Platyrhacidae

Subfamily Platyrhacinae

Tribe Platyrhacini

Tribe Euryurini

Subfamily Aphelidesminae

It will be noted that the Aphelidesminae

was given a rank equivalent to what would

now be regarded as a superfamily including

the Platyrhacidae and Euryuridae (a posi-

tion which I believe in the light of present

knowledge to be too exalted). That Silvestri

was equally impressed by the characters of

his synonymical genus T'rachelorhacis is

evidenced by the new family which he

proposed for the reception of 7. rzvicola.

In admitting that the distinction generally

accorded Aphelidesmus is probably well-

founded, one has to recognize that as much

or more difference obtains between Huryurus

and Amplinus as between those two genera

on one hand and A phelidesmus on the other.

Giving these differences a coordinate degree

of recognition requires the establishment of

a third subfamily. These groups may be

distinguished and diagnosed as follows:

KEY TO THE SUBFAMILIES OF EURYURIDAE

1. Tibiotarsus of male gonopod substantially

expanded into a broad sheath, which shields

or actually encloses solenomerite branch and

which often has one or more small processes

of its own

Subfamily I. Aphelidesminae Brolemann

Tibiotarsus of male gonopod generally long

(reduced in one genus), very seldom broad-

ened, never forming a protective element for

solenomerite (when one is present)........ 2

4This name is proposed as new, being much

more limited in its scope than Euryurinae as used

by Pocock in the Biologia.

MILLIPEDS OF FAMILY

EURYURIDAE Da

2. No definite solenomerite present; telopodite of

gonopod simple, tibiotarsal joint not set off

by a conspicuous articulation

Subfamily II. Euryurinae, n. subf.‘

A long slender solenomerite present, arising at

base of tibiotarsus; telopodite of gonopod

with a definite joint or line of separation

between femur and tibiotarsus

Subfamily III. Amplininae, n. subf.

The genera belonging to these groups

may be in turn separated by the following

keys:

KEY TO THE GENERA OF THE APHELIDESMINAE

1. Tibiotarsus of gonopod twisted almost 360° on

axis of telopodite, seminal canal making a

complete circuit around gonopod before

entering on solenomerite, latter concealed or

partly enclosed by tibiotarsus, which forms a

30 5721(0)) Ree eee mg Aphelidesmus Brolemann

Tibiotarsus of gonopod not rotated on its axis;

seminal canal running directly to solenomer-

ite; latter not shielded or concealed by

LLC GAIESUIS he ne Mie uO EE Erm Go Ko gun iat 2

2. Solenomerite short, upright, arising from near

base of tibiotarsal blade; latter without

secondary processes

Protaphelidesmus Brolemann

Solenomerite long, arising from middle of

femoral portion of gonopod; tibiotarsal

portion composed of two laminate processes

which conceal tip of solenomerite

Varyomus Hoffman

KEY TO THE GENERA OF THE EURYURINAE

1. Gonopod relatively long and slender, tibio-

tarsal portion present and with a small

subterminal process........ Euryurus Koch

Gonopod short and robust, tibiotarsal portion

rudimentary, represented only by a short

digitiform lobe........ Auturus Chamberlin

KEY TO THE GENERA OF THE AMPLININAE

1. Preanal scale distally truncate or concave,

lateral tubercules large (23 species)

Pseudamplinus Hoffman

Preanal scale subtriangular or broadly rounded,

the lateral tubercules very small or absent. .2

2. No subantennal swellings present (9 species)

Pycnotropis Carl

Prominent subantennal swellings present... .3

3. Male gonopods with a secondary tibiotarsal

process in addition to the larger main blade;

caudolateral corners of the keels rather pro-

longed into spiniform processes (5 species)

Polylepiscus Pocock

Male gonopod without a secondary tibiotarsal

bipaiiGhicres: Ad onde estrone (PGs base oda whine ae) 4

4. Femoral portion of gonopod very stout and

short, exceeded in length by the correspond-

ingly elongated solenomerite............... 5

58 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

Femoral portion of gonopod long and slender,

exceeding the solenomerite and often tibio-

barsus Inslemgeila yi 58 Se eee te eet ee i

5. Tibiotarsus of gonopod generally slender, un-

modified, distally acuminate (2 species)

Seminellogon Chamberlin

Tibiotarsus of gonopod distally modified... . .6

6. Tibiotarsus slender, parallel-sided, bladelike,

bifid distally (8 species)

Thrinoxethus Chamberlin

Tibiotarsus broadly sigmoid, widest at mid-

length, distally trifid (1 species)

Sigmogonotropis Hoffman

7. Tibiotarsus generally distally acuminate,

occasionally expanded but never with spinous

processes or teeth (5 species)

Amplinus Attems

Tibiotarsus broadened, blade-like with a

conspicuous process at its midlength and

several small subterminal teeth (1 species)

Colomborus Chamberlin

Of the genera admitted to this key, there

is considerable uncertainty in my mind

concerning the status of the last five enu-

merated. Very little in the way of annectant

forms would be required to necessitate

consolidation of all of these nominal genera

back into Amplinus. For the time being,

however, they appear to be reasonably

discrete and easily recognizable groupings;

the gonopods being more distinctive than

one would appreciate from the inadequate

characterizations in the key.

In addition to the pores of the 18th and

19th segments of Semznellogon, another

character of unestablished taxonomic value

is the presence or absence of subantennal

swellings. I have never seen any specimens

VoL. 44, No. 27

of Pycnotropis; the information regarding

this difference is derived from Attems.

Considering the fallability of other char-

acters employed in that author’s key to

the euryurid genera, this one must be held

in suspicion until it has been more thor-

oughly investigated.

REFERENCES

ATTEMS, Cart GRAF. Diplopoden des Belgischen

Congo. I. Polydesmoidea. Rev. Zool. Bot. Afr.

17 (3): 2538-878. 1929.

. Fam. Strongylosomidae. Das Tierreich 68:

1-300. 1938.

. Fam. Leptodesmidae, Platyrhacidae, Oxydes-

midae, Gomphodesmidae. Das Tierreich, 69:

1-487. 1938.

BROoLEMANN, HENRI W. Essai d’une classification

des Polydesmiens. Ann. Soc. Ent. France 84:

523-608. 1915.

CHAMBERLIN, Raupu V. The millipeds of Central

America. Proc. U. 8. Nat. Mus. 60 (8): 1-75.

1922.

. Results of the Bryant Walker Expeditions of

the University of Michigan to Colombia, 1913,

and British Guiana, 1914. The Diplopoda.

Occ. Pap. Mus. Zool. Univ. Michigan. no.

133: 1-148. 1923.

. On a collection of millipedes and centipedes

from northeastern Peru. Bull. Amer. Mus. Nat.

Hist. 78 (7): 478-585. 1941.

. Neotropical chilopods and diplopods in the

collections of the Department of Tropical

Research, New York Zoological Society. Zoo-

logica 35 (2): 133-144. 1950.

HorrMaNn, RicHarp L. A new genus of Central

American milliped (family Euryuridae),

with notes on the American genera. Proc. U.S.

Nat. Mus. 102: -235-243. 1951.

MALACOLOGY .—Leiostracus (?) kugleri, . sp., a new bulimulid mollusk from

Venezuela. LoraHar Forcart, Museum of Natural History, Basle, Switzer-

land. (Communicated by Harald A. Rehder.)

Since 1922 Dr. H. G. Kugler and other

Swiss geologists have been sending most

interesting scientific collections from Vene-

zuela and Trinidad to the Museum of

Natural History in Basle (Switzerland).

During the war, 1939-1945, when normal

communications between South America

and Switzerland were interrupted, Dr. Kug-

ler sent malacological collections from

Venezuela to the U. 8. National Museum

in Washington. Dr. H. A. Rehder recently

entrusted this material to the author for

determination. Shells of a species of Buli-

mulidae were identified with those the

Museum in Basle received as early as 1926,

and of which the revision established that

they. belong to a species hitherto unde-

scribed. The species is dedicated to Dr. H. G.

Kugler, to. whom science owes much for the

scientific exploration of Venezuela and

Trinidad.

Leiostracus (?) kugleri, n. sp.

Diagnosis —The shell is solid, elongate-turricu-

late, narrowly umbilicated; its color is white

with ochraceus stripes, which are faded in worn

shells; the apical whorls are yellowish to whitish.

Fepruary 1954

The nepionic whorls are almost smooth; the

following whorls have more or less distinct ir-

regular riblets.

The aperture is elongate-ovate, its base some-

what angular. The columellar lip is expanded

with a straight vertical edge. The internal part

of the aperture and the expanded lip are brownish

colored.

The shells show nearest morphological rela-

tions to those of Leiostracus cinnamomeo-lineata

(Moricand) from the Brazilian Province Bahia,

of which paratypes (Mus. Basle 1489-a) have

been compared.

Fig. 1.—Letostracus (?) kugleri, n. sp.: Holo-

type, X2 and natural size.

PROCEEDINGS:

THE ACADEMY a9

Holotype —Mus. Basle 4950-a.

Type locality —Venezuela, Est. Faleén, Distr.

Colina, Porta Juela near Cumarebo—leg. Dr.

H. G. Kugler and Dr. L. Vonderschmitt 1926.

7? paratypes——S8 (Mus. Basle 4950-a’) from

the type locality; 36 (Mus. Basle 4950-c and

4950-d) and 27 (U. S. Nat. Mus. 508834 and

508855) from Est. Faleén, Distr. Zamora, Cuma-

rebo Field—leg. Dr. H. G. Kugler 1933-1949;

6 (Mus. Basle 4950-b) from Est. Faleén, Distr.

Acosta, near Rio Tocuyo—leg. Dr. H. G. Kugler,

1929. ;

Measurements of the shell (in mm).—<s follows:

: Aperture ]

Specimen peg Height Z ae

Width | Height Whorls

Holotype (Mus.

Basle 4950-a)...... aS 20 4.1 7.8 gl¢

Paratype (Mus.

Basle 4950-a’). 9.4 25.7 5.9 0.5 RiZ

Paratype (U.S. Nat.

Bil. creates 8.3 | 22.1 3.6 8.9 gl;

Because only shells of this species are knowns

its classification in the genus Levzostracus Albers,

1850, is based only on conchological feature.,

PROCEEDINGS OF THE ACADEMY

462D MEETING OF THE BOARD OF MANAGERS

The 462d meeting of the Board of Managers,

held in the Library of the Cosmos Club on March

16, 1953, was called to order by the President at

8 p.m., with the following in attendance: F. M.

SETZLER, F. M. Deranporr, J. R, SwaALLen,

H. 5. Raprteyve, J. A. Stevenson, A. G.

McNisu, W. H. Gitsert, F. W. Poos, H. A.

Bortuwick, C. A. Berrs, A. H. Scort, L. A.

SPINDLER, F. W. Hoven, Sara E. Branuam,

W. W. Dieu1, and, by invitation, E. H. Waker,

Kari Herzretp, W. W. Rusey, and J. C.

Ewenrs.

In the absence of Chairman Davis of the

Committee on Meetings, Mr. Setzler reported

that Dr. Gordon Macgregor, of the Technical

Cooperation Administration, Department of

State, would speak at the April meeting of the

Academy.

Dr. Ruspey, Chairman of the Policy and Plan-

ning Committee, reported informally that the

Committee unanimously approved the affiliation

of the Washington Section of the International

Association for Dental Research. The report was

accepted by the Board. Dr. Rusery also reported

that four members of the Committee thought the

appointment of a special committee to consider

ways and means of improving the JouRNAL was

not desirable, but suggested that the Board of

Editors review the reports of the past 10 years

in this connection and bring recommendations to

the Board of Managers; one favored the ap-

pointment of a special committee; and one was

noncommital. After a brief discussion to the

effect that the report leaves the subject as it was

before, the question was set aside pending a

formal report of the Committee.

In the absence of Chairman McPHErson, Mr.

SETZLER reported that the Committee on the

Encouragement of Science Talent had arranged

a dinner at the Cosmos Club on March 17 for 17

persons, including counselors from Washington,

Maryland, and Virginia, members of P.T.A.

councils, members of engineering groups, and

the President, to build up enthusiasm with

P.T.A. organizations in the metropolitan area in

60 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

junior high schools as well as in senior high

schools, not only among the students, but also

the parents, with a view toward contributions to

defray expenses of the Science Fair.

The Secretary reported the death of CHARLES

Moon on January 31, 1953.

The result of the ballot vote for the affiliation

of the Washington Chapter, American Society

for Metals was announced. A total of 406 ballots

were returned, of which 399 approved affiliation,

5 were against, and 2 were blank.

The Treasurer read a letter from EUBANKS

CaRSNER requesting to be placed on the retired

list. The request was approved.

The Treasurer reported that $745 had been

received in 1952 and 1953 for the Science Fair

Fund. With the $200 contributed by the Acad-

emy, a total of $945 was available. Deducting

bills already paid, partly for the 1952 Fair, there

is a balance of $736.20 for the 1953 Fair.

Further consideration was given to _ the

purchase of a motion-picture projector for the

Assembly Hall of the Cosmos Club. It was

brought out that six of the affhhated societies

held their meetings there: Botanical Society of

Washington, Philosophical Society of Washing-

ton, Geological Society of Washington, Wash-

ington Society of Engineers, and the District

of Columbia Section, American Society of Civil

Engineers. Since only these societies would be

concerned, it was suggested that a special com-

mittee be appointed consisting of the Vice

Presidents of these societies and the Treasurer

of the Academy.

463D MEETING OF THE BOARD OF MANAGERS

The 463d meeting of the Board of Managers,

held in the Library of the Cosmos Club on April

13, 1953, was called to order by the President at

8 p.m. with the following in attendance: F. M.

Serzper, F. M. Deranporr, J. R. SwWALLeEN,

H. S. Rapprieye, J. A. Stevenson, A. G. Mc-

Nisu, H. A. Borruwick, G. F. Gravart, C. A.

Berts, L. A. Sprnputer, M. A. Mason, Sara E.

BranuaM, R. G. Bates, W. W. Diext, and, by

invitation, Warson Davis, W. N. Fenton,

Karu Herzretp, W. W. Rusey, and A. T.

McPHERSON.

President SETzLER announced the appoint-

ment of a special committee to determine the

advisability of the Academy and its Afhliated

Societies purchasing a modern arc-equipped

motion-picture projector and presenting it to

the Cosmos Club for use in the Assembly Hall.

vou. 44, No. 2

The members of the committee are: CLIFFORD A.

Berts (Chairman); I. C. GarpNnmr, Philosophical

Society; A. Netson Sayre, Geological Society

of Wasuineton; Harry A. Bortuwick, Bo-

tanical Society of Washington; Martin A. ©

Mason, District of Columbia Section, American —

Society of Civil Engineers; and Howarp S.

RaAPPLEYE, representing the Academy.

Dr. Joun G. THompson, nominated for Vice

President representing the Washington Chapter,

American Society for Metals, recently affiliated

with the Academy, was elected.

Chairman Davis, of the Committee on Meet-

ings, announced that Lynn Pootz, Director of

Public Relations, Johns Hopkins University,

would present an illustrated lecture on Science

on television at the May meeting of the Academy.

Chairman Frnton, of the Committee on

Monographs, reported on the progress of the

index to the JoURNAL.

A. T. McPumrson, Chairman of the Commit-

tee on Encouragement of Science Talent, an-

nounced that the Seventh Annual Science Fair

would be held from April 30—May 38, at American

University. Winners in grades 9-12 will auto-

matically become members of the Junior Acad-

emy. He also reported that a total of $955 had

now been received, which will probably be suffi--

cient to cover the cost of the Fair.

The following interim report of the special

committee appointed to consider the purchase

of a motion-picture projector, was presented by

the Chairman, Currrorp A. Brrts:

The matter of the proposed purchase of a

modern arc-type 16-mm motion-picture projector

by the Academy and those of its affiliated societies

which use the Cosmos Club Assembly Hall has

been investigated by the committee. The follow-

ing can be reported at this time:

1. The Board of Directors of the Washington

Society of Engineers, meeting on April 1, au-

thorized the use of its share in the slide projector

which is in the Cosmos Club projection room

unused because the Club has a new one.

2. The Botanical Society, through H. A.

BorTHWICK, reports no funds available.

3. Pending formal confirmation by Dean

Mason, it is understood that the D. C. Section

ASCE has no surplus.

4. Howarp Rapp.eye, for the Academy, feels

that the potential benefits are not commensurate

with costs.

5. In deference to I. C. GARDNER, who is in

Europe, and A. NELSON SayRE whose group has

not met, final report will be deferred. Prospects

are not very propitious, however, for the original

proposal.

Frespruary 1954

Four deaths were reported by the Secretary:

-Doverias H. Camppety on February 23, 1953;

“Tuomas A. Jaaccar on January 17, 1953;

Cuarutes W. Bacon on March 19, 1953; and

GeorceE R. Wair on April 9, 1953.

A letter from Mayne R. Cor was read by the

Treasurer, requesting to be placed on the retired

list. This was approved, effective as of December

31, 1951.

The following statement, prepared by a

special committee of the Committee on Policy

and Planning, was presented by the Chairman,

W. W. Rosey: |

The Board of Managers of the Washington

Academy of Sciences at its meeting of April 13

instructed me to transmit the following report to

you.

The Board discussed the feeling of deep concern

and unrest that is now widespread among

scientists of Washington and elsewhere as a result

of the recent dismissal of the Director of the

National Bureau of Standards. The scientific

and technological agencies of the Federal govern-

ment have made important contributions to the

health, welfare, and safety of the nation—

contributions that have been possible only because

these agencies have traditionally maintained an

atmosphere of disinterested investigation free

from political and commercial pressures. Unless

this tradition is maintained, the scientific agencies

will be unable to attract and retain scientists of

the caliber required for the satisfactory per-

formance of their duties.

The recent action of the Secretary of Commerce

raises fundamental questions about the scientific

work and administrative policies of the National

Bureau of Standards. The members of the Board

of Managers note with approval that Secretary

Weeks has invited advice regarding the scientific

performance and broader purposes of the Bureau

from the Visiting Committee and from a special

committee appointed by the presidents of the

National Academy of Sciences and of national

societies in seven fields of science and technology.

In order to allay the growing concern about the

future of science in Government, it is respectfully

urged that the Director of the Bureau of Standards

be retained in his present position until these two

committees have reported, at which time it

should be possible to assess the merits of the case

properly.

The statement was unanimously approved by

the Board with the recommendation that it be

sent to appropriate persons. It was then moved

and earried unanimously that the statement be

sent to President Eisenhower in the form of a

telegram and released to the press. It was sug-

gested that copies of the statement be sent to

DetLev W. Bronk, President of the National

PROCEEDINGS:

THE ACADEMY 61

Academy of Sciences, Hucu L. Drypren, Chair-

man, Interdepartmental Committee on Scientific

Research and Development, members of the

committee appointed by the National Academy

to look into the objectives of the National Bureau

of Standards, the American Association for the

Advancement of Science, and the Vice Presidents

of the Washington Academy.

464TH MEETING OF THE BOARD OF MANAGERS

The 464th meeting of the Board of Managers,

held in the Library of the Cosmos Club on May

18, 19538, was called to order by the President

at 8 p.m., with the following in attendance: F. M.

SETZLER, F. M. Deranporr, J. R. SwALen,

H.S. Rappieys, J. A.Stevenson, H. A. REHDER,

J. P. M. Morrison, H. A. Borruwicx, G. F.

Gravatt, C. A. Brerts, GLEN Stocum, F. W.

Hoven, H. G. Dorsry, M. A. Mason, R. G.

Bates, and, by invitation, E. H. Waker,

Watson Davis, W. W. Rusery, and A. T.

McPHERSON.

Chairman Davis, of the Committee on Meet-

ings, announced that the October meeting of

the Academy would be held at the National

Institutes of Health and the November meeting

would be held jointly with the Anthropological

Society of Washington.

The Secretary read the following formal re-

port of the Committee on Policy and Planning

on (a) whether a special committee on improving

the Journal should be appointed, and (b) whether

the application from the Washington Section of

the International Association for Dental Re-

search for affiliation with the Washington Acad-

emy should be approved.

The Committee on Policy and Planning has

conducted a mail ballot on these two questions

and the Committee chairman has talked per-

sonally with four of the five other members.

On the second of the two questions all six

members of the Committee are unanimous in

recommending that the Board of Managers

approve (and pass on to the Academy membership

for ratification) the application from the Washing-

ton Section of the IADR for affiliation.

On the other of the two questions referred to

the Committee, it is not possible to report any-

thing like a unanimous recommendation. Two

Committee members definitely recommend that a

special committee Not be appointed. One Com-

mittee member definitely recommends that a

special committee SHOULD be appointed. A fourth

member of the Committee weighs the pros and

cons of appointing a special committee and ends

up with no recommendation either way. The fifth

62 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

member ignores the question of a special com-

mittee but, from his suggestion that ‘‘the Editors

print what they may find available,’ I judge

that he considers a special committee unnecessary.

This accounts for all members of the Committee

except the chairman. Trying to strike a balance

between these divergent recommendations, I am

inclined to recommend that a special committee

on improving the Journal Not be appointed at

this time but that the present editorial board be

asked to review the whole problem and report to

the Board of Managers not later than the first

meeting next fall.

The report was accepted by the Board.

Chairman McPuHrrson reported that the

Science Fair held at American University from

April 30-—May 3, 1953, was very successful.

There were 1,052 exhibits with 59 secondary

schools participating. Four students were se-

lected to represent the Washington Science Fair

at the National Science Fair held at Oak Ridge

May 7-9. Students have already begun work on

projects for the next science fair and applica-

tions for additional space over and above that

allotted this year have been received.

President SmTzLER expressed his gratitude to

Dr. McPuerson for his effort in connection

with the Science Fair.

The Secretary reported the death of HERBERT

E. Grecory on January 23, 1952.

Senior Editor Morrison reported a suggestion

by Dr. McPuHeErson, that the Washington Acad-

emy of Sciences consider a prize contest for

articles for publication in the Journal, similar to

that of the New York Academy of Sciences. It

was pointed out, however, that the prizes awarded

by the New York Academy were made from

special outside memorial funds set up for the

purpose, rather than from current funds of the

Academy. The proposal was tabled for further

study.

Dr. SmrzuprR, on behalf of the Board of

Managers, expressed his appreciation for the

assistance given by Watson Davis and Science

Service in preparing the telegram sent to Presi-

dent Eisenhower and the press release in regard

to the National Bureau of Standards.

465TH MEETING OF THE BOARD OF MANAGERS

The 465th meeting of the Board of Managers,

held in the Library of the Cosmos Club on Octo-

ber 12, 1953, was called to order by the President

at 8 p.m. with the following in attendance: F. M.

SETzLER, F. M. Deranporr, J. R. Swa.ueEn,

H. 8S. Raprueyve, J. A. Srevenson, J. P. E.

Morrison, A. G. McNisu, W. H. GiLpeErt,

VOL. 44, No. 2

F. W. Poos, H. A. Bortuwick, G. F. Gravatt,

C. A. Brrts, A. H. Scort, L. A. SpinpiEr, F. W.

Hove, M. A. Mason, W. W. Dieu, J. G.

TuHompson, and, by invitation, E. H. WALkKmR,

Watson Davis, A. V. Astin, and A. T. McPuer- |

SON.

In reporting on the progress of the Index to |

the Journal, President Snrzipr stated that it —

was expected that it would be published in |

December 1953. The price was set at $7.50, with ©

a 20 percent discount to members. This was done ~

prior to the Board meeting in order that a notice

could be carried in the October issue of the

Journal which went to press on October 7. The

action of the committee on the Index was ap-

proved by the Board.

Chairman Davis of the Committee on Meet-

ings reported that the November meeting would

be a joint meeting with the Anthropological So-

ciety of Washington and that the speaker would

be Dr. Cuirrorp Evans. Suggestions for the

December meeting were requested.

The question of presenting the Awards for

Scientific Achievement at the annual dinner

meeting was discussed, since it was felt that

holding a separate meeting for the purpose was

not enough of an occasion. It was suggested that

the awards might be presented at the annual

meeting without responses, or responses limited

to about five minutes, or the responses spread

over several meetings; and that the annual

meeting could be held separate from the dinner

meeting. It was the opinion of the Board that

there was no objection to combine the annual

meeting with the dinner meeting. Dr. Asrrn

said that the Committee on Awards would make

specific recommendations at the next meeting of

the Board.

Dr. Astin, Chairman of the Committee on

Awards for Scientific Achievement, reported

progress in selecting candidates for the 1953

awards. He brought to the attention of the Board

that Dr. Lyman J. Bricges would celebrate his

80th birthday next May and that special recog-

nition would be appropriate. It was suggested

that a special number of the JournaL might be

prepared for the occasion.

In the absence of Dr. Herzretp, Chairman

of the Committee on Grants in Aid for Research,

the Secretary announced that the allotment for

grants from the American Association for the

Advancement of Science for 1953 is $255. With

a balance of $99.50 from the 1952 grant, there is

a total of $354.50 available for the current year.

Fespruary 1954

In the absence of Dr. Rusry, Chairman of the

Committee on Policy and Planning, the Secre-

tary reported that the application of the Wash-

ington Section of the Institute of the Aeronautical

Sciences for affiliation with the Academy, was

unanimously approved by the Committee. The

report of the Committee was approved by the

Board.

Chairman McPHErson, of the Committee on

Encouragement of Science Talent, stated that

$1,200 would be needed for the next Science Fair.

Letters were written to the Vice-Presidents re-

questing the support of the Affiliated Societies.

Keita JOHNSON announced that the next Science

Fair would be held April 9-13, 1954, a time

which would enable participating students to

arrange their exhibits without missing time from

school. He also stated that the Prince Georges

County Science Fair was being continued, and

that the Arlington Science Fair would be held

prior to ours.

President SETZLER announced that bids for

printing the Rep Book had been received from

two printers. The Board authorized acceptance

of the lower one, from the Waverly Press. E. H.

WALKER suggested reprints of the first part of

the Rep Book containing information about the

Academy should be arranged for use of the Com-

mittee on Membership, and answering requests

for data on the Academy.

The Secretary reported the result of the ballot

vote on the application of the Washington Sec-

tion of the International Association for Dental

Research for affiliation with the Academy. A

total of 316 votes were cast, of which 298 were

for affiliation, 15 against, 2 were blank, and 1

undecided. Dr. Epwarp G. Hampp,~ nominated

by the Washington Section of the I.A.D.R. for

Vice-President of the Academy, was elected.

The death of Winit1am H. Hoover on Sep-

tember 11, 1953, and that of ERMINE CowLeEs

CasE on September 7, 1953, were announced.

C.S. Cragor, EvtiotQ. ApaAms, and ARTHUR F.

Beal requested that they be placed on the retired

list. The requests were approved, effective De-

cember 31, 1952, for C. S. Cragoe and Elliot Q.

Adams, and December 31, 1953, for Arthur F.

Beal.

Senior Editor Morrison reported that there

was $2,823.76 remaining in the budget for the

JOURNAL for the rest of the year. Since disburse-

ments were less than the budget, the December

number of the Journal will probably be enlarged

to 48 pages. Sixty-two papers have already been

PROCEEDINGS:

THE ACADEMY 63

published this year, and 40 manuscripts are on

hand. Papers are placed in separate categories,

and priority maintained in each category. Thus a

paper in the Physical Sciences might be published

in a minimum time, while one in Biology would

take approximately 6 months at the present time.

A communication from the Greater Washing-

ton Educational Television Association, Inc.,

requesting that the Academy appoint a repre-

sentative to the Advisory Council of the Associa-

tion, was received. The request was referred to the

Committee on Policy and Planning.

466TH MEETING OF THE BOARD OF MANAGERS

The 466th meeting of the Board of Managers

held in the Library of the Cosmos Club, No-

vember 17, 1953, was called to order by the

President at 8 p.m., with the following in at-

tendance: F. M. Srrzumr, J. R. SwWAuuen, H. 8.

RapplLEeYg, J. A. STEvENSoN, H. A. REHDER,

W. H. Giitpert, H. A. Bortuwick, G. F. Gra-

VATE, A. H- Soorr RS. Dini, Ff. W. Houes,

SaRA EH. Branuam, W. W. Dieu, E. G. Hampp,

and, by invitation, Hrernz Sprcut, WATSON

Davis, A. V. Astin, W: W. Rusey, and A. T.

McPHERSON.

President SmrTzLER reported that final proof

of the Index to the JouRNAL had been returned

to the printer and publication was expected by

the end of the year. He also reported that the

galley proof of the Red Book has been returned

to the printer, and publication was anticipated

in December.

Watson Davis, Chairman of the Committee

on Meetings, stated that Dr. Marvin J. KE.ty,

President of the Bell Telephone Laboratories,

and Donatp A. QuARLES, Assistant Secretary

of Defense for Research and Development,

would address the Academy at the December

meeting on Science in Government, and that Dr.

ALAN T. WATERMAN would be the speaker at the

Annual Meeting in January.

Dr. Astin, Chairman of the Committee on

Awards, moved that the Awards for Scientific

Achievement be presented at the Annual Meet-

ing, and that the printed program include pic-

tures and brief biographies of the recipients.

The motion was carried unanimously. It was

suggested that the awards be framed for presenta-

tion and the recipients requested to prepare a

paper of their own choosing for publication in

the Journal. It was also suggested that the

President appoint an Academy Committee to

consider plans for celebration of the 80th birth-

day of Dr. Lyman Briaes.

64

In the absence of Dr. Hprzreitp, Chairman of

the Committee on Grants-in-Aid for Research,

President SETzLER read the following report of

the committee:

A total of $354.50 is available for the current

year. Two applications (for Grants-in-Aid) have

been submitted, one by Dr. HERBERT C. Hanson,

Research Professor of the Department of Botany,

Catholic University of America, the other by Dr.

ALFRED WEISSLER.

Dr. Hanson has been studying the relationships

of grassland communities to environmental

conditions, particularly soils. About 200 samples

of soils have been collected but analysis of the

texture has to be made. This is a routine procedure

but requires considerable. time for so many

samples. Dr. Hanson asks for $100.00 to pay

student assistants for the analysis of the soil

samples, tabulating and analyzing the data on the

vegetation of individual stands, organizing these

stands into community types and determining

relationships. Dr. Hanson’s project is recom-

mended by Dr. Edward G. Reinhard, Head of the

Department of Biology.

The second application comes from Dr. Alfred

Weissler who is head of the Washington Office of

Ordnance Research. Dr. Weissler wishes to

continue on his own work in the application of

ultrasonic waves to chemical problems. He has

worked in this field at the National Research

Laboratory before 1951 and wishes to continue it

on Saturdays at the University of Maryland. Dr.

Francis E. Fox of the Catholic University has

placed some equipment at his disposal but he

needs a transducer which the Brush Company sells

for $350.00. Dr. Weissler’s project is recommended

by Dr. Richard K. Cook of the Sound Section at

the National Bureau of Standards.

The Committee recommends as follows:

1. That a grant of $100.00 be given to Dr.

Herbert C. Hanson in accordance with his request.

2. That a grant of $250.00 be given Dr. Alfred

Weissler on his request, and that this grant may

be used in full or in part for renting ultrasonic

equipment if it should not be possible to buy it

in full with the money provided or by raising

additional funds.

The report was accepted by the Board.

A. T. McPuHerson, Chairman of the Commit-

tee on Encouragement of Science Talent re-

ported that the Academy is cooperating with

the D. C. Council of Engineering and Archi-

tectural Societies in a comprehensive program for

developing interest in engineering and science

in the Junior and Senior High Schools of the

Greater Washington area. An engineer and a

scientist are assigned to each school to assist the

teachers of science, to get acquainted with gifted

students through science clubs, and to speak or

provide speakers for school assemblies and P.T.A.

meetings. A Speakers Bureau has been set up to

JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

provide speakers on special topics; it will co-

operate with a speakers bureau for chemists —

already established by the Chemical Society.

A joint meeting of the Washington Junior

Academy of Sciences and the local Science Clubs

is to be held on November 21, 1953, at the Na-

tional Museum under the direction of Miss—

MarGareET Patterson, Executive Secretary of

Science Clubs of America.

Societies and individuals interested in the

promotion of science talent are invited to send

contributions for the Eighth Washington Science

Fair to Howarp S. RappeLeye, Treasurer of the

Academy.

The Secretary reported the result of the ballot

vote on the application of the Washington Sec-

tion, Institute of the Aeronautical Sciences for

affiliation with the Academy. A total of 193 votes

were cast, of which 190 were for affiliation and 3

against. Dr. R. J. SreeGer, nominated by the

Washington Section of the IAS for Vice-President

of the Academy was elected.

The deaths of the following members of the

Academy were announced: JoHN R. MouteEr,

February 28, 1952; Grorcr R. Putnam, July 2,

1953; FreppERIcK E. Wricut, August 25, 1953;

and Epwarp B. Veppmr, January 1951.

Mr. RappLeye presented the resignation of

O.S. AamMopr and S? F. SnreszKo which were ac-

cepted, that of Dr. Aamopr effective December

31, 1952, and Dr. Snreszxo effective December

aie G53: 3

Payment of insurance on shipments of re-

prints by the Academy rather than by the authors

was approved by the Board.

Mr. RaAppLeye reported that six contributions

to the Science Fair, amounting to $150, had been

received to date from affiliated scoieties.

Dr. ReErxHpDER presented data and plans for

advertising the Index to the Journal. The es-

timated cost was about $600. The Board ap-

proved the expenditure of this amount for the

purpose.

The Secretary read a communication from

the Academy Conference of the A.A.A.S., re-

- questing a contribution from the Academy at the

rate of $1 for each 100 members, to defray office

expenses of the Conference. A contribution of $9

was approved by the Board.

Dr. ALLEN V. AstTIN presented to the Academy

the report of the ad hoe Committee for Evalua-

tion of the Present Functions and Operations of

the National Bureau of Standards, and a Report

of the Committee on Battery Additives of the

National Academy of Sciences. He also expressed

his appreciation for the action taken by the

Academy.

Jason R. Swauen, Secretary

VoL. 44, NO. 2m

Officers of the Washington Academy of Sciences

SME i Me ais Coes wv ees Francis M. Dreranporr, National Bureau of Standards

MUNEIRTLE-GICCE, 0 ee MARGARET Pitrman, National Institutes of Health

Nh Meee cs Up w Gletials waleia Jason R. Swauuen, U. 8. National Museum

mreasurer.......... Howarp S. Raprteye£, U. 8. Coast and Geodetic Survey (Retired)

SE A a a JouN A. STEVENSON, Plant Industry Station

Custodian and Subscription Manager of Publications

Haravp A. Renper, U.S. National Museum

Vice-Presidents Representing the Affiliated Societies:

mmenepoical society of Washington............0....cnccee neues S. E. ForsusH

macaropological Society of Washington..................... Wiuuram H. GILBERT

mological Society of Washington......................00e eee. WiuuiAM A. DayTon

mum society Of Washington...) ... 0.0.0... .0. eee eee ee ee ee JoHn K. TAYLOR

macmmoarogical Society of Washington.................c.cc cee ekeeeee FW. Poos

Memmi Creopraphic Society.............0..0 ccc cee ce wees ALEXANDER WETMORE

peeorenl Society of Washington...................c00se bees ARTHUR A. BAKER

Medical Society of the District of Columbia.................. FREDERICK O. CoE

DI AtSTOVIGH! SOCIELY. .... 6... ce ek ee cee ees GILBERT GROSVENOR

Seema society of Washington.................c0.e0ee neces. Lee M. HutcuHins

Washington Section, Society of American Foresters.......... GrorGcE F. GravatTr

Beerumeson pociety of Hngineers. .. 2... 2... ce ee ne ee C. A. Betts

Washington Section, American Institute of Electrical Engineers. ARNOLD H. Scott

Washington Section, American Society of Mechanical Engineers. .Ricuarp S. D1Lu

Helminthological Society of Washington........ .............. L. A. SPINDLER

Washington Branch, Society of American Bacteriologists......... GLENN SLocum

Washington Post, Society of American Military Engineers...... FLtoyp W. HouceH

Washington Section, Institute of Radio Engineers..... HERBERT GROVE DorRsSEY

District of Columbia Section, American Society of Civil Engineers...M. A. Mason

District of Columbia Section, Society for Experimental Biology and Medicine

Water C. Hess

- Washington Chapter, American Society for Metals........... JoHN G. THOMPSON

Washington Section, International Association for Dental Research

Epwarp G. Hampp

Washington Section, Institute of the Aeronautical Sciences...... F. N. FRENKIEL

Elected Members of the Board of Managers:

IRIE i bcc teks on de aeiee awk R. G. Bates, W. W. DrEHL

C02 ASCE oS A a M. A. Mason, R. J. SEEGER

oi DI Lr rrr Ae A. T. McPHERsoN, ’A. B. GuRNEY

MEMEO) ONNOMGGETS.... 0... cece ee All the above officers plus the Senior Editor

mmmrusewaziors and Associate Mditors............056 0c ee eae [See front cover]

Pe reeumee Commitice........-..... F. M. DeranporF (chairman), MARGARET PITTMAN,

J. R. Swauuen, H. 8. Rappieye, J. A. STEVENSON

Committee on Membership....HE1nz SpecutT (chairman), Myron S. ANDERSON, CLARENCE

Cottam, Rocer W. Curtis, JoHN Faser, J. J. Faney, FRaANcois N. FRENKIEL,

Wess HayMakeR, CLARENCE H. Horrmann, Louris R. Maxweiu, Epwarp G.

REINHARD, JOHN A. SANDERSON, Leo A. SHINN, Francts A. SMITH, ALFRED WEISSLER

Committee on Meetings Teg he ee Dorvanp J. Davis (chairman), ALLEN V. ASTIN,

GrorcE A. Hotrir, Martin A. Mason, Wituram W. Rusry

Committee on Monographs (W1Lu1aM N. FENTON, chairman):

1 pee ee a i Wiuu1aAM N. Fenton, ALAN STONE

iol 1) 20 eG) G. ArtHuR Coopsr, JAMES I. HorrmMan

11 LET eins gi [57 iA re Haratp A. REHDER, WiLL1AM A. DayTon

Committee on Awards for Scientific Achievement (RoBERT C. Duncan, general chairman):

For Biological Sciences........... Byron J. OLSON (chairman), Sara EK. BRANHAM,

Let M. Hurtcuins, FrepERicK W. Poos, BENJAMIN ScHwaRtTz, T. DALE STEWART

For Engineering Sciences...KuLiotr B. Roperts (chairman), CiirrorpD A. BETTs,

JosEPH M. CaLpWELL, MicHaEL GoLpBERG, EARLE H. KENNARD,

ARNOLD H. Scott, Horace M. TRENT

For Physical Sciences......... FRANK C. Kracexk (chairman), Witi1am H. Avery,

Ricuarp 8. Burineton, Natuan L. Drake, Luoyp G. HENBEsT,

Epear R. SmitH, BENJAMIN L. SNAVELY

For Teaching of Science...M. A. Mason (chairman), A.H. CLarx, Ker1tuH C. JOHNSON

Committee on Grants-in-aid for Research.............. HERBERT N. Eaton (chairman),

Mario Mouuari, Francis O. Rict, J. LEON SHERESHEFSKY, JAMES H. TayLor

Committee on Policy and Planning: (FRANCIS B. SILSBEE, chairman):

RUMMY MGs 2s else ariag vines ea ce Py airs Sse L. W. Parr, Francis B. SrusBEE

LE IT Ie 05 | CR eaten ee ee las OP CRITTENDEN, A. WETMORE

LS LET ee 2s) SY A nr ie ee ee eae JoHN E. Grar, Raymonp J. SEEGER

Committee on Encouragement of Satewioe Talent (A. T. McPueErson, chairman):

TO, CSTE ec NS 2 eee ae ea ee tee eee eet McPHERSsoN, W. T. Reap

LP aCe Tia as (0 56a a Austin H. Cuiark, J. H. McMriuen

Mey january POGR yk ss es es 2 oa kere ek oe L. Epwin Yocum, WILLIAM J. YOUDEN

feeprescataiue on: Counc) of A: ALAS... ooo ee adie nee oe Seed tl ee Watson Davis

Mommiitice Of AUGUOTS «. oc. 0c. ccc. os cece eee c cuas JosEPH P. EH. Morrison (chairman),

GALEN B. ScHUBAUER, EGBERT H. WALKER

Committee of Tellers...GzoRGE H. Coons (chairman), SAMUEL Levy, Waxpo R. WEDEL

CONTENTS

MatTHEMATICS.—Inequalities restricting the form of the stress-deforma-

tion relations for isotropic elastic solids and Reiner-Rivlin fluids.

M:; Baxer and J.‘L. BRIcKsmn 2 5.4.0 7.84 22

PALEONTOLOGY.—The development of the hinge of Veniella conradi

(Morton) and some conclusions based on its study. H. E. Voxss..

MycoLtoey.—Some Discomycetes new to Alaska. Epita K. Casu.....

% ZooLocy.—Description of Hocyzicus concavus (Mackin) with a review of

other North American species of the genus (Crustacea: Conchos-

traca,).: NT. MaAPPor) oes 24 teks ccc? es ee

ZooLtocy.—Further studies on American millipeds of the family Euryu-

ridae. (Polydesmida). Racnarp L. HorrMan.........- 2 eee

Mataco.tocy.—Levosiracus (?) kuglert, n. sp., a new bulimulid mollusk

from Venezuela. Loreaar PoRCcART.). 222.2272 ee

PROCEEDINGS: THE ACADEMY. 2242.4 csch cco) ence cee. oe

This Journal is Indexed in the International Index to Periodicals,

Page

&. fw

eDAWBS

Vou. 44 Marcu 1954

No. 3

JOURNAL

OF THE

WASHINGTON ACADEMY

OF SCIENCES

BOARD OF EDITORS

JoHN C. EweEers

U.S. NATIONAL MUSEUM

FENNER A. CHACE

U.8s. NATIONAL MUSEUM

ASSOCIATE EDITORS

J. 1. HorrMan

CHEMISTRY ~

Dean B. CowlEe

PHYSICS

ALAN STONE

ENTOMOLOGY

2

PUBLISHED MONTHLY

BY THE

R. K. Coox

NATIONAL BUREAU

OF STANDARDS

BERNICE SCHUBERT © °

BOTANY

Puitiep DRUCKER

ANTHROPOLOGY

Davin H. DUNKLE

GEOLOGY

WASHINGTON ACADEMY OF SCIENCES

Mount Royvaut & GUILFORD AVES.

BALTIMORE, MARYLAND

Entered as second class matter under the Act of August 24, 1912, at Baltimore, Md.

Acceptance for mailing at a special rate of postage provided for in the Act of February 28, 1925

Authorized February 17, 1949

Journal of the Washington Academy of Sciences

This JOURNAL, the official organ of the Washington Academy of Sciences, publishes:

(1) Short original papers, written or communicated by members of the Academy; (2)

proceedings and programs of meetings of the Academy and affiliated societies; (3)

notes of events connected with the scientific life of Washington. The JoURNAL is issued

monthly. Volumes correspond to calendar years.

Manuscripts may be sent to any member of the Board of Editors. It is urgently re-

quested that contributors consult the latest numbers of the JoURNAL and conform their

manuscripts to the usage found there as regards arrangement of title, subheads, syn-

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Manuscripts should be typewritten, double-spaced, on good paper. Footnotes should

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Illustrations in excess of the equivalent (in cost) of one full-page halftone are to

be paid for by the author.

Proof.—In order to facilitate prompt publication one proof will generally be sent

to authors in or near Washington. It is urged that manuscript be submitted in final

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Unusual cost of foreign, mathematical, and tabular material, as well as alterations

made in the proof by the author, may be charged to the author. :

Author’s Reprints.—Reprints will be furnished in accordance with the following

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Remittances should be made payable to ‘‘Washington Academy of Sciences” and

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Changes of Address —Members are reared to report changes of address promptly

to the Secretary.

JOURNAL

OF THE

WASHINGTON ACADEMY OF SCIENCES

Vou. 44

March 1954

Nows

BIOCHEMISTRY.—The reciprocal effects between calcium and phosphate ions

upon the growth, composition, and structure of castor bean, Ricinus communis

L.! Frank D. VENNING, Swingle Plant Research Laboratory, University of

Miami. (Communicated by J. R. Swallen.)

The present investigation was undertaken

to determine the effects of calcium and

phosphorus deficiency on the growth and

development of the main axis of castor bean,

Ricinus communis L., and to determine the

possible influence of the interaction between

calcium and phosphorus ions on its growth

and development. Comparisons between the

composition and structure of plants grown

with and without calcium, with and without

phosphorus, and with and without both

ealctum and phosphorus are given, and

possible cause and effect relationships are

pointed out. New data are presented con-

cerning growth phenomena as expressed by

the axis of castor bean, and correlations be-

tween such growth phenomena and the

chemical composition of the tissues involved

are offered.

MATERIALS, METHODS, AND PROCEDURES

Methods of Culture of the Experimental Plants

Certified seeds of Ricinus communis L. var.

cambodgensis J. B. S. Norton were germinated

in greenhouse flats of sand, watered with tap

water, and 12 days after germination were trans-

planted to four hydroponic culture series: (1)

Controls (2) minus calcium, (3) minus phospho-

Tus, and (4) minus both calcium and phosphorus.

For each culture series 10 2-gallon glazed

stoneware jars were employed, filled with No. 10

1 This study was undertaken as a part of a

course at the State University of Iowa, and the

author is indebted to Dr. Walter F. Loehwing

and Dr. Robert L. Hulbary of that institution for

their personal interest and suggestions. The study

was completed in its present form at the Swingle

Plant Research Laboratory of the University of

Miami.

65

mesh washed quartz gravel for root anchorage

and aerated by a regulated flow of washed com-

pressed air introduced into the bottom of each

jar. Three seedlings were planted in each jar,

providing 30 plants in each series. A modified

Knop’s solution was used as a nutrient medium

in all four series (Table 1), and the jars were kept

filled with the solution to a level just beneath

the surface of the gravel. In addition, micronu-

trients of boron, zinc, manganese, copper, and

iron were supplied to all series of plants. The orig-

inal pH of the solutions is noted in Table 1.

At the time of transplanting the seedlings to

the cultures the nutrient solutions were diluted

to supply a concentration of 0.01 per cent of

total solutes; beginning a week later the level

of total solutes in all series was gradually in-

creased by adding small amounts of nutrient

solution containing 0.1 percent of total solutes

from time to time as required by the plants,

until at the end of a 4-week period a concentra-

tion of 0.1 percent of total solutes was achieved

in the nutrient solutions of all four series of plants.

The series were thereafter grown at 0.1 percent

of total solutes for the remainder of the experi-

ment; there were no observable adverse effects

on the plants of a nature which would suggest

that the osmotic pressure of the solution was

unfavorable.to the growth of castor bean.

Weekly checks of the pH of the nutrient solu-

tions in the cultures were made, and it was

noted that the pH in the control series and to

some extent in the series lacking both calcium

and phosphorus tended to rise above 7.0, pre-

sumably because of unequal uptake of ions. The

cultures were artificially maintained in a pH

range between 6.5 and 7.0 by the addition of

small amounts of HCl when indicated, as some

66

foliar chlorosis develops in castor bean when the

pH rises above 7.2. It was determined that the

chlorosis was caused by insufficient iron uptake

in the alkaline medium, which was quickly over-

come when the pH was maintained slightly below

TAU

TABLE 1.—CoMPOSITION OF THE

NUTRIENT SOLUTIONS

1. ContRou (pH 5.35)

2¢ KNO3 2g KH2POs

2g MgSO. 8g Ca(NO3)2

Distilled water to make 14 liters.

2. Minus caAtcium (pH 5.4)

2g KNO; 2¢ KHe2POs,

2g MgSO. 4¢ Mg(NO3)2

4g NaNO; Distilled water to 14 liters.

3. MINUS PHOSPHORUS (pH 5.9) ;

2g KNO; 1g KH2SO4

lg KCl 2¢ MgSO.

8g Ca(NO3)2 Distilled water to 14 liters.

4. MINUS CALCIUM AND MINUS PHOSPHORUS (pH 6.1)

2g KNO; 4g Mg(NOs3)2

2g MgSO. 1g KCl

lg KH2SO4 4¢ NaNOs

Distilled water to 14 liters.

All four series of plants were grown simultane-

ously in the same greenhouse under comparable

conditions of hight, humidity, and temperature.

Since the castor bean is of tropical origin and

these investigations were begun in the north at

latitude 42° in the month of January, the photo-

period was extended to 12 hours per day by

means of 16 20-watt fluorescent lamps hung 6

feet above the greenhouse bench. This extension

of the photoperiod allowed the plants to develop

under conditions more nearly approximating

those which would be considered “normal” for

castor bean.

At the beginning of the experiment the day

temperatures were maintained at 75°F., and

night temperatures at 65°F., the day temperature

coinciding with the period of illumination.

Fifty-two days after their introduction into the

culture media small flower buds were noted on

several plants in all four series, and the tempera-

tures were raised to 75°F. night—85°F. day, and

maintained there during the balance of the ex-

periment, a total of 133 days.

Sampling: Anatomical Data

Growth within any particular series was rela-

tively uniform for all plants in that series. Be-

cause of this uniformity the plants were not

sampled at random but were sampled selectively:

The three plants having the longest, shortest,

and the most nearly medium-sized axis were

selected from each series, and the anatomical

JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

vou. 44, No. 3

and morphological data were based on their

development. Within the four series the dates

of sampling were staggered to give more nearly

comparable results, and were as _ indicated:

Controls: after 113, 118, and 122 days in the |

cultures; Minus calcium: after 116, 120, and 123 |

days; Minus phosphorus: after 116, 121, and -

124 days; and the series Minus calcium and |

minus phosphorus after 117, 122, and 125 days

in the nutrient cultures. At this time all elonga-

tion had ceased in the lower portions of the axis,

and secondary radial growth was relatively slow.

The seven basalmost internodes were selected

for study because they were all differentiated at

approximately the same time in all four series,

and the eighth node had been differentiated

before the inception of any floral primordia. The

tissue of these lower internodes was thus the

most comparable, physiologically, between the

series. |

The tissues composing the axis of castor bean —

are arranged as a series of fairly regular concen-

tric cylinders, with the exception of a small

region at. each node where the leaf trace leaves

the stele, and which was ignored for the pur-

poses of this study. Several transverse and longi-

tudinal free-hand sections were made near the

center of each internode, stained in a 1:10,000

solution of neutral red in distilled water, and

mounted in glycerin for microscopic study. Eight

transverse measurements were made of the width

of each of the major tissue regions within each

internode of each plant. As successive nodes are

not at an exact right angle to the sides of the

internode, the length of a particular internode

was determined as the average of its longest and

shortest length. From averages of the data so

obtained it was possible to determine the trans-

verse-section area and actual volume of all major

tissues within a particular internode.

The sections were also used for volumetric

studies of the pith cells. Tangential, radial, and

longitudinal measurements were made of 50

pith cells from each internode. Those pith cells

nearest the center of the internode are the largest

in all three dimensions. They gradually decrease

in size as one moves toward the periphery of the

pith; the smallest cells bordering on the medul-

lary sheath are from one-third to one-fourth as

large in any one dimension as those nearest the

center (the immediate center of the more mature

internodes is hollow). In order to secure cell

measurements which would afford a relatively

a

Marcu 1954

average picture of the development of the pith

within a particular internode, and because of the

general progression of size in the cells as described,

these cells were not sampled at random; all of

the pith cells which lay along two vectors which

intersected at right angles in the center of the

internode were measured along their greatest

radial and tangential axes. There were usually

between 12 and 13 pith cells along a vector from

the medullary sheath to the hollow center of the

internode. The 50 longitudinal measurements

were made in a similar manner, i.e., the same

number of cells was measured along vectors ex-

tending at right angles from the medullary

sheath to the center of the internode.

Sampling: Histochemical Data

Standard histochemical testing procedures as

described by Chamberlain (1932), Johansen

(1940), and Foster (1949) were employed to

determine the presence and extent of a number

of substances in the seven basal internodes. It

was usually possible to determine in which tissue

a particular substance was present, and also

possible, within the limits of sensitivity of the

tests, to determine relative quantities of a par-

ticular substance. Five plants from each series

were employed for this purpose, and the tests

performed on fresh freehand sections usually

made from the center of the internode, but

frequently from the upper and lower portions of

each internode as well. A part of these determina-

tions were made during the period of anatomical

sampling, and the remainder were performed

within the subsequent week; the plants had thus

been growing in the cultural solutions for a period

of 126 to 133 days. &

OBSERVATIONS

General Growth and Development of the Four

Series of Plants

During the period of vegetative growth the

axis of castor bean is monopodial. One leaf is

produced at each node; the axillary buds re-

main dormant and the axis is unbranched. With

the advent of flowering the habit becomes sym-

podial; in addition to the leaf a terminal inflores-

cence is produced at each node, and the axillary

bud develops the successive internode. The stem

is thus of indeterminate growth, and the plant

is a perennial under favorable environmental

conditions.

VENNING: MINERAL DEFICIENCY IN CASTOR BEAN 67

During the entire vegetative phase of growth

of the plants studied in this experiment, differen-

tiation of nodes and leaves took place at the

same rate in all four series; i.e., successive nodes

and leaves became visible on all plants after the

same time interval had elapsed. During this

phase the lower internodes of all plants were

approximately of the same diameters and lengths,

but sensitivity to differences in nutrient supply

was soon apparent in the leaves. The control

series, with a complete nutrient supply, had

fohage with the largest leaf area, deep green in

color. The leaves of plants deficient in calcium

had an area approximately two-thirds as great

as those of the control, somewhat wrinkled in

texture, and yellow-green in color. Plants lack-

ing phosphorus had a leaf area about three-fourths

as great as the control, and developed the deep

blue-green hue frequently associated with phos-

phorus deficiency. The plants deficient in both

calcium and phosphorus more closely resembled

the control than did either of the series deficient

in calcium or phosphorus alone. The leaves were

of comparable color and size to those of the con-

trols.

The uptake of nutrient solution during the

vegetative phase of development paralleled

general size and leaf area of the plants. It was

greatest in the controls, next largest in the series

minus both calcium and phosphorus, and least

in the series minus calcium.

Fifty-two days after their transfer to the

nutrient solutions, inflorescences were noted on

several plants in each of the four series, and

within a few days the first inflorescence had

appeared on almost all of the plants. The first

inflorescences were differentiated at either the

eighth or ninth nodes on all plants.

After entering the flowering phase, profound

changes in development took place between the

four series of plants. The control plants continued

to develop vigorously, and fruits were set and

matured from the lower flower spikes while new

foliage and inflorescences were continuously

differentiated at each successive node. The leaves

produced at the flowering nodes: were somewhat

larger than those from vegetative nodes; the

blade width averaging slightly over 30 em. Nutri-

ent uptake in this series gradually increased

throughout the course of the experiment. At the

time of sampling, the roots were noted to be

clean and: firm, much-branched, and in active

growth.

68 JOURNAL OF THE WASHINGTON ACADEMY OF

In the series minus calcium, flowering brought

about an abrupt cessation of elongation in the

axis. For a time nodes and internodes continued

to be differentiated at the same rate as those of

the control, but subsequent elongation of the

internodes was greatly inhibited, both in the

newly-formed ‘‘floral” internodes and in the

younger internodes which had been differen-

tiated prior to the appearance of the first inflor-

escence. Increase in stem diameter was also

retarded, but not to as great a degree. The inflor-

escences produced were rudimentary, and _ be-

came increasingly so at each successive node. A

few of the male flowers of the first inflorescences

produced pollen, but for the most part the male

flowers were sterile; and no fertile female flowers

were produced in any of the inflorescences.

Foliage differentiated at flowering nodes was

much reduced in size; most leaves had a blade

under five centimeters wide. This foliage was

pale yellow or almost white, brown-spotted,

curled or wrinkled, and ephemeral. The leaves

on the vegetative portions of the axis were gradu-

ally abscised, and as defoliation increased, the

rate of growth of the apical portions of the

axis fell behind that of the control and finally

ceased altogether. Uptake of nutrient solution

decreased sharply after the initiation of flowering,

and continued to decrease during this phase of

development. Many of the plants in this series

remained alive until the termination of the ex-

periment, but all had ceased any visible develop-

ment. At the time of sampling the roots were

noted to be soft, relatively few-branched, and

considerable sloughing of root tissue was evident.

In the series deficient in phosphorus, the first

inflorescence bore fertile male and female flowers,

and young fruits were set. As development of

the first fruits began, the lower leaves developed

dark brown spots, became flaccid, and were

promptly abscised, so that the ‘vegetative’

portions of the axes of these plants were almost

completely defoliated within a week. Elongation

of the younger “vegetative” internodes and

newly-formed ‘floral’? internodes was inhibited,

but not to so great a degree as in the plants

lacking calcium, and their lateral expansion was

only shghtly inhibited. The foliage produced at

the flowering nodes was smaller than that formed

previously, with a blade width between 10 and

15 em. The rate of differentiation of the subse-

quent internodes was slowed, and if young fruits

were forming on the first inflorescence, the female

SCIENCES VOL. 44, NO. 3

flowers were abscised from all subsequent in-

florescences. On the lower inflorescence, if several

fruits were developing, all but one were abscised;

and this remaining fruit required a longer period

to mature than did the earliest fruits of the

control. At maturity the seeds were found to be |

undersized, imperfectly formed, and contained —

almost no endosperm. The embryo was rudi- |

mentary, and the seeds were nonviable. Plants of |

this series which matured a fruit as described —

died when the capsule matured and dehisced. |

A few plants in this series did not enter the floral

phase of development; these continued vegetative

growth for the duration of the experiment. As

new leaves were differentiated the older foliage

underwent the discoloration and abscission as

described for the plants which flowered, but the

abrupt loss of foliage did not take place; the

new leaves were larger, attaining a width of ©

20 cm. The uptake of nutrient solution decreased !

with defoliation, and remained at a relatively —

low level thereafter. The roots of these plants

were not so extensive as those of the control,

but appeared to be healthy and in slow growth

at the time of sampling.

The series lacking both calcium and_ phos-

phorus reacted to flowering in a manner similar

to the plants grown without phosphorus, but

with several important differences. At first they

showed none of the outward effects seen in the

series deficient in calcium. The lower inflores-

cences set fruits, and discoloration and abscission

of the lower leaves followed, but defoliation took

place more slowly in this series than in that lack-

ing only phosphorus. Subsequent inflorescences

did not set fruits, but the elongation of the inter-

nodes was inhibited less in this series of plants.

Two or three fruits were ultimately matured on

the lower inflorescence; they were slow to ripen,

undersized, with small seeds, but endosperm was

present, the embryo appeared normal though

reduced in size, and several were viable. During

maturation of the fruits the rate of differentiation

of the upper nodes and internodes was retarded

below the control, and the new foliage produced

during the maturation phase resembled that of

the plants in the series minus calcium. The leaves

were about 10 cm broad, somewhat wrinkled

and chlorotic, but never as much so as those of

the plants lacking calcium alone, and they were

fairly persistent. Plants in this series did not die

at the maturation of the fruits, but subsequent

differentiation of nodes, as well as stem elonga-

Marcu 1954

© OND OR WwW

uN

(le @ 98a

Fig. 1.—Diagrammatic transverse-section area

of a segment of the axis of castor bean, indicating

the general tissue regions in the axis and their

arrangement: 1. Hpidermis, composed of a single

layer of small brick-shaped cells which bear on

their outer face a cuticular layer. 2. Outer

cortex, or hypodermis, is made up of relatively

small cells of isodiametric shape, for the most part

collenchymatous, containing abundant chloro-

phyll. 3. The inner cortex is parenchymatous,

the cells highly vacuolate, more or less isodia-

metric, chlorophyll-bearing, and of larger individ-

ual volumes than the cells of the hypodermis.

4. Sclericycle is several cells deep, an interrupted

layer of mechanical tissue with greatly thickened

cell walls, and whose protoplasts are dead at

maturity of the elements. Ontogenetically it is

probably a part of the primary phloem. 5 0:

The phloem contains two distinct regions: The

outer, older, primary phloem (5) contains a high

proportion of phloem parenchyma, quite large

amounts of chlorophyll, and is probably no longer

active as conductive tissue. It is for the most part

of primary derivation. The inner, younger, second-

ary phloem (6) is principally composed of phloem

parenchyma, sieve tubes, and companion cells;

is derived from the vascular cambium. 7. The

vascular cambium is continuous around the stem,

and appears as a thin cell layer one cell in thick-

ness. 8. Secondary xylem parenchyma is the

tissue lying to the inside of the vascular cambium,

and which has not as yet undergone differentiation

into one of the more complex xylem elements.

9. Xylem is represented by the large black area in

the diagram. The bulk of the secondary xylem is

composed of small. elongated cells with pitted

lignified walls, tapering ends, and living proto-

plasts. These mechanical and storage elements,

the xylem prosenchyma, are derived from the sub-

sequent differentiation of cells which arise from

the activity of the vascular cambium. 10.

VENNING: MINERAL DEFICIENCY IN CASTOR BEAN 69

tion, was greatly retarded. Uptake of nutrient

solution became more and more retarded as the

leaves were abscised, and, while the uptake was

greater than in either of the series lacking only

one macronutrient, was far less than the control

plants. Root development at the time of sampling

was comparable to the plants grown without

phosphorus, and there were no evidences of

deterioration of root tissue.

Structure and Development of the Basal Internodes

The tissue of the internodes of the main axis

of castor bean is at first solid, but unequal rates of

differentiation and expansion of the inner and

outer tissue regions result in a hollow area ex-

tending down the center of each internode as

it matures. All axial tissues are differentiated as

a series of concentric cylinders of fairly regular

outline as shown in Fig. 1. At the time of sam-

pling, both primary and secondary tissues were

present in the axes of all plants. As is seen in

Fig. 1, the axis of castor bean is of regular and

conservative structure, free from growth anom-

alies. Under the conditions of extreme deficien-

cies of calctum and phosphorus in this experi-

ment, no changes in anatomical ‘‘patterns’’ or

tissue arrangement occurred. The tissues as

described in Figure 1 were developed in all

series of plants; the response to variations in

nutrients was expressed by differences in quan-

tities of tissues and their proportions to one

another. Such differences in quantities reflected

differences between the activities of the meri-

stems, both apical and vascular, and also reflected

differences in the amount of secondary differ-

Pith rays are usually only one or two cells wide,

may have a thin deposition of lignin in their walls,

and usually extend from the medullary sheath

across the vascular cylinder to the cortex. They

are represented in the diagram by white lines in

the xylem. 11. The first formed conducting

elements and protoxylem parenchyma, both of

which are primary in origin, lie to the inside of the

secondary xylem cylinder as a series of small verti-

cal ridges which project into the pith, the so-called

protoxylem points. The largest number of vessels

(represented by white circles) are found in these

protoxylem points. 12. Medullary — sheath:

The outermost layers of the pith, one to three cells

deep and abutting on the inner face of the xylem

cylinder. They differ from the remainder of the

pith cells in their smaller transverse diameter,

greater length, and intercellular spaces much

reduced or lacking. 13. The pzth is the inner-

most tissue. The cells are parenchymatous, iso-

diametric in transverse view, and highly vacuo-

late. 14. Hollow in the center of the axis of the

more mature internodes.

70 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

entiation into specialized tissues from cells

originated by these meristems.

As the quantity of primary vascular tissue is

small, the bulk of the vascular tissue is secondary

and the area of xylem and phloem affords an

index of cambial activity within the axis. These

data are presented for all series of plants in

Tables 2 to 5. For purposes of measurement,

the medullary sheath was included with the

pith; xylem was measured from the medullary

sheath to the cambium; phloem from the cam-

bium to the outside of the sclericycle (as these

cells are probably primary phloem derivatives);

and the cortex included all tissue between the

sclericycle and epidermis.

As shown in Tables 2 to 5, the larger areas of

secondary tissues are found in the first basal

internode, and the proportions of vascular to

nonvascular tissue decreased in each successive

internode. The diameter of the pith is succes-

sively increased; whereas the transverse-section

area of cortical tissues is approximately constant.

In the control plants there is a marked ten-

dency for each successive internode to elongate

to a greater degree than the previous internode;

this trend continues until “‘floral’”’ internodes are

differentiated; from the first node bearing an

inflorescence upward the internodes are of more

or less equal length.

In the series lacking calcium, cambial activity

was greatly reduced, particularly in the upper

internodes. In the fifth and sixth internodes,

most of the secondary xylem indicated in Table 3

was xylem parenchyma, cells of cambial origin

which had not undergone any differentiation

into functional xylem elements. There was little

secondary xylem in the seventh internode. On

the other hand, the primary xylem was well

differentiated into conducting elements in these

upper internodes. A small amount of phloem in

the upper internodes appeared to be derived

from the vascular cambium. The length of the

internodes in this series decreased progressively

upward.

The first three internodes of the plants grown

without phosphorus showed similar develop-

mental patterns to those of the control, with a

progressive increase of length of the internode.

The length of the remaining vegetative inter-

nodes tended to become constant, with decreased

length occurring in the ‘floral’ internodes.

The transverse-section area of vascular tissue

progressively decreased, but the elements were

vou. 44, No. 3

for the most part well differentiated into conduct-

ing and mechanical tissue.

The tissues of plants grown without both

calcium and phosphorus more nearly resembled

the controls than did either of the series grown

without calcium or phosphorus alone. Progressive

increases in internodal length were similar for

the first five internodes; thereafter the internodes |

became progressively shorter. Secondary vas-

cular tissues were well differentiated in all seven

internodes.

The average volumes of the various tissues in

the first seven internodes were computed from

TaBLE 2.—TissuE AREAS AND LENGTHS IN THE

SEVEN BasaL INTERNODES OF THE CONTROL

SeRrIzEs!

AREA OF: (Areas in mm2)

LENGTH OF

; INTERNODE

Pith | Xylem | Phloem| Cortex

‘ mm.

INTERNODE |

Plant les. 51226 43.10 13.12 8.09 17.5

RigntiZiesseece 15.49 53.18 13.97 8.86 16.5

Rianitroeeeeeee 10. 24 58.05 15.31 9.69 22.5

Average..... 12.66 | 51.44 14.13 8.88 18.83

INTERNODE 2

2 lemita lee 13.91 41.10 13.39 7.60 14.5

lanier 18.16 49.68 14.25 7.55 24.0

Planter 16.51 51.81 13.91 8.38 17.5

Average..... 16.19 47.52 13.85 7.84 18.67

INTERNODE 3

lantern 17.17 36.42 10.82 8.98 17.0

Plante eee 20.20 43.64 13.04 7.43 19.5

Pisnitioecee er 15.87 47.88 13.23 7.72 25.0

Average..... 17.75 42.65 12.36 8.04 20.5

INTERNODE 4

iantaliseeee 30.74 26.78 lw 8.43 24.0

Rlamti2 eee 36.21 36.71 13.18 8.86 25.3

Rlantromeeee 24.25 42.62 12.81 fama! 29.0

Average..... 30.40 | 35.37 12.40 8.17 26.1

INTERNODE 5

lant seek 37.61 21.47 10.57 8.51 35.0

lant 2s ares 41.69 31.03 lets 9.59 39.0

lantroee 3). 1183 B25} 11.95 8.11 38.0

Average..... 37.14 28.92 1152233) S8aie y/o 88)

INTERNODE 6

lant = 39.70 21.80 10.75 9.29 32.0

Plante2ineeeere 38.32 22.85 10.02 7.85 40.0

Riantigens.e 38.44 31.69 11.07 8.78 47.0

Averagé..... 38.82 25.45 10.61 8.64 39.66

INTERNODE 7

Plaritaliy ae a. 45.81 17.16 10.50 8.98 40.0

Plant 2he mace 45.86 20.33 9.73 8.35 38.5

Blante3ijon.o- 38.90 25.26 10.32 7.66 56.0

Average..... 43.52 20.92 10.18 8.33 44.83

1 Internode 1 is basalmost.

Marcu 1954

the data of Tables 2 to 5, and are presented

numerically in Table 6, and diagramatically in

Figs. 2 to 5.

The length of any axis segment, or internode,

is primarily determined by the number of tiers

of cells within the unit, and the degree of elonga-

tion which they have undergone; whereas the

transverse-section areas of primary tissues, such

as the pith and cortex, are determined by the

width of the apical meristem, and the subsequent

degree of intercalary growth and expansion of

the cells. In all four experimental series, the size

of the individual secondary vascular elements

TABLE 3.—TissuE AREAS AND LENGTHS IN THE

SEVEN BASAL INTERNGDES OF THE SERIES

Minus Catcrum!

AREA OF: (Areas in mm?)

LENGTH OF

INTERNODE

Pith | Xylem | Phloem} Cortex

mm.

INTERNODE 1

Planbpl ye: ... 8.69 6.72 4.03 | 4.81 1085

Riagnteee. a... 19.15 | 10.82 oh MSHA 20.0

etamibrore. | ete! 13.85 | 10.40 62427 ie 7-23 30.0

Average..... 13.89 9.31 5.90 6.77 20.17

INTERNODE 2

IBlamtetecs = cc. 10.48 6.44 4.92 | 4.72 9.0

Plate 22 28 issu; 18.08 7.09 7.19 | 6.80 iS

Plant os. 12.06 8.63 5.99 | 5.99 25.0

Average..... 13.54 7.38 6.06 | 5.84 16.5

INTERNODE 3

Blame e. os 11.01 5.29 4.42 | 4.79 116165)

Planta) 2S: 19.95 4.66 rez, Gs02 16.5

Plants 2... 18.17 8.26 6.51 6.65 20.0

Average..... 16.38 6.07 5.75 5.99 16.67

INTERNODE 4

Rlantwle fe... 16.32 3.76 4.74 | 5.61 19.0

12 ha 74 ae 21.40 1.01 5.30 | 6.89 15.0

IPlantioie.s- .. 21.20 3.68 6.18 | 6.09 15.0

Average..... 19.64 2.81 5.41 6.20 16.33

INTERNODE 5

Riantate,. i.e. 16.64 1.86 4.46 | 5.23 10.0

Rlamtigi +. << 18.19 0.00 3.90 | 5.54 11.0

liens) eee 24.32 Seu 6.52 6.69 10.0

Average..... 19.72 1.66 4.96 | 5.82 10.33

INTERNODE 6

Blamteteer = «3.2 14.10 0.62 3.15 | 4.70 65)

awit 2a ee 11.60 0.00 Wah I) 9 BRAG 6.0

iPiantraeece ..- 29.28 0.00 6.31 7.36 10.0

Average..... 18.33 0.20 4.01 5.44 7.83

INTERNODE 7

tanta... : 11.97 0.00 BATE |) 33GB - 6.0

Plamt2 0... 9.25 0.00 2.17 | 4.69 3.0

Plamen. e 24.10 0.00 5.24 | 6.46 10.0

Average..... - 15.11 | 0.00 | 3.88 5.02 6.33

1 Internode 1 is basalmost.

VENNING: MINERAL DEFICIENCY IN CASTOR BEAN 71

was approximately the same, but the size of

primary elements, such as the cortex and pith

cells, was dissimilar. The rate and degree of

activity of the apical meristem is directly reflected

in the number of nodes produced, and the actual

number of cells in the primary tissues of the

internodes. As the pith had the largest overall

volume of any tissue in the axis, and is the first

tissue to terminate meristematic activity and

undergo vacuolation during the ontogeny of the

internode, its cellular makeup was selected as

an index of the activity of the apical meristem

at the time the lower internodes were differen-

TABLE 4.—TissuE AREAS AND LENGTHS IN THE

SEVEN Basaut INTERNODES OF THE SERIES

Minus PHospHorus!

AREA OF: (Areas in mm?)

LENGTH OF

INTERNODE

Pith | Xylem | Phloem| Cortex

mm.

INTERNODE 1

PRamth i i975 14.80 6.12 4.79 | 4.70 16.5

Planti2- 2s oe 17.54 19.10 8.73 by 15.0

Plantis2 2 3-0 20.00 | 10.70 Mein les 100 15.0

Average..... 17.45 11.97 7.09 5.61 15RD

INTERNODE 2

Plant le .5 2. 15.29 6.92 4.76 | 5.24 15.0

Plant: 2ees 5 oe 19.68 16.70 6.87 (ale 18.0

Rlantiopaes 22.68 10.44 (22 eO0o 16.0

Average..... 19.22 ists 6.28 6.48 16.33

INTERNODE 3

Plamnigee see 18.66 5.55 4.20 | 4.93 16.0

Planta2eas fee 23.78 {3253 7.43 6.99 24.0

Plantiseee oe 26.17 7.44 a297 |e Ole 20.0

Average..... 22.87 8.84 5.87 | 6.22 20.0

INTERNODE 4

Planitia: 22.09 4.70 4.45 | 5.40 17.0

Planti2inesacce 24.50 8.21 5.85 | 5.81 Dae

Plante ee 27.96 5.83 6272) ||) 5278 23.0

Average..... 24.85 6.25 5.67 5.66 20.83

INTERNODE 5

Plantes. ae 20.55 2.55 3.41 | 5.85 17.0

Plamtezie see 27.20 8.74 4.99 | 6.17 18.5

Blanton. a 30.80 3.97 5.09 | 7.61 Pi 5

Average..... 26.18 5.09 4.50 6.54 21.0

INTERNODE 6

Rismitel 22 see 21.47 2.18 3.00 | 5.28 17.0

Plantese on 28.53 6.13 4.49 | 5.59 21.0

Plantes ees cee 27.87 2.63 Bia nee 9 28.0

Average ..... 25.96 | 3.65 | 3.75 | 5.76 22.0

‘INTERNODE 7

Plantily eee 19.15 1.42 2.08 5.00 15.0

Pyeanite 2ee te ee 29.40 4.12 ERB || aici} 20.0

Plants -2.be 24.37 1.90 BaD eae OOO 25.0

Average..... 24.31 2.48 3a7PA | Gait) 20.0

1 Internode 1| is basalmost.

72 JOURNAL OF THE WASHINGTON ACADEMY OF SCIENCES

tiated. The average volume of the pith cells in

all four series is presented numerically in Table 7,

and diagramatically in Figs. 6 to 9.

The average size of individual pith cells of

the control plants tend to become increasingly

larger in the first four internodes, then the size

decreases slightly and tends to become more or

less constant. In the plants lacking calcium the

average cell volume is greatest in the first inter-

node, slightly exceeding the control. These cells

show a general trend to become more reduced

in volume in each successive internode, and the

average cell volume is much less than that of

TABLE 5.—TiIssuE AREAS AND LENGTHS IN THE

SEVEN BasaAL INTERNODES OF THE SERIES MINUS

CALCIUM AND Minus PHospHoRus!

AREA OF: (Areas in mm?)

LENGTH OF