bola a, Pye ng geet gy i ha Nt
<= ae ee ated
rei ee ee eee
ee anaes
centre te EE et Oe ener ws
0 peta ena cpearge ioe ara ae)
ee
Ge ae oes
eee
barnett Sr ees
be
A oP a
oe
ie es
a Ee
A
Mees
oS ot
Cita Tie
—_ iy oy (eireecs | ar aah tes, Te 4: 96, pled i eins
ie o hee UE ee hs +s OR a art at pe ete j &*
is A ven
ae Ay
i 7
' }
ct - he epee | - |
: a 1 ae ee | _ »
ee hae ir | ae
; a or if 1 ;
a elsNind a a
Na sc it Jah nae ry 7
oe ue
FG ails i ae
SMITHSONIAN
MISCELLANEOUS COLLECTIONS
VOR. 159
©cecccese®®
“EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES,
AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN”’—JAMES SMITHSON
(PusiicaTion 4468)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1961
PORT CITY PRESS, INC.
BALTIMORE, MD., U. S. A.
ZA HSOWgs
AUG 28 1961
LIBRARY
ADVERTISEMENT
The Smithsonian Miscellaneous Collections series contains, since the
suspension in 1916 of the Smithsonian Contributions to Knowledge,
all the publications issued directly by the Institution except the An-
nual Report and occasional publications of a special nature. As the
name of the series implies, its scope is not limited, and the volumes
thus far issued relate to nearly every branch of science. Papers in
the fields of biology, geology, anthropology, and astrophysics have
predominated.
LEONARD CARMICHAEL,
Secretary, Smithsonian Institution.
(iii)
ay ai ie ; al ve
Fite ee ee A ;
re
/ i ; K
TO.
BY.
CONTENTS
PEABoby, FRANK E. The oldest known reptile, Eosauravus copei
Williston. 13 pp., I pl., 3 figs. May 7, 1959. (Publ. 4377.)
WETMORE, ALEXANDER. The birds of Isla Escudo de Veraguas,
Panama. 27 pp., I pl., 3 figs. July 8, 1959. (Publ. 4378.)
GREGOIRE, CHARLES. Further observations on distribution of pat-
terns of coagulation of the hemolymph in Neotropical insects.
23 pp. Aug. 18, 1959. (Publ. 4379.)
Hanp ey, Cuartes O., Jr. A review of the genus Hoplomys
(thick-spined rats), with description of a new form from Isla
Escudo de Veraguas, Panama. I0 pp., 1 fig. July 3, 1959.
(Publ. 4380.)
Cooper, G. ArTHUR. Genera of Tertiary and Recent rhynchonel-
loid brachiopods. 9o pp., 22 pls., 1 fig. Nov. 23, 1959. (Publ.
4382.)
BoarDMAN, RicHarp S. A revision of the Silurian bryozoan
genus Trematopora. 14 pp., 2 pls., 1 fig. Oct. 29, 1959. (Publ.
4383.)
Gazin, C. Lewis. Early Tertiary Apheliscus and Phenacodaptes
as pantolestid insectivores. 7 pp., 2 pls. Aug. 12, 1959. (Publ.
4385.)
Snoperass, R. E. The anatomical life of the mosquito. 87 pp.,
30 figs. Nov. 4, 1959. (Publ. 4388.)
AssotT, C. G. A long-range forecast of United States precipita-
tion. 78 pp., 12 figs., g charts. Mar. 23, 1960. (Publ. 4390.)
WILLIAMS, JEROME; JOHNSON, E. R. FENIMorE; and Dyer, AL-
BERT C. Water transparency observations along the east coast
of North America. 181 pp., 2 pls., 13 figs. Oct. 26, 1960.
(Publ. 4391.)
WETMoRE, ALEXANDER. A classification for the birds of the
world. 37 pp. June 23, 1960. (Publ. 4417.)
(v)
SMITHSONIAN
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 1
Charles D. and Mary Waux Walcott
Research Fund
THE OLDEST KNOWN REPTILE,
EFOSAURAVUS COPEI WILLISTON
(WitH 1 Prate)
By
FRANK E. PEABODY
Department of Zoology
University of California
Los Angeles, Calif.
(PuBLicaTIon 4377)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
MAY 7, 1959
yy EXON UDA
hp ry
Pot
oy
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 1
Charles D. and Mary Waux Walcott
Research Fund
THE OLDEST KNOWN REPTILE,
EOSAURAVUS COPE! WILLISTON
(W1TH 1 PLATE)
By
FRANK E. PEABODY
Department of Zoology
University of California
Los Angeles, Calif.
(PusiicaTion 4377)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
May 7, 1959
SMITHSONIAN
INSTITUTION MAY 7
THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.
FOREWORD
Dr. Frank E. Peabody died on June 27, 1958, leaving several manu-
scripts in various stages of completion. The one published here, based
on a specimen in the collections of the United States National Mu-
seum, was complete except for final touching up of some of the il-
lustrations ; these illustrations have been finished by Miss Madeline M.
Peabody with the help of Dr. Theodore H. Eaton, Jr. The manuscript
has been edited by Dr. Eaton and myself. Dr. Peabody’s paper presents
a welcome clarification of the relationships of an important, and here-
tofore much misunderstood, early reptile.
PETER P. VAUGHN
i a Aire
aim
Charles D. and Mary Vaux Walcott Research Fund
THE OLDEST KNOWN. REPTILE:
BOSAURAVUS COPED WILLISTON
By FRANK E. PEABODY
Department of Zoology
University of California
Los Angeles, Calif.
(WitH ONE PLATE)
One of the most tantalizing examples of Carboniferous tetrapods
is the posterior part of a small skeleton from Linton, Ohio, described
by Cope in 1897 as the earliest known reptile. Some 60 years later, and
after many taxonomic vicissitudes, the specimen seems in danger of
slipping into obscurity among the microsaur Amphibia. Meanwhile
no more reptiles have been found at Linton or in earlier horizons.*
Various students have described Cope’s specimen, but most have
tended to discount its importance because the anterior part of the
skeleton, including the skull, is missing, and have tended to accept the
early descriptions with little question. Present high interest in the
origin of reptiles during the Carboniferous prompted a restudy of
Cope’s historic specimen. It was found that strong lighting from a
very low angle, and directed from various positions, revealed much
new detail that can be demonstrated by photographic enlargements.
The result is a new interpretation, particularly of the vertebrae and
tarsus, which reaffirms the reptilian affinities of the specimen and
furthermore strongly suggests a captorhinomorph relationship.
I am indebted to Dr. Peter P. Vaughn of the United States National
Museum for permission to borrow Cope’s specimen, and to Miss
Madeline M. Peabody, my sister, for assistance with the illustrations.
1[Cephalerpeton ventriarmatum, apparently a captorhinomorph reptile (see
Gregory, 1950), is known from the nodule beds at Mazon Creek, IIl., which
represent a somewhat earlier horizon (but still within the Allegheny series).
—Ep.]
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 1
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
SYSTEMATIC DESCRIPTION
EOSAURAVUS COPEI Williston
PLATE I; TEXT FIGURES I-3
Isodectes punctulatus Corr, Proc. Amer. Philos. Soc., vol. 36, pp. 88-90, pl. 3,
fig. 3, 1897; WuiLutston, Journ. Geol., vol. 16, pp. 395-400, text fig. 1, pl. 1,
1908; Moonie, Proc. U. S. Nat. Mus., vol. 37, pp. 11-16, pls. 4-5, 1909.
Eosauravus cofei WILuisTon, Bull. Geol. Soc. Amer., vol. 21, p. 272, 1910; CASE,
Carnegie Inst. Washington Publ. 145, pp. 31-32, text fig. 8, IQII.
Tuditanus punctulatus Romer, Bull. Amer. Mus. Nat. Hist., vol. 59, pp. 134-135,
1930; Bull. Mus. Comp. Zool., Harvard Coll., vol. 99, p. 300, 1947; Amer.
Journ. Sci., vol. 248, p. 641, 1950; HuENE, Palaontologie und Phylogenie der
niederen Tetrapoden, p. 163, 1956.
?Tuditanus Romer, Osteology of the reptiles, p. 483, 1956.
Type—uvU.S.N.M. No. 4457; posterior $ of a reptilian skeleton pre-
served belly-down on a slab of coal from Linton, Ohio.
Horizon.—Allegheny group, Middle Pennsylvanian (Westphalian).
Diagnosis.—Small reptile with a minimum of 28 presacral vertebrae
of generally captorhinid structure, with broad, swollen neural arches,
low neural spines, zygapophyseal facets in horizontal plane, and small
intercentra; free ribs on all vertebrae except distal caudals; distal
caudal vertebrae with low neural arches and probably without haemal
spines, centra occasionally fused forming relatively stiffened axis;
one principal and one accessory sacral rib; hind limb with prominent
internal trochanter, with relatively short epipodial (=zeugopodial)
segment having relatively massive fibula; primitive, well-ossified
tarsus of basic captorhinid or pelycosaurian plan with separate median
and lateral centrale and with a 6th distal tarsal (=postminimus) ;
phalangeal formula 2-3-4-5-4, terminal phalanges blunt-ended. No
gastralia present; possibly with body scales, having striae radiating
from anterior margin of scale. No obvious aquatic adaptations of
well-ossified skeleton. Anterior skeleton unknown.
Taxonomic notes.—The taxonomic history of Cope’s specimen is so
devious and confusing that a short explanation is necessary to sup-
plement the synonymy listed above. Cope (1897) described the
posterior skeleton and believed it to be conspecific with another small
vertebrate represented by a skull and anterior two-thirds of a skeleton.
The latter had been described by Cope (1874, p. 271) as Tuditanus
punctulatus, but in his 1897 paper, it was referred along with the
posterior skeleton to the genus Jsodectes. Williston (1908) and
Moodie (1909) offered new descriptions of the posterior skeleton,
treating it as distinct from the anterior skeleton, but tending to over-
look the fact that the anterior skeleton is the type of Jsodectes
NO. I THE OLDEST KNOWN REPTILE—PEABODY 3
punctulatus. (Moodie’s plate description (p. 28) in fact refers to the
posterior skeleton as “the type specimen of Isodectes punctulatus,”
which, of course, it is not.) Later, Williston (1910) and then Case
(1911) established the posterior skeleton as a new genus and species,
Eosauravus cope: Williston. Unfortunately, the European genus
Sauravus to which Williston related the posterior skeleton is clearly
an amphibian with nectridian vertebrae, so the name Eosauravus is
inappropriate morphologically but remains valid taxonomically.
Romer (1930) restudied the Linton fauna and, in a commendable
attempt to reduce the large number of artificial species, referred Cope’s
posterior skeleton again to the anterior skeleton now designated as
Tuditanus punctulatus. The synonymy of Tuditanus with Isodectes
had proved to be wrong since the latter genus now appears to be a
captorhinomorph (Gregory et al., 1956, p. 2), and the former genus is
a microsaur. Romer’s decision apparently rested mainly on the im-
probability that there might be more than one reptile at Linton, and
that there was the distinct possibility that the smaller, less ossified
anterior skeleton merely represents a more immature individual than
the posterior skeleton. The two specimens were regarded by Romer
as reptilian with no recognizable ordinal characters. Later, Romer
(1947, p. 300) suggested that the two specimens together represent
either a seymouriamorph or cotylosaur on the basis of a stemmed in-
terclavicle, seemingly broad-arched vertebrae, and a pes with a pha-
langeal formula 2-3-4-5-4. Still later, Romer (1950, p. 641) dis-
counted the importance of the stemmed interclavicle and phalangeal
formula, and, while noting a presumed high presacral count of verte-
brae, long, slender body proportions, apparent lack of caudal chevrons,
and long postorbital region of the skull, concluded that Tuditanus
punctulatus (based on anterior and posterior skeletons) “is not im-
probably a microsaur.” This conclusion, undoubtedly influenced by
increased understanding of microsaurs, was followed by both Piveteau
(1955) and Huene (1956) in their valuable compendia of vertebrate
paleontology. Meanwhile, Romer (1956, p. 483) apparently turned
once more toward Williston’s opinion of the posterior skeleton as
shown by the lone entry “[ Reptilia] Incertae sedis. ?Seymouriamor-
pha. ?Tuditanus Cope 1874 (Eosauravus Williston 1910).” Thus at
present, the posterior skeleton designated as Eosauravus copei by
Williston, is in an obscure position both taxonomically and phyloge-
netically. The anterior skeleton is best considered a probable micro-
saur amphibian under the designation Tuditanus punctulatus. In any |
case it is difficult to demonstrate distinctive reptilian characteristics in
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
T. punctulatus, and especially difficult to demonstrate any real affini-
ties with Eosauravus copet.
Description —The specimen consists of the posterior two-thirds of
a postcranial skeleton preserved belly-down on a coal stratum. Neither
the opposing slab, probably containing a dozen thoracic vertebrae and
caudal neural arches, nor adjoining blocks of matrix containing the
tip of the tail, some terminal phalanges of the left pes, and the anterior
end of the skeleton, were collected. The remaining parts of the skele-
ton have undergone very little deterioration since Cope’s time, judging
from the excellent photograph presented by Williston (1908) and
republished (with inaccurate retouchings) by Moodie (1909).
The presacral, sacral, and anterior caudal vertebrae lie on their right
sides (as observed by Cope, 1897) in such a manner as to cover the
proximal tips of the right ribs while the proximal ends of the left pre-
sacral ribs are pressed against the upper (left) surfaces of their cor-
responding centra. The outline of successive neural spines is clearly
visible on the right side between successive ribs. The caudal vertebrae
posterior to the rib-bearing caudals are preserved with ventral side
down and have lost their neural arches, thus exposing the neural canal
as a longitudinal groove in the dorsal surface of the centra. Un-
fortunately, Moodie (1909, pl. 5) illustrated the entire column as
though it were oriented with the dorsal side uppermost (figure repro-
duced by Case, 1911, fig. 8). The result is an erroneous picture of the
vertebrae from anterior caudals forward. Cope’s illustration (1897,
pl. 3) shows the correct orientation, but is only slightly suggestive of
the true form of the vertebrae.
The true form of the presacral and anterior caudal vertebrae may
be reconstructed with reasonable accuracy from a composite of details
exhibited along the column. Specifically the impression of the anterior
presacrals clearly shows the contour of the centrum; the first 5 pre-
sacrals and anterior caudals preserve details of swollen neural arches
as well as of the centra and intercentra. The position of intervertebral
foramina is clearly indicated by a series of circular pits. Figure I is
presented as a reconstruction based on composite detail.
There seems to be little doubt that the neural arch is low and broad
as mentioned by Romer (1947, p. 300), has a low spine, and has a
perceptible swelling above the posterior zygapophysis ; also that small
intercentra are present. The latter are indicated between the first
several presacral centra, between the 1st and 2d caudal centra, and
by a haemal wedge between the 3d and 4th caudal centra. In the pre-
sacral series the left ribs appear to have been crushed precisely against
THE OLDEST KNOWN REPTILE—PEABODY
I
NO.
"(o1) esyUadIOJUT ‘(4IS) Sqit [eroes ‘(su) outds yesnou ‘(9S)
g]qissog ‘“wowmdads ad4} Jo Jrejop 9}1sodui09 WoIF pozONns}sUOIOI ‘1agoI SnapinDsoy JO UoTseI
sayeos Apoq
Jepnes Josajue pue ‘eroes ‘sequiny—'I ‘o1,f
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
the intervertebral area; thus the supposed intercentra here may be
parts of the ribs. However, there appears to be a distinct intercentral
space coincidental with the position of the ribs, and, in any case, the
evidence for intercentra in the anterior caudals is clear and unobscured
by ribs.
Evidence of 24 presacral ribs on the left side is fairly clear, although
the first presacral is difficult to see, and only the distal tip of the 24th
presacral is preserved on the edge of the slab. The general pattern
and number of the presacral ribs and the far anterior position of the
left manus have led to the opinion (Romer, 1950, p. 641) that the
presacral count is significantly higher than the 25-27 vertebrae usually
found in the most primitive reptiles. However, the 5 most anterior
ribs are clearly more massive than following ribs, and the distal ends
are slightly spatulate—all indicative of an extreme anterior thoracic
position related to serrati muscles of the pectoral girdle. Also, the
successive positions of the distal ends of the 5 anterior ribs suggest
a progressive shortening in a forward direction as might be expected
in a smooth transition to the cervical region. Accordingly, a recon-
struction will show that the total number of presacrals may have been
as few as 28. The forward position of the left manus as an indicator
of a far anterior position of the pectoral girdle is probably misleading.
The girdle probably shifted forward or to the right side away from its
life position lateral to the 5 anterior ribs.
Rib heads are obscured in the presacral series generally, but the 3d
to 8th left presacral ribs appear to have a proximal expansion com-
mensurate with the elongate diapophysis of the neural arch. Certainly
these ribs are not single-headed as in lizards, but bear a general re-
semblance to captorhinid ribs.
The pelvic girdle and sacral vertebrae are distorted beyond certain
recognition of salient features, although the spacing of vertebral seg-
ments and disposition of lumbar and caudal ribs suggest the presence
of two sacral vertebrae. A short, thick element lying across the ad-
ductor fossa of the right femur may be a right second sacral rib; an
obscure spatulate structure immediately anterior to the anteriormost
left caudal rib may be the first or principal sacral rib. Except for a
general outline of the acetabular regions of the girdle, little can be
demonstrated here except that the mass lying between the heads
of the femora probably constitutes a pelvic girdle and sacrum of
primitive reptilian plan. According to my interpretation, Moodie
(1909, pl. 5) included the internal trochanter of the left femur in his
outline of the left acetabular region, thus giving the left pelvis a more
NOL THE OLDEST KNOWN REPTILE—PEABODY 7.
distinct outline than is warranted. A thin plate lying anterior to the
head of the right femur may represent the left ilium broken over to
the right. Although the thin plate may be regarded as a patch of over-
lying matrix such as obscures the centrale of the left tarsus (see be-
low), there is a definite anterior border that looks much like the an-
terior edge of an iliac blade. Nowhere is there evidence of a long
posterior process of the ilium like that of Eogyrinus.
The anterior 4 or 5 caudal vertebrae are associated with 3 pairs of
sharply curved ribs. In addition, there are short structures faintly
shown on the left side that are not curved and probably represent short
haemal spines nearly in the correct position. Also, there is a distinct
haemal wedge between the 3d and 4th caudal centra. Certainly there
is enough evidence to question seriously earlier observations (Cope,
1897, p. 89; Romer, 1950, p. 641) that there are no haemal spines in
the tail.
The caudal series becomes twisted, possibly 180 degrees, at the posi-
tion of the 7th vertebra, which appears to be lying on its left side.
Posteriorly the series is oriented with ventral side down—an unusual
position if neural and haemal arches were at all well developed here,
or if there was any lateral compression of the centra. Under these
conditions the vertebrae would be almost certainly lying on one side
or the other as in the anterior column. However, the caudal centra
appear broader than high, and occasional fusion of neighboring centra
seems to have occurred. All features of the tail, including the orienta-
tion, suggest some specialized function—perhaps a prehensile action
in the dorsoventral plane. A special aquatic function does not seem
possible, insofar as a lateral sculling motion is concerned, although
the fused vertebrae may suggest a stiffened axis serving as the founda-
tion for a rudder.
Part of the left manus (omitted in Cope’s figure, 1897, pl. 3) lies
disarticulated near the anterior end of the vertebral column. Enough
is shown to indicate that the carpus was definitely as fully ossified as
the tarsus, and less surely that the phalangeal formula was comparable
to the reptilian count in the pes.
Both limbs are complete except for the loss of some terminal
phalanges on the left side. The left femur is preserved with the dorsal
surface uppermost—the right femur with the ventral surface upper-
most. Thus the whole contour of the bone can be recognized in com-
posite. The femoral head, internal trochanter, adductor fossa, and
distally the tibial and fibular condyles resemble those of primitive
reptiles such as ophiacodonts and captorhinids. The trochanter is
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
especially prominent and extends proximally nearly in line with the
femoral head. Ossification is fully developed in the femur as well as
in the more distal elements.
The tibia and fibula are short, stout bones of generally primitive
contour ; the fibula appears relatively more massive than is usually
the case in the tetrapod limb. The distal end of the left tibia appears
to have slipped slightly upward from the life position and now rests on
the neck of the astragalus. Otherwise, the left femur, tibia, and fibula
are in normal articulation.
The right pes is twisted so as to obscure details of the tarsus, but
details of the digits help to complete a restoration of the left pes. The
left pes is preserved dorsal side uppermost and exhibits one of the
most perfect preservations of tarsal structure known from the Car-
boniferous, indeed, from the Paleozoic, as will be demonstrated pres-
ently. The pes has been given several superficial descriptions (Cope,
1897; Williston, 1908; Moodie, 1909) which fail to recognize the ex-
tent of ossification in the tarsus, but nevertheless establish two proxi-
mal elements in the tarsus and a phalangeal formula of 2-3-4-5-4. A
main difficulty lies in the interpretation of tarsal elements distal to the
presumed astragalus and calcaneum, especially in the medial region
of the tarsus where no one has been able to recognize central elements.
Moodie’s figure (1909, pl. 5), republished by Case (1911), is particu-
larly misleading in that the tarsus appears to have an enigmatic pat-
tern, doubtfully reptilian. (Also, in Moodie’s figure a nonexistent
element is added distal to the lateralmost distal tarsal, although none
is shown in Moodie’s retouched photograph—his pl. 4). My photo-
graphs (pl. 1A, B), taken under low-angle light from first one direc-
tion and then from the opposite direction, demonstrate the wealth of
detail making possible text figure 2. The two proximal bones of the
tarsus are clearly the astragalus and calcaneum which enclose between
them a perforating foramen, not previously noted. The astragalus
has a small but definite tibial facet directed mostly preaxially. There
is no evidence of tripartite structure such as exhibited by Captorhinus
(Peabody, 1951). The preaxial border between the astragalus and the
first metatarsal clearly exhibits two bones that must be a median cen-
trale and distal tarsal 1. A thin veneer of matrix obscures part of
the dorsal surface of these bones, but the oblique lighting (pl. 1A)
clearly brings out their contours in the preaxial border. Lateral to
these bones and median to the large distal tarsal are at least 2 and
probably 3 separate bones that are identifiable as the lateral centrale
and distal tarsals 2 and 3. A slight proximal jamming (see fig. 2) has
NO. I THE OLDEST KNOWN REPTILE—PEABODY 9
forced distal tarsals 2 and 3 slightly out of position. The existence of
two separate centralia seems certain although the separation between
the lateral centrale and distal tarsal 2 is not clear—probably because of
a slightly overriding relationship due to jamming. A unique feature
of the tarsus is a postminimus or distal tarsal 6 in the postaxial border.
Such an element is unknown in reptiles but is found in the tarsal pat-
tern of the urodele, Salamandrella, by Holmgren (1933, p. 217).
Fic. 2—Left pes of Eosauravus copei showing primitive reptilian pattern
with separate median and lateral centrale, and with unique postminimus or distal
tarsal 6 on postaxial border.
There is no doubt that the tarsal pattern is generally comparable to
primitive captorhinids and pelycosaurs.
The metatarsals are all well developed as indicated in figure 2. No
special features seem to be present except for a generally robust os-
sification (like that of more proximal bones) that contrasts markedly
with a seemingly delicate ossification of the phalanges.
The phalanges may be confidently restored with a 2-3-4-5-4 formula,
using the evidence from both feet. The terminal phalanges are not
acutely pointed and cannot be considered as definitely bearing claws.
The relative length of the 5th digit suggests no obvious aquatic adap-
tation—in the obviously aquatic Mesosaurus, the 5th digit is longer
than the 4th. This condition may also be noted in nothosaurs.
10
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Fic. 3—Left limb of Eosawravus copei reconstructed in
fully extended position.
NO. I THE OLDEST KNOWN REPTILE—PEABODY II
The surface surrounding the skeleton seems devoid of structures re-
sembling gastralia as indicated by early descriptions. However, a
problematical object that may be an unidentified bone from the an-
terior skeleton lies just to the left of the distal tail. Possibly of more
importance, an enlarged view of the surfaces near the skeleton reveals
a number of delicate, ovoid areas with fine striae radiating from a
point near one border. The striated areas occur only close to the
skeleton ; an example of the striated areas can be seen clearly between
the right ribs in plate 1C. It is possible that these striated areas repre-
sent body scales developed from the epidermis of Eosauravus. No
bone is indicated in the delicate impressions
Conclusions.—It is concluded that Cope’s historic specimen from
Linton, Ohio, is surely a reptile that has evolved beyond the sey-
mouriamorph level. The broad-arched, cotylosaurian vertebrae pos-
sess small intercentra, and the narrow space between successive pleuro-
centra is in decided contrast with the wide, unossified gap seen in
seymouriamorphs. Here, the pedicel of the neural arch has a marked
overhang above the intercentral gap. The tarsus has a characteristic
reptilian astragalus and calcaneum, with enclosed perforating foramen
in the usual position. The astragalus is fully developed with no
indication of a compound origin as in the relict Captorhinus aguti of
Early Permian age (Peabody, 1951). The whole structure of the
pes is of basic reptilian pattern except for the 6th distal tarsal or post-
minimus. The latter may be considered an amphibian feature rather
than a supernumerary element that widens the pes surface in correla-
tion with aquatic adaptations—an untenable point of view considering
the general lack of characteristics suggesting aquatic habits of
Eosauravus. The combination of vertebral and tarsal characteristics
is consonant with other features of the skeleton; together they
strengthen the evidence that the astragalar bone, originating from a
fusion of tibiale, intermedium, and proximal centrale of the amphibian
foot, may be regarded as a reliable osteological indication of the at-
tainment of the amniote level of organization—at least until conflict-
ing evidence is found.
If it be granted that Eosauravus is a reptile, there is a question as to
its subgroup affiliation. Current evidence strongly suggests that
early ophiacodont pelycosaurs and captorhinomorphs are very close
to the root of the reptilian stock. The tarsus of Eosauravus is exceed-
ingly primitive in the possession of separate median and lateral cen-
trale, and of the postminimus. Only early pelycosaurs have separate
centralia—they are fused in Captorhinus and Limnoscelis. No reptiles
I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
presently known have a postminimus. The nature of the vertebrae of
Eosauravus would indicate that its affinities probably lie with the cap-
torhinomorphs. No pelycosaur is presently known to possess vertebrae
of a pure cotylosaur type such as is evident in Eosauravus. In view
of the primitive pattern of the tarsus, a position near the base of the
captorhinomorphs is indicated.
Establishment of a true reptile of captorhinomorph affinities deep
in the Middle Pennsylvanian helps to clear away some of the uncer-
tainty surrounding the time of origin of reptiles. The varied reptiles
found in the Upper Pennsylvanian of Kansas (Peabody, 1954) and
more fragmentary remains from elsewhere indicate that the evolution
of pelycosaurs and captorhinomorphs (if petrolacosaurs be considered
an offshoot of the captorhinomorphs as suggested by Vaughn, 1955,
p. 446) was well advanced. Eosauravus appears to have been at an
evolutionary stage which could be ancestral to any known later reptile.
The particular adaptations of Eosauravus to life in a coal swamp
are difficult to assess. Moodie (1909, p. 12) suggests that the reptile
was aquatic or semi-aquatic mainly on the basis of an “expanded
foot” similar to the broad foot of the obviously aquatic mesosaurs.
However, the foot of Eosauravus and the rest of the preserved skele-
ton have little to suggest even semi-aquatic habits, but do allow the
possibility that this small reptile spent most of its time in the “upper
story” of the coal forest at Linton.
REFERENCES
Case, E. C.
1911. A revision of the Cotylosauria of North America. Carnegie Inst.
Washington Publ. 145, pp. 1-122, 52 figs., 14 pls.
Corr, E. D.
1874. Supplement to the extinct Batrachia and Reptilia of North America.
I. Catalogue of the air-breathing Vertebrata from the Coal-measures
of Linton, Ohio. Trans. Amer. Philos. Soc., vol. 15, pp. 261-278.
1896. The Paleozoic reptilian order Cotylosauria. Amer. Nat., 1896, pp. 301-
304, 1 pl.
1897. On new Paleozoic Vertebrata from Illinois, Ohio and Pennsylvania.
Proc. Amer. Philos. Soc., vol. 36, pp. 71-91, 3 pls.
Grecory, J. T.
1950. Tetrapods of the Pennsylvanian nodules from Mazon Creek, Illinois.
Amer. Journ. Sci., vol. 248, pp. 833-873, 11 figs.
Grecory, J. T., PEAzopy, F. E., and Price, L. I.
1956. Revision of the Gymnarthridae, American Permian microsaurs. Pea-
body Mus. Nat. Hist., Bull. 10, pp. 1-77, 33 figs., 1 pl.
Hotmcren, NILs.
1933. On the origin of the tetrapod limb. Acta Zool., Stockholm, vol. 14,
pp. 185-295, 106 figs.
NO. I THE OLDEST KNOWN REPTILE—PEABODY nS
HUuENnE, F. von.
1956. Palaontologie und Phylogenie der niederen Tetrapoden. xii + 716 pp.,
690 figs. Jena.
Moopir, Roy L.
1909. Carboniferous air-breathing vertebrates of the United States National
Museum. Proc. U. S. Nat. Mus., vol. 37, pp. 11-28, 7 pls.
1916. The Coal Measures Amphibia of North America. Carnegie Inst.
Washington Publ. 238, x + 222 pp., 43 figs., 26 pls.
Peapopy, F. E.
1951. The origin of the astragalus of reptiles. Evolution, vol. 5, pp. 339-344,
2RigSeT) pie
1952. Petrolacosawrus kansensis Lane, a Pennsylvanian reptile from Kansas.
Univ. Kansas Paleont. Contr.: Vertebrata, art. I, pp. I-41, II figs.,
3 pls.
1954. Pennsylvanian reptiles of Kansas. Bull. Geol. Soc. Amer., vol. 65, p.
1293 (abstract).
PIVETEAU, JEAN (Eb.).
1955. Traité de paléontologie. Vol. V, Amphibiens, reptiles, oiseaux. 1113
pp. Paris.
Romer, A. S.
1930. The Pennsylvanian tetrapods of Linton, Ohio. Bull. Amer. Mus.
Nat. Hist., vol. 59, pp. 77-147, 26 figs.
1947. Review of the Labyrinthodontia. Bull. Mus. Comp. Zool., Harvard
Coll., vol. 99, pp. 1-368, 48 figs.
1950. The nature and relationships of the Paleozoic microsaurs. Amer.
Journ. Sci., vol. 248, pp. 628-654, 4 figs.
1956. Osteology of the reptiles. xxi+ 772 pp. 248 figs. Univ. Chicago
Press.
VAUGHN, PETER P.
1955. The Permian reptile Aracoscelis restudied. Bull. Mus. Comp. Zool.,
Harvard Coll., vol. 113, pp. 305-467, 15 figs., 2 pls.
WILttston, S. W.
1908. The oldest known reptile—Isodectes punctulatus Cope. Journ. Geol.,
vol. 16, pp. 395-400, I fig., 1 pl.
1910. Cacops, Desmospondylus; new genera of Permian vertebrates. Bull.
Geol. Soc. Amer., vol. 21, pp. 249-284, 12 pls.
EXPLANATION OF PLATE 1
Left pes and lumbar region of Eosauravus copei seen under low-angle
illumination.
A. Pes, lighted from distal direction, showing clearly: Two elements—median
centrale (c. 1) and 1st distal tarsal (dt. 1)—lying between astragalus (a)
and Ist metatarsal; and 6th distal tarsal (dt. 6) lying between calcaneum
(cal) and 5th metatarsal (mt. 5).
B. Pes, lighted from proximal direction, showing 3 distinct elements (indicated
by black dots) lying median to large 4th distal tarsal.
C. Presacral vertebrae of lumbar region, lighted from anterior direction, show-
ing low neural spine (ns), presence of intercentrum (ic), and striated patches
(sc) possibly representing body scales.
14
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOUS 139; NOS 1, PEs 2
(For explanation, see p. 14.)
Hr
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 2
THE BIRDS OF ISLA ESCUDO
DE VERAGUAS, PANAMA
(With One PLatTE)
By
ALEXANDER WETMORE
Research Associate
Smithsonian Institution
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JULY 8, 1959
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 2
He BIRDS OF ISLA ESCUDO
DE VERAGUAS, PANAMA
(WitrH ONE PLATE)
By
ALEXANDER WETMORE
Research Associate
Smithsonian Institution
(Pus.icaTion 4378)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JULY 8, 1959
THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.
iris BIRDS OP ISLA, ESCUDO, DE VERAGUAS,
PANAMA
By ALEXANDER WETMORE
Research Associate, Smithsonian Institution
(WitH ONE PLATE)
Isla Escudo de Veraguas lies in the southern Caribbean Sea at
lat. 9°00’ N., long. 81°34’ W., distant a little more than 18 kilometers
from Coco Plum Point on the base of the Valiente Peninsula, Prov-
ince of Bocas del Toro. The island is roughly rectangular, with a
projecting point at the southeast and a somewhat irregular shoreline
on the western and northern sides. It is a little over 4 kilometers
long by less than 14 wide, with the long axis running east and west.
A sand beach extends along three-fourths of the southern side, around
the flat, open southeastern point, and across the eastern side, past
the mouth of a small stream, to end against a cliff, 12 meters high,
of sandy, indurated clay. Similar bluffs separated by short stretches
of beach mark the shoreline along the west and north. The northern
side is broken by a small bay with a sand beach at its head. On the
west the sea has cut back into the land, leaving several small islets,
some of them barren except for grass and other low herbage, and
some with a crown of brush and trees. Wave action is steadily erod-
ing the low cliffs, forming small caves, and in some cases arches that
pass through projecting points to the sea on the opposite side. The
shallow bank surrounding the island indicates that this process has
served to reduce it in size. The land back of the southern beach,
elevated sufficiently above high-tide line to form a flat, is fringed
with coconut palms on the sea side. Behind these extends low jungle
in which scattered trees rise 15 to 20 meters tall. Toward the center
the surface is lower and is swampy, with two or three trickles of fresh
water, discolored by swamp peat, that drain to the sea. There is a
small stand of mangroves at the mouth of the stream that enters the
sea above the southeastern point.
Columbus during his fourth voyage sighted the island on Oc-
tober 17, 1502, when he came out of the Laguna de Chiriqui through
Canal del Tigre (Tiger Channel) (Morison, 1942, vol. 2, p. 350).
He gave it the name El Escudo as it appeared to resemble an escudo,
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 2
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
or shield. In the following years the island became a landmark for
navigators along this stretch of coast, and is mentioned from time to
time in ancient documents, the name being abbreviated often to Scudo,
Scuda, or sometimes modified to Skoday (Anderson, 1911, p. 371).
Presently it was designated Escudo de Veragua, and finally the latter
part of the name became Veraguas. In the last voyage of Sir Francis
Drake (Hakluyt, 1904, pp. 239-240) it is related that his ships came
to Escudo on January 10, 1596, where they anchored on the southern
side, remaining until January 23. The island was described as “not
past two leagues long full of wood, and hath great store of fresh
water . . . and that very good.” Many of the men soon fell sick,
and Drake himself contracted the illness that caused his death on
January 28 when they were near Portobello. He was buried at sea
off that harbor.
In occasional seventeenth-century accounts of buccaneers and other
voyagers there is casual reference to Escudo de Veraguas as a place
of shelter or a source of water. Dampier’s observation (Dampier,
1697, p. 39) made in 1681 that “We past by Scuda, a small
Island (where ’tis said Sir Francis Drake’s Bowels were bury’d)”
repeats a tale, apparently of common belief, that cannot concern this
island since Drake’s death and burial, off Portobello, came more than
200 kilometers to the east. Escudo was visited by Indians, since
Dr. Matthew W. Stirling of the Smithsonian Institution informs me
that in the town of Bocas del Toro he was shown artifacts found on
the island, proof that aboriginal people had lived there, at least from
time to time. But there may be confusion with some larger place in
the report (Anderson, p. 272) that records a considerable Indian
population, divided under two caciques or chiefs. The land area, with
due allowance for a reasonable amount of erosion since these early
times, is too small to have permitted permanent residence for many
persons.
At present men come at intervals to gather the coconuts, or occa-
sionally to fish, search for turtles, or to hunt the introduced wild pigs.
There is no permanent human resident, and the wildlife, except for
the pigs, is tame.
I was able to visit Escudo de Veraguas through the kind assistance
of George Munch, manager of the Almirante Division of the Chiriqui
Land Co., which has its headquarters at Almirante, Province of Bocas
del Toro. We left Almirante on February 28, 1958, shortly before
midnight, on the diesel launch Talamanca, entered the sea through
the pass of Boca del Toro, and before dawn anchored in the lee on
the southeastern end of the island. Accompanied by Ziska Hartmann
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 3
and Jorge Burke, I was ashore near the southeastern point shortly
after 7 o’clock and during the forenoon worked through the southern,
level section parallel to the beach nearly to the western end. As the
sun rose higher the humidity and heat of the dense jungle, where no
breeze could enter, became oppressive, so that it was pleasant at the
end to walk back to our cayuco along the open beach.
At dawn the following morning the breeze blew from the mainland
to the south, so that waves were breaking on the beach. We went off
before 7:00 in a choppy sea, and finally landed near the mouth of the
small stream. I crossed first into the ridge area at the northeast, but
finding this difficult travel and unproductive I sought more level
ground. Through this I crossed again toward the western end parallel
to the northern shore. The sky was overcast, one shower of rain came,
and at times it was difficult to see birds in the heavy jungle shadows.
Though there were no trails, the low jungle was open and easy to
penetrate. Where the growth became dense the ground was covered
heavily with vines. On the north and west the surface rose Io to 25
meters in broken, steep-sided ridges, separated by little valleys. Here
there was much undergrowth of the spiny pita (a plant of the pine-
apple family) which, with the steep, slippery slopes, and the swampy
floors of the small valleys between, made it difficult to get about. The
taller trees that grow along the crests of these ridges from the sea
give a misleading appearance of true high forest.
On this final day we returned to the launch a little after 11:00 and,
as the sea was rising, left for Almirante, returning through Crawl Cay
Channel.
The only record of any earlier visit of a naturalist to the island is
the skin of a white-crowned pigeon in the collections of the University
of California at Los Angeles. From the end of February to early in
April 1936, Dr. Loye Holmes Miller of the Department of Zoology
of that Institution, on sabbatical leave, accompanied by a graduate
student, Frank Richardson, as assistant, visited the Laguna de Chiri-
qui, living on a barge that served as a base for a Navy Hydrographic
Office detail engaged in a survey of the area. Dr. Miller informs me
that on March 2 Richardson accompanied a shore party of Navy per-
sonnel to Escudo and brought back a white-crowned pigeon. No other
specimens were taken.
While Escudo de Veraguas lies well offshore, it is located on a bank
where the sea is shallow. A narrow trench of 24 to 35 fathoms lies
to the west and southwest, but elsewhere the depths are considerably
less. Since it is estimated that sea levels dropped from go to 120
meters during the last period of extensive glaciation in Wisconsin
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
time during the Pleistocene, it is apparent that then the island was
part of the mainland. A similar connection should have come during
part or all of the three preceding periods of maximum glacial ice.
Return of warmer temperatures in the interglacial periods, which
melted the ice, again raised the water level, placing Escudo once more
as an island, remote at sea. It is reasonable to suppose that the resi-
dent wren and the manakin, as well as the peculiar spiny rat of the
island, were established there during one of the periods of land con-
nection, since they are jungle creatures that do not range far from
cover, nor are the birds of kinds that would be readily windblown
by violent storms. Whether the characters of size and color that now
mark them were theirs in whole or in part on their arrival, or whether
these are distinctions that have developed during isolation, cannot
be said, except that it seems probable that the peculiarity of greater
size may have become intensified, since this condition is found regu-
larly in populations that seem to have been restricted for long periods
to small islands. The manner of development of the differences that
mark the blue-gray tanager is not easily understood since in mainland
regions these birds appear to roam far. It would appear that they may
not cross fairly wide water barriers, since another insular form is
found on Isla Coiba off the Pacific coast of Panama (Wetmore, 1957,
p. 94).
Though there were few species of resident birds on Escudo de
Veraguas, individuals were fairly numerous. The songs of the bay
wren, joined occasionally by the raucous notes of a small flock of
parrots, were regular bird notes of the jungle, aside from which there
were only the subdued sounds of the wind in the higher treetops, and
of the wash of waves against the shoreline. The smaller birds were
encountered mainly in the more level areas, where at times they were
detected with difficulty in the dim shadows that prevailed in the
thickets when the sky was overcast. Occasionally I noted large spiny
rats of the genus Hoplomys. One that I shot on the ground proves
to be a form new to science.
ANNOTATED LIST
Family PELECANIDAE: Pelicans
PELECANUS OCCIDENTALIS Linnaeus: Brown Pelican, Alcatraz
Pelecanus occidentalis Linnaeus, Systema naturae, ed. 12, vol. 1, 1766, p. 215.
(Jamaica. )
Several were fishing around the island on the morning of March 2.
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 5
Family SuLipAE: Boobies
SULA LEUCOGASTER LEUCOGASTER (Boddaert): Brown Booby, Piquero
Moreno
Pelecanus Leucogaster Boddaert, Table des planches enluminées, 1783, p. 57.
(Cayenne. )
Scattered groups rested on small islets off the western end of the
island, selecting those that were rocky or covered with short herbage.
They were nesting here, as I noted several large down-covered young.
At sunset adults came in from the open sea, flying low above the water,
singly or in groups of three or four. As our launch passed, a number,
part of them fully grown young, came flying out from the islets to
circle about with evident curiosity. There were no frigate-birds here,
and so the boobies were free from molestation. I estimated that about
200 individuals were present.
Family CHARADRIIDAE: Plovers, Turnstones
CHARADRIUS SEMIPALMATUS Bonaparte: Semipalmated Plover, Chorlito
Semipalmado
Charadrius semipalmatus Bonaparte, Journ. Acad. Nat. Sci. Philadelphia, vol. 5,
August 1825, p. 98. (Coast of New Jersey.)
A flock of 14 ranged the beach at the southeastern end of the island.
Family ScoropacipaAE: Snipe, Woodcock, Sandpipers
ACTITIS MACULARIA (Linnaeus): Spotted Sandpiper, Playerito Coleador
Tringa macularia Linnaeus, Systema naturae, ed. 12, vol. 1, 1766, p. 249. (Penn-
sylvania. )
One seen on March 1.
NUMENIUS PHAEOPUS HUDSONICUS Latham: Whimbrel, Zarapito
Trinador
Numenius hudsonicus Latham, Index ornithologicus, vol. 2, 1790, p. 712. (Hud-
son Bay.)
One seen on the beach March 1.
Family CoLuMBIDAE: Pigeons, Doves
COLUMBA LEUCOCEPHALA: White-crowned Pigeon, Paloma Cabeciblanca
Columba leucocephala Linnaeus, Systema naturae, ed. I0, vol. 1, 1758, p. 164.
(Bahama Islands.)
Two were seen March 1 in the top of a thickly leaved tree. A male
in the collection of the University of California at Los Angeles was
shot on March 3, 1936, by Frank Richardson, now of the Department
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
of Zoology of the University of Nevada, at the time student assistant
with Dr. Loye Holmes Miller (see p. 3).
Family PsrrracipAE: Parrots, Macaws
AMAZONA AUTUMNALIS SALVINI (Salvadori): Red-fronted Parrot, Loro
Frentirrojo
Chrysotis salvini Salvadori, Catalogue of the birds in the British Museum, vol.
20, 1801, p. 271. (Lion Hill Station, Canal Zone, Panama.)
Three pairs were seen in the early morning of March 1, and a
female was collected. The same small group was observed the fol-
lowing day.
Family TRocHiLipAE: Hummingbirds
AMAZILIA TZACATL TZACATL (De la Llave): Rieffer’s Hummingbird,
Colibri Colimorena
Trochilus Tzacatl, De la Llave, Registro Trimestre, vol. 2, No. 5, 1833, p. 48.
(México. )
Several were observed among the lower shrubs back of the beaches.
Family ALCEDINIDAE: Kingfishers
MEGACERYLE TORQUATA TORQUATA (Linnaeus): Ringed Kingfisher,
Martin Pescador Grande
Alcedo torquata Linnaeus, Systema naturae, ed. 12, vol. I, 1766, p. 180.
(México. )
One was recorded on March 2 near the mouth of the small stream
at the southeastern end.
Family PipripAE: Manakins
MANACUS VITELLINUS (Gould): Gould’s Manakin, Matraco
Pipra vitellina Gould, in Hinds, R. B. (editor), Zoology of the Voyage of
H.M.S. Sulphur under the command of Captain Sir Edward Belcher, R.N.,
F.R.G.S., etc., during the years 1836-42, vol. 1, pt. 3 (Birds, pt. 1), October
1843, p. 41, pl. 21. (Panama = Panama City, Panama.)
The manakin (fig. 1) was fairly common, ranking next to the wren
in abundance. The birds were found among the branches of the
smaller trees, where they were quiet, moving about rather slowly,
often remaining motionless for several minutes at a time. I regretted
that there was no indication of display among the males, as their
larger size should make the noises that accompany these activities
definitely impressive.
The bird of Escudo de Veraguas was so different from the repre-
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—-WETMORE Ai
sentative of this species around Almirante Bay that I recognized it as
an unknown race when the first specimen came to hand. It is described
in the following paragraphs:
MANACUS VITELLINUS AMITINUS, subsp. nov.
Characters ——Similar to Manacus vitellinus cerritus Peters? but
definitely larger; bill distinctly larger and heavier; tarsi and toes
Fic. 1—Gould’s manakin, Matraco.
heavier ; adult male with lower back, rump, and posterior ventral sur-
face, including the sides and under wing coverts, darker green ; female
and immature male somewhat darker green throughout, with the
abdomen less yellowish.
Description.—Type, U.S.N.M. No. 468919, male adult, from Isla
Escudo de Veraguas, Prov. Bocas del Toro, Panama, March 2, 1958,
collected by Alexander Wetmore (orig. No. 22241). Entire crown
1 Manacus cerritus Peters, Proc. New England Zool. Club, vol. 10, September
22, 1927, p. 9. (Almirante, Bocas del Toro, Panama.)
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
to nape, including the lores, wings (except inner lesser coverts), upper
back, and tail black; sides of head, throat and neck, including hind-
neck, bright apricot yellow, becoming lemon chrome as the yellow
collar meets the black of the back; lesser wing coverts, except the
outermost, lemon chrome; lower back, rump, and upper tail coverts
yellowish oil green; breast, sides, flanks, abdomen, and under tail
coverts between warbler green and olive-green; an indefinite light
wash of lemon yellow on center of breast and abdomen; outer under
wing coverts Roman green, inner ones yellowish citrine; under sur-
face of inner webs of primaries and secondaries, except toward the
tips, dull white. Bill dull black; tarsus and toes fuscous; claws dark
neutral gray (from dried skin).
Measurements——Males (3 specimens), wing 59.3-61.4 (60.6), tail
39.2-42.0 (40.2), culmen from base 14.0-14.8 (14.5), tarsus 23.8-24.5
(24.1) mm.
Females (2 specimens), wing 59.5-60.0 (59.7), tail 38.2-38.3 (38.3),
culmen from base 14.7-14.7 (14.7), tarsus 21.0-21.5 (21.2) mm.
Type, male, wing 59.3, tail 39.4, culmen from base 14.6, tarsus
23.8 mm.
Range.—Isla Escudo de Veraguas, at sea off the base of Peninsula
Valiente, Bocas del Toro, Panama.
Remarks.—The greater size of this handsome bird as compared
with mainland forms is evident on comparing the measurements with
those listed in succeeding paragraphs. In bulk the island birds appear
nearly one-third greater. In drawing the description comparison has
been made with cerritus since the shades of yellow on head and neck
of these two are more nearly in agreement. In terms of present distri-
bution Manacus v. vitellinus is assumed to be the form of the main-
land opposite Isla Escudo, since it is the one recorded at Cricamola
on the shores of Laguna de Chiriqui, opposite Peninsula Valiente.
Manacus v. cerritus is known to range south only to the southern
shores of Almirante Bay so that if the water barrier is disregarded,
cerritus and amitinus are separated by an intervening population of
typical vitellinus.
The name is taken from the Latin amitinus, a cousin.
To determine clearly the affinities of the manakin from Escudo a
survey has been made of the related members of the genus Manacus
found in Panama, particularly Manacus vitellinus, of which an excel-
lent series is at hand from the entire range including Colombia. It
became evident immediately that cerritus, described by James L. Peters
as a distinct species, was in fact a geographic race of M. vitellinus,
as the supposed specific characters break down when the entire area
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 9
occupied by this bird is given review. It may be noted also that the
display of males of cerritus, as I saw it in January and February 1958,
was similar to that of typical vitellinus.
Following is a summary of the subspecies of vitellinus based on this
examination, with the races arranged in geographic sequence from
west to east.
MANACUS VITELLINUS CERRITUS Peters
Manacus cerritus J. L. Peters, Proc. New England Zool. Club, vol. 10, Sep-
tember 22, 1927, p. 9. (Almirante, Bocas del Toro, Panama.)
Characters.—Similar in color pattern, and in colors in general, to
Manacus v. vitellinus. Male, with throat, sides of head, and band
across hind neck and upper back more yellow, less orange, varying
in some to completely bright yellow; lower breast, abdomen, sides,
flanks, and under tail coverts more greenish yellow ; rump and upper
tail coverts brighter green; female, and male in immature plumage,
darker green throughout.
Measurements.—Males (9 specimens), wing 51.8-54.2 (53.3), tail
31.2-35.8 (34.2), culmen from base 11.1-12.3 (11.7), tarsus 20.0-22.6
(21.5) mm.
Females (3 specimens), wing 54.0-55.7 (54.9), tail 33.1-34.4
(33.9), culmen from base 11.8-12.5 (12.0), tarsus 20.2-21.4 (20.9)
mm.
MANACUS VITELLINUS AMITINUS Wetmore
Characters—Generally similar to M. v. cerritus, but decidedly
larger ; darker green.
Range.—Isla Escudo de Veraguas, Province of Bocas del Toro,
Panama.
Full details of differences, and of measurements, are given in the
description above.
MANACUS VITELLINUS VITELLINUS (Gould)
Pipra vitellina Gould, in Hinds, R. B. (editor), Zoology of the Voyage of
H.M.S. Sulphur, under the command of Captain Sir Edward Belcher, R.N.,
F.R.G.S., etc., during the years 1836-42, vol. 1, pt. 3 (Birds, pt. 1), October
1843, p. 41, pl. 21. (“Panama’= Panama City, Panama.)
Characters Similar to M. v. cerritus, but male decidedly orange
on foreneck, throat, sides of head, and band across base of neck;
posterior under surface more greenish; rump and upper tail coverts
grayer green.
Measurements.—Males (47 specimens), wing 50.4-55.7 (52.3), tail
He) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
25.8-31.5 (28.3), culmen from base 11.0-13.0 (11.8), tarsus 20.4-
22.4 (21.4) mm.
Females (46 specimens), wing 50.7-54.9 (53.2), tail 27.3-31.7
(29.3), culmen from base 11.1-12.7 (12.0), tarsus 18.3-20.7 (19.4)
mim.
Range.—On the Pacific slope from the foothills of eastern Veraguas
(Santa Fé) eastward through the western part of the Province of
Panama (La Campana, Chorrera), throughout the Canal Zone, and
eastern Panama, to extreme eastern Darién (Jaqué, Rio Jaqué, Cana) ;
on the Caribbean slope from central Bocas del Toro (Cricamola),
through northern Veraguas (Guaval on Rio Calovevora), northern
Coclé (El Uracillo), the Province of Col6n (Chilar, Portobello) and
the Comarca de San Blas (Mandinga, Permé, Obaldia) ; entering
Colombia on the western side of the lower Rio Atrato (Unguia,
Chocéd) and along the shores of the Gulf of Uraba at Acandi, Choco,
on the western side, and Necocli, Antioquia, on the east.
This is the first published report of this race for Colombia. Speci-
mens from Acandi and Unguia, both near the Panamanian boundary,
are like typical examples from Panama. A series of 7 males from
Necocli on the eastern shore of the mouth of the Gulf of Uraba
averages faintly paler, more yellowish green below, and very faintly
more yellowish orange on the head. They thus show an approach
toward the paler milleri of the Sint Valley to the east, but are to be
placed with vitellinus.
Gould published the description of this manakin twice, first in the
Zoology of the Voyage of H.M.S. Sulphur, where it appeared in
October 1843 as indicated above. The bird was displayed with 8 other
new species from this voyage at a meeting of the Zoological Society
in London in July 1843, but publication in the Proceedings did not
come until December. In the first publication, in October, Gould
states that “The specimen here figured was procured by Mr. Hinds at
Panama, and is the only one I have seen.” The introduction to the
Voyage of the Sulphur indicates that the vessel made surveys along
the entire Pacific coast of the Republic, but it appears clear that the
locality “Panama” refers to the vicinity of Panama City, which is the
only place mentioned that lies within the range of vitellinus. This is
accepted, therefore, as the restricted type locality.
MANACUS VITELLINUS VIRIDIVENTRIS Griscom
Manacus vitellinus viridiventris Griscom, Bull. Mus. Comp. Zodl., vol. 69, April
1920, p. 179. (Jiménez, near Buenaventura, Valle, Colombia.)
Characters.—Similar to M. v. vitellinus, but male with lower breast,
abdomen, sides, flanks, under tail coverts, rump, and upper tail coverts
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE II
definitely darker green ; yellow of anterior part of body, including the
neck band, somewhat less orange, more yellow ; female darker green,
in this resembling female M. v. cerritus, from which it differs in
being somewhat less yellowish on the abdomen, and duller green above.
Measurements —Males (14 specimens), wing 50.6-53.7 (52.2),
tail 26.3-30.6 (28.7), culmen from base 10.8-12.5 (11.6), tarsus 20.4-
22.7 (21.5) mm.
Females (6 specimens), wing 53.0-54.3 (53.5), tail 28.1-30.1
(29.5), culmen from base 11.6-12.4 (11.9), tarsus 19.1-20.0 (19.6)
mm.
Range.—Western Colombia, from northern Chocd (Rio Jurado,
Rio Jurubida, Nuqui) and northwestern Antioquia (Villa Artiaga,
Dabeiba) south through western Caldas (Santa Cecilia) and Valle
(Puerto Muchimbo, Jiménez), including the upper Cauca Valley
(Riofrio, Cali).
This race has been supposed to range into extreme eastern Darién
at Cana but specimens from that locality agree best with typical
vitellinus,
MANACUS VITELLINUS MILLERI Chapman
Manacus vitellinus milleri Chapman, Bull. Amer. Mus. Nat. Hist., vol. 34,
Dec. 30, 1915, p. 645. (Puerto Valdivia, Antioquia, Colombia.)
Characters—Much paler than M. v. vitellinus; male with head
(except for the black crown) and band across hindneck bright, light
yellow, without orange; rest of lower surface much paler, being
grayish green with a wash of yellow; rump and upper tail coverts
paler ; female, definitely paler below, being whitish on abdomen, and
duller, grayer green above.
Measurements—Males (11 specimens), wing 49.7-52.9 (51.6), tail
26.8-30.4 (28.6), culmen from base 10.8-12.2 (11.5), tarsus 20.6-22.3
(21.3) mm.
Females (6 specimens), wing 52.5-54.5 (53.7), tail 28.8-30.8
(29.6), culmen from base I1.0-12.0 (11.6), tarsus 19.0-20.0 (19.4)
mm.
Range.—Northwestern Colombia, from the valley of Rio Sint
(Nazaret, Socarré) in western Bolivar, south to the middle Cauca
Valley in northern Antioquia (Taraza, Puerto Valdivia) ; recorded
from Remedios in east central Antioquia at the head of Rio Ité, a
tributary of the lower middle Rio Magdalena.
In the series at hand this race is typical on the middle Rio Cauca in
northern Antioquia. In some specimens from the lower Rio Sint,
taken at Nazaret, Tierra Alta, Socarré, and Quebrada Salvajin, most
of the males have the head somewhat more orange, and the breast
I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
and abdomen somewhat darker, varying in the direction of vitellinus.
They are thus somewhat intermediate, but are definitely near mulleri.
It has been suggested that Manacus aurantiacus (Salvin) found on
the Pacific slope of western Panama would eventually prove to be
conspecific with M. vitellinus, but my studies to date do not bear out
this supposition. Brighter color, particularly in the male, and smaller
size mark aurantiacus uniformly throughout its range from south-
western Costa Rica through Chiriqui, southern Veraguas, and both
sides of the Azuero Peninsula in Veraguas, Herrera, and Los Santos.
Manacus vitellinus vitelinus from near Santa Fé, Veraguas, and
La Campana and Chorrera in the western section of the Province of
Panama, where intergradation, if present, should occur, show no
variation from the normal pattern of that race. From present infor-
mation the two groups appear to be separated by a savanna area in
which neither is found. The two appear so completely distinct that
there is no basis for uniting them.
Aldrich (1937, p. 95) separated the population of the western side
of the Azuero Peninsula as Manacus aurantiacus flaviventris, as a
series from that area appeared brighter colored than those available
at the time from western Chiriqui and southwestern Costa Rica.
During the course of my own field investigations I have accumulated
a considerable series from Veraguas and eastern Chiriqui, and have
examined additional material from western Chiriqui and Costa Rica.
A study of this extensive material indicates that the supposed differ-
ences do not hold. Males in fresh plumage from both areas are
strongly orange, but as the season progresses there is fading, particu-
larly in the dry months when sun is more intense.
The following measurements may be useful for comparison with
those of the races of Manacus vitellinus.
Males (25 specimens), wing 44.8-47.8 (46.3), tail 26.0-30.2 (28.7),
culmen from base 11.2-12.2 (11.7), tarsus 19.5-20.6 (20.1) mm.
Females (21 specimens), wing 47.8-50.0 (48.7), tail 209.0-30.9
(30.3), culmen from base 11.3-12.3 (11.8), tarsus 18.2-20.5 (19.1)
mm.
Family TyrANNIDAE: Tyrant Flycatchers
TYRANNUS MELANCHOLICUS CHLORONOTUS Berlepsch: Tropical King-
bird, Pechi-amarillo Grande
Tyrannus chloronotus Berlepsch, Ornis, vol. 14, 1907, p. 474. (Temax, Yuca-
tan.)
A female was collected and several others seen along a stretch of
sandy beach, where they rested on the open ends of branches, or on
the tops of low shrubs.
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 13
Family HirUNDINIDAE: Swallows
PROGNE SUBIS (Linnaeus): Purple Martin, Golondrina Turquina
Hirundo Subis Linnaeus, Systema naturae, ed. 10, vol. 1, 1758, p. 192. (Hudson
Bay.)
On the return journey on March 2 I noted an occasional purple
martin flying northward, low over the water, near the mainland coast
from the vicinity of Plantain Cay to Chiriqui Point. These swallows
are known as migrants through México and Central America, but little
is reported regarding them in Panama. The only published record
that has come to my attention is by Zimmer (1955, pp. 4, 5) of an
immature male of the southwestern subspecies, Progne subis hesperia
Brewster, taken at Cocoplum, Bocas del Toro, October 27, 1927.
At Almirante on February 18, 1958, during a forenoon of nearly
constant rain, a band of 8 purple martins came to rest in dead branches
of a tall avocado tree beside our house. At intervals others arrived
until finally between 35 and 40 were present, resting in close forma-
tion. When the rain ceased and the sky became lighter two hours later
they disappeared. From then until March 6, I recorded purple martins
in northward flight, singly or in scattered, straggling groups, across
Almirante Bay, along its shoreline, or over the outer beach near
Boca del Drago. Occasionally a few came to rest in the tree beside
the house. It appears that there is a regular flight in migration along
the Caribbean coast.
The female of a pair taken on February 18, in its darker color on
the under surface and in wing length of 148 mm., represents typical
Progne subis subis. The male, with the wing 149.7 mm., agrees in
size with that race.
Family TRoGLoDYTIDAE: Wrens
THRYOTHORUS NIGRICAPILLUS Sclater: Bay Wren, Cucarachero Castafio
Cabecinegro
Thryothorus nigricapillus Sclater, Proc. Zool. Soc. London, pt. 28, May 1860,
p. 84. (Nanegal, 4,000 feet elevation, Ecuador.)
This wren (fig. 2) was the most common land bird on the island,
found in pairs scattered through the undergrowth. Though they were
encountered most often in low tangles, where creepers were matted
and cover was dense, they ranged also out into more open areas, and
at times worked up through branches and creepers into the tops of
the taller trees. They were quite tame, often appearing within 6 feet
or so. On our second day ashore the sky was overcast and it was
often difficult to see these birds in the darkly shadowed coverts. We
were usually notified of the presence of a pair by the series of repeated
I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
notes that made up the clear song. This resembled closely that of
Thryothorus nigricapillus costaricensis as heard at Almirante, but
seemed to be higher in tone and somewhat less varied in repertoire.
One pair worked busily at a nearly completed nest located near the
tip of a leafy branch about 6 feet from the ground in heavy under-
growth. This was a ball, nearly round, of palm and other slender
Fic. 2.—Bay wren, Cucarachero Castaiio Cabecinegro.
fibers, with the ends projecting all around as a rough fringe. The
entrance was in one side.
The larger size and paler color of this island population in com-
parison with the birds of the adjacent mainland were easily evident
in the field. A description of this previously unknown race follows.
THRYOTHORUS NIGRICAPILLUS ODICUS subsp. nov.
Characters —Similar to Thryothorus nigricapillus costaricensis
(Sharpe)? but larger, with longer, heavier bill; in color paler brown.
2 Thryophilus costaricensis Sharpe, Catalogue of the birds in the British Mu-
seum, vol. 6, 1881, p. 217. (Valley of the Rio San Carlos, Alajuela, Costa Rica.)
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 15
Description—Type, U.S.N.M. No. 469015, male adult, from Isla
Escudo de Veraguas, Bocas del Toro, Panama, taken March 1, 1958,
by Alexander Wetmore (original No. 22230). Throat, upper fore-
neck, malar region, loral area, a line on the margin of upper and
lower eyelids surrounding the eye, a superciliary line extending back
from the center of the eye, and the auricular region white, with some
mixture of black on loral area and along upper eyelid; crown, hind-
neck, side of neck, side of head, except as noted above, and a line
separating the white malar area from the throat, deep black; back,
rump, and upper tail coverts auburn, the tail coverts with short central
bars of black along the shaft; wing coverts auburn, with irregular
shaft lines and subterminal bars of dusky neutral gray; tertials and
outer webs of secondaries auburn, barred heavily with dusky neutral
gray; outer webs of innermost primaries auburn, changing on the
outer ones to hazel, the brighter color finally reduced to a narrow
edging on the ninth and tenth; concealed webs of remiges fuscous-
black ; rectrices dusky neutral gray, barred narrowly with hazel ; breast
and center of abdomen ochraceous-tawny; sides and flanks hazel;
under tail coverts ochraceous-tawny, barred heavily with black; axil-
lars ochraceous-tawny; under wing coverts ochraceous-buff, mixed
with white ; edge of wing white. Maxilla dusky neutral gray ; mandi-
ble pale smoke gray, becoming smoke gray at the base; tarsus and toes
fuscous-black (from dried skin).
Measurements.—Males (5 specimens), wing, 75.2-79.2 (77.0), tail
58.6-62.3 (60.2), culmen from base 21.8-24.2 (23.2), tarsus, 28.4-
21-8 (29.7): mm.
Females (6 specimens), wing 70.2-72.8 (71.6), tail 54.5-58.8
(56.8), culmen from base 21.0-22.3 (21.5), tarsus 26.2-28.7 (27.2)
mm.
Type, male, wing 75.2, tail 58.8, culmen from base 24.0, tarsus
29.1 mm.
Range.—Isla Escudo de Veraguas, at sea off the base of the
Valiente Peninsula, Bocas del Toro, Panama.
Remarks.—tThe actual difference in measurements will be indicated
by consulting the summary of a series of Thryothorus nigricapillus
costaricensis, the nearest relative, both physically and geographically,
that is given in the review of the species that follows.
The name of the new race is from the Latin odicus, musical, appro-
priate because of the pleasing song.
The complete and definite dissimilarity in the lower surfaces found
in this group of wrens between the chestnut-breasted, white-throated
groups of the Caribbean slope of Nicaragua, Costa Rica, and Bocas
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
del Toro, and the forms with the anterior under surface barred closely
with black and white that range from eastern Darién through western
Colombia to Ecuador, long led to their separation under two specific
names. The series of specimens now available justifies their union
under the specific name nigricapillus, though it may be supposed that
the two terminal groups must have been separated for a long period
to have become so completely different. In costaricensis, the darkest
of the Central American races, and the one farthest removed from
those of South America, remote common ancestry with the other is
indicated in the rather indistinct black bars found on the breast and
sides in the juvenile plumage. This marking may persist in the fol-
lowing plumage, especially on the abdomen, but many are plain chest-
nut on the posterior lower surface except for the bars on the lower tail
coverts that are common to many of the numerous species of the
genus Thryothorus. Proceeding eastward along the Caribbean coast
of Panama from the valley of the Rio Calovevora, on the boundary
between the provinces of Bocas del Toro and Veraguas, the wrens of
the species under discussion become paler brown, with sides and flanks
barred with black, except for occasional plain individuals. This group
—the race castaneus—is found through the lowland Caribbean drain-
age of the Canal Zone.
Continuing eastward there is an abrupt change near Portobello
and in the foothills of the Cerro Azul in which the plain white of the
throat extends down on the upper breast, the brown on the sides
becomes paler, and there are strongly marked black bars on sides,
lower breast, and abdomen in most individuals. This style—the race
reditus—crosses to the Pacific slope along the base of the Cerro Azul,
and at Chiman has reached the coastal lowlands. On the Caribbean
slope it continues almost to the Colombian boundary in the Comarca
de San Blas, and on the Pacific side to about the western boundary of
Darién near the Golfo de San Miguel. There is then rather abrupt
transition to birds with lower surface heavily barred—the race
schotiti. Markings on the white throat are faint or absent, and the
brown is restricted to the flanks and under tail coverts. In the valley
of the Atrato the barring reaches its maximum and here the throat
in most specimens is heavily marked. The plainer throat persists to
the eastward in Colombia along the Rio Sinu, and on the middle and
upper Rio Cauca. In southwestern Colombia, beginning in the Depart-
ment of Cauca, the throat bars begin to lighten still more and to
disappear, and farther south, in Narino, the upper breast also becomes
less heavily marked. This style leads over to typical nigricapillus of
Ecuador, in which throat and upper breast are white, without bars, and
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 17.
the flanks and under tail coverts are lighter brown. In all the changes
that have been described these wrens have remained uniformly chestnut
above, with black crowns.
One possible explanation of this interesting gradient might be that
the plain, chestnut-breasted forms had become established fairly early
in the Central American area where they have continued with modifi-
cation toward the elimination of barring. In the South American area,
on the other hand, the barring became intensified. Through a subse-
quent spread of range in the latter population, the two groups have
been brought in contact, with resultant hybridization that has caused
the mixing that has been described.
The races recognized as reditus and castaneus represent two stages
in this process. It would appear that the schottii group has been the
one in active expansion because of the extensive range that it now
occupies. It is interesting that the chestnut-breasted group is not
found farther north in Central America, though there would appear
to be no ecological barrier to prevent this.
Hellmayr (1934, p. 180) includes another group, Thryothorus
semibadius Salvin, found in tropical lowlands of the Pacific slope
from southwestern Costa Rica to western Chiriqui, also as a race of
nigricapillus, but this does not seem justified. The bird in question is
more finely barred, with 3 narrow dark bars on the individual feathers
of the breast, and the crown chestnut, concolor with the back; also
it is smaller. In the schottu-nigricapillus group, which semibadius
resembles superficially, the black bars are heavier, there are 2 bars
on the individual feathers of the breast, the crown and upper hindneck
are deep black, and the size is larger. There is no indication whatever
of hybridization between semibadius and the adjacent Thryothorus n.
costaricensis. While juveniles of the costaricensis-nigricapillus group
show spots or a slight wash of brown on the pileum and hindneck, the
crown cap remains plainly defined. Thryothorus semibadius would
appear to be an older offshoot of the ancestral stock that has produced
the forms with barred breast, and from its limited range one that may
be on its way to extinction.
The following summary, based on extensive series throughout the
entire range of these birds, outlines findings as to their relationships
and distribution. It should be noted that museum series of skins almost
invariably include immature individuals that are not fully grown,
especially in the development of the wings. These are easily detected
and have been omitted in the measurements that are given under the
different forms.
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
THRYOTHORUS SEMIBADIUS Salvin: Salvin’s Wren, Cucarachero
Castafio Cabecimoreno
Thryothorus semibadius Salvin, Proc. Zool. Soc. London, November 1870, p. 181.
(Bugaba, Chiriqui.)
Characters—Crown and hindneck chestnut, concolor with the
back; under surface white, barred, except for the throat, narrowly
with black, the breast feathers having three black bars; size smaller.
Measurements.—Males (8 specimens), wing 61.4-65.5 (63.3), tail
42.0-49.2 (46.3), culmen from base 18.7-21.0 (19.9), tarsus 23.4-24.0
(23.7) mm.
Females (4 specimens), wing 61.2-64.7 (63.2), tail 43.7-47.4
(45.8), culmen from base 19.9-20.8 (20.3), tarsus 23.1-24.5 (23.8)
mm.
Range.—Tropical zone of the Pacific slope from southwestern
Costa Rica in the valley of the Rio Pirris to western Panama in the
Comarca del Bart (Puerto Armuelles), and the lowlands of extreme
western Chiriqui (Divala, Bugaba).
THRYOTHORUS NIGRICAPILLUS Sclater: Bay Wren, Cucarachero
Castaiio Cabecinegro
Thryothorus nigricapillus Sclater, Proc. Zool. Soc. London, pt. 28, May 1860,
p. 84. (Nanegal, 4,000 feet elevation, Ecuador.)
Characters.—Crown and hindneck deep black, in sharp contrast to
the chestnut of the remainder of the upper surface; under surface
chestnut, auburn, chestnut-brown, clay color, or white, barred more
or less with black ; in the races that are white below, with 2 black bars
on each breast feather ; size larger.
THRYOTHORUS NIGRICAPILLUS COSTARICENSIS (Sharpe)
Thryophilus costaricensis Sharpe, Catalogue of the birds in the British Museum,
vol. 6, 1881, p. 217. (Valley of the Rio San Carlos, Alajuela, Costa Rica.)
Characters—Throat and upper foreneck white, rest of lower sur-
face auburn to hazel; sides in some specimens with a few bars of
black, which usually are indistinct.
Measurements.——Males (17 specimens), wing 66.5-72.0 (69.3), tail
51.0-56.8 (54.3), culmen from base 20.4-22.7 (21.3), tarsus 24.5-27.8
(25.9) mm.
Females (9 specimens), wing 62.5-67.2 (64.6), tail 47.8-54.0
(50.2), culmen from base 19.4-21.7 (20.5), tarsus 23.2-25.6 (24.5)
mm.
Range.—Caribbean slope from southeastern Nicaragua (Los Saba-
los, Rio Escondido, San Juan del Norte) through eastern Costa Rica
(Rio Frio, Guayabo, Bonilla, Jiménez, Reventazén) to central Bocas
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 19g
del Toro, Panama. Specimens from Cricamola at the eastern end of
the Laguna de Chiriqui are intermediate toward castaneus.
Sharpe described this bird from a single specimen that he said was
collected by Adolphe Boucard in Costa Rica, without giving a more
definite locality. Boucard (1878, p. 51) in an account of his collec-
tions made in Costa Rica listed this wren as Thryophilus castaneus
Lawrence, with the statement “Several specimens, from San Carlos;
killed in February.” In his itinerary he says that this locality was in
the Valley of the Rio San Carlos, a tributary of the Rio San Juan on
the Atlantic slope. I have therefore designated this area as the type
locality.
THRYOTHORUS NIGRICAPILLUS ODICUS Wetmore
Characters —Similar to T. n. costaricensis but larger, with longer,
heavier bill; paler brown.
Measurements.—Given above.
Range.—Confined to Isla Escudo de Veraguas, off the base of the
Valiente Peninsula, Bocas del Toro, Panama.
THRYOTHORUS NIGRICAPILLUS CASTANEUS Lawrence
Thryothorus castaneus Lawrence, Ann. Lyc. Nat. Hist. New York, vol. 7, June
1861, p. 321. (“Atlantic slope near the Panama Railroad’= Lion Hill,
Canal Zone.)
Characters —Similar to T. n. costaricensis, but paler brown on
ventral surface, with the white of the throat extending farther down
on the foreneck, in some reaching the upper breast; more definitely
barred with black on sides and flanks, in some specimens with the
bars extending across the lower breast and abdomen.
Measurements.—Males (14 specimens), wing 66.2-70.7 (68.9), tail
49.4-53-7 (51.7), culmen from base 20.3-22.0 (21.0), tarsus 24.9-27.5
(25.8) mm.
Females (17 specimens), wing 63.1-67.0 (64.8), tail 46.3-53.4
(49.2), culmen from base 19.3-21.9 (20.2), tarsus 23.4-25.7 (24.7)
mm.
Range.—Caribbean slope from the valley of the Rio Calovevora in
eastern Bocas del Toro, through northern Veraguas, northern Coclé
(extending inland on the northern slope in the higher foothills to the
headwaters of the Rio Coclé del Norte and the Rio Indio), and
western Colon (Chilar, Rio Indio, Colén, Marajal), to the Canal Zone
(Gatun, Lion Hill, Barro Colorado Island, Frijoles).
Back of Fl Valle, Coclé, I found these birds at 2,000 feet elevation
along the upper course of the Rio Mata Ahogada, ranging on its
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
higher branches to 2,500 feet. The divide here between this stream,
which flows into the Pacific, and the Rio Indio of the Caribbean side
is low so that rainfall in the heads of the valleys is sufficient to main-
tain the type of green-leaved undergrowth that these wrens frequent
across for a short distance on the Pacific side. The birds here do not
range below 2,000 feet elevation where the scrub growth changes to
the semiarid type characteristic of the Pacific lowlands of this area.
This is the only point known to me at which the race castaneus crosses
to the Pacific slope. Records of Salvin (1867, p. 134) and of Salvin
and Godman (1880, p. 88) for Santiago de Veraguas are not sup-
ported by specimens in the Salvin and Godman collections now in the
British Museum (Natural History) and are certainly in error.
The type specimen of castaneus, described by Lawrence, came to
him in a collection made by James McLeannan and John R. Galbraith
during the winter of 1860-1861. The collectors were located at Lion
Hill, but it must be borne in mind that it is certain they covered a
considerable area along the line of the railroad in the course of their
work. In the present instance Lawrence (1861, pp. 315-316) states
that their specimens were taken ‘‘on the Atlantic side of the isthmus”
except for half a dozen species that he lists, which do not include the
bird here under consideration. Though the type specimen of castaneus
is labeled only ““Panama” with the initials of the collectors, the desig-
nation “Lion Hill” found in current literature may be accepted as the
restricted type locality.
THRYOTHORUS NIGRICAPILLUS REDITUS Griscom
Thryophilus nigricapillus reditus Griscom, Bull. Mus. Comp. Zool., vol. 72,
January 1932, p. 358. (Permé, Comarca de San Blas.)
Characters.—Similar to T. n. castaneus but with white of breast
more extensive; sides, abdomen, and under tail coverts paler, duller
brown; more heavily and extensively barred with black.
Measurements——Males (15 specimens), wing 67.0-70.5 (68.9), tail
47.5-54.3 (52.0), culmen from base 19.3-21.9 (20.9), tarsus 24.0-26.5
(25.5) mm.
Females (11 specimens), wing 63.2-67.7 (65.4), tail 45.0-51.4
(48.5), culmen from base 19.0-21.5 (20.1), tarsus 23.1-26.3 (24.7)
mm.
Range.—From eastern Colon (Portobello) eastward on the Carib-
bean slope through the Comarca de San Blas (Mandinga, Permé,
Puerto Obaldia), crossing through the western Cerro Azul to the head
of the Rio Pacora on the Pacific slope, ranging eastward in the Prov-
ince of Panama along the Pacific side of the Serrania de Majé
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 21
(Quebrada Cauchero, on the base of Cerro Chucanti), reaching tide-
water at Chiman, and on the Rio Majé (Charco del Toro).
This race constitutes the definite intergrade between the western
group with bright brown breast and little or no barring, and the
eastern and southern population with completely barred breast.
Transition between castaneus and reditus on the west is fairly abrupt,
an intermediate condition being evident in one specimen from near
Frijoles in the Chagres drainage. Birds from near Colon are definitely
castaneus, while those from near Portobello, 30 kilometers to the
east, are reditus. At the eastern end the type locality at Permé is
barely within the range, since skins from Puerto Obaldia, about
15 kilometers farther east, are intermediate toward schottii, which
is the race found on the coast at Acandi, Chocd, Colombia, 25 kilo-
meters beyond Puerto Obaldia.
THRYOTHORUS NIGRICAPILLUS SCHOTTII (Baird)
Thryophilus schottii Baird, Review of American birds in the Museum of the
Smithsonian Institution, vol. 1, August 1864, p. 123 (in Key) ; September
1864, p. 133. (Rio Truand6é, Choco, Colombia.)
Thryophilus nigricapillus connectens Chapman, Bull. Amer. Mus. Nat. Hist.,
vol. 31, July 23, 1912, p. 157. (Cocal, 5,000 feet elevation, Cauca, Colombia.)
Characters.—White of throat and foreneck extending down over
breast, sides, and center of upper abdomen; lower surface heavily
barred with black, in typical form the bars covering the throat, but
in intermediate stage the throat partly or wholly plain.
Measurements.—Males (16 specimens), wing 64.0-66.9 (67.3), tail
44.6-51.6 (48.2), culmen from base 19.5-21.5 (20.4), tarsus 24.4-26.8
(25.4) mm.
Females (10 specimens), wing 59.9-65.6 (63.0), tail 43.0-47.8
(45.6), culmen from base 19.0-20.8 (19.6), tarsus 23.0-25.0 (24.1)
mm.
Range.—Darién, eastern Panama, from the lower Rio Sambu
(Jesusito), and the lower Rio Tuira (Cituro, on Rio Cupe) inland
to 600 meters elevation near Cana, and south to the valley of the
Rio Jaqué; continuing in Colombia throughout Chocd (from the
Pacific coast across to Acandi on the Gulf of Uraba), and western
Antioquia in the Atrato valley (Villa Artiaga), and western Valle
(Buenaventura and San José), to western Cauca (Cocal) ; east into
southern Bolivar in the upper Sint Valley (Socarré, Quebrada
Salvajin), and northern Antioquia in the lower Cauca Valley (El
Pescado), and the valley of the Rio Nechi (Regeneracion, El Real,
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Hacienda Belén), crossing to the Rio Magdalena drainage on the
Quebrada Enanea (Volador).
Remarks.—The typical form of this race, with the throat and fore-
neck distinctly barred with black, is found mainly in the Choco. In
southwestern Colombia, through western Cauca, the throat barring
disappears, and in Narifio the breast appears whiter as the barring
on this area is reduced. The birds of this region are intergrades of
unstable character between schottii and nigricapillus. The influence
of the reditus style of markings produces similar intergrades on the
opposite side of the range, beginning in northern Chocé at Acandi on
the Gulf of Uraba, and extending across to the upper Sint Valley
and the lower Nechi. Specimens from this area are identical in appear-
ance with those of western Cauca which Chapman named connectens.
Under these circumstances there is no basis for recognition of such a
race, as the supposed characters, unstable at best, are duplicated on
the opposite side of the population of typical schotiu. The birds
described are allocated as intermediates to schottti, except for those
of Narifio which are placed best with typical nigricapillus.
THRYOTHORUS NIGRICAPILLUS NIGRICAPILLUS Sclater
Thryothorus nigricapillus Sclater, Proc. Zool. Soc. London, pt. 28, May 1860,
p. 84. (Nanegal, 4,000 feet elevation, Ecuador.)
Characters —Similar to T. n. schottti, but averaging lighter brown
on back, flanks, and under tail coverts ; throat, foreneck, and center of
upper breast immaculate, with the barring reduced on the sides.
Measurements.—Males (13 specimens), wing 62.6-67.1 (65.1), tail
44.2-50.8 (48.1), culmen from base 19.1-20.9 (20.1), tarsus 24.0-25.5
(24.6) mm.
Females (6 specimens), wing 63.9-66.8 (65.5), tail 46.6-50.7
(48.7), culmen from base 19.2-21.6 (20.0), tarsus 23.0-25.3 (24.2)
mm.
Range.—lrom western Narifio (intermediate) in Colombia south
through the tropical zone of western Ecuador, nearly to the boundary
with Pert.
Remarks.—As indicated under schottiu, specimens from Narifio are
intermediate.
Family MimipaE: Mockingbirds, Thrashers
DUMETELLA CAROLINENSIS (Linnaeus): Catbird, Pajaro Gato
Muscicapa carolinensis Linnaeus, Systema naturae, ed. 12, vol. 1, 1766, p. 328.
(Virginia. )
Three were noted, and one female was collected.
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 23
Family PAruLIDAE: Wood Warblers
DENDROICA PETECHIA ERITHACHORIDES Baird: Golden Warbler,
Canario Manglero
Dendroica erihtachorides (= erithachorides, typographical error, corrected in
index) Baird, Report of explorations and surveys ... for a railroad from
the Mississippi River to the Pacific Ocean, vol. 9, pt. 2, Birds, 1858, pp. 283,
976. (Cartagena, Colombia.)
These warblers (fig. 3) were found scattered through the taller
trees where they were fairly common, though each of the four taken
appeared to be alone. It should be noted that on Escudo they were not
restricted to the limited growths of mangroves found near the sea, as
is the case on the mainland, but ranged throughout the forest growth,
as appears to be the regular habit of this warbler when found on small
islands. On the present island they ranked third in abundance among
the smaller land birds. The four taken include three adult males which
are similar to a small series from the shores of Almirante Bay on the
nearby mainland. A female that had just begun the molt from the
gray juvenile dress to the yellow adult plumage had the skull fully
ossified, indication that this character as a criterion of age is not re-
liable in tropical areas, where the life cycle of an individual bird is not
necessarily arranged on a calendar year basis.
The series from Escudo and from Almirante Bay agree fully with
type material of this race, which is interesting since specimens from
Limon, Costa Rica, about too kilometers to the north, are Dendroica
p. bryanit.
Family THRAUPIDAE: Tanagers
THRAUPIS VIRENS (Linnaeus): Blue-gray Tanager, Azulejo
Loxia virens Linnaeus, Systema naturae, ed. 12, vol. 1, 1766, p. 303. (Surinam.)
Blue-gray tanagers were fairly common in the taller trees, a num-
ber being seen and three collected. It has been unexpected to find that
they are so different from the widely distributed race of the mainland
that they merit description as an additional subspecies.
THRAUPIS VIRENS CAESITIA subsp. nov.
Characters Similar to Thraupis virens diaconus (Lesson)* but
darker, particularly below; central lower surface nearly uniform in
shade from throat to under tail coverts; sides definitely darker ; bill
longer and heavier.
3 Tanagra (Aglaia) diaconus Lesson, Rev. Zool., June 1842, p. 175. (Realejo,
Nicaragua. )
24
SMITHSONIAN MISCELLANEOUS COLLECTIONS
Fic. 3—Golden warbler, Canario Manglero.
VOL. 139
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 25
Description—Type, U.S.N.M. No. 469168, female, Isla Escudo de
Veraguas, Bocas del Toro, Panama, March 2, 1958, collected by
Alexander Wetmore (original No. 22248). Crown mineral gray, with
a faint wash of gnaphalium green, which is stronger on hindneck ;
back and scapulars dull greenish glaucous-blue, changing to light
glaucous-blue on rump; upper tail coverts bluish gray-green, washed
with greenish glaucous-blue at tips; shoulder patch formed by lesser
and middle coverts, grayish violaceous blue ; primaries and secondaries
dusky neutral gray, with outer webs, except for the tips of the
primaries, dull Venetian blue; outer webs of scapulars dark gobelin
blue; central rectrices and outer webs of others dark gobelin blue,
with inner webs of all but the central pair dark neutral gray ; median
under surface between court gray and gnaphalium green, with center
of abdomen faintly whitish; sides gnaphalium green; edge of wing
glaucous-blue ; under wing coverts light gull gray to white. Bill dull
black, except for a wash of hair brown toward base of gonys; tarsus,
and toes dusky neutral gray (from dried skin).
Measurements—Females (3 specimens), wing 87.5-90.1 (88.4),
tail 60.1-62.8 (62.3), culmen from base 16.4-18.0 (17.1), tarsus 20.4-
20.7 (20.6) mm.
Type, female, wing 90.1, tail 62.8, culmen from base 18.0, tarsus
20.4 mm.
Range.—Isla Escudo de Veraguas, at sea off the base of the
Valiente Peninsula, Bocas del Toro, Panama.
Remarks.—The fact that this widely distributed tanager was repre-
sented by a distinct form on this small island was not detected until
I began examination of specimens in the preparation of the present
report. The three specimens, all females, were taken merely as a
matter of routine during my visit. Comparison has been made with a
series of recently collected skins, consisting of 15 females of Thraupis
virens diaconus, and 21 of T. v. cana. In none of these is there dupli-
cation of the characters on which the race caesitia is based. Attention
was first drawn to the island form by the large bill, this measuring
13.8 to 15.7 (14.6) mm. in the 15 diaconus, and 13.7 to 15.7 (14.7)
mm, in the 21 cana.
Hellmayr (1936, p. 214) expressed doubt as to the validity of the
race diaconus, and recently Blake (1958, p. 566) has combined this
form with cana. In comparing an extensive series taken throughout
the range of the two subspecies in question I find, however, that while
the two are similar in general, diaconus is darker on the back, and
slightly duller blue on the rump, in addition to averaging somewhat
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
darker in color below. These characters hold in birds of Central
America south through the Isthmus of Panama, with intergradation
in extreme northwestern Colombia. In making comparison it is
necessary to separate adult from immature birds, since the distinctions
listed are masked when this is not done. I believe the confusion
regarding the two races has been due to lack of understanding of this
fact.
The name given to the new race, in connection with its darker
coloration, is from the Latin caesitius, meaning bluish.
LITERATURE CITED
ALpricH, Joun W.
1937. Annotated list of birds, in Aldrich, J. W., and Bole, B. P., Jr., The
birds and mammals of the western slope of the Azuero Peninsula
[Republic of Panama]. Sci. Publ. Cleveland Mus. Nat. Hist., vol.
7, pp. 27-130, Aug. 31.
ANDERSON, C. L. G.
tot. Old Panama and Castilla del Oro. Pp. i-xiv, 1-550, 7 maps, 35 pls.
Washington.
BLakeE, Emmet R.
1958. Birds of Volcan de Chiriqui, Panama. Fieldiana: Zool., vol. 36, No.
5, PP. 499-577, June 25.
Boucarp, ADOLPHE.
1878. On birds collected in Costa Rica. Proc. Zool. Soc. London, 1878,
PP. 37-71, I pl.
DAMPIER, WILLIAM.
1697. A new voyage round the world, describing particularly, the Isthmus
of America, several coasts and islands in the West Indies, the Isles
of Cape Verd, the passage by Tierra del Fuego, the South Sea
coasts of Chili, Peru and Mexico; the Isle of Guam one of the
Ladrones, Mindanao, and other Philippine and East India islands
near Cambodia, China, Formosa, Luconia, Celebes&c. New Hol-
land, Sumatra, Nicobar Islands; the Cape of Good Hope and Saint
Hellena. London, pp. (1-8), i-vi, 1-550, 5 maps.
Haxtuyt, RIcHarp.
1904. The principal navigations voyages traffiques & discoveries of the
English nation made by sea or over-land to the remote and farthest
distant quarters of the earth at any time within the compasse of
these 1600 years. Vol. X, pp. 226-245. Glasgow.
HELLMAYR, CHARLES E.
1934. Catalogue of birds of the Americas and the adjacent islands. Field
Mus. Nat. Hist., pt. 7, pp. i-vi, 1-531, Nov. 15.
1936. Idem, pt. 9, pp. i-vi, 1-458, Oct. 6.
Mortson, SAMUEL ELroT.
1942. Admiral of the Ocean Sea, A life of Christopher Columbus. Vol. I,
pp. i-xlvi, 1-448, 14 maps, 25 ills.; vol. 2, pp. i-viii, 1-445, 18 maps,
2 illus. Boston.
NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 27
SALvIN, OSBERT.
1867. On some collections of birds from Veragua. Proc. Zool. Soc. London,
1867, pt. I, pp. 129-161, 1 pl., June.
SALVIN, OsBERT, and GopMAN, FREDERICK DUCANE.
1879-1904. Biologia Centrali-Americana, Aves, vol. 1 (text), pp. i-xliv,
I-512.
WeEtmMorE, ALEXANDER.
1957. The birds of Isla Coiba, Panama. Smithsonian Misc. Coll., vol. 134,
No. 9, pp. 1-105, 4 pls., 15 figs., July 8.
ZIMMER, JOHN T.
1955. Studies of Peruvian birds. No. 66, The swallows (Hirundinidae).
Amer. Mus. Nov. No. 1723, pp. 1-35, Apr. 29.
ie
“ me
Mi
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE Iso NO 2) (Pia
1. Western end of Isla Escudo de Veraguas, from the south.
2. Southern shore of eastern end of Isla Escudo de Veraguas.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 3
PUR THER OBSERVATIONS ON
DISTRIBUTION, OF PATTERNS OF
COAGULATION: OF (‘THE -HEMORYMPH
INSINEOTROPIGAN INSECTS
By
CHARLES GREGOIRE
Department of Biochemistry, Institut Léon Fredericq
University of Liége, Belgium
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
AUGUST 18, 1959
THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.
FURTHER OBSERVATIONS ON DISTRIBUTION
OR PADIERINS OF COACUEATION OF TEE
HEMOLY MPH IN NEOTROPICAL
INSECES:
By CHARLES GREGOIRE
Department of Biochemistry, Institut Léon Fredericq
University of Liége, Belgiwm
The present paper is a contribution to a long-term inquiry on dis-
tribution of patterns of hemolymph coagulation in various arthropods,
especially in insects.
The reactions of the main elements involved in the process of co-
agulation of the hemolymph—a category of unstable hyaline hemo-
cytes (coagulocytes: Grégoire and Florkin, 1950) and the plasma—
differ in various insects. These differences, appreciated by phase-
contrast microscopy, have been classified into four patterns of micro-
scopic pictures (Grégoire, 1951).
The characters of these patterns may be described as follows:
Pattern I. Inception of the plasma coagulation in the shape of
islands of coagulation around the hyaline hemocytes.—Selective alter-
ations in the unstable hyaline hemocytes (shrinkages of the cell body
and occasionally of the nucleus, sudden expansions, bulging of blisters
and of blebs) result in exudation or in explosive discharge of cell
material into the surrounding fluid. Coagulation of the plasma starts
in the shape of circular islands of granular consistency around the
altered hyaline hemocytes. The islands of coagulation develop to a
certain size; then their increase stops. At the beginning of the
process, the islands are scattered and separated by fluid channels.
When the coagulation proceeds farther, the plasma in these channels
clots into a granular substance in which the islands preserve generally
their original size and shape.
The mechanism involved in pattern I is identical to one of the types
of coagulation described by Hardy (1892), Tait (1rg10, 1911), Tait
and Gunn (1918), Numanoi (1938), and Grégoire (1955b) in crus-
1 This is No. 9 in a series of papers entitled “Blood Coagulation in Arthro-
pods” published in various journals.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 3
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
tacean blood, in which a special category of cells, the Hardy’s ex-
plosive corpuscles, corresponding to the insect hyaline hemocytes or
coagulocytes, plays a selective part in the inception of the coagulation
of the plasma.
Pattern II, Extrusion of cytoplasmic expansions by hyaline hemo-
cytes, with development of cytoplasmic meshworks. Reaction in the
plasma in the shape of veils—On contacting the glass, a category of
fragile hyaline hemocytes undergoes alterations that differ from those
characterizing pattern I. These corpuscles extrude threadlike cyto-
plasmic expansions, sometimes of considerable length. These ex-
pansions are highly adhesive to solid particles (dust, chitinous
debris), other hemocytes, and physical interfaces (bubbles). These
alterations result in formation of cytoplasmic meshworks of various
complexity, on which the other kinds of hemocytes are passively
agglutinated.
The reaction in the plasma after these cellular changes occurs in
the shape of transparent, elastic, and contractile veils, developed within
the cytoplasmic systems built up by the hyaline hemocytes, or in their
vicinity.
In various insects the alterations in the unstable hemocytes are not
followed by changes in the plasma, and the modifications of the
hemolymph in vitro consist only of a cellular reaction.
Pattern III, Patterns I and II combined.—Association of the re-
actions taking place in patterns I and II characterizes the picture in
pattern III. In the same film of hemolymph, hyaline hemocytes send
out cytoplasmic expansions (pattern I1) while islands of coagulation
(pattern I) appear around the body of these corpuscles. When they
develop within the veils, which characterize the reaction in the plasma
in pattern II, the islands form circular, denser areas centered by the
altered unstable corpuscles.
Pattern IV. No modification in the hyaline hemocytes, or altera-
tions not followed by visible reaction in the plasma, in the optical con-
ditions of phase-contrast microscopy.—lIn the pictures of this pattern,
hemocytes resembling in their cytological characters the unstable
corpuscles involved in the other patterns do not visibly alter. They
appear as pale vesicles containing a few dark particles. In several
insects, these corpuscles are the remnants of darker refractile, hyaline,
frequently oenocytoid-like hemocytes, which undergo clarification
after explosive discharge of a part of their cytoplasm. In the vicinity
of these inert or altered hyaline hemocytes, no change can be detected
under the phase-contrast microscope in the consistency of the plasma.
NO. 3 HEMOLYMPH COAGULATION IN INSECTS 5
Specimens from more than 1,000 species of insects and of other
arthropods have already been tested about the pattern of coagulation
of their hemolymph or blood (Grégoire, 1951, 1953, 1955a, b, 1957,
unpublished observations on palearctic insects (1957-1958) ; Grégoire
and Jolivet, 1957). Predominance of one of the patterns has been
observed in several taxonomic groups. In other groups, owing to the
scarcity of the data available, or to large variations in the results, the
pattern representative of a species or of a group at a supraspecific
level could not be established.
The aim of the present study was to fill some gaps in the data. Four
hundred Neotropical insects, belonging to 215 species, including 185
species not yet investigated, were collected and studied during visits to
Tingo Maria, Peru (Estacion Experimental Agricola), August 1956,
and to the Smithsonian Institution’s tropical preserve on Barro Colo-
rado Island (Canal Zone Biological Area), October 1956.
MATERIAL AND METHODS
The samples of hemolymph were mostly thin films prepared as soon
as possible after capture. The hemolymph issuing from severed or
punctured appendages (antennae, legs, wings, joints of the wing-
cases) was placed immediately in contact with the edge of a cover
glass lying on a slide and was allowed to spread out into films.
A phase-contrast optical equipment WILD M/1o was used for the
observations (see Grégoire, 1955a, p. 105, and 1957, pp. I and 3).
RESULTS
DISTRIBUTION OF THE PATTERNS OF COAGULATION OF THE HEMO-
LYMPH IN INSECTS (TABLE I)
Detailed descriptions of the four patterns of coagulation of the in-
sect hemolymph, used in the present study, have been given elsewhere
(Grégoire, 1955a, p. 104; 1957, pp. 4-6 and text figs. I-4).
In the table, the names of the species are followed by the numbers
of specimens studied (adults, unless otherwise stated) and by the
patterns of coagulation provisionally found predominant or repre-
sentative on the basis of the study of several samples of hemolymph
obtained from these specimens. Incidental findings of other patterns
are reported under “Comments.”
In order to avoid duplication, the patterns recorded in the present
study in 50 insects belonging to Neotropical species previously in-
vestigated (Grégoire, 1957) are reported in the notes, preceded by
the date “(1957).”
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
The patterns of coagulation have been represented in the table by
the following symbols:
@: pattern I: inception of the plasma coagulation in the shape of
islands of coagulation around the unstable hyaline hemo-
cytes. Various degrees of extension of the process in the
films.
©: pattern II: development of cytoplasmic meshworks by hyaline
hemocytes. Reaction in the plasma in the shape of veils.
©: pattern II incomplete: emission of cytoplasmic expansions,
characterizing the reactions of the hyaline hemocytes in pat-
tern II, but unaccompanied by formation of veils in the
plasma.
@): pattern III: patterns I and II combined.
—: pattern IV: no visible coagulation by phase-contrast micros-
copy.
(): pattern incidentally or exceptionally recorded in limited fields
of preparations exhibiting predominantly another pattern.
(?): microscopical characters of a pattern not clear-cut or equivocal.
Artifacts possibly involved.
Other abbreviations used: sp., species; spm., specimen; T., speci-
men captured and studied at Tingo Maria; B., specimen captured and
studied on Barro Colorado Island.
Gradations in the intensity of the reactions, especially with regard
to pattern I, are indicated by the following symbols: I poor (scarce
fringes of clotted plasma around a limited number of altered fragile
hyaline hemocytes, without extension of the coagulation; I (scattered
islands of coagulation of various sizes, with moderate coagulation of
the fluid outside the islands) ; I*, I**, 1*** (islands around all the
hyaline hemocytes, substantial and general coagulation of the film).
NO. 3 HEMOLYMPH COAGULATION IN INSECTS 5
TABLE 1.—Patierns of coagulation
Patterns of
coagulation
representative
Number of or predominant
Material specimens in samples Comments
Orthopteroid Complex
DICTYOPTERA
BLATTODEA ? #
Periplaneta australasiae (Fabricius)
(adulbiand larval CDs) 04 Jk ee 2 @ >
Archimandrita tessellata Rehn (B.). 1 @
PHASMATIDAE *
Pseudophasma menius Westwood &
RCE EPUB ieee HOS Lael Wav Parr'ata aliens 6 I © 7
Prisopus cerosus Westwood (B.)... 1 eS *
Prisopus ariadne Hebard (B.)..... I ) uF
3 ulidet.isp. (2 adults, 1 larva) (T.). 3 @ G25) aes)
ORTHOPTERA
PENTIGONIIDAE +
Scudderia paronae (Griffini) (T.)... 1 ® 7
Eupeucestes crassifolius (Haan) 9%
ly eae cht accis sletera als Wiehe \annrargs 2 © res)
Undet. larva (Phaneropterinae)
DD aid aaa Ta ORD Va IU A I ®
Acanthodes aquilina (Linnaeus)
CIDE) Reret ere ett cman cease aa I 6 x
Micracentrum: sp. ©, (B. \lea/sccwe se 33 I @
Neoxtphidion conocephalus saltator
Goaussieey OUT) oe ie ealeitias I ge he
Moncheca pretiosa (Walker) (T.).. 1 @ ih
EUMASTACIDAE
PGrOniaSia ees NOD.) sate < qaceess des, s 3 ©
GRYLLIDAE*
Paragryllus temulentus Saussure
Cee A sare ieee Meh ION, So coe I @ a
GRYLLACRIDAE +
Abelona salvini (Saussure and Pictet)
eC ITA EPR Nice GRABS op au I @
1 Det. by Dr. C. Willemse.
2 (Grégoire, 1957) Epilampra azteca Saussure (B.): I ***,
L a ies Neoconocephalus affinis (P. de B.) 2 (B.): I (**); Caulopsis microprora He-
‘bar Pisce Ls
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
TABLE 1.—Patterns of coagulation—continued
Patterns of
coagulation
representative
Number of or predominant
Material specimens insamples Comments
ORTHOPTERA (continued)
FROSCOPILDAE
Apioscelis verrucosa Brunner Von
Weattenwyl) OCT) aie) I
ACRIDIDAE =+
Orphulella concinnula Walker (T.).
Tetrataenia surinama (Linnaeus )
OS it es) pert Seccmnete eee eres
Leptysma imsularis (Bruner) (T.)..
Wind etespyn( Al BM As cals atu gees mie eee a
Dicaearchus (gen. sp. nov.?) (T.)..
Kegua crenulata.Stoll (B.) 2.2...
DERMAPTERA ?
PENG ASD Cleat: )is sftusuaiye. iabemraldied: > 2
Hemipteroid Complex
HEMIPTERA
REDUVITDAE *6
Saica meridionalis Fracken and
PSIMOH (MESS ees ec ars Sims eiepeuete ee eee I -—
Stenopoda cinerea Laporte (B.)....
poor
eS
poor
poor
22k
2K
OK
Se eH HW
Ci")
(O poor
or @)
= (@ °)
ws
|
Rasahus sulcicollis (Serville) (B.)..
Zelusisp:? (nymph) (Ps). cb 2 3 ss
Zeus spat (nympi)e Cl Wisc daists sxe
Castolus subinermis (Stal) (B.)....
Moniuna lobaia'Stal (1.). 2% os 8b:
Montna fjumosa (Stal) (T.) 2.025.
Brontostoma notatum Stal (B.)....
Doldina bicarinata Stal (T.).......
PYRRHOCORIDAE ©
Largus balieatus Stal (T.).........
Dysdercus incertus Distant (T.).... 12 a (S)
Ss Se HN SS Se
=
4 (Grégoire, 1957) Copiocera specularis Gerstaecker: I; Osmilia flavolineata (de Geer,
(T.): I poor; Xyleus rosulentus Stal, 3 larvae (T.): I (**); Schistocerca paranensis Bur
meister CL poor:
5 Det. by Dr. me Catz!
6 (Grégoire, 1957) Saica apicalis Osborn and Drake (B.) : —; Zelurus spinidorsis (Gray,
(B.): — (II poor or incomplete) ; Panstrongylus rufotuberculatus (Champion) (B.): —
Panstrongylus geniculatus (Latreille), 3 spm. (B.): — (1
NO. 3 HEMOLYMPH COAGULATION IN INSECTS 7
TABLE 1.—Patterns of coagulation—continued
Patterns of
coagulation
representative
Number of or predominant
Material specimens in samples Comments
HEMIPTERA (continued)
PYRRHOCORIDAE (continued)
Dysdercus ruficeps (Perty) (T.)... 1 —_ (Se)
Dysdercus sp.? (nymph) (T.)..... I --
COREIDAE®
thine. decorata Stal (T.) ..... 432+ I -—
Spartocera fusca (Thunberg) (T.). 1 —~
Plapigus foltaceatus (Blanchard)
CODY C Clee ciel. dey wean ere 2 _—
Anasa haglunds Stal (T.).......... I —
Hypselonotus striatulus (Fabricius)
LEA er SYS wie eae atahe dig wate ts I —
Paryphes adelphus mutans Horvath
CTE) Sore Nea ey, iets wo aiken. S I —
Hyalymenus tarsatus (Fabricius)
CER Riawoate woke aka sei BAe ied aigok i —
Leptocorisa filiformis (Fabricius)
(1S Ue URS AR a to ec NE PU ee PaO 2 —
Zoreva dentipes (Fabricius) (T.)... 5 - Co)
Zoreva spintfera Stal. (T,). 3.2.5 5. 2 -——
GELASTOCORIDAE®
Nerthra peruviana (Montandon)
CUE Nl Sissel aes iatal ss Save. oscar tiace.s ahs I -- CO)
PENTATOMIDAE™
Symphylus deplanatus (Herrich-
SACRO ails tlaiaes\'a cm aiatn migra I —
Augocoris gomesti Burmeister (T.). I —
Macropygium reticulare (Fabricius)
Meveyaralaiteh ser lcitelealraileioest arse) stele) s 2 ==
Euschastus crenator (Fabricius)
Reyes Ohne Miser ah Bem Le Ts ie) ica 950 2 —_
Euschistus sp.? (nymph) (T.)..... I — ( ?)
Loxa picticornis Horvath (B.)..... I -~
ETEORIEEUS SPI Ess) secu ci u 2 aye erynre = I —_
Edessa .afiuis, Wallas (T.) .. 3.5.1 2 _-
Edessa polymita Distant (B.)...... I _
Ba cssa.spe SEU Ce CLs aces aes es I -~
7 (Grégoire, 1957) Mecistorhinus piceus (Palisot de Beauvois) (T., B.), 2 spm.: —;
tdessa rufomarginata De Geer, 4 spm. (B.): —; Acrosternum scutellatum Distant (T.): —;
Veodine macraspis (Perty), (B.): —.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
TABLE 1.—Patterns of coagulation—continued
Patterns of
coagulation
representative
Number of or predominant
Material specimens in samples Comments
HEMIPTERA (continued)
PENTATOMIDAE (continued)
dessa Spe (Ba) its iniete lea bie erie oe I a
Fdessaispa: (oyimph) (Ls)... .8-o-. I —
Bidessn spire Gaympl) (Cy cine I —
MIRIDAE *
Mimoncopeltus, n. sp. (T.).......- I —
HOMOPTERA
CICADIDATE =
Carineta sp., near boliviana Distant
A Gea DN oescreee aps ak ciaiiaNonaleiare Fe ete I @
FULGORIDAE*»??
Copidocephala ornanda (Distant)
CBG yen des ciste se alse mies I & te
Odontoptera sp. (CB iis. seein. ss sae: I @ ses:
Diareusa annularis imitatrix (Ossia-
UNitTSamy (GBS) frecnceancieyci ctsiaiecee sci I 2) a
Gen. and sp. unknown (B.)....... i & +
CL DAE ®
Gen. and sp. unknown (B.) 0.0.05 i — (?)
DICTVOPEAARIDAE *
Nera florens Stal (Bi)... oc)s66 << a 2) a
Taosa herbida (Walker) (B.)...... I @ .
Gen. and sp. unknown (B.)........ I Cs] ree
MEMBRACIDAE &
Star Oleows spy DiC Tha) jee ase weet = oats I @ o
CERCOPIDAE®
Cephisus siccifolius Walker 9 (B.).. 1 @ oo
AMOS Oops Tors (s)he eta ia elas 2 @ +
Ai Gusta Eo: Ot Al Lin) aia ai, scsisin aie 2 © sok
Homaspis sp. ae tis (1). .cieae ie eae I @ (@ ?)
AROMAS PIS iS: tae 2c (W.)) sais see ss ahs I @ RK ( @) FRX)
MOMS PIS ISD... 52 3) Pulled). «greta Oss 014 I @ Kei
MOmaspis SD: EAs (UA) cies alee pier I @ he
Momaspisison sess hE). ee eee I @ +F
MOMS PIS Sv aE Geils e)) seem eee I @ ce
8 Det. by Miss Louise M. Russell.
9 Det. by Dr. D. A. Young; Diareusa by Dr. V. Lallemand.
10 (Grégoire, 1957.) Nae elegans (Olivier), 2 spm. (B.): I ***; Cathedra serrate
(Fabricius) (B.):
NO. 3 HEMOLYMPH COAGULATION IN INSECTS 9
TABLE 1.—Paiterns of coagulation—continued
Patterns of
coagulation
representative
Number of or predominant
Material specimens in samples Comments
TOMOPTERA (continued)
CERCOPIDAE (continued)
Monmaspes Span 7 96 Clo) wns. 3. 3s 2 @ ***(@))
MOMISPIS Sp: SEO oy CLs) 8. vad s oes I @) aa
omasyis sp. 220 6 CB:). 2.256222. I ~~ (@ ?)
CICADELLIDAE?
Tettigellinae
Diestostemma nigropunctata (Signo-
BEE Me elise ticks eka ollclfolsioi'e «ats olde wose I @ **(@)
Deestostemma:sp, -O CL...) sai. sie ys ss I 8 4k
Baleja flavoguttata (Latreille) (B.) 1 ® re
Spatmknow On) sepa esate ha anf, I @
Oncometopia sp. +: I, sex anomaly
Ba Alsias Aree lic Ph) es Gallia stare ads I @ poor (@))
Oncometopia sp. #£2, normal? (T.). 1 —
SpyeuniknOwan CB.) jars /-! .ieiatesslecs s 08s 6 @ 7 LO
Wes MAAR E A Wee Mate fel cire 8/5. oy ol) dats) e\aseven we) s
“Gypona”’ decorata Fowler (B.).... 2 @ poor
Gypona atitlana Fowler (B.)....... I — (?);@
probable
Gypona hebes Fowler (B.)........ 3 © poor to **;
(@);— in
I spm.
ALGIERS IPM (1 a)ip Belarc 12: je) 536, 2%! sh. 0h ove 3 I ~ (?); dry spm.
PORENESD Ua) e050 RGN, Licata 3 — (@ °)
in I spm.
Gyponana sp: Ga B2)\. 3. Assen eain « I @ probable.
Negostana’ sp. sete (BSS o2bod cc I ©
Wegostana' sp. 422 9 (Biche. . fei 2 —(@ ?) @ probable
PEATIDAE =
Anormelis nigrolimbata (Fowler)
CEP Tae aris Otis sixes eiaieie shea 8 — (@ poor)
Flatormems sp. (2) (B.).......... 2 — (?)
Paradascalia metvi (Distant) (B.).. 6 — (@ ?)
ES SIDA?
Oroneqia isos (CB)... bith gacretes pe I _
11 (Grégoire, 1957) Carthaeomorpha rufipes Melichar, 3 spm. (B.): I **; (—?) in 1 spm.
IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
TasLe 1.—Patterns of coagulation—continued
Patterns of
coagulation
representative
Number of or predominant
Material specimens insamples Comments
COLEOPTERA
ADEPHAGA
CARABIDAE *
Hurpalinae sp.et (B.) 6s. se. ees I sas
Fharpalinae sp. 32°(B.)\.. os os ese ss I —
Harpalinae sp. 3£3.(B.) 656.2 ee I G
WARE Spt e GE) oe sin oe erecta sek ao I O poor (—)
Migr arspume 2 (B.) ssi is aislds selene I () (e) probable
PACD IM IST Ala) id a's) 8 wleday a nie ane atare = Ve I —
POLY PHAGA
PASSALIDAE 1224
Passalus (Neleus) interstitialis
ESC lISChIc Es) verve etanelel aeaiiaye'= le inp I o—
Vetwrins Sp. (QB. )\.'. derais/saeiciels «5 ais I — (@)
SCARABAEIDAE *
Coprinae (Scarabaeinae)**
Cama sp. \(W.)ia's siclc lal lerals erereie'es I @
Uroxys gorgon Arrow (B.)......-. 2 = (a)
Rutelinae **
Mesomerodon spinipenne Ohaus
Ca pes oe eoceetine olay sls aoe ayia tava 2 ©) *
Pelidnota chlorana Erichson (T.)... = 2 @)
Anomala virescens Burmeister (T.). 2 (B) (Ss)
Pigonala SSP. TCL) sacle ee ole eel ake ares 3 @)
Dynastinae ** 3°
Gen? near Bothynus: CU.)i.d..< atarereawe I O
CEBRIONIDAE **
Zeta TKO WIISI( LE <))\s ceo she olaheyesedei= I 8 *
ELATERIDAE
Chalepleprdius ep iC Bi) so exe ose ayoyspues I © (@)
SCMUIOTES ‘SP. CT: Dis aisrsse lets ciordieieqinteg I ©
LY CIDAE *%
ME NILEST SDM Wea) ale pnts id tatiess te Allee aes tere I z (Os)
LAMPYRIDAE
POHAUS |S. MLS) os sats oie wale 0 spe me 2 ?
12 Det. by G. Fagel.
13 Det. by O. L. Cartwright.
14 (Grégoire, 1957) Veturius platyrhinus Westwood (B.): — (III?).
15 (Grégoire, 1957) Aspidolea singularis Bates (B.): II.
16 Det. by T. J. Spilman.
17 Det. by Dr. Ch. Jeuniaux.
NO. 3 HEMOLYMPH COAGULATION IN INSECTS LE
TaBLe 1.—Patterns of coagulation—continued
Patterns of
coagulation
representative
Number of or predominant
Material specimens in samples Comments
POLY PHAGA (continued)
VIVE GY LIDAE A°
Mekiomma spr CB: )). seks e6 6s «stn I @ or @
ENDOMYCHIDAE”
Probably Amphiz sp. (T.)......... I Oo
COCCINELLIDAE 7
Bi palachaa: Sp (ls) (cies wher ale -nai'e!2ys 32) I —
Monomeda marginata (Linnaeus)
GUEE )Raiacve toy srcvenetavace siciettle pis.«: ates, ste I —
PROLEYVLIDAE *°
Erotylus, prob. spectrum Thomson
CULE Rg ciate ses eiaiate' da erchare a el asen ale I @ (@ ?)
Prob. Homoeotelus sp. (T.)......-. I 2 *, (@) probable
Gent unknown (CEs) a. 6. beets cases I Co
Gene naknowa (Dia si< 4. 5.sr0he.0'6 510: I oO
TENEBRIONIDAE *
Strongylium auratum Laporte (T).. 1 © probable
MELOIDAE
CERAMBYCIDAE*
Prioninae
Sienadontes Sp. (E.) as vias.i3 alg srenis I @ ae
ymad es sp, (Wa) whe daceie lence oye wiera o's I & *KE(@)
Lamiinae
Pe swt POTS. (1D. Viajes :s cae «ss ones I 8 (@ ?)
TOMES! MUD )iicia ain s).2 si l'aide a oe) I @ poor
Oreodera glauca (Linnaeus) (B.).. 1 @ ¥r
Acanthoderes bivitta White (B.)... 1 @) **(@).
ihogocherus sp. S21) (Be) ce vcieien'« I @ ge
MaGgOCherus Spi:FE2 (OB. ) p)ssieicert -s,« I @ kK
Golabotea spe (CEs) a5 sess doce. a 506s I gS a
Charoules spit (Es) 2385 ssie cases I e@© tok
yaroudes Sp. se 2 CN \leineictiae oe eae I @ ee
18 Det. by Dr. J. G. Rozen,
19 Det. by Dr. J. G. Rozen.
20 Det. by Dr. E. A. Chapin.
21 Det. by T. J. Spilman. (Grégoire, 1957) Zophobas prob. atratus (Fabricius) (B.):
III ** probable.
22 Det. by T. J. Spilman. (Grégoire, 1957) Epicauta grammica (Fischer von Waldheim),
Secum. (8.): 1 *** (IIT).
23 Det. by George B. Vogt. (Grégoire, 1957) Taeniotes scalaris (Fabricius) (B.): I (III).
I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
TABLE 1.—Patterns of coagulation—continued
Patterns of
coagulation
representative
Number of or predominant
Material specimens in samples Comments
POLY PHAGA (continued)
CHRYSOMELIDAE br. sense *4
EUMOLPIDAE: poss. near Priono-
LOPE Ses (Ho) teen gyno vie wien eMetseke I -—
Chrysomelinae
Woryenona sp. is) 2 dessa eid eo I — (a)
Seal odesi( ey Spee ile) cca ee erect I (@)
Gosmogramma isp) \(T.)) ss. hae ee ee I CS?) (—)
Galerucinae
Drab rotica spe Kis) 5 aioe) ve ielaele ee I ao
VAM ME GOON SIO ily) cet ale ove ee Gleate eee I ——
Alticinae
Gedionychus sp. 420 (0). oo ess 2 _—
Oedionychus sp.) Fe2(T))). ui a'ss's I —
Hispinae
Oediopalpis guerini Baly (B.)...... 2 —
Cassidinae
Cyclosoma tristis Boheman (T.).... I o—
Echoma sp., prob. aulica Boheman
CHa vale ele a coletave eiaue itebeeutio rs I 2 poor
CURCULIONIDAE
Wig Pactusisp! sek (ler ie ve es 4 —
Naupactus sp. ge 2: (Us) cn. so 56s I —
COM PSUSNSP hea) eve ee eee ee 3 —
Pett Pits Spy. eT 1B. )ieus ain eu aco /ssaere I —
Hiciipus sp. see (UBS) eo cue es te 2 —
MiCiOINGSIUS SPig( D\. \is ac sieves Gets oe I —
Panorpoid Complex
NEUROPTERA-PLANNIPENNIA
MANTIS PID A Ey 26, 24
Climaciella semihyalina (Serville)
GEER Che oie! aaivucie wiateueeatianets I = (@ ?)
24 Det. by George B. Vogt.
25 Det. by Miss Rose Ella Warner. (Grégoire, 1957) E-xophthalmus jekelianus (White),
2 spm. (T., B.): —.
26 Det. by Miss Sophy Parfin.
27 (Grégoire, 1957) Mantispa phthisica Gerstaecker (B.): —.
NO. 3 HEMOLYMPH COAGULATION IN INSECTS 13
TaBLe 1.—Patterns of coagulation—continued
Patterns of
coagulation
representative
Number of or predominant
Material
YEUROPTERA-SIALODEA *8
CORY DALIDAE
Corydalus sp., near armatus Hagen
CM oe Ae a cae
FRICHOPTERA
BY DROPSY CHIDAE **
Prob. Leptonemaisp: ¢(B.)s 2.8:
EE PIDOPTERA
VAD AE sp, Cadult 2? (Be...
SAUNT AE sp. (atva) Ci... «
MCI DAE sp. Carva)! (1.) ... -..-
JIPTERA
LARVAEV ORIDAE *°
Ormiophasia bushkiu TNS.
TY MENOPTERA
ICHNEUMONIDAE *
WNetela spe OB sob ooo eet
FORMICIDAE
Aelecaspyae i) 9) (Bi). 58s sevice fone
Asteca spy se2 0 (Ba )isi iets tates seats:
Pachycondyla crassinoda (Latreille)
SCL SR uiccics, nach diutegs (lope ayn ete
Dinoponera sp. (worker) (T.).....
Labidus coecus (Latreille) ¢ (B.)..
VESPIDALE 25:2
Poehymnenes, SpoCl..)'. vases os ees
Polistes major weyraucht Bequaert
(aN ea creer tera. MEM lui da «
POMMPILIDAE *2°
28 Det. by Dr. A. B. Gurney.
29 Det. by W. D. Field.
30 Det. by C. W. Sabrosky.
31 Det. by Miss Luella M. Walkley.
in samples
O1O=|
~@@ | |
@ Oo
Comments
KK
very poor (—)
*
poor or @
possibly @
#4 (©)
possibly ©
82 Det. by Dr. M. R. Smith. (Grégoire, 1957) Paraponera clavata (Fabricius) ¢ (B.):
[ **; Camponotus sericeiventris Guérin, br. sense, 4 workers (B.): possibly III.
33 Det. by K. V. Krombein.
34 (Grégoire, 1957) Polistes canadensis panamensis Holmgren, 4 spm. (B.): I.
35 (Grégoire, 1957) Anoplius a-amethystinus (Fabricius) (B.): III.
14 SMITHSONIAN MISCELLANEOUS COLLECTIONS
TABLE 1.—Patterns of coagulation—concluded
Patterns of
coagulation
representative
Number of or predominant
Material specimens in samples
HYMENOPTERA (continued)
SEH ECIDAE §8
Sceliphron fistulare (Dahlbom) (B.). 1 B
Stictia maculata (Fabricius) (B.).. 1 @
ODONATA
AGRIONIDAE *
Megaloprepus coeruleatus (Drury)
EA eisai ans tienatal a tote eadatataubesitars I --
ARACHNIDA *
Araneae
THERAPHOSIDAE
Eury (Brachypelma) sp. (B.)...... I _
THOMISIDAE
Epicadus heterogaster (Guérin)
Cees Ai Dieta dice anita abc.e Baier meletetalt I —_
OPRTLIONES *
COSMMETID ES | Se 95 2 eho sslinss. sft sekanelererers I —
PEDIPALPIDA **
Tarantula palmata barbadensis Po-
COGIG MOIS! J Wie ulate slam isuttel pave eo oye I —
TAO DEDAE =,
Amblyomma humerale Koch § (B.). I ——
86 Det. by Dr. A. B. Gurney.
37 Det. by Dr. J. Cooreman.
MICROSCOPY
VOL. 139
Comments
2K
The microscopical features of the reactions which characterize the
coagulation of the hemolymph in several supraspecific groups of
insects (Orthopteroid Complex, Heteroptera, Homoptera, Scara-
baeidae, Cerambycidae, Hymenoptera, Lepidoptera) have been de-
scribed elsewhere (Grégoire, 1955a, pp. 109, III, 115, 118, 123 ; 1957,
pp. 7, 27, 28; Grégoire and Jolivet, 1957, pp. 28-33). They were also
observed in the corresponding groups of the present material. A few
particular reactions will be briefly mentioned below.
Phasmoptera.—As repeatedly pointed out (Grégoire, 1951, 1955a,
1957; Grégoire and Jolivet, 1957) the various categories of hemocytes
are passively embedded in the coagulum initiated by the alterations
NO. 3 HEMOLYMPIL COAGULATION IN INSECTS 15
in the fragile hyaline hemocytes or coagulocytes. Modifications of the
plasma induced around the former corpuscles are exceptional. Such
modifications, recorded previously in two specimens of Neotropical
stick insects (Grégoire, 1957, p. 7), were observed in Prisopus cerosus
(table 1) around macronucleocytes of small size (stem cells), sec-
ondarily to the typical formation of islands of coagulation around
the unstable hyaline hemocytes.
Heteroptera——Granular precipitates, unrelated to the presence of
hemocytes in the vicinity, recorded previously in the same group of
insects, were observed in the present material in Montina lobata,
Saica apicalis (Reduviidae), Macropygium reticulare, 3 species of
Edessa (Pentatomidae), Anasa haglundi, Zoreva dentipes (Coreidae).
A tentative interpretation of these occasional findings has been given
elsewhere (Grégoire, 1957, p. 7).
Coleoptera—The sequence in the alterations in the fragile hemo-
cytes and in the plasma, characterizing pattern III (see Grégoire, 1957,
p. 2 and text fig. 3), appeared with great clarity in the two specimens
of Elateridae mentioned in table I.
In the samples of hemolymph from Compsus sp., Heilipus sp.,
Exophthalmus jekelianus (Curculionidae), characterized, as shown in
the table, by the absence of detectable alteration in the plasma, in the
conditions of phase-contrast microscopy, a category of highly labile
hemocytes, unrelated to the unstable hyaline hemocytes, underwent
considerable modifications in their shape: immediately upon with-
drawal and spreading out into films of the hemolymph, these hemo-
cytes appeared spindle-shaped, with two straight expansions on both
sides of the cell body. The expansions became progressively flexuous
and exhibited continuous trepidations and jerks. They reached great
lengths, bent suddenly at right angles, and sent out lateral ramifica-
tions in various directions. Simultaneous development of such changes
in neighboring hemocytes resulted in constitution of loose meshworks
in wide areas of the preparations. Similar labile hemocytes have been
reported in African weevils (Grégoire and Jolivet, 1957, p. 32) and
in Diptera by Grégoire (1955a) and Jones (1956). In the present
material they appeared in Ormuophasia bushkui (Diptera).
Much smaller bipolar corpuscles, of unknown origin, unrelated to
the labile elements described above, developed similar modifications.
A detailed study of these corpuscles will be reported later.
Arachnida. Araneae—In Epicadus and in Eurypelma, a category
of hemocytes with coarse refractile granules scattered in their cyto-
plasm and highly sensitive to foreign surfaces underwent disintegra-
tion immediately upon shedding of the blood, in contrast to other
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
categories of more resistant blood cells, such as macronucleocytes of
small size (stem cells) and other kinds of granular hemocytes. A
similar “differential sensitiveness’” has been formerly observed in
extensive material of spiders (see Grégoire, 1955b).
DISCUSSION
DISTRIBUTION OF THE PATTERNS OF COAGULATION IN THE VARIOUS
TAXONOMIC CATEGORIES OF INSECTS
Detailed accounts on the relationships between pattern of coagula-
tion of the hemolymph and taxonomic category have been given in
previous papers (Grégoire 1955a, pp. 132-137; 1957, pp. 28-32;
Grégoire and Jolivet, 1957, pp. 34-37). In this respect, the informa-
tion obtained in the present material supports our former conclusions.
With one exception (Carthaeomorpha rufipes, see below), the pattern
detected in the samples of hemolymph collected in the present study
(table, notes) from 50 specimens belonging to 30 neotropical species
already investigated (1957), were identical to those recorded pre-
viously.
1. Orthopteroid Complex.
That broad group constitutes a highly homogeneous category with
regard to the pattern consistently recorded at the specific and at the
supraspecific levels.
2. Hemipteroid Complex.
Hemiptera.—With the exception of Nepidae and Belostomatidae,
studied previously (Grégoire, 1955a; Grégoire and Jolivet, 1957),
all the specimens from 14 other families of Hemiptera investigated,
including Reduviidae, Pyrrhocoridae (see 1955a), Coreidae, Gelasto-
coridae, Pentatomidae, Miridae of the present (38 species) and of
former materials, exhibited consistently the pattern IV.
Homoptera.—The present material includes 41 species not investi-
gated previously (Grégoire, 1955a, p. 110; 1957, pp. 15 and 16).
Pattern I was predominant in Cicadidae, Fulgoridae, Dictyopharidae,
Cercopidae, Cicadellidae, and was recorded in the only specimen of
Membracidae captured, a family not yet investigated. In a few
Cercopidae (see also 1955a, p. 110) and Cicadellidae, pattern I was
associated with pattern II (= pattern III).
A substantial coagulation of the hemolymph, developing rapidly,
sometimes instantaneously, characterized these families, with the
NO. 3 HEMOLYMPH COAGULATION IN INSECTS L7
exception of Cicadellidae, and was especially conspicuous in
Fulgoridae.
In Cicadellidae, the amount of clotted material varied greatly and
appeared scarcer than in the other groups listed above.
Pattern IV was observed in the samples of Cixiidae, Flatidae, and
Issidae. However, in Flatidae, pattern I was found in Carthaeomorpha
rufipes (table, note 11), a species in which pattern IV had been
recorded previously in the only specimen available (Grégoire, 1957,
p. 16). Pattern I appeared also incidentally in Anormelis mgrolim-
bata and in Paradascalia nietvi. Pattern IV, observed to occur pre-
dominantly in the few samples examined till now, is then questionable
as being representative of Flatidae, a family which requires further
investigation.
3. Coleoptera.
The patterns predominant or representative in several groups
formerly investigated were seen again in the present material: pat-
tern II in Scarabaeidae (Rutelinae, Dynastinae), pattern III in
Elateridae and in Tenebrionidae, pattern I in Meloidae (note 22),
Cerambycidae (very substantial coagulation), pattern IV in Cur-
culionidae.
Pattern I, alone or associated with pattern II (= pattern III) was
recorded in specimens of Cebrionidae and of Lymexylidae, two fami-
lies not represented in our former data.
In the other groups listed in the table, scarcity in the material, large
variations at the individual, specific, and generic levels, already noticed
previously, do not permit conclusions about the pattern predominant
or representative of these groups.
In this and in former studies (Grégoire, 1957, p. 22; Grégoire and
Jolivet, 1957, pp. 22 and 23), absence or scarcity in clotting substances
was observed in several specimens of Eumolpidae and of Cassidinae.
In the present material, pattern III was recorded in one (Stilodes)
out of 3 specimens of Chrysomelidae s.s., a family involving genera
with obviously predominant patterns (see 1955, p. 114: Chrysolina,
7 species: patterns I and III; Timarcha, 5 species: patterns I and
TU),
4. Panorpoid Complex.
The present results are in agreement with former data with regard
to Mantispidae (pattern IV: see Grégoire, 1957, p. 23), Sialodea:
Corydalus sp. (pattern I, instantaneous reaction: see 1955a, p. I15:
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Sialis flavilatera L.) ; Trichoptera: Leptonema (pattern I: see 1955a,
p. 116: Limnophilidae sp. and Anabolia nervosa Leach) ; larvae of
Lepidoptera (pattern IT, see 1955a, pp. 116-118; 1957, p. 23; Grégoire
and Jolivet, 1957, p. 25), and adult Diptera (Ormiophasia bushku:
pattern IV, see Grégoire, 1955a, p. 121).
As already pointed out, pattern I frequently characterizes insects
belonging to relatively archaic orders (Plecoptera, see 1955a, p. 107;
Megaloptera, 1955a, p. 115).
5. Hymenoptera.
Patterns I and III are representative in several families of this
order (Grégoire, 1955, pp. 122-123; 1957, pp. 24-26; Grégoire and
Jolivet, 1957, p. 25). However, individual and specific variations may
mask the representative pattern of the genus or of the family when
only limited material is available.
In the present (note 32) and previous materials (1957, p. 24) of
Formicidae, a substantial pattern I characterizes the genus Paraponera.
Patterns I and III were also recorded, though not consistently, in
several specimens of the genus Camponotus (1955a, p. 123; 1957,
p. 24; Grégoire and Jolivet, 1957, p. 25).
On the other hand, no coagulation could be observed (pattern IV)
in seven females of Azteca sp., from which the films of hemolymph
were collected and prepared without interference of any artifact.
The present observations on Vespidae (note 34), Pompilidae
(note 35) and Sphecidae are in agreement with those made previously
(pattern I and/or III: 1955a, p. 123; 1957, pp. 25-26).
6. Odonata.
As in former studies (1955a, p. 107; 1957, p. 26), pattern IV was
recorded in the only (adult) specimen of this order collected in the
present material.
7. Arachnida.
Coagulation of the blood was not detected in the present and
former specimens of Pedipalpa, Ixodidae (1955b, pp. 497-498).
Pattern IV was also recorded, in this and in previous studies, in
specimens of Opiliones and of Brachypelma (Theraphosidae,
Araneae), while other specimens of the latter genus exhibited pat-
tern II, sometimes substantial, sometimes incomplete (see 1955),
p. 495).
NO. 3 HEMOLYMPH COAGULATION IN INSECTS 19
ON THE DISPARITIES IN THE REACTIONS OF COAGULATION OF THE
HEMOLYMPH RECORDED AT THE SUPRASPECIFIC, SPECIFIC,
AND INDIVIDUAL LEVELS
1. In contrast to the taxonomic categories characterized by a pat-
tern of coagulation representative or predominant, other groups,
especially Carabidae (Grégoire, 1955a, p. 111; 1957, p. 16; Grégoire
and Jolivet, 1957, p. 12), exhibit such variations that, in spite of
increased samplings, a representative pattern did not appear clearly
in these groups at the family level, but provisionally at the generic
or specific levels.
In that respect, incidental coincidences may be deceptive and sug-
gest erroneously that a pattern is characteristic of a genus, when it
may actually represent an incidental failure of the true pattern to
appear with all its particularities in a set of specimens being pro-
visionally, at the time of capture, in similar abnormal conditions. For
instance, in three specimens belonging to three different species of the
genus Agra (Carabidae), pattern II, incomplete in two of these speci-
mens, was predominantly observed in the present study, while
formerly, in three other species of the same genus, pattern I had
been consistently found (Grégoire, 1957, p. 16). Pattern III, possibly
dissociated in the individual samplings into its two components
(patterns I and II), might be the representative pattern of the genus
Agra. Other examples are furnished in Hymenoptera in the genera
Eciton (1957, p. 24) and Azteca (table), in which the predominant
patterns are possibly not the actual ones.
In families such as Lycidae, Lampyridae, Coccinellidae, Chryso-
melidae (Cosmogramma), and Cassidinae (Cyclostoma), the observa-
tions were handicapped by the presence in the hemolymph of particles
floating in considerable numbers, a finding already noticed (1955a,
p. 106; 1957, p. 27; Grégoire and Jolivet, 1957, p. 30).
2. Divergences at the specific or individual level recorded in genera
characterized by a pattern predominant or representative, appear, for
instance, in specimens of Cicadellidae. However, the pattern char-
acterizing the group was found incidentally in the samples (see under
comments in the table).
At the individual level, pictures of another pattern were recorded
incidentally in limited fields of preparations exhibiting a predominant
pattern (Reduviidae: Stenopoda, Rasahus, Dysdercus; see also 1955a,
p. 109; 1957, p. 13; Grégoire and Jolivet, 1957, pp. 10-11).
Tentative interpretation of these divergences have been presented
elsewhere (1955a, pp. III, 124, 126; 1957, discussion; Grégoire and
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Jolivet, pp. 36 and 37). Artifacts of preparation are responsible for
a part of the pictures recorded. Nutritional balance of the specimens
at the time of capture, seasonal and pathological conditions, able to
alter the sensitivity of the unstable hemocytes or the amounts of the
coagulable substances in the hemolymph, are among the factors which
might explain these discrepancies: change in the pattern of coagula-
tion has been observed in infected insects belonging to species or to
groups characterized in their normal conditions by another pattern
(Acrididae, Dermaptera, Cerambycidae) (see Grégoire and Jolivet,
1957, p. 36). Similarly, in a specimen of Gypona hebes from the pres-
ent material, exhibiting pattern IV (table, comments), the unstable
hemocytes responsible for the inception of the coagulation contained
unusual coarse granules, absent in the other normal specimens in
which the pattern representative of the group was observed.
The present results support former conclusions (1957, p. 30) that
the patterns of coagulation are not individual particularities, but
rather characterize species, more frequently supraspecific categories.
DIVERGENCES BETWEEN NEOTROPICAL MATERIAL AND INSECTS FROM
THE OLD WORLD
In 10 specimens belonging to 6 species of Neotropical Passalidae
(1957, p. 18, and here, table 1), pattern I was recorded exceptionally
in one sample from a single species, while this pattern, unmixed or
associated with pattern II (= pattern III), appeared in the 5 African
species (25 specimens) available (Grégoire and Jolivet, 1957).
Pattern I, absent from the samples of Neotropical Coprinae (4 spe-
cies, 8 specimens), was found, alone or associated with pattern II
= pattern III), in 12 (29 specimens) out of 17 African species
examined (Grégoire and Jolivet, 1957), and was questionable in three
other species (5 specimens).
These data might suggest the possibility of discrepancies, with
regard to these two families, between Neotropical and Old World
material. However, as already pointed out (Grégoire, 1957, p. 32),
large individual variations characterize these families, especially
Passalidae. Numerous samplings from insects of both origines, and
belonging to genera and species more closely related than those
available, are required before any conclusion might be drawn about
the existence of such discrepancies.
SUMMARY
1. Coagulation of the hemolymph from 400 (mostly adult) speci-
mens, belonging to 215 Neotropical species of insects, and including
NO. 3 HEMOLYMPH COAGULATION IN INSECTS 21
185 species not yet investigated, has been observed on films in vitro
by phase-contrast microscopy. In that material, the pattern of coagu-
lation predominant in the samples or representative for the species
or for the supraspecific taxonomic category has been recorded.
2. The material contained insects from 14 families poorly (Dictyo-
pharidae, Cercopidae, Cicadellidae, Flatidae) or not (Gelastocoridae,
Membracidae, Cixiidae, Issidae, Cebrionidae, Lymexylidae, Erotyli-
dae, Hispidae, Corydalidae and Larvaevoridae) represented in pre-
vious studies.
3. Additional information obtained for the present paper was con-
sistent with former data, with regard to the pattern predominant or
representative, in the Orthopteroid Complex, in several families of
Heteroptera (Reduviidae, Pyrrhocoridae, Coreidae, Pentatomidae,
Miridae), of Homoptera (Cicadidae, Fulgoridae, Dictyopharidae,
Cercopidae, Cicadellidae), of Coleoptera (Scarabaeidae, Elateridae,
Tenebrionidae, Meloidae, Cerambycidae, Curculionidae), of Hymen-
optera (Formicidae, Vespidae, Sphecidae).
4. In the families not represented in former investigations, pat-
tern I was recorded in specimens of Cebrionidae and of Lymexylidae
(Coleoptera).
5. Pattern I was also observed in specimens of Corydalidae (Sialo-
dea) and of Hydropsychidae (Trichoptera), in agreement with pre-
vious results on palearctic representatives belonging to these groups.
6. Divergences in the reactions of coagulation observed in the
present and in a former study between Neotropical and African
Passalidae and Copridae (Coleoptera) require further investigations
on more extensive material, owing to the large variations existing in
these groups of insects.
7. The reactions of the blood in vitro observed in five specimens of
Arachnida (Araneae, Ixodidae, Opiliones, Pedipalpa) are briefly men-
tioned in relation to previous results on more extensive material.
ACKNOWLEDGMENTS
I am grateful to Dr. Carl B. Koford, former resident naturalist of
the Canal Zone Biological Area, for invaluable help in supplying me
rapidly with adequate equipment. I also wish to express my thanks
to Mrs. A. Gomez, administrative assistant at the station, who, as
usual, was very cooperative in arranging for living accommodations
in the Canal Zone.
I am greatly indebted to Dr. Ing® Oswaldo Vargas Gonzales,
head of the Seccién Entomologia de la Estacion Experimental Agri-
cola en Tingo Maria (Peru), for authorization to carry on my work
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
in his department. I acknowledge with sincere appreciation the assist-
ance of Dr. Ing® Matias Reyes Valdivieso for rapid and clever
adjustment of my optical equipment to one of the microscopes of
the department.
I wish to thank Dr. Remington Kellogg, Director of the United
States National Museum; Dr. Waldo L. Schmitt, former head curator
of the department of zoology, for authorization to have the material
determined in the Museum; and Dr. J. F. Gates Clarke, curator of
the division of insects, for collecting and mailing the data. I also
wish to express my gratitude to the following specialists for identifica-
tion of the specimens: O. L. Cartwright, Dr. E. A. Chapin, Dr. J.
Cooreman, G. Fagel, W. D. Field, Dr. A. B. Gurney, Dr. Ch. Jeuni-
aux, K. V. Krombein, Dr. V. Lallemand, Dr. J. C. Lutz, Miss S. Par-
fin, Dr. J. G. Rozen, Miss Louise M. Russell, C. W. Sabrosky,
Dr. M. R. Smith, T. J. Spilman, George B. Vogt, Miss Luella L.
Walkley, Miss Rose Ella Warner, Dr. C. Willemse, Dr. D. A. Young.
REFERENCES
Grécorre, CH.
1951. Blood coagulation in arthropods. II. Phase-contrast microscopic
observations on hemolymph coagulation in sixty-one species of
insects. Blood, vol. 6, pp. 1173-11098.
1953. Blood coagulation in arthropods. III. Reactions of insect hemolymph
to coagulation inhibitors of vertebrate blood. Biol. Bull., vol. 104,
PP. 372-393.
1955a. Blood coagulation in arthropods. V. Studies on hemolymph coagu-
lation in 420 species of insects. Arch. Biol., vol. 66, pp. 103-148.
1955b. Blood coagulation in arthropods. VI. A study by phase-contrast
microscopy of blood reactions in vitro in Onychophora and in
various groups of arthropods. Arch. Biol., vol. 66, pp. 489-508.
1957. Studies by phase-contrast microscopy on distribution of patterns of
hemolymph coagulation in insects. Smithsonian Misc. Coll., vol.
134, Pp. 1-35.
GREGOIRE, CH., and FLorKin, M.
1950. Blood coagulation in arthropods. I. The coagulation of insect blood,
as studied with the phase-contrast microscope. Physiol. Comp. et
Oecol., vol. 2, pp. 126-139.
Grécorre, Cu., and JoLivet, P.
1957. Coagulation du sang chez les arthropodes. VIII. Réactions du sang
et de l’hémolymphe in vitro, étudiées chez 210 espéces d’arthropodes
africains. Inst. Parcs Nat. Congo Belge. Expl. Parc Nat. Albert,
sér. 2, fasc. 4, pp. 1-42.
Harpy, W. B.
1892. The blood corpuscles of the Crustacea, together with a suggestion as
to the origin of the crustacean fibrin-ferment. Journ. Physiol.,
vol. 3, pp. 165-190.
NO. 3 HEMOLYMPH COAGULATION IN INSECTS 23
Jones, J. C.
1956. The hemocytes of Sarcophaga bullata Parker. Journ. Morphol., vol.
99, pp. 233-258.
NuMANOI, H.
1938. On crustacean blood coagulation. Japan. Journ. Zool., vol. 7, pp.
613-641.
sear
1910. Crustacean blood coagulation as studied in the Arthrostraca. Quart.
Journ, Exper. Physiol., vol. 3, pp. 1-20.
1911. Types of crustacean blood coagulation. Journ. Mar. Biol. Assoc.,
vol. 9, pp. 191-108.
Tait, J., and Gunn, J. D.
1918. The blood of Astacus fluviatilis: a study in crustacean blood, with
special reference to coagulation and phagocytosis. Quart. Journ.
Exper. Physiol., vol. 12, pp. 35-80.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 4
A REVIEW OF THE GENUS HOPLOMYsS
(THICK-SPINED RATS), WITH DESCRIP-
TION OF A NEW FORM FROM ISLA
ESCUDO DE VERAGUAS, PANAMA
By
CHARLES O. HANDLEY, JR.
Associate Curator, Division of Mammals
United States National Museum
Smithsonian Institution
(PUBLICATION 4380)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JULY 3, 1959
THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.
A REVIEW OF THE GENUS HOPLOMYS (THICK-
SPINED RATS), WITH DESCRIPTION OF A
NEW FORM FROM ISLA ESCUDO
DE VERAGUAS, PANAMA
By CHARLES O. HANDLEY, JR.
Associate Curator, Division of Mammals
United States National Museum
Smithsonian Institution
A specimen of the thick-spined rat, Hoplomys gymnurus Thomas,
that Alexander Wetmore shot in a thicket on Isla Escudo de
Veraguas on the morning of March 1, 1958, is probably the only
mammal from this Caribbean island that is preserved in a museum.
Other rats that Wetmore saw in coconut palms on the same day
apparently were of another genus. No other mammals have been
reported from this locality except feral hogs. Although Indians once
lived on the island, human beings are now only transients there.
Escudo de Veraguas is a low island, about 1 mile wide and
2.5 miles long, in the Caribbean Sea, 11 miles off the base of the
Valiente Peninsula, Province of Bocas del Toro, north coast of the
Republic of Panama. Wetmore (Smithsonian Misc. Coll., vol. 139,
No. 2, 1959) has given a detailed account of the history, geography,
and zoological position of the island.
Other echimyid genera, Diplomys and Proechimys, are known to
occur on certain islands in the Gulf of Panama and elsewhere, but no
insular populations of Hoplomys have been reported. The Escudo de
Veraguas Hoplomys differs in so many respects from other known
populations of the thick-spined rat that it has prompted a brief review
of the genus.
Many of the National Museum (US) specimens reported here were
collected in cooperation with the Gorgas Memorial Laboratory,
Panama. I express my thanks to Carl Johnson, director, and other
members of the laboratory staff for numerous courtesies and assistance
in fieldwork. Some of the specimens were collected by C. M. Keenan
of the Army Preventive Medicine Survey Detachment, Ft. Clayton,
Canal Zone. Richard Van Gelder kindly permitted the study of speci-
mens in the American Museum of Natural History (AMNH), New
York.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 4
A
«J ¥! f \
mic i |i
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Genus HOPLOMYS J. A. Allen
1908. Hoplomys J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 24, p. 649.
Genotype.—Hoplomys truet J. A. Allen.
Distribution—The genus has a limited distribution in Central
America and northwestern South America. It is monotypic. Published
records of collecting localities are mapped in figure 1. Hoplomys is
known to occur at medium elevations (800-3,100 ft.) on the Caribbean
slope of the highlands of Nicaragua and Costa Rica; near sea level
on the Caribbean coast of Panama; at medium elevations (600-4,000
ft.) on the Pacific slope of eastern Panama, Colombia, and Ecuador ;
and near sea level in extreme southwestern Colombia and north-
western Ecuador. The distribution of Hoplomys in South America
appears to be limited by the Western Andes. J. A. Allen’s record for
Puerto Valdivia on the Rio Cauca (Bull. Amer. Mus. Nat. Hist.,
vol. 35, p. 207, 1916) is erroneous (the specimen is a Proechimys).
Proechimys cayennensis hoplomyoides Tate (Bull. Amer. Mus. Nat.
Hist., vol. 76, p. 179, 1939) from Mt. Roraima, Venezuela, appears
not to be a Hoplomys, although a relationship has been suggested
(Moojen, Univ. Kansas Publ., Mus. Nat. Hist., vol. 1, p. 324, 1948).
In the Caribbean lowlands of Panama, where Proechimys is abun-
dant and Hoplomys seemingly rare, I have trapped individuals of
both genera under the same log on successive nights. At medium alti-
tudes in the mountains of Panama where Hoplomys is fairly common,
Proechimys apparently does not occur.
All the Hoplomys that I have collected in Panama were caught in
banana-baited live traps under large decaying logs—in fairly open
mature rain forest, in grassy clearings and adjacent streamside
thickets, and in dense, hillside Heliconia thickets. Goldman (Smith-
sonian Misc. Coll., vol. 69, No. 5, p. 124, 1920) found Hoplomys
associated with fallen trees and rocks in Panamanian forests.
Diagnosis ——Dorsum, flanks, and rump, in both adult and juvenile
pelages, with spines 26 to 33 mm. in maximum length and 1.5 to 2.0
mm. in maximum diameter, tending to obscure soft fur. Tail shorter
than head and body, scaly, and sparsely haired. Ears scantily haired.
Hind feet long and narrow; fifth toe scarcely longer than first; claws
relatively straight, but claw of second toe slightly expanded. Skull
prominently ridged, and supraorbital shelf beaded ; rostrum relatively
broad at tip ; auditory bullae relatively small; and infraorbital foramen
without subsidiary canal on floor, and with external wall thin in
lateral view. Cheek teeth with oblique folds; counterfold formula
normally 4/4-4/4-4/4-4/4, rarely 4/4-4/3-4/3-4/3.
NO. 4 HOPLOMYS (THICK-SPINED RATS )—-HANDLEY 3
V aviation Specimens of Hoplomys have never before been availa-
ble in series. Fourteen specimens, seven of which are adult, recently
collected on Cerro Azul, Panama, now permit a fairly good estimate
RiO COCO
BIJAGUA
SAVALA GATUN
RfO INDIO
FT. SHERMAN
CERRO AZUL
RiO CANGANDI”
CANA
MT. TACARCUNA
SUERRE
STA. TERESA
PERALTA
ESCUDO DE VERAGUAS
ALTO BONITO
BAGADO
RIO TAMANA.
LA GUAYACANA
BARBACOAS
BUENAVISTA
SAN JAVIER
CACHABI-
Fic. 1.—Distribution of Hoplomys gymnurus. All known specimen localities
are indicated.
of individual variation in the genus. Eleven specimens from Darien
and nine from the Canal Zone, are also helpful. In addition, random
series of up to 75 specimens per sample of the closely related
Proechimys semispinosus have been used to evaluate the variations
seen in the smaller series of Hoplomys.
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Size, flatness, and ridging of the skull increase with advancing age
in Hoplomys and Proechimys. Tooth wear appears to be a reliable
criterion of age. Full adult pelage usually is attained after M3 appears
and before it becomes functional. Only juvenile and adult pelages have
been distinguished. Specimens in which all cheek teeth are functional
are considered to be adults. Generally, the largest, flattest, most
heavily ridged skulls have the most worn teeth. Apparently these
rodents continue to grow after all teeth are functional. Thus, there
is considerable size spread among adult skulls. For this reason only
maximum and minimum figures are given in the table of measurements
(p. 6).
Body sizes appear to be uniform throughout the mainland range
of the species, but larger on Escudo de Veraguas. The skull is narrow
in the south—Ecuador, Colombia, and Darién—somewhat broader
in central Panama, Costa Rica, and Nicaragua, and broadest of all
on Escudo de Veraguas. Likewise the nasals and cheek teeth are
smaller in the southern populations. Size of the auditory bullae
increases northward from Ecuador to Nicaragua, but the bullae are
largest and most inflated anterolaterally in the Escudo specimen.
Several features of the zygomatic arches vary geographically. The
maxillary roots of the zygomata flare less widely and less perpendicu-
larly from the longitudinal axis of the skull (so that the zygomata are
more convergent anteriorly) from the Canal Zone southward than
they do in the north. From Cerro Azul southward the maxillary roots
tend to flare up, away from the ventral plane of the skull, rather than
paralleling that plane as they do in the north. The jugal has a hooklike
posteroventral process in most Canal Zone and Cerru Azul specimens,
but not in others. Most of the specimens from Ecuador, Colombia,
and Darién, and a smaller percentage of the central Panamanian
specimens have a small conical projection on the dorsal edge of the
zygoma at the jugal-squamosal suture. I failed to check this character
in the Costa Rican and Nicaraguan specimens. There is hardly a
trace of it in the Escudo individual. The nasals, broad and posteriorly
truncate in the island specimen, are usually narrower and posteriorly
acute in mainland populations.
Among mainland populations of Hoplomys flatness and ridging of
the skulls of mature individuals are similar to these features in mature
individuals of Proechimys semispinosus. None of the available
Hoplomys or Proechimys closely approaches the Escudo specimen in
flatness or ridging, despite the fact that the island specimen, judged
by tooth wear, is a prime adult, not as old as many individuals with
NO. 4 HOPLOMYS (THICK-SPINED RATS )—-HANDLEY 5
which it was compared. The degree of reduction of dorsal doming and
ventral depression of the brain case of the Escudo specimen is reflected
in the convergence of greatest and condylobasal lengths of the skull,
and in the more posteriorly oriented (as opposed to ventrally
oriented) foramen magnum.
The thick spines that distinguish Hoplomys are longest and
strongest just behind the shoulders on the upper midback, from
which point they diminish in size in all directions. The spines possibly
vary geographically in size. They appear to be longer and stronger
toward the southern part of the range of Hoplomys. The Escudo
specimen, although it is larger than any other, has the smallest and
weakest spines. Maximum spine length varies as follows (mean,
followed by extremes): 6 Ecuador 29 mm. (28-31), 4 Darién 30
(28-33), 11 Cerro: Azul 28) (26-29), 5 Canal Zone’ 28 (27-30),
1 Escudo de Veraguas 26.
Coloration of the spines is individually variable. All specimens
have all spines proximally white and distally colored. The tips of those
of the dorsum are always black, but the flank spines usually are tinged
with orange or banded with orange and black distally. Occasionally
the flank spines are colored like the dorsal spines.
Coloration of the soft hairs of the dorsum is geographically variable.
At the southern extreme they are reddish orange, especially on the
shoulders. The soft hairs of the Escudo specimen are similar but
darker and brighter. Costa Rican and Nicaraguan examples have the
hairs more orange, and those from Panama and northern Colombia
are more yellowish on the average. The presence or absence of black
ocular and crown areas appears to be individually variable throughout
the range of Hoplomys, but only the Escudo specimen has the soft
hairs blackened to form a distinct middorsal stripe from snout to base
of tail.
All populations have the underparts dominantly white, and all have
some individuals that show encroachment of agouti hairs of the side
neck onto the throat, suggesting an incipient collar. This is well
marked in the Escudo specimen; one from Rio Indio, Canal Zone,
has a complete collar. Nine of the 14 Cerro Azul specimens have
clear orange collars, and several of them have a band of clear orange
hairs separating the agouti hairs of the flanks from the white hair of
the belly. Neither of these features is seen in samples of other popu-
lations. Coloration of the forefeet (usually white on the inner side,
colored on the outer, occasionally colored throughout), and coloration
of the cheeks (clear orange, buff, gray, or agouti) are individually
variable.
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
+5°bS
6°0S
QbvS-E'1S
QZ5-L'25
z'bS—9'2S
E-gS-L-2S
0°9S-0'FS
oes
6'gS
zvs
oss
0'z9
y}su2]|
[eseqo[Apuos
+r09
¥'gS
Q'Z9-1'°9S
£S9-S°6S
£°19-9°6S
£99-9'09
“£E9-1'19
£68
1°99
S09
1'z9
zL9
here er eke
Yy}sue]
yso}yeatH
Le-17
be-zz
Cz-€z
of-z~e
IZ
bz
ve
€z
(yo}0u WO1}) (Arp “n'9)
1eq
gS—oS
pS-1S
1S—0S
og-1S
£S-+S
19-vS
6S—-1S
zg-OS
19
zs
cs
gs
q00F PUT
SgI-gSI
OgI-OF 1
OO
bee-rgi
88
SSz-SgI
bZ1
brz—o0z
Sgi
OBIqd}I9A
Pet
Soz—-getz
oLz-tbz
goSz-£hz
Sgz—-bez
Sez—-Ctz2
SZz-g1z
ogz-6Se
Sge-Izz
GZze
obz
oz
£1¢
qjsuea] Apoq
pue peayy
oSb—-96€ Ta ozetier ekeeevats (qnys I Ayuo ) Soy[euloy v
oSb-—z6e ailevel/anelle (e)iei-o/ewiere (24s I Ayuo) soyeur ¢
Jopenoy “MN pue eiquiojog “A'S
6or 6.0) © (a 8c. \6 © e,ele>s ome eee eee ere Soyeway Z
61S-¢br ©) [sire isn (ole) ea, s s!ia) a) felie re liake) eusas el wice\\areie. So] eu 9
eueueg ‘usted
Scr Byejim: (@50) (ele) \0) ele e:elje\lee ekeleies eee So] eUloy c
o£S-—for Wiis; 6sloxe oreiey a dastene teitotloleistevacal sus) w eve soyeul ¢
eureueg “nzy O1109
ber eijei.e) wale |e Jereye) pier of aiielie (eis! eo) scene) .e oe SoTeway ie
6¢S-tSt mesic lsguelal elites suasoreWerevegene lamers diet etele soyeul £
vueueg ‘ouo07 [eue)
Bettie: teeeees oven I
ey vISOD “VYeIIg esrIay, “eS
OoVr G OrO) Ch OnOstia) G0 202'@ne:(0);4) 008) sie) eles . a]eUuloy I
eee ior Beers veeeeees aren 7
enseiedin ‘enselig
gor 2/6 Qupun ees) 0 ace) spe eveheve oe. sere ee . oyeur I
eureued ‘Sense1a A. 2p ophosa
Yy8uz] [e}0T,
‘SAIIIUY UM UL UdQAH 24D S}UaMaANSDIML WNUXDIL PUD mn ay J
sninuws skwojdoyy yjnpp fo Spuamadnsvam Joudéd puo [DUsAIXY—I AIAv IL,
HOPLOMYS (THICK-SPINED RATS )—-HANDLEY
NO. 4
og
gZ
S-Z-b'°Z
age-i'Z
£°9-S°Z
Eg-S°Z
PlLel
gl-eL
£2
bZ
ZZ
9°8
4 yapearq
ayeleg
ia 9°8
ev 6g
o'S-o't 6'9-£'2
grer = 9'6-9°8
zs-2'V o°01-S'6
g'S-S‘b Z'01-0°6
9S°-9' 9°6-1°6
o'S—9'V S°6-0'g
gS 36
LY 16
LY $6
I'9 I’or
qW33u9] y MOI 300}
Use 1OF AILT[IXCJY
DAISIOUT
+6:0z
F'0z
ofz-Z°61
¢-€z-6'0z
S'7eZ-1'1ze
oSc-r'1z
1°Sz-o0'7~
Z'Qz-0'1z
IZ?
Cze
Cec
g'Sz
q33u9] [ese Ny
giz
g'0z
I'1z-£'0z
¥'7Z-Q'0z
v1z—-9'0z
Lze-0'1z
g'1z-9'1z
Q'ez-1'1z
£77
g'0z
r'bvz
qypessq
[eprloyse yy
F'O1
Lol
PeiI-LeI
ZLEI-1'€1
Pei ei
Ivi-S'E1
I°€I-1'f1
ge1-o'f1
I'vi
CCI
o'vi
zSI
MIRE t
[e}iq1010}UT
“pamuyuoj—I ATaVT,
‘sq JO Wooaye jo sursieur 19yno0 usemzog |
£92
0'9z
bgz—-h' Lz
982-892
1°6z-€°9z
6°0£-L'9z
9'6c-b'6z
0'0£-7'9z
Toe
£°92
£32
Eze
qypessq
d1}EUIOBAT
“qysuey, AL[OOATY »
bette teen ee eeeeeeeees greumay I
cee eee ee ee eee ese eee eee ayeul I
Jopendy “MN pue erquiojod “A'S
eee eee wee wee ew eee
SoTeUloy Z
see eeeeeeeeeeeesereees sore g
eweueg ‘Usieqd
ere eee ee ee eee eee eee soyemoy c
Sitelole elfeiipib rails terete Valtadslaheteteltenie soyeu S
eweueg qnzy O19
"+" SoreUaz Z
wee ee ee eee ee eee eee eee sayeur ©
eureueg ‘ouoZ yeued)
reese ee eeeeeeeeeeeeeres QrpUT T
BINT eBSOD ‘eyesag esatay, “eS
ee weer eee eer eee eee wees od] euloy I
“* oyeut I
enseiesin ‘enselig
Ce ee
Pees scene eeeeeeeeeeeees oreUT T
eureueg ‘senseiaA op opnosy
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
The large size of the Escudo specimen, the massiveness, broadness,
and heavy ridging of its skull, the inflation of its auditory bullae,
and its distinctive coloration all seem to be beyond the possibility of
individual variation. This suggests that the Escudo animal is taxo-
nomically distinct from mainland populations. That it is conspecific
with them is indicated by its alignment with some of the morphologi-
cal clines observed in the mainland populations. Wetmore (of. cit.)
has named three birds (a wren, a manakin, and a tanager) collected
on Escudo de Veraguas that differ from their mainland counterparts
in greater size, in addition to differences in color.
Classification—The genus Hoplomys is represented by one species,
which includes four subspecies :
HOPLOMYS GYMNURUS GOETHALSI Goldman
1912. Hoplomys goethalsi Goldman, Smithsonian Misc. Coll., vol. 56, No. 36,
p. 10 (Rio Indio, near Gatun, Canal Zone, Panama).
Characters —Size medium; skull of medium width and ridged;
brain case domed and slightly depressed ; foramen magnum ventrally
oriented ; cheek teeth large; auditory bullae medium; zygomata con-
verging conspicuously anteriorly, and maxillary root tending to flare
up slightly from ventral plane of skull; jugal with hooklike postero-
ventral process and small conical posterodorsal projection ; nasals long,
narrow, and posteriorly acute; dorsal spines long and strong; soft
hairs of dorsum appear uniform yellowish orange in mass effect.
Specimens examined.—Panama: Cana, 2,000 ft., 5 US; Cerro
Azul; 2:100 it:., 14 US; Ft? Sherman,.4 US; Gatun, 3 (2 AMNE,
1 US) ; Cerro Tacarcuna, 2,650 ft.,6 AMNH; Rio Cangandi, 200 ft.,
t US; Rio Indio, 2 US. Colombia: Alto Bonito, Antioquia, 1,500 ft.,
1 AMNH;; Bagado, Choco, 600 ft., 2 AMNH.
Additional published records——Colombia: Rio Tamana, branch of
the Rio San Juan, Choco (J. A. Allen, Bull. Amer. Mus. Nat. Hist.,
vol. 35, p. 207, 1916).
HOPLOMYS GYMNURUS GYMNURUS Thomas
1897. Echimys gymnurus Thomas, Ann. Mag. Nat. Hist., ser. 6, vol. 20, p. 550
(Cachabi, N. Ecuador, alt. 560 ft.).
Characters.—Size medium or small; skull narrow and ridged ; brain
case domed and slightly depressed; foramen magnum ventrally
oriented; cheek teeth small; auditory bullae small; zygomata con-
verging conspicuously anteriorly, and maxillary root flaring up from
ventral plane of skull; jugal lacking posteroventral process, but with
NO. 4 HOPLOMYS (THICK-SPINED RATS )—-HANDLEY 9
prominent conical posterodorsal projection; nasals short, narrow,
and posteriorly acute; dorsal spines long and strong; soft hairs of
dorsum giving reddish-orange mass effect, slightly darkened on
shoulders.
Specimens examined—Colombia: Barbacoas, Narinmo [75 ft.],
8 AMNH;; Buenavista, Narifio [1,200 ft.], 1 AMNH; La Guayacana,
Narino, 800 ft., 2 US. Ecuador: Mindo, Rio Blanco [4,000 ft.],
1 AMNH;; San Javier, 60 ft., 7 (1 AMNH, 6 US).
Additional published records —Ecuador : Cachabi, 560 ft. (Thomas,
Op Cit pe 551).
HOPLOMYS GYMNURUS TRUEI J. A. Allen
1896. Echimys semispinosus Alfaro (not Tomes, 1860, Proc. Zool. Soc. Lon-
don, p. 265), Primera Exposicion Centroamericana de Guatemala, Museo
Nacional, San José, p. 41 (Suerre, Costa Rica).
1908. Hoplomys truei J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 24, p. 650
(Savala, Matagalpa Prov., Nicaragua).
Characters —Size medium; skull of medium width and ridged;
brain case domed and slightly depressed ; foramen magnum ventrally
oriented; cheek teeth large; auditory bullae large; zygomata con-
verging less anteriorly than in goethalsi, and maxillary root in plane
of ventral surface of skull; jugal without hooklike posteroventral
process; nasals long, narrow, and posteriorly acute; dorsal spines
relatively short and weak; soft hairs of dorsum giving uniform dark
orange mass effect.
Specimens examined.—Nicaragua: Lavala [ = Savala, 800 ft., along
the inner border of the low east coast region], 2 AMNH; Rio Coco
[800 ft.], 2 AMNH; Vijagua [= Bijagua, probably 1,500 to 2,000 ft.,
on eastern slope of highlands in Matagalpa Prov.], 3 AMNH. Costa
Rica: Santa Teresa Peralta [3,100 ft.], 1 AMNH; Suerre, 1,500 ft.
[near Jiménez], 1 AMNH.
Additional published records.—Tate (Bull. Amer. Mus. Nat. Hist.,
vol. 68, p. 401, 1935) supposed that True’s record (Proc. U. S. Nat.
Mus., vol. 11, p. 467, 1889) of Echinomys semispinosus in Nicaragua
was the first published reference to a Hoplomys. The specimens,
still in the U. S. National Museum, however, are Proechimys.
HOPLOMYS GYMNURUS WETMOREI subsp. nov.
Holotype —U.S.N.M. No. 307057; adult male, skin and skull;
collected March 1, 1958, by Alexander Wetmore; Isla Escudo de
Veraguas, Prov. Bocas del Toro, Panama; original No. 1479.
10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Characters.—Size large; skull broad and heavily ridged; brain
case flattened dorsally and ventrally; foramen magnum posteriorly
oriented ; cheek teeth large ; auditory bullae large and inflated antero-
laterally ; zygomata converging less anteriorly than in goethalsi, and
maxillary root in plane of ventral surface of skull ; jugal without hook-
like posteroventral process, or conical posterodorsal projection ; nasals
long, broad, and posteriorly truncate; dorsal spines relatively short
and weak; and soft hairs of dorsum giving dark reddish-orange mass
effect (between Burnt Sienna and Sanford’s Brown of Ridgway, 1912,
Color Standards and Color Nomenclature), with black middorsal
stripe from snout to base of tail. For measurements see table 1.
Specimen examined.—The holotype.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 5
Charles DB. and Mary Waux Walcott
Research Fund
GENERA OF TERTIARY AND RECENT
RHYNCHONELLOID BRACHIOPODS
(Wit 22 PLatTEs)
By
G. ARTHUR COOPER
Head Curator, Department of Geology
United States National Museum
Smithsonian Institution
{ nov 25 1959
\
AS
we dif Pe fi RN
CITY OF WASHINGTON :
PUBLISHED BY THE SMITHSONIAN INSTITUTION
NOVEMBER 23, 1959
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 5
Charles D. and Mary Waux CA alcott
Research Fund
Gene Or. TERTIARY AND: RECENT
RHYNCHONELLOID BRACHIOPODS
(WitH 22 PLATES)
By
G. ARTHUR COOPER
Head Curator, Department of Geology
United States National Museum
Smithsonian Institution
(PUBLICATION 4382)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
NOVEMBER 23, 1959
SMITHSONIAN ,
INSTITUTION NOV 2 3 195¢
THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.
CONTENTS
Page
AEE ORUICELOEN i repare cleats eri ae ware) at oisn-w 6 (a,c) sta dk Sater y wataieranet dveraralcecteratartvarc <laleie I
icksro pled ettien tsi oratsieiots: aims ahaa aro aco! patecs/a pcaetenoy ave alenatata arate sapepal ia were Wa 2
RbynchorielloidwmOrpnolo ey ooh os \o:4.0 arse) a) 0, sayy eowiessie/sreysialepele)« a alavaidieisls/oa\6 3
Béalccharactersmme occichi aw cinos.c'08 tereie a pete are ero ctaMaree a Dabaiereie alovers 3
Interior characters of the ‘pedicle valve. io).255 cc emaie cas cw escc ss 4
Intemornmcharactersvotthie: prachialluvalvesmc- see cence seeeiciiae ciel 5
Gacditial Processes tae aie Fite. ain oe isvePe sic ob geaes oat ote t Sr ete BUG ies rahe ale 5
VANE ERP lALEST eyerciess crarctercvorsie fo,aio-sca syn arse: Se Giete sere e rt reste Pese rate atte elaiare 6
GEUG AN Mhate a cicteisys els wins, Sletehem.a wos da SOR ERT ate acta oleae cemiore 6
Median ridges, median septa, and camerae..............ccceeees 9
teTI1O hs GHALACEEN SY tgs tos a/aldiaicid coe: ola tain ahd she aise one Wee ANE Oe maura ee 10
Olrtrtsiseta pation ye fe icteis ialete See. 6.05 <isio's se Sewers es usar aaa casa clasaitelee 10
Sol inte everer sre eioes Seiko ae raved oh, RO ACL tore RE EIS eLearn II
Rhynchonelloidelassification® 27.0555 0 vo nae eee eee ce inicio aie’s ects oieveleietetah 12
Family and generic arrangement and characters............seeeeececees 14
Hanatlys Ory PlOPOLlUA, tac mise e hac occ rere ensrsimie es Sica Bain ae eta luelaliers 17
COA PEO BON Gr ule Sista del tthe PANES Seok See 17
INUIT eRe calatars ei neie oie ie wheel oe Oa Te Breasts AAS SN IRE cae 2
Heratlye MS aSIOMUAG «ste ss reealeis oe oa e seis eis elec clarns cis ave tiayeter she ta ate otere 25
Sabtanilye*Basiliolinaey 4 ence cose tee ae em ne eee te anaes wrotstac 25
PUSH OUEST Heia wiaiss Acta takes Sasa Na ah, SL) osteo ake engh buts te; aves oo ka a Tera ns arene 25
ORG HAEWN IAS ope savere toh « ais Sarak srs aVo aia osstinlsteswvetars:avayalelale/sistores teetely «skaters 30
IN COC ULE TUS) tarsi stace See che sa ieiAioi nies oi Sake Tomei ni vise nied oloaatteveteteroveariets 34
INICOLIVN CHADS cris otates soe eee hae Na cl ete RIE Oe, SEO Tee Peete el sieket steaks eae 34
CAVE UMICHITDN Sic niovals/ ciate SRS atts eater ere lo aie sheer he are air eoreieieeee 35
ROD OUATMIE es elena ele ta etalon erate IS & fata heer aren Petraes 37
SREP TANI A Ti les ara cd ASE a Male RS OPE SBP A thus kwaebs ie otares 38
StubtamilyeA phelestinaer.pavecr cach pees oie herahe oak steno teteceiatske aks 41
VAD WWE LE SUR W ore) ce tee yor ors chav el olds Sor cust Ya SATE aoe AG euwceraee ehaielyy Govelorolenateeienait 4I
Subtannly eA etherina ems cats src teacetersern oa ete cence osatelatsiokaret sis otelecolenctohels 42
A CEIVED as MMe a Taare ONC Late eC TS A eC NTN eee ee Gietoatareie chavecrenstereeee 42
IZ GLAGORLANACHIG, WR. ie RTE eS ee I eee ete ae ee ee A4
Bamilyettemuthynidaeyen ave ttvcicicccs ck cicus oils ete eae cherote ic te soe 45
CIALLY LES eps TRS GETS TS or TELE or as lavas rey Sssioatena Fale ts 45
INO FOS CUGE Haren oxy oy oes okT is shes SHEEP ae neee Ae eile ake eaeioetaie see aie 48
UR COMLOPA VACHE itor ine cooks ss ropa oins eels Slee eyba elo eiesleel ee Sloe 50
SEL VITCHEG Meee oe were cee eA aie ie ee ieee Sata eon sicaletste Bi Seiats 52
Pamiuly PHrielewdae iy. Bogs. ee feo oa se oie ad cai a oie Wine c eibiele s etelore rete 53
PROSCL CLE 5 5) cPak do eles faci nie otike Aa We ad ia aah eata a heataere Biola ae ele Ie 53
COMBO SBEY TES 2 avocs So raleveroyaiore Gio’ svaiars et alot meri Marea ele ort earela’ ae 50
COM IR CLOIUTS a EIS OG lake ia io) os aare she acs Sisto eto esa Iee oS a eke 58
ratty eAIS PAIL MY MEMIMGAE acc ays (x/acc-e s)«/s ss vcore gate aapehoiss aha 8 xiehatonalajalsls, sd) che 50
TLASPOWAT AY NCMBO acc coc occa sce Ses sae deta eulsslewleees «savas s 50
IS DHE MATING te ac aals accra dirs shoes tes aratoetalay oe ata arate. Seamer ait tee cletece 62
Baty, Porytastiarit ae ah.g. atejora.0,2/eieret 0 3 « xpanetovs Sava Sate ae eherwletd aibr® syaelels 64
PEP AAIN AE TR at hey Pca tats, Sas aro i> 5 stig Chee pedestal ge ese eels aks ns OG 64
RR TRVIAECHISDECICS 7 cries oickess ahora reesreist sso cs af nro, oe 0s ty ecaeta eam a Re ee aaeral as 66
REL ETC OSH rere oreo VS eee EPA Ree owes Sears EES ne ors (SPST ecco are ea eI RC aietie ole et a iene 67
Perera AM TREGEN MOE! PALES ogc. oes) S516 cis wre oo eel ave te 3 sos eve waaay ston eer afecaues slahaie 73
ui Wis bes ete Pa
ae hak Hey ie Coatin
isin <a Why coat oan m
Mr al a CA oF wey
Ih Fe (aha CEASE AAD
eee ns a ROR ‘Ae Ne
“ean gy Nant ;
‘ Han an tt Lite Noten thea yee heat
ne ALE
sees
teat ey
ie Wz pve iad ‘bn
RC ae
ws - Ks Biel aa ema
pike rien ow a
ON fh * ee a ‘ taba saa
he! 10 a te nag
i / )
ey
ea nay
PAO MANE we hens 4%
a "igh caldera ye ei sa eh Nabalioe teem ee ie sr
BIEN Vas) 8
aig ee Se el
a veutinn,
FARM ait Wiens ;
Masa sints Ashe)
Na aA cays pee Rs
the
ren sa
Pe shah dsy. Od, 4
gi aa 9 :
i i ;
my Elkins ee
wi Beh. catjann diane)
i % t, ie “yay
ea
i ; ; 7 § MF J
1 y Ly ‘i A 4
» eee ae
er ayaa ;
aed nN
Charles DB. and Mary Vaux Walcott Research Fund
GENERA OF TERTIARY AND RECENT
RHYNCHONELLOID BRACHIOPODS
By G. ARTHUR COOPER
Head Curator, Department of Geology
United States National Museum
Snuthsonian Institution
(WitH 22 PratEs)
INTRODUCTION
For several years the writer has been studying the few brachiopods
known from the Tertiary formations in the eastern United States.
Most of the species are terebratuloids but a few rhynchonelloids ap-
pear in the collection. The Palmer collection of Cretaceous and Terti-
ary brachiopods from Cuba was also made available. This collection,
too, contains a few rhynchonelloids. This group of brachiopods seems
to be unusual in Tertiary deposits and the same is true of the rhyncho-
nelloids of modern seas. In the study of the Tertiary forms it proved
necessary to compare them with modern representatives. In making
these comparisons it became evident that modern rhynchonelloids
have been well described in only a few instances and very few of them
have ever been adequately illustrated. Inasmuch as the collection of
Recent brachiopods in the National Museum contains a good repre-
sentation of modern rhynchonelloids, the opportunity presented itself
to correct the deficiencies outlined above.
In addition to the American Tertiary rhynchonelloids, some species
from outside North America are also included. The National collec-
tions do not contain many representatives from foreign Tertiary de-
posits but some good specimens are available from the Mediterranean
region and elsewhere. These made possible the figuring and descrip-
tion of some new or little-known genera.
Although this monograph adds considerably to our knowledge of
Recent and Tertiary genera of rhynchonelloids it does not include all
the known species or all the possible genera. A number of species are
known from European deposits but the interior details have never been
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 5
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
described, nor were specimens available to use in this study. Conse-
quently it is possible to assign to their proper genera only some of the
known species. It is also known that the interior of a number of
species differs from that of any of the genera discussed herein, but
these species are represented by too few specimens to make generic
description possible. Much therefore still remains to be done in the
study of the Tertiary and Recent rhynchonelloids.
Possibly the biggest handicap in the study of modern and Tertiary
rhynchonelloids is the fact that, except in a few instances, the speci-
mens are quite rare. Several of the Recent species are known from one
or two specimens only, yet their morphological details are unique or
sufficiently different from known genera to make it impossible to
include them in any of the established categories.
Some of the Tertiary species are sufficiently numerous for good
descriptive work but their describers seldom made any effort to obtain
interior details. Davidson (1870) did not describe the interior of any
of the Italian Tertiary rhynchonelloids, probably because emphasis in
his day was on description of the species. Later authors seemed to
be content to assign many of the modern species to Hemithyris regard-
less of whether or not the interior or exterior details were in ac-
cordance with the generic characters of the type species. In present
times emphasis is now placed on interior details because it is on them
that the family and frequently the generic characters are based.
ACKNOWLEDGMENTS
In any study of this sort it is necessary to ask help of one’s col-
leagues. I am grateful to all the scientists listed below for their help.
Dr. Helen M. Muir-Wood, Deputy Keeper, Department of Palaeon-
tology, in charge of the brachiopods in the British Museum (Natural
History), furnished a specimen of Compsothyris, photographs of
Rhynchonella grayi Woodward, photographs of serial sections of FR.
polymorpha (Massalongo) and R. bipartita (Brocchi), and casts of
the specimens serially sectioned. Dr. A. Vandercammen, Subdirector
of the Laboratory, Royal Institute of Natural Sciences, Belgium,
furnished a specimen of Mannia nysti Davidson, a rare Belgian
species.
Mrs. Ellen J. Trumbull of the U. S. Geological Survey made avail-
able specimens of west coast Tertiary species. Dr. J. H. Peck, Senior
Museum Paleontologist of the Museum of Paleontology, University
of California, made available a fine suite of topotypes of Eohemithyris
which made possible preparations of the inner details of that interest-
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 3
ing genus described and illustrated herein. Dr. L. G. Hertlein, Uni-
versity of California at Los Angeles, lent paratypes of Eohemithyris.
Dr. E. Montanaro Gallitelli, University of Modena, Italy, presented
the National Museum with specimens of Rhynchonella polymorpha
(Massalongo) that made it possible to prepare the interior details of
the cardinalia described and figured in this monograph.
RHYNCHONELLOID MORPHOLOGY
Throughout geological time the rhynchonelloids have been charac-
terized by triangular to subpentagonal form, with prominent beak and
a strong median fold on one valve, usually the brachial valve, and a
deep sulcus on the other. Nearly all the genera are provided with a
conspicuous beak having a foramen modified by deltidial plates. In
some genera, especially a few of the Recent, Tertiary, or Mesozoic
ones, the deltidial plates are elaborately auriculate, a feature unusual
in other brachiopods, The rhynchonelloid shell is commonly costate ;
smooth forms are usual in Recent, Tertiary, and Mesozoic families
but rare in Paleozoic representatives. The distinctive feature of the
rhynchonelloid interior is the more or less long curved crura and
hinge plates which characterize the cardinalia. Details of these latter
features have long been neglected.
BEAK CHARACTERS
It is not here the intention to discuss these characters for all the
rhynchonelloid brachiopods but to point out the significant features
shown by the genera discussed herein. Of modern and Tertiary
rhynchonelloids only Cryptopora and Mannia do not have a small
round or elongate-oval foramen. In the two genera mentioned the
foramen is elongate-triangular and is restricted only slightly by at-
tenuated deltidial plates, which, unlike most other modern and Tertiary
genera, form an elevated rim on the sides of the delthyrium.
The nature and completeness of the deltidial plates usually define
the form of the foramen. In some genera the deltidial plates are dis-
junct, that is, they do not meet on the anterior side of the foramen.
In such cases the foramen is said to be incomplete. An excellent ex-
ample of this type is Hemithyris. When the deltidial plates meet on
the anterior side of the foramen, they are spoken of as conjunct and
the foramen is completely enclosed. Examples are Basiliola, Aphelesia,
and Aetheia. These two conditions of the deltidial plates, disjunct or
conjunct, are given considerable weight in genus making by some
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
workers. Yabe and Hatai (1934), for example, distinguished their
genus Neohemithyris (=Basiliola) from Hemuthyris chiefly on this
basis.
Another feature given importance in the study of the rhynchonel-
loids is the position of the foramen in relation to the beak. A common
condition is one in which the entire foramen is surrounded by the del-
tidial plates and is called a hypothyrid foramen. In other genera the
foramen has migrated posteriorly because of pedicle pressure and has
thus resorbed or worn away the portion of the deltidial plates on its
posterior side. In this condition, which is called submesothyrid, the
foramen is bounded posteriorly by the beak and anteriorly by the del-
tidial plates. A further condition is called mesothyrid and results from
continued posterior migration of the foramen, which has resorbed part
of the beak and is bounded posteriorly by part of the curving umbo and
the deltidial plates anteriorly. This condition is rare in modern and
Tertiary rhynchonelloids.
One of the most characteristic rhynchonelloid features is the rims
or winglike extensions that adorn the deltidial plates of some genera.
Perhaps the most exaggerated modern examples are those of Gram-
metaria and some species of Cryptopora in which the deltidial plates
bear prominent lateral extensions. The more common condition is that
of Basiliola in which the lateral and anterior margins of the deltidial
plates in contact with the foramen are reflected dorsally in the direc-
tion of the brachial valve and form a conspicuous lip around the fora-
men. This may have helped, in conjunction with the pedicle collar,
to form a tube which strengthened the hold of the valve on the pedicle.
INTERIOR CHARACTERS OF THE PEDICLE VALVE
Most of the Recent and Tertiary rhynchonelloids have the beak and
pedicle regions strengthened by a pedicle collar. An elaborate collar
is developed in Basiliola. The deltidial plates are conjunct and auricu-
late. Their inner margin grows laterally along the sides of the del-
thyrial cavity to meet on the floor of the valve. In B. pompholyx
(U.S.N.M. 274135) the anterior margin of the collar protrudes an-
terior to the edge of the deltidial plates and is elevated above the
valve floor (pl. 12, figs. 9, 10). In B. beecheri the anterior ends of
the deltidial plates are thickened and expanded inward to form a flat
area that rides over the umbo of the brachial valve when the valves
are opened and closed (pl. 14, A, fig. 2).
Hemithyris possesses a pedicle collar but it is not complete because
the deltidial plates are disjunct. The inner edges of the deltidial plates
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 5
are extended ventrally to the valve floor where they join to form the
collar, but this is never closed at the anterior end. In genera with dis-
junct deltidial plates the collar is seldom prominently developed. In
some instances as in Frieleia it forms a callosity at the posterior apex
against which the pedicle rests. It is suggestive of the pedicle callist of
the Orthoidea in the Paleozoic. In Cryptopora no pedicle collar is de-
veloped, but a small apical plate elevated above the valve floor serves
the same purpose, the pedicle evidently lying against it.
The dental plates are another part of the apical region of the rhyn-
chonelloid of importance in classification and generic definition. Den-
tal plates are generally present in rhynchonelloids from the time of
their origin. They are present in all but two of the Recent and Tertiary
members discussed herein. Usually they are strong and erect plates
which define narrow but distinct umbonal cavities. In a few genera
such as Rhytirhynchia, Aetheia, and Patagorhynchia the dental plates
are reduced to mere vestiges or are absent. The only specimen of
Patagorhynchia available for dissection, that figured on plate 6, A,
failed to show any trace of dental plates. Aetheia which is usually
described as lacking these structures seems to have vestiges of them.
Rhytirhynchia has fairly distinct dental plates in the Okinawa Pliocene
species but they are mere vestiges in the Recent R. sladeni (Dall)
from the Indian Ocean.
INTERIOR CHARACTERS OF THE BRACHIAL VALVE
The definitive family characters of the brachiopods are in the
brachial valve. This is the more conservative of the two valves and
thus retains its diagnostic features while parts of the pedicle valve
which is fixed to some solid object may be evolving. The most im-
portant characters of this valve are the cardinalia which embrace the
cardinal process, the hinge plates, crura, and septa. Except for
Thomson’s (1927) work, no attempt has been made to apply the fea-
tures of the cardinalia to the classification of Recent and Tertiary
rhynchonelloids. Parts of the cardinalia have been used in defining
families and subfamilies of the Paleozoic rhynchonelloid genera.
These attempts have been based on the presence or absence of a car-
dinal process. The type of crura and hinge plates, however, have not
been used even though they offer the greatest possibilities.
Cardinal process—In the modern and Tertiary brachiopods this
structure does not attain a high state of development and makes little
impress on the classification. In some Paleozoic genera the cardinal
process is a simple vertical blade, suggesting inheritance from an
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
orthoid ancestor. In the Devonian the cardinal process of some genera,
especially the robust forms that have passed under the name Uncinulus
(=Sphaerirhynchia), have elaborate cardinal processes. Some of
these appear to be secondary characters and difficult to evaluate in the
present meager state of our knowledge. The cardinal process is not
highly developed in the few Mesozoic forms, the interiors of which
have been described. In modern and Tertiary forms the most promi-
nent cardinal process is that of Plicirhynchia, a robust and thick shell.
The cardinal process of several genera such as Notosaria (pl. 6, B,
fig. 16) and Hemithyris (pl. 4, E, fig. 9) appears as a triangular
roughened area at the apex. In the younger shells it is scarcely visible
but it is fairly prominent in old or obese specimens. The majority of
the modern and Tertiary forms have no cardinal process, the diductor
muscles being inserted in a pit under the apex. The presence of a car-
dinal process in rhynchonelloids of this age is thus a ready means of
distinction.
Hinge plates——These structures are an important part of the car-
dinalia and the combination of them with various kinds of crura makes
recognizable patterns. The sockets, which are corrugated in nearly all
of the genera discussed herein, are defined by a prominent ridge that
curves anterolaterally from the apex or the cardinal process to form a
narrow cup defining the socket. This ridge may be high or low, thick
or thin, and to its inner side is attached the outer hinge plate or the
crus, depending on the genus. The outer hinge plate may not exist in
some genera or it may be a fairly broad plate between the socket ridge
and the crus. To it are attached the muscles that rotate the animal
on its pedicle. The outer hinge plates are especially well developed in
Basiliola (pl. 12, fig. 15) and Neorhynchia, but not present in
Aphelesia.
The inner hinge plates are seldom well developed but appear in
several genera. These are extensions medially from the inside edge
of the crura. They are best developed in Frieleia (pl. 15, A, fig. 10)
where they are so strong that they unite in the middle of the valve to
create a small apical chamber somewhat reminiscent of the septalium
(or cruralium?) of certain Paleozoic and Mesozoic genera. The inner
hinge plates are also developed in exaggerated form in Aetheia but
in a way different from Frieleia. In Aetheia they are not flat or slightly
concave plates but are great swellings that extend medially from the
crura and plug the whole apical region. The degree of development
of either of these hinge plates may play a role in genus definition.
Crura.—The crura are the most distinctive part of the rhynchonel-
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 7
loid shell and are usually moderately long, somewhat curved plates ex-
tending into the body cavity. To them the body wall is attached and the
brachia are attached to the anterior body wall at their extremity.
Among the Mesozoic rhynchonelloids several distinctive types of crura
have been named. The five types distinguished do not cover the pos-
sibilities among the Rhynchonelloidea because the crura of many of
the Paleozoic genera have not yet been described and illustrated. Fur-
thermore all these types are not recognizable in the Recent and Terti-
ary forms.
Rothpletz (1886, p. 86) was the first to name types of crura. He
distinguished the following (translated from the German) :
1. Radulifer type—Generally consisting of two dental plates in the
larger [pedicle] valve, a median septum in the small [brachial] valve,
two hinge plates joined at the beak of the small [brachial] valve, and
two narrow crura curved toward the large [pedicle] valve, which at
their free lower ends are provided with barbs. One can compare these
crura with the radula [Schabeisen] of the Greek athletes and I there-
fore name rhynchonellas with such crura radulifer. (Rothpletz, pl. 11,
figs. 20 and 21.)
2. Falcifer type-—The crura, with otherwise like structures, rarely
have the form, as with Jacunosa according to Quenstedt’s researches,
the form of broad, sharp septa which are extended parallel with the
plane of symmetry of the shell and possess a sickle shape (hyncho-
nellae falciferae). (Rothpeltz, pl. 11, fig. 19.)
3. Septifer type—There can be, however, such sickle-shaped crura
so broad they make contact with the edge directed toward the small
[brachial] valve, are grown with it and consequently appear like actual
septa extending from the shell (Athynchonellae septiferae). (PI. 8,
figs. 46-48.)
Thirty-four groups or “Sippe’” of rhynchonelloids were recognized
by Rothpletz but the interior details of 19 of them were unknown at
this time. Of the remaining 15 Sippe, 3 belong to the falcifer group
(Trilobita, Lacunosa, and Varians), 2 belong to the septifer type
(Inversa and Trigona), and 10 are placed in the radulifer group
(Amalthei, Variabilis, Concinna, Plicatissima, Tetraédra, Inconstans,
Difformis, Plicatella, Psittacea, and Spinosa). Some of these Sippe
have been made into genera but generally little relationship exists
between the interior details of many of the species placed in each
group.
Wisniewska (1932, p. 6), in her fine work on rhynchonelloids from
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
the Jurassic of Poland, more clearly defines these types and adds a
fourth, as follows:
1. Radulifer type—Crura narrow, recurved toward the ventral
valve, widening gradually toward their extremity. This type, charac-
terizing the genera Septaliphoria, Rhynchonella, and Cyclothyris, was
given the name “radulifer” by Rothpletz.
2. Falcifer type—Crura with a large suspended crural plate, touch-
ing the bottom of the valve only near its summit. This is the “falcifer”’
type of Rothpletz characterizing the genus Lacunosella.
3. Septifer type—Crura short with the crural plates supported at
the bottom of the valve and extending for about one-third the valve
length. This is the “septifer” type of Rothpletz affirmed by us only
in the genus Septocrurella.
4. Arcuifer type-——Crura with large bases separated from each
other and curved so as to turn their concave sides toward the middle,
the extremity turned toward the ventral valve and terminated by a sort
of small crural plate in the form of a hammer. This type of crura,
seen in the genus Monticlarella, may be called “arcuifer.”
Muir-Wood (1934, p. 526; 1936, p. 14) added a fifth type as a
result of her work on Mesozoic brachiopods:
Calcarifer crura—‘‘... The crura consist of two flattened,
curved, posteriorly concave laminae which project from the hinge-
plate into the cavity of the pedicle valve. These laminae each unite
with a second curved lamina which appears to be suspended from it
and projects dorsally like a spur. A ventral extension of the second
lamina terminates in a hook-shaped process, the apex of which is
directed medianly.” Kallirhynchia and Rhynchonelloidella possess
this type.
Among the Tertiary rhynchonelloids considered herein five types
of crura are distinguishable, three of which have been identified
among Mesozoic genera and have been named. Two types have not
been named or described among the Mesozoic genera.
Of the three named types Hemithyris belongs to the group having
radulifer crura. These are long, slender, and curved but have a hori-
zontally flattened, bluntly pointed distal extremity. The Hemuithyris
crus is strengthened by a narrow ridge on the anterior side. The
radulifer type of crus is not common among Recent and modern
genera.
The second type of crus known in the Mesozoic and present among
modern and Tertiary genera is that characteristic of the Basiliolidae
and named falcifer type. The Basiliolidae are all characterized by
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 9
having broad-bladed, gently curved crura that are convex outward and
gently concave inward. This crus is generally attached to the hinge
plate by its convex side and may or may not be separated from the
socket ridge by outer hinge plates.
The third type of named crus is that characterizing the Erym-
nariidae and called septifer type. This is an extremely rare type of
crura known in a few genera only.
A fourth type of crus is recognized in the Cryptoporidae. The
crura are long and slender and appear to be continuous with the distal
end of the socket ridge. The distal extremity of the crus is com-
monly flattened, expanded, and serrate or digitate, some examples
suggesting a tiny hand with outspread fingers. The name “manicu-
lifer” is proposed for this type.
The fifth type of crus in the modern and Tertiary genera is gen-
erally shorter than the others, laterally compressed, somewhat flat in
section and attached to the hinge plate or socket ridge so that the
short direction is nearly vertical or slightly oblique. This type is best
seen in Frieleia, but Grammetaria, Hispanirhynchia, and Compso-
thyris have similar crura. This type is here designated as “spinulifer.”
Median ridges, median septa and camerae.—A conspicuous feature
of many rhynchonelloid stocks is the median septum. Some groups
however, such as the Basiliolinae, are devoid of septa in the brachial
valve. The most conspicuous septum in any modern rhynchonelloid
is that of Cryptopora in which, although short, it is so elevated that
it almost touches the inner surface of the opposite valve. The septum
of Frieleia is also considerably elevated.
In the descriptions below a distinction is made between median
septa and median ridges. The term septum is reserved for the thin
blades, like those of Cryptopora or Frieleia that stand boldly and
abruptly above the inner valve surface. These are in contrast to the
ridges such as that of Aetheia, which is low, short, and stout, and that
of Aphelesia which is low, long, and slender. Dorsally aseptate rhyn-
chonelloids are commonly provided with low and inconspicuous median
ridges, some in the form of a myophragm but others buttressing the
cardinalia.
In a few genera of Recent and Tertiary, Frieleia for example, the
median septum joins folds from the inside of the crural bases to form
a small chamber at the posterior. In some instances the chamber re-
mains open but it is frequently closed by deposit of shell material on
its inner walls to form a thick apical callosity. All degrees occur in
Frieleia from the open chamber to the solid callosity between the
Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
hinge plates. These do not seem to constitute a septalium in the true
sense of the word as defined by Leidhold (1928, p. 11) who says that
the median septum divides to produce the chamber. Wisniewska
(1932, p. 6), on the other hand, states that the septalium of the Meso-
zoic rhynchonelloids is formed by internal inflection of the hinge plate
to meet the median septum. This seems to be the method of formation
of this structure in Frieleia rather than division of the median septum.
The method suggested by Wisniewska seems certainly to be the
case in Septaliphoria in which it is possible in some specimens to see
the median septum between the lateral walls of the apical chamber.
The specimen illustrated (pl. 21, C, fig. 6) shows the plates converging
to the median septum and bounding a small chamber. In other speci-
mens the plates bounding the chamber meet the floor of the valve
rather than the median septum (see Wisniewska, 1932, p. 26, fig. 6).
In Camarotoecha (pl. 4, D, figs. 6-8) the entire structure seems to
be different from the Jurassic forms and strongly reminiscent of the
orthoids. The sides of the chamber buttress the crura which can be
seen buried in excess shell tissue surrounding the plates (Kozlowski,
1920, p. 146, fig. 43, A). Division into hinge plates is difficult. The
structures of the modern and Tertiary forms with camera seem more
like the Mesozoic species than like the Paleozoic.
EXTERIOR CHARACTERS
It is usually difficult to evaluate the generic characters of the ex-
terior of brachiopods and all workers are not agreed on this evalua-
tion. It is, however, quite clear that ornamentation and folding pat-
terns are generic in character.
Ornamentation.—Buckman (1917) and Rothpletz (1886), who
made attempts at the classification of rhynchonelloid brachiopods,
mostly used the exterior to make genera or species groups which
might ultimately become genera. Both of these classifications fail be-
cause ornamentation and folding are repetitious in many unrelated
stocks. Buckman attempted to make his genera on the basis of a
scheme of ornamentation development: those that are smooth and
then develop costae, those that are capillate and develop costae and
ornate or spinose forms. These characters were combined with shell
outline and anterior folding. Buckman, however, failed to determine
the characters of the cardinalia.
Rothpletz (1886) arranged many rhynchonelloid species into groups
or “Sippe” having similar external characters. Although he determined
the nature of the crura of some species he did not reveal the details
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER Tsk
of all of them. Consequently he placed many species together which
are utterly unlike internally.
In modern and Tertiary rhynchonelloids a smooth exterior is com-
mon, a capillate or costellate exterior is also fairly common, but a
strongly plicate exterior is unusual, occurring in only a few genera.
Rhytirhynchia is similar to Basiliola but differs exteriorly by its an-
terior costation. Notosaria and Tegulorhynchia are the only com-
pletely strongly costate modern rhynchonelloids. Rhynchonella grayi
Woodward, the true generic affinities of which are with the Eocene
genus Eohemithyris, is partially costate but not so strongly costate as
the Patagonian genus Plicirhynchia,
Folding—The anterior commissure of most rhynchonelloids is
uniplicate but some of them are sulcate or even more complicated. The
production of a fold is thought to be related to the feeding habits of
the brachiopod, the median fold helping to channel the excurrent
stream at midvalve.
Sulcation, brachial valve with sulcus, pedicle valve with fold, is not
a common feature of the brachiopods but crops up again and again
in many unrelated stocks, producing confusing heterochronous and
isochronus homeomorphs. Neorhynchia is the only known Recent
sulcate rhynchonelloid, but another modern deep-sea form, Abys-
sothyris, is a terebratuloid having a shape identical to that of Neorhyn-
chia. If it were not for the punctae of Abyssothyris it would be almost
impossible to tell the two genera apart on exterior characters alone.
Rhynchonelloids of almost identical form to Neorhynchia are known
from all the periods of the Mesozoic era and from the Paleozoic back
at least as far as the Devonian. It is difficult to suggest any reason
for the reversal of folding from the usual uniplicate condition because
the two types must have functioned in the same manner. It is a com-
mon feature of the young brachiopod to have a more robust pedicle
valve more or less prominently folded in the ventral direction and with
a somewhat sulcate brachial valve. Perhaps sulcation is merely a re-
tention of youthful shell characters into the adult stages.
Several of the modern and Tertiary brachiopods have rectimarginate
anterior commissures. This, too, is a youthful character. Buckman
emphasized the folding of brachiopods and used this feature as a
major part of his classification. It is evident, however, from the above
remarks and known brachiopod history that folding is of value only as
a generic character. When combined with ornamentation features as
Buckman advocated, valid genera have been established. These how-
ever can only be placed in their proper families by determination of
their beak and cardinalia characters.
I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
RHYNCHONELLOID CLASSIFICATION
Very few comprehensive works have ever been written on the
rhynchonelloids. The first to have attempted a detailed classification of
these difficult shells was Rothpletz (1886), who divided them into
seven species groups and numerous subdivisions of these groups based
on exterior details. Although Rothpletz carefully defined the interior
of some of the Jurassic rhynchonelloids, using strictly modern meth-
ods, he did not bring the information into his classification. Some of
Rothpletz’s groups and subdivisions bring together species now
known to have nothing in common except exterior form. Besides over-
looking details of anatomy in his classification, Rothpletz also com-
posed unlikely assemblages from various parts of the geological
column.
Bittner’s (1890, 1892) great works on the Triassic brachiopods
defined in exquisite detail some of the rare and unusual spiriferoids
but neglected interior features of the rhynchonelloids except for a
few forms. The Triassic rhynchonelloids are a prolific lot and will
amply repay in new information a modern, detailed study. Bittner
added only a few genera but left many for the future. He, too, was
content to work chiefly on exterior details even though the method of
serial sectioning was well known and even used by him in some cases.
Hall and Clarke (1894) described many rhynchonelloid genera but
never made a serious attempt at classification. They did, however,
show the importance of internal characters and described these details
in many Paleozoic genera.
Weller (1910) used the serial-section method to make known the
details of many rhynchonelloid genera, but he did not go beyond
genus making. His work was important for showing that a combina-
tion of interior and exterior details is necessary for the correct elucida-
tion of rhynchonelloid descent. He indicated several genera that had
interior details like those of Camarotoechia but were quite unlike that
genus in exterior details. He had, therefore, no other choice than to
create new genera for them.
The greatest strides in the understanding of the rhynchonelloids
came in Buckman’s classic work on the Jurassic brachiopods of Burma
and Great Britain. Buckman also proliferated genera more than any-
one before him. In his work he relied almost wholly on exterior
characters, first on the kind of ornamentation and then on the type
of folding of the anterior commissure. These features were supple-
mented by some details of the interior such as the septa and the
muscle scars which were exhibited by a process of calcining the shells.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 13
Unfortunately most of the interior characters developed by Buckman
are of secondary importance compared with the cardinalia, which he
did not develop. He made no effort to learn the details of these fea-
tures by serial sectioning as since used by many British authors.
A necessary task of the future is to determine the cardinalia
characters of the Buckman genera and then to sort these genera into
families based on these characters. It seems likely that most of Buck-
man’s genera will prove useful because the exterior features of most
of them are distinctive. It will probably be found that some of these
ornamentation patterns will be repeated in combination with various
cardinalia patterns. The result will be a further, but necessary, pro-
liferation of genera, but a considerably better understanding of
Jurassic genera will be forthcoming. In this connection the writer has
determined the interior features of a number of Jurassic rhynchonel-
loids by etching the shell from limestone. These show the cardinalia of
Septaliphoria in combination with exterior characters indicating more
than one genus. Another interesting feature is variations of the sep-
taliphore interior that promise to be of great interest. Silicified Meso-
zoic rhynchonelloids occur in South America, Israel, Africa, and else-
where, and should be sought and prepared because they offer the best
opportunity for understanding interior details.
Leidhold (1921, 1922) wrote several papers elucidating the interior
of the rhynchonelloid shell. His work in 1921 defined the interior of
Septaliphoria and two other genera, but he did not make any strides in
classification of these brachiopods.
Schuchert, in Schuchert and LeVene (1929), took a stride forward
in rhynchonelloid classification when he separated the Camarotoechii-
dae from the Rhynchonellidae. Unfortunately, however, he did not
define the characters of the family. Even with this family divided into
three subfamilies Schuchert has many forms of unlike structure clas-
sified together. He states that the “Classification into families [of the
Rhynchonellacea= Rhynchonelloidea] is not yet satisfactory.” No at-
tempt was made to subdivide the Mesozoic and later forms except to
group them according to geological periods, and to recognize the
Dimerellidae of Buckman.
Thomson (1927, pp. 145-164) discussed Recent and Tertiary rhyn-
chonelloids in detail and made many interesting observations on them.
He also assigned the genera to two families. The peculiar and primi-
tive Cryptopora was assigned to the Dimerellidae where it seems very
unhappy and the rest of the genera were put in the Rhynchonellidae
where they are likewise out of place.
I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Pettit (1950, 1954), in revising the Cretaceous rhynchonelloids of
Great Britain, described some details of their interior but his work is
disappointing in this respect. In some instances the interior was de-
scribed by serial sections when direct preparations should have been
less time consuming, easier to make, and far better understood. Owen
(1955, p- 369) recently described a method for making serial sections
of brachiopods preserved in chalk. In the writer’s opinion the serial-
section method should be a last resort when all others fail. Chalk
brachiopods are easy to prepare directly. The serial-section method is
destructive of material and the interior characters may be obscured by
old age growth and inner injury. Sectioning is far less satisfactory
than direct observation unless it is the only course that can be taken.
Rzhonsnitzkaia (1956, p. 125) presented an abstract and outline
of a new classification of the order Rhynchonellida of Moore 1952.
This classification is more elaborate and complete than any hitherto
published but the families are not defined and the characters on which
they are based are not stated. Family splitting of the rhynchonelloids
has been so long needed that the characters of some of Rzhons-
nitskaia’s new families and subfamilies are quite obvious. For a few,
however, they are not so clear. Among the younger rhynchonelloids
the only new category introduced is the Hemithyrinae, which will
probably receive general acceptance, and is here elevated to family
status.
FAMILY AND GENERIC ARRANGEMENT AND CHARACTERS
This brief survey of rhynchonelloid classification indicates that
fundamental work is still to be done on the group. These shells are
difficult, but they can be made to yield good interiors by simple methods
of manual preparation or by serial sectioning. The writer attempts
below to group into families the Recent and Tertiary rhynchonelloids
on the basis of their interior details combined with features of the ex-
terior. The cardinalia characters in their over-all pattern are, in ac-
cordance with his work on the orthoids, triplesioids, pentameroids, and
several other groups, regarded as of family rank. Some details of the
cardinalia are generic but mostly they help to define families. The
generic characters are found in minor interior details combined with
ornamentation features and beak characters of the pedicle valve. This
is well shown by the number of genera in the Paleozoic that have the
internal characters of Camarotoechia but vary in external form and
ornamentation: Paraphorynchus, Camarotoechia, and Pugnoides are
examples. The principle is well exemplified by the families described
below.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 15
Family CryptoporipAE Muir-Wood, 1955.—Primitive rhyncho-
nelloidea having a large deltoid foramen slightly restricted by elongate,
triangular, elevated deltidial plates; crura long, maniculifer, continu-
ous with the socket ridges ; median septum strongly elevated ; cardinal
process a lobate thickening between the socket ridges; one pair of
nephridia.
Genera: Cryptopora and Manmia.
Cryptopora.—Triangular in outline, exterior smooth.
Mannia—Exterior spinose, with spoon-shaped expansion of median septum
of brachial valve (validity of genus in question, see text, p. 22).
BASILIOLIDAE Cooper, new family—Smooth or semicostate rhyn-
chonelloids having conjunct deltidial plates and small auriculate
foramen ; pedicle valve with dental plates varying from nearly obsolete
to strong, pedicle collar well developed; brachial valve with broad
falcifer crura supported by outer hinge plates or the socket ridge; no
median septum but a median ridge may be present.
Subfamilies: BASILIOLINAE, APHELESIINAE, and AETHEIINAE.
BASILIOLINAE Cooper, new subfamily.—Basiliolidae with crura at-
tached to broad outer hinge plates; no median septum.
Genera: Basiliola, Eohemithyris, Neorhynchia, Rhytirhyncha,
Probolarina, and Streptaria.
Basiliola—Brachial valve much deeper than the pedicle valve; pedicle collar
elaborate; exterior smooth; anterior commissure strongly uniplicate.
Eohemithyris—Valves subequal in depth, smooth to semicostate; anterior
commissure uniplicate.
Neorhynchia.—Deltidial plates disjunct; exterior smooth but anterior commis-
sure sulcate; incipient inner hinge plates.
Rhytirhynchia—Outline like that of Basiliola but anteriorly costate; anterior
commissure sulciplicate.
Probolarina—Beak elongated; deltidial plates well exposed; anterior half
strongly costate; elaborate pedicle collar; anterior commissure uniplicate.
Streptaria—Exterior smooth to semicostate; anterior with sides twisted;
foramen with reflected rim; dental plates reduced; pedicle collar poorly
developed.
APHELESIINAE Cooper, new subfamily.—Basiliolidae with crura at-
tached directly to side of socket ridge; thick median ridge present in
brachial valve.
Genus: Aphelesia.
Aphelesia—Smooth to anteriorly costate; anterior commissure uniplicate.
AETHEIINAE Cooper, new subfamily—Basiliolidae having a minute
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
foramen, concave deltidial plates, reduced to obsolete dental plates
and thick inner hinge plates.
Genera: Aetheia and Patagorhynchia,
Aetheia.—Elongate triangular in outline and exterior smooth.
Patagorhynchia.—Costellate and anteriorly imbricate.
Family HeMirHyrRIDwaAE Rzhonsnitzkaia, 1956 [proposed as a sub-
family].—Rhynchonelloidea with strong, slender, curved radulifer
crura attached to small outer hinge plates by their posterodorsal face
or to thick socket ridges; crura distally pointed and horizontally
flattened.
Genera: Hemithyris, Notosaria, Tegulorhynchia, and Plicirhyn-
chia.
Hemithyris——Beak long, surface striate to costellate; deltidial plates disjunct.
Notosaria——Beak short; exterior costate; nonimbricate; deltidial plates dis-
junct.
Tegulorhynchia.—Costellate to costate, strongly imbricate to spinose; deltidial
plates conjunct?; medium septum reaching apex.
Plicirhynchia—Long beak, posterior striate to costellate, anterior costate to
plicate; deltidial plates conjunct.
FRIELEIIDAE Cooper, new family.—Usually capillate to costellate
valves with triangular outline, strong dental plates, and brachial valve
with short, straight, laterally compressed spinulifer crura supported
by short plates that unite with the median ridge or septum to form a
small chamber.
Genera: Frieleia, Compsothyris, Grammetaria.
Frieleia—Elongate shells with the pedicle valve having the greater depth
and the brachial valve with a high median septum; anterior commissure recti-
marginate to ligate; deltidial plates disjunct; inner hinge plates extravagantly
developed.
Compsothyris—Roundly triangular in outline; valves of subequal depth;
median septum only moderately elevated and deltidial plates disjunct; anterior
commissure gently uniplicate; inner hinge plates incipiently developed.
Grammetaria.—Elongate, costellate shells with rectimarginate anterior com-
missure, low, thick median septum, and conjunct, strongly auriculate deltidial
plates; inner hinge plates incipient.
HISPANIRHYNCHIIDAE Cooper, new family.—Triangular, capillate
rhynchonelloidea having a weak median ridge or no median ridge in
the brachial valve; crura spinulifer ; anterior commissure rectimargi-
nate to ligate.
Genera: Hispanirhynchia and Sphenarina.
Hispanirhynchia.—Deltidial plates disjunct and median ridge of brachial valve
low and thick; inequivalve, the pedicle valve being the deeper; inner hinge
plates strongly developed.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER a7,
Sphenarina.—Deltidial plates conjunct; subequivalve; brachial valve with no
median ridge; slight or no development of inner hinge plates.
ERYMNARIIDAE Cooper, new family.—Rhynchonelloidea having sep-
tifer crura.
Genus: Erymnaria.,
Erymnaria.—Exterior smooth, inequivalve; anterior commissure uniplicate
to twisted ; deltidial plates conjunct; the brachial valve having the greater depth.
Family CrypTorormpAE Muir-Wood 1955
Genus CRYPTOPORA Jeffreys, 1869
Plates't2; A, By 2A, 5,°C, 2%; Di text figure TA
Cryptopora Jeffreys, Nature, vol. 1, p. 136, 1869 (inadequately described, not
figured) ; Thomson, Geol. Mag., n. s., dec. 6, vol. 2, pp. 387, 388, 392, 1915;
Thomson, New Zealand Board Sci. Art, Manual 7, p. 146, 1927.
Atretia Jeffreys, Proc. Roy. Soc., vol. 18, No. 121, p. 421, 1870 (inadequately
described, not figured) ; Ann. Mag. Nat. Hist., ser. 4, vol. 18, p. 250, 1876;
Proc. Zool. Soc. London, p. 412, 1878; Davidson, Trans. Linnaean Soc., ser. 2,
vol. 4, pt. 2, p. 173, 1887. Not Atretium Cope, 1861.
Neatretia Fischer and Oehlert, Exped. Sci. Travailleur et Talisman, p. 122, 1801.
Mannia Davidson, Geol. Mag., dec. 2, vol. 1, No. 4, p. 156, 1874(b).
Small, translucent to transparent, subtriangular in outline with the
greatest shell width anterior to the middle; subequivalve; anterior
commissure rectimarginate to broadly sulcate; surface smooth. Beak
of the pedicle valve moderately long, pointed, nearly straight ; foramen
large, incomplete, not restricted ; deltidial plates rudimentary, forming
a ridge on the delthyrial edge, auriculate to alate. Shell fiber mosaic
coarse.
Pedicle valve interior with small noncorrugated teeth; apex with
thickened plate elevated above valve floor ; teeth supported by strong,
divergent dental plates. Muscle scars not well impressed.
Brachial valve interior with small, smooth or roughened sockets
bounded by high socket ridges; socket ridge overlying crural base;
crura of maniculifer type, long and slender, slightly curved, expanded
distally and commonly with the distal edge deeply digitate. Cardinal
process small, bilobed and transverse. Median septum high anteriorly
but sloping steeply to the valve floor posteriorly and disappearing an-
terior to the apex; anterior face of septum steep. Adductor scars
lightly impressed. One pair of metanephridia in the fleshy body of
the animal.
Type species (by monotypy).—Atretia gnomon Jeffreys. Ann.
Mag. Nat. Hist., ser. 4, vol. 18, p. 251, 1876.
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Comparison.—This is a very distinctive little brachiopod and can-
not be confused with any other modern form. It is characterized by
a yellowish to white and shiny, transparent to translucent shell having
peculiar deltidial plates, long, slender maniculifer crura and a short,
high, slender median septum. The only described genus similar to it
is Mannia which is said to differ in the form of the septum and the
possible presence of spines on the exterior. (See Mannia.)
Cryptopora has frequently been compared with the Triassic genus
Dimerella but the two are actually very different. The median septum
of the brachial valve of Dimerella has a different form, the deltidial
region of the Triassic shell is different, and the dental plates are much
less strongly developed than those of the modern genus. The exterior
of the two genera is also quite different, the Triassic shell being wide
with a fairly wide hinge and costellate exterior. The modern genus on
the other hand is narrowly triangular and smooth.
Geological horizon.—Cryptopora was recorded from the Eocene
(Salt Mountain formation) by Toulmin (1940, p. 229). It is also
known from the Oligocene of Cuba and Miocene of Europe (see below
and Mannia).
Thomson (1927, p. 147) cites Rhynchonella discites Dreger from
Vienna, R. lovisati Dreger from Sardinia, and Hemithyris parvillima
Sacco from Italy, all from the Miocene, as possible fossil examples of
Cryptopora. Thomson also cites Terebratella acutirostra Chapman, a
possible synonym of C. brageri from the Miocene of Victoria, Aus-
tralia, as another fossil species. The geological range is therefore
from Eocene to Recent.
Distribution.—In the North Sea and North Atlantic south to Cuba
in waters ranging from 75 to 2,200 fathoms. In the Southern Hemi-
sphere it occurs off New South Wales in 17 to 100 fathoms, and on
southern Agulhas Bank, South Africa, in 500 to 565 meters or about
275 fathoms.
Assigned species——The one Eocene form known was not named
but species are known from the Miocene and in modern waters :
Atretia gnomon Jeffreys, Recent, North Atlantic.
A. brazeri Davidson, Recent, east Australia.
Cryptopora boettgeri Helmcke, Recent, southern Agulhas Bank, Africa.
C. rectimarginata Cooper, Recent, East coast Florida, Cuba.
? Rhynchonella discites Dreger, Miocene, Vienna.
? R. lovisati Dreger, Miocene, Sardinia.
? Terebratella acutirostra Chapman, Miocene, Australia.
? Hemithyris parvillima Sacco, Miocene, Italy.
Mannia nysti Davidson, Miocene, Belgium.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 19
Discussion —This genus differs strongly from other modern and
Tertiary rhynchonelloids except Mannia which is discussed below.
The form of the median septum and crura are unique and the deltidial
plates are formed differently from those of the other rhynchonelloid
genera.
The deltidial plates of Cryptopora are disjunct throughout life. The
foramen is not greatly restricted by these plates because they usually
grow at a high angle to the edge of the delthyrium rather than being
a continuation of it. The foramen is thus incomplete and not circular
but is deltoid and roughly parallel to the delthyrial margins.
The deltidial plates are small and elongate triangular, forming on
the delthyrial margin at a high angle and commonly reflected laterally
to overhang the dorsolateral slopes of the beak. In Cryptopora rec-
timarginata Cooper, new species, the deltidial plates are strongly
alate, the projections being located near the posterior of the plate and
narrowly rounded, bluntly pointed or rarely serrated. In the older
shells the blunt points disappear.
The apex of the pedicle valve is occupied by a small elevated tri-
angular plate against which the pedicle rests. A plate similar to this
appears in other genera, such as Hemithyris. Aside from the strong
dental plates the pedicle valve reveals no other structures. The shell
is so thin that muscle scars cannot be seen easily. A suggestion of a
low myophragm appears in some specimens.
The cardinalia of the brachial valve are unusual. The socket plates
are small and delicate, appear to be continuous with the crural bases
and lie above or posterior to them. The socket plates are attached
directly to the shell wall and buttressed for a short distance by a small
supporting plate. The crura are long and welded with the crural bases
and supporting plates in such a way that they appear to make one
plate. The main part of the crura are strong but slender and are
bowed outward to a considerable degree in older specimens, less so in
the young ones. The distal extremity is flattened laterally and the free
end serrated or frayed into a number of small prongs. The whole
suggests a tiny hand with outstretched spreading fingers or a flattened
fist.
The diductor muscles were attached to a bilobed boss or cardinal
process at the posterior apical part. This is somewhat thickened in old
shells, the thickening spreading to the base of the crura and uniting
with an extension of the median septum.
The most conspicuous feature of the brachial valve is the median
septum. It is highest at about midvalve but descends rapidly posteri-
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
orly to disappear before reaching the apex in young shells. In old
specimens a low extension of the septum extends to the apex where
it unites with a thickening from the cardinal process. The septum
thus makes a narrow wedge extending ventrally almost to the inner
wall of the pedicle valve.
I have not observed the radial striae reported by Dall (1920, p. 293)
in young shells.
The fossil species assigned doubtfully to Cryptopora may be the
young of other species. The gaping foramen and rudimentary deltidial
plates are suggestive of young rhynchonelloids. Meznerics (1943,
p. 23) points out that Sacco believed H. parvillima to be a juvenile of
H. de buchii= Streptaria buchi.
I have examined a specimen of Manma nysti Davidson from the
Miocene of Belgium. As explained in the discussion under Mannia,
this specimen has the features of Cryptopora but does not conform
completely with the description given by Davidson. The description
of this genus is evidently inaccurate and the two genera are exact
synonyms (see discussion under Mannia).
CRYPTOPORA RECTIMARGINATA Cooper, new species
Plates 1, B, 2, A
Atretia gnomon Dall (not Jeffreys), Proc. U. S. Nat. Mus., vol. 57, p. 293,
1920 (U.S.N.M. Cat. Nos. 83131, 274138, 274130, 94367, 3360804).
Shell small, translucent to white, subtriangular in outline, with the
greatest width anterior to the middle; sides gently rounded; anterior
margin strongly rounded; valves subequal in depth; anterior com-
missure rectimarginate ; surface smooth.
Pedicle valve slightly deeper than the brachial valve; lateral pro-
file gently convex, most convex in the posterior third and flattened
in the anterior third; anterior profile broadly convex, slightly more
convex than the brachial valve in this profile. Beak pointed, forming
an angle of about 85°, suberect ; deltidial plates erect, thickened along
their distal margin, commonly extravagantly auriculate, the auricula-
tions directed laterally.
Pedicle valve interior with thick apical plate well elevated above the
valve floor ; teeth small, wide ; dental plates stout, slightly divergent an-
teriorly, approximately vertical to the valve floor. Muscle field anterior
to delthyrial cavity.
Brachial valve with gently convex lateral profile, the maximum con-
vexity located just anterior to the umbo and posterior to the middle;
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 21
anterior profile broadly and gently convex; posterolateral slopes
moderately steep; anterior slope long and flattened.
Brachial valve interior with long, approximately parallel crura;
socket ridges stout, grown together with the cardinal process which
forms a thickening between the socket ridges at the apex; median
septum stout, short anteroposteriorly, narrow in profile; adductor
scars deeply sunk and forming an elongate track on each side of the
median septum.
MEASUREMENTS IN MILLIMETERS
Brachial
Length length Width Thickness
EVOLOLY [IE a Sere nicjeiee ey sie atsisns:brewiataie.eveue rae’ 5.2 4.4 4.4 1.8
Paratype: Ui SANGIN: 274TA Sd s wie cis cise 5.3 4.7 4.6 1.6
Types.—Holotype, U.S.N.M. 274143a; figured paratypes, U.S.
N.M. 274143b, c, d, 274168a, 336895a, and 336896a.
Horizon and locality—Recent, Eolis Station 340, at 209 fathoms
off Fowey Light; several other stations off Fowey Light at depths
ranging from 85 to 205 fathoms ; off Sand Key at 75 to 120 fathoms ;
off Sambo Reef at 135 fathoms; off Western Dry Docks, at 80 and go
fathoms; and off Key West at 110 fathoms; all off Florida.
Comparisons.—This species is characterized by its narrowly len-
ticular profile, the rectimarginate anterior commissure, auriculate
deltidial plates, thick cardinalia and short, stout median septum.
Cryptopora rectimarginata differs from C. gnomon in several re-
spects. The latter is quite strongly sulcate, whereas the Florida species
is rectimarginate ; C. gnomon is a thicker shell than C. rectimarginata
and has a longer median septum and does not have the auriculations
on the deltidial plates. Helmcke’s species C. boettgeri likewise does
not have the auriculations on the deltidial plates and also has a longer
median septum than C. rectimarginata. The Australian C. brazieri
differs from C. rectimarginata in having a flattened brachial valve, no
auriculations on the deltidial plates, and a longer and more delicate
median septum.
A specimen from the Oligocene (Cojinar formation), from Sagua
la Grande in Las Villas Province, Cuba (U.S.N.M. 459424a) is
strongly suggestive of C. rectimarginata because it has auriculate
deltidial plates and the form and profile of the Florida species.
Cryptopora rectimarginata appears to be a shallow-water species
ranging in depth from 75 to 209 fathoms. Cryptopora gnomon, on the
other hand, is a deeper-water form. Depth ranges given for specimens
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
in the U. S. National Museum collection are from 650 to 2,200
fathoms. The Australian species is a shallow-water form found in 100
fathoms. Cryptopora boettgeri from off Agulhas Bank, South Africa,
is from deeper water, 500 meters (275 fathoms).
CRYPTOPORA GNOMON Jefireys
Plates 5; C, 21, D
Crytopora gnomon Jeffreys, Nature, vol. 1, Dec. 2, p. 136, 1869.
Atretia gnomon Jeffreys, Proc. Roy. Soc., vol. 18, No. 121, p. 421, 1870; David-
son, Trans, Linnaean Soc., ser. 2, vol. 4, pt. 2, p. 173, pl. 25, figs. 6-13, 1887.
Neatretia gnomon (Jeffreys) Fischer and Oehlert, Exped. Sci. Travailleur et
Talisman, p. 122, figs. I1a-c, 1891.
This is a deep-water form that differs markedly from C. rectimar-
ginata in its nonalate deltidial plates, more strongly folded anterior
commissure, and other details. Figures are introduced for comparison
with the Florida species.
Types.—Figured specimens U.S.N.M. 94367, 44911, c, d.
Horizon and locality—Recent, 780 fathoms, off Cuba; 1,525
fathoms at U. S. Fish Commission Station 2221, south of Marthas
Vineyard, Mass.
Genus MANNIA Davidson, 1874
Plates 1, A, 21, F; text figure 1, B
Mannia Dewalque, Prodrome d’une Description Géol. Belg., p. 432, 1868 (not
described or figured); Davidson, Geol. Mag., dec. 2, vol. 1, No. 4, p. 156
(extract p. 6), 1874(b) ; Thomson, New Zealand Board Sci. Art, Manual 7,
p. 2906, 1927.
This interesting little brachiopod [type species (by monotypy),
Mannia nystt Davidson, 1874] was described by Davidson who indi-
cates that some points of its anatomy are still to be learned. The
affinities of Mannia, as well as its anatomy, are not clearly under-
stood because some workers have regarded it as a rhynchonelloid but
one of the best informed students of brachiopods, J. Allan Thomson,
thought that it is a terebratuloid. Its rhynchonelloid affinities, how-
ever, now seem clear and unquestionable. Because of the rarity of this
species little is known of it but restudy of a good specimen and photo-
graphs of the types now make its features clear.
According to Davidson’s description, Mannia is similar to Crypto-
pora externally as well as internally. The beak region is elongated
and pointed and the pedicle opening is elongate triangular. The pedicle
opening is bordered by attenuated, triangular deltidial plates as in
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 23
Cryptopora. An external difference between the two, on the other
hand, is suggested by Davidson’s report on the exterior of Mannia of
“concentric scaly lines of growth, from which scattered adpressed
spinules seem to arise.” The specimen figured by Davidson is very
small. Its measurements are given in lines: length 2 lines=4 mm.,
width 14 lines=3 mm.
The pedicle valve interior is not well known, but Davidson (1874b,
p. 157) speaks of a “narrow vertical plate” dividing the larger por-
tion of the beak into two parts. However, no indication of a median
A B
Fic. 1.—Partial side views of the brachial valves of A, Cryptopora, ca. X 8,
and B, Mannia, ca. & 16, showing maniculifer crura.
septum can be seen in the beak region in Davidson’s figure 10a, plate 7.
Inside the brachial valve the cardinal process is medial and the crura
are long but the socket plates are small. In one figure the crura are
convergent ; in the other they are divergent, but in both they are similar
to the crura of Cryptopora. Davidson reports them as being broken,
which is probably the reason why they are illustrated as not flattened
and expanded distally.
The median septum of the brachial valve is illustrated by Davidson
as like that of Cryptopora in being short and very high. Unlike Cryp-
topora, however, its distal extremity is embellished by “two small tri-
angular plates united posteriorly, separate and angular anteriorly.”
These form a small spoonlike trough which was interpreted by Thom-
son (1927, p. 297) as a terebratelliform structure. Thomson goes on
to state that the shell of Mannia will ultimately prove to be punctate,
but Davidson said emphatically that it is impunctate and referred it
to the Rhynchonellidae.
24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
I am indebted to Dr. A. Vandercammen and the officials of the
Institute Royal des Sciences Naturelles de Belgique for a very fine
specimen of Mannia nysti from the Upper Miocene (Diestien-Sables
de Deurne) from Wommelghem, just east of Antwerp, Belgium.
When delivered to me the specimen had both valves attached. Its
exterior was photographed and then the valves teased apart with only
a small fracturing of the anterior margin of the pedicle valve (see
pl. 1, A). Inasmuch as this specimen is essentially a topotype and
from the only horizon from which the species is known, it gives an
authentic check on Davidson’s description and figures.
My obligation to Dr. Vandercammen is still greater because he also
sent me notes on Davidson’s type lot and an enlarged picture of his
types. These and the fine little specimen now make it possible to cor-
rect Davidson’s description and to refigure the genus with unretouched
pictures. The combined result of this reevaluation is to demonstrate
beyond reasonable doubt that Manmia is Cryptopora.
The specimen from Wommelghem is small, having a length of 2.5
mm. and a width of 1.8 mm. The outline is elongate triangular, the
beak sharply pointed and the deltidial plates strongly elevated. The
exterior appears to be completely smooth, without any trace of con-
centric, scaly lines or “adpressed spinules.” The anterior commissure
is rectimarginate. The interior shows a coarse mosaic of shell fibers,
a distinctly rhynchonelloid character.
The striking feature of the pedicle valve of this specimen is the
strong elevation of the deltidial plates and the large size of the apical
plate. The interior of the brachial valve is generically exactly like
that of modern Cryptopora but specific differences may be readily
noted, especially in the crura. These are bowed as in the modern
species but the distal expansion is greatly exaggerated in its size and
flatness. Furthermore the serrations on its distal extremity are nu-
merous and minute (text fig. 1, A).
Of features recorded by Davidson as characteristic of Mannia I
was unable to confirm the presence of a median septum in the apex of
the pedicle valve. The triangular plates on the distal extremity of the
median septum of the brachial valve were not confirmed and the
details of the exterior are not in accordance with Davidson’s descrip-
tion and figures.
The information and data furnished me by Dr. Vandercammen in-
cluded a photograph of Davidson’s type specimens and the label ac-
companying them. Five specimens are shown, one complete specimen
exactly like that from Wommelghem sent to the U. S. National
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 25
Museum, three brachial valve interiors, and one pedicle valve in-
terior. The outside and inside of these specimens are exactly like
those of the specimen illustrated on plate 1, A, of this monograph
except in one instance. Examination of the pictures of the brachial
valve interiors under a magnifier failed to show any expansion of the
distal end of the median septum. A possible mixture of species has
occurred because the median septum of the largest of the three brachial
valves extends anterior to midvalve and nearly to the apex in the
opposite direction, is not abruptly elevated as typical, and seems to
have different crura. This appears to belong to some other genus
but it is difficult to say what it is. The other two are quite characteristic
of Cryptopora. The complete specimen is illustrated from the dorsal
side and shows the characteristic large foramen bordered by narrow
deltidial plates. I am unable to distinguish the details of the pedicle
valve interior from the picture.
Davidson was a keen observer but it is difficult to escape the conclu-
sion that his figures of Mannia are a misrepresentation of specimens
of Cryptopora. I have therefore placed Mannia in the synonymy of
Cryptopora.
BASILIOLIDAE Cooper, new family
BASILIOLINAE Cooper, new subfamily
Genus BASILIOLA Dall, 1908
Plates) 11,8} 12, 13,. By 14,°A, ©
Basiliola Dall, Bull. Mus. Comp. Zool., Harvard Coll., vol. 43, No. 6, p. 442,
1908; Thomson, New Zealand Board Sci. Art, Manual 7, p. 154, 1927; Jack-
son and Stiasny, Siboga-Exped., Monogr. 27, p. 10, 1937.
Basiola Thomson, Geol. Mag., n. s., dec. 6, vol. 2, p. 390, 1915 (Lapsus calami).
Neohemithyris Yabe and Hatai, Proc. Imp. Acad. Japan, vol. 10, No. 9, p. 587,
1934; Hatai, Sci. Rep. Téhoku Imp. Univ., ser. 2 (Geology), vol. 20, p. 210,
1940.
Outline elongated subpentagonal to rounded subpentagonal ; widest
at about midvalve; strongly inequivalve, the brachial valve being
greatly swollen but the pedicle valve gently convex ; anterior commis-
sure strongly uniplicate but the fold on the brachial valve low and in-
conspicuous; surface smooth. Beak of pedicle valve small, nearly
straight, short; foramen small, complete circular to elongate-elliptical,
submesothyrid; deltidial plates small, conjunct, moderately to elabo-
rately auriculate.
Pedicle valve interior with strong and complex pedicle collar ; teeth
small and corrugated, supported by short receding dental plates which
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
define shallow and narrow umbonal chambers. Muscle field short and
narrow, the diductors surrounding the large adductors and with small
adjustors in a posterolateral position. Pallial marks strongly im-
pressed, the vascula media extending from the anterolateral end of
the muscle field to converge anteriorly on the long tongue. Lateral
branches few.
Brachial valve interior with deep corrugated sockets bounded by
strong socket ridges; crura of falcifer type, moderately long crescentic
in section, scimitarlike and attached to the socket ridges by prominent
outer hinge plates. Inner hinge plates absent. Vascula media widely
divergent.
Type species (by original designation) —Hemuthyris beecheri Dall,
Proc. U. S. Nat. Mus., vol. 17, p. 717, pl. 31, figs. 1-4, 1895.
Comparison.—Basiliola is characterized by its smooth shell, elabo-
rate pedicle collar, conjunct and auriculate deltidial plates, small round
to longitudinally elliptical foramen and the broad outer hinge plates
of the cardinalia. Basiliola differs from Rhytirhynchia, which it other-
wise resembles, in lack of anterior costation. It differs from Aetheia
in the nature of the foramen and the lack of inner hinge plates in the
cardinalia. Aphelesia is similar externally to Basiliola and has a similar
foramen but its cardinalia are quite distinct in lacking outer hinge
plates. Basiliola differs from Probolarina by its smooth exterior.
Geological horizon.—Basiliola is known from Pliocene to Recent.
Distribution—The known Pliocene species of Basiliola are from
Okinawa. Yabe and Hatai (1935) identified one Okinawa form as
Neohemithyris lucida and identified its age as Pleistocene. Cooper
(1957) described specimens of this species from the same place as
new, and another, not named, in addition. Furthermore, the U. S.
Geological Survey now dates the beds producing these specimens as
late Pliocene. So far as known these are the only fossil basiliolas
known.
Basiliola occurs in modern seas around the Hawaiian Islands, Japan,
Fiji, Borneo, Malay Archipelago, the Celebes, and Philippine Islands.
Bathymetric range-—Each of the species assigned here to Bastliola
has a different bathymetric range and different temperature tolerance.
Basiliola beecheri ranges in depth from 143 fathoms down to 313
fathoms, and the temperature range is 43.8° F. to 60.8° F. Basiliola
pompholyx usually occupies deeper water, from about 150 fathoms
(275 meters, Jackson and Stiasny, 1937, p. 10) down to 1,105 fathoms,
and with a temperature tolerance between 43.3° F. and 52°F. Ba-
siliola elongata occurs in 24 fathoms but the temperature is not known.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 27
Another specimen, possibly the same species, is from 153 fathoms.
Basiliola lucida is from waters of 56 fathoms down to 122 fathoms
and the temperature range is from 51° F. to 63° F.
Assigned species ——The following species are assigned to Basiliola:
Hemithyris beecheri Dall, Recent, Hawaiian Islands.
Basiliola pompholyx Dall, Recent, Philippines.
Rhynchonella lucida Gould, Recent, Japan.
Basiliola nitida Cooper, Pliocene, Okinawa.
B. elongata Cooper, new species, Recent, Philippines.
Discussion.—Basiliola with its strongly unequal valves and small
foramen bounded by auriculate deltidial plates is usually easy to recog-
nize. The shells range from hyalescent when living to opaque in the
older or dead shells. The color ranges from pale yellow-brown to
brownish gray. The anterior commissure is usually strongly folded
in the dorsal direction, a long tongue from the pedicle valve fitting into
the deeply reentrant brachial valve. Although the anterior uniplica-
tion is strong, the fold on the brachial valve is not, as a rule, well
defined. Except for the uniplicate commissure the valves are not
otherwise plicated, nor do they have any radial markings.
Aside from the smooth shell and uniplication the only other distinc-
tive exterior feature of Basiliola is the beak. This is generally not
much elongated but is bluntly pointed. The foramen is usually small
circular, longitudinally oval, or elongated elliptical. The anterior side
of the foramen is usually bounded in all the species by a moderate
to elaborate flange or auriculation. In B. beechert and lucida this is
present but not as exceptionally developed as in B. pompholy.x.
The deltidial plates are conjunct and often so tightly joined as to
approximate a symphytium. The anterior margin of the deltidial
plates commonly rests on the umbo of the brachial valve. In old
shells the movement of the umbo against the anterior margin of the
deltidial plates leaves a smooth area. In some specimens an extension
grows anteriorly from the anterior margin of the deltidial plates along
the surface of the umbonal slope of the brachial valve. This usually
is part of the pedicle collar.
The chief character on which Dall based his genus is the pedicle
collar and which is elaborate in many specimens. It is best seen in
B. pompholyx (pl. 12, fig. 10) although it is well developed in the
other species. The collar is built as a plate from the anterior edge of
the deltidial plates as mentioned above and extends around the inside
of the apex. The collar in many specimens is clear of the valve floor
but in others shell substance has been added under the free antero-
ventral edge.
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
The muscle field of the pedicle valve is generally small in all species
but it is also deeply excavated into the shell. The individual scars are
usually strongly impressed. The diductor scars are small for such
large shells, are somewhat rectangular in form, and surround the
moderately large adductor patch. The adjustor scars are small and are
located just anterior to the front edge of the dental lamellae. Posterior
to the main part of the muscle field and within the delthyrial cavity the
floor of the valve is considerably thickened. Here two small scars, the
accessory diductors, are located.
The genital area is small and is situated on the sloping sides of the
shell just anterior to the dental plates. The teeth are small, corrugated,
and are supported by thin dental plates that are nearly obliterated in
some specimens by thickening of the inside of the shell and filling of
the umbonal cavities.
A prominent feature of Basiliola is the strong development of the
pallial markings. One main pair of pallial trunks, the vascula media,
originates between the diductor and adjustor muscles. A subsidiary
pair of trunks, the vascula genitalia, originates at the same place but
extends posterolaterally to surround the genital area. The vascula
media extend slightly anterolaterally to just beyond midvalve where
they branch. The main vascula media then extend slightly antero-
medially to terminate on the outside of the tongue. The other branches
at midvalve extend laterally where they divide. One branch swings
posteriorly near the valve margin to die out just before reaching the
teeth. The other branch extends anterolaterally. Short vascula ter-
minalia are given off from the outside anterior part of the vascula
media and their lateral branch, the vascula arcuaria.
The cardinalia of the brachial valve are characterized by the wide
and flat outer hinge plates to which are attached concave scimitarlike
falcifer crura. These are concave inward and are blunt at their distal
extremity. The crural blades are slightly oblique or nearly vertical as
viewed from the posteroventral side. The distal edge of the crura
facing the pedicle valve are usually finely serrate.
No median septum is present but in some specimens the elongated
adductor field is divided medially by a faint myophragm. The adductor
scars are small and elongated. The posterior pair is much smaller
than the anterior pair. The genital area is small like that of the pedicle
valve and is surrounded anteriorly and laterally by vascula genitalia
that connect with the posterior end of the vascula media. The major
trunks in the brachial valve are like those of the pedicle valve. The
vascula media originate at the outer ends of the adductor field and
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 29
extend anteriorly, generally following the outer slope of the median
trough formed by the fold. The vascula media branch at midvalve and
the branches form a course similar to those of the opposite valve.
The genus Neohemithyris as defined by Yabe and Hatai in 1934 is
identical to Basiliola, The authors of this genus emphasize the con-
junct deltidial plates and the nature of the foramen. Specimens of
Rhynchonella lucida Gould, type species of Neohemithyris, have been
compared with B. beechert and proved generically identical. In fact
the Japanese species suggests immature B. beecheri.
Specimens (U.S.N.M. 499321) from Vanua Mbalavu, Lau, Fiji,
referred by Ladd and Hoffmeister (1945, pp. 329-330) to Neohemi-
thyris lucida, are young forms having the characters of Basiliola. The
genus Basiliola thus proves to have a far wider range in the Pacific
than hitherto believed.
BASILIOLA ELONGATA Cooper, new species
Plate 14, C
Not Basiliola pompholyx Dall, Proc. U. S. Nat. Mus., vol. 57, p. 292, 1920
(U.S.N.M. 235844 and 300266).
Shell thin, of about medium size, elongate oval in outline; greatest
width anterior to the middle; beak acute, forming an angle of about
80°. Anterior margin nearly straight ; anterior commissure moderately
uniplicate ; valves subequal in depth, the brachial valve being slightly
deeper than the other; surface marked only by concentric lines of
growth.
Pedicle valve gently convex in lateral profile, with the maximum
convexity in the posterior half; anterior profile broadly but gently
convex; umbonal region moderately and narrowly swollen and with
steep lateral slopes; sulcus originating just anterior to the middle,
broad and shallow; tongue short and abruptly truncated; flanks
gently inflated and with gentle slopes. Foramen elongate-oval, fairly
large; deltidial plates conjunct and with a marked lip on the anterior
side of the foramen.
Brachial valve fairly evenly and moderately convex in lateral pro-
file ; strongly convex in anterior profile ; umbonal and median regions
inflated ; umbonal slopes steep ; fold originating at about midvalve, in
the anterior third slightly elevated above the surrounding flanks which
are moderately swollen and steep sided.
Pedicle valve interior with small teeth and short, inconspicuous
dental plates; diductor field moderately large, flabellate not strongly
inserted ; genital areas narrowly crescentic ; pallial marks not strongly
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
impressed. Brachial valve interior with long, stout, slightly curved
crura and moderately wide outer hinge plates.
MEASUREMENTS IN MILLIMETERS
Brachial Greatest
Length length width Thickness
POLS by De Ao) c)uceorn chet alnies decease actos 14.8 13.7 13.4 8.9
Bisared (paratype srs sicuiccinies as cise 15.8 13.8 niga 9.0
Types.—Holotype, U.S.N.M. 235844a ; figured paratype, U.S.N.M.
235844b.
Locality —U. S. Bureau of Fisheries Station 5146 at 24 fathoms,
off Sulade Island, Tapul Group, southern Philippines.
Discussion—tThis pretty little species is listed by Dall (1920,
p. 292) as B. pompholyx but it is a proportionally much longer shell
and with completely different outline. Compared with the growth lines
of specimens of B. pompholyx corresponding in length to those of
B. elongata the two species prove to be quite distinct.
Basiliola elongata is intermediate between B. beechert and B. lucida.
It is larger than the latter but smaller than the former although its
outline is similar. The Hawaiian shell is stouter and has somewhat
stronger shoulders than B. elongata. The Japanese B. lucida is also
less elongated than the new species.
Inside the brachial valves of these three species Basiliola elongata
has the longest crura whereas B. pompholyx has shorter crura rela-
tive to B. elongata and B. beecheri.
It is interesting to note that the bathymetric range of most of the
specimens of B. pompholy-x listed by Dall (1920) is deeper than 300
fathoms except for the new species and specimen U.S.N.M. 300863
which appears to be referable to B. elongata.
The specimen mentioned by Jackson and Stiasny (1937, p. 10) as
very small and coming from Kei Island is suggestive of the new
species. This is cited by them as a juvenile.
Genus EOHEMITHYRIS 1! Hertlein and Grant, 1944
Plates 5; A; 8; B; 75))B;) 20; (Ap B22, 1A:
Eohemithiris Hertlein and Grant, Publ. Univ. California at Los Angeles, Math.
and Phys. Sci., vol. 3, p. 55, 1044.
Subpentagonal in outline, thick shelled, coarsely fibrous, translu-
cent to transparent ; subequivalved, the brachial valve having a slightly
1 The spelling of Eohemithiris is here corrected to Eohemithyris to make it
coincide with the corrected spelling of Hemithyris. Inasmuch as Eohemithiris
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 31
greater depth than the pedicle valve; anterior commissure uniplicate,
the fold being broad and gentle; beak erect, small, inconspicuous ;
foramen small, round, slightly auriculate, submesothyrid; deltidial
plates conjunct. Surface smooth but with obscure anterior costation
in old individuals.
Pedicle valve with narrow delthyrial cavity bounded by short dental
plates plastered against the shell wall ; vascula media short, prominent.
Brachial valve with short, slender socket ridges bounding narrow
corrugated sockets ; outer socket plates moderately broad and shallow,
attached to long and exceptionally broad falcifer crura ; adductor field
subcircular, deeply inserted ; pallial trunks not impressed.
Type species (by original designation) —Eohemithiris alexi Hert-
lein and Grant, 1944, p. 55.
Comparison.—The entire anatomy of Eohemithyris is most like
that of Basiliola and other members of the Basiliolidae. It differs,
however, in being more nearly equivalve, whereas Pasiliola is strongly
inequivalve ; the deltidium of Eohemithyris is only slightly auriculate
and does not have the elaborate development of the pedicle collar
seen in Basiliola. The pallial marks of Basiliola are more strongly
developed in all species but those of the type species of Eohemithyris
are much abbreviated. The outer hinge plates attaching the crura to
the socket ridges are narrower in Eohemithyris than in Basiliola.
Comparison of Eohemithyris and Rhytirhynchia is essentially the
same as that with Basiliola. The brachial valve of Rhytirhynchia is
much deeper than that of Eohemithyris but the anterior costation of
the former is much stronger than that seen in Eohemithyris which
seems to be a rare feature.
Eohemithyris is quite suggestive of Aetheia in outline and beak
characters but differs from it in interior details. No development of
inner hinge plates appears to have taken place in Eohemithyris.
No other members of the Basiliolidae compare closely with Eohemi-
thyris.
Geological horizon.—Eocene (Domengine and Capay formations).
Assigned species—Four species are assigned to this genus, two
fossil and two Recent:
Eohemithiris alexi Hertlein and Grant, Eocene, California.
Eohemithyris? gettysburgensis Cooper, Miocene, California.
Hemithyris colurnus Hedley, Recent, Australia.
Rhynchonella grayi Woodward, Recent, Fiji Islands.
was thought by its authors to be similar to Hemithyris and an early relative of
it, the correction of spelling in the latter by Bronn is essential in the former.
The spelling of Eohemithiris is corrected to Eohemithyris in Zoological Record
for 1950, p. 21.
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Distribution.—The fossil species are from the Eocene and Miocene
of California but one Recent form is from off southeastern Australia,
and another off the Fiji Islands.
Discussion—The name chosen for this genus is unfortunate and
inappropriate because the interior details now make it clear that it is
totally unrelated to Hemithyris as its name implies. Relationship to
the Basiliolidae seems clear in the broad falcifer crura, the details
of the deltidial plates, foramen, and smooth exterior. Eohemithyris
is the oldest known member of the Basiliolidae but its roots are prob-
ably deep in the Cretaceous. It is also interesting that species are living
today.
Hemithyris colurnus Hedley is here assigned to this Eocene genus.
This Australian species has never been satisfactorily placed and some
objections may be raised to assigning it to Eohemuthyris, an Eocene
species now known only from California. In spite of the time gap
indicated, close comparison of the California and modern Australian
species leaves few anatomical points of difference. The exterior of
H. colurnus is essentially identical to that of Eohemithyris alexi.
Both are thick-shelled forms with translucent to almost transparent
shells, especially when they are wet. They are both coarsely fibrous.
The beak characters of the two are identical. It is not possible to
make a comparison of the pedicle collars of the two species because
it is very difficult to determine these details in Eohemithyris. Actually
some uncertainty exists as to whether the fossil species has a pedicle
collar, but the area of the beak is so thickened that some sort of
tubular arrangement must be present.
Inside the pedicle valve the dental plates of the modern species
may be somewhat less prominent than those of the fossil form;
dental plates are definitely present in both however. It is to be ex-
pected that those of the older species might be better developed than
those of the modern form. The delthyrial cavities and muscle areas
of the two seem identical; the pallial trunks of the modern form are
better impressed but this may be a matter of preservation rather than
one of generic distinction.
Inside the brachial valve the crura and hinge plates are almost iden-
tical, no features of generic value having been detected. The outer
hinge plates are of about the same size, narrower than in Basiliola
but much wider than in Aphelesia. The adductor field of the modern
species is deeply impressed as in Eohemuithyris alexi but the pallial
marks of the Recent species are more plainly impressed. The sock-
ets of H. colurnus are strongly corrugated but the corrugation of the
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 33
Eocene species is not so strong in the specimens examined. This
however could hardly be regarded as a generic difference. Other
species of Eohemithyris can be expected in other Tertiary deposits
and should be looked for.
Rhynchonella grayi Woodward is another species that has never
been correctly placed generically. Through the kindness of Dr. Helen
M. Muir-Wood and the authorities of the British Museum I am able
to furnish exterior and interior views of this species. It is clearly
very similar to Hemithyris colurnus but is more strongly costate in
the anterior third. Eohemithyris alexi and colurnus are both ob-
scurely to definitely costate in the anterior part. Davidson’s figures
of R. grayi greatly exaggerate the plication. The interiors and beak
characters of R. grayi are clearly identical to those of E. colurnus
and E. alexi, except for the swellings of shell material on the hinge
plate, consequently the species is assigned to Eohemithyris. The shell
profile and beak characters exclude FR. grayi from assignment either
to Basiliola or Rhytirhynchia. Lack of inner hinge plates separates
R. grayi from Aetheia which, except for the anterior costation, it
otherwise resembles in its exterior characters.
EOQHEMITHYRIS? GETTYSBURGENSIS Cooper, new species
Plate 8, B
Shell large, subpentagonal in outline, slightly wider than long ;
sides narrowly rounded ; widest slightly anterior to midvalve ; anterior
commissure strongly uniplicate; surface marked only by concentric
lines of growth.
Pedicle valve less deep than the brachial valve, moderately convex
in lateral profile and with the strongest convexity in the posterior
third; anterior profile nearly flat but with the median region slightly
concave ; beak low, incurved ; umbo moderately swollen; sulcus origi-
nating on the umbo, shallow and narrow but deepening and widening
anteriorly to occupy slightly more than half the width at the anterior ;
flanks somewhat flattened and with gentle slopes to the margins;
tongue moderately geniculate, moderately long and broadly rounded.
Brachial valve gently and fairly evenly convex in lateral profile ;
anterior profile moderately strongly domed; fold originating at about
midvalve, low, flattened, and prominent only at the anterior; flanks
bounding fold slightly depressed, gently rounded. Umbonal region
only slightly convex.
Interior —Strong, short dental plates visible in pedicle valve;
small, short socket ridges visible in brachial valve but no trace of a
median septum or ridge seen through the moistened shell.
34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
MEASUREMENTS IN MILLIMETERS
Brachial Maximum
Length length width Thickness
FIGIGE DE. asst Gira s keae ensues sete 24.3 22.2 28.0 13.0
Type.—Holotype, U.S.N.M. 549382.
Horizon and locality—Miocene, Station 69, on coast, 44 miles west
of Gettysburg, Washington.
Discussion—This is a large and distinctive species unlike any
figured by Hertlein and Grant (1944) in their monograph on the
Tertiary and Recent brachiopods of the west coast of the United
States. Tentative assignment to Eohemithyris is made because the
exterior is smooth, dental plates are present, but a median septum or
ridge is absent. The species differs from FE. alexi in its greater size
and more pronounced fold and sulcus.
Genus NEOHEMITHYRIS Yabe and Hatai 1934
Plate 13, B
Neohemithyris Yabe and Hatai, Proc. Imp. Acad. Japan, vol. 10, No. 9, p. 587,
1934; Hatai, Sci. Rep. Tohoku Imp. Univ., ser. 2 (Geology). vol. 20, p. 210,
1940.
Yabe and Hatai (1934, p. 587) described their new genus Neo-
hemithyris with type species (by original designation) Rhynchonella
lucida Gould as resembling Hemithyris “in shape, folding, beak
characters and microstructure’ but differing “only in possessing an
entire foramen and conjunct deltidial plates in the ventral valve.” In
the brachial valve a cardinal process is absent and no median ridge
is present. Although these characters do distinguish Neohenuthyris
from Hemithyris they do not differentiate the Japanese shell from
Basiliola with which it seems to be identical. Consequently I have
placed Neohemithyris in the synonymy of Basiliola. [For further dis-
cussion see under Basiliola; see also pl. 13, B, figs. 6-23, for illus-
trations of Rhynchonella lucida Gould type species of Neohemithyris
(=Basihiola).]
Genus NEORHYNCHIA Thomson, 1915
Plate 2, B
Neorhynchia Thomson, Geol. Mag., n. s., dec. 6, vol. 2, p. 388, 1915; Dall, Proc.
U. S. Nat. Mus., vol. 57, p. 290, 1920; Thomson, New Zealand Board Sci.
Art, Manual 7, p. 149, 1927; Hertlein and Grant, Publ. Univ. California,
Math. and Phys. Sci., vol. 3, p. 57, 1944.
Pentagonal in outline, with the greatest width at midvalve; valves
unequal in depth, the pedicle valve having the greater depth ; anterior
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 35
commissure deeply sulcate; surface smooth. Beak of pedicle valve
short, nearly straight, and bluntly pointed ; foramen of moderate size,
hypothyrid ; deltidial plates disjunct.
Pedicle valve interior with small corrugated teeth supported by
short dental plates which define a small delthyrial chamber ; muscle
area small.
Brachial valve interior with corrugated sockets bounded by over-
hanging socket ridges ; crura short, falcifer type, crescentic in section,
attached to socket ridges by broad outer hinge plates. Inner hinge
plates small and inconspicuous. Median ridge short and reaching the
apex. Adductor field small.
Type species (by original designation).—Hemithyris strebeli Dall,
Bull. Mus. Comp. Zool., Harvard Coll., vol. 43, p. 441, 1908.
Comparison and discussion—The important and striking difference
between Neorhynchia and all known Recent and Tertiary rhynchonel-
loid genera is the sulcation of the anterior commissure. Rhynchonel-
loids of similar habit are known from the Mesozoic. They are also
known from the Devonian, Mississippian, and Permian as well.
Sulcation is a folding tendency that has appeared many times in dif-
ferent stocks of the rhynchonelloids.
Neorhynchia is most closely related to Basiliola in the presence of
the wide outer hinge plates and falcifer crura having a crescentic sec-
tion. The presence of incipient inner hinge plates in Neorhynchia is
another difference between the two genera.
Assigned species.—Only one species is so far known in this genus:
Hemithyris strebeli Dall, Recent, Pacific.
Distribution—The known specimens of this species are all from
great depths: 2,084 fathoms at 35.1° F. in mid-Pacific and 2,035
fathoms at 35.3° F. off the Galapagos Islands, both on Globigerina
ooze.
Genus RHYTIRHYNCHIA Cooper, 1957
Plate 11, A
Rhytirhynchia Cooper, U. S. Geol. Surv. Prof. Pap. 314-A, p. 8, 1957.
Subcircular to suboval in outline and with the maximum width at
midvalve; strongly inequivalve, the brachial valve being swollen and
deep; anterior commissure sulciplicate, surface smooth except ante-
rior which is paucicostate. Beak small, rounded, inconspicuous ; fora-
men rounded, submesothyrid to mesothyrid; deltidial plates short,
conjunct.
Pedicle valve with thick, coarsely corrugated teeth and moderately
developed to remnantal dental plates; pedicle collar well formed;
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
muscle field short and narrow, somewhat longitudinally rectangular in
outline; diductor scars small; adductor scars large and surrounded
anteriorly by the diductors. Vascula media strong, converging an-
teriorly on the tongue.
Brachial valve interior with deep sockets bordered by overhanging
socket ridges ; crura attached to socket ridges by narrow outer hinge
plates ; crura falcifer, long crescentic in section, convex outward; in-
ner hinge plates incipient, forming a slight thickening on the inside of
the crura near their proximal end; median ridge or septum absent;
adductor field small, rounded in outline with large anterior scars and
small posterior ones; vascula media prominent, diverging widely at
the anterior end of the adductor field.
Type species (by original designation).—Hemithyris sladeni Dall,
Trans. Linnaean Soc. London, ser. 2, Zool., vol 13, pt. 3, p. 440, pl.
26, figs. 7-12, I910.
Comparisons.—This genus is most like Basiliola in form and out-
line but differs in having anterior costation. Inside the pedicle valve
the dental plates are reduced to remnants or are wanting in the mod-
ern species. In the brachial valve the development of outer hinge
plates in Basiliola is usually greater than that in Rhytirhynchia but
otherwise the details of the valves are the same. Incipient inner hinge
plates appear in Rhytirhynchia.
Rhytirhynchia in its anterior costation suggests Eohemithyris which
likewise has anterior costation in old adults. In the latter this seems
to be a rare feature but the two genera are not likely to be confused
because their lateral profiles are different, that of Rhytirhynchia hav-
ing an extremely deep brachial valve, whereas Eohemithyris has both
valves nearly equal.
Geological range.—Pliocene to Recent.
Distribution —Rhytirhynchia occurs as a fossil in the Pliocene
of Okinawa and today lives in the Indian Ocean south of the Saya de
Malha banks.
Assigned species ——Two species are now assigned to this genus, one
living and one fossil:
Hemithyris sladeni Dall, Recent, Indian Ocean.
Rhytirhynchia hataiana Cooper, Pliocene, Okinawa.
Discussion—This genus is essentially a semicostate Basiliola. In
the one modern species the dental plates are remnantal but in FR. hatai-
ana from the Pliocene of Okinawa the dental plates are moderately
developed. This is a small and delicate form in which internal
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 37
thickening of the shell is not great. The degrees of development of
the dental plates are, in this case, not regarded as generic in character.
PROBOLARINA Cooper, new genus
(Gr. probolos, projection)
Plate 17, A, B
Subpentagonal to subtriangular in outline, with the greatest width
at or near the middle; inequivalve, the brachial valve having the
greater depth and convexity ; anterior commissure uniplicate; surface
semicostate, the posterior third to half smooth, anterior half to two-
thirds costate. Beak moderately long, pointed, nearly straight; fora-
men small, longitudinally elliptical, hypothyrid to submesothyrid and
with strongly auriculate margins. Deltidial plates prominent, wholly
visible, conjunct throughout their length and anteriorly resting on the
umbo of the brachial valve.
Pedicle valve interior with strong pedicle collar, small teeth sup-
ported by vertical dental plates separated from the side wall by narrow
umbonal chambers. Details of the musculature not available.
Brachial valve interior with narrow sockets bounded by erect but
not greatly thickened socket ridges; crural bases attached to socket
ridge by a prominent, flat outer hinge plate; crura falcifer, long, scimi-
tarlike, crescentic in section and convex outward. No cardinal proc-
ess. Muscle and pallial marks not visible in available material.
Type species—Rhynchonella holmesu Dall, Trans. Wagner Free
st. Scie. Vol. 9, pt. G.p. 1530, pl. 50, fes.10,,.12) (0b Li), £903.
Comparisons.—This genus is most like Rhytirhynchia in its exterior
characters but differs importantly in the interiors as well as in details
of the exterior. Rhytirhynchia has almost nude valves except for the
costation at the very anterior margin. In Probolarina on the other
hand the costation affects more than two-thirds of the valve, only the
umbones being free of costation. The deltidial plates of the two
genera are conjunct and both are auriculate but those of Probolarina
are more developed and more elaborately auriculate than those of
Rhytirhynchia.
Inside the pedicle valve of Probolarina a strong pedicle collar
strengthens the beak and strong but thin dental plates buttress the
teeth. In Rhytirhynchia on the other hand the dental plates are rudi-
mentary in the type species and can be seen only as a trace on the sides
of the shell. In R. hataiana Cooper from the Pliocene of Japan mod-
erately developed dental plates are present but they are not to be
compared with the strong and vertical plates of Probolarina.
38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
The cardinalia of the brachial valves of the two genera are very
similar and a median septum or ridge is lacking from both of them.
The crura of both genera are of the same falcifer type and the outer
hinge plates are developed to about the same degree.
Assigned species.—At present two species only are assigned to this
genus:
Rhynchonella salpinx Dall.
R. holmesii Dall.
Distribution—Eocene (Castle Hayne), North Carolina.
Discussion—Tertiary brachiopods are a rarity in the United States
and that is especially true of the genus Probolarina. The two species
of this genus are represented by a few specimens only. They are
also quite different in form but the beak characters and the cardinalia
of the two appear to be identical. It is interesting that the cardinalia
of Probolarina are so like those of Rhytirhynchia, a modern inhabit-
ant of the Indian Ocean and represented in the fossil state in Okinawa.
PROBOLARINA HOLMESII (Dall)
Plate 17, B
Rhynchonella holmesii Dall, Trans. Wagner Free Inst. Sci., vol. 3, pt. 6, p. 1536,
pl. 58, figs. 10, 12 (not 11), 1903.
In Dall’s description of this species it is stated that one of the
figured specimens is a young individual. The other specimen figured
is somewhat fragmentary, probably belonging to a different and
undescribed species. I here select the smaller of the two specimens
as the type of R. holmesiti, U.S.N.M. 109298a. This specimen is
clearly a young form of those figured on plate 17, B. Specimen
U.S.N.M. 549359 is a well-preserved adult of R. holmesu.
STREPTARIA Cooper, new genus
(Gr. streptos, twisted)
Platenro wpe
Pentagonal in outline with the greatest thickness near midvalve;
valves unequal in depth, the brachial valve having the greater depth;
anterior commissure uniplicate to twisted, either right or left; sur-
face marked only by concentric lines of growth, occasionally with
obscure marginal costae. Beak short, deltidial plates conjunct,
foramen hypothyrid to submesothyrid, small and usually with promi-
nent elevated rim.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 39
Pedicle valve interior with remnantal dental plates and strong
corrugated teeth. Other details not yet known.
Brachial valve with deep corrugated sockets, long falciform crura
attached to fairly broad outer hinge plates ; inner marginal rim pres-
ent on crura but no inner hinge plates ; socket ridges thick and curved.
No median septum. Other details not yet known.
Type species —Terebratula De Buchit Michelotti, Cenn. Brach.,
Acefali foss. Italia, p. 4, 1938.
Comparison and discussion.—This genus is characterized by its
smooth exterior, small, short beak, twisted to uniplicate anterior
commissure, nearly obsolete dental plates, and long falciform crura
attached to broad outer hinge plates. In the latter character and the
smooth, uniplicate shell Streptaria is like Basiliola but it differs in
beak characters, lack of a pedicle collar, and the small development
of the dental plates of the pedicle valve.
Specimens of this genus are similar to Erymnaria in the smooth
exterior, beak characters, and the twisted anterior margin, but the
latter genus possesses two strong, diverging septa in the brachial
valve—a character unlike any other known rhynchonelloid from Terti-
ary rocks or Recent seas.
Assigned species—This genus is known in Mediterranean and
West Indian rocks.
Terebratula De Buchii Michelotti, Miocene, Italy.
Rhynchonella deformis Seguenza, Miocene, Italy.
R. eocomplanata Sacco and var., Eocene, Italy.
Streptaria streptimorpha Cooper, new species, Eocene, Cuba.
Distribution—The known species of this genus are from the
Tertiary of Italy, southern Europe, northern Africa, and Cuba.
Discussion.—One of the interesting features of Streptaria is the
twisted anterior margin. This character occurs in rhynchonelloid
stocks from Paleozoic to Tertiary times. It has been seen in many
different stocks and undoubtedly is an aberration of the anteriorly
produced folding that facilitates the passage of nourishing currents
into the valve and their elimination with waste from the valves.
Streptaria and Erymnaria form isochronous homeomorphs in this
respect. The Ordovician genus Streptis is like Streptaria in having
shells twisted to right and left but also has normally uniplicate indi-
viduals or species.
Cuba has produced another species of Streptaria which is not de-
scribed because of insufficient material. Three specimens of this
undescribed shell are known from the Eocene of Camaguey Province
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
in which the valves are normally folded (uniplicate), but a third
specimen has a wider sulcus on the pedicle valve which shows a
definite twist. This species has abbreviated dental plates and the
same cardinalia as the Italian forms and S. streptimorpha.
STREPTARIA STREPTIMORPHA Cooper, new species
Plate 19, B
Of medium size for a rhynchonelloid, subcircular in outline; sides
narrowly rounded ; greatest width at the middle ; anterior commissure
twisted. Surface marked only by concentric growth lines.
Pedicle valve gently convex in lateral profile; broadly and slightly
convex in anterior profile; depth less than that of the brachial valve ;
umbonal and median regions slightly swollen; sulcus indefinite, shal-
low; beak short, blunt, forming an obtuse angle (about I10°).
Foramen hypothyrid, small, oval; deltidial plates forming low rim
around foramen.
Brachial valve deeper than the pedicle valve, moderately convex
in lateral profile but strongly domed in anterior profile; umbonal
region somewhat flattened; median region and flanks strongly swol-
len; fold ill defined because of twisted commissure.
Interior —Pedicle valve with remnantal deltidial plates and no
pedicle collar. Brachial valve with long falciform crura attached to
the socket ridges by fairly broad outer hinge plates. Median septum
absent. Other details not yet known.
MEASUREMENTS IN MILLIMETERS
Brachial
Length length Width Thickness
PPOLORY Pe Ns YER. Mideted esi ollatene 17.0 15.2 18.0 10.6
Types.—Holotype, U.S.N.M. 549386a ; figured paratype, U.S.N.M.
549386b.
Horizon and locality—Eocene, deep cut north of Grua 9, Ramal
Juan Criollo, Camaguey Province, Cuba (Palmer locality 1640).
Discussion.—This species is characterized by its rounded form,
small foramen, and broadly twisted anterior commissure. It is sug-
gestive of Streptaria de buchii (Michelotti) from the Mediterranean
region but differs in its rounded form, less narrow twist to the an-
terior commissure, smaller foramen, and lesser development of the
foraminal lip.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 4I
APHELESIINAE Cooper, new subfamily
APHELESIA Cooper, new genus
(Gr. apheles, smooth)
Plates, 7, 8,8: C. 22,
Outline subtriangular to subpentagonal; widest anterior to the
middle; strongly inequivalve, the brachial valve being deep and swol-
len, the pedicle valve gently convex; anterior commissure strongly
uniplicate but fold of brachial valve defined only at the anterior;
smooth on most of the surface but the anterior with incipient costa-
tion. Beak of pedicle valve moderately elongated, nearly straight to
suberect, pointed ; foramen complete, elongate-oval, small hypothyrid ;
deltidial plates thick, conjunct, moderately auriculate; beak apex
thickened internally.
Pedicle valve interior with elongated, corrugated teeth supported
by thick dental plates. Muscular field large and flabellate, extending
to about midvalve with the diductor scars surrounding the adductors ;
adjustor scars small and laterally disposed.
Brachial valve interior with deep corrugated sockets bounded by
strong overhanging socket ridges; crura of falcifer type, long, cres-
centic in section, broad, scimitarlike and cemented directly to the
socket ridges with no outer hinge plates developed ; inner hinge plates
lacking ; crural supporting plates thick. Median ridge low and thick;
adductor field narrow and elongated. Pallial trunks not deeply im-
pressed.
Type species—Anomia bipartita Brocchi, Conch. Foss. Subapp.,
vol. 2, p. 469, pl. 10, fig. 7, 1814.
Comparisons.—This species is generally referred to Hemithyris
but it actually does not have either the exterior or interior features
of this genus. Aphelesia is completely smooth or with slight and very
obscure costation. It does not have the numerous and regular sub-
dued costellae or striae of Hemithyris. Furthermore the foramen of
Aphelesia is small and the deltidial plates are conjunct. The foramen
of Hemithyris is large and not enclosed anteriorly because the deltidial
plates are disjunct.
The interior of the pedicle valve of each of these genera is quite
similar except for the fact that the dental plates of Hemithyris are
somewhat more prominently developed and with deeper umbonal
chambers than those of Aphelesia. Important differences appear on
the inside of the brachial valves where the cardinalia of the two
genera are quite distinctive. In Hemithyris the crura are of radulifer
42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
type, long and slender and with only a slight development of outer
hinge plates. The crura however are flattened horizontally as one
observes them from the posteroventral side and the distal extremi-
ties are quite thin. In Aphelesia the crura are long, broad, of falcifer
type, and the bases attached to the socket ridges with no development
of outer hinge plates. The distal ends of the crura, unlike Hemi-
thyris are scimitarlike, are laterally compressed and their distal ex-
tremities serrate. In cross section these crura are crescentic and the
convex surface faces laterally. The crural blades are broad and thick
and thus quite unlike those of Hemithyris.
The crura of Aphelesia are like those of Rhytirhynchia, Neorhyn-
chia, and Basiliola but differ from all of them in the absence of outer
hinge plates which are so prominent in the other genera. Aphelesia
differs from these genera also in other important respects.
Geological horizon.—Eocene through Pliocene.
Geographic distribution—Mediterranean region.
Assigned species.—At present it is difficult to assign the several
species of Mediterranean Tertiary rhynchonelloids to their proper
genus because the interiors are poorly known or completely unknown.
Anomia bipartita Brocchi, Pliocene, Italy.
Terebratula plico-dentata Costa, Miocene-Pliocene, Italy.
Rhynchonella (Hemithyris) saccoi Patané, Pleistocene, Sicily.
Hemithyris acuta Meznerics, Miocene, Hungary.
Rhynchonella bipartita pseudobipartita Patané, Pleistocene, Sicily.
Discussion—These species have been assigned to Hemithyris at
one time or another but the exterior characters preclude such a place-
ment. The little that is known of the interior also excludes these shells
from assignment to Hemithyris. The beak characters and cardinalia
of Aphelesia as shown by A. bipartita are quite unlike the same fea-
tures of Hemithyris. The exterior of most of these shells is smooth
or nearly so. Some exhibit anterior costation but it is generally not
regularly developed. None of them have the fine striate exterior of
Hemithyris. The latter, too, has disjunct deltidial plates and an
elongate beak, whereas the beaks of the Italian species are short and
the deltidial plates conjunct. The crura of Hemithyris are long,
curved, and slender, quite different from the broad-bladed Aphelesia
bipartita.
AETHEIINAE Cooper, new subfamily
Genus AETHEIA Thomson, 1915
Plates 4, A, 9, B
Actheia Thomson, Geol. Mag., n. s., dec. 6, vol. 2, p. 389, 1915; Thomson, New
Zealand Board Sci. Art, Manual 7, p. 156, 1927.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 43
Thomsonica Cossmann, Rey. Crit. Pal., vol. 24, No. 3, p. 137, 1920; Finlay,
Trans. New Zealand Inst., vol. 57, p. 532, 1927.
Outline elongate-oval to triangular with the greatest width at the
front; inequivalve, the brachial valve having the greater depth; an-
terior commissure broadly uniplicate, the brachial fold low and in-
conspicuous. Surface marked by concentric lines of growth only.
Beak small, erect; foramen minute, submesothyrid; deltidial plates
conjunct, forming a concave plate.
Pedicle valve interior with thick teeth attached directly to the shell
wall; dental plates absent; muscle field short and narrow, the di-
ductors small but surrounding the adductor scars. Vascula media
strong, branching about two-thirds the shell length from the beak.
Brachial valve interior with deep corrugated sockets bounded by
long vertical socket ridges to which the long crura are cemented;
crura of falcifer type, crescentic in section, convex outward ; inner
hinge plates thick and filling the intercrural space. Cardinal process
small and transversely triangular. Median ridge short and low, but
thick, united with the cardinalia. Adductor field large, with large
anterior scars.
Type species (by original designation).—Waldheimia (?) sinuata
Hutton, Catalogue of the Tertiary Mollusca and Echinodermata of
New Zealand in the collection of the Colonial Museum, p. 36, 1873=
?Terebratula gualtert Morris, Quart. Journ. Geol. Soc., London,
vol. 6, p. 329, pl. 28, figs. 2, 3, 1850.
Comparisons—This interesting genus has exterior and interior
features that set it aside from nearly all other rhynchonelloids. It
is unlike all other known Tertiary and modern rhynchonelloids ex-
cept Patagorhynchia in the extremely small pedicle foramen and con-
cave deltidial plates. It differs from Patagorhynchia in being smooth
rather than marked by squamose costellae. Internally it differs from
all other known Tertiary and Recent rhynchonelloids except Frieleia
in the great development of the inner hinge plates which grow and
swell between the crural bases to plug the entire posterior.
Geological horizon.—Upper Cretaceous to Miocene.
Distribution—New Zealand.
Assigned species:
Terebratula gualteri Morris.
Waldheimia ? sinuata Hutton.
Discussion—This genus presents some peculiarities not seen in
most of the Tertiary and Recent rhynchonelloids. The small foramen
is submesothyrid, an unusual position for this group of animals. The
44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
deltidial plates are conjunct but they do not overlie the umbo of the
brachial valve as in Hemuithyris and several other genera. They are
concave and lie ventrally to the umbo of the brachial valve.
The teeth are large, corrugated and not buttressed by dental plates.
Instead of dental plates a thickening extends posteroventrally but
does not meet the floor. I have never seen immature specimens and
therefore cannot say whether or not dental plates existed in the
young as in some other genera.
The pallial marks of the pedicle valve are like those of His-
panirhynchia and Basiliola. In the one specimen showing these marks
the course of the sinuses appears as an elevated ridge rather than a
depression. This bifurcates near midvalve as in the genera mentioned,
The great thickening of the apical region of the brachial valve ob-
scures many of the details of the cardinalia that can only be cleared
up by a study of young specimens. These are not available in the
National collections. The true nature of the crural bases is not known,
whether they attach to the median ridge or to the valve floor or
whether they have supports that extend dorsally.
Thomson (1927, p. 157) assigned Hemithyris colurnus Hedley
and H. sladem Dall to his genus Aetheia even though they differed to
some extent from the fossil genus. Because of its anterior costation
and the bulbous brachial valve the latter of these two species is here
placed in Rhytirhynchia, and the former, because of its nearly equally
deep valves, among other characters, is placed in Eohemithyris.
In their correction of brachiopod homonyms in 1951, Cooper and
Muir-Wood suggested that Thomsonica Cossmann, 1920, should be
substituted for Aetheia because the latter name is preoccupied by
Aethia Merrem 1788 (Aves). It is now the sense of the Zoological
Commission as outlined in the Copenhagen Proceedings (Hemming
1953, Article 34, paragraph 153, p. 78), that these two names are not
in conflict. It is therefore necessary to return to Aetheia and reject
Thomsonica as Thomson did in 1927 (p. 156).
Genus PATAGORHYNCHIA Allan, 1938
Plates 6, A, 21, B
Patagorhynchia Allan, Rec. Canterbury (N.Z.) Mus., vol. 4, No. 4, p. 199, 1938.
Subcircular to subpentagonal in outline; inequivalve, the brachial
valve having the greater depth; anterior commissure uniplicate, the
fold of the brachial valve being moderately strong. Surface costellate,
lamellose to imbricate. Beak short, nearly straight, bluntly pointed ;
foramen minute, submesothyrid, deltidial plates conjunct and form-
ing a concave plate.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 45
Pedicle valve interior without dental plates. Other details not
known.
Brachial valve interior with short crura and with the inner hinge
plates thickened and filling the intercrural space.
Type species (by original designation) —Rhynchonella patagonica
von Ihering, Anal. Mus. Nac. Buenos Aires, ser. 3, vol. 9, t. 2, p. 334,
pl. 3, figs. 11a, b, 1903.
Comparisons.—Patagorhynchia is comparable to two genera from
the Southern Hemisphere: Tegulorhynchia and Aetheia. The first
genus is ornamented like Patagorhynchia but there the similarity
ends because the South American genus has completely different beak
characters and the interiors are wholly unlike. Close comparison is
possible with Aetheia on the inside and in beak characters but Aetheia
is a smooth shell and externally not to be confused with Patago-
rhynchia. The Argentine shell has the minute foramen and concave
deltidial plates like the New Zealand shell. Inside the pedicle valve
no dental plates were observed in Patagorhynchia. The interior of
the brachial valve is not well known and the published illustration
of it is poor. It does indicate, however, cardinalia with moderately
long crura, concave inward, probably of falcifer type and a thicken-
ing of the inner hinge plates to fill the posterior space between them
with shell substance. The illustration indicates a condition even more
extreme than that seen in Aetheia.
Geological horizon.—Eocene (Patagonian).
Distribution—Argentina and Chile.
Assigned species——Only one species, the type of the genus, is
known.
Discussion—Allan (1938) discussed the interior details of Pata-
gorhynchia, especially those of the pedicle valve. Specimens of
pedicle valves in the Canterbury Museum enabled him to determine
some characteristics not before seen, such as the strong concavity of
the deltidial plates and the fact that they do not exhibit the suture line.
He also described the great thickening formed by coalescence of the
deltidial plates with a platform made by a thickening at the base of
the teeth.
Family HEMITHYRIDAE Rzhonsnitzkaia 1956
Genus HEMITHYRIS d@’Orbigny, 1947
Plates.3;;Aj By 4 EF
Hemithiris d Orbigny, Paléont. France Ter. Crét., vol. 4, p. 342, 1847; Hertlein
and Grant, Publ. Univ. California at Los Angeles, Math. and Phys. Sci., vol.
3, P. 41, 1944.
46 . SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Hemithyris d’Orbigny, Bronn, Neues Jahrb. Min., Geog., Geol. u. Petrefaktenk,
p. 246, 1848; Thomson, Geol. Mag., dec. 6, vol. 2, p. 387, 1915; Thomson, New
Zealand Board Sci. Art, Manual 7, p. 149, 1927; Grabau, Sci. Quart. Nat.
Univ. Peking, vol. 3, No. 2, p. 112, 1932; Hatai, Sci. Rep. Tohoku Imp.
Univ., ser. 2 (Geology), vol. 20, p. 194, 1940.
Outline triangular, greatest width at or anterior to the middle;
inequivalve, the brachial valve having the greater depth and con-
vexity ; anterior commissure broadly to narrowly uniplicate; surface
obscurely to moderately costellate, the costellae broad and separated
by fine striae. Beak of pedicle valve prominent, elongate, and sub-
erect. Foramen incompletely hypothyrid; deltidial plates disjunct ;
apical region thickened and buttressed by a short median ridge.
Pedicle interior with strong, somewhat elongated, corrugated teeth ;
dental plates vertical and strong, the umbonal cavities becoming par-
tially filled by adventitious deposit in old specimens. Delthyrial
cavity occupied by the pedicle. Apical plate thick, commonly some-
what elevated. Muscle field anterior to the delthyrial cavity, sub-
flabellate, the diductor scars surrounding the adductors. Adjustor
scars lateral to the diductors. Lateral areas bounding muscle field
pitted; pallial marks obscure.
Brachial valve with deep, corrugated sockets defined by strong
crural supporting plates; socket ridges prominent; crura of radulifer
type, long, slender, curved, forming horizontally flat blades distally,
widening posteriorly to form a narrow hinge plate and strengthened
anteriorly by an oblique ridge running from the outside of the tip to
the inside of the hinge plate; crural supporting plates buttressing
hinge plate; inner hinge plate absent or incipient. Cardinal process
absent, the diductor muscles being attached to an apical, roughened
pit. Median ridge low, defined chiefly at midvalve and disappearing
posteriorly in the umbonal chamber. Adductor field small consisting
of a large pair of triangular anterior scars and a pair of small,
elongate, subrhomboidal scars situated on the outside posterior to
the anterior set.
Type species (by subsequent designation, d’Orbigny, 1847).—
Anomia psittacea Gmelin, Syst. Nat., vol. 2, p. 3348, 1790.
Comparisons.—The exterior and interior details of Hemithyris are
distinctive and have no known close counterparts among the Tertiary
and Recent brachiopods. The genera nearest like Hemithyris are
Aphelesia and Notosaria. The former differs from Hemithyris in
being exteriorly smooth and in having broad, concave, falcifer crura.
The external form of Notosaria is suggestive and the beak charac-
NO. 5 RHYNCHONELLOID BRACHIOPODS—-COOPER 47
ters are similar but that genus has well-marked costae and the crura
are shorter.
Geological horizon.—Miocene to Recent.
Distribution—All of the northern seas from the Arctic south to
Japan in the Pacific and south to the coast of Maine in the Atlantic.
Assigned species——The following species, fossil and Recent are
placed in Hemithyris:
Anomia psittacea Gmelin, Pleistocene to Recent, Northern Hemisphere.
Hemithyris psittacea alaskana Dall, Recent, Alaska.
Rhynchonella woodwardi (A. Adams), Recent, Japan.
H. braunsi Hayasaka, 1928, Pliocene to Recent, Japan.
FH. peculiaris Nomura and Hatai, 1936, Recent, Japan.
Discussion —The most distinctive features of Hemithyris are the
ornamentation and cardinalia. The type species is strongly marked
but other species assigned here are very delicately or obscurely orna-
mented. Hemithyris is really better described as striate rather than
costellate. The surface is marked by radial grooves or impressed
lines, fairly uniform in H, psittacea but discontinuous and irregular
in H. woodwardi. The spaces intervening between the lines are flat
and broad and thus simulate costellae. In H. woodwardi the im-
pressed lines are very delicate and so irregular that broad, smooth
patches of shell are separated by the striae. These cannot be con-
strued as costellae. This type of ornamentation was seen in this
study in only one other rhynchonelloid, Plicirhynchia. In this Ar-
gentinian genus the region around the umbones is marked as in
Hemithyris but the striae do not extend to the costate portion of the
valves.
The crura are unusual because they are long and slender and are
usually flattened in a dorsoventral direction rather than laterally as
in most of the other genera. They are thus unlike the crura of any
other modern rhynchonelloid. The flat blade is attached to the outer
hinge plate on its posteroventral surface. In side view the edge of
the outer hinge plate forms an oblique ridge and the crus lies at
angle under it. The distal end of the crus is usually pointed, the point
being on the inside of the plate. This type of crura is generally
classified as belonging to the “radulifer” group.
The cardinal process is seldom conspicuous. It is a triangular area
at the apex, roughened horizontally and usually divided by the cleft
in the hinge plate which extends to the apex. It is quite like that of
several other genera such as Notosaria.
Median septa or ridges are never well developed in Hemithyris,
48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
even in old and obese specimens. This makes a ready distinction and
helps to show that septate specimens from the West Coast Tertiary
of the United States are not referable to Hemithyris where some of
them have been placed.
The name Hemuithyris has been applied to many Tertiary and
modern species without regard to geographic realm or biological con-
siderations. Many smooth species have been referred here and some
plicated species have also been given this name. The ornament of the
genus is so distinctive that confusion with other genera should not
occur. Hemithyris as now defined appears to be confined to the
Northern Hemisphere. Most of the species assigned to this genus
from the Southern Hemisphere clearly belong to other genera. As
explained above it is difficult to place the rhynchonelloid species gen-
erically from descriptions which do not include the interior details.
It has thus proved impossible to reassign many of the species now
described as Henuthyris or placed in that genus.
NOTOSARIA Cooper, new genus
(Gr. notos, south)
Plates; 422.16
Subpentagonal in outline, usually widest at the middle ; inequivalve,
the brachial valve having the greater depth and convexity; anterior
commissure uniplicate; brachial valve fold usually low. Surface
costellate ; costellae crossed by growth lines and growth varices only.
Beak short to moderately long, nearly straight to suberect ; foramen
large, incomplete, hypothyrid; deltidial plates vestigial to prominent,
disjunct ; beak with thick, transversely striated apical plate.
Pedicle valve interior with large corrugated teeth supported by
short receding dental plates; muscle field large, wide and flabellate,
lobate anteriorly and leaving adductor scars open to the anterior.
Pallial marks consisting of numerous anteriorly directed channels.
Brachial valve interior with deep, coarsely corrugated sockets and
overhanging, thick socket ridges; crura of radulifer type moderately
long, horizontally flattened and attached to socket ridges without
outer hinge plates. No inner hinge plates. Pallial marks as in the
pedicle valve. Cardinal process transversely widely triangular,
thickened and somewhat elevated. Median ridge short, low, not reach-
ing the apex.
Type species——Terebratula nigricans Sowerby, Proc. Zool. Soc.,
p. 91, 1846.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 49
Comparisons —This genus was hitherto placed under Tegulorhyn-
chia and was generally regarded as the reference species for that
genus because it is a Recent as well as fossil form, the interior details
of which are well known. Significant differences between this and the
type species of Tegulorhynchia make it impossible to keep the two in
intimate association. The exterior ornamentation of the two is quite
distinct, Tegulorhynchia having the strongly squamose or spinose ex-
terior whereas Notosaria is costellate but with only fine growth lines.
Another exterior difference of importance is the presence in No-
tosaria of disjunct deltidial plates. Tegulorhynchia has conjunct
deltidial plates and an entire foramen. Inside the pedicle valve the
pedicle collar of the modern form is not well developed. A major
difference appears inside the brachial valve of Tegulorhynchia. In
that genus the median septum extends posteriorly to unite with ex-
tensions from the crural base to form a thickened plate at the pos-
terior. This is illustrated by Chapman and Crespin (1923) for
T. coelata, plate 12, figure 17. In shells of modern Notosaria the
median septum is not extended to the apex.
Geological horizon.—Miocene to Recent.
Distribution—New Zealand and Kerguelen Island; Belgium.
Assigned species:
Rhynchonella nigricans Sowerby, Miocene to Recent, New Zealand.
R. nigricans pyxidata Davidson, Recent, Kerguelen Island.
R. nysti Davidson, Pliocene, Belgium.
Hemithyris sublaevis Thomson, Miocene, New Zealand.
Discussion—It may come as a surprise that the group of shells
so long associated under the generic name of Tegulorhynchia could
be separated, but the differences in ornamentation, beak characters,
and cardinalia are sufficient. The interior differences of significance
are in the pallial markings and cardinalia.
As indicated on plate 6, B, figures 12 and 14, the pallial markings
in both valves of Notosaria are entirely different from those figured
by Leidhold (1922, pl. 11) for Tegulorhynchia déderleini (this
monograph pl. 22, C, figs. 16 and 17). In Notosaria the vascula media
cannot be easily distinguished on the inner shell surface and the
pallial marks make numerous short trunks extending anteriorly and
anterolaterally from the muscle and ovarian fields. The latter is also
not distinctly impressed but seems to be a quite narrow crescent in
the pedicle valve but somewhat wider in the brachial valve. The
pallial trunks of Tegulorhynchia as figured by Leidhold are like those
common to many other genera illustrated herein.
50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
The cardinalia of the two genera are similar except for the median
ridge. In Notosaria the median ridge is short and thick and is usually
on a small callosity between the posterior adductors. The ridge does
not extend to the apex which is generally smooth. In Tegulorhynchia
on the other hand the median septum is short but extends to the apex
where it meets short extensions from the crural bases which form a
small apical callosity. The cardinal process of Notosaria is well de-
veloped but that of Tegulorhynchia can scarcely be distinguished.
Rhynchonella nysti Davidson from the Pliocene of Belgium was
referred by Thomson (1927, p. 154) to Tegulorhynchia with the re-
mark that Davidson (1874a, p. 7) had compared the species to
Tegulorhynchia nigricans. Comparison of the interior and exterior
details corroborates this assignment and comparison. The beak
characters of a pedicle valve in the national collection (U.S.N.M.
549417a) has the characteristic foramen and deltidial plates of No-
tosaria. The cardinalia, too, are like those of Notosaria as shown
by a brachial valve (U.S.N.M. 549417b). The sockets are large
and the socket plates broad and strong. The cardinal process is a
thickened triangular callosity like that of the New Zealand species.
These features combined with the exterior ornament clinch the as-
signment. hynchonella nystt is costate and the costae bifurcate at
places on the valve as in the New Zealand shell, a feature unusual in
the Rhynchonelloidea. This occurrence, as Thomson remarks, leads
to interesting speculation on the paleogeographic distribution of
Notosaria. It is possible that the Austral members of the genus
originated in European waters and thus constitute a clearly distinct
stock from Tegulorhynchia as its anatomy suggests.
Genus TEGULORHYNCHIA Chapman and Crespin, 1923
Plates 5, D, 21, E
Tegulorhynchia Chapman and Crespin, Proc. Roy. Soc. Victoria, n. s., vol. 35,
pt. 2, p. 175, 1923; Thomson, New Zealand Board Sci. Art, Manual 7, p.
152, 1927.
Transversely triangular to subpentagonal in outline; inequivalve,
the brachial valve having the greater depth; anterior commissure
rectimarginate, the brachial valve having a moderately well-defined
fold; surface costellate and lamellose, the lamellae being produced
into hollow spines in some species. Beak of pedicle valve long and
pointed ; foramen complete in the type of the genus (Allan, 1940,
p. 279). large, hypothyrid ; deltidial plates usually conjunct.
Pedicle valve interior with strong corrugated teeth supported by
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 51
short, receding dental plates; muscle field large, located anterior to
the delthyrial cavity; diductor scars of moderate size, surrounding
the large adductor scars; adjustor scars large; pallial sinuses sparse,
with the vascula media short and branching near midvalve, one
branch continuing anteromedially, the other laterally and posteriorly
(plier, Ey fig.15).
Brachial valve interior with small cardinalia having strong and
elevated socket ridges, no outer hinge plates and no inner hinge
plates ; crura short, of radulifer type, stout, curved; median septum
short, low, and meeting the crural bases at the apex; diductor at-
tachments a pit at the apex. Adductor field small.
Type species (by original designation).—Rhynchonella squamosa
Hutton, Cat. Tertiary Mollusca and Echinodermata of New Zealand,
P. 37, 1873.
Comparison.—The squamose to spiny ornamentation of this genus
makes it one of the most conspicuous of modern and Tertiary shells.
The distinctive ornamentation is, however, only one means of dif-
ferentiation from other genera. Interior differences also exist in the
form of the moderately short crura and apical callosity formed by
the crural bases. This and the difference in the pallial markings are
means of distinction from Notosaria which is most like Tegulorhyn-
chia.
Geological horizon.—The type species and most other species of
Tegulorhynchia are found in the fossil state. The genus ranges from
Oligocene into the Recent where it is represented by T. doderleim
(Davidson).
Distribution—The fossil species occur chiefly in the Southern
Hemisphere in the southern part of Australia and New Zealand.
One fossil form, identified as T. ddderleini occurs in the Pliocene of
Okinawa.
The geographic range of the one living species T. déderleini is
from Japan south to Borneo.
Assigned species——The species of Tegulorhynchia are:
Rhynchonella squamosa Hutton, Miocene, New Zealand.
R. tubulifera Tate, Miocene, Australia.
R. déderleini Davidson, Recent, Japan to Borneo.
Hemithyris antipoda Thomson, Miocene, New Zealand.
? H. depressa Thomson, Miocene, New Zealand.
H. squamosa Buckman (not Hutton), Miocene-Oligocene, Antarctic.
H. imbricata Buckman, Miocene-Oligocene, Antarctic.
Tegulorhynchia thomsoni Chapman and Crespin, Miocene, Tasmania.
T. coelospina Chapman and Crespin, Miocene, Tasmania.
52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
T. coelata (Tension-Woods), Oligocene to Miocene, Tasmania and
Australia.
T. masoni Allan, Miocene, New Zealand.
Discussion.—As here limited, the name Tegulorhynchia is applied
only to imbricate species with cardinalia and hinge characters like
those of T. squamosa.
Genus PLICIRHYNCHIA Allan, 1947
Plate 7, A
Plicirhynchia Allan, Journ. Paleont., vol. 21, No. 5, p. 493, 1947.
Subtriangular to subpentagonal in outline with the maximum width
near the middle; inequivalve, the brachial valve having the greater
depth; anterior commissure uniplicate, the fold of the brachial valve
being conspicuous and fairly long; surface ornate, the posterior half
being marked by fine radial lines and striae, the anterior half strongly
costate. Beak of the pedicle valve long, narrowly pointed, and nearly
straight; foramen complete, large, longitudinally oval, hypothyrid ;
deltidial plates thick and conjunct.
Pedicle valve interior with thick corrugated teeth supported by long,
stout dental plates; pedicle collar long; muscle field large and
flabellate, enclosing the adductor scars.
Brachial valve interior with corrugated sockets bounded by thick
overhanging socket ridges; crural bases attached directly to the
socket ridges; crura of radulifer type, long, horizontally flattened ;
inner hinge plates strongly developed. Cardinal process thick and
bilobed. Median ridge small.
Type species (by original designation) —kRhynchonella plicigera
von Ihering, Rev. Mus. Paulista, vol. 2, p. 270, text fig. 7, 1897.
Comparisons.—This genus is characterized by its peculiar exterior
ornament, the posterior and umbonal regions being finely costellate
but the anterior half strongly costate. The only other semicostate
genus marked like this is Probolarina, but from that genus it differs
in beak characters, cardinalia, and the presence of a cardinal process.
Geological horizon —Plicirhynchia occurs in the Eocene (Pata-
gonian) of Argentina.
Assigned species—Only one species is assigned here with assur-
ance:
Rhynchonella plicigera von IThering.
? Hemithyris plicigera Buckman, not von Ihering.
Distribution—Known only from Argentina and possibly from the
Antarctic.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 53
Discussion.—The exterior characters of this genus, except for the
anterior costation, are most like Hemuithyris. The general form of
the shell, the obscure, fine costellation of the umbonal region, and
the large foramen are suggestive of the northern genus. The deltidial
plates are conjunct, however, and there the resemblance ends.
The specimens available for study of the interior are not good
because the muscle marks are obscure and the cardinalia partially
broken. Nevertheless some important details can be distinguished.
Inside the pedicle valve the pedicle collar is fairly long and slightly
elevated above the valve floor. The dental plates are solid but of the
receding type. They are separated from the valve walls by moderately
deep and wide umbonal cavities. The muscle field is large and reaches
to about midvalve, possibly somewhat beyond in old specimens.
Inside the brachial valve the cardinalia are stout and thick. The
cardinal process is a thick, bilobed boss at the apex. The socket
ridges are thick and the crura are long, slender, and flattened hori-
zontally. These features are well shown in specimen U.S.N.M.
549423a. The crural bases are attached directly to the socket ridges
without outer socket plates. An inner thickening along the edge of
the crural bases suggests some development of inner hinge plates.
Median ridge small and inconspicuous. The cardinalia appear to be
related to those of Hemithyris, Notosaria, and Tegulorhynchia.
According to Jaanusson (1951, p. 196) the shell referred by Buck-
man (1910, p. 12) to Hemithyris plicigera should be referred to
Plicirhynchia. Jaanusson also points out in connection with Hemithy-
ris dibbleei and H. reagani, both of Hertlein and Grant (1944), that
these do not belong to Hemithyris but, because of their anterior
costation, may be assigned to Plicirhynchia. Unfortunately the in-
terior details of these two species are unknown and the suggested
assignment can only be tentative (see discussion of Hemithyris).
FRIELETIDAE Cooper, new family
Genus FRIELEIA Dall, 1895
Plates 4, B, 14, B, 15, A, 21, A
Frieleia Dall, Proc. U. S. Nat. Mus., vol. 17, p. 713, 1895; Thomson, Geol. Mag.,
n. s., dec. 6, vol. 2, pp. 389, 392, 1915; Jackson, British Antarctic (“Terra
Nova”) Exped., 1910, Nat. Hist. Rep., Zool., vol. 2, No. 8, pp. 192, 193, 1918;
Thomson, New Zealand Board Sci. Art, Manual 7, p. 157, 1927; Hatai, Sci.
Rep. Tohoku Imp. Univ., ser. 2 (Geology), vol. 20, p. 219, 1940; Hertlein
and Grant, Publ. Univ. California at Los Angeles, Math. and Phys. Sci., vol.
3, DP. 57, 1944.
54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Elongate oval to subtriangular in outline, with the greatest width
at or anterior to the middle; thin shelled; inequivalved, the pedicle
valve having the greater depth and convexity; rectimarginate to
ligate ; surface smooth to obscurely and minutely costellate. Beak of
pedicle valve short, nearly straight to suberect; foramen incomplete,
elongate oval, hypothyrid; deltidial plates thick, disjunct but nearly
united ; apex marked by a small triangular plate elevated above the
valve floor.
Pedicle valve interior with long, curved, corrugated teeth buttressed
by prominent and strong dental plates; muscle and pallial marks
lightly impressed; diductor field subquadrate, small, surrounding
adductors ; vascula media branching at about midvalve, the branches
diverging anteromedially and anterolaterally.
Brachial valve interior with deep corrugated sockets margined by
thick overhanging socket ridges; crura, of spinulifer type, thin, long,
divergent, attached directly to the socket ridges; inner hinge plates
small, rounded, disjunct or coalesced at the posterior to form a
central bilobed plate; median septum long, slender, elevating pos-
teriorly and united to the inner hinge plates to form a small V-shaped
chamber which may be filled by callus in old specimens. Median
septum rising to a crest just anterior to the apex; cardinal process
a small, triangular, transversely striated pit at the apex; adductor
scars long and narrow, posteriorly situated.
Type species —Frieleia halli Dall, Proc. U. S. Nat. Mus., vol. 17,
p. 714, pl. 24, figs. 6, 9-13, 1895.
Comparisons.—This is a thin-shelled, fragile brachiopod with both
valves somewhat sulcate in the Recent species and characterized by a
great development of inner hinge plates and a camerate apex in the
brachial valve. It is unlike all other described genera in these re-
spects.
Geological horizon.—Possibly Miocene to Recent. Species from
the California Tertiary now referred to Hemuithyris or other genera
may belong here.
Assigned species:
Frieleia halli Dall, Recent, West Coast North America, Japan.
? Terebratula nitens Conrad = Hemithyris astoriana Dall, Miocene,
Oregon.
Hemithyris pellucida Yabe and Hatai, Recent, Japan.
Distribution—Known from Alaska to California, Japan, and Kam-
chatka in waters ranging from 21 to 1,059 fathoms.
Discussion—The important exterior features of this genus are
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 55
the obscurely costellate surface, the rectimarginate anterior com-
missure, and the disjunct deltidial plates. In most specimens the
median portion of one or both valves is marked by a depressed line or
flattening that produces an emargination of the anterior. The deltidial
plates are usually strongly developed but have not been observed to
meet.
In the apex of the pedicle valve a small triangular plate appears
which is elevated above the valve floor. This forms a partial sheath
for the pedicle which rests against it. The teeth, as in most modern
rhynchonelloids, are corrugated and are supported by well-developed,
erect dental plates defined by deep umbonal cavities. The muscle
field is small, with a large subquadrate flabellate diductor field sur-
rounding the adductors anteriorly. The adjustor scars are small and
are located just anterior to the front of the dental plates. Faint
pallial marks preserved in one specimen show the vascula media
as in Hispanirhynchia.
The most interesting parts of Frieleia are the cardinalia. The
diductor muscles are attached in a small, triangular, horizontally
striated pit at the apex. In some specimens this is much thickened to
form a cardinal callus. The socket ridges are strong and curved. To
them are attached small triangular outer hinge plates. The hinge
plates bear the crural bases and crura. The crural bases are further
strengthened by inner hinge plates that extend dorsally to unite with
the median septum to form a small chamber. I am unable to detect
any substantial resemblance of this structure to that of Camarotoechia
or even to the camerate Mesozoic rhynchonelloids.
The median septum in Frieleia is a narrow, strong, elevated plate
that is a myophragm and a crural buttress. It is interesting to note
that the inner hinge plates, in decking over the space between the
crural bases, do not form a septal chamber as in Camarotoechia but
fill in the space solid. In some specimens the inner hinge plates coa-
lesce in such a way as to form an undivided but concave hinge plate.
The adductor field is divided by the median septum and is long and
slender. The anterior scars are elongate, tear shaped in outline. The
posterior pair is smaller and lies posterolateral to the anterior pair.
Frieleia has not yet been definitely identified in the Tertiary of
California or Japan. It has distinctive characters and is one of the
few modern or Tertiary brachiopods having a prominent median
septum. Several species occurring on the Pacific Coast of the United
States may be referable to Frieleia, especially if the definition were
to be broadened to some extent. The so-called Hemuthyris astoriana
56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Dall (=Terebratula nitens Conrad) has interior characters strongly
suggesting Frieleia, especially the strong median septum in the
brachial valve. The species is fairly strongly uniplicate, however,
which is not in accordance with the current definition of Frieleia.
All the specimens of H. astoriana available for study, including the
type specimen, are badly exfoliated. The exterior is therefore not yet
wholly known. The exfoliated shells have fairly strong radial costel-
lae, but these may be only a feature of the exfoliated shell. A cross
section of the beak of the brachial valve reveals a small triangular
chamber. No modern specimens of Frieleia are uniplicate. The
specimens of H. astoriana are here referred to Frieleia with a query.
They are nearer that genus than they are to Hemithyris. Ultimately
it may be necessary to erect a new genus for uniplicate Frieleia if
specimens good enough for detailed description are brought to light.
Genus COMPSOTHYRIS Jackson, 1910
Plate 16
Compsothyris Jackson, British Antarctic (“Terra Nova”) Exped., 1910, Nat.
Hist. Rep., Zool., vol. 2, No. 8, p. 188, 1918; Thomson, New Zealand Board
Sci. Art, Manual 7, p. 161, 1927.
Rounded triangular in outline with the greatest width at about the
middle; valves subequal in depth, the pedicle valve having a greater
depth than the brachial valve ; anterior commissure broadly and gently
uniplicate, the brachial fold inconspicuous. Surface marked by fine
radial costellae. Beak of pedicle valve nearly straight to suberect,
bluntly pointed; foramen incomplete, of moderate size, elongate
elliptical, hypothyrid (permesothyrid according to Jackson, 1918) ;
deltidial plates disjunct.
Pedicle valve interior with small teeth supported by strong dental
plates ; muscle field located well anterior to the delthyrial cavity, small ;
diductor scars small, surrounding the adductor pair. Pallial marks
not impressed.
Brachial valve interior with narrow corrugated sockets bounded by
strongly overhanging socket ridges; crura of spinulifer type, short,
attached to the socket ridges by narrow hinge plates. Inner hinge
plates incipiently developed. Median ridge or myophragm slender,
moderately elevated and reaching the apex where it is divided and
supports the proximal ends of the crural bases. Adductors closely
crowded against the myophragm, the right and left pairs being tear
shaped in outline.
Type species (by original designation).—Rhynchonella racovitzae
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 57
Joubin, Résultats voyage S. Y. Belgica, 1897-1898-1899, Zool., Rapt.
Sci. Commiss. Belgica, p. 5, pl. 1, figs. 1-4, 190I.
Comparisons.—The genera to which Compsothyris can profitably
be compared are: Frieleia, Hispanirhynchia, and Grammetaria. The
differences between Compsothyris and Frieleia are chiefly exterior
characters but the cardinalia also vary importantly. Frieleia is not
so strongly and evenly costellate as Compsothyris and the shape and
folding of the two are quite different. In Frieleia it is common for
both valves to have a sulcus and for the front to be emarginate.
Compsothyris is faintly uniplicate. Furthermore, Frieleia is much
more triangular than Compsothyris and has a great development of
inner hinge plates in the cardinalia of the brachial valve, a feature
not shared by the Antarctic shell.
Hispanirhynchia differs in shape from Compsothyris, having a
compressed profile and strongly triangular outline. The two genera
are similarly marked on the exterior however. Inside the brachial
valve only a slight development of inner hinge plates appears in
Compsothyris, and the small chamber at the apex is not obliterated by
shell growth in the adults as it is in Hispanirhynchia.
Compsothyris differs from Grammetaria in the form of the valves
and in the less elaborate deltidial plates. Compsothyris is nearly
circular in outline whereas Grammetaria is strongly triangular. The
deltidial plates of Grammetaria are elaborately auriculate but those of
the Antarctic genus are small and not auriculated. Inside the brachial
valve no inner hinge plates are developed by Grammetaria but the
small apical chamber present in the young is sealed off in the adult
by the sides growing shut. In adult Compsothyris this chamber
remains open.
Geological horizon.—Not known in the fossil state.
Distribution—Ross Sea area and western Antarctic in depths
ranging from 45 to 300 fathoms.
Assigned species—So far only one species can be definitely as-
signed here but two others may belong:
Rhynhonella racovitzae Joubin, Recent, Antarctic.
? Hemithyris striata Thomson, Recent, Antarctic.
? Rhynchonella valdiviae Helmcke, Recent, Indian Ocean.
Discussion.—Jackson (1918, p. 193) expressed interest over the
fact that the features of Compsothyris and Frieleia were suggestive
of certain Paleozoic genera, especially Camarotoechia. Comparison
with interiors of Camarotoechia (see pl. 4, D, figs. 6-8), however,
show the relationship to be quite remote because the structures in
58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
the two genera, which look similar, actually are developed differently.
In the Paleozoic genus the median septum is strong and high and
supports a V-shaped chamber having strong walls. This is in turn
covered by a flat plate connecting the crural bases. This plate is ap-
parently built, at least in part, posteriorly because it does not gen-
erally close the chamber but leaves a small round cavity at the apical
end. This chamber of Camarotoechia is more like the structure in
Septaliphoria. The chamber of this Jurassic shell is, however, also
different from that of Camarotoechia, although the two look alike.
The chamber of Compsothyris is a much more delicate affair and, it
seems to me, not related to any Paleozoic form.
GRAMMETARIA Cooper, new genus
(Gr. gramme, line)
Plates 4, C, 9, A
Outline elongate triangular with the maximum width at the ante-
rior; valves subequal in depth, the pedicle valve having a slightly
greater depth ; anterior commissure rectimarginate ; surface marked by
fine costellae. Beak small, bluntly pointed, suberect ; foramen incom-
plete, rounded, hypothyrid ; deltidial plates auriculate, conjunct.
Pedicle valve interior with small corrugated teeth, supported by
strong vertical dental plates. Muscle field small and subcircular ;
diductor scars small; adjustor scars large, posterolaterally placed.
Brachial valve interior with corrugated sockets bounded by strong,
thick socket ridges; crura short, of spinulifer type, triangular in sec-
tion but laterally flattened distally, attached to the socket ridges by
very narrow, inconspicuous outer hinge plates; median ridge thick,
not quite reaching the apex in the adult, but in the young forming
a low, wide V-shaped chamber with the crural bases; V-shaped
chamber covered by shell substance in the adult. Adductor field
elongate triangular, the anterior and posterior scars on each side of
the median ridge tear shaped in outline; posterior set of adductors
located outside the anterior set.
Type species—Hemithyris bartschi Dall, Proc. U. S. Nat. Mus.,
vol. 57, p. 289, 1920.
Comparison.—This genus is most suggestive of Frieleia and
Compsothyris among described genera, but it has important differ-
ences from both of them. In the first place, Frieleia has a strong
tendency to ligation while Compsothyris is broadly uniplicate. Gram-
metaria, on the other hand, is rectimarginate. The deltidial plates of
the pedicle valve of Grammetaria are elaborately auriculate but such
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 59
features have not been seen in the other two genera. The deltidial
plates of Grammetaria are conjunct but those of the other two genera
are disjunct. However, those of Frieleia nearly meet.
The interior of the brachial valve is the significant part of each
of these genera. In Frieleia the crural bases are attached to the
median septum but in addition a strong development of inner hinge
plates may create a small chamber at the apex. In Compsothyris the
crural bases are likewise supported by the median septum but no
comparable development of the inner hinge plates is known. In
Grammetaria the very young are similar to Compsothyris in having
the crural bases united to the median septum but the apical V-shaped
chamber thus formed is much broader and shallower than that of
Compsothyris. In the adult of Grammetaria the spaces between the
septum, shell wall, and the broad chamber are filled to form a thick
apical callosity between the crura. Thus the low septum ends in a
callosity at the rear of the adult shell.
Assigned species—At present only Hemithyris bartschi Dall is
known in this genus.
Distribution —Hemuthyris bartscht is known only from Philippine
waters from depths of 161 and 298 fathoms.
Discussion.—Only two specimens of this genus are known but they
indicate a brachiopod having several unusual characters. Although
the deltidial plates of the adult specimen are broken, probably in sepa-
rating the valves, those of the young specimen are quite definitely con-
junct even though broken slightly at their line of junction. This is
one feature that distinguishes this genus from Frieleia and Comp-
sothyris. Another feature of considerable interest is the development
of the brachial interior from a camerate brachial valve to one having
only a callosity at the posterior. The small camera of the young is
buried in callus as the valve grows and is completely obliterated. This
takes place to a lesser extent in Hispanirhynchia. The crura of
Grammetaria are more like those of Frieleia in not having prominent
outer hinge plates developed. In Compsothyris modest but definite
outer hinge plates are present, making the crura more suggestive of
Basiliola than of Frieleia.
HISPANIRHYNCHIIDAE Cooper, new family
Genus HISPANIRHYNCHIA Thomson, 1927
Plates 10, 13, A, 21, G
Hispanirhynchia Thomson, New Zealand Board Sci. Art, Manual 7, p. 159, 1927.
Outline elongate triangular with the greatest width in the anterior
third; inequivalve, the pedicle valve being deeper than the brachial
60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
valve; anterior commissure rectimarginate to ligate to slightly uni-
plicate ; surface marked by concentric lines of growth and fine obscure
radial costellae; beak of the pedicle valve short, suberect ; foramen
incomplete, large, hypothyrid ; deltidial plates small, disjunct.
Pedicle valve interior with well-developed but incomplete pedicle
collar and thick teeth supported by small, somewhat receding dental
plates; muscle field small, rounded, with small diductor scars sur-
rounding large adductor scars. Vascula media prominent, originating
between diductor and adjustor scars, extending anteriorly to branch
about one-third the length from the front margin.
Brachial valve interior with corrugated sockets and overhanging
socket ridge to which the short, bladelike, spinulifer crura are attached
by small and narrow outer hinge plates. Inner hinge plates small and
narrow. Median ridge low, thick, extending to the apex. Adductor
field small, divided by a low median ridge; anterior adductors rounded,
posterior pair elongated. Vascula media widely divergent.
Type species (by original designation).—Rhynchonella cornea
Fischer, in Davidson, Trans. Linnaean Soc., ser. 2, vol. 4, Zool., pt. 2,
p. 171, pl. 25, figs. 2-4, 1887.
Comparisons—This is one of several triangular or nearly trian-
gular genera with faint radial ornamentation. It differs from Frieleia
and Compsothyris in not having a strongly camerate apex in the
brachial valve of the adult. It differs from Grammetaria in its less
prominent radial markings, nonalate deltidial plates, and the develop-
ment of the cardinalia which are camerate in the young of Gram-
metaria. Aetheia can be readily distinguished from Hispanirhynchia
by its small foramen, concave deltidial plates and the great develop-
ment of inner hinge plates on the interior.
Distribution—In modern seas Hispanirhynchia is known from off
the coasts of Morocco, the Sudan, and the Canary Islands. It is also
known from west of Cape Finistere, northwestern Spain. It is gener-
ally found in deep water, from 5774 fathoms ? off Cape St. Vincent,
Portugal, to below 1,000 fathoms off the coast of Spain. One species,
H. ?craneana (Dall) doubtfully assigned, taken off Cocos Island,
Panama, came from 117 fathoms.
Geological horizon.—Possibly present in the Eocene of Cuba.
Assigned species——Two Recent species are assigned to this genus:
Rhynchonella cornea Fischer, Recent, North Atlantic.
? Hemithyris craneana Dall, off Panama, Pacific Ocean.
Hispanirhynchia sp., Eocene, Cuba.
2 See note by Jackson (1918, p. 192, footnote).
NO. 5 RHYNCHONELLOID BRACHIOPODS—-COOPER 61
Discussion.—External features of importance in this genus are the
beak characters, the ornamentation, and the anterior commissure. In
the type species the deltidial plates are well developed and disjunct
but in some specimens approach each other very closely. Dall de-
scribes the deltidial plates of H. ?craneana as “obsolete” but the speci-
men has definitely been damaged in the beak region. In some old
specimens of H. cornea these plates are also lacking, possibly due to
abrasion.
The shell of young specimens of Hispanirhynchia is generally
translucent and a pale brown. Adults are opaque and a deeper brown
in color. The surface is minutely costellate, the costellae extremely
fine and very closely crowded.
Specimens of H. cornea are generally rectimarginate but Thomson
(1927, p. 159) speaks of some as being ligate, that is, with a gentle
depression in each valve which will produce an emarginate anterior.
The anterior commissure of H. ?craneana, on the other hand, has a
slight wave in it toward the pedicle valve, thus producing a faint
sulcation.
The interior of the pedicle valve of the mature to old shells usually
shows the details to perfection because the muscles and pallial marks
are deeply impressed. The pedicle collar is well developed and may be
elevated above the valve floor. The teeth are strong and corrugated.
The dental plates are strong and separated from the lateral shell wall
by deep cavities. In old shells these tend to become nearly obliterated
by deposition of shell substance inside the cavities.
The muscle field is small. The flabellate diductor scars are small
and surround a fairly large adductor patch anteriorly. The adjust-
tor scars are deeply impressed at the anterior edge of the dental
plates. Accessory diductor scars are not visible in the delthyrial cavity.
The vascula media take off anterior to the adjustor scars and along the
outside of the diductor scars. The main trunk branches at about mid-
valve, one branch extending laterally, the other anteromedially. Both
of these branches bifurcate again to produce distributaries anteriorly
and laterally.
The genital area is small and located on the shell wall just anterior
to the dental plates. This area is smaller than that in Basiliola and
Rhytirhynchia.
The cardinalia of this genus are interesting because the young show
features that are buried by excess shell in the adult. The insertion of
the diductor muscles appears as a small, triangular, horizontally stri-
ated pit at the apex. No swollen cardinal process is present as in
62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Plicirhynchia. The socket ridges are thick and curved; the crura are
attached to them by small, triangular outer hinge plates. The crura
are laterally compressed blades, not concave in section and their distal
end is serrated. Prominent inner hinge plates are formed at the apical
end of the shell which attach to the floor of the valve. With the
septum they form a poorly defined chamber but in old specimens the
inner plates become thickened at the apex and fuse to form a thick
callus. In such cases the extensions of the inner plates to the valve
floor are obscured.
The adductor field is small and elongated. The anterior pair of scars
is the larger and the posterior pair more elongated, at least in the old
shells. The field is divided medially by a short, thick median ridge
which extends no farther than the anterior end of the adductor field.
At the anterior end of the ridge a small scar appears in old specimens.
The genital areas are small like those of the pedicle valve. The vascula
media originate at the inside ends of the anterior adductors and sur-
round the small scar mentioned above. These pallial trunks divide
near midvalve in a manner similar to that of the pedicle valve.
Comparison of Hispanirhynchia ?craneana with middle-aged speci-
mens of H. cornea show slight differences but they do not appear to be
great enough to exclude the species from Hispanirhynchia. The outer
hinge plates of the brachial valve appear slightly wider than those of
the Atlantic shells and the inner hinge plates are not so well developed,
but they are there.
Rhynchonella sicula Seguenza, here made the type of the genus
Sphenarina, was early identified with H. cornea, but examination of
the interior of the Sicilian shell makes it clear that the two have little
in common but shape and ornament. Sphenarina has no median
septum and its beak characters are different from those of Hispami-
rhynchia.
SPHENARINA Cooper, new genus
(Gr. sphenos, wedge)
Plates 5, B, 8, A
Shell triangular in outline, with the greatest width in the anterior
third ; subequivalve, the pedicle valve having a slightly greater depth
than the brachial valve ; anterior commissure rectimarginate; surface
marked by minute radial lines. Beak long, suberect ; foramen small,
circular, hypothyrid; deltidial plates conjunct, elaborately auriculate.
Pedicle valve interior with short pedicle collar and well-developed
dental plates with wide umbonal cavities.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 63
Brachial valve interior with prominent socket ridge to which the
short crural bases are attached without outer hinge plates. Crura
moderately long, of spinulifer type, nearly straight, compressed to
slightly crescentic in section and with distal extremities flattened; no
inner hinge plates. Posterior of crural bases attached to floor of valve
by short plates ; cavity between plates occupied by callus, thus making
the apex solid. Median ridge or septum absent; adductor field
elongate.
Type species—kRhynchonella sicula Seguenza, in Davidson, Geol.
Mag., vol. 7, No. 76, p. 461, pl. 20, fig. 6, 1870.
Comparisons.—This is a wedge-shaped form with fine radial orna-
mentation comparable to Hispanirhynchia, Grammetaria, and Comp-
sothyris. It differs from all these in the nature of the cardinalia. On
the inside of the brachial valve the cardinalia of Sephenarina differ
from all three in the almost total absence of a median ridge or septum
and in the fact that the plates supporting the crural bases at the apex
meet the floor of the valve directly.
Geological horizon.—Pliocene of the Mediterranean region.
Assigned species.—The following species are placed in this genus:
Rhynchonella sicula Seguenza.
R. soricina Defrance= R. sicula Seguenza.
? Hemithyris eotrigona Sacco and variety obliquatella Sacco.
Discussion——This species has commonly been referred to Hispani-
rhynchia because of the close similarity of form and ornamentation.
In fact Jeffreys (1878, p. 413) identified dredged specimens of the
latter as identical with the Italian species. Examination of the beak
and brachial valve interior of FR. sicula will dispel the idea of identity
almost immediately.
The material of S. sicula showing interior details is scanty. The two
specimens in the National Museum from which the above description
was drawn were prepared by needles, a delicate operation considering
the thin shell of the species and the fragile nature of the cardinalia.
The length of the crura is moderate and the ends are flattened later-
ally, strongly suggesting the crura of Frieleia.
The cardinalia of Sphenarina are suggestive of those of Hispani-
rhynchia but the median septum is lacking. The plates supporting the
crural bases thus rest directly on the valve floor rather than on the
median septum. A young specimen dissected shows no trace of a
septum and no evidence of supporting plates for the crural bases.
64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
ERYMNARIIDAE Cooper, new family
ERYMNARIA Cooper, new genus
(Gr. erymnos, fenced)
Plates 18, A, B, 19, A, 22, B
Outline irregular triangular to rounded pentagonal, usually with the
greatest width at or anterior to the middle; inequivalve, the brachial
valve having the greater depth; anterior commissure irregular, twisted
or regularly uniplicate; surface smooth or marked by concentric
growth lines and short, irregular costae occupying the anterior third or
half. Beak of pedicle valve short, deltidial plates conjunct, slightly
auriculate ; foramen small to moderately large, oval, hypothyrid.
Pedicle valve with short dental plates defining a deep delthyrial
cavity; muscle field small, with small adductor scars surrounded by
subflabellate diductor scars. Vascula media short.
Brachial valve with large, deep, corrugated sockets; socket ridges
elevated and strong; outer hinge plate broad; crura of septifer type,
curved, supported by two long septa that extend along the valve floor
for about one-fifth the valve length. Vascula media thin, moderately
long.
Type species —Terebratula polymorpha Massalongo, Schizzo geog-
nostico sulla valle del Progno o Torrente D’Illasi, con un saggio sopra
la flora primordiale del M. Bolca, Verona, pp. 18, 19, 1850.
The septifer type of crura are not well known but have been recog-
nized in the Jurassic. Rothpletz recognized two groups or Sippe hav-
ing septifer crura. One of these is the Inversa-Sippe in which the
species have a sulcate anterior commissure and are smooth or semi-
costate ; the other group is the Trigona-Sippe in which the shells are
rectimarginate to uniplicate and are wholly costate.
Septocrurella of Wisniewska is a paucicostate genus having a sul-
cate anterior commissure and the septifer type of crura. Ahynchonella
deluxa Oppel, which is similar exteriorly to Septocrurella sanctaclarae
Wisniewska, also has the septifer type of crura.
No Cretaceous rhynchonelloids having this structure are now known
to me, but the fact that septifer genera appear in the Jurassic and
Focene indicate the strong likelihood that specimens with this structure
occur in the Cretaceous. It is interesting to note that the known
Eocene septifer genus is smooth pauciplicate but is uniplicate rather
than sulcate.
Comparison.—The exterior form of two species of Erymnaria is
like that of Streptaria in having the strongly twisted anterior com-
NO. 5 RHYNCHONELLOID BRACHIOPODS—-COOPER 65
missure, but there the similarity ends. The interior of Erymnaria is
so unlike that of Streptaria that confusion of the two is not possible.
Geological horizon.—Eocene of Italy and Cuba.
Distribution—Two species of this genus are known in the Eocene
of northeastern Italy and one at the same horizon in Cuba. Only one
specimen is known from the latter occurrence but the interior details
visible through the shell make the identification with this genus quite
certain.
Assigned species—Three species of this genus are now known:
Terebratula bolcensis Massalongo, Eocene, Italy.
T. polymorpha Massalongo, Eocene, Italy.
Erymnaria cubensis Cooper, new species, Eocene, Cuba.
Discussion.—The genus is characterized by having strong support-
ing plates buttressing the crura and constituting the septifer type of
Rothpletz. It is the only Tertiary genus known to me having this
peculiar structure. The supporting plates of the crura make two
long, dark suture lines diverging from the beak. In several specimens
the socket plates are also visible as dark lines on the inner filling of
the shell. In such cases the socket plates occupy the outside and are
shorter than the crural supports.
ERYMNARIA CUBENSIS Cooper, new species
Plate 19, A
Shell of about median size for a rhynchonelloid, slightly wider than
long; subpentagonal in outline; widest at midvalve; sides narrowly
rounded ; beak forming an angle of 100° ; anterior margin truncated.
Anterior commissure strongly uniplicate ; surface marked only by con-
centric growth lines.
Pedicle valve evenly and gently convex in lateral profile ; nearly flat
in anterior profile with margins abruptly bent dorsally ; umbo some-
what narrowly swollen ; median region flattened ; sulcus originating at
about midvalve, broad and shallow ; tongue moderately long, narrowly
rounded, and bent nearly at right angles to the lateral commissure ;
flanks bounding sulcus narrow, gently convex, and moderately steep.
Beak small, rounded ; beak ridges not prominent ; deltidial plates con-
junct; foramen moderately large, longitudinally elliptical, and with
the anterior margin having a small lip.
Brachial valve deeper than the pedicle valve; gently convex in lat-
eral profile but narrowly domed in anterior profile, the sides long and
steep. Umbo and median region swollen; fold originating anterior to
midvalve, low and gently rounded, scarcely protruding beyond the
flanks ; sides steep, gently inflated.
66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
MEASUREMENTS IN MILLIMETERS
Brachial
Length length Width Thickness
EALOIGEV DE. ava Nre whessiiers Lisette Reena 14.1 12.8 15.0 10.0
Types.—Holotype, U.S.N.M. 549385.
Horizon and locality.—Eocene, 80 meters northeast of school,
Chucho Machin, Matanzas Province, Cuba.
Discussion.—Only a single complete specimen is known of this in-
teresting species, but it is well preserved and some of the interior
details are visible through the transluscent shell. It is most like T.
bolcensis (Massalongo) in its symmetrical form and folding, but
differs in having a broader and less narrowly folded anterior commis-
sure, a larger foramen, and the crural supporting plates seem to be
somewhat shorter than those of the Italian species.
That this species belongs to the Italian genus seems certain because
the crural supporting plates and socket ridges are clearly visible
through the thick but transluscent shell as narrowly divergent septa.
Visibility was made better by washing the beak and umbo of the
brachial valve with dilute acid to thin the shell.
UNPLACED SPECIES
Rhynchonella lamothei Dautzenberg (1909, p. 271). This is a com-
pletely costate (16 costae) species from the Pliocene of Algeria. It
has a sulcus on the pedicle valve with 7 costae and a prominent fold
with 6 costae. No details of the hinge or interior were described. It
is unlike any other Tertiary rhynchonelloid.
R. (Hemithyris) vinassai Boni (1933, p. 86). Miocene, Monte Val-
lassa, Italy. This is a semicostate form suggestive of Aphelesia bi-
partita but the interior details are not figured.
Rhynchonella washingtoniana Weaver (1912, p. 55). Weaver's
figures of this species indicate a brachiopod with a type of ornamenta-
tion never seen in rhynchonelloids. Examination of specimens from
the Cowlitz River proves the shell to be punctate and the ornamenta-
tion to be that of the genus Terebratulina. The species is thus not a
rhynchonelloid.
Rhynchonella meneghiniana Davidson (1870, p. 463). This is a
small completely costate species from the Eocene of Bolca, Italy. It is
quite distinct from any other Tertiary species but nothing is known
of its beak characters and interiors. It may be related to R. poly-
morpha (=Erymnaria) which may be strongly costate.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 67
Hemithiris dibbleei Hertlein and Grant (1944, p. 46). Eocene of
California. No details of the interior of this species are given but it
is semicostate. In exterior view it accords with Plicirhynchia but this
is a much younger genus located in a completely different faunal realm.
Hemithiris reagani Hertlein and Grant, (1944, p. 54). Oligocene,
California. This species is also semicostate like that above and might
be referable to Plicirhynchia, but no details of the interior are known.
“Rhynchonella” supraoligocaenica Gorges (1952, p. 5). This spe-
cies is from the upper Oligocene of Germany. It is a large, smooth
form suggestive of Aphelesia bipartita. The interior is, however,
unknown and the species cannot be assigned with confidence.
“Rhynchonella”’ valdiviae Helmcke (1940, p. 290). This species is
found near New Amsterdam in the south-central part of the Indian
Ocean. It resembles Compsothyris in form, ornamentation, and beak
characters. The color is brownish gray and the shell transparent as in
Compsothyris. Dental plates are present in the pedicle valve. The
cardinalia consist of spoon-shaped, curved crura truncated at the end.
The brachial valve is provided with a “very weak median-septum, the
front end of which is about even with the ends of the crura. The
septum is highest in the middle.” The figure given by Helmcke (fig.
37) does not show the septum clearly. The species strongly suggests
Compsothyris, but it is not possible to be sure until better details of the
interior are known.
This species also suggests Hemuithyris striata Thomson from off
Shackleton Glacier, Davis Sea, Antarctica, by its rounded outlines and
fine costellae. These two species are assigned to Compsothyris with
a query.
REFERENCES
Acer, D. V.
1957. The true Rhynchonella. Palaeontology, vol. 1, pt. 1, pp. I-15, pls. 1,2.
ALLAN, R. S.
1938. A new genus of Tertiary rhynchonellids (Brachiopoda) from Pata-
gonia. Rec. Canterbury (N.Z.) Mus., vol. 4, No. 4, pp. 199-201.
1940. Studies on the Recent and Tertiary Brachiopoda of Australia and
New Zealand. Pt. 2. Rec. Canterbury (N.Z.) Mus., vol. 4, No. 6,
Pp. 277-207, pls. 35-37.
ARCELLIN, F., and Rocué, P.
1936. Les brachiopodes bajociens du Monsard. Trav. Lab. Géol. Fac. Sci.
Lyons, fasc. 30, Mem, 25, pp. 1-107, pls 1-19.
BITTNER, A.
1890. Brachiopoden der Alpinen Trias. Abhl. K. K. Geol. Reichsanst., Bd.
14, pp. i-iv, I-325, pls. I-41.
1892. Brachiopoden der Alpinen Trias. Nachtrag I. Abhl. K. K. Geol.
Reichsanst., Bd. 17, Heft 2, pp. 1-88, pls. 1-9.
68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Bont, A.
1933. Fossili miocenici del Monte Vallassa. Boll. Soc. Geol. Ital., vol. 52,
fasc. I, pp. 73-156, pl. o.
Buckmal, S. S.
1910. Antarctic fossil Brachiopoda collected by the Swedish South Polar
Expedition. Wiss. Ergebn. Schwedische Siidpolar-Exped. 1901-1903,
Bd. 3, Lief. 7, pp. 1-40, pls. 1-3.
1917. The Brachiopoda of the Namyau beds, northern Shan States, Burma.
Mem. Geol. Surv. India, Palaeont. Indica, n. s., vol. 3, Mem. 2, pp.
i-iv, I-290, pls. I-21.
CHAPMAN, F., and CresPiIn, IRENE.
1923. The Austral Rhynchonellacea of the “Nigricans Series,” with a spe-
cial description of the new genus Tegulorhynchia. Proc. Roy. Soc.
Victoria, n. s., vol. 35, pt. 2, pp. 170-193, pls. 11-13.
Cooper, G. A.
1957. Tertiary and Pleistocene brachiopods of Okinawa, Ryikyi Islands.
U. S. Geol. Surv. Prof. Pap. 314-A, pp. i-iv, 1-20, pls. 1-5.
Cooper, G. A., and Murr-Woop, H. M.
1951. Brachiopod homonyms. Journ. Washington Acad. Sci., vol. 41, No. 6,
pp. 195, 190.
CRANE, A.
1886. On a brachiopod of the genus Atretia, named in MS. by the late Dr.
T. Davidson. Proc. Zool. Soc. London, Apr. 6, pp. 181-184.
Dati, W. H.
1877. Index to the names which have been applied to the subdivisions of
the class Brachiopoda. U. S. Nat. Mus. Bull. 8, pp. 1-88.
1895. Report on Mollusca and Brachiopoda dredged in deep water, chiefly
near the Hawaiian Islands, with illustrations of hitherto unfigured
species from northwest America. Proc. U. S. Nat. Mus., vol. 17,
No. 1032, pp. 675-733.
1910. Report on the Brachiopoda obtained from the Indian Ocean by the
Sealark Expedition, 1905. Trans. Linnaean Soc. London, ser. 2,
Zool., vol. 13, pt. 3, pp. 439-441, pl. 26.
1920. Annotated list of the Recent Brachiopoda in the collection of the
United States National Museum, with descriptions of thirty-three
new forms. Proc. U. S. Nat. Mus., vol. 57, pp. 261-377.
DAUTZENBERG, P.
1909. Description d’une espéce nouvelle de Brachiopode du Pliocéne Al-
gérien. Journ. Conch., vol. 56, pp. 271, 272, pl. 11, figs. 15-17, 1908.
Davinson, T.
1870. On Italian Tertiary Brachiopoda. Pt. 3. Geol. Mag., vol. 7, No. 76,
pp. 460-466, pls. 19 and 20.
1871. On Japanese Recent Brachiopoda. Proc. Zool. Soc. London, pp. 300-
B12. pis.s0al:
1874a. A monograph of the British fossil Brachiopoda. Vol. 4, Teritary,
Cretaceous, Jurassic, Permian, and Carboniferous supplements ;
etc., Paleontograph. Soc., pp. 1-16.
1874b. On the Tertiary Brachiopoda of Belgium. Geol. Mag., dec. 2, vol. 1,
No. 4, pp. 150-150, pls. 7, 8.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 69
1880. Report on the Brachiopoda dredged by H. M. S. “Challenger” during
the years 1873-1876. Voy. Challenger, Zool., vol. 1, pp. 1-67, pls. 1-4.
1886. On a living spinose Rhynchonella from Japan. Ann. Mag. Nat. Hist.,
January, pp. 1-3.
1886-1888. A monograph of Recent Brachiopoda. Trans. Linnaean Soc., ser.
2, vol. 4, Zool., pt. 1, pp. 1-73, pls. 1-13, 1886; pt. 2, pp. 75-182, pls.
14-25, 1887; pt. 3, pp. 183-248, pls. 26-30, 1888.
FiscHer, P., and OEHLERT, D. P.
1891. Brachiopodes. Exped. Sci. Travailleur et Talisman, 1880-1883, pp. I-
140, pls. 1-8.
GorceEs, J.
1952. Neue Invertebrata aus dem norddeutschen Oberoligozan. Palaontol.
Zeitschr., Bd. 26, Nos. 1, 2, pp. 1-9, pls. 1, 2.
GouLp, A. A.
1861. . . . descriptions of new shells collected by the United States North
Pacific Exploring Expedition. Proc. Boston Soc. Nat. Hist. for
1859-1861, vol. 7, Brachiopods, pp. 323, 324.
HALL, J., and CLarkKeE, J. M.
1894. An introduction to the study of the genera of Palaeozoic Brachiopoda.
Pt. 2. Paleont. New York, vol. 8, pp. 177-210.
Harat, K.
1940. The Cenozoic Brachiopoda of Japan. Sci. Rep. TOhoku Imp. Univ.,
ser. 2 (Geology), vol. 20, pp. 1-413, pls. 1-12.
HELMCKE, J. G.
1939. In Kukenthal and Krumbach, Handbuch der Zoologie, Bd. 3, 2
Halfte, Lief. 13, pp. 139-262, figs. 152-271.
1940. Die Brachiopoden der Deutschen Tiefsee-Expedition. Wiss. Ergebn.
Deutsch. Tiefsee Exped. 1898-99, Bd. 24, H. 3, pp. 215-316, figs.
I-43.
HemMING, F.
1953. Copenhagen decisions on zoological nomenclature, additions to, and
modifications of, the Régles Internationales de la Nomenclature
Zoologique, 14th International Congress of Zoology, Copenhagen.
Hert etn, L. G., and Grant, U. S., IV.
1944. The Cenozoic Brachiopoda of western North America. Publ. Univ.
California at Los Angeles, Math. and Phys. Sci., vol. 3, pp. i-vi,
1-236, pls. I-21.
Hutton, F. W.
1873. Catalogue of the Tertiary Mollusca and Echinodermata of New
Zealand in the collection of the Colonial Museum. Brachiopoda,
pp. 35-37. Colonial Mus. and Geol. Surv. Dept.
JAANUSSON, V.
1951. Brachiopoda from the Tertiary Nahuel Hupai formation of the
Patagonian Cordillera. Bull. Geol. Inst. Univ. Uppsala, vol. 34,
pp. IQI-200, pl. 1.
Jackson, J. W.
1918. Brachiopoda. British Antarctic (“Terra Nova”) Exped. 1910. Nat.
Hist. Rep., Zool., vol. 2, No. 8, pp. 177-202.
7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
JacKkson, J. W., and Strasny, G.
1937. The Brachiopoda of the Siboga Expedition. Siboga-Exped., Monogr.
27, pp. 1-20, pls. 1, 2.
JEFFREYS, G.
1869. The deep-sea dredgings. Explorations on H. M. S. “Porcupine.”
Nature, vol. 1, p. 136.
1870. Preliminary report of the scientific exploration of the deep sea in H.
M. Surveying vessel “Porcupine” during the summer of 1869.
Proc. Roy. Soc., vol. 18, No. 121, p. 421.
1876. On some new and remarkable North-Atlantic Brachiopoda. Ann. Mag.
Nat. Hist., ser. 4, vol. 18, pp. 250-253.
1878. On the Mollusca procured during the “Lightning” and “Porcupine”
Expeditions, 1868-70. Pt. 1. Proc. Zool. Soc. London, pp. 393-416,
pl. 23, figs. 5, 6.
Jousin, L.
1901. Brachiopodes. Résultats voyage S. Y. Belgica, 1897-1898-1899, Zool.,
Rapt. Sci. Commiss. Belgica, pp. 1-11, pls. 1, 2.
KozLowskI, R.
1929. Les Brachiopodes Gothlandiens de la Podolie Polonaise. Palaeon-
tologia Polonica, t. I, xiv + 256 pp., pls. 1-12.
Lapp, H. S., and HorrMetsTer, J. E.
1945. Geology of Lau, Fiji. Bernice P. Bishop Museum Bull. 181, pp. i-vi.
4-390, pls. 1-62.
LEIDHOLD, C.
1921. Beitrag zur genaueren Kenntnis und Systematik einiger Rhynchonel-
liden des reichslandischen Jura. Neues Jahrb. Min., Geol., Palaont.,
Beilageband 44, Heft 2, pp. 343-368, pls. 4-6.
1922. Rhynchonella doederleini Davids., eine kritische Brachiopoden-Unter-
suchung. Neues Jahrb. Min., Geol., Palaont., Beilageband 45, pp.
423-470, pls. 11-15.
1928. Beitrag zur Kenntnis der Fauna des rheinischen Stringocephalen-
kalkes insbesondere seine Brachiopodenfauna. Teil 1, Abh. Preuss.
Geol. Landesanst., N. F., Heft 109, pp. 1-99, pls. 1-7.
MasSALonco, A.
1850. Schizzo geognostico sulla valle del Progno o Torrente d’Illasi, con
un saggio sopra la flora primordiale del M. Bolca. Verona. Brachi-
opods, pp. 18, 10.
Mezwnerics, I.
1943. Die Brachiopoden des Ungarischen Tertiars. Ann. Hist.-Nat. Mus.
Nat. Hungarici Par. Min., Geol., Pal., vol. 36, pp. 10-60, pls. 2-6.
Murr-Woon, H. M.
1934. On the internal structure of some Mesozoic Brachiopoda. Philos.
Trans. Roy. Soc. London, ser. B, vol. 223, pp. 511-567, pls. 62, 63.
1936. A monograph on the Brachiopoda of the British Great Oolite series.
Pt. I. The Brachiopoda of the Fuller’s earth. Palaeontograph. Soc.,
1935, PD. i, ii, 1-144, pls. 1-5.
1955. A history of the classification of the phylum Brachiopoda. British
Museum (Natural History), London, pp. i-viii, 1-124.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER qe
Owen, E. F.
1955. The use of sugar solution in the differentiation of the internal struc-
tures of Upper chalk brachiopods. Proc. Geol. Assoc. London, vol.
66, pp. 369-370.
Parmer, R. H.
1948. List of Palmer Cuban fossil localities. Bull. Amer. Paleont., vol. 31,
No. 128, pp. 1-178.
PaTAnge, G. M.
1929. Brachiopodi postpliocenici delle marne a briozoi dei Dintorne di
Augusta (Sicilia). Palaeont. Italica, vol. 29-30, 1923-1928, pp. 80-
127, pls) 17, 18:
Pettit, N. E.
1950-1954. The Rhynchonellidae of the British chalk. Pt. 1. Palaeonto-
graph. Soc., vol. for 1940, pp. i-vi, 1-26, pls. 1, 2, 1950; pt. 2, pp.
27-52, pl. 3, 1954.
RotHpLetz, A.
1886. Geologisch-palaeontologische Monographie der Vilser Alpen, mit
besonderer Beriicksichtigung der Brachiopoden-Systematik. Palae-
ontographica, Bd. 33, Lief. 1-3, pp. 1-180, pls. 1-17.
RZEHAK, A.
1904. Rhynchonella polymorpha Mass. im Karpathischen Eocan Mahrens.
Verhandl. K. K. Geol. Reichsanst., No. 7, pp. 182-184, Apr. 12.
RZHONSNITZKAIA, M. A.
1956. Systematization of Rhynchonellida. Resumenes de los Trabojos Pre-
sentados, XX Congreso geologico Internacional, pp. 125, 1206.
Mexico.
Sacco, F.
1902. I. Brachiopodi dei Terreni Terziarii del Piemonte e della Liguria,
Pp. 3-50, pls. 1-6.
ScHUCHERT, C., and LEVENE, C. M.
1929. Brachiopoda (generum et genotyporum index et bibliographia).
Fossilium Catalogus, 1, Animalia, pp. 3-140.
SEGUENZA, G.
1867. Intorno ai brachiopodi miocenici delle provincie Piedemontesi. Ann.
Acad. Aspiranti Nat., 1866, ser. 3, vol. 6, pp. 53-67, pls. 1-3.
Tuomson, J. A.
1915. The genera of Recent and Tertiary rhynchonellids. Geol. Mag., n. s.,
dec. 6, vol. 2, pp. 387-302.
1918. Brachiopoda. Australasian Antarctic Expedition, Sci. Rep., ser. C,
Zoology and Botany, vol. 4, pt. 3, pp. 5-76, pls. 15-17.
1927. Brachiopod morphology and genera (Recent and Tertiary). New
Zealand Board Sci. Art, Manual 7, pp. i-iv, 1-338, pls. 1, 2.
Toutmin, L. D.
1940. Eocene brachiopods from the Salt Mountain limestone of Alabama.
Journ. Paleont., vol. 14, pp. 227-233, pl. 28.
Weaver, C. E.
1912. Preliminary report on the Tertiary paleontology of western Wash-
ington. Washington Geol. Surv., Bull. 15, 80 pp.
72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
1942. Paleontology of the marine Tertiary formations of Oregon and
Washington. Univ. Washington Publ. Geol., vol. 5, Brachiopods,
pp. 12-18.
WELLER, S.
1910. Internal characters of some Mississippian rhynchonelliform shells.
Bull. Geol. Soc. Amer., vol. 21, pp. 497-516, figs. 1-18.
WIL.IAMS, A.
1956. The calcareous shell of the Brachiopoda and its importance to their
classification. Biol. Rev., vol. 31, pp. 243-287.
WISNIEWSKA, M.
1932. Les rhynchonellides du Jurassique sup. de Pologne. Palaeontologia
Polonica, t. 2, No. I, pp. i-viii, 1-71, pls. 1-6.
Woopwarb, S. P.
1855. Description of a new species of recent Rhynchonella (R. grayi,
Woodw.). Ann. and Mag. Nat. Hist., ser. 2, vol. 16, p. 444.
Yase, H., and Harat, K.
1934. The Recent brachiopod fauna of Japan. (1) New genera and sub-
genera. Proc. Imp. Acad. Japan, vol. 10, No. 9, pp. 586-580.
1935. On some Brachiopoda from the “Ryttkyi limestone” of Kikaizima
and Okinawa-zima, Rytkyt Islands and southwestern Formosa
(Taiwan). Japanese Journ. Geol., Geogr., Trans. and Abs., vol. 12,
Nos. 3 and 4, pp. 93-102, pl. 14.
EXPLANATION OF PLATES
PLATE I
Cryptopora
Page
A. Mannia nysti Davidson = Cryptopora nysti (Davidson).............. 22
Figs. 1-14. 1, Dorsal view of a complete specimen showing triangular
outline, 10, hypotype U.S.N.M. 549422a. 2, Same view as preced-
ing, X 20, showing strongly elevated deltidial plates and large tri-
angular foramen. 3-5, Respectively partial side, side, and ventral
views of the same specimen showing the profile and large deltidial
plates, X 10. 6, Interior of the pedicle valve of the same specimen
showing teeth and deltidial plates, 10. 7, Interior of the brachial
valve of the same individual showing high, narrow median septum,
and crura with flattened distal extremities, X10. 8-10, Respectively
interior, partial side, and side views of the same valve, X 20, showing
median septum and details of the cardinalia. 11, 12, Interior and par-
tial side views of another pedicle valve interior, X 10, showing teeth
and deltidial plates, hypotype U.S.N.M. 549422b. 13, 14, Partial side
and anterior views of the preceding specimen, X20, showing dental
plates, apical plate and deltidial plates.
Upper Miocene (Diestien-Sables de Deurne), Wommelghem, east
side of Antwerp, Belgium.
B. Cryptopora rectimarginata Cooper, NeW SPeCi€S...........eeeeeeeeeee 20
Fig. 15. Dorsal view of a complete specimen showing alate deltidial
plates and apical plate, & 15, paratype U.S.N.M. 274143d.
Recent, Eolis Station 340, at 209 fathoms, off Fowey Light, Florida.
Fig. 16. Another specimen showing the alate deltidial plates and apical
plate, X 15, paratype U.S.N.M. 274168a.
Recent, Eolis Station 320, at 80 fathoms, off Western Dry Docks,
Florida.
Figs. 17, 18. Respectively tilted to the side and full views of a specimen
showing elaborate alae on the deltidial plates, 15, paratype
U.S.N.M. 336806a.
Recent, Eolis Station 378, at 165 fathoms, off Fowey Light, Florida.
Fig. 19. Interior of a pedicle valve with tooth broken on right side
but showing alae with scalloped edges on deltidial plates and the large
plate just anterior to the apex, 15, paratype U.S.N.M. 336895.
Recent, Eolis Station 377, at 190 fathoms, off Fowey Light, Florida.
PLATE 2
Cryptopora and Neorhynchia
T. Cryptopora rectimarginata Cooper, new SpecieS............eccceeeeees 20
Figs. I-11. 1-3, Respectively dorsal, side, and anterior views of a com-
plete specimen, <6, showing rectimarginate commissure, holotype
73
74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Page
U.S.N.M. 274143a. 4, Partial side view of another individual showing
the alate deltidial plates, X 6, paratype U.S.N.M. 274143b. 5, Pos-
terior of the specimen shown in figure 4, illustrating the alate deltidial
plates, X 8. 6, 7, Respectively anterior and full views of the interior
of the pedicle valve, 6, showing strong dental plates, paratype
U.S.N.M. 274143c. 8-11, Respectively partial side, full, anterior, and
posterior views of the brachial interior of the preceding, showing the
long slender maniculifer crura, the short, strongly elevated median
septum and the bilobed cardinal process, 8, paratype U.S.N.M.
274143¢.
Recent, Eolis Station 340, 209 fathoms, off Fowey Light, Florida.
B. Neorhynehia:strebelt Dall yore. cca gee vives seca fete es cee oa tea ae ele 34
Figs. 12-23. 12-15, Respectively ventral, dorsal, anterior, and side views
of a complete specimen, * 2, showing the sulcate anterior commissure,
paratype U.S.N.M. r10741a. 16, 17, Respectively anterior and full
views of the posterior pedicle valve, X 4, showing dental plates, corru-
gated teeth, and disjunct deltidial plates, holotype U.S.N.M. 110741.
18, 19, 21, 22, Respectively full, posterior, anterior, and partial side
views of the posterior of the brachial valve of the holotype, Xx 4,
showing broad outer hinge plates, low median ridge, small inner
hinge plates, and falcifer crura. 20, 23, Partial anterior and partial
side views of the paratype showing socket ridges and short crura,
SG:
Recent, U. S. Fish Commission Station 4721, 2,084 fathoms in
Globigerina ooze, 35.1° F., mid-Pacific.
PLATE 3
Hemithyris
AY Hemithyns woodcwards ((DayidsOm)s <x es cicis oss w/o'sre ce cele /acaieraveis epslclelerers 47
Figs. 1-11. 1-3, Respectively side, anterior, and dorsal views of a com-
plete specimen, X 1, hypotype U.S.N.M. 111083a. 4, Dorsal view of
the same specimen, 2, showing fine radial striae. 5, Beak of a ped-
icle valve, 3, showing large foramen and discrete deltidial plates,
hypotype U.S.N.M. 111083b. 6, Interior of the same pedicle valve
tilted to show the vertical dental plates, & 2. 7, 8, Respectively ante-
rior and full views of a brachial valve showing cardinalia, inconspicu-
ous median ridge and muscle scars, & 2, U.S.N.M. 111083c. 9,
Posterior view of the same brachial valve showing socket ridges,
outer hinge plates, and radulifer crura, X 4. Io, Partial side view of
the same brachial valve showing the long slender crura, & 4. 11, Pos-
terior view of another brachial valve showing the cardinalia and the
scars of the diductor muscles at the apex, 4, hypotype U.S.N.M.
111083d.
Recent, Hakodate, southern Hokkaido, Japan.
Beemithyrispsittacea \(Gimelin) saseeneer mecca c eee eee eee 45
Figs. 12-21. 12-14, Respectively anterior, side, and dorsal views of a
complete specimen showing the long beak, & 1, hypotype U.S.N.M.
111004a. 15, Dorsal view of the preceding specimen, X 2, showing
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 75
Page
the closely crowded costellae separated by narrow, shallow striae.
16, Interior of the pedicle valve of another specimen tilted to show
apical and dental plates, 2, hypotype U.S.N.M. 111004b. 17,
Posterior of the preceding pedicle valve, X 3, showing large, incom-
plete foramen, large teeth, and small disjunct deltidial plates. 18, 10,
Respectively anterior and full views of the brachial valve interior
showing cardinalia, myophragm, and muscle scars, 2, hypotype
U.S.N.M. 111004b. 20, Posterior of the preceding specimen tilted to
show socket ridge, corrugated socket, and long, slender, curved,
radulifer crura with the strengthening ridge on their under or dorsal
surface, X 3. 21, Posterior view of the cardinalia of the same speci-
men showing socket ridge, small outer hinge plates, and distally
flattened crura, X 3.
Recent, Coal Harbor, Unga Island, Shumagins, Alaska.
PLATE 4
Aetheia, Frieleia, Grammetaria, Camarotoechia, and Hemithyris
An Acthersgualters: (NOLES \oss vain 2h dein aus ats gate sim desia poemtaneen as 42
Fig. 1. Rubber impression of a pedicle valve interior prepared to
show the muscle scars and pallial marks, 2, hypotype U.S.N.M.
369298a (see pl. 9, B).
Miocene (Duntroon greensand), Otago, New Zealand.
Ba lraeletas Halle Tal 50 oie aah cls ish resco nie 8) shoys ajaesravadei ghar aiahn iejnini aie Shaveysees 53
Figs. 2, 3. 2, Posterior of a brachial valve, 6, showing the hinge
plates surrounding a plug of the median septum, and the inner hinge
plates engulfing the septum, hypotype U.S.N.M. 111021a. 3, Posterior
of another brachial valve interior, X 6, with well-preserved cardinalia
and elongate adductor scars, and showing inner hinge plates not yet
coalesced, hypotype U.S.N.M. 111021b.
Recent, 559 fathoms, in ooze, 38.4° F., U. S. Fish Commission Sta-
tion 2871, off the coast of Washington.
Cc. Grammetoravarischi: (Dall )is.\s's0. cjc.cisters oypjalals «ia oles realty tases 58
Figs. 4, 5. 4, Dorsal view of a young but imperfect specimen, 6 above
and X10 below, showing elaborate auriculations on the deltidial
plates, paratype U.S.N.M. 274134. 5, Same specimen with brachial
valve tilted to show cardinalia and dental plates, 6, and also show-
ing septum united with hinge plates and long curved crura.
Recent, at 161 fathoms, 57.4° F., on sand, U. S. Bureau of Fish-
eries Station 5735, off Jolo, Philippine Islands.
Di Gamaroloechta Species), vas cs so) acid ahelete Sie nielsleia Siete aici de ve sO RIA HOT Slate akasre 10
Figs. 6-8. 6, 7, Two views of the posterior part of a silicified brachial
valve, X 4, showing crura and median hinge plate welding them to-
gether, figured specimen U.S.N.M. 134812e. 8, The same specimen
tilted to show apical chamber (septalium?) and crural plates, x 4
(see p. IO).
Devonian (Norway Point formation), junction French and Truckey
roads, SW1/4SW1/4 sec. 4, T. 31 N., R. 8 E., Alpena County,
Michigan.
76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
E. Hemithyris poitiacea (Gmelin) (oes eae ee ee ie tee een teen
Figs. 9, 10. 9, Posterior of a large brachial valve showing cardinalia
with the narrow outer hinge plates and slender radulifer crura with
the transverse cardinal process at the apex, 3, hypotype U.S.N.M.
IIIO13. 10, Same specimen, X 3, tilted to the side to show the slender
radulifer crura with their flattened distal extremity and strengthening
ridge on dorsal side.
Recent, 13 fathoms, Upernavik Harbor, east coast of Greenland.
Figs. 11, 12. 11, Posterior of another brachial valve, x 3, tilted to show
the long, slender crura and slight development of outer hinge plates,
hypotype U.S.N.M. 549379. 12, Side view of the pedicle valve inte-
rior belonging to the preceding and showing the inner face of the dental
plate and tooth with its corrugations, X 2.
Recent, north end of Nunivak Island, Alaska.
PLATE 5
Eohemithyris, Sphenarina, Cryptopora, and Tegulorhynchia
A. Eohemtthyris alexis Hertlein and Grafit..............0ccccccwnescces
Figs. 1-5. Respectively anterior, posterior, side, dorsal, and ventral
views of a somewhat crushed paratype cracked at the beaks, > 1,
UCLA. 7257.
Eocene (Domengine formation), from section line, 2,600 feet south
of the northeast corner of sec. 20, T. 28 S., R. 19 E., M. D. B. and M.,
near headwaters of west branch of Agua Media Creek, McKittrick
Quadrangle, Temblor Range, California.
BeSphenarinastcula (Sestenza)'h. Sakae cee ec weac ates cae ectemeaes eons
Figs. 6-15. 6-10, Respectively dorsal, posterior, ventral, side, and ante-
rior views of a well-preserved complete specimen, I, showing the
strongly triangular outline, hypotype U.S.N.M. 549353a. 11, Dorsal
view of the preceding showing the foramen and obscure radial lines,
X 2. 12, Posterior of the same specimen, X 4, showing alate conjunct
deltidial plates. 13, 14, Two views of the interior of another specimen
showing the cardinalia, <4, with their small outer hinge plates,
dental plates and lack of a median ridge or septum in the brachial
valve, hypotype U.S.N.M. 540353c. 15, Enlargement of the shell
surface, <6, showing the delicate radial lines, hypotype U.S.N.M.
549353.
Pliocene, Milasso, Messina, Sicily.
CG. Cryptopora ‘gnomon \( Jetireys) ec keiak eater ees eases cee ceee keene
Fig. 16. Brachial valve tilted to the side, & 8, to show the high median
septum and the long maniculifer crura with the distal end handlike
and with fingers extended, paratype U.S.N.M. 94367.
Recent, 780 fathoms, off Cuba.
D. Tegulorhynchia.
Teguiorhynchia ‘squamasa’ (CEMUton) ic... ere inte ss oa he's we eictee s tan eee
Figs. 17-24. 17-20, Respectively ventral, dorsal, anterior, and side views
of a complete specimen, 1, hypotype U.S.N.M. 89855a. 21-23,
Respectively dorsal, anterior, and side views of the preceding speci-
30
62
50
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER Ti
Page
men, X 2, showing the imbricating ornament. 24, A partially exfoli-
ated pedicle valve showing the impression of the muscle scars, X 2,
hypotype U.S.N.M. 80855b.
Miocene (Ototaran), Broken River, Trelissick Basin, Canterbury,
New Zealand.
T equlorhynchia, daderlemus (Davidson)... «0000 <iej Adal dds cceeinenes's 51
Figs. 25-31. 25, Dorsal view of a complete but small specimen, X 2,
showing imbricate ornament, hypotype U.S.N.M. 549317b. 26, 27,
Tilted and full views of the interior of the pedicle valve showing the
muscle field and dental plates, * 2, hypotype U.S.N.M. 549317d.
28, Beak and teeth of the pedicle valve, & 4, showing nearly conjunct
deltidial plates, hypotype U.S.N.M. 549317f. 29-31, Respectively full
view of the cardinalia, lateral view, and with shell tilted to show short
median ridge and hinge plates, 4, hypotype U.S.N.M. 549317c.
The crural points have been broken off.
Miocene or Pliocene (Shinzato tuff), high road cut along Highway
64, about 0.1 mile west of sharp bend in road about 0.3 mile east of
Yashitomi, Okinawa, Rytkyi Islands.
PLATE 6
Patagorhynchia and Notosaria
A. Patagorhynchia patagonica (von Thering) ..........cccccecceecsevsss 44
Figs. 1-4. Respectively dorsal, posterior, side, and anterior views of a
complete but imperfect specimen showing imbricated ornamentation,
X 1, hypotype Princeton University.
Eocene (Patagonian), Lake Pueyrredon, Argentina.
Figs. 5, 6. Interior of the pedicle and brachial valves, & 1, showing
concave deltidial plates and thickened cardinalia. (After von Ihering,
1903, pl. 3, figs. IIa, 11b.)
Eocene (lower Patagonian), north of Seco River and San Julian,
Argentina.
BOON OIC MOTiCORe CSOWELDY) <o unc ce ne cas ce cade se nbetecuoue seats 48
Figs. 7-17. 7-9, Respectively side, dorsal, and anterior views of a com-
plete specimen showing commissure and folding, 1, hypotype
U.S.N.M. 111018b. 10, The beak of the preceding specimen enlarged,
X 3, to show apical plate, small disjunct deltidial plates, and large,
incomplete foramen. 11, 12, Interior of pedicle valve respectively tilted
and in full view, X 2, to show apical plate, teeth, muscle scars, dental
plates, deltidial plates, and numerous pallial sinuses, hypotype
U.S.N.M. r111018a. 13, Posterior of same, X 3, showing foramen,
small disjunct deltidial plates, and large teeth. 14, 15, Brachial valve
in full and tilted views to show cardinalia, transverse cardinal process,
inconspicuous median ridge, muscle scars, and numerous pallial
sinuses, X 2, hypotype U.S.N.M. r111018a. 16, Posterior of same,
X 4, showing cardinal process and curved crura. 17, Same tilted to
show crura in side view, X 4.
Recent, Stewart Island, New Zealand.
78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
PLATE 7
Plicirhynchia and Aphelesia
Page
A. Phcirhynchia plicigera (von Uhering) ...........0--cccccscccesesenses 52
Figs. I-11. 1-3, Respectively anterior, side, and dorsal views of a com-
plete specimen showing the long beak and anterior costation, X I,
hypotype U.S.N.M. 549346a. 4, Exterior of the dorsal side of the
same specimen showing the posterior obscure striation and the strong
anterior costation, X 2. 5, Posterior of the same specimen, X 4, show-
ing the large foramen, conjunct deltidial plates, and the fine striae on
the umbo of the brachial valve. 6, 7, Full and tilted views of interior
of a pedicle valve showing the deltidial plates and strong vertical
dental plates, 2, hypotype U.S.N.M. 549346b. 8, 9, Full and tilted
views of a fragmentary brachial valve showing cardinal process and
incomplete crura, 2, hypotype U.S.N.M. 540346c. 10, 11, Full and
partial side views of the cardinalia of the same brachial valve showing
the radulifer crura, thick socket plates, and cardinal process, X 4.
Eocene (Patagonian), Mazaredo, Patagonia, Argentina.
B. Aphelesia bipartsia: (Brocehiya: Weare eee someone petals viele a oroiete eters strane 4I
Figs. 12-22. 12-14, Respectively side, anterior, and dorsal views of a
complete specimen showing small beak, small foramen, and anterior
fold, X 1, hypotype U.S.N.M. 549349a. 15, 16, Interior of a pedicle
valve in full view and tilted to show the small teeth, small dental
plates, oval foramen, conjunct deltidial plates, and muscle field, X 2,
hypotype U.S.N.M. 540349b. 17, Posterior of the same pedicle valve,
x 4, showing the oval foramen and conjunct deltidial plates. 18-20,
Respectively partial side, anterior, and full views of the brachial
valve belonging to the preceding specimen (pedicle valve), X 2,
showing the cardinalia. 21, 22, Respectively side and posterior views
of another brachial valve, <3, showing the broad falcifer crura,
hypotype U.S.N.M. 540380.
Pliocene, Messina, Sicily.
PLATE 8
Sphenarina, Aphelesia, and Eohemithyris
A. SPRERGIING SUCHE \ CS CRUCIAL Yee id ital cca taxa cl spwieishs a oie: » aioisrminjalavstetene/aieliateleyat 62
Figs. 1-7. 1-5, Respectively dorsal, ventral, anterior, side, and posterior
views of a complete and nearly perfect specimen, X 1, showing tri-
angular form, rectimarginate anterior commissure, and nearly erect
beak, hypotype U.S.N.M. 173728. 6, The same specimen enlarged,
2. 7, Interior of another specimen showing cardinalia, X 10, with
their long, slender, spinulifer crura, narrow outer hinge plates, and
narrow, elevated socket ridges, hypotype U.S.N.M. 549381a.
B. Eohemithyris? gettysburgensis Cooper, new Species............eseees 33
Pliocene, Messina, Sicily.
Figs. 8-12. Respectively anterior, posterior, side, dorsal, and ventral
views of a large and complete specimen, <1, holotype U.S.N.M.
549382.
Miocene, on coast 43 miles west of Gettysburg, Washington.
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER
GA phrele ste UEPGFIEE <@ESC OCGII)) 5: 5\sh0540 ain mre cites a aievesel ev alvieschelayeyaceteie:sie/s/ouais =: 6
Figs. 13-18. 13, Imperfect brachial valve showing falcifer cardinalia,
lack of outer hinge plates, deeply entrenched muscle scars, and short
median ridge (myophragm) in full view, <3, hypotype U.S.N.M.
549380a. 14, Same tilted slightly toward the observer to demonstrate
absence of outer hinge plates, X 3. 15, Same in side view, showing
the serrated edge and convex outer surface of the falciform crus,
X 4. 16, Same in side view, 3, showing falcifer crura and socket.
17, Same tilted to show posterior face of cardinalia with the short
socket ridges, lack of outer hinge plates and falcifer crura, * 3. 18,
Same tilted to show cardinalia and serrated distal extremity of crus,
X 3.
Pliocene, Messina, Sicily.
PLATE 9
Grammetaria and Aetheia
Az Grommectariacbartschs CD ally ex sce scets: sca tpezsjadsco.s opcvebitars ye taers.cle: btolocs scolar she's
Figs. 1-12. 1-4, Respectively dorsal, side, anterior, and posterior views
of the holotype U.S.N.M. 239269, 1, showing the rectimarginate
commissure. 5, 6, Ventral and dorsal views of the holotype, X 2,
showing the fine radial costellae. 7, Posterior of the pedicle valve of
the holotype showing the corrugated teeth, large foramen, and auric-
late but broken deltidial plates, & 4. 8, Anterior view of the same
pedicle valve showing the vertical dental plates, & 2. 9, 10, Full and
slightly tilted views of the interior of the brachial valve of the holo-
type showing the cardinalia with their spinulifer crura, 2. I1,
Partial side view of the cardinalia of the preceding brachial valve
showing the strong socket ridge, corrugated socket, and short, slender
spinulifer crura, X 4. 12, Posterior view of the same, brachial inte-
rior, X 4, showing socket ridges, small inner hinge plates, and plug
closing apical chamber.
Recent, U. S. Bureau of Fisheries Station 5621, off Jolo Island,
Philippines ; 298 fathoms, in Mollucca Pass off Makyan Island.
Bs AciheragualtersCMOteis))) ac) ajnjs\oare sis © aie x's 2rs ale, Sidon arei aietale eieis'eiatere.avesee
Figs. 13-22. 13-15, Respectively dorsal, side, and anterior views of a
complete specimen, I, hypotype U.S.N.M. 89828a. 16, Dorsal view
of the preceding specimen, X 2. 17, Posterior of the preceding, x 3,
showing the minute foramen and concave deltidial plates. 18, 10,
Respectively full and anterior views of the pedicle valve interior of
another specimen showing deeply impressed muscle field, & 2, hypo-
type U.S.N.M. 3609298a. Note absence of dental plates. 20, Interior
of an imperfect brachial valve, hypotype U.S.N.M. 80828b, x 2,
21, Full view of the preceding specimen showing the thick socket
ridges, corrugated sockets and thick coalesced inner hinge plates,
and long falcifer crura, X 4. 22, Same as preceding, & 4, but tilted
to show the slender bladelike falcifer crura, socket ridge, and corru-
gated socket.
Miocene (Duntroon greensand), 1 mile north of Kakanui, north of
Otago, New Zealand.
58
42
80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
PLATE I0
Hispanirhynchia
Hispamrhianchta’ cornea ‘CRischer) .. sc asicsoc cents cee oleee eases Coreaee es 59
Figs. 1-21. 1-3, Respectively dorsal, anterior, and side views of a com-
plete specimen, 1, hypotype U.S.N.M. 130327a. 9, Pedicle valve in-
terior of same tilted to show small dental plates, X 2. 4, Exterior of
preceding hypotype showing fine costellae, 2. 5, Anterior of pedicle
valve of another specimen, XX 3, showing fine, subdued costellae,
U.S.N.M. 130327b. 6, 8, Two views of the interior of the preceding
pedicle valve, X 2, showing teeth, dental plates, small muscle field,
and pallial marks. 7, Interior view of another pedicle valve showing
teeth and pallial marks, X 2, hypotype U.S.N.M. 130327c. 10, Beak
of preceding, & 4, showing teeth, foramen, and remnantal deltidial
plates. 11, Interior of the brachial valve of a young specimen
showing strong development of inner hinge plates, *X 2, hypotype
U.S.N.M. 130327d. 13, Same, <6, showing inner hinge plates in
great detail. 12, Same specimen as preceding, X 4, tilted to the side
to show the spinulifer laterally flattened crus with serrated distal
extremity, the inner hinge plates, and corrugated socket. 14, Same as
preceding tilted to show junction of septum and inner hinge plates,
< 4. 15, Posterior of an old specimen showing deeply impressed ad-
ductor field, 3, hypotype U.S.N.M. 130327c. 16, Same, ca. X 6,
tilted to show inner hinge plates and median ridge. 17, Interior of
another brachial valve showing pallial sinuses, small genital areas,
and cardinalia, & 2, hypotype U.S.N.M. 130327a. 19, Same tilted to
show direct view of cardinalia and septum, X 2. 18, 20, Same speci-
men, respectively side view showing bladelike spinulifer crura and
view showing strong socket ridges and inner hinge plates, ca. x 6.
21, Interior of another brachial valve tilted to show socket ridges,
inner hinge plates and rostral chamber, X 3, hypotype U.S.N.M.
130327e.
Recent, 240 fathoms off the coast of Mogador, Morocco.
PLATE II
Rhytirhynchia and Basiliola
A; Rhytirhyachiausladent.> (Dall) osents assaeiaie: sya oy0in.0 15s caps Seales slaispeigr eel aolnie 35
Figs. I-11. 1-3, Respectively anterior, side, and dorsal views of a com-
plete specimen showing anterior costation, 1, lectotype U.S.N.M
111086. 4, 5, Respectively full and tilted views of the interior of the
pedicle valve showing pallial sinuses, small conjunct deltidial plates,
deeply incised muscle scars, and pedicle collar, X 2. 6, Posterior of the
pedicle valve of the lectotype, X 3, showing small foramen, conjunct
deltidial plates, and corrugated teeth. 7-9, Respectively anterior, full,
and partial side views of the interior of the brachial valve showing
cardinalia, deeply impressed adductor field, and pallial sinuses, XX 2.
10, Partial side view of the falcifer crura showing serrate distal ex-
tremity and corrugated socket, X 4. 11, Posterior view of the same
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 81
Page
brachial valve, X 4, showing curved socket ridges, modestly developed
outer hinge plates, incipient inner hinge plates, and falcifer crura.
Recent, Sealark Expedition, 1905, Station Cl, 123 to 158 fathoms,
south of the Saya de Malha Banks, Indian Ocean.
B. Bassitola’ beechers: CD allys v5 sec h cme eee ae eee Caen aks 25
Figs. 12-16. 12-14, Respectively side, anterior, and dorsal views of a
complete specimen showing the smooth exterior fold, & 1, hypotype
U.S.N.M. 334678. 15, Dorsal view of the same specimen, XX 2. 16,
Posterior of the preceding specimen showing conjunct, auriculate
deltidial plates, 3.
Recent, U. S. Bureau of Fisheries Station 4130, 283 to 309 fathoms,
46.1° F., near Kauai Island, Hawaiian group.
Figs. 17-23. 17, 18, Respectively tilted and full views of the pedicle
interior of another specimen showing deeply impressed muscle scars
and pallial sinuses, X 2, hypotype U.S.N.M. 334679. 19, Posterior of
the preceding pedicle valve, X 4, showing corrugated teeth, conjunct
and auriculate deltidial plates. 20, 21, Respectively full and anterior
views of the brachial valve of the preceding pedicle valve showing the
cardinalia and pallial marks, & 2. 22, Posterior view of the cardinalia
of the preceding specimen showing socket ridges, corrugated sockets,
broad outer hinge plates, and falcifer crural plates, <4. 23, Side
view of the preceding specimen showing the broad, distally serrate,
falcifer crura and the corrugated sockets, X 4.
Recent, 147 to 198 fathoms, 49° F., off west coast of Hawaii.
PLATE 12
Basiliola
Basiolaiambholos Vallix Saxktal. Ao: Sates Wale Alay wana s tae s 27
Figs. 1-6. 1-5, Respectively dorsal, posterior, anterior, ventral, and side
views of the lectotype, 1, showing robust form, smooth exterior
and strong dorsal fold, U.S.N.M. 229301b. 6, Beak of the preceding
specimen, X 3, showing small round foramen and auriculate deltidial
plates. :
Recent, U. S. Bureau of Fisheries Station 5592, 305 fathoms,
43.3° F., gravel and mud bottom, Sibuko Bay, south of Silungan
Island, Borneo.
Figs. 7-15. 7, Posterior of the pedicle valve of another specimen, X 4,
showing corrugated teeth, and conjunct and auriculate deltidial plates,
hypotype U.S.N.M. 274135. 8, 9, Respectively full and tilted views of
the preceding pedicle valve, X 2, showing dental plates, muscle area
and pallial sinuses. 10, Interior of the apex of the preceding pedicle
valve, X 4, showing pedicle collar, auriculation of deltidial plates,
corrugated teeth, and small genital areas. 11, 12, Tilted and full
views of the interior of the brachial valve of the preceding specimen
showing cardinalia, pallial sinuses, and muscle scars, X 2. 13, 14, Two
views of the apex of the preceding brachial valve tilted to show the
cardinalia in partial side and partial anterior position, the strongly
corrugated sockets, and the broad outer hinge plates, X 4. 15, Poste-
82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Page
rior view of the cardinalia of the same brachial valve as the preceding,
> 4, showing the broad and flat outer hinge plates. The crura are
shorter than normal because of slight breakage at the distal extremity.
U. S. Bureau of Fisheries Station 5487, 585 fathoms, 52° F., on mud,
off Panaon Island, Philippines.
PHATE SIG
Hispanirhynchia? and Neohemithyris (=Basiliola)
A... Hispanirhynéina?.. species ii ss socds nea 9 Bis ooisere Roe OER eee 60
Figs. 1-5. I-3, Respectively ventral, dorsal, and anterior views of a
somewhat crushed specimen, X 1, figured specimen U.S.N.M. 540361.
4, Dorsal view of the preceding specimen, X 2, showing foramen. 5,
Beak of the preceding, & 4, showing foramen and disjunct deltidial
plates.
Eocene (lower), 200 meters south of the south side of the River-
side Yacht Club, west side Almendares River, in Reparto Kohly, Ha-
bana Province, Cuba.
B. Bastholas lucida CGould) (0 ee Ree a Re eee 34
Figs. 6-23. 6-10, Respectively posterior, anterior, dorsal, side, and ven-
tral views of a complete specimen, 1, showing small size, rounded
form, and nearly smooth exterior, hypotype U.S.N.M. 110826a. 11-13,
Respectively anterior, side, and dorsal views, x 2, of the preceding
specimen showing the same features. 14, Interior of the pedicle valve,
2, showing pallial marks indistinctly, hypotype U.S.N.M. 110826b.
15, Apex of same specimen, X 4, showing corrugated tooth and con-
junct deltidial plates. 16, The same specimen tilted to show the
pedicle collar and dental plates, X 4. 17, 18, Interior and tilted views
of another pedicle valve showing the pallial marks, foramen, teeth,
and deltidial plates, & 4, hypotype U.S.N.M. 110826c. 109, Interior
of the brachial valve, & 2, hypotype U.S.N.M. 110826b. 20, Posterior
part of the same specimen showing the cardinalia with the long falci-
fer crura, X 4. 21-23, Respectively side, tilted anterior, and full views
of another brachial valve, & 4, showing the long falcifer crura, the
small elevated inner hinge plates, corrugated sockets, small genital
areas, and pallial marks, hypotype U.S.N.M. 110826c.
Recent, U. S. Fish Commission Station 4936, rocky bottom at 103
fathoms, Kagashima Gulf, Kyushu, Japan.
PLATE 14
Basiliola and Frieleia
A. Bastliola beechers (Rial) eee faite etletonte eisseiarees to tatetertns te ete oiesetterene 6 25
Fig. 1. Interior of the pedicle valve of an obese specimen, X 2, showing
thickened marginal rim and pallial marks, hypotype U.S.N.M. 334667.
Recent, U. S. Fish Commission Station 3864, 163 to 198 fathoms,
55.9° F., Pailolo Channel, Hawaiian Islands.
Fig. 2. Dorsal view of the apex of a pedicle valve showing the deltidial
plates with their reflected rim and the anterior smooth area of the
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 83
Page
pedicle collar which slides over the umbo of the dorsal valve, X 4,
hypotype U.S.N.M. 274136.
Recent, U. S. Fish Commission Station 3811, 238 to 252 fathoms,
70.5° F.?, south coast of Oahu, Hawaiian Islands.
B. Frieleia? nitens (Conrad) = F.? astoriana (Dall)..........0.0cc0ceee 55
Figs. 3-6. 3-5, Respectively side, dorsal, and ventral views of the holo-
type, U.S.N.M. 3487. 6, Ventral view of another specimen showing
broad sulcus and radial lines on exfoliated shell, 1, paratype
U.S.N.M. 3487a.
Miocene, Astoria, Clatsop County, northwest Oregon.
C. Basiliola elongata Cooper, new species... 16:2... s.cscccccoccecuceccuce, 20
Figs. 7-21. 7-11, Respectively dorsal, anterior, ventral, posterior, and
side views of the holotype, 1, U.S.N.M. 235844a. 12-14, Respec-
tively dorsal, side, and anterior views of the holotype, < 2, showing
smooth surface, elongate form, and growth lines. 15, Interior of the
pedicle valve of the paratype U.S.N.M. 235844b, X 2. 16, Beak region
of the same pedicle valve, X 4, showing the fused deltidial plates and
the reflected rim around the foramen. 17, Same pedicle valve, x 3,
tilted to show the pedicle collar, dental plates, and small genital region.
18, Interior of the brachial valve of the same paratype, X 2, show-
ing elongated falcifer crura. 19, 20, Side and anterior views of the
preceding showing the broad falcifer crura, concave inward, and with
serrate distal extremity, the small reflected inner hinge plates, and the
broad outer hinge plates, & 4. 21, Interior of the apex of the same
brachial valve, 6, showing the falcifer crura, broad outer hinge
plates, small inner plates, and corrugated sockets.
Recent, U. S. Bureau of Fisheries Station 5146, 24 fathoms on
coral sand, Sulade Island, Tapul Group, Philippines.
PLATE I5
Frieleia and Eohemithyris
a rrelera mole MANN ate steals Sead ste et heck, Sane Mee la al le ee 53
Figs. 1-5, 12-14. 1-3, Respectively anterior, brachial, and side views,
X 1, of a complete specimen showing the narrow sulcus in each valve
and the rectimarginate anterior commissure, hypotype U.S.N.M.
110830a. 4, Interior of the pedicle valve tilted to show the strong
dental plates and small teeth, < 2, hypotype U.S.N.M. I10830b. 5,
Apical region of the preceding, X 4, showing the disjunct deltidial
plates and incomplete foramen. 12, Brachial valve tilted to show
apical chamber of the cardinalia, X 2, hypotype U.S.N.M. 110830c.
13, Another brachial valve tilted to show the apical chamber, X 2,
hypotype U.S.N.M. 110830b. 14, The same, x 4, showing the apical
chamber and cardinal process.
Recent, U. S. Fish Commission Station 4797, 682 fathoms, off
Avacha Bay, Kamchatka.
Figs. 6-11. 6-8, Respectively full, partial side, and tilted views of a
brachial valve showing cardinalia and median septum, X 2, hypotype
U.S.N.M. 540348a. 9, 11, Apical part of another brachial valve in full
84
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
and anterior views showing the cardinalia, x2, hypotype U.S.N.M.
540348b. 10, The same, X 4, showing the large inner hinge plates
covering the apical chamber and the small transverse cardinal
process.
Recent, U. S. Fish Commission Station 2923, 522 fathoms, off
San Diego, California.
B. Bohemithyris:colurnus | (tiledley )) io steer Gases shen cas Sere lees teases
Figs. 15-26. 15-17, Respectively brachial, anterior, and side views of a
complete specimen showing the anterior costation and uniplicate ante-
rior commissure, I, hypotype U.S.N.M. 333012b. 18, Enlargement,
< 2, of the beak of the preceding specimen showing the small sub-
mesothyrid foramen and conjunct deltidial plates. 19, 20, Two views
of the interior of a pedicle valve, X 2, one in full view, the other
anteriorly tilted to show the minute dental plates, small genital areas,
and pallial sinuses, hypotype U.S.N.M. 333012a. 21, Apical part of the
preceding specimen showing beak, foramen, and conjunct deltidial
plates, X 4. 22, 23, Full view and slightly tilted view of the brachial
valve of the preceding specimen showing the cardinalia, pallial sinuses,
and genital areas, X 2. 24, 25, Partial side and full views of the cardi-
nalia showing the distally serrate, falcifer crus, corrugated sockets,
and thickening over the crural bases, X 4. 26, Posterior part of the
preceding tilted to show the concave ends of the crura, the small
genital areas, and the interior thickening, x 4.
Recent, 115-135 fathoms, off Gabo Island, Victoria, Australia.
PLATE 16
Compsothyris
A: Combsothurts racovttage ( Joubina) vcs): citer jas ox sis0.¢ bisa «eile Ore nies
Figs. 1-17. 1-4, Respectively anterior, dorsal, ventral, and side views of a
complete individual, showing faintly uniplicate commissure, < 1, hypo-
type U.S.N.M. 549343. 5, Dorsal view of the preceding specimen
showing fine closely crowded costellae, X 2. 6, 7, Interior of the
pedicle valve of the same specimen, X 2, showing small foramen and
small dental plates. 8, Beak of the preceding valve, X 4, showing
small corrugated teeth and small disjunct deltidial plates. 9-11,
Respectively full, slightly tilted, and strongly tilted views of the
brachial interior of the same specimen showing cardinalia, median
septum, and muscle scars, X 2. 12, Same brachial interior tilted to
show the socket ridges and distally serrate spinulifer crura, x 4.
13-15, Three views of the cardinalia variously tilted to show socket
ridges, narrow outer hinge plates, and crura, X 4. 16, Same brachial
valve strongly tilted to show junction of crural supporting plates with
median septum, X 4. 17, Exterior of the pedicle valve, X 6, showing
the very fine radial costellae.
Recent, British Antarctic Expedition 1910, Station 316 of Terra
Nova, 190 to 250 fathoms, 30.5° F., off Glacier Tongue, 8 miles north
of Hut Point, McMurdo Sound, Antarctic.
Page
32
56
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER
PLATE 17
Probolarina
Aa Prapowurina saris (COadll pvcn/ctie sess cece pele cmela canis a csoas s.asiias.as
Figs. 1-19. 1-5, Respectively, posterior, ventral, side, dorsal, and anterior
views of a small but complete individual, <2, hypotype U.S.N.M.
5409357a. 6, 7, Dorsal and ventral views of another hypotype showing
variation of costation, X 1, U.S.N.M. 549355. 8-10, Respectively dorsal,
ventral, and anterior views of a specimen larger and more strongly
costate than the two preceding ones, X 1, hypotype U.S.N.M. 549357b.
11-14, Respectively side, anterior, ventral, and dorsal views of a small
specimen with few costae, X 3, holotype U.S.N.M. 109293a. 15, Apical
portion of a large specimen showing the small submesothyrid fora-
men, conjunct and auriculate deltidial plates, X 4, hypotype U.S.N.M.
549354a. 16, Interior of a brachial valve, X 3, showing cardinalia,
hypotype U.S.N.M. 549356d. 17, 18, Partial side and full views of the
apical part of the same specimen, X 6, showing the concave falcifer
crura and large outer hinge plates. 19, The same tilted anteriorly to
show the concave crura and lack of median ridge, X 6.
Eocene (Castle Hayne formation), at the city quarry near the
cemetery, Wilmington, North Carolina.
Ba Progolarina: Doles Call) 55 aesaissnicierainlavess sssto «0.556; o0e: wide Win, 6) Saad gets Os
Figs. 20-36. 20-24, Respectively posterior, anterior, dorsal, ventral, and
side views of a complete specimen, 2, hypotype U.S.N.M. 549350a.
25, The same, X 3, showing the ornamentation and long beak. 26, 27,
Beak of the same specimen, X 5, showing conjunct and strongly auric-
ulate deltidial plates. 28, Small specimen showing foramen and con-
junct deltidial plates, & 4, hypotype U.S.N.M. 549359b. 20, 30, Apical
part of another specimen showing conjunct and auriculate deltidial
plates, X 6, and the same tilted to show the dental plates and pedicle
collar, X 6, hypotype U.S.N.M. 549350e. 31, The same tilted to the
side to show the pedicle collar, X 4. 32, Apex of another pedicle
valve showing strongly auriculate deltidial plates, 6, hypotype
U.S.N.M. 549359f. 33, Interior of the brachial valve, X 3, showing
cardinalia and absence of median ridge, hypotype U.S.N.M. 549350g.
34, 35, Apical part of the preceding tilted to show concave falcifer
crura, X 6. 36, Same in full view to show the outer hinge plates, x 6.
Horizon and locality same as above.
PLATE 18
Erymnaria
A. Erymnaria. bolcensis, (Massalongo) qs 422.00) «:cisjstersieidlawie’se «iblasis'qeu e 6
Figs. 1-17. 1-5, Respectively dorsal, side, posterior, anterior, and ventral
views of a decorticated specimen showing symmetrical form, X 1,
U.S.N.M. 549383b. 6, Posterior view of preceding, X 3, to show
dorsal umbo, trace of the crural supporting plates, and the shorter
socket ridges outside of them (photographed under water). 7, Anterior
of same specimen, X 2, showing symmetrical fold on brachial valve.
86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Page
8-12, Respectively posterior, anterior, side, dorsal, and ventral views
of another symmetrical specimen, X 1, hypotype U.S.N.M. 549383a.
13-15, Respectively dorsal, anterior, and side views of the preceding,
x 2. 16, 17, Two views of cross sections of the beak of the brachial
valve U.S.N.M. 549383c, ca. X 8, respectively 2.6 mm. and ca. 6.0 mm.
anterior to the beak, showing crura, crural supports, and socket ridges
(see pl. 22, fig. 9, and explanation).
Lower Eocene (Spilecciano), Spilecco, Verona, Italy.
B. Erymnaria polymorpha (Massalongo) ..........0...eeceecenscceecenes 64
Figs. 18-25, 31-34. 18-22, Respectively ventral, dorsal, side, anterior, and
posterior views of a large but imperfect specimen showing twisted
anterior commissure and traces of pallial marks, 1, hypotype
U.S.N.M. 75888a. 23, 24, Dorsal and anterior views of the same speci-
men, X 2, showing trace of vascula media and twisted anterior com-
missure. 25, Dorsal view of the same specimen taken under water
and tilted away from the observer to show the long crural supporting
plates as black lines and the shorter socket ridges, & 3. 31-33, Respec-
tively side, anterior, and dorsal views of the preceding specimen, X 2,
showing costae along the anterior margin. 34, Another specimen tilted
away from the observer and taken under water to show the trace of
the long crural supporting plates and the shorter socket ridges
diverging widely from the beak, x 4, hypotype U.S.N.M. 75888b.
Lower Eocene, Spilecco, Verona, Italy.
Figs. 26-30, 35, 36. 26-30, Respectively anterior, ventral, side, posterior,
and dorsal views of another specimen, not decorticated like the pre-
ceding, and showing, in addition to the twisted commissure, short
radial costae, X 1, hypotype U.S.N.M. 549384a. 35, Posterior of a
brachial valve excavated to show cardinalia and crural supporting
plates, X 6, hypotype U.S.N.M. 540384b. 36, Another brachial valve
interior showing cardinalia with their fairly broad outer hinge plates,
and crural supporting plates, < 6, hypotype U.S.N.M. 5409384c.
Lower Eocene (Spilecciano), Spilecco, 400 meters southwest of
Purga di Bolca, Monti Lessini, Verona, Italy.
PLATE 19
Erymnaria and Streptaria
A Eramnaria cubensis Coopenan. Js. cka2taee eins soe cele ales tates ac cee 65
Figs. 1-10. 1-5, Respectively dorsal, anterior, ventral, posterior, and side
views of the holotype, X 1, U.S.N.M. 540385. 6-8, Respectively side,
dorsal, and anterior views of the same specimen, X 2, showing folding.
9, Posterior of holotype, X 3, showing conjunct deltidial plates and
foramen. 10, Holotype tilted and photographed under water to show
crural supporting plates and the shorter, more widely divergent
socket ridges, X 3.
Eocene, 80 meters northeast of school, Chucho Machin, Matanzas
Province, Cuba.
B. Streptaria streptimorpha Cooper, new species...........-...eeeeeeeeee 40
Figs. 11-21. 11-15, Respectively anterior, ventral, side, dorsal, and pos-
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 87
Page
terior views of the holotype, X 1, showing twisted anterior commis-
sure, U.S.N.M. 549386a. 16, Dorsal view of the holotype showing
smooth exterior, X 13. 17, 18, Side and anterior views of the holo-
type, X 2, showing twisted commissure. 19, Posterior of the holotype,
3, showing deltidial plates and foramen. 20, 21, Posterior of another
specimen, < 4, showing short dental plates and cardinalia with falcifer
crura (see discussion), paratype U.S.N.M. 549386b.
Eocene, deep cut north of Grua 9, Ramal Juan Criollo, Camaguey
Province, Cuba.
C. Streptaria buchi (Michelotti) ........ cece ee cece eee e cence eer ceeeeees 38
Figs. 22-31. 22-26, Respectively ventral, dorsal, anterior, side, and
posterior views of a complete specimen showing twisted anterior com-
missure, X 1, hypotype U.S.N.M. 549352a. 27-20, Respectively ante-
rior, dorsal, and side views of the preceding specimen, X 2, showing
twisted commissure. 30, Posterior of the same specimen, X 3, show-
ing large foramen with elevated rim. 31, Posterior of the brachial
valve of another specimen prepared to show the strong socket ridges,
wide outer hinge plates, and falcifer crura with thickened ventral
edge, X 6, hypotype U.S.N.M. 549352b.
Middle Miocene, Messina, Sicily.
PLATE 20
Eohemithyris
A. Eohemithyris alexi Hertlein and Grant..........ccceesecceceseccceees 30
Figs. 1-16. 1-5, Respectively dorsal, posterior, ventral, side, and ante-
rior views of a complete and undistorted specimen, showing form and
true profile of valves, X 1, hypotype U.C.M.P. 15524. 6-9, Respec-
tively, anterior, ventral, dorsal, and side views of the same specimen,
<2. 10, Dorsal view of another nearly perfect specimen showing
rounded outlines and low fold, 1, hypotype U.C.M.P. 15526. 11,
Posterior of the same specimen, X 3, showing small foramen. 12,
Posterior of another specimen, X 3, showing small foramen and con-
junct deltidial plates, hypotype U.C.M.P. 15541. 13, 14, Dorsal and
ventral views of a specimen from which the shell has been scraped to
show the muscle fields, & 2, hypotype U.C.M.P. 15542. 15, 16, Full
view and partial side view of the cardinalia, x 6, showing narrow
outer hinge plates and broad, long falcifer crura, hypotype U.C.M.P.
15545.
Eocene (Domengine formation), from just below the 1/4 section
marker toward the top of the 25-foot last sandstone “reef” on the ridge
on the east side of the North Fork of Media Agua Creek, grid.
coord. 142001-139004, McKittrick (15’) Quadrangle, Kern County,
California (see text for further information).
Bo Baheathres, Gave CNV GOGWALG) i, «ai. 5:5:»/orctors fargrs! oyapeieve a olel atten ©\e abno aislels 33
Figs. 17-23. 17, Dorsal view of the exterior, ca. X 2, showing anterior
costae and minute foramen, holotype British Museum (Nat. Hist.)
ZB280. Specimen coated by ammonium chloride. 18-20, Respectively
88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.
side, dorsal, and anterior views of the same specimen, ca. X 2. 21,
Posterior of the same specimen (coated) showing small foramen, ca.
X 24. 22, Interior of the pedicle valve of the holotype, ca. X 23,
showing small foramen, muscle field, and pallial marks. 23, Interior
of the brachial valve of the holotype showing cardinalia with falcifer
crura, thickened inner edges of crural bases, muscle field, and pallial
marks, ca. X 4.
Recent, Fiji Islands.
Photographs by permission of the Trustees of the British Museum
(Natural History) through Dr. H. M. Muir-Wood Deputy Keeper,
Department of Palaeontology.
PLATE 21
Frieleia, Patagorhynchia, Septaliphoria, Cryptopora, Tegulorhynchia,
Mannia, and Hispanirhynchia?
A. Hemithyris astoriana Dall= Frieleia ? mitens Conrad............2000%
Fig. 1. Section of the apex of the brachial valve showing the small
chamber and median septum, X 8, paratype U.S.N.M. 3487a.
Miocene, Astoria, Clatsop County, Oregon.
B. Patagorhynchta patagomeca (Thering) 2. v2.30 sciedea ee selene o ejeeiartee see
Figs. 2-5. Respectively anterior, dorsal, side, and ventral views of a
fairly well-preserved specimen showing costellate ornamentation,
x1, figured specimen U.S.N.M. 549387.
Eocene, 1 to 2 miles southwest of Ancha Terriza, Rio de los Cier-
vos, Provincia de Magallones, Chile.
C, Sepralt phoma species: sec cle aa ae tee ain ea oe oc e ee ack eee ae vtelore eatersie
Fig. 6. Interior of the dorsal valve tilted to show septalium and median
septum, X 2, figured specimen U.S.N.M. 129896a.
Jurassic (Lower Callovian), in the railroad cut 300 meters east
of the station at Chatillon-sur-Seine, Department of Cote d’Or,
France.
D. \Cryptopora gnome C(ICHLEYS )le/.(. x o's< o's creyse esis ais alecals,ss sacnislecce eels syelute
Figs. 7-14. 7-9, Respectively anterior, side, and ventral views of a com-
plete specimen, X 6, hypotype U.S.N.M. 44911a. 10, Anterior of an-
other specimen, <6, showing the sulcate anterior margin, hypotype
U.S.N.M. 44911c. 11, Dorsal view of the preceding, < 8, showing
the nonauriculate deltidial plates. 12, Posterior of the pedicle valve,
<8, showing deltidial plates, apical plate, and teeth, hypotype
U.S.N.M. 44911d. 13, 14, Full and partial side views of the brachial
valve of the preceding interior showing median septum and maniculifer
crura.
Recent, U. S. Fish Commission Station 2221, 1,525 fathoms, 36.9° F.,
south of Marthas Vineyard, Massachusetts, in gray ooze.
EB. Tegulorhynchia doderlews ‘((Davyidson)). ws sans. 2 2s ce ccvancsete«
Fig. 15. Interior of the pedicle valve, X 1, showing pallial sinuses, after
Leidhold, Neues Jahrb. Min., Geol., Palaont., BB. 45, pl. 11, fig. 1b
[here reduced 1/2], 1922. Original in Institute of Zoology, Strasburg
University.
Recent, Sagami Bay, Honshu, Japan.
139
Page
56
44
10
22
51
NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER
PERV C1190) AY SEs sD A VAG SOME oycyercicuekteveleversisjevaictokece le PAV alsietate Wreianal siataretaeks reeset s
Figs. 16-20. 16, Drawing of the dorsal side, ca. X I. 17, Exterior of the
dorsal side, ca. X 7, showing large foramen and long, elevated delti-
dial plates. 18, 19, Interior of two brachial valves showing septum
with spoonlike plate and long maniculifer crura. 20, Cross section
through a complete individual showing relationship of valves, median
septum, spoonlike plate, and crura. All from Davidson, Geol. Mag.,
dec. 2, vol. 1, No. 4, pl. 7, 10-13, 1874.
Miocene (Diestien), 3 miles east of Antwerp, Belgium.
G.. Elaspanwhynchia °° craneana: (Dall)... Sear Fe Eee cues Hoe iia we
Figs. 21-26. 21-23, Respectively dorsal, anterior, and side views of the
holotype showing gently sulcate anterior commissure, X 1, U.S.N.M.
122861. 24, The holotype, 2, showing the beak area. 25, The beak
region of the holotype, X 4, showing teeth (the deltidial plates prob-
ably have been broken away). 26, Posterior of the brachial valve
showing spinulifer crura and small outer hinge plate (these struc-
tures have been damaged).
Recent, U. S. Fish Commission Station 3362, mud at 117 fathoms
and 36.8° F., off Cocos Island, Pacific Ocean off Panama.
PLATE 22
Eohemithyris, Erymnaria, Notosaria, Aphelesia
A. Bohemihyris: alex: Hertlein' and: Grant... 56 cic ew cln ele cee cceccseens
Figs. 1-3. 1, 2, Full and partial side views of the cardinalia to show
broad, falcifer crura, ca. X 5, hypotype U.C.M.P. 15545. 3, Drawing of
the posterior of a pedicle valve, ca. & 4, showing the conjunct deltidial
plates and small, round foramen, hypotype U.C.M.P. 15524.
Horizon and locality as in plate 20, figures 1-16.
B. Erymnaria polymorpha (Massalongo) ..........cccccscccnseccescccvcs
Figs. 4-9. Sections through a slightly distorted individual, ca. X 4,
hypotype U.S.N.M. 549384e. Sections measured from pedicle beak
respectively: (4) ca. I mm., (5) 1.7 mm., (6) I.95 mm., (7) 2.25 mm.,
(8) 2.6 mm., and (9) ca. 4.0 mm.
Lower Eocene (Spilecciano), 400 meters southwest of Purga di
Bolca, Monti Lessini, Verona, Italy.
Figs. 10-15. Sections through another individual showing long crural
supporting plates, ca. X 4, British Museum (Natural History) B 8088.
Sections respectively 0.3 mm. apart except figure 13 which is 0.4 mm.
from figure 12.
Eocene, Castelvecchio, Vicentin, Italy.
C. Tegulorhynchia (= Notosaria) nigricans (Sowerby).............20+:
Figs. 16, 17. Diagram of the interior of the pedicle and brachial valves
of the adult showing pallial sinuses, ca. & 1.5, after Williams (1956,
fig. 7, No. (4) on p. 276).
Recent, New Zealand.
61
30
64
48
go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
DA phelésia biparttia, .GBcoccht ) icine 2% kx teiccmie wiciel Mele eee tatale ele aiie 41
Figs. 18-25. Sections through the valves of a full-grown adult, Xx 2,
British Museum (Natural History) not numbered, showing sockets
and teeth. Sections respectively from the beak of the pedicle valve:
(18) 1.0mm., (19) 1.6mm., (20) 2.0mm., (21) 26mm., (22)
2.8mm., (23) 3.2mm., (24) 3.6mm., and (25) 4.0mm.
Miocene, St. Lorenzo, Tuscany, Italy.
Photographs of figures 10-15 and 18-25 by permission of the Trustees of the
British Museum (Natural History) through Dr. H. M. Muir-Wood, Deputy
Keeper, Department of Palaeontology.
cH a
iyi
Dau
Pita cri Mine a a,
ach ay ie a)
hy rh i j iu f ea ' vi ra
tS is i ua 7 i ry i ; iy
fe int! hate a a ( ia
Mab ve ; i vat a Wat i
Ca i h ah ae Ne Onn iu
' } i Poe . ; gt Gea Ron at Ht i aa
i
ny AN Lt ae ne ji ‘
r i At ‘ HLH ‘ | " WV ae it tl) \ :
ras i | i } Ne i Ms eh a
Hea td . An acae ee
a ) ina
7 i / |
j j ‘* ’
‘ i \ s : Pies
7 x t (put : i
Fae { a i +H ¥ iit
mt oa Ay ret y hal i
if { ; Biss Neg
4i'3{) H f i F ’
iv Wie 4 ! ;
Ri ‘al i eel | 7 i |
Hie x : 1 , i i My
} , y Mi a Ae
. mT i ‘ { ay i /
a ah a i PAWEL LST ea
, MT 7 j ;
| " A ie , , yt f
Pi } f anh f ‘ i My
you . r B ; Ma
}
ur ny, '
Al; \ i.
Von i y = 3 veal nt \tn Ny
f 4
i i my YA \ Te Se My
1 4 My
ii t q
Ap,
Wn J ‘ N
‘ ;
. ‘
l
is |
1 t y ‘
: J Sa
Pe Pn
wa
sally
i i rl
ih
| iy
‘ i |
\ Me
' i hat 7
. j i} yi
| Crd pet
1h
4
’ 7 ue ‘ i j
\ 4 cm. - t ¥ ‘
\ 1
iP 1 ys f i 7 y
J ce ,
1 fi
7) i ‘ . oy
\
Vi
leat
\
VOL. 139, NO. 5, PL. 1
THSONIAN MISCELLANEOUS COLLECTIONS
CRYPTOPORA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES {39 NO 5. PEs
CRYPTOPORA AND NEORHYNCHIA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 3
A
HEMITHYRIS
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 4
AETHEIA, FRIELEIA, GRAMMETARIA, CAMAROTOECHIA, AND HEMITHYRIS
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
THSONIAN MISCELLANEOUS COLLECTIONS
VOL. 139, NO. 5, PL. 5
i
Ho
x
cas
Ay re
EOHEMITHYRIS, SPHENARINA, CRYPTOPORA, AND TEGULORHYNCHIA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, ‘NO: 5, PIS
PATAGORHYNCHIA AND NOTOSARIA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
VOL2 139) |NO2 5; Pia
THSONIAN MISCELLANEOUS COLLECTIONS
PLICIRHYNCHIA AND APHELESIA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL
17
SPHENARINA, APHELESIA, AND EOHEMITHYRIS?
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 9
GRAMMETARIA AND AETHEIA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 10}
HISPANIRHYNCHIA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
MITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 11
RHYTIRHYNCHIA AND BASILIOLA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139; NO: 5, (PES if2
BASILIOLA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 13
HISPANIRHYNCHIA? AND NEOHEMITHYRIS = BASILIOLA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
VOL. 139, NO. 5, PL. 14
SMITHSONIAN MISCELLANEOUS COLLECTIONS
BASILIOLA AND FRIELEIA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
MITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 15
A
FRIELEIA AND EOHEMITHIRIS
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
VOL. 139, NO. 5, PL. 1
SMITHSONIAN MISCELLANEOUS COLLECTIONS
COMPSOTHYRIS
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 139, NO. 5, PL. 17
PROBOLARINA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 18
ERYMNARIA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 19
ERYMNARIA AND STREPTARIA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 20}
EOHEMITHYRIS
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
MITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 21
A
FRIELEIA, PATAGORHYNCHIA, SEPTALIPHORIA, CRYPTOPORA, TEGULORHYNCHIA,
MANNIA, AND HISPANIRHYNCHIA?
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 22
EOHEMITHYRIS, ERYMNARIA, NOTOSARIA, AND APHELESIA
(SEE EXPLANATION OF PLATES AT END OF TEXT.)
RS Oe Ee pe eee ey
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 6
Charles D. and Mary Waux THalcott
Research Fund
A REVISION OF THE SILURIAN
BRYOZOAN GENUS TREMATOPORA
(WitTH 2 PLATEs)
By
RICHARD S. BOARDMAN
Associate Curator of Geology
United States National Museum
Smithsonian Institution
(PuBLICATION 4383)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 29, 1959
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 6
Charles D. and Mary Waux Talcott
Research Fund
A REVISION OF THE SILURIAN
BRYOZOAN GENUS TREMATOPORA
(WiTH 2 PrateEs)
By
RICHARD S. BOARDMAN
Associate Curator of Geology
United States National Museum
Smithsonian Institution
HSOW SS I
ATV Wee
tI NGTON 32?
ee0ee8®
(PuBLIcaTION 4383)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 29, 1959
THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.
Charles D. and Marp Vaux Walcott Research Fund
A REVISION OF THE SILURIAN BRYOZOAN
GENUS TREMATOPORA
By RICHARD S. BOARDMAN
Associate Curator of Geology
United States National Museum
Smithsonian Institution
(WitH Two P ates)
INTRODUCTION
The genus Trematopora Hall, 1851, is placed in the order Tre-
postomata of the Bryozoa and is the type genus of the family
Trematoporidae Ulrich in Miller, 1889. The type species of Tre-
matopora is T. tuberculosa Hall, 1852, from the Rochester shale in
New York (type by subsequent designation, Ulrich, 1882, p. 241).
The name Trematopora was established in an article by the editors
of the American Journal of Science and Arts (Hall, 1851, p. 400)
in which parts of Hall’s manuscript for volume 2 of the Paleontology
of New York (1852) were quoted. The species of Trematopora listed ~
following the diagnosis of the genus were nomina nuda and were not
published by Hall until the next year in volume 2.
The development of the generic concept of Trematopora has been
controlled partly by the study and preparation techniques employed by
the various authors, each advance in technique adding refinements to
the original very generalized description. All the work of Hall and
Hall and Simpson (1851-1887) was done on external characters with-
out the use of thin sections. In fact, Hall’s primary types of the type
species were sectioned for the first time for the present paper. Owing
in part to the external homeomorphy common in the Trepostomata,
Hall included in the genus many forms now placed in other genera,
families, and orders. At various times Hall considered such diverse
genera as Tremaiella Hall, Orthopora Hall, Chaetetes Fischer (part),
and Callopora Hall (part), as subgenera of Trematopora.
Ulrich (1883, p. 257) was the first to section some “authentic speci-
mens of Trematopora tuberculosa Hall.” These sections are in the
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 6
x
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
U. S. National Museum collections and are conspecific with Hall’s pri-
mary types of the species. After seeing the sections, Ulrich greatly
restricted the concept of the genus and indicated the great range of
forms that Hall had included in the genus. The concept established
by Ulrich in 1883 has remained essentially unchanged to the present
time and was the type-genus concept for the family Trematoporidae
in 1889. Under Ulrich’s definition of the genus, 12 species and sub-
species have been assigned to Trematopora, ranging in age from
Middle Ordovician through Middle Silurian.
The primary type specimens were made available for sectioning and
study by N. D. Newell of the American Museum of Natural History.
Helpful suggestions were made by Helen Duncan and W. A. Oliver,
Jr., of the U. S. Geological Survey, and N. Spjeldnaes, of the Uni-
versity of Oslo. Thin sections were prepared by T. M. Robison of the
U. S. Geological Survey. Photography was done by J. Scott, and the
text figure was drawn by L. B. Isham, both of the Department of
Geology of the U. S. National Museum.
INTERPRETATION OF SKELETAL MICROSTRUCTURE
The skeletal structures of most trepostomatous Bryozoa are com-
posed of finely laminated calcite (fig. 1 and pl. 2). These laminae are
assumed to have been deposited parallel to the surface of the secreting
tissue (Cumings and Galloway, 1915, p. 361). Therefore, trends of
the laminae within skeletal structures such as walls and diaphragms
are considered to reveal something of the disposition of the original
secreting tissue and the mode of growth of the skeletal structures.
In longitudinal thin sections of T. tuberculosa, laminae are com-
monly oriented parallel to the zooecial walls (fig. 1) in the endozone
(immature or axial region of authors) and to the thinner walls and
mesopore diaphragms in the inner region of the exozone (mature
region of authors). This type of microstructure is here designated
longitudinally laminated structure. Such an orientation of laminae
is assumed to indicate that the depositing tissue was parallel to the
walls and diaphragms, but it does not indicate whether the laminae
were deposited on one or both sides of the structures.
Another type of structure is characterized by laminae that are
curved or angled transversely to the walls and diaphragms as seen in
longitudinal sections. The transverse laminae form V- or U-shaped
patterns with apices pointing distally and aligned along the median line
of a wall or diaphragm. This type of microstructure is here designated
transversely laminated structure. In T. tuberculosa, this structure is
No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN 3
found in the walls of zooecia and mesopores in the outer region of the
exozone, and in the inner region in some of the thicker mesopore walls
and in the vicinity of central pores in the mesopore diaphragms
(fig. I).
Assuming that secreting tissue was oriented parallel to the laminae,
transversely laminated structure indicates that the tissue must have
been wrapped around the growing edges of walls and diaphragms
Inner Region Outer Region
<—- Endozone ——>\< of Exozone ——><— of Exozone ———>
1
!
i
!
I
I
!
1
y
MESOPORE +
Fic. 1.—Idealized diagram of T. tuberculosa in longitudinal view illustrating
the variety of laminated structures commonly occurring in the species. Two
mesopores and an intervening zooecium are shown in profile. Few central pores
of the mesopore diaphragms are intersected in a longitudinal section.
on both sides of the median lines. Thus, transversely laminated struc-
ture presumably indicates deposition from both sides of a wall or
diaphragm.
Such interpretations applied to the skeletal laminae of T. tuberculosa
and correlated with other morphologic characteristics of the species
suggest that the exozone is divisible into two parts, an inner and an
outer region (fig. 1) based on fundamentally different modes of
growth of the mesopores. The physiologic significance of the two
modes of growth is a matter for speculation. The taxonomic signifi-
cance of these characters must await comparable studies in related
genera.
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
In the inner region of the exozone, the mesopores are beaded (series
of rounded chambers). This beading is produced by the mesopore
walls curving transversely to the axis of the mesopore to form a
diaphragm. The succession of longitudinally and transversely lami-
nated structures along the walls and diaphragms is interpreted to
indicate deposition from both sides of longitudinal as well as trans-
verse structure throughout the inner region. Thus, at least a part of
the depositing tissue of the mesopore remained behind the diaphragm
and within the chamber being formed. The pore in the center of the
mesopore diaphragm presumably would be necessary to allow the soft
parts within the chamber to communicate with the outside environ-
ment. There is no indication as to whether the soft parts in the
mesopores during the formation of the inner region consisted of any-
thing more than a depositing tissue or mantle, but the distal diaphragm
would have acted, temporarily at least, as a covering for any soft parts
within the last chamber. Continuity of laminae from the distal side of
a diaphragm to the wall of the succeeding mesopore chamber implies
that at least the proximal part of the chamber was formed as the dia-
phragm developed (see middle diaphragm of the inner region in the
upper mesopore of figure 1). For a more complete discussion of
figure I see species description.
The thick-walled outer region of the exozone contains transversely
laminated structure in the walls of zooecia and mesopores and longi-
tudinally laminated structure in the thick diaphragms. The mesopores
in this region are not beaded. The pattern of continuity of the laminae
of walls and adjoining diaphragms (fig. 1) indicates deposition was
limited to the outer surface of the walls and diaphragms, con-
temporaneous deposition of laminae taking place on the outermost
surfaces of the zooecial wall, around the median line or boundary to
the mesopore wall, and back to the distal side of the diaphragm. There
is no evidence that deposition occurred on the proximal side of the
diaphragm within the mesopore chamber.
The formation of the diaphragms in the inner and outer regions
of the exozone is quite different. Diaphragms in the inner region were
formed by continued distal growth of mesopore walls that merely
curved through an angle of 90 degrees to form transverse structures.
Diaphragms in the outer region were formed by an additional, trans-
versely oriented sheet of depositing tissue that was continuous with at
least the depositing tissue of the mesopore side of the walls and ac-
tively deposited calcite at the same time that the mesopore walls were
being formed. This transverse sheet of tissue apparently had no
counterpart in the inner region of the exozone.
No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN 5
Evidence concerning the position of the soft parts in the outer
region of the mesopore is inadequate. Configurations of the laminae
give no indications of deposition behind the distalmost diaphragm.
The common occurrence of single, centrally located pores that either
partly or completely penetrate the thick outer diaphragms suggest some
activity within outer chambers. The majority of these central pores
appear to have been cut through the laminae of the diaphragms. Their
termination within outermost diaphragms suggests that activity within
the outermost chambers might have been choked off by the growth of
the thickened diaphragms.
SYSTEMATIC DESCRIPTION
Genus TREMATOPORA Hall, 1851
1851. Trematopora Hall, Amer. Journ. Sci. and Arts, ser. 2, vol. 11, p. 400.
1852. Trematopora Hall, Paleontology of New York, vol. 2, p. 1490.
1881. Trematopora Hall, Nicholson, Genus Monticulipora, pp. 232-234.
1882. Trematopora Hall, Ulrich, Journ. Cincinnati Soc. Nat. Hist., vol. 5, p.
241.
1883. Trematopora Hall, Ulrich, Journ. Cincinnati Soc. Nat Hist., vol. 6, p.
257.
1887. Trematopora Hall, Hall and Simpson, Paleontology of New York, vol.
6, p. Xiv.
1893. Trematopora Hall, Ulrich, Geol. Minnesota, vol. 3, pt. 1, p. 308.
1911. Trematopora Hall, Bassler, U. S. Nat. Mus. Bull. 77, pp. 267, 268.
1882. [non] Trematopora Hall, Ulrich, Journ. Cincinnati Soc. Nat. Hist., vol.
5, DP. 153.
Type species —Trematopora tuberculosa Hall, 1852.
Emended definition—Zoaria are ramose, conspecific overgrowth is
common, and monticules range from level to tuberculated. Externally,
zooecia are elliptical to subcircular in cross section and walls are
slightly elevated above intervening mesopores. Mesopores form
shallow, subpolygonal depressions between zooecia.
The exozone is divided into an inner thin-walled region containing
the earliest chambers of the mesopores and an outer thick-walled
region. In the inner region, both mesopores and zooecia are polygonal
to subpolygonal in cross section and mesopores are beaded and contain
diaphragms with single central pores. In the outer region of the
exozone, zooecia become elliptical to subcircular in cross section and
the mesopores contain thickened diaphragms and are not beaded.
Diaphragms are thin and few in zooecia.
In the outer region of the exozone, walls of adjacent zooecia are
divided by sharply defined zooecial boundaries, as seen in longitudinal
sections. Laminae on either side of a boundary converge to form a
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
V-shaped pattern that has extremely long limbs trending nearly
parallel to the boundary and curving very slightly just before the
boundary is intersected. Laminae of walls of zooecia and adjacent
mesopores are more broadly curved approaching the median boundary
and form a broad U-shaped pattern having limbs of varying lengths.
Acanthopores are common in the zooecial walls.
Discussion.—Based on an examination of thin sections of primary
types of species previously assigned to Trematopora now in the U. S.
National Museum collections, the following species are considered cor-
rectly assigned to the genus:
T. halli Ulrich 1883, Niagaran group, Waldron, Ind.
T. whitfieldi Ulrich 1883, Niagaran group, Waldron, Ind.
The holotype section of T. spiculata Miller 1877, Niagaran group,
Waldron, Ind., is not adequate to determine generic affinities. This
species is retained in the genus until a more detailed study of addi-
tional material can be made.
The following species originally placed in Trematopora do not
satisfy the generic definition proposed here and are not considered to
belong to the genus. Their proper generic assignments must await
restudy of both the species themselves and the available genera.
calloporoides Ulrich 1890, Cincinnati group, Alexander County, III.
cystata Bassler 1911, Kuckers shale (C2), Reval, Esthonia. This
species is the type of Aostipora Vinassa 1920).
debilis Ulrich 1890, Girardeau limestone, Alexander County, III.
kukersiana Bassler 1911, Kuckers shale (C2), Reval, Esthonia.
primigenia Ulrich 1886, Decorah shale, Minneapolis, Minn.
primigenia var. ornata Ulrich 1886, Decorah shale, Minneapolis,
Minn.
None of the Ordovician species investigated displayed the two
regions of the exozone or pores in the mesopore diaphragms. Thus,
the genus is limited presently to the Middle Silurian.
A close taxonomic relationship seems to exist between Trematopora
and some or all of the Silurian and Devonian species that have been
placed in the genus Leioclema Ulrich. These species of Letoclema
are largely incrusting and possess many of the morphologic characters
now defining Trematopora. In general they have elliptical zooecia
with few thin diaphragms, abundant mesopores with closely spaced
thicker diaphragms and an irregularly discontinuous inner region of
the exozone containing beaded mesopores. Pores in the diaphragms
of the mesopores are rare but do definitely occur in the following
species.
No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN 7
Leioclema asperum (Hall) 1852, Rochester shale, Lockport, N. Y. (Only
Bassler’s plesiotypes of 1906 available.)
L. confertiporum (Hall) 1883, Hamilton group, New York.
L. decipiens (Hall) 1883, Hamilton group, New York.
L. passitabulatum Duncan 1939, Traverse group, Michigan.
The region now considered to be the inner region of the exozone in
species of Leioclema from the Hamilton group of New York was
interpreted as the outer part of the endozone (Boardman, in press)
and diaphragm pores were overlooked.
TREMATOPORA TUBERCULOSA Hall
Pl. 1, figs. 1-4; pl. 2, figs. 1-3
1852. Trematopora tuberculosa Hall, Paleontology of New York, vol. 2, p. 149,
pl. 40A, figs. Ia-g.
1883. Trematopora tuberculosa Hall, Ulrich, Journ. Cincinnati Soc. Nat. Hist.,
vol. 6, p. 259, pl. 13, figs. 2, 2a, 2b.
1906. Trematopora tuberculosa Hall, Bassler, U. S. Geol. Surv. Bull. No. 292,
p. 43, pl. 13, figs. 15, 16; pl. 17, figs. 1-3; pl. 25, fig. 8.
TYPE DATA
Lectotype (Hall, 1852, pl. 40A, fig. 1a) and the two paratype zoaria
from syntype suite No. 1747, American Museum of Natural History.
MATERIAL STUDIED
In addition to the primary types, 55 fragmentary topotype zoaria
were studied. The topotypes are from U. S. National Museum col-
lection No. 2998 and cat. No. 43618 collected by E. O. Ulrich. U. S.
National Museum catalogue numbers of illustrated topotypes are
137847 to 137850.
OCCURRENCE
Rochester shale member of the Clinton formation, Lockport, N. Y.
DESCRIPTION
Zoaria.—Zoaria are ramose and branches are circular to elliptical
in cross section. Branch arrangement was affected by branches rising
from conspecific secondary growth superimposed on the normal bi-
furcating pattern. Branches of secondary growth produced irregu-
larities in branch arrangement, commonly causing branches to
anastomose and form erratic and confused zooecial growth at surfaces
of contact. These irregularities in branch arrangement were formed
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
by random bends at ramose extensions of overgrowths beyond the
distal tips of primary branches, and irregular branch angles in lateral
secondary branches. The zoaria were further complicated by repeti-
tions of thin- and thick-walled growth in the outer region of the
exozone (mature region) without the formation of intervening basal
laminae. This apparently rejuvenated growth formed localized swell-
ings on the zoaria, and combined with adjacent patches of overgrowth
to form some of the secondary branches.
Monticules.—Monticules are prominent tubercles. The apertures
of some monticular zooecia are restricted or closed by a distal thicken-
ing of the walls, and the walls and outer diaphragms of monticular
mesopores are somewhat thicker than those of surrounding mesopores.
Monticular mesopores generally contain one to several more dia-
phragms than intermonticular mesopores.
Longitudinal View: Endozone.—In the endozone (immature or
axial region), zooecial walls are longitudinally laminated and do not
show the dark granularity that is common in the Trepostomata. The
zooecial walls range from undeviating to irregularly undulating. Ina
few specimens the endozone is interrupted by a zone arching distally
across the branch that is marked by variable thickening of the zooecial
walls. Normal thin-walled zooecial growth generally continues distally
from the thickened walls of the arched zones with some bifurcating
but without other break. At apparently random levels within a colony,
some or all of the zooecia within the endozone have been eroded and
the tubes filled with mud. Subsequent growth was initiated from
adjacent zooecia and the eroded areas were covered by a basal lami-
nation of the overgrowth that continued the colony distally.
Exozone: Inner region of mesopores—The boundary between the
endozone and exozone is defined by the points of origin of the
mesopores. The inner region of the exozone extends distally for one
to several mesopore diaphragms, but generally not more than four.
The mesopores begin proximally with walls and diaphragms that are
slightly thicker than the zooecial walls of the endozone. Mesopore
walls curve broadly through 90 degrees into transverse positions
relative to the length of the mesopores, thereby forming diaphragms.
The broad curving results in constrictions of the mesopores at the
positions of the diaphragms to form cystlike chambers. In this inner
region of the exozone, mesopore walls commonly are longitudinally
laminated, but many, especially the thicker ones, develop transversely
laminated structure, either intermittently or throughout their length.
In the inner region, mesopore diaphragms regularly display centrally
No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN 9
located single pores that penetrate the diaphragms at right angles.
In longitudinal thin sections that pass through these pores, diaphragms
display transversely curved laminae that continue uninterrupted to
the pores. The curved laminae immediately adjacent to the pores
define the rounded boundaries of the pores.
If walls of adjacent mesopore chambers are longitudinally lami-
nated, generally the wall of the earlier chamber is connected directly
with the curved laminae on the proximal side of the intervening dia-
phragm and the wall of the later chamber is connected with the
distal side of the diaphragm. If walls of adjacent mesopore chambers
are formed by transversely curved laminae, the diaphragm and ad-
jacent walls will appear to be a continuous unit, or the diaphragm is a
direct continuation of the proximal wall and the wall of the distal
chamber is discordantly joined to the distal side of the diaphragm.
Rare, isolated mesopore diaphragms display complete continuity with
the walls of distal chambers.
In longitudinal thin sections, mesopore diaphragms in which the
pores were not intersected appear longitudinally laminated. Com-
monly the diaphragms are compound; the proximal half of a dia-
phragm is continuous with the wall of the preceding mesopore
chamber, the distal half is continuous with the wall of the succeeding
chamber. Other variations in diaphragm-wall relationships are less
common ; the two parts of the compound diaphragm can be unequal
in thickness, or in extreme development a diaphragm loses its com-
pound appearance and is wholly continuous with the preceding or very
rarely the succeeding chamber walls throughout or at either end.
Outer region of mesopores.—In the outer region of the exozone,
mesopores are not beaded and the walls and diaphragms display ex-
treme thickening. This greatly thickened skeletal growth can begin
on the distal side of the last thin diaphragm, the laminae covering the
central pore of the thinner diaphragm and curving distally into the
mesopore walls, or it can begin by an abrupt thickening of the meso-
pore walls. Diaphragms in this outer region are extremely variable
in thickness and spacing. A single diaphragm, greater in thickness
than the diameter of the enclosing mesopore, can correspond in thick-
ness and position with a series of irregularly and closely spaced dia-
phragms in adjacent mesopores. Most diaphragms are planar, but a
few are strongly curved and join adjacent diaphragms before reaching
the mesopore wall. The last diaphragms that were formed are in the
distal ends of the mesopores so that in external view the walls and
diaphragms of the mesopores combine to form the very shallow polyg-
onal depressions between the zooecia.
Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Many of the thick diaphragms also display centrally located pores
that do not penetrate through to the distal sides of most of the thickest
diaphragms. Laminae of the diaphragms generally stop abruptly at
the pores without changing direction or flexing, so that the pores have
no lining or apparent influence on the structure of the diaphragms. In
other thick diaphragms the laminae trend in a proximal direction in
varying amounts and there is a noticeable decrease in diaphragm thick-
ness approaching the pore. The pores in the outer region also differ
from the central pores of the inner region of the exozone by being
consistently smaller in diameter. In addition to the pores, mesopore
diaphragms and walls in the outer region contain small, dark, sub-
spherical to elongated cavities formed by the concentric arrangement
of laminae about imaginary centers. These cavities seem to be ar-
ranged at random in the walls and diaphragms.
Zooecia.—In the outer region of the exozone, undistorted wall
structure of adjacent zooecia is rarely seen because of intervening
mesopores and acanthopores. Zooecial boundaries are well defined,
dark, slightly serrated lines or zones in two dimensions, formed by
the abutting ends of laminae from adjacent zooecia. In walls formed
by a zooecium and adjacent mesopore, or by adjacent mesopores,
boundaries are more coarsely serrated and are commonly discontinuous
along their length.
Diaphragms are not present in most zooecia and not more than
two were seen in any one zooecium. If present, diaphragms are very
thin, planar to slightly curved, and extend distally into the zooecial
wall. Single, hollow, subspherical cystlike structures occur in the
zooecial voids of a very few zooecia, more commonly in the monticules.
The cyst walls are thick and are constructed of laminae that merge
with the laminae of the zooecial walls. Irregular spinelike proc-
esses are common in the zooecial walls in the thick-walled outer
region. These mural spines have their origins at or very near the
zooecial boundaries and trend in general toward the zooecial voids at a
high angle to undisturbed laminae in the walls. Zooecial wall laminae
surrounding the spines are flexed about the spines in a series of ir-
regular superposed cones and some of the laminae are pierced. The
spines extend far enough to cause inflection of the walls but none were
observed to break through the wall laminae and stand in relief in the
zooecial voids. The cores of the spines appear structureless or hollow.
Tangential View.—In tangential sections passing through the
outer region of the exozone, zooecia range from irregularly elliptical
to subcircular to petaloid in cross section. Major axes of the ellipses
are approximately parallel to branch length. The rare petaloid ap-
No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN Il
pearance is caused by extreme inflection of zooecial walls by adjacent
acanthopores and mural spines. Acanthopores are large, laminated,
possess well-defined central tubes, and are generally located at points
of closest proximity of adjacent zooecia. Mural spines appear to begin
outside the broad band of striated-appearing tissue lining each
zooecium and project inwardly toward the zooecial void, strongly in-
flecting the laminated tissue but not breaking through to the void.
Mesopores are numerous, subpolygonal to subcircular. In very shallow
sections that pass through the outermost and thickest diaphragms,
mesopore boundaries are concealed and interspaces between zooecia
appear solid. Many of these solid interspaces do not show the smaller
central pores that are the rule in sections that pass through earlier
parts of the outer region.
In deeper tangential sections that pass through the inner region of
the exozone, zooecia are polygonal to subpolygonal and approximately
equidimensional. Mesopores are also polygonal to subpolygonal and
have fewer sides than the zooecia, merely appearing to fill the spaces
between zooecia. Pores in mesopore diaphragms here are several
times larger in diameter than those in the outer region. Acanthopores
are considerably smaller in diameter than they are in the outer region.
QUANTITATIVE DATA
The following tables are based on sections of two fragments from
the lectotype, three fragments from the two paratype zoaria, and 49
fragments from 33 topotype zoaria. Sections from 55 zoaria of
Trematopora tuberculosa were examined. All measurements are in
millimeters. The axial ratio is the ratio of the diameter of the endo-
zone to the corresponding branch diameter.
TABLE 1.—General measurements
Lectotype AMNH Paratypes and
1747 topotypes
Frag. A. Frag. B. Minimum Maximum
Diameter of zoaritim: food. 62 weeks 6.5 4.9 3.0 7.2
Width of endozone............00200. 5.3 3.3 28 5.6
No. zooecia in 2 mm. (longitudinal
GIGECHOR)) hee sialon cue te amet a 53 63 6 8
Average major axis of zooecial void
Per tragment, jcciwcsvoac soca casteleleas 0.14 0.15 0.14 0.22
Average minor axis of zooecial void
Deri tragmenty oasis teihsbo steed eres 0.12 0.12 0.09 0.14
Acanthopores per zooecium .......... 0.73 0.63 0.50 0.5
Mesopores per zooecium............. 1.6 — El 2.0
I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
TABLE 2.—Ontogeny
Average No.
diaphragms Width of
in mesopores exozone Axial ratio
2 0.3-0.6 0.87-0.92
3 0.5-1.0 0.74-0.90
Braswe Acs lectotyperdacsciaicieacice 4 1.2 0.82
4 0.9-1.4 0.71-0.86
5 1.0-1.4 0.66-0.82
6 1.4-1.8 0.75
Fira cmmbcmlectOtyperercieeiciciaele sete a 1.6 0.67
2 I.1-1.6 0.68-0.70
DISCUSSION
The number of mesopore diaphragms and the width of the exozone
are not particularly sensitive indicators for ontogenetic development of
the mesopores and zooecia in T. tuberculosa. The variation in dia-
phragm counts and in width of exozone within a longitudinal thin sec-
tion is unusually large because of a marked variation in the number
of chambers developed in adjacent mesopores in the inner region.
Also, the unusual variation in thickness and spacing of mesopore dia-
phragms in the outer region of the exozone makes diaphragm counts
less reliable.
T. tuberculosa differs from both T. halli and T. whitfieldi in having
the larger branches, tuberculated mesopores, and a broader exozone in
mature specimens. Both T. halli and T. whitfieldi are smooth,
rhomboporoid-sized species.
REFERENCES
Basster, R. S.
1906. The bryozoan fauna of the Rochester shale. U. S. Geol. Surv. Bull.
No. 292, 65 pp., 31 pls.
1911. The early Paleozoic Bryozoa of the Baltic Provinces. U. S. Nat.
Mus. Bull. 77, 348 pp., 13 pls.
BoarpMAN, R. S.
Trepostomatous Bryozoa of the Hamilton group of New York State.
U. S. Geol. Surv. Prof. Pap. 340, in press.
Cumincs, E. R., and GAtLoway, J. J.
1915. Studies of the morphology and histology of the Trepostomata or
monticuliporoids. Bull. Geol. Soc. Amer., vol. 26, pp. 349-374, pls.
10-15.
HALL, JAMES.
1851. Jn Silliman, B., Silliman, B., Jr., and Dana, J. D., eds. New genera of
fossil corals from the report by James Hall on the Paleontology
of New York. Amer. Journ. Sci. and Arts, ser. 2, vol. II, pp.
398-401.
No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN 13
1852. Paleontology of New York. Vol. 2, 362 pp., 101 pls.
HA.L, JAMEs, and Simpson, G. B.
1887. Corals and Bryozoa. Paleontology of New York, vol. 6, 292 pp., 66 pls.
Miter, S. A.
1889. North American geology and paleontology. 664 PP., 1,194 figs. in
text. Cincinnati.
NicuHotson, H. A.
1881. The genus Monticulipora and its sub-genera. 235 pp., 6 pls.
UricH, E. O.
1882. American Paleozoic Bryozoa. Journ. Cincinnati Soc. Nat. Hist.,
vol. 5, pp. 121-175, 232-257.
1883. American Paleozoic Bryozoa. Journ. Cincinnati Soc. Nat. Hist., vol.
6, PP. 245-270.
1893. On Lower Silurian Bryozoa of Minnesota. Geol. Minnesota, vol. 3
pt. 1, chap. IV, pp. 96-332, pls. 1-28.
I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
EXPLANATION OF PLATES
IPPATE T
Figs. 1-4. Trematopora tuberculosa Hall.
1. Longitudinal view of paratype, A.M.N.H. 1747, <5, showing primary
branch with growth direction upward and branch from secondary
overgrowth with growth direction to lower right.
2a. External view of lectotype zoarium, A.M.N.H. 1747, X 2, showing tuber-
culated monticules.
zb. Longitudinal view of lectotype, < 20, showing beaded mesopore cham-
bers in inner region of exozone.
2c. Tangential view of lectotype, 20, showing aspect of both inner and
outer regions of exozone. Note thin-walled polygonal tubes of inner
region of monticule in upper left.
3. Tangential view of paratype, A.M.N.H. 1747, X 50, showing the smaller
central pores in mesopore diaphragms of outer region of exozone.
4. Longitudinal view of topotype, U.S.N.M. 137847, X 5, showing zooecial
growth at surface of contact of anastomosing branches. U.S.N.M.
collection 2998.
PLATE 2
Figs. 1-3. Trematopora tuberculosa Hall.
1. Longitudinal view of topotype, U.S.N.M. 137848, * 100, showing lam-
inated structure of a beaded mesopore and two diaphragm pores in
the inner region of the exozone.
2a. Longitudinal view of topotype, U.S.N.M. 137849, X 100, showing con-
figuration of laminae of mesopores and small diaphragm pore in outer
Tegion of exozone.
2b. Longitudinal view of same specimen, X 100, displaying a mesopore with
diaphragm pore of inner region covered by first diaphragm laminae of
outer region. Note discontinuous and ragged boundary between
mesopore wall and zooecial wall above.
3a. Longitudinal view of topotype, U.S.N.M. 137850, X 100, showing first
a diaphragm pore and then a compound diaphragm between beaded
chambers in the inner region of the mesopore.
3b. Longitudinal view from same zoarium, X 100, illustrating the structure
of the wall of adjacent zooecia.
3c. Tangential view from same zoarium, X 100, showing the general aspect
of the outer region of the exozone, including acanthopores, mural
spines, and a small pore in the center of a diaphragm of a mesopore.
The dark intermediately sized spots are the randomly arranged cavi-
ties noted in species description.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 6, PL.
TREMATOPORA TUBERCULOSA
(See explanation of plates at end of text.)
IITHSONIAN MISCELLANEOUS COLLECTIONS ‘ VOES 1397 NOn Gy REZ
TREMATOPORA TUBERCULOSA
(See explanation of plates at end of text.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NO. 7
Charles DB. and fMlary Waux CHalcott
Research Fund
BARLEY (TERTIARY. APHELISCUS AND
PHENACODAPTES' AS PANTOLESTID
INSECTIVORES
(WitH Two PLatTEs)
By
C. LEWIS GAZIN
Curator, Division of Vertebrate Paleontology
United States National Museum
Smithsonian Institution
eee Ov e ea yr ty \
4 is { 1 ft L ad te! ¢
(Pustication 4385) \\ /
\LBRAaBy
eo
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
AUGUST 12, 1959
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NO. 7
Charles D. and Mary Waux Talcott
Research Fund
EARLY TERTIARY APHELISCUS AND
PHENACODAPTES AS PANTOLESTID
INSECTIVORES
(WitH Two PLatTEs)
By
C. LEWIS GAZIN
Curator, Division of Vertebrate Paleontology
United States National Museum
Smithsonian Institution
(PusiicaTion 4385)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
AUGUST 12, 1959
ITHSONIAN ana i '9 4
STITUTION Aug 1'2 1959
THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.
Charles D. and Mary Waux Walcott Research Fund
BARELY TAlERELARY, APE LISOCUS AND
RHENACODAP LES cAS PANTOLESTEID
INSECTIVORES
By C. LEWIS GAZIN
Curator, Division of Vertebrate Paleontology
Umted States National Museum
Smithsonian Institution
(WitH Two PtatEs)
INTRODUCTION
Examination in 1954 of Phenacodaptes material in the Paleocene
collections at Princeton University, believed pertinent to a review of
Eocene artiodactyls then underway, led to rather inconclusive results.
Dr. Jepsen’s tentative suggestion (1930, p. 519) of such a relationship
may, nevertheless, have merit.1 More recent studies of the Knight
faunas, however, involved certain pantolestids, and comparison of
these among a wide range of both Eocene and Paleocene forms has
convinced me that Cope’s Apheliscus and Jepsen’s Phenacodaptes are
closely related and that both are pantolestids, although perhaps some-
what less closely related to the Pantolestinae than to the Pentaco-
dontinae. Their relationships would seem possibly best illustrated by
placing them both in the Apheliscinae as a subfamily of the Panto-
lestidae.
I am indebted to Dr. Glenn L. Jepsen for permitting me to borrow
and illustrate specimens of Phenacodaptes sabulosus in the Princeton
collections, and to Dr. George G. Simpson and Mrs. Rachel H. Nichols
for sending me materials of Apheliscus imsidiosus and Pentacodon
inversus from the collections of the American Museum. The pencil
drawings of specimens shown in the accompanying plates were made
by Lawrence B. Isham, scientific illustrator for the Department of
Geology in the U. S. National Museum.
PREVIOUS INTERPRETATIONS OF RELATIONSHIP
Apheliscus insidiosus was described by Cope (1874, p. 14) from
the lower Eocene San Jose beds in New Mexico. He described it
1 After this manuscript was submitted for publication, Dr. Jepsen showed me
a note that he had placed in the collection drawer some time ago suggesting that
Phenacodaptes be compared more carefully with Apheliscus.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 7
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
first as a species of Prototomus and included it together with “Pro-
totomus” jarrovii (=Pelycodus jarrovii) in the carnivores with
Prototomus (=Sinopa) viverrinus. In 1875 (p. 16), however, he
proposed the name A pheliscus, regarding it as “nearly allied to Panto-
lestes,’ although at the same time he thought that the molar teeth
suggested a relationship to Anaptomorphus, noting, nevertheless, that
the premolars were “totally different.’ Cope’s statement that the last
lower molar lacked a heel would seem highly significant, but, if the
meaning is here properly interpreted, it is surely an error, as may be
seen from his illustration (1877, pl. 45, fig. 18). In addition to the
described condition of the talonid of the third molar, Cope noted as
distinctive in comparison with Pantolestes only the simplicity of the
inner anterior tubercle of the lower molars.
Matthew (1918), in naming the family Apheliscidae, was very dubi-
ous as to its affinities, and while referring it to the Insectivora, con-
sidered that it might well be condylarthran, primate, or creodont. It
should be noted, however, that at the time of his writing, such genera
as Aphronorus, Bessoecetor, and Phenacodaptes were not known.
Only large and comparatively aberrant Pentacodon, which he had
recognized as a pantolestid insectivore (1909), and the Eocene mem-
bers of the Pantolestinae were available for comparison.
Simpson (1937a) demonstrated the most logical arrangement for
the pantolestids and pentacodonts, while adding the Paleocene genera
Bessoecetor and Aphronorus to their respective subfamilies. He
noted, moreover, the resemblance of Apheliscus to the Pentacodonti-
nae in characters of the fourth premolars, but considered, however,
that the molar structure was widely different. Nevertheless, his sug-
gestion that Apheliscus might be an offshoot of the same stock seems
particularly pertinent and certain of the lacking evidence for such an
hypothesis may lie in Phenacodaptes. The family, however, was re-
tained incertae sedis, questionably in the Insectivora in his 1945
classification.
Saban (1954), evidently following Simpson’s suggestion, included
the Apheliscidae with the Pantolestidae in the superfamily Pantoles-
toidea. His including Shikama’s Endotheriidae, created for the Man-
churian Jurassic Endotherium, as a subfamily of the Pantolestidae,
however, seems surprising. McDowell (1958) rejected certain fea-
tures of Saban’s classification and in discussing the family Aphelisci-
dae regarded it as incertae sedis, but noted that the teeth are “recon-
cilable with those of Mirodectes.’ McKenna, on the other hand, in a
field conference guidebook (1955) has the Clark Fork species Aphelis-
cus nitidus listed as a condylarth.
Older but more recently described Phenacodaptes sabulosus is from
NO. 7 TERTIARY APHELISCUS AND PHENACODAPTES—GAZIN 3
the Silver Coulee or Tiffanian horizon of the Polecat Bench forma-
tion in the Big Horn Basin. The possibility of a relationship to artio-
dactyls was tentatively suggested by Jepsen (1930) because of resem-
blances noted to such genera as Diacodexis and Bunophorus, evident
in certain features of the molars. Simpson, however, in his classifica-
tion of the mammals (1945) cited Phenacodaptes as a condylarth
under ?Mioclaeninae incertae sedis.
COMPARISON OF APHELISCUS AND PHENACODAPTES
A lower jaw of Apheliscus, referred to A. insidiosus, in the Na-
tional Museum collection (No. 19162) from the Gray Bull beds in the
Big Horn Basin, exhibiting P.-M, inclusive (see pl. 1, fig. 1), shows
that the form and relative proportions of the lower premolars are
strikingly like those in Phenacodaptes (see pl. 1, figs. 3, 4). The rela-
tively small size of P, and particularly of P; in comparison with P,
is quite alike in the two. P, is a little more slender in Apheliscus and
the distinctive talonid seen in this tooth of Phenacodaptes is more
sectorial and essentially better developed or exaggerated in Apheliscus.
Both have a strongly developed primary cusp and only slight evidence
of a paraconid. There is no metaconid on P, in the known material of
Apheliscus. It is usually absent, but may be weakly developed in some
specimens of Phenacodaptes. The lower molars differ noticeably in
the anteroposteriorly shorter trigonid and more elongate talonid
in Apheliscus (see pl. I, fig. 2) ; moreover, they are relatively more
slender than in Phenacodaptes. There is, nevertheless, a rather
marked similarity in many details, particularly in form of the cusps
and crest surrounding the talonid basin, and in the shape of the basin.
The compressed trigonid of Apheliscus is rather less like that in
Phenacodaptes, although the paraconid is absent or very much re-
duced on the posterior two molars of both forms. In M, of Phena-
codaptes, however, this cusp is moderately well defined as an anterior
crest from the protoconid, whereas in Apheliscus only a slight median
cuspule remains.
The upper cheek teeth of Apheliscus insidiosus (see pl. 2, fig. 1)
may appear a little less like those of Phenacodaptes (see pl. 2, fig. 2)
than perhaps do the lower teeth, although both exhibit the compara-
tively small and subequal second and third upper premolars and en-
larged fourth. The more noticeable differences between the two in
upper teeth include less development of the cingulum, particularly
on P*, and the transversely narrower molars of Apheliscus. More-
over, the hypocone, though distinctive on M, and M, of Phenaco-
daptes, is weak or absent in Gray Bull Apheliscus. It is important to
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
note, however, that the upper teeth seen in Clark Fork Apheliscus
nitidus seem intermediate in most, if not all, of these respects. A com-
parison of Matthew’s figure (1918, fig. 24) for the Clark Fork speci-
men, which Simpson (1937b) made the type of A. nitidus, with P*
and M?* in Phenacodaptes sabulosus, here shown in plate 2, figure 2,
reveals little to distinguish them. The Sand Coulee lower teeth of
Apheliscus, figured by Matthew (1918, fig. 24) also seem to show
a little less compression of the trigonid than more typical Gray Bull
specimens.
The foregoing comparisons strongly suggest that Phenacodaptes,
or at least a very closely related form, gave rise to Apheliscus. The
succession may well have been Phenacodaptes sabulosus—A pheliscus
nitidus—A pheliscus insidiosus. In the course of this postulated de-
velopment it would seem that the principal tendency was toward the
transverse narrowing of the teeth, both upper and lower series ; the
loss or weakening of the cingulum in the upper series; the increas-
ingly Pentacodon-like development of P*; the relative increase in
length of talonid of the lower teeth, Ps as well as the molars ; together
with the shortening of the lower molar trigonids.
RELATIONSHIPS OF APHELISCUS AND PHENACODAPTES
The most nearly comparable development to that illustrated in the
Phenacodaptes—A pheliscus line would seem to be among the panto-
lestids. The suggested comparison is perhaps not so close to the
Bessoecetor—Propalaeosinopa—Palaeosinopa—Pantolestes succession as
it is to the middle Paleocene Pentacodontinae. The premolar develop-
ment would seem rather like that in both Aphronorus (see pl. 2, figs. 3
and 4) and Pentacodon (see pl. 2, figs. 5 and 6), except that there
tends to be no metaconid on P, or tritocone (uncertain for Pentaco-
don) on P* in the apheliscids. Aphronorus, moreover, differs in having
somewhat higher crowned, more definitely insectivore teeth. The
upper molars of Aphronorus show better developed and more laterally
directed anteroexternal and posteroexternal styles and the lower molar
trigonids are a little higher and show better development of the para-
conid.
Much larger Pentacodon has a more enlarged fourth premolar, but
the upper molars (not previously illustrated) do not show the distinc-
tive outer styles seen in Aphronorus. Also the trigonids of the lower
molars do not appear to be so elevated, but, like Aphronorus and unlike
the apheliscids, show a prominent and forward-placed paraconid. The
talonid construction, nevertheless, is much alike in the two subfamilies,
except for relative length.
NO. 7 TERTIARY APHELISCUS AND PHENACODAPTES—GAZIN 5
The mental foramen, the position of which, as Simpson (1937a,
p- 120) notes, has been unduly emphasized, may warrant comment.
It exhibits a comparatively small posterior opening somewhat farther
forward in the Apheliscinae than in Pentacodontinae or Pantolestinae.
It is variable in Phenacodaptes and is observed in positions beneath
the anterior part of P, to the posterior part of P;. A larger opening
is noted beneath P, or Pz. In a specimen of Apheliscus (U.S.N.M.
No. 19162), these foramina were noted beneath posterior portion of
P; and beneath P,. In Aphronorus the posterior foramen may be
small and varies in position from beneath M, to the posterior part of
P,. An equally large or larger anterior opening is seen below P.. In
Bessoecetor foramina were noted beneath the posterior part of both
M, and Pz, and in one specimen, U.S.N.M. No. 9442, anterior fora-
mina were observed below the posterior portions of both P, and Ps.
In Bridger Pantolestes I have seen only the well-developed foramen
beneath M,.
Among the Insectivores outside of the Pantolestidae I find a rather
more remote relationship to the mixodectids indicated. There would
appear to be rather less to suggest affinities with other orders. Among
these, however, perhaps the condylarths should be considered. The
relatively low trigonids of the lower molars seem indicative of a pos-
sible condylarthran relationship, and a form such as Choeroclaenus
among the mioclaenine hyopsodonts is not too different from Phenaco-
daptes but there is, nevertheless, a more inflated appearance to the
molar cusps and the premolars would appear to have little or nothing
to recommend them. The possibility that the Phenacodaptes—A phelis-
cus line represents condylarth development rather paralleling that of
pentacodonts cannot be entirely disregarded, but the same reasoning
might apply equally well were they to be regarded as belonging to such
other orders as primates, creodonts, or artiodactyls. Comparison with
Pentacodon and Aphronorus appears rather more pertinent and better
accounts for a number of minor details of similarity not easily
dismissed.
REFERENCES
Corr, Epwarp D.
1874. Report upon the vertebrate fossils discovered in New Mexico with
descriptions of new species. Geogr. Expl. and Surv. West of 1ooth
Meridian (Wheeler), Appendix FF, Ann. Rep. Chief of Engineers,
1874, pp. 1-18.
1875. Systematic catalogue of Vertebrata of the Eocene of New Mexico,
collected in 1874. Geogr. Expl. and Surv. West of tooth Meridian
(Wheeler), pp. I-37.
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
1877. Report upon the extinct Vertebrata obtained in New Mexico by parties
of the expedition of 1874. Rep. U. S. Geogr. Surv. West of 1ooth
Meridian (Wheeler), vol. 4, pt. 2, pp. i-iv, 1-370, pls. 22-83.
JEPSEN, GLENN L.
1930. Stratigraphy and paleontology of the Paleocene of northeastern Park
County, Wyoming. Proc. Amer. Philos. Soc., vol. 69, No. 7,
pp. 463-528, figs. 1-4, pls. I-10.
MatTrHew, WILLIAM D.
1909. The Carnivora and Insectivora of the Bridger Basin, Middle Eocene.
Mem. Amer. Mus. Nat. Hist., vol. 9, pt. 6, pp. 201-567, figs. 1-118,
pls. 42-52.
1918. A revision of the lower Eocene Wasatch and Wind River faunas.
Part IV. Entelonychia, Primates, Insectivora (part). Bull. Amer.
Mus. Nat. Hist., vol. 34, pp. 429-483, figs. 1-52, pls. 42-52.
McDoweELt, SAMUEL B., Jr.
1958. The Greater Antillean insectivores. Bull. Amer. Mus. Nat. Hist.,
vol. 115, art. 3, pp. 115-214, figs. 1-46, tables 1-2.
McKenna, Matcoim C.
1955. Age of the Four Mile local fauna, northeast Sand Wash Basin, Colo-
rado. Wyoming Geol. Assoc. Guidebook, roth Ann. Field Confer-
ence, Green River Basin, pp. 105-107, fig. 1.
SABAN, ROGER.
1954. Phylogénie des Insectivores. Bull. Mus. Nat. Hist. Nat., ser. 2,
vol. 26, No. 3, pp. 419-432.
Simpson, GEORGE G.
1937a. The Fort Union of the Crazy Mountain field, Montana, and its
mammalian faunas. U. S. Nat. Mus. Bull. 169, pp. 1-287, figs. 1-80,
pls. I-10.
1937b. Notes on the Clark Fork, upper Paleocene, fauna. Amer. Mus. Nov.,
No. 954, pp. 1-24, figs. 1-6.
1945. The principles of classification and the classification of mammals. Bull.
Amer. Mus. Nat. Hist., vol. 85, pp. i-xvi, 1-350.
NO. 7 TERTIARY APHELISCUS AND PHENACODAPTES—GAZIN 7
EXPLANATION OF PLATES
PLATE I
Apheliscus and Phenacodaptes from the early Tertiary of Wyoming
Figs. 1, 2. Apheliscus insidiosus Cope: 1, Right ramus of mandible (U.S.N.M.
No. 19162), lateral and occlusal views. 2, Left ramus of mandible (A.M.
No. 15696), lateral and occlusal views. All four times natural size. Gray Bull
lower Eocene, Big Horn Basin, Wyoming.
Figs. 3, 4. Phenacodaptes sabulosus Jepsen: 3, Right ramus of mandible (P.U.
No. 13926), lateral and occlusal views. 4, Left ramus of mandible (P.U.
No. 13391), lateral and occlusal views. All four times natural size. Silver
Coulee (Tiffanian) upper Paleocene, Big Horn Basin, Wyoming.
PLATE 2
Apheliscinae and Pentacodontinae from the early Tertiary of the
Rocky Mountain Region
Fig. 1. Apheliscus insidiosus Cope: Left upper cheek teeth (A.M. No. 15696),
occlusal view. Four times natural size. Gray Bull lower Eocene, Big Horn
Basin, Wyoming.
Fig. 2. Phenacodaptes sabulosus Jepsen: Right upper cheek teeth (P.U.
No. 13977), occlusal view. Four times natural size. Silver Coulee (Tiffanian)
upper Paleocene, Big Horn Basin, Wyoming.
Figs. 3, 4. Aphronorus fraudator Simpson: 3, Right upper cheek teeth
(U.S.N.M. No. 9561, Ps from U.S.N.M. No. 9564), occlusal view. 4, Left
ramus of mandible (U.S.N.M. No. 6177, type specimen, with molars restored
from U.S.N.M. No. 9289, Pi: to Ps from U.S.N.M. Nos. 9537 and 9201), lateral
and occlusal views. All four times natural size. Fort Union middle Paleocene,
Crazy Mountain area, Montana.
Figs. 5, 6. Pentacodon inversus (Cope): 5, Left upper cheek teeth (U.S.N.M.
No. 15502), occlusal view. 6, Left ramus of mandible (A.M. No. 17038),
lateral and occlusal views. All twice natural size. Torrejon middle Paleocene,
San Juan Basin, New Mexico.
hy eee
we five.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE 139; NO: 7, IP, 2
APHELISCUS AND PHENACODAPTES FROM THE EARLY
TERTIARY OF WYOMING
(See explanation of plates at end of text.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL 1397) NOS 7) Pez
APHELISCINAE AND PENTACODONTINAE FROM THE EARLY TERTIARY
OF THE ROCKY MOUNTAIN REGION
(See explanation of plates at end of text.)
et
J
Se ae ae ee Se ee
= gy NT Le Te .
oo &
fs a
Ree AIA Sy
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 8
THE ANATOMICAL LIFE
OF THE MOSQUITO
By
R. E. SNODGRASS
Research Associate
Smithsonian Institution
ase OOseng,
"0
f
i
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
NOVEMBER 4, 1959
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 8
THE ANATOMICAL LIFE
OF THE MOSQUITO
By
R. E. SNODGRASS
Research Associate
Smithsonian Institution
(Pustication 4388)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
NOVEMBER 4, 1959
THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.
CONTENTS
Page
Ug AOVaLE TSS OTD RIS EHO GO OGD SO DCO TIERCE Bier ha casio 6 bt a ETAT aoe I
Pee mila revere sets ainlafocrererecer a ere och ois nia eraye To ole ee SNA NERD eceteatoaat alate 2
Mhew headin yee eR yaAR ae rh atta eae o's ie eae eiao ete aeeie 4
The teed NSsOLRANS rytecrcte oie sistas. ctovere Gee iwik svejaictomrmoreiarsiere aateterere Il
Me MIAMI eos « Sig diee's isis g eire mien lava lo rotemrcteree ta ate psrerershotale 12
he preatal. Cavaty sec vi Noes cone Gels © a acatiomarcie sels maleye t's 15
The epipharyngeal apparatusss si: julc tocar cab eelee © eimcieie 15
AM hekmiandi bles Aves arate wisiersie: aeierel nee ei entterenmen evoke ome tater 16
ARS arias Te hai see eyetey bel otepoueere ceousuaro bie. 3 cfpanseoaraed wate rate ets 17
The labrum and hypopharynx< ws... cc ss cieai epee eas 19
SDE SP MATIV UR |, sisi crt eecicre tase ore ober tresoiers a) ays: eSNG autora iarete sie eles 21
Warvaltteeditigs toss ne oa ceteris tere dd abin teeter he enteric. 22
ADRS T CHOTA Ca cA Ae Se ey ere crea ete alevsicia wla-c suslale ERT Selstene ane tials 24
ible ANdaMieny «5 HIV Tha ete aka les, «ced SRM ere ete ee 26
Intennalganatoriy, Gfevtcs ceeeve feelin ie cies stoi s olaernkenstebas sie eiereiaiece 21
‘Whew itrachealmsystenns wasn e elo, oe oetee le elaevaats aoe helve ZT
‘phe dorsal bloodiaviessel yemrverecrers asicva eee eeere a cores 32
aPhewalimentarya Catiale were se tawe tale: « th ete ces eerclale ey arre erate 32
MheeMalpightanetupulesescaas te cmice shoo eae tees a2
AbheNsalivatymolatid serra sire cels ino Richtee talet ote erons eet 33
AREY MER VOUSH SY.SECIM csyersiey sie reis oreiks ho ohn ae ane aaieh s 34
PRE} FEPrOGUCtiVe OL GANS) «We iale.././«,« ssa isin, Sedscereme lene soins ohans 34
I OOG PESCE VES a tole satan Iai5 "Lo clay idee ousie s,s o/s ercem arlene ate tetnelalees ovate 34
WIE, CMTE APU Tees core c.ciakare ecole scree e aati d clave wee Oars Meme ate cicaaaaats 34
he mpupal developments... Lee sees cies a os stale oe ee lore eile aati 35
Aheematine ppupareyidc solace setae cs oko ec sltieteteis. oavaveteomro tts 39
GeneralVexternall stature ia acie.sie cia. oni teeta 39
The head and amiauth) Parts ch iciustss + os.0 Ghereake aiucinaen’s clueve 43
PINE MEH O tela eh se eerie caterer ete rar seni ehcee cceaic Ace eRT Re ere ete 44
PBheyabdomenly arene setae wiaciete cis Nace ce Nee eee ore etree 45
Ache pupal’ metamoOnpuOsiss,. sheaves. cess hon be eee en ae ee 46
Tela ul he via ttl tice evreratch Shae Hove Mahe tela 8 a ap aaa ie a Se ad 48
AEM HEA sive crcveoe Such eve ca ciey ciate eek l ale rol Gre soa eases oor ETTORE AITO SI
PENENOTSANSIOT ACCUM 21a vies us cies) a) ocho bis eu Aue ice SiG aleie c 55
PUG PU OMOSCIS ala 3 cle cis te anor a Leratienet sere. 270) Sachets tomamtene naire Ss cars 55
a exci Daria DUBIN Ne feces cake Saks es cis eters Mere or 60
hekpharyngeal: punipaces isosceles os ERE eee 63
MUNG PROP AS erate aid erarei te acd ute: RAN. 0 |, UNINC 5 le i, 64
sihrerabd omen srs easter asa aicrsiew io sols oleae RO CRIEC Rae icheee 67
internal anatomiye «sscrec sh as Lae a 63) 52.4.0 :0's geen inetd 72
PHS CALCINACOE We (ORE ANGY (20 eaves! o1sne' sce 3 0 sel ammeto nelle eniaks Mats 73
Dhevalimentany. (canal ours ciers crores iaye soo Ao eles ots hee 73
mher salivary) wands ivy sad Hos) SA ORR A oe 76
Dhe reprodtictiver-sy ster). tie: 4 scsi c'e sea en ae claialeieers.cie its 76
IRGLEL ER COSMET ieee SED crac a aired a tee Khaleel isle cheer Oe oh aieiticlats 80
i
iy
y,
Toe
den
'
WEY,
Fela y
[6 ee MR Pier ye ed eae
Wa ee Mi Ae PRA NR eo
e } ¥FY Shia ae A eAN fa .*
Na ee A; aM i COUN ME EWE S44
‘ i a i) H er, ay esiomd rete 1a
ha ‘ Cul) Aaa, 3° i ‘ea, oP ection na VMN 4
Mt a MI ye LR 2! Ot Oe any DORIS
1% erty) 7 Sree SMA Le Reo Me
Pm ea ey i
me ; ae eh Te ree whe ayia Y's
FR i , % ey ie hoe bb! MY f .
‘hy VE A ey Oy, xy de) yay
ma eee tail bike! ack dA AW Gt in
i ‘ a ih& : Reale Variehk fe ghee , Oy
bh : ig ; ht t : i ‘ 1 DF
rh. |, hake tobe ODS 6a 8 On LR ea we oe ee ey ob
cr j : Ale
; alk powekie take ad
Dre - Ns ip
¢ \
8 ee MCU REPMN NT ad
mic dy ey a) a 8 YORE As hy
nh 4 ! ee) pa Sik ae ve eayeUN a w
eiieramaea 9) eta
Best hss PR eg eee Mil W ed 9 So asta ¥
PE DIRS eM A wea ye ie bees a
to Reh iin eee Mae yt
Ribs eh ‘
Re OP Ree he Rae be
a j el y
ST AT ee Em eS
VOR Ts yy (ot LM
Eee
, i
6 ae Ea Pee ee ie bk
ee Wi are aM eae
Pi Saye bh pa MT ale ae
A R » ad pay Te
TS OR Ree a Te ERO ER te me rey
: ; } i
FeETER
Pa a ee a MO Ae Me ae con) | pe bidi, me
be ; ret a
ea We aed Pee PNRM ork ae WE
OU A t U teL 9 ee Ee
Pt Peet Wal wR Fon NO ag a hoy ae
Pe CL eh ere Tad eae Ree
ee ee
SORRY AA hen 1S ee Pel ys eee
: Vi MAES AAT ht
a8 DAVOS 7 F400 ON ele ORY
ets ah Ae ae OR NS 4 ee Sala eH oe
+ ' uf AS
7 aye ee ra ay 8
eek Phe a ew ie
eu eas te
ins 4! yikes
Hew RN made yo RatamN ey
(iF ay ava Nh Pak oaks ALN A ilk i)
: ee eS Ty k aly eas
i gg bhi oe
R2SRS2SSStR se
Tre ANATOMICAL’ PIPE OF THE MOSOUTITO
By R. E. SNODGRASS
Research Associate
Smithsonian Institution
INTRODUCTION
Mosquitoes are not popular with warm-blooded animals, but from
their own standpoint they have been highly successful insects, until
recently when they have been attacked with poison sprays and have
had their larval habitats drained. Success, however, is always to be
admired whether in man or an insect, and it is instructive to see how
it has been achieved. The mosquitoes have attained their place in the
world by the evolution of highly specialized anatomical characters. A
study of their anatomy may help some in our war against them, and
it will give a most interesting example of how insects have evolved
structures fitting them for particular ways of living and of feeding
that have made them so successful in the struggle for existence.
The family name of the mosquitoes is Culicidae, and they belong
to the order Diptera, or two-winged flies, which in turn are members
of that large group of insects in which the young, or larvae, are very
different from their parents in form, structure, and habits, and must
undergo a renewed growth to attain the adult state. We are so ac-
customed to seeing young animals grow up gradually into adults that
it seems very remarkable that an animal can completely change its
shape and structure in the middle of its life. The young mosquito, for
example, hatches from the egg as an active larva having no resem-
blance to its parents but fully adapted in its structure for living and
feeding in the water. During its life the larva sheds its cuticle four
times. At each of the first three ecdyses it comes out a little larger
than before, but with little change otherwise. On shedding the fourth
cuticle, however, a very different creature, the pupa, emerges. The
pupa has all the adult organs, though in an incomplete state of de-
velopment, and is clearly a preliminary adult. With a final moult and
ecdysis the completed mosquito appears, equipped for an entirely dif-
ferent life from that of the larva.
It is commonly said that the larva is metamorphosed into the adult
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 8
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
during the pupal stage. Actually it simply returns to its parental adult
structure after having undergone during its evolutionary history a
metamorphosis by which it took on a form and structure suited to a
way of living quite different from that of its parents. The embryo,
from its very beginning in the egg, develops into a larva. The egg,
therefore, contains two distinct hereditary factors, one that first
produces the larva, another that later generates the parent adult. When
the larva does not differ too much from its parents, the adult may be
formed mostly by a new growth of the larval tissues ; but as the differ-
ence becomes more extreme, the larval tissues go into a state of dis-
solution and the adult is built up of embryonic cells that multiply but
do not become organized during the larval stage. The transformation
of the mosquito is intermediate between these two conditions.
Inasmuch as the word metamorphosis means simply a “change of
form,” we may say that the larva in its aberrant evolution has under-
gone a divergent metamorphosis, and that as an individual it resumes
the parental form by a convergent metamorphosis.
Since the egg has the potentiality of developing into both the larval
and the adult form, there must be some influence that allows the larva
to develop first. The inhibition of adult development is effected by a
hormone, known as the juvenile, or status quo, hormone. When
the larva is mature and has served its purpose in the life of the insect,
this hormone ceases to be effective, and the adult development pro-
ceeds under the stimulus of another hormone. This at least is the
usual story of endocrinal regulation of insect growth and transforma-
tion, but, as will be seen, the mosquito does not comply fully with
the rules of hormone control in its growth from larva to adult.
Before going on with anatomical descriptions of the larval, pupal,
and adult stages of the mosquito, a few terms should be defined as
they will be used. An instar is the insect between any two consecutive
moults. Moulting is the physiological process of separating the old
cuticle from a new cuticle being formed by the epidermis beneath it.
The new instar begins its development when the moult is completed,
but remains inside the old cuticle until it is fully formed. Then it
breaks the cuticle and comes out. The emergence of the insect is its
ecdysis (coming out). Moulting and ecdysis, therefore, refer to two
different events, and are not synonymous terms, though many en-
tomologists have not distinguished them as such. In life-history
studies the “instar” is usually regarded as the insect between ecdyses,
but since development begins inside the old cuticle, an instar is really
the insect between moults. The concealed intracuticular period of the
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 3
instar has been appropriately named by Hinton (1946, 1958a) the
pharate, or cloaked, stage of development. The pharate stage of the
pupa in the larval skin is commonly called the “prepupal stage of the
larva,” but the larva has already ceased to be a larva, so the expres-
sion does not conform with the facts. The mosquito will demonstrate
a number of other errors commonly made by entomologists.
The problem of explaining how an animal in its evolution has be-
come adapted structurally to its environment and a special way of
living is complicated in an insect such as the mosquito that lives two
entirely different lives. If adaptation affects two or more organs
separately, the matter is relatively simple, but when it involves
coadaptation in all parts of the animal, it is hard to understand how
evolution by means of natural selection has brought it about. On the
other hand, the technique of “special creation” is entirely incompre-
hensible.
The writer began this work on a very meager acquaintance with the
anatomy of mosquitoes, especially of the larva and pupa. For its
completion he is deeply in debt to others, in particular to Dr. Alan
Stone and Dr. Richard H. Foote at the U. S. National Museum for
literature and the identification of species; to Dr. Paul Woke of the
National Institutes of Health at Bethesda, Md., for an abundance of
live larvae; to Dr. Ernestine B. Thurman, also of the Institutes of
Health, and Dr. Jack Colvard Jones of the University of Maryland
for much supplementary information and a critical reading of the
manuscript ; to various authors for copied drawings; and to Mrs. R. E.
Snodgrass for the typing. For morphological interpretations the
writer assumes entire responsibility.
T Loe CARVA
Mosquito larvae hardly need an introduction. They are the familiar
aquatic “wrigglers” or “wigglers” that everybody knows turn into
mosquitoes. Anatomically the most specialized parts of them are the
head, the feeding organs, and the respiratory system. A number of
good papers have been written on the larval anatomy, and the facts
of structure have been well-enough described, but the writers, par-
ticularly on the head and feeding organs, mostly disagree as to the
homologies of the parts, and consequently the different terminologies
used must be very confusing to students. Hence, in the following text,
the larval head and organs of feeding are given a disproportionate
amount of space in an effort to arrive at reasonable interpretations
and an acceptable terminology. Otherwise than in the head and feed-
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
ing apparatus, the principal specialization of the larva pertains to the
respiratory system. The only functional respiratory apertures are a
pair of dorsal spiracles near the end of the abdomen, the lateral
spiracles being closed except at ecdysis when the tracheal linings are
partly pulled out through them.
THE HEAD
The head of a mosquito larva projects forward from the thorax in
line with the axis of the body, bringing the mouth parts to an anterior
position. In most adult insects the head hangs downward on the
thorax, so that the face is anterior and the mouth parts ventral. In
the prognathous mosquito larva the face becomes dorsal and the mouth
parts anterior. In going from adult to larva, therefore, instead of
reversing the meaning of “dorsal” and “ventral,” it will be better to
speak of the upper and lower surfaces of the larval head, though
“anterior” and “posterior” in either larva or adult will be directions
relative to the axis of the body.
The typical shape of the mosquito larval head is oval or ovate,
whether seen from above (fig. 1 A,B,C) or from the side (E), but
the upper surface is more rounded than the lower. In some species,
however, the head is almost rectangular in form (D). Anteriorly
the head bears laterally a pair of large mustachelike brushes, and
usually between them a small median brush, the three being supported
by the labrum. Shortly behind the lateral brushes arise the slender,
tapering, unsegmented antennae (E, Ant). Posteriorly on each side
of the head is a large dark spot (£) varying in size with the age of
the larva. These spots are the pigmented compound eyes of the adult
developing in the epidermis beneath the larval cuticle. Behind or
below each compound eye is a small, simple, presumably functional
larval eye (O). The lateral area of the head between the antenna and
the eye is the gena (Ge), that behind and below the eye the postgena
(Pge). Posteriorly the head abruptly narrows to the occipital fora-
men, which is rimmed by a darkly sclerotized band, the postocciput,
set off by a postoccipital sulcus. The membranous neck is usually
cylindrical (fig. 1 A), but in Anopheles (fig. 3 C) it is narrowed where
it joins the thorax, evidently to facilitate the turning of the head upside
down while feeding.
The upper surface of the head (fig. 1) is differentiated into a large,
shieldlike central area, narrow lateral areas bearing the antennae and
the eyes, and a slender transverse anterior sclerite at the bases of the
brushes. This sclerite (A,B, Lm) is the dorsal wall of the labrum,
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 5
as contended by Cook (1944a), though some writers have regarded
it as a “preclypeus.’”’ The groove behind it (A, cls) then is the
clypeolabral sulcus. The large central area of the head is bounded by
lateral lines (CL) that diverge forward from a very short occipital
cleft and become continuous with the clypeolabral sulcus. These
lines, commonly called “frontal sutures,” are merely lines of weakness
Fic. 1.—The larval head, dorsal and lateral.
A, Aedes aegypti. B, Culex sp. C, Anopheles quadrimaculatus. D, Toxo-
rhynchites rutilus. E, Culex sp., lateral. F, Anopheles farauti, head exuviae.
Ant, antenna; Apt, cephalic apotome; CL, cleavage line; cls, clypeolabral
sulcus; E, compound eye; Ge, gena; Hstm, hypostomium; Lm, labrum; Md,
mandible; Mx, maxilla; O, group of larval simple eyes; Pge, postgena.
in the cuticle where the latter will split at ecdysis (F, CL) to allow
the emergence of the next instar, and are best termed the cephalic
cleavage lines. In most young insects the cleavage lines take the form
of a Y, which has been known as the “epicranial suture.” In the
mosquito larva the stem of the Y is the short occipital cleft. The
frontal arms follow such very different courses in different insects
that they can have no morphological significance (see DuPorte, 1946;
Snodgrass, 1947), and hence do not define any specific part of the
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
head. The part of the head wall cut out at ecdysis may be termed the
cephalic apotome (F, Apt).
The space between the arms of the cleavage lines and the
clypeolabral sulcus in the mosquito larva has been variously called the
“frons,” the “clypeus,” and the “frontoclypeus.” The respective areas
of the frons and the clypeus may be identified in other insects by
specific groups of muscles that arise on them. In the mosquito larva
the clypeal muscles arise anteriorly, the frontal muscles posteriorly on
the central head area, but there is no external demarcation between
the two regions. This area, therefore, is frontoclypeal in a limited
sense, but it is not the entire clypeus or the entire frons. Ordinarily
the clypeus extends laterally to the bases of the mandible, and in adult
insects the frons is the facial area between the antennae and the eyes.
The whole aspect of the mosquito head is changed at the transfor-
mation to the pupa and the adult.
The larval antennae are slender, unsegmented shafts bearing vari-
ously distributed spines and tufts of long hairs. Each terminates in
a small apical papilla. The antenna of the pupa, being eventually much
larger than the larval organ, does not develop within the latter but in
a pocket extending posteriorly from the base of the larval antenna.
The eyes of the larva are each a group of simple lateral eyes; their
structure in Culex pipiens has been described by Constantineanu
(1930) and by Sato (1951b). According to Sato each eye consists
of three parts, each with its own retinular cells. One part is central
and has three retinulae, a second part is dorsal and has a single retinula
of eight cells, the third part is a long band of about 40 cells surround-
ing the other two parts dorsally, anteriorly, and ventrally. Con-
stantineanu, on the other hand, describes five parts in the eye of Culex,
as in some other nematocerous larva. Probably the three retinulae of
Sato’s “central part’? are regarded as three eyes. The larval eyes
have no lenses, the ordinary head cuticle being continuous over them.
They are present from the beginning of the larval stage and persist
into the pupa, or even into the adult.
The presence of large, darkly pigmented compound eyes visible on
the surface of the head gives the mosquito larva, as also the corethrid
larva, a very unusual appearance. The compound eyes of other re-
lated Nematocera are developed likewise in the larva, but, because of
the absence of pigment until the pupa stage, they are not apparent ex-
ternally.
The undersurface of the larval head (fig. 3 C,D,E,F) is more dif-
ficult to understand than the upper surface. The mandibles and
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 7
maxillae are articulated on a transverse margin between the bases of
the antennae. The long ventral cranial wall behind them is sclerotically
closed by the union of the postgenae (C, Pge) along an incomplete
median suture (C,D,E, ms). This same condition occurs in certain
other insects, and to understand how it has come about we shall have
to digress on some comparative studies.
The hypognathous position of the insect head in which the mouth
parts are ventral (fig. 2A) is clearly primitive, because the mouth
parts, being modified legs, thus hang down from the head in the posi-
tion of the thoracic legs. The prognathous condition has been attained
in some cases by a mere turning forward of the head on the neck,
involving a ventral elongation of the occipital foramen on the under-
side of the head (fig. 3 B). More commonly, however, the foramen
remains in the vertical plane, as in the mosquito larva (fig. I E), and
the underside of the head is lengthened by a ventral elongation of the
postgenae (Pge).
With the elongation of the postgenae the entire labium, as in some
beetles (fig. 2 B), may be simply enclosed between them, with a gular
addition (Gu) to the submentum. This condition, however, does not
occur in the larval mosquito, though some writers have so interpreted
the mosquito head structure. More commonly, the postgenae come
together medially and displace the labium. A primary stage of this
transformation is seen at C, which might represent the head of a
caterpillar or an adult honey bee, in which lobes of the hypostomal
margins of the postgenae are intruded between the occipital foramen
and the base of the labium. In other cases the lobes become united
(D), forming a bridge between the foramen and the labium. An elon-
gation of the bridge then produces the condition seen in the beetle
larva at EF, in which the labium is still fully exposed. Finally, as in the
larvae of Chironomidae (F), the labium has become greatly reduced
and is hidden from below by a median hypostomal lobe (Hstm) of the
united postgenae.
This same process of closure and elongation of the postgenae and
the reduction of the labium can be traced among nematocerous fly
larvae. For example, in the primitive rhyphid larva of Olbiogaster
(fig. 3 A) described by Anthon (1943b), a pair of small postgenal
lobes are approximated behind the submentum (Smt) of the labium.
In others, as in Trichocera and Philosepedon figured by Anthon
(1943a, figs. 7, 10) the postgenal lobes are united in a bridge; the
labium, though much reduced, is still mostly exposed. In the mosquito
larva (fig. 3 C) the united postgenae form the long underwall of the
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
cranium and the greatly reduced labium is concealed above a median
postgenal lobe (stm) between the maxillae (Vx).
The darkly sclerotized dentate lobe between the maxillae has com-
monly been regarded as a part of the labium, “mentum” or ‘“‘submen-
Fic. 2—Structure of the posterior or ventral wall of the head in different
insects, mostly diagrammatic.
A, Generalized structure of the posterior head wall and the mouth part at-
tachments. B, The labium enclosed between the postgenae. C, The labium
separated from the occipital foramen by intruding hypostomal lobes of the post-
genae. D, The labium entirely separated from the foramen by union of postgenal
lobes forming a postgenal bridge. E, Postgenal bridge lengthened. F, Larval
head of Chironomus, postgenal bridge produced in a toothed lobe, labium dis-
placed dorsally.
For, occipital foramen; Gu, gula; hs, hypostomal sulcus; Hstm, hypostomium,
hypochilum; Lb, labium; Md, mandible; ms, median postgenal suture; Mt,
mentum; Mz, maxilla; Pge, postgena; pos, postoccipital sulcus; Prmt, pre-
mentum; pf, posterior tentorial pit; Smt, submentum.
tum.” To cut a long argument short, however, we have only to look
at a tipulid larva (fig. 3 B) to see that the lobe is formed by the union
of two processes extended forward from the anterior median angles
of the postgenae, which themselves are not united in the tipulid. Above
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 9
this lobe are the united labium and hypopharynx (fig. 7 A, Lb, Hphy).
We may, therefore, following Anthon (1943a), Hennig (1948, 1950),
and Lawson (1951), appropriately call this lobe the hypostomium
(Hstm), as it is termed also by Chiswell (1955) in the tipulid larva.
Though Schremmer (1949) called it “mentum” in the mosquito larva,
he later (1950) expressed doubt of the correctness of this designa-
tion, concluding that the lobe is rather a part of the cranial wall. More
recently, Gouin (1959) has termed the dentate lobe the hypochilum
(underlip).
From the base of the hypostomium there arises in some species a
thin fold bearing a fringe of pectinate hairs or blunt teeth (fig. 15 A,
Aul). The fold is the aulaeum (curtain) of Cook (1944a), but it has
been variously named. Shalaby calls it the “glossa” on the assumption
that it is formed by the union of a pair of labial glossae, a highly im-
probable interpretation since the hypostomium itself is no part of the
labium. However, Shalaby has given detailed illustrations of the
pectinate hairs of the lobe in Aedes aegypti (1957a) and Culex
quinquefasciatus (1957b), and its armature of eight blunt teeth in
Anopheles quadrimaculatus (1956). In Psorophora ciliata (1957c) he
says the fold is absent.
In most mosquito larvae two dark lines in the ventral wall of the
head diverge posteriorly from the basal angles of the hypostomium.
In some species the lines are short (fig. 3 C,D, 7), in others (E,F) they
extend back to the posterior tentorial pits (pt); in Chironomus
(fig. 2 F) they are absent. These lines when present are the external
marks of internal ridges; their variable development suggests that the
ridges are secondarily formed to strengthen the head wall. The surface
area between the lines, however, has commonly been regarded as the
basal part of the labium, probably because the structural pattern they
produce resembles that of the head shown at B of figure 2. It has been
suggested even that the median suture is the line where the two origi-
nal labial appendages have united! Cook (1944a), for some obscure
theoretical reason, calls the area in question the “maxillary segment,”
though the maxillae have no relation to it. That the ventral closure
of the head results entirely from the union of the lateral cranial walls
is clearly indicated in illustrations by Hennig (1948, figs. 31-37) of
larval heads of Sciophilidae, in which are shown various degrees of
approximation and union of the postgenal margins.
In most adult insects the lower edges of the cranium are reinforced
by submarginal internal ridges formed by external grooves known as
the subgenal sulci. The part of each groove on the postgena behind
Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
the mandible is distinguished as the hypostomal sulcus (fig. 2 A, hs).
Posteriorly these grooves become continuous with the postoccipital
sulcus (pos) that surrounds the occipital foramen. In the mosquito
larva the lower ends of the postoccipital sulcus have extended forward
in the postgenal region carrying with them the minute rudiments of
the posterior tentorial arms, the position of which is marked externally
by a pair of pits (fig. 3 E,F, pt). The anterior tentorial arms are
extremely slender bars arising from the cranial margins mesad of
Fic. 3—The larval head, undersurface.
A, Olbiogaster sp., Rhyphidae (from Anthon, 1943b). B, Tipula sp. C,
Anopheles quadrimaculatus. D, Toxorhynchites rutilus. E, Culex sp. F,
Aedes aegypti.
r, r, grooves in lower head wall. Other lettering as on figure 2.
the antennae that extend back to the posterior arms. Possibly it is
the great lengthening of the postgenal regions of the head that has
brought the posterior arms to their forward position.
The postgenal bridge is known also as the hypostomal bridge be-
cause it is the hypostomal margins of the postgenae that come together
to form it. Lawson (1951) contends that the sclerotized ventral wall
of the head behind the mouth parts cannot be derived from the post-
genae because, he says, “the hypostomal sutures form the ventral
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS II
boundaries of the postgenae.” This is clearly making anatomy con-
form with definitions. The lower parts of the postgenae are mechani-
cally strengthened by ridges formed by the submarginal hypostomal
grooves. The narrow strips below the grooves, therefore, are simply
the marginal parts of the postgenae, so it is immaterial whether we call
the bridge resulting from their union hypostomal or postgenal. The
grooves are sometimes absent, and the ridges may be marginal on the
postgenae. In the mosquito larva the anterior edge of the ventral
cranial wall on which the mandibles and maxillae are articulated is the
united hypostomal margins of the confluent postgenae.
From the free cranial margins just mesad of the antennal bases,
a slender bar on each side (fig. 7 B, hb) extends mesally, downward,
and somewhat posteriorly through the preoral epipharyngeal wall to
the base of the hypopharynx (Hphy). Each bar runs close before the
mandible of the same side and goes below the narrow lower lip of
the mouth (Mth). In Dixa, as shown by Schremmer (1950), similar
structures are present but are much wider than in the mosquito larva.
The mandibles have their anterior articulation on these rods, a very
unusual condition, since the anterior mandibular hinges are typically
on the basal angles of the clypeus. The rods have been called “cibarial
bars,” but there is no defined cibarium in the culicid larva. Since the
rods appear to serve principally as suspensoria of the hypopharynx,
they are here termed hypopharyngeal bars. They are the Verbindungs-
leisten of Schremmer (1949). Since the hypopharyngeal bars carry
the anterior articulations of the mandibles, Menees (1958b) reason-
ably argues that the parts of the bars laterad of the articulation are
extensions of the clypeus. His identification of the posterior parts
with the “hypopharyngeal suspensorial bars of generalized insects,”
however, is less convincing, since these bars enter the mouth angles
and give attachment to the hypopharyngeal muscles, though each may
have a lateral preoral branch.
THE FEEDING ORGANS
One of the remarkable things about insects is the way their feeding
organs are variously adapted to feeding in different ways on different
kinds of food. Nothing comparable occurs among the vertebrates,
their only adaptation to the nature of their food is in the size, strength,
and dentition of their jaws or in the length of the neck. Yet the feed-
ing organs of all insects are made up of the same fundamental parts.
There is an upper lip known as the labrum, a pair of mandibles, a
median tonguelike hypopharynx, a pair of ma-illae, and a lower lip, or
I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
labium, composed of a united pair of second maxillae. The mandibles,
maxillae, and labium, furthermore, have been fashioned from three
pairs of legs, since the original arthropods had no other organs for
feeding than their legs. The insect mouth parts, therefore, are all
outside the mouth; the space between them may be termed the preoral
food cavity, but by a long-perpetuated error it has commonly been
called the ‘‘pharynx.” For want of a revised nomenclature we still
speak of the upper wall of the preoral cavity as the epipharyngeal
surface, and call the tonguelike lobe that projects below the mouth the
hypopharynx. This is just a part of our heritage from the early insect
anatomists, who had only vertebrate names to draw from, and applied
them to insects on a functional rather than a morphological basis.
The true pharynx is a part of the stomodaeal section of the alimentary
canal behind the mouth.
The labrum.—The labrum of the mosquito larva includes the small
transverse sclerite on the dorsal wall of the head before the clypeus
(fig. 1 A, Lm), and a larger membranous undersurface that bears
laterally the two vibratory feeding brushes (fig. 4B), and usually
a small median brush. The median brush is the “palatum” of mosquito
students, another example of misuse of a borrowed vertebrate name,
which in this case properly refers to the roof of the mouth cavity.
The lateral brushes of the labrum are the organs by which those
larvae that feed on particles create currents in the water directed
toward the head, and drive a stream of water back to the mouth along
the epipharyngeal surface. The individual hairs of the brushes are
finely pectinate and serve also as combs for retaining particles filtered
from the water.
The vibratory movement of the brushes is produced by a pair of
strongly musculated sclerites on the under side of the labrum. Simi-
lar sclerites are present in the larvae of Chironomidae (fig. 4 G, Tor),
which have no brushes, but the posterior ends of the sclerite are
produced into strong pointed processes (Mes) projecting freely from
the epipharyngeal surface. These toothed sclerites were therefore
called by most earlier writers ‘“premandibles.”’ Chaudonneret (1951),
however, has shown that this term is entirely inappropriate. Cook
(1944b) named the sclerites “messores’” (harvesters) and carried
the term over to the mosquito larvae, in which he has been followed
by several recent writers, though the culicid sclerites are unarmed.
It must be noted that the insect labrum is commonly equipped with
four muscles, one pair dorsal, the other ventral, all of which arise on
the frons. The ventral muscles are usually attached on a pair of
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 13
sclerites in the lower labral wall known as the tormae. In a tipulid
larva (fig. 4A) the tormae (Tor) are simple sclerites, each giving
attachment to a long muscle (mc/) from the frons. There is, there-
fore, no apparent reason why the similarly musculated sclerites of
the mosquito larval labrum (C,D, Tor) should not be the tormae.
On the other hand, Cook (1944b) has contended that the sclerites are
Fic. 4.—The larval labrum and tormae.
A, Tipula sp., underside of larval labrum. B, Culex sp., labrum of young
larva, anterior. C, Anopheles quadrimaculatus, left labral brush and torma,
undersurface. D, Aedes aegypti, same. E, Toxorhynchites rutilus, anterior
view of larval head. F, Same, labral brush with torma and muscles. G,
Chironomus plumosus, underside of larval labrum.
Ap, tormal apodeme; B, labral brush; Hstm, hypostomium; Lm, labrum;
mcl, mels, tormal muscle or muscles; Md, mandible; Mes, messorial teeth of
torma; Myr, maxilla; Tor, torma.
a, connective sclerite between torma and brush; b, c, detached sclerites of
cena wall; d, e, anterior and posterior articulations of torma; f, epipharyngeal
ar.
not the tormae because he finds in chironomid larvae another pair
of muscles attached more dorsally and laterally on the labrum, which
he insists are the true tormal muscles. These muscles, however, would
appear to be the usual dorsal muscles of the labrum, which may have a
lateral position. Furthermore, Cook adds that the ventral muscles are
not tormal muscles because they arise on the clypeus, but what he calls
14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
“clypeus” is the entire frontoclypeal area of the upper head wall be-
tween the cleavage lines. The sclerites in question, being in the
ventral wall of the labrum and giving attachment to the ventral labral
muscles, practically identify themselves as the tormae, and they have
been regarded as such by Anthon (1943a), Schremmer (1949, 1950),
and Menees (1958b). If it is desirable to keep the term “messor,”
it might be restricted to the free prongs of the tormae where they
occur (fig. 4G, Mes).
The tormae of an Anopheles larva (fig. 4C, Tor) are elongate
sclerites lying mesad of the brushes. Each torma is connected by its
tapering anterior end with the base of the corresponding brush;
posteriorly it is hinged to a small sclerite (c) in the cranial margin.
A connective plate (a) lies between the torma and the base of the
brush. A single muscle (mcl) from the frontal region is attached by a
long tendon to a small point anteriorly on the lateral margin of the
torma. Cook (1944a) ascribes a second posterior muscle to the torma
of Anopheles, but this muscle, as shown by Farnsworth (1947) and by
Schremmer (1949), belongs to a V-shaped sclerite of the epipharyn-
geal wall between the posterior ends of the tormae.
In the culicine mosquitoes the tormal apparatus is somewhat more
complex than in Anopheles. In Aedes aegypti (fig. 4D) the tormae
have the same relation to the brushes and the cranial margin as in
Anopheles, but each torma is specifically hinged posteriorly (e) to
a detached triangular plate (b) of the cranial wall, and anteriorly (d)
to the end of a transverse epipharyngeal bar (f). Since both this bar
and the connective plate (a) underlap the torma, the anterior part
of the latter appears to be sunk into the lower wall of the labrum.
Posteriorly a strong apodeme (AP) arises from the dorsal surface of
the torma and curves mesally. On this apodeme are attached two
large muscles (mcls) from the frontal region of the head. Con-
traction of the muscles evidently rocks the torma mesally on its
articular points and thus gives a backward and mesal stroke to the
connected brush. The reverse movement of the brush, as other writers
have noted, results from the elasticity of its basal connections. Ac-
cording to Cook (1944a) in specimens of Theobaldia [Culiseta]
killed and fixed with the brushes retracted, on cutting the muscles
the brushes quickly spring back to the expanded condition.
The Aedes tormal mechanism is probably characteristic of the
Culicinae. The same structure and musculature is shown to be present
in Culex by Thompson (1905) and by Chaudonneret (1951), and in
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 15
species of Theobaldia [Culiseta], Lutzia, and Armigeres by Cook
(19442).
In the predaceous larvae of To.xorhynchites the brushes are sup-
ported on prominently projecting lateral lobes of the labrum (figs.
1D, 4E). The brushes are narrow, stiff, and falciform, and appear
to be grasping organs, but as observed by Breland (1949) and by
Horsfall (1955) they are not used for obtaining prey. Just mesad of
the base of each brush is a small, slender sclerite (fig. 4 E, Tor).
Dissection reveals that this sclerite has a connection with the base
of the brush (F) and gives attachment to two large muscles, leaving
no doubt that it is the torma.
The preoral cavity—The undersurface of the labrum is continuous
with the so-called epipharyngeal surface below the clypeal region,
which extends back to the mouth. In most adult insects the part of
the preoral cavity above the base of the hypopharynx becomes a special
food pocket, the cibarium, opening directly into the mouth. In the
mosquito larva the shortness of both the labium and the hypopharynx
leaves the entire preoral cavity open below, but still it serves as a
channel for water carrying food particles to the mouth. In the tipulid
larva, however, there is a short cibarial pocket (fig. 7A, Cb) above
the hypopharyngiolabial lobe just in front of the mouth. In the adult
mosquito and other sucking insects the closed cibarium becomes a
preoral sucking pump. In the mosquito larva the pharynx assumes
the sucking function.
The epipharyngeal apparatus—Lying in the epipharyngeal surface
between the posterior ends of the tormae is a structure that serves to
comb food particles from brushes on the mandibles. Since it is
musculated, and hence functions actively instead of passively, this in-
strument has been termed by Schremmer (1949) the Epipharynx-
apparat. Other writers have called it the “palatal bar,” the “epi-
pharynx,” and the “epipharyngeal armature.” It includes a transverse
bar and groups of setae or other structures arising in front of the
bar. The crossbar is usually bow-shaped or V-shaped with the arms
diverging forward to the posterior ends of the labral tormae. The
setal accompaniment of the bar is quite different in different species.
In Aedes (fig. 8 A) the epipharyngeal apparatus is relatively simple.
The bar is slender, gently curved forward, and its ends appear to be
connected with the tormal apodemes. Arising in front of the bar
are two large brushes of stiff hairs that converge posteriorly beneath
the bar. At the sides of the brushes arise a pair of large, tapering,
hair-bearing processes directed posteriorly, and at the base of each are
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
two small clawlike structures. In Culex (B) the bar is strongly de-
veloped and angulated, its ends, as in Aedes, appear to be attached
to the apodemes of the tormae. In front of the bar are two large
oval masses of setae curving inward and posteriorly. From above
these setal masses two brushes of long hairs project posteriorly.
Medially there arise two pairs of short tapering processes that project
beneath the bar, and from each angle of the bar a slender, bladelike,
sharp-pointed process extends posteriorly.
The epipharyngeal apparatus of Anopheles as described by Schrem-
mer (1949) is again quite different from that of either Aedes or
Culex. The bar is V-shaped with an acute angle. Several brushes
arise in front of the bar, but particularly developed are two long, wide
combs of flattened, sharp-pointed bristles that extend posteriorly
from a pair of triangular sclerites in front of the bar. These are the
Klingenborsten of Schremmer, who says they are used for cleaning
the food particles from the combs of the mandibles. In Anopheles
maculipennis, as shown by Schremmer and by Farnsworth (1947), a
large muscle from the clypeal region of the head is attached on each
end of the epipharyngeal bar. These muscles the writer has not been
able to find in Aedes and Culex, but the close connection of the bar
with the apodemes of the tormae possibly coordinates the movements
of the epipharyngeal apparatus with the movements of the labral
brushes. In all three genera a pair of very slender, closely adjacent
muscles is attached medially on the bar. Contraction of the lateral
muscles of Anopheles, according to Schremmer, protracts the ap-
paratus from the epipharyngeal wall, the median muscles are retrac-
tors. Thompson (1905) makes no mention of lateral muscles attached
on the epipharyngeal bar in Culex, but he notes the presence of the
median retractors.
The mandibles—Both the mandibles and the maxillae lie on the
underside of the head, where they are implanted obliquely in the
membranous area that turns upward from the hypostomal margins
of the postgenae to the hypopharyngeal bars (fig. 7B, Md, Mx), the
mandibles being above the maxillae.
The typical culicine and anopheline mandibles are flattened lobes
(fig. 5 D,E,F) with their mesal ends produced into strongly sclero-
tized toothed processes and a lower seta-bearing lobe. The dorsal
margins bear large comblike fringes of long setae directed mesally.
The tips of the mandibles on opposite sides do not meet when the
mandibles are closed, but come against the hypopharynx, which lies
between them (fig. 7B, Hphy). Each mandible has a posterior basal
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 17
articular point (fig. 5 E,F, a@) that articulates with a process of the
hypostomal margin just laterad of the base of the maxilla (fig. 7 B, a).
Its anterior articulation (fig. 5 D, c) is with the hypopharyngeal bar
(fig. 7B, hb). The mandibles move in the transverse plane by strong
abductor and adductor muscles. The principal function of mandibles
of this type is the collection by their setal combs of food particles from
Fic. 5.—Larval mandibles.
A, Tipula abdominalis, right, ventral. B, Lutzia sp., right, ventral. C, To-xro-
rhynchites rutilus, left, dorsal. D, Anopheles quadrimaculatus, left, dorsal. E,
Culex, sp., right, ventral. F, Aedes aegypti, right, ventral.
a, posterior (ventral) articulation; c, anterior (dorsal) articulation.
the labral brushes, but the incisor points are said to break up larger
particles that collect on the hypopharynx.
The mandibles of predaceous larvae, such as Culex vorax (fig. 5 B)
and Toxorhynchites (C), are strongly toothed jaws, the points of
which come together in adduction (fig. 4 E). Most larval Nematocera
have jawlike mandibles (fig. 5 A), though they present many varieties
of structure. C. vorax is a culicine mosquito, and its mandible (B)
might be derived from the culicine type, but the mandible of To..o-
rhynchites (C) is a typical biting insect jaw.
The maxillae—The maxillae of the mosquito larva (fig. 6 B-F) are
so greatly simplified that they have lost the appearance and structure
of an ordinary insect maxilla. They are borne on the transverse
hypostomal margins of the postgenae at the sides of the hypostomium,
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
where they lie below the mandibles (fig. 3 C-F, Mx). The principal
part of each maxilla is a flat lobe (fig. 6 D, St) of different shape in
different species, bearing brushes of long setae or combs of shorter
ones. Laterad of this lobe is a second cylindrical or fusiform lobe
regarded as the palpus (P/p) varying in size relative to that of the
mesal lobe. At the base of the palpus is usually a small sclerite (+)
in the articular membrane.
A
ry
=
be
Fic. 6.—Larval maxillae, right, ventral.
A, Tipula abdominalis. B, Culex vorax. C, Culex sp. D, Aedes aegypti.
E, Anopheles quadrimaculatus. F, Toxorhynchites rutilus.
Plp, palpus; St, stipes; x, sclerite at base of palpus.
In other nematocerous larvae, as in Tipula (fig. 6 A), the maxillary
palpus (PIp) is a small lateral appendage of the main maxillary lobe,
as it is also in the culicid Culex vorax (B). In most mosquito larvae,
however, the palpus appears to have somehow become separated from
the rest of the maxilla (C-F). The main maxillary lobe in some
nematocerous larvae, as shown by Anthon (1943a), may bear on its
distal margin mesad of the palpus two variously developed out-
growths, which are identified as the galea and lacinia. The main
maxillarly lobe, therefore, appears to be the stipes (St). The nature
of the small sclerite (1) at the base of the palpus is uncertain. Cook
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 19
(1944a) calls it the “palpiger,” but it might be referred to the cardo,
though no muscles are attached on it.
The principal functional features of the culicid larval maxillae are
their setal brushes and combs, which serve to collect food particles
from the labral brushes. In the predaceous To.xorhynchites the
maxillae (fig. 6 F) are similar to those of other species, but they are
greatly reduced in size (fig. 4E, Mx). The palpi are presumably
sensory organs, but their disparity in size, as between Culex (fig. 6 C)
and Anopheles (E), for example, is difficult to explain. The princi-
pal movements of the maxillae are in the transverse plane.
The labium and hypopharynx.—In most adult insects the salivary
duct opens between the bases of the hypopharynx and the labium. In
some larval insects, as in caterpillars and hymenopterous larvae, the
labium and hypopharynx are united in a single suboral lobe traversed
by the duct of the salivary, or silk, glands, which opens at the tip
of the composite lobe. The same is true of some nematocerous fly
larvae, as is well seen in the tipulid (fig. 7 A, SIDct). In the mosquito
larvae the combined labium and hypopharynx are reduced to a flat or
somewhat protruding vertical surface between the mouth and the
hypostomium, with the salivary duct opening on it. The salivary
orifice, therefore, separates the dorsal hypopharyngeal component
from the ventral labial component.
The hypopharynx (fig. 7 B, Hphy) is supported by the hypopharyn-
geal bars (hb) from the lateral cranial walls; immediately above it is
the wide mouth (Mth) opening into the pharynx. The labial area
below the hypopharynx (D,E, Lb) is variously developed, usually
strongly sclerotized and armed with spines or teeth. Other writers
have well illustrated the details of the labial structure in different
mosquito species. Some have attempted to analyze the larval labium
into the parts of a typical insect prementum, but their results are
not fully convincing. At C of figure 7 is shown the labiohypopharyn-
‘geal complex of Toxorhynchites rutilus in dorsal view, in which the
salivary duct (S/Dct) is seen opening between the two component
parts. Attached laterally on the base of the labium are the tendons of a
pair of muscles from the ventral head wall, as in the tipulid larva (A).
Inasmuch as all the cranial muscles of the insect labium are inserted
on the prementum, the labium of the mosquito larva is evidently the
prementum; the hypostomium and the ventral head wall, as already
shown, being no part of the labium. Menees (1958a), however, has
argued that the ventral head area behind the hypostomial lobe must be
the labial submentum because the labial muscles have their origins on
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
it. He thus assumes that these muscles are the retractors of the pre-
mentum. The premental retractors, when present, do arise on the
submentum, but they are always median in position. The muscles of
the mosquito larval labium are lateral muscles, and therefore should
pl i
y We nencls- ifs
iL FrGne LZ
LZ
i mi
Z/f
AG ee ra
it
fA
iit Ati i
Fic, 7.—Labiohypopharyngeal complex of larvae, and associated structures.
A, Tipula sp., section of larval head. B, Culex sp., posterior part of head,
anterior. C, Toxorhynchites rutilus, labiohypopharyngeal complex, dorsal. D,
Aedes aegypti, same, anterior. E, Culex sp., same, anterior
a, posterior articulation of mandible; Ant, antenna; c, anterior articulation
of mandible; Cb, cibarium; Clp, clypeus; dlcb, dilator muscles of cibarium;
Fr, irons; FrCon, brain connective of frontal ganglion; FrGug, frontal ganglion;
hb, hypopharyngeal bar; Hphy, hypopharynx; Hstm, hypostomium; Lb, labium;
lbmcl, muscle of labium; lbrmcl, labral muscle; Lm, labrum; Md, mandible;
Mth, mouth; Mx, maxilla; phmcls, pharyngeal muscles; RNv, recurrent nerve;
S/Dct, salivary duct; Stom, stomodaeum.
be one pair of the usual two pairs of cranial muscles of the prementum,
which in other insects commonly arise on the tentorium. The same
muscles in the tipulid larva (fig. 7 A, /bmcl) certainly have their
origins on the head wall, since there is no sclerotization between the
postgenae (fig. 3 B). The labial muscles of the mosquito larva, there-
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 2]
fore, do not identify the head area on which they arise as any part
of the labium.
The interpretation of these parts has been still further confused by
Shalaby (1957d), who regards the median ventral head area as the
labial submentum and mentum, the toothed hypostomial lobe as the
paraglossa, the fringed lobe below it the glossa, and the entire complex
above the toothed lobe the hypopharynx. Comparative studies, as
already shown, give no basis for any such interpretation. Moreover,
the adult labium is formed entirely from the rudiment beneath the
cuticle of the larval labium (figs. 9 F, 15 A, pLb) and involves no
part of the ventral head wall of the larva.
The larval labiohypopharynx is evidently retractile, but it plays no
active part in feeding. Its principal function is said to be that of
an “anvil” on which the incisor points of the mandibles strike to break
up food particles.
Elaborate studies of the developmental changes in the mouth parts
of larval instars of Anopheles, Aedes, Culex, and Psorophora have
been made by Shalaby (1956, 1957a, 1957b, 1957c).
The pharynx.—The pharynx of larvae that feed on water-borne
particles is a small, flattened, ovate or heart-shaped, thin-walled sac
(fig. 8 C, Phy) opening directly from the wide mouth (fig. 7 B, Mth)
and tilted upward and posteriorly in the head. From its posterior
ventral surface is continued the thick-walled oesophagus (fig. 8C,
Oe). The ventral wall has an outer layer of semicircular muscles
(E, cmcl) the dorsal wall is crossed by four wide muscle bands (C,
tmcl) ; the extrinsic musculature includes dorsal and ventral dilator
muscles from the head wall. The lateral margins of the pharynx are
strengthened by two narrow, concentric, riblike thickenings on each
side, convergent to the narrowed posterior end. Internally each of
these ribs bears a long brush of fine hairs (D), suggestive of the
brushes in the mouth of a baleen whale, and in fact they serve the
same purpose, namely, that of filtering the food matter from the in-
gested water. A pharyngeal filter apparatus very similar to that of
the mosquito larva is shown by Anthon (1943a) to be present in the
larvae of several other nematocerous families. The pharynx of the
predaceous culicid larva of Toxorhynchites, however, is a simple
funnnel-shaped enlargement of the anterior end of the oesophagus,
and has no filter brushes. In any case, the larval pharynx is not to be
identified with the sucking pharynx of the adult mosquito, which lies
in the posterior part of the head (fig. 24 A, PhP), and the larva has
no cibarial pump.
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Larval feeding.—tThe process of feeding by nonpredaceous larvae
is not a mere matter of having food particles washed into the mouth
by streams of water from the vibrating labral brushes. It involves
cooperative action on the part of the labrum, the epipharyngeal
apparatus, the mandibles, the maxillae, the labiohypopharynx, and the
pharynx. The whole feeding process has been minutely described by
Schremmer (1949) for the Anopheles larva, in which it is more readily
Fic. 8—The larval epipharyngeal apparatus and the pharynx.
A, Aedes aegypti, epipharyngeal apparatus. B, Culex sp., same. C, Anopheles
maculipennis, pharynx and its muscles, dorsal (from Schremmer, 1949). D,
Same, filter-bearing rib of pharynx. E, Same, cross section of pharynx (from
Imms, 1907).
cmcel, circular muscle; fb, filter brush; Mth, mouth; Oe, oesophagus; Phy,
pharynx; tmcl, transverse muscles.
observed than in other species because the head is held with its under-
side turned upward against the surface of the water. Briefly, Schrem-
mer’s account is as follows.
The movements of the lateral brushes of the labrum create currents
in the water that converge to the front of the head and are directed
medially by the middle brush. With the backward stroke of the lateral
brushes the mandibles and the maxillae are closed upon them, and
as the brushes again go forward particles that may be adhering to
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 23
them are scraped off by the combs of the mandibles. Accompanying
the opening of the mandibles, the epipharyngeal apparatus is pro-
truded by action of its muscles and its bristles remove the food
particles from the mandible combs. These freed particles and others
that may be adhering to the epipharyngeal surface are then collected by
the long basal brushes of the mandibles (fig. 5 D) and, with the closure
of the mandibles, are pushed into the mouth of the pharynx. Though
the mandibles and the maxillae close at the same time, the maxillae
open first and the mandibles following remove whatever particles
may be adhering to the maxillae, which lodge on the hypopharynx
and with the next stroke of the maxillae are thrust into the pharynx.
Large particles collected on the hypopharynx are broken up by the
toothed lobes of the mandibles, which strike on the hypopharynx like
hammers on an anvil.
The pharynx, by muscular expansion of its walls, functions as
a sucking organ for drawing in a stream of water accompanying the
mechanically ingested food particles. A contraction then follows in
which the dorsal wall is deeply infolded by the action of the dorsal
transverse muscles (fig. 8 E), reducing the pharyngeal lumen to two
lateral channels containing the filter brushes (fb). At the same time
the water is driven toward the mouth and the food particles are
filtered out by the brushes. The water is then discharged through the
open angles of the mouth, goes above the mandibles and escapes past
the sides of the head. Schremmer made further experiments on a
Culex larva by impregnating the water in a dish with carmine particles.
After feeding by the larva, the carmine was found massed in the
brushes along the sides of the pharynx. When the pharyngeal brushes
have worked as filters for some time and have become well loaded, the
pharynx makes a strong contraction which suddenly removes the
carmine particles from the brushes and lodges them in small clumps
at the mouth of the oesophagus, into which they are finally taken. The
mosquito larva swallows no appreciable amount of water, its water
balance being maintained by the anal lobes.
The extreme specialization of the mouth parts and the pharynx in
the filter-feeding mosquito larvae gives a striking example of how
independent of the adult structure an insect larva may become in its
adaptation to a new way of feeding. In various mosquito genera,
however, the larvae of some or all species are predaceous on other
small aquatic animals, particularly on other mosquito larvae. Notable
in this group are members of the subgenus Lutgia among the Culicini,
and of the genus Toxorhynchites. In these forms the mandibles are
24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
strongly developed jaws (fig. 5 B,C), the toothed lobes of which come
together or overlap for grasping and biting. Yet these larvae have
labral brushes and some of the other special features of particle-feed-
ing larvae, so it is difficult to say whether they represent a partway
stage in the evolution of filter feeding, or have been secondarily
adapted for feeding on whole live prey. In some species the larvae are
particle feeders in the first instar and become predaceous in their later
instars. It would appear, therefore, as said by Bates (1949), “that
the predacious habit has developed independently in the larvae of a
number of mosquito groups, involving distinct adaptations both of
structure and behavior.”
THE THORAX
The larval thorax has a simple oval form, in which the interseg-
mental lines are but faintly marked as grooves of the cuticle, and
there is no external trace of appendages. In the fourth instar the
thorax becomes conspicuously enlarged (fig.9g A). Beneath the cuticle
on the ventral side are now plainly visible the extroverted wings and
legs of the future pupa, and on the dorsal side the pupal respiratory
trumpets. On removal of the cuticle (C) the legs are seen to be long,
fully segmented appendages (E) closely folded in loops against the
sides of the thorax. The forewings (W-) are large pads corrugated
in their basal parts (D) to allow expansion; the smaller hindwings
(Ws) are more slender and tapering free folds of the metanotum. It
has been shown by Imms (1908) that the rudiments of the wings,
legs, and respiratory trumpets are formed in a young larval instar of
Anopheles as integumental folds in pockets of the epidermis (B). Ap-
parently they are extruded beneath the cuticle at the beginning of the
fourth instar. This early eversion of the wings and legs occurs also
in other nematocerous larvae, such as Dixa, Corethra, and Chirono-
mus, shown by Miall and Hammond (1900) in Chironomus.
On each anterior lateral angle of the thoracic dorsum of Anopheles
larvae there is usually to be seen a pair of minute, tapering, trans-
parent lobes arising from a common base (fig. 9 A, no). These struc-
tures are known as the “notched organs.” They are retractile and
hence are not visible on all specimens, or only their tips may project.
Between the lobes of each pair is a funnel-shaped depression that ends
in a strand, which is said by Chang and Richart (1951) to be at-
tached to the neighboring dorsal tracheal trunk. These writers con-
tend, therefore, that the organs are the “prepupal respiratory trum-
pets.”” However, when the cuticle of a fourth-instar larva is removed,
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 25
the lobes and the funnel come off with it, showing that the organs are
larval structures. Furthermore, the trumpets of the “prepupa’’ (i.e.,
the pharate pupa) are present beneath the larval cuticle. They appear
to arise from the pupa just beneath the larval organs, but they project
forward or mesally until the pupal ecdysis, when they stand out from
the thorax.
Fic. 9.—The larva, and developing pupal appendages.
A, Anopheles punctipennis, fourth instar larva, dorsal. B, Anopheles maculi-
pennis, section of wing and leg buds in early larval instar (from Imms, 1908).
C, Aedes aegypti, thorax of mature larva with cuticle removed, exposing extro-
verted legs and wings. D, Same, pupal wings of larva. E, Same, third left pupal
leg of larva. F, Anopheles maculipennis, pupal labium developing inside larval
labium (from Imms, 1908).
al, anal lobes; e, dorsal brush of larva; L, leg bud; pLb, pupal labium; s/O,
salivary orifice; Sp, spiracle; W, wing bud; We, Ws, pupal wings of larva.
The nature of the “notched organs” of the Anopheles larva is not
clear. Their position on the dorsum of the thorax suggests that they
might be remnants of anterior spiracles such as are present on larva
of many other flies, including some Nematocera. Since these spiracles
of successive instars are not formed in the usual manner within the
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
preceding spiracle, but as independent branches from a persisting
spiracular atrium, it is perhaps possible that the pupal trumpets are
in this manner related to the “notched organs” of the larva. Chang
and Richart contend that the latter serve to keep the anterior part of
the Anopheles larva afloat while feeding at the surface, but experi-
ments have shown that the organs can be cut off without any apparent
effect on the suspension of the larva (Jones, unpublished obser-
vations).
THE ABDOMEN
The larval abdomen (fig. 10 G) appears to have only nine segments,
and it is usually represented as nine-segmented, with the respiratory
apparatus on the eighth segment and the terminal segment enumerated
as the ninth. However, there is reason for believing that a true ninth
segment is combined with the eighth. Christophers (1922) con-
tended that though “much of the apparent eighth segment is actually
this structure, the greater part of the spiracular apparatus must
be assigned to the tergite of a hitherto unrecognized ninth abdominal
segment.” Convincing evidence of this interpretation is the fact that
the rudiments of the male genitalia are formed beneath the larval
cuticle at the base of the terminal segment, and that in the adult male
the genital claspers are carried on the posterior margin of a small
but distinct ninth segment (fig. 27 B). Though this segment is not
evident as a distinct annulus in the larva, it must be represented by
some part of the apparent eighth segment immediately anterior to the
genital rudiments. In the pupa, as will be shown (fig. 16 D,E) a small
ninth-segment ring (JX) lies behind the eighth segment and carries
the tail fins and the small anal lobe. The anal segment of the larva
(fig. 10 B) must therefore be the tenth, as it is in the pupa and the
adult.
The fully segmented abdomen of the mosquito embryo is shown by
Telford (1957) in Aedes and by Menees (1958a) in Anopheles to
have Io segments. Telford says the tenth segment, or telson, dis-
appears with the ingrowth of the proctodaeum, but since a tenth seg-
ment is present in the adult, the “telson” must be an eleventh segment.
In some larvae, as seen in Mansonia (fig. 11 A) a small lobe (XJ)
protrudes from the end of the tenth segment, which would appear to
be the evaginated anus-bearing telson. Even in the embryo, then,
the ninth segment is not differentiated from the eighth. It appears as
a distinct ring first in the pupa and as a definite segment in the adult.
The first seven segments of the larval abdomen have no distinctive
features, except that in Anophelini (fig. 9 A) the last five or six of
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 27
them bear on the back pairs of small palmate brushes (¢) that suspend
the larva from the surface of the water in its usual horizontal feeding
position. The respiratory apparatus on the dorsum of the ninth seg-
mental region contains a pair of large open spiracles, which are either
Fic. 10—Larval respiratory organs.
A, Anopheles maculipennis, tracheal system, dorsal (from Imms, 1907). B,
Anopheles quadrimaculatus, end of abdomen, spiracular apparatus open. C, Same,
spiracular apparatus closed. D, Same, details of spiracular apparatus. E, Culex
pipiens, end of abdomen. F, Aedes aegypti, same. G, Culex quinquefasciatus,
larva in feeding position. H, Culex pipiens, end of respiratory siphon.
al, anal lobes; dTra, dorsal tracheal trunk; Sp, spiracle; VIJI-X, abdominal
segments.
flush with the surface (fig. 10 A, Sp) or carried out on the end of a
respiratory tube (E,F,G). The tenth segment contains the func-
tional anus at its end, and bears four lanceolate, thin-walled apical
appendages, or anal lobes (A,B,E, al). Flat dorsal and ventral brushes
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
of long, spreading hairs are usually present on the end of the tenth
segment, and perhaps serve as a rudder during swimming. Though
mosquito larvae are commonly known as ‘“‘wrigglers’” or “wigglers”
they swim by lashing movements of the abdomen, which drive them
forward, backward, or sideways. The active larvae of Culex zigzag
through the water like tumbling acrobats. Anopheles, however, is a
true wriggler ; it swims either on the surface or under the water by
quick lateral movements of the abdomen and propels itself backward.
The dorsal spiracles of the abdomen are the only breathing aper-
tures of the mosquito larva. The lateral spiracles are closed except
at the ecdyses, when they are temporarily opened to allow the tracheal
linings to be pulled out. Since the dorsal spiracles open into the dorsal
trunks of the tracheal system and the lateral spiracles into the lateral
trunks (fig. 10 A), the dorsal spiracles cannot be supposed to be a
pair of lateral spiracles that have moved up onto the back. It may be
conceded that spiracles can change their position, but they cannot
change their tracheal connections.
The spiracles of anopheline larvae lie in the floor of a shallow,
basinlike peritremal structure elevated on the back, the margins of
which are variously produced into lobes (fig. 10 B). In Anopheles
maculipennis (D) there are two large, thin posterior lobes, a pair of
small tapering lateral lobes, and a single anterior lobe supported on
a transverse basal bar. The spiracles (Sp) lie anteriorly; behind
them is a median V-shaped sclerotization on the floor of the basin,
and on each posterior lobe is a weak submarginal sclerotization. As
the Anopheles larva feeds stretched out against the surface film of
the water the peritremal basin projects just above the water with the
spiracles freely exposed to the air. When the larva submerges, the
whole apparatus folds up and the lobes clamp tight together (C).
Imms (1908) describes three sets of paired muscles that effect the
closing of the lobes, which retain a bubble of air between them. When
the muscles relax the lobes open. Curving around the end of the
ninth segment beneath the ends of the posterior lobes is a narrow semi-
circular bar that supports on each side a small plate bearing a comb
of strong recurved bristles (D), or in some species is armed with
spines or teeth.
In the larvae of Culicinae and Toxorhynchitinae the spiracles are
carried out on the end of a tube, or siphon, varying in length and
thickness in different genera (fig. Io E,F). The spiracles are at the
end of the tube and are surrounded by lobes similar to those in
Anopheles, but necessarily much smaller (H). When the larva is at
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 29
the surface it hangs from the end of the siphon with the spiracles ex-
posed to the air (G). Two strands of slender muscle fibers traverse
the tube and converge to attachments on a strong apodeme from the
terminal apparatus.
An extensive comparative study of the peritremal structure has
been made by Montschadsky (1930) from a taxonomic standpoint.
His illustrations are not realistic since they appear to have been drawn
from flattened specimens, and the sclerotic parts are overemphasized
by an unnaturally dark tone, but they show the great specific variation
in the pattern of the peritremal lobes.
Glands associated with the spiracular apertures have been described
by Keilin, Tate, and Vincent (1935). The secretion is oily and serves
to give a hydrofuge quality to the peritremal surface, which prevents
wetting and the entrance of water into the spiracles.
Though the respiratory siphon is primarily constructed for breath-
ing air at the surface of the water, in species of Mansonia and a
species of Ficalbia it is modified for insertion into the roots of aquatic
plants. The siphon tapers distally and the apex is armed with spines,
teeth, and hooks, which, operated by the inner muscles of the tube,
enable the larva to insert the tip of the organ into the plant. In
Mansonia indubitans (fig. 11 A) the siphon is large, conical in shape,
and narrowed at the distal end. The apex is not sharp, but is armed
with a pair of strongly toothed movable lobes (B), which can be re-
tracted and brought together, or protracted with the teeth turned out-
ward. The siphon in this case is a cutting and not a piercing instru-
ment. It contains only one tracheal trunk, formed by the union of
the dorsal body trunks in the eighth abdominal segment, and there
is a single median, ventral spiracle between the bases of the toothed
lobes. These larvae live entirely submerged and obtain their air from
the air channels of the plant, to which they remain attached.
According to Iyengar (1935a, 1935b) species of Mansonia in India
attach themselves only to the water plant Pistia stratiotes. To insert
the siphon the larva moves backward with the siphon held horizon-
tally and thrusts the tip against the root. It then wriggles actively
backward, while it operates the apical armature with muscles attached
on a rodlike apodeme, until the end of the siphon penetrates the root
deep enough to enter an air chamber, when apical hooks anchor the
larva to the root. The adult female lays her eggs only on submerged
leaves of the Pistia plant, thrusting her abdomen into the water to do
so, and where Pistia is not present she will lay no eggs.
While most other mosquito larvae spend most of their time at the
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
surface of the water, any of them can stay below without apparent
discomfort, and some do so indefinitely. It was formerly supposed
that the four thin-walled tracheated lobes borne on the end of the
tenth abdominal segment were gills serving for underwater respira-
tion. Wigglesworth (1933), however, has produced evidence that
Fic, 11.—Respiratory tubes of larvae and pupae that get their air from the
roots of aquatic plants.
A, Mansonia indubitans, terminal segments of larva. B, Same, apical part of
siphon, ventral. C, Mansonia sp., thoracic respiratory horns of pupa, dorsal. D,
Same, right horn, mesal. E, Mansonia richiardii, pupal respiratory horns, dorsal
(from Wesenberg-Lund, 1920-21). F, Ficalbia hybrida, terminal part of pupal
respiratory horn (from Bonne-Wepster, 1932).
An, anus; Tra, trachea.
these lobes are water-absorbing organs rather than gills. By immers-
ing larvae in a water culture of the flagellate protozoon Polytoma,
which is highly sensitive to the amount of oxygen in the water, he
found that the flagellates first assemble at the posterior end of the
larva and then spread all over the body surface. Soon, however, they
move away in a mass, indicating that oxygen is being consumed by
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS ie
the general integument of the larva as well as by the anal lobes. The
submerged mosquito larva, therefore, breathes through its skin, and
some other aquatic larvae are known to do the same.
From experimental ligaturing of the body of the larva in different
places, Wigglesworth furthermore showed that the larva absorbs
water from the posterior end of the body, presumably through the
thin, permeable anal lobes. During feeding, the larva does not swallow
the water taken into the pharynx with its food, this water, as already
noted, being discharged from the mouth. The anal lobes thus serve
to maintain the physiological balance of water in the larval body.
INTERNAL ANATOMY
Inasmuch as the principal specializations of the mosquito larva have
to do with feeding and breathing, there is little in the internal or-
ganization that is essentially different from that of other insects.
The tracheal system.—The tracheal system of most insects in-
cludes a pair of lateral tracheal trunks running lengthwise through
the body, with which the lateral spiracles are connected. Many insects,
however, have also a pair of dorsal longitudinal trunks. In dipterous
larvae, including the mosquito larva, that breathe through dorsal
spiracles, the dorsal trunks are particularly large (fig. 10 A, dTra),
and the lateral trunks connected with the closed lateral spiracles
are mutch reduced. The dorsal spiracles of the ninth abdominal
segment are evidently secondary respiratory apertures to allow the
larva to breathe at the surface of the water, since it is hardly to be sup-
posed that a primitive lateral spiracle could migrate dorsally and
change its tracheal connections. In general the last pair of lateral
spiracles is on the eighth segment. In the larvae of higher Diptera
there is also a pair of secondary anterior dorsal spiracles on the thorax.
The fine end branches of the insect tracheal system in general go to
the cells of the body tissues, which are thus directly oxygenated. In
the larva of Anopheles, Imms (1907) describes a series of small tubes
from the longitudinal trunks in the eighth abdominal segment that
break up into fine branches going to the posterior end of the heart.
Imms suggested that these branches may oxygenate the blood in the
heart, but Jones (1954) says they end on the heart wall.
At each larval ecdysis the cuticular intima of the tracheal tubes is
shed with the outer cuticle. In the mosquito larva, according to Wig-
glesworth (1949), the intima of the main tracheal trunks breaks be-
tween the segments, and the pieces attached to the shed cuticle are
drawn out through the lateral and the posterior dorsal spiracles of
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
the new instar. The lateral spiracles are then closed again, since they
are not functional in the larva for respiration. In the same manner,
at the ecdysis of the pupa the tracheal trunks in Culex are said by
Hurst (1890) to break up into segmental pieces, which are pulled out
through the temporarily opened spiracles. The soft inner tissue of
the respiratory siphon is withdrawn into the body where it is finally
absorbed. The siphon itself is shed with the larval cuticle, and its two
tracheal trunks break off at the base.
The tracheal system of the young larva on hatching is filled with a
liquid. According to Frankenberg’s (1937) observation on Culex,
air enters the tracheae only when the end of the respiratory siphon
comes above the water surface. One of the dorsal longitudinal trunks
fills first, and then the other. The air is drawn into the tracheae as
the embryonic liquid diffuses through the tracheal walls.
The dorsal blood vessel——vThe dorsal blood vessel of the mosquito,
particularly in Anopheles quadrimaculatus, has been elaborately de-
scribed by Jones (1954). Structurally it differs in no essential respect
from the vessel of other insects, except for a dilatation, or sinus, of
the aorta in the thorax. The larval organ is a simple muscular tube
extending along the midline of the back from the eighth abdominal
segment into the head. The part in the abdomen, known specifically as
the heart, is perforated along the sides by eight pairs of segmental
openings, or ostia. The part in the thorax, called the aorta, is im-
perforate. In the head the aorta goes beneath the brain, where it is
open ventrally allowing the blood to be freely discharged into the head
cavity, whence it flows backward through the body to reenter the heart
through the ostia. The larval heart, Jones says, always beats forward
at an average of 85.2 pulsations a minute, but it has no nerve connec-
tions. Along the sides of the heart are attached the usual fan-shaped
segmental groups of muscle fibers, the so-called alary muscles, that
support the heart on the body wall.
The alimentary canal.—In the mosquito larva the alimentary canal
(fig. 12) is a relatively simple tube. It consists of the usual three
parts of the arthropod digestive tract, an ectodermal stomodaeum, an
endodermal mesenteron, and an ectodermal proctodaeum. The sto-
modaeum begins in the head with the pharynx (Phy), which is fol-
lowed by a narrow oesophagus (Oe) that goes through the neck into
the thorax, where it enters the first part of the mesenteron, known as
the cardia (Car). (This term, borrowed from vertebrate anatomy,
has no literal significance in the insect.) Within the cardia the oesopha-
geal walls are reflected to form the usual entrance funnel of the
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 33
stomodaeum into the mesenteron. The cardia is followed by a long,
straight tube, the stomach, or ventriculus (Vent), that extends back
into the seventh abdominal segment. The anterior end of the ventricu-
lus bears a circle of eight large pouchlike diverticula, the gastric
caeca (GCa). The dark mass of food particles in the ventriculus is
contained in a thin tubular peritrophic membrane (PMb), shown by
Wigglesworth (1930) to be secreted by the cell walls of the cardia
surrounding the stomodaeal funnel. The proctodaeum, or intestine,
is differentiated into a short anterior part (Alnt), and a longer
posterior part, or rectum (Rect). The anterior intestine begins as an
expansion against the end of the ventriculus, and then narrows to a
tube that makes an S-shaped bend to the saclike anterior enlargement
Fic. 12—Lengthwise section of a Culex larva, showing the alimentary canal.
Alnt, anterior intestine; An, anus; Car, cardia (anterior part of ventriculus) ;
GCa, gastric caeca; Hstm, hypostomium; Mal, Malpighian tubules; Mth, mouth;
Oe, oesophagus; Phy, larval pharynx; PMb, peritrophic membrane; Lect,
rectum; S/G/d, salivary glands; Vent, ventriculus (stomach).
of the rectum, which finally proceeds as a narrow tube to the anus
(An). ae
For a detailed study of the general structure, histology, and move-
ments of the larval alimentary canal of Anopheles the reader is
referred to a forthcoming paper by Jones (in press).
The Malpighian tubules—The excretory Malpighian tubules of
the larva (fig. 12, Mal) are five in number. They arise from the an-
terior end of the proctodaeum, first going forward into the sixth
abdominal segment, and then turning posteriorly to end in the sub-
terminal segment around the rectal sac.
The salivary glands——The larva has a pair of small salivary glands
of various shapes lying ventrally in the thorax (fig. 12, SIG/d). The
ducts unite in a common outlet duct that enters the head and opens on
the labiohypopharyngeal surface just below the mouth (fig. 15 A,
SIO). The glands usually consist each of two parts of different shape
34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
separated by a constriction. The histology of the glands in Anopheles
larvae has been described by Jensen and Jones (1957). In Anopheles
albimanus the globular anterior part of each gland consists of 12 to 15
large cells; the pear-shaped posterior part contains 50 to 60 much
smaller cells. The glands of opposite sides are connected by a strand
of nephrocytes. In other genera the relative size and shape of the
two gland parts differ in various ways.
The nervous system.—The central nervous system of the larva in-
cludes a brain and suboesophageal ganglion in the head, and a ventral
chain of segmental ganglia in the abdomen united by paired connec-
tives. The last ganglion is that of the eighth abdominal segment.
The reproductive organs——Rudiments of the reproductive organs
are present in the young larva in a very elementary state; they
slowly develop during the larval life.
Food reserves.—The insect larva has no idea of the meaning of its
life or of what is to become of it. Its hereditary factors automatically
determine its destiny by converting it into a pupa and finally into an
adult. Yet, physiologically, the larva is loaded with responsibilities.
Not only must it maintain its own existence, but at the same time it
must provide for the future nutritional needs of the pupa and for its
transformation to the adult. In the mosquito pupa there is a minimal
breakdown of larval tissues to furnish food for the developing adult
organs. The active mosquito pupa, moreover, is not a “resting stage,”
and, since it cannot eat, it is dependent upon the larva for everything
except the air it breathes. An important function of the larva, there-
fore, is the storage of food reserves in its body to maintain the pupa
and to insure the development of the adult. Only when the winged
adult finally emerges from the pupal skin can the mosquito again take
food and become once more an independent, self-sustaining insect.
The elaboration and storage of food reserves in the body of the
fourth-instar mosquito larva is the subject of a special study by
Wigglesworth (1942). The stored materials include principally pro-
tein, fat, and glycogen, which are shown by experiments to be rapidly
consumed when the larva is subjected to starvation, and replenished on
subsequent feeding. Normally, it is to be supposed, the stored prod-
ucts are passed on intact to the pupa, but Wigglesworth does not go
into this phase of the subject, or follow the utilization of the reserves
by the pupa. The matter, however, is well-enough known in other
insects.
Peewee
The active pupa is familiar to all students of mosquitoes after its
ecdysis from the larva. The fact, however, that it is already fully
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 35
formed shows that it became a pupa while still within the larval cuticle.
It will therefore be of interest to follow the transformation processes
that convert the larva into a pupa.
THE PUPAL DEVELOPMENT
As before noted, the primary buds of the pupal wings, legs, and
respiratory trumpets are formed at an early larval period in pockets
of the epidermis beneath the cuticle, as are also those of the antennae
and the labium, and rudiments of the compound eyes are present in
the first instar.
Fic. 13.—Pupae in natural floating position against the surface of the water.
A, Aedes atropalpus. B, Anopheles punctipennis.
The development of the compound eyes of the mosquito has been
described by Zavrel (1907), by Constantineanu (1930), and by Sato
(1951a, 1953a, 1953b). The eye rudiments are first evident in the
first larval instar as thickenings of the epidermis just in front of the
larval eyes. With development of the eye pigment, the compound eyes
become visible externally in the second instar or the early part of the
third instar. From then on they increase in size as the ommatidia are
slowly differentiated in the epidermis. During the larval stage the
ommatidia are covered by the unmodified cuticle, but in the pupa the
cuticle over each ommatidium becomes convex and the corneal facets
are thus defined. After emergence of the adult the lenses become bi-
convex, and the ommatidia are completed in from 3 to 12 hours, but
the lenses may continue to thicken during the first 24 hours of adult
life.
The early development of the wings and legs in the mosquito larva
is nothing unusual. The leg buds are always formed in the embryo,
and all immature insects have legs, whether external or internal. Like-
36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
wise the young of all winged insects have wing rudiments developing
either externally or internally. The unusual thing about the mosquito
and related Diptera is that the legs, wings, and pupal respiratory
trumpets are fully extruded beneath the cuticle of the thorax at the
third larval moult instead of at the moult to the pupa (fig.9 C). The
wings are still in the form of pads (W2, Ws), but the legs (E) are
already fully segmented appendages.
At a somewhat later period of the fourth instar, the larval cuticle
is separated from the abdomen except at the posterior end (fig. 14),
and beneath the cuticle on the back of the first segment are now seen
the two small suspensory brushes of hairs characteristic of the pupal
abdomen. The thorax and the abdomen inside the moulted larval
Fic. 14.—Fourth-instar larva of Aedes aegypti with larval cuticle (/Ct)
moulted over the thorax and most of the abdomen. The inner cuticle (pCt)
is that of the pupa.
cuticle, therefore, pertain to the future pupa. The head cuticle of the
larva has not yet been moulted, so that the larva in the fourth instar
still feeds with its own mouth parts. It breathes with its posterior
respiratory apparatus, and uses for locomotion the muscles now in the
pupal abdomen.
The condition found in the mosquito is simpler than that described
by Hinton (1958b) in Simulium. Here the fully formed pupa still
within the larval skin is active for several days before ecdysis. Its
activity is due to the fact that many of the former larval muscles, in-
cluding those of the head, remain attached by tonofibrillae to the
moulted cuticle of the larva. The pupa of Simulium is thus able to
use the larval organs, and it not only continues to feed, but spins its
own cocoon. In the mosquito there is no evidence of muscles retain-
ing their attachment on the moulted larval cuticle; the insect feeds
with the larval mouth parts until the latter are cast off at the final
pupal moult.
The larval musculature of the thorax and abdomen is said by
Thompson (1905) in Anopheles to go over into the pupa and the
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 7
adult with little alteration. The rudiments of the future wing muscles,
however, are present in the thorax during the last larval instar. Hulst
(1906), on the other hand, reports that in the larva of Culex there
takes place an extensive histolysis and histogenesis of the body mus-
culature, beginning when the larva is two-thirds grown. Some imaginal
muscles thus appear first in the larva, particularly those of the wings
and legs, prior to the advent of the pupal stage. Destruction of the
larval muscles of the abdomen, however, Hulst says, is not complete
even in a late stage of the pupa. In the Simuliidae, according to
Hinton (1959), “the indirect flight muscles and the tergal depressor
of the trochanter develop quite independently of the larval muscles
in all post-embryonic stages.”
Histological changes in the alimentary canal beginning in the larva
have been described by Samtleben (1929), by Berger (1938) for
Culex, and by Richins (1945) for Aedes. The replacement of func-
tional cells from regenerative cells in the ventriculus during larval life
is generally in other insects not a metamorphic process but the usual
procedure of replacing worn-out digestive cells by new cells. At the
fourth ecdysis to the pupa, however, Berger (1938) says, rapid
changes take place. The alimentary canal of the pupa, well illustrated
by Hurst (1890), differs from that of the larva, but is still not that
of the adult. The short pupal stomach is said by Richins to be formed
from only the posterior part of the larval stomach. According to
Samtleben no specific pupal epithelium is formed for the pupal
stomach.
Considering the precocious development of so many of the imaginal
organs, the fourth instar of the mosquito larva presents the anomalous
condition of being part larval and part pupal. In other words, the
pupal development begins within the larva long before its completion
at the pupal ecdysis. It ends with the formation of the pupal head,
mouth parts, and tail fins.
In most young insects the endocrinologists find that the larval struc-
ture is maintained by the inhibitory action of the corpus allatum hor-
mone on the adult development until the end of the larval life. The
early origin of pupal organs in the mosquito larva and the continu-
ance of their development through the larval period shows, however,
that the juvenile, or status quo, hormone does not necessarily function
as a complete inhibitor of adult development. In the mosquito it ap-
pears to be selective in its action, allowing the growth of pupal parts
that do not interfere with the normal activities of the larva, while it
maintains to the end of the larval period such parts as the head, feed-
38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
ing organs, and respiratory apparatus that are essential to the life of
the larva.
The corpora allata of the mosquito larva are described by Boden-
stein (1945) as a “corpus allatum complex” composed of two small
cellular bodies of elongate form, tapering posteriorly, attached laterally
on the aorta just behind or within the neck. Anteriorly they adhere
closely to a transverse trachea and are connected with each other by
a loose chain of cells. Each body is entered by a slender nerve from
the brain. Since the bodies contain different kinds of cells it is possible
that they include elements of the usually separate corpora cardiaca. In
higher Diptera the aorta is surrounded by a cellular ring, which is
thought to include the corpora allata and corpora cardiaca, but accord-
ing to Bodenstein the nature of the cells in the mosquito larva is not
certain. The larval complex goes over into the adult in reduced form
as two small, rounded bodies lying on the sides of the aorta.
If the fourth-instar larval mosquito behaves as other larvae have
been shown to do when experimentally given an extra dose of juvenile
hormone, it should go over into a fifth larval instar. In this case the
larva issuing from the fourth-instar cuticle would have external legs
and wings! We can only wait the results of some endocrinologist who
may make the experiment.
When at last the cuticle of the larval head is moulted, taking with
it the larval antennae and mouth parts, the corresponding pupal organs
are rapidly developed within the still-unshed larval cuticle. The recon-
struction of the mouth parts involves an extreme change from the
specialized organs of the larva to the equally but differently specialized
organs of the adult. The development of the pupal mouth parts has
been described by Thompson (1905) for Culex, and by Imms (1908)
for Anopheles.
The pupal labrum begins its growth as a fold of the epidermis at
the anterior end of the dorsal wall of the head that first extends pos-
teriorly beneath the cuticle (fig. 15 B, pLm). The fold elongates
(C, Lm) and finally turns forward and downward over the other
mouth parts. The buds of the new mandibles and maxillae are formed
directly from the epidermis retracted into the bases of the larval
organs. An early stage of their development still within the larval
cuticle is seen at C of the figure taken from Thompson. The labium
and the hypopharynx of the larva, as already shown, are greatly re-
duced and united in an area between the mouth and the hypostomium,
the two components being separated only by the opening of the
salivary duct. In Aedes the labiohypopharyngeal complex as shown by
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 39
Salem (1931) forms a distinct lobe below the mouth (fig. 15 A, Hphy,
Lb), as it does also in a tipulid larva (fig. 7 A). The rudiment of the
pupal labium within the larval labium (fig. 15 A, pLb) is said by
Imms (1908) to be a pair of hollow lobes confluent at their bases.
There is no separate rudiment of the adult hypopharynx. It is shown
by Thompson (1905) that the hypopharynx is still united with the
pupal labium (fig. 15 B) when the larval cuticle (/Ct) is moulted.
Later, as will be described, the hypopharynx of the adult female is
separated from the labium. In their final stage of development the
pupal mouth parts have become greatly lengthened and are closely
pressed together in a long curved proboscis (D).
Rudiments of the pupal tail fins are formed beneath the cuticle of
the fourth larval instar behind the respiratory apparatus, and the
primary buds of the male external genital organs appear beneath the
cuticle of the same instar behind the sternal region of the ninth
abdominal segment.
THE MATURE PUPA
The pupa at ecdysis (fig. 16 A) is fully formed in all its outer parts
and thereafter does not change externally. It is clearly a preliminary
adult with the appendages in a halfway state of completion. The pupa
can hardly represent a former active stage in the life of the mosquito,
since its mouth parts are unfitted for any kind of feeding. The pupal
thorax has already assumed the approximate size and shape of the
adult thorax. In Simuliidae, Hinton (1959) says, the definitive
thoracic structure is developed during the pharate stage of the pupa.
General external structure.—The head and thorax of the mosquito
pupa are combined in a large cephalothorax, from which projects the
slender abdomen (fig. 16 A). When at rest the pupa floats at the sur-
face of the water (fig. 13), but it does not hang from its respiratory
trumpets (as it often does in pictures). The back of the thorax and
of the two anterior abdominal segments comes against the water,
while the rest of the abdomen hangs downward as ballast. The open
ends of the respiratory trumpets project just above the surface of the
water, and two small brushes of spreading hairs on the back of the
first abdominal segment help keep the pupa suspended. The floating
position of the pupa is necessary for the future emergence of the
adult, and is maintained by bubbles of air enmeshed in the folds of
the legs and beneath the wings.
The source of the air that maintains the buoyancy of the pupa,
according to Hurst (1890), appears to be a pair of large open spiracles
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Mada Mx MxPlp
Fic. 15.—Development of pupal mouth parts and an adult leg.
A, Vertical median section through labiohypopharyngeal lobe of larva of
Aedes, with contained rudiment of pupal labium (from Salem, 1931). B, Section
of anterior part of head of Culex larva, with pupal labrum and labium forming
inside the unshed larval cuticle (from Thompson, 1905). C, Head of Culex
pupa removed from larval cuticle, with pupal mouth parts in early stage of
development (from Thompson, 1905). D, Pupal head of Aedes aegypti, lateral.
E, Same, anterior. F, Fully developed pupal mouth parts of Aedes aegypti.
G, Distal part of a pupal leg with adult leg formed within it.
Ant, antenna; Aul, aulaeum; C/p, clypeus; Hphy, hypopharynx; Li, first leg;
Lb, labium; /Ct, larval cuticle; Lm, labrum; Md, mandible; Mx, maxilla;
MxPlp, maxillary palpus; pLb, pupal labium; pLm, pupal labrum; P/p, palpus;
ae salivary duct; S/O, salivary orifice ; SoeGng, suboesophageal ganglion;
, tibia,
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 4I
on the sides of the first abdominal segment of the pupa covered by
the metathoracic wing pads. The tracheal system of the pupa, how-
ever, is so weakly developed that it would hardly seem capable of
supplying the amount of air carried by the living pupa. Manzelli
Fic. 16.—The pupa.
A, Aedes aegypti, male pupa, lateral. B, Same, terminal part of female
abdomen. C, Culex sp., thorax and base of abdomen, dorsal. D, Same, end of
female abdomen, dorsal. E, Same, apical structures of male abdomen. F, Same,
phallic organ of male pupa, ventral.
br, suspensory brush of first abdominal tergum; H, head; Ns, metanotum;
PhL, phallus; Prb, proboscis; Rect, rectum; S, sternum; 7, tergum; #f, tail fin;
Tmp, respiratory trumpet; W+2, mesothoracic wing; Ws, metathoracic wing;
I-X, abdominal segments.
(1941) described and figured the pupa as “enclosed in a sac-like struc-
ture,” which he says “has long been seen by all mosquito workers and
is usually known to them as the pupal shell.” This is a curious state-
ment, since no such structure exists. Furthermore, the “shell” is said
42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
to enclose a large air cavity, but on pressing a pupa in alcohol the air
issues as free bubbles from beneath the legs and wings.
The pupa has two features that are peculiarly its own. First are
the trumpet-shaped respiratory tubes projecting from the back of
the thorax (fig. 16C, Tmp), and second, a pair of thin, oval fan-
shaped tail fins, or paddles, borne on the end of the abdomen (A, tf).
Because it is necessary for the pupa to float with the back of its
thorax against the surface of the water, with the abdomen hanging
down, it had to discard the posterior spiracles of the larva and have
its breathing apertures forward. The trumpets are connected with
the anterior ends of the dorsal longitudinal tracheal trunks, and their
open ends project just above the surface of the water.
It is a curious fact that in species of Mansonia and Ficalbia, the
larva of which gets its respiratory air from the roots of aquatic plants,
the pupa does the same thing by means of its thoracic trumpets. The
trumpets in these species are drawn out into a pair of long horns
directed forward from the thorax. In Ficalbia hybrida each horn
ends in a pair of tapering blades (fig. 11 F), but in species of Man-
sonia each terminates with a strong, curved spine. The spines of
Mansonia richiardiu (E) are convergent and are said to be applied
close against each other as inserted into the plant. In the species shown
at C of the figure the spines are divergent, and, as in other species,
each is bordered anteriorly and posteriorly (D) by a very thin, trans-
parent, faintly striated flange. A trachea (C,E, Tra) is attached to
the base of the organ, but does not penetrate the latter. The cylindrical
basal stalk contains a wide lumen, which narrows abruptly where it
enters the spine and opens by a minute aperture at the tip. Wesenberg-
Lund (1920-21), however, says of M. richiardu that “the trachea runs
through the whole tube,” and Grossbeck (1908) figures a tube of
Culex perturbans with a trachea going through it to the tip of the
spine. It seems very unlikely, however, that the thoracic respiratory
tubes in any case contain tracheae. They are merely elongated trum-
pets, and a typical trumpet is an open funnel with the trachea opening
into its base (fig. 17 C, Tra).
As the pupa of Mansonia emerges from the larval skin, according
to Galliard (1934) as quoted by Marshall (1938), it brings the tips
of its horns together and searches for a neighboring root. Then it
violently works its way out of the anchored larval skin and at the
same time inserts its horns. When the adult is ready to emerge, the
pupa breaks away from the plant and comes to the surface where it
floats by reason of two tracheal air sacs in the thorax. The winged
mosquito thus escapes into the air in the usual manner.
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 43
It is truly remarkable that the same kind of structural adaptation
for the same purpose has occurred twice in the life of the same indi-
vidual, affecting two different organs. Furthermore, with the acquisi-
tion of a new structure designed for a new use, the insect must be
twice endowed with a new instinct for using the modified organs. It
is enough to make us wonder if we really understand the nature of
biological adaptation.
Though pupae that breathe free air ordinarily float at the surface
of the water, they can escape danger by darting around on the surface
or submerging quite as actively as the larvae by snapping movements
of the flexible and well-musculated abdomen. The large tail fins are
organs for increasing the motor efficiency of the abdomen. Func-
tionally they are comparable to the tail fan of a crayfish. The pupa
when swimming progressively on or below the surface kicks back-
ward with its abdomen and propels itself forward, but the crayfish
does just the opposite. When the pupa swims downward in the water,
however, it goes tail first, and thus maintains its floating position.
If it remains inactive it passively rises to the surface, otherwise it
swims up by abdominal movements.
The head and mouth parts ——The head of the pupa (fig. 16 A, H) is
closely attached to the lower anterior angle of the thorax, with its
true dorsal surface directed anteriorly. It retains nothing of the
structure of the larval head. The long, many-jointed antennae curve
upward and backward beneath the lower edges of the wings. The
large, black compound eyes (fig. 15 D, E) are conspicuous beneath
the cuticle, and between them the clypeal region (E, Clp) makes a
prominent bulge on the face. Posteriorly the head is produced into
a long, tapering proboscis that lies beneath the thorax with its end
upcurved behind the lower legs (fig. 16 A, Prb). The component
elements of the proboscis are closely adherent (fig. 15 E), but are
easily separated (F). Along the lower side is the relatively thick
labrum (E, Lm) which is continuous from the clypeus. Flanking the
labrum are the very delicate slender mandibles (Md), and bordering
the mandibles are the maxillae (Mx). The wide base of each maxilla
bears a free, tapering palpus (F. Plp). On the posterior (upper) side
of the proboscis is the soft, slender, tubular labium ending in a bifid
tip (F. Lb). There is no free hypopharynx in the young pupa.
As we have seen, the hypopharynx is not separated from the labium
in the larva, and the two parts go over still united into the pupa, with
the salivary duct enclosed between them. In most adult insects the
hypopharynx is an independent suboral lobe, and the salivary duct
44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
opens behind its base in front of the labium (fig. 23, SIO). The fe-
male of the mosquito and other adult Diptera possesses a free hypo-
pharyngeal stylet, but it is traversed by the salivary duct. According
to Thompson (1905) the hypopharynx of the female mosquito is
differentiated by cellular growth from the median line of the anterior
(lower) surface of the labium during the pupal stage. Since the
hypopharynx, when it becomes a free stylet, contains the salivary
duct, it would seem that in its separation from the labium it must take
a part of the labium with it. In the male the hypopharynx is not
separated from the labium, and the salivary channel remains in the
labium. Dimmock (1881) says that in the male of Culex “the hypo-
pharynx is, throughout its whole length, joined to the labium,” and
Hurst (1890) observes that it is “inseparable from the labium.”
The fact that the hypopharynx of Diptera contains the salivary
duct has given rise to the idea that this stylet is a new formation not
homologous with the hypopharynx of other insects (see Demerec,
1950, pp. 375, 376). Yet the stylet in Diptera has all the usual rela-
tions of the hypopharynx to surrounding parts, and its base forms
the floor of the preoral cibarial pump (fig. 24 E), just as in the
cockroach (fig. 23) and other generalized insects.
The cuticle of the pupal mouth parts represents the organs as
they are developed in the pupa. Inside the cuticular sheaths a renewed
growth of the epidermis produces the final adult form of the stylets,
just as the adult legs are formed within the cuticle of the pupal legs
(figs. 15 G, 17 A). The segmented maxillary palpus of the adult, for
example, is clearly seen inside the simple palpal sheath of the pupa
(fig. 15 F, Plp), and within the end of the pupal labium (Lb) are
visible the labellar lobes of the adult.
The thorax.—The large thorax of the pupa is indistinctly seg-
mented, but it bears the legs and wings, and carries on its back the
respiratory trumpets (fig. 16 A). The legs and the wings of the pupa
have been taken over directly from the larva. The legs have increased
in length and their joints are more distinct (fig. 17 A), but they are
closely folded in loops against the sides of the thorax as in the larva.
The mesothoracic wings are much larger and more winglike in shape;
the hind wings are still triangular lobes of the metanotum. Within
the cuticle of the pupal appendages are plainly seen the developing
appendages of the adult. The venation of the forewing is already
laid out (D). Within the hindwing may be seen the club-shaped
halter (E, Hit), which, whatever may be its evolutionary history, is
not formed in ontogeny by a gradual modification of the wing.
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 45
The abdomen.—The abdomen of the pupa (fig. 16 A) resembles
that of the larva except for the lack of the respiratory apparatus, the
presence of the tail fins (tf), and the reduction of the tenth segment
(X) to a small anus-bearing lobe. The dorsum of the first segment
has a special pattern of sclerotization (figs. 16 C, 17 F) and bears the
two brushes of spreading hairs that keep the base of the abdomen sus-
pended at the surface of the water. It is suggested by Hurst (1890)
that these brushes, besides serving as suspensoria, probably also are
Fic. 17.—Pupal characters and an adult leg of Aedes aegypti.
A, Left third leg of pupa with adult leg formed inside the cuticle. B, Same
leg of adult on emergence, same magnification. C, Right respiratory trumpet,
mesal. D, Left mesothoracic wing with adult wing inside the cuticle. E, Left
metathoracic wing with halter forming inside it. F, Metanotum and first two ab-
dominal segments. G, End of abdomen with tail fins.
Hilt, halter; Ns, metanotum; Tra, trachea; Ws, metathoracic wing; I, IJ, first
and second abdominal segments.
sensory organs responding to vibrations in the water. The pupa be-
comes immediately active on any disturbance of the water, even to a
tap on the containing vessel.
The pupal tail fins, as usually drawn in illustration, appear to be
attached to the end of the eighth abdominal segment (fig. 17 G). If
they are pulled away from the eighth segment, however, they are
seen to be carried by a transverse dorsal bar entirely separated from
the tergum of the eighth segment (fig. 16 D,E, XT), which, in fact,
is the tergum of the ninth segment. On it is supported also the small
tenth segment (X). In the male pupa (E) the ninth segment is a
46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
complete narrow annulus (JXT, XS) as in the adult male (fig. 27 B),
and below the small tenth segment projects a pair of large lobes (fig.
16 E, PhL) ona common base arising from the sternal arc of the ninth
segment (F). These lobes are the genital appendages of the male as
far as they are developed in the pupa. Male and female pupae, there-
fore, can be distinguished by the presence (A) or absence (B) of the
genital lobes (PHL), though in the male the lobes might be mistaken
for the tenth segment, since the latter is mostly concealed above them
(Ay Xx).
THE PUPAL METAMORPHOSIS
The pupal life of most mosquitoes is very short, two or three days
or less, though with some species it is much longer. During this time
the contour of the adult is modeled by new growth of the epidermis
beneath the pupal cuticle, while the mouth parts, wings, halteres, and
legs take on the adult structure within their pupal sheaths. At the
same time reconstruction of internal organs takes place inside the
body. The degree of reconstruction necessary to change the larval
organs into those of the adult, however, is much less in the mosquito
than in many other insects, especially in the higher Diptera.
The mosquito pupa breaks with the tradition that a pupa is a “rest-
ing stage” in the life of the insect. When an ordinary pupa is broken
open it is seen to be full of a creamy mass of soft material resulting
from the disintegration of the larval tissues. The inside of a mosquito
pupa is as clean as that of the larva or the adult, and its organs appear
to be intact. Whatever reorganization is going on takes place mostly
inside the alimentary canal and the refuse is not thrown into the body
cavity.
The abdominal muscles are so well preserved that the pupa is an
extremely active stage of the mosquito, and the thoracic muscles are
so well developed that the pupa might be expected to fly if its wings
were more mature. As already noted, Hulst (1906) has described the
process of muscle histolysis and histogenesis as beginning in the larva,
but he is not explicit as to what larval muscles are destroyed or when
the imaginal musculature is completed. In Culex, according to Hurst
(1890), the muscles of the pupa are those of the imago; the principal
muscles are present in the young pupa, but they increase greatly in
size. A casual examination of the abdominal musculature in the larva,
pupa, and adult shows little difference between the stages, except for
the greater size of imaginal muscles. However, we need a more de-
tailed comparative study of the muscle pattern and more information
on the replacement of individual muscles.
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 47
The larval head musculature appears to be largely replaced by an
imaginal musculature. According to Thompson (1905) there is an
extensive histolysis of the larval head muscles, accompanied by a
regeneration of muscles appropriate to the adult, which takes place
in the eighth to tenth hour of pupal life.
The pupal tracheal system is weakly developed and is difficult to
see in dissections. According to Hurst (1890) tracheae go from the
base of each thoracic trumpet to various parts of the head and body,
and a transverse trunk connects the two trumpets. A pair of longi-
tudinal trunks runs back to the rear end of the body, giving off
branches to the internal organs and to the site of each spiracle. Only
the spiracles of the first abdominal segment remain open.
In his study of the heart of Anopheles quadrimaculatus, Jones
(1954) reports that no evidence was found that the heart is “de-
stroyed, reconstructed, or otherwise drastically modified during meta-
morphosis.” In young pupae, according to Jones, the heart beats in a
forward direction as in the larva, but later it may cease beating for
prolonged periods of time. Circulation of the blood, therefore, ap-
pears to be unessential for the regenerative changes taking place in
the pupa.
The alimentary canal of a young pupa, as described and illustrated
by Hurst (1890) in Culex, might be supposed to be a functional organ
if the pupa could feed. It more resembles the digestive tract of the
larva than that of the adult, but since the adult feeds on a very dif-
ferent kind of food from that of the larva, the alimentary canal under-
goes a complete reconstruction in the pupa, details of which have been
described by Hurst (1890), Thompson (1905), Samtleben (1929),
and Richins (1938). The oesophagus is least affected insofar as its
epithelium goes over intact from larva to adult, but the larval pharynx
is lost, and an enlargement in the back of the head forms the post-
cerebral sucking pump of the adult. In the thorax the dorsal and
ventral diverticula of the adult grow out from the oesophageal wall.
The larval gastric caeca are absorbed and not replaced in the adult.
The larval epithelium of the stomach, according to Richins, de-
generates completely and is cast off into the stomach lumen, as a new
epithelium is formed by permanent regenerative cells. Transforma-
tion in the proctodaeum is brought about partly by histolysis and
histogenesis of the epithelium and partly by regrowth. The five
Malpighian tubules of the larva go over into the adult without change.
In the rectal sac of the pupa are formed six invaginations of the wall
that become the rectal papillae of the adult. The salivary glands of
48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
the larva degenerate and each is replaced by three slender tubules
generated from cells in the neck of the larval gland.
The central nervous system undergoes little change in the pupa
other than growth and union of some of the ganglia. The first ab-
dominal ganglion of the larva is drawn into the thorax, where it
fuses with the metathoracic ganglion, and later the four ganglia now in
the thorax condense into a single mass. The last abdominal ganglion
of the larva unites with the ganglion of the seventh segment. In the
adult mosquito, therefore, there are only six separate ganglia in the
abdomen (fig. 30 C). In the head, as described by Woolley (1943)
for Aedes, the brain and the optic lobes grow rapidly by peripheral
formation of new cells. The circumoesophageal connectives shorten
and the suboesophageal ganglion unites with the brain around the
oesophagus.
Though the visible changes that take place in the nervous system
are slight, there must be a considerable reorganization of the internal
structure. The behavior and instincts of the adult mosquito are en-
tirely different from those of the larva. Since the activities of the in-
sect resulting from sensory stimuli are determined by established
neuromuscular pathways and synapses in the central nervous system,
the system that serves the larva must be entirely reorganized into one
appropriate for the activities of the adult. Of this, however, we know
little or nothing in any insect.
Ill. THE ADULT
The adult mosquito fully formed within the pupa has now only to
cast off its pupal mold to gain its freedom in the garb of a mature
winged insect. But this is not easily done since the confined mosquito
has no instruments for cutting or breaking the pupal cuticle. More-
over, the wings, legs, antennae, and mouth parts are enclosed in tight-
fitting sheaths, from which they must be slowly extracted. However,
much as we might wish that the mosquito should remain a prisoner in
the pupal skin, nature has made provision for its liberation.
As noted by several observers, the first evidence that the adult is
about to emerge is the appearance of a film of air beneath the pupal
cuticle on the back of the thorax. A slight retraction of the adult ap-
parently breaks the connections of the pupal trumpets with the tracheal
system and thus allows air to escape beneath the cuticle. Usually a
short piece of trachea remains attached to the base of each trumpet.
According to Marshall and Staley (1932) rhythmical movements now
begin in the sucking pump of the adult which draw the air forward,
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 49
forming a bubble at the base of the proboscis. This air is then
pumped into the stomach as a long narrow bubble that extends back to
the fourth abdominal segment. Pressure by the distended abdomen
now pushes the thorax forward until it ruptures the pupal cuticle in
a median slit along the back from the neck to the end of the meso-
thorax. Outside air then enters the cleft and is rapidly swallowed,
going back in the stomach as far as the sixth abdominal segment and
greatly distending the abdomen. Knab (1909), in describing the role
of air in the ecdysis of insects, says of the mosquito that on emergence
from the pupa it is distended with air far beyond its natural size, the
integument being stretched to its utmost. According to the writer’s
observations on emerging mosquitoes the degree of distention is highly
variable, even with individuals of the same species.
Pupae of Aedes aegypti, before the adult ecdysis, are observed to
have the abdomen extended straight back from the thorax, and during
the emergence it is held, or floats, in this position with the tail fins
against the water surface (fig. 18 A). When the pupal cuticle splits on
the back of the thorax, the thorax of the adult bulges out and pushes
apart the lips of the cleft. This produces a transverse split over the
back of the pupal head, so that the pupal skin can now be widely
opened anteriorly (B) to allow the egress of the adult. At the same
time the cuticle on top of the pupal head between the eyes breaks
out and folds forward as a free flap beyond the antennal bases (A,B).
Behind the antennae the anterior tentorial arms project internally as
a pair of slender tapering rods (B).
Inasmuch as the legs of the adult, as well as the wings and mouth
parts, are enclosed in tight-fitting pupal sheaths, the mosquito cannot
use its appendages for freeing itself. Yet, when the head and thorax
are free, the abdomen follows and the entire adult slowly rises verti-
cally from the pupal skin as if pushed out from below. The legs and
wings are at first closely pressed against the body, but as the legs
are freed they at once become active, and appear to be reaching for
the surface of the water. The mosquito seems to know instinctively
that now and henceforth it must support itself on its legs. It will be
noted that the legs of the emerged adult are greatly longer than their
pupal sheaths ; the hindleg of an Aedes (fig. 17 B), for example, may
lengthen to two and a half times the length of the corresponding folded
leg of the pupa (A). When the end of the abdomen and the wings
are finally out of the pupal thorax and the legs are all free, the new
insect confidently steps out onto the surface of the water and calmly
walks away from the discarded pupal skin. It may come to rest on
50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
some nearby floating object (as a bit of cardboard in the aquarium),
but usually in a very short time it is able to fly, and immediately is
gone. Sometimes, however, mosquitoes in culture appear to have
much difficulty in finally extracting their legs; often they fall over
on the surface of the water, and some perish in this position with
their tarsi still held in the pupal sheaths. It is probable that in such
cases the larvae were not properly nourished.
ic. 18—An adult female of Aedes aegypti emerging from the submerged
pupal cuticle; and the open thorax of the discarded pupal cuticle of Anopheles
quadrimaculatus.
A remarkable thing about the mosquito is that, after its whole
previous life spent in the water, on emergence from the pupa it is
at once at home in the air. Without a flutter of the wings or any
practice trial, it makes a perfect takeoff, flight, and distant landing.
During the pupal stage, therefore, the mosquito has not only been
equipped with a complete mechanical apparatus of flight, but in its
nervous system a mechanism of control has been fully elaborated.
Compare this with the difficulty the young human has in learning even
to walk, but of course his ancestors did not always walk upright on
two legs.
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—-SNODGRASS 51
The newly emerged mosquito (fig. 19) is really an elegant insect as
it stands high on its long slender legs, the abdomen held straight back
beneath the neatly folded wings, and the long proboscis extended
from the head. The sexes of most species are readily distinguished at
once by the antennae, those of the female having usually circles of
short hairs, those of the male being large spreading plumes.
Fic. 19.—Aedes aegypti, adult male.
THE HEAD
The head of an adult mosquito has little likeness to that of the larva.
It is an oval capsule (fig. 20D) with the facial region carrying the
antennae directed forward, and the long proboscis extended from its
lower end. The sides are largely occupied by the great compound
eyes, which almost meet dorsally and ventrally (A,B, E). The large
bases of the antennae arise so close together on the face that the
frons is reduced to a narrow verticle bar between them (A, Fr),
but its lower end forks into diverging arms that support the clypeus
(Clip). A median coronal sulcus (cs) on the vertex extends down-
ward on the face through the frons. The strongly convex clypeus
(A,C, Clp) forms a prominent lobe just above the base of the
proboscis. The undersurface of the head (B) resembles that of the
52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
larva in that it is completely closed from the occipital foramen to the
base of the proboscis. The head is attached to the thorax by a slender
membranous neck and is supported by a pair of lateral cervical sclerites
(D,E). The head of the male is similar to that of the female, but
is a little smaller. The internal head skeleton consists of a pair of
simple tentorial arms extending from anterior pits above the lateral
angles of the clypeus (A, at) to posterior pits (B, pt) on the ventral
margin of the occipital foramen.
From the front of the face arise the long antennae (fig. 20 D,E).
The hairy flagellum of each organ is borne on a large globose base
(A, Pdc), which is the pedicel, or so-called torus, but when the pedicel
is removed (right) it is seen to be itself supported on a narrow ring
(Scp) that represents the usually much longer scape of other insects.
The slender shaft of the flagellum is divided into 14 sections (errone-
ously called “segments”), 13 of which carry each a whorl of hairs.
In general the sexes are readily distinguished by the number and
length of the flagellar hairs, which in the male (fig. 22 A) give the
antennae a plumose appearance in contrast to the short-haired female
antennae (D,E). The two types, however, intergrade, females of
some species having bushy antennae, and some males short-haired
antennae. In the female the hairs arise from clear areas near the bases
of the flagellar units (B) ; in the male (C) they are borne on promi-
nent, darkly sclerotized, subapical expansions of the units. Tulloch
and Shapiro (1951) have shown from electron microscope studies that
the flagellar hairs are armed with rows of minute teeth; in Culex
quinquefasciatus they estimate there are at least 16 rows along each
hair. These writers, however, are in error where they say the hairs
“arise at the junctions of the flagellar segments.”
The large globose pedicel of the antenna in each sex contains a
highly developed sclopophorous sense organ, present also, though
usually much smaller, in the antennal pedicel of most insects. The
organ was first described in Culex as an auditory organ by Johnston
(1855), who did not at all understand the nature of the structure in
the pedicel, but it has since been known as Johnston's organ. Sub-
sequently Child (1894) made good histological studies of the organ
in various insects, including the mosquito, and his illustrations are
now famliar in most entomological texts. A more recent comparative
study of the organ in Culex, Aedes, and Anopheles is given by Risler
(1955). The component sensory elements in the pedicel are attached
to a plate or prongs on the base of the flagellum, and thus evidently
register movements of the flagellum.
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 53
7 A
i A4YTT Ay y
we
SMM \|
Lm re Hphy Md Mx
1
Fic, 20.—Head and mouth parts of an adult female mosquito, Aedes aegypti
except G.
A, Head and base of proboscis, anterior, B, Same, posterior. C, Clypeus and
base of proboscis, lateral. D, Head, lateral. E, Same, with mouth parts
separated. F, End of labium, dorsal. G, Cross section of proboscis of Anopheles
(from Vogel, 1921). H, Distal ends of mouth part stylets. I, Basal parts of
maxilla. J, Proximal parts of right maxilla and labium, posterior.
at, anterior tentorial pit; Cd, cardo; Clp, clypeus; cs, median cranial sulcus;
E, compound eye; fc, food canal; For, occipital foramen; Fr, frons; Hphy,
hypopharynx; Hst, hypostome; Lb, labium; LObI/, labellum; Le, lacinia; LG,
labial gutter; Lig, ligula; Lm, labrum; mel, muscle; Md,Mds, mandible,
mandibles; Mx,M-xae, maxilla, maxillae; MaPlp, maxillary palpus; Nv, nerve;
Pdc, antennal pedicel; P/p, palpus; Prb, proboscis; pt, posterior tentorail pit;
sc, salivary canal; Scp, antennal scape; St, stipes; Thc, theca; Tnt, anterior
tentorial arm; Tra, trachea; Vx, vertex.
54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
While it is probable that the organ of Johnston in the antenna of
most insects registers the movements of the flagellum, the elaborate
experimental work of Roth (1948) leaves no doubt that the highly
developed organ in the male mosquito is responsive to the effect of
sound waves on the flagellum. This, of course, does not imply that
the mosquito has an auditory “sense” ; mechanical reaction to stimuli
is all that we can attribute to the insects. Male mosquitoes are at-
tracted to the females in flight by the tone produced by their wings.
Roth showed that males with intact antennae, when subjected to the
sound of a tuning fork at 480 vibrations a second held behind a
suspended piece of cloth, fly to the source of the sound where they |
exhibit typical mating activities though no females are present. Even
after complete removal of the flagellar hairs, males still respond to
more intense sounds apparently by vibrations of the shaft alone, but on
complete removal of the flagella they give no reaction. Roth’s tests
were made particularly on Aedes, but males of other genera were
found to react similarly. Females of Aedes aegypti gave no evidence
of being attracted to sounds, “though they may give shock-reaction
to certain intensities.”
Further experimental work of Roth (1951) on females of Aedes
seems to show that the antennae function as directional distance
thermoreceptors and probably also as chemoreceptors. Females de-
prived of their antennae are unable to locate a host from a distance.
The antennae and the palpi are said to be the chief organs responding
to stimuli that induce probing by the proboscis. The receptor organs
of the antennae, however, are not described, but along the shaft of
the female antennae (fig. 22 B) are numerous hairs, and on the male
antenna (C) a ring of very short hairs encircles the distal end of each
flagellar section. The antennae of insects in general are known to be
the principal seat of chemoreception.
The compound eyes of the mosquito are so large that they almost
encircle the head. Sato (1950, 1953a, 1953b) reports that by actual
count there are from 440 to 462 facets in the eye of a male Culex
pipiens, and 503 to 566 in the female; and that in Aedes japonicus the
male eye contains 440 to 462 facets, the female eye 504 to 527. The
surface area of the eye in each genus is larger in the female than in
the male. The internal structure of the compound eye in Culex is
described by Constantineanu (1930) and by Sato (1950).
An extensive experimental study of the visual responses of flying
mosquitoes made by Kennedy (1939) on unfed females of Aedes
aegypti shows that the mosquitoes react negatively to light, and are
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 55
attracted to dark objects. Experimentally they orient toward black
stripes on a white background, and continue to do so when the stripes
are rotated about them. When confronted by two black stripes, they
face one or the other and not the intervening space. In a wind tunnel
freely flying mosquitoes move against the current.
THE ORGANS OF FEEDING
The feeding organs of the adult mosquito include the proboscis
and two sucking pumps, One of the latter is a preoral cibarial pump
beneath the clypeus, the other is a pharyngeal pump, being a part of
the alimentary canal behind the brain in the back of the head. In
describing the feeding organs of the adult it will be better to take
the female first, because in most mosquitoes she is the biting and
bloodsucking member of the species and has the mouth parts fully
developed. In the nectar-feeding male some of the parts are much
reduced or absent.
The proboscis—The slender, rodlike proboscis in the female
mosquito is usually composed of all the mouth parts possessed by in-
sects that feed on solid food, namely, a labrum, a pair of mandibles, a
hypopharynx, a pair of maxillae, and a labium, but the parts are all
structurally modified in adaptation to the mosquito’s way of feeding.
The relation of the parts in the undisturbed proboscis is best seen
in a cross section (fig. 20G). In the deeply channeled upper side of
the labium (LD) are enclosed the labrum (Lm), the mandibles (Md),
the hypopharynx (Hphy), and the maxillae (Mx). The labrum itself
is practically an inverted tube, since its margins are curved downward
and may overlap. The enclosed labral canal (fc) is the food conduit.
The hypopharynx contains the salivary canal (sc). By careful
manipulation with a dissecting needle all these parts can be separated
as shown at E.
The labrum (fig. 20 H, Lm) is the thickest and the strongest of the
stylets. It is movable by muscles from the clypeus attached on its
base (fig. 24D), but the muscles simply elevate and depress the
labrum, which is firmly hinged on the clypeus. The term “labrum-
epipharynx” often applied to the labrum is quite unnecessary, since in
its general form the labrum is a flat lobe of the head and therefore
has an upper and lower surface. In the mosquito the decurvature of
the lateral parts converts the labrum into a tube through which the
ingested liquid food is drawn up by the sucking apparatus at its
base. At the sharp-pointed distal end (fig. 20 H, Lm) the walls of the
channel diverge to make an opening like that of a hypodermic needle.
56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
The mandibles are the slenderest of the stylets, but they vary some-
what in thickness and shape in different species. In Aedes here
illustrated (fig. 20 H, Md) each is slightly enlarged toward the taper-
ing distal end. The base of each mandible is movably connected with
the lower part of the cranial wall by a small suspensory sclerite, and
a slender muscle from the tentorium is inserted on the mandibular
base. The mandibles are thus retractile for a short distance, and, when
retracted, their withdrawn tips give free entrance to liquid into the
open end of the labral food canal. Protraction results from the elas-
ticity of the suspensory mechanism on relaxation of the muscles.
The single, median hypopharynx, present as an independent stylet
only in the female, is a simple, flattened rod (fig. 20H, Hphy)
traversed by the salivary outlet canal (sc), which opens on its acute
tip. The hypopharynx is not individually movable; its anterior wall is
continued basally into the floor of the cibarial pump.
The maxillae are less reduced than the other mouth parts, and are
well equipped with muscles. The principal part of each maxilla (fig.
201) is a long, flattened, sharp-pointed blade (Lc) armed with re-
curved teeth near the end of its outer margin (H, Mx). From the
base of the blade projects a usually short four-segmented palpus (1,
Pip). The maxillary blade has been regarded as the galea by some
writers (Robinson, 1939; Snodgrass, 1944), but it is more reasonably
interpreted by Schiemenz (1957) as the lacinia, which is usually the
operative part of a generalized maxilla. From its base a long, strongly
sclerotized, apodemelike rod extends backward in the head and gives
attachment to muscles (J). This rod is evidently the stipes, or more
probably stipes (St) and cardo (Cd) combined, sunk into the head,
since in some related flies, such as Phlebotomus (fig. 22 G), it is
superficial on the back of the head and articulates on the cranial
margin.
The maxillary musculature of Aedes (fig. 20]J) includes a long
retractor arising on the head wall close to the posterior end of the
tentorial arm (Tnt) inserted on the distal end of the stipes, and two
protractors attached proximally on the stipito-cardinal rod. One of
these muscles arises on the tentorium, the other, very curiously, on the
base of the labium. A lateral muscle from the tentorium and a short
mesal muscle both attached on the base of the lacinia are regarded
by Schiemenz (1957) in Theobaldia [Culiseta] as an abductor and
adductor respectively of the maxilla. A short muscle from the stipes
is inserted on the base of the palpus, and each palpal segment contains
a small muscle inserted on the segment distal to it.
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—-SNODGRASS 57
The long, gutterlike labium of the mosquito is the so-called pre-
mentum of a generalized labium, the usual basal part of the labium
being absent, though a small postmental sclerite may be present in
other Nematocera (fig. 22 G, Pmt). The prementum in Diptera is
known as the theca because it ensheaths the other mouth parts.
Apically it bears two small movable lobes, the Jabella (fig. 20 F, Lol),
and ends between them in a slender median projection, or ligula (Lig).
The labella appear to be two-segmented, and evidently represent the
labial palpi because each is provided with an abductor and an adductor
muscle from the prementum. The only muscles attached on the base
of the labium are the two already noted that arise on the maxillary
stipites (J) and probably act as protractors of the maxilla, since the
labium is firmly fixed to the head.
The styliform mouth parts within the labial theca adhere to one
another in a compact fascicle. They are usually said to be held to-
gether by an oil liquid, but Bhatia and Wattal (1957) have described
rings issuing from the margins of the labrum that surround the
hypopharynx, mandibles, and maxillae and bind these stylets to the
labrum. However, no other investigator has reported the presence
of any such structures, and the writer has failed to see them in
Aedes, Culex, or Anopheles. The incurved lower edges of the labrum
enclose only the food canal.
When the female mosquito is about to take a meal of blood, she
places the tip of the proboscis against the skin of the victim (fig.
21 A), closely holding the end of the stylet fascicle between the labial
labella. The movable maxillary stylets are the active piercing organs.
Acting alternately, first one is protracted and holds its position in
the flesh by means of its recurved teeth, then the other is forced in
beyond the first and takes a deeper hold. The labrum, mandibles, and
hypopharynx penetrate along with the maxillae. The retractor muscles
of the maxillae, instead of pulling the stylets out of the wound, where
they are held by the maxillary teeth, bring the head down closer to the
feeding surface. The labrum, still holding the stylet fascicle between
the labella, is thus forced to bend backward (B) and the bend becomes
greater the deeper the stylets penetrate (C). When finally the stylets
pierce and enter a small blood vessel, or let out a pool of blood, the
mandibles are drawn back from the end of the labrum to allow the
blood to enter the food canal in response to the suction of the cibarial
pump. Saliva discharged from the hypopharynx in some species serves
to prevent coagulation of the blood. A more detailed account of the
feeding act and of accompanying movements by the maxillary palpi is
58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
given by Robinson (1939). After feeding, the maxillary stylets are
retracted, the female braces herself against the skin of the victim with
her legs, and forcibly pulls out the fascicle of stylets, which again
is ensheathed in the straightened labium.
In discussing the feeding of mosquitoes, we must not overlook
the fact that not all females are bloodsuckers. A prominent exception
to the rule are species of Toxorhynchites, in which both sexes feed on
Fic. 21.—Successive stages in the penetration of the stylets of a female
mosquito feeding on blood. (B, C, from Gordon and Lumsden, 1939, with neck
plate added.)
nectar or other plant juices. In this genus (fig. 22 D) the proboscis is
very long, tapering, and strongly decurved. The maxillary palpi pro-
jecting from the base of the proboscis are long and four-segmented.
The laciniae by contrast are weak and taper into filaments reaching
only a little beyond the end of the first palpal segment ; evidently they
play no part in feeding. A slender labrum extends to the tip of the
proboscis, but mandibles appear to be absent.
Then there are species of Malaya (=Harpagomyia) that get their
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 59
food from ants. In these the proboscis is curved forward at its lower
end (fig. 22 E) ; the distal part is thickened and armed with long hairs.
The elongate labella terminate with a pair of small transparent lobes.
The species of Malaya are minute mosquitoes, much smaller than
ordinary ants. As described by Jacobson (1911) they sit on branches
AE
\
Wi
\y
YY
l
Fic. 22.—Various types of antennae, palpi, proboscides, and labia of adult
mosquitoes.
A, Culex sp., head of male. B, Same, part of female antenna. C, Same, part of
male antenna. D, Toxorhynchites rutilus, head and proboscis of female. E,
Malaya jacobsom, head and proboscis of female. F, Culex, distal end of male
labium, showing salivary duct. G, Phlebotomus verrucarum (Ceratopogonidae),
head and proboscis, posterior.
Cd, cardo; For, occipital foramen; LObI, labellum; Pmt, postmentum; S/Dct,
salivary duct; St, stipes; Thc, theca (prementum).
inhabited by ants, and when an ant runs between the legs of one of
them the mosquito thrusts the end of its proboscis between the open
mandibles of the ant, which accommodatingly gives up its dinner to
the mosquito. The proboscis of the adult Malaya lacks mandibles and
maxillae. According to de Meijere (1911) these members are present
60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
in the pupa, but the imaginal parts formed inside of them are short
and disappear.
The mouth parts of the male mosquito are much simplified by the
great reduction of the mandibular and maxillary stylets and the entire
absence of a hypopharyngeal stylet. The male proboscis, therefore,
consists principally of only the labrum and the labium, but the maxil-
lary palpi are usually highly developed and may be much longer than
the proboscis (fig. 22 A). Mandibular stylets when present are seldom
longer than half the length of the proboscis and are usually much
shorter. Marshall and Staley (1935) report that they are present in
all genera examined except “Aedes and Ochlerotatus.” These writers
found maxillary stylets to be present in representatives of all genera
examined, but the length is highly variable, even in species of the same
genus. The labium is a deep trough, as in the female, and ends with a
tapering median ligular lobe between the labella (fig. 22 F). It will
be recalled that the hypopharynx of the male mosquito is not separated
from the labium, as in the female. The hypopharynx thus retains in
the adult male the larval condition of union with the labium. The
male “labium” is, therefore, really a labiohypopharynx. The hypo-
pharyngeal component in Anopheles is identified by Vizzi (1953) as
a sclerotic plate on the floor of the labial gutter. In sectional figures
he shows the salivary canal in an apparent median thickening of the
plate. In Culex (fig. 22 F) the salivary duct (S7Dct) is a thread-
like tube that traverses internally the floor of the labial gutter and
opens on the tip of the ligula, but it appears to be free in the labial
lumen,
The cibarial pump.—The structure here termed the cibarial pump
lies just beneath the clypeus at the base of the proboscis, and is the
organ that sucks the liquid food up through the canal of the labrum.
The same pump is present in all Diptera and is the sucking apparatus
of other liquid-feeding insects, such as the Hemiptera. It has long
been erroneously called the “pharynx,” and even some recent writers
continue to call it such on the pretext of not wishing to confuse stu-
dents. It is possible, however, that some students might prefer to
know the facts. The organ in question is entirely outside the mouth,
as no true pharynx could be, but admittedly it is difficult to understand
its anatomical status in the mosquito. We must therefore turn to
some other more generalized insect for light on the nature of the
preoral sucking organ, and for this purpose the cockroach will be par-
ticularly illuminating.
In a vertical lengthwise section of the head of a cockroach (fig.
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 61
23) the mouth (Vth) is seen to lie beneath the upper end of the clypeal
region (C/p) of the cranial wall. Below the mouth projects the large
tonguelike lobe commonly termed the hypopharynx (Hphy), which
has a long base sloping up to the mouth. On this basal part of the
hypopharynx is a depression that forms the floor of a pocketlike space
(Cb) in front of the mouth beneath the inner wall of the clypeus. The
masticated food passed back from the mandibles is stored in this
pocket before it is swallowed. The pocket, therefore, is named the
Th /
Lib S10
Fic. 23.—Vertical section through the left side of the head of a cockroach,
exposing the preoral cavity.
Cb, cibarium; Clp, clypeus; Fr, frons; hf, hypopharyngeal fulcrum; Hphy,
hypopharynx; Lb, labium; Lm, labrum; Mth, mouth; Phy, pharynx; PrC,
preoral cavity; S/Dct, salivary duct; S/O, salivary orifice; y, oral suspensory
arm of hypopharynx.
5a,5b, dilator muscles of cibarium; 6,7, frontal muscles of stomodaeum; 13,
adductor of hypopharynx; 14, abductor of hypopharynx.
cibarium (food container). On its dorsal wall are attached strong
muscles (5a, 5b) from the clypeus. The hypopharynx can be pressed
against the inner clypeal wall by muscles (13) attached to arms (y)
from its base. The cibarium then becomes a closed chamber that can
be dilated by the clypeal muscles, and probably serves as a sucking
organ when the cockroach drinks liquids. In insects that habitually
feed on liquid food, the cibarium becomes elaborated to form a
permanent sucking pump.
62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
When we turn now to the mosquito, a section of the head (fig.
24 A) will show beneath the bulging clypeus (C/p) a small elongate
capsule (COP), which is the primary sucking pump. The basinlike
lower wall is strongly sclerotized and, in the female, is directly con-
Fic. 24.—The sucking apparatus of an adult mosquito.
A, Diagrammatic section of female head. B, Culex sp., section of pharyngeal
pump (from Thompson, 1905). C, Aedes aegypti, pharyngeal pump exposed
by removal of anterior head wall. D, Same, muscles of labrum. E, Same,
cibarial and pharyngeal pumps, left, cibarial pump opened to show lumen and
dilator muscles.
at, anterior tentorial pit; Br, brain; CbP, cibarial pump; For, occipital
foramen; FrG, frontal ganglion; lvr, labral lever; MaPIlp, maxillary palpus;
Oe, oesophagus; PhP, pharyngeal pump; sc, salivary canal; S/P, salivary pump;
SoeG, suboesophageal ganglion; Tt, tentorial arm; y, oral arm of cibarial pump.
Other lettering as on figure 23.
tinuous with the supper surface of the hypopharynx (Hphy). The
dorsal wall (E) is the so-called epipharyngeal surface from the
labrum (Lim) to the mouth (Mth), and is thin and flexible. On it is
attached a great mass of dilator muscles (5) from the clypeus. The
NO. 8 ANATOMICAL LIFE OF TITE MOSQUITO—SNODGRASS 63
inner end of the organ opens through the mouth (Mth) into the
narrow first part of the alimentary canal, and at each side of the mouth
projects a small process (y) on which are attached two antagonistic
muscles (A, 13, 14), as in the cockroach. All these features so
closely duplicate those of the cibarium in the cockroach as to leave
no doubt that the preoral sucking pump of the mosquito is the ciba-
rium. In the mosquito, however, the organ has been made into a much
more efficient sucking apparatus than that of the cockroach by the
union of the edges of its lower hypopharyngeal wall with the epi-
pharyngeal wall, thereby converting the lumen into a closed cavity.
The clypeal muscles on contraction lift the flexible dorsal wall and
expand the lumen, drawing in the liquid food from the canal of the
labrum. On relaxation of the muscles the dorsal wall snaps back by
its own elasticity and drives the liquid from the pump back through
the mouth.
On the epipharyngeal wall of the cibarial pump are small spines
and papillae of various kinds, some of which are sense organs. A com-
parative study of these structures and an armature of ventral teeth at
the mouth entrance has been made by Sinton and Covell (1927), and
Chwatt and Major (1945) in the anophelines, and by Barraud and
Covell (1928) in anopheline and culicine species. The epipharyngeal
sense organs are described by Day (1954).
The pharyngeal pump.—From the mouth at the inner end of the
cibarial pump the stomodaeal section of the alimentary canal begins
as a narrow tube (fig. 24 A,E) that curves upward and backward in
the head, going between the brain (A, Br) and the suboesophageal
ganglion (SoeG). Behind the brain it expands into a large, bulblike
structure, which is the pharyngeal pump (PAP). The walls of the
organ when relaxed are deeply concave above and on each side, as seen
in cross section at B. Into the concavity of the dorsal wall is inserted
a pair of large muscles (A,B,E, S) from the dorsal wall of the head
behind the brain, and into each lateral concavity a large flat muscle
(zr) from the side of the cranium. Contraction of the muscles dilates
the lumen of the pump; on their relaxation the walls spring together
again by their own elasticity. From the rear end of the pump, the
narrow oesophagus (Oe) proceeds through the neck into the thorax.
A cibarial and a pharyngeal pump like those of the mosquito are
common to bloodsucking nematocerous flies. Presumably the two
pumps work in alternate phases to keep the ingested blood flowing
freely back into the stomach. In the nectar-feeding male mosquito
the sucking apparatus is less strongly developed than in the female.
64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
THE THORAX
The thorax of a winged insect may truly be said to be the most re-
markable anatomical mechanism developed anywhere in the animal
kingdom. It is remarkable both for its efficiency as a flight mechanism
and for its structural simplicity. In insects with two pairs of wings
the two wing-bearing segments have essentially the same structure,
and are equipped with duplicating sets of muscles. In the Diptera,
however, in which the flight function has been taken over entirely by
the first pair of wings, the mesothoracic wing muscles have to do the
work of the muscles of both winged segments in four-winged insects.
Consequently, the mesothorax of the flies has been greatly enlarged
and the metathorax much reduced. The knobbed stalks known as
halteres borne on the metathorax are undoubtedly reduced wings,
since, as seen in the mosquito pupa (fig. 17 E), they are developed in
flat wing lobes of the metanotum. They are still important accessories
of flight, being vibratory organs for maintaining the equilibrium of
the flying insect, but their musculature is very simple, and the usual
wing musculature of the segment has been eliminated.
In the adult mosquito (fig. 25) the mesothorax appears as a great
wedge inserted between the narrow prothorax and metathorax. It
alone retains the structure typical of a thoracic segment. Two princi-
pal plates, an anterior notum (AN,) and a posterior postnotum
(PNz), cover almost the entire dorsum of the thorax. The strongly
convex postnotum, furthermore, is deeply infolded posteriorly be-
neath the narrow metanotum (Ns) and extends into the first ab-
dominal segment as a bilobed phragma (fig. 27D, Ph). A narrow
paranotal fold (pf) borders the edge of the notum between the first
spiracle and the wing. The pleural area tapers downward and becomes
continuous with the sternum (S,) between the first and second legs.
A typical pleural sulcus (PIS,) extends from the base of the middle
leg to the wing fulcrum at the base of the wing (W). The area before
the groove is episternal, that behind it epimeral. The episternal area
includes a major episternal plate (Eps,) continuous below with the
sternum, and a smaller preepisternum (eps,). The epimeron (Epme)
is a simple quadrate plate. Below it is a small triangular plate (Ss),
which in the mosquito appears to be a postcoxal lobe of the sternum ;
but a plate in the same position in higher flies is the detached meron of
the coxa. In some species the episternum is divided into an upper
and a lower part (fig. 27 A).
The prothorax is so reduced and modified that it is difficult to
interpret its parts. The notum (fig. 25 Nz) includes a narrow plate
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 65
across the back beneath the overhanging front end of the mesonotum,
and apparently a larger posterior plate on each side. This posterior
plate, however, tapers narrowly down to the coxa so that its lower part
must be epimeral. The episternum then is represented by a short plate
(Eps,) between the first notal plate and the coxa. A plate in the side
of the neck (CvP1) that supports the head is unquestionably a cervical
sclerite.
Fic. 25——Thorax of Psorophora, with head and base of abdomen (from the
author’s illustration in Howard, Dyar, and Knab, 1912).
AN, wing-bearing notal plate; CvPl, cervical plate; Epm, epimeron; Eps,
episternum; eps, preepisternum; H/t, halter; N, notum; PI, pleuron; PIS,
pleural sulcus; PN, postnotum; puf, paranotal fold; S, sternum; Scl, scutellum;
Sct, scutum; W, wing.
Subnumbers 1,2,3 designate parts of prothorax, mesothorax, and metathorax.
The metathorax is even more simplified than the prothorax. The
notum (fig. 25, Ns) is much narrowed across the back, but it expands
on the sides where it carries the halteres (H/t). From the notum the
pleural region continues downward on the side, tapering to the hind
coxa. Close to its posterior margin is a faint line that perhaps repre-
sents the pleural sulcus. A narrow strip (PNs;) between the meta-
notum and the first abdominal segment, more plainly seen in Aedes
(fig. 27 C,D, PNs), is clearly the metapostnotum, since it gives at-
tachment to the first abdominal muscles (G).
The wings of the mosquito have a simple pattern of venation, shown
at A of figure 26, in which the veins are named according to the Com-
66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
stock-Needham system. Mosquito taxonomists, however, usually
designate the veins behind the subcosta by numbers. In this scheme
R, is vein 1, Rg and Rs are branches of vein 2, R,,5 is vein 3, M and
its two branches are vein 4, Cu and its two branches are vein 5, and A
is vein 6. The veins are densely clothed on both sides of the wing with
long, slender, fusiform, or scalelike setae (omitted in the figure).
While the simple venation of the mosquito wing is of a fairly
generalized pattern, the basal wing structure has little resemblance
R Se
\ i ‘ R
Cup Cu, M3 My+2 ae
Fic. 26.—The wing and halter.
A, Culex, left wing and halter, wing partly flexed on basal lobe, scales re-
moved to show venation. B, Anopheles, base of wing, flattened. C, Culex, halter
enlarged, posterior.
A, anal vein; C, costa; Cu, cubitus; M, media; R, radius; Sc, subcosta.
to that of most other insects, and would appear to be specialized by
elimination of the usual axillary sclerites. When the wing is flexed
(fig. 26 A) a fold near the base sets off a triangular basal lobe by
which the wing is attached to the thorax. During flexion the wing
turns posteriorly over the basal lobe, which is then covered from above
by the fully flexed wing, and gives the wing the appearance of being
supported on a lobe of the thorax. The principal sclerotization of the
wing base is a long, anterior jointed bar (B, r) that supports the
radial vein, and bends at the joint when the wing is flexed (A).
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 67
Otherwise the membrane of the whole basal area is occupied by irregu-
lar thickenings or weak sclerotizations that are hardly sclerites and
seem to have no mechanical significance. They are better developed
in Anopheles (B) than in Culex (A). The same structure in modified
form is present also in some related Nematocera, but not in Tipulidae.
The wing mechanism of extension and flexion is not understood,
but all the direct muscles of flight appear to be attached on the basal
lobe. The indirect flight muscles are as fully developed as in any
other fly. They include great masses of dorsal longitudinal fibers and
lateral vertical fibers that almost completely fill the thorax. The
weight of the flight muscles of Aedes has been calculated by Hocking
(1953) as from 16.5 to 18.7 percent of the total body weight, which,
however, is small as compared with Tabanus in which the flight
muscles are 23 to 35 percent of the body weight.
The rate of the wing vibration in flight, measured in wing beats per
second, is given by Sotavalta (1947) for females as 165 to 196 for
Culex pipiens, 165 to 247 for Anopheles maculipennis, 241 to 311 for
Aedes cantans and Aedes punctor. With males the rate is consistently
higher, from 330 to 587 beats per second by Anopheles and Aedes.
Hocking (1953) has measured the flying speed of five species of
Aedes. In ordinary cruising flight they go from 75 to 110 centimeters
per second, but for short distances they can make 220 to 252 centi-
meters in a second.
The legs of the mosquito have no unusual features, except for their
length and relative slenderness. Each leg (fig. 17 B) has the usual
six segments of an insect leg, a coxa, trochanter, femur, tibia, tarsus,
and pretarsus. The long tarsus is subdivided into five tarsomeres. The
pretarsus has two decurved claws but no arolium. In some species, as
in Culex, the foot is provided with a pair of small padlike pulvilli; in
others there is only a heel-like hairy swelling at the bases of the claws.
Most mosquitoes, however, whether they have foot pads or not, are
able to cling to smooth vertical surfaces, such as window panes or the
walls of a glass jar.
THE ABDOMEN
The abdomen of the adult mosquito (fig. 27 A) is broadly joined to
the thorax and tapers posteriorly. The tergal and sternal plates are
separated on the sides by membranous areas containing the spiracles,
which are present on segments I to VII. In each sex the abdomen has
Io segments, as in the pupa, but in the females of some species the
eighth segment is ordinarily retracted into the seventh, and in the
male the ninth segment is concealed within the eighth.
68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Fic. 27.—Details of the adult thorax and abdomen.
A, Aedes aegypti, male thorax and abdomen. B, Same, end of abdomen ex-
tended. C, Same, thorax and base of abdomen, dorsal. D, Same, postnotum of
metathorax extended as a phragma into base of abdomen, ventral. E, Same,
tenth abdominal segment of male, lateral. F, Same, undersurface (dorsal). G,
Same, showing dorsal muscles of first abdominal segment. H, Culex, end of
female abdomen, ventral. I, Same,. lateral. J, Composite diagram of female
terminalia, lateral (from Gerry 1932). K, Same, ventral, with ventral arc of
sigma (c) turned forward (from Gerry 1932).
a, cowl; An, anus; b, dorsal arc of sigma (ninth sternum?) ; c, ventral arc of
sigma; Cer, cercus; Gir, gonotreme (opening of genital chamber) ; H/t, halter;
N, notum; pgpl, postgenital plate; Ph, phragma; Pmr, paramere; PN, post-
notum; s, lateroventral prong of tenth segment; S, sternum; Scl, scutellum;
Sct, scutum; ¢, tergum of tenth segment; 7, tergum.
Subnumbers 1-3, thoracic segments; J-X, abdominal segments.
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 69
The male mosquito is readily distinguished from the female by the
presence of a pair of large, two-segmented genital claspers, or
parameres, projecting from the end of the abdomen (fig. 27 A, Pmr).
Though the ninth segment is ordinarily concealed by retraction into
the eighth, on pulling out the end of the abdomen (B), it is seen to
be a small sclerotic ring (JX) carrying the parameres. The anus-
bearing tenth segment, or proctiger (X), is mostly hidden between
the bases of the parameres, and is apparently ventral in position. In
fact, the whole terminal part of the male abdomen beyond the seventh
segment, except in newly emerged individuals, is turned upside down,
so that the tergal plates are ventral and the sternal plates dorsal. The
inversion takes place slowly during the first 24 to 48 hours after
emergence from the pupa.
The tenth abdominal segment of the male is a flattened anal lobe
with an expanded base projecting from above the inverted tergum of
the ninth segment (fig. 27 E,F). In its base are two dorsolateral
sclerites (¢) that may be regarded as tergites. On the ventral (upper)
surface are two marginal bars (s), the ends of which project as a
pair of free, toothed prongs. These bars have commonly been re-
garded as sternites, but Christophers (1923) says they are the cerci
united with the anal lobe.
The external genital organs of the male insect, because of their
generic and specific variations, are important diagnostic features for
taxonomists. In the mosquito they include primarily the paired lateral
claspers and a median intromittent organ, carried by the ninth ab-
dominal segment. Various names are given to these parts by different
specialists, but the organs have essentially the same origin in all insects,
and there is no need for special terms in the several orders, and
certainly there is no excuse for specialists in one order to use different
names for the same parts in different species. For simplicity the
claspers are here termed the parameres, and the intromittent organ
the aedeagus. Various secondarily developed accessory parts, of
course, must have more specific names.
In the insects in general the male genitalia take their origin from
a pair of primary phallic lobes that develop in a late instar of the
nymph or larva on the posterior part of the ninth abdominal segment
at the sides of the future gonopore. Later, each lobe divides into two
parts, a mesal mesomere and a lateral paramere. Eventually the
mesomeres unite around the gonopore to form the aedeagus, and the
parameres become the claspers.
The development of the genital organs in the male mosquito has
been shown by Christophers (1922) to proceed in the usual manner.
7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Early in the fourth instar of the larva paired thickenings of the epi-
dermis appear behind the region of the ninth sternum. These “genital
plaques” soon take on the form of budlike outgrowths, which are the
primary phallic lobes (fig. 28 A, PhL). With further development
the lobes elongate and unite at their bases, forming the genital ap-
Fic. 28.—External genital organs of the adult male, and their development.
(A,B,C, from Christophers, 1922.)
A, The primary phallic lobes that appear in a late instar larva behind the
sternal region of the ninth abdominal segment. B, Later stage of same, each
primary lobe divided into a mesomere and a paramere. C, Still later stage,
mesomeres united around the gonopore to form the aedeagus. D, Adult genital
apparatus of Anopheles quadrimaculatus, lower surface (dorsal). E, Parameres
and claspettes of Aedes pullatus.
Aed, aedeagus ; Bmr, basimere; bp, basal plate; c/sp, claspette; Gpr, gonopore;
IXT, ninth abdominal tergum; Mmr, mesomere; PhL, primary phallic lobes;
Pmr, paramere; Tmr, telomere.
pendages as they appear in the pupa (fig. 16F). At this stage the
lobes are termed ‘‘proandropodites” by Christophers (1922), but this
term literally translated would mean “primitive male parts of legs”
(as “coxopodite” means the “coxal part of a leg’). Since there is
no real evidence that the male genital organs of insects represent
primitive legs, the genital organs of the pupa are simply the developed
phallic lobes. Within them are formed the definitive genitalia of the
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 71
adult. From the base of each lobe inside the pupal cuticle, as de-
scribed by Christophers, is cut off a small median lobe (fig. 28 B,
Mmr), and the lateral part becomes the rudiment of the clasper
(Pmr). Finally, the two median lobes unite around the gonopore
to form the aedeagus (C, Aed), while the lateral parameral lobes
elongate to become the two-segmented claspers of the adult (D,E).
In the mature condition the genitalia take on a great variety of
forms and are complicated by the development of accessory parts.
All this is a great boon to taxonomists, but it often creates difficulty
for the morphologist. Anopheles quadrimaculatus (fig. 28D) gives
a good example of one type of structure. Each paramere is divided
into a large basimere (Bmr) and a long slender telomere (Tmr).
The telomere is movable on the basimere by strong antagonistic
muscles arising in the latter. The slender aedeagus (Aed) lies between
the bases of the parameres and is connected with the basimeres by a
pair of small basal plates (bp). The basimeres are equipped with
long spines, and proximally each bears a membranous median lobe
(clsp) united with the one from the opposite side. Each lobe is armed
with strong spines and is known as a claspette, or claspette lobe. In
other genera the claspettes are more commonly independent ap-
pendages of the parameres, as seen in Aedes (E). The claspettes,
according to Christophers (1922), are cut out from the parameres
by secondary incisions of the latter.
For illustrations of generic and specific variations in the male
genital structure the student must consult taxonomic papers, but the
nomenclature will be confusing. In the current terminology of
mosquito specialists, the aedeagus is called the “mesosome” or “phallo-
some,” the basal plates (bp) that connect it with the claspers are the
“parameres,” and the claspers are the “side pieces.” In this scheme
the term “paramere” is entirely misapplied, since it was first given to
the claspers, and moreover, “side piece” is a direct English translation
of “paramere.” The segments of the claspers are known also as the
“basistyles” and “‘dististyles,’ but as shown by their development the
claspers have no relation whatever to legs or abdominal styli. The
terminology given on figure 28 is recommended for its simplicity and
because it can be applied, on the basis of development, to the male
genitalia in all the principal orders of insects (see Snodgrass, 1957).
The terminal parts of the female abdomen are much simpler than
those of the male, but their homologies are more difficult to under-
stand. Beyond the eighth segment projects a small lobe (fig. 27 1)
representing the combined ninth and tenth segments. The dorsum of
72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
the ninth segment is a transverse basal arc (JX) usually containing a
small tergal sclerite. Beyond it is the tenth segment (X) bearing a
pair of lateral cerci (Cer) and the terminal anus (4m). Ventrally is
a lobe known as the postgenital plate (pgpl) because the gonotreme
(H, Gtr), or opening of the genital atrium, is situated at its base
above the sternum of the eighth segment (VJJ/). The nature of the
postgenital plate is doubtful ; it looks as if it should be the projecting
sternum of the ninth segment. On its base there is generally a trans-
verse fold known as the cowl (K, a) because it is sometimes reflected
to form a hoodlike pocket. Surrounding the gonotreme above the
end of the eighth sternum is a sclerotized ring (b, c) named the sigma
by Christophers (1923). In figure K the ventral arc of the sigma (c)
is turned forward; normally it is directed posteriorly (J, c). The
sigma thus, as described by Christophers, resembles the lips of a half-
opened clasp purse, in which it is represented by the metal frame-
work of the purse. Some writers, however, without adducing specific
evidence, regard the dorsal arc of the sigma as the ninth sternum.
According to Christophers the whole structure is formed as a scleroti-
zation in the intersegmental membrane of the gonotreme.
All parts of the female terminalia are subject to much variation, as
shown in comparative studies by Macfie and Ingram (1922), Christo-
phers (1923), Davis (1926), Gerry (1932), Gjullin (1937), Roth
(1946), Rees and Onishi (1951), and Hara (1957). The student,
however, will be somewhat confused by the different ways the parts
are represented and named. The drawings J and K on figure 27, taken
from Gerry, are composite diagrams showing all the parts that have
been described, but they probably do not present the exact structure
in any one species.
The gonotreme surrounded by the sigma above the eighth abdominal
sternum leads into a small infolded pouch, the genital chamber, or
atrium. In its anterior wall is the female gonopore, which is the open-
ing of the median oviduct. Behind the gonopore the globular sperma-
thecae (one, two, or three in number) open through the dorsal wall of
the atrium, and into a posterior pouch of the dorsal wall, the caecus,
opens the single accessory gland, called the “mucus gland,” but the
nature of its secretion is not known (fig. 30 B).
INTERNAL ANATOMY
A thorough study of the internal anatomy of the mosquito has not
been made, but the parts of principal interest will be the alimentary
canal and the reproductive organs. The muscular and tracheal systems
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 73
have no features peculiar to the mosquito, and even the unusual char-
acters of the reproductive organs are common to other Diptera. The
simple nervous system is that of the larva with an elaboration of the
brain and the optic lobes in the head, a transposition of the first
abdominal ganglion to the thorax, and the union of the eighth ab-
dominal ganglion with the ganglion of the seventh segment. In the
abdomen of the adult, therefore, the first ganglion is in the second
segment (fig. 30C, Gngl/), and the last is a composite ganglion
(Gng VII+VIII) in the seventh segment. The tracheal system has
lost the large dorsal trunks of the larva, and the lateral trunks along
the spiracles have been enlarged.
The circulatory organs.—In the adult mosquito, as described by
Jones (1954) in Anopheles, the dorsal blood vessel has in general the
same structure as that of the larva. The part in the abdomen, how-
ever, is more distinctly “chambered” because of segmental swellings
before the ostia. An aortic sinus is said by Jones (1952) to be present
in the adult as in the larva and pupa of Anopheles, Culex, and Aedes.
The sinus is a dilatation of the aorta in the dorsal part of the thorax,
with the corpora allata-cardiaca attached to it laterally. Anteriorly
the sinus is continued into the cephalic aorta. The adult heart, ac-
cording to Jones, beats predominantly forward, but periodically re-
verses the direction of the beat. The heart has no innervation from
any source and therefore its pulsations are myogenic, that is, engen-
dered by the muscles themselves of the heart wall. Lateral alary
muscles support the heart, but they do not vibrate, and when cut the
heart keeps on beating.
A vibratile muscular membrane across the cavity of the mesothoracic
scutellum appears to be an accessory pulsatile organ, as in some other
insects. A frontal bulblike organ between the bases of the antennae
has been described by Day (1955) as a sense organ, and by Clements
(1956) as a pulsating organ for driving blood into the antennae.
If it is a sense organ, it is a newly discovered one as Day claims;
if it is a pulsating organ it is not unique since a pulsatile organ in the
same place is present in various other insects.
The alimentary canal——The alimentary canal of the adult mosquito
(fig. 29 A) in its general form is quite different from that of the larva.
From the pharyngeal pump in the head (PhP) a short, narrow
oesophagus (Oe) extends into the front of the thorax, where it joins
a wider tube, which is the beginning of the stomach, or ventriculus
(Vent). Shortly before its junction with the stomach the oesophagus
gives off three pouches, known as the oesophageal diverticula, two
74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
of which are dorsal and one ventral. In Aedes aegypti the dorsal
diverticula (A, ddv) are small, flat, elongate sacs with slender necks
diverging forward and laterally from the oesophagus (C). The single
ventral diverticulum (4, vdv) has a long, slender neck which ex-
pands into a large sac in the anterior half of the abdomen. This
ventral diverticulum corresponds with the usual ‘“‘crop” of other
Diptera.
Fic. 29.—Alimentary canal and salivary glands of the adult female of
Aedes aegypti.
A, Sectional view of body showing alimentary canal and salivary system
(glands on left displaced). B, The salivary glands. C, The oesophageal
diverticula, ventral. D, Rectal sac and papillae.
Alnt, anterior intestine; CbP, cibarial pump; ddv, dorsal diverticulum;
Hphy, hypopharynx; Lb, labium; Lm, labrum; Mal, Malpighian tubules; Oe,
oesophagus; PhP, pharyngeal pump; Rect, rectum; rp, rectal papillae; S/Dct,
salivary duct; S/GI/d, salivary glands; S/P, salivary pump; vdv, ventral di-
verticulum; Vent, ventriculus.
The ventriculus (fig. 29 A, Vent), which is the functional stomach
of the insect, for most of its length in the female mosquito is a
narrow tube that extends upward through the thorax and then turns
backward into the abdomen where it ends in a saclike enlargement that
joins the intestine. The first part of the latter, or anterior intestine
(AlInt), is a short, slender tube thrown into a small loop. Its anterior
end, the pyloric region, joins the ventriculus by a funnel-shaped ex-
pansion. At the other end the anterior intestine is continued into the
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 75
posterior intestine, or rectum (Rect), which is much enlarged an-
teriorly and tapers back to the anus. The inner wall of the pyloric
funnel is armed in some species with numerous small spines directed
posteriorly. These pyloric spines have been described and well il-
lustrated by Trembley (1951) in species of Anopheles, Aedes, and
Culex. In the anterior end of the rectum are six small, soft, conical
rectal papillae (D, rp) projecting inward from the rectal wall. Five
Malpighian tubules (A, Mal) arise from the pyloric region of the
intestine as in the larva.
The oesophageal diverticula are said to be empty on emergence of
the mosquito from the pupa. Within an hour after ecdysis, however,
according to Marshall and Staley (1932), the air that was pumped
into the stomach begins to pass forward into the diverticula, and in
12 to 22 hours the stomach is empty.
The function of the oesophageal diverticula in relation to food
intake has been studied by a number of investigators, but, though
using the same experimental methods of feeding, the latter have not all
come to the same conclusions. The subject has recently been well re-
viewed by Trembley (1952) and by Megahed (1958), and good
bibliographies are given by both these writers. In general it is found
that ingested blood goes directly to the stomach, while fruit juices and
sugar solutions go first into the diverticula, to be later delivered to
the stomach. According to Trembley, blood in small amounts may
occasionally go to the diverticula, and sugar solutions sometimes go
direct to the stomach. The work of Megahed on Culicoides gives
essentially the same results, the stomach being ordinarily the receptacle
for blood, the diverticula for concentrated sugar solutions, but water
and dilute sugar solutions go direct to the stomach. Most observations
seem to apply to the female insect. Day (1954), however, in experi-
ments on male mosquitoes, found that the sexes react similarly : “blood
went to the mid-gut and sugar to the diverticulum in the male in spite
of the fact that males do not ingest blood under natural conditions.”
The “switching mechanism” that determines whether the ingested
food goes into the stomach or the diverticula, Day (1954) has pro-
posed, is governed by the different kinds of sense organs in the wall of
the cibarial pump (buccal cavity). If receptors of one type are
stimulated by sugar it may be supposed that they cause a relaxation of
sphincter muscles of the diverticula; if others are sensitive to blood
components, they may effect a relaxation of the cardiac sphincter of
the stomach. In the neck of the ventral diverticulum, Day notes the
presence of a group of spines, which would appear to assist in keeping
76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
blood corpuscles out of the diverticulum when the circular muscles in
the neck of the diverticulum are contracted.
The salivary glands.—The salivary glands of the mosquito consist
each of three lobes (fig. 29 B), of which the middle lobe is shorter
than the other two. The glands lie at the sides of the anterior end of
the ventriculus (A, S/G/d; the left gland is displaced in the figure).
The two ducts extend into the back of the head, where they unite in a
single outlet tube (fig. 24 A, SIDct), which ends at the base of the
hypopharynx in a small syringelike swelling that acts as a salivary
ejection pump (S/P). On the elastic dorsal wall of the pump is in-
serted a dilator muscle (18) from the floor of the cibarial pump. The
salivary pump discharges through the salivary canal (sc) of the hypo-
pharynx in the female; in the male the duct traverses the labium
(fig. 22 F). The salivary secretion in species of Anopheles, according
to Metcalf (1945), contains both an anticoagulin and an agglutinin,
but in other pest species neither appears to be present.
The salivary glands are of particular interest in connection with the
transmission of disease by mosquitoes. They offer the only avenue of
escape for disease organisms from the body cavity of the mosquito into
the blood of an alternate host. The sporozoites of malaria, for ex-
ample, that penetrate into the salivary glands are carried in the saliva
of the biting mosquito directly into the vertebrate host, which is
necessary for the completion of the complex life history of the
malaria parasite, Plasmodium. This suggests the question of how it
became obligatory for some parasites to divide their developmental
history between two different animals, but the known facts give no
answer. Mosquitoes do not bite each other, and there is no way by
which the malaria parasite can be normally transferred from one
vertebrate to another.
The reproductive system.—The organs of reproduction in the
Diptera include the parts common to all insects, but their structure in
two respects is exceptional. Each testis appears to correspond with a
single testicular tube in other insects; the egg tubes of each ovary are
extremely small, and all are enclosed in a cellular sheath.
The male organs of the mosquito include a pair of testes (fig. 30 E,
Tes), a pair of testicular ducts, or vasa deferentia (Vd), which enlarge
posteriorly to form a pair of seminal vesicles (SV) that in some
species are united (D). The vesicles end in a very short common
ductus ejaculatorius (Dej), which receives a pair of large accessory
glands (AcG/d) and then opens directly into the base of the aedeagus
(Aed). In the normal condition the reproductive organs lie beneath
the alimentary canal, but, with the inversion of the terminal segments
of the abdomen, the relation is reversed (fig. 30 A)—the ejaculatory
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS TG.
Fic. 30.—Reproductive organs and the abdominal nerve cord of the adult
mosquito.
A, Culex quinquefasciatus, end of male abdomen, left side removed, exposing
the inverted reproductive organs and rectum. B, Diagram of female reproductive
organs, dorsal (adapted from Christophers, 1901). C, Aedes aegypti, male,
ventral nerve cord of abdomen and genital outlets, dorsal. D, Same, male re-
productive organs, ventral. E, Culex quinquefasciatus, male reproductive organs
after inversion, lower side (dorsal). F, Same, with part of intestine, upper side
(ventral).
AcGld, accessory gland; Aed, aedeagus; Atr, atrium, genital chamber; Bmr,
basimere; Dej, ductus ejaculatorius; Gng, ganglion; Gtr, gonotreme; Int,
intestine; Odc, oviductus communis; Odl, oviductus lateralis; Ov, ovary; Ovl,
ovariole; Pmr, paramere; Ptgr, proctiger; Rect, rectum; S, sternum; Sh, sheath
of ovary; Spt, spermatheca; SV, seminal vesicle; T, tergum; Tes, testis; tf,
terminal filament of ovary; Tmr, telomere; Vd, vas deferens.
78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
duct, the seminal vesicles, and the accessory glands now lie above the
intestine. Since the testes are not affected by the inversion, the long
vasa deferentia as in Culex (E) cross each other, but when the ducts
are united as in Aedes (D) a simple twist takes place at the junction
of the ducts.
The testis of most insects consists of a number of individual tubes
in which the spermatozoa are formed as are the eggs in the ovarian
tubes, and, except in the apterygotes, the tubes of each testis are
enclosed in an investing sheath. The testes of the mosquito are
elongate, pear-shaped bodies (fig. 30 D,E,F, Tes) continuous with the
ducts. Each testis, however, appears in its entirety to be a single
testicular tube. The same is true of the testes in other Diptera. In the
narrowed upper end of each organ is a mass of undifferentiated cells ;
the rest of the lumen is filled with spermatocytes and spermatozoa
in various stages of development. The mature spermatozoa are ex-
tremely long and threadlike; when liberated from the testis they
exhibit active undulatory movements. The spermatozoa are stored
in the seminal vesicles preliminary to mating, and the accessory glands
probably have a prostate function, giving the spermatozoa a liquid
medium in which they are discharged.
The reproductive organs of the female mosquito, represented
diagrammatically at B of figure 30, include the parts characteristic of
the female organs of insects in general. These are a pair of ovaries
(Ov), the lateral oviducts (Odl) from the ovaries, and a median
common oviduct (Odc) with which the lateral ducts are joined. The
common duct opens by the primary genital aperture, or gonopore, into
a small pocket above the end of the eighth abdominal sternum. This
pocket, the genital chamber, or atrium, being a secondary inflection
of the body wall between the eighth and ninth abdominal segments, is
therefore not a part of the primary genital passage. The external
opening of the atrium may be designated the gonotreme (Gtr). Into
the dorsal wall of the atrium just behind the gonopore open the ducts
of the spermathecae (Spt), which are usually three in number, though
in Anopheles there is only a single spermatheca. Behind the
spermathecal openings arises an accessory gland (AcGld), the func-
tion of which is not known in the mosquito. In other insects accessory
glands usually secrete a cement for attaching the eggs to a support,
or a material to form an egg covering.
The atrium serves as a copulatory pouch at the time of mating, and
the spermatozoa from the male are stored in the spermathecae. Then
when the eggs leave the oviduct they are received in the atrium and
no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 79
are here fertilized by sperm discharged from the spermathecae.
Finally the eggs are passed out through the gonotreme at the time of
laying.
The ovaries of the mosquito differ in several respects from the
usual structure of these organs in other insects. A typical insect
ovary consists of a group of slender tubes known as ovarioles opening
into the end of a lateral oviduct. The ovarioles taper upward and
end in filaments that unite in a common strand attached to tissues
in the neighborhood of the heart. A mature ovariole contains a series
of ripening egg cells of successively larger size, with the mature egg
in its lower end. Each egg is accompanied by a number of nutritive
cells, or so-called nurse cells, which are absorbed by the egg as it
matures. Each egg and its nurse cells are contained in a compartment
of the ovariole known as a follicle. The follicles appear as swellings
along the ovariole, increasing in size with the growth of the egg.
The egg cell and the nurse cells are formed by division of the undif-
ferentiated cells in a chamber, the germarium, in the upper end of
the follicle. The eggs do not pass down the ovarioles; each ovariole
grows from the germarium as an egg leaves the lowermost follicle and
the latter disintegrates.
In the mosquito ovary (fig. 30 B) the ovarioles (Ovl) are very
short and are arranged in rows along an axial cavity of the ovary. As
in other Diptera, each ovary is invested in a thin membranous sheath
(Sh) in which there are fine muscle fibers, and the sheath itself ends in
a terminal filament (tf) attached to tissues along the sides of the
heart. The muscle fibers of the ovarian sheath in Anopheles are said
by Nicholson (1921) to be striated, but Jones (1958) finds that those
of Aedes do not show a distinct striation in live, unstained whole
mounts at 1,000 magnification under phase optics.
Each ovariole consists of a large egg-containing follicle with a small
projection on its free end representing the germarium and one or
two minute undeveloped follicles, The structure of the egg follicle of
Culex has been described by Nath (1924), and an account of the de-
velopment of the ovary and the development and nutrition of the eggs
in the ovary of Anopheles is given by Nicholson (1921), by Christo-
phers, Sinton, and Covell (1928), and by Mer (1936). The develop-
mental processes described in the mosquito differ little from those in
insects generally.
Many female mosquitoes need a meal of blood for the production of
eggs. The eggs of Anopheles and Aedes are fully developed in two to
three days after the female has fed. It is said by Roy (1936) that in
80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Aedes there is “a definite quantitative relationship between the weight
of the blood meal and the number of eggs produced.” As noted by
Christophers, Sinton, and Covell (1928), the eggs in the lower follicles
of all the ovarioles mature at the same time, so that as many eggs are
ready for laying as there are ovarioles. When these eggs are deposited
the eggs in the next follicles above mature, and so the production of
fresh lots of eggs “seems to have no limit other than the life of the
mosquito.”
REFERENCES
ANTHON, H.
1943a. Der Kopfbau der Larven einiger nematoceren Dipterenfamilien. Spolia
Zool. Mus. Hauniensis, vol. 3, pp. 1-61, 11 pls.
1934b. Zum Kopfbau der primitivsten bisher bekannten Dipterenlarvae:
Olbiogaster sp. (Rhyphidae). Ein Beitrag zur Phylogenie der
nematoceren Dipteren. Saertryk af Ent. Medd., vol. 23, pp. 303-320,
3 pls.
Barraup, J. P., AND COVELL, G.
1928. The morphology of the buccal cavity in anopheline and culecine
mosquitoes. Indian Journ. Med, Res., vol. 15, pp. 671-670, 8 pls.
Bates, M.
1949. The natural history of mosquitoes. 379 pp., 6 text figs., 16 pls. New
York.
BERGER, C. A.
1938. Multiplication and reduction of somatic chromosome groups as a
regular developmental process in the mosquito, Culex pipiens.
Contr. Embryol., No. 167, pp. 211-232, 10 text figs., 1 pl.
BuatiA, M. L., Aanp Watrat, B. L.
1957. Description of a hitherto unknown structure in the mouth parts of
mosquitoes. Indian Journ. Malariol., vol. 11, No. 2, pp. 183-180,
4 text figs., 1 pl.
BovENSTEIN, D.
1945. The corpora allata of mosquitoes. Connecticut Agr. Exp. Stat. Bull.
488, pp. 396-405, 3 figs.
BonNE-WEPSTER, J.
1932. A mosquito with larval habits like Taeniorhynchus. Bull. Ent. Res.,
vol. 23, pt. I, pp. 60-72, 3 figs.
Boyp, M. F.
1949. Malariology. 3 vols., 65 contributors. Philadelphia and London.
BRrELAND, O. P.
1949. The biology and the immature stages of the mosquito, Megarhinus
septentrionalis Dyar & Knab. Ann. Ent. Soc. America, vol. 42, pp.
38-47, 3 figs.
Cuane, S. L., anp Ricwart, F. E., Jr.
1951. The anatomy and function of the so-called “notched organs” of Nuttal
and Shipley on the thorax of larvae of Anopheles quadrimaculatus.
Journ. Nat. Malaria Soc., vol. 10, No. 4, pp. 287-292, 4 figs.
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 81
CHAUDONNERET, J.
1951. Reflexions sur les premandibules des larves de culicides (Dipteres
nematoceres). Bull. Soc. Zool. France, vol. 76, pp. 399-404, 1 fig.
CuiILp, C. M.
1894. Ein bisher wenig beachtetes antennalen Sinnesorgan der Insekten, mit
besonderer Berticksichtigung der Culiciden und Chironomiden.
Zeitschr. wiss. Zool., vol. 58, pp. 475-528, 2 pls.
CHISWELL, J. R.
1955. On the last instar larva of Tipula livida van der Wulp (Diptera,
Tipulidae) with notes on the fronto-clypeal region of larval
Tipulinae and caterpillars. Proc. Roy. Ent. Soc. London, A, vol.
30, pp. 127-136, 20 figs.
CHRISTOPHERS, S. R.
1901. The anatomy and histology of the adult female mosquito. Reports to
Malaria Committee of Roy. Soc. London, ser. 4, 20 pp., 6 pls.
1922, The development and structure of the terminal segments and hypo-
pygium of the mosquito with observations on the homologies of the
terminal segments of the larva. Indian Journ. Med. Res., vol. 10,
PP. 530-572, 7 text figs., 4 pls.
1923. The structure and development of the female genital organs and
hypopygium of the mosquito. Indian Journ. Med. Res., vol. 10,
pp. 698-720, 3 text figs., 2 pls.
CHRISTOPHERS, S. R.; Stnton, J. A.; AND COovELL, G.
1928. Dissection of mosquitoes to determine age and condition of females.
Govt. of India Health Bull. No. 14, pp. 61-74, 1 pl.
Cuwatt, L. J., anp Major, H.
1945. The morphology of the pharyngeal armature in Anopheles gambiae
and Anopheles gambiae var. melas from southern India. Ann. Trop.
Med. Parasitol., vol. 39, pp. 124-128, 3 figs.
CLEMENTS, A. N.
1956. The antennal pulsating organs of mosquitoes and other Diptera.
Quart. Journ. Micr. Sci., vol. 97, pp. 429-433, 2 figs.
CoNSTANTINEANU, M. J.
1930. Die Aufbau der Sehorgane bei den im Siisswasser lebenden Dipteren-
larven und bei Puppen und Imagines von Culex. Zool. Jahrb., Anat.,
vol. 52, pp. 251-346, 25 text figs., 12 pls.
Cook, E. F.
1944a. The morphology of the larval heads of certain Culicidae (Diptera).
Microentomology, vol. 9, pt. 2, pp. 38-68, 17 figs.
1944b.On the morphology of the larval head of Chironomus (Diptera:
Chironomidae). Microentomology, vol. 9, pt. 2, pp. 69-77, 4 figs.
Davis, N. C.
1926. Notes on the female hypopygia of anopheline mosquitoes, with special
reference to some Brazilian species. American Journ, Hyg., vol. 6,
March Suppl., pp. 1-22, 38 figs.
Day, M. F.
1954. The mechanism of food distribution to midgut or diverticula in the
mosquito. Australian Journ. Biol. Sci. vol. 7, pp. 515-524, 2 text
figs., I pl.
82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
1955. A new sense organ in the head of the mosquito and other nematocerous
flies. Australian Journ. Zool., vol. 3 (3), pp. 331-335, 3 figs.
DE MEIJERE, J. C. H.
1911. Zur Metamorphose der myrmecophilen Culicide Harpagomyia splen-
dens de Meij. Tijds. Ent., vol. 54, pp. 162-167, 1 pl.
DEMEREC, M.
1950. Biology of Drosophila. 632 pp. New York.
Dimmock, G.
1881. Anatomy of the mouth-parts and the suctorial apparatus of Culex.
Psyche, vol. 3, pp. 231-241, I pl.
DuPorte, E. M.
1946. Observations on the morphology of the face in insects. Journ. Morph.,
vol. 79, pp. 371-417, 7 pls.
FARNSWORTH, MARjoRIE W.
1947. The morphology and musculature of the larval head of Anopheles
quadrimaculatus Say. Ann. Ent. Soc. America, vol. 40, pp. 137-151,
3 pls.
FRANKENBERG, B. v.
1937. Die erste Fiillung der Tracheen bei Culex und Corethra. Zool.
Jahrb., Zool. und Physiol., vol. 58, pp. 175-180, 3 figs.
GALLIARD, H.
1934. Notes sur la biologie et l’anatomie de la larve de Taeniorhynchus
(Mansonia) richiardii Ficalbi. Ann. Parasit. Hum. Comp., vol. 12,
pp. 465-471.
Gerry, B. I.
1932. Morphological studies of the female genitalia of Cuban mosquitoes.
Ann. Ent. Soc. America, vol. 25, pp. 31-75, 6 pls.
GyuLuin, C. M.
1937. The female genitalia of the Aedes mosquitoes of the Pacific Coast
States. Proc. Ent. Soc. Washington, vol. 39, pp. 252-266, 3 pls.
Gorpon, R. M., AnD LumMspeEn, W. H. R.
1939. A study of the behaviour of the mouth-parts of mosquitoes when
taking up blood from living tissue; together with some observations
on the ingestion of microfilaria, Ann. Trop. Med. Parasitol., vol.
33, PP. 259-278, 13 figs.
Goutn, F. J.
1959. Morphology of the larval head of some Chironomidae (Diptera,
Nematocera). Smithsonian Misc. Coll., vol. 137, pp. 175-201, 15 figs.
GROSSBECK, J. A.
1908. Additional notes on the life history of Culex perturbans. Ent. News,
vol. 19, pp. 473-476, I pl.
Hara, J.
1957. Studies on the female terminalia of Japanese mosquitoes. Japanese
Journ. Exp. Med., vol. 27, pp. 45-91, 48 pls.
HeENnNIG, W.
1948, 1950. Die Larvenformen der Dipteren. Pt. 1 (1948), Pt. 2 (1950).
Berlin.
Hinton, H. E.
1946. Concealed phases in the metamorphosis of insects. Nature, vol. 157,
PP. 552-553.
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 83
1958a. Concealed phases in the metamorphosis of insects. Science Progress,
No. 182, pp. 260-275, 5 figs.
1958b. The pupa of the fly Simulium feeds and spins its own cocoon. Ent.
Month. Mag., vol. 94, pp. 14-16, 1 fig.
1959. Origin of indirect flight muscles in primitive flies. Nature, vol. 183,
PP. 557-558, I fig.
Hock1ne, B.
1953. The intrinsic range and speed of the flight of insects. Trans. Roy. Ent.
Soc. London, vol. 104, pp. 223-345, 20 text figs., 6 pls.
HorsFati, W. R.
1955. Mosquitoes, their bionomics and relation to disease. 723 pp. New
York.
Howakrp, L. O.; Dyar, H. G.; anp Knap, F.
1912. The mosquitoes of North and Central America and the West Indies.
3 vols. Carnegie Institution of Washington,
Hutst, F. A.
1906. The histolysis of the musculature of Culex pungens during meta-
morphosis. Biol. Bull., vol. 11, pp. 277-304, 2 pls.
EursT Gere
1890. The pupal stage of Culex. Dissertation Univ. Leipzig. 26 pp., 1 pl.
Manchester.
Imus, A. D.
1907, 1908. On the larval and pupal stages of Anopheles maculipennis,
Meigen. Pt. I (1907), Journ. Hyg., vol. 7, pp. 291-318, 2 pls. Pt. II
(1908), Parasitol., vol. 1, pp. 103-133, 2 pls.
IvenGaAR, M. O. T.
1935a. Biology of Indian mosquito larvae that attach themselves to roots
of water plants. Proc. Roy. Ent. Soc. London, A, vol. 10, pp. 9-11.:
1935b. Eggs of Ficalbia minima Theo., and notes on breeding habits of three
species of Ficalbia. Bull. Ent. Res., vol. 26, pp. 423-425, 4 figs.
JAcosson, E.
1911. Nahere Mitteilungen tiber die myrmecophile Culicide. Harpagomyia
splendens de Mej. Tijds. Ent., vol. 54, pp. 149-161, 3 pls.
JENSEN, DINNIEMAUD V., AND JONES, J. C.
1957. The development of the salivary glands in Anopheles albimanus
Wiedemann (Diptera, Culicidae). Ann. Ent. Soc. America, vol. 50,
pp. 464-460, 8 figs.
Jounston, C.
1855. Auditory apparatus of the Culex mosquito. Quart. Journ. Micr. Sci.,
vol. 3, pp. 97-102, 1 pl.
Jones, J. C.
1952. Prothoracic aortic sinuses in Anopheles, Culex, and Aedes. Proc. Ent.
Soc. Washington, vol. 54, pp. 244-246, 1 fig.
1954. The heart and associated tissues of Anopheles quadrimaculatus Say
(Diptera: Culicidae). Journ. Morph., vol. 94, pp. 71-124, 4 pls.
1958. Observations on the anatomy and physiology of the ovaries of Aedes
aegyptt L. Anat. Rec., vol. 132, p. 460.
The anatomy and rhythmical activities of the alimentary canal of
Anopheles larvae. Ann, Ent. Soc. America. (In press.)
84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
KeiLin, D.; Tate, P.; AND VINCENT, M.
1935. The perispiracular glands of mosquito larvae. Parasitol., vol. 27, pp.
257-262, 2 figs.
KENNEDY, J. S.
1939. The visual responses of flying mosquitoes. Proc. Zool. Soc. London,
vol. 109, Ser. A, pp. 221-242, 5 figs.
Knap, F.
1909. The role of air in the ecdysis of insects. Proc. Ent. Soc. Washington,
vol. 11, pp. 68-73.
Lawson, J. W. H.
1951. The anatomy and morphology of the early stages of Culicoides
nebulosus Meigen (Diptera: Ceratopogonidae-Heleidae). Trans.
Roy. Ent. Soc. London, vol. 102, pp. 511-570, 46 text figs., 1 pl.
Macrig, J. W. S., AND INGRAM, A.
1922. On the genital armature of the female mosquito. Ann. Trop. Med.
and Parasitol., vol. 16, pp. 157-188, 23 figs.
MANZELLI, M. A.
1941. Studies on the effect of reduction of surface tension on mosquito pupae.
Proc. 28th Ann. Meeting New Jersey Mosquito Extermination
Assoc., pp. 19-23, 3 figs.
MarsHALl, J. F.
1938. The British mosquitoes. 341 pp., 172 text figs., 18 pls. London.
MarRSHALL, J. F., aNp STALEY, J.
1932. On the distribution of air in the oesophageal diverticula and intestine
of mosquitoes. Parasitology, vol. 24, pp. 368-381, 8 figs.
1935. Generic and subgeneric differences in the mouth-parts of male mos-
quitoes. Bull. Ent. Res., vol. 26, pp. 531-532, I fig.
MEGAHED, M. M.
1958. The distribution of blood, water, and sugar solutions in the mid-gut
and oesophageal diverticulum of female Culicoides nubeculosus
Meigen. Bull. Soc. Ent. Egypte, vol. 42, pp. 339-355.
MENEEsS, J. H.
1958a. The maxilla and labium of the larva of Anopheles quadrimaculatus
Say. Bull. Brooklyn Ent. Soc., vol. 53, No. 2, pp. 25-38, 2 pls.
1958b. The facial areas, labrum, epipharynx, hypopharynx and mandibles of
the larva of Anopheles quadrimaculatus Say. Bull. Brooklyn Ent.
Soc., vol. 53, pp. 124-140, 2 pls.
Mer, G. G.
1936. Experimental study on the development of the ovary in Anopheles
elutus Edw. (Dipt. Culic.). Bull. Ent. Res., vol. 27, pp. 351-359.
MeEtca_r, R. L.
1945. The physiology of the salivary glands of Anopheles quadrimaculatus.
Journ. Nat. Malaria Soc., vol. 4, pp. 271-278.
Miati, L. C., anp Hammonp, A. R.
1900. The structure and life-history of the harlequin fly (Chironomus).
196 pp., I pl., 129 text figs. Oxford.
MontscHaDsky, A.
1930. Die Stigmenplatten der Culiciden-Larven. Eine systematische Studie.
Zool. Jahrb., Syst., vol. 58, pp. 541-636, 6 pls.
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 85
Natu, V.
1924. Egg-follicle of Culex. Quart. Journ. Micr. Sci., vol. 69, pp. 151-175,
2 pls.
NicuHotson, A. J.
1921. The development of the ovary and ovarian egg of a mosquito,
Anopheles maculipennis, Meig. Quart. Journ. Micr. Sci., vol. 65,
PP. 395-448, 4 pls.
Rees, D. M., ANp ONISHI, K.
1951. Morphology of the terminalia and internal reproductive organs, and
copulation in the mosquito, Culiseta inornata (Williston). Proc.
Ent. Soc. Washington, vol. 53, pp. 233-246, § text figs., 1 pl.
Ricuins, C. A.
1938. The metamorphosis of the digestive tract of Aedes dorsalis. Ann. Ent.
Soc. America, vol. 31, pp. 74-87, 1 text fig., 3 pls.
1945. The development of the midgut in the larva of Aedes dorsalis Meigen.
Ann. Ent. Soc. America, vol. 38, pp. 314-320, 1 pl.
Risver, H.
1955. Das Gehororgan der Mannchen von Culex pipiens L., Aedes aegypti
L. und Anopheles stephenst Liston (Culicidae). Zool. Jahrb., Anat.,
vol. 74, pp. 478-490, 12 figs.
Rosinson, G. G.
1939. The mouthparts and their function in the female mosquito, Anopheles
maculipennis. Parasitology, vol. 31, pp. 212-242, 9 figs.
Rotu, L. M.
1946. The female genitalia of the Wyeomyia of North America. Ann. Ent.
Soc. America, vol. 39, pp. 292-297, 1 pl.
1948. A study of mosquito behavior. American Midl. Nat., vol. 40, pp. 265-
352, 59 figs.
1951. Loci of sensory end-organs used by mosquitoes (Aedes aegypti (L.)
and Anopheles quadrimaculatus Say) in receiving host stimuli.
Ann, Ent. Soc. America, vol. 44, pp. 59-74, 6 figs.
Oven Ds NE
1936. On the role of blood in ovulation in Aédes aegypti, Linn. Bull. Ent.
Res., vol. 27, pp. 423-429.
SALEM, H. H.
1931. Some observations on the structure of the mouth parts and fore-in-
testine of the fourth stage larva of Aedes fasciata (Fab.). Ann.
Trop. Med. and Parasitol., vol. 25, pp. 303-419, 14 figs.
SAMTLEBEN, B.
1929. Zur Kenntnis der Histologie und Metamorphose der Mitteldarms der
Stechmiickenlarven. Zool. Anz. vol. 81, pp. 97-100, 5 figs.
Sato, /S:
1950. Compound eyes of Culex pipiens var. pallens Coquillett. Sci. Rep.
Tohoku Univ., ser. 4 (Biol.), vol. 18, pp. 331-341, 3 figs.
1951a. Development of the compound eye of Culex pipiens var. pallens
Coquillett. Sci. Rep. Tohoku Univ., ser. 4 (Biol.), vol. 19, pp. 23-
28, 3 figs.
1951b. Larval eyes of Culex pipiens var. pallens Coquillett. Sci. Rep. Tohoku
Univ., ser. 4 (Biol.), vol. 10, pp. 20-32, 6 figs.
86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
1953a. Structure and development of the compound eye of Aedes (Finlaya)
japonicus Theobald. Sci. Rep. Tohoku Univ., ser. 4 (Biol.), vol. 20,
PP. 33-44, 6 figs.
1953b. Structure and development of the compound eye of Anopheles
hyrcanus sinensis Wiedemann. Sci. Rep. Tohoku Univ., ser. 4
(Biol.), vol. 20, pp. 45-53, 6 figs.
SCHIEMENZ, H.
1957. Vergleichende funktionell-anatomische Untersuchungen der Kopf-
muskulatur von Theobaldia und Eristalis (Dipt. Culicid. und
Syrphid.). Deutsche Ent. Zeitschr., N.F., vol. 5, pp. 268-331, 59 figs.
SCHREMMER, F.
1949. Morphologische und funktionelle Analyse der Mundteile und des
Pharynx der Larve von Anopheles maculipennis Meig. Oster-
reichische Zool. Zeitschr., vol. 2, pp. 173-222, 14 figs.
1950. Bau und Funktion der Larvenmundteile der Dipterengattung Diva
Meigen. Osterreichische Zool. Zeitschr., vol. 2, pp. 380-413, 12 figs.
SHacasy, A. M.
1956. On the mouth parts of the larval instars of Anopheles quadrimaculatus
(Say). Bull. Soc. Ent. Egypte, vol. 40, pp. 137-174, 37 figs.
1957a. On the mouth parts of the larval instars of Aedes aegypti (L.)
(Diptera: Culicidae). Bull. Soc. Ent. Egypte, vol. 41, pp. 145-177,
38 figs.
1957b. On the mouth parts of the larval instars of Culex quinquefasciatus
(Say). Bull. Soc. Ent. Egypte, vol. 41, pp. 269-208, 38 figs.
1957c. The mouth parts of the larval instars of Psorophora ciliata (Fab-
ricius). Bull. Soc. Ent. Egypte, vol. 41, pp. 420-455, 39 figs.
1957d. The homology of the labium-hypopharynx complex of the larva of
Culex molestus Forsk. (Diptera: Culicidae). Ann. Ent. Soc.
America, vol. 50, pp. 265-270, 5 figs.
Sinton, J. A., AND COVELL, G.
1927. The relation of the morphology of the buccal cavity to the classifica-
tion of anopheline mosquitoes. Indian Journ. Med. Res., vol. 15,
PP. 301-309, 4 pls.
Snopocrass, R. E.
1944. The feeding apparatus of biting and sucking insects affecting man
and animals. Smithsonian Misc. Coll., vol. 104, No. 7, 113 pp., 39
figs.
1947. The insect cranium and the “epicranial suture.” Smithsonian Misc.
Coll., vol. 107, No. 7, 52 pp., 15 figs.
1957. A revised interpretation of the external reproductive organs of male
insects. Smithsonian Misc. Coll., vol. 135, No. 6, 60 pp., 15 figs.
SoTavALTa, O.
1947. The flight-tone (wing-stroke frequency) of insects. Acta Ent. Fen-
nica, vol. 4, 117 pp., 18 figs.
Tetrorp, A. D.
1957. The pasture Aedes of central and northern California. The egg stage:
gross embryology and resistance to desiccation. Ann. Ent. Soc.
America, vol. 50, pp. 537-543, I pl.
Tuomeson, M. T.
1905. Alimentary canal of the mosquito. Proc. Boston Soc. Nat. Hist., vol.
32, pp. 145-202, 6 pls.
No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 87
TREMBLEY, HELEN L.
1951. Pyloric spines in mosquitoes. Journ. Nat. Malaria Soc., vol. 10, No. 3
PP. 212-215, 3 figs.
1952. The distribution of certain liquids in the esophageal diverticula and
stomach of mosquitoes. Amer. Journ. Trop. Med. Hyg., vol. 1, pp.
693-710, I fig.
Tuttocu, G. S., anp SHapIRo, J. E.
1951. Electron micrographs of antennal hairs of mosquitoes. Bull. Brooklyn
Ent. Soc., vol. 46, pp. 76-78, 4 figs.
Vizzi, F. F.
1953. The mouth parts of the male mosquito Anopheles quadrimaculatus
Say. Ann. Ent. Soc. America, vol. 46, pp. 496-504, 2 pls.
VocEL, R.
1921. Kritsche und erginzende Mitteilungen zur Anatomie des Stechap-
parats der Culiciden und Tabaniden. Zool. Jahrb., Anat., vol. 42,
pp. 259-282, Io text figs., 1 pl.
WESENBERG-Lunp, C.
1918. Anatomical description of the larva of Mansonia Richiardii (Ficalbi)
found in Danish freshwaters. Saertryk Videns. Medd. Dansk Natur-
hist.. Foren., vol. 69, pp. 277-328, 37 figs.
1920-21. Contributions to the biology of the Danish Culicidae. D. Kgl.
Danske Vidensk. Selsk. Skr., Nat. Math. Afd 8, vol. 7, 210 pp.
2I pls.
WIGGLESwortTH, V. B.
1930. The formation of the peritrophic membrane in insects, with special
reference to the larvae of mosquitoes. Quart. Journ. Micr. Sci., vol.
73, PP. 593-616, Io figs.
1933. The function of the anal gills of mosquito larvae. Journ. Exp. Biol.,
vol. 10, pp. 16-26, 4 figs.
1942. The storage of protein, fat, glycogen and uric acid in the fat body and
other tissues of mosquito larvae. Journ. Exp. Biol., vol. 19, pp. 56-77,
Il figs.
1949. The physiology of mosquitoes. In Boyd, Malariology, vol. 1, pp. 284-
301, 8 figs.
Woottey, T. A.
1943. The metamorphosis of the nervous system of Aedes dorsalis Meigen.
Ann. Ent. Soc. America, vol. 36, pp. 432-447, 9 pls.
ZAVREL, J.
1907. Die Augen einiger Dipterenlarven und -Puppen. Zool. Anz., vol. 31,
PP. 247-255, 13 figs.
Ci ire»
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 9
Roebling Fund
A LONG-RANGE FORECAST OF
UNITED STATES PRECIPITATION
By
Cc. G. ABBOT
Research Associate, Smithsonian Institution
(PusricatTion 4390)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
» ©. MARCH 23, 1960
+i,
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 9
Roebling Fund
A LONG-RANGE FORECAST OF
UNITED STATES PRECIPITATION
By
Cc. G. ABBOT
Research Associate, Smithsonian Institution
(Pusication 4390)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
MARCH 23, 1960
THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.
Roebling Fund
A LONG-RANGE FORECAST OF UNITED STATES
PRECIPIFATION
By C. G. AxBBor
Research Associate, Smithsonian Institution
FOREWORD
A hidden family of harmonic regular periods exists in weather.
The periodic members of this family persist with unchanged lengths
for scores of years. By determining their average forms and ampli-
tudes for intervals of a thousand months, successful forecasts may
be made for years to come; or backcasts may be made for former
years and compared to former events. Agreement of such backcasts
with the records warrants confidence in future forecasts.
These claims seem preposterous to most meteorologists. Therefore,
before proceeding to explain the method and to give forecasts to 1967
for 32 cities of the United States, illustrative forecasts for the years
1950 to 1958 will now be shown and compared to the records of that
interval graphically.
Figures 1, 2, and 3 show forecasts (dotted) and the observed march
of precipitation, 1950-1958. These curves represent 3-month running
means, and are expressed in percentages of normal precipitation. Fig-
ure I represents precipitation at Madison, Wis., and figure 2 at Nash-
ville, Tenn. The curve at the top of figure 2 will be described later.
Figure 3 shows forecast and observation for Sacramento, Calif.
I have computed for several cities coefficients of correlation be-
tween my forecasts and the observed precipitation for the years 1950
through 1958. They are as follows: Washington, D. C., 52.3 percent ;
Cincinnati, Ohio, 57.3 percent ; Nashville, Tenn., 59.0 percent ; Inde-
pendence, Kans., 52.0 percent; Madison, Wis., 56.6 percent; Sacra-
mento, Calif., 69.0 percent.
These coefficients indicate that my forecasts are over halfway to-
ward perfect long-range prediction of weather. There still remain
undisclosed variables that produce the discrepancy of about 40 percent
between my coefficients and perfect correlation.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 9
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
‘suvall SuIUUNI YUOW-L£ WOIJ [[Y ‘2AINd [[N} Guede four] peyop
‘\svoo10} foul] [eWOozti0y AAvay ‘feuION “gS61-O0S61 “uoHepdIoeI1d [eIOU WOIF soInjIedap pee JO JUOAD puUv JSBIVIO “SIM “UOSIPEIY—I “DIy]
a 1S Fs ag
a
eee
Pict
as rT eee
Le — + VAI
be on ae
boise fr bo |p
Very
pou [ere ba
FORECAST OF U. S. PRECIPITATION—ABBOT
9
NO.
‘suvaL SuIUUNI YJUOUI-£ WOIJ [TY ‘dAIND [[NJ YusAs foul] po}Op YseoaroF f aUT] [e}UOZIIOY AAvOY ‘eur
-ION *9S61-0S61 ‘uorepdi901d JeuIOU Wor, sainjiedsp A][YyJWOU Jo Jada pure jsvoIIO “WUaT, ‘o]]IAYSeN—z “SI
6561
£15)
PER
eS6t
C
A
7161
2561 9S61 SS6I vS6l £s6l
zS61 1S6I
pa EM ES:
eee) ere ce
et lb afb
ATUL VA pT
PURE
ae Aer
16! O16! 6061 g06! 2061
SECA UESE SAV ra Et
A
906) S061
Ab ie
| CHEETA “CANE
ROMA CEE CECI
BORE eR EE) Me epi
PU EE
COCCI CTD TAT
NECA CS ONS PAT NENT
SICA
' ACCC NPA
CREE ECR Ce EP
COLOR EEE EEE GO Pre a
MM
NS og
vo6l
SMITHSONIAN MISCELLANEOUS COLLECTIONS
foul] yewwozioy AAvay as "gS61-0S61 ‘uonepdIoaid jeutoU Woy sammjziedop A]Yy}UOUW # JUDAD PUL JSBIVION “Jed ‘ojuswIVIseGS—E ‘91.7
SS osét,
La
i}
t
238 ai aa /
ea
eA Ee
seen HY
of SHE
ain?
NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 5
FORECASTS OF PRECIPITATION FOR 32 CITIES, 1950-1967
This project was sponsored by the Association for Applied Solar
Energy of Phoenix, Ariz., and the Smithsonian Institution of Wash-
ington, D. C. Funds for the costs of electronic computations were
supplied to the Association by the Valley National Bank and the
Arizona Public Service Company. About 7,000 tables of precipitation
were electronically computed by Jonathan Wexler, a student at the
Arizona State College at Tempe. He ingeniously programmed the
machine for this special purpose. Monthly records of precipitation
at 32 stations from about the year 1870 were taken from publications
generously furnished by the United States Weather Bureau.
TABLE 1.—List of stations
1. Abilene, Tex. 17. Nashville, Tenn.
2. Albany, N. Y. 18. Natural Bridge, Ariz.
3. Albany, Oreg. 19. Omaha, Nebr.
4. Augusta, Ga. 20. Peoria, Ill.
5. Bismarck, N. Dak. 21. Port Gibson, Miss.
6. Charleston, S. C. 22. Rochester, N. Y.
7, Cincinnati, Ohio 23. Sacramento, Calif.
8. Denver, Colo. 24. Salisbury, N. C.
9. Detroit, Mich. 25. Salt Lake City, Utah
10. Eastport, Me. 26. San Bernardino, Calif.
11. El Paso, Tex. 27. Santa Fe, N. Mex.
12. Helena, Mont. 28. Spokane, Wash.
13. Independence, Kans. 29. St. Louis, Mo.
14. Little Rock, Ark. 30. St. Paul, Minn.
15. Madison, Wis. 31. Thomasville, Ga.
16. Montgomery, Ala. 32. Washington, D. C.
Secretary Leonard Carmichael of the Smithsonian Institution as-
signed Mrs. Lena Hill and Mrs. Isobel Windom to assist me in
preparing forecasts. He approved grants from funds given for the
study of solar radiation and weather by the late John A. Roebling.
I am greatly indebted to Miss M. A. Neill for careful preparation of
my manuscript.
I selected 32 cities distributed with approximate uniformity over
the United States. The cities chosen are listed in table 1.
THE METHOD
As I suppose no one hitherto has ventured to predict values of
precipitation, at definite places, for as much as 8 years in advance,
I now indicate briefly how it is done. I quote apposite passages from
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
my former papers,’ with slight changes dictated by later experience.
Periods in sun and weather.—The sun’s radiation which we see
and feel, like that of many other stars, is variable. Solar output of
radiation seldom exceeds 2 percent in its variation. However, its
variation comprises as many as 60 regular periodic pulses, ranging
from 1 month or less to 273 months or more. All are exact submul-
tiples (or aliquot parts) of 273 months, as 91, 39, 7 months, and many
more. They range in amplitude from 1/50 to 1/4 percent. All go
on simultaneously, like overtones of a musical note.
As many as 30 of these exact periods have been found in monthly
weather records which have been kept from 1870 and earlier. They
occur in records both of precipitation and temperature. Far from
being confined to fractions of 1 percent, as in solar radiation, in pre-
cipitation they individually range from 5 to 35 percent of the normal
average. In temperature they range from 1° to 3° F., and these limits
refer to 3-month smoothed records. Owing to the large number of
these weather periods, some in plus, some in minus phases at any one
time, their combined influence is not usually startlingly great.
Normals.—Long records of weather ordinarily state “normal”
monthly values found by taking the monthly averages of all the years
tabulated. I have found considerable differences in normals if com-
puted separately for years of high and low sunspot frequencies, re-
spectively. I therefore compute separate monthly normals for years
above and below an average of 20 Wolf numbers in sunspot fre-
quency. From these normals I tabulate the departures in tempera-
ture, and the percentages of normal precipitation.
The monthly values have too wide jumps to be most useful. I
smooth the record by 3-month consecutive means. Thus for February
I use (January+February+ March) X1/3, and similarly for other
months.
Lags.—Supposing, contrary to meteorologists’ opinion, that the
variation of the sun is the real cause of the variation of the weather,
since it has identically the same periods, I point out that well-known
variations of insolation suffer variable lags in their weather influence,
depending on place and time.
Lags of solar effects, as they differ with locality, indicate that the
state of the atmosphere is an important factor. The atmospheric
1a, Journal of Solar Energy, Sci. and Eng., vol. 1, No. 1, January 1957;
b, ibid., vol. 2, No. 1, January 1958; c, Smithsonian Misc. Coll., vol. 122, No. 4,
August 1953; d, ibid., vol. 128, No. 3, April 1955; e, ibid, vol. 128, No. 4,
June 1955; f, ibid., vol. 134, No. 1, September 1956; g, ibid., vol. 138, No. 3,
February 1950.
NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 7
condition varies not only with locality but with time of the year,
prevalence of sunspots, and march of population. To partially meet
these difficulties, I tabulate separately for three periods of the year:
January-April; May-August ; September-December ; also with Wolf
sunspot numbers above and below 20; also with lapse of time before
and after the midpoint of the record. These divisions of the available
monthly data lead to computing 220 tables at each station before
undertaking a forecast.
Forecasts by periods —My forecasts are made by adding the effects
of 27 regular periodic cycles in precipitation. These cycles, like the
harmonics of musical sounds, proceed simultaneously, and are in-
tegrally related to a fundamental cycle. This fundamental is 273
months. The harmonics employed are as follows:
TABLE 2.—Periods used for forecasting
Fraction Months Fraction Months Fraction Months
1/3 gl 1/12 22-3/4 1/27 10-1/9
1/4 68-1/4 1/14 19-1/2 1/28 9-3/4
1/5 54-3/5 1/15 18-1/5 1/30 9-1/10
1/6 45-1/2 1/18 15-1/6 1/33 8-3/1
1/7 39 1/20 13-13/20 1/36 7-7/12
1/8 34-1/8 1/21 13 1/39 7
1/9 30-1/3 1/22 12-9/22 1/45 6-1/15
1/10 27-3/10 1/24 11-3/8 1/54 5-1/18
1/1! 24-9/11 1/26 10-1/2 1/63 4-1/3
The harmonic family referred to was discovered in the variation
of the measures of the solar constant of radiation. Figure 4 shows
26 of over 60 periods discovered in solar variation.” Identical cycles
were later found in precipitation and temperature by study of long-
continued weather records. While the periods of the harmonics are
invariable, both in the sun and weather, and their phases are invari-
able in solar radiation, their phases shift in weather, depending on
atmospheric influences, as will be described below. On account of
these phase changes, depending on several variables discovered in my
studies of precipitation begun with Peoria, Ill., about 10 years ago,
the harmonic family in weather is obscured and hidden, and is as yet
unrecognized by most meteorologists. Nevertheless it is verified by
an enormous mass of evidence, as will appear below.
No observations required—Many meteorologists and others sup-
pose that my method of long-range weather forecasting depends on
solar observations, but this is not so. The harmonic family referred
2 See in reference, footnote 1, e, above, figure 3 and table 3.
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
‘cS61-0Z61 JO SuOT}eAIIS
-qO JUP}sUOI-Ie[OS URIUOSY WIS WoIZ [PY ‘sporsod poyefot A][e1sojUT SulsoJsojUL o}yeurpsroqns Jo poseo[o
[[@ ‘yjSuey ur syjuoW ve - Zo/¢Ze wor Bursuer ‘UOTyeLIeA IejOsS Ul sported xIs-AyUOMT—V “Oy
est Mant Pe ist ak
SOMA TI Nae Naar TP Me
CPR elie tet er Te
ep SE
EPA eS ane
EbS ahh re ee
ere al test | SZ
A Baar Ws at As
are ae Pee | a
eae AP PCa pie
ee EN Ce TE a el
Cod
Pi a Ee NT
CEPT TCE a PSA Ti PW TAA Da’
NO. Q FORECAST OF U. S. PRECIPITATION—-ABBOT 9
to was indeed discovered by the study of over 30 years of daily “solar-
constant” observations of the Smithsonian Astrophysical Observatory.
But now that the harmonic family has been found in weather, no
observations of any kind are required. It is only necessary to employ
a long record of monthly mean values of precipitation, or temperature,
to make long-range predictions. These are approximately verified if
no unusual alterations of atmospheric conditions make the averages
from long records inapplicable.
Sports—As my forecasts depend on the assumption that the av-
erage conditions of the periods over a thousand months will be pro-
jected into the future, it is important not to include wild “sport”
values of precipitation in the thousand-month basis. Hence I have
diminished sporadic very high values to about two times normal, and
have raised sporadic drought values of less than 4o percent of normal
to exceed that limiting low value. These limits refer to 3-month
smoothed records. For most of my 32 stations these changes are very
rare. But in two or three of the desert stations possibly one value in
ten was changed to avoid spoiling the representative character of the
basis. The considerable measure of success of my forecasts is the
main defense of the method used to produce them. If the degree of
success is found to be valuable, no doubt those who in future will
use the method will greatly improve it by modifications dictated by
reason and experience.
Backcasts—Since my forecasts are made by adding the average
effects of 27 harmonic periods over an interval of about 1,000 months,
the 12 months of record for any one year can produce only about 1
percent of influence on the forecast for that year, even if those 12
months are among the thousand months employed as a basis. There-
fore all forecasts or backcasts are equally sound, whether they relate
to time before, within, or after the thousand months of record.®
The preceding paragraph is important. The forecasts for 32 cities
all extend from 1950 to 1967. The degree of similarity between the
forecasts and what happened up to 1958 is the index of their prob-
able agreement from 1959 to 1967.
The 273-month period.—Daily solar-constant observations pro-
ceeded from 1920 to 1952 at Montezuma, Chile. This interval is not
long enough to determine the master period accurately. But the
10-1/9-month period in weather is a strong one and has long been
followed in Washington precipitation. I determined its amplitude for
several periods differing slightly from 10-1/9 months. For this pur-
8 See discussion of backcasts at a later page.
IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
pose I used about 790 monthly mean values of Washington precipita-
tion, all observed when Wolf sunspot numbers exceeded 20. These
values were smoothed by 3-month consecutive means, which of course
reduces the ranges of percentage departures from normal to about
two-thirds of their actual monthly values. Table 3 and figure 5 show
the results.
Figure 5 clearly shows that a value of the master period between
273 and 275 months is definitely indicated.
I have preferred 273 months rather than 275 months because it is
an integral multiple of the strong periods 7, 13, 39, and 91 months.
It cannot be much more than 1/3 percent from the true master period.
TABLE 3.—Percentage amplitudes of proposed periods
Period Ranges
Months Percent
2752 \NO5.7. STOS:4| O25" 100.7. 166.9) 00.3 \, 07.3.1, 107.0, Gk.0U pon 9.4
27
273.0 05:7. 95.8 \ 03.4... 06.1, 00.3) 102.0, 103.7 108.0). 104:8 {tOL.1, (34.6
27
275.0 109.8 102.4 103.3 99.3 95.4 929 962 97.6 988 104.5 16.9
27
277.0 94.6 104.4. 106.2. 10T.3 105.8 105.5 (946.. 97.5 » 96.0. 03:3. 12.90
27
The subordinate periods—Of the 27 periods used in forecasting,
12 exceed 15-1/6 months in length. Owing to arrangements used to
treat changes of phase, which will be described, 42 tabulations for
each city are made of these 12 periodicities. Almost without excep-
tion the curves representing these 42 tables betray overriding har-
monics of the period in question, from two to eight in number. These
overriders must be evaluated and eliminated before the period in
question stands free.
I show in table 4 and figure 6 the treatment of one only of the four
tables representing the 39-month period in precipitation at Helena,
Mont. Eight tabulations of successive runs of this period over the
interval of years 1891 to 1917 give the mean values and average de-
viations from the mean in percentages of normal precipitation. Then
five harmonics of 39 months are successively removed, yielding the
smooth-curve deviations from 100 percent given in column S, and its
deviations from what remains after the five removals of harmonics.
In the final column of table 4, and the final smooth curve of figure 6,
we see the real periodicity of 39 months. The average deviation from
NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT II
curve a is 29.6 percent, and that from curve 0 is 2.1 percent. The
reduction of 93 percent in deviation is due to removing exact har-
monics of 39 months.
Overriding periods—As another example I quote from footnote
I, g, cited above, showing figure 4 of that reference (here figure 7).
T NORMAL PRECIPITATION AMPLITUDE
yp
PERCEN
Fic. 5.—Demonstration of 273-month master period in weather.
From the mean of 16 repetitions of the periodicity of 45-1/2 months
in Natural Bridge precipitation, the true 45-1/2-month period is
cleared of four overriding harmonics.* The reader will note what
similarity to true sine curves is attained in both the above examples,
4 Refer also to the clearing of overriders from the period of 68-1/4 months at
St. Louis. Note 1, g, figure 3.
12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
when overriding harmonics are computed and removed. From the
examples given (out of about 10,000 cases available in my files) the
following 10 exact harmonics of 273 months are exposed as follows:
1/4, 1/7, 1/8, 1/12, 1/14, 1/21, 1/28, 1/35, 1/49, 1/56.
I 3 5 13 17
33 37
Fic. 6.—Helena, Mont. Thirty-nine-month period in precipitation as cleared of over-
riding subordinate integrally related periods. Original tabulation, a; cleared curve,
b, with smoothed curve above. Note approximate sine form. Range, 27 percent of nor-
mal precipitation.
While most removals of harmonic riders are done to clear periods
exceeding 15-1/6 months, many curves representing periods between
g-1/10 and 15-1/6 months required removal of harmonics of 1/2 or
1/3 of their length. An algebraic theorem affords a check on mistakes
of computation when clearing half periods.
Let a periodic curve be represented by equally spaced ordinates
a,b,c .*.': k,l, mi, and, proceedimp tuttuer, 1, Orgy... ok yee:
NO. 9 FORECAST OF U. S. PRECIPITATION—-ABBOT 13
The mean form of the supposed overriding period of one-half
length is:
Gari bite O Cita) sie rai
2 2 2 nea 2 2 2
Eo
&
2
=
3
t
When this half-length curve is written twice, and subtracted, we have:
a—n BS ay c—p k—-sx I—y m—zZ
) ? Ya ar ,’ d
2 2 2 2 2
?
b
and following that:
z=
|
Qa
°
|
Ss
>
|
a
8
a
K=
~
t
—m
eee ) ) -
2 2 2 2 2 2
So the last half of the long curve, when cleared of the period of one-
half of its length, is exactly like the first half, but with reversed signs.
Grouping of periods——All weather influences caused by changes in
solar rays are subject to lags. For instance, June and noonday are
times of highest solar altitudes, but the warmest months and hours
occur later. The lag is longer the longer the period of the solar radi-
ation change. These lags are due to atmospheric conditions, and vary
from locality to locality, from month to month, from times of great
sunspot activity to quiet solar times, and as population and foresta-
tion change. Hence, though the family of periods integrally related
to 273 months proceeds with perfect regularity in measures of the
solar constant, in weather the same family of periods is affected by
changes of phase, depending on the locality, the population, the sun-
spot frequency, and the time of the year. The periods are the same
in weather that they are in solar radiation, but owing to complex
atmospheric influences on the lags the weather phases are so altered
from time to time that these periods are unrecognizable without a
segregation of the data, governed by consideration of these modifying
influences.
It is not possible to anticipate and allow for these phase changes
precisely. I content myself as follows:
(a) The year divided: January to April; May to August; September to
December.
(b) Solar activity divided: Wolf numbers > 20; Wolf numbers < 20.
(c) Secular time divided: first half of tabulated records; second half thereof.
All these divisions of data hold for periods up to 15-1/6 months,
or 15 groupings for these periods. The segregation according to the
Wolf numbers holds from 18-1/5 months up to 39 months, but not
the segregation for times of the year.
14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
4HoKA papas L\ A
‘ ori ea Sa
i A AL Ar
cag
oS Een ae
[Aaa
ko Sue ee eS ad
al)
M. babsad” Pct lh ot Nes cit aP cle |
MeN a NU oes a
Ha WES A Sei Lael etd
Va So fh a tat} |
pao a TION ey, oil aa EN sala 4a
De hy EOMNIIN CE eo
YS A" SE SH |)
YI I aa on NE VAM CTT eNom
pot
eal iv Tg ay a ee alee
YH pec eNIOAIMOD TS e eesleeer ee ee re
yi Pe 0 eh Sa a
si — A Ki gi Aid an ae Sa Wk NA a a
a ff
apap
ERS a aan
HPA A TE
Pen Vee
ad opal che
/\
ena a
TPR al al
ede
Fic. 7.—Natural Bridge, Ariz. Forty-five-and-one-half-month period in pre-
cipitation cleared of overriding subordinate integrally related periods. Range
reduced ninefold by clearing.
NO. 9 FORECAST OF U. S. PRECIPITATION—-ABBOT 15
TABLE 4.—From three-month running means of precipitation, Helena, Mont.
39-month period = p
Elimination of subordinate periods. Interval 1891-1917
Mean percentages of the normal. Original mean and departures after
removing subordinate periods
Average
Original devia-
mean of __ tions.
8deter- Percent X 4
mina- ° Removed periods Final
tions normal (ooo, Smooth ~/8—S mean
a ppt D2 PS iP se ue) 7a PLS 15) A cleared
102 30 o +2 -—9 —I3 —I7 —I2 —5 88
99 40 —8 —8 —I0 —II —II —13 +2 87
92 29 —24 —26 —18 —17 —I7 —I4 —3 86
82 35 —35 —32 —20 —I5 —I2 —I4 +2 86
82 29 —37 —27 —I5 —I5 —I7 —I4 —3 86
89 32 —16 —9 —9 —II —I5 —I3 —2 87
112 34 —2 o —9 —I3 —I3 —I3 0 87
III 31 +12 —2 —I12 —I3 —I3 —I2 —I 88
III 29 —6 —4 —I5 —I4 —II —I10 —I 90
98 38 —9 —II —13 —8 —I0 —8 —2 92
109 23 Onis Ee tele 3 —6 cao 94
109 30 —8 —12 o -—-4 —-4 —4 oO 96
112 40 —I —Ir -+I1 oO O —I +1 99
110 40 —9 —7 —7 -6 -—3 0 —3 100
107 18 Oo Gata Said +2 +1 102
III 19 +11 +9 +44 44 oO +4 +4 104
112 34 +16 +19 +8 +6 +46 +6 oO 106
119 26 +7 +17 +15 +11 +11 +7 3 107
97 34 —13 —6 +2 +1 +4 +9 —5 109
100 23 oO) Ay) 1-0) Ob 7 +9 ae 109
102 27 o —4 +8 +13 +9 +11 —2 III
116 20 +o +11 +11 +11 +411 +11 0 Ill
146 16 +25 +13 +14 +10 +10 +12 —2 112
152 20 +35 +28 +18 +17 +20 aig +7 113
156 33 +37 +33 +22 +23 +19 +13 +6 113
122) Jk 34 +17. +7 +5 +10 +10 +13 —3 113
117 21 +3 +5 +13 +13 +13 +12 +1 112
108 16 —I —I +11 +9 +12 +11 +1 III
123 27 +6 +4 +16 +12 +10 +9 +1 109
117 21 t+i1o +13 +13 +12 48 +8 0 108
109 28 o +10 +1 +2 +42 +4 —2 104
125 37 +8 +15 —3 +2 +2 +2 oO 102
115 26 Spe EA mez) ang a —I —3 99
128 32 ea 5 ft) 97
89 32 =O 7a hyd oO) 4 —4 0 96
90 33 —I0 —I2 o +1 +1 —6 +7 04
81 30 —I5 —24 —I2 -7 —7 —7 oO 93
106 50 —6 —10 —I0 —I0 —7 —9 +2 QI
123 30 +13 +3 —6 —8 —1I0 —I0 oO 90
Mean da 29.6 percent. Mean A 2.1 percent.
Average deviation before clearance 29.6 percent. After clearance 2.1 percent.
Norz.—Thus the removal of overriding harmonics reduces the average deviation by 93
percent. Of about 10,000 such removals of overriding harmonic periods, probably 4,000 gave
fully as satisfactory end results as the 39-month curve at Helena did for the years 1891 to
1917.
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Hence for these longer periods there are about four divisions to a
period. The secular time segregation holds beyond 39 months, two
divisions each for four periods.
The grouping just indicated leads to computing many tables for
each station:
Up to 15-1/6 months, 15 X 12 = 180 tables
Thence to 39 months, 8X 4= 32 tables
Thence to 91 months, 4X 2= 8 tables
otal, i seiacacis ss eauyeasisice 220 tables
Shifts of phases—The numerous groups used for the shortest 15
periods leads to tabulations with so few columns that the mean values
of individual periods are of little weight. To remedy this defect, I
assume that the forms and amplitudes of periods up to 15-1/6 months
in length, and in the same grouping as regards Wolf numbers, will
be similar, though in different phase relations. I therefore make
superposed graphs of the six tables of one period for each of the two
stated conditions of sunspot activity. From inspection checked nu-
merically I am then able to shift the individual curves of the graphs
to the same phase relations. Then I take a mean for all six tables
and use that generalized mean in forecasting. But when using it in
forecasting, I must shift back the generalized mean to the proper
phase, as will appear by an example later. Figure 8 gives an example
of these shiftings in phase.
NOMENCLATURE, SYMBOLS, AND TIME
As stated above, 27 periods, all aliquot parts of 273 months, are
to be used in the forecasts. But, as just stated, these are used in
several groups, depending on the length of the periods. Lags, depend-
ing on atmospheric conditions, dictated tabulations of 12 independent
groupings for the periods of shortest length, that is a1, D1, C3, @2, De, Co,
as tabulated for the period of 9-1/10 months of SS>2o in tables 5
and 6. Besides these, there are six tables a’, b's, c's, a2, b's, c's, for
SS<2o0. However, for periods above 15-1/6 months this extended
grouping brings too few columns into the tables to be capable of yield-
ing satisfactory mean values. Hence for periods 273/15 to 273/7, the
distinction between months of the year is dropped, thus reducing the
number of groupings from 12 to 4 for these 8 periods. For the re-
maining 4 periods, 45-1/2 to gt months, the distinction SS>2o0 or
SS<20 is also dropped, reducing their groupings to 2. So there are
three different arrangements of assembly, as just explained (12X15)
NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 17
= 180+ (8X4) =32+(4x2)=8, making 220 separate tabulations
in all.
In tables of periods 1/18 to 1/63 of 273 months, there are many
cases when the number of columns for @,, b1, ¢1, @2, D2, C2, and a’, b’1,
ae
ean | Mean
Es
if
a
AS
o
vee
Pl cade
i
aa heebre
eee”
SE
Sy
:
\ i
a
on
ays
deg eter
he
agate
RIAs EON
tpi
Cree
ia
repent
A=
oe
Cicer
\
Fic. 8.—Sixfold grouping of periods to form generalized means.
c’,, a2, b's, c’, are too few to give a trustworthy mean. Accordingly,
as I stated above, I have made the assumption that in form and
amplitude groups of SS>2o0 will be fairly similar, though of different
phases, and in form and amplitude groups for SS<2o will also be
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
fairly similar, though differing in phases. Making this assumption,
I combine into one table a,b,c,d2b2c2, merely changing the phases to
give best accord, and similarly I combine a’,b’;c’,a’,b’c’s into a single
TABLE 5.—Grouping of six tables when SS > 20. Period of 9-1/10 months.
Eastport, Me.
First half of the records, 1891 to 1920
7 aia aay ee en Na 2 lg eee PR Pe gy gael am
64 92 55 62 130 158 79 84 90. 92 87
71 47 56 92 164 127 QI 66 85 80 88
79) 54> OB S04 ERO Ce OZ 27" Eo 7e igo gG 08
103 62 98 179 156 Sr 320 69 95 109 108
103 SO) 106} Lr Bivc E10. 80 120 96 = 108 93 108
OI 78 E22 111 TOO 640133 2: 123)... 10s... TOT 108
51 Ja TAA) 20S 55 120 66 127 107 86 94
G0) 47 AEIO 88-5 70 134) 54 TOO Fel A 90
88 59 80 61 81 78 78 79 70 «4116 79
75
Teee clade ke Maced Mises: Recess Tehek- fede d odee soak ts
86 115 72 160 81 132 70 ~=+167 64 105
II4 120 67h 62 PTO rst 76 04 70 105
85 04 134 103 90 128 82." 175 88 102
86 04 164 66 102 126) 103 77 85 100
SOOO SOU 77) 75.) ETA 97) 80. 770 92
G5 ' 04 IIs. 93)) 94. 128 108. | 72/75 94
69 3865 04 82 83°°"926"" "530" ‘100 84 92
75 70 109 Baie tOO. | TIO. (56S 06 86 08
ZU 92 BO 57 AP 05) SE Ta) 178") 04. W203 97
75 78
1891 1894 1807 1903. 1906 1916 191 Means
Sept. Oct. Oct. Nov. ov. Oct. Oct. C1
126). "67 80 06 123 Wg.) VATS 08
100 72 85 04 131 88 104 96
99 7it20)) 1OAy | 1148 95 87 105
105 68 E46. 130 FSH TOO ETO 115
104 So) 150 142) 143) IO TAO 123
117 Sa 147) 128) tI6) Ose 172 124
Bi ii, Zonas es) 80. OOD Onn TIO 95
89 63). \112 70 96 150 92 06
90 73 73>) LES Ae IOT 7134 55 92
table. I give samples of this simplification here in tables 5 and 6 for
SS>20. Figure 9 shows the matter graphically.
The final combinations of two sets of six tables each, with phases
shifted to harmonize, is given in figure 9 and table 7, both from East-
port data.
NO. 9
FORECAST OF U. S. PRECIPITATION—ABBOT
19
TABLE 6.—Grouping of six tables when SS > 20. Period of 9-1/10 months.
Eastport, Me.
1926
Aug.
63
136
145
150
92
85
70
70
1927
May
95
135
185
189
180
178
167
119
oI
1925 1928 1937 1940 1941 1947 1950 1956
Feb. F Mar. Apr. Jan. Feb. Feb. ar.
apes. Adil 7G. AS Tae Tie. Tas
68) 72) (166: 7 so 126 9206) 129
G4, |, 462). (89) 63)). 50) "196 887 (a9
Go 166, (7 75. 68 | TOG 107
83). 109) 63) 426.) 54 3651107, 459
Of, 129 "74, S17)" Ok 134 162) ¥05
03 1201 G5, 128 03) 88" a4. “SS
1240 TIO) 102") oss e1OLN 508) 102) 193
122-87 —. 67-844 | 105° 69). (FT, TOL
1929 1930 1936 1939 1945 1946 1948 1949 1951
Aug. ay June July July May Aug. May Aug.
92) 55° 97 S72).400" 134, 9104: (90 “150
OS -95. 161) 7824 66. 83 0 GAR O77 ah rss
S7ert07) 77.) 73h) WMO: (568-104). 54 130
VA SETOO) “IOT!” WGagets7, (80) 130) 104, 50
20) SS Oa TOSS e7ON 12) 0138) ln 200
TOS, Ate Ite, GhioEsO Fe Tez eT 5s. MES
TOS Sh O04) Ol 127. nila lez Ter i TAS
TOK, 05 0) Oly 1 50k) DO UTIOW eLTOM 1366 107,
bony 76. 52. )79 166 145 ‘108! 13. 66
84 154
Mie itor, Dee Sek Be. Gee; Gee or tne.
TION 972. S25.) FTA" S60 NEO) “ZO | FAC WISE
Ore 76>-nIP—So4-456> 73 | S420 84
Oo S4: “Tee - 66° 950 GS as GE. c08, 965
$E2/7).03. OS 550) 7O 7A, SQu i TOR Oye
965093 54 Gooy 76) 62, S87" 88 "1Z0
Sa )a12. G2 102) 104. 025 «9 8r). 14r sas0
SI Os Ga 10s" "IS1. Tog) | Shiro eras
oP BO ek! | TIOM BST) OS) GOR Io ae
7 OS BO. ETF... TAG), 04, ST 22 TAG
112
Second half of the records, 1925 to 1956
Means
aa
84
86
86
104
112
103
97
95
1952
May
130
12I
131
99
105
81
95
110
136
1955
June
60
80
87
65
113
102
126
Means
89
86
87
107
107
106
103
The meaning of the symbols on figure 9 is as follows:
ok, no shift.
T , shift backward.
) , shift forward.
Subscripts, number of months shifted.
Means
ba
97
105
116
115
108
108
101
102
20
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 139
TasLe 7.—Phase adjustment. The 6-1/15-month period
Division = Time before and after 1900.
Category = Records when Wolf sunspot numbers
Phase shifts indicated:
Basis of forecast, over 1,000 monthly records smoothed by 3-month running means. Forecasts employ 27 periods
all exact submultiples of 273 months.
= 20.
ok, 4.n, YN, drawn dotted below.
Phases shift with changing atmospheric conditions, but periods remain, and are of the exact lengths found i
solar variation. It requires 220 tables electronically computed to make a forecast for one station.
a1
105
102
100
106
105
103
biok
95
95
94
92
103
114
CatTEecory 2 ASSEMBLY
cifs
108
106
95
azok beok cook Mean
100
93
92
97
106
105
96 95
97 88
93 80
94 88
100 91
101 99
Fic. 9.—Phase shifts
100
97
92
97
101
105
6/593
99
A
+1
—2
az.
—I
+2
+6
CaTEGoRY 1 ASSEMBLY
as biyi ciok azok bef2 cook Mean ZZ
120 95 100 105 96 117 107. +
108 86 ~=100 gI 940) 110 98
110 86 100 90 92 100 95 —
92 97 102 87 84 88 gt —
105 gt 104 gI 90 93 93 =
110 98 103 102 94 97 IoI +
6|586
98
2
in sixfold grouping of periods.
NO. Q FORECAST OF U. S. PRECIPITATION—-ABBOT Zi
Times.—The growth of population, destruction of forests, multi-
plying of oil engines, automobiles, and airplanes alter the properties
of the atmosphere and thereby shift phases of periods. Hence, as
stated above, I divide the thousand months of records into first and
second halves and compute the phases and amplitudes within the two
parts separately.
TABLE 7a.—The sixfold groupings.* The 9-1/10-month period. Eastport, Me.
Values in percentages of normal precipitation
A. WoLF SUNSPOT NUMBERS BELOW 20
a4 b1 ok ci 4 a2 ok boY3 c2 ok = =+6 A
101 83 94 90 63 106 537 89 —9
88 88 oI IOI 66 107 547 90 —8
88 79 98 107 73 107 552 92 ae
75 82 81 108 78 100 509 88 —I0
97 97 93 110 80 106 583 97 —I
104 95 III II5 QI 110 626 104 6
106 III 112 105 109 122 685 114 16
106 107 109 118 98 116 654 109 II
IOI 114 97 107 76 109 604 IOI 2
Mean 98
B. WoLF SUNSPOT NUMBERS EXCEED 20
a ok bil 3 car az ok bz ok cof2 = =+6 A
87 92 92 84 97 89 541 oo 10
88 98 98 86 06 86 552 92 —8
08 97 96 86 105 87 569 95 aad
108 105 105 98 116 107 639 106 6
108 105 II5 104 II5 107 654 109 9
108 102 123 112 108 106 659 110 10
04 100 124 103 108 100 629 105 5
90 92 95 97 101 98 573 96 4
79 94 96 95 102 99 565 94 —6
Mean 100
* The shifting of phases is indicated by arrows as in figure 9 and table 7. The accompany-
ing subscripts indicate the number of months shifted up or down.
Not only so, but considerable differences of amplitude between the
two halves are sometimes found. As forecasts are for present and
future time, weights, as 2/1, 3/1, or 4/1, are given to favor the second
half when considerable differences in amplitude of periods between
the two halves appear. It matters not whether the later amplitudes
are the less or the greater, the larger weight is ascribed to amplitudes
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
of the second half. If a backcast were to be made to long ago, the
weights would of course.be reversed.
At some chosen date all periods must be in the same phase and
preferably in zero phase. I chose 1957-0 as this zero date. To insure
that any particular period will be in zero phase with 1957-0 it is
necessary to compute ahead from the start at about the year 1870.
This may be done as follows. Take the period 8-3/11 months for
example.
From 1870 to 1957, 87 years, there are 1,044 months. About 126
periods of 8-3/11 months would cover this interval. But a date must
be chosen which is an exact integral multiple of 8-3/11. The nearest
is that which gives 121 periods in the interim. Multiplying, we find
that 121 periods require 1,001 months, or 83 years 5 months. Sub-
TABLE 8.—Repeated 8-3/11 months and round numbers
I 8.2737 8 7 57.9089 8
2 16.5454 9 8 66.1816 8
3 24.8181 8 9 74.4543 8
4 33.0908 8 10 82.7270 9
5 41.3635 8 II 90.9997 8
6 49.6362 9
tracting these figures from 1957-0 we find 1873-7. Thus a suitable
starting point is August 1873. But it was assumed that the record
begins about 1870-0. If so, 43 months would be lost. One therefore
counts backward from 1873-7 five periods, and therefore begins with
March 1870.
We now come to considering periods ending in fractions of a
month. We may make tables of accumulation for them. Again using
the period 8-3/11 months, table 8 results.
For most of the periods of inexact months, tables to 91 months
suffice. But for such as 12-9/22, 13-13/20, 24-9/11 and 27-3/10 the
tables must be carried on to 273 months.
RESULTS OF FORECASTS
Having treated of most of the features of the method, the re-
mainder of this paper will disclose the results of these forecasts of
precipitation. As I have stated, I discovered discrepancies sometimes
as great as 10 percent between the published monthly normals and
new normals obtained by separating years when Wolf sunspot num-
bers are respectively above and below 20. As my new normals may
NO. Q FORECAST OF U. S. PRECIPITATION—ABBOT 23
be of value to other investigators of periodicity I first give in table 9
the two sets of normals for the 32 cities I have investigated.
The cities are in alphabetical order. The months in the first column
apply for all cities. Precipitation is given in inches. Columns A and
B give monthly normals for times when Wolf sunspot numbers are
respectively Jess and more than 20.
Departures; observation minus forecast 1950-1958.—There are 20
cities showing (1950-1958) departures in level of 4 percent or more
from the values given in table 9. This is to be expected. One could
not suppose the mean precipitation, 1950-1958, would be identical
with the average precipitation, 1870-1958. Table 9a gives all the
cities where such differences of 4 percent or more occurred.
When I come to give tables and maps of forecasts, 1959-1967, I
shall not use table ga to correct the maps, but shall quote the results
as they are determined from table 9. Persons interested may apply
the values of A, table ga, as corrections in level to the forecasts, using
them in reverse of the signs given in table ga.
Sunspot effect on normals.—Lest readers think the differences
between mean precipitation values attending high and low sunspot
frequency are merely due to the sparsity of evidence, considering the
irregularity of precipitation, I call attention to the numbers of months
entering into the mean values of table 9. For nearly all of the stations
approximately a thousand months participated. That indicates about
600 for high sunspot frequency, about 400 for the low. Dividing by
12, there were about 50 values per monthly mean for sunspots ex-
ceeding 20 Wolf sunspot numbers, and about 33 per month for the
low sunspot frequencies.
Referring to table 9, the yearly sums show seven cities where sun-
spot frequency makes no more than 1 percent difference in the totals.
For seven other cities low sunspot activity brings more precipitation,
with an average difference of 5 percent. For the remaining 18 cities
precipitation averages 5-1/2 percent higher at high sunspot frequency.
While the discovery and elimination of these differences by computing
new normals was of importance in my forecasting, seasonal differ-
ences made the elimination of the sunspot effect imperative. Thus at
Salisbury, N. C., precipitation averages 17 percent higher with low
Wolf numbers, January-April; 9 percent lower, May-August; and
I1 percent higher, September-December, for Wolf numbers below 20
than for those above 20.
Credibility of forecasts.—It is difficult to compress within the limits
of a paper, aimed to be available at moderate price to all who desire
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
o'oS 1°08
GV ea
Te SeeOles
Oe eee
OF Stee. (SHS
oH (ORY
O:Se ecag
Tisva. aeGs
Oe SOS
Oia Ev,
zo Og
To SS
CSE ean
oe
Baoan
Cid ese
cio, 74:0
go SO
Or go
ons 2:0
Ve eT
Lie sOsr
Coteau
Vite a
Tie Qual
Ogleeeeiek
So 86: So
So) = So)
A
“O]O)
‘I9AUIq
a Vv
TelGeec.O%
Co TeeK
Cit Oal
Cem cae
iC e Ore
Cece Cae
St Sa
Baie SS
We = Oe
(EEG ANITA
Oe lke
Cote val
QI Ol
eee
‘STM
‘uOsIpe I
ogo Vze
ieee Oe
Ke Aw
oC ae
ge 92
OC SIZE
Vibe Gi)
OS Se Lie.
Cm CLe
OS Os
Clee eV,
of 9g2
O:Comege:
ee ee
yeutioutg
@ Vv
rb ov
Sf CY
Ce ae
Tee ele
Gig Sane,
Gite ose
Ee Seis
os 012
O77 ail
O17 = 05
Cove Tes
OAs
oi VV,
uu ——"
AV
‘O0Y eT
ov ov
Teer =19ic
ae ese
Co 29'S
Cie eS
TO eas
co) e:9
gk LV
1s e012)
Giggs
Coe ice
Vie ls
Ge eae
Sng
cen
a Vv
o'9f 69f
Hl Oi
OZ OZ
OC mn gic
Oe Gy,
CiceeOre
Litre Se
vS gt
aba ca
Sei OW,
Vo Ve
Sel meee
Orley cal
—_
“SUBYT
‘gouapuodapuy
oor SOI
So 6FO
OO {S10
go.) 6(Ol
ela bal
Opten Zeal
Creme eae
Te :S
Leva Vacs
Orig 1
60 660
So 60
co +o
poe
tgoreustg
a V
ViEr Vit
Zo Go
Zi0 9:0
ZO O10)
Bl O10
Zo gO
Buy — (oko)
Vice Ore
Cie ike
60) =O!
JEL ASK 0)
go SO
go 690
US
‘UOT
*eusjo ty
grb ceV
He == ie
ge Ve
IA Re
Vieowee Ta:
6r rT
rs = gor
Ci Sav,
Coe Ric,
PGi fe
ye YP
1 OFZ
Lies Vaes
e+,“
eH)
‘ejsnany
a V
00g = gS°Z
4yo v0
ZE-0 gbo
gZ0 6S°0
O0'I gQo'l
gSI oP!
OZ41F LOL
vrvo § gg'o
Zz0 OO
Izo £20
gz'0 €z'0
cto gfo
gro «ff
eee
“xo TL
‘osed 1a
ob for
rZ 9°9
(BO) ON
Zot: eat
Len eel
VO =S:0
Vi00€:0
Cie nel
Isv wee
Oe L2
Cat Vv
oS -Vy
£9 39
ee“
"3210
‘Aurqiy
a Vv
‘oz < JOqunU JOM — g ‘Oz > JOquINU JOM = V
sayous us LO6r ybnosys OLgr 4azfo uorynjrdr9a.g Cjyzuous joUu4oN—O TIAV,
6OSe ov
ace eOie
Oe) ee
OC aL
Ci Ore:
oitg ets
Laan Oi
Li Cae Oe
Qe Se
LE AGS
6a Sv
AG one)
Goa Ose:
——
sUle
y10djseq
ose ce
CS hae
Ogee Lic
Oi GeO
PC cae:
Goo ae
SE SOse
Oper cee:
O;Cameese:
+ a
Se
Ate A
WG VPA
anes
neeany
a V
6'1¢ Orc e@eoceesee
fara Ate
Vice,
Cie) eae
Qe Be
Se Oe
ToC ee
Gs “6iz
VGemavac!
gz Ve
Ci
tA Ake
CeO
=a
osteo
Gee ze
Ove Vi
OW sen
Vive ae ne
iGee lee,
Orcas Sol
Olea Zr
Oi Ore
OS SV.
Bio oaseLic
Tel Sisk
9:0) Ss1:t
60 £0
UY
*xoL
‘susTiqy
a V
.
.
.
IeoK
Jaquia.0q
JaquiaAON
* Jaqopo
Jaquis}das
"* 4snsny
eeeee Ayne
eeeee oun
ART
eeee pudy
"7° prey
Aveniqa.t
- ATenuel
esee
Ieax
Jaquis.a(qq
JOquIsAON
* 33qg0O
Jaquis}das
°° 4snsny
eeere Ane
eoeee oun
Ae
ceee [udy
teers preyy
** Areniqay
so ATENUL |;
25
S. PRECIPITATION—ABBOT
FORECAST OF U.
NO. 9
Gob rer
Oise
ge ge
Se AS
APL iy
Siva,
ib... EF
ge oF
OGe wit
ice bee
ee I+
Gizs. biz
So oe
ee.
Santen’
zob S'gh
coe AV
Oise tec,
lojra (OSS
Lb = ONS,
OS eles
oH 4a
oy tv.
Teenie
to eh
OV ASS
Ca,
Loa Asie
eae
See eies
a Vv
S1S gos
of oF
QS 1S
fz gz
gb LY
20 a a)
39. 89
$7. es
gf — VE
Oia cre,
or vr
ee VAY
OVS
e_
20
‘ay[qaseuog L
e241 gSI
GE -™0'2
QI gi
go £0
ZO 10
oo O70
oo 600
TO 22.0
go SO
Sl SSH
GZ 22
of Lz
Geet
e~,-—
“HTeO
‘oyUsMIeIDeS
a Vv
gle Ose
OS O10
Sore Soa
Li Ole
1 Set ae
SE Se
Sige wie
ev ov
CS. eS
CCCs
Oa ren
OI go
Ow 610
— ~~
“UU
‘neg 3S
Qe oz
Vivier Lie
SZ 2012
Li mec
Via = Oe
LiZ aaa Gie
Lele eS,
ge L2
gz SZ
See Lie
gz eae
ies
Oa Lic
SEE ane
‘A'N
§194SOqOOY
a Vv
6ge oe
V2) V2
61s) Siz
OZ Sie
Te Cenc one
Vien’
Caen cis
Sivan Wav.
Site av,
©) Ceacne
Cibeeseaes
Se Ne
Giz OLE
e+
OW
‘sino’y 4S
Ges ors
QV ies
Ic vac
Oi Oe
Gig are
ov ve
Le hay.
See SY
i? — (OAS
Sve 19
Cae cs)
ZC SeeOS
CS GE)
nd
"SSITN
‘uosqiy 10g
a Vv
Sor SI
Ate tA
Oe or
Coleaevel
60 60
So £0
go g0
Vale Sen
Vale cal
Tate StOnd
Seba torr
Gulaemmeal
OZ. “OS
ef"
"yseM
fgueyods
OVE OFF
QuleemOrl
LiGa Vie
Oe) BVT
7 GUS:
E Zo ae
Di Common
ices
Ohm BaAv,
Tocemcae,
On eZic
Cul matal
Semantic:
Veg oe
at
a V
Vac leer
Zo £0
COmeZ.0)
Ol] 20:1
Oplameayal
Ce ONS
Tee VAes
OO weer
Caso
OT AiO
go 6390
LO a0
LO a0
ese
“xo WN
‘Oy ByUeS
O2Ze 1:92
2:0) Our
Cul Or
re men Aea
OS. 1O'E
OS TEE:
Oe 1ee.
= Sev.
C Cacie.
Loe ane,
Lol Ou
60 8 6g'0
go £0
ese’
"Iq9N
‘eqewoO
a V
VeSteeeiat
Size hace
Gols Hat
go 690
TO oO
TcO}e iGO
oOo 86©0'0
TOme co
go vo
Cote Ser
LiGeeVac
EE Ze
Cio Cae
a a
soiAne ues
ySzZ giz
ge ge
Si gt
Opkame deo
Wave Ou
Lee Lie,
So endie
0s) 0
go so
Ovlee tel
Sa = tats
Sz ge
Oe. Ze
ee,"
“ZW
‘asplig [einjeny
a
Vv
ocr gv eee
Sel aval
Val eee
Velour oul
60 90
go 40
coe a.0)
go 630
Oe Oe
(ope (oP it
Out Out
Vale nl
Val mou
ey ,-—-——"
qe
OHeT HPS
gtrh 6:or
OF SPV.
Lo Oe
(Ot A
of VE
OS ae Oe
Os SOW.
Sts OS
LBL es
Oi) Vy,
O;S) cis
Tay
Os = O17
ee —_
‘uuay,
‘aT[LAqseN
a Vv
.
tes eax
Joquiss9q]
JIqUIdAO NT
* raqgopo
Jaquis}das
"* ysneény
eens Ain[
eeeee aunt
eeee ARW
eree Judy
-e° OIC
Areniqo 4
* Arenuel
Sire tea
Jaquiao9q
JaquisAoN
* sraqopoO
Jaquie}das
‘* 4snsny
ee eee An
eeeee sunt
eens AeW
eeee Judy
eee eee Yoel
eeeee Aren1qa.y
eeeeee Avenue f
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
it, the results and comments representing this project. Even with 32
stations, the United States is so vast in area and so varied in con-
trasting conditions that with the fullest use of my results no adequate
country-wide coverage of the expected precipitation to 1967 can be
made. As stated above, confidence in the forecasts must depend
largely on the fidelity with which the first half of the forecast, 1950-
1958, inclusive, fits the observed record.
Table 10 presents in parallel columns for all 32 stations the monthly
percentage departures of forecasts and observed records, 1950-1958,
from the normals given in table 9.
That readers may see from a graphical standpoint to what degree
the forecasts represent the events, I present figure 10. It gives the
march of forecasts and events from 1950 to 1958 for Cincinnati, one
of the best, and Denver, a less favorable station.
TABLE Qa.—Percentage departures (O-F) 1950-1958, from table 9
GHEY Festi ate fea e's oo Abilene Augusta Bismarck Charleston Cincinnati
TA. aes claie atayele Hass —I2 —I7 —6 —Il +4
CIE Yair eciorers oieiale Detroit Eastport Helena Independence Little Rock
Gi UA arora musics doe oce —4 +23 —II —I7 +4
Gaby ice vals se arta Natural Bridge Peoria Sacramento Salisbury Salt Lake
Te NW sre etcintateieavere —7 —6 —4 —5 —7
CH ere a elec nid ais San Bernardino SantaFe St. Louis St. Paul Thomasville
GN ie wana oniermss +10 —I17 —8 —II —9
Figure 10 shows for a more favorable and a less favorable station
a graphic view of data taken from table Io.
A glance at figure 10 shows for both cities an obvious similarity of
the features of the forecasts and of the events for the majority of
months covered. There are, to be sure, differences in amplitude of
features observed and forecasted. In many cases the forecast, built
on average conditions of about 1,000 previous months, hits the fea-
tures found in the observed record from 1950 to 1958 on the exact
months. But in the better station, as well as in the worse, there occur
relative displacements of features common to both forecast and event.
These displacements are rarely as great as 5 months for any station,
but may extend through durations sometimes as great as several years
before returning to agreement.
Displacements of features.—Several years ago I published the ac-
count of a forecast for 104 years of St. Louis precipitation, including
a comparison with the observed records. I quote from my discussion *
* Text continued on page 44.
NO. Q FORECAST OF U. S. PRECIPITATION—-ABBOT 27
SRS Reeeees Sees
HA SO EERTn Ko
wt LA A Fae ee
CICA HEIN. ACC UANT He
“ATTEN RC
CECT ASN a tots
“ARECACEAE PT
ie My a) gis Wiech Ny ella ale
(0 a la a i
HHA-R ET --L
eee LaLa a
ACEC CREEE EE
FLAY LUT ET
OG
Be Ca SETA IP
SEEEEPCECEP AVE
eee al ele Walaa ti fi 1 ens a
HeLa oele ee PN ol AS] 2 RPT Ea ee
feet ttt ji
SUNG a Wa dt ae lai ell
1952, ps7
Fic. 10—Comparison of forecasts and events, 1950- Le Upper curves, Denver, Ooi: lower
curves, Cincinnati, Ohio. Forecasts, dotted lines; events, full curves.
oi a
Pe nS ag
ECO
THiS
POPPE EF
Percent NorRMAL Precipitation §
= +
ir
nr
al armel
aa Sel LS
eee
f1— _ vir fer f+ ovr Z1r w— Z1r ght “28+ 41 “1 ci— 66 III oO1+ 6g £Z gz— 9. OI
4v— fg EI ge+ gvI oz Oov— oor ofr 69+ f12 hi Gi— gg Zor iIf+ 101 of zi+ Zor 6
gOfe— gor cvi ogI1+ 6S1 Chr zI— Sg £6 OI— -SS1-- 121 of— Ze €zi vi-- 901 26 of-+ ter v6
Se-+ 641 Si c— Sr of1 E1-+ gg SZ el— 4g 651 ¥S— 96 OSi=— VE-- Fri orl, “17-4-— ori ~ ‘Sor
Si Ol ral —O— “OlI eel = ic) $6. 72") -6y— 08). 06. 9S—. 66 ~ZS1 —- ZE-- oft ~=6 G66. oS 651 €or
ge+ ggrI zor Ib— gf Zit Zi-- 10 62 of&+ 9gzI 96 ty— zg ack, I= 90T= GOL gf--= ori - ‘gor
4I— ctr Ori If— a9 16 VI— gf z6 vzi+ 991 ey f— £8 06 es— a S6 ve-+- &1 601
AC /0GTs <ctl = ve— oS £8 zI— 16 for g+ ggI 18 f1— 2 28 fo— 1 94 GI Se
ov— 16 ir 41— €£ 06 ov— 2 611 S1+ oor &@g If— £09 140) g—— ZF SP ZE— Zor ri
of— Zor thr z— ZL 62 gI— £3 ior be— +S 98 og9— oS Oltemsecspeule v1 or— 66 SrI
v— QO OI1 II+ 9g SZ 6f&— zZ Ill vy— ov vg Gr—> £9 £6 gz+ LL 1S SI— gir €€1
eI-+ ofl gII ge+ 16 £9 Ze— (9S 83 Zi—- 02 £6 oz— gf 36 9—. 19 2g Or ae cole cil
ge+- oor ocr 25+ fer 99 oc—— LZ 46 f— | €or, (901 z— o£ 46 ce-+- Zr zg Ii-+ er 21
AY Oct cll clo OO lee ne --— OZ 92 Ape lete Onl 16 —e 04 10l> =I¢-|> COl 19 Toe cole col
Scie) cele e ROL. O-- grr —6€1. ze—: £9... 63-1 -0 16 16 os— 19 [ol=-Si—=. «Olt 1 FI ei ET
6&— £2 O11 z&— ozri 2S1 gI— Sg ror €1-+- £2 v9 OV=—) JcOL. Ast. f= SVi- Gyr= f-——=901 “TIT
St gol oor 6+ O6rr OI oz— 96 Q1I f€1-+ 9 vZ ae— COL yi vr Ovi (Sie ——=- ta £3
6s— gz Sg wvi— Szr 6&1 of— 6o1 G6€1 o1— £6 fol of— 9g Yel. -OI—— Sor {iZte Src r- GOL
61+ 69 os Z- S€1- ger--Se— vor “o6gr - S— =~ Seg 06 VI-F gel” Vir> “ee— "er 0vnze--~ get “vor
£S-= 06 Le c= Sil -0cl— “I= S01 011: 61 —-..£ ZZ Z— 96 601 L— 0141 —-9+-- O11 G01
WS-+- z2el 99 f+ g1r Sir v&— 06 vZI gI— 99 tg gs chr 9 6f€— SZ VII zy— og cZ1
zot oS1 22 re-- £15 2g QOI— VOI OzI . Z€— 99 gII gi+ 2 9s Sl=—= V5) 69 99— cZ gel
g1+ 46 1g LE OQ 1S gf— 99 WOIS SI5-- 921 1221 (90--: oor. ee 6f+ 18g ev or— gf Lz1
zS— 12 €zI og+ o1r oS C—O Sg za-- Sgr €€r eet ¢¢ 0z Te-|-= Ole oe iS3— €or v1
eee cee OSL ehOr S2oi-- Sp ergh—= 0S - gy 810-4.) Sie ters 6 — —.{86e> Op oo45 e699 = — ES1- “Ser
zs— £0 OI SI0-- esr lithe ic" Oy = 00L,__ oe-|- (68 Cir oz—__Sy Sg Vet Zr ‘gol O1--. Zer -Z11
V PpeAtos yseo Vv peates 4seo Vv peAtos yseo Vv pears 4yseo V PpeAres ysvo Vv peAtes seo V PeAatos yseo
"qO —--240q "GO 40 qO 0g qO = --10g qO = -A40T -qOQ --840 qQ 40g
0199 23: (6) Es. "xed ‘N "ey ‘3210 AN
‘roAusq ‘IyeuuOuId ‘uojsapieqa ‘yo1eusig: ‘ejsnony ‘Aueqiy ‘Aueqiy
V
QS6I-OS6" ‘uo1uDA4aS5qGO PUD 4SDIZAO.J—OI TIAV ],
61+ ¢Z 9S
Sp— Lb tA)
bL— €P ZII
6z— SE tg
11+ ¢V ce
be+ re oO
c— I ov
SE— —=S9 oo
oL— 66 691
cZ— 6 ZZI
Gi 08 Se ut
Ze— ZL VIL
c4— SS¢ O£I
Z5— "Te gzI
zoI— € Cor
cc— z ZS
f+ 1 gt
6S-— Gfr 0g
€zi+ Sze zor
6£1+ foz 9
Cl+ ofr 2
te+ 101 LZ
oz— 68 601
Tee 601
e— SS LZ
I— 06 16
V peAtas yseo
-qQO -a10q
‘xoy
‘auaTiqy
ee:
owe
‘Qa
‘uel
eS6r
29q
“AON
"0
*¥d3S
“sny
Ane
oun
Ae
“adv
“ey
‘Qu
‘uel
IS61
‘29q
“AON
‘OQ
39S
“sny
Ayn
oun
Ae
‘Idy
“Te
‘eu
‘uel
oS6r
28
Si+
II—
61-++
cc—
=
gze—
oe
c6
bZ
€Z
oS
€Z
1Z
Iv
be
62
ZEI
efI
LS
1g
£11
Lz1
gol
zol
Cor
LEI
£11
601
£8
ZO
ZS
tas
ZS
99
62
99
ze
re
Lz
LL
zZ
62
06
LL
bg
zzl
6r1
QSI
OOoI
LL
$S
48
zSI
€St
OgI
6z1
601
€Z
<8
£g
gil
OIl
8e—
€c+
ve
9e+
ge-+
{p+
ge
eet
zi+
vr
f+
Zz+
co—
6I—
LL
tzI
OzI
OgI
ZOI
Se1
£6
QoI
Vz,
62
oZ
69
09
zs
ev
gt
9S
z9
66
FOL
o£
/
a
FOI
zo1
L£z1
6£1
Ogi
ff1
€9g1
991
eS
Sol
gL
LV
SP
ve
FI
£z
os
ss
<9
LL
€1I
€ZI
1Z
98
VII
gti
gsI
ZgI
cI
Tot
63+
$£-+-
zg+
6ce-+
oi-+
oc+
I1+
e1—
9+
ZI—
oz—
1I+
be-+
€b-+-
or-+-
Z1+
III
IzI
PII
voi
oZ
£6
ZO
£0
£6
PIT
Tz
Lb
zs
GZ
PII
ZEI
gz
gol
ZOl
FOI
601
LOI
96
gol
08
14
1Z
OOI
48
00
101
gZ
c¢
ZS
GZ
z6
ZI1
ocr
ci+
oz+-
69+-
1é¢+
Se+
Gz
$8
fer
LL1
PHI
gII
SZ
bZ
gs
bg
£3
FOI
90
/
a
29
8
cz
CZI
sgt
6S1
Val
69
ov
OI
zz
Se
ov
oZ
6c
€or
96
ZO
OgI
691
tor
£OoI
£6
ZZ
9
9
VI
eZ
1Z
QZ
16
£Z
8
QZ
ZLI
g0z
89
QzI
£6
6
$9
36
62
of
1/7
(panuyuo?)
6—
s—
e+
g+
1$-+-
ge
1z+
c—
ZI—
€r+
I—_—
git
Lo—
vi—
zI—
eb
Ser
oI1+
zp
og—
zsS—
61+
Le—
eo—
oo
Sle
og —
zg—
7O—
+6
69
6S
Zs
69
88
£6
58
8
bZ
£9
£11
Zol
rg
69
£8
ZO
QZ
10)
$21
Col
ZI1
gol
QII
£OI
LL
ZS
br
gi
<9
cL
£8
+6
19
89
S6
IOI
zzi
Cc1
PYI
QSI
66
ZOl
cil
£11
£6
IZI
LZz1
Ori
thr
ce
gz+
09+
og-+
Sra
4i+
zI+
be—
Sk
6I—
re—
£9+-
49+
ori+
bl+
6£-+-
yS+-
oc~—
Oca
9f£—
cc—
f9—
€1-+-
gar
fies
QZ
vol
ZOl
££1
OZI
Zz
ZQI
CzI
POI
QII
6z1
zel
QII
£8
QzI
10
IZl
£o1
QZ
Vel
£6
fer
Ifl
Zol
6r
ve
6z
1Z
46
©
OTT
16
601
£1
IVI
IQI
641
OfI
oS
IOI
AU
gSI
QfI
Cer
LVI
CZI
gS
of
QI
oS
gs
62
ISI
O£I
CeI
611
IZI
£8
rg
rg
C6
€r1+
oI—
oI-++
gf-+
gf+
Se-+-
Lz+
1Ii+
cc—
bi-+
ee-+-
ge+
ec+
e+
ote
nz lL
IOI
oll
48
06
98
£OI
tel
Tl
For
POI
C11
16
+6
S6
/
Z9
PEI
PLI
£1z
L4YV1
LEI
IVI
Ori
oI
£8
89
ZZ
c8
gor
ACT
I0I
Sor
£6
98
1g
98
€Z
Ig
OII
For
gor
QzI
QzI
QZ
or
or
£or
Crr
gri—
6g—
ee
ee
3o—
Sor—
SoI—
1Z—
fgo—
co—
1z+
bi+
gi
z
ci+
nate
ge+
1+
of+-
99—
Lz—
ee—
£4L—
6€-+-
ci+
88
zs
09
62
98
SII
Se1
6v1
140)
0g
99
QV
LV
Ori
evi
QrI
ee
SV
£1
ve
ec
gtz
Zg1
ZzI
90
OOI
Gz1
ZgI
SSI
£6
gz
ee
of
Z9
9
£8
LL
vzI
6Il
OgI
Lol
66
101
PzI
Zo1
Cer
9S
of
Zz
89
‘29d
“AON
pO
ydaS
‘any
Ayn(
aunf
Ae
‘ady
“we
‘Pu
ze (i
Poor
29d
“AON,
0
33S
“any
Ang
oun
Are
“aIdy
“Ie
‘qq
‘uel
EL61
229q
“AON
‘0
*ydaS
“Ssny
Ane
oun
fLorat
29
cht ger 98
0 vit 11
6r— SZ vz1
os— ¢¢ €o1
Ir-+ gzi 8g
Sy-+ Sf1 06
Z2-+ tz &g
€1— +S 19
of+ 2 gt
gIi+ 638 14
6h+ gor ZS
oc— &§ CoI
o9o— 9 LZ
¥g— 19 1
QI— OzI gtI
oO gris gv
6h+ Sgr oOf1
ce+ “Zvi ell
gf+ €z1 Sg
fze— €2 96
6€— 94 ZI
9— 101 ZO!
QO+ II OII
o+ “411 III
Vv peAtes 4yseo
"qO 10g
‘fo
*raAuad
obp— 8 vz
41— vZ 16
SI—_ €9-—s«g
SI— 94 16
I1— +6 S6
S+ 101 96
II— oOzi_ 1€1
ZI— II O€1
€+ ez OI!
or+ ef1 £6
ve+ ozI 98
OI— z6 zol
1z— Z $6
oI+ vir vor
gb+ egi F£1
oz+ Sgr SOI
o— Zr €gI
gS— gir vZI
yS— gO1 OO!
gr— 3 zel
4S— 601 991
ve— SI géZI
6— O6br QSI
gt 6€1 I€1
Vv peAtes yseo
"GO -940
o140
“1qeuUlDUI?D
or— FS +6
1S— 6S O1l
6f£— 9 £o1
Sh— £2 gII
oz— 68 601
Ze— £9 VII
Zi— +8 IOI
zz~— 98 gol
Sc— 1g 901
1e— €2 Vol
Ol 83 g6
6— 2 £g
€I— 99 64
Gia 16 98
6+ zo €g
gz— 64 Sor
tz— 3 gol
z— 146 611
Ly— 624 QzI
goh— 12 ZI
a= “90 IZI
Sain. “09-98
Izc+ +6 €Z
of+ Sg or
Vv pearas 4seo
"GO 210g
TS
“ag
‘uoysa[1eyD
Le+
c1—
gI+
er1+
6+
ee+
bi+
ti+
Vv
¥9
Lb
19
£Z
88
62
cZ
+6
gz
sgl
IvI
SZ
94
gII
gzI
Gzi. gor fo—. 19 vin “Zi— Sh. zét v=. £6 246 S+ 69
oIr Ser So— #S.- 6&1 oc— 6 tir Se— 16 gir i= 2
601 £6 1iZ— 92 6FI gIi— 4g Sor 9g— 06 96 c— 2
62 62 oS— 98 oft, £—- S0r “gor S-{- 0g Ig Sz— of
601 96 z&— 9f GOL, ci. Z vZ {1+ 46 9 gr— &zZ
O1r o1r “Ze— zs 62 aod £9 1s— 9 QII z2Q— gz
gzi_ 6 ce— ww 62 ZE+ 39 1¢ L6— V2 141 I19— gl
88 vZ €1— £9 gf ee+ 62 gt 6s— 9 zyr vE— gf
601 g6 f— gf 1g zi+ +6 zg I— Off -1£1 Z¥— ZV
19 LS ve-+- 611 $3 o¢— 16 “ir 4gf-+- i for i¢— AI
GzI 901 9gI+ oor fg 61+ ZEI gir iIv-+ gti ZO oZ— 36
IOI zor v+ S9 19 €1+ OFr ofr VE— 9g ocI e— 63
gsi gf o— 19 £9 z—. oS gi fe— <2 Cor ob— SE
2OL O11 Ole. V2 gs OI--. 991 car SZ-- oor 92 es—
VeI vei ze 2g Sg e— oS: 281 96+ S21 62 1v— SZ
+6 vir «IS— 66 O1I €z— E11 off gat Cgr $6 1g— S6
Zor var —-ZZ—. 95 cfr tvze— 11 LE1 e+ 6g S9 S1+ vor
z6 601 €Z— z9 cfr bi— ol i271 g— 2 zQ z+ Sor
Yor Cer — 10— 29 Orr G--_ = oer fel] 19— = tr vor 9gi-+ of!
ghr 14 gf-+ f€z1 Sg Zi---Z11 “S01 2y———59 Lol z— gel
gor 68 Gci— gir €Sz. €€-+ Ser 26 gi— 0g 96 ogI— z6
c6 oor 46— z6 6gr -“€4-— G1t - O11, §4= II Vol. 9f1—-"a9
zg OIL g6— 22 oft Ly— €Z ofl c— 20 t6 641— SZ
LL 146) ZE— GZ Slt = Oc» cO 88 I— 101 zor géi— 12
peAtes yseo Vv peatss 4seo V peAates jsto V peates 3svo Vv peAtes
"qQ = -e40g "qO 8405 "qQ --840g "GO 840g “40
ed ‘N “ey ‘2210 ‘AN *xOL
‘yoIeUIsig ‘ejsnony ‘Aueql|y ‘<ueqry ‘suztIqg
V
panuyjuorI—Ol TIaV J,
"Ie
“qo
‘uel
CCOr
30
Zi+
gf—
cc—
£1—
iE
zo—
Iv—
Zy+
o1I+
orI-+
og+
f1+
6£1
ool
6z1
zil
SvI
IzI
6£1
6f1
FoI
OI!
ZB
611
gz
QzI
z9
zl
oSI
161
Foz
grI
Sor
gor
ori
£g1
PSI
SPI
601
94
gs
£9
zor
gil
trI
Ivt
SzI
vz1
£11
£o1
1g
+6
tg
OOI
c6
bz+
6e—
1I—
Lz—
oz+
€1+
zI+
6+
ea:
ZI—
Z1+
09+
zo+
gi+
c—
>
oS-++
rS+
Zzi+
£i+
oZ-+-
Tf
1I+
Zol
$S
96
601
991
zgl
gst
ezi
16
gS
£v
Zt
QSI
SEI
IfI
zSI
OSI
ISI
61
Cer
ZiI
go
£6
Zor
gel
ov
6r1
ori
£11
S6
oS
09
62
140)
Zi1
gel
gSI
601
es
gs
A 4
1Z
ge—
6—
oa:
eo—
6f—
SO ——
e+
oc+
6v-+
3i—
gr+
6¢+
Rial
e+
oI+
se+
A
€1+
92+
o1+
zl
3
62
vri
Pst
vol
IzI
Zol
9z1
OI!
gz
cel
VII
QII
SII
£9
9
98
68
zo
gII
gil
EI
evi
EI
CII
IZI
CzI
gZ
1Z
+6
£71
e7Z1
6£1
CzI
Zol
ZZ
zs
ci+
we+
SI
bs—
99—
cs —
19—
99—
at
gI+
L+
98—
Si—
ceo—
$+
lI—
11+
91+
gi+
£+
6s—
of—
IvI
zSl
zel
cv
oS
62
o£
oI
gil
gz
QZ
C11
III
CII
62
+6
gol
ss
“9
gzI
Oo!
VII
66
gII
ci
cel
IfI
OI!
g6
OoI
IZ1
FOI
of1
orl
oll
£8
QZ
1g
611
IfI
(panuiu02)
gf— 9f
zs— gf
goI— 92
ozi— Lf
vO— $8
Zs— £6
a On
f£— 98
z+ 26
Iz+ €II
o1— S6
€i+ £6
6S+ zi
6r+ O6rI
6£+ z6I
gf+ 621
ozc+ 66
re— 2s
€z— £6
gi+ oor
S1+ Sol
gz— 09
cSr— «x
lo— 4
zZ
06
eel
ZS1
zS1
oSI
KASS
Sor
08
£6
€11
16
62
98
gil
28
LOI
bzi
ve
£+
ov—
so—
19—
Zi+
6+
Ze+
zI+
a+
9+
6I—
63—
zQ—
1S—
zgt
69+
09+
te+
zI—
6£—
col
16
c6
ZS
ee
eZ
gil
Zol
ZOI
€f1
vol
36
69
£8
8
£6
Lz1
Fil
tVI
CrI
gol
99
)
46
611
ZII
62
So
16
ZII
gSI
SgI
€¢1
zo
SV
cv
v8
16
gII
SH
9s—
vs
1Z—
ay
o1—
or
gi+
ci+
os—
os—
£o—
cz—
39=
Qe—
ef—
ica
1g
$8
tol
08
rg
92
£8
08
58
£o1
ei
zS1
gf 1
601
1Z
19
vs
09
Fol
£6
gs
Lv
08
$Z
£Z
gZ
£g
Ig
evi
Pel
gol
co
ZeI
gol
ozI
+6
gol
III
TOI
£6
vzI
zil
ec
Of!
£11
101
6I—
so+
€z+
e+
£6+
cy-+
61—
6I—
so
6zI-+
o1i1+
¢Z+
t-
Iz+
ve
£g
Cer
ori
6rI
Vil
601
ZoI
vz1
ggt
Sol
zi
SSI
zgl
SgI
601
09
gv
ISI
obz
6zz
Lox
g6
VII
€z1
zol
oZ
58
LZ
z6
99
¥8
grI
ogI
£91
6£1
zS1
gII
z6
v9
6£
59
gil
III
611
vol
c6
£6
22d
“AON
0
"ydag
“sny
Ayn
oun
Ae
“idVv
"Ie
“Pu
‘uel
gSOr
29d
"AON.
‘pO
*yd9S
‘sny
Aynf
oun
Ae
“idy
"Ie
"Pal
‘uel
£561
31
AO
oz— 94 96
ss— SZ o€1
gl— SZ esi
oo1— ZZ Zg1
elZ— FEI 902
4y— 621 QAI
Si+ vbz1 601
z+ $9 £9
0 Lo = La
gf+ of zt
ZI— 9 zg
sfe— £9 96
gg— 65 SzI
SZ— 39 €r1
v6— 9 19I
601— 9S Sg
eL— zi 961
vhe— cfr LL1
Z+ gor OS!
tz— Lg III
gf-+ zor +9
z+ 8g Zs
Of-+- 98. 8&
vy+ 64 Ze
Ot SQ, 68
f<— logo o£
Vv peAtes yseo
qO 405
—
"elV
‘£1903 UOT
Ae ee
9+ II gol
z+ zor of
oz+ 611 £6
6z+ gS1 621
z+ “yi SbI
{c+ grr SzI
f+ 96 £6
4e-+ +6 49
I= 8 88
y+ f€11 601
ZI+ QcI OII
€c+ Sti czzi
gz-+ ozi 26
Sv-+ of: S&S
gI+ for Zg
6— v3 £6
ov— ZS for
S— bZ zg
ov+ of1 96
6S+ 691 O11
ss+ ¥o1 601
Zi+ «zi Vfol
9— OOI 901
g— gII gil
SI— If1 obi
Cit Ser oz
Vv peAtess 4svd
"qO = 40
Noe
“SIM
“uosIpey
eB
69—
II—
ev
19-+-
ce+
e1—
ie
o=—
6+
Def
or1+
V
oak vv Sea ee —e eS <6 Fe ete ye Ma INE ae cy ao TF G4
gor €1r €€— 69 vor ZiI— 46 itr g— oe 12 elf-+- Shr £2 ZE-+- 611
zII 6z1 €v— Sg gol -1f— 26 €z1 g— zhi oS: 69+ gr 66 vS-+ Sz1
yor Sr €r— v6 461 9f— SZ IST. -€€—- "09; - £6. 905-000 Fr, - So OFA
lel cyl Le VeI ~AOl “eS OOl hon = 1 > 22 94 S1+ 6Sr hr 06+ gor
vel 601 9o— thi oSt L6— 92 E240 Y= = 1 Lz Qa Orel o ect meaelibal
66 Zol oz— 21 v1 iIv— fr vZ1 S+ SE of 61-- S€r o1r f+ 621
6f1 ~ 26 z— gor or g— whi 21 Z4E— Sg ZOlL 6Qv-- OS1t «111 ZI-+- Oot
ofr £6 Of igk= Stl — S— <eyr orl. =c&— V0 96 cb+ O6br Zor zt+ OI!
fEr «£6 QZ— QII 9grI Iz2— 69 OII 9Q1I— 6V Sor 6+ cfr fz 96+ Zor
62 69 II— er €z1 fz— g6 IZI goi— o£ g41 6r-+ thi S6 cf-+ 6
2s 89 gI— 12 23 Ze+ ir £8 Sz— ovr Sor ZS+ oS1 £6 €z-+ col
GZ 62 lZze+ £6 99 oS+ 601 66 vy+ i102 “451 €o+ O6f€1 924 Zz+ 611
gZ 4g 9gz+ 66 eZ f+. 69. 99 i+ Sr €€1 Sh+ vir 69 #«xzI+ II
LL gor v-+ 08 94 z— 19 £9 gI— 3g zor 46+ SOI 9g§9 ze Vil
v2 Sci €f€— 62 z9 OI— tz t£I L— 36 $g+ 61 601 gI+ 2zzI
Qs GAC {I0)}——> (0) ZZ zee— “Zz71 O6S1 IiI— Iv zZ OyG > Tey EST IRS ea
ofl «Imi immW— If cZ Co—— Sci Gil eS a9 PS €— 111 vir o1+ v1
LAM; Sel. 0-2 LZ £6 ev— or zS1 ti— Sg 66 e— zor vor grh— 6
Col wiecl. v= St 19S) e1—= Gor = 221 =-0 Vl, VIl Q=— II ‘zl Q9— 63
QII 16 61-- vor Stl gz— «11 6£1 6+ 2g €Z vi+ zor gbhr 69— 08g
S6 gor «1+ 491 ovr O6b— PL zl e— 10 +) of— Zor 24E1 ~-§c—" "10
+6 fll. Si— gor 1. ~— oor col tg— 12 Sor Z€— 411 St €— _ gor
Cit 6b iI—220 for 1+ ZoI gor 66— ZI Ilr 9v— 2g SS Sap ra
oor wet sa. SZ coy A 6— 16 ool osS— Sz GZ LOM, = “Sar Shs 121
Ql {197 -— I=+ 9S ZS Ya 26) 9a. Gi Gr 19 ce+ o11 Ig tr-+- £21
ZQ0 221 — Oi. 18 S9 c+ &2 1Z of-+ gor 94 ov-+ 11 €Z zo+ £61
peAtTss 4sevo Vv Pears yseo Vv peAtes ysvo Vv peAtes 4yseo V PpeAtes JSP Vv peAatos
"qO 40g "qO --340T "qO --40g "qO “940 "qO “840 "40
“y1V “SURyT “JUOTT *xoL sUIL IA TONAL
‘yspoy 2I}IWT ‘gouapuedapuy ‘eus[aH ‘ostg Iq ‘y10djseq *ylOI1jIq
panuyjuoI—Ol AAV],
70
2g
1Z
zol
z6
vol
ZolI
bz1
evi
SSI
OVI
611
III
ZI1
of!
eI
If1
yseo
32
8+
a>
fz+
Sth
cb
9+
61—
ool
Iz+
aoe
net
ee
ci+
oe
-
co—
Z1—
99—
1g—
or—
gf—
1hF—
Zo—
r+
6e—
se
ol—
s3—-
1s—
£o—
I7—
ol
96
66
Qo
£6
1ZI
QZ
£6
bz1
601
Cz1
98
CzI
16
1g
zs
LL
IZ
GZ
CH
eZ
99
1Z
gl
ZI
QP
19
©
ov
LY
£8
59
8
Sor
CII
gel
trI
tz1
FOI
cri
991
£1
ZII
OfI
op
QZ
zg
zl
IZI
TH
Fi-+
Cr+
Z1+
ge-+
o9-+
of-++
Se+
61+
17+
Sit+
£+-
sc
£S+
or+
jm
1Z
Sol
gol
OzI
£o1
IVI
CVI
vLI
OzI
101
bs
58
08
99
Ze
LS
vs
ZII
£6
ZO
grI
QII
QII
gor
So
6s
Zt
VOI
orl
Wh
<9
58
Cz1
CZI
£QI
€Z1
zgI
cvI
84
19
6£
gs—
€z+
1
bS+
fr-+
cs+-
6r-+-
(0)
gI-+
9+
i1i+
6I—
Cc+-
z+
oi
I—
SI—
£+-
9+
ze+
Acs
ZS
Sor
63
08
ze
ee
89
16
92
Lol
QI
Z6
ss
ev
cs
6b
ov
$6
LSI
IVI
VEI
QII
Cv
Zo1
z6
9
88
9S
Z9
aon
SII
Sor
99
II
9z
$S
£11
ZII
QII
LOI
OIl
ZVI
III
£Z1
£6
62
18
oS
of
7
or+
17+
oh+
co+
ce—
ZOI—
gs—
o9—
19—
ec—
co—
SZ
OI!
66
OII
19
gs
o£
+6
z6
69
6£
ov
6S
+6
III
£6
89
59
bZ
16
66
601
9s
o£
£9
br
Iz
Iv
cP
6e
Lb
£3
IPI
991
6g1
Ovi
Tor
QZ
bg
1Z
ez
ZI
QZ
£9
QII
ZOI
Qs
gol
601
£g
QZ
88
FoI
vz1
101
cll
IZI
For
<8
89
19
6b
(panuryuo2)
zI—
£1—
oO
Zi—
ZIi—
z+
Se+
e+
ChH_]|_
gr
€Z
6g
OzI
I€1
bri
For
94
<9
oS
96
zOl
Zo1
ev
6z
19
62
OOI
08
Fol
OOI
Fol
gs
98
6g
LEI
Qrl
Zvi
62
ss
1Z
GZ
r9
99
ov
ZI
gv
oZ
bZ
96
601
10
ZIl
O£1
19
1Z
ZII
£L1
zi
601
QOl
1Z
ee—
oe=
Vee
zb—
fo1—
€c+
6b-+
ce
osS—
og—
£6—
c6o—
bz—
6I—
oO
6e—
gz+
oz+
I-+
or+
L+
nar lL
19
61
LY
£o1
Sol
98
LY1
gSI
gSI
ov
£z
cy
cv
69
LE
Lv
of
ZS
ZO
VLI
Sol
6FI
Lv
ov
1¢
1¢
£z
cz
19
orl
og
att
140)
Ly
66
LY
691
rrI
191
IvI
6v
9z
ce
6I
oz
£6
OII
OSI
Zg1
gor
SgI
gol
Lol
62
of
If
09
O£1
€Z1
rT
$Z+
fo+
tht
ret
Le—
fy—
vI—
zwbv+
og-+
¢Z+-
99+
89+
6€+
6c+
Z1+
Ab—L
€1I
bS1
gel
gII
WI
zZ1
6gI
QrI
gf
611
Of
OzI
Iz
QII
ozI
br
fer
Zur
101
Ifl
foI
OSI
Of
OII
C6
18
Sor
66
Ifl
IZI
OcT
ge
16
zZOl
6VI
IQI
Cciz
Cee
ZOI
6€
£9
vs
¢s
LL
611
OgI
Sgr
cI
62
1S
ce
OV
£v
£3
gz
Cor
gII
IOI
vor
TO
+
of+
Z1+
9+
os—
eo—
17+
to+
46+
ori+
ozi+
zS+
or
zS—
€o—
os—
6z—
61+
git
Si—
1Zz—
FI—
Ib—
6z—
vo—
LL—
c6o—
Cn
gf
If!
PEI
ori
gZ
£8
69
58
ort
Cer
gol
Z9
Iv
ev
os
69
£8
£6
III
III
+6
a
1Z
g6
00
QZ
06
o£
el
no
c6
ZI1
PEI
IfI
SII
Id
£1
Ss
ZZ
58
c6
€1I
6II
ZII
Zl
ZO
£6
IOI
zo
ZO
ZIl
O£1
611
£v1
PSI
LVI
Zg1
Cor
‘dy
“Ie
‘Pa
‘uel
ESOI
00q
"AON
po
*3d9S
“sny
Aqnf
oun
KPTAT
o£— 6 1Z1
Sc— €Z1 go1
Oz— ggl goz
Z+ Lo¢ 002
g+ 691 £91
zI+ zi 601
vo+ €or 6£
0 ZS LS
ze+ 2 es
I+ $6 +6
oI+ £6 £3
61— ss bZ
ce— 1D 9Z
69— gv 411
zl— ov gil
of oF IzI
oL— ZS Zz1
Ly— $3 Ser
Ge+ ZE1 II
zs+ .S1 66
cp— €z1 gol
obh— ZZ €z1
zh— 99 gol
Vi=—— 08 +6
V peAtos yseo
-qO _-240,
SS
“elV
‘AIQMIOSJUOPT
vyr— 1g SzI
ve— gl zol
s— 69 bZ
II— 92 63
gt+ 911 88
Qz- gel Zor
gI+ gII oor
€I— 611 Zei
zgi+ SfrI E21
€— om E€1!1
11+ 2 £9
of— gr gf
Ly— 62 gf
e1— £9 0g
€z— zo 4&8
ge— 09 98
op— ¢€S £6
prs— LL I€I
yS— 16 Sv
ce— zir ZVI
61— +6 €11
zi+ 6o1 £6
of— 2 £o1
z+ 16 69
VY peatss 4yseo
"(QO ~104
= a
‘DOSIPE
be— Sor 621
vi _96- —€8
i+ 42 964
€i— 6S ZZ
gI+ Sg £9
Sot ter 65
ogo+ fz £9
1— {£6 go
€{I— 72 So
Z+ £1 vezi
z— O6£1 IVI
€1— zi ov
1r— 12 ZII
zo— vv Of1
1s— 8 gti
6s— 9 vr
O—— 56 101
6z— 39 £6
os+ ter FZ
6f+ oz 2g
vZ— SzI 661
ocI— 6 viz
90— 118 Ly
9of— cg gsi
V padres 4seo
"qO 9-240
TU
IV
‘ooy eT
Zo— vg tSt “YR 12 «Shi Ge eg ZS er" 16> «=6Sbr CI-F Gor 06
of— 06 gtr oS— £4 Zer of— SV SZ §-F- 10r- —- -£6 Se—" 99° E11
6€— So tor vL— 6S cfr Lo— 1 Ig o1+ £6 £g og— z 2b
og— SE £6 gfE— 611 “ZSt wt— gz gs 6+ 38 62 £s— +6 LVI
gI— Sr £9 gfe— 36 off If— gh 62 i S6> for’ - Z¥—~66- —orr
ve— SS 62 oc+ gIi 36 oI+ Szr Sir g— 69 46 Q— 9OI 2x1
oi— 9 LL o1+ 92 99 OI— ZII ezi vi— C11 “zt I+ g9 rhe}
S&z— £9 88 z+ &2 1Z ¥— 96 eer 62 OTT, ~-SI ~— OP=|=. 081. 09
6z— 6b 92 ze+ 19 cv 6— gor ZI of+ 621 66 19-- oS: S6
zI— gr gf Se+ 6b vz 1S §6., OVI. Vi--s 901 20: . OS-— ZI. ee
9I— of c e— ¢S gS li— lil cei. 2e--. 921. 66 45+ €or 9S
oI+ tb re f— 68 z c+ «mr 601 2+ zor S6 f— ¢Z 92
SG— _2e- ezi> vi--- OP! “zel_ -€5—= "gt 16- —£2--" -O1t-< 06°" 2b—- tf S01
ti— £6 I1I oz— Zz1 LZb1 og— SYP Sor I— &9 99 L—— Sita cer
“4o— 11 oSt bvL— 92 esr ZE— oS £6 zi— 4g 66 jzI— “£11 621
of— of1 ogi 6z1i— lz gSI g99— 2S gir t— dg OIr Si+ 6z1 #11
Sgs— tS 6€1 ozg— 06 O11 Oo 760 6—i‘ié‘i KI — «COKHQBsi«éKtO:Cs«QE—-—sod08M=—s*=EPIT
i— 19- Zit i= £11 —_ Prt Si—= 101 = Ol) ~6— 0g - 69 1v~— v2 SII
S810 gor “Se-- 121 -96 St QII OIL 9gI— og 92 zo— IP for
Oi==— ell tel _fi— 26. Sor Sh—- ov. 16 ve— S2 66 4&—= ZS 6
Z— S6 zol t+ or vil €f— 9S If€l gI+ 06 ZZ 1Z— 9 Ser
Z4Ii— €S of S+ «mr gor voti—ot fr oof 611 6S 62— £6 ZzzI
6b+ 98 Ze t~— gf zor Lz— 06 Zit vo-+ 66 ce z+ 96 bZ
ga. 26 63 6z— 2 19 y+ 99 zQ ofZ+ S11 SP sé+ 6 09
Vv peAtos 3seo Vv peatss 4svd V peaAtes 4svo Vv PeAtos jseo Vv PpeaAtos 3sed
GO = --340g GO P40 qO 0g 4O 07 qO -910g
“suey “JUOWW “xOL suleyy “qOlT
‘gouspuedapuy ‘eus[ayy ‘osed Iq *yr0djseq *yo179q
qd
panurjuoI—Ol ATaV],
‘29d
“AON
pO
*"yd0 S
‘any
Ang
oun
Ae
“idy
"Ie
‘Qa
‘uel
9S 61
00
“AON
po
42S
‘ony
Aq
oun
Ae
“1idy
“Ie
“Qe
uel
SS6I
34
zg
6£—
o—
gt—
gr+
gs
£¢-+
ver+
yei+
£o1-+
I:
Ig—
SO
69
gS
16
ZZ
ZIl
98
ZIl
06
IZI
ZII
ZOI
¢Z
vol
101
Zgi
oVvi
LZYI
&8
IvI
g41
Zgl
OOI
59
66
£11
gel
O41
ogl
eel
gtl
88
SZ
62
gil
ZI
691
6£1
ori
+6
6v
ZI
6S
gvI
£91
e+
Sl
bi—
oe
Ly—
ve
£s—
=
1g—
ave
zI-+
b+
£9-+-
of+
a=
oz—
GZ
98
ZOl
c8
19
gs
1Z
63
gli
ev
+6
gil
140)
z6
36
ori
CVI
LEI
gol
6S
gf
6¢
OII
ZI1
CII
68
ZZ
+6
Z6
ZIl
Sz1
IVI
gel
66
bz
o£
$9
gs
¥8
Ze— os
6or— ZS
Zy-+ of1
cs+ zti
zo1+ ZOQI
bo+ gti
ve+ €S1
o9+ IZ4I
6+ ItI
fz— Col
£5— gZ
Ig— 08
be— 6£1
Z+ = g9gl
Si+ 681
ze+ gzi
41+ 66
os+ £01
Z£Z+ gti
Zo1+ 9gI
Zo1+ S61
Sot+ 091
I+ zi
z— gol
gol
6g
LL
bZ
O11
III
Zel
ofl
Ser
191
fol
I9I
vr
gol
©8
es
19
62
66
Zzl
Ofl
git
Le—
Zo1i—
o£+
98+
ys+
(——
of
of
6S
6S
6gI
ozI
ZVI
48
€z1
Zol
zs
62
zol
III
£g
1Z
66
£g1
GZ1
ost
gII
6g
Iz
L9
£6
eer
621
ze
140)
+6
48
101
GZ
g6
601
Ce
6£1
1Z1
gZi
€or
gz
Sol
v9
29
+6
£11
(panunuo?)
11+
oI—
19—
tg—
L£L—
g¢—
[=
€1—
61+
£b+
Sy+
Sots
Sea=—
61—
g1+
09+
ov+
6+
of--
ZS-+
g—
fo
gol
£6
gs
gv
ZZ
611
ZI1
Zi1
gol
LE1
Vil
gZ
vs
Zo1
£1
691
of1
gor
Zil
S11
gt
£11
28
c6
Lol
O11
Ifl
Ovi
czl
¥zI
of!
£8
06
€Z
gz
ZzI
601
o£
o£
$8
1g
1zI
cer
ue
6g1+
gZ1-+
ghi+
7s+
oz—
oz+
691-+
csz+
oze+
eg+
bgt
zei+
ziI+
09+
ov-+
LO—
Z6—
IZ—
e+
19+
ze
vol
g6z
€or
ZI
OSI
Cer
€Zz
GZe
Qzv
ogI
O11
oI
IZ1
zol
zvi
ori
99
6z
£6
gor
oct
LOI
LZz1
L6
gz
621
€11
FoI
OzI
gol
LL
se
6S
£6
4)
ool
C1
gz1
48
140)
bz
Sr
y—
sI—
ZI—
f1—
6+
1S+
Lyt+
Sr+
6¢-++
89+
co+
rg+
c1—
blL—
6r—
cz—
I—
£+
ge+
Mats
Pate
ALT
Ill
e721
€z1
vz1
OvI
Qe
6z1
£o1
ZEr
691
6g1
Z11
62
58
zol
oll
18
58
88
bl
IzI
9z1
EI
zzl
gzI
CII
16
$3
t9
69
$Z
Col
ofl
€SI
Zz1
CII
8
LL
LS
o£
gIl
Zo—
£1—
1z+
oI—
61—
1Z—
99—
ZQ—
ei
ze+
Sv-+-
6z+
gi+
11—
3i—
88
89
601
oil
ZZ
09
gt
Ze
62
16
kat
ogi
OVI
gzI
gzI
611
Ce1
Fil
FoI
£o1
62
16
88
Sor
VII
Sor
6g
16
06
£o1
Ill
06
10
ZI1
gf1
ze
£11
GZ
48
601
20d
"AON.
‘PO
*4daS
‘any
Ajnf
ounf
AW
“idy
“Ie
‘qa
"uel
gor
09d
“AON,
pO
3d9S
‘sny
Anf
oun
a
“idV
"Ie
‘qed
‘uel
L561
35
be—
Lz—
1g—
Iv—
Lz—
Lo—
or-+
gé+
6z+
c--
e1—
gI—
V
Oo OR = Se ORE Ss f9f6) v9 SI— 99 £g t= -c0P Oot “Qi fel ave —. O2I- Ret, -g4--- Oot ier
tcl. Ost 0g-|> “Ovr 39cK, 1Oe-— 001, 02 Z4y-- €or oS g— 1g Zo— £6 OOr QOI— gir EI gi+ of: ef1
Off Se41 =to-- S0l =cbi.- +95- -<O2I. =<QI1. -6— 06 Sor th— gZ zl gy— zg ofr 45+ €or gor ~2b+ Sor E11
IOI gor gf4+ 41 96 Ve— ELIT oS iVI, sOt— =I Z11 ov— S6 Str g&— 99 QOL “19--~ £91 cor ZS-- 921 tz1
89 Eels 40f-- 291 17l =1z— por -Szt S-+4- £6 88 €f— 66 eer -vI— “19 SZ t¥-- “001 ~ ZS w+ ZzI Sol
ze 26 Se+ 66 v2 vS— z9Q QIr iI2+ 92 gs v-+ zor 936 OLE Ser SVit- 2O1-- ett Solr sei— = 66 III
gs vg fe ZV SI 6— £9 z6 OS-- O21 10 €€-- vil, 1g Sz+ “ZbI zzI 9z— 66 Szi ov— SZ SII
Bee a O 0 0 OTs S201 205, “£0717 “eal <00.. - WV fet. GL. SOL-| 8S) “Fil. “Of — —v6n “bel “Ig— 2701. =nEt
vZ 1Z ce+ cf oO vS+ ir £9 Z4$-+- zz1 &9 Ii £01, SQll ste Styl, -Yvi: Of — 19 O1I of€— vor ofFi
$6 46 Zit os Iv We+- 7221 QZ Se+ vor 69 {go— 29 OSI G6¥-+- 621 of: oc— er Sr -2— C11 ova
28 O1I ZI— 69 98 Sf+ 621 v6 v¥— 18 Sg fo— gO TOl. <GG-- -Vfc° “S21 ~£2-- 25° oho, C-- 2 62
oor fr ghr— <s oor oS+ tI 6 Wie SZ1i. fl, Ses SOll. seri. =SZ-- ihe. oor -oZ-- "Orr Zp. I= = 90 £0
49 QzI €6— 1S Vi se (SET “So W= Oct €Z%, We “gel. O81 ZEc- es VOI. “2451 — GO-F re SI i= ——— els SET
ZS Zo1 oZ— 4g aot se OOK (ROT 08 “gor. Zor OS VOL VSI: —I--. 901. Zot —Jo-{s 19 O VI— VII. ger
zQ~ ~=—sé@6G £e— -4S1 ~oO1 ~¢— ~6f1 141 So— ZQ zSI €6— o£ €91 g— Lb ss ze-+ 9S vz oz+ OSI O€1
gf cor S€+ cic ZZ4i ge— grr of1 Ib— 62 ozI 66— & zsi tvi+t v9 oS Zi-- ol oO C= COl = YOn
98 Zir 72Si+ 122 611 o1— Shr SSr1 zze— +6 oir Ly— of Zit QoQ vS ci+ ¢z 8 Cel = et
vZ cit 6095 19l- =10l ice — 90 Of. =9I—— “Gor Ser o— 09 fey. —1¥s-o Sat. 2 6g— Iz OM aca yel == Oor
gf €or z+ vor 0g Iz— 006 GE ee AT CAC (A 100) LOW e1Or | el Te ice osS— 624 Ser Sv-+ 641 Er
4 g6 gv— z 69 Ox > 211 201) evict cri: “9zi -1€-— for teZ St-- -1fI. 901- -91— +9 oor zZ£+ OgI gol
cle ee 99— ¢P 601 9¢-- -ZII 16 99+ 241 gor gze+ 46 69 Lé-- cel S6 or+ 0g ov 6S+ tri SQ
36 = 09 fy— 12 vir 4+ Sor 9g9 Sf-+- zx 6 Sé-+- SEI ~09 S&Z—— 19. “FOI 308 —— fo £5 S€-- 16. of
zor £2 zoI— SZ Zoi. SE 601 “V2, Se-- vEI 601. Gh+- 601-9 =6— ~09. $6 ~6§— ge... -Zo.. =VE4---SZ Iv
19 oS £6— 106 vgr oS+ ri v6 f+ oz Z11 f+ “41 Gor z+ 06 88 ov— gv vO 6€+ zor £9
99 62 BOs OL “900 - V5-}- 001 =21L “gc—= GEE =<€o1 2z— -ZEr s6h1- o1-- Zot. ~26 fgo— 94 O£1 oz+ 91 SEI
a 89 Se— 6 Oc 817: yl ver —OI—. Szi_ Sr o9-+- oze oft ei—_ 26 6ol. es— Zor O6S1. ve+ 691 Sor
peAres 4seo V pedAtes = 4seo VV peAtes 4seo V pedtss = 4seo Vs ipearas = seo V pedates 4seo VV pedties 4svo VY pedaros 4yseo
qO -a0g "GO 40g "QO -940g "QO = --840 -qQ -240 "qO -2404 "GO 840 "QO = --240
ee es oS 9 SSS | ——__ re
‘O'N “HED “AUN “SSI “II 3q2N ‘Z11V ‘uUaT,
‘XINQsi[eS ‘oyUseIOeS ‘Iaysoqo0y ‘wOsqly 10g *el10ag ‘eqeuo ‘asplig [einjeyy ‘OT[TAGSeEN
>
panuyuor—ol alavy,
36
eb—
of—
19—
SP
e+
ce
6e-+-
1$+
6z-++
ge+
it
62+
IZI
gs
cs
TZ
So
06
bZ
09
VII
IzI
I0I
eg
6S
62
z0
gs
Fol
ISI
bri
QII
€v
gr
LV
gZ
18
ZIl
ZZI
€Q1
QzI
gil
611
40)
cs
ce
QI
If
ZZ
SLT
£6
ZO
OzI
III
VOI
VII
O£I
III
Crr
zo—
oz—
os—
fz—
9z—
oh
cy—
zg—
c1i—
z7Q—
€Z1+
1Z£+-
voe+
fal
grI
gII
z9
°
VI
VII
Ori
LZz1
8
Lv
Iv
99
E21
6zI
go
gS
Sor
CrI
VII
gel
1S
¢z
gz
z9
16
gst
Qzz
abe
991
Zg1
611
LV
€z
gol
Ifl
eZI
gSI
601
grr
eer
gol
£o1
£Z
Te
a
IVI
sCr
re+
or1+
12+
oo
ci—
ov—
6z—
7o—
Vite
ee
ez+-
fz-+-
it
Ofl
OII
If]
88
88
59
36
oll
+6
gf
08
1g
60
ZII
VII
z6
gd
c6
601
10
08
68
SZ
68
bl
98
$$
gZ
Ca
Cer
goz
Loz
ZI
oor
89
rg
16
gil
£8
94
ov
1Z
oI
O6zI1
ze1
Lz
ZI1
Sol
£3
gs
/45
99
98
zor
OZI
£z1
Sol
Cor
fg.
z+
€1—
er—
£S—
1I—
1€+
be+
gI+
ob-++
zo
Z1+
iets
gt
6¢+
e+
oI—
6f—
6c-+-
61+
c=
1Z
ZZ
gII
£LI
vzI
ZZ
III
OII
LZz1
eS
62
9S
19
6S
9
94
ZII
Og!
zZI
OFI
gol
ZzI
$6
SZ
ce
se
6V
ev
ZS
or
Cu
Il
O41
IgI
£8
08
94
601
eS
ty
ZI
o£
09
oZ
1S
Ol
vZ
Ze1
661
v1
or1
68
OII
Sol
Sor
06
89
o£
gI
ZS
(panuizuo )
gz
6+-
Zi—
SS--
zgo+
06+-
y—
8+
zi+
Sie
be+
Sv+
ci+
9+
cco—
ee
Goa
Roa
re—
vo—
1I—
I—
zi+
oI-+
gz+
oe-+
oz+
oI-+
c—
ze+
Vote
Qh
GzI
gol
&8
ori
gti
QgI
Vil
Ovi
IzI
ovI
Zor
ZII
o£
6S
ee
oY
16
Sor
Zol
1g
vor
36
0g
89
1Z
08
10
OzI
66
TTT
66
66
16
QZ
gII
zZeI
16
£Z
49
gs
¢s
g¢
£6
ZII
£1
IVI
cri
CII
LL
gs
89
1S
1S
¥9
+6
88
[6
hh.
se+-
Se
88+
16-++
se-+-
oz+
gI-+
8e—
of¢—
bo—
Si—
be+
Si+
o>
Gel
ce-+-
69+-
6e+
Cea _i_
ov
89
QrI
10
59
£o1
+6
gol
Cer
gz
Of1
£6
ce
of
€S
6S
66
III
£8
zo
Zz1
601
98
£6
O11
Qt
£o1
arr
LL
48
ZS1
661
OgI
Z91
zl
66
£C1
1ZI
OOI
ge
cy
OI
Ze
4g
ozI
€z1
gz
IfI
89
Zil
SII
$6
88
La
vl
bl—
geI—
0
e+
4
Z£Z£+
bri-+
og+
zs-+-
vo—
zi+
e+
95—
Soa
Ze—
ez—
of+
fg—
611I—
1Z—
Sr—
ci—
zi+
ee+
5
16—
zZQ—
9+
Me Gin
99
Be
vZ
vol
ZE1
bri
ZO1
QVvI
gsi
08
fer
€Z
ez
£1
of
19
gr
oe
19
59
es
z9
Ori
vE1
€£1
19
66
£9
or
ori
gst
08
o£
VI
£9
£3
98
goI
bri
IZI
94
6zI
1Z
oS
£S
99
£¢
IfI
gSI
eel
OII
06
LZ
bE1
IOI
gel
FEI
ool
GzI
re
An f
161 fo
11+
Z1+
be-+-+
ge
SZ--
ght
Iv+
z+
e+
er+
ez—
¥9
OOI
101
19
ey
<9
06
£6
06
£11
£11
ov
tz
gz
SZ
£g
Zl
ZzOl
gII
+6
gol
So
19
62
£o1
46
£9
ev
feo
z6
III
vz1
££1
S6
ov
94
gII
SE
S6
46
ZOl
+6
ez
°
ce
Ig
08
SII
Ig
oSI
QII
Vil
ZOl
29d
“AON
yO
*¥daS
‘ony
Ant
oun
ACT
‘Idy
“Ie
‘Pa
‘uel
vSOI
09q
“AON
0
*yd9S
‘any
Ane
oun
ABW
aidy
“Ie
‘Qa
uel
ESOI
29d
"AON
‘0
"yas
‘any
Amn
aunt
fLeavtar
37
1s+
gi+
L--
=
88 ivi 6 Ge= Ge G1 ty SO: Pe— 6 der -OF 26° eS te We G2 fe Zor w- gtr 4
z60=— «61 = 6I— = 00 62 6I— 19 og z2i— Szr “401 I— 4 SZ 9+ 8 62 of— If 19 OI— zor 2zilI
gbr ort z6+ OS1 Lg vI— Ig $6 gI— z6 o1r tw— Sb _ 63 6+ oor 16 45— le +g vy— Sb 6g
Cre Oni (ebr-- 9hr 92 c+ 621 ter z+ 26 0g zI— _—-z v2 0 16 16 tr— tr 48 4E— Of 28
oor ger oft ror 18 Si-+ Gyr oft Zi-- 26 SE 65 96 46 Gr+ gir 66° w— oF 2° Le— oF bh
94 = 89 gt— oOo ge G+ Wr vor 15+ o£ Sz 6v-+ oor 1S cc+ 9gII 19 gz— 09 98 Z— 9 S9
gor €2 I— 2 £s l— lg ¥6 gz+ 1g es z+ £6 1Z 19+ gg Le gr— gr v6 6+ 9 #6
7g ~«=9 Z+ or 96 OI— ZII 7zI 9QI— gg vor I+ Zé gZ 61— gv 49 vi— SS 69 z— ££ Sg
66, - 2S cc— vor 61 Se+ zbr 411 SI— 101 o1r t— OL og wW— 62 €Z Sh— SE gf gst 63 1g
ci 19) COZ 92 VS, «OE OWI 101 =61c— ven Swi = yy— Sh: 66 of— ge vor Zi+ SS gt 1€+ oviI 601
z6 gZ 69— 96 Sor ZI+ o1r £6 QOI— 611 SEI o6— I€ IZI Qv— cw gg L— se a QI+ ZI gzI
49 99 If 941 4Z4t t+ v6 £06 Sv— 24g v1 o6— of ozI go— 6b “11 ef+ 1g OF €— gz 621
ey gs vi—_—«SZr:«= gt Sok — «ZZ Zor gr— 19 601 9Qf— oz 96 ve— g& ze “41+ 16 2 Sz— € gor
62 62 OI—1f1 “Zhi So Zer sir g—= 92 ¥g Z1I— 69 98 z— oI gv f9+ £8 oz os— zZ Zz
66 gor Gf-+ of: «mur €— Sr gtr 1h— of 411 i+ 24 94 zI— oS z9 1z— ZE gs 61— gor ZI
gor 66 go1i+ Szi 61 f+ €Z1 oft zs— gf ger _GE=-- Ger 06 zy— 9 gor £6— 19 vSt Le— gir Shi
4g. QO1 {101+- 101- 0 t— (46 Ol. “OI-> Sir Sor tey- eg 98 €1-- SO- 29 4Z— 611 961 SI— OO i121
£6 40t-- S==— 5 0 c+ 4g Sg 61— 66 gIr I+ £3 £9 £1— 98 66 6£+ 6€ ogr vI— Sg 66
io. 0M ck iv. -G ve—= iy €0:. O1-- 611. 601 ‘of 09 og: or4- €9 €2 vrI-- Sgr 12 vi=—- Zo 1g
96 OIL 6€+ 11 gZ fz— 69 z6 4 =6OoI— ~=£6 Zor tz— v6 gIr of— oor ofr o+ os: £& {1+ 4g 4
ie She L- Zit Olt; Ol—. 1OL’ 331 If Or zor fe-= 99:- E11 ch eg Vel wWhi=— 26> Of 2 lek gel
33 «Ow 6e— 6 ovr o1— Sor Sir g+ s9 LL o1+ vor v6 6+ vir Sor vl— 6V fzr cze— Lbr 691
cae Ob GS OR de Oke OQ «6890. «CS eer «CH OE — 06. et eg. CE SCA «eI
Gye Zo. Go FOL OSas Ie 2 gO TEE. ghee. 2h eh SER 86. tee ge Ol 09H eg. ON Oe FOES «OF
peaArss 4sEed V peares jyseo VY -pedtas = 4seo VY paariss = 4seo V ~peares cast VY peates 4seo Vs ~peates §=-4sto VY peates §=4seo
"qO —--40g qO 8407 qO 10g "qO = --340T "qO 8107 "qO = -840 QO 0g QO 40g
‘0 'N “HED A‘N ‘SSI IL "192N ‘Ziy “aue TL,
‘<anqsi[es ‘oyuaMIeIOeS *194}S9yIOY ‘uosqiy 10g ‘el10ag ‘eqeug ‘a3plig [einjen ‘ST [TAGSe NT
panuiyuoI—Ol FAV],
‘29d
"AON
0
19S
“sny
Aqnf
oun
Ae
“Idy
“ey
“Pal
‘uel
OSOr
29q
“AON,
0
*¥d3S
‘any
Ant
oun
ARI
“1dV
“ey
‘Pa
uel
SSOI
38
L£b+
vo—
L6—
ZI—
6+
£S--
19+
bo+
61+
9S+
os+
£f+-
gi+
Co
7—
II—
ci+
zI+
e+
oI+
62
48
58
Sor
gol
gS1
EI
zel
$6
LSI
gtr
CZI
vol
Cor
SII
cz1
£6
QzI
Sei
ze
ISI
SgI
Pr
€1I
66
£Z
LL
IOI
88
96
III
vol
9zI
OII
O11
gil
£z1
OIl
6o01— gS
IVI— gz
goI— zy
ofc— 22
s1— +S
OzI+ OSI
Iv1+ zogl
g6-+ zgz
Sgt gt
€z1+ 10
of+ O6zz
Z— ogi
fe— 9
11+ +6
gi+ 2g
6h+ og
oO fz
vyr— oO
6— £6
gt— zhi
v~— LI
v¥o— ZZzI
gf— 66
1z— IZ
Z9I
691
Qri
gor
gf
Iz
98
¢SI
gét
£61
Zgl
gir
£8
If
fz
gol
Og!
g61
ogi
Cer
z6
6z—
or
z+
g+
z+
ov+
4
ce+
so+
69+
ee+
61—
gt—
Le—
II—
ci+
cl
46
gol
gr
IFI
6z1
OzI
gil
IZI
CSI
OSI
OzI
63
£9
o£
ZS
gZ
101
SII
+6
62
SZ
rg
gz
PSI
QrI
Cer
gol
08
/
[7]
98
48
18
£8
96
£8
gor
v8
48
g6
ZI1
6g
08
09
94—
Geo
ert
oS-++
bo+
12
ZI+
£+
89—
So1—
Irp—
gs+
Le—
oI—
grt
16+
cet
fees
6r>—
ov
LY
ogl
18
98
Sor
LEI
ZII
OzI
+6
£6
QO
Ift
OII
vol
88
zB
c9
09
4g
06
VSI
Fg
ZgI
OFI
ZOl
gol
LL
98
ZS
oP
£3
LZ
zi
ori
(panuyyuor)
zi+
lz+
62+
II-+
ob—
Z9—
19—
Sz—
ge—
be—
oI—
62
67
gII
gst
gor
Lb
99
gol
gel
grI
v6
SZ
94
£g
c6
fol
8
ZOI
101
Zol
CII
ISI
O£f1
fer
16
62
£6
Of1
ZeI
fer
Z91
€o1
VSI
ozI
98
19—
9e—
—
eg+
og-+
ss+
Z1+
ze-+
gf
co+
ge+
y—
99+
12+
9+
g+
zo—
¢o—
1S
1f
0g
vg
ZZi
£1
LZII
62
£6
Zo1
ZOI
G6
€z1
zel
Pr
vol
bz1
bri
£11
czI
OI
gor
Lz
69
z6
gol
63
6g
£Z
29
29
ofl
LVI
O11
16
6r
gz
gzI
gZ
611
Ivt
gor
crr1
ZO1
9s—
66—
oI+
gz+
gi-+
zZI—
z-+-
91+
oS-++
Ly—
ze—
1z+
fI—
zi+
Se+
ee+
ce+
aie
£S—
aa
6z
ze
Ser
Cer
OzI
z6
IzI
cSt
ZE1
ZEr
ZOI
¥9
PEI
OzI
III
18
v1
OSI
QZI
ZZ
LE
58
Ifl
SfI
£zI
6
vZ
€¢1
€S1
IzI
48
III
III
101
£11
££
66
VII
gil
9zI
£6
49
06
9s—
Lo—
ceo—
zI—
eo
6e+
6e+
6e-+
1z+
th—
a
f+
ve+
zf-+
zI+
6I—
g1+
6g+
£L£+
gI-+
6s—
zI—
or-+-
89
09
08
gol
Zvi
LEI
gor
LOI
Fol
£6
9S
6z1
zgl
PSt
vzZI
63
gor
zS1
Og!
ZII
58
z1
Sor
ver
Lz1
CII
gil
OII
zz
QzI
ZI
ZI
gor
£9
£g
101
thr
Of1
SzI
29d
“AON
0
*4d9S
‘any
Ayn
ounf
Ae
“Idy
“Ie
‘Pa
‘uel
gSOr
"29q
"AON
2 O
39S
‘any
Ajnf
oun
Aey
“Idy
“Ie
‘ga
"uel
L561
39
e— vir ofr 6— Sor bil VII 3 Iz+ Ig og
¥--- gcl gil Oo Gor - 60% F5-— gti 9 0z-+ 0g rs
eS+ €or 111 62+ off 01 oz+ eziII 2z6 61+ 68 oZ
6f+ Str 96 Ze-+ vl II 12+ OzI Sor 9gI+ vol gg
Ol—== 20l. cll: “G-|-- Gel Oct 1-- O11 €or vi— Gor E21
co— SS 48 £09 69 gi+ Zzr 601 of— +6 Of1
6e— 55) v6 II-- 924 S9 V— er Zzt ZE— 96 QZ
1z+ S11 v6 v+ = 96 zo gI+ PEI go1l €I— Zor ozI
eg+ 4er1 SY 9z— 96 eI €+ zit 601 te— 96 zi
06+ ghr gS g9— £6 I9Ql QOI+ gQtI zziI iIt— Ig ZZ
ace ago £0” = GR—— OOl- gor Si-- 6&1 ter Z2Z— SS. —cex
9+ oor *¥6 Of— “Oll- O91 Vi-> zi Sor fe-- OziI 901
S-- So oR £g— 30 Igt €v-+ Zr +6 “I> eer “SII
I-50 146) 6g— +S tvI -Os-}-— fe8 - £9 4I-- O11 ZOl
gI+ zor 9g 2c—— 97. OOl Zi- oot 6. €2— 62 66
Oro wath 102 gz— SZ I0I 7zé— Ig for 9+ 96 os
wot 6b1 Zg 1i— 9S 6g ez— £9 Sor t1— €Z vg
Oll---ser OF Qz— 69 £6 gt— 39 gOoI €I-+- €11 OOo:
OfLI+ gvI gI ZI— 99 £9 6S— zo It~Z— 11 ver
I9+ vor €&P V+ 2 89 6r— Lo QII 7zze— OII_ IPI
az+ €11 £6 8+ Ig £Z 19— £2 gtr gf— FZ zi
Iz+ oor 64 I-- 10- —0g -0S— £6. -€v1 ~ ey—. 06 «sg Ex
gy-+ 611 12 f+ mr ger Sz— €&9 gor 61— g6 £11
Sf+ 63 vs Ss; 21 401 OI— VOI ofr e— C11 S11
lz— gf S9 ve— 2 $6 IS—— (00s Zen ¢eS-|- ter eer
gi— 49 coe. ce—"_ of ZO ze— £6 SzI 96+ Foz gor
VY pears 4seo VY pedtas = 4seo Vs ~peAtos §=4seo V padres 4seo
"qO —--840g "GO = --8405 qO P10 "qO 840g
Ooo RES) “CUI “ON
*WOJBUIYSe A ‘a][IAseuoy J, ‘neg 4S ‘SIROT 3S
Sea
of-+
7Oo
ze+ +0 z9
EZ-= 191. — ‘gor
Iz— gti 6S1
61+ Ogi I9g1
oz+ fr FI1I
gr+ 6zI 1g
zI+ 12 6S
ZI+ gf 99
ve+ L9 ee
of+ 9S 92
gI+ 2f +S
42+ 98 68
cs+ ozi 14
oz+ Sir $6
f1— 611 2ZéI
iat Oye eater
6+ gII 601
Sc- e11 9g
v= 300 o£
vit ZE1 a1
oc— “zt Z¥I
gof— gor zt
9-921 Oct
Se-- Shr - oat
4g+ g61 III
oz+ zi Zor
Vs padres §=4yseo
-4dO0 3107
“SEM
‘gueyods
d
panuyuoI—ol AIaV J,
S— ZS zQ
Ge er By
vi— ¢S £9
gI— 6s
gI— ze QP
tv+ Lg bP
L4E+ 98 or
gz— 006 QII
vo— vS gII
Se—— 65 £8
t+ 99 9
w— OV 1Z
cse— ov SZ
gr— tz oZ
zb— dai vS
19— + S9
zL— 62 101
Se Ge €Z
9— 90 vol
€+ om JZo1
oSs— 66 Or
S6— OF vrI
IvI— ZI €S1
1g1— Zo ggI
Og=="2£0 = GPE
Q— OOI 9gOI
Vv PeAtos 4seo
qO -710g
pee Sees
‘Oq eyUeS
N=.
Ea ee =O
Zi-— g&r ret
6o— grr ZS1
0 S61 S61
QO+ tor gSI
TI— Yer Si
vI— Zool IzI
gI— Zzol ozI
I+ gp LV
Og-+ OOI oz
IQ eit 15
goI+ 611 €1
ov-+ 09 VI
OI-+ 6F 6e
I+ a Iv
Sy+ oor G&S
S89 94
gr— 99 FVII
QlI— z gII
fg— 0 £3
Zg— ZI 66
of— gz gs
f1-+ of Ze
Ste— ov Ig
se 10 08
GE— "259 vor
£— Ig 88
VV pearles seo
"qO = -240g
ee
“se,
‘OUIpIeustog ues
Mar {[ sv aa
Sot zor £6
gt oZ1 e&1
zi-- €or ISI
4t— ‘OSt 21
19— +8 SvI
92— OZI OrI
Of=-. er) con
gf4+ Zvi 69
vS-+ 06 of
Se+ S95 of
tz+ tg 09
6z+ +3 ss
II-+ 62 89
SsI— g9
VI— oOO1 FII
es— 2 Lz
OI— €z1 6f1
£l— 92 6r1
SI— of1 StI
gt— SZ €1I
QI-= “PIT 66
0 69 69
Z+ 08g £Z
gI+ 69 1S
ve+ 06 99
v-- -ZO% COL
V peariss 4seo
-4O -I10 7
qea)
“ame Hes
- CIN
‘Qa
‘uel
eSOr
‘29q
“AON
pO
*yd9S
‘any
Ajnf
oun
AW
“Idy
“Ie
‘Qa
‘uel
IS6I
‘20q
“AON,
0
*¥d9S
“sny
<qnf
oun
Ae
“idy
“IV
‘qe
‘uel
oS6r
40
Ze—
9r—
or
Sc—
Se+
vz-+-
9f—
6S+
ov+
6z+
et
ze—
1g+
z+
os—
ZI—
gI—
ge-+
zv-+-
or+
Si—
61—
6£-+
s+
12+
Zz+
z7eo—
9c—
bot
9S
co
EZ
LL
gr
09
oS
94
06
1Z
1Z
gL
631
£8
6g
rg
$9
fOr
OII
PSI
CzI
OzI
ool
FoI
Ser
ger
FOL
val
OI!
101
Orr
£6
IVI
6II
Zol
gol
58
O£I
16
Z6
1g
CZ
LL
QOI
OII
O£1
gol
C21
08
59
LL
Se1
zi
Lz1
zl
ci+
1Z--
gs
ye-+
ght
Iv+
6v-+-
gt
OI-++
£+
Z1+
a
fg—
ge+
eS
61—
oI—
oz-+
6z+
e+
Se
vz—
99
ve
ov
gv
gs
89
SZ
cP
LZz1
OzI
ZI
QfI
QfI
98
fer
+6
Ov
c6
£or
6S
SP
<9
OII
Sor
Sol
Ill
19
vv
oe
gt
LV
bg
es
oer
99—
Z9—
fo1—
¥Qg—
os—
Ot
Qc
o1-+
os-+
Ze+
gt+-
oZ+
zg+
ert
of-+-
If-+-
Cr+
€g+
OZ
of+-
9+
ve-+-
ze+
62
<9
1Z
£8
10
101
SII
SII
£6
O11
OII
92
£Z
58
zol
Crt
Qc
OSI
£e1
601
Cor
FoI
b9
or
vs
GZ
GZ
101
SIL
58
68
6g
SII
ZeI
Zvi
66
of—
6sS—
9t—
7—
6f—
ce—
oz—
6+
6z-+-
viI-+
Zl—
yS—
gr
Clas
1
ve+
be-+-
ce
cs-+
gi—
Tote
ap
zI—
gs—
89—
£9—
Z9—
QZ
ZII
PEI
gor
£9
£Z
Ae)
Qt
SV
IV
ev
gs
vs
zs
Iz
SF
Zs
£L
101
£6
63
LZ
&
“9
ev
o£
98
69
60
LAs)
6z1
141
Ost
eri
gol
gol
QoI
cg
6£
gI
Lz
ZoI
£Z
bZ
FOL
6f1
Qe
£S1
ofl
JIY
(pamuyu0d)
oI—
re—
61—
oz-+
Ze+
Lo—
61+
of-+-
6S+-
Zbe+
gi+
ZI—
Geen
gz—
br—
Si—
cot
bS-+
6r-+-
act
oz+
ZI—
gI—
94—
ge—
Lz—
Satis
pt
14,
69
gor
Qri
SgI
ori
101
29
<9
OZ
O11
LZ
Cf1
ZZ
40)
£6
zO
£6
69
OI!
evi
ial
CVI
evi
If1
tg
or
ge
62
£g
60
Ig
fol
LZ
QzI
grI
€gi
28
i
€z
101
OLI
QZ
OOI
£e1
O£I
ZII
LY
59
+6
OI!
ScI
OgI
of-++-
bi-+
z+
Cri—
1€+
Sp+
oz-++
ce—
Z9—
ogo—
os—
010
ai
gv
PS
Sor
6z1
ori
6rI
101
QzI
Zl
oZ
Lz1
IzI
gol
gs
ZIl
Oll
gr
ZZ
49
£Z
6S
gs
ov
ss
59
29
AD
os
1S
©8
OgI
ozz
voz
ORI
+6
gs
1Z
QZ
Sor
VOI
98
Lzt
gt
QII
QZ
Lv
>
co
6S
rg—
IlI—
zoI—
cei—
16—
gs—
601I—
zoI—
IzI—
Zg—
g6—
6Z—
OSes
rlZ—
1s—
61+
61+
gz-+
zo+
oe+
of—
69—
¢Z—
€$-+-
ge+
¥£+-
=
6S+-
ge—
ZI—
if pp ate
gz
O6zI
QZ
£oI
691
gol
6z
cz
cw
61
61
9z
£8
zol
08
ev
49
1zz
10Z
GLI
Ce1
O£1
61z
Zg1
€gi
g4I
bez
1Sz
Z0z
gZ1
SII
€Z
°
oz
£9
OII
ZIl
evi
ZzI
Lz1
191
ofl
O£I
z6
1g
yy
Ze+
61+
c1—
ss—
ZI—
1Z—
vg—
bg—
gt—
ze+
££-+-
r+
cb
of-+
gt+
gs-++
9£+
zb+
Sp-+
ci—
£--
of—
oz—
34—
sf—
eg—
zZI—
zh
6S-+-
or_lL
€Z
88
6ZI
OL1
Sez
VSI
£O1
1S
6S
ZZ
eZ
9S
ov
vol
6II
z6
58
£Z
10
vor
ozI
g¢
gz
92
ber
OfI
1S
101
IZ1
ver
Sez
Zg1
SEI
£6
oS
ce
o£
VI
Ol
QI
ev
ZS
Lb
as
00
88
88
ve1
ori
fer
101
16
06
zB
LG
rC
‘20
“AON
0
7309S
“sny
Ane
oun
Avy
‘aidy
“Ie
‘Pa
wel
PSOI
29q
"AON
pO
*ydaS
“ony
Ayn
oun
Ae
‘Idy
“Ie
Pu
‘uel
ESOI
‘20(q
“AON
0
1093S
“sny
An
oun
Lowmr
41
£+ 26 Sg
4I— zol Oil
+ QO0I 9O1
cs— 16 96
I+ 66 gf
4I— 26 601
gz— 6 €z1
ve— 94 Oo
é— 7 68
6z— 96 SzI
of— OO1 O€1
S9— 69 vel
Po xn 1g
0 £3 £8
vi+ 16 LL
Z4b-+- Igt v1
11+ Sor YvS1
c+ zor Zg1
Z43— vol I9l1
oI— 98 96
or+ zg cb
1+ oor 69
gor+ 941 OZ
OI-+ &Z 6s
VY pedaries 4seo
OQ 2404
Ue —’
2 'd
‘UO}SUIYSe A
gv— £z 69
ve— 2 98
Q-—- =Ozi ZII
SZ+ 9z1 gO
oO gll 911
1i— £4 gol
€z+ gzi Sol
f— gz 621
z— VOI 921
fy— or oS!
OI— zi EI
Z8— 601 oOf1
1S— 69 ozI
gy— 19 601
Sz— zl £6
Sis 789 98
vI— 23 96
ve— OZ ool
I+ 01 901
4~— vol i€1
ehi— 26 sez
fo— zZ Sgl
6Z— z9 IVI
6+ 2g 94
VY pedtes 4svo
"GO 940g
——
‘a][IAseu0g J,
fy— £9 gor z+ 66 49 OlI— OV gSI Ig— o£ 1Si ez EZ 1S Sz— 33 €11
gr— 1S 66 oI+ zo zg GZ— 92 oor 09—— Se vir 61I— VE es 6— oor 601
ve— OL oor f€1— 6¥ zg 9g— v9 ost 6S— VI £2 49— gz 56 62— 19 £6
vit+ gor 26 H&— =or 94 O-- S45. FE =So== “OF. z6 QoI— gz ver or— 1S 16
1v+ 611 QZ ve+ 6o1 Sg I— “er Shi. Se— 0s $6 voi+ Sez Iz21 S9— II gZ
Gz-- SII 06 ve+ 9QII 2g vEe-+ i191 “421 be— 14 sé o1i+ Sez SII g— ve zy
Si+ Sir oor 19+ It 04 e+ £2 cs z€— 0g gir mf1t-+ 2Se.- ovr —e2f-+- “SZ cv
1z+ vor &€ vi— 1g ) 41+ OV ze 6z— ¢S zg 1v— GZ oir S€+ 46 zg
z— SL £6 6&— 2 £11 6+ of te ‘Ge— aE 209 S1— <S2- “99 r+ 22-165
99— 5S ozI 6f— 98 Sei S— £9 89 LO— =e 62 ov— gt £6 6— &S v9
Ze—" 59 zol I— 69 oZ Si+ Zor 26 Zo— €& 06 IS— zg €f1 i— 9g 4
61— ¢€Z zo o1-+ 9S gh Iz— cfr €SI get gol og v€— cor ofr g€— 111 OVI
zi— + &Z Sg QI— gf vS z— gSr ogi o4+ 96 gz oS+ i191 Sor S¥— gtr €g1
zI— of 88 e+ ZL SZ z+ 61 91 1+ 9 Lv oI— 3 v6 vo— zol 9ggI
z— £9 fOr t-- -i0rn — 419 te+ tar oor tvh— ZF 1g oz— 99 98 cse— o1 S¥VI
oI— vor VII g+ Zit vor g— of: gti gi— ZZ s6 vi—- 9Z 6&— Z e1l
€1— 96 601 €— ozr zi €1— Zer ofr Sh— $6 fv1 z— gf og o+ for £6
Ly— 08 Lei (65-166 gsi Zi— oor Zir -. €5— 62 zf1 61+ 9g6 62 Iz+ O11 68
L4S— zg 6£1 9oS— £6 Ovi Z— gir Sei 92— 02 ohr goi1+ ver gZ wv-+- 21% 52
o9— €&Z Zor bvS— £9 Lit 0 OOI OO1 og— gZ gsr €&+ der 6 vi+ ZOoI g8
6— 99 ZL or— 9 vor or+ 101 19 Sgs— 92 €St “=DLI-- fit 0 Z11— +9 IgI
oI— 12 Ig ov— 16 “Er goz4- Sir 29° oS— 6S 601 oO ev ev zg— £2 SSI
co— $2 Gol zi 16 fer S— Sf og £e— Z4— oor J=— 94 of “i= FL “alr
ve— SZ 66 Zi— oor “Zr ¢sS— LZ ofr ge+ 2g 19 cSvy— 2% Zzr €— 2g OIL
V peAtss seo Vv PeAtos 3seo Vv PpeAtes 4seo Vv peaAtes 4Seo Vv peAtss 4svo Vv peAtes seo
"qO 840g "GO 9-840 "qO “240 "dO = --340T QO 07 qO = -a40
aut ‘OW “qsSeM xOW 'N HITED qe
‘med 3S ‘sInoy 4S ‘queyods ‘aq eyes ‘ouIpieusog ues ‘axe’T Hes
ad
panuyuoI—Ol Fav],
22d
“AON
‘pO
*3daS
‘any
sme
oun
key
aidy
“Ie
‘Qa
‘uel
9SOr
22d
“AON
‘pO
*3d9S
‘sny
Ame
oun
ACW
“Idy
“ey
‘Pat
42
$Z—
LL—
61—
s+
4I—
o1+
eg+
z9+
Sr+
11+
ye+
61+
59
9
£9
Of!
gor
96
ori
Sol
1S1
£1
LEI
gor
1g
<9
oS
19
98
£g
88
£8
+6
obi
IvI
cI
SII
SzI
Sz1
98
£9
£o1
gol
IfI
601
gil
601
c6
gol
£9
11
IZ
vor
6r—
36—
¢Z—
17—
gI+
ze+
ce+
3i—
3 to
$9+
ss-+
ee1+
bg+
€Z+-
€c-+
eet
o¢-+
bi+
>
c—
vo—
6b
gs
09
ZO
Sg
OII
QzI
zSI
EI
zi
88
$Z
OII
PII
Pg
SE1
LZz1
16
gil
Ifl
IZI
rg
97
El
LYI
SEI
gol
+6
gol
ZII
St
ori
101
&g
+s
6S
1S
1S
PS
IOI
Zol
£11
I1+
git
s+
=
Sh—
Lv
gS
bZ
69
£9
€Z
6S
6€
ne
Le
gr
$S
58
Ozl
gII
€zI
£zI
For
18
ss
19
89
z6
oor
IOI
III
gol
£o1
QII
vzI
£8
16
Ill
IZI
oSI
1QI
Cer
eIl
VII
98
9Z
1Z
16
gol
e+
e+
co—
Iv+
of+
6e+
ZI+
11+
cc—
Ly—
or1+
os
cb—
Sg—
re
03+
ef1+
bS+-
gs-+
4+
1¢+
89
gol
16
£S1
6£1
If
92
48
1Z
cZ
LL
CII
gil
98
Qs
zgl
Coz
Ibe
VSI
gr
66
621
$9
V4
FOI
Qz1
ZII
z6
6S
94
czi
col
Cor
OIl
QzI
PEI
ff
QzI
CZ
9/+
$+
LL—
th—
gI-+
1€+
te+
ge+
ss+-
of-+
Ze+
ZgI
QzI
66
Lv
c6
ZII
Sel
OII
16
PEL
ovi
OVI
I0I
Io!
IzI
gol
vel
gel
101
v6
62
OII
cz1
€or
66
ool
£6
to
ey
cv
+6
Czr
bri
f+
zI—
ty—
or—
1Z—
16—
€c+
€11+
zoI-+
ev
or+
bri+
4$-+-
bi
gh—
€z+
6z¢+
o¢+
z1I+
€11+
16+
e+
oZ
$6
QzI
oS!
gIl
v6
ge
14
SzI
991
SSI
96
vel
ZO0z
£61
Z91
Cor
88
ZO
grI
6rI
gSI
Cor
£9
Lol
1Z1
€Z1
Sgr
6Z1
ZIl
)
es
es
88
Qf1
eS1
LY1
QZ
ZII
£s
of
£9
zol
IvI— 02
ofI— £1
101+ zz
Ccib-++ tor
Sof+ 96f
O8t+ géI
ss— of
oS+ ger
601+ 9goz
16+ ebz
oI+ oI
€1+ ri
z— SOI
ozi-+- ozz
So+ zogl
be+ FEI
ZII— 0
vS+ Z&1
oviI-+ g1z
QgI+ tz
9I+ 36
gz— 9S
9— 4
Z+ 16
191
£1
IfI
16
26
58
/
aX
L6
zSI
PSI
Ifl
LZ
ool
£6
OOI
ZII
£o1
/
~
gs
€or
rg
£3—
c6—
IlI—
SzI—
IlI—
ZQ—
61I—
Sr++
9Z+
z+
er—
eS—
1g—
z+
SZ-+-
L£L+-
IzI+
49+
ol—
Se
Vil
CII
140)
z6
£9
OOI
IOI
ecr
gr
€gi
LSI
£11
z6
SEI
Cer
6£1
Lz1
gII
Lh
06
+6
36
gor
69
29
06
€z1
SSI
gil
43
44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
of discrepancies from pages 2-3 of my paper cited in footnote 1, d,
above:
8. Of 100 years of St. Louis precipitation forecasted, 70 seem fairly satisfactory
and yield high correlation coefficients with the events. The failure of the other
30 is reasonably explained.
9. As shown by Dr. W. J. Humphreys in his “Physics of the Air,” figure 227,
great volcanic eruptions, which throw high columns of vapor and dust, pro-
foundly modify weather. He cites the first four cases in the following list [here
my table 11], and I add several more.
TABLE 11.—Great atmospheric disturbing causes
Approximate
dates Volcanic eruptions
TOHOMeeials Gale er aciale levelalote Cotopaxi and others.
PGSG= TOO: oes: a ers ovale Krakatoa and others.
TOOT=1OO4) oh iat- ers eeciets Pele, Santa Maria, Colima, and others.
LOUD Hye epe sree este steta ciate Katmai.
TOZATANAILOZB xenon Many great eruptions.
TOZOM pe rheres ae se cic Great eruptions.
TOAT OU eee eye par eNotes Niuafoo Island.
10. Of 30 unsatisfactory years, in 100 years of synthesis of St. Louis precipi-
tation, these lie in groups as follows: 1854 first half; 1856 to 1860; 1887 to
1889; 1900; 1901; 1905 to 1907; 1912 last half; 1913 first half; 1915 to 1917; 1020;
1923 to 1926; 1930; 1940 to 1950. It will be seen that many of these unsatis-
factory intervals fall either soon after tremendous volcanic eruptions occurred
or there was tremendous use of explosives in war or explosions of atomic bombs.
As has been pointed out, atmospheric changes alter the lags in the weather effects
of all solar impulses, and of course unequal periods have unequal lags. These
unusual atmospheric disturbances may very well have mixed up the timing of
terrestrial responses to the 23 periods so as to cause the events to differ from the
predictions.
At some future time it may be possible to connect theoretically the
displacements found in my forecasts with causes producing atmos-
pheric alterations of importance in weather. As yet I have been
unable to name with certainty causes operative to produce these occa-
sional displacements. For the practical inquiries of farmers, however,
it is of importance to estimate the magnitude of forecasting error
rather than the cause attending such discrepancies.
As a step toward that, I cite the case of Spokane, Wash., figure 11.
A computation made in 1957 derived a “correlation coefficient” of
59+5 percent over the interval March 1950 through October 1956
between forecast and event in Spokane precipitation. In simple lan-
guage this means that my forecast represented the observed precipi-
tation 59 percent perfectly for almost 7 years.
45
ABBOT
S. PRECIPITATION
FORECAST OF U.
NO. 9
*S9AIND Po}JOp ‘sjudAa { sSaAInd [[NF ‘sjseovIOY “D *q
‘UO}SuIYse AA ‘SAAIND JOMOT f*yseEAA ‘oUeYOdS ‘saasnd Jadd~, “gS61-OS61 ‘syuaAa pue s}sedaIOF Jo UOSIIedUIOQ—TI “OIA
a3Ad2SaC} Ntvzj 40 SHINO ByOW 2% OOO! 49 sqgaoosay NO S1S3y NoO1L9I10d3aaq S,HINO\|W/ wy7
°
SNWA\J PNINNNA HINO S fF
WAON Wous SIUNLavd3g LNIIAAg
46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
When records through 1958 became available two considerable dis-
crepancies between forecast and event were noted. In the months
January and March 1950 heavy precipitation (over twice the normal
even in 3-month running means) raised the February observed curve
far above the predicted curve. Both curves, as has been said, are
smoothed by using 3-month running means in all computations, hence
the February effect. Not until April did the two curves come close
together. Yet there was a difference of only 6 percent of normal
precipitation between the averages of their heights, January-April,
1950.
Beginning October 1956, and extending through August 1957,
there was a shift of 5 months, leaving the predicted curve in the rear,
and exposing opposed high and low values of the prediction and
event. When the two curves were averaged over this interval of 11
months, the predicted curve was 116 percent of the normal and the
observed curve 96 percent of the normal.
To sum up: At Spokane, in the 9-year interval, my forecast gives
for over 7 years a correlation with observations of 59 percent. Two
intervals of marked discrepancy occurred. The first, of 4 months,
culminating with February 1950, was obviously caused by extraordi-
nary precipitation in two almost adjacent months. It produced a
difference of only 6 percent between the averages over these 4 months.
The second discrepancy, extending 11 months, was of unknown cause.
It involved a 5-month shift of phases and produced 20 percent dif-
ference between forecast and event in average precipitation over those
II months.
Having set forth those discrepancies I remark that this is in the
infancy of my method of forecasting, before any help has come to
me from theoretical meteorologists. It may be that some of them
will discover the causes of occasional displacements of features be-
tween forecast and event. If so, it may reduce error of forecasts
greatly. Then, too, my method assumes that the average behavior of
periods in weather in a thousand months that are past will be followed
in the months to come. It perforce neglects changed conditions which
may arise from unpredictable storms, volcanoes, or even from man’s
interposition, as from forest destruction, invention of new powerful
devices, wars and the like. Even a minor atmospheric change may
alter the time of a feature in precipitation by a month. All these
factors tend to lower the coefficient of correlation.
The 273-month period in weather features——It will have occurred
to some readers that if one were backcasting from April 1927 or from
July 1904 he would employ the same tabular data that I have used
NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 47
in forecasting from January 1950. Hence one might infer that the
precipitation following these earlier dates should parallel that follow-
ing January 1950.
There is indeed a partial similarity, as I pointed out many years
ago, between the march of weather at successive intervals of 273
months. But the correspondence is very imperfect. This appears in
figure 2, where the precipitation at Nashville following July 1904 is
compared to that following January 1950. However, I call attention
to the close agreement of the two curves for the last three years of
the comparison. I have computed for several cities, including Nash-
ville, the coefficients of correlation of the observed precipitation fol-
lowing April 1927 and July 1904, and compared with the forecast
made to follow January 1950. These coefficients have fallen between
18 and 22 percent, while, as stated in my foreword, the correlation
following January 1950 ranges from 52 to 59 percent.
This difference is easily explained. Over 40 percent of perfect 100
percent correlation is unpredictable as yet. There are several causes.
(a) There is occasional unusual precipitation, as occurred in January
and March 1950 at Spokane. (b) There are displacements of fea-
tures as yet unexplainable. (c) The graphs I have published show
large discrepancies in amplitude between forecast and event of ob-
viously identical features. (d) Unpredictable events occur to alter
weather from the averages of 1,000 months.
In the march of precipitation from April 1927 and from July 1904,
the vicissitudes of the later years up to January 1950 cannot have
affected the observed precipitation of the earlier times as they have
done that following 1950. As such vicissitudes account for 40 per-
cent and more in coefficients of correlation, the tabulation suited to
January 1950 can only roughly forecast what follows these earlier
dates.
FORECASTS, 1959 TO 1967
Table 12 gives for 32 stations for the interval 1959-1967 the ex-
pected monthly mean percentages of the normal precipitation tabu-
lated in table 9. The reader will recall that all forecasts are made
from 3-month running means taken from published monthly mean
values, and expressed in percentages of the normal values of table 9.
Expressing these forecasts in a more usable form, table 13 gives
average percentages of the normal for the intervals January-April,
May-August, September-December, of each year, 1959 to 1967,
inclusive.
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
G+ $+
Open
gh+ b+
9961 1961
pi+ e+
pS— 1€+
v6— + Si+
cl—- v—
oSs— 6 —
So se—
6z+ 6I—
19-- €c—
Zy+ e£e—
os+ I —
€r+ €e-+
1+ 2+
$96I og6r
De Ee
€+ 6r+
9o4- 2
Oo-- co—
O03 = se
oI+ zI—
lZz+ £€S-+-
era scr
“E+ 39+
bee) 58
6z— 6f-+
to6r 66x
‘OOD
‘1aAUueqd
Teo
6Z+ €—
vl+ ce—
996r r96r
tit+ tbh—
ZI— += Iv—
f+ Sze-+-
eve; = ¥O=-
Of-- _ &2--
VGa- = east
Baap ies
6Z-- ~2—
POs Ve
12-99
lz+ 0o9—
$961 og6r
c+ 7z—
4Z£=-- 1%—
1+ r=
ev—= —of--
a= 0a
€c— 6¢+
vS— SE
gy— vi—
Te A00—
Sze— sr—
ie
Qe =Sis-
$96r =66S6r
Enea
orto
‘IyeuUToUID
Ze+ “E+
QS-- O1-+
49+ S1—
9968 I961
o9+ Le—
7st+ gr—
12+ Sy—
OI-=>-Si--
Z£— 1+
SI— oz+
t= “go--
of Sb
z+ b+
1f-- de--
OI-+ I1+
$96r og6r
vit 6+
oO 1+
b— Se+
¢— i+
€c-+- €&-+
w+ ce+
Ze £9 4
9§— Ze—
z~— 2—
I+ 6z—
¢— tv—
yz— 2S—
voor 661
ee
‘O'S
‘uojsalieqa
6 a[qeL ‘q pue Y suWNjod Woy soinjiedsp sSejuIdINg
a fee
g— 4+
e+ ¢1—
9961 1961
£1-- <3 —
or+ 9 —
VO=vi=t
9 — OI—
te+ Lz—
1+ SI—
Z+ e+
1I— 2 —
O¢— ~62--
gze+ €& —
I1v+ = ge+
gI+ I —
$96r og6r
C= als
oz+ ozc+
“Z1-- ~ LZy--
QZ— ~2c-|-
pS— y+
99- 2+
£e— 9c—
z+ So—
SOnr a=
fo+ ze—
QZ+ 9f—
w+ 2+
bo6r 86561
OT
dN
yp1eUsig
Og TI=F
of gf
A (0,
996% 1961
€+ m+
4— i1z2—
ZE— 6¢—
Oo 0g
61— cb—
Z+ zeb—
ge- bve—
op+ 24+
w+ v—
zZt 2@—
gr+ = =FI—
ov+ oO
$961 0961
z— 9 —
oz— f+
GI" "892-|=
gf— gt+
vVI— ZiI+
e+ 61+
Si-- ge--
¢+ 12+
cz— viI—
fc— oF —
lz— vhr—
SOLS
$961 6561
ee
"ep
‘ejsnany
41+ 6—
oS
y-+- ce+
996% I961
4i+ 61+
IlI-+ giI-+
Zl ~SE--
Zo-+- oS-+-
Zz-+- 9S--
Iz+ 99+
Si 50s
iz— 61--
Eos SLea.
Soa eae
ZI viI—
tO
SQ6I o9g61
te+ of-+
8 te <984-
€1— o6ob+
g— 6r+
ZE— 6 +
oI— I—
4s— ov—
Ii— _—- 1Z—
vo zQ—
990 Se=
Iv 2 —
w+ LIi—
bO6I 6f6r
——
“2310
‘Aueqry
uoynji1gi9a4g¢ JSVIaAOJ—TI AVL
ot
csIi+ +¥—
ve+ c+
9961 I96r
i+ 72+
ozc+ e+
6f+- 6 —
fc+ o7c—
€— ti2e+
I— Z£+
Qi se 81s
i a il
oc+ 61+
e+ ve+
£-+ 92-+-
2— 9giI+
$961 og6r
ied ae
eee
4Ir 9+
gi— £ +
z— 6 —
42— GS +
vb— e+
ge— —s: TI—
ZI— + gI—
¢+ 1&—
ai—— te
£- oc—
bo6r 8666561
en ae
ACN
ueqly
gE+ 98+
99+ b+
o£ ev+
9961 I961
61+ 9gI+
61— or+
zvy— o9g+
6rb— 9S+
e+ ¢+
gofe+ z2—
cs+ oI—
gh+ oO
goI+ oz—
gor-+ 9€—
6g-+ Si—
e+ +4 —
$96r og6r
mee eae
Z2— oc—
6S— €z—
Zo— 6¢—
Zs— 6b—
I— 6+
6S-+ S —
O0-ee Sha
Zy+ ce—
e¢+ Z4—
b+ 7z—
iv— ti-+
b96r 6S6r
J
*xoL
“aualiqv
scinces “SBN
e@eeveee ‘qoJ
eeeeeee ‘uel
eeeeeece ‘29q
eeeeee “AON
eeereeee ‘po
eeeeee *ydaS
sq}UOyy
49
FORECAST OF U. S. PRECIPITATION—ABBOT
NO. 9
zI+
zi+
Bt
be+
of-+
ze+
€1-+
Bort
Sis
zI—
Cc1+-
ge-+
1961
99+
bl+-
1v+
re—
oe
6z—
Ib—
oI—
oo
be-+-
oI—
II—
of—
zZI—
f1—
cz—
c=
£961
Ii-—
cz—
re—
61—
zl—
16—
6£—
yl—
¢l—
e961
re—
11+
6 +
€z+
Zz+
zb+-
tear
gI-+
gé+
Or
oe
Go
1h
95—
fgo—
£961
ce—
II—
or—
—
Zz—
ios
ZI—
gi+
Zz+
gt+
Sec
cse+
$Z+
99+
Z4¢-+-
£961
gI-+-
vi—
Lz—
€ +
vI—
y+
fe-+-
£ +
II—
to
Sr—
2961
1z—
gI—
git
zs+
wy+
zs+
Si
8 —_—
f9+
bh
Se
git
L£=—
oI—
2961
C=:
Soci
9f—
ss—
2961
Zz+
bi-+
ge-+
gi+
ce-+
of-++
Zz+
Z1+
o1+
bose
ae
Vey
L961
Scc+
zc+
Or
fo
oI—
zi+
1v+
1b-+-
Soe
Ib—
Sie
eo—
£961
gI+
Ib-+-
€c-+-
11+
(panuijzuo?)
I-
€+
Sce+
ce+
Si+
bi-+
QF
Tost
gi—
O--
£961
€1+
2 =
bi—
zo—
3e—
ZI—
Qe
g1+
rp —
€c—
re—
61—
61—
of-+
gi+
Z1-+
Ic+
£961
2961
e1+
or-+
6z-+-
oc—
2961
61+
f1-+-
€S--
gb+-
o1-+-
or
bi+
7—
gt—
gr—
6z—
61+
61—
Vase
€1-+
o1—
Ii-_
£961
61+
Z1+
z+
L+
eat
ort
Et
vi-+
Le-+
a
oi1+
zi+
6 +
be+
gz+-
o1-++
Ic+
c¢—
gz—
2961
oar
or—
8i—
zI—
gI—
Or
SiS
Zi+
LI—
SF
c1—
ZI—
6 —F
Vick
£961
oI-+
€1-+
61+
ci+
II—
a —
£1—
Woo
Lz—
11+
Si+
bet
vi-+
obv-++
€r-+
Z1+
2961
61+
Sp
w—
va
oI+
17+
feet
c1—
y—
61+
1€-+-
63+
6¢-+
£961
ae
LI—
be+
Si+
8e—
Sze—
of—
c=
Sc+-
£¢+
2901
1g+
gI-+
eeeeee
eee eeee
-99q7
+ “AON
pO
"yd2S
‘any
inf
oun
Kew
“Idy
"IVI
‘Pa
‘uel
‘20
"AON
pO
*yd9S
‘sny
An
oun
Ae
“Idy
‘ILI
Fees gout
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
50
g€+ 9+
Bele ea
OV-— 2S-F
9961 I961
I— gt
Sc Ol—
tc—) Qi
Go Gear
ee Ce
e+ + +
Olea {poeta le
cet+ = te+
69+ 1+
Oar 19ea
09-- vi—
gt of—
$96r og6r
0) ie
po aes:
Oe hea
Of=— 0451
te==- <20--
Bien em Ovaie
Ol Ons
Cte a
Ota to
ea een oa
Zoo Le
ic Ze
bo6r 6561
rs
"ely
‘£1903 UOT
19—— seo
gr— f+
AEE) oe
z— o-+
9961 1961
ot
Oo el
to Oe
99+ 91+
dea 6Qeat
€ — c+
fci— v—
LZ— gi—
Te a
fea Gr
teal ae —
QI Zn
$96r og61
te+ tvi—
£4 te+
es Sv
~— se+
Soop. Gar
Bie Olle
VI— 9c--
css— gI-+
of— 9 —
Ol Ol
ae a=
zi— 3s—
b96r 86661
—
RSENS
‘UOSIpe
6z+ € —
Ze+ =SI—
GRaj= GE—
or-+ of—
9961 IQ6r
€1-+ 61+
of— E+
vo— cet
vo— LZy+
fg9— Ss +
€y— ogi+
4— 9+
QU Bea
vo+ o1+
69+ VI-+
O97 = Gua
Wamp Ole
$o6r 0961
sz— gI—
gr— e+
og— 62+
Qh Vest
OS —- ou
Ole sera:
9e-- hI—
e+ ZI—
Lo-- ve—
or-- o1—
¢+ tvi+
Sr
bo6r =: OS OT
a
“2V
‘90y eT
vi— gi-+
Se+ 1¢—
e+ vI—
9961 I961
a
11+ 9+
Caan Ova
of+ vr-+
ge+ 6€+
Ze-+- = 19--
of-+- 6€--
re+ or+
6c+ 6+
Ol siec—
oi+ ¢e—
Lia
$96% og6r
es— 6 —
1~— 91+
Gea 00a:
Zz+ ofZ+
ee+ 7+
vi GS-+
g+ or+
bi+ ci+
6+ 9gI—
€— os—
te— Sl—
orp— Ll—
bo6r 6S6r
ee
‘suey
‘gouapuadapuy
panuyuoI—e1 ATAVY,
y— vS+
Cleo.
fc-- ZE=--
9961 I96r
~¢— oi+
ev+ 8 €1+
6€-+- 09+
b€+ 69+
Obs elvae
Of sect
$o6r og6r
6+ oz—
bi— e+
2+ vS+
Sit gst
Vil c ieee cO ate
Z-+ oe+
Ly-+- €z2--
w+ 9gI—
6I— I19—
es— zg—
zs— 1f—
1v— tvi—
to6r 8666S 6r
.
“‘qUOyW
‘eualaH
46— 9e--
Qo set
Sy— S€+
CO oe
996I 1961
iit Ol
6vI— 3g —
gr+ = 1I—
vor+ 1So—
gori+ zi—
v6+ 69+
S6+ tvor+
96+ gét
Zo-- O04.
e+ Wt
OS4> *Zb--
$96I og6r
Lyle lave
of + +
a2-- Ot
zo+- SI—
O24 ee esate
G7 eee rae
bz f9+
OC aia
oie
6— z+
6+ 65+
€-+ ght
hoor §=66S OT
sexay,
‘oseg IH
OI o1-+
ti— oz+
ef— $i +
9961 IQ6I
6+ 6r—
6r+ vo—
gy+ re—
of+ £1—
oi+ tv —
b-— oc+
g— e+
9I— VvI--
ef ON
£9— 0z—
OQ eee Ga
GO VS —
$96r og6r
So) Og
oI+ 7—
ce+ g—
f+ gI+
Ze+ obv+
€I- z+
Se+ o1+
OIF = Ors:
y+ 6—
emg es
L— oz—
oz~— 41—
b961 OS6r
ET
oureyW
‘ya0d\seq
I— Iv—
of— re—
Ce ee
4i+ SiI—
9961 I961
i+ € —
of+ gc-+
get ght
ie ee eet
Sit nO tae
oft Oe
fe) or —
Lice ge
625 ic —
of— 2 —
€e— ze+
$96r og6r
gs— ss-+
89——~ sha
£S— 1+
€1— vet
Z4I+ “Zi+
Z+ 1I—
eSt+ €1—
r+ g—
pe+ 2z~—
(A
Se— ¢I—
6r— te—
bO6r 6S6r
uu, —Y
“QoL
‘plorjaq
eeeeeee “idy
veers ey
teeeeee cggg
veseeee sues
eeeoese ‘29
eeeeee “AON
ereesee ‘po
eeeeee *¥d9S
eeesee “ony
eoeeeee Ajn(
eeoseee oun
eeeeee Ae
see iON
eeeees “Ie
eoeeeee ‘qoyy
eevee eee ‘uel
teseeee Q9q
reese “AONT
reeeees 199
reese adag
eeeeee ‘sny
eooeeee Ajnf
oun
eeevre AeW
susevekened’ Jen Cy,
teeeee pW
eoeeeee ‘qa
eeeoeeee ‘uel
sqquOW
51
FORECAST OF U. S. PRECIPITATION—ABBOT
NO. 9
ze+
Ze+
€S+
1Z+
zb+
1Z7—
7—
vi—
Os
oI—
2961
oc+
gt+
oc-+-
Sls
zi+
t1i+
S9—
Ly—
9 —
gi+
gi
2961
€r+
ci+
ob-+
oz-+
ge+
£b—
or1+
fz+
ci+
fet
61+
be+
z+
6¢-++
Lz-+-
2961
Gra
ci+
1+
Ti+
aie
[ie
zI—
or+
be-+
6€-+-
ve-+-
ei1+
oz+
2961
eet
gz+
Scot
oz+
ei1-+
0z-++-
ge+
Ib-+
s+
Z1+
8e—
2961
gi-+
1g-+-+
€1+
8e—
Lz—
z+
€z+
Iz+
vo+
€z+
si—
61+
I—_—
Done
bi+
2961
6 +
2s
ci+
Teste
1I—
zZg—
gi-+
bi+
Sr+
to+
Zz+
9e-F
[o—
SI
4o—
2961
Li
coat
bi+
ci+
(panuijuo2)
Coa
zf+
Zy+
6r-+
34°
oz—
b=
ci+
1g¢+
or+
Zi+
ze+
£961
zb+
2o-+
ze
Z1—
eo
Co:
ve—
gi
Ls
ve—
fects
41+
Qeat
be-+
1c—
est
Tp
1+
£961
(0) oe
gi—
€c+
cp
t1+
ce+
of-++
ce—
Si—
e1—
Ve
ov—
2961
oI—
1zZ--
6 +
(oe
ots
Tea
9 +
1Z7—
or—
Le—
6z—
9 —
ge+
1v-+
€ +
c1—
6z—
4961
61—
Ip—
g4—
SQ++
ze-+
or1+
f9—
[7—
ysS—
og—
or—
g1+
1 +
ve+
Zi--
6z-+
1I—
ss—
£961
ZQ—
[9—
ve—
61+
ge+
or+
be-+-
6r—
or—
6r—
ss—
2961
c1—
be—
7e—
v=
ve—
I—_
tact
Gat
6z-+
1g-+-
oz+
Z—
0
Lz—
osS—
g5—
ev—
£061
9I— 61—
cit v—
Z-+- 21+
0 A
ee— 9+
Zy— oS+
of— 9+
Ol) se
6— 4£-—
6i= Ze
Le a
iu— €-+4
496% 2961
9S— zi+
gof&— zS-+
1+ ei+
GO-- aia
ooI-- 9g —
zoi+ ti+
61—
ci—
zi+
€c+
4961
gz-+
of+
LZz-+
z1i+
66-++
zg+
aa
re—
II—
Gees
ga
Sp
gz+
8 _—
be—
89-—
sy
gh—
£961
ci—
zI—
zI—
C=
LE =
i
ZI—
Z1I—
WP os
2961
6z+
gi-+
Ze+
0z-++
9z+-
gz+
eeereee 20,
eeeses "AON
erreree ‘po
eoeeee *yd9S
eeeeee ‘sny
eee eeee Ayn[
eeeere oun
eeeeee ACW
oioteiele:ss eet
eereee “Ie
e@eeeeeve Qu
eoeveeee ‘uel
eeeeee “any
eeeeeee An
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
52
aA —a —_- — — 4 pada -_=
oI+ 6 — Zt e+ gp+ 4— QI— Sz-+ ¢— ort gI— gf+ gI+ zw— 4£— e+
zo+ ¢i— 6€-+- SZ+ i6-- 2— Of-+- 97+ gi— + =$Z+ 4+ 1€+ vi+ 61— bi-- 12+
99+ SI— o6+ 11+ 99+ Le— zI+ € — gz— 39+ Sy+ zit Z— z— 6S+ 6€-+
z+ oO 411+ vii+ gv+ gz— 6+ i+ ez— $+ gI+ Zi+ 9I+ gg— 6r-+ c+
9961 I961 9961 I961 996% 1961 9961 I961 996I I961 996r I961 9961 I961 996r 196%
ZI— + 9f— £6+ 66+ Ly+- 3t— z+ s+ Si— 4+ Zze+ 0 ¢— — oO Z—
z— tf— IZI-+ 06+ 41+ sz— ie—- -“Vi—= Set 200-1 zo— I+ OI— of€+ gf— gI—
gz— vI— L£Z+ 9+ 1I-+- e+ e— Si Oe 29-1 v6— E-+- Zs— £0-- f— I—
f€+ 3-— 8— 92+ t-+ Sbt-+ €+ ov— fz— gI+ f9— St-- zl— 6¢-+ 9—- gt
See Oe fe= 2e— 2 -- ~S9-+- ve-+- So— 61— ¢£— os— 41-1 €Z— -62-+ b+ z+
g+ re— fo == 492+ PSE ape— ig) le Zt— <0 gg—- iI— iit 0
Seo Sia S45 — 1+ 11+ Iz+ 93c— vI— gI— v— Q- 4L— ¢v-+ gI- vI—
Sea eae ia — =F Sia 16.-F oh— vI— QI ae gI— gI+ e— L/¢— fZe+ zi—
gr-- 9 + Ege cee Rea =8e-/ C-- e— Lie ee oO ate of or— zl+ ve—
HC am WS zo+ 62+ 6 1 — 6-4. 6c-- oO gi+ Ze Oz-|- €9-- 65— SE-- =O.
of-+- o1-+ g1i+ &¢+ Si Si ~— I€-+ + tvI— ge+ Sze+ w+ €z— 6-+- 1+
41-- Ti+ Ori-+ 9obv+ I— g— of— 1+ f1-+- 6e— S1-- fe= fi-- —& -- g@+ z+
$96r 0961 S$Q6I og61 $96Ir og6r $96I o9g6r [96% og6r $96r og6r SO6r 0961 $96I og6r
et pvt ezi+ ¢o+ e+ e1+ a 0) oit+ P— ov-+- =1b— Iiz— O1-+- fzc— b+
y— €2+ 6s+ 6+ 4— oI+ cse— go— zI— Ss 6 + ss+ + S1-- S€-- b— gf+
“Zz— ¢S+ 2+ e+ ¢— 61+ 2+ 9 — 61- +F¥— 1+ oz+ gb+ =€s+ Iz— €9+
yE— 19+ goI— rr— 6— 9+ vi— zI— I+ 12— 6I— o6of+ zo+ SP wzo— bl+
SI— 0Z+ f6— £6— gi— 66+ vyr— e2h— ve+ 12— zi+ 61+ 0 Gre sf— oz+
oO 1€-++ gr— tvzi— f1— 3+ Srp— £so— ce+ oz+ €€-+- fe+ b+ lz— gI+ 9 —
zi— 2+ ¥g— of— oI— _-95— oz+ 6r— €r+ of+ co+ f+ g9— oI-+ 6z+ P+
6z— vze+ gz— 6b— €1— 7— ¢— tve— re+ 6c-+ Q— e+ vi+ Zy+ z— se—
zw— 12— Se=- -se— of— Lr— Sgo— zI— Iz+ 12— g+ si+ 46+ 9b+ cc— ch—
Iz— £e— ob+ + — gz— ef — Z4Z— LI— 9+ tr 6— 9gz+ 46+ z+ 6b— + og—
6— 72— vS+ gz+ 6s— 0oz— zS— 7+ vi+ ob— oO $2+5 83+ sr— SZ— ~7—
te— £1— gg-+ 1S-++ zeZ— 6+ “i+ «1+ “2+ ¢e— + 4— vo+ Sh— e+ + —
bo6r §=66S6r bo6r 86661 boor 6961 boor 6S6r to6r 6961 bo6r 86961 bo6r 6f6r to6r 661
‘O(N “yreO ‘A'N “SST “HII “Iq9N ‘wy “‘auay,
panuyjuor—el AAV],
e's Sa — a | fa ~ G's ——_ Pine —-
eeeeee
eeeeee
eoeeree
ee eeee
"IeW
"qq
"Ie
* “qoq
. ‘uel
sqUO WY
53
ABBOT
S. PRECIPITATION
FORECAST OF U.
NO. 9
11+
61+
€z+
61+
¢—
oI—
Oe
ce
of-+-
Sel
ce+
gs+
4961
c¢e+
€z-+
6z-++
¥g—
95—
Z2s—
Q7—
gi—
-
fe-+
Zz-+-
Oi
61I—
gt—
Seis
bi+
Ou
II—
ceo
£961
zz+
¢ +
8e—
er—
ee—
1Z7—
¢i—
61—
gf —
—
9h—
2961
gI—
zS-+
og+
gh+
gz+
Ze+
b+
yy—
gt—
ve—
vi—
1g—
c1—
er
41+
vE-+-
62Z+
SZ
4961
g-+
92+
v2—
v6—
ZI—
1yi—
zhi—
76+
g5+
se—
88—
gzI—
66—
2L—
¢s—
1g—
LE--
9f+
Ta-|-
£061
99+
er-+-
zs-+
e+
IZ—
ce—
g —
z6—
So—
ceo
yfZ+
fo+
2961
gz-+
9—
Zo—
o£—
of—
¢{—
¢o—
Pest
Ip+
Se+
€e+
oI—
zI—
4 ae
¢s—
1S—
be—
oI—
4961
8 —
gI-+-
gi+
zb—
II—
oils
c=
Fi—
6¢-+-
oS+
or-+
cai
(ia ig
1z+
co
£061
oI—
acl
11+
CiL—
6z—
8e—
gi—
oI+
Ons
2961
9—
gi+
2+
1p+
oI—
ZI—
oI—
oz—
co—
Zo
g —
€1-+
eo—
ce—
Ly—
zI—
c+
2961
bi-+
ce—
ei—
goI—
€zI—
gl—
02—
oI+
9e+
6z-++
2961
ze+
gI-+
eb
Lo+
6¢-++
f1—
Le—
(panuyuor)
9 —
6z-+
ob+
ge-+
o¢+
Si+
ee+
ey+
os$-++
ze+
Cr+
zs-+
4961
TZ
ri+
S-=
61+
be+
oz-+
6e+
6¢-+
fe+
OF=-
cn
e+
vi—
lz—
ceo—
ois
De se
Cre
f96r
41+
ei
6z—
2v—
9z2—
61I—
Tot
eos
9 —
Beale
1v+
2961
vi+
Bok
cc—
ov—
gi—
Sz—
¢ —
6z-+
ge+
ee
o1+
OF
1¢—
8e—
¢ +
er
of-+
e+
e+
2961
Lz—
of-+
ze+
I—_—
€ +
zeo—
61+
gb—
Ze—
co—
¢1-+
zo+
gf-+
ve—
¢f—
6g—
so—
oI—
£961
gs+
ce
Leet
Di
8e—
be—
—
Zz+
z—_—
1Z7—
1s—
2961
c1—
Lo
o1+
1¢-+
2961
vI—
one
gist
zI+
8e—
So—
zeo+
Lo
zs—
act
oz—
“lees
61-++
Sce+
ge
ci+
£961
ee
Of
c6—
¥g—
88—
61I—
of —
S9—
99—
viI—
2961
of—
oh—
or+
gS-+
84+
98+
z+
1Z-++
£9+
bS-+-
bi--
fc+
orI+
296r
of-+-
gz-+
Z1-+
oo
€c—
eee eee
Gis.6 6 a) e)s. 2)
eee eee
eee eee
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
54
aes
p= 20
9— e—
Oe ee Oe
Go eel
9961 I961
zI— oOI+
SI— € -+-
or— bvy—
6+ 6s—
Oba aa
ee-- Vet
Tea Ava
gs+ 42+
ce+ z+
re+ f4i—
Gece = Sse
One ver:
$961 og61
ert git
Tee
Vie te
f= Seq:
esa) 2
Zo+ gI—
Z4i+ vi—
co—- - <e—
19g— GE—
pZ— th—
z~— ot —
O—= €1-|-
bo6r 6561
—
Fu
*U0}SUIYSE AA
UZa- 2s aI
Geo St:
ce-+ i+
fi1— of+
ee— oi1+
9961 I96I
of —— Ze
CZ — On
C= 05 —
z— g3I—
Ve—— gl
oe+ 61+
Pees teat
16+ voi-+
6£+ 99+
of+ f+
ea ea
SI— oz+
$961 o9g6r
6I— gt
ve— VvI—
Z£+ eh—
zi+ £45—
gs-- It—
AG = Beate
to+ s€+
gr+ gz+
€+ z+
OI— Si+
sSi1+ g-—
6— I12—
t9o6r = OS 61
UY
2
‘a[IASeuoy T,
ao OO.
0 er
op
OV-- Vv =
Bia oi
996% I961
Z4I— 9—
pe— € +
is— 11+
gy— I+
Sz— 9e+
6I— of+
Zi— 92+
b= ace:
Loom
R05; ces
Ze-- Oc
$96r 0961
ori+ vI—
I— 1€—
6r-- ss—
ceo+ v—
te+ E+
zep+ of+
6z+ Sf+
€+ te+
oI— +viI—
Or-- ~“tz—
git 9¢—
bQ6I 6S6I
“uu
‘neg 4S
Rie Ais
fis> Of--
zeZ+ Ie+
Oe ea
1f-- Qe—
9961 1961
ei
€c— €e--
ame a
Og ais
6zc— +
OS arte
Gc aaa eb ete
o9+ ti+
6v+ o1+
Veo) aes =f
gc— vI—
Bea = eal
$o6r og61
Talo ce
tr+ 0
10-5 (Ol
cr+ 6I1—
6g€— £—
€y— vi-+
cIi— —-¥ +
[= oO -+-
I+ e¢I+
gs— 4E+
CVeeei la 1
VB Glee
too6r 8=66S61
—
OR
‘smoy “3S
papnjIu0J—zZI ATaV
C= ea
ia Ope
Bee OR
gz+ 92—
St-+ ie-+
9961 1961
e+ €-+
tof L+
Sipe eaeele
Zit mI —
ve— Sv-+
of— 7cS+
€Ss— zr+
vo— Zb--
vVE— ge+
OTs = Sie
g9+ sé+
09+ 91+
SQO6I ogor
2 OFF
vo-+- 06--
ef = €gr
a ey
T= Oe
0 S05
cit GS +
z+ 9f—
glt+ 9e—
69-- 1S—
z6+ of—
6Z+ 1m6—
vOOr OS6r
Sa -
"qse MM
‘guexods
CL AO
96— oc—
gi teat
le— t+
Oi hes
9961 I9Q61
62-}-— “SE
zgt+ gh
Sv-- 16—
€z— zg+
QgI— Ib+
Le
Le
4i+ ¢f—
6h+ Lg—
sy eal
g6-- SZ—
gor Or
SQ6r 0961
ob+ 92+
Vi== 102=-
o7— £1--
VO—=— *2e--
WV eS
See
c= 2.55
€z— gz—
eI— —st—
git 9€—
gy =e
to-- 1-{-
v96r 6S6r
Se ae
“HHO
‘ouIpieulag ueS
Or. ih
oe oer
Z2— S$ —
goI— c—
O1I— 61—
9961 I9Q6I
14— o9f—
6&—" or —
zw— ve—
te— ce—
i oc—
€Z+ tv —
io 9c—
ob+ lz—
1z+ tr—
gz— rs—
ev— + ob—
eS— ~—
SQ6r O96!
£e€— of-+
66— 6S-+-
SS— €€-+
Z— ob+
Ze+ Set
4y+ €1+
61+ 41+
OI— SI—
Le— ~—
oI— Sz—
t~e— ¢y—
or— oS—
b96r OS61
ees nena
‘xO ‘N
‘aq eyueS
re wo
61— + ss—
O 6S—
i, Se
9961 I9Q6I
oI+ i+
1S-- 62-1
Vos St -
€S+ of€+
O14: — GVA
Zt cet
tr— 61+
ce— =ei1—
9+ gi—
Ov+ f+
So gi--
SQ6I ogor
e+ of€+
II— ¢b+
gi— 64-4
Ors Sach
re+ b+
6r+ iI +
Se Or
11+ ¢e—
e+ o6of—
le—- +F¥—
b961 © 6S 61
yen
‘aye’] aes
eevee idy
RIS Sse TET
eeveeee ‘Qt
eeeeevee ‘uel
‘20q
ree ees AON
teeeeee 9G
eveeee *ydaS
eeeeee “ony
eeereee Ayne
oun
eeeeee ACTIN
eee ‘Idy
reese rey
eeereees ‘qoyy
eeerereee ‘uel
.
.
.
sq}uOy
55
ABBOT
PRECIPITATION
FORECAST OF U. S.
9
NO.
bi—
zi+
e¢o-+-
zg+
vs
co+
g —
ze—
or
bi+
L961
61+
co—
e1—
ZI—
61—
ze+
9 —
gI+
¢o+-
es-+
1€+
a —
br—
9 —
6z—
re—
69-++
£961
fe -1=
zi+
ce—
or
6e—
Le—
of—
€z+
gr+
zo
296r
gI-+
Zo+
9 —
8e—
2) —
te—
9+
f9o—
yS—
1b—
eae
1 hs
Ct
€e-+-
Ip+
19+
vS+
61+
L961
DUS
£o—
ga—
£o—
6z—
LE
of—
oz—
11+
al
e1—
e1+
Set
Sie
zI—
a
zI+
£961
9$-+
f9+
get
Die
ziI—
2961
ez—
of—
ae
Ly—
gf—
ci—
¢+
6e—
cz—
er+
gI-+-
€c+
zI-+
I-_
oles
lI—
=
£961
of—
Fri—
¢f—
f9—
IbF—
Mele
Si-+
2961
oS-+
ig—
61—
eat
6z+
1s--
oc
Zi+
Iv-+-
cr
ZS++
Oc-+
or1-+
6;
Or
oP
oz~—
1g¢—
49061
gz-+
09+
zo+
€ =F
zQ—
1g—
oI—
Iv+
92+
1S-+-
61+
Oar
Le—
19—
=
TO
£061
Sr—
1z—
8e—
ZI—
€I—
OC
cc—
of—
cI—
Ibp—
296r
cL
gf+
er-+
1¢+
zo+
Ze-+
Si+
ci+
61—
Zi+
€1+
ci+
8e—
Ze—
Gat
9 +
(ee
29061
0)
61+
zg+
69—
oz—
biI—
gt
eet
of
Wp
€96r
1s—
61+
Qe-F
gb
oh
(0)
tc
1s—
f9—
1Q—
6z—
2Q6r
6-3
me
QI
Oats
9 ==
O=-
cIi—
c=—_
OF
bi+
cet
ge+
£+
61+
cot
acts
Cla
£961
ot
oo
Ze—
Zi—
ge
bor
CoA
Zi
61+
61+
c=
£961
or1+
Sc+
Le--
Scc+
O0c—
2961
co—
ol
cet
Zz+
i
Sle
eo+
ge
opt
pS--
ge+
Ze+
41+
cc+
7 —
Sc—
c=
£961
I~-_—
ae
9)
gI-+
gI-+
Yoo
ss—
cc—
1s—
Olias
oI—
ze+
One
6e+
9e+
Si+
6 =-
fe+
£961
e+
be+
S =f
ge
of-+
Qe—
LS—
89—
1f—
61—
296r
® cls
98+
£S+
09-+-
ev+
be+
br
Z1+
erp
OE ohe
Sete
Ch
1g¢—
Ly—
zb—
6e—
ci—
4061
Scot
Zy-+
s+
cL
6z—
lo—
br—
oz~—
II—
oI+
oI—
oz+-
crt
11+
8e—
cz—
£961
zi+
€c+
oi-+
Ze+
gl+
os+
eo+
95—
¢/—
bre—
Or
61—
2961
€c+
r+
pe
1g—
of—
Fi—
eooeeeee
‘Ie
‘Pu
‘uel
‘20
"AON
pO
*yd9S
“ony
Ang
oun
Ae
eee “Idy
oe “Ie
. "qo.7
uel
20
“AON
0
*¥daS
“ony
Anf
oun
56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
At the end of text are 27 maps of the United States with the 32 cities
as listed above, and each accompanied above the circle by a number
identical with the appropriate number in the column headings in
table 13. Below is the predicted departure from normal. Each group
of three maps covers the three intervals per year of four months each
named with table 13. Large areas of approximately equal departures
from normal precipitation are clearly noticeable on the maps. These
area similarities may aid farmers remote from the 32 cities to estimate
the precipitation probable in their locations.
APPLICATIONS
Periods control long-range weather.—I have sought to present to
meteorologists evidence of two important propositions. First, that
there exists in weather a family of periods, all exact submultiples of
273 months. These periods are hidden from immediate recognition
because their phases are shifted according to the state of the atmos-
phere. When, however, the long monthly records are grouped and
reduced with reference to time of the year, sunspot activity, and
march of population, the family of periods is clearly disclosed with
constant length, and with approximate sine-curve forms.
Second, long-range precipitation is to nearly 60 percent governed
by this family of periods. By evaluating the average forms and
amplitudes of these periods from thousand-month records, precipita-
tion and temperature may be forecasted for years in advance, with
considerable approximation to the event.
Whether these forecasts will appear to interested parties as trust-
worthy guides to help in planning their future operations must depend
on the agreement attained between forecasts and events, 1950-1958.
I therefore prepared table 14 which gives for 32 cities the 4-month
forecasts and observations, 1950-1958, and the differences in per-
centages of normal precipitation, A, in the sense observed minus fore-
casted. Their means are given disregarding signs.
Agricultural requirements.—For agricultural purposes a foreknowl-
edge of seasons rather than of individual months is most desired.
Hence I give in table 14 4-month mean values computed from table ro.
But it is the difference between forecast and event which would be
the controlling factor in estimating the value of the forecasts.* The
average differences, A (observed minus scale-corrected forecasts) are
5 As differences in level of observed precipitation, 1950-1958, from the averages
of 1,000 months, are disclosed in table ga, I refer to that table for possible cor-
rections of level which might be applied to values for some stations in table 14.
NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 57
entered at the bottom of the columns of A in table 14. These averages
will be needful to the use of table 15 which is to follow.
Assuming that the degree of success attained in the forecasts from
1950 through 1958 will be attained from 1959 through 1967, I have
prepared table 15 from which the probable sizes and numbers of
discrepancies between forecasts and events in 4-month mean values
over the entire interval of 9 years, 1959-1967, may be estimated.
Selected from table 14, four groups of cities, 25 in all, are tabulated
in table 15. The first group of I1 cities have average 4-month mean
discrepancies, 1950-1958, of about 20 percent between forecasts and
events.
The second group of six cities have mean 4-month discrepancies
of about 26 percent, the third group of five cities, 30 percent, and the
fourth group of three cities, 4o percent. All the percentages relate to
normal precipitation given in table 9, with the scale correetions from
table ga used in table 14.
The six columns of table 15 give, respectively, the numbers of cases
in table 14 when the discrepancies between forecast and event, 1950-
1958, are (a) less than one-fourth, (b) one-fourth to one-half, (c)
one-half to one times, (d) one to one and one-half, (e) one and one-
half to two, and (f) over two times the average discrepancy of the
group.
If the same degree of success is reached 1959-1967 as was reached
1950-1958, the interested person of a city in Group I would expect
the numbers of discrepancies (O-F) among the 4-month means stated
in the mean values at the bottom of the columns of table 15 to occur
in the entire interval of 9 years with magnitudes in percent of the
normals as stated at the top of the columns of the first group. If he
were located at a city of Group 4, the percentages would be twice
as large, because the numbers heading Group 4 are twice those head-
ing Group I. But the numbers of cases would be the same.
Stated numerically, a person residing where the mean departure of
forecast from observation, given in table 14 for 4-month intervals
from 1950 through 1958, was about 20 percent of normal precipita-
tion, may expect the following numbers and magnitudes of departure
from the forecast of 4-month means during the entire 9 years, 1959-
1967, given in table 14.
Numbers of departures..... 4.6 4.5 6.1 6.0 2.8 3.0
Magnitudes in percent...... 0-5 6-10 II-20 21-30 31-40 >40
If he resided where the mean departure given in table 14 was greater,
the numbers of departures as just given would be unchanged, but 7
+ Text continued on page 67.
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
58
bi-+
oz+
e+
e+
ze—
9I+
zs—
zi+
6+
6€+
vi+
gI—
1I—
SI—
Zo—
og—
e+
gI+
€r+
17+
1z—
Sr
gi+
ie
19+
“ojo
‘raAuoq
€z+
es-+
LZ—
b+
61—
g+
ae-t=
—
oc—
€S-+
6b—
o=
be+
zb—
ogo
‘1yeuUToUIZD
Z
vi—
IZ—
of—
9+
ee—
zI—
1I—
of-+
Ze+
be—
gis
1g€-+
zi+
Se+
6f—
MES:
‘uojso[ieyaD
9
2961-6561 uwounpidisacd jousou wosf Saanjangap pajaipa4q—E€l ATV],
‘%
92+
6z-+
ci+
S--
ee—
Cr+
Zi+
1z+
Ze—
ci+
oI—
ci—
oI—
er+
Ze+
II—
82—
cee
s
9+
z1+
Si—
O=
99+
61—
bi+
g9+
tz—
e+
€z—
c—
oI—
zi+
tp—
of—
tr—
I1+
zI+
gi+
61—
Sc—
c=
€e-+
zo+
ze—
29
‘ejsnony
v
UvoW YJUOW-INOY
c1—
61—
co—
Iz+
11+
ci+
£g-+-
e+
ZI—
e+
6z—
es+
1I—
6I—
Tt
61+
Sa:
of-+-
gh+
e+
Iv-+-
LZe—
ge—
"3310
‘Aueqiy
£
£+
Sie
ci—
‘A'N
‘Aueqry
z
or—
Ze+
ez—
ce+
gz+
Lp—
bi+
£+
ZI—
ie
Zo+
oz—
9
8e—
7S
ev+
61I—
ZI— °
cI—
SI—
"xOL,
eusTiqy
eevee
“AMUSMOUSMOSMOSAMOAMOAMOAMOAMYO
2961
9961
S961
to61
£961
zg61
1961
0961
6S61
59
ABBOT
FORECAST OF U. S. PRECIPITATION
NO. 9
gr+
£1—
oI—
Si+
ge-+
Zo+
gt+
—
er
==
gz+
z+
1s—
‘ey
‘£19UI0ZUO TT
gt
Vote
Iz+
11+
oz+
gz-+
6+
oO
z+
6g—
z+
ee+
of—
‘SIM
‘UOSIPeIT
SI
SS
ee
6=
oO
o=—
Sr
bh—
Iz—
19-++
os—
£+
ge+
ce—
6e—
£¢-+-
61+
1z—
gI—
1€+
gI—
Iz—
ye+
61+
I-P
or1+
Z—-
S—
Vv
‘yooy 2107
VI
ze+
oI+
cbr—
ci+
€1-++
ete
6I—
Z1+
Fi—
Ge+
£1—
1v—
ve—
€+
osS—
i=
oz+-
O=—
ce
Cr+
61—
9+
Z¢-+-
95—
“sue
‘gouepusdapuy
1
(panuruos)
9s+
z+
0z+-
fe+
f=
6=
ge+
lo+
y—
e+
ge+
1y—
get
gt+
zI+
bbh—
1¢+
cz—
zb—
*qUOW!W
‘eua[eH
zi
6s—
oo1+
1$+
£9+-
gt+
p—
89—
e+
o=
cr—
€1+
oy
lz—
zI—
oz-+
£1—
Zz+
or+
G2
€S+
of—
“xo
‘ostd IF
II
1g¢—
oO
61+
9i+
Z
or—
gi+
6z—
oI—
7+
£I1—
aUIeyy
‘yaodyseq
or
gr—
€€-++
ie
Scc—
of+-
6r—
oa
oe
II—
€z-+-
bi+
6z—
ze+
9+
z—
ce+
y—
bi— °
“Qo
‘yorqd
6
eee ee wens
£961
9961
S961
Fo61
£961
zgo1
1961
0961
6S61
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
gi+
s+
cf+
Pate
ge—
or-+
c1—
£+
vh+
1I—
vI—
61—
8+
¢1—
e+
1I—
6c—
1v—
of+
bi-+
6=
tz—
€1—
or+
ce+
Ze+
e—
‘O'N
*Aangsies
ve
gt—
vi—
1+
gli—
£9+
1Z+
zg+
61+
zs+
gs—
98+
69+
1f¢—
gt+
oi+
Soa
zi+
“FHED
‘oyUIWIEIIES
€z
€z-+-
zeo—
ce—
9+
zS-+-
61+
4I—
Pe
ger
I+
be+
==
¢z—
“A 'N
‘J9}SaOOY
ze
t1— lz+
61— of-+
gI— cr+
cc— y+
og— Se+
6+ gi—
9= Sin
41+ gi—
e— b+
Z— S—
gI— be+
br— 61+
oz+ of-+
Zze+ oI—
oI+ SI—
91+ ziI—
Z— I7—
€1+ 6+
ze+ II—
oO sI—
ci+ 09+
gI— b+
Ly— Z1I—
bi+ zI—
L— c—
zb— vi+
Ca of —
“SSI, IIL
‘UOSqID) 210g ‘el1oag
Iz
oz
pamyuor—El FIV L
ci+
SI—
9z+
Se
Si—
€1+
gy
Lz—
oz+-
le+
41+
cts
Se—
z+
6s—
Iz+
oI—
gi—
I+
p—
be+
17+
9+
6+
£+
Cc+
o1+
“Iq2N
‘eqeuo
61
61+
p—
e+
9+
cv—
11+
ce—
89—
6v-++
z+
t+
98+
y—
cr—
Ie+
9s—
ZQ—
Ly—
~—
Iz+
¢s—
12+
8+
ve—
ZE+-
gi
c—
‘ZuYy
‘asplig [einjen
gi
gr+ ee ceceeee
[a= cece
6z-++ oe
Si+ ereee
og+ °
ZI— °
c+ °
61 a
Iz+ °
II— °*°
pa ee
6+ eee
¢o— °
Dia =
oz-+ °
oI— *
get
1
Q— ttre
ip
ce. s°
p— crteeeeee
4f—°
‘uudL
‘ay[raqse N
“I
GMOUAMOMTMOSTMOSMOAMOAMOAKMOAAMDYO
£961
9961
S961
to61
£961
zg61
1961
0961
6S61
61
FORECAST OF U. S. PRECIPITATION—ABBOT
NO. 9
9+
ce+
ZI—
ozc—
oI—
61—
Ze-+
oc+
L+
zi+
gh+
be+
6+
ys—
==
gf —
17+
z+
II—
cco—
tc—
gz+
I—
ze--
viI—
cc—
Oa RTROe
ze
ge—
9+
ve-+
ch—
I--
ve
or—
oS-++-
lz+
o=—
vo+
Oo
oS
e+
3==
Se—
af
ce+
Lo—
ol
LE+
se—
ge
S++
9z—
o1+
e+
"ex
‘OT [Asemoy T,
1f
bi—
zi+
oI—
of—
8e—
gI+
Le—
SI—
Zy--
91+
be+-
ats
9I—
bi+
6z-++
I—
9+
12+
II—
Oca
If-+
Sz—
“UUIPL
“(neq 4S
of
gz-++
ge+
1I—
gf+-
ge—
gf+-
6f—
aoe
oI—
fe
bz—
Ly—
git+
9+
yo—
ge—
1z—
Ib—
Z1-+-
6b-+-
9+
11+
f+
z+
6—
Pafe
oI+
ow
‘sInoy 39
oz
f+
o—
oa
e+
8I—
11+
Zy+-
er—
Ze+
19+
S1+
6b-+-
cc+
fe—
SI—
o1-+-
gi-+-
9s—
£+
oe:
9e—
+
ov+
€z-+-
s9+
c1—
6b—
“GSeM
‘gueyods
gz
Z€+-
Ze+
SI—
e+
gt+
69—
be—
6S-+-
Sz—
gr
€z--
Sz—
Ze—
Z1+
61+
c1+
11+
£5-+
8z+-
cl—
of—
IZ~@—
1v—
(0)
o1+
se—
‘xOTW 'N
‘QT eyes
le
Tete
€z+
b+
f1—
rg—
Sr—
obv-+-
3
$Z+
II—
6e—
ge+
ZI—
€1+
zi+
ce+
of—
SI—
oI-+
gI—
61-+
99+
3i—
Z9—
Sc+
oz+-
oI—
“HHED
‘OuIpIeusiag UPS
ge
QI-+- eereeee
ve— °
of— °
6c+ °
we— °
——
tr-+-°
vi— °
te+ °
91— °°
ze-+
c— .
6z— °
ci+ °
SI— *
oz-+ °
e+ °
ce— °
ziI-+ oe
OC =
or— °
z+ °
e+ °
fi— °
Qorr
or-++
61— °
qe
‘aye eS
Sz
H“MOUSMUSMOKTMNOSAMOKMNOAMOAMOSAOYO
2961
9961
S961
vo61
£961
zgO1
1961
0961
6S61
992 “gi
Sp— eS1 Zor gi— 601
W-- og- Zer _11-— ei
o9+ og ofr €&— SZ
gI— cei VII 6c+ Fil
vit Sor 611 2S+ gor
gr-+ €6 rr SE of
Si— oir 26 gz— OO!
gz+ 2g o11 6— AZII
1z+ 99 68 €1-+ vol
ov— O6£1 66 II+ gl
Sc+ Zor zr Sb— 9gg1
Q— fil col —¥zZ——— Zo
ee+ ob €2Z I1z— IVI
f— 72 69 gy— gSI
cc— 6£ OF I— €Z
ge— cli vor gf+ OI
ce+ 2g Zor e+ 2S
Zz— ghi izI1 oz— 601
S— 16 98 6— 901
gz— 66 14 1v— vv1
II— obi 621 f+ 121
0 OOr oor fi— £1
vi— £1 ozi Sz— 16
Olsj- os slie Tél — si = 09
QI— ZOI 98 zi— ZV
1S+ gS 601 c+ SII
sss Cite eee 1s+ 624
yseo =paAtoS Vv
£0 “2107 -4O TO 40g
‘o]OD oro
‘19AUuIq ‘yeuUuloUlD
yseo = psAtas
16
¢S1
zZ
vi
og
Sor
vZ
gol
ZII
6fI
IzI
gel
ozl
OII
ZZ
gr
6L
68
46
€o1
vel
gzi
99
Col
SEI
OZI
Of!
-4O
oe g'0z
oi+ zZ ZQ z+ 01 gol
Iz— II £6 gs— Szi 69
€e-- C11 Soft. €— 101 96
cst+ 1g QZI gz— I€1t Sor
vi-+ gir gzi SiI-+ &4 88
of-+ 9S 98 cf€— 101 99
¢€— S6 z9 zi+ $6 Zo1
c—— 10 98 z+ 88 OIl
o-— 2g 8022 VI-- 59> 60
9+ 44 Coes ve Orie val
v~— QOI 2g 61+ g6 Lit
€ir+ 142 9 =%bv— v6 06
vL+ gf ZII 9og— Zor Iv
I1f— or 62 Zt Qel = gel
ZZ— QB zg Zz+ gv $2
6I— $6 94 zI1+ z2f F898
4i+ of & €— gor Sor
S+ gf Coe (g— Crre Ser
ee+ zy %8 =Qe— PS ge
t~e— zi 66 LZi— ZL Sa
of— If 111 gi+ Zor SzI
oz+ o£ 06 1z— Szi- v01
e— £6 1g ©=6bS+ 399 zz1
II— zg 1Z i= 40 £8
€1— oor 4g 6+ gf 48
lI— ZII “101 €1— 4g ZZ
gz— 3 ZS 19+ gir 621
Vv 4yseo peAtos V yse0) = pa Aas
£0 7301 GO, FO 70f -40
aus, "xd “N
uojsapieyy ‘yoreusig
£°9z
FA0
v+
€c+
14+
zI+
Fi—
¢s—
oI—
6z+
zI-+
£3—
Si+
1v+
Iz+
zI—
e+
—
6I1—
ge—
(Os
ge—
oI—
oI—
ce—
ce+-
zb+
Rene ie
96 ve
Sg 6g
SZ gd
zg est
bl 98
Sg 1Z
cet £9
69 6S
19 06
gs o£
ror ZZ
O41 6g
eS 89
ev tg
eg bg
96 «#8
$6 Z6
VII Ill
88 69
Zit 6
06 66
Zi1,- 16
gf 89
0g o£
ozt -50
ZZ Ill
eZ v9
yseo = pe Atos
2107 -4O
sonyeA ULOU YUOW-InN0F
0'9z
Tate
ceo—
gz+
er—
Zo+
z+
6f—
ti+
or
Date
ae
o—
vi—
of+
erp
tz—
cS--
6¢+-
ol—
p—
viI+
be+
ez—
1z+
Qo
ZI—
Ly+
S'1z LYve
£6 +6 S1+ 22 £8 gt-+
gor &Z gz— O11 72 zS+ g9
98 atte S-— -Olipeesile=s0-j-7 = OVT
ver, 10 vi— gor v6 os+ +6
1Q gor of— 6o1 64 fv+ 1g
601, el te ZV —= 7 ie OF fo+= 10
SzI 98 g— 306 06 o>. €S
vS So O6b— O6zl og 6&— 99
Cll] ghee Ii (Slim vel roc Met
Orr oSi 65+ 6g gbr 6z— 98
IZi O11 zez— 06 89 gi+ 66
vor g6 f— 66 96 19— SEI
igi) ZOE i4- 00 46 g9— eri
ovr ofr of-+ 12 rom s9Z— Tel
69 lit 6— O0ZI- tir 7— $9
cH 161i. I— ~~ Zo 16 oz+ O69
6S vir e€+ £2 OIL SI-+ gor
Sg ver _Lb-- cil OS 9E—
ver vS Gcc+ 2 66 of+ 72g
23 £Q 4+ £6 (NON ae (As
L9 1g z+ zor vol tI11I+ 09
Zol get OF-- cor> crt st— “Ir
gs se ZiI— «1 vir S€— OI!
gZ 46 I-- 9zte6en 1 SS—~ 211
1s1 oZ I— gor Zor zi— gS
111 6 1is— gir 9 zoI+ 49
LL OzI pe Col O1n--ei— 59
yseo psAtos yseo = pd AIss
2107 GO, YO -210f -4O 10 “210
‘3910 “ACN “XO
‘Aueqiy ‘Aueq iy ‘aualiqy
* §S61-0S61 “UuoyDasasgo pud 48DII40,J—VI AAV],
3yse9 Bete
IZI
ozI
OVI
eSt
tzl
VSI
69
€z1
09
gil
of
1Z
gt
SZ
29
ov
691
£Z
\o
in
O77
a
Sc61
vSO1
£61
zS61
1S6I
SMOOUIAMOGAMOSMOKTMOAMNOSAMOAADYD
°o
in
a
~
62
oof
¢f—
FI—
O+
oS—
48+
zI—
c=
oc+
e+
to—
oI—
Ze—
i
61+
£+
£+
=
ee—
og—
ze—
rey
9f+-
Fi+
gr—
og—
e1+
£0
cri
VII
S£I
oS
QI
VOI
26
9Z
gol
QII
£z1
ZS
of
ZS
vrI
cer
gol
£8
OgI
59
ZII
CLI
46
1S
yse0 peaires
3107
ElY
eZ
OOI
FOI
ell
ZE1
gol
ZIil
£8
FoI
62
9
c6
OI!
bZ
“gO
‘k1auI03{UO TL
ghz S°Z1 261 o6e vz
zOl 9g OO 96 96 SIi— za solv Sz— zor ZZ 4gt+ z1r 60oe St— 6+I
for 69 19+ 96 4S1 of+ gor gtr Si— Iz1 gor O6r+ Ger gfr £+ Of:
£6 gv fy— f€h1 oor ti2+ €&2 140) or-+ 69 601 9goI+ zoI goz gf+ 96
611 66 1+ “Zy1 ght “Zze— iz 6 Ot zeir gir Zor+ 9S for Se&— €c1
Ir of: gS+ €&Z rer of oer - Zz -QP3-" 99 gir Z4I— 111 6 62— E21
69 09 €f+ C11 obvi Sbh+ 99 11r 6+ = g6 Zor <L1-- SZ zo oI— S6
eo 92 - 0r— 6 “Fg Of— Por . 99 is— ei 19 — g9 46 Le— ol
Sor ocr <¢€-- o£ gor €— 09 ZS 6+ 2% 16 €1— 601 96 Qz— Of£1
140) 16 I— ger ocr S-- of tw Iz+ & 140) ZI— €zI gor €— 611
zg vS vo— ofr ZZ QI— gol <6 zy— of1 6 yS— zor gp z+ 98
Ger eg Gre: = 0 for gi— €or Sg 2+ 126-— vor s€t— 2or- 6g -0=—-_ "a8
96 z6 gg— gr Ss6 Sz-+ 9S £g oO +g tg oS— I11 19 of-+ 92
6£ oor S+ 92 £g fze— gel 101 O zg zg v— 4 o£ 1f-+- Sor
C21 --abl <1 2B iS- tie 9 02 ve+ 06 tir “2— oor E21 65—- Sie
Ig 06 6I— gir £6 1€+ of 19 zi+ SQ LL 1Z— ZE1 99 ee+ C11
1S) 09's 205-9 7o 0 Ohh )S 6tis«é'FE te+ z& oS vE-+- SI 6b oz+ 66
gd Z 1f+ 62 09 f— Z24 bZ c— 94 1Z £Z— I91 g8 se— 6S1
2. 6gtr 6o— = Shr— +s‘ EL sa _ 4B 98 6+ 38 46 of— of: oor zI+ SEI
£9 Lg b— Sor 101 th— £6 or tze— 66 SZ oI+ gI gz ze— Ofl
€or S€r o1-i- oS 99 z+ ov cy zZ— 99 99 1+ thr SZr oO OzI
wii for v-— Vor gor g--’ 69 LZ 91+ 1Ol 99+ 961 zoz SE+ 86
fer Str Si-- Zexr vir Z— «walt wer 4o— or 26 9— 45 1s t--- ZS1
6g 96 gz+ 26 OCI a+- wir Gir y= ci git €2— Ger ZO vi-+ ze1
yor If: o&f— 06 o£ fZz-+ 6S 98 of+ gS 88 o+ 6€1 Svi zb+ oor
€6.- 109 _Oe— Ofr ull fe— 6S gt gI— Of1 i211 61I— gf ZS 6+ 9gf1
vor “vr OI-+ ZII ez1 oz-+ ozI ort vI— zII 96 61— 96 LL fe— €o1
aie (OA te 10) vSr Sr gi+ gS gZ II-+ 0g 16 ey 1g LV 4 Sz1
3889 = peATOS AV #820 = psAlOS Vv ys®2 9 =psAtos oe yseo =psAtos #889) =psAIOS 3Sse5
2107 CO FO 710 GO 4-0 710f -GO 2101 GO ‘o 2107 GO XO 7210,
“STM “yIV ‘suey “yuo xOL sUuIeyy
‘aosIpey “yoo 21331] ‘aouapusdapuy ‘euajaH foe Iq Giodjseq
£02
gI—
Iz+
II—
gt+
oz-+-
ze+
6—
oz+
L-
Ze+
£1—
ep+-
c—
(panutjuor)
VII
£e1
PEI
QII
+6
£6
vor
gII
88
QZ
gol
gel
gst
Ser
6II
vzI
LY1
g6
OzI
£f1
gSI
grI
evi
Lyi
SzI
gor
PpeAres
“40
Loe
6— 66
v=. 108
os— 66
1v-+ oI
Iz+ ool
S-=08
ef<— Ozl
I+ zo1
fr+ gf
9— MFII
Iv— Vol
Z— 46
61+ I01
e+ GZ
IlI+ 8
gt— 4g
6I— gol
f£— 16
of— zzl
gZ— SI
1+
ofZ+ 22
gt+ 99
Sz7+ £8
Q- Sir
ZE— 621
zs ee VII
3se0
10
-210.5
STON
*yror}9q
67
SvI
IzI
88
£o1
gil
gol
£9
Lg
ozI
gZ
OIL
6
6g
88
88
94
gII
LVI
VOI
ZIl
€z1
ZO
gor
ie
ZS61
gS61
SCOr
a
wW
OV
=
MOLMOAMOATMOAMOGAMOAMOAADYO
i zs
S 2
V oS61
1Se P'gt F'61 961 v-LZz Q°9z Sze Lie uray
vo— FFI 08 v6— bhi of QI— OVI vel O6z— E1I 2g II— 1g o4 Le— ~ 63 zg ty— Szr zg e— 21 624 9
1+ S6 o11 11+ 6V €or e+ 96 €21- 46+ €2. of1 I1b+ 2c fre £5 2 Stir c+ ozr ‘ver E+ 98 tote J
ge of bir Si-+--2fr Zgr 8+ Sg oft g&— Ger 16 of— gzI 9S Sc— ter 66 v— 11 O11 6— £6 V gs61
es+ 06 €r1 Si+ 14 98 fze— £6 oZ z—- of1 blr Sze+ £6 gir Se O11 II Ol—' gir gor iI¢e-- Ick gyi 9
z+ 601 III oOz— &@g So s— 96 16 of+ Z9 46 g99— SSr 68 vE+ 26 QOZI Q+ O21 Ser IV+ og fer gf
c= ein Gir ci— gyr O11 c= 19 £g 6I— SzI gol tvz— zzI- 986 Zy— obi £6 s— LZ eZ f— gzr fer vy 4S6r
C— Aer at SS—) OY. I0I OI— 96 88 €{1— €11 oor I+ 49 89 f+ +9 23 vE— Qf a zI— 96 +g 9
ce+ 94 for 1+ 15 v9 1~— ofr Sir oz+ v5 vZ Zo+ S 16 gz+ 9 zo cS— Zor 2s Z— g9 19 q
z+ 09 26 oyV— oor vin vV+ Zit ier ve— er gor iIb— 66 gr 6s— 96 Le Onre “Sh ES fi- 11 ver vy 9861
0 Qe 128 te eit Sit g--= 21 O81 ZE— 601 ZZ s— 19g QZ gz— of a I— of 69 o&— Ser $6 4)
SI— gor 16 GS-2 99 ZI— 06 eZ f— o1 Zor o+ tz 0g eee 00) 16 6+ v&r €Z1 g— v6 98 q
f+ 72 & ge— If: £6 6— oo1 16 €+ 0g £g 61+ £6 vit te— Sr 6 gI— 62 19 oc— O6br fEzI vy SS61
0 Son Sen Of — onl 8 zO— OgI gir 6— Zor g6 v~+ 16 Sin, -ov— Of1 7g csf¢— vor 69 vVe— SIt 16 9
fy+ Ib v3 Ze— ZV oz Si— $6 0g 6— 11 vor Sb- oor Shr ZS— ZyI 00 ooI-+ 9S OS Sig— 59 ¥9 gq
g— Zor 66 ve+ 6 Sit. I-- £0 140) Of -O8 98 w+ O62 Ii. Z+- Sin ofl Q=—} Git «wr S-—==Zor gor -y vér
1s— zi Qf gz— o£ rAd gi+ SZ £6 Ti-- S&S 99 9o— 9S zs L+ oz 96 to— 69 s¢ op+ ¢ SV 9
Se-- oF 18g I9+ € v9 Qc— 9QZI OOI1 O vor vor z— gzr 96 o=— 19 gs os— vor 5 6v-+ ov S6 q
oz+ £2 66 1v— gf1 £6 e+ ZL 68 €1+ Szri gi o1+ og 06 cI— zir £6 6b>— 601 09 Se— Thr Cor y-eson
fz— Col 0g gf— gII 0g zi— +6 zQ z— 6 09 of-+ ¢S £g z+ cor vor gz— orl gII i1z2— vor & 2
gt— vir gf 611+ vor £ze i1€— Zor gf c+ 6P PS z+ £3 Cor 6€+ 624 QIlI ov— 11 IZ 6— 2@Z £9 q
Tie) Zot. OVI. “Ve— Pol ofl O0f-- VZ yor + 19g tg z+ 66 IOI Fi— off zzr ve+ gir zr “Z+ ozt “et v esr
6) Gt -- £9 ca-\- Zon. O55 t= Scr Zor S— § 16 98 €z— gQI1 £6 zz— gor +g 4ge+ zor 6g ge+ er ovr 9
I= 92 SZ ve+ O1 ve Lea 09 for €S+ v9 Zit Z-+= 301 901 ge Zar SSr g&— Ser Zo Qo¢— 9QZI OO gq
of— Fir gf es— “411 vo re+ 6 gzI of— 6f1 G6or oe— Chr Zr b+ 181 VOI O6f+ 2 Sor 6— 11 Sor vw IS61
6z— for vZ eo-— yr Soz Qr— off cer of&— gzr 26 of— €€1 £9 Vice gs ZZ g+ 02 gz Gis Ob Zen. -9%
11+ 64 06 6z— 68 09 6+ £6 Q90OI Oc+ Izt Iwi <Sf€+ IZ gor Sz+ +6 {11 6— GZ 99 zS+ 96 gyI qg
o+ 9 04 Zo— gSI_ 16 GE-— Or OVE c— rr 621 oz— Zr Si z— £6 S6 fy— gor ‘9 i+ ior z&r VY oS6r
Vv ysed poeAtos Vv 3St9 8 psAlos Vv #889) =—peAtOS V yse@9 = peAtos Vv 3St2 posAtos V 3889 =psAJOS V yse2 = =peAtos Vv sed) =pdAtos
w@ 2107 -{O H-O 40" qO 4-O 407 -qO ZO 404 -qO CEO) ERG ESO GO) AOE Seto = Ge) EehOlGt IO) iO 404 GO
‘ON “FeO ‘AN “SST “HI "392N “ZIV ‘uuay,
‘AINQstjeg ‘OJUSUEINES *19}89q00Y ‘uosqiyy 10g *e110ag ‘eyeuo ‘asplig [einjen ‘OT[IAGSe Ny
panuyuoI—VI ATV L
64
491 LSz 6'0£ Oz 6°61 6°62 Oey £1e uroyy
gS— of1 zZ zo— 611 ZS Iz— 62 98 I— +3 €g o Gir Gir S— O11 «ir gz+ hr Zor 96— dZzI Ie 5
f— eer 611 vze-+ £06 6II 9z— zOo 99 ov+ 9 Scr “Z+ SOI ZzII 9OvV— gzI zg gg+ 6 gr = £9— 19 QI a
oS-- ror mr S— om Sor ‘SS— 6 oe vi— z6 94 of+ 6 oft 49+ gP Ger or-- Shr — 691 ~~ SP-F 22 Zit We -QSOr
6+ g01 411 fg+ SYP gtr. r—. Ger 22 z— QII 96 oi— Zor £6 €/+ 101 vr 6€+ or SS1 €1— oor £4 9d
lz— 26 S9 ob+ 14 411 o1+ 01 Zir 6S+ Zor og91 18+ 09 Ill e+ 6 96 cs+ 66 Sr bZ+ az gvI qd
zz— 601 24g Iz— £6 cL II— GZ 99 ecs+ 62 zeI I— ZII oor 46+ EP oFI zI— g6 98 I— vir ¢1r vy S61
£— 101 96 I+ zg £g gi— 06 bZ z+ 939 o£ 99— zSI 98 sS— 06 ce zS— £6 Iv gI— +6 gZ a)
II— zoI 16 Cl-je Olen ZEt Ob ZZ €1I 62-+ og 6or gi+ Zg Sol oI— 62 £9 oo+ 621 S6r S+ O67 vs q
vVE— O11 Sg Ol-—sObine Ll Se 20e 20) O---- 00m = eZ I— 6g 9g 6— 09 Te SV— =cel 92h VV —= mat Ig V gS61
gt S6 for gz— S6 £9 Z— 16 tg Z— 8 98 Z+ 6Sr oor €2-+- oF £69 z~— O01 gf 6€— Shi 901 9
cI—— oft, Zor 2—— 66 z6 v~z— Lor & of— vzi v6 6— ozri i111 “Zy— gzI Ig 62+ £6 ZI ge-+ 0g gol gq
€z+- 09 £8 g9— or gf SI— 4g ZZ 17~— ZOI 938 e+ = 00 z6 ri— 69 SZ 4+ o£ LL fo— Ler VL Vv $s6r
f€— €11 og tr+ LL vS— Ser 12 OV— AS as0l =“veE— Cer 266 fe-- 6f ZZ gli+ f61 SZ ee+ £3 o7I gO
poe Oa 10 I— +S es fZ~— Ser 36 OI— cz zQ co— Sva “Eor ~Ze4> yor hi, 02-3 9c 36 Cy eee te OO Ta
6+ 99 44 oS+ 61 69 1v+ 69 OI -OI-= “tf 6 gz+ 39 +6 Z£+ +9 16 IlI— tee €11 oI— ZZ ZO Vv vs61
61— gor Zg fy+ eZ SII go+ €1 Ig sf— 98 oa) gz— II 16 SI+ 98 fol 7zg— gel WwW cy+ 11 os J
61+ £9 zor 11+ ver E€zr gS+ So cz1 6+ ZF ov y+ 06 ¢6 g— oo1 2z6 of+ 9 of So+ Iv vor q
b+ zi Sz Sc— tir 601 iIt+ ez €11 €2+ 9 06 Gci+ ver O6f1 tie+ “4 89 br— “Zir €Z c— £6 88 Vv esr
Eca-sell sel Gt 49 6) Ge— tg 9S €€— -16 95 of— gtr 99 ZE— 09 Gb s2e- Sti “ely “25-—— 00n ses S
6— 621 ozr 2+ £01 o11 9— €or yS— gzi vZ l— 64 ZZ 1g<— 66 89 €I— Sor zo 1v-+ 99 601 gq
Ps Obie CEI Sc Ii Orr e6-- 2) — 61 gi-- 00, fg. sri-b 2 Sg a—= fo. = fo" “Or S0el, 1Oll Sgt PeZge yh vecwon
MieeelOte Vile = set <0 Olle <Gc-= 1607 ~~ 0¢r= Vj EO — 66 Olt s0cl eat) 2 Ol ages va G-a=- 2b Test “OI 2Vi- er 5)
1 Vee iilg Iter LOL) 00 oz- “Zor Zeit oe— Sit SO zet= oF =90' =t— 202, =e4° SSc-- sy0n 300) —Wes-p een ir 6g
fo) v6 +6 SZ— 991 16 e+ 16 1 Oo O01 oor off of oot g— €S “Sp of ZE Zo 61-- 00° Gf Y I5G1
ozc+ OIL Of€1 Szc— 8 z9 Z— 16 tg c+ GZ 03 oI+ vir tz1 SE— oS IZ eh— i101 66 f€— ocr £6 “)
GI— £1 OQII 11+ 9 gZ fv— gir SZ f@— QzI 101 i1I— oOzi 601 OI— Sor 68 cc— 62 bz z— 606 £6 q
Za- 69 0% f<— 68 98 SI— III 06 OV-F OIF “OSI S€-> Pit (67r “PZ—FZer eS gf— I01 £9 z+ 99 98 V os61
Vv yseo peAtes V ysto pears Vv yseo pdaiss V }sB0 pdAtos Vv yseo = paatas V yst0 pdaAtas Vv yseo = paAias v qsvo paAtos
oa =o “Ur ‘OW “yseM "XO “N “FeO qn
‘wo}SUIYSe MM ‘ay]IASeuoy ‘neg 38 ‘sIno'y "3S ‘gueyods ‘aq eyUeSG ‘oUIpPIvuIog Ueg ‘ayey ues
65
66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
TABLE 15.—Expected numbers of discrepancies of forecasts between assigned
limits
Numbers expected of 4-month intervals in 9 years, 1959-1967, when
(O-F) has certain values
Group 1. Mean (O-F)=20 percent
<5 6-10 10-20 20-30 30-40 > 40
Bismarck As ota eta. 6 3 8 5 I 4
Charleston join! see eb 4 oO 10 8 I 4
CCITICHATTA CTY chee clap als ais a's 4 4 7 3 4 5
Independence .......... 4 5 4 8 2 4
Madison acisjtaccnkicrse tvs 3 5 8 4 4 3
INa‘Shivilleteemcr crete 2 z, 2 8 5 3
PortGibsonuac cece 8 2 6 5 4 2
Rochester y auc aisle sietiele's 5 5 5 7 3 2
SMOkane oes selciss aheeaie 3 6 7 6 2 3
Spr Eos ie eid ae ovelete 6 4 3 6 4 4
Washington! 2.330.056 6 5 8 4 2 2
Group 2. Mean (O-F)=26 percent
<6 7-13 14-26 27-40 41-52 > 52
Albany? Oreg.) 4.6 ei sa 4 5 7 5 4 2
POUISE A ele aches mais I 7 9 6 I 3
Denver. oh aah eae 3 2 10 7 3 2
METEGlemROGI jaysrreecd rears 5 6 4 6 I 5
Peoria. ss Heidislesusdnses 4 3 6 7 4 3
SAUISEUIBY tera larns c eers wi bie 5 5 B 8 3 3
Group 3. Mean (O-F)=30 percent
<7 8-15 16-31 32-46 47-62 >62
DEtROItE cis ae eit cteke wove 5 5 4 8 2 3
Natural Bridge, Ariz.... 6 4 4 9 2 2
Salt Wake os cja.0 sictesore's ss 6 2 4 6 4 5
Santan Mel sivas. cs iiets astonsis 5 5 7 4 2 4
Sti (Paul iijccrcsae,crrsinarevats 3 5 8 6 4 I
Group 4. Mean (O-F)= 40 percent
<10 II-20 21-40 41-60 61-80 > 80
FGINBASO eecorebetas uieisvaeieies 3 5 5 3 7 4
SacramentOn iescsiicts oO 4 II 6 2 3
San Bernardino ........ 3 2 9 7 2 4
Sumsvohiesuenneaces 104 106 159 152 74 80
MEATS iehsjcleersteie eres 4.2 4.2 6.4 6.1 3.0 3.2
amitsyeeieeseree ce <i AS 4-1 1-3/2 3/2-2 2
NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 67
their magnitudes would be greater in proportion as the mean de-
parture of his place bears to 20 percent.
As actual cases, farmers living near Albany, Oreg., or Augusta, Ga.,
both by table 14 lying in the 26-percent class of table 15, may expect,
according to table 15, during the 9 years 1959-1967, the numbers of
4-month averages found in table 14 to differ as follows from the 27
mean 4-month departures from normal precipitation they will actually
experience: Four cases less than 7 percent; four cases between 7 and
13 percent; six cases between 14 and 26 percent; six cases between 26
and 39 percent ; three cases between 39 and 52 percent; and four cases
over 52 percent. Farmers living near one of the cities of the 20-per-
cent class might expect this same division of the 27 cases for the
4-month mean departures from normal precipitation, but these de-
partures would be smaller in percentages in the ratio se It will be
2
for their judgment to dictate whether it is worth while to procure
from the Smithsonian Institution, and make use of this paper, “A
Long-range Forecast of U. S. Precipitation.”
COUNTRY-WIDE TRENDS IN PRECIPITATION
The maps of the United States presented below show large areas
over which similar forecasts prevail. This should be helpful to in-
terested persons who reside at a distance from the 32 cities for which
forecasts were made.
I have been interested to search further to see if similar trends of
precipitation sometimes prevail over the whole United States. Table
14 gives the actual departures of 3-month consecutive means of pre-
cipitation as averaged over three 4-month intervals per year, 1950-
1958. A working table of these results was prepared, giving the 32
departures from normal of the cities employed in each line of a table
of 27 lines, 3 lines per year for 9 years. Recording separately plus
and minus departures, sums were taken for each line. These plus and
minus departure-sums were plotted in figure 12, lower two curves.
Plus sums are given in full lines, minus sums in dotted lines.
The plus and minus departure curves run generally in opposite
directions, and in some 4-month intervals are widely separated. In
such cases of wide separation the 4-month intervals were strongly
heavy in precipitation if the high points are on full lines, and strongly
drought-prevailing if dotted. With this explanation it is seen that the
autumn of 1951 and winter of 1952 were wet periods generally for
the whole United States, and similarly from the summer of 1957
68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
NO. Q FORECAST OF U. S. PRECIPITATION——ABBOT 69
through the summer of 1958. On the other hand from the summer
of 1952 through the autumn of 1956 the country was generally dry.
This interpretation of generality over the country is justified by
the fact that the high points of figure 12 depend on observations of
identity of signs for more than 20 out of 32 cities, in 15 cases. Some
peaks are supported by 28 cities out of 32.
When both curves are near the heavy horizontal line the precipita-
tion of the country as a whole was nearly normal. That is, through
1950 and the first four months of 1951, and for portions of the years
1953, 1954, and 1957 precipitation generally averaged nearly normal.
The curves of figure 12 show plainly that the entire country is subject
to nearly simultaneous trends of precipitation, depending, as they do,
on nearly universal agreement of observations of departures in 32
cities over an interval of 9 years.
With this result established, turn to the two upper curves on fig-
ure 12. These are plotted similarly to those below, but are from
table 13 which gives the 4-month mean departures from normal pre-
cipitation forecasted 1959-1967.
Reading these upper curves: After the dry winter of 1959 there
should follow a short well-watered interval, and an interval of nearly
normal precipitation before a rather well-watered period in 1960.
Then, following normal precipitation in 1961, should come pretty dry
conditions in the winter and early summer of 1962. A long period of
normal rainfall follows from the autumn of 1962 through the summer
and autumn of 1964. A very wet winter of 1965 follows, and fairly
normal precipitation thereafter, except for the dry summer of 1966.
The last preceding paragraph concerns the country as a whole.
For details of forecasts for individual stations, the predictions may
be found in tables 12 and 13, and in the 27 maps of the United States.
MAPS
Twenty-seven maps of the United States follow, with circles show-
ing location of 32 cities. Numbers above the circles refer to the cities
given in table 13, which are numbered correspondingly. Numbers
below the circles give percentage departures from normal precipitation
as forecasted as means for 4-month intervals in table 13, 1959-1967,
A, B, and C, for each year. Three maps form one chart. The nine
charts are dated from 1959 to 1967.
7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
\ 41 a9 2 at
NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 71
72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
s
=36
@
+8 +38
+.
noo
NO. Q FORECAST OF U. S. PRECIPITATION—-ABBOT VS
28
-56
z 12 <6
+7 30 &
-15
30 22 3
O °
° 7 45
iS
ro 2)
= i.
23 sri aes g 20 @
a * a +2!
~80 2 9 -19
13 Al
Ps, “50 17 24
fe) Oo
ay
os 8. i i -16
© aR.
74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
( y g |
es “28 eA}
23 a IZ 20
433 ° -59 =I qi B
aul 29 453
12 64
Pa —4l 24
17
7 %6-——__12 3 we 3
+12 a Se 4
\ 4 %
ie +2
W 2) ° 1
= << 47 #10 %
!
°
Se
26 é ne
-33
2 2 a
oO
~19 +15 Su Pr
°
+21
Os aN (
9 ee 3
-¢ 3 S
: i E
15 9 an nae
5 +20 ¢
23 a
° is ee 209
-88 8 +34 -Ie 32
“16
28 g 27
+19 45
a“ !
\{ Re “
23
fe}
+25
3 ig
“ul -39 2
+17
° Z
28
23 19
-29 2o
2 is 34 Pe ere
o
9 3g 423
T 29 -19
2 +18
729 24
26 18 ay 5 Y
e 3 4
1 Sy I
-35 3
uw ie ue
~69 -I7 at 3
oO -
+2 as 38
1963
NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 75
76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
FORECAST OF U. S. PRECIPITATION—ABBOT Th
NO. 9
78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
: VOLUME 139, NUMBER 10
: WATER TRANSPARENCY
OBSERVATIONS ALONG THE EAST
COAST OF NORTH AMERICA
(WitH 2 PLaTEs)
By
JEROME WILLIAMS
E, R. FENIMORE JOHNSON
AND
ALBERT C. DYER
FREE reg eae NO FET ee eee
{ 1 1 ss Z
oe
(Pustication 4391)
RS ae rey
= a re
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 26, 1960”
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 10
WATER TRANSPARENCY
OBSERVATIONS ALONG THE EAST
COAST OF NORTH AMERICA
(WitTH 2 PLATES)
By
JEROME WILLIAMS
E. R. FENIMORE JOHNSON
AND
ALBERT C. DYER
(Pusiication 4391)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 26, 1960
THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.
WATER TRANSPARENCY OBSERVATIONS
ALGNG THE EAST: COAST (OR
NORTH AMERICA
By JEROME WILLIAMS, E. R. FENIMORE JOHNSON AND
ALBERT €. DYER!
(WITH 2 PLATES)
INTRODUCTION
Marine biologists have long been interested in the transparency of
natural waters as an important parameter in the determination of
both the amount and type of plant life at various depths. Owing to
this interest, many transparency surveys, in the oceans [4, 8, 16, 19,
22, 23],? in lakes [7, 33], and on pure water [27], have been made.
In recent years, however, this interest in water clarity has spread to
other fields, such as underwater photography (1, 9, 20, 24, 33) and
television. In addition, there is a growing movement among workers
in the field to utilize transparency as a “tag” for water masses in the
study of such things as circulation patterns [23, 25].
During the years 1947-51 the yacht Elsie Fenimore made a rather
extensive survey of water transparency conditions along the east coast
of North America from Labrador to the Gulf of Mexico, including
some stations around Newfoundland and the British West Indies.
Even though the data herein presented are admittedly far from com-
plete and a number of other studies have been made of the area [3,
5, 10, II, 12, 13, 14, 15, 17, 18, 21, 28, 31] this study represents, from
a geographical standpoint, the most extensive single piece of work
done on the subject to date. For this reason, if for no other, it seems
desirable to publish this information in the present form so that it
may become available.
To make the data as universal as possible the unit chosen was the
so-called Equivalent Secchi Disc Reading. Since it is obviously im-
possible to use the Secchi Disc [32] for measurement of water trans-
parency if the water mass to be measured is at a great depth, this
water mass is hypothetically brought to the surface for measurement.
Thus the Equivalent Secchi Disc Reading may be said to be the dis-
1 Mr. Williams is associated with the Chesapeake Bay Institute; Mr. Johnson
is a research associate in the Limnology Department, Academy of Natural
Sciences of Philadelphia; and Mr. Dyer is connected with the Fenjohn Company.
2 Numbers in brackets indicate references in the bibliography.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 10
EEO EO
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
tance at which a Secchi Disc would just disappear if it were immersed
in water and if that water were at the surface.
As an example, if an Equivalent Secchi Disc Reading were given
as 10 feet for water at a depth of 100 feet, this would mean that if
the water mass at a depth of 100 feet were brought to the surface a
Secchi Disc would disappear from view at a distance of 10 feet in
this transposed volume of water.
The Secchi Disc is admittedly a crude indicator of water trans-
parency, since it was originally used by marine biologists to measure
the so-called extinction coefficient. This is a measure of the amount
of light reaching a horizontal surface at some depth. Unfortunately,
the extinction coefficient is not only a measure of the water transpar-
ency but also a function of such things as sea state, cloud cover,
altitude of sun, and other factors. Even so, however, the Secchi Disc
reading is probably a reasonably good indicator of water clarity if
it is taken with the sun fairly high in the sky and if it is viewed
through a glass-bottom viewer or hydroscope [30].
In addition, the Secchi Disc reading is an easily understood unit,
generating an intuitive feeling for the existing conditions, so that
it has become fairly universal in its use as an indicator of water
transparency.
Of course, the actual Secchi Disc reading gives an average value
of the transparency of the surface layers, so that if a layer of mark-
edly different water exists somewhere from top to bottom, it will not
be seen. For this reason, other instruments which measure trans-
parency of relatively small volumes of water were used in conjunction
with the disc. These will be discussed in a later section.
The writers wish to express their appreciation to Dr. Ruth Patrick,
Curator, and Miss Margaret Le Mesurier, Librarian, of the Depart-
ment of Limnology, Academy of Natural Sciences of Philadelphia, for
their indispensable aid in the preparation of this manuscript. Ap-
preciation is also expressed to the Smithsonian Institution for material
aid and advice in this project and publication of the paper, and to the
Academy of Natural Sciences of Philadelphia for its contribution of
personnel and materials in the carrying out of this program. We
regret that space does not permit the listing of over 50 other persons
and institutions to whom we are indebted for advice and assistance
rendered.
INSTRUMENTS
The instruments utilized in the accumulation of the data presented
herein can roughly be divided into two classes: (1) those that meas-
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER 3
ure the medium in its natural environment and (2) laboratory-type
instruments in which a water sample is removed from the medium
and examined in the shipboard laboratory. The first type is usually
considered the more reliable when dealing with natural waters, since
the transparency properties seem to change rather markedly when a
sample is taken out of its natural environment, and therefore this type
is discussed first.
I. IN SITU INSTRUMENTS
A. Secchi Disc ($l. 1, fig. 4)
The Secchi Disc, owing to its ruggedness and ease of use, was the
most often used of any of the devices to be listed. The disc used
was 74 inches in diameter and was painted a flat white, having a
reflectance coefficient of about 0.8. It was obtained from the Oceano-
eraphic Institution at Woods Hole, Mass. A specially designed hydro-
scope (pl. 1, fig. 3) was occasionally used in conjunction with the
Secchi Disc to eliminate water-surface effects. Generally the Secchi
Disc was observed by means of a glass-bottom bucket. Readings were
made from the sunny side of the ship, except where otherwise noted
in the data tables, and the recorded value is the distance from the
bottom of the hydroscope to the disc, i.e., the distance traveled by the
reflected light from the disc surface through the medium in which it
is suspended.
B. Point Source Light
On a number of occasions the transparency of water was measured
by observing the distance at which a point source of light can be seen.
This method of measurement may be seen to be similar to that of the
Secchi Disc.
Although a true point source of light is well-nigh physically impos-
sible, the tungsten filament of a 1,000-watt diver’s lamp approximated
this well enough for the range of transparency encountered in the
near coastal and inland waters. It unfortunately fails badly in the
ultraclear sections of the open ocean, where it diminishes in size and
eludes the observer before reaching extinction through absorption.
In turbid waters the point source shows up as an incandescent spot
surrounded by scattered light having the appearance of luminescence
in which the visual range is the point at which it disappears into the
background of scattered light. In clearer water, on the other hand,
the background of scattered light, if it can be seen at all, is seen only
when the point source is close to the observer and disappears while
4. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
the incandescent spot is still plainly visible. The energy from this
spot is so reduced by attenuation that the structural shape of the
filament can be clearly seen. The visual range is then taken to be the
distance to that point at which the filament completely disappears.
Most of these observations were made horizontally with the lamp
and the objective of the hydroscope both placed 5 feet below the
water surface. For the sake of completeness, observations were made
both during the day and at night. Plate 1, figure 1, shows the point
source of light being observed through the hydroscope.
C. Illuminated Letter
This observation method involved the use of a low-powered lamp
enclosed in a small housing with an opal glass window, in front of
which was mounted a rotatable disc which had a series of cutout let-
ters. The whole rig was mounted on a pole which could be extended
approximately 5 feet below the surface and was observed by means of
the hydroscope. The procedure adopted consisted of bringing the illu-
minated letter toward the hydroscope in a horizontal direction until
the observer could make a positive identification of the nature of the
letter.
D. Underwater Objects
To obtain some idea of the horizontal visibility available at various
stations, black and white balls approximately 6 inches in diameter
were lowered about 5 feet below the surface of the water and observed
with the hydroscope. The horizontal distance at which the balls dis-
appeared from view was recorded.
E. U.S. Navy Hydrophotometer Mk. II (pl. 1, fig. 2)
To obtain a measure of the variation in transparency with depth,
standard U.S. Navy hydrophotometers were used quite extensively.
They consisted of two principal parts; a control box and an under-
water unit connected by an electrical cable. The underwater unit may
be lowered to any desired depth and the transparency at that depth
is indicated at the control box. It is very similar in its operation to
a number of earlier instruments [6, 29, 33].
The underwater unit consists of two heads separated by a fixed
distance of 0.5 meter, one head containing a photocell, P;, and the
other containing a collimated light source and another photocell, P»
which is connected so that its output is in opposition to the output of
cell P;. In operation the light shines both on P, and P, and the com-
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER 5
bined output of the two cells is adjusted by means of light irises so that
the meter in the control box reads 100 percent when the underwater
unit is in air (air is assumed to be a nonattenuating medium). Then,
as an attenuating medium such as water is placed between the light
and photocell P,, the meter will read some fraction of 100 percent.
Actually, since there is a light loss of about 4 percent per glass-air
interface owing to the different indices of refraction of glass and air
which does not occur when the device is submerged because of the
similarity of glass and water indices of refraction, the reading in air
should be set to 92 percent instead of 100 percent [34].
There is a definite temperature effect on the device, but in view of
the sources of error existent in the other methods of measurement
and the length of time required for an internal temperature change
to occur, it is felt that this temperature dependence is negligible. This
temperature effect is reported in the National Bureau of Standards
Text No. 43P-1/47.
F. Hydroscope
This instrument is essentially an underwater telescope having a 15°
field of view with interchangeable heads for either vertical or hori-
zontal viewing for Secchi Disc or other visibility range readings.
Plate 1, figure 3, shows the device which is approximately 15 feet
long and uses a lens system of unit magnification. The viewing head
is equipped with a focusing eyepiece, a rubber face pad to exclude
external light, and two positioning control handles.
In use, the hydroscope is supported in a ball-and-socket mount on
a platform extending from the side of the ship, with the objective
head of the instrument extending 5 feet below the water surface.
II. LABORATORY TYPE INSTRUMENTS
A. Peraquameter (pl. 2, fig. 1)
This device is very similar in principle to the illuminated letter
described above, except that the letter to be identified is placed in a
long tube (11 feet long) which is filled with the water of interest by
means of a pump. The observer looks into this tube and is able to
move the image of the letter, by means of a movable mirror, until
positive identification is possible.
The peraquameter was used when visual range, using the illumi-
nated letter, was found to be under 22 feet.
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
B. Scattering Meter (fl. 2, fig. 2)
To measure light scattering due to suspended particles in natural
waters, Dyer developed a device which essentially consisted of a light
source that sent a beam of light through the sample. At right angles
to the beam, a photocell was placed, and the amount of scattering was
then a function of the output of this photocell.
The sample cell used was first a 24” x 25” x 1” rectangular glass
container, but this was later changed to a 3” x 3” x 2” plastic cell to
handle a larger sample and at the same time defeat the problem of
condensation on the outside of the cell due to cold-water samples.
The electrical circuit was so designed that the output current of the
photronic tube affected the grid current of an amplifier tube, thus
causing changes in the plate current of the amplifier for small changes
in the output of the photocell. A microammeter with scale ranging
from o to 100 was selected as an indicator of the degree of scattering
and was connected in the plate circuit of the amplifier. The circuit
was adjusted so that the output current could be zeroed for any given
beam intensity with the sample cell empty. For operating con-
venience, a reflecting rod was so mounted that it could be swung into
a fixed position in the light beam in order that a check could be main-
tained on the source light output by means of its effect on the output
of the photronic cell. The entire unit, including batteries, was mounted
in a glass-fronted metal case for convenience.
As finally evolved, the device proved capable of covering the entire
range of turbidity from Delaware River water to the finest obtainable
grade of triple-distilled pharmaceutical water.
METHODS OF DATA ANALYSIS
For the sake of uniformity it seemed desirable to convert all the
hydrophotometer readings to “Equivalent Secchi Disc Readings,” as
defined in a previous section. To do this required some relationship
between actual Secchi Disc readings and hydrophotometer readings,
which was not readily available. Williams, however, has developed an
expression involving the extinction coefficient as a function of the
Secchi Disc reading, and since the hydrophotometer transparency
measurement is similar to the extinction coefficient measured under
ideal conditions, it was decided to use this approach.
Let:
B, = Illumination at the sea surface.
B.= Brightness of the Secchi Disc as seen by the eye.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE 2397 NO; 10; IRE.
1, Hydroscope in use. 2, United States Navy hydrophotometer Mk II.
3, Specially designed hydroscope. 4, Secchi Disc.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES 1397 NOS 0; RE 2
I, Peraquameter. 2, Scattering meter.
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER 7
B,= Brightness of the surrounding water at the hydroscope depth. (This
is the background against which the Secchi Disc is seen.)
Bon = Actual brightness of the disc at the disc.
Boo = Actual brightness of the surrounding water at the disc depth.
R;, = Reflectance of the sea surface.
Ra= Reflectance of the Secchi Disc.
Uw = Relative amount of light going in an upward direction compared to
that going in a downward direction at the hydroscope depth.
D= Length of attenuating medium interposed between the eye and the
object.
d= Depth of the glass-bottom bucket or hydroscope.
k = Extinction coefficient.
When the Secchi Disc is observed, it can be seen as long as the
brightness of the disc is greater than that of its surroundings. In
other words, the contrast produced by the disc against its background
allows the disc to be seen as long as this contrast is above the thresh-
old value for human visibility.
Contrast is usually defined in the following manner :
Object brightness — Background brightness
Contrast= :
Background brightness
where the absolute value signs are used to keep the quantity positive
when contrast is produced by a dark object on a light background.
In this particular case, there are two distinct contrasts to be dealt
with—the apparent contrast, or that which the eye sees, and the actual
contrast, or that which actually exists at the disc level.
Using the symbols defined above, the apparent contrast C4 may be
expressed as:
28 Be —= Bo
(1) a= BIA
and the actual contrast, Cr, by:
ere Bop =. Bop
(2) a Boo
It turns out that diminutions of contrast through an attenuating
medium follow this relationship:
(3) Ci=Caer?
or, substituting the values for C4 and Cp from (1) and (2) in (3)
we get:
Bo — Bs _ Boo — Boo -kD
(4) Bian aos a
Since B, is the brightness of the disc at the eye, this means that only
the amount of sunlight reaching the eye from the disc is involved.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Let us derive an expression for B, in terms of some of the other vari-
ables. If there are Bs units of illumination striking the sea surface,
R,B, units will be lost owing to reflection, and B, (1—R,) will be the
amount of light actually entering the water surface. At a depth of
(d+D) the light value will now be B,(1—R,)e*t).
Since only Rg of the light reaching the disc is reflected from it,
the light just leaving the disc would then have a value equal to
B,(1—R,)e*@+)) Ra, which is Bop.
(5) Boo = B.(1— Re) e* Ra
Traveling back upward, the light would be further attenuated over
the distance D, so that at the bottom of the hydroscope the brightness
value would now be equal to B,(1—R,)e*#@+)) Rye", One more
reflective loss occurs at air-glass-water interface which may be as-
sumed to be equal percentagewise to the original surface reflective
loss so that the object brightness at the eye turns out to be:
(6) Bo= Bile— Ree OR aes)
= B.Ra(t — Re) *e
Using the same methodology for calculation of the background
brightness, we get the following:
(7) BoB. eel — he) ee
(8) Buy Beet hee
When (5), (6), (7), and (8) are substituted back in (4), the fol-
lowing is obtained:
B;Ra(1 ny IR) ease Brace jah at Rees tad,
B,Uw(1 — Rs)*e* Ts
Cr Baka Eo Ren SSRs (1 RS Rene™
Buen R,)ere
which, upon simplification becomes:
(9) aC Uw _ Ra— es) er
Uw Uw
Clearing fractions and transposing :
ea (ieee us) pan Wile
Ra Ra
Letting z = A, and simplifying, gives:
em) __ (1 — A)e”’ —A=0
or, multiplying by e**? to give positive exponents, we get:
Ae™ 4. (1 —A)e”™—1=0
which, when solved for e*? gives:
e =0
RD ee
Cy) Oi "3aH iE
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER 9
or in terms of natural logarithms:
kD=In a
er ae
(11) k= D In Us
which in common logarithms is:
(12) — =S log oe (for D in meters)
— 7:54 jo, Ra
(13) = D log U. (for D in feet)
Equations (12) and (13), then, express a relationship involving R,
the extinction coefficient, D, the Secchi Disc reading, Ra, the reflec-
tivity of the disc used, and Uy, the relative amount of light traveling
in an upward direction compared to that traveling downward. Let
us look at each one of these variables a little more closely.
If we define a term £, sometimes called optical density, as:
iG ee
where %7T=percent transmission, we may express k in terms of £
by:
E
Feit
since k is given in terms of natural logarithms. Since £ values and
%T values are conveniently tabulated in readily available tables, we
may easily obtain a k value for any %T value we may have as given by
the hydrophotometer. In this manner we may reduce any hydro-
photometer reading to its equivalent Secchi Disc reading or vice versa
by substituting the k or D value in equation (12) or (13).
The D is, of course, the Secchi Disc reading which may be either
read directly or calculated from the hydrophotometer reading. For
the disc used Ry was about 0.8.
The relative amount of upwelling light, U,, however, was not meas-
ured and values were assumed for this quantity, based on other data
taken by Williams in Chesapeake Bay and by the calculated values
from the large number of stations where both Secchi Disc readings
and hydrophotometer readings were taken.
If equation (12) is rewritten:
k=
SIs
where
*% = 2.3 log a
IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
or, since Rg=0.8,
4 = 2:3'— 0.7 + log at
A plot of x vs. D may now be made, where -~ is calculated from
stations at which hydrophotometer readings which give k and Secchi
Dise readings which give D were taken simultaneously. This plot
shows a marked variation of Uy as the Secchi Disc reading is
changed, and is the graph which was used to determine unknown Uy,
values when the S.D. readings were known, both for stations which
had hydrophotometer and Secchi Disc readings and for those which
had only S.D. data.
By means of this methodology, then, it was possible to calculate
equivalent Secchi Disc readings for each hydrophotometer reading
taken.
DISCUSSION OF DATA
In the two appended tables, all the data taken on the Elsie Fenimore
are tabulated. Table 1 includes the hydrophotometer and Secchi Disc
data presented by seasons and in geographical order from North to
South. Winter is considered to include the months of January, Feb-
ruary, and March; spring—April, May, and June; summer—July,
August, and September ; and fall—October, November, and Decem-
ber. The various stations may be easily located by number on the
series of charts (figs. 1-13, preceding the tables), which show the
latitude and longitude of each of the stations mentioned.
Table 2 includes all the other data taken, utilizing the various de-
vices of Dyer plus a few others which were also used. These data
are presented in simple geographical order, proceeding from north
to south.
The data as a whole, although being among the most extensive
available at the present time, have many limitations and shortcomings,
and these should be kept in mind while any attempt at utilization is
being made.
The hydrophotometer readings were taken with utmost care. How-
ever, the calibration in air was apparently not standardized, the ad-
justment varying from 92 to 96%T in air instead of 92 percent as
previously mentioned. This would have the effect of making all read-
ings above go percent highly suspect since a small change in %T at
this end of the scale is associated with a large change in the Secchi
Disc reading.
This is probably also the reason for the significant number of read-
ings which are above 100 percent, and hence change from quantitative
readings to qualitative. This 92 percent reading in air as being the
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER If
equivalent of a 100%T reading in water was apparently unknown
to the data takers, which is not surprising since the instruction book
written for the U.S. Navy Hydrophotometer Mk. II specifies a cali-
bration setting of 100 percent in air.
The Secchi Disc readings in general are undoubtedly quite reliable.
However, any taken when the sun was low in the sky or in the shade
of the boat are probably doubtful.
In table 2 are given the remainder of the data taken with instru-
ments other than the hydrophotometer or Secchi Disc. These data
have been tabulated separately, since their meaning is not as well
understood as those in table 1.
An attempt was made to deduce some sort of a regular pattern of
transparency in the area covered, but no regular pattern appears to
exist. This may be due to the fact that all stations were not taken
simultaneously (a physical impossibility), although this is not nec-
essarily so. Previous experience indicates that local conditions, espe-
cially in more shallow coastal regions, almost completely determine
transparency conditions at any one point in space and time. Thus the
turbidity will vary from one place to another, one depth to another,
one time to another with seemingly constant environmental conditions.
These data seem to emphasize this seemingly unpredictable nature of
transparency in natural waters.
In general, however, the data do show the following expected
changes in transparency:
1. An increase in transparency with distance from the coast.
2. A seasonal change in transparency, with the winter months
seeming to provide the greatest turbidity.
3. An increased turbidity around heavily industrialized areas.
These three are, of course, to be expected, as outlined by Williams
[35] in a set of general rules for predicting transparency based on
geographical location, weather conditions, proximity of polluting
sources, etc. But there are so many variables to be considered simul-
taneously that these generalizations are often invalid.
This information is therefore presented not as a basic scientific
study to determine the causes of transparency variations, but rather
to present actual conditions existing at particular points in time and
space.
12
10.
II.
12.
13)
14.
TS?
16.
17.
18.
19.
20.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
BIBLIOGRAPHY
Anonymous. Electric camera for deep sea photography. Sci. Amer., vol.
116, No. 20, p. 483, 1917.
Submarine camera. The Engineer, vol. 129, p. 658, 1920.
ArmstronG, A. R., and Ewrnc, C. W. Studies of the measurement of
submarine illumination in Virginia waters in Chesapeake Bay. Virginia
Fish. Lab. Contrib. No. 13, 1044.
ATKINS ET AL. Measurement of submarine daylight. Journ. Cons. Int.
Expl. Mer, vol. 13, No. 1, pp. 37-57, 1938.
BiceLow, H. B. Oceanography of the Gulf of Maine. Bull. U. S. Bur.
Fish., vol. 40, pp. 511-1027, 1924.
. Birce, E. A. A second report on limnological apparatus. Trans. Wisconsin
Acad. Sci., vol. 20, pp. 512-551, 1922.
Birce, E, A., and JupAy, CHAUNCEY. Penetration of solar radiation into
lakes as measured by the thermopile. Bull. Nat. Res. Counc., Washington,
vol. 68, pp. 61-76, 1928.
Boas, F. Beitrage zur Erkenntnis der Farbe des Wassers. Inaug. Diss.,
Kiel, 188r.
Boutan, L. Mémoire sur la photographie sous-marine. Arch, Zool. Expéd.
Gén., vol. 21, pp. 281-324, 1893.
Bumpus, D. F., and Crarxe, A. H. Transparency of the coastal and
oceanographic waters of the western Atlantic. Woods Hole Oceano-
graphic Inst. Techn. Rep., vol. 10, p. 10, 1947.
CiarkE, G. L. Light penetration in the Caribbean Sea and the Gulf of
Mexico. Journ. Mar. Res., vol. 1, No. 2, pp. 85-94, 1938.
Light penetration in the western North Atlantic. Woods Hole
Oceanographic Inst. Collected Reprints, Contribution 101, 1936.
. Observations on the penetration of daylight into mid-Atlantic and
coastal waters. Woods Hole Oceanographic Inst. Collected Reprints,
Contribution 26, 1933.
Observations on transparency in the southwestern section of the
North Atlantic Ocean. Journ. Mar. Res., vol. 4, pp. 221-230, 1941.
Variations in the transparency of three areas of the Atlantic
throughout the year. Ecology, vol. 20, pp. 520-543, 1930.
CLaRKE, G. L., and James, H. R. Laboratory analysis of the selective
absorption of light by sea water. Journ. Opt. Soc. Amer., vol. 29, pp. 43-
55, 1939.
Darsy, H. R.; Jounson, E. R. F.; and Barnes, G. W. Studies on the
absorption and scattering of solar radiation by the sea. Carnegie Inst.
Washington Publ. 475, 1937.
DESYBOS, DonALD P. Transparency by black and white Secchi Disc of the
water of Long Island Sound, Block Island Sound, and Newport Bight,
cruise Stirni IJI. Cornell Univ. Status Rep. No. 18, 1952.
HELLAND-HaAnson, B. Physical oceanography and meterology. Michael
Sars North Atlantic Deep Sea Exp., I910. 1031.
Hunter, G. R. Underwater camera world. Pop. Photogr., vol. 17, p. 42,
1045.
. Hutcuins, L. W. Buoy data at Woods Hole Oceanographic Institution.
(Unpublished. )
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER 13
22.
23.
24.
25,
26.
28.
20.
30.
a1.
32.
33-
34.
35.
Jenkins, F. A., and Bowen, I. S. Transparency of ocean water. Journ.
Opt. Soc. Amer., vol. 36, pp. 617-623, 1946.
Jertov, N. G. Optical studies of ocean waters from reports of the Swedish
Deep Sea Expedition. Vol. 3 (Physics and Chemistry), No. 1, pp. 3-59,
1951.
Jounson, E. R. F. Undersea cinematography. Journ. Soc. Motion Picture
Eng., vol. 32, pp. 3-17, 1939.
JosEpH, J. Durchsichtigkeit Registrierungen als Oceanographische- sich
Untersuchungsmethode. Deutsche Hydr. Zeitschr., vol. 13, Heft 1, p. 60,
1959. |
Undersuchungen tiber Ober- und Unterlichtmessungen und wtber
ihren Zusammenhang mit Durchsichtigkeitsmessungen. Deutsche Hydr.
Zeitschr., vol. 3, Heft 5/6, 1950.
List, Ropert J. (preparator). Smithsonian Meteorological Tables, 6th rev.
ed., Smithsonian Miscellaneous Coll., vol. 114, pp. 214-218, 1951.
MarkKHAM, J. J., and Perry, G. R. Oceanographic stations of the New
London Laboratory from May 1943 to November 1944. Columbia Uni-
versity, 1944.
Pettersson, H. A transparency meter for sea water. Medd. Goteborgs
Hogakolas Oceanogr. Inst., Series B, Band 3, No. 8, pp. 1-17, 1934.
Poort, H. H., and AtxKins, W. R. G. Photo-electric measurement of
submarine illumination. Mar. Biol. Assoc. U. K. Journ., vol. 16, pp. 279-
324, 1929.
Powers, C. F.; Deusier, E. E.; and Backus, R. H. Transparency by
black and white Secchi Disc of the water of New York and Newport
Bights. Cornell Univ. Status Rep. No. 15, 1952.
Seccur. Esperimento per determinare la trasparenya del mare. Cialqi Sul
Moto Onaoso del Mare, 1866.
Wuirtney, L. V. Microstratification of inland lakes. Trans. Wisconsin
Acad. Sci., vol. 31, p. 155, 1938.
Wuttams, J. A small portable hydrophotometer. Techn. Rep. Chesapeake
Bay Inst. In press.
. How to find clear water. Water World, May-June, 1956.
14
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 139
Ge
Son
50°
49°
48°
47°
ON
_
°
5
ae
| se
ik
| A
|
|
| ae
i
i
i
Ly
J
5
a
a
i
L
a
a
|
i
Bee
of]
a
LU
4
]
|
C
J
a
i
5
i
a
C
i
q
|
a
YI
a
|
i
5
a
LI
m1
{|
|
|
i
5
Z
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
aA ee
cays eae
15
$1°
50°
49°
4s°
47°
16
SMITHSONIAN MISCELLANEOUS COLLECTIONS
e
Sable Island
VOL. 139
4
46°
45°
44°
"43°
141°
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER 17
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
44°
42°
41°
40°
39?
'?
UI
a
|
4
ie
at
a
UW
|
a...
a
C
4
UI
ae
| | 5
C
r
UW
a
a
a
sof]
g
a
C
Z)
(5° 74° hon 72° 71° 70°
Fic. 5.
NO. IO
2
WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER 19
78°
77° 76° 75° 74°
J
[TT ETT TE TTT T Ter TTT Te eT ere rr rr rer rrr rr rr ier TN
Fic. 6.
73
é
Biot
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
85° 84° 83° §2° 81° 80° oe
NCEE
~o
°
no
ao
°
Beal etal e ee afela alah ase ele lee lela gale pals lolol ay eal | palep etl apayate ea
76°
26 ATT TTT TT TT PEER EEE EERE See re rela]
85° 84° 83° 82° 81° 80°
Bom
se
31°
29°
PRO
ie
26°
21
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
Fic. 8.
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
91° 90° g9° 88° 87° 86°
Fic. 10.
23
30°
aon
28°
4 Dae
26°
baa
24
SMITHSONIAN MISCELLANEOUS COLLECTIONS
oo
+ Alacran
R eef
90° 89° 88° 87°
VOL. 139
86°:
ooo
20°
19°
18°
Vis
16°
25
WILLIAMS, JOHNSON, DYER
WATER TRANSPARENCY
Io
NO.
°
wm
N
LITT TTT TT
°o °
=
N
mm
N
Dp
\J
92°
93°
94°
95°
96°
Ti
. 12
Fic
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
26
98°
94°
96°
oT
98°
92°
93°
9
96°
o7>
98°
Fic. 13.
27
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
‘a's
Tenjy
Sg
v6
$6
9°6
gol
ZOo1
gol
IOI
QZ
zs
v9
for
gol
gol
OI
6°01
II
wilt
Crt
Sey
C1r
£9
ci1>
‘as
“ainby
Sz
oI-g
ov
Oz-gI
Oo
suipear
“O1pAPT
~
N
~
_
~
°
g-0
q-o
yideq
(panuyjuos)
OoOIl gr /6/E 190,94 S°bS gf eee gen eereemy ciel gases ga segctumrelonje sae SE ANETO Gl eoys uMOIg
o£iz gb/zz/z u80,¥S FL 144,95 o8e oe ee ee eee e newer eee eee weer ee eeeeee sees eee ‘og
aes gr /tz/z uR0,VS Vs n4¥, 98 Sf Cee eee meee eee eee wee ewnnne SUE ER IIONO EO]
SzgI gb /1/€ 180,¥S V4 vL¥ 95 BF So een eee ae aan heh ag pike at ee peek ee OG
= gb/Z/E u80,¥5 v2 ve¥ 9S .8f See Ee ee caren ener es cto al OCLC ET ACI adey
OoZI QV/O/E ,SS,fe,S2 ,cV,ZIoOF “°° =" "2" "ss *="" Avg sIEMPIOG ‘IWsry Uyol- dys BO
omy, =e (AM) CN) worsdr19s9q7
apnyisu0'T apnqiyze'y
Se
UOI}L}s JO UOT}LIO'T
MALNIM “V
pawpnys SuoDjs 4D
shuippas ISuT WII9G yuajwamba 0}; sbupvsa sajamozoygoaphy fo uoyonby—t alav ye
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
28
‘a's
enjoy
S11
II
Z01
€or
gL
al
gl
1°
SQ
38
5-6
6
88
1'g
1°9
Sg
C-SI
S-E1
CVI
Qri
gvI
ZI
ZI
£91
£91
gZ
1°
cas:
“aInDy
ov
Ze
ce
ee
SI
£1
gI
gI
oz
zz
Oz
cz
ce
gI
gI
Oz
oS
os
tS
cS
SS
Z9
<9
09
09
SI
gI
SuUIpess
‘o1pAH
jov
ie
joe
ve
81
jz
9
oO
da
i8V
yidoq
ofzi gr/S/ze $:10,$2 iQhge Siete ela eee ster ao era ee Ne TITS Ws] S[TPI2AQ YO
Zier gh/gz/I (8794 (05,.g& *“seg ayxvadesay FysrT log Apooyg Jo *M a!u g’o
OOOI gb/S/z2 1S PZ IS ,8F Baie oi ae eae ee ashenateseenaeecnean 7 (AKIO) ) jeoys SITIDIIN
OOIL gr /6/E {90,2 $,VS gf Seether cieneicae cnee seers ionte eS TBM BTS (T {eous uMOIg
omy a7eq CM) CN) uordriosaq
apnzyisuoy epnzyyeq
SS SSS
WOI}e}S JO UOT}LOOTT
(panuiju09) MALNIM “V
(panuiyu0d)—l ATAV I,
29
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
6°02
6'0z
oz
S°SI
SEI
C1
cs
cs
19
cs
gs
19
19
19
19
19
19
SEI
VzI
VI
vz1
C1
Cat
Sam
ZI
89
ZZ
GZ
bZ
LL
bZ
rZ
cL
o£
ZS
oS
oO
+6
+6
co
S6
C6
5)
c6
c6
c6
oS
cr
cv
cv
ov
ov
ge
eh
199
199
SIgI
0060
ooSI
gv /t/z
gr/6/€
gb /S/z
(panurquos )
25,2
$1;90,S2
LS€,bZ
2VQE woereee sees dS FST S][ejIeaOQ Jo “YS som cor
CLZV QE coos esses soqieyy Joyemyeoig ‘aremejed ‘someT
BVRE Sere ee ee eee TS WYSIT woy}e,J-S jo Ss So] Tut gz
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
30
HS
8
‘a's
Tenpy
IOI
£O1
£-o1
C1r
VzZI
‘a's
“alInby
ZI
gI
£z
cz
6z
6z
6z
If
ze
ze
6z
of
SI
oz
£z
Sz
lz
Qz
of
of
1¢
gz
Qe
of
ov
4
sSurpeal
‘orpATT
184
yidoq
OfrL gh /6z/1 (02,94 {OI Bf Cee eee eee eres eserseeeseeeessesesesseeseese ‘od
O£e1 gh /6z/1 (02,94 jO1 Qf pote achniessaelioy 81eAe nen Gel eka cc eos ney aneussouuereet se seve OFT: Jepsy yO
Sze1 €v/3/1 ,,0£,82,94 ,O€,O1,g€ “cccc cesses ss py ‘pueysy suowojos ‘souejeg qwIog
Sigt gb/v/z ea vl VoQE Co aS JST Sy[eyzoaAC Jo “qs soajlu zor
OntEy, s3e@ CM) CN) uord1osaq]
apniisuoy apnyyqeT
ee ee
UO0I}e}S JO UOT}LOOT
(panuiquor) MALNIM “V
(pantijuor)—1I ATAVY,
31
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
6°6
66
6°6
or
6°6
ofZI
ofgI
oest
gb /6z/1
gb /6z/1
gb /6z/1
(pamuijuor)
{07,94
j07 94
(02,94
OI,8f POCO OTOH OSE HEHEHE OEE HEE HEE HOO HHO HEE EEE ‘og
OI,8e COP eee ese r sree eeereseesseeeeeeseseesese ‘oq
OI gf Ce ‘od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
‘a's
jenpy
I'O1
l'OI
£01
ol
oI
£01
£01
1‘Ol
ol
99
al
v6
OI
OI
6°60
66
lO
IOI
oI
I'O1
ol
Iv
z'9
6
36
‘as
“AINDY
Lz
1¢
1¢
ze
of
of
ze
ze
if
o£
ol
€1
cz
of
of
6z
62
1¢
1¢
of
1¢
of
z
8
€z
Qz
surpesl
‘o1pAH
irs
iS?
jv
if
joe
ibe
81
zl
9
0
99
109
vs
18h
ev
i9&
j0£
jz
81
jel
9
0
99
09
iS
is?
yidoq
ofé6l gb /6z/1 507,94 /O1,8f Sere errr reese esreereeeeesesereesseeves ‘od
oO£gI gb /6z/1 507,94 OI .8& Bia stp esse cathe aA ANE (ATT) Suiog repay HO
ofZ1 gr /6z/1 {02,94 OI gE Sse ee seen ee Ors eine ewe ee eT OT lepay yO
oun, = ae CM) CN) uondri0saq
apnysuoT epnyyeyT
a
wolzeYS JO UOILOO'T
(panuyuor) YALNIM “V
(panuyuo0r)—1 AIavy,
33
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
6°6
IOI
£o1
£0o1
£o1
£o1
£O1
OI
OI
v6
6z
1¢
of
of
of
CI
oz
cz
of
Ze
2
ze
Ie
of
ze
SI
fz
Le
of
2
ce
ze
2
of
of
Sr
cz
ofze
o£1z
ofoz
gb /6z/1
gh /6z/1
gr /0/1
(panuyuo?)
02,94
{07,92
‘
0
094
/O1,8e Ceo eeeerooseseeseseeeeeseeeseseeees ee ‘od
OT,8f eee eres eee eee eee ee eee eeeeseeseeereesee ‘oq
FOLRRe Coe Cee eee eeeereeeesereseeeeeeeeeeereeee ‘oq
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
34
‘a's
Tenpy
OI
SII
£91
£91
ZI
S61
Z°g1
gI
gl
OI
66
66
1'OI
OI
OI
IOI
9°6
66
36
36
g°Z
go
1'O1
SOI
OI
66
36
‘Gas
samby
I¢
sulpeal
‘o1psAH
81
je
9
0
199
199
vs
ig?
jov
it
joe
qideq
gIQI gb/b/z 9)z0, $4 aot SER OO AE Aaron oe et sc ater ease aT COU GSTOr MUO TEL (G)
offz gb /6z/1 08,94 01 .9£ Cee e erence errs senccees sol estaba lots Saye
ote = gh/6z/1 (0% 94 Po oge ere EVER EES Sane: "91, (ATA) ‘Wtod 1epeD HO
ow y, a7eq CM) CN) uorjdis9saq
apnyisuoT epnyyeyT
nn SSeS
UOl}e}S JO UOT}EIO'T
(pamujuor) YALNIM “V
(panuyuor)— I alav L,
35
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
“LS
ZI
9
199
vs
BY
9&
of
v~
81
ZI
9
of
rz
81
ZI
9
4
3?
9
o£
vz
Bt
A
9
SrZo
OOIT
CiIzI
cer1
gr/v/z
gh/t/z
gh/t/z
gb/v/z
(panunuos)
27,94 [a0 4e ec aseloyouy onsvoyessy
$:S0,$2 O54 “orc tpeoyS JoyrenZ) JoyUrAA Yo Ainy woyjej-o1
Z01,SL 9)20,98 “cc cc ccc pues] onseazessy YO ‘aAinD woy}e7-S
¥;00,S2 V:91 Sf ie lois fovea (ove Seer eisioieis: tse oe eS EC, 1m) yearn yO
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
=e ie
Oo
je
a's
npy
99
1g
g4
1°
giZ
gs
VL
ciI>
S1>
Ge
Se
Ge
ce
I'v
GES
*AInDy
HH He HO
c
surpeot
“O1pATT
qidoq
cezi = €v/O1/1 OT,9Z 4,9E,SO,4E occc erect tse ess Keg ayeodesayy) ‘yeoys: afquiryy
VIII ev/0/1 JOls04 {fOb0Z cE “2 ee renie tons Ke oseodesous) GerT, JON TO
ofe1 gb /£/z ,00,52 nGaaae Neg eee tense elie det oae ea eM ea eee UTE TOIT qqop yO
$160 gb/v/z ,40,S2 Lv LE corre rete teeter cece esses seeesoompe UShT WEI BO
740) gb /V/z ZS eG. LG» isieitir.cnieh woe ninven es clusiiee + SRB IOUIU YE ans eoieEsy,
out], ayeq CM) CN) worjd110saq
apnyisuoy opnyieq
a
UOl}e}S JO UOT}LOO'T
(panuyuo7) YALNIM ‘V
(panujuod)—1 adv I,
37
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
49
c1I>
bz
ez
zz
Iz
oz
,O1
ist
ait
91
ar
al
£1
zl
9
4
Vz
£z
OC
IZ
Oz
{OL
81
al
91
SI
VI
or
ZI
‘
‘
oO
0
Sv10
CSroo
oort
gr/f/z
gh /£/z
£V/o1/1
(panwijuos)
Gat OZ
G'11,94
{01,94
GE GOL) eee sro eels iate ce alae lopre sear tegen CT
{S£,S0,2€ “7 keg ayeadesayy ‘spunory a[ppryy s0ysass0;T
19° ,90,LE Gre <6 676056 elullale ele evel eaie slecetelale se) ejje/uinis el eibin ‘oq
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
‘a's
jenyoy
Ig
Eg
Ig
Dahy
Ig
8
Ig
‘a's
"ainda
6
Zl
VI
SI
gI
gl
61
02
IZ
Of
gI
61
oO
cf
8
vI
SI
gl
gI
gl
SI
gI
gi
gI
gl
ZI
gI
suipear
“o1pA HT
Oo Crco gr /f/z G;11,92 ae Oye Coe eer ere sees eeseseseesesesereseeese ‘od
OL
81
41
91
pet!
vi
jel
ZI
9
Oo Svzo)8=—_ hb /£/z G-11,9Z ,S€,S0,Z€ -*°**** Keg ayvadesayy ‘spuno1r o[ppryy eysestoF{
yidoq = OU, aed CM) CN) worydtz9saqqT
apnyisu0T apnyzeyT
UWOI}RYS JO UOIWLOO'T
4
(pamujuor) YALNIM “V
(panunyuor)—I aTdv I,
39
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
g
QZ
cr
cs
9°9
t~aowrnn
or
gi
Oo
Bt
Sez
Sriz
Svoz
gr/z/z
gh/z/z
gh/z/z
(pamujuog )
Sj11,94
$;11594
$;11,94
nae, G05 Sie lsioinvereisveleveiovorsicierenes
oe ,90,4£ B06 elev sia veloleleters lereralelevalelelesiaienaianciots
ae Ones: Vlelelolelecierers lois ipierslareiee
see eee
og
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
40
vS
a“
aS.
jenpy
(B65
oI
£-O1
relia
VZI
VI
Cri
oor
GST
“G1
gI
Se
gv
3
v2
gl
Pi
oe
‘as
“andy
$g-€g
ZI
VI
OI
O
I
SuIpeos
“OIpALT
q-0 00z0
j24-,vz
Bl
jel
9
oO Ooo1
aM
9
O OLO1
ve
13"
jev
i9&
joe
ive
it
jel
19
O OOII
ive
81
jel
19
oO cre]?
£z
ce cre
4
4
dog = omy,
0S /S/z
0S/b/z
oS /P/z
gh /E/z
gr /z/z
gh/e/z
ajeq
Sites QES€ **UOILYS PseNnr) yseod OdtWOOKUIKOIY) fo “ay safius FEr
Gorge G;41 ,9£ occ e snare eee DIOS yonyaing jo "7 SopTU Eee
y85,01,92 p0e,¥S,9£ Core eee ee eee eens oho ncl en escncanien suas SUIS LTV AG *yao1g IVT
Le j00,4E Pe aS SASS SSC EN ESSER ape teneey* Ae APTS WST ayeodesoyy
$511,942 Way aOace eee e eee esos esse ees eessesesrsesesesevee ‘oq
G,11,92 ,S€,S0,Z€ ***:****keg ayeadesayy ‘spunory s]{ppryy soysoss0F{
CAA) CN) uoiqdr1osaq
apn}yisuoT opnyijzeyT
ss
UOlZe}S JO UOTZLIOT
(pamuyuor) YALNIM “V
(panmjuod)—I ATVI,
41
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
v9t
“3,2
+6
Lez
Lz
£97
c'0z
£01-g9
83
55-88
88
gs
Ailiti<
£01
£7
190-0
09-0
q-0
q-0
q-0
128-0
q-0
q-o
{32-0
,00-0
/3Y-0
gq-0
oocom
SPgt
OfII
o£go
ofez
SeZI
Srol
Soor
OOII
$160
Siz1
S£60
ZV30
oogI
ob6o
oSSo
1100
Sccr
oSZI
VEe1
0030
0090
O£9QI
ACA
0S /z1/z
0S /6/z
0S /6/z
0 /g/z
€v/1z/1
0$/g/z
£v/1z/1
0$/g/z
€v/1z/1
05/8/z
€v/oz/1
€v/1z/1
0S/9/z
0$/9/z
0$/9/z
0$/9/z
€v/v1/1
0S /S/z
0$/S/z
0S /S/z
0S /S/z
vp /€1/1
ev /€1/1
(panunuos)
S'9z,18 G;€z,0€
$;90,18 S°20,,08
S;1¥ 08 VS,0€
/87,08 SFO SALe!
OV ,08 Goaig le
VS,02 j00,z£
°V,62 ,,0€,0V,z€
/9V,04 cV ce
OV 62 WV ze
VS 62 ZS ef
jea,00.- Ob avoee
2a004 je ave
CE8 92 0S ze
190,01,,84 ive 01 oct
GjSE,42 Zien ee
20,22 S Annee
0b, zh bf
01,94 Size ve
Bj2o SZ QS ,ve
Sjoz SZ )80,S©
“Zizz.Sh SZio1.S€
8£,94 1Z,5€
{51,92 S/0E Jae
eee eee ee” pipe cay ng nee ee SISOS “ely ‘rod Ae
eee ee eee se eee “dys sry] suyof aS HO
eee ee ee eee ey) ‘spur]sy purytioquing jo “aA
Di iersisierersie ieee WsrT yeuuerars yO
‘diy ysry yeuuears ‘eare yeuueaes
os
.
eeee
.
.
.
oer ee eee 29) ‘yeuueaes jo ysey
“"*9z% “ON Aong vag uO}saTIeYD
Gavereamucsses a) Ss ‘uojsop rey)
"*satyjef opisjno ‘ease uojsapieyD
SO ones eke each eacin) aS ‘UOjsoy Tey)
reteeeeeeeeeeee log
"I9JUMNS 110.7 Yo ‘ease uojsapseyy
Soild eel e heres TT EULLO SY] ode) HO
“yYSryT ueg surdryy wor sop 2Vve
“ress diys FysryT ueg Surk1y YO
06:6. 0;8 6 uo 5 0b. @ e's 8 8D ge ele a Bee Aong IITA
"*4seOo eUTOIe) YON ‘A}ID peoyoio0jy
SNSICNe oie serene teh gn meea TOM OORT adey yO
tteeeeeeeeesssagmT 909810 HO
Peet eee eeeeeeeeeenee sog
Gianereelivieielevetersiy SPIOL ET adey HO
ele see © 0 bie 6 e's 6 8) 0 ele JIANT oorwue J
‘yeuey) JOARY OSung pue s0}es Ty
eS gl 59
nt “2g1 £9
3 21x z‘O1 39
= SS ci
a 4
9
y, z’9 8
s 99 oI
oO ie g°Z SI
A — S11i< = + 001
a 1'g gi
oO OI o£
n O71 Lb
5 z'61 gg
o g°St 4S
<i v1 €S
| — SI S
ey gv €
a gZ SI
a Ser ov
z GOI 69
a £31 Lo
a v1 Ss
B “Lz1 ov
m — 621 Lb
=
a Tea g-£
a _ oI of
‘a's ‘as Surpear
[enjoy “AINDg ‘OIpATT
Q
wr
tz-0
o£-o
yideq
SPeL
OOf!
00gI
Ivol
ooo
ofo1
$160
our,
0S/0£/€
0S /gz/€
oS /€1/z
oS/f1/z
0S /€1/z
0S /z1/z
oS /€1/z
a7eq
$:z1,%6
0£,%6
Ligh 08
$;1Z,18
12718
8£,18
12718
CM)
apnysu0'T
{80,62 RNAs eerie Maree Deis Oa ATES) TO ‘yueg pleoH
{01,02 Sietoperecieiel vere chenc lens meitre oe eA OTC e®3S “xOL, ‘WO}S9ATe)
£:90S ,6z Ce EOS HIG B ‘Quisnsny 4S HO
VI,0€ Mee ASeeSOan elena! Sey Sie elaksne Se Steerer Be eeeeeO TT ‘OpId A. o}U0g yO
jO1,0€ eee eee ee eee eee eee eens pieCRA DBT ‘yoeog aunydaN HO
jO1,0€ speben Grek ete eh hoe Xess eer ceo Sent opel eae mlesceeee (iT ‘IOATYT suyof 1S
G;€z,0€ CO crcecar phasvicke seals Nexeesietek SRT yrodAeyy ‘QoueI}US sorqjof
CN) uorjd110s9q
apnqziqey
Seana,
SSS SN
WOl}eIS JO UOT}BIOrT
(panutju09) YALNIM ‘'V
(panuyuor)—I aTdv J,
43
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
use Px
oT
z9
99
CL
gr
19
gri
gr
Saar
fg
OIL
gr
CII
Lilt
“11
O'll
Z9g1
eos
gv
oz
Ger
£-v1
QVI
Lila
“QI
Ser
II
“Sr
£91
0
8
OI
£1
cs
c6
cs
cs
ZS
09
$°66
zo
06
ov
Iv
Iv
Lo
gZ
zO0
oZ
oS
es
ss
79
49
oZ
oS
Ze
gs
199
99
199
;vS-o
jo0-,FS
Sv
zv-0
col
96
00
v8
82
;7L-0
06
‘
184
99
(09
vs
igh
jv
9t
joe
j¥z-0
ov
cb
é
‘
(panuyuos)
SuOI}Ipuod ASSO} JopuN JO JAaSsaA Jo apis Apeys UO Udye} SUIPVat ISI] IyIIBG Sa}eOIpUy x
oSZo
$1So
ofoz
O£I0
orez
0S/gz/E
oS /r1/z
oS /1£/£
oS /1¢/€
0S /1£/€
$;10,56
$/95 68
jZ£,e6
eizv 68
PRC men ae iaectcn cs ‘''xoT Yaodse1g Yo faAiny woyyey-S1
9e 8e Cee reer e sere er ese eeeesese eee eare aT: fade) IS]e yO
Of 82 Coeereesreerreeeesreseeesees ****yoAuey) IddIssisstjV PIO
OV 82 Pere errrerercevrereeseseserevese "* 99 *OOTXOT jo yJny
20,0 Cee eee eee eee ee **90ueI1]U 1ddIssisstjv JO ‘MN keg
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
44
6€
it
ne Pl
‘a's
[enpy
ZI Lv
83 oe
Z ZI
gi So
SEI os
Zo1 S¢
6 £%
9°9 ol
9°9 or
SII<-g3 -+001-96
19 S6
19 56
1S £6
SiSe: £8
nl< OO1
89 96
ss r6
av 06
SII< +001
tS £6
1S £6
19 S6
LL 46
£01 S€-0F
Egc-g'rz = zQ-QZ
‘cs sulpear
*amnbay ‘O1pPAH
oO SVgl
9 ofSi
oO CIOL
joe
jve-0 ooz
84-8"
jv
9
joe-0 = 00D0I
)OsI-0 oStz
i8Y
ev
if
0&0 =S1EI
99-87
BY-0. ofzI
yideq omy,
0S /£1/z
0S /Z1/z
ev /Sz/1
0S /1z/E
0S /Zz/€
0$/£z/€
oS /r1/z
0S /1£/¢
ayeq
240,08 ,,S€,50,92
Z;Z0,08 ,,S£,S0,9z
u22,20,08 WE, I £92
,00,,46 00,42
5€,96 zh dz
joe s6 8 86SiEV Le
01,08 bb cz
$/6S ,06 LE Qe
CM) CN)
apnysu0T epnyyey
s Sece.e spoinisistsrsr pny ‘oqie FT SOPe[ SIVA 10d
stress ser ‘poomATOyy YO ‘ease ojepsopne Ty
sinks see ele veat wins pisielel sista sie, e( iO Nn jo ypny
Soluiswanenelelie “YStIUD snd107y yo ‘OIIXOTT jo yn)
stresses systray) sndioy Yo ‘aAsind uroy}e.J-oo1
hee eae ae eocne sis eee ren eT ey ‘yoeag JO}UL MA VO
settee eae “yeoyS drys yo ainy woyjye y-O1
uorjdriosaq
—_—=?.? o>. ———
(paningnoa) ALN IMS V.
(panwjquor)—1 aTav J,
uolze YS JO U01}L90"T
45
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
sé
48
ol
49
8
oogI
o£go
OzI1
SSor
Se61
Ivgo
orgo
0002
oob1
£b/g/E
£¢/6/€
ev/z1/£
ev /€1/€
£¢/g1/€
e¢/L/z
€b/6/z
0S /L1/z
0S/Z1/z
0$/L1/z
(panuyuos)
u21,40,08 MORAN OR Germne Sok ss sane aut eien cc euel tera ahs
u21,40,08 PUOCORGe cots hee eer ee meen
Gap OsOge OR SORGeR © 2. oirioeinre helenae ares
121 ,40,08 OCT ORG Bi ee ese eis ie she aha oieae cea
81,2008 ,06,50,92 “ic ttt
y0°,40,08 FOR COMES L22t le Sees emer
,0£,40,08 yOL ,S0,9e SNeMabelieliale arene Shelieleleisie-erelt
Z:L0,08 ,SE,S0,92 “itt
z;Z0,08 Se 809% eoereeeer ee eee eee ee ee eee eee
ZL0,0Q ,,SE,S0,9% crores ites ttle teen
.
oq
od
od
od
oq
oq
‘so0q ‘ayepsiopney 310.7
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
at _ =e oO 0060 = €t//gi/z ,40,08 BjpOoGe ~* - oo oem encase SUTIN ‘S[epiopney 307
/0 cet = oO Cré0 £b/E/z 421,40,08 OL 90,92 Chee oreo nee e scree este e eee cecceete tes ‘og
a = a oO QIgI £v/v/z 721,40,08 pOF ,80,9e eee eee eco eer assesses tere eee cece e ese ‘oq
Ss s. ji Oo grz1 tvr/v/z 721,40,08 OC ,80,9e ee eee Cee mm ee ee ewe eee eee eee were ween ‘od
oie aca a. oO SSgo0 cv /v/z 421,420,038 y0° 50,92 ee ee eee ewer eneces ‘og
ie = =——s oO Corl £v/S/z~ 121,40,08 OL 80,92 Coe eee eer eer seeseeseersesesesseeesseseve ‘oq
ye hating ao oO Itol £V/S/z 421,£0,08 yOe 80,92 Seer e ere eee seo eee eee e esses eee oreseseses ‘og
HEC p so oO S£g0 £v/S/z 421,L0,08 10° 80,92 Comme eee eee ere e sees ereseeeereerseseses ‘og
XS ested c oO 6£g0 £v/9/z 121,40,08 0€ 80,92 Ce eee eee eee eee ee sere esereseessresene ‘og
(49 tz ‘ne oO Qzg30 £v/3/z 121,40,08 WF 80,92 eee eee ere rere see ener eesesesseeeeeeeee ‘og
RL cs ae oO Crri €v/€z/1 151,20,08 yO 80,92 eel e)le iu lele)eeie6ceteie lelexeleieie-aietensretsaloteieetetedecs ‘od
Fir 2 ex Oo SI1g1 €v/gz/1 481,40,08 y0£ 50,9 ef e208 /[@, ella (e/a ere: @h6)0),6) 6) 646046) 816761616) st onat ep ielnisle ‘oq
Fat pat bars oO Lyr1 €b/92/1 151,40,08 {FE S092 Ssevelsleletekalalterieleveceielviavara)s/eisievelevacanchenetnictereie ‘od
(0 sue. Se oO OVvItI €¢/gz/1 481,420,083 OL 80,92 Ou @kelleers ere) edeielrareienhiex6 welled ara\/elelalelelevellehelarare ‘og
8 co =e Oo Cc60 €v/9z/1 1S1,40,03 0 80,92 Sl ONOia 1616 (0) eLOl ee 0010516 e:0\616)es8lle.6 eer els) eietereaiauets ‘od
x01 —e ae Oo SSe1 Er /o€/1 451,40,08 {0£ 50,92 oenetetelereieeislerelaleluei ates) elecelereieieieleevarevsiesele ‘od
eb a se! Oo Soo1 €v/o£/1 u81,40,08 OL 80,92 Cece ceneccrecceseccceeseccececcece ‘og
Vv a ale, Oo zrgi er /o1/z 151,420,038 70 ,S0,9e Soe eee wee rere ese reer ese esreseseseses ‘og
2 — ES oO grgl £v/o1/ze 421,240,038 yOL 80,92 CP ee meee errr eres eeresesrereesessseee ‘og
8 = a: oO grSi ev/o1/z 21,40,08 OL ,80,9e Coe mee m meee rere sree erereeeseeeesesees ‘og
fol eae —_—_ oO gShrI ev/oi/z~ 121,40,08 y0£ 0,92 eee eee eee ee eee re eeeeeeesesesesesresves ‘od
jol aaa ‘aes oO Creel €v/o1/z 721,40,08 yO 80,92 See eee eee eres e sees eseeeesseeeeesesese ‘od
8 me mai Oo cril £r/o1/z 121,40,08 0£ 80,92 Coe ee eee eee eee seer eeeseeseseseesesese ‘od
4 pi c=, Oo grol £¢/o1/z 721,40,08 yOL 80,92 Com ee eee eee see eee eee seers eeeeesese ‘0d
We ms = oO S€go ev /o1/z 721,40,08 10£ 80,92 eee eee eee eee see ese eeseseeseseeseeeeeses ‘oq
YI —- oO ZSE1L ty /zi/ze 751,400 {OL ,S0,9% See eet ewes eer r eee eens seseesresesevens ‘og
VI _— _ 0 OfSi £V/Ei/e Sa ,20j08 (Of S0,02 2205 oitsiccsssseescesigtctnn “pod. ‘oepryepney jog
aes) AES Surpeot qideq ourL 97eq CM) CN) vorydi10saq
jenjpy “aInDy ‘OIpAH epnyisu0T apnyiey_
Senne nn nc TITIIEIEIIIIE SEINE
UWOI}e}s JO UOT}LIO-T
(pamuyuor) YALNIM ‘V
(panuyuor)— I ATaV J,
47
IO WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
NO.
uv ve
109
SI
£6
OI
fol
jo1
‘40
jf
jl
Hain
fel
for
Cer
¢-Z¢
gv
9
eo) © Oo 0 oO) OS
cei
OOfI
Sriz
Sgr
OfrI
oo61
oogI
ooZI
OogI
OoST
oorl
Oo£l
ooz1
oOorl
OOOI
oS /1z/€
ep /€1/€
oS /Z1/z
oS /£1/z
0S /Z1/z
€v/g1/z
ev/gi/z
€v/o1/z
£b/g1/z
€v/gi/e
€v/gi/ze
€v/gi/z
€v/g1/z
£r/o1/z
ev/g1/z
( panurjuos)
$790,40
20,08
z;S0,08
Z;S0,08
Z;S0,08
40,08
40,08
40,08
40,08
49,08
40,08
40,08
40,08
40,08
,40,08
$;b0,9z SOB GOOROOC COD OOO 0ODAMOQO HL) qhay Bas ‘olzelg 0}UeS
SSo,9¢ “ott t ttt pouueYyD diyS yNog “vare sepsopneyT
gj$0,92 *
$°S0,92 -
$;S0,92 °
$;S0,92 °
$;S0,92 °*
$:S0,02
¢;S0,92 °
G°S0,92 °
$:S0,92 °
$°S0,92 °
$;S0,92 °
$;S0,92 °
G;S0,92 °*
ss orga SOO Gy
See eny
‘sope[sI0Aq 310g
SopR[SIIAW 310g
“od
‘od
‘od
‘od
‘od
‘od
‘od
od
od
‘od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
48
Sl £-g¢-L7z og-SZ g-0 oozr oS/Lze/z VEE 18 BER GGs SUSE Le ne Ban, ‘Sezer Je 0
— SII< Ss + O01 0 StSo oS/Sz/z Zj0£,1g Bi HOE See eee teins CBO ASEZUEIEIN IC)
£1 gv jbz
Les v9 1
at ZS jz
eI tv 9
ae er gh Oo Ofer 0S/z/€ 02.28 (OO, 8O SE e NES oN esau TOGNE ET BURAR ET
gti ae joe
Lez SZ bz
gr ae 1
£1 QV je
6°6 6z 9
— 96 Qz oO Ofer OS/E/E ,,0£,02,z28 MOGEe sors o res ss ete os sss 330g) [PMUSDsOqIeET VULAELT
= cnm< +-oo1 Oo Sooz 0S/91/€ £z,L6 SOORf eas oot reese rec ce eee Spe Scie = OOM STN jo yny
—_ SII =—s— + 001 oO ofzo 8 0S/61/£ 940 VE Sunn esses tie COTTE) MOsKO FO F[HXy
oll ane< -_-|-90T oO of60 0$/€z/z Of £2 [EOE cos pores SS sen ee see F eS PONG) pup iOsan suo
= SI11< = + OOI q-0 o€g1 oS/ez/e Lov, 4Z ache Tite Ty tis fos 229 ss OURIsT sOlpmy, “alae o1ppHIN
—- SII< = ++ 001 gq-0 of01 0%/61/z2 lee le PaO Mag apie oo pe Coen re susesrsiehegs "sess" JoqIepT nessenN souesUq
_ SI1I<—s + 001 {991-0 off0 0f/61/z $718 42 Banas oes ONE eres roe sei es Apa anne meats)
/S9 =o B= oO 9£60 er /1/z ,Z0,08 BOO eis eset ma ssearemer nes Sates. oes senha oe Ge sare acs oeag PooMA [OFT vO
Fag aS a oO QIOI £v/g/ze S;70,08 ,10,92 eee eereeeee eee eee eeeeeeeeeeeees ‘0d
me =< cee o ces th/g/z $;S0,08 PLO ROE as Ne sia ehaisibp Sicis sivieedy selene Sey ar
Le = ars oO 160 £v/3/z $;S0,08 ,10,,92 eoeceee reser ees eee eee eee eee ee ee ‘og
ioe = — O oogI £r/6/z $;S0,08 {10,92 eee eer eee eee eee eee eee eee ee eee ee ‘oq
)sS a= eae Oo ofII £¢/6/z ¢:S0,08 ,10,,92 Seem ee ere esse eee reese esesesece ‘0g
Of aS a oO orri fb/z1/€ $;S0,0g TO MOR, eine ng ee yovag poomAToyy Yo ‘eaie s[epsopney
8 a cal oO oofir eh/11/€ 40,08 WO,0o 220202 tors ce ess? pes. Inn) “vere ojepsopneT
‘as ‘a's Surpear qidaq our a7eq CM) CN) uordr10saq
jenpy “aInDy ‘oIpAHT epnyisuoe'T opnyjzeTy
eee
U0N}T}S JO UOTILIO'T
(panuyuor) YALNIM “V
(panuijuord)— 1 Alavi
49
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
£91
gI
gl
£91
£91
= QrI
_ cir<
— c1ll<
{OV cli<
£1
£71
£1
GI
AO LL LLZI
AXE £°9¢-cv
be ob-£o1
19
‘
Lo1
S°S6
4c 19
— cli<
— Crli<
jcvl c11<
jvit c1i<
ZI
VZI
ZI
Zo1
—_ gO1
09
59
o£
19
09
cs
+oo1
+oo1
+oo1
gv
or
oS
9
09-06
63-66
S6
S6
$80
c6
+001
+001
+ool
+ool1
£v
SV
£V
c¢
be
4
a
jel
9
oO
OZI—,ZO1
{90-hz
jve-0
oS—O
q-o
{001-0
84
24
99
109
,VS-0
i¥g-0
g-o
9
é
o0zI
ofSt
o00zz
Ooo!
O£II
OO£I
orri
Oz£I
oOzeI
CIe1
£ez1
QzIl
o£o1
££60
o£g0
Sr10
ooV1
ooVvi
of61
o$/z1/¢
0S /S1/€
0S/9/€
0$/S1/€
0$/3/€
0$/91/€
oS /Z1/€
0S/9/E
0$/Z/€
0S/Z/€
0S /S/€
0S /tz/z
0S /t/€
0S /tvz/z
0S /Lz/z
(pamuiyuos )
1809,80,,96
150,96
£0, £6
i856
05,10
V1,46
j£¥,L6
S:1F,6g
S-17 68
S-1b,68
1 008
S522
£198
£:01 62
PEE ,19
120,21, 01
Asi Geos
/O1,01
giz, 61
G;€z 02
Cone
j4loee
Gfz ce
G;£2 ee
Goo,ce
|v cz
OV ez
jOV ,2z
10S ,zz
$;£0,€z
se OOIXe, “JOGIe_T ZNID e19aA
oe ‘
JoqIepY ZNID eIIA Yo ‘oxo
ti teee teen ee eeeeees “Og
.
.
*ssgyooduiey Jo Jpn
"**-9ypeduiey jo Avg
‘***purysT SoqoT YO
‘oorxeyy ‘oordwey yo
reeteees s0q
.
sree 10g
‘aseIOyoUY jJooy ueIOL[y
eee eeee yueg aysodure7)
“s*yueg euleyeg j}eaIn
"o** TQUURYD Ue}EON A
5 LL eUUEYD SSe Oro rn
reeeee s0q
VOL. 139
MISCELLANEOUS COLLECTIONS
SMITHSONIAN
50
ust
uo
udt
u8
OI
a“
cI
é
‘a's
Tenpy
c1>
o>
S1>
Iy-S:1>
oe
cI>
6
IOI
1'O1
II
£°O1
ZO1
Gri
gv
£92
Lie
z61
‘a's
“AIND a
4-9
II-O1
PI-zI
II-Ol
onood
z—-0
oo
€z
I€
I¢
Ze
ze
ce
oor
z6
08
GZ
89
Suipeas
“OIpATT
q-0
q-0
q-o
d
be
(81
jel
9
oO
1
inv
9
ve
81-0
mdoq
ofiz
ofoz
of61
O£g0
CIzI
ofol
o£60
o£Zo
Orel
oozI
Oori
out L
Ly/ez/9 81,985,524 ,08,2¢,6€ °
LZv/€zZ/9 ,,81,95,S4 ,,02,22,6€ -
4b/€z/9 ,Vz,20,94 ,9€,22,0€ °
L£v/S/¥ ,,£8,90,94 Sizz,6€ -
Ly /92z/S ,,€S,90,92 sew sOo.
L4V/S/v ,VS,9S,S2 ,,9€,92,6€ °
“IOAN Seijesseg ‘aseioyouy jwIog Aieuripio
wmeceoresiveeeee "IMO AOIN) “IOATY “Sesyesses
sisssesaeesiscsleususcn Kar oyeodesoyy uiog []2MoTT
steteesssKeq ayeadesayy ‘julog [JaMoF YO
neat iene eae ODUCT sp104q HO
Ly /¥/¥ ive, iQf,0£ . Biajess Wie sie tipi see SF is Cale Caos ‘Q138eD MON
ONTNdS “a
0S /o1/€ uf ,¥Z,%6 PIE RORQL ae ee ieee cane SOM EET OIKS IT ONT
oS /11/€ 00,56 SG TROL (ess
a7eq CM) CN)
apnyisuoT apnyyey
eles sre. eKeie oN Tenis ese eeeeene hc SUD OGUIES) jo yny
uorjd1419saq
ee
(papnj2u09) MALNIM “V
(panuizuor)—I Alav J,
UOT}e}S JO UOI}LIOT
5!
WILLIAMS, JOHNSON, DYER
IO WATER TRANSPARENCY
NO.
Ze
v
ONNN THRO NMOWMNO MMM MM MONWMO MNMMNHYHNTNOM + Ww
8
ey
9
81
ZI
9
81
ZI
4
C1iz
S10z
S161
SIgI
c1Z1
SIQI
o£g0
L£y/Lze/S
Ly/Lz/S
Ly /Lze/S
L£b/92/S
(panuijuos)
u91,85,S2 07,27 ,0F Oee ereie\s eo nuchslelaidys ekelauahuliste: alalnleinis ‘og
aL Sie GZ, 02 ,27,0€ CO e ee ee ere renee eererneerersee ‘od
pal Otol 0S ,27,,0£ heroine lair aekeis eile, ieirs) wife) caiielie Pita OG
181,935,954
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
52
cl
““
u8
‘a's
[enjoy
gS
4
cc
See
ce
cae
Si
CT
cI>
o>
ca
S1>
ciI>
TTS
>
cr >
>
o>
c1>
oT
ciI>
Gg
cs
oS
as
“AIND|
TMNMMODADODADODADOODODOOCCOCOOC OR HHH HUN HMH
supers
‘OIpAPT
qideq
oorl
OO£I
oozI
Oorl
CIe1
S1iz
aury,
6r/61/¥
6b/61/¢
6¢/61/P
6r/61/¥
LV /€z/9
Ly /Lz/S
aieq
AW Ka et4 ycv 41 ,0€ CC ‘0g
Fase e, ucV 41 ,0€ eee eee eee ee eee eee essere eeeeessees ‘oq
PWG uc¥ 41,08 See e eee ee eee eee eee ee eeeeeeeesoes ‘od
PASTecese {VaDeOe. oo clini TOAT TT STE MEGE IU sEoOr ais
40,94 uQl CZ, 08 Ce ee “keg ayeodesay) ‘\uiog []2moyy
uSl,QSoS4 ,,0%,27,0€ “°° *** **TOATY seapesseg ‘asesoyouy julog AreuIpiO
CA) CN) uondrsosaq
apnzyizuoy aepnjzyey
(a
UOIL}S JO WOTPLOOrY
(panuyuod) ONIAdS “A
(panuijuod)—I Alav
53
WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
IO
NO.
a“
a“
a
FI
VI
CI
cl
ot
I'v
Iv
I'v
Iv
I'v
Iv
Cs
ce
cI>
Ce
se
se
ce
coe
ce
ce
S-¢
cI1>
I'v
I'v
MOANNANANAANMYTOH HH OOH HOTDOONNAANANAMNM
o£
ve
ist
cl
00gI
Cozi
CogI
ooSt
0+ /01/¥
6¢/61/7
64 /61/¥7
6b/61/¢
(panuyuos)
gS €z,SZ
UE
)SS,879SL
)SS,8ZS4
)SS,8Z SL
neV 41 ,0£ fe) oli6\1e) 6 (etruus <ave eo) else] e 11 shes wensisie® ivellvireiie ° ‘og
ycv 41 ,0€ Sieveie) elie! olivisisisl 2veye ©1610 00600 © ce 0 0 se) ee ‘oq
VL Oe eie:e//alevee) stele! see] (elersie see eee e eee eens . ‘od
pV al OF Sem ww ee wm ee eee eee wee ewe sere esens ‘oq
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
54
MODONNNHOCOCOCOCOOCOOCOCON_NMMO0OC CO
sulpear
‘o1pAH
qidoq
00Z2
OO1z
0002
Ooo!
OOgI
amity,
6+/61/P
67/61/¥
6/61/7
6¢/61/7
6r/61/7
a7eq
ad eeoad u@V 41,08 COP ee eee meee ener eeee rr ervesseses ‘og
waneeon’ ye LI ,6F Coe eee erro reer eeereeeeeresrerneseeee ‘od
BOG SoAGe FeV EL Oe eee ee eee eee ese ree eee ereeeoseseses ‘oq
Wasueceae VgGre0e ee ee ee ‘oq
Pospeoaae yeV 61, 0€ eeeereceeeeeesease neta TOATNT dIeMelaq qysrT uyof drys
CM) CN) uondisoseg
apNnzsSuoeT epnziyey
a)
UWOr}e}S JO wWores0r]
(panuyuod) ONIAdS “a
(panuijuod)—I ATaV I,
55
WILLIAMS, JOHNSON, DYER
WATER TRANSPARENCY
IO
NO.
ie
99
Z01
l'OI
£83
89
Git
£-o1
g4
gZ
Ait
£3
1g
1g
CIr
IQ
88
ci
c1>
c1>
ciI>
Ship
ca
or
ooooo$o
0060
00go0
0090
oori
Sv60
$160
S1g0
00go
Ly /vz/9
L£y/bvz/9
6¢/3/S
£¢/61/¥
(panuyjuor)
07,94 ,0V,40,0€ TO eee eee em Hee eee Hee wee HEHE OS ‘og
j07,94 ,0V,40,0€ Seem meme ee eee eee eee eee eereeeneeeeon ‘og
07,94 ,0V,L0,6€ Se Heh aL at OL 1 ayeodesay quiog ueMS
Waseeeae nev 41 ,0€ a) @:.0;10)/0) 9:00 (6:0) B ©.0 a) #) 6/6) 8.60! 610) @ 8) 006),e) 0) eee a ‘oq
88
9°9
VzI
II
VOL. 139
tes
9
18
co
v ZO
Sol
5:6
& 38
G's
99
6°01
ft 34
‘as ‘a's
jenpy “andy
SMITHSONIAN MISCELLANEOUS COLLECTIONS
=
cD
oO
56
cw
07
OL
cr
Ze
6z
v~
OI
o£
Zz
vz
of
S
fl
If
€z
ez
OL
gl
ee
gz
zz
¢
OL
of
£z
oI
suipesr
‘OIpATT
9
0
vz
4
wt
‘cl
9
oO
Vz
‘
it
jel
9
O
Ve
é
81
cl
9
0
he
81
zl
9
0
ve
81
zl
9
oO
yadaq
oOoSI
Oori
OO¢ I
oozI
oor
OOOT
ouuty,
Ly /¥z/9 j07 92 ,0V ,40,0€ CC ‘og
Ly /¥z/9 407,94 OF ,40,0€ See eee ee eee eee rere eee eeseeeeee enone ‘og
L£y/vz/9 10Z 604 2 OV ZOSOE ©" a2 ees teins aess +e? = Keq oseodussys) quod ueMS
ayeq CM) CN) uordrsosaq
epnzisu0'T epnyyey
ee lll nn SF
wo01jze}S JO UONBIOT]
(panwyuor) ONTAdS “a
(panwiyuod)—'1l ATAV L,
57
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
FI
ie
: Ro CORO
1N In 1n tt NOD NO
Zo1
£1
S83
99
SSI
OIL
z0
Sg
9°9
z6
gr
6°01
mANANOTTTOMAWMN
eH Ne
of
ve
I
ei
9
‘
Vz
31
cl
9
‘
i9£
of
vz
2!
ZI
é
ve
81
ZI
9
bz
81
ZI
9
vz
81
ZI
SIzI
O£II
Ofo1
ooZI
0091
£b/L/¢
Ly /gz/S
Lb/1/9
Ly /vz/9
(panuyuos )
07,94
07.94
402,94
{07,92
,0V ,40,0€ er ‘0g
OV ,Z0,6€ Ce ‘oq
,0V ,L0,0€ Ce ey ‘oq
,0V ,Z0,0€ See ew me eee Oe wee ewe ee mew ewe we eee Oe ‘og
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
"a ‘Ss
Tenypy
‘a's
“AInDY
Oz
vz
cz
Oz
Oz
Oc
O72
O72
Oc
S¢
oo
ono a9
0
suipeot
‘O1pAHT
81
zi
9
qdaq
OOIz
0002
Ooo!
OfVI
SrZo
orzl
ofzi
oumry,
£b/32/S 9;Sz,94 1;S0,6€ See em eee wr eee eee ee ee eer esreseseseses ‘oq
£b/62/S 9)$z,92 1;S0,6£
"otesssss-a8eroyUYy qniD YyNoZ ‘purysy uosqry
LV /S/V {07,94 ,0V,40,6€ Cee eee eee ee eee eer eee ee eeeseeeeeses ‘oq
L£b/2/V B64 ZOV,;4O502 “0 < ott c cect hoses * Kee oxpodesoyy GUIOg UeMC
ayeq CM) CN) wordrs9saq
apnysuoyT apnyyeqT
—_—. TFT hv
(panuyuod) ONIAdS “A
(panu1juo2)— AAV,
UO0T}E}S JO WOT}LIO'T
59
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
5
tn
SO
g'6
II
ell
gol
Sol
gO
ve
VzI
SLI
Cer
ge
SS-oS
{51-0
OOoIl
O@ZI
gboz
gr61
QrZ1
gro1
oost
6¢/oz/P
Lb/6z/S
L£b/€z/9
Lr /£z/9
Lh /92/S
(panunuos )
{90,$2 SVG EQe. oso 2 tieie sissies oier se TOAT OAV MELOG] = [EONS UMOlE
pOealoGe ,VE,LS.9£ °° oot soso Og AID) AIIIOIN ‘Avg uI0ISea
20,97,94 yOe £0,608 Cee meee eee eee eee eee eee ee eeeee ‘og
u20,92,0L ,,08,£0,0€ *-°seso°*°ess**e"""amtog uUlejunOyY “Joany Ayjoseyy
$0;92,94 j80,6€ **"****(Aeg puruy) Aeg oxyeadesayy ‘pueysy uosqis
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
Sol
1 g°6
4
ZO01
Sol
LOI
+0
o1S CIl
a“ ‘
‘a's ‘a's
Jenpy “AINDT
be
Lz
6z
gE
Ze
o£
LG
ZI
9c
Sz
of
ve
ce
£¢
of
ve
£z
OV
SI
0z
IZ
cz
ce
£¢
ce
ce
ov
Surpes1
‘o1pAHT
81
zl
9
oO oor 6b /oz/V (90,54 SbS gf Ce ‘od
a
jv
it
j0£
vz
81
jel
9
oO Oo£t 6P/0z/V 190,94 Gib .gf eee eee eee eee terns ee ‘0g
a
jv
it
joe
ive
81
ja
9
oO ooziI 6b /oz/V {90,S2 GGG. = 72 rs teen ncee tne ee T9ATNT QIEMETOGe [POUS UMOITE
yideq = wT, aed CM) CN) wor;d1s9s9q
apnyisu0T opnyyeyT
nn ss
U0I}L}S JO UOIJEIO'T
(panuijzu0r) YNTAdS “A
(panuijuod)— I ATav J,
61
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
=n
wm
49
ve
ee
eis
00g
ooZI
0091
Oost
ob /oz/¥
6b/oz/P
6/02 /V
6b /oz/V
(panuiuor)
{90,84
(9084
190,84
190.84
SiS gf Ce ‘og
S;rS gf eeeceeresr ere eerste esses eeeoeseeeeeese ‘oq
GS-5 gf Seem meme rem e wet e eer ree eer eraeraee ‘od
S7rS gf Ce od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
62
u9 £1
‘as
jenpy
O7zI
Qe
S€1
SEI
£-S1
Vil
II
OI
S11
O'7ZI
VI
o:Sr
Ligh
€-Z1
LL1
gi
OIL
OIL
S11
C11
Sit
CCI
SEI
S°3
88
S-6
601
‘as
“AINDY
cz
gz
gf
SuIpeot
‘oIpAH
i9t
joe
jz
81
yl
9
oO
a
ev
if
joe
yidoq
ohhr § L¥/9z/9 82,94 (OS g¢ ‘‘Aeg ayeodesayy Jysry wiog Apoojg jo "AA aiIwW I
OOIIT 6v/zez/v 4 FLSRQo BO SRO EEE NO IEIO OOO Koh gta yeoys SITIDQIIN
Syor Zy/1f/S Sg'S1 92. -GS:2S.9F “eccs*sessscccersss Keg oyvadesal) GUIOg URIS].
oogI 6¢ /oz/t 190,$4 SVS .9f eee ceria ee aie cies She TOT AT dIVMLIC eoys uUMOIg
omar L a7eq CM) CN) woryd1i0soq
epnyisuoyT epniniey
|
UOl}E}S JO UOTEIOT
(ponuiju0d) ONTAdS “a
(paniijzu02)— 1 aAldv [,
63
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
u9 01
OI
LI
LE
bgt
CII
CET
VI
S‘h1
S-rI
Shr
oz
UiS/AN
St
g°SI
TI
Tsoi
gti
Sf
Ql
O11
ZI
£21
Cer
CVI
II
Il
601
9"
£8
QI
ZI
evi
-
“©
99
ov
oS
es
rs
+S
vS
Z
ZO
0S
G°ZS
zS
gS
ss
oS
civ
cv
ZO
£9
oS
vs
Ze
Ze
gf
ae
IZ
59
<9
£s
oool
¢S60
Oorl
oozI
6V/zz/P
£v/1€/S
LV /62/S
Lb /1€/S
(panuijuos)
S:LbbZ
S:21,9Z
82,94
Se,0
9;gb gf Recses Vic's Saber TeO TS SIDI ek ‘AS So]IU O'~
Z6P gf TS ee ecg eae Le AE | ayeadesoyy) ‘FOATT oA MA
OS 8e Seer ere eeereeeeneeeeseee ‘od
OS .8° Coe e rece eeesereseeeesees ‘oq
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
64
-~
IY
io
8
‘ce
jenjoy
SV 10
gv zo
14 16
cv 06
O71 LY
SIGE oS
eI os
Gel oS
Sor os
Gio os
gel 1S
gel 1S
Ocl LY
SOI ce
col Ze
VzI cv
gr aa
ge QZ
£97 08
£gz zg
£°9% z
S62 £8
S62 £8
S62 £Q
S62 £8
bgt 99
‘as SuIpeo1
“AInND “o1pAHL
joe
ve
81
wwdeq
Srr1 1S/S1/S ,0V,SE,¢4 iBV.8e eee ecg coir aS WSrT WoYyey-IALT
o€zl 6b /zz/V G;10,$4 QV o8E spesPeteteseseie) cheval shelenensensy ciel su sehe ica nere TITUS JST S][eJ19AO
oool 6F/zz/r G;Z4V v2 ORO ARE ee eres Sica ee "***TROYS SINAN “A X “AS Solu gre
out y, a1eq CM) CN) uodr11osaq
opnysuoT epnyryeT
ee
UOI}E}S JO UOT}LIO’T
(panuyuor) ONIAdS “a
(panuyuod)—1 ATav I,
65
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
wl 81
FI
Qvi
Lz1
£1
£1
fv
Lzi
Lien
OcI
35
S:ZE
Sof
£°9z
£92
ee
gbe
ove
oz
ez
be
ale
ov
cv
0
(panuyuos)
ofol
of£60
ofgo Lr /o€/S WVZ,21,,9L URL LY gf SH oF osc lnia oielseare Neko ce ene TELE TAT 4S jo AJIT A,
o£Zo 6v/zz/¥ ,0V,SE VL BV 8e ee eee eee ee ee eee ee eeeeeeeeeoeeeeeeeene ‘od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
«991
‘a's
yenpy
gSe
g°S¢
See
ze
ze
£92
£92
£92
Vzi
£1
£1
Ger
OIL
611
zi
VZI
Zit
SII
SII
£71
Conn
“11
“11
€-7c1
£1
Ger
‘a's
“ainby
£8
48
98
58
58
£3
£8
£8
oY
gv
gv
oS
cv
ZV
ev
cV
ge
ov
ov
by
ov
Iv
Iv
th
gv
oS
Surpeat
‘o1pAH
i9f
o£
Vz
81
zi
4
9
oO
a
eI
yydeq
S730
ofr1
Ofer
oozI
O£II
ofor
omy,
6b/z2z/V yOV Ve APSE coos dys JST woyyey-aany "MA sow V
Ly /of/S uVZ,2Z1,94 u81,4P .8F CeCe eee eee seeeeeseeseeeseeeeeeeeeeseese ‘od
Lv /o£/S uVE,21,94 u81,4P .8& eee eee eee eee ee ee se seeeseeeesees sees esrese 0d
Lb /oe/S uVZ,21,94 ARLE RE aratarefainrevatererererstesereretel acim: “Mase SOPMOU 1S Jo AJUIOT A,
aed CAN) CN) uoryd11989q
apnyisuoyT epnyjzeT
: !
UOT}E}S JO UOIWBROO'T
(panuyuor) ONIYdS “A
(panuyuor)— 1 AIavV
67
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
9
81
S-Ze
Ofor
6¢/3/S
of60 Lr/Lz/9
OIOI
Ly /€z/v
(panuryuos )
“4SZ.9L VigE,g& occ °oc seo sess Keg oyvodesey.) ‘purlsy sdieys yO
0° 01,94 ue¥ IV3f 29:0) 010: (0¢ 8 Sie 9/0) 00:0 06,0: 0 s)e:0)0\0 0 60/0 0 lejeite 0.8 o)ele PIN ‘pIOJXO
5" eV oRe rene Gus quay SiTej4oa) aS somun. eon
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
68
i218
6,6
a“
Or
“
u8
ae
jenqoy
GZx
OzI
Ger
Cr
+6
G°f1
VI
Vzi
OIL
eI
£1
Sn
Ger
Cri
C1
ci
I>
ci
Cr
c1>
I>
ciI>
cr
C1
I>
iis:
*amnby
OvOvOuo Onl o) soilG lc)
°
suIpeot
‘o1pATT
81
j21
9
o
q-0
dog
o£go 6¢/2/S 07,92 O1,8f Coe ee eee Ooo eee HHH EEE HEHEHE EEE EEE EE HEED ‘od
o¢Zo 6v/L/S 07,94 OI 8e PSaoia Sia gisloisieieie ile esele alse RSs eee SOOT (WTA) I?
Oort 6b/32/P 80,22 ¥Z,88 COCO HET ET EH HE HHO EE HHO EES ‘od
0040 = 6¥/g/S 20,42 WWZQE corre OANY SeMOJOg 4Ulog Jepsg JoddQ YO
0091 1$/S/S {80,22 Vz ,38 Ce eeeoeeeseeeoeeeeeeseeeeeeeeseeeses JOANT deuI0}0g
omty 31eq CM) CN) uorqdt1osaq
apnysuoT apnyyeyT
a a nt
UOE}S JO UOTLOOT
(panu1zu09) YNIAdS “a
(panuyjuor)—1l Fav,
69
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
a“
a“
6,6
3,6
v6
SEI
gti
Oz
S06
OI
OIL
ger
SC1
CPI
SEI
Gol
£1
VI
ZL1
ol
6°01
Cau
C'1r
Sai
VI
SEI
O'ZI1
Vcl
Cer
ISI
£O1
Cer
Zit
GIL
O7ZI
O'zI
oS
gs
o£
gz
of
A
1S
oS
vs
oS
6v
zs
b9
of
of
ov
by
zs
oS
LV
cr
os
gS
QZ
ze
oS
ov
Ly
Ly
OfIl
O£ol
o£60
6r/2/S
6F/2/S
6b/2/S
(paniyjuos )
(0% 994
07.92
0294
/OL,8f erat elevacelevlereleXers (el ciekel eel ere) ee: sve eieieisrer ee eauienene) tier are ‘od
OI 8e Cec ceceeccerse reese eseeseeseseresssesess ‘og
OI ,8e es ieleieieleiele 66 ee 6°08 0/6 6 8070/6 ele eles) 06) 68041006) e 107m elelere ‘oq
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
7O
a“ £6
‘as
Tenpy
GiSt
g rl
“G1
gr
£1
£1
gi
ZI
go
Z01
Vz
oO.
I'S1
S:S1
Qh
ge
Ser
“S1
6:0z
883
38
O'1I
v1
Q eI
VI
VI
SEX
‘as
“ainda
ET
1S
rs
Zs
oS
superar
‘o1pAH
it
joe
ibe
81
qidoq
Off
O£ZI1
Of1I
ouny,
6b/Z2/S 0% ,92 {OI ,8f PCO mee eee ee eee eerneresrerseeeseeeseessesese ‘0g
6b/2/S {02,94 jO13f cone eee eee eee eee ©), (0.6 0s(0) 0) 6.074\\e-10 0 0/16) 0a\ enue 608.08 ‘od
6¢/2/S {02,94 /O1,8f ens a ee ee OE (QUT) IP
aqeg (MA) CN) uoljdtaz0saq
apnysuo0 Ty epnyyey
OO
UOl}e}S JO UOT}LIOT
(panuyuod) ONIAdS “a
(panuijuor)—I aI1av J,
re
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
Og
«8.8
£3
88
39
£o1
ol
ZO1
Ser
9
of
v~
81
ZI
4
$160
OOIT
S1go
Oo£T
OfrI
6+ /3z/P
67/9/S
67/3z/¥
6r/€z2/P
6b/4/S
(panuyuoz)
a“ or,rr, 94
uO Vr, gZ
ev, 94
8)z0,$2
102.94
S'II,9f ee er ‘od
SHII,BE vores TOATY DeWMOJOg ‘puesy sUuO\sxPLTG YO
jVIgt pei hi SS SOOO A CYNE | deuIo}Og ‘keq uopIg
L1,9£ (Sele sieieisieleiviee see TBOUS SIMEaT BO ANG OAS AA
OI ,8e Seem eee Here ee oe eeeen see eeeseeseeeeeesesens ‘od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
72
Ie
in
-
9
8
a's
npy
‘a's
*arnby
Fe
gi
suipear
‘O1pATT
81
jz
gidoq
Cc1Z1
oeSt
Siri
¢160
out
6b/Ez/b
6b /2£z/V
6¢/9/5
6b /gz/V
a3eq
$°S0,S4 yoaege Mab crate seers ish tie sese ee eps ee Ol ae (SOM) Song apasiy AA,
(1Z,94 {10,88 Cr ee ‘od
12,92 {10,88 Binesioheicreiereneie ieee cicie enhance see ei eheceeensiccea a MGT) «ds» Ia
yOV bb 94 CII,Qe “oct OAR DeWIO}JO ‘puR]syT U0}syoR[G YO
CM) CN) uoiydr10saq
apnyisu0T apnyyzeyT
——————————————————————————— nnn
UOTZRYS JO UOIZLIOT
(pamuyu0d) ONIAdS “A
(panuyuor)—1 AIaVv IL,
73
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
ZI
nll ,G
ZI
Vz1
O'ZI
Ge1
£1
eO1
Vit
SII
GIr
gl
gI
ZI
gI
Sg
SO
£01
VI
SV
z6
oI
PQ
71
LL1
SIr
S61
SOI
bgt
bgt
0z
oz
SV
Lv
o
os
29
09
o£
ov
ov
59
29
oz
£z
gz
ze
zs
¢S
ve
€z
o£
99
9
ov
oZ
69
99
oZ
oZ
OOIl
Ooo!
S1g0
ofo1
S1Zo
6y/9z/¥V
6¢/9z/¥
6r/Lz/F
6b/Lz/b
6b/be/P
(panuijudd )
,0V 00,94
w0¥ 60, gf
(22,94
01,94
OreGZ
/8£,50,2E
/S£,50,2E
Oe Ligce
[8S ,LE
3 ,4E
BNET OI RO eeNe Oar eierer eo AeA eae areca) (6
“s+ Keg ayevadesoyy ‘spunois d[ppiyj, soyses1oFy
sis seis cite OD NETS) NO ‘IOALQT YIOKR YHOPT
vrreeeeees Keg ayeadesoyy) ‘ysry dery JOA, HO
"y2[UT onseajoouryy YO “TLZ (MTA) 4ong IPA
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
74
%o
é
‘a's
jenjoy
CIl
Vzl
V'ZI
Ser
“11
g°0
ZO1
VI
VzI
611
Vzl
ZI
Val
Vel
QI
Later
Our
SvI
Lt
ZI
OZI
SII
GIL
v1
f91
SI
£zI1
iS
*AInDY
ov
SP
cP
ov
Iv
lz
ge
vs
cv
zy
cv
ev
SV
SV
cs
Iv
zp
vs
29
29
LV
ov
ZV
gs
09
59
vy
SuIpest
‘o1pAH
81
21
yidoq
ooSI
Oori
OO£1
OozI
OOITI
aun,
6b/9z/¥ ,0¥,60,92 ,SE,S0,ZE °°
OV/Oz/T- 7, 0V 00594 7,46,50,0e —*
6¥/oz/v ,,0V,60,92 ,,S€,S0,2E °°
6V/oz/v ,,0V,60,92 ,,S€,S0,Z€ °°
Or/9z/F ,0V,00,94 ,S€,S0,4€ **
22, CM) CN)
apny}isuoT apnyyeT
ee
(panuyuor) ONIAdS “a
(panuyjuod)— I aldv
beste eseeseeeeseees cog
Lesteeeneeedenseees “ger
Lesseneeeeeseeeeees Ogg
seers a ecmin emer
‘Aeg ayeodesoyy ‘spunossy s[ppry, xoysessopy
uordr19saq
UW01ZR4S JO UOT}LIOT
/
75
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
II
II
OO!
CII
OZI1
VI
gr
G°St
ol
VZI
Ser
FI
£91
“LSI
Cael
OVI
SPI
ZS1
SAT
ZI
ZI
VzI
S-E1
£1
£91
£91
O11
ZI
Ger
SEI
£91
eg
Zit
of
vz
81
ZI
9
é
joe
be
81
ZI
9
‘
of
te
81
ZI
9
of
te
81
ZI
9
of
tz
BI
cl
9
of
4
4
0002
ooo1
o0gI
ooZI
o0gI
6b/9z/P
6b/9z/b
6h/9z/¥
6b/9z/¥
6h/9z/¥
(panuijuos)
y0V 60, g4
,0V 00,94
,0¥ 60,94
OV 60 ° gf
,0V 60,92
nee ce *
Sejs0540.
a“
SeSo,Ze °
a“
Sf,50,L£ *
4Sf,50,2E °
.
od
od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
70
SII< oo1 00
aie ool vg
€or 66 34
LL £6 4
1S £6 99
G°Ze-7'Z~ = 9Q-I8 09-,hz
Lez SZ Q1-,2Z1
See QI S9 {9-0 Crro 0S/1/¥ 80,08 Sah 92 Ss hligua) aie) Shia sieves rig oe Silene se OBIT T: tddississtjq yO
—- acre -++o01 q-0 oogo0 0S/9/¥r v:v~,S8 W106," <P cneGhseenannns=*' "Stig CES ode) 10 pIeMunOS
— S11<-gg _ OOI-g6 q-0 oogt OS/g/F Cizb,fg ZOU Ge os eee esses HOS QUEMS JO" PIVAYSO
gh 1-02 SS-oZ jOZI—,92
_ Lez SZ 134-0 oof1 0S/1/P S;€1 88 CVE OG. 2 ne ee rere citieeies sce e eee roiKOTN sO. pac Miso
— C'6c-bz £3-22 q-0 SPgI 0$/Z/V Zj01 48 QiZI OL (otter testes rer esseseesccmr ga “GrooEstag JO ‘a[S
le a —— Oo ozgI £r/9z/¥ j1Z 18 \27,08 eee ewer wee ee ee reese ee eee eeeeeeseeesee sess ‘oq
BI aad = Oo Sre1z fr /gz/P 1Z,18 EC EO eS CO RON AR et Ser oe COTE “ely Gad fey
6'o1 gf q-9
a Orr os Oo o£60 oS /£/r £1,248 bz ,0€ SEsis Saleol fakes esesea sac cac eeu eh ‘ejooesudg ‘apisyooq
Sof rg vs
ze Sg igv
"of vg ev
ze Sg oe
£-9z zg joe
£92 0g vz
ale 1g Bl
Z°lz Ig jel
eg <8 9
jee Sof vg oO Sper 6r/bz/v (GV JOO REE, 2 es een Reh Tee e Senne Gite ager coeaGuselo)
IGS ‘a's Sulpear qidoq SUahy, SEC CM) CN) mor d119saq
Tenpy “andy ‘o1pAFT apny}isu0T opnyiey,
Se eee
017238 JO UOTICIO’T
(pamujuor) ONIAdS “a
(panuijuod)—I ATdV I,
77
WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
Io
NO.
(panuyuos)
—_ 4 16 9 otri
— cv 06 Ql oir
— Sv 16 1 oozI
eae LZ LO BI 00g0 S¢/z2z/S Sa VS 9&8 yO1,S£ ve alae vavelialeleneuanensnarete cel ekerelieresevejuve.e)(ske see tells else ‘oq
19 $6 joe
ss v6 ive
ia £6 QI
1S £6 zl
ez cs +6 9 ozri
S2Z £96 1 Scer
89 90 joe
89 96 ve
89 90 Bt
tg £56 jl
= 89 90 9 —-SOEL
— Sz £16 BI €f11
—_ Q8 g0 Bl zOIl
— Sz £16 Ql Sfo1
— L£L 46 Bl OnGon: -S/2Ze/S. {SO VSeEQ: — OL SEavo mS Ver a “s*seSnyioy, Aq ‘Aoy Uepser
— S11<-zZ o01-¥96 BI opm St/oc/S {0V,Z5.¢9 {Ob Seve = see xe Sale ce ec sesnjioy, Aiq ‘juueyd “MAS
_— Gni< -- 001 81 Goan © SiV/0c/S.- 7 OS COneGe el Beek ee ***suoujyey Il ul Yue sesnzo],
81 59 82
/0 “SI 6S—ZS jyz-0 Sool 0%/z1/V S;rS 18 BOY Verret ge aa a “-ysaAQ Aasy GUST TeoyS YWUIS
ZI = == oO Zcole =Cvleciy, 4S, 20209 = ,Ol SO.0en on oe ee yeaa ee SINT aaToO GL ‘Q[epsopney 310.T
— c-¢e-ce 9g-S8 q-o C1ec 0S/11/V G:€7,28 j£1,92 o) orolske eh ssekel stoke fo) cose Nenee eaeke eee ae ULC | Ol} Jeqrues HO
— GSe-€9z 9-28 j7v-0 vito 8 0S/6/F 98 28 GE phe, eicisie ovietsinieinia.s ie viele os RONG Ba Gee Ae et eG Use U(e)
99 ol joe
v6 Sz he
11 ShI vs ‘gio Stor “OS/11/P —-“Siof.eg. - “5/446 "++" Keg orogs|[IF] Jo yynou ‘keg edwey
v6 Sz q-—vz
Z01 ae jbz—81
es Z zl {Q1-0 0060 0S /11/V 70V 97.28 G95 42 SfeKodel sxel che telexevohene leurac isco nce! habeus meer OC LEIET “ely “edwie [,
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
78
‘as
jenpy
£o1
£Oo1
cli<
Sti<
Sli<
S1i<
c1i<
SIi<
Sir<
Sli<
Sli<
Sli<
S1i<
S°S6
€o1
OII
£o1
£o1
LL
89
LL
LL
LL
ev
ov
eV
ey
‘as
“aInbay
66
OOI
OO!
OOI
OOI
OO1
OOI
-+-o0oI
-+-OOoI
-+-ooI
+001
+001
$80
66
$66
66
66
£6
96
46
£6
£6
06
6g
06
06
surpear
‘O1pATT
21
qideq
oLol
Sr60
beri
Ofer
OO£I
Szz1
OOO!
OfE1
Cozi
SIII
OI0I
Sz60
O£F1
oun,
Sv /S2/S
SP/be/S
Sv/Ez/S
Sv/zz/S
27eq
429,45,23 yOL,SE vz Ce ee eee eee s esses ereeeseseeseeeeeseses ‘oq
425,528 OL SE ve Ce ee ey ‘oq
wan VS5e8 ZOUSO Ve Peewee ere e ree reese sreereesresreseseses ‘oq
pan VaneQ OL SE abe ss cries siaielee o essieiesisie es pao AATCTS ho eOpIeS)
CM) CN) uordiiosaq
apnysuoy apnyje'T
ss
u0l}v}s JO UOT}ROO7]
(panuyuor) ONTAdS “A
(panuyu0od)—1 alav
79
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
QrI
al
z'61
1S
1S
1S
Lat
gI
oz
6:02
cs
19
Cri<
cli<
SOF
19-QV
LL
S11<-€o1
c1i<
Cri<
SsII<
SII<
cli<
cri<
ori<
Srt—
OII
c1i<
OII
OI!
£O1
ss
£1
89
£6
£6
£6
9
59
o£
ZZ
v6
[6
OOI
OOI
£26
6-26
46
oo1-66
+001
-+-oo1
+001
+oo1
+001
+001
+0oo1
-+-Oo1
£66
OOI
£606
$66
66
zoZi
ooZI
ofSt
SI€1
CrZI
ocSI
€z£1
ost
ofSt
ovet
Cecl
CCII
Sb/Sz/S
sh/gi/s
Sv/Z1/S
St/9oz/S
Sr /Sz/S
(panuijuos)
= lf le SN tree cog
JAsescee. jOLaeene 2° ss" e888 0g
GasiPaece. AOL sesve a2. re8) 0g
BSGYS co. One ves eos reese cog
u25VS 23 401 SE, be © O10 er ew Oleje erclere 6 6/0 0-8) 6) 0 6 6 016 0. esis ole oeietale ‘od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
80
‘a's
Trenpy
0z
ze
€°Z1
oz
cz
Giz
“£g1
fz
C61
SIZ
6'0z
£-fz%
Lz
Zg1
QrI
£91
£91
gl
Le
€g1
“G1
Sar
cli<
ci
Sli<
S1i<
cli<
‘as
“aInDy
-+-oo1
-++ooI
+oo1
+001
+oo1
suipeo1
‘o1pAH
ZI
9
ve
81
zl
9
ve
81
el
9
ve
81
jel
9
ve
81
el
9
vz
st
jz
9
921
tl
jz
9
q-0
qideq
4
offz
ofze
o£So
ofvo
o£€o
o£zo
ofol
ofiI
amir,
Sv/oz/S 018 ove CC 0d
Sv/12/S 018 ove Bee fore ey eepea stale eee engi cau ney olemae SAS TET SoM Aayy
SP/9gz/S ;Z0,£8 Ol ve “*'sesnyioy, Aiq ‘Avy Uspiey ‘aAInD wWoY}e,J-001
oS /€1/v Ziz€,18 1 bz SS ee eS ODEO TOO SOLE (SOUS ea GS eT AY
23eq CM) CN) uoryd110saq
epnysuoyT opnyjey
nn
U0I}e}S JO UOT}EI07T
(papmj9u02) ONTAdS “A
(panuyuor)—1 Adv],
8I
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
for
1S
St
6°02
£-€z%
z'O1
zOl
S61
6'0z
oz
zO6l
Vg
zOl
z61
v1
zO1
£c
lz
oor
S61
6°0z
S'1z
St
v'gI
6°02
SIZ
St
sor
66
£6
ZZ
QZ
89
ZZ
oZ
89
99
89
89
99
zSZ
ZSL
ZS
69
>
©
£Z
gs
cl
£2
gs
$19
9
81
ve
BI
cl
9
itz
st
cl
9
itz
BI
Zl
vz
81
ZI
9
vz
i8t
zi
ive
81
zr
te
8!
0z60
00go
OfgI
O£Z1
O£gI
ociz
ofoz
oOf61
gh/1z/g
Sh/2Z1/S
Sv/oz/S
Sv/oz/S
(panuyuogs)
(22,98 V1 pon ais ee oto. eae Te era ODE IG eal ‘keg pey
GAWWNAS “D
u9C,2V eg, OF ,FE 02 Soe cc cece 2. Joo) SSRRNIO TF ATG 1eOuG ceosagon,
018 VG Seal ohn iors Fao aia Ce ake oxanore see eee Chale ale) 6) 0) wer élele ‘oq
018 VEE cha) oXeKosaleiterek ke xeYeToxevKoxereiereVolololalexeisvelersvolenelefeler sels ‘od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
82
co
wn
AOS)
jenpy
ca v6
cy 06
z'61 89
z'61 39
z'61 99
281 £9
cs +6
cs v6
gv zo
ay 06
cv 06
a 06
ov 06
ov 06
av 06
Soe, £8
ose £8
ose 48
c-s¢ £3
S°SE 28
LL £6
88 go
88 go
88 go
88 go
£o1 66
£o1 60
‘a's surpeot
“ainby ‘o1pAH
joe
bz
81
jel
wydoq
0090 «gt/iIz/g $;Z€,98 JOLoIS crc ttre t tess puRTpuNnojMaNT GleIS eS] aJog
00g! gb /zz/g eV55 Zj0f 1S Cee reser ereesreseeessesseeeeeesesesseesee keg Joost g
cost gb/fz/g ,,0z,9z,SS jOV IS corset tess ee sess pURTpUNOFMAN, YUIOg Yue
0z60 gr /1z/3 \22,98 S518 SS SES 918 9 60S 08 00:8 C000 ea Rs eo wens ON EICE awe CaploOm
omy 27ed CAN) CN) worjdts0s3q
epnyisuoT apnyiyey
Xn EUnet
WOI}e}S JO UOT}VOOT
(panuyuor) YANWNAS “D
(panuijuor)—I ATavy,
83
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
4
3
WwW
Se Fs ¢ og
A
S°ZE
S°Ze
o°Ze
S:Ze
S°Ze
zyv
cv
88
£o1
19
89
89
19
Be
ss
SS
ss
SOABSSARRS
66
c6
96
$6
140)
140)
v6
v6
199
ofoz
OfzI
gh /oz/g
8h /9z/8
(panu1juo?)
Ol, ee 4a
Oe ,05,95
yOe LV 0S ee eee eee ee Ti PUCTPUMOT NON ‘\uiog syn
yOV,1S,08 eyaieus Si eure eve rnie oleh es eiejeieheiie/4-S*>- DIel DUNOL AS NI ‘xo adey
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
84
iv
OI
S-Se
Sof
vz
SOI
QI
C21
suipeot
‘O1pAPT
yidaq
of60 gb/Zz2/g sz 11, 9S
ofg0 gb/2Zz/g 81,98 ;IE,OG cosets ssereeeocecscs* NueDUNOTMON “IOqIeFy syono.7
of£oz gb /oz/3 je ceees 0¢,£V,0S eee eee e eee eee ee ee ese ese *puelpunofMoa Ny uiog duo
oury, 27eq CM) CN) uorjdtsosaq
opnjyisu0T opnyyey
nn
U0I}2}S JO UOIZBIO'T
(panuyuor) YAWWNS ‘OD
(panuiyjuod)—I aIav
85
cy
c-Ze
c-Ze
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
0°
=A
oO¢St
crzI
gr/Lz/3
gr/2Zz/g
(panuyjuos)
oR
Lv.ss
OS Ob crrtttt teeter terete ees
eee eee eee
~ PBeIst TED
KAO] ec te To uueU ‘purys] oqieg aS
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
86
~
°o
wn
+,08
ides
jenpy
‘a's
*aiInby
superar
‘O1pAH
oO OOIL gF/6z/3
199
vs
8Y
ie
9
joe
bz
Bt
el
9
0 Ofol gr7/gz/Z
84
124
99
199
rs
ev
iE
joe
ve
81
zl
9
0 OLII gh/gz/g
184
jel ofS1 gbr/Zz/g
qideq awry a1eq
s8VeS ie,ae OV BO Pe Sie eg a ene es a DUE a weypeM JIVO
ov vs S;ov ,6V CC e eee eee ese s sees eseeeseseeseses Joqiepy dESUITIIM T
{Ol OS."s OV 8V.OY * tet Sess “DH UeIDUNOTAON dol] NOnpeoOre
j2e555 {0S OV SES eape ah SS Satie wie: Sie 8, setae soa sere ee DUIS eS)
CM) CN) uolydiiosaq
epnyisuoyT apnyijzey
Os OS Oo
U01}e4S JO UOI}LIOT
(panuyuor) YAWWNAS ‘D
(panwjzu0d)—1 aTavy,
87
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
%
S-gS
S°gS
62
0g
Ig
18
18
£6
z6
10
zo
z6
£6
zO
zO
z0
zo
£6
£6
AQ}
16
16
06
10
16
16
16
zO
40)
£6
140)
+6
09
orgi
OorI
gh/8z/3
gr/6z/g
(panunuo?)
tvs
julaed
uS¥,SE,6 FOO I GOD SIO Ca OSGI foe Fey Agq-awio)-wopyjas IT
2,0b “orseeesse sees puRIsy YUNY Jo °S salut o1 *xoiddy
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
88
of
89
89
89
89
£L
LL
89
19
19
gr
eV
S-ee
96
96
)
96
£6
£6
96
S06
So
zO
006
98
28
120)
140)
£6
zO
zo
ZO
ZO
06
06
06
63
06
06
68
sulpest
‘o1pAH
ops
wt
9
oO
184
te,
199
499
Ws
i8Y
jv
i9e
joe
ibe
31
ra
9
0
deg
00g0
C11z
onsty,
gv/0z/g ,,0€ 90,8
gr/41/g 142085
aed CM) CN)
apnjyisuoT epnyyeTy
OO, hv ae
(panuyuo9) YAWWNAS ‘O
(panuyuod)—I alav L,
10,07 Dies Se ie cee eS DUE TPUNOTMON ‘WIV Jaquinyy
40,00 ******"******puR[punOJMAEN ‘“YSrT peep wog opwWyT
wor}d119s9qq
uo1ze4s JO UOTILIO'T
89
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
Le
oN
NN
SAFSARABSARASAASAASASER
2° aaaa
SSBRRRS8SSRS5
$1Zo
o£60
ofoz
gr/L1/3
8b/2£1/3
gr /6z/3
(panuyuos)
ple Qos Oster nbesVeTav 0d ae *puelpunojmaNn ‘1oqieyyZ onbseg xny 0g
\22,08 yov ve LV Ce rr ay * pue]punojyma Ny ‘Key ede)
408 OVER Bee Cane "ree eeesss*DUeIDUNOTMIN ‘ejsTAeUOg ode
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
go
S°ZE 88 jz
aZé 88 9
jee or 6g 0 cesr = gb/v/6 $;6S 2S EIQE OV. Botts Oe eiss serene sites UE DUNOPMONT “SObN oUe>)
88 g6 184
88 go 124
88 g6 199
ZL 46 409
69 90 is
69 96 i
LL £6 iv
88 gO sgt
88 go jot
88 gO ive
42 £6 81
£L £6 jel
LL £6 9
29 19 S6 0 0060 = gi //6 GOS ca OS, Cec cr =o a ciie *neleisicis Onin S* 2 DUE PUNOLMON, -aeoaG Ges)
8t 59 da
v°L1 £9 BI
gI $9 jz1
“gi £9 9
SI gl $9 oO OfII GV /£/G= leer es: CV Ec Lh rice: sins OUR PUNOIMON JOGIEET Ss una, 3S
LYvz gZ qd
“bz gZ joe
Lvz gZ ve
Lvz 92 8! S1Zo gh/Z1/g ,,1£,90,65 ,ZE,vE,Zb -*°*°°°°*****puelpunoymoan ‘JoqieZy onbseg xny 10g
‘a's ‘a's Bulpear yidoq = ow, ered CM) CN) uorydr319saq
jenpy “ainda *O1pATT epnisu0T apnyizey
ee
U0I}24S JO UOT}LIOT
(panuyuor) YAWWAS ‘D
(panuyuor)— 1 AAV L
gI
NO. IO WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
SSF
gZe
ze
ze
gzet
gze
9
oO
184
4
199
199
WS
BY
jer
if
joe
vz
‘
81
eI
9
O
ibe
Bl
jal
9
Oo
o£60
CizI
ofgi
(panuiuos)
gb/€1/g ,,SP,08,09 ,S1,0S,Sh “ttt tte eMOIg PAON ‘OxeT IOP sesg
gr/£1/3 ,1V,09 0° ,50,9V DID OOO RAED D OOO OOD OL ORS BAON ‘I0,P selg
Qr/SI/9. .j20\7s09: — 16,90, 9 <2 2° or So ee EOS EAON ‘roqiepZ AaupAS
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
92
-
co
WwW
Te
61
a's
npy
a
zy
zy
gv
gv
19
gr
cy
g'6£
ze
£-9z
€-9z
Z'Qz
Z'Qz
Lez
LL
19
QV
g'6£
S:Se
SEE,
See
ze
Sof
BEES
Sie
see
‘as
“arnby
00
06
06
zO
z6
S6
zo
06
6g
58
08
08
£8
38
SZ
46
$6
ZO
63
48
98
98
58
bg
98
98
98
Surlpesl
‘o1pAH
199
iol
yideq
O¢ri gb /S/6 jee aa ve Sv SERRE SS SESSA S OPER ESS SS Ep aI yules
OLE §h/zI/g ,SV,25,09 ,OF,SE,SV ****°°*ss°*ss*"s"""-BnoIG AON “sjeoyS pealesioyy
0860 9f/f1/s ,S¥;0S,090 {S1,0S,Sh “°° °°" tomes enOOGoRAON “SNe IO .p Sstrq.
omy, a3eq CM) CN) uoijdi19s9q
apnyisuoyT spniyieyT
a ee)
UO1}e4S JO UOT}LIO'T
(panuyuor) YAWWAS “O
(panuijuor)— 1 alav
93
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
81
if
g'6£
OfII
Srri
gb/11/8
gr/z1/3
(panuyuos)
y01,£V,19
190,15 ,09
85,01, sisie oi vieleinisis via sien seine s.> aaa BAN ‘Joqieyy Axyunoy
440,12 ,SP sire Sibi (areseli ciske,evekesciete store eerBTA OO ME AON ‘osue) ade)
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
94
199
ee,
GES)
yenyoy
cv
L6€
L6£
cv
cr
0z
Lbe
£°6z
Ltz
LYve
‘as
“aInbD ay
S6
$6
S6
v6
+6
+6
v6
g6
£6
140)
140)
£6
£6
+6
ZO
£6
06
06
6g
68
10
16
o£
gZ
£g
gZ
94
Surpesr
‘O1pATT
oO
qideq
Ozol
C161
ofZI
amt
gr/or/g
gr/z1/8
gr/o1/g
33eq
u©5 82 <9 each We wisiate, ee vexercrsisversigscae chre eres B TI OOS eAON ‘oes doyue x
ueV,2£,19 Z0,Sh *oseree renee teserecnsssoqsery ATUNOD Yo Aon "ees
WOLSP,I9 ,,SS,01, Sh “crest ttt tt ttt eOIg BAON “TOqreTy AsjUNOD
CM) CN) uo1jdrsosaq,
apnyisuoy apnyijzey
eee ee ee
UW01}L4S JO UOI}PEIO’T
(panuyuor) YAWWAS ‘D
(panuijuor)—1 AAV,
95
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
92
Sof
cS-ee
ZV
88
88
LL
LL
89
so
£6
“©
10
10
z6
z6
oor
oor
ool
oo!
oor
Oo!
Ss!
(panuiyuos)
00g0 gr/o1/g w2¥ OV 29 pO Se ovP eeoeeceeeneeeonese eres eee ee ece *B1}09S PAON ‘queg uyof
OzOI —- V/O/B_,BIPEE9 ZOOE,PY “reser rere esses HD0q [IO Aq soqueHT xepyEY
oozI gb/6/g uVZ,8E £9 u9°,1¥, bP Ce oerereeeererecresecscecesen *xeylyepy ‘ulseg plojpog
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
Id
98
‘a's
jenpy
Lz
£92
£62
cee
S61
S61
S02
Ze
882
88
LiL
LL
19
19
19
19
19
19
19
LL
LG,
ZL
LL
88
88
89
89
‘as
“andy
a4
08
£8
98
69
69
1Z
vZ
0g
86
£0
£6
so
so
So
so
so
s6
s6
£6
£6
£6
£6
36
g6
96
96
suipear
“oIpAH
wi
ior
9
O
Vz
St
jel
9
O
184
jo
99
09
vs
iB
ev
9
of
iva
31
jz
9
Oo
84
24
199
199
yideq
é
S1z1
ofol
o£90
00g0
amt
Bb/9z/L ,Z1,08,89 ,0Z,R1, PY “sees t tts KOG IIH onfg ‘puejsy Buoy YO
gb /Le/Z uo¥ o£ 89 40° Vz, bv Spo Seeks ae Misuse ae cee ee COULLE TN ‘roqiey IH ond
gb/6/g 02,08 £9 ,Qb,If,Vh -* ccc eIONG BAON “sJOqIePZ xepyeP ooues1qUA
gr/o1/g uo¥ OV,29 yOl Sev ee eee eee eee eee eee ee eeeeeees *B1J09S eAON ‘queg uyof
aq (A) CN) wordi19saq
opnyisu0T opnyzyze'y
nN ——
UO0}}E4S JO UOT}LI07T
(panuyuor) AANWAS ‘DO
(panuijuor)—I ATaV L,
97
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
Al
4
IZ
cz
cz
Lez
2
Sz
6'0e
S61
61
S61
Zz
be
Lbze
£-9z
£91
gI
VI
VI
Ze
G-ZE
ze
ce
Lbe
Lrze
Ltz
E22
2
S1z
ce
S-0z
QI
€g1
Cir1
OOo!
gbh/5/8
gb /Sz/Z
(paiiju02)
uel OF .89
uel, 489
y02,Q1 VV reeks aaa eNO aC ADE cates SAE lI IH ng ‘purysyt Suo'Ty HO
121,91 VP Ce ee ee
‘* YoRay ulssowos35]
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
S62
gz
ve
fz
L7z
gre
vz
£-9%
coz
fz
L7z
ve
ez
“= Lz
ov
ay
Q'6E
G°ZE
S-ZE
G-Ze
gcc
cee
SSE
SLE
g'6F
S-ZE
Ig eV
4
‘a's ‘a's
suipesr
‘o1ps£H
je4
199
9
irs
iY
jee
if
joe
ve
Bt
jel
9
0
qideq
o0zz gr/9/3 y0,10,89 11, VP a rea oe aaa ae cece Sede (9.0) | }I9Ss9q yunoyy
0091 gb/S/3 wVS,¥1,39 ABL Ib Ce eee e reer eee e eer eee ce ree seseee keg s,uewyoues J
omar y, 23eq CM) CN) u01}d110saq
opnyisuoT apnyijzey
eee)
UOT}L}S JO UOTJBOOT
(panuyuor) YANWNAS ‘OD
(panu1juod)—I ATaV J,
99
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
Tt
VI
at
SrvI
£91
Z91
QI
88
SIl<
QZ
19
19
1S
1S
19
19
19
gS
19
STt<
“81
1°61
S61
1°61
oz
I’61
g'Z1
9°41
g°41
QI
ZI
ZI
QI
IOI
oor
vz
1
zl
9
oe
99
199
Oo
oori
oofo
Sver
gv /Sz/Z
gr/6/8
gb/Sz/Z
(panuijuod )
“ eV ,9Z, 89
j0F,0£ ,69
n&1 12,89
{OOOO 00 nt nite rn een SoC Ie yy) pueys | eee
OC ,O1 VV gishigh ies sera relekere iether chey*Anadeae cee ET OOS eAON ‘Q8poT ssoly
neo ,01, PV eee eee ee rere essere eeeesseeeeeeseseseeeee keg oyotief
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
100
Te
81
82
a's
npy
v~
Loe
Lvz
ve
LVzZ
Lbz
L7z
cz
ez
cz
Size
1°61
ZI
EGE
£92
GSE
Gee
StS
SZE
SII
OIL
CTL
Ger
Gel
Qvi
V'z1
“LSI
‘as
“AINDY
gs
SuIpest
“OIpAH
i9F
joe
yideq
o£g0
Crz1
0002
OoVl
aunty,
gh/9gz/2
gr /bz/Z
gb /Sz/Z
33eq
n 9,47 539 0,40, vv eo ere eee eee eee eee eeeeessese ‘song B3S pueysy sueMS
131,00,69 GO Ut nis Soe Sig io ntesesten slay side Sinan on nt kc eRe eq@asdOnar
uc&¥ {97539 790,30 ,"V Sea ei A ge ean ee RT CS (OV (| puryst SUeMS
CM) CN) uorqdts0saq
dpnyisuoT apnyyey
FF
UO01ZL}S JO UOI}LI07]
(pamuyuor) YAWWAS “O
(panuiyuor)—1 Alavy,
Tol
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
Oc
199
G-1z
Zz
bz
£°9z
ale
£-9z
ale
LV
Lbvz
Lvz
LL
88
88
zOl
zol
zor
zol
col
zor
col
Sll<
ii
ori
Srr<—
oz
ace
S'6z
£972
£°9z
SSz
£92
L bz
Sr60
OfzI
8r/9z/Z
gr/9/6
(panuiyuos )
15Z 89
9/02 ,29
jbo,'V Boe ee ee ee eee ee eee ee ne eo NPE IH ang
S:9, "PF eee reer eee err eeeereee eee eee eeereeeeee ee yueg plese
Or’
nO
HH
4
Oo
>
SMITHSONIAN MISCELLANEOUS COLLECTIONS
102
je
a's
npy
Lrz
Lvz
Lbz
Lvz
geez
42
f-gz
gcse
got
£92
£°gz
ze
ze
ze
S°ZE
ose
gor
zy
a
cy
Sv
cv
1'61
gI
ZgI
SOI
‘as
*AInDa
Surpeai
“OIpATT
199
199
Ws
BY
jev
ie
of
bz
8t
ZI
4
é
9
o
184
eh
199
499
qidoq
Cz
oofo
Sv60
omy,
gb/ez/Z 131,00,69 uBlQsoeP SrOeseAe ight 6 wir eons donee cholerate.eteccieyere 2)” 2a) SULIT PUG) ysng OMT,
gr/Z2/8 0,17 ,49 yOe,10, VF Nee ee eee ee ee eee ee a a DEE jo Aeg
gh/ge/Z 5% 89
a7eq CA) CN)
aepnysuoyT asapnjijzeT
vo, vv Ole 60 & SS 6 6 0 60 6 00) 6 6 6h woes ele 6: ele ie es 6 eue: ete
uorjd1iosaq
‘ACG IH eng
Sr
(panuyuor) YAWWAS ‘9
(panuyuor)—1I ATAV
UO0IZe}S JO UOT}BIO'T
103
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
BY
i9f
of
vzZ
st
Zl
4
0060
CrSt
00g0
Crst
gb/zez/Z
gr/Z/8
8h/8/8
gb/iz/Z
(pamuyuos )
1Z£,09
191 ,40,99
91,4099
y08,TE,69
jOS eV i eS DT Deon yo ‘Keg suyof
yatjoc ev soo eee eee mere oer eeeeeeeeeeeeeeeeee ‘od
Wal OScey ete orl ss eee ONG CAON JOUIER UnOmaE
LSE SRO Eire lebepe rene che Ache neAns aceon gene Sse OU ENT ‘roqiey pinbewog
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
104
co
i9t
‘a's
jenpoy
ve
vz
V~
va
Lye
Lbe
Lye
19
$°3S
gv
1S
1S
1S
1S
S°9S
S°gS
$°9S
S°gS
19
19
89
oy
ay
Ge2e
ze
ce
Lbvz
‘as
‘andy
SuIpeat
*OIpATT
jel
9
0
8
184
ae
99
09
ibs
yid3q
0060 = 8h/8/g ,£1,60,99 ,zv,9r,€7 “°° °°" e00S BAON ‘aoURIJUq JOqIe_T YMowIe_
0020" §¥/Z/e- ,0V,c£ 99. av, QVeer 22° ens AON “GINS INI, Jos
0060 gb/ez/Z £09 jOS,cV Dose sere: one cor ee eee eoe ee eee ee *pinbewog yo ‘Keg suyof
oul y, 23eq CM) CN) uorjdi1osaq
apnyisuoT apnyizeyT
———E——————————EEE ;
UOI}E}S JO UOTJLIOT
(panuyuor) YAWWNAS ‘D
(panuyuor)—1 TIav
105
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
if
ZI
£97
GSS
gv
ose
ose
cee
ze
ze
ze
Ze
£92
Lbvz
S62
ZV
oz
G-1z
oz
C61
Giz
Cer
gl
€QI1
S61
Lee
LS.
Zz
L7z
Lz
vz
199
oSfI
0077
o£oz
OfZI
gb /1z/Z
8?/8/8
gv /oz/Z
(panuyuos )
u99,8£, 69
ZF V9
p01 ,c1,02
[ESTP EP Sete ere ase
“
nav £V£¥ eee eee e cree eseee
eee eee weet eee
mrs © 8 Melee mea oannele. SSS AT weed HO
of 2b fF wraps Sesedetel ci ehslehe Sones clot Aiea ieee ae eee OOS BAON Tol WOg
apiIsolo,y YyNowye.y
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
106
0g
‘a's
jenpy
89
19
19
19
19
19
ZV
See
£92
S*6z
Seae
ev
gI
“Lt
ZL
ZI
VSI
£-S5
QV
ZV
a
a
Ze
£o1
g ri
£91
SZ
‘as
“andy
58
09
Ss
09
Sb
SurIpear
“OIpATT
uidaq
Ooo!
of61
oS€1
awry,
gb /12/Z
gr /61/Z
gb/1z/Z
ayeq
08,469 BEEP cece eceeseeccccererecessossmupST WaaIL) J3INO yO
195,¥1,04 25,08 £7 seen cece eee ceeecceeeseeesesseeeees IOGIBIT PULTIOg
190,809 uVS,1V,€F igs eiensisre@ieneivisneisielesierensisie Se orth eet TOON weed HO
CM) CND uord110saq
apn}yisuoT opnyizeyT
Qe EEE neneent
UOoTze}s JO UOT}VIO7T
(panuyuor) YAWWNS ‘OD
(panuyuor)—1I Fav,
107
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
Vee
fgz
Lbz
Lz
g'0z
Lz
SOI
£-9z
Ltz
£-fz
L7z
oz
Zg1
gI
gI
Ser
SE
LL
SIvi
ofSt
SIQI
gh /61/2
gb /61/Z
gb /61/2
(panuiyuo?)
y0e,41,04
uVS,L2,0L
wVE 21,02
u¥Z,9Z,£P PSS S Sie eiORie sie sine ples vise COS iR etre any YOOX ayeu MA
uVS,42,£V SLeTa caierey ers iene Telers prose efe sete file Ce eat OO ST Auoyjuy PIO
Betyeoser. Sieg ai oie tev eatac ping esey arexese0 <, ie sneer Lay peoy puryyiog
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
108
oo
nA
‘a's
[enjoy
ev 06
gv z6
gv z6
ze S8
19 Te)
19 S6
19 So
S:5S bo
19 S6
19 S6
ev 06
S°ZE 88
ze S98
aise 28
SZ 88
cy 00
Sof tg
“bz 94
ze sg
ze sg
Sof bg
LL £0
£L £6
19 So
89 96
‘Gs sulpea.
“aInDy ‘o1pA
81
ZI
9
oO
oO
184
24
99
199
jPS
iB
eV
ie
joe
ile
81
ae
9
o
(84
(cL
99
199
WS
iB
jov
yidog
4
oOfIl gr /61/2 yO¥ 82,04 190,21 ,£V A eilel\eileije (¢)\ej(e)\e 1610/06: 6\\e, © '¢ 10 (0 6 6 siecle panishenel S \S TTS) peop Plead
OLVIS” BY/8/8) {Vaj0VQ59 ROOT Ch. 2-2 sie we tieiewieinns "BI0IS BAON ‘Spay jsamyjnoS
Oo€1 gh /61/Z WED IAS uQ1,0Z£P shave ssl anelehohsleiaceherejelsiens)ielleijsl etal slieiceishelaierspene eee astodiog adey
Civtr gr /61/Z O°, L104 uVZ,92,€P Sp GA aria eresiey aay 65 O46, mua. a, a) Sire, eC) epeveya, eye, are, «08 asprT YOY ae
ou, qed CAA) CN) uord140saqq
epnzisu0T epnyyzeyT
a eee
U01}e}S JO UOT}EOO'T
(panuyuor) YAWINAS “OD
(panuijuor)—1 alav J,
109
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
ile
Sof
gcse
gce
See
See
sec
£°9c
fz
£-s%
£92
G°Sz
Lye
g’zz
zz
zz
Giz
vz
£92
£92
£92
£°97c
“yv
ov
cce
ze
ze
ov
S-gS
19
19
S-gS
ooo!
ood
Ofer
ofSt
gb /61/2Z
gh/gi/4
gh/gi/4
gbh/gi/é
(panuyuod)
yOL,VE 04 vz,ro,£b STaPe eee SOrersTeii: 6 eke Shera sormieserereies See TTS LOT aspo] OX
uBejeV 04 1Vz,VO,€P CC Ce ce ‘0d
nBZ,e¥ 04 iVe,VO,£V CC ‘od
uQZ EV 04 vz ,¥o £0 sisi o sora rshSialgis OnreTaeNAeY soe Rere lel gee, ‘1OqIe FT ynouts}.10q
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
IIo
cl
RX
‘a's
jenpy
S-oS
SV
4
cv
£92
£-9z
41
19
QV
Gee
ze
ee
S-6z
Sof
19
19
C56
gv
SV
G25
S°ZE
GSE
ze
£92
€-Zz
£92
S°6z
1GES:
“andy
88
88
98
58
28
18
28
£3
suipess
‘OIpATT
(84
24
99
199
irs
8?
jev
i9t
of
ve
81
yl
9
4
yidaq
CVvil gb /g1/Z Wee OL 442,05 eV Sprecher ec etka SS ODT ysodAinqMoN yO
0060 gb/61/Z ~vz,Ib,0£ ,VS,Z0,€¥ GiesPibia acelin cbaasor ete siccnaes essa oRe ee E CN ‘yynouls}10g yO
OOOIL gr /61/2 Oe ,VE 04 uVe,VO,€V ee ops A 3spoT yIOK
amy eq (CM) CN) uordr19saq
opnyisu0'T epnyizey .
Se
U01}E}S JO UOT}LIOTT
(panuiu09) YAWWNS “OD
(panuijuor)—1 ATav I,
IIil
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
6z
21
L7z1
£vI1
gI
10
Bil
Vel
QrI
gI
88
88
LL
89
89
19
QV
ov
ce
Sec
£92
£92
Sof
ze
c1<
cm
zol
88
88
LL
LL
199
S11I
SI1o1
SrvOoI
(panuyuo?)
OP/S1/ 2° Ve-OR0L. OC Oe ei ee ea SN ‘loqiey{ Jojsaonojy
° a“
gh /gi/Z u81,9£ 04 790,300 oi Sheleerevave:elenerexsieiehe)isieusieiessiesele eho oese SOS DiTAl ‘uuy ade) yO
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
II2
ie
48
8
0
‘as
jenpy
ZV
av
£92
£92
£97
6°02
Zz
cz
OIL
O'Z1
Vl
C1l
S11
Sor
ZI
aAt
SEI
SII
S-Er
£1
VI
ZI
O01
O7ZI
O7ZI
VI
QrI
‘as
“andy
06
06
<8
8
c8
Ca
bZ
bZ
zy
Ly
oY
ov
ov
ov
ev
Sv
oS
ov
oS
gv
zs
29
of
LY
Ly
Zs
ss
surpear
‘o1pATT
jv
if
joe
ve
81
wl
9
0
ibe
Bl
jel
9
0)
vz
81
el
9
oO
1
jel
9
0
ive
81
jel
9
oO
yidaq
o£60
cist
CIvI
Ozel
CIZI
awry,
8h/81/4
Qb/S1/Z
gh /S1/Z
ajeq
yV 1,08 02 yVI,Ve ev ose fake ene lt ene) ese acess Shee, S18ie) SreAeue She see SOCIAL ‘uuy adey HO
4VZ,0V 04 4072,9£ eV eee eee ee eeeeeeeeseeeeeeeeeeeeeeeeeses ‘og
Vz ,0V 04 ,02,9¢ eV eee errr ee eeeee eee esesesee **SSeyy f10qieyy 19}s9dNO]+)
CAM) CN) uorjdt19saq
epnyisuoyT apnyjey
ed
U013e}S JO UOT}BI0T
(panuyuor) YAWWAS ‘DO
(panuiyuor)—1I ATAVL,
PE
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
‘
£z
for
Lz
f9z
19
19
gv
cs
QP
QV
gv
199
coool
ooSI
gr/Z/6
gb/o1/Z
(panunyuos)
iAP .59
“ 9£,9£ 04
/S0,eV see geese Le ceneken chelates okekese took gn ecn near LE sumMOIg jo ynosg
u9£91 ocV
ee eee oats acai peed tance eames CELL JST u0}soq yO
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
II4
“en
=
OT
cz
g bz
£-9z
£91
41
41
€°Z1
£91
vZ
94
08
09
29
29
£9
29
09
30
16)
c6
+6
v6
z6
z6
06
06
06
388
88
06
06
OOI
OOI
OOI
66
suIpeat
‘OIpAPT
199
yidaq
ofzi gb/6/Z 40£,08 S42 E517 mi S)sia alsin leis Sia !s(e/ainisle\t/e/n\s[sjs\aiersie.e # tine sslares Spee GDA Beatle
$160 gv /o1/Z Se ,0V 04 wav OV Iv sien shol seh el she) ee) Sielereie ech) ese iapeis iS cease es TELLC@) pop aden
OIZI gb /o1/Z uVZ,87,04 ye ,10,cV Cece ee eee eeeseesreeeeeseeeerresess seg s}jasnyoesseyy
OOOL gb/L/6 4¥,,S9 S0,2r seeee see eee eees seme eee encees *‘yueg suMOIg JO yynos
oun 97k CM) CN) uoi3diiosaq
apnyisuoT apnyiey
eet
U01}L4S JO UOT}LIO'T
(panuizu09) YAWINAS ‘D
(panuyuor)—1 alav
II5
NO. IO WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
91
oz
Oz
Oz
£91
£91
ZI
f91
{or
“I
ZI
fg1
£91
gi
gI
oz
Lez
6'0z
Le
LOS
£91
ZI
gI
Ver
Vz1
VI
gr
ZI
“291
gI
Oz
oZ
oZ
o£
OZ
09
09
29
09
29
29
09
0)
59
oZ
SZ
ZZ
GZ
GZ
o£
“9
cv
oy
zs
cs
29
£9
59
o£
S1zo
OofT
o£go
8r/3/4
8h /8/é
gr /6/2
(panuyuos)
yOe,VI IZ uV£,61,1V Coeee eres eeseseesesreeeeesesrersesee punos pieAout A
07,02 yS,37,1P rr ‘od
02,02 ySe,8,1P Oe ee errs precxour, sey} ey
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
116
‘as
yenpy
gv
g vi
1ST
St
Gat
Z91
gl
oz
oz
GIr
Gir
O21
ge1
Gil
Ol
VzI
Gel
gi
gI
gI
zol
oz
oz
oz
C61
zO6l
SOI
‘a's
“andy
69
89
69
SulIpest
‘o1pAH
BV
jv
9
of
Ve
st
jel
9
Oo
4
Bt
ee
9
Oo
Bt
21
9
0
(84
eh
199
/99
vs
gv
i
it
of
ve
4
qidoq
4
Svor 9b/3/2 ,00,"V,04 yOV,1f Iv ewe ewer seer ree eesesesseersseseseeees punos piesout A
ofZ1
OOIL gb /L/Z Oe VE 12 ,OV,11,1V oso Weis ie. execeisieselcie\e (ene! seiste. orice ve OO TEED pueysy yoorg
C1zo 9b /3/2 OVI 1S yVe Ol, Iv eee were teense eee e weet eee eee ew eee reese punos preAoUl A
omy 93eq CM) CN) uo1j}d119saq
UOT}eYS JO UOT}LOOT
(panuyuor) YANWAS OD
(panuyjuod)—1 Fav,
117
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
92x
ze
Sof
Sof
S°Ze
G°Ze
ev
1s
1S
1S
1S
Lite
6:02
0z
QI
£21
QI
gl
QI
SI
gI
GILT
oI
3'6
ZOo1
OI
Ol
OI
OI
£o1
Ol
OI
gb
00g0
OfF1
ofZt
gr/3/6
gr/9/Z
gh /Z/2
(panunyuo)
12,69
08 1,22
u Sf PPL
/OV OF eee et eee ww we meee ee ee Pee eras JYySIT yoxonyue Ny
y0L,25,0V eee eee eee eee eee eee eee meee ewe eee uojduieyaspig yO
1ev vO, IP okersheperexsssie selene noel svc scue ke: keer sacs TUTE OG pues] :olg
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
118
ye
iS
a's
npy
v'z1
VZ1
SL
Ce
ae |
c1I>
ciI>
ciI>
9°9
99
z9
i>
ciI>
ce
VI
S-E1
eI
SB
Cer
gi
ZI
Ger
Ev.
89
19
cs
G-se
[GES
“AINDY
os
a)
z9
os
es
96
$6
¥6
48
Suipea1
‘O1pATT
Bt
zt
9
0
84
24
199
09
yideq
ofSo
SPEI
of£g0
00g0
aunty,
gr/9/4
Qh /S/Z
8h/9/Z
8r/3/6
a3eq
nOV LV 2 n9e,1£,0F wieiojeliehes chez siemens) eleseueledereiche) tune ehe-n tar cre PMCS ONT: Ie HO
725,00, ¢L nol VE OV cisteieseicusveleleiese}elej erehe/ehekoxe ties enndohese (ele keke PURIST: Quo) HO
QV 4To&S nee ae OV SEEN YS TSS tess cee ye ey ree peel DUETS: aIL yO
12,69 OV ,0V Sar ehohexsie ish scsrele re: ojeieysiste sionsneshen sresens TUS WSrT yoxonjueN
CM) CN) uorydri9saq
apnyizuoT opnyyey
ee SS ed
U01}e}S JO UOI}LIOT
(panuyuor) YANWOAS ‘D
(panuijuor)—I AIaV I,
119
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
be
‘
~
ite)
iP
£92
LEE
oz
oz
oz
oz
0z
GLE
6°0z
Lez
oz
oz
gI
Lee
VzI
OI
C1
Gates
c1>
car
£-9z
£-9z
gI
grt
SEI
OI
Sg
+'6
VzI
V'eI
Vel
08
GZ
o£
/
02
VA
~
oZ
GZ
ZZ
GZ
oZ
OZ
59
GZ
ofVvi
O£II
orZo
ooor
gh/b/£
gh/S/2
gb/S/Z
gh /S/2Z
(pamuiyuos )
19S ,£4
0985084
702,00, V2
98082
n2¥ ,9V,6£ SR aay” iri n ieee Ue eneke) Or ae seaery Seal eee LTE ECS WSIT yesouleg
OL OF ,OV Ce YOY AinqsmaysS
y0V,9Z,0¥ See eee ee ree Se te see ee OO ET Apues jo SIM
,0V ,9Z,0V oa eEa hat neces S Lotsa oheae bese eenee se SCAT O SIT pur]}09S
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
120
GS
jenpy
£97
£92
ce
£92
ze
ze
Ce
ov
ay
19
Cnn—<
ClI<
Gin
Sri<
Cnn<
Cli<
St1<
Cimi<e
Cli<
Cie
Sli<
Grice
88
Oz
L7z
e902
GS
“andy
08
SuIpear
“o1pA HT
je
199
i99
qidaq
OOOoI
0z00
OfVI
oul y
gh /F/2
gr/6/6
gr /t/Z
23eq
08,V1 VL BI ,0€ See eet Ch giracus nee) a cese a seaea ccuascteke | "'N ‘AWD oUeyTVy yO
St 6;9€ ,6£ ee ere e eee eee er ee ee eeeroeeeseeseeeee uoduey uospnyy
OSReG uSV,9V,0€ shob mace sueje GueseneiessletNe iene ebeh selena OTuyS WS yesouleg
CM) CN) uorjdiiosaq
apnjyisuoT opnyijyey
co ——
U01}e4S JO UOTJRI0'T
(panuyu0r) YAWINAS *D
(panu1juor)—'I ATAV I,
121
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
““
FI
z'6
88
0°60
£01
S-1>
Ol
IOI
II
Oo!
SOL
OI
1g
Cie
I>
cI>
cI>
piss
Sis
c1>
o>
Pv
cI>
S9
Iv
rv
ze
€°9z
ofor
o£g1
0091
Oofl
O£II
L£y/31/Z
gy /E/Z
1$/1€/2
Ly /9z/2
gb /1/Z
(panuyuos )
130,V8 V4
130,¥5,¢2
LizZ,94
85,824
osjeceae
na¥ 95 8° eevee reer er ee erases ereeeeeeereeeeerereeeeee ‘od
nV 95 08° eeroeceerereeeeeeeeeereer sees eres se SOU UE ET ACT ode)
$;6S ,g& See ees eke UP eb ch ace hence een OLG |) TLE, keg ayeodesayy
yev 41,08 corr c ct saArYy aremejag 4ysiy uyof diyg YO
1z¥,L1,6€ Seis aired ote shin eile ATE OAC MEIC qysrT uyof drys
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
122
‘a's
penjoy
oSI
Tes
VI
QI
LSI
a
Ser
Cer
VzI
VzI
SIl
Lol
£01
a0)
1°
1'g
88
S°6
+6
v6
z6
i)
$6
9°6
OI
£01
Sg
‘a's
“AInD”
ZS
9S
vs
6S
gs
oS
os
oS
cv
cr
ov
cc
Ze
Gz
gI
gl
Ze
Qz
Sz
Sz
ve
9z
9z
Le
of
Ze
oz
super
‘OIpATT
midoq
OOo1IO
oo4I
Ooo!
Of Ir
Ofor
out
L£y/9z/Z
gh /1/Z
£y/91/Z
Lv /31/2
27eq
{90,82 US.98 Peet m eee e eee sees sess eseesesesees 0d
190,42 S-bS gf a steininisialeiniaiaierPinnien eee TOATRT Oe BIS GT ‘Teoys uMOIg
180,¥S,¥4 uA¥ 98 o8e wee er ewer errr secre eee ee reeves eereseres ‘og
130,VS,02 v4¥,9S .3f IAs NO GS RRR OYE Aree ER ge Pee eR TOUR ET Ae ade)
CM) CN) worjdrs1osaqq
apnysuoyT apnyyeqT
Oe
U0T}e}S JO UOT}VOO'T
(panuy109) YANWWNS ‘OD
(panujuo0r)—I AIdV L,
123
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
II
FI
Sr
Fr
£1
r°Si
Q vi
Si
SSI
“Sr
SSI
QrI
VI
CVI
Qri
Qri
VzI
VzI
Vel
FI
Qvi
G°SI
QvI
£1
QV
GSI
c'S1
FI
VI
ZS1
£91
£91
LL
Oort
oofo
00ZO
L£b/S2/Z
Le/oz/Z
L£y/92/Z
(panuyuos)
{90,54
90,54
(9082
SS gf eee eeeeeeereeeseeereeeeeeesreeeseeene ‘od
S-7S gf ee ‘0g
S-S .g& See ewww meee reer e es ee ee eeeeeeeeeene ‘od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
124
je
a's
nypy
Q'V1
SI
Q bi
SI
EVI
VZI
61l
Cale
SII
Ol
OIL
O1I
£1
sor
8
VI
CPI
Cri
OVI
£1
ZI
ZI
ZI
ZI
CSI
Vel
O1l
‘as
“aInbDy
surpeat
‘OIpAH
joe
jbz
wideq
0091 Ly /Sz/Z 90,S2 S-FS 9 Bigiepe maeksredost Ae tener Don ss pilay Sener ee See ees ee tO GT
ooSt Lb /Sz/Z 90,54 S75 9 Ce Ce ee veseise =O
oOorr Ly /Sz/Z {90,4 G-bS gf SoS peeeoe ay it ane buen ene nac tein MAT NTN IBMPTIG ‘yeoyus uMOIG
sully, 23eq CM) CN) uorydi19saq
epnyrsuoT epny yey
SSS ee
U0T}e}s JO WOT}BO07T
(panuyuor) YAWWAS ‘O
(panuijuor)—1 alTav I,
125
WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
IO
NO.
iG
Bat
CSI
ISI
S°f1
VI
6721
Eon
Lek
CVI
Q'rI
1st
ISI
gr
St
CVvI
VI
6O'II
OIL
ZI
PI
VI
FI
Sy
CPI
Qvi
Ist
CEL
cl
Zol
Cali
S’VI
QrI
ogi
ooZI
Ly /Se/Z
Ly /Sz/Z
( pamjuo?)
190,84
190.84
190,94
Gis gf eer cece rroseeesreeseeeseeeeeeeseeere ‘oq
G-VS .gf Seer ees ee sore eeeseeesseeesesseeesee ‘oq
S;bS ,gf Cee reeeeseesresseeeeseseeeseseeeeee ‘oq
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
126
ICES:
jenpy
af
OI!
(ka
€-gI
e-o1
SSI
GSI
Zl
“SI
1'S1
CVI
oVI
C-v1
V1
£91
£91
eg
gl
St
cS1
SI
St
gel
SEI
CCI
TEST
“91
gli
ey
suIpeot
‘ospATy
9
Oo oozz LV /S2/L {90,84 S-bS gf eee meee reese eeeeeeeeeeeseeseeeeeeees ‘od
d
jvS
13
ev
19k
joe
iva
Bl
jel
9 :
Oo OOIZ Ly /Sz2/Z 190,54 Sir ge eee eee eer eee ee eee eeeeeeeeeeeeesene ‘oq
d
vs
18
jev
it
j0e
ive
1
jel
9 :
Oo 000z Lv /S2/Z 190,94 G;vS gf Cee e cree eee eee reser eeseseereeeesee ‘og
d
vs
By 0061 + Zb/Sz/Z 190,94 GS QE ccrcs cress sees es IOANY arEMeEjaqd ‘eoys umoig
qyideaq = wy, aye CAN) CN) uorydisosaq
epnyisuoT epnyize'y
<r
WO1}e}S JO UOT}LIO'T
(panuyu0s) YAWWAS ‘D
(panuiyuor)—I aATav J,
127
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
ISI
I°St
SSr
gl
S71
491
£01
£91
“SI
gr
Q ri
Ss
VI
V1
QI
ZI
21
gI
GSI
CVI
£-€1
Q f1
gl
“SI
gli
gI
QI
GSI
St
Q v1
VI
it
oot
oo£ze
Ly/Se/L
Lb /S2/Z
(panuyuos)
190,54
90,82
Sib .9gf @eoeeroesssceeseeseeseersesserevesese "od
S*bS gf ee ce ‘og
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
128
al
“9,9
9
Ges
jenpy
gI
gl
gl
ZI
ZI
Vgi
8
8
g4
VL
aZ
Mee
1°g
gs
+6
£O1
Vol
I'ol
g'O1
GzI
6cI
Ceo
Il
O'O1
OO!
II
Zo1
‘a's
“aInD ay
6S
6S
6S
<9
c9
99
ZI
an
gl
v1
C1
VI
QI
SI
c~
Ze
1¢
Ig
vE
Lv
LV
bP
ZE
gf
of
Ze
ce
suipest
‘o1pAH
9
joe
ve
8!
ZI
4
O
sof
of
ve
4
81
ZI
‘
9
oO
of
ibe
81
jel
9
O
jv
98
joe
jhe
81
jel
9
oO
‘
4
qideq
vrgo Ly /S2/Z AS Ares ene a eae eee Wee nese heel ens ees COD GE yeoys SOIQIW
Sv60 18/62/Z ,21,11,94 SATS Q es oo nee eh ee oh bine KEG Molseu) Aes Els
zSol 1$/6z/Z Szv1,94 G1S.9e Coe eeeereeeeeeesreeeeeeseeeeseeeese ‘oq
oSg0 1S/1€/Z Szv1,9Z GISGQE “co tose settee sess Kong 4uIOg UeMIys|LT yO opus Ol,
omy, 278d CA) CN) worsdrs9s9q
apnyIsuoT opnyjzey
[Se ee EEE
UoT}eYS JO UOT}LIO"T
(panuyuod) MAWIANS ‘DO
(panunuo0d)—I ATV J,
129
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
11¢
“ ‘
uO
199
6'0z
6'0z
6:02
zor
gI
QI
r°St
eZ
zZ
89
89
oZ
ZI
te
lz
6z
re
be
se
ce
Sv
gs
£9
vs
oF
LE
ze
o£
oZ
ZZ
cL
GL
89
59
9S
bz
ol
ZI
9
8!
=
jel
Oost
0060
Sogo
SIoI
OfZI
gh/t/Z
18 /6¢/L
1GS/1€/Z
16 /1£/L
L£b/o1/Z
(pamujuos )
u81,08 VL
Sziz1,9Z
bz ,94
£708 ,bZ
pVEJOVQE “roosts ttre 9 Kee SJeMETeC] [EOUS solo, BO
405,00 gf @).eiaye) elel le Sie seh ajceis aie e¥ ere 0) 9 fe 20ue1qUS JOART oAM pue SOTA
408 ,OF 8 STaveacNo leds ocuey encren ne Monte ECR CNA UES yao pur]poo yO
j0S gf ececee eo eeconseeeen "WSrT yuIOg Apoo|g jo “M oyu E
G:0S ,gf ee ey yeous SITIDQIW
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
130
{0
zi
£1
‘a's
jenpy
GSI
GSI
LSI
“:S1
GSI
Q v1
S:S1
£o1
gl
gl
gl
£91
Zgl
“SI
QVv1
£-o1
£91
ZI
ZI
€-Z1
“LSI
€91
gbz
go be
oS
C-Se
‘as
“ainby
Surpesr
‘o1pAH
gv
cv
‘
if
joe
yydoq
OIFI gr /6/6 S105 eZ IBV .8E Spas ephag seaie ceckc ial eactste caciee one ae LS WSIT S][PJ1IAO HO
Oo£or LV /S2/Z S;9S ,vZ OV gE eo eee eee essere eeeseeeeeeeeeeeeeeeeeseeeeesese drys
WSIT S][ejsoaQ pue [Boys salgjopy vsomjoq AemyyeTT
oost gV/E/Z ,91,0S,VLZ ,,VZ,OV,gE --* cote ttt AO areMelEG ‘JeOYS sa1ID.IV BO
omy, ayeq CAA) CN) uondis90saq
opnyisuoT opnyyey
YY - w~Y
UOT}eYS JO WOT}LIO'T
(panuyuor) YAWWOAS ‘DO
(panwiyuor)—1 AAV,
131
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
Te
~
ze
II
0z
gI
£-9z
ze
ze
Sof
ze
g-Ze
88
oni<
cIi<
Grr
cli<
STl<
oni<
pal <<
Sr
£1
OZI
67Z1
£1
Lz1
O7zI
Cer
V1
VI
v1
£71
Bait
OIL
VI
(ASK
199
o£So
CIeI
gh/h/Z
Qr/6/6
0$/€z/6
Ly /S2/Z
(panuyuos)
0¥,S6,r2
,0V;S€,bZ
LSE pL
$;10,S2
BV OBE Cee e seers eeeeeerereseseseseeeeeseeee ‘od
BV Re eRe ene ney eved ie AURME LS ater rae Cee WSVT WOE J-IALT yO
BV o8F Pee O eR tee S26 eres eae aS WSsrT WOY}eJ-dATy
BV8F ©)'8 0.010) 6.8) © 0:60:06. 6 610 00 00:0) 5 6 60/06 0 00\0 ss 6 ‘oq
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
132
o1,€
a“
‘as
jenpy
SSz
ES
£7
fz
fz
LZ
Cig
cz
ez
0z
AT
GLI
£-O1
v2
vZ
gZ
Oz
gI
gI
ZI
£91
QI
gl
€or
£91
QI
gI
‘a's
*ainbay
Surpeor
‘oipAH
qdaq
oozi gb/f/ZL 481,10,$4 i¥S,9¥ 3 Seaceselpscletelovel pleuslcieisicteseseicholern oT sry] S][PIIOAQ HO
Of91 G6r/ez/b S1;90,S4 SZZV,g& scorers c cess ce ess JOqIeEy Joyemyeoig “jeq ‘semey]
ofSo gb/r/Z ,0V, Sr BV .9f Sess fi Dish ok oe! exriareceyeix er elo) texas CTT Gs qwSrT woyle.J-IALT yO
out y, aed 1) CN) worydt19s3q
apnyisu0T apnyzzey
nn
U01}e4S JO UOT}LIOT
(panuyuor) YAWINAS *D
(panujuod)—1 AIAV L,
133
WILLIAMS, JOHNSON, DYER
IO WATER TRANSPARENCY:
NO.
61
ZS
ZLz1
Se1
£1
Tear
gl
£91
£-o1
Oz
bz
Cee
Cer
geI
ger
Son
V'ZI
Sci
Ez
£°9z
£-9z
€-gz
£°9z
Sof
£°Qz
Gz
S'0z
LiL
b'gI
Oz
cz
g bz
£-9z
oO
¢160
S191
Czz1
L£y/1/Z
L£b/91/Z
L£b/€z/Z
(panunyuo? )
07,94 pZ1LSoQe wisivielere’eisizie in siermianetee sr er MRGy ayeadesay) uiog 9A02
zSvL jorgt wilb reireh elke ioieliaNefeloifertelceheioilefe; eller enevelels drys WSrT S]JBJIOAO “AS
Se, vl beget aseinis)uloxoiessiaeicns cuneterecs © slecichy ITs WSJ woyyeq-aALT YO
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
134
‘a's
enjoy
Gas
Lok
gl
S-oz
VI
VzI
ZI
“G1
“S1
ZI
Lien.
gi
z Ol
6:0z
gl
£1
VZI
Q v1
671
Sea
au
Sai
gI
gi
“L931
S:0z
“1
EGS
‘and
49
suIpeor
“OIpATT
OZz1 Ly /1/Z {07,94 TOL eeeRe Soe eee eee eee eee ee eee eee eeeeeeees ‘od
OzII Lv/1/Z 07,94 Helen Re: eee eee eee eee eee eee eeeeeeeeeeseeee ‘od
S160 3=— Zh /1/Z (02294 =ZET PS LQE 2° tS eee tin eee spe Syeauesaya) JUlO gq OAOs
our y, 97eq CM) CN) worydrs9soq
apnyisuoT epnyijzeyT
Se TEIEEEInnIIIEIEEEEEEEEnnnEN nee
UOI}L}S JO UOT}LIOTT
(panuyu0r) YAWWAS ‘D
(panuyuod)—I aTav IL,
135
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
£-o1
QI
gI
gI
gI
£-Z1
GLI
Liat
Lie
SET
ZI
S21
gI
gI
COI
G21
Le
o°St
OCI
£1
SFI
v'gI
£931
L931
S‘oz
S21
Vz
grr
vI
Qri
crer
geri
OzrI
OzeI
£y/91/Z
Ly /1/Z
Ly/1/Z
(panuijuo? )
20,94
02,91
{02 , 94
lege wien se kevetmdss® elena siete aneeale.<ua marke ore TORT YOIMUs J yO
PLES Qe Piplele ee 6. eels oe) 6016.66) 0) 6 00) ees eins (ees) ele e.s ‘od
FEL CCAS Cece merece eer e ese ee ssene coer ercce ‘od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
136
9
9
‘a's
Tenjpy
£01
£o1
£o1
£01
£-O1
Sor
ZO1
Ol
Ol
£01
£01
O01
cy
1S
1S
gh
SSE
ev
aw
£92
£02
gbe
£-9z
gre
ghz
£92
‘as
*aInD ay
ze
ze
Ze
Ze
Ze
£¢
ce
of
o£
Ze
Ze
gf
00
£6
£6
ZO
£8
06
06
08
08
QZ
08
gZ
gZ
08
suipes1
“oIpAH
if
‘
bz
Bt
Fai
Oo
yidaq
ofZ1 LV /2/Z y¥5,00,92 {£0,8© Ce ‘od
ofg1
ogSt L4y/z/Z ,,VS,00,92 EOLQE tere ABI CuI], SUOLMO[OS Jo A. Opras/ OFT
Shir ZY/91/Z S-zo,SZ Sgfo,gf sco peoyS JozsenG J9WUIMA “AN [PA (MTA)
61€1 Lbv/o1/Z ¥;00,S2 POI,BE “crete KONG aps ‘YUeg [IND yeaIH YO
our y, ave CM) CN) uorzdt10saq
apnyisuoT apnjieT
a nd
UOT}2YS JO UOTFBOO'T
(panuyuor?) YANWAS “OD
(panu1uor)—I Adv J,
137
WILLIAMS, JOHNSON, DYER
IO WATER TRANSPARENCY
NO.
cor
gol
Z01
lol
ol
£01
Sor
SOI
gol
or
go
ol
ol
OI
£01
Q'01
Sor
SOI
£01
£°O1
or
COI
£01
OI
oI
£O1
Cor
Sor
g'01
Zol
Z01
ee
be
ce
If
of
ze
ee
ee
ve
o£
gz
gz
of
o£
of
Ze
ve
£¢
ee
Ze
ze
of
ee
Ze
o£
o£
ce
£e
£e
ve
ce
ce
ofez
ofzz
oL1z
ofoz
of61
Ly/z2/Z
Le/z/L
Ly /z2/Z
(panunyuos )
1¥S,00,94
uVS,00,94
u¥S 00,94
;£0,8¢ eoecereceo ree ee ee eee eeeeeeeeeeees ‘od
{£0,8° oer error ere eereeereeeseeeeeeeeres ‘oq
{£0,382 wore eee errr eee ee eee eee eeeeesens ‘oq
VOL. 139
MISCELLANEOUS COLLECTIONS
SMITHSONIAN
138
cl
cl
‘a's
renpy
oe
6:02
Ze
cz
ghz
£:9z
£92
£°9z
£92
c-0z
ve
£91
gi
zOI
£1
Son
G11
eo1
€°Z1
B41
zOI
ori
gr
Z01
901
[oI
‘as
“aInbDy
SuIpes1
‘o1pAH
ist
yadeq
ooor Z¥/o1/Z $;$0,S4 25,4€ “°° e****yeOyS JazrenZ) JowIA\ Yo ‘aiming woy}e4-01
O£Il L£y/S1/2 Zj01 SZ 9}Z0,9f ooo eee reer oe ee reer eeeeeeeeeee ‘oq
oo1r = LZb/g1/Z Z;01,S2 Q;Z0,Qf “cc cc cette puRISy anSeoyessy Jo ‘oAinD woy}e-S
off Lb/z/Z ,¥S,00,92 /£0,9F rresesce cesses qyusry duiny suowojos jo “q aru OY
amity, aed (MA) CN) uo} dt1989q
apnyisuoT opnyizeT
a
U0T}E}S JO UOT}BIO7T
(panuyuor) YAWWNS 'D
(panu1ju0d)—I AIaV I,
139
WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
IO
NO.
18
BE
S-o£
£-Qz
ce
oz
Sz
oI
off1
ofzI Lbe/€/Z
OfII
ofor Lb/€/Z
ooor §=Lb/S1/Z
(panuyue?)
195.094
(Baigs4
$*S0,$2
1QV.4e Pees eeeesseeeseseeeseeeesses ‘od
QV ZF coco seg ayevodesayy ‘puv[s] sorsuey “aq opr 1
aa oae Sew me mw ener renee eee eeenee ‘od
Sol
L2z
Lez
gSe
g be
S62
£92
ze
ze
ze
— oe:
Sg
ZO1
SII
SII
Lee
oes MS
99
88
0
z6
z0
_ 76
Iv
9°9
QZ
Sg
‘as ‘as
VOL. 139
o
MISCELLANEOUS COLLECTIONS
SMITHSONIAN
jenpy “andy
140
oI
CI
oz
sulpeot
“o1pATT
ve
1
ei
yidaq
— fb/S1/Z
Shgo Lh /91/4
SSZo = Lb/g1/Z
ofri LV /€/L
oger = Lb /€/Z
our, 27eq
(P
SO 84 9b Ze eee eer ere eer eee ereeeee peice 9TUT anseojooury) yO
,40,54 Vouk Susie ole <ehek sh cheeke sieusiant seit Shas eee UCL ysl orl HO
81,84 QV ,4E °° ***yoTUT anSeajoouryy HO “TLZ (MTA) 40ng IP
185.094 igh 4 asia ipssielesehe iets) n\wiaakelas essere: Tek usionee CGT
495,54 QV ZE soossseses Keg oyeodesayy ‘purls] Jomuey “ay api 1
CM) CN) uorjdr10saq
apnyisuoyT epnqyyey
eee
U0172}S JO UOI}LIO7T
anuyuod) YAWWNNAS ‘DO
(panuijuor)—'I AAV],
I4I
WILLIAMS, JOHNSON, DYER
IQ WATER TRANSPARENCY
NO.
OI
oI
8
Cale
CII
Ziti
II
601
ir
Vz1
z9
g4
z'6
8!
=
yl
oorl
ooo!
OO£T
Ofe1
Lb/ex/Z
Lbe/S/L
Lb/v/Z
(panuyuo?)
,0V 60,94
{01,94
Giz1,92
nSe,S0,4€ °° °° ** keg ayeodesayy ‘spunoss apply Voysess0}F]
Sole Keg axeodesoy) 4usiy der, yOM BO
)1V,Ze shee ole toh thet cbehsceh Stes sie kelcsens: Noncn vita cine PACT) Cll keg ayeodesoy)
VOL. Ig9
SMITHSONIAN MISCELLANEOUS COLLECTIONS
142
Or
Ol
OI
(GES
jenpy
Ce
Z
Sg
O11
OIL
G-€1
SI
OI
OI
O7cI
ZI
VI
CVI
S°g
OIL
€-1
LZI
Val
Lic
OIL
Ger
VZI
O7ZI
CVI
vZ1
Ol
-mnbig
Zurpesr
“o1pAH
oO
ze
vz
st
ZI
9
oO
ze
vz
81
ZI
9
oO
Ze
v2
8l
el
9
O
ze
je
81
ZI
9
oO
ce
vz
uidaq
4
4
4
é
é
4
4
oo41 §=Lh/21/Z
0091
oost Lb/e1/Z
oovl
oozl Lb/z1/Z
oo1r Lv/z1/Z
oun = ya
,0V 60,94 AekspO (AS COOC OCS SEES ESHEETS O EEE HE He ‘od
,0V 00,94 Ese cOnLe Cee rere ese r ere sees eeesevreses ‘og
y0¥ 00,94 pee edge’ Ce rr ‘og
w0V,00,94 ,S€,50,L8 ****++***Keg oyvadesayd ‘spunosy o[ppry eoysess0 zy
CM) CN) uondr10saq
apnyisuoyT — epnytyery
eS)
U01}e4S JO UOT}EIO'T
(panuyuor) YAWINNS OD
(panuijyuor)—1 alav
143
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
€1
Syl
ZI
6O1
Cult
VZI
O'ZI
$6
36
OI
O11
ZI
eI
OI
Sor
1°
ZO1
Cer
VZI
OI
Z01
Il
SrA
Lt
Lion
1
S°g
CII
Lo
£1
o00ze
ooIa
0002
ooo1
00g
Lb/er/Z
Ly /z21/Z
(panunuos )
,0¥ 60,94
,0V 60,92
nacOoae Pe ‘od
pao Oeke: ee em meee eee meee ee ewes eeneee ‘od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
144
i592
o£
é
‘as
jenpy
ao v6
ss v6
19 S6
Sone 28
Sof rg
£-9z 0g
£92 zg
19 So
oe b0
19 S6
19 c6
gv z6
Sof bg
gre gZ
rAd 06
ss v6
ss +6
19 S6
19 S6
gv zo
£-92 zQ
Sof 1g
II Ze
wil gt
iia ov
6'ZI Lb
Crt ov
‘a's SuIpeot
‘ainbay ‘OIpATT
18
eA
9
yidaq
ooSt Lb /¥1/L £352 00, 4€ ee ee eee eee e eee reer reese eeeeeeeseesreesne ‘od
Oorl Lh /v1/Z LL, ROO BLE Ao Ses re seess esens)se..8 eee eee eee eee eee eee ee ‘0d
oof1 Ly /v1/Z L£:SL 00,4 speigusk titi © Ruse Sbieres eyeiaNeelasanerevsien sist WSrT ayeodesayy
ooze Lb/z1/Z ,0v,60,94 ,S€,S0,Z€ ******** keg ayeadesayD ‘spunosy s]pprIy{ Poysess0F{
omy, aed (AA) CN) uorydrsosaq
epnjyisuo'T epnyney
nn
uol}e}s JO UOT}eI07T
(papyjru0d) YAWIWAS ‘OD
(panui4yuo0d)— I AIdv J,
145
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
a“
3
Sg
Zil
Ol
S11
6'zI
GSI
Ev1
L7z1
{z1
€1
G°S
Sg
v0
ON
ev
0g
08
+6
ZO
ZO
+6
c6
rg
c8
08
08
98
z6
oS60
OIvI
Crri
0091
1$/S/11
1$/1I/II
Lb /1/11
Lb/vr/Z
(paiijuod )
190,¥5 ,b2 u4¥ 9S 8° C65 666 60 Cee Os oe 6 6 6 660 04 66 6 bee Aong BIS ‘ACTIN odey
£2294 $°6S ,gf eee eee eee eee ee “-OSplig Aeg oyeodesoy)
00602) FOV, 20,08 0 S08 onsale ss Keer OYOGesoYa)) AIO ge UENS
NWOALNV ‘Gd
£32 j00,4€ eee e eee eee eer ee eeeeeeeeeeeeeeseeneeae 0g
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
146
je
~
wn
=
=
by
“
a“ 09
a's
npy
Sg
Sg
99
Sg
z6
38
8
z6
96
9°6
96
96
Z
Z
Sg
Sg
VL
CL,
g4
gs
eal
Z
al
el
La
Zi
z9
‘as
“ainbay
Oz
0c
Ol
0z
ve
cz
LI
ve
Le
Zz
LE
ke
ZI
ZI
0z
Oz
VI
£1
gl
SI
£1
ZI
£1
er
9
9
8
sulpest
‘O1pAPT
9
0
d
jel
9
0
d
ZI
9
Oo
ve
81
el
yydeq
ofoz
oSgo
0zgo
SozI
OIZI
ovel
o0S60
out]
1$/6z/O1
1$/1£/o1
1S/1€/o1
1$/V/11
1$/V/11
1S/V7/1I
1S/S/11
aqyvq
180,¥8 v2 v4¥ 98 .8& re
uBOVEVe 495.98 °°
u80,VS v2 u4¥ 9S,8€ fa
u80,"5 .bZ v4 9888 oh
UO VaGve- 4ArOI.86
180,"S VL w4¥ 95.88 et
TBO aela WAV Oteee
CA) CN)
apnitsuo'T opnyyey
eee eee eee sone
seeessyo0p ‘1oqiepy Ae aded
sseeeess Kong vag ‘Ae ode)
uorjydiiasaq
ee
u01}e}S JO UOIZBI0'T
(panuyuor) NIWOLAV ‘Gd
(panwyuor)—'I ATV,
147
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
v1
OI
ghz
bgt
ZI
gt
Zl
Cz
Os
6°07
oz
QI
C75
LG
oz
oz
S61
GSI
9°01
g'ol
V'VI
os
Zi
of
ve
81
ZI
‘
(panuijuo? )
Of1I I8/z1/1r. Sz;"1,94
‘
Bost. “15/en/1r Seri 02
€191 mS/€1/1r Sz;r1,92
oS1r 19/S/11 90,S2
of60 6/2/11 SZizz,9Z
G19 Qf Oe ‘oq
GIS .g€ **Aeq oyvodesoyy ‘Aegq usojseq uIog ueWwYsiIT yO
GPS ge Sea ae hee cae ot LOAN dIVMBIIG eoys uMOIG
UGG Ge oot cece cere Kpey oyeodesoy®, “Aeg Sssnoyyog
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
148
ius
OI
VI
u8S
+71
ual
as
renpy
“St
“G1
Lal
ZI
VI
SEI
ZI
Z 91
Zg1
QI
“ZS1
£-E1
ZI
S9
89
8
AL
£9
Sg
£91
£91
S61
“31
zz
S-0z
bgt
z'61
‘as
“andy
gs
9
c9
zs
oS
29
19
19
6S
oF
r9-09
6
Buipest
‘O1pAPT
i8t
1
9
0
81
a1
9
Oo
i381
zi
9
0
qidoqg
GSoSt 19/S1/11
S€g1 19/1/11
oozi 1S/0€/o1
oSor 19/9/11
OShi 1S/f1/11
Srgo 19/S1/11
outy 23eq
jaljig,02, 1U0S.e8
ial lcs9e. 1h, 0s .9e
18,02 iage
$798 ,¢Z mlalege:
Wor, LlaOe GIS gf
PTL LleOS SIS EQS
CM) CN)
apnysuoy apny yey
(panuyuor) NWOLAV ‘d
(panuyuod)—'t ATaV
cee eeseeeesecereececceeses tO
secereceses Keg ayeadesayy ‘uIOog JUsy Jo seq
veo e eee e cere eeseecceeseeeeesoess TBOUS SaLIjIY
SiS tebese nee pene eter elcaerek cece cieccde ee AAA SUIT OTAS [eoys S][eJI2AQ
renee eee eee eeeeeseeseeceeeeres AQT
SA eebenees cpp siehe ose ne ic arse ACD GT U19}seq ‘Keg Meys
uorjdi10saq
Be a
uolzeyS JO UOT}ZIOT
149
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
“
£1
FI
4
c,01
IVI
II
+6
+6
+6
Z01
S41
¥'gI
£91
£91
£-o1
Liat
zor
Z:91
QI
PgI
OI
gr
O'ZI
O7zI
Lea
Lei
L2z1
O7zI
Teo
V°QI
P'QI
REL
ISI
Sets
VZ1
eS—zS
Lo
cz
cz
cz
cc
ac
99
gq-0
ooSt
orgl
oOfeSt
oSIl
O0O0zI
6z£1
£V/9/11
LV /z2/11
4V/9/11
£V/9/11
LV /2/X1
1$/S1/11
(panunuo? )
Z:€0,94
££0,94
v;0,92
€:z1,94
(87.94
Ral LE LOZ
Sg;re gf eee eee reeses eres eteerereeeeeeeeeene ‘0d
Sg're gt ashame a rs OPaReRe pets ae AACS TAG ‘o8pliquie) yo yueydoyy
SEVE QF Braet aac SOU [| | “DIN ‘o8priquiey
S56 ,9£ Siecols/ uate leieiwisisiseieioe Nee te OLS) ‘keg oyeodesoy)
OS,g& **Aheg oyvodesayy ‘Yysry yIog Apooyg Jo "AA soyiu Er
y11,0S,8& Coe meee reese ee etree ereeeeereveee ‘od
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
150
58 = a oO CSor Zv/o£/11 wV OF .94 80.82 ee eee ee eee ree eee see eee esse esses 0g
ae om as O of60 zb/of/11 ,Sb,0€,94 130,98 Part ssaceur ico Sc Sky Semolog <quiog Asanq 116
8 aa =e oO Sbbr eh/6z/11 0% .94 9E,QIQQE “TARY OeWOJOg 4JuIOg Iepsy YO
/0 —— oe oO OI£I1 cv /9z/11 0 82,94 ,0£,O1,8° eee e reer eee ees ese e rere e rere eens 0g
u¥,0 Sis oe oO 6€Z1 Zv/9z/11 yO° 82.94 y0£,01 8° Cee reer een roe en ence er cern eee cece ‘0d
8 — —; oO SCor cv /9z/11 HOE 82,94 OL OL Qe Cee erro ee ence nsec eee eee ees ecceee oq
8 mo oe oO Ccocr ov /Lz/11 10 82,94 W0£,01 8° Cee ee ee eee see eee eee seers sreeeees ‘0g
8 a —r oO 6rr1 abv /Lz/11 yO’ 87.94 y0L,01 Se CO eee cece reser cc ccc ers cccc cere es ‘og
i —— = Oo VorL ov /Lzo/11 10° 82,94 OL ,O1,8e eee eee eee eee ee ete see eee sees ee ee ‘0d
0 2S, = oO Ooel ov /Le/11 y0° 82,94 yO ,01,8f eee e reece eee eee see eee eee eee eee ‘0g
8 facanes = oO CCII ov /Le/1t nO 82,94 Oe OL Be eee eee eee eee r reser cece eres ees ce 0g
0 eal = oO CSor ov /Lz/11 HOE ,82,94 yO ,O1,8e eee eecec errr erro eee ese seco ese ee ee 0g
8 a as Oo 9S60 zb/Lz/11 HOE {8704 yOE OI of sliaiietetevensloperadeleney cla vata/ ele asLalsiicienseacenetansy's ‘oq
WV ees = oO Z000 cv /Le/11 OL 8294 OL ,O1 QE Cee ee ree eee ees ese seeeeerseeeces ‘od
128 — ae oO €0go ev /Zz/11 Oe 82,94 ,0£ OL QE Cece e reese r ees sereeserecsec estes ‘od
ig ieee — oO Crri cv /9z/11 yOL 82,94 70,01 ,8F Coe eeereeeesreercece res eecrcce cece ee ‘od
(8 rene ea O oort ZV /92/11 70° 82,94 0 OL, 8e Cee em eee eee reese eee essere eee eee ee ‘od
8 = —— oO Coe1 ov /3z/11 nOE 87094 y0£,01,8F aiekeheilede otelelene.oneietelohe Se Sas eer
x el ee oO ESil eV /9z/11 WOE BZ,94 10 ,O1,8f eer eee eee ee eee CC ee ee ‘oq
i ome as oO CSor eV /3z/11 OE 82,94 70£,O1,8f ee eee sce oreo ere essence Secs bie olan selene ‘og
8 =, —— oO OOOL eV /Qz/11 OL 82,94 y0e OL QE S)'0\-e ee) 6.0: 6, /e)(6) (0) (e611) 8/01 \6)6).6' (6) 4,0) 4) =) lela) |0 18:19) «16 ‘od
3 aes — oO SfoOL ev /6z/11 70° 8294 POSH OMAQ Ga cacuceetoneens cco ceel Kose Neha lestusie Rensergeie cena ‘od
8 == — O 8€g0 ch/Oz/Il ,0€,92,94 ,0€,61,gf “*** +7" "+ “py ‘PuRTsy suoWOJOS ‘souaneg juIog
ine a = oO gavr 2v/of/1r ,,S€,00,22 OZ BE ********* ANY DEUIOJOg ‘Y9aIDQ sadog jo yo YO
iGes: ‘a's Surpeat qideq oury, 27eq CM) CN) uoldt1z0saq
Tenpy “AInDa ‘o1pAH opnyisuoT epnyyzeTqT
ey
UOT}ejs JO U0TJLIOT
(papnj2u09) NWOLNAV “d
(papnj2u0d)—1 aAlav I,
I51
IO WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
NO.
wm WMO
st IND
OF 0 OD Oo nwo +
Zl
VI
(49}}eu
Ja}e0S JO
s}iun) sut
-19}}80S
Ja}
-enbe
“18g
o001 jOll
006 pele
ooSI S01
o0zr VII
ze
so[pueo (a01n0s
“33 GqstT ~— yurod)
quaprour) durey
19}9UI0}
s04d.
pajsy SuoyDis
u9,0¥
tI
Vey
129
Ol Te
uZs9e
(os
19}}°T
uei9e
o1,£€
“a
Ger
“und
202
“avd
ze
(ea)
3, 38N
( panuyuod )
yidoq
Z;11,9S
O/gI,98
o£,z£,48
“
{08 OGa8
¢;Z£,9S
oer SS
yOZ 8% 098
(e698
(A)
opa}isu0T
0;6z,08 ‘°° *****puR[punoymaN ‘JoqreFY ayNoy PO
O;1€,08 *purppunoymaN AD “YAN HO ‘Joqrepy oyonoy
jOSSPs0S. 2 voreee sks SPU PURO Leh \uiog aR YO
POTTS OS.-<% os Seo = ae ‘+++*pueypunoyman ‘xo ade YO
O08 25 > es PURI PUROT MON 4rerqS IST I]9a
ZOEnTG ee Se DEEIPUROTSSN, ‘Keq Os!
jov1S “sre = pUBTPUNOF May \uiog youely YO
SP, 18 elele.ese a6) 0 (0) 9) 0) e s1e.e * IOpeIqey] ‘keg pow ‘roqie yy
CN) uorjdr10saq
opnyy7eT
ne
UO1}e}S JO UOT}LIOT
yo saqoarz fo Kouaandsunay Bumdsaquor UoUnULso fut fo saga, snoiunA—z AAV,
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
152
mmm
NonwrnS
‘© 00 00 oOo NO Oo 1n
(19}}eu
13}}¥0S Jo
s}iun) 3ut
-19}}B0S
o0f9
00g
OOST
0002
oocI
Jojou1
-enbe
-1eg
so[puro
~33 (aq3t
quaprour)
19}9UI0}
oud
rg
199
u98
(901n0s
qutod)
duey
uaa
yee al 9,4 I
AG If 198 I
j0S uVO£
133197 (lea)
a1 I8N
9
9
ZZ
i9e
9
ZZ
j9e
Ps
of
9
EL,
9
ZZ
9f
9
ZZ
i9e
9
qidoq
400,47 85
O;f1 VS
O}lisee
oS7 eS
oor VS
o£,0€ ,vS
“
0;0z SS
o;Zh,$S
(“M)
apnyisuoT
00,4000
4S¥,S£,0¢
0;ZE Or
Gz,Ze ,OV
“
Sor ,6F
ov £+,6P
0;0S ,6V
0;Z1,08
CN)
opnyiey
“*pue[punopamon “WYySrT pray Wog 31 HO
eee ee eee eee oe eee ec eee ere reese see eee ee
PEE
-punoymoN ‘loqrepy Ag-atu0Dg-woppaIs IIT
eee errr eee eee eee eee eee eee eeeeee
puepunoy
-MON ‘puR[sT yun jo “Ss salu or ‘xoiddy
“+ *puR]puNOJMIN ‘purysy weypem, PYO HO
tr reees ss epeTpuNoOJMON ‘TOGIeFT o}eSUry]IM T,
seeeesssspDuRIpunopMaN 4YysIT noyjes01g YO
**pur[PUNOJMIN JO “S alu I ‘pueysy [IND YO
treeeeesssspupIsyT oqieg 3G Usamoq jouUeYy)
wor}drs9osaq
ON
(panuyuod)—z ATAV I,
UOI4e}S JO UOT}eI0'T
153
IO WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
NO.
+TwNS
mom
VI
cl
nutans one
0 0
Oo0gI
0002
oozZ
0007
$z9
8
Pe ul ,81
yil,ae ae
(hh uz RE
vl u9,9
,O1,9F
01,01
vei81
wi8t
9
0
>
ie
9
Oo
je
it
9
Oo
81
oO
vz
4
9
Oo
Kq4
(98
9
0
es
if
9
4
oe
9
Oo
5
if
(panurjuo? )
S-6S 2S
j00,9£ 05
4
Bi9e ZS
jlo; oes
ul£,30,08
00,2708
o:Zh 28
y0F 90.85
£:ge,0P pele ele.0 76 bs Obe Chee eererssn e7e/e"5 eis) em pur]punojmaN
‘sap E11 JULISIP—]L Bg Sulsvaq ‘Qoey aded
“
QS,SS,oh creer rt teste puRTpUNoFMaN, “Wess 30qGeD
Sie e J eV. Cielo ese)eleslevealere)e se eleieiee iF a DiETPUNOT MO NI
‘sgyu 1 yURIsIpP—J,41z Sutsvaq ‘reads edeD
Ae
weve
(aero
over >"
reses+spuRTpuNOJMON “TOqIeFT SuYyof “3g
‘purrpunoyMoN ‘1oqiepy onbseg xny Hog
ejaeretaiateretenste: ‘ pue]pUuNoJMIN ‘Key ode) YO
++ +5 ++ DULeTPUNOFMIN, ‘eystavuog ode) YO
,00,10,0F *pueypunojMmMeN ‘ulIy Joquiny 4uiog xo, HO
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
154
II
gi
OI
oI
wN\o
2
8
(29}7;e0r
3937898 jo
s}iun) sur
-193}89S
0082
u¥ OF
0009 He
Jejour Sojpueo (a01n0s
-enbe = ="33. (3481, jutod)
-1eg queprour) duey
19}9010}
Od:
u9 EE
u2iBe
uQiee
Pal
B44
19}}9'T
7 Uy oD
y01,41
ee
190 I
ce
(1leg)
IL -IEN
eZ
i9e
9
99
i9t
9
ZZ
i9F
9
cl
i9&
9
9
o£
9
24
if
9
ZZ
ge
qideq
jeceas je SP
y2V,25,09 ,0€,SE,SP
wav 0S ,09 jan Paosa
,00,1F,09 ,0€,S0,97
yc0,Z1 009 a“ 1£,30,9¥
190,80 ,09 wel, 81 o9V
yOL 10,09 400,42, or
G;6S 2S £:9€ ,oV
CM) CN)
epnysuoT opnzyeT
.
.
.
.
.
enaraetecens "2109S eAON ‘ueg dItot 4S
**B1}09$ PAON ‘s[eoyS pesyasioxT YO
"reeese"enOIG BAON “IQ.p seig
"ses **-RpNOIG BAON ‘9YeT IO,p seig
"s+s*s*pr09S BAON ‘JOqaeFT AaUpAS
"****eI}09G PAON ‘hong eaG ‘IoqIeFZ AsupAS
ee ee eee eee eons
eee eee eee eee ree eeeos
®1}09S
eaon ‘Aong vag ‘1oqiez_ AsupAS yo sayrw or
Peet eeeeeeeeeeerereeeceeees pirerpunoyMaN
SapIu E11 JULIsIP—]J ,gge Sulsesq ‘aoey odeD
uondriosaq
— , -
(panuyu0r)—ze ATaV
U01}B}S JO UOT}VO0'T
155
N 0 69 19 mOwromnmanm
oe
WILLIAMS, JOHNSON, DYER
+N +
IO WATER TRANSPARENCY
wnt +
NO.
(panuiguo? )
9 ,0Z,08£9 ,QV,If,by **** "BOIS PAON “JOqIeE] XepyeET suey
mor 86 40,28 ult VE 0 paviOVeeo. “Ouse yy “8 EHO Se On ‘queg uyof TaSqAA
JaA0 oO URL PESE9 —,Z0,6E FH *** yO TIO Aq “eOIG BAON ‘soqieyy XeyyeH
00zg LSE yor ntl oO uVZ,QE0£9 98,1, vb °°°°" °° I}OIG PAON, ‘xepyey ‘uiseg psojpogd
ges ueeV oO ul2 BloP9 445 ev WW eee cence ene res eset eee seseeeencene song eas
oa ze < ‘JOqIeET Joys ‘eIO0IS PAON “Yoel saxUeA YO
ooSz neice uV Qe nll Zz 0 {2v,Z£,19 ,00,20, Sh **eI00G BAON “JoqIeyy AryUNOD Yo kong aS
nc,OV uBio Oo yOl,eV 19 yan Oloa’. ale) vile sa6) 6eSee, 5) sro. eleva’ ae) ee sieie5s le ‘od
0002 Of uV 92 aeyle 0 jOUCVIO. YSSjOIGSY <° "8 Sek er SENOS MEAON ‘roqieyy Asjunoz
oozS 2,08 We nLljee 0 {00,1500 LO le sy 0 ee o0S EACN fosuey adey YO
9 WOE OE ,€9 ,,0€,01,0V “°° "********eNOIG BAON OBpaT ssoly YO
VOL. 139
NOOO MAS
~
N
a
©
0008 ull tz u9 £1 wil 2 0 u©1,12,89 £501 ay, pip eo SORT S3q usxUNS Jo ‘Aeg oystse f
Baie’ u9 le o y£,10,89 00,11 hh *‘asnoyyysry YoY 34989q “IJV JO ‘'N soy +1
00g9 aS le ce 0 —,W8,P1,89 81,11 bh «+ Aeg suemyouerg ‘purysy yNq apWwT BO
oohL —_ 8,62 uesal 400 oO 721,068,989 ,02,Q1,Vh -°°°+7°° °° °° Keg PED onjg ‘pueysy Su0oT HO
MISCELLANEOUS COLLECTIONS
©
©
oogg Wee, Ojte. 11,0 0 (2106.99 81,01 ohh o:7 °° 81 reds “Yoesg UIssoUlDs3q
o0zr LiFe yO1,0z Z,z1 oO 77l/0e.89 G2, 8l,VV ose keg Ip ong pues; ou07 TO
“acd “nda
SMITHSONIAN
OQ Oon
©
oozb ub £% av VI v8 0 us¥ £8.89 0% be vy °° °° Sales Ss oureyy “JoqieHy TPH eng
(49}}eUr — Jayour Safpueo (a91n0s 43339. (ea) yydaq CM) CN) . uorzd110seq
sayjeos jo) 60-enbe=—=— ="35: (94.31, = urod) ‘HCFEN apnyisuo0'y epnzyey
s}iun) sur = -1eg yuepiour) duwey ———— err’
-49}}89S 19}9WI0} U01}e}S JO UOTIBO0'T
No) “oud
re (panuyuod)—z Fav
157
NO. I0 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
0g
oI
gI
oN
FI
VI
S69) SF
0009
oors
0002
00z9
oorg
oozZ
Pp £1
“cd
Vev
“cd
u929
9,42
u Bie
z,Ov
bond
I 61
wil 8
vor
“od
43,41
u9,41
u& Be
Teor
dled
adit I
0,4
F601
“avd
C11
“ard
198
ual
as
9
oO
4e4
if
9
oO
ig
9
oO
4c
j9e
9
Oo
184
i9f
9
oO
eh
ge
9
oO
24
ge
9
0
9
oO
ve
9
Oo
(panunyuo?)
y0¥ IE 09
u81,00,,09
ye ,12,L9
700,87,89
9,02 <9
98 4E.89
181 ,00,69
a“ ZV ,9z ° 89
u2¥ 97.89
plsjeaiccy SoGigats then gsi te e8)259/ 28 SOUTB TAT ‘loqiepy pinbewog
u81,85 o£¥ CC eTaD,
08,10, bb
,00,V0,,v¥
S:90, bb
08 L0,¢b
00,8007
u9& 80, .¥V
u9E 20 PV
(MITA T-S) ‘TaSyM PouueyD ysng OM
alee "6 @eid\ere © 6 6 elekmiele * Apuny jo keg ‘ynoj;L
a\elelexelereteve “keg IH nT ‘purest suo'y HO
poate oc is STE, pjerowig jo ‘N Soyrut of
USES En SAO GH e9S pueysyT SUeMS
“PA VIO, 1as4M Aong seq joosqousg
Stee letehse cise} CS OGIE LT puryst SUeMS
en
%
é 99
, 3 it
5 € 9
> 46 ov oO y00,2V,19 ,0€,2V,€0 **""****" e095 PAON ‘TOL 310g JO }sS¥aYyNOS
gl 2
2 y¥ 0% ejb oO
° FI 9
5 y 4,01 yayel Wee o yOl,Z1,0Z ,SV,V EV *s***seeerreese* Sure ‘apIsoloq-ynowyeq
fx) €1 199
i SI 9e
o £1 9
= 00zg ales 0c u¥ £1 oO 4£1,00,99 ue¥ OV eV eee reese ere ener eee sees "y90Y yey pe 1H#
5 IPPd ‘et09G eAON ‘a0ueI}Ua Joqiey] YJnouse x
& 3 Vs
e 6 9
4 oor9 +04 S,QE Sv EE 0 OV FF ,99 ,SV,9P,€F °° ****** B09 BAON “diy IST] JoyxonT YO
o gh
D S jw
a e 9
Za o0zZ u89¥ jof 01,51 0 ,00,Z£,09 ,00,05,€h ********""**"yySrT pinbeuag yo ‘keg suyof
< of 9
7 jjA® u9,¥ 19 oO u91,40,99 Aol Osnev ey eee weer een eee ‘0d
3S
Dn Ze 9
a A) uv,¥ KA 0 191,40,99 ,S1,0S,€¥ *°****"*****eMooG BAON ‘s0qIeT]T YJnowsez
g ol 9
a WERKE Lt, le 0 OL, Le 009 - 1S es 6r V eee tiriens oo ae: JOGseEy pinbemod
(19yeu I330Nr s3]pueo (301n0s 39}}9'T (leg) qideq CM) CN) uor}d110s9q
qoyyeos jo) -enbe = ="35. (3481, = utod) ‘WH VEN apnyisuoT opnyjey
syiun) Bul «-1eg = =6juapiour) = dwueq SSS SS
-493}89S J9}oUI0} uO0T}eYS JO UOT}COOT
sold!
co
LA (panuyuod)—z ATavV
reat
159
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
ons ‘©
II
Si
ve
eI
£z
gl
II
or
DE
ZI
II
€1
oogy
0002
oore
00772
00zze
0009
0088
oozZ
ayae
“ard
Cee
“nde
VIZ
“acd
Ler
Clie
€,6€
“ans
¢ 21
bhesd
v8.81
u9,21
“acd
b rr
“aed
jue
VII
“aod
z,oVv
“and
8
016
“ j
“and
L v1
Mia
(panunuos )
nav ,Qz,02
u¥S,0V,S9
Wo eee OG
00,4104
VS,2z,0L
uVe,21,04
y0F,ZE 69
195,¥1,04
199,8£ 09
190,21 £0
130,07 ,£V
Ql ,OZeF
¥z,972, £7
“ 17-0
iVS,Le,0
)SE,2£ 8
100,88 EF
28,68 8F
U¥S,1P,€F
SSUSIEA- 2.800018) U0) Ce IE sae S ONG ELTIG) peasy pled HO
vib pr iid case peed sages ane
BAON ‘a8pr] omynog YO (YT) 1ASHA\
“8 ddz%, (MTA) Song astodiog edep
eee terete Hehe lee theuege ee cleo SPO IO ayeuM HO
.
.
.
.
.
“TASUM «2%, (MTA) POY AUoWY PIO
ee eee eee er eeee *WUSIT peop purpji0g HO
.
.
.
.
.
veeeeeeeeeeeeeeeessssDuBIsT UdaID 12INO HO
cece ccc ccce ‘aseioyoUuy Joqieyy puelyiog
"MG 91, (MTA) ASYM POM Wejueg YO
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
160
Oc 1h
ve
CI
eI
Lz
IZ
ZI
Ol
Z
ZI
gI
SI
VI
Ss
Z
(49}}euL
49}}e9S Jo
s}iun) sur
-19}}00S$
0034
0088
oozZ
Joyo so[purs
vende = -"3 (qq31]
-Ieg yueprout)
49}9U10}
“Oud
pG,0e
ySr
“ed
€,9e
“
Z,0£
“and
u Qi
wl 8e
(901n0s
yurod)
dureT,
u8,0e
u,91
6,92
“
6,91
a“
II
to}JoTJ
Ais
ue
6Z1
u8i6
(leq)
"a, -32N
8
jv
2
0 uB1,9F,04 90,882 °°*..2, (OTH) IaSUM “Sse ‘uuy odeD YO
84
i9e
9
0 u2Z9E04 = 42,0S,2V sree essey, GaodAinqmoaN Yo soapiw z'6
joe
9
O Ve, IV 04 y¥3,20,cV Ce sr Suene Aeneas cee CET ‘Ni ‘yjnour
“10g HO “YM 2, ANG [ASYM 3POM SHOT
(84
jv
9
O =, 08, VE,04 —_,be,VO.€0 °°" TK Fe, (ATA) TASH O8peT 440K
jor
9
oO 1 BZ,£V 04 1VZ,VO EV See eee ee eee reese see sees ssesene ‘od
jor
9
oO u8Z,£ 04 FV Oe Yee sees eeis ees steers ee erees ‘og
ve
9
oO WAUeVeOe y¥oVO.eV Sn tonr eh ache “HN “toqiepy YNoUIs}I0Og
84
jbS uv (B04 90,21 ,€V RS SEe BP SOS NN deka ceenomoNe Sek s Sarees LT] '9) peasy pled HO
qidoq CM) CN) uorjdr1osoq
apnyisu0T apnyyeyT
$$ sss
UO1}e}S JO UOT}LIOT
(panuyuor)—z AlavV
161
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
(panuyuos )
6 ve
6 9
ooSZ v0,81 ,O1,z1 ese O yit,0V,04 ,SP,0F,1F *****Teued pod ode) ‘souvsjus Aeg spiezzng
zI jeh
or oe
ZI 9
00zg Bejae. 100,61 40,0 0 u¥Z,32,04 ,6€,10,c7 **""*'''*** keg syjasnyoesseyy ‘Aong [ASyAA
v 4
5 i9f
£ 9 AV 99 {80,24 Pas cheese? ie ole nl wie ol oasis HUT CmOFaNN CT CYEREO MER
v 184
II i
eI 9
o0z4 u8 ZF u9,8t u£ 01 0 u9E,9E,04 =, 9E,9TeF “ts s**t dIYS WS] uojsog Jo “qy sayiu 9°9
8 8?
1 9
oozZ {Ole jet ROS 0 nV1,O€,02 iVi,ve ev Cem meer emer e meee ener en eaneassssnens AY ae
(ATA) JO ‘W Sop € “sseyp ‘uuy odey YO
€1 Bl
cz oO
SI Ql
Oz Oo yV7Z,0V,02 y07,90,eV ee ‘Od
zz 81
oz oO
Iz Ql
vz oO 1VZ,0V,,02 OT, OF eV Dee em wwe wm we ew wee eee eee ‘od
IZ 81
ez 0
ez BI
Sz 0 yVZ,0V,02 - ,,0Z,9€,27 “*°° °° °S “ON UeD “Sse ‘IoqIepZy Joj}s9dNO]y)
VOL. 139
MISCELLANEOUS COLLECTIONS
SMITHSONIAN
162
or
N
SI
v1
S
9
9
gz
0%
ez
o£
Sz
cz
oz
VI
£1
(19}}e a
49}}89S jo
s}iun) sul
-19}}B0S
iJ9}om
-enbe
-1eg
0006
0002
o0zg
0062
Sa[pueo
“3 GUI
yueprour)
419}0UI04
“Oud
“aed
aCe
ail
ue Ol
43,02
(90.1no0s
yurod)
dweyT
u9,¥I
Pa
uQiae ies
ajee
sol u¥9
ja Le Hs
419}99'T 4 a ee
ist
joe
9
0 woe, ,0£ ,0V,1€,17 *****punog psedourA ‘pur[sy] euameyseN YO
9
oO
9
o yor Ve,14 ,OV,11,1” ***puesy yorg ‘(puog eS yeoiy) soqsey{
we
joe
9
oO yor,vI,1Z ,v&,61,1F **punosg pieAourA pue pur[sy xo,g usaMyoqg
8
joe
9
oO 00,0704 he QCo1¥ Coe ee ese tee eee soe eee eee preAour A seyqIeyy
‘1OqiefFT UMOJeSpy Yo ‘suor ..O1,, (MTA)
ev
ve
2
oO 00,07,04 12 ,32,1V Sere ere eee eee ee seseeese presour A. seyIeyy
‘oqiepT UMO}IeSpy Yo ‘Buo0ry .O1,, (MTA)
vs
joe
9
O y0E,0S 02 400,18 IV *plojped MON yo ‘Aq SpreZZN fra (MTA)
qideq CM) CN) uondrsosaq,
opnyisu0T opnyyeyT
———————— — ——— eeeeeeeeeeeeerererer— ee” oon
UWOre}s JO UOTWVIO'T
(pamuyuor)—ze AIav
163
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
se
ze
OI
of
II
or
IZ
oz
of
Ol
VS
“ed
u9¥
094 II
19,9
Ailee
uB8i8e
yOl,Le
ut 3,€
lie
Ly
ave
ue
wO1,81
pil Zt
9
oO
(panuijuod )
19e anges
07,002
00,98,,£2
0V,£b,£2
145,00, 72
uBV 41 oe4
12,69
jOE,f1 cL
ySf,vh 12
0£ ,07,0V Poem ree eee eee eer e reer esters esseee ate Pa
(MT4d) Ang WSsryT yoy AanqsmeyS YO
nOV,9Z,0F “°c e ttt ts HOOR Apues jo ‘AA s8es0=uy
yOV,OZ,0F “erst esses dS FYsYT puryjoog Aurora
‘jauueyD yyNosg ‘yoeoidde soqieyy YIOR MON
u9e,1e OV
yo1,Vo,00
u£f,S£,0F
{OV,,0V
0° ,25,,0V
uvv ¥O,1V
teeeeeessKemepoy req yo ,‘b,, Long [asyA\
"purest Aouor Jo ‘sMoIIeN JOqIePZy] YOK MIN
.
“purysy eat HO .'b, (ATA) Song TasuAA
slieteie exsterss “dys ST yox/onjueN yo SoyTut 9
“ss uoydueyasplig yo [Pq .Vz2,, (ATA)
Fettteeeesseeeeseeeeeepumog purIs—T Aol
aouRIyUa “YS so Fe ‘suor) .£,, (NTA) HO
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
+
\o
4
eZ
89
6r
es
LV
oS
PS
es
09
29
£9
40
se
ZI
Ol
NO HHO
HH
Z
ol
(1933 eUr
4a}}B0Ss JO
sun) sur
-19}}89S
ooSz
yOL,eV
u8,0V ue le jel
49jour so[pueo (991n0s 19}}97T (ued)
cenbe = "33. 9431] yutod) ‘47EN
-1eq quspriour) duey
191910}
soldi
joe
181
oO yasjeloae
joe
81
0 an eeoae
of
81
0 Uaajecone
joe
i831
9
O yesgeegne
Ql aa ecees
jes
9
9
O 407 V1, be
134
i9f
/9 Szjlz,22
of
9
0 WO0/OSi5E4
ev
le 00a 62
yideq CM)
apnyisu0T
2¥,L1,68
i2v,L1 68
nev £1,608
y2V 41 OF
ycv ,61,0€
400,81 ,08
eee
.
ee
‘oq
- oq
+ oq
--og
"TIAL atemejaq ‘Wwsry uyof dig yo ‘1 ‘eis
See eee eee seer essere eseesesesesesesseen AWD
onuepy Yo ‘suo (ATA MO) Jo A sopra Fr
6;9£ ,6£ eee eo ee ee oer eeseeseeeese een eee uoAury uospnyy
7 2¥,9V ,0€ eee eee
yO ,0F OV ©, uae) 6 ¥e10)|6/001e),¢/:8 .0).s (0\(e ele) 6) ele \-sin (0/6 \e181 00
CN)
apniey
ei oted Nene ncatdeyens uA mene ILLS ST yesouleg
«ly IPA
(MTA) Song 3ST 3poy Ainqsmeys YO
uoljd110s3q
eee
(panunyuor)—z ATAV I,
U01}z2}S JO UOI}BIOT
165
=R
DY
>
NSO
OH
, ay}
AM
NC
W ARE N E
SP
TR
R
TE
A
Io
NO. W
ooo
gr
ie
“
oor9
“
2
uF
ez
“
ez
cz
I
OO1
o0gf
o0gy
00S9
02)
nunu
(pa
Lizz 94
‘op
‘op
0z,94
‘op
‘op
‘op
‘op
‘op
‘op
‘op
‘op
‘op
‘op
$565 gf
‘op
‘op
ov ,20,6£
‘op
‘op
‘op
ayy
axyeades
d
** Split
“+ og
eee
"21S
‘Ol
yo
BMS
9 Yuiog u
desoy
ayeo
FROM
* 0G
ace oa
0g
“+ -0q
Po “olay
+ 0q
5 -0q
Onn
VOL. 139
MISCELLANEOUS COLLECTIONS
SMITHSONIAN
166
ZI je
OL vz
CL ooryv oO ‘op ‘Op ooo eer eee eee ere eeee reese ne ‘od
gi 44
ZI ire
ger oors Oo ‘op ‘Op sissies eee, sieie s syeleiseierelt 22055 SO Gy
41 ev
VI joe
Il oors Oo ‘op ‘op ES A ee LE LONER UY Cf
£0z ev
er ve
£71 0099 Oo ‘op ‘op eee eer eee eee eee eee eee eens ‘og
3t ev
£21 vz
1 0089 O 190,92 GPS Qf occcce es AEG areMmeElaq eoyS uMor1g ‘Z ‘eS
v weak q-0 G2;zzZ,.94 ZiGS QE cseeeeesee* KO ayeadesayy ‘leq asnoyyong
ze Be oO ‘op ‘op eee eer eee eee eee eee ee ee ‘od
ZI A oO ‘op ‘op eoeeree eee eee eee ee eeee ‘od
OI BS iV9 Oo ‘op ‘op eeoeereeeereereeere reese ee ‘od
8 uit i
ov q-0 ‘op ‘op eee ereereereeeeeeeeeeee ‘od
Zz 0 ‘op ‘op Peete eee e ee eeeeeceees Og
Vz 8 0 ‘op ‘op beet eee eeee seer seers cog
6 009 oO ‘op ‘op eeoeoeeee eee een eeeeenee ‘0d
ZI oosSv oO 190,V5 V2 72,95 ,8e eC ‘od
gl oozr 0 uQ0,VS "2 ,2¥,9S,8€ °° (pod eulreyy Ayoyey) Joqrepy Aepy ode
(49}}euL = Joyour so[pueo (a91n0s 19379T (lea) yyWoqg CM) CN) uolzdi1osaq
Jayeos Jo «-enbe= = -"34 (3431, = yurod) VU CVEN apnyisu0T apnyyeT
syiun) sur -seg yuspiour) = dwey SS ae
-29}}B9S 19}IuI10} UoT}eYS JO WOT}BIO'T
“Ud
(pamiyuor)—z Alav |,
167
(panuiyuos)
"8,11 ‘uIOog Uaa49S y
Iv
6¢
£z
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
QI
II
ol
gI
PI
II
¥gI
£1
Or
J9II yl
2 ootr oO Sz;¥1,94 SAIS.@e- © sky “***Song ylog uewysiIyT, Yo aim 0%
60er oO ‘op ‘op Pees e eee eeseeeeeerers cog
oogz Oo ‘op ‘op reteeee eee eeeeereeees cog
ocr oO Sz;v1,94 S*1S gf © 6. 0.6 0c: ole v-s)e ho © a0 eae ere eee eee eee eee eee Aeg
ayeadesoy) ‘Avg Uslojseyq ‘UIOg UeWYSIT, YO
oO ‘op ‘op Peer eeeeeeeeeeeseerers og
ogi O ‘op ‘op Fetes eee esses eee eeeeees cog
000 0 ‘op ‘op bette esses eee e seen ee eees cog
00z9 Oo ‘op ‘op Detter eee seers eee eee eres Og
zy
o£
zi
port wOL 0 ‘op ‘op ‘od
jw
ve
9
6,2 08 0 ‘op ‘op Petes eee eeeeeee eee eeeeee cog
jv
‘op ‘op ‘0g
ooIr ‘op ‘op ‘oq
oors ‘op ‘op ‘oq
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
168
u9
00
,VOL
yoil
wool oogt
ooos
ia
ge
vz
zz wiv
oory
€¢ oogt
£V o00£
Ooze
4¥9 wl
VI
6
Eg oor
OI
«98
O8Z
S oS9
(19}};em Jojem s3[puro (901n0s 49}}9T (ea)
Jayyeos JO. «0 -enbe=—-"33.: (9431, «=. quod) ‘HW CYEN
syiun) dul = -ieg = yuspioulr) duey
-429}}80S 4Ja}auI0}
“Oud
og
31
jel
9
0 $;24,94
oO $z;z1,94
0 ‘op
409
joe
oO 82,94
oO VZ,94
oO ‘op
oO ‘op
o ual 4 12,94
0 ‘op
i9f
Sl
oO ‘op
q-0 18,04
Oo ‘op
Oo ‘op
o weil I 094
yideq CM)
apnysuoT
405,00 ,.8f
405,00 .8&
‘op
{05 .8&
OS 8e
‘op
‘op
ll Ole
-op
‘op
15,98
‘op
‘op
S15 ,9€
CN)
opnyney
eC keg oyvodesayy
‘Keq ulojseq ‘ dueI}UA JIATY OAAA pUe STITT
eee eeeeer ere errors eee ese eee Aeg oxyeodesay)
‘Aeq Uso}sey ‘aouRIJUD YooIQ purjpoom YO
Steen ceceeeecsecess sO
eee ee ene eee ee ee eeeee sees seseeeeeee
(JO “M
gyi go) Aeg oyeadesayD ‘YysIT yuIog Apoolg
see we wee ee Aeg
ayvodesayy Gysry yIog Apooyg Jo “AA aw F
siete ee eeeeee ee eses cog
Fete t eee etee eee e ss cog
Ce Aeg ayeodesoyy
‘Keg ulojseq JO “A Salt OI yuIog Joy Jo "|
skein /aigrae sec niecaus/e exes th 6 menemagescuaesclSiAicies +0) (ay
eer e rere ree ee eee ee eeeeeese
Bibel eb 6: Aor Umm wom Serene VAaSTe sceguetove eee hes F180) (ay,
Se JEOYS ea IN.
oq
settee eee eeeeeeeeeeess sO
*+++Keq oyeodesoyy ‘Aeq ussjseq ‘Aeq MeyS
eae eee ese eee eee eee eee ee
uorjdriosaq
Se Sr rors mm
(panuyuod)—z Fav I,
U01}e}s JO UOTZBIO'T
169
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
IZ
cI
£11
zi
tI
SI
wat u8e
oo$
it
oozS §86SzI<
Oof1 ene, uQiee
81 ol
006 wh9e lial
oogt = ,01,8% 9,91
00z 221
a“
a“
Js
v,6
Py £1
4
(paniujuo? )
q-0 ‘op
2
oO $1:90,S4
Oo “Se vZ
199
ie
9
Oo ‘op
184
if
9
O ‘op
199
of
9 ‘op
Oo
O yOV SEL
vs
9 ‘op
0
99
9
0 $;10,S2
129
9
9
0 SZ) b2
jv
ve
9
0 181,05 VL
‘op
SL:Lb.9f °°
a) 8) 60) 06 eleva le 6) 6.6) 8,8 ee) a “dys wsrT WOY}ey-OAT YT
8h .8& *
SV .8E *
BY BE °
9i8h BE °
iVZ,OVQF °
efarerarereraanior Joqieyy Joyemyeoig “jad ‘seMoT
ween eee e eee ee eee eeeeeeresees “Og
.
°
.
.
.
.
.
.
.
.
.
.
.
‘dys WSr]T woyyeJ-ALy YO
Binh eiolo/ehexelngaiersieialnl"ioisoce\epun fe e/elolssnscQ) (ay
pie eee seed ic nthe esas Oo TT WS] WO} J-IALT
spsarelss rere eleneeeieicge OLS JST S][VJIOAO HO
soho eeieusiiel isis itasrcae ier arne na LUG JST S][VJ1IAO
**fong [eoysS Saigo Jo “q X “YS saylur gz
seereeees Keg aremejaq eoyS seo YO
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
170
09
gZ
~
9
99
ain
+cat
(19}}e
49}}89S JO
S}iun) Sur
-19}}89S
o000S
O00zI
0004
v1 S1
oor
ute 0 I
oo0gI 111,0V
i9jom saypueo (901n0s 19349]
-enbe = -"}3: (3431, + yutod)
“leq yueprur) duey
4939010}
“Oud
u9S
39
u88
1,92
(ed)
‘WEN
Oo 400,02 94
9
Oo ‘op
ibe
0 ‘op
0 480,44
oO v;¥0,92
joe
0
O £721,924
24
9
9
oO j25,UL
je
ev
oO 25, V4
199
joe
9
2 1Q1,10,54
i9f
9
oO jov v2
yidog CM)
apnysuoT
uel ZQe oooc ccc keg ayeadesoyy “ulog aA07>
‘op Peete eee eeeee seers s cog
‘op Pretest eeeeeeeeeess cog
PZ ,8E ieln gai abehiecuehel ghee SAsueneter otek (AUNT ay JULOg pur]
-Arey) JuIOg repay soddn yo ‘saany sewojog
SEPE QE -10qIey{ ‘py ‘espliquieg ‘Aeq oyeadesayD
S;6€,g£ a CDs
SM JedS yo) yuejdoyD ‘keg oyeodesayy
VBE -° °° dIYS IST S][eJs9AQ Jo “YS say zor
j2V9F suasieys kone ac piess ise Clee *ie a COMBS) IIEMETIC] HO
uVS,9V,8€ ***Aeq asemefaq ‘diyS WWSIT sejsAQ YO
Abge -°* dys WS] woyeq-aary Jo “yy sym 7
CN) uoid110s9q
opnyyey
HS
UOI}e}S JO UOT}LIO'T
(panuyuod)—z Alav y,
171
-_ _
H
aN HIN
NOK t+TO HIND MINH tine +t+tO
IO WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
NO.
(panuijuo?)
4)
o£
oosP nee Bie 0 ‘op ‘op ween eeeeeeeeeeeees sO
vs
of
oogt AOeL yee Oo ‘op ‘op sete eee eeeeeeeess OM
bs
joe
ooor jvI io)0 Oo ‘op ‘op ste eeeeeeeeereesees “Od
rs
ie es 0.9 78,
oors Woj9L 01,0 Oo ‘op ‘op gisletete eueimieneleleleteie od
bs
ie 6 Shee) 0.8) OF
006z v4,91 HA 0 ‘op ‘op woo ececesesee od
t
WwW
~
Oo
om
- II 07,94 OT Be arb etecenene eecvceeee eve) blelefelwreciete «© keg ayeodesayy
a — =z ‘QAI Juaxnyeg HO ,O91, CATA) Id
oorz Oo ‘op ‘op eee e reese se seeeeeeesee ae
ooze oO ‘op ‘op Se) sce) 56) efeleve ers) © 6p) 6 We 6) 6) 618165 ia
ooze oO ‘Op ‘op © eee ¢ 60,6 6 6 06a 6 © 8.0.6 0 00 8 0 a oq
ood oO ‘op 0) ) aispelieleveneriel/si sieleke cain gree copier ell
ooze Oo ‘op ‘op oe @)e)¢ 6-6 © 90, 6/5 6 5 6 © 010) 0 0s) vials a 0d
ooor oO ‘op ‘op ewer eee eres ee eee eseeeeere od
ooze Oo ‘op ‘op eoececeerrceesererreseeeesee od
ooor Oo ‘op ‘op eee rece r es eseeeeeeeseseee oq
000z oO “op ‘op wee teneeeeeceeeeeeeeeeess OC
009 0 40£,8%.94 ,08,61,8€ °°°° "°° “PIN “PURIST SUOUIO]OS ‘Qoustjeq jUIog
ooze 0 u9£,00,42 ,,00,02, gf **"** *FOATY IeUIO}OT ‘yao1g sodog yynouw YO
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
172
Ze
Il
£1
ZI
Le
gI
gl
gL
(49}}eUL
19}}89S JO
s}iun) sul
-19}}89S
000z
oors
0002
oor
$6 ual
a“
u09
OOO0£
oorvy ae 9
49}0ur
-enbe
-1eq
so[pueo
35 (G3t
quoprour)
41939W10}
04d
(a01n0s
jurod)
dure]
t3}eT (ied)
41 12N
Vz
4
ve
O yOV VP ,94
9
0 j2V,94
9
ve
0 v:00,S4
9
ive
0 ‘op
ve
9
0 93z0,S2Z
0 ‘op
199
joe
0 07,94
VS
joe
O ‘op
Vs
joe
O (07,94
yideqg CM)
apnyisuo0T
onl Age
VI,ge
Pj91 gf
‘op
/O1 Be
CN)
opnyzjey
Pete tenses te eeeeeeeees cog
"Ts ess**DURIST DUO} HIV Po ‘AIAN dews0j0g
Soeeie SP eaeats EAS ere Se TONES, deUI0}0g ‘Keg uoyoIg
“"TISUM «HH, (MTA) We TMD yey HO
““TISUM «SIAL, (MTA) [eoys 4ormuey HO
Pee e eee seer e tees eeeeeeeeeseeeees BOTS
purjsy xormuag “(STAT) Aong apsiy Ay
Kiger nrecrea ation Oat
eo eree ese ee eee eee eee ese eeeee se eesens
«9I,,
(ATH) “Keg oayvodesayy ‘uiog sepey yO
bet eteeeeeeeneeeee cog
@- G8) e) 4) 01660) 8 278) 86.0. 8%e).6) ee /e oe Aeg ayeadesoyy)
‘ART Juexnjeg yo O91, (ATA) IP
uorjdt1oseq
——eeeEeEeEeEeEeEeEeEeEeeEeEeEeEeEe———— SS SSS ee
(panuyju0d)—z ATAV L,
WO11e}S JO UOT}LIO'T
173
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
ZI
VI
II
o00zZ
008
0091
oSg
OO£T
oSor
oozs
009
o00S
ooor7
ul,9e 10,41
6,61 ,0O1,11
“ard
Vee ,,01,¥1
“re
(panunuo?)
OV 00,94
$711,924
‘op
9 I 094
12294
01,94
01,94
81,84
¢;S0,$2
‘op
112,94
“l SV,9£, g4
S£,S0,2E
S,80,4E
‘op
u9E 80, LE
OS tae Ze)
Oe 0S ,4£
[eT LE
BY 4E
‘op
ASE
‘op
esaineaneatanie eamiateyhsnanltanser8 8 Aca Oy iT
‘keg ayeodesayyD ‘spunoiy a[pprif 2Oysestoyy
beeen tent eee eee eeeseeeees sO
trrersresces Keo gyevadesayy) ‘speoysg squiiyy,
@ccencecs "ION qeiy yo ‘IOART yIOK qynoyy
siaceaie tos wc EED ayeadesoy ‘der JOM yO
"ss***Kegq oyeodesayy ysiyT dery YOM YO
anbeajoouryy YO ,TLz, (MTA) 40ng IP
Ur (ee 61010 s7eNe) els eleleia ie: e sess ISUM 9 (SOM)
yeoyS JojrenG) sJoJUTA, Yo sammy woy}ey-01
goqren() sow, 9# (SOA) ong apsry A,
Dette eeeeee eee eeeees 19g
O,8& *** JOA eWMOJOg Jo YNow “(YTA) A, IPA
700,R0,88 “Tt rOATY oVWMOJOg ‘uIog AouIg HO
ON ZI iB
a I joe
r II 0 ‘op ‘op Sisls areal mre eankyiasere ieee io FSO AU EIR) (GE
° v jvS
e € joe
€ 9
o0zI O L:SZ LE Baas BD TS +ss-diys yyBry eyeadesoyD
° uo Pa} O ‘Op me) 9) eatasgrbeunieaeasatgsyeieunaesutale SOG
3 8 ve
o . . @" 68. (0)'8. (6) 6) 6 6 0 © (pO G8 8868 (8. &
a $ 99% 9,01 0 op op od
a) g ve
S ¢ uv Bz ne Oe Oo ‘op ‘op eee eee seeeroeseeeeeee ‘od
= s ue Crt Os .
D> gz 008z uV Ae Ol Oo ‘op ‘op ee og
o
: 3 ive
4 > 085 a op Op 0 tteteeteseseseresess cog
S oI ike
a 4 oO ‘op ‘Oop 6G) @ 016) 0,014: P16) a. 00 fe), 860,19, 28, O28 ‘og
5 g€ oor9 e
- ef ‘ eovoeveveeveeee 82 8 &
a S ooZS oO ‘op ‘op sisienties od
Se ie aR enti
< 9 0009 0 ‘op ‘op ee od
7, bz
a at d e@eoeeveveesee ee .
9 Z o00$ oO ‘op ‘op eee eescee od
Z 9 Re . ree .
a Z ooz$ oO ‘op ‘op mielesereiniersiece) si oeielers oq
a 4 hz
= £ oo0gs 0 ,0V,60,92 ,S€,S0,Z€ **heg oxeadesay ‘spunosy a[PpryT 204SastOFy
(Jeyeur = Joyo So[pueo (a01nos 1a}qaT (ile a) yidoq CM) CN) uorzdttosaqy
Jaye0s jo «-enbe = = =} (3431, = yurod) V1 CVEN apnyiZu0T opnziey
syiun) sur = -1eg quspiour) duwey EF
-13}}89S 13}9UU0} U0I}2}S JO UOI}EI0T
“Oud
=
> (pamuyuod)—ze AAV I,
175
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
nN
£1
u89
19,8
07
uO
ooor
OOO!
osz
o00z
OOII
SzI
oogs
oorz
0002
ooor
00g
oozZ
ev
199
u90e
of
cooooooooceo
°
°
oooo 90
°
(panutjuo?)
$190.18
G;1F,0g
137,08
OF ,08
¥S,02
fh 62
gr 6L
WS ,02
‘op
25,04
G;£S 94
190,03 084
GSE 522
120,42
o;0r 94
01,94
GeZaoad
Sioz 52
Lzze,SL
o;S¢ 92
G;11,84
$761 ,S4
489,01 94
$:2z,0€
ja 0r
Gesie
{40018
j00,c£
o£ oF zt
jovert
ZjSV ze
‘op
o£ Sb ,zf
OS 2
“
a“
yVe Ol ct
TLELEE -
G2a,e6
o;zP ve
Size ve
0:35 , ve
B08
GZ;91,S¢
O;1Z,SE
0;9£,,S€
S41 59£
yOe,VS9€
ceccevcrreseusoreserd ug qysry suyof IS YO
See nee a sarnema teen) ‘pueys] puejiequing jo ysey
wale yojors\ehsvel Stet pec loean CFL] Ss ST yeuurares yO
seseesesdrus Jury yeuuears ‘vole YeuURAeS
asexerelatetetel0 eqn ia tedere ii: ake sada Xa) ‘yeuueaes jo ysey
Sheehan eed Fat inet ene ‘ON Aong BIS uojysop1ey)
Ciel eletoisi sl Kereinistelets yt E TT ie yo 2) AS ‘uoj}so[1ey)
ola/at onset e\cscae aol ieus hence OC VEE] oom) Ae ‘uojsofseyy
wee e ce een ee eeeeeeeereteeeres “OK
teeeeeeeee ss raqUIMS WO Yo ‘vaie uojsajreyy)
ee Oe ai z'O JuRsIP—J,OgI Sutsvsq
TASUM «MOZ, (MTA) Urewoy ede HO
Or sia eneveareuel s leneie sie, «cel chee pea LLU are que}
-sIP—,9re Sutseaq “diyS ys] ued SUIAL
estes vee eeeeeteeeeeeerrres QAIND WOUIEA
-Ol jo a8spq ‘drys qwsry ueq Susy YO
“say ¥% “sip .ZM,, (ATA AG) fond AM
steer eiel ere cow” eres aeae EO) =) "N ‘AUD peaya1oo0jyy
O10: «| 6)06 oe © © 6) 6 6. a) 0:6) 66 sisjoevici 2 SNOMOO es aee) yO
Sica lig yousdoke = cage ekasapelemeaan “+ +4aquy ayooR1I9Q HO
"+ SaqIt Z JURIsIP—T,Of0 Sulresq [ASU
aq 8 Ud (MTA WO) serneH ede HO
treseeeeees scot Q juRIsIP—] oz Sutsreaq
‘gsnoyysry sessyepE aded ‘sesayyep adey YO
wee eeeeeeeeseeeeeseeerreerees IOADYT OOPWET
ere eyepaliese hk a ‘aeale eles eal eoW sie ndiere a. (6010 616s ese e Soy tut
#61 UONeIS “H “D OolumooeUVsIy) JO 4sey
Sinica + soqtu Ez ‘punos yonjyiiny jo sey
She lekereiers yee (evar ene, ieiep se lexelehenerete. am Ig ‘aseq Sut
-ule1y, xouuy yseq wnydury “eA “Ya2ID 9]IT
VOL. 139
MISCELLANEOUS COLLECTIONS
SMITHSONIAN
oO
N
=
colt wn min culo
lt
Ss &
awroan
(19}}eu
419}}89S Jo
sjiun) 3ul
-199}899
joome
yO1,Sl
uBid I
ne al
VI
19,0
nejel
jeom
jooxm
u9e
joox
OL vi
uke
u@9
uv
«89
iJ9}ou
-enbe
-1eg
0002
oo0of
o007
00z9
ooze
S3[pueo
“33 (43IT
queprour)
19}ouI0}
ud
119
ov
ZT
(99.1n0s 19319T
jurod)
duey
oO
198
9
00
oz
o
j9€
ZI
Oz
4
0
4
4
4
o£
0
jof
g-0
O
0z
0
4
4
071-32
4-0
82
i831
0
Vv 0
VI
O
(lea) qideq
VFN
(panu1yuod)—e AAV L,
$/95,68
180,08
Ze ,c0
eizb 68
G:z1,v6
v;hz $3
e:zb.€g
(6£,+6
5/£1,88
Z;01,28
8£ 18
292 018
£1,248
(AM)
epnyisu0'T
i8£ 82
5/8P 82
OV 82
{20,02
80,62
V1,62
£91,602
/O1,02
G;Ve ,6z
Q;z1,0€
jO1,08
$;£z,0€
7,08
CN)
apnyiey
**eLq ‘odea asje.q Yo ‘uokurs iddississty_ plo
eee ee
srrseesessssaoueiua IddIssIssIf, JO
Sp Ey ngage oa) <2) (0 jo Jn
"s+ -990uRI}Ua IddIssissIfy Jo “AN Aeg
AAUME ie, CMODO) Jed One
gio ynoqe “xayT ‘UOJsaATey) YO yue_ pyeoxy
Vos Senos ss ete. Seren ues odey jo ptemyynos
te eceeeseeeeeesssepunoS 2ouUeMS PIeMISIA\
AP See SEPA SUT AN Aas (M14)
anu Otf ‘ong vag ‘sexay, ‘UO}saATey
“eTy ‘IGoJY Jo premyynos
vss? ely “ejooesued JO “qiS
Spite ries? Soc SIAN suyof 1S
‘ssopisyoop “epy “asodAeyy
“sep ‘ejooesuag ‘apisxo0q
worydt19saq
a
UOT}e}S JO UOT}LIO'T
~——-—+-—-
177
WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
10
NO.
TINH Oe iN ot
7iN
nm
oor9
oS9
ozit
uBOl
Be
u981
oo0f
vi
00g’
ue I
009S
At
oor9
aay
0097
“ Il 9
1302
(pamujuod)
481,40,08
‘op
ye ,49,08
z;Z0,08
Gj£7,28
425 ,20.08
346
195028
S£,90
Of $0
/91,08
S;0€ .28
S65 ,06
¢;10,$6
$;0z 08
OL {89,92
‘op
Oe 29 69%
phe 20.9%
L192
yOr LE 9%
oLe
SE 42
2 42
Cty ,4e
WY 42
G:ZV £2
{LE 8%
LE.92
92.82
sie Rieiele aia bie ost sie acmretein ioe CS (an
és potinin ta eeaiaee= "="! “OCT
STeiny Diane! ss “TSN ‘90d “ely ‘gyepiopne’| }104F
reeseeuiseq Suruiny ‘10qreyy sope[sioay 0d
cocceccceceseceeccsseooeDuUBIST Jaques HO
sep “Yovod poomAyjoH{ yo “vale aepsopne Ty
Wielsveiere) sfe'e sie lecermealsrsieererecis oleae ea ODT OVNI) jo yn
sue eve dase baat: oS 7 aa eUNe OMG
qs) ‘Ww F yp song vag Avg edwey, YO
reseeeeeeerstyD sndioy Yo ‘foorxayy Jo FIND
reseeeensriys sndioj Yo ‘JAIN WoY}e,-OOt
sjelsleieveisieresielere)* lens he KOET yeoys Pup Aq
asop JasuM (M14) “Plt ‘yovog WIM PO
seeeeeeeeesssg3psoyouy euluerend) Jo pus
"MN yo ‘Keq os0qsyTH yjnow ‘Aegq edwuey
eve are) eens Wats Leh ohe) Sucuaretoyls ere. 6) She yasare Aq asopo TASUM
«fy CMTS) TOUS 44S HO ‘Qarnd woyye-Ot
reseseeeeexg gy Gada Yo ‘oAmny woyyeq-Si
Be ner eue eye: a feretavenoletexous eye Ke leneKeL.e].crenmgen ius Aq asojo-
TISUM «8, CATH) Aong TeOuS P2zPH
ot
in
ro)
I uBiet
O%
z;S0,08 SISO Oe Se SOE ee Meee Soler SRG OLD Wel, )
‘Q0uesIqUS Joqiey yo ‘10qie Py SOpels19AY 10g
‘op ‘op onstesbtaveresat Ree: NaS RO Sarees
‘op ‘op acer sc oreseiaeny opens iepenoreass oe eee Ly (ay
‘op ‘op eRrorntliyy aN iaike ase eO ey
‘op ‘op Sybisie/ anh exoSsia} cA0\ek ovals) CSS cay yan
‘op ‘op NeSatodosec TO aye oseess Cele Sai Fay
‘op ‘op COAG SSRIS ie alas is ame m ITY
‘op ‘op cunt tat slants iat aI saat OT
‘op ‘op Deeeeeeeeee seen seer esses cog
‘op ‘op Pt tteeee ese eee sees eee es Og
‘op ‘op Preeeeeee eer e sess eeeeees cog
‘op ‘op Matos Resi aa oteeist nex mites “0G
‘op ‘op Shiels ae tar eee ee RIES gy
‘op ‘op Sia wip mig ereiass oi? aisle escheeie eines 0) (af
‘op ‘op pePeRPEGuGure rs Ie ela Le PSS tO
‘op ‘op Seer ela eee eee cy
‘op ‘op ephi Peeey baseicitess tog
ozis ‘op ‘op i ‘og
oos ja 20209 OF S092 Se EEC NSO Gn ea, .o[EpsJopHed. 0 n)
(49}}eur = Joyo sa[pueo (901n0s 19339] (leg) yydoq CM) CN) uol}d119soq
Joyeos jo. 0 -enbe=— -"33. (9431, = utod) ‘3 ‘JEN apnyisu07T opnyje7y
s}iun) sur -1eg quapiour) duey ——— -~
-19}}89S 123910} U0I}Z2}S JO UOI}EIOT
-o"ud
co
~ (panuyuor)—z aATaV I,
0089
VOL. 139
ooze
o0zI
oorg
OgZI
oogr
o0gr
QRZ
oof
oor
0009
oogr
0090S
0061
oorg
00z
Siz,
00gr
oo1s
oor9
000g
oor9
SMITHSONIAN MISCELLANEOUS COLLECTIONS
oooo0o0coococCoCoOcCOoe99900 000
179
att hy
NO. 10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
(paiiyuo?)
jov oO
QO Oo
gol oO ‘op ‘op beeen eee teense eee e reese cog
jov oO
Ql Oo ‘op ‘op ee ee ‘od
(02 q ‘op ‘op Peete nets eee eeeeee eee eeeees Og
jos o wSS¥S,c3 ,O01,8€,b2 scrcct epy ‘sesnzioy, Aq ‘Ay uapsedy
W918 000s O SS ,18 eb ve costs sam Aayy ‘Fysry yeoys yyWUIS
0 12,44 80,S@ **fong vag Aq asojo ‘10qie}y NesseN sdueIjJUY
LV jee
iV 9
oozz Oo
jbl |e
VL 9
ooge 0
je 81
94 9
oo0zz 0
45,20 1
Wile 9
oozS oO ezey uC eg -*eieisierein'e's"s eu leleievase eis tele oA era ay NT ees
{1d 0 $346 sOGSGE. °° ose Vereen elie SENS Ta Ol PASUMOnG S20)
j08
gee oO Sr1S5 42 G25 Sc ce eel ates eael eek Neen naan Cll dn11S yearn
oogr Oo $:V0,08 10,92 eee eee eee ee eee ee eee rere eeeeeeeeeeee yoeag
poomAT]OFY HO sop zc “ease “ep ‘oyepsropneT
o0o09S Oo ‘op ‘op Ce ‘og
ooze oO ‘op ‘op eoeee eee reer eee eee eee ‘oq
0090S Oo ‘op ‘op eee eww ee eee eee ee eee ‘og
ooze 0 $;$0,08 10,92 *"yoeag PoomATOFZ YO vase “vpy ‘apepsopneT
oozb ) $:90,,£6 GsHO Ou Ss eerie cies ei KONG Bas OMeIg, BUES
jae ‘op ‘op eeeeeee see ee eee eeee ‘oq
VOL. 139
SMITHSONIAN MISCELLANEOUS COLLECTIONS
180
oe
=
e
£1
+A FR
ey
(293721
139}3898 Jo
5ylun) 3ul
-193380G
oor’
ooz~
oStz
oSoS
00z9
0009
uBi4
FI oor9
u801
iepll
uBio
oooe
00z9
Joyo so[puro
-enbe =~}. (34,81
“ted jueptout)
4J9}9umI0}
oud
192
u9 0%
se
i9F
jeol
40,21
(SLz
ie
igh
(2041n0s 13}}9T (lea)
yurod) ‘WEN
durey
9
oO
9
oO
9
oO
9
Oo S-1F,6g S/£2,2z
O 21,68 ZV ce
Oo {S1544 {OV ze
oO £1.98 OV ,2%
0 £701,642 1/08 ,22
0 ‘op ‘op
O Hil ,138 ho 0
0 z;0£ 1g giro, tz
{aL
9
Oo 0% ,.c8 ,00,,€2%
0 j£2,46 $'60,£z
0 46 ove
O j0f 42 40, ¢2
) Lov 42 0% ve
fo) £:2£ 18 jleaVe
O ‘op ‘Oop
0 oe VS 23 —_,, O1, SE, FZ
qideg CA) CN)
epnyisu0'y opnyyey
eee e ese eee eens
eoeceeereecncecce
ee eee eee oe eeee
‘ISePIOYIUY Jooy UeIDeLY
eee eee ee yueg aypedue)
“coo*suRG BUeye” jet)
seceees TQuUeYy) Ueyeon_
veeeess TQUUeYD SP[OYIIN,
teteeeeees cog
*eqny ‘sezuejeyy ye YOod
cata see eqn, ‘sezurjyeyy HO
ee eee eee Jodie yy CUBA TT
‘od
teres eeesOorXaT, JO FIND
PC
‘shed Avg SuOT jo ue900 ot} JO onsSuoy,
TreeessessspurysyT SoIpuy “YSiq PPI
eeeveee
eevee ece eee sees
eevee eee
sreeess TROUG UPOTIOWYy
eevee eee ‘og
sl elieieheiace "eT ‘sesnj10 7, AIq ‘KO uapjzer)
wor}dr14s0s9q
a ae
(papnjIu0d)—z ATaV J,
UOT}E}S JO UOT}LVIOT
181
NO. I10 WATER TRANSPARENCY—WILLIAMS, JOHNSON, DYER
ain
N
“iN
o1,S
18,01
a“
z,O1
‘
VI
1
oooe
ooor
ooor
ooo0Z
oozs
ooos
0099
ooge
o000S
190
jooe<
uBiO
CI
272
4
u9 BE
~
oor ecoooeoooooo”g
°
sv,rz,r6
60,56
180,809.96
180,90
£0,£0
137.86
oS ,16
jb1,46
ev ,46
‘op
‘op
uiZ,8081
G;11,61
4a0,21,01
S1,O1
01,01
9)Sz,61
G;€z 02
0:72, 12
j41,c%
‘op
‘op
we cecvecicesecsecscseessr"** JOANT SOI[EOD
-ezyeo) ‘cH Jag JoOqieH ‘OolXe CMM
‘sop S “eprieg Bo0y Yo ‘gysoduiey JO JN)
seresssopisyoop ‘OqIeF{ ‘OOrxXey ‘ZMIZ) PIDA
.
.
.
.
.
.
seecooeooees1OqIepy ZNID eloA HO ‘OOLXO
seve eeeees COG
ssssgupaduiey Jo Jpn
*s+sgyoodwedy jo Aeg
ver PREIS, FOdoT 30)
‘oorxoyy ‘oorduiey, YO
reetees cog
reeeees cog
or
=e es : = nee
Skies eae . P. ees = - 7 #- ee = 2 = E
eet = Se sere
ao
—
ae
aSF-et-
>
1
J
c3c-se ted
, er
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER I1
(Enp oF VoLUME)
A CLASSIFICATION FOR THE
BIRDS OF THE WORLD
By
ALEXANDER WETMORE
Research Associate, Smithsonian Institution
(Pusiication 4417)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JUNE 23, 1960
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 139, NUMBER 11
(Env oF VoLuME)
& CLASSIFICATION FOR, THE
BIRDS OF THE WORLD
By
ALEXANDER WETMORE
Research Associate, Smithsonian Institution
SOeCOCeeg
(Pustication 4417)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JUNE 23, 1960
SMITHSONIAN ,..0. \
INSTITUTION YUNA & ttf,
THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.
AV CLASSIBIGATION: KOR THE BIRDS; OF) THE
WORLD
By ALEXANDER WETMORE
Research Associate, Smithsonian Institution
The principal additions to current information that affect the ar-
rangement of the family and higher groups in birds since the previous
paper on this subject by the author was published (1951, pp. 1-22)
have come in the fossil field and deal in part with the earliest known
forms of the Jurassic and Cretaceous periods. While there has been
much discussion of family limits among the Passeriformes, with
considerable spread of opinion as to family limitations, in the main
these have been expressions of individual viewpoint, without com-
pletely firm support in the new information offered. Valuable new
data that are accumulating from many sources relative to this order,
where they are completely decisive, in the main suggest better align-
ment of existing families through shift of genera from one group to
another. The great majority of the many species still require detailed
anatomical study.
Under the revision of the International Code of Zoological Nomen-
clature as adopted at the Fifteenth International Zoological Congress
held in London in July 1958, now in press, a new rule provides that
family names are to be based on strict priority in publication. There
is no attempt to follow this requirement in the classification presented
herewith since the final draft of the Code was not yet in print when
the paper was under preparation. It is apparent, however, that ac-
ceptance of this new proviso, while intended to establish stability, in
the beginning will bring many changes in current family and higher
group designations in the class Aves.
The following notes that discuss the more important changes are
added to material from the introductory section of the revision of 1951
where this remains pertinent. In the classification at the end of the
text the fossil groups are enclosed in brackets to enable their ready
recognition on the part of students familiar mainly with the family
and other categories of living kinds.
Archaeornithes—The recent careful study of the specimen of
Archaeopteryx in the British Museum (Natural History) by Sir
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 11
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Gavin de Beer has added greatly to knowledge of this bird through
application of modern methods of examination. De Beer (1954, pp.
39-41) has outlined clearly the resemblances found in the two nearly
complete specimens preserved in London and in Berlin and has shown
that most of the differences between them that have been described
either have been misinterpreted or do not rate the value that has been
assigned to them. His conclusion is that “proposed generic and even
specific distinction between them calls for very critical examination.”
In his final statement on this part of his study (lLc., pp. 50, 57)
he unites both under the name “Archaeopteryx lithographica Meyer.”
In brief review, formal recognition of the two specimens as repre-
sentative of separate species came when Dames (1897, p. 829) named
the one in Berlin Archaeopteryx siemensii. Petronievics (in Petro-
nievics and Woodward, 1917, p. 5) considered that differences be-
tween the two were of sufficient weight to separate siemensii tenta-
tively as the type of a new genus, Archacornis. In a later study
Petronievics (1921, p. 10), after further consideration, was definite
in establishing the two in distinct genera and added that they might
“vielleicht sogar zu zwei verschiedenen Familien gehoren.” In a more
detailed account (1925, pp. 67-69) he placed the two in separate
families, which he maintained later in a further review (1950, pp.
118-120).
The major points on which Petronievics based his two families have
disappeared through the information supplied by de Beer. There re-
main, however, distinctions of size and relative proportion, the London
specimen being about Io percent larger in general dimension, with
the foot about 25 percent greater. De Beer regards these size char-
acters as individual, to be attributed either to age or to sex. Steiner
(1938, p. 292), who also has considered the two identical, says that
in his opinion the Berlin specimen was a young individual and a
female, in contrast to the London example which he believes was a
mature male.
While my personal study of this problem has been confined to views
of the London fossil and the nearby cast from Berlin in the British
Museum, additional comparisons of casts of the two in the U.S. Na-
tional Museum, and examination of published figures, it appears to
me that the foot of the Berlin bird not only is smaller but also has the
toes of different proportion in relation to one another and to the
tarsometatarsus. The wing elements in the two specimens appear
quite similar, but the entire leg in the Berlin bird seems more slender.
It is possible that these ancient birds, like some reptiles, continued to
grow in size for a longer period than is true with modern species,
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 3
a factor, however, which must remain hypothetical. Steiner’s supposi-
tion that the London specimen is male and the Berlin fossil female
is equally speculative, since if sex is assumed, the reverse might be
true. While the male is larger than the female in most living birds,
this is not the universal rule, and as reptilian characteristics persist in
these earliest known avian forms it must be remembered that in
reptiles it is common for the female to be larger than the male. As a
further contribution to available information there should be noted
the analysis of the primary wing feathers by Savile (1957, pp. 99-101),
which points out an apparent difference in wing formula between the
London and the Berlin birds. This recent observation if accepted
would indicate rather wide separation, but, on the other hand, if denied
would serve to bolster the conclusions of de Beer.
A third specimen found in 1956 near the point where the first ex-
ample was discovered shows mainly wing and leg bones and vertebrae,
in addition to feather impressions. It has been described in detail
by Heller (1959, pp. 1-25), who finds that it agrees in size and char-
acters with the one in London, so that there are now two of the
larger form known. |
It is important to have a modern study, like that of de Beer, of
the Berlin specimen, to add to the data assembled by Dames. As
matters stand, the three known skeletons present an appearance of
differences sufficient to mark them as two distinct species on the basis
of criteria found in the osteology of living birds. These data, for the
present, appear to warrant recognition of two genera, Archaeopteryx
represented by two specimens and Archaeornis by one, which, how-
ever, should be united in one family, the Archaeopterygidae.
Ichthyornithes——A recent study by Gregory (1952, pp. 73-88) has
severed the long-standing association of Hesperornis and Ichthyornis
in a superorder separated from all other birds known from the New
World through the possession of teeth. In brief, Dr. Gregory has
shown that the toothed lower jaw fragments allocated to the skeleton
of Ichthyornis dispar Marsh, unduly large in proportion to the rest
of the skull and the skeleton with which they have been associated,
in reality are not avian but are those of a small mosasaur. Two other
jaw fragments placed by Marsh with Ichthyornis anceps and I. victor
are similar, so that all these specimens, which have the teeth in sockets,
are identified as reptilian. This leaves Hesperornis as the only group
of Cretaceous age in which teeth are known. To give a balanced
treatment that will emphasize the important characters of the birds
concerned it has seemed appropriate to establish a suborder Ichthy-
ornithes for the Ichthyornithiformes, separated from all other birds
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
by the possession of biconcave vertebrae. I have given a somewhat
more detailed discussion of this matter elsewhere (Wetmore, 1956,
pe 2)i
The penguins.—The question of the weight to be given the peculiari-
ties of uniform pterylosis, extreme specialization of the wing as a
flipper for submarine progression, and incomplete fusion in the meta-
tarsal elements, as well as such other details as erect posture in stand-
ing and walking and the anatomical adjustments involved, found in the
penguins, is one that has merited careful review. It seems reasonable
after this examination to retain the Impennes as a superorder, at least
until we have further evidence through fossils as to their line of
evolution. It is necessary, however, to remove the fossil family
Cladornithidae, since Simpson (1946, pp. 24-25) has found that the
two genera Cruschedula and Cladornis placed in this family have no
apparent relationship to the Sphenisciformes. These two, described
by Ameghino from the Deseado formation of Patagonia, now placed
in the Oligocene, are based on fragmentary, considerably flattened
metatarsi. The descriptions and figures that have appeared thus far are
not sufficiently definite to demonstrate characters of importance in
classification. However, from what we now know these ancient birds
cannot be considered as ancestral penguins of terrestrial habit, as has
been supposed. The only suggestion that has come to me is that pos-
sibly they may belong in the order Pelecaniformes, in which I have
placed the family tentatively in a suborder Cladornithes (see p. 25).
The Neognathae.—One important result of recent studies has been
the allocation to the Neognathae of the orders formerly separated as
the Palaeognathae. For years I have felt that recognition of the
Palaeognathae, as a separate group apart from other birds, on the
basis of a supposed peculiarity in the palate, stood on flimsy ground.
The studies of McDowell (1948, pp. 520-549) demonstrate that the
structure of the palaeognathous palate, in which the palatine and
pterygoid bones are articulated by a squamous suture, is variable from
order to order and that in fact the details of this union differ con-
siderably in the several groups. For example, McDowell points out
that in Dromiceius the palatine and pterygoid are not in contact, while
in a number of families placed in the Neognathae, as in the Anatidae,
to name only one, the two bones are in articulation. As there is no
clear-cut separation, the former Palaeognathae must be combined with
the Neognathae.
The supposed bird Caenagnathus collinsi described by R. M. Stern-
berg (1940, p. 81) from the Belly River series of beds of Upper
Cretaceous age in Alberta has been carried tentatively in our avian
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 5
classification, though it has been my belief from the beginning that it
was reptilian. It is known from a lower jaw, beautifully preserved,
without appreciable deformation and practically complete except for
part of the lower section of one ramus. The resemblance to birds is
found in the lack of teeth, fused symphysis, and the considerable size
of the mandibular foramen. While these are characters found in
birds, there is nothing peculiar included since all are duplicated in some
of the groups of the Reptilia. The fossil resembles Reptilia in the
form of the articular surface, the forward position of the coronoid
area, the conformation at the symphysis, especially on the upper
surface, the upward curvature in that area, and in the general texture
of the bone. In none of these is there exact duplication in Aves,
except partially in the form of the symphyseal region. The whole
appearance of the bone strongly suggests a species related to the
Ornithomimidae among the therapod dinosaurs. In view of this the
“Order Caenagnathiformes” is now omitted from the avian classifica-
tion, since it is felt that its continued tentative inclusion may promote
misunderstanding as to its status.
The family Eleutherornithidae is introduced for the fossil Eleu-
therornis helveticus Schaub, from the Eocene of Switzerland, de-
scribed from a fairly well preserved pelvis. Apparently this is repre-
sentative of an ancestral group from which the living ostriches may
have come. Its greatest importance is found in its indication of re-
lationship with carinate groups though of unquestioned ratite stock. It
is thus important as definite indication that the struthious birds are
descended from flying ancestors, not from some distinct cursorial line
that always has been flightless, as some have contended.
The genus Podiceps.—The differences of opinion that prevailed for
years as to the application of the generic name Colymbus have been
adjusted currently by an arrangement under which Gavia has been
accepted for the loons and Podiceps for the grebes. There is, how-
ever, discussion still as to the proper spelling of the ordinal and
familial names for which Podiceps is the base. The uncertainty arises
from misunderstanding of the derivation of this generic term. The
colloquial name applied to these diving birds in the English of the
16th to the 18th centuries (and later) was “arse foot,” or “arsfoot,”
from the posterior position of the leg. The term is found in the early
dictionaries of Johnson, was carried in the later editions of Todd and
Walker, and is still found in a footnote in Webster’s 1953 volume,
with indication there that the word now is obsolete. Some early
authors who wrote in Latin rendered this term appropriately as
“‘Podicipes,” as for example Willughby (1676, p. 258), and Ray (1713,
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
pp. 125, 190), where the horned grebe is listed as “Colymbus sive
Podicipes minor.” Catesby (1731, p. 91) wrote of the pied-billed
grebe under the heading “Prodicipes Minor Rostro vario,” but he
corrected the spelling of the first word in the legend for the plate
that faces the text, which is labeled “Podicipes &c.” This account
by Catesby was the sole basis on which Linnaeus (1758, p. 136) estab-
lished his specific name for the pied-billed grebe. And it is here that
present-day confusion has its beginning, since Linnaeus called the
bird “Colymbus Podiceps,’ and in citing the reference to Catesby
wrote it ‘“Podiceps minor, rostro vario.” While he corrected Catesby’s
error in spelling he thus made another of his own, which remains
in our current name Podilymbus podiceps (Linnaeus) for the pied-
billed grebe. Following Linnaeus, John Latham (1787, p. 244) pro-
posed the genus Podiceps, in which he included several species of
grebes, with basis for the name on Linnaeus, as he makes reference
to “Colymbus Lin.” The error in spelling was recognized by several
early authors, as in a note attributed to Oken (1839, p. 674) and one
by Gloger (1854, p. 430). Correct usage for a family name based on
Podiceps (=Podicipes) was indicated by Newton (1896, p. 381).
That this history, well known up to 40 years or so ago, has been for-
gotten by many is shown by recent action of the International Com-
mission on Zoological Nomenclature (1957, pp. 300-304) which it
appears should have further review. The data supplied by the Com-
mittee to Dr. Grensted, as classical adviser, were misleading, as
there was no indication for his information that “Podiceps” had been
derived from “Podicipes.”
As the terminal root in Podiceps is a contraction of the Latin pes,
pedis, it would appear that the correct form for the family name is
Podicipedidae (not Podicipidae or Podicipitidae), and for the order
Podicipediformes (not Podicipitiformes or Podicipidiformes).
The Procellartiformes.—Family segregation in this order has been
oversimplified in some recent discussions, probably through misunder-
standing of the group characters, possibly also through somewhat con-
fusing names that have been applied to familial and generic categories.
Verheyen (1958, pp. 11-14) has placed the Pelecanoididae in an order
with the Alcidae, as indicative that the auk group is allied rather
closely to the Procellariiformes. The resemblances that he cites appear
due to convergence, as the basic form of the diving petrels is definitely
that of the shearwater-petrel group. Aside from this, the Diomedeidae
and the Pelecanoididae have been accepted without apparent question,
but the remaining species have been combined by some under a single
family name. Lowe (1925, pp. 1436-1443) has shown that the genera
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE Ti
included in the Hydrobatidae have a simplified condition in the
quadrato-tympanic region of the skull in which the opening of the
upper tympanic recess is small, and is so located that it separates the
squamosal and opisthotic facets. In addition, the posterior border of
the sternum is truncated and entire, and basipterygoids are absent or
are represented only by small spines. In the Procellariidae, on the
other hand, the foramen of the upper tympanic recess is greatly
enlarged and lies anterior to the two facets for the quadrate, which
are joined by a bridge of bone; the posterior border of the sternum
is notched ; and basipterygoid processes are present. These constitute
distinctive characters at the family level.
The Pelecaniformes.—lIn the arrangement of suborders in the order
Pelecaniformes we encounter in marked degree the standard difficulty
of logical placement in linear alignment of groups that really stand in
three-dimensional relationship. Lanham (1947, pp. 65-70) has made
a summary of the major anatomical characters of the group in which
he points out the differences that set off the Phaéthontes and the
Fregatae from the Pelecani. There is no question that the first two
carry primitive characters, which may be presumed to be similar to
those found in ancient ancestral stocks, since in these resemblances
they are more like other types of birds, notably the Procellariiformes.
From this style the families of the suborder Pelecani have become
widely divergent. Although the tropicbirds and the frigate-birds both
have retained a part of what may be regarded as a basic pattern, they
are so distinct in other respects that it appears to be more reasonable
to relate them individually as branches from a common stock rather
than to combine the two on one line, separate from the Pelecani. The
Phaéthontes possibly may have separated earlier than the Fregatae.
Among interesting differences other than those of internal anatomy,
it may be noted that the tropicbirds have the young covered with down
at birth and that the adults possess series of air cells under the skin on
the forepart of the body like those found in pelicans and boobies. The
frigate-birds have young almost naked at hatching, and the emphy-
sematous condition is mainly lacking. In view of this I prefer to
continue to align these groups on either side of the Pelecani.
Though there is no question that the cormorants and snake-birds
are closely allied, they differ in such degree that they should be retained
in separate family status. The snake-birds are marked by a peculiar
conformation of the cervical vertebrae through which the beak be-
comes a triggered spear in feeding. The bridge of Donitz on the ninth
vertebra is an important part of this arrangement. The stomach also is
unusual in possessing a curious pyloric lobe, lined with a mat of hair-
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
like processes. And there is only one carotid artery while in cor-
morants there are two.
The description of Osteodontormis orri by Hildegarde Howard
(1957a, pp. 1-23) from the Monterey formation in the Miocene of
California adds a third species to the strange Odontopteryges, whose
common character is found in the sharply pointed, dentate projections
developed on the margins of upper and lower mandibles as continuous
parts of the bony structure of the jaws. This suborder was placed
tentatively (Wetmore, 1930, p. 3), following Lydekker (1891, pp.
57-58), in the Pelecaniformes, but this was not definite, as the char-
acters of Odontopteryx have been interpreted by some as indicating
closer alliance to the petrel-albatross group. In July 1956, at the
British Museum (Natural History), through the kind attention of
Dr. W. E. Swinton, I had the privilege of studying the type skull of
Odontopteryx toliapica Owen, which came from the London clay of
the lower Eocene, on the Isle of Sheppey, Kent, England. It was
possible thus to ascertain certain details not clear from the published
accounts. Asa result of this study it is my opinion that the characters
clearly indicate relationship with the Pelecaniformes.
Without repeating unnecessary detail, available in Lambrecht’s
great volume (1933, pp. 304-307), it was interesting to note the
strongly marked craniofacial hinge at the base of the bill, like that of
gannets and cormorants, and also the impressed line along the side
of the premaxilla, and the definite closure of the external narial
opening, as in the Sulidae. The distal articular end of the quadrate
suggests that of Phaéthon, though somewhat more flattened, with the
whole articular surface narrower, and the separate segments more
nearly in line than in any living species of the various pelecaniform
families. The lachrymal appears to have been slender and is firmly
anchylosed on its upper margin to the frontal as in Phalacrocorax.
The rounded cranium suggests that of pelicans, rather than the more
flattened form of other families of the order. The sum of the char-
acters indicates a bird of gannetlike diving habit that, when slippery
aquatic prey was seized, could hold it firmly in the sharp dentations of
the mouth.
Dr. Howard in her interesting study of Osteodontornis has elevated
the group to the rank of an order, on the consideration that it “may
represent an early connection with procellariiform—pelecaniform
stock” (1957a, p. 22). It has seemed to me appropriate to emphasize
the evident pelecaniform character by retaining the two families recog-
nized in subordinal status in that group, since the resemblances that
point toward the Procellariiformes appear to be much less definite and
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 9
possibly may be subject to other interpretation. It is desirable now to
place the Odontopteryges at the beginning of the order because of their
antiquity. The known history of the group, which begins in the early
Eocene, indicates probable ancestry in Paleocene time. The pointed
projections on the jaws, assumed to have been sheathed in the in-
tegument of the bill, were without question used in seizing prey. The
disappearance during Miocene time of such a holding apparatus may
indicate that the bony projections were not completely successful for
their purpose, perhaps because of their hollow centers, as accidental
breakage in them would not be restored. The fine serrations restricted
entirely to the ramphotheca, found in the straight-billed species of the
pelecaniform order (tropicbirds, gannets, boobies, and anhingas),
may be regarded as a functional replacement.
The change in position made to the beginning of the order covers
only the Odontopterygidae and the Pseudodontornithidae and leaves
Cladornis and Cruschedula still unsettled as to relationship. As ex-
plained above (p. 4), Ameghino described both as forms of penguins,
but Simpson says that they have no connection with this group. As the
suborder Cladornithes, they are located in their former uncertain
position at the end of the Pelecaniformes.
Suborder Ardeae—The general resemblance of the boat-billed
heron (Cochlearius cochlearius) to the night herons has been the
occasion of differences in allocation of its rank in classification from
that of a subgenus of Nycticorax to full family status. In a recent
review of the Ardeidae, Bock (1956, pp. 31-35) has treated it as a
separate genus in a “Tribe Nycticoracini” allied to Nycticorax. Super-
ficially the boatbill is like a black-crowned night heron, but in detail
there are outstanding differences. The enlarged bill is obvious, and
there are four pairs of powder-down patches, instead of the three
found in the other herons. In the skull, the bill has been changed
from the spear point usual in herons to a broad scoop with the roof
of the mouth smoothly arched. The lower jaw is widely bowed to
fit this change, and the symphysis is greatly reduced in length. The
palatines are so greatly broadened, and so inflated on the outer
posterior margin, that they have little resemblance to the ordinary
heron form. The quadrate has the orbital process shorter and thicker
and the mandibular articulation narrowed ; the lachrymal is small ; the
eye opening considerably enlarged to house the exceptionally large
eye; and the external nasal opening considerably reduced. The palatal
musculature is decidedly stronger than in the true herons.
In life boatbills act like night herons, as they roost and nest in
groups and are mainly nocturnal. When hunting at night, I have
IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
found them feeding in shallow waters, often in riffles where they
scoop at their living prey, rather than spear at it as is the custom with
the typical herons. The eyes, wood brown by day, at night reflect the
jacklight with a faint orange sheen, which I have not observed in
other herons. The eggs are pale, nearly white, and often are lightly
speckled with brown, so that they resemble those of the tiger bittern,
Tigrisoma lineatum, rather than those of the night herons, which are
deep blue.
While there is no fossil record for the boatbill, I regard it as an
ancient sideline from the typical herons that, judged from its present
restricted range in the American Tropics, has not been too successful.
It may seem attractive to unite Cochlearius with the true herons,
but from long acquaintance I regard their characters, briefly outlined
above, sufficient to maintain a separate family status.
In view of the fact that the structural characters of the Balaenicipit-
idae have been summarized clearly by Stresemann (1934, p. 809), it
seems strange that the status of this family has been a matter of ques-
tion. The single species shows affinity both with storks and with
herons, in addition to outstanding peculiarities of its own. Miss
Cottam (1957, pp. 51-71) has made a careful summary of the osteol-
ogy from which she deduces a pelecaniform relationship, but this
appears to be due to convergence rather than to actual relationship.
The great enlargement of the skull has occasioned superficial re-
semblances to pelicans, but these, and others seen elsewhere in the
skeleton, are subordinate to the general sum of all characters, which is
ciconiiform.
Phoenicopteri—The position of the modern flamingos, which show
characters that point on one hand to the Ciconiiformes and on the
other to the Anseriformes, has been a matter of some variance in al-
location, Mayr and Amadon (1951, pp. 7, 33), with only brief
discussion, have set them up as a distinct order, but general opinion
has carried them as a suborder allied to the herons, storks, and their
relatives. The latter course remains justified when the fossil genera
Palaelodus and Elornis of the upper Eocene to Miocene of western
Europe are considered (Wetmore, 1956, p. 3). This group of flamingo
relatives was identified in North America when Alden Miller (1944,
p. 86) described Megapaloelodus connectens from the lower Miocene
of South Dakota, a species to which remains from the upper Miocene
of California also are referred (Loye Miller, 1950, pp. 69-73; 1952;
pp. 296-298). The group may be recognized as the family Palaelodidae,
on the generic name Palaelodus Milne-Edwards (1863, pp. 157, 158).
(There has been confusion relative to the proper spelling, since Milne-
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE Dt
Edwards in his important later work [1868, p. 58] used the form
Paloelodus.)
Howard (1955, pp. 3-23) has described a still different form of
the flamingo group as Telmabates antiquus from the lower Eocene
(Casamayor formation) of Chubut in Patagonia. While this species
resembles the Palaelodidae in shortness and other details of form in
the leg, it may prove to be representative of a separate family on
characters found in the vertebrae and wing, as suggested in the
original description. It is regarded for the present as of subfamily
status in the Palaelodidae.
Suborder Cathartae.—The superfamily Neocathartoidea, and family
Neocathartidae, for the curious vulture Neocathartes grallator (Wet-
more), discovered in the Upper Eocene fossil beds of Wyoming, in-
troduced a new element in our known avifauna in the form of a
small-winged, strong-legged vulture that evidently was terrestrial with
limited powers of flight. It had about the same relation to the other
American vultures that the secretarybird has to the hawks and falcons.
Its inclusion also requires a separate superfamily, the Cathartoidea,
for the previously known cathartine families.
Galliformes.—The Numididae, which have been placed by some as
a subfamily of the Phasianidae, differ in completely lacking the
tuberosity or plate on the inner side of the second metacarpal that is
sO prominent in pheasants and grouse. It should be recorded, however,
that Hudson, Lanzilloti, and Edwards (1959, p. 64) note that Numida
shows no peculiarities in the leg musculature when compared with the
Phasianidae. The Tetraonidae, in contrast with the Phasianidae, have
the pelvis relatively much broader and different in proportion, and
the tarsus relatively shorter in relation to the length of the tibiotarsus.
With these differences in mind it seems reasonable to retain the three
groups in family status, at least until more detailed knowledge of their
anatomy as a whole warrants change.
Gruiformes.—In the Turnices the two genera of bustardquails,
Turnix and Ortyxelus, have no hind toe, the wing is eutaxic, only the
left carotid is present, and the eggs are rounded oval. The plain-
wanderer of Australia, Pedionomus, has a small hind toe, the wing is
diastataxic, right and left carotids are found, and the large eggs are
pyriform. It seems desirable to continue these as separate families,
rather than as subfamilies of one group, an arrangement that Strese-
mann (1933, p. 760) has accepted.
It has long been known that Mesites Geoffroy for the curious
roatelos of Madagascar is antedated by the same name used by Schon-
herr for a group of beetles. It has been in error, however, to replace
I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
this with M/esoenas Reichenbach 1862, since the conflict had been noted
seven years earlier by Prince Bonaparte who gave the group the name
Mesitornis (Bonaparte, 1855, p. 484). The suborder becomes Mesi-
tornithides and the family Mesitornithidae.
In the course of study of the fossil Andrewsornis abbotti from the
Oligocene of Patagonia, Bryan Patterson (1941, pp. 50-53) has re-
viewed related groups to the end that he has added the family
Psilopteridae for the South American fossil genera Psilopterus and
Smuiliorms. Further, he has placed Phororhacos and its allies as a
superfamily Phororhacoidea under the suborder Cariamae. His
further observations on these matters are to appear later in a more
comprehensive paper.
The family Cunampaiidae, for the fossil Cunampaia simplex, named
by Rusconi (1946, p. 1) from the Oligocene of western Argentina,
while placed in the Cariamae, still remains of uncertain status.
The allocation of the phororhacid group to its new position and
its demotion from subordinal status requires recognition of a super-
family Cariamoidea for the living Cariamidae and the fossil group
Hermosiornithidae. The common name for the Cariamidae in most
English writings has been “Cariama,” being the form instituted by
Marcgrave in 1648 in his Historiae rerum naturalium Brasiliae, when
he rendered the Tupi name “cariama” as cariama. This was copied by
subsequent authors, including Linnaeus in his twelfth edition, and so
came finally into English usage, beginning with Ray’s translation of
Willughby’s Ornithologiae in 1678. Seriema, a modification of the
Indian word gariama, is used in Brazil, and with that spelling has
come into the Engish language, where it should replace the other form.
Charadruformes.—Differences of treatment at present are found
mainly in the superfamily Charadrioidea and the suborder Lari, in
which the groups have been regarded by some as of family value
and by others have been allocated to the rank of subfamilies. The var-
ious studies that have been made have not been complete from a
taxonomic point of view except for part of the species, and the con-
clusions derived from the data available appear in the main more
philosophical than concrete. The picture therefore still remains
confused.
In view of the diverse specializations that are apparent, and the
obvious long evolutionary history, it appears better to me to continue
to acknowledge the main segregations as families, at least until the
subjects involved have been more thoroughly investigated. A family,
Rhegminornithidae, covers the fossil Rhegminornis calobates Wet-
more, described from the lower Miocene of Florida. This was as
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 13
large as a medium-sized curlew, of peculiar form as regards the
foot, the only part of the skeleton known, which shows certain char-
acters that seem to point toward the jacanas, though the bird is to
be placed in the Charadrioidea.
It should be noted that the family affinity of the turnstones and the
surfbird, long considered members of the plover family, is not certain
as some studies (Lowe, 1931, pp. 747-750) place them in the Scolo-
pacidae. (See also Bock, 1958, pp. 85-86.)
In the Lari the terns and the gulls are regarded as one family,
though there are some reasons that make further examination of this
treatment desirable. The Stercorariidae possess a 2-notched sternum,
large caeca, a cere, and a complex rhamphotheca. In the Laridae
ambiens and biceps slip are present, the sternum is 4-notched, there
is no cere, and the rhamphotheca is simple in form.
In further discussion of proposals relative to this group it is
pertinent to observe that a logical scheme of classification should
attempt to outline relationships in living and fossil species through
examination of all available data, considerations in which modern
studies of behavior find increasingly useful part. There are pitfalls
and hidden traps, however, when attempt is made to establish affiliation
through any single method of approach, as inevitably inconsistencies
appear. I fully agree with Martin Moynihan (1959, pp. 22-23, 35-38)
that the skimmers (Rynchops) represent an early separation in the
ancestry of the gull-like birds and find it pertinent that this is shown
in their behavior pattern. At the same time these birds present out-
standing peculiarities that should be considered in assigning them ap-
propriate status in relation to their relatives. The bill, compressed
to knifelike form, with great elongation of the ramphotheca of the
lower jaw, is unique, and the method of feeding, where the lower
mandible cuts the water surface with the bird in flight, is equally
strange. The structural modifications in the form of the skull from
that found in skuas, gulls, and terns also are too extensive to be
ignored. The elongated blade of the lower mandible anterior to the
symphysis of the rami is intriguing but less important than the pro-
found changes elsewhere. The palatine bones are greatly expanded,
the orbital process of the quadrate is reduced to a short, pointed
spine, the impression for the nasal gland is much reduced, the frontal
area is inflated and produced posteriorly, with compression of the
lachrymal, and consequent reduction in size of the cavity for the eye,
to enumerate the most outstanding differences in the osteology. Ex-
ternally, the pupil of the eye is a vertical slit similar to that of a cat,
and thus unlike that of any other group of birds (Wetmore, 1919,
I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
p. 195). Other peculiarities have been described in the musculature.
The sum of these characters justifies treatment of the Rynchopidae
as a distinct family in their suborder.
The fossil humerus, type of Mancalla californiensis Lucas, that
was the first intimation of a flightless auk on the west coast, while
unique for many years, now has been supplemented by abundant ma-
terial from which an additional, smaller species, Mancalla diegense
(L. H. Miller), is recognized. It has been possible also to construct
a composite skeleton of the larger one that is sufficiently complete
to give a clear picture of its form and characters. The evident peculiari-
ties of the genus Mancalla are found in the wing, as elsewhere the
skeleton resembles that of other alcids, except for differences of a
generic and specific nature. In comparison of the wing with that of the
great auk, now extinct, that formerly ranged the coasts of the North
Atlantic, the humerus of Mancalla is generally similar, the forearm
appears proportionately shorter, and the hand more elongated. Ulna,
radius, metacarpal, and phalanges so far as present are more slender.
The head of the humerus in Mancalla differs decidedly in the relative
angles of different elements, and also in the conformation of the
distal articular surface. The general indication in the west-coast bird
is of a proportionately longer wing, with the slighter bones to be ex-
pected in a form of lesser bulk. Loye Miller (1946, pp. 34-36) and
Loye Miller and Howard (1949, pp. 222, 225) have likened the
specialization seen in the wing to that found in penguins and explain
any similarity to the great auk, Pinguinis impennis, as due to con-
vergence. On this basis they have separated Mancalla from the other
auks in the family Mancallidae. While I followed this, with some
reservation, in the last revision of the fossil list (Wetmore, 1956, pp.
3, 80-81), a further review of the subject raises definite doubt, since,
except for some specialization in the wing, Mancalla, as said above,
is like other alcids. The change in the wing is no greater than that of
Pinguinis, though the divergence is in a different direction. It would
seem sufficient to place Mancalla in a well-marked subfamily, rather
than in a separate family.
Finally, the proposals of several authors to separate the auks in a
distinct order appear to require further study.
Strigiformes.—Old World ornithologists in the main regard the
owls as belonging to a single family, but while all are deceivingly
similar in general aspect, Ridgway (1914, p. 598) years ago sum-
marized the considerable structural characters that separate the
Tytonidae and the Strigidae. It is necessary here only to point out the
more outstanding differences of the barn owls in lack of the manu-
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 15
brium, the different form of the posterior margin of the sternum,
which is entire or 2-notched, the straight outline of the palatines, and
in the ventral pteryla where the outer branch joins posteriorly to the
main tract. The Strigidae possess a manubrium, the sternum is
4-notched, the palatines are greatly expanded posteriorly, and the
posterior end of the ventral pteryla does not join the main tract at
the posterior end.
Apodiformes.—Lucas (1889, pp. 8-13; 1895, pp. 155-157) long ago
demonstrated the differences between the true swifts and the crested
swifts, though his work seems latterly to have been overlooked, in view
of the recent inclusion of the two in one group, as by Stresemann and
by Mayr and Amadon. The skull in the Hemiprocnidae is quite dis-
tinct in the general form of the cranium and in the development of
the nasals, vomer, and palatines. The hypotarsus has a tendinal
foramen (like that found in hummingbirds), and the plantar tendons
have the flexor longus hallucis connected with the branch of the flexor
perforans digitorum, which extends to the fourth digit. Coupled with
this there may be noted the curious nest, which, fastened to the side
of a branch, is barely large enough to contain one egg, and the further
fact that these birds perch regularly on branches and twigs in trees.
As Apus Scopoli, published in 1777, is recognized now in place of
Micropus Meyer and Wolf, 1810, for the type genus of the swifts,
the terms in the classification change to order Apodiformes, suborder
Apodi, and family Apodidae, which replace the former terms Mi-
cropodiformes, Micropodi, and Micropodidae, respectively.
Coraciiformes.—The proposal of Mayr and Amadon (1951, p. 35)
to include the rollers in one family, the Coraciidae, with three sub-
families, goes back to the arrangement of Dresser in his monograph
of the group (1893, pp. xviii, 85, 101). Sclater (1865, pp. 682-688),
however, many years ago, pointed out the pelvic powder-down tracts,
the small manubrium, and other peculiarities of Leptosoma, and set
it apart in a distinct family. The anatomy of the syrinx and feet was
further elaborated by Forbes (1880, pp. 464-475). The family
Leptosomatidae therefore should be recognized.
The groundrollers, Brachypteracias, Atelornis, and Uratelornis,
usually have been included as a subfamily of the Leptosomatidae, but
Stresemann (1934, p. 829) places them in a separate family, the
Brachypteraciidae. There seems to be reason for this in their general
appearance, though their anatomy is not well known. Brachypteracias,
in its skeleton, differs from Coracias and Eurystomus in the much
greater depth of the outer notch on the posterior border of the
sternum, in the much broader and stronger pelvis, the heavier femur,
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
and the greater curvature of the shaft and reduction of the crista
superior of the humerus. I have not seen the skull. The habit of life
is markedly different. Although anatomical material of the other
genera is not presently available, it seems reasonable to accept Strese-
mann’s proposal. These peculiar birds certainly are not closely allied
to Leptosoma.
Lack of information on the anatomy of the woodhoopoes must be
the reason for recent nonrecognition of the Phoeniculidae as a family
separate from the Upupidae, since the two are quite distinct and
have been so recognized for many years. The external differences
are readily apparent. In the skeleton in Phoeniculus (of which I have
seen several examples) the posterior part of the nasal area is ossified,
there being only a small, narrow, elongated nasal opening ; the ecteth-
moid is much reduced; the anterior end of the pterygoid is broadly
expanded ; the sphenoidal rostrum is swollen at the anterior end, where
the expanded ends of the pterygoids join it; the quadrates are de-
cidedly larger; the keel of the sternum is greatly reduced, being only
half as high as in Upupa; the furculum is broader ; the pelvis is nar-
rowed, and considerably enlongated posterior to the acetabulum, with
the ischio-pubic fenestra greatly enlarged; and the tarsus is heavier
and broader, with two definite fenestra below the head. There are
other minor details. In all of the above the characters of Upupa are
directly opposite. The two groups appear to me to be sharply set off
as distinct families.
Passeriformes.—This order, with more living species than all the
others combined, and far fewer fossil forms known, presents many
difficult problems in logical arrangement. The major groups are clear,
whether we rank them as suborders or superfamilies being a matter
of opinion. But the limits and status of numerous families contained
in these larger categories are uncertain since the internal anatomy is
known for so few kinds that details of difference are poorly under-
stood. Superficial resemblances, on the other hand, are so obvious in
many cases that they cause confusion. Under the circumstances it
continues to seem appropriate to me to accept the family grouping
that has been current for many years, except in those cases where
acceptable studies clearly indicate change. Supposition in these matters
has led to various proposals for changes, some part of which un-
doubtedly will prove correct. It is equally probable that a part,
possibly the considerably larger part, may prove to be unfounded when
details are more clearly known. If change is accepted under these cir-
cumstances it may prove unwarranted, necessitating further shift,
perhaps a return to the original status. Since this can only prove
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 17
confusing I prefer the conservative course. In the remarks that
follow I shall discuss only a few matters on which I have more or less
concrete ideas.
In the superfamily Furnarioidea, von Ihering (1915, pp. 145-153)
united the Furnariidae and the Dendrocolaptidae, since he was unable
to separate two groups on the basis of the form of the posterior border
of the nasal opening. The variation that he showed seems valid, but
there are numbers of other points of supposed difference in the osteol-
ogy and other structural details, so that his suggestion is far from
established. Pycraft (1906, pp. 133-159), though seemingly uncer-
tain in the beginning, finally retained the two families. It may prove
that some genera are wrongfully allocated at present between the two
groups, so that their shift, when we have sufficient information, will
clear our understanding.
In the Tyrannoidea, the family Oxyruncidae is known through ex-
ternal characters that seem to warrant separation. If the sharpbills
have other affinities it is doubtful that these are within the family
Tyrannidae, where some have placed them.
In the family Cracticidae, recognized by Australian ornithologists,
the skull, according to Pycraft (1907, pp. 355-365), mainly from
examination of Gymnorhina, has the zygomatic process of the
squamosal bifurcate, the postorbital process large, the orbitosphenoid
ossified, the interorbital septum with a single opening, the prefrontals
unusually large, and the form of the palate peculiar. In his phylo-
genetic tree Pycraft places the group on a common stem with the
Artamidae, and not far from the Paradisaeidae. His account is diffi-
cult to summarize in concrete form.
The family Grallinidae is likewise recognized officially by Austra-
lian ornithologists for Grallina cyanoleuca, the magpie-lark. The
principal study of the osteology is that of Shufeldt (1923, pp. 16-19,
pl. 6) but his account is mainly descriptive and without definite con-
clusion. Amadon (1950, pp. 123-127) has placed Corcorax and
Struthidea here tentatively, though this seems subject to further proof.
Stonor (1937, pp. 475-490) has outlined excellent reasons for recog-
nition of the Ptilonorhynchidae, finding that they differ from Para-
disaeidae, with which they have been united, in having an apterium
in the center of the dorsal feather tract, the tip of the vomer convex,
larger, more developed maxillo-palatines, the margin of the palatines
angular, smaller ectethmoid, much larger lachrymal, and _ slender,
greatly elongated orbital ramus of the quadrate. The genera Loria
and Loboparadisea, usually included here, he transfers to the Para-
disaeidae. His conclusion is that “the Ptilonorhynchidae constitute
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
a singularly complete and isolated family of the acromyodian passerine
birds and show no special relationship to any other, being sharply
marked off by the structure of the skull, the colour-pattern, and the
bower-building habit.” (It should be noted that the names on Stonor’s
figs. 6 and 8 have been transposed, fig. 6 being Semioptera wallacei,
and fig. 8 Amblyornis subalaris, not the reverse as printed on pp. 481
and 483.)
Oberholser (1917, pp. 537-539) has set up a distinct family Irenidae
for the fairy bluebirds (Jrena), and Delacour (1946b, p. 3) a family
Aegithinidae for the leafbirds, which would cover Irena, Aegithina,
and Chloropsis.
The proper allocation of the genus Chamaea for the wrentits, at
present accepted by the A. O. U. Committee on Classification and No-
menclature as a separate family, the Chamaeidae, is one of consider-
able uncertainty. Delacour (1946a, pp. 18, 25, 35) has suggested
that the group be located in the family Timaliidae in a special sub-
family in which he includes also such diverse genera as Chrysomma
(Moupinia), Panurus, Conostoma, and Paradoxornis (combining un-
der this name Suthora, Psittiparus, Neosuthora, and Cholornis). This
is an obviously heterogeneous assemblance, in which Chamaea has
slight resemblances to the first only. From Moupinia poecilotis
(placed in Chrysomma by Delacour) the wrentit differs definitely in
weaker, less arched bill and in differently proportioned feet. It has
no close similarity to any of the others that are mentioned. Although
the relationships of Chamaea are obviously uncertain, it is retained
as a family pending other information.
In consultation with Herbert Deignan, expert in matters that relate
to the birds of eastern Asia, the Campephagidae have been placed
near the Pycnonotidae, an arrangement that agrees with that adopted
by Charles Vaurie in his recent volume on the palearctic region (1959,
p. 181), and the Paradoxornithidae are brought nearer the Timeliidae.
The fossil family Palaeoscinidae, proposed by Hildegarde Howard
(1957b, p. 15) for the species Palaeoscinis turdirostris, has been in-
serted provisionally near the Pycnonotidae. The specimen on which
this name is based is a skeleton found in Santa Barbara County, Calif.,
compressed in a slab of Miocene limestone of the Monterey forma-
tion. The type, in which most of the bones are outlined, is one of those
attractive silhouette impressions that delight the eye but that often
pose difficulties in classification through lack of clear-cut characters
on which to judge relationship. In the present instance Dr. Howard
has concluded that “affinities of the Palaeoscinidae lie with the
Pycnonotidae, Bombycillidae, Corvidae and Cinclidae” of the suborder
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 19
Passeres. Affinity with the Bombycillidae may be queried, as the fossil
differs from Bombycilla in the proportions found in the hind limb,
where both metatarsus and femur are longer in comparison with the
tibiotarsus, and the toes appear longer, as well as of different propor-
tion. The corvid affiliation also seems uncertain because of the slender
form of Palaeoscints, since the skeleton of the crows and their relatives
is strong and robust.
Separation of the two genera of leafbirds, Aegithina and Chloropsis,
in a family distinct from the Pycnonotidae is justified on the basis of
characters found in the skull. The entire palatal structure is slighter
than in Pycnonotus and allied genera, with the central plate of the
palatine reduced in area, and the transpalatine produced posteriorly.
The sphenoidal rostrum is slender, as is the orbital process of the
quadrate. In Pycnonotus the palatine is broad, the transpalatine proc-
ess distally is only slightly angular without posterior projection, and
both the rostrum and the orbital process of the quadrate are strong
and heavy. Herbert Deignan informs me that the group, recognized
by several authors, seems to have been first separated by Cabanis
(1847, p. 326), who designated it as the subfamily “Phyllornithinae”
based on Phyllornis Temminck, 1829. This generic term is antedated
by Chloropsis Jardine and Selby, 1826, so the family name based on
this genus will be Chloropseidae, rather than Aegithinidae which dates
from G. R. Gray in 1869 (p. 312).
The fairy bluebirds, genus Jrena, often have been placed with the
leafbirds but have no close connection with that group. The main
external peculiarity of Jrena is found in the smooth, enamel-like tip-
ping found in adult males on the feathers of the central dorsal area
from the center of the crown back over hindneck, back, rump, and
upper tail coverts, and on the elongated under tail coverts. As this
is a secondary sexual character, not present in females, it has no value
at the family level. In the osteology, the skull differs from Chloropsis
and Aegithina in the completely open external narial opening, the
ossification of the vertical plate between the nares, the more inflated
lachrymal, and the more elongate maxillo-palatines. In the sternum
the depth of the notch on either side of the posterior margin relatively
is decidedly less, and in the pelvis the antitrochanter has the dorsal
margin much produced laterally. The general resemblance in these
matters is to species of the genus Oriolus. It may be observed further
that the feathers of breast and back in the aberrant species Oriolus
traillii and O. mellianus have smooth exposed ends that suggest the
condition found in male Jrena. In view of these resemblances, and in
lack of important differences, it seems sufficient to include the fairy
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
bluebirds in the family Oriolidae, as the subfamily Ireninae, which
incidentally dates from G. R. Gray (1869, p. 288) and not from the
name Irenidae set up later by Oberholser (1917, pp. 537-539).
Suggestions for the union of the Bombycillidae, Ptilogonatidae,
and the Dulidae in one family are not substantiated by examination of
the skeleton. Dulus, the palmchat, is widely different from the other
two, a structural distinction that is further emphasized by its curious
communal nesting habits. The first two seem more closely related but
are separated clearly by characters found in the ectethmoid region
of the skull, and in the manubrium, to mention only two points that
are easily apparent. Delacour and Amadon (1949, pp. 427-429) con-
sider Hypocolius closely allied to Ptilogonys.
While Zimmer (1942, p. 10) believed that the family Vireolaniidae
should be included in the Vireonidae, separate family rank in my
opinion is definitely justified. In addition to characters assigned by
Pycraft (1907, pp. 378-379) for the shrike-vireos I have found that in
the pterylosis the dorsal tract on the lower back is divided, the arms
being broad at the ends, and separated from the narrowed line that
continues onto the caudal area. This is completely different from the
usual rhomboid in the vireos, and may indicate that the family eventu-
ally should be removed from the vicinity of the Vireonidae.
The family characters of the peppershrikes, likewise outlined by
Pycraft in the reference given above, are easily apparent on examina-
tion of the skeleton.
The family Callaeidae has been separated by Stonor (1942, pp.
1-18) on the weakened keel of the sternum, the great development of
the lower limb coupled with reduced powers of flight, and the presence
of a mouth wattle, for three peculiar genera, Callaeus, Heterolocha,
and Philesturnus of New Zealand.
Continuing discussion relative to the group of families to be placed
in elevated position at the end of the list has led to publication of
several useful studies and interesting statements. Beecher (1953, pp.
270-333) from examination of the musculature of the jaw, aided
by other anatomical features, has proposed two major divisions of the
suborder of the song birds, within which he has diagramed radiating
lines of family and subfamily relationship. While he shows a variety
of connections that in many cases vary widely from ideas current at
present, he places the crow group in the assemblage with simpler
muscle development in the area of the jaw, in contrast to those of
higher status with a more complicated arrangement.
Tordoff (1954a, 1954b) in a study of the skull, particularly the
palatal structure, of species allied to the Fringillidae, has proposed the
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 2I
union of part of the honeycreepers and the wood warblers in one
family, the tanagers, with part of the coerebine assemblage with some
of the fringillids in the Fringillidae, and removal of the cardueline
finches to the Ploceidae, placing that family at the end of his list. His
detailed studies afford much valuable information. I agree with him
that shifting of certain genera to families in which they are not classi-
fied at present will lead to better alignment, but I am not prepared
from present information to completely dismember the Coerebidae
without further study. Coereba, for example, has a stomach peculiar
in its small size; Diglossa differs in the form of the bill, in which
the gonys is extended posteriorly behind the level of the nostril, so
that it differs from all other oscinine species, to cite only two easily
seen characters.
Mayr and Greenway (1956, pp. 2-5, 8-9) discuss problems of
sequence in some detail and cite the approval of a committee appointed
at the International Ornithological Congress held in Basel in 1954 to
allocation of the Corvidae at the higher end of the list, as has been
long customary among most ornithologists of Europe. In further
consideration of these matters, I published a note on the humerus of
the Corvidae (Wetmore, 1957, pp. 207-209), which called attention
particularly to the proximal end of the bone, where the pneumatic
fossa in Corvus, for example, has a form not only generally similar to
that of the New World flycatchers and their allies, which are recog-
nized as low down in the linear classification, but also to the wood-
peckers, the Coraciiformes, and the trogons. There is transition
from this simpler form to the style found in such groups as the
Icteridae, Thraupidae, and Fringillidae, where the fossa is enlarged,
and is more complex, as it is partly divided by a bladelike process pro-
jecting from the internal tuberosity. (In the paper cited I neglected
to refer to an earlier study by James T. Ashley [1941] on the humerus
of the Corvidae, which outlined the same differences, and on which
Ashley considered the crow group to have more primitive status. )
Amadon (1957) recently has outlined the three major groups of
oscinine families, with the conclusion that the one most highly ad-
vanced includes the 9-primaried New World groups, while the section
containing the crows is placed low at the beginning. There is general
agreement with this in the classification outlined by Delacour and
Vaurie (1957).
Storer (1959) in a clearly stated summary of these recent contri-
butions, in which he includes a more recent statement by Mayr (1958),
writes that in a classification for a text on the biology of birds now in
preparation he has placed the 9-primaried groups in the highest place,
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
and indicates that this is the procedure that is gaining in acceptance
in parts of the world other than America.
The former family Melithreptidae becomes the family Melipha-
gidae, since the name of the type genus is now accepted as Meliphaga
Lewin, 1808.
In a similar way the family Compsothlypidae for the wood warblers
becomes the family Parulidae, since the former Compsothlypis
Cabanis, 1851, is replaced by the older Parula Bonaparte, described
in 1838.
The order of arrangement in the Passeriformes as said above is in
part necessarily arbitrary, through the easily perceptible and often-
remarked fact that we are required to list the groups in linear order in
a two-dimensional alignment when actually they stand in three-dimen-
sional relationship to one another. A further element that may be re-
garded almost as a fourth dimension is found in some of the extinct
groups known only as fossils that have no close relatives alive today.
The sequence in the following pages is the one that best represents my
present understanding, based on personal studies over a period of
more than 50 years. I continue to place the Fringillidae at the end of
the list, because of my feeling that this group is the modern expres-
sion of a main core or stem that through the earlier Tertiary periods
has given rise to more specialized assemblages that we now recognize
as distinct families. Further specialization is apparent in some parts
of the existing fringilline assemblage that, if undisturbed, may lead
to further differentiation, should these variants be able to persist for
the necessary millenniums in our rapidly changing world. Adjacent
to the Fringillidae I place the other groups that obviously are closely
allied to them. Attempts to arrange the avian families with the Cor-
vidae and their allies in the terminal position, because of supposed
more advanced development of the brain, appear to me quite uncertain,
particularly in view of our decidedly limited information in this field.
Should this idea be coupled with belief in superior mental reactions
in the corvine assemblage, I would consider this more an anthropo-
morphic interpretation than one supported by scientific fact.
In the formation of group names the suffixes -idae and -inae for
families and subfamilies are accepted rather universally so that they
do not require examination. In view of the limited number of species
covered in ornithology I see no point in the introduction of tribes as
another category between the subfamily and the genus. This may be
useful to entomologists with their tens of thousands of species but
seems unnecessary and cumbersome with birds. In some of the more
comprehensive avian genera there are groups of species more closely
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 23
allied to one another than to their fellows, but the taxonomist may
discuss these at need as groups without imposing another burden on
a classification that now is highly divided. For the group names above
the family level, I believe it preferable to use suffixes that allow im-
mediate identification of the rank, coupled with a stem that, like the
family name, is based on a current generic term. Where ordinal and
subordinal names are both formed as Latin plurals there is possibility
of confusion.
SYSTEMATIC LIST
Fossil groups in brackets
Class Aves, Birds.
[Subclass Archaeornithes, Ancestral Birds (fossil). ]
[Order Archaeopterygiformes, Archaeopteryx, Archaeornis
(fossil) .]
[Family Archaeopterygidae, Archaeopteryx, Archaeor-
mis (fossil).]
Subclass Neornithes, True Birds.
[Superorder Odontognathae, New World Toothed Birds (fos-
sil).]
[Order Hesperornithiformes, Hesperornithes (fossil).]
[Family Hesperornithidae, Hesperornis (fossil) .]
[Enaliornithidae,! Enaliornis (fossil) .]
[Baptornithidae, Baptornis (fossil).]
[Superorder Ichthyornithes, Ichthyornis and Allies (fossil).]
[Order Ichthyornithiformes, Ichthyornithes (fossil). ]
[Family Ichthyornithidae, Ichthyornis (fossil).]
[Apatornithidae, Apatornis (fossil).]
Superorder Impennes, Penguins.
Order Sphenisciformes, Penguins.
Family Spheniscidae, Penguins.
Superorder Neognathae, Typical Birds.
Order Struthioniformes, Ostriches.
[Family Eleutherornithidae, Eleutherornis (fossil).]
Struthionidae, Ostriches.
Order Rheiformes, Rheas.
Family Rheidae, Rheas.
Order Casuariiformes, Cassowaries, Emus.
Family Casuariidae, Cassowaries.
Dromiceidae, Emus.
[ Dromornithidae, Dromornis (fossil).]
1 Position provisional.
24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
[Order Aepyornithiformes, Elephantbirds (fossil and sub-
fossil) .]
[Family Aepyornithidae, Aepyornis (fossil and sub-
fossil).]
[Order Dinornithiformes, Moas (fossil and subfossil).]
[Family Dinornithidae, Dinormis (fossil and subfos-
sil).]
[ Anomalopterygidae, Anomalopteryx, Emeus,
and Allies (fossil and subfossil).]
Order Apterygiformes, Kiwis.
Family Apterygidae, Kiwis.
Order Tinamiformes, Tinamous.
Family Tinamidae, Tinamous.
Order Gaviiformes, Loons.
Family Gaviidae, Loons.
Order Podicipediformes, Grebes.
Family Podicipedidae, Grebes.
Order Procellariiformes, Albatrosses, Shearwaters, Petrels,
and Allies.
Family Diomedeidae, Albatrosses.
Procellariidae, Shearwaters, Fulmars.
Hydrobatidae, Storm Petrels.
Pelecanoididae, Diving Petrels.
Order Pelecaniformes, Tropicbirds, Pelicans, Frigate-birds,
and Allies.
[Suborder Odontopteryges, Odontopteryx, and Allies (fos-
sil).]
[Family Odontopterygidae, Odontopteryx (fossil).]
| Pseudodontornithidae, Pseudodontornis, Os-
teodontornis (fossil) .]
Suborder Phaethontes, Tropicbirds.
Family Phaéthontidae, Tropicbirds.
Suborder Pelecani, Pelicans, Boobies, Cormorants, Snake-
birds.
Superfamily Pelecanoidea, Pelicans and Allies.
Family Pelecanidae, Pelicans.
[Cyphornithidae, Cyphornis, Palaeochendides
(fossil) .]
Superfamily Suloidea, Boobies, Cormorants, and Allies.
Family [Pelagornithidae, Pelagornis (fossil).]
Sulidae, Boobies, Gannets.
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 25
[Elopterygidae, Elopteryx, Eostega, Actiornis
(fossil) .]
Phalacrocoracidae, Cormorants.
Anhingidae, Snake-birds.
Suborder Fregatae, Frigate-birds.
Family Fregatidae, Frigate-birds.
[Suborder Cladornithes, Cladornis and Cruschedula (fos-
sil) .]
[Family Cladornithidae, Cladornis, Cruschedula (fos-
sil).]
Order Ciconiiformes, Herons, Storks, and Allies.
Suborder Ardeae, Herons, Bitterns.
Family Ardeidae, Herons, Bitterns.
Cochleariidae, Boatbilled Herons.
Suborder Balaenicipites, Whale-headed Storks.
Family Balaenicipitidae, Whale-headed Storks.
Suborder Ciconiae, Storks, Ibises, Spoonbills.
Superfamily Scopoidea, Hammerheads.
Family Scopidae, Hammerheads.
Superfamily Ciconioidea, Storks.
Family Ciconiidae, Storks, Jabirus.
Superfamily Threskiornithoidea, Ibises.
Family Threskiornithidae, Ibises, Spoonbills.
Suborder Phoenicopteri, Flamingos.
[Family Agnopteridae, Agnopterus (fossil).]
[Scaniornithidae, Scaniornis, Parascaniornis
(fossil).]
Phoenicopteridae, Flamingos.
[Palaelodidae, Palaelodus, Megapaloelodus,
Telmabates (fossil).]
Order Anseriformes, Screamers, Ducks, Geese, Swans.
Suborder Anhimae, Screamers.
Family Anhimidae, Screamers.
Suborder Anseres, Ducks, Geese, Swans.
[Family Paranyrocidae, Paranyroca (fossil).]
Anatidae, Ducks, Geese, Swans.
Order Falconiformes, Vultures, Hawks, Falcons.
Suborder Cathartae, New World Vultures.
[Superfamily Neocathartoidea, Neocathartes (fossil).]
[Family Neocathartidae, Neocathartes (fossil). ]
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Superfamily Cathartoidea, New World Vultures.
Family Cathartidae, New World Vultures.
[Teratornithidae, Teratornis, Cathartornis (fos-
sil).]
Suborder Falcones, Secretarybirds, Hawks, Falcons.
Superfamily Sagittarioidea, Secretarybirds.
Family Sagittariidae, Secretarybirds.
Superfamily Falconoidea, Hawks, Falcons, and Allies.
Family Accipitridae, Hawks, Old World Vultures,
Harriers.
Pandionidae, Ospreys.
Falconidae, Falcons, Caracaras.
Order Galliformes, Megapodes, Curassows, Pheasants, Hoat-
zins.
Suborder Galli, Megapodes, Curassows, Grouse, Pheasants.
Superfamily Cracoidea, Megapodes, Curassows.
Family Megapodiidae, Megapodes.
[Gallinuloididae, Gallinuloides (fossil).]
Cracidae, Curassows, Guans, Chachalacas.
Superfamily Phasianoidea, Grouse, Pheasants, Turkeys.
Family Tetraonidae, Grouse.
Phasianidae, Quails, Pheasants, Peacocks.
Numididae, Guineafowl.
Meleagrididae, Turkeys.
Suborder Opisthocomi, Hoatzins.
Family Opisthocomidae, Hoatzins.
Order Gruiformes, Cranes, Rails, and Allies.
Suborder Mesitornithides, Roatelos, Monias.
Family Mesitornithidae, Roatelos, Monias.
Suborder Turnices, Bustardquails, Hemipodes.
Family Turnicidae, Bustardquails.
Pedionomidae, Plainwanderers.
Suborder Grues, Cranes, Limpkins, Trumpeters, Rails.
Superfamily Gruoidea, Cranes, Limpkins, Trumpeters.
[Family Geranoididae, Geranoides (fossil).]
[Eogruidae, Eogrus (fossil).]
Gruidae, Cranes.
Aramidae, Limpkins.
Psophiidae, Trumpeters.
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 27
Superfamily Ralloidea, Rails.
[Family Orthocnemidae,? Orthocnemus, Elaphrocne-
mus (fossil). ]
Rallidae, Rails, Coots, Gallinules.
Suborder Heliornithes, Sungrebes.
Family Heliornithidae, Sungrebes.
Suborder Rhynocheti, Kagus.
Family Rhynochetidae, Kagus.
Suborder Eurypygae, Sunbitterns.
Family Eurypygidae, Sunbitterns.
Suborder Cariamae, Seriemas and Allies.
[Superfamily Phororhacoidea, Phororhacos and Allies
(fossil).]
[Family Phororhacidae, Phororhacos and Allies (fos-
sil).]
[Psilopteridae, Psilopterus and Allies (fos-
sil).]
[Brontornithidae, Brontornis, Liornis, and
Allies (fossil).]
[ Opisthodactylidae, Opisthodactylus (fossil) .]
[Cunampaiidae, Cunampaia (fossil).]
Superfamily Cariamoidea, Seriemas and Allies.
[Family Bathornithidae, Bathornis (fossil).]
[Hermosiornithidae, Hermosiornis, Procari-
ama (fossil).]
Cariamidae, Seriemas.
Suborder Otides, Bustards.
Family Otididae, Bustards.
[Order Diatrymiformes, Diatryma, Omorhamphus, and Allies
(fossil).]
[Family Diatrymidae, Diatryma (fossil).]
[Gastornithidae, Gastornis, Remiornis (fos-
sil).]
Order Charadriiformes, Shore Birds, Gulls, Auks.
Suborder Charadrii, Shore Birds.
Superfamily Jacanoidea, Jacanas.
Family Jacanidae, Jacanas.
Superfamily Charadrioidea, Plovers, Sandpipers, and Al-
lies.
2 Position provisional.
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
[Family Rhegminornithidae, Rhegminornis (fossil).]
Rostratulidae, Painted Snipe.
Haematopodidae, Oystercatchers.
Charadriidae, Plovers, Turnstones, Surfbirds.
Scolopacidae, Snipe, Woodcock, Sandpipers.
Recurvirostridae, Avocets, Stilts.
[| Presbyornithidae, Presbyornis (fossil).]
Phalaropodidae, Phalaropes.
Superfamily Dromadoidea, Crabplovers.
Family Dromadidae, Crabplovers.
Superfamily Burhinoidea, Thick-knees.
Family Burhinidae, Thick-knees.
Superfamily Glareoloidea, Pratincoles, Coursers.
Family Glareolidae, Pratincoles, Coursers.
Superfamily Thinocoroidea, Seedsnipe.
Family Thinocoridae, Seedsnipe.
Superfamily Chionidoidea, Sheathbills.
Family Chionididae, Sheathbills.
Suborder Lari, Gulls, Terns, Skimmers.
Family Stercorariidae, Skuas, Jaegers.
Laridae, Gulls, Terns.
Rynchopidae, Skimmers.
Suborder Alcae, Auks.
Family Alcidae, Auks, Auklets, Murres.
Order Columbiformes, Sandgrouse, Pigeons, Doves.
Suborder Pterocletes, Sandgrouse.
Family Pteroclidae, Sandgrouse.
Suborder Columbae, Pigeons, Doves.
Family Raphidae, Dodos, Solitaires.
Columbidae, Pigeons, Doves.
Order Psittaciformes, Lories, Parrots, Macaws.
Family Psittacidae, Lories, Parrots, Macaws.
Order Cuculiformes, Plantain-eaters, Cuckoos.
Suborder Musophagi, Plantain-eaters.
Family Musophagidae, Plantain-eaters, Touracos.
Suborder Cuculi, Cuckoos, Roadrunners, Anis.
Family Cuculidae, Cuckoos, Roadrunners, Anis.
Order Strigiformes, Owls.
[Family Protostrigidae, Protostrix (fossil).]
Tytonidae, Barn Owls.
Strigidae, Typical Owls.
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 29
Order Caprimulgiformes, Oilbirds, Goatsuckers.
Suborder Steatornithes, Oilbirds,
Family Steatornithidae, Oilbirds.
Suborder Caprimulgi, Frogmouths, Goatsuckers.
Family Podargidae, Frogmouths.
Nyctibiidae, Potoos.
Aegothelidae, Owlet-frogmouths.
Caprimulgidae, Goatsuckers,
Order Apodiformes, Swifts, Hummingbirds.
Suborder Apodi, Swifts.
[Family Aegialornithidae,? Aegialornis (fossil).]
Apodidae, Swifts.
Hemiprocnidae, Crested Swifts.
Suborder Trochili, Hummingbirds.
Family Trochilidae, Hummingbirds.
Order Coliiformes, Colies.
Family Coliidae, Colies.
Order Trogoniformes, Trogons.
Family Trogonidae, Trogons.
Order Coraciiformes, Kingfishers, Bee-eaters, Rollers, Horn-
bills.
Suborder Alcedines, Kingfishers, Todies, Motmots.
Superfamily Alcedinoidea, Kingfishers.
Family Alcedinidae, Kingfishers.
Superfamily Todoidea, Todies.
Family Todidae, Todies.
Superfamily Momotoidea, Motmots.
Family Momotidae, Motmots.
Suborder Meropes, Bee-eaters.
Family Meropidae, Bee-eaters.
Suborder Coracii, Rollers, Hoopoes.
Family Coraciidae, Rollers.
Brachypteraciidae, Groundrollers.
Leptosomatidae, Cuckoo-rollers.
Upupidae, Hoopoes.
Phoeniculidae, Woodhoopoes.
Suborder Bucerotes, Hornbills.
Family Bucerotidae, Hornbills.
3 Position provisional.
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Order Piciformes, Jacamars, Barbets, Toucans, Woodpeckers.
Suborder Galbulae, Jacamars, Barbets, Toucans.
Superfamily Galbuloidea, Jacamars, Puffbirds.
Family Galbulidae, Jacamars.
Bucconidae, Puffbirds.
Superfamily Capitonoidea, Barbets, Honeyguides.
Family Capitonidae, Barbets.
Indicatoridae, Honeyguides.
Superfamily Ramphastoidea, Toucans.
Family Ramphastidae, Toucans.
Suborder Pici, Woodpeckers.
Family Picidae, Woodpeckers, Piculets.
Order Passeriformes, Perching Birds.
Suborder Eurylaimi, Broadbills.
Family Eurylaimidae, Broadbills.
Suborder Tyranni, Ovenbirds, Tyrant Flycatchers, and Al-
lies
Superfamily Furnarioidea, Ovenbirds, Woodhewers, and
Allies.
Family Dendrocolaptidae, Woodhewers.
Furnariidae, Ovenbirds.
Formicariidae, Ant-thrushes.
Conopophagidae, Antpipits.
Rhinocryptidae, Tapaculos.
Superfamily Tyrannoidea, Tyrant Flycatchers, Pittas, and
Allies.
Family Cotingidae, Cotingas.
Pipridae, Manakins.
Tyrannidae, Tyrant Flycatchers,
Oxyruncidae, Sharpbills.
Phytotomidae, Plantcutters.
Pittidae, Pittas.
Acanthisittidae, New Zealand Wrens.
Philepittidae, Asities, False Sunbirds.
Suborder Menurae, Lyrebirds.
Family Menuridae, Lyrebirds.
Atrichornithidae, Scrubbirds.
Suborder Passeres, Songbirds.
Family Alaudidae, Larks.
[ Palaeospizidae, Palaeospiza (fossil) .]
Hirundinidae, Swallows.
Dicruridae, Drongos.
No. II
CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 31
Oriolidae, Old World Orioles.
Corvidae, Crows, Magpies, Jays.
Cracticidae, Bell Magpies, Australian Butcher-
birds,
Grallinidae, Magpie-larks.
Ptilonorhynchidae, Bowerbirds.
Paradisaeidae, Birds of Paradise.
Paridae, Titmice.
Sittidae, Nuthatches.
Hyposittidae, Coralbilled Nuthatches.
Certhiidae, Creepers.
Paradoxornithidae, Parrotbills, Suthoras.
Chamaeidae, Wrentits.
Timaliidae, Babblers.
Campephagidae, Cuckoo-shrikes.
Pycnonotidae, Bulbuls.
[Palaeoscinidae,* Palaeoscinis (fossil).]
Chloropseidae, Leafbirds.
Cinclidae, Dippers.
Troglodytidae, Wrens.
Mimidae, Thrashers, Mockingbirds.
Turdidae, Thrushes.
Zeledoniidae, Wrenthrushes.
Sylviidae, Old World Warblers.
Regulidae, Kinglets.
Muscicapidae, Old World Flycatchers.
Prunellidae, Accentors.
Motacillidae, Wagtails, Pipits.
Bombycillidae, Waxwings.
Ptilogonatidae, Silky Flycatchers.
Dulidae, Palmchats.
Artamidae, Woodswallows.
Vangidae, Vanga Shrikes.
Laniidae, Shrikes.
Prionopidae, Woodshrikes.
Cyclarhidae, Peppershrikes.
Vireolaniidae, Shrike-vireos.
Callaeidae, Wattled Crows, Huias, Saddlebacks.
Sturnidae, Starlings.
Meliphagidae, Honey-eaters.
4 Allocation to this position is tentative.
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
Nectariniidae, Sunbirds.
Dicaeidae, Flowerpeckers.
Zosteropidae, White-eyes.
Vireonidae, Vireos.
Coerebidae, Honeycreepers.
Drepanididae, Hawaiian Honeycreepers.
Parulidae, Wood Warblers.
Ploceidae, Weaverbirds.
Icteridae, Blackbirds, Troupials.
Tersinidae, Swallowtanagers.
Thraupidae, Tanagers.
Catamblyrhynchidae, Plushcapped Finches.
Fringillidae, Grosbeaks, Finches, Buntings.
December 31, 1959.
LITERATURE CITED
AMADON, DEAN.
1950. Australian mud builders. Emu, vol. 50, pt. 2, Oct., pp. 123-127.
1957. Remarks on the classification of the perching birds [order Passeri-
formes]. Proc. Zool. Soc. Calcutta, Mookerjee Mem. Vol., May 1,
pp. 259-268.
ASHLEY, JAMES F.
1941. A study of the structure of the humerus in the Corvidae. Condor,
vol. 43, No. 4, July 15, pp. 184-195, figs. 50-56.
BEECHER, WILLIAM J.
1953. A phylogeny of the Oscines. Auk, vol. 70, No. 3, July, pp. 270-333,
18 figs.
Bock, WALTER J.
1956. A generic review of the family Ardeidae (Aves). Amer. Mus. Nov.,
No. 1779, July 27, pp. 1-49.
1958. A generic review of the plovers (Charadriinae, Aves). Bull. Mus.
Comp. Zool., vol. 118, No. 2, March, pp. 27-97, 6 figs.
BONAPARTE, PRINCE.
1855. (Remarks, in proceedings of Académie des Sciences de Paris.) Rev.
et Mag. Zool., ser. 2, vol. 7, Oct., pp. 482-485.
CaBANIS, J.
1847. Ornithologische Notizen. II. Arch. Naturg., vol. 1, pp. 308-352.
CATESBY, MARK.
1731. The natural history of Carolina, Florida and the Bahama Islands:
Containing the figures of birds, beasts, fishes, serpents, insects and
plants: Particularly the forest-trees, shrubs, and other plants, not
hitherto described, or very incorrectly figured by authors. Vol. 1,
Pp. i-xii, I-100, 100 pls. London.
CotTAM, Patricia A.
1957. The pelecaniform characters of the skeleton of the shoe-bill stork,
Balaeniceps rex. Bull. Brit. Mus. (Nat. Hist.), Zool., vol. 5, No. 3,
July, pp. 1-71, 1 pl., 4 figs.
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 33
DaAMEs, W.
1897. Uber Brustbein, Schulter- und Beckengiirtel der Archaeopteryx.
Sitzb, preuss. Akad. Wiss. Berlin, vol. 38, Aug. 9, pp. 818-834.
DE BEER, SiR GAVIN.
1954. Archaeopteryx lithographica, a study based upon the British Museum
specimen. Pp. i-xii, 1-68, 16 pls., 9 text figs. British Museum
(Natural History), London.
DELACOUR, JEAN.
1946a. Les timeliines. Ois. Rev. Frang. Orn., n.s., vol. 16, pp. 7-36.
1946b. Notes on the taxonomy of the birds of Malaysia. Zoologica, vol. 31,
pt. 1, Apr. 29, pp. 1-8.
DELACcoUR, JEAN, and AMADON, DEAN.
1949. The relationships of Hypocolius. Ibis, vol. 91, No. 3, July, pp. 427-
429, pls. 19-20.
DELACOUR, JEAN, and VAURIE, CHARLES.
1957. A classification of the Oscines (Aves). Los Angeles County Mus.
Contr. Sci., No. 16, Oct. 31, pp. 1-6.
DRESSER, HENRY EELES.
1893. A monograph of the Coraciidae, or family of the rollers. Pp. i-xx,
I-111, 27 pls. London.
Forpes, WILLIAM ALEXANDER.
1880. On the anatomy of Leptosoma discolor. Proc. Zool. Soc. London,
1880, pp. 464-475, 5 figs.
GLoGER, CONSTANTIN LAMBERT.
1854. In J. Reinhardt, Bemerkungen zur Ornithologie Gronlands. Journ.
fiir Orn., vol. 2, No. 11, Sept., pp. 423-443.
Gray, G. R.
1869. Hand-list of genera and species of birds, distinguishing those contained
in the British Museum. Part I. Pp. i-xx, 1-404. London.
GREGORY, JOSEPH T.
1952. The jaws of the Cretaceous toothed birds, Ichthyornis and Hes-
perornis. Condor, vol. 54, No. 2, Mar. 26, pp. 73-88, 9 figs.
HELLER, FLortAn.
1959. Ein dritter Archaeopteryx-Fund aus den Solnhofener Plattenkalken
von Langenaltheim/Mfr. Mit einem Beitrag von Klaus Fesefeldt.
Erlanger Geol. Abh., vol. 31, June 10, pp. 1-25, 15 pls., 2 figs.
Howarp, HILDEGARDE.
1955. A new wading bird from the Eocene of Patagonia. Amer. Mus. Nov.,
No. 1710, Mar. 11, pp. 1-25, 8 figs.
1957a. A gigantic “toothed” marine bird from the Miocene of California.
Santa Barbara Mus. Nat. Hist., Dept. Geol. Bull. No. 1, Feb. 1,
pp. I-23, 8 figs.
1957b. A new species of passerine bird from the Miocene of California. Los
Angeles County Mus. Contr. Sci., No. 9, June 28, pp. 1-16, 2 figs.
Hupson, Georce; LANzILiott, Patricia J.; and Epwarps, GLENN D.
1959. Muscles of the pelvic limb in galliform birds. Amer. Midl. Nat.,
vol. 61, pp. 1-67, 20 figs.
JHERING, HERMAN VON.
1915. The classification of the family Dendrocolaptidae. Auk, vol. 32, No. 2,
Apr., pp. 145-153, pls. 11-12.
34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE.
1957. Opinions and declarations, vol. 13, pt. 22, Direction 75, June 21,
PP. 291-308.
LAMBRECHT, KALMAN.
1933. Handbuch der Palaeornithologie. Pp. i-xix, 1-1024, 4 pls., 209 text
figs. Berlin.
LANHAM, URLEss N.
1947. Notes on the phylogeny of the Pelecaniformes. Auk, vol. 64, No. 1,
Jan., pp. 65-70.
LATHAM, JOHN.
1787. General synopsis of birds, Supplement, vol. 1, pp. i-iv, 1-298 (with 8
pages of additions, and index), 13 pls.
LINNAEUS, C.
1758. Systema naturae per regna tria naturae, secundum classes, ordines,
genera, species, cum characteribus, differentiis, synonymis, locis.
Ed. 10, vol. 1, pp. 1-824.
Lowe, Percy R.
1925. On the classification of the Tubinares or Petrels. Proc. Zool. Soc.
London (Jan. 14, 1926), pp. 1436-1443, 2 figs.
1931. An anatomical review of the “waders” (Telmatomorphae), with spe-
cial reference to the families, sub-families, and genera within the
suborders Limicolae, Grui-Limicolae and Lari-Limicolae. Ibis,
Oct., pp. 712-771, figs. 19-30.
Lucas, FrReDERIC A.
1889. The main divisions of the swifts. Auk, vol. 6, No. 1, Jan., pp. 8-13,
1 fig.
1895. Additional characters of the Macropterygidae. Auk, vol. 12, No. 2,
Apr., pp. 155-157, I fig.
LYDEKKER, RICHARD.
1891. Catalogue of the fossil birds in the British Museum (Natural His-
tory). Pp. i-xxviii, 1-368, 75 figs. London.
Mayr, ERwnSsT.
1958. The sequence of the songbird families. Condor, vol. 60, No. 3, May
20, Pp. 194-195.
Mayr, Ernst, and AMADON, DEAN.
1951. A classification of recent birds. Amer. Mus. Nov., No. 1496, Apr. 2,
pp. 1-42.
Mayr, Ernst, and GrEENWAY, JAMES C., JR.
1956. Sequence of passerine families (Aves). Mus. Comp. Zool. Breviora,
No. 58, June 29, pp. I-11.
McDoweELL, SAM.
1948. The bony palate of birds. Part I. The Palaeognathae. Auk, vol. 65,
No. 4, Oct., pp. 520-540, 6 figs.
MILLER, ALDEN H.
1944. An avifauna from the lower Miocene of South Dakota. Univ. Cali-
fornia Publ., Bull. Dept. Geol. Sci., vol. 27, No. 4, June 22, pp. 85-
too, 8 figs.
Miter, Loye.
1946. The Lucas auk appears again. Condor, vol. 48, No. 1, Feb. 4, pp. 32-
36, fig. 3.
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 35
1950. A Miocene flamingo from California. Condor, vol. 52, No. 2, Mar.
15, pp. 69-73, fig. 16.
1952. The avifauna from the Barstow Miocene of California. Condor, vol.
54, No. 5, Sept. 22, pp. 206-301, 2 figs.
Miter, Loye, and Howarp, HI_pEcArRDE.
1949. The flightless Pliocene bird Mancalla. Carnegie Inst. Washington,
Publ. 584, June 22, pp. 202-228, 6 pls.
MiLtneE-Epwarps, ALPHONSE.
1863. Mémoire sur la distribution géologique des oiseaux fossiles et descrip-
tion de quelques espéces nouvelles. Ann. Sci. Nat., ser. 4, vol. 20,
Pp. 133-176, pl. 4.
1868-1871. Recherches anatomiques et paléontologiques pour servir a V’his-
toire des oiseaux fossiles de la France. Vol. 2, pp. 1-632. Paris.
MoyniHAN, MartIN.
1959. A revision of the family Laridae (Aves). Amer. Mus. Nov., No.
1928, Mar. 5, pp. 1-42.
NeEwtTon, ALFRED.
1896. A dictionary of birds. Pp. i-xii, 1-1088, 700 figs. in text. London.
OBERHOLSER, Harry C,
1917. Diagnosis of a new pycnonotine family of Passeriformes. Journ.
Washington Acad. Sci., vol. 7, Oct. 19, pp. 537-541.
[OKeEN, LorENz.]
1839. In Isis von Oken, vol. 33, pt. 9, p. 673.
PATTERSON, BRYAN.
1941. A new phororhacid bird from the Deseado formation of Patagonia.
Field Mus. Nat. Hist., Geol. Ser., vol. 8, No. 8, Oct. 31, pp. 49-54.
PETRONIEVICS, BRANISLAV.
1921. Ueber das Becken, den Schultergiirtel und einige andere Teile der
Londoner Archaeopteryx. Pp. 1-31, 2 pls. Genf.
1925. Ueber die Berliner Archaeornis. Beitrag zur Osteologie der Archae-
ornithes. Ann. Géol. Pén. Balkanique, vol. 8, fasc. 1, pp. 37-84,
6 pls. (Issued also as a separate, repaged, with addition of a table
of contents. )
1950. Les deux oiseaux fossiles les plus anciens (Archaeopteryx et Archae-
ormis). Ann. Géol. Pén. Balkanique, vol. 18, pp. 105-127, 25 figs.
(On pp. 89-103 the paper is presented in Bulgar.)
Petronievics, BRANISLAV, and WoopwarpD, ARTHUR SMITH.
1917. On the pectoral and pelvic arches of the British Museum specimen
of Archaeopteryx. Proc. Zool. Soc. London, 1917, No. 1, Apr. 20,
pp. I-6, pl. 1.
PycrarFt, W. P.
1906. Contributions to the osteology of birds. Part VIII. The “Tracheo-
phone” Passeres, with remarks on families allied thereto. Proc.
Zool. Soc. London, 1906, pp. 133-159, figs. 49-52.
1907. Contributions to the osteology of birds. Part IX. Tyranni; Hirun-
dines; Muscicapae, Lanii, and Gymnorhines. Proc. Zool. Soc. Lon-
don, 1907, pp. 355-379, figs. 101-104.
Ray, JoHNn.
1713. Synopsis methodica avium. Pp. 1-198, index, 2 pls. London.
36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139
RipGway, Rosert.
1914. The birds of North and Middle America. U.S. Nat. Mus. Bull. 50,
pt. 6, Apr. 8, pp. i-xx, 1-882, 35 pls.
Rusconi, CARLOS.
1946. Ave y reptil Oligocenos de Mendoza. Bol. Pal. Buenos Aires, No. 21,
May 2, pp. 1-3, 2 figs.
SAVILE, DoucLas Barton OSBorRNE.
1957. The primaries of Archaeopteryx. Auk, vol. 74, No. 1, Jan., pp. 90-
101.
ScLaTEr, Puivip LuTtey.
1865. On the structure of Leptosoma discolor. Proc. Zool. Soc. London,
1865, pp. 682-688, 8 figs.
SHUFELDT, Robert W.
1923. Comparative osteology of the Australian mud-builders. Emu, vol. 23,
pt. 1, July, pp. 4-19, pls. 4-6.
Srmpson, GEORGE GAYLORD.
1946. Fossil penguins. Bull. Amer. Mus. Nat. Hist., vol. 87, art. 1, Aug. 8,
PP. 1-99, 33 figs.
STERNBERG, R. M.
1940. A toothless bird from the Cretaceous of Alberta. Journ. Pal., vol. 14,
No. 1, Jan., pp. 81-85, 6 figs.
STEINER, HANs.
1938. Der “Archaeopteryx’”—Schwanz der Vogelembryonen. Vierteljahrsch.
Naturf. Ges. Ziirich, vol. 83, Beibl. 30, Festschr. Karl Hescheler,
Nov. 3, pp. 279-300, 5 text figs., 1 pl.
Stonor, C. R.
1937. On the systematic position of the Ptilonorhynchidae. Proc. Zool
Soc. London, vol. 107, ser. B, pt. 3, Sept., pp. 475-490.
1942. Anatomical notes on the New Zealand wattled crow (Callaeas), with
especial reference to its powers of flight. Ibis, ser. 14, vol. 6, No. 1,
Jan., pp. 1-18, 10 figs., tables 1-3.
Storer, R. W.
1959. The arrangement of songbird families. Condor, vol. 61, No. 2, Mar.
31, PP. 152-153.
STRESEMANN, Erwin.
1927-1934. Aves, in Kiikenthal und Krumbach, Handbuch der Zoologie,
vol. 7, pt. 2, sect. 8, pp. i-xii, 1-890, 944 figs.
TorpoFF, HARRISON B.
1954a. A systematic study of the avian family Fringillidae based on the
structure of the skull. Univ. Michigan Mus. Zool. Misc. Publ.
No. 81, Apr. 26, pp. 1-42, 77 figs.
1954b. Relationships in the New World nine-primaried Oscines. Auk, vol.
71, No. 3, July, pp. 273-284, 2 figs.
VAURIE, CHARLES.
1959. The birds of the Palearctic fauna, a systematic reference: Order
Passeriformes. Pp. i-xii, 1-762. London.
VERHEYEN, RENE.
1958. Contribution a la systematique des Alciformes. Bull. Inst. Roy. Sci.
Nat. Belgique, vol. 34, No. 45, Dec., pp. 11-14.
NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 37
WETMORE, ALEXANDER.
1919. A note on the eye of the black skimmer (Rynchops nigra). Proc.
Biol. Soc. Washington, vol. 32, Dec. 31, p. 105.
1930. A systematic classification for the birds of the world. Proc. U. S.
Nat. Mus., vol. 76, art. 24, Jan. 8, pp. 1-8.
1951. A revised classification for the birds of the world. Smithsonian Misc.
Coll., vol. 117, No. 4, Nov. 1, pp. 1-22.
1956. A check-list of the fossil and prehistoric birds of North America and
the West Indies. Smithsonian Misc. Coll., vol. 131, No. 5, Jan. 25,
pp. I-105.
1957. The classification of the oscine Passeriformes. Condor, vol. 59, No. 3,
May 24, pp. 207-209, 1 fig.
WILLUGHBY, FRANCIS.
1676. Ornithologiae, libri tres in quibus aves omnes hactenus cognitae in
methodum naturis suis convenientem redactae accurate describuntur,
descriptionis iconibus elegantissimis & vivarum avium simillimis,
aeri incisis illustrantur. Pp. 1-307 (with introduction and index),
77 pls. London.
ZIMMER, JOHN T.
1942. Studies of Peruvian birds. No. 41. The genera Hylophilus, Smarag-
dolanius, and Cyclarhis. Amer. Mus. Nov., No. 1160, Jan. 30,
pp. 1-16.
1
| pe
|i i
ie Sete OY |) iis
Hite emer nn tee
=
vy,
” i
i: f ;
,
e
tae
YY, “
Yn
NII
il
9088 01 421 5289
SMITHSONIAN INSTITUTION LIBRARIES
|
|
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11