}

■t'"'

JANUARY, 1953

RVCIFIC

SCIENCE

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

IN THIS ISSUE: Banner — The Crangonidae, or Snapping Shrimp,
of Hawaii • NEWS NOTES • INDEX

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PACIFIC SCIENCE

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

VOL. VII JANUARY, 1953 ' NO. 1

Previous issue published October 3, 1952

CONTENTS

PAGE

The Crangonidae, or Snapping Shrimp, of Hawaii.

Albert H. Banner 3

NEWS NOTES 145

INDEX 147

Pacific Science, a quarterly publication of the University of Hawaii, appears in Janu-
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Crangon diadema (Dana)

The Crangonidae, or Snapping Shrimp, of Hawaii^

ALBERT H. BANNER^

INTRODUCTION

The decapod family Crangonidae, or the
family Alpheidae of the earlier workers and of
the present European workers, is a family of
shrimp known popularly as snapping or pistol
shrimp. The members of the family seldom
exceed 30 millimeters in length, are usually
of stout shape, are noticeably laterally com-
pressed, and, though their color is variable,
are most often reddish- to grayish-white in
color. The outstanding characteristics of the
most common genera of the family are the
large size and asymmetrical development of
the chela of the first pereiopod. It is the vio-
lent closure of the enlarged chela that pro-
duces the clicking or snapping sound which
has given the members of the family their
common name.

Considering how common the snapping
shrimp are in Hawaii, surprisingly little work
has been done on them. The earliest record of
members of this family from the Hawaiian
Islands was made by Randall in 1839 when he
reported Crangon ventrosa (Milne-Edwards) as
Alpheus laevis Randall. Dana in 1852 [this vol-
ume was distributed on Eebruary 4, 1853, ac-
cording to Haskell (1942: 79) and therefore
for questions of nomenclatorial priority the
later date should be used] reported upon the
collections made by the United States Explor-
ing Expedition and listed three species, all
new, from the Hawaiian Islands. Stimpson in
I860 reported four species from Hawaii. Only
two species were reported from Hawaii by
Bate in the Challenger Reports in 1888. Lenz
in 1901 reported a single species from Laysan

1 Contribution No. 27, Hawaii Marine Laboratory.

2 Department of Zoology and Entomology, Univer-
sity of Hawaii, Honolulu, Hawaii. Manuscript received
July 5, 1951.

Island, a species previously known from the
Hawaiian Islands. Coutiere in The Fauna and
Geography of the Maidive and Laccadive Archi-
pelagoes (1906) reported five species from the
Hawaiian Islands — three on the basis of re-
ports of previous workers, one on a specimen
in the Museum of Paris, and one without fur-
ther reference (the last species is discussed on
page 142). In 1909 Coutiere, in his treatment
of the synalpheids of the United States Na-
tional Museum, listed four species of the
genus Synalpheus collected in Hawaiian waters
by the steamer "Albatross.” By far the most
extensive studies on the Hawaiian crangonids
were those of Edmondson. In 1925 he re-
ported on 17 species collected by the "Tana-
ger” in the western islands and shoals of the
Hawaiian Archipelago; in 1930 he reported
two new species of a genus previously un-
known to Hawaii; and in 1933 in his book.
Reef and Shore Fauna of Hawaii, he gives a brief
description of 13 species.

The geographical limits of the present study
coincide with the geographical limits of the
Hawaiian Archipelago (Fig. 1). Most of the
collecting has been done around the larger or
windward islands of the archipelago, which
are, from east to west, Hawaii, Maui, Kahoo-
lawe, Lanai, Molokai, Oahu, Kauai, and Nii-
hau. Extending over a thousand miles to the
west and north of these large islands is a chain
of shoals and normally uninhabited small is-
lands known as the leeward islands; these are,
again in their order from east to west, Nihoa,
Necker, French Frigate Shoal, Gardner Pin-
nacles, Raita Bank, Maro Reef, Laysan, Lisi-
anski. Pearl and Hermes Reef, Midway, and
Kure, or Ocean, Island.

The present study was begun in the year
1937-38, when quite extensive collections

C 3 }

4

PACIFIC SCIENCE, Vol. VII, January, 1953

were made of the species that could be col-
lected in the very shallow water of the reef
flats (in water up to about 5 feet deep). The
specimens in the collections of the Bernice P.
Bishop Museum also were examined at that
time. My departure from the Islands in the
fall of 1938 prevented the completion of the
study, and the work was put aside until my
return to Hawaii in 1946. At that time a new
series of collections was made, the new col-
lection in large part being made in the waters
beyond the reef flats, the specimens being
taken by '"skin-diving” in water up to 20 or
25 feet deep. The specimens taken by the
United States Fish Commission Steamer "Al-
batross” in the Hawaiian Archipelago in 1902
were obtained on loan from the United States
National Museum; these were a valuable con-
tribution to the study as they represented to a
large extent a deeper fauna that was impossi-
ble to collect without dredging. Additional
specimens in the Bishop Museum were ex-
amined. Some deep-water specimens dredged
by the "Makua” of the Territorial Board of
Agriculture and Forestry and by the "Salpa”
of the University of Hawaii were available.
Finally, I was loaned a small but very impor-
tant collection of crangonids taken off the
island of Hawaii by R. W. Hiatt. Several
thousand specimens from these various
sources were examined.

I believe that this study is quite complete
for the crangonid fauna of the Hawaiian Is-
lands. It is true that collections were not made
from all the reefs on all the islands, but enough
collections were made on Oahu, Maui, Ha-
waii, and the leeward islands of the archi-
pelago to show that the fauna does not seem
to vary to any major extent from island to is-
land. On Oahu all possible types of habitats
have been examined, from muddy bays to the
clean coral growing on the outer sides of
wave-swept reefs. The deep-water dredgings
of the "Albatross” and other ships have sam-
pled extensively the deeper fauna from the
islands. On the other hand, the several spe-
cies in the collections that are represented by

a single or several specimens indicate that
other rare species not collected may exist in
the islands. However, it is unlikely that spe-
cies not discussed here will be found in the
course of routine collecting.

Measurements

In the numerous measurements used in this
paper, the following standards are adopted.
For total length, the body is measured from the
tip of the rostrum to the end of the telson-
following the middorsal line of the body. If
the abdomen is flexed, a slightly greater meas-
urement is obtained than if it were straight,
but the difference is not great. For the append-
ages, the breadth is taken at the broadest point
unless otherwise specified; the length is meas-
ured from articular surface to articular sur-
face (this is especially important in the carpal
articles, as the articular surface is toward the
inferior side of the meral-carpal joint, and, if
the maximum length to the edge of the su-
perior portion of the article were taken, it
would be found to vary greatly with the de-
gree of flexion of the joint). The lengths of the
antennular articles are taken in the mid-line of
the article in dorsal view, with the exception
of the first article, where the length is taken
from the most proximal portion visible dor-
sally to the middle of the distal end. The
height or breadth of the large chela is taken to be
the diameter from superior to inferior surface,
the thickness as the diameter at right angles to
this height, the length of the chela as including
the fingers; unless otherwise specified the
length of the fingers is given as a proportion of
the length of the entire chela. The breadth of
the merus of the chelipeds, in those cases where
the cheliped was definitely triangular in cross
section, is taken as the width of the outer face
of the triangle; the length as from the middle
of the ischial-meral articulation to the meral-
carpal articulation. Finally, throughout the
paper the relative lengths of the carpal articles of
the second legs are expressed as ratios based
on the length of the first article, to which the
value 10 is assigned; it is possible to carry

Hawaiian Crangonidae — BANNER

5

175*

165*

55*

30*

Kure

20*

Midway

179*

Pearl and flermcs Kecf

I

Lisianski ,

. Laysan

Maro Reef

— N

Raiia Dank

Gardner Pinnaele?

Necker

"French Friga'tc Shoal

300 Nautical Miles

Nihoa

Niihau^

Kaula

Kauai

Oahu

Molokai

Lanai»Jf^Ma

Kahoola we

•Hawaii

m

160*

Fig. 1. Map of the Hawaiian Archipelago, showing principal islands, reefs, and shoals.

these ratios to two or more significant places,
but in most cases the figures are rounded to
the first significant place (for example, 10 :
7 : 5 : 5 : 7) because so much variation was
found in the relative lengths of the articles
within one species that it is felt that any fur-
ther refinement of figures would be scientifi-
cally inaccurate. The measurements of the tel-
son are self-evident. Throughout the study,
fractions are expressed as their decimal equiv-
alents and rounded with the exception of one
fourth and three fourths.

Acknowledgments

I wish to thank the following institutions
and individuals for their assistance in this
work: The Bernice P. Bishop Museum; the
United States National Museum; Dr. C. H.
Edmondson, Professor Emeritus of the Uni-
versity of Hawaii and Curator of Zoology at
the Bernice P. Bishop Museum, for his help
and cooperation in all phases of the work, es-
pecially during the initial year’s study; Dr. R.
W. Hiatt, Professor of Zoology at the Uni-
versity of Hawaii, for the use of collections of
the shrimp made by him; Dr. Fenner A. Chace,
Jr., of the United States National Museum for

his cooperation and advice on several points;
and Dr. L. D. Tuthill, Editor of Pacific Science,
for advice on several taxonomic points.

I also wish to acknowledge the help of the
following artists employed by the University
of Hawaii. Miss Inger Achton drew Figures
6^-/, j-l. Mrs. Florence Lambeth drew Fig-
ures 14^-^ , \ld-e, l^a-d, /, /, j, m, and 24
a-h. Evan I. Gillespie drew the frontispiece,
redrew Figures 1, 12^-c, A2a-h from their
original sources, and completed the following
drawings after I had delineated them: Fig-
ures 16 a-j; lla-c,f-j; 18^, g, h, k, 1; I9a-g;
20a-m; l)Aa-i; 37 ; Alg-h; Aha-k; AAa-l; 45
a-g; A6a-i; Ala-k; A^a-h; 50^-/.

DISTRIBUTION

The members of the family Crangonidae
are commonly circumtropical in distribution
although in the Pacific Ocean they have been
reported from as far north as Sitka, Alaska
{Betaeus harrimani Rathbun, 1910), and as far
south as Hermite Island, Tierra del Fuego {Be-
taeus truncatus Dana, 1852). The center of dis-
tribution for the family in the Pacific appears
to be the East Indian Archipelago, from which
de Man (1911) reported 113 species and 20

6

PACIFIC SCIENCE, VoL VII, January, 1953

varieties. Though he noted that the members
of the family were largely limited to the upper
littoral 2one of the sea, four species of Syn-
alpheus and eight species of Crangon were
dredged from depths greater than 100 meters
during the Siboga Expedition. Two species
have been reported from tropical fresh -water
lakes.

Like most of the marine fauna of the Ha-
waiian Archipelago, the crangonids belong to
the Indo-Pacific faunal group and not to the
Western American. Those species that are not
endemic range southward and westward, in
many instances as far as the Red Sea. Crangon
ventrosa (Milne-Edwards) is the only excep-
tion to this statement: its range extends from
the Red Sea through Hawaii to the Gulf of
California.

In the present state of knowledge of Crus-
tacea in the Pacific it is useless to speculate on
the distributional patterns of the nonendemic
species. There are too many islands and archi-
pelagoes where no collecting of marine inver-
tebrates has been done at all; on the few major
islands where some collecting has been done,
it was done usually in a random fashion, and
the lists of species cannot be regarded as even
approaching completion. The few exceptions
to this in the tropical Indo-Pacific region are
the area around the mouth of the Red Sea,
those portions of the Indian Ocean that were
visited by J. Stanley Gardiner, and the portion
of the southwestern Pacific visited by the Si-
boga Expedition. It is likely that the report on
the Bikini collections, now in preparation by
Fenner A. Chace, Jr., of the United States Na-
tional Museum, will add another area. But
whether the range of a species extends from
Hawaii to the Marianas or to the Marquesas
or to New Caledonia is not known.

Without the knowledge of the geographic
range of the species, speculation on paths of
distribution would be without basis. As the
adults are bottom dwellers, and as the larvae
are planktonic, it is safe to assume that most
of the species that reached Hawaii were carried
here as larvae. However, with the preponder-

ance of the currents in this portion of the
Pacific flowing from Hawaii toward the west-
ern Pacific, it is difficult to understand how
these larvae, without any marked powers of
locomotion, were able to reach Hawaii from
the closest island groups, over a thousand
miles away. It is likely that they were carried
here by fortuitous combinations of eddies and
temporary shifts in the superficial currents in
changing weather conditions.

It is likewise fruitless to speculate on the
number of endemic species. According to the
present records 19 of the 44 species recorded
from these islands, or some 45 per cent, are
known only from the Hawaiian Archipelago.
However, no one knows how many of these
species may occur at Johnston Island or Can-
ton Island or in Tahiti or Samoa; no one
knows how many of these species may reach
to the northern Marianas where conditions
somewhat similar to those in Hawaii exist. It
is to be expected that the isolated position of
Hawaii would produce some speciation, but
how much cannot yet be determined.

ECOLOGY

In Hawaii the members of the family seem
to be confined largely, if not exclusively, to
the eulittoral zone, which is deeper in tropical
than in temperate waters. Many of the species
are found exclusively in the shallow water of
reef flats and in waters less than 50 feet deep
at the outer side of the reef; a few have never
been collected except on the reef flat itself,
where at low tide the water varies from only a
few inches to several feet in depth. The spe-
cies inhabiting the more shallow waters are
found in five types of habitats: in dead corals,
where they live in cracks, folds, old worm
holes, or even in tubes of algae which they
construct for themselves; in living coral heads,
among the branches of the coral; in sandy or
muddy portions of the inner reef and of bays,
where they hide about the bases of rocks and
partially buried objects; in the dense growth
of algae found on exposed wave-beaten
coasts; and commensally with sponges and

Hawaiian Crangonidae — Banner

7

tube worms. Lack of data prevents the tabu-
lation of the habitats of the deeper- dwelling
species, but the few I have collected in dredge
hauls from deeper water were either living in
dead coral boulders or in sponges.

Although the general habitats of the spe-
cies are known, very little is known of their
actual ecology. For example, what are their
food habits.^ They are reported to be carnivor-
ous, yet the digestive tracts of specimens kept
in aquaria with Ulva turn a bright green. In
the case of a dozen species living in one dead
coral head, are there zones typical of each spe-
cies, and, if so, what factors determine this
selection? Does a pair found living together
in a single tube represent a pair mated for life?
What are the food and defensive relationships
of this pair in which the smaller male has the
larger chela? What is the biological signifi-
cance of the sexual dimorphism of the small
chela of some species? Why are the two close-
ly related species, Crangon crassimanus (Heller)
and C. pad fica (Dana), which seem to inhabit
the same type of coral in the same zone, al-
most never found together in the same coral
head? What are the limitations of the distri-
butional range of C. diadema (Dana)? To my
knowledge, none of these problems has ever
been investigated.

VARIATION WITHIN SPECIES

A number of species reported upon in this
study are noted to show great variation, a var-
iation which transcends the limits previously
accepted to be of subspecific and, in some
cases, of specific worth. These variations are
encountered particularly in the armature and
proportions of the appendages, characteristics
that have been accepted as stable within quite
narrow limits for a single species. Some of the
species from Hawaii, especially those that
have been collected in large numbers, like
Synalpheus paraneomeris (Coutiere), Crangon
hrevtpes (Stimpson) , and C. dypeata (Coutiere) ,
show great variation; many of those that are
less plentiful, such as S. charon (Heller) and C.
edmondsoni Banner, show less variation. There

are occasional species that appear to be rather
constant and do not show as much variation
as would be expected from observation of re-
lated species collected in similar numbers,
e. g., C. pad fica (Dana), C. nanus Banner, and
C. collumiana (Stimpson).

This apparently unusual degree of variation,
reported in detail in the text, may be due to
three factors:

The first possibility is the lack of discrimi-
nation on my part, resulting in my confusing
several discrete but narrowly separated spe-
cies and considering them as a single species.
However, whenever during this study any
considerable variation of a species was noted,
that species was examined critically, and, in
instances of complicated variations, tables of
measurements were prepared to facilitate the
drawing of conclusions. Characters which ex-
hibited a more or less continuous spectrum of
variation were not considered as of specific
worth. Although other workers have some-
times used a combination of variable charac-
ters as a basis for distinguishing species, I do
not believe that, even if the procedure is valid,
the knowledge of the family is sufficiently ad-
vanced to warrant the use of such criteria.

A second possible cause of the variation is
a species radiation in Hawaii. This would be
caused by the immigration of one or a few
species to an isolated location and then the
rapid evolution of the species to fit into the
unoccupied ecological niches. When suffi-
cient time has elapsed, this process should
produce many new but closely related spe-
cies; when the process is just starting, it should
be noticeable as variations within a species. It
is true that the Hawaiian Archipelago has
fewer species than other parts of the Pacific;
for example, de Man (1911) reported 38 spe-
cies of the genus Synalpheus from the area vis-
ited by the Siboga Expedition, whereas in Ha-
waii the genus is represented by only six
species, three of which are common. There-
fore, those three species in Hawaii may be in
the process of evolution to fit the ecological
niches filled by the 38 species of the Nether-

8

PACIFIC SCIENCE, Vol. VII, January, 1953

lands Indies. In Hawaii this type of species ra-
diation has been observed in groups as diverse
as the birds, the drosophilids, and the land
snails. However, possibly because the marine
forms have not been as completely isolated,
or possibly because the marine animals have
not been as thoroughly studied, the phenom-
enon ha,s not been especially noted in the
animals inhabiting the reefs and shores.

The third possibility is that the variation is
not unusual or unique to Hawaii and occurs
in all areas but has been overlooked to a large
extent by other workers. They may not have
had a large enough series of specimens of one
species to give a picture of the variation, or,
having the specimens, they may have been too
imbued with the type-specimen concept to
discuss the variations in their papers. It is true
that in many cases the earlier workers did dis-
cuss these variations; de Man, for example,
presented long numerical tables on the varia-
tions in some of his species. Yet in his keys,
de Man (1911: 192) used characteristics that
could easily overlap, e.g.,

i. Rostrum usually 3-times (2.75-3.25) as long as

wide. . . . Dactylus of third legs 3-times as long

as thick at the base. . . .

ii. Rostrum 2 2-times as long as wide. . . . Dactylus of

third legs 2.51 times as long as thick at its base. . . .

Coutiere has erected species upon differ-
ences with even less allowance for variation.
De Man states in his introduction (1911: 133) :

Coutiere also first called attention to the great impor-
tance of relative measurements . . . and it was just by
means of these new characters that often specimens of
small size of Alpheus and Synalpheus proved to be species
that were still unknown, while formerly such specimens
usually would be regarded as juvenile forms or at most
varieties.

Coutiere stated (1909: 36) that some of these
species were even based largely on "geo-
graphic characteristics." For example, "5.
paulsoni and the other Indo-Pacific forms are
not represented [in America] by identical
forms, but the differences are at times so
slight that, without indication of locality, the
identification would be very difficult."

The lack of a sufficient number of speci-
mens of a given species to show the extent of

variation often renders it necessary to erect
new species for what may later be found to be
divergent members of the same species. This
has been done in the past, and I fear that in
this paper it may have been done again. For
example, Crangon pseudopugnax Banner is sep-
arated from C percyi (Coutiere) chiefly by the
differences in the base of the rostrum, and the
character seems to be quite clear-cut and de-
cisive. However, of the two species only nine
specimens are known from Hawaii, and per-
haps, if a large number of specimens were col-
lected, it might be found that the two species
described are merely the extremes of a vari-
able single species, with specimens of inter-
mediate form of the rostrum more plentiful.

Only by extensive studies can it be deter-
mined which of these postulated causes for
the observed variation is the correct one. To
arrive at a solution of the problem it will be
necessary to study not only a large series of
specimens from Hawaii but also many speci-
mens of these species from other portions of
their ranges. Specimens from the southwest
Pacific especially should be studied, as that
area appears to be the center of distribution
for the Indo-Pacific members of the family.
Through such a study some of the criteria
based on small differences in the relative pro-
portions, like those of the antennular or carpal
articles, will perhaps have to be discarded.
With any such change in the criteria, some of
the species now recognized will fall into sy-
nonymy. It may also be necessary to find new
and more stable characteristics upon which to
base specific descriptions.

In any case, the work done in this present
study is sufficient to cast doubts on the valid-
ity of some of the species that have been de-
scribed on the basis of slight differences in
what appear to be variable characteristics.

Family CRANGONIDAE Weber

Carapace smooth, provided with cardiac
grooves; rostrum reduced, antennal and bran-
chiostegal spines always absent; in known Ha-
waiian species carapace more or less project-

Hawaiian Crangonidae — Banner

9

ing over eyes. Antennular base cylindrical,
with basal article not longer than sum of other
two; scaphocerite rarely longer than pedun-
cles. Mandible always bipartite, with palpus
of 2 articles. Distal article of first maxilla bi-
furcate at its extremity. Epipodites of first and
second maxillipeds always undivided. Chela
of first thoracic leg predominant, always large.
(In the Hawaiian species the chelae are usual-
ly asymmetrically developed.) Carpus of che-
liped short, cup-shaped or hemispherical. Sec-
ond legs chelate, weakly developed. (The
multiarticulate carpus in Hawaiian species is
always composed of four to five articles.) Fol-
lowing legs short, compressed, with spinous
propodi (in Hawaiian species the dactyli are
either simple or biunguiculate) . Propodus of
the fifth leg with more or less well-developed
"brush” of bristles placed in transverse-
oblique rows. Abdomen usually with grad-
ual curve, without any pronounced bending
at third segment; sixth segment broad and
short; size and shape of pleura showing sexual
dimorphism. Pleopods of usual form, show-
ing only slight sexual dimorphism. The bran-
chial formula always includes five pleuro-
branchs. (Description adapted from Coutiere,
1899.)

Of the 18 (?) previously known genera of
the family Crangonidae, only 4 are represented
in the fauna of the Hawaiian Archipelago. In
addition, one new genus has been described;
this genus, however, is not endemic to the
Hawaiian Islands, as it includes one species
from the southwest Pacific which de Man
tentatively placed in the genus Alpheopsis Cou-
tiere. The great majority of the species in Ha-
waii are members of the genus Crangon Weber
as they are in all other investigated regions of
the Indo-Pacific.

The generic name Crangon and the family
name Crangonidae replaced the widely used
names Alpheus and Alpheidae as the result of
the work of Rathbun in 1904. At that time she
published the results of an examination of a
paper by Weber, published in 1795, in which

he used the name Crangon for species called
Alpheus by Fabricius in 1798. Her interpreta-
tion of this, according to the International
Rules of Zoological Nomenclature, made it
necessary to change the genus previously
known as Alpheus Fabricius to Crangon Weber
and the genus known as Crangon Fabricius to
Crago Famarck. She presented the case to the
International Commission on Zoological No-
menclature, and they confirmed her views in
Opinion 17: "Weber’s Nomenclator Entomologl-
cus 1793 complies with the requirements of
Article 25, hence the genera in question are to
be accepted. ...” Apstein in 1915 proposed
that the name Alpheus be a nomen conservandum^
but the proposal was rejected (although not
specifically for Alpheus versus Crangon^ by the
International Commission’s Opinion 74. The
whole controversy is reviewed by Hult (1938).
As a result of these rulings most American
and Australian workers are using the name
Crangon instead of Alpheus, whereas most Eu-
ropean workers are still using the name Al-
pheus. As this paper goes to press there is an
appeal to the International Commission for a
suspension of the rules to permit the re-estab-
lishment of the name Alpheus.

The position of the family Crangonidae dif-
fers but slightly in the various modern sys-
tems of classification of the lower decapods.
However, because the names and definitions
applied to the subsidiary classifications differ
with the various schemes, that of Balss (1927)
has been selected arbitrarily for use in this pa-
per. Balss followed, with only minor changes,
the scheme of classification earlier devised by
Borradaile. In it the Crangonidae are placed
in the suborder Natantia, the division Eucyph-
idae (which seems to be the equivalent of the
tribe Caridea of others), and the tribe Palae-
moniida (equivalent to the superfamily of the
same name used by other workers) . Except for
differences in names as pointed out above, all
present workers appear to be in accord on the
classification of the Crangonidae to this point.

Within the tribe Palaemoniida there is a
difference of opinion as to the relative position

10

PACIFIC SCIENCE, VoL VII, January, 1953

of the family with respect to the three or fout
other families. Primarily on the basis of the
characteristics of adults of the lower genera of
the family, Coutiere (1899) placed the cran-
gonids below but adjacent to the Hippolyti-
dae. Gurney (1938), on the basis of charac-
teristics of the larvae, concluded that the
family was most closely related to the Palae-
monidae and of higher phylogenetic position.

KEY TO THE HAWAIIAN GENERA

1. Eyes at most only partially concealed by

projection of margin of carapace, and
visible at least anteriorly; large chela
without a plunger on the dactylus fit-
ting into a cavity on the finger oppo-
site, but instead usually bearing a se-
ries of strong teeth 2

Eyes completely concealed by projec-
tion of anterior carapace and visible
only in anteroventral aspect; large
chela almost always with a plunger
that fits into a cavity on the. finger op-
posite, never serrate 4

2. Sixth abdominal segment without ar-

ticulated pleura; large chela carried
flexed toward body, with the palm
excavate to accommodate merus; tel-

son with tip emarginate

Jousseaumea (p. 10)

Sixth abdominal segment usually with
pleura articulated; if large chela is car-
ried flexed, then merus is flattened to
accommodate palm; telson with tip
either straight, slightly convex, or ex-
tended to form a triangle 3

3. Large chela carried extended; posterior

margin of telson straight to slightly

arcuate Alpheopsis (p. 14)

Large chela carried flexed; posterior mar-
gin of telson projecting as a strong
triangle Neoalpheopsis (p. 20)

4. With pterygostomial margin pro-

duced into a definite angle; without
anal tubercles; with only 2 epipod-

ites. . . .Synalpheus (p. 26)

With pterygostomial margin rounded,

not angular; with anal tubercles; with

7 to 8 epipodites Crangon (p. 46)

Jousseaumea Weber

Carapace extending forward as rostrum of
variable size and as orbital teeth forming
broad eave-like projection that conceals eyes
in dorsal view.

Antennular peduncle with strong stylocer-
ite. Scaphocerite broad, oval, with squamous
portion equal to or longer than lateral spine.
Carpocerite of antennular peduncle short and
broad, sometimes shorter than merocerite.

First chela very asymmetrical, carried flexed
at carpus so that distal end approaches body.
Fingers of large chela serrate with large teeth
joining exactly. Merus long.

Carpus of second legs composed of 5 ar-
ticles. Following legs robust with merus un-
armed, ischium bearing 2 spines, dactylus
simple. Sixth abdominal segment without ar-
ticulated pleura. No anal tubercles on telson.
Posterior margin of telson emarginate.

Branchial formula: 5 pleurobranchs, 1 ar-
throbranch, 8 epipodites.

DISCUSSION: The nine species recorded for
the world by de Man (I9II5 1922) have been
increased to eleven by the addition of the two
Hawaiian species described by Edmondson
(1930). Of these, three species occur in the
Red Sea and the Gulf of Aden, two species in
the Banda Sea south of the Celebes Islands,
one 'Trom the coasts of America,” one from
Puerto Rico, and the other two species from
the Hawaiian Islands. There are no records in-
dicating the occurrence of any of the species
at other than the type locality.

HAWAIIAN SPECIES OF JOUSSEAUMEA
1. Rostrum long and broad, extending
from basal angles anterior to middle
of eyes to apex beyond end of second
article of antennular peduncle

/. mauiensis (p. 12)

Rostrum short, extending from base be-
tween eyes to or slightly beyond end

of first antennular article

J. hrevirostris (p. 12)

Hawaiian Crangonidae — Banner

11

1 mm

2 mrr

_i0.5mm

Fig. 2. Jousseaumea mauiensis Edmondson, a, b, Anterior region, dorsal and lateral aspects; c, large cheliped,
lateral aspect; d, large chela, ventral aspect; e, small cheliped; /, second legs; g, third legs; h, telson. {a, b, scale A;
c-g, scale B; h, scale C.)

12

PACIFIC SCIENCE, Vol. VII, January, 1953

Jousseaumea mauiensis Edmondson
Fig. 2 a-h

Jousseaumea mauiensis Edmondson, Bernice P.

Bishop Mus., Occas. Papers 9(10): 5-7, fig.

2a-g, 1930.

DESCRIPTION: Rostrum nearly equilaterally
triangular, with base arising anterior to cor-
nea of eye and only slightly shorter than con-
cave lateral margins; tip reaching to middle of
third antennular article. Orbital teeth acute
and about 0.2 to 0.25 as long as rostrum. Ros-
trum with a rounded keel extending back to
its base. Pterygostomial angle rounded.

Antennular peduncle short and heavy, with
third article longer than visible portion of
either first or second article; second article al-
most twice as broad as long. Stylocerite acute,
reaching slightly beyond end of rostrum. Bi-
furcation of outer flagellum occurring near
base; secondary flagellum short.

Antennal peduncle with basicerite bearing
a poorly developed lateral tooth. Scaphocer-
ite slightly longer than antennular peduncle,
with squamous portion exceeding poorly de-
veloped lateral spine in length. Carpocerite
short and heavy, reaching almost to end of
antennular peduncle. Flagellum large in diam-
eter basally, only as long as carapace.

Large chela 0.3 length of body, with basal
articles almost as long as chela. Chela large,
subcylindrical, with palm 1.3 to 1.5 times
length of fingers. Fingers curved and com-
pressed, with faces in opposition armed with
4 large and 2 smaller exactly fitting teeth;
tips curved, acute, and crossing each other.
Merus flattened on side toward palm. Small
chela about 0.25 length of larger chela, not
swollen, and without teeth.

Carpal articles of second legs with ratio
10 : 2.6 : 1.6 : 1.6 : 3.3.

Merus of third legs 3.5 times length of
ischium, 1.3 times length of carpus, 0.75 times
length of propodus, 2.7 times length of dac-
tylus. Merus over 4 times as long as broad,
unarmed; propodus bearing about 8 movable
spines and about 6 shorter tufts of bristles;

dactylus moderately heavy, curved, simple,
with acute tip.

Telson 1.5 times as long as broad at base;
tip 0.4 as broad as base. Terminal emargina-
tion 0.1 total length of telson, rounded and
bearing 3 to 4 pairs of plumose setae. Both
pairs of terminal spines almost as long as
width of tip.

DISCUSSION : (See discussion under J. hrevi-
rostris) .

DISTRIBUTION: The type specimen came
from the island of Maui; additional specimens
have been collected about Oahu at Hanauma
Bay, Kahala Beach, off Waikiki Reef, off
Nanakuli, and at Kawela Bay, at depths from
the reef flat to about 20 feet. Edmondson
(1946: 256) states that this species is commen-
sally associated with a large polychaete of the
genus Eurythoe.

Jousseaumea brevirostris Edmondson
Fig. 3 r-g

Jousseaumea brevirostris Edmondson, Bernice

P. Bishop Mus., Occas. Papers 9(10): 7-9,

figs. 2a-e, 1930.

DESCRIPTION : Carapace with dorsal keel for
over 0.5 its length, continuous anteriorly with
rostral carina. Rostrum triangular, reaching
slightly beyond end of first antennular article.
Eyes normally concealed by orbital hoods in
dorsal and lateral view; orbital hoods contin-
ued as small acute orbital teeth, teeth about
0.3 length of rostrum. Area between base of
orbital teeth and dorsal carina slightly con-
cave.

Antennular peduncle short and heavy, with
visible portion of first and second articles sub-
equal in length, about 1.5 times as long as
broad. Heavy stylocerite with acute tip, reach-
ing almost to end of second antennular arti-
cle. Flagella short, with shorter branch of bi-
furcated upper flagellum not as long as last
2 antennular articles together.

Basicerite heavy, bearing acute lateral tooth
and short spiniform process (in lateral view)
above articulation of scaphocerite. Scaphocer-

Hawaiian Crangonidae — Banner

13

2mm

Fig, 3. Jousseaumea brevirostris Edmondson, a, b, Anterior region, dorsal and lateral aspects; c, d, large chela,
lateral and ventral aspects; e, second legs; /, third legs; g, telson.

14

PACIFIC SCIENCE, Vol. VII, January, 1953

ite short and broad, with weak lateral spine
just reaching end of antennular peduncle,
with squamous portion noticeably longer.
Carpocerite short, heavy, reaching slightly be-
yond end of second antennular article.

Large chela rounded, subcylindrical, length
about 3.3 times breadth at maximum diam-
eter; surface of palm toward merus excavate
permitting flexure of appendage. Fingers over
0.4 length of chela, distally compressed and
somewhat curved; dactylus and fixed finger
with 8 large triangular teeth in opposition,
tips acute and crossing. Merus slightly longer
than palm, slender, rounded on outer side,
flattened on side toward carpus and chela.
Ischium also slender, 0.5 as long as merus.

Small chela lacking.

Carpal articles of second legs with ratio
10 : 3 : 2 : 2 : 4.

Third legs with ischium 0.5 length of me-
rus; merus 4.1 times as long as broad, un-
armed; carpus over 0.6 length of merus, with
superior margin projecting, acute; propodus
0.75 length of merus, with 3 weak spines and
several bristles; dactylus 0.25 length of merus,
somewhat curved, slender, simple.

Telson almost twice as long as broad at
base; tip almost 0.4 breadth of base; anterior
surface strongly convex, posterior surface
more flattened. Terminal cleft of telson about

I. 5 times as deep as wide, only 0.1 of total
length of telson. Inner terminal spines equal
in length to breadth of tip.

DISCUSSION: The description given above
and the figures are those of the type specimen
previously described and figured by Edmond-
son. The slight differences to be noted on a
point by point comparison of the two descrip-
tions are merely differences in interpretation.

The two species of this genus from Hawaii,

J. brevirostris and J. mauiensis^ can best be sep-
arated by two characteristics. First is the na-
ture of the rostrum, which in J. brevirostris is
short, reaching only to near the end of the
first antennular article and correspondingly
narrow at its base (rostrums of both species
approach the dimensions of equilateral trian-

gles), whereas in J. mauiensis it is long, reach-
ing beyond the end of the second antennular
article, and broad at its base. Furthermore, in
J. brevirostris the rostrum and anterior cara-
pace carry a carina that is lacking in J. maui-
ensis. Second is the nature of the terminal cleft
of the telson, which in J. brevirostris is narrow
and deep, about 1.5 times as deep as broad at
the outer edge, whereas in J. mauiensis it is
broad, less than half as deep as broad at the
outer edge. Other slight differences can be
found in the length of the stylocerite, the ratio
of the lengths of the articles of the antennular
peduncles, the merus of the third legs, and
other characteristics.

DISTRIBUTION: The two specimens known
were collected from Waikiki Reef (the type
locality) and from the nearby reef at Kahala,
Oahu.

Alpheopsis Coutibe

Frontal border of carapace produced into
rostrum and projection that screens eyes from
above; orbital teeth present or absent. Cornea
of eyes visible from front and maybe from
sides.

Antennular peduncle short and stout; sty-
locerite variable. Scaphocerite usually broad
without heavy lateral spine. Carpocerite long.

Large chela carried extended, without
grooves or sheaths. Carpus globular, merus
roughly triangular. Chelae show slight asym-
metry; either entire and subcylindrical or with
lines and depressions. Fingers compressed,
either without teeth or with simple arrange-
ment of exactly fitting teeth.

Second thoracic legs with carpus of three
articles (H. idiocarpus Coutiere) or five.

Following legs robust, without teeth on
merus; propodus weakly spinose; dactylus bi-
unguiculate or simple. Propodus of fifth legs
with or without ’’brush” of bristles.

Sixth abdominal segment without articu-
lated pleura only in A. biunguicidatus Banner.
Posterior border of telson rounded.

Branchial formula: 5 pleurobranchs, 1 ar-
throbranch, 8, 7, or 6 epipodites.

Hawaiian Crangonidae — Banner

15

DISCUSSION: The description of this genus
as given by Coutiere (1899: 330) is modified
to accommodate those species described sub-
sequently and further to permit the assign-
ment of a new species of doubtful relation-
ship, A. hlunguiculatus Banner, thereto.

The genus is confined largely to the west
Pacific and the Indian Oceans; aside from A.
chilensis Coutiere, found off South America in
the Pacific, the two species listed below are
the only members of the genus that have been
recorded as far east as the central Pacific
Ocean.

HAWAIIAN SPECIES OF ALPHEOPSIS

1. Dactyli of third to fifth legs simple. . .

A. equalis (p. 15)

Dactyli of third to fifth legs biunguicu-
late (?) -A. hlunguiculatus (p. 18)

Alpheopsis equalis Coutiere

Fig. 4 a-o

Alpheopsis equalis Coutiere, Paris Mus. d’Hist.

Nat., Bui. 2(8): 380, 1896.

Alpheopsis equalis var. truncatus Coutiere, Soc.

Philomath. Paris, Bui. IX, 5(2): 89, 1903.
Alpheopsis equalis and Alpheopsis equalis var.
truncatus Coutiere, Fauna and Geog. Maid,
and Laccad. 2(4): 868, 1905. [A fuller de-
scription and discussion, with figures, of
both forms.]

Alpheopsis aequalis de Man, Siboga Exped.,
39ai (2): 177, 1911.

DESCRIPTION: Frontal border of carapace
produced into acute rostrum extending about
0.7 length of visible portion of first antennu-
lar article. Termination of rostrum, in lateral
view, varying from acute to obliquely trun-
cate. Frontal border projecting beyond cor-
neas of eyes, masking them entirely in dorsal
view and almost or entirely in lateral view;
marked by 2 pairs of slight convexities, one
pair dorsad, one pair laterad of eyes. Ptery-
gostomial angle rounded or produced into
acute tooth.

Antennular peduncle short and heavy with
second article shorter than visible portion of

either first or third. Stylocerite with tip ex-
tending almost to end of second article of
antennular peduncle.

Scaphocerite short and broad, as long as
antennular peduncle, slightly shorter than pe-
duncle of antenna. Lateral spine of scaphoce-
rite not pronounced and equal in length to
squamous portion. Lateral spine of basicerite
acute but not well developed.

Basal articles of third maxilliped with strong
spiniform bristles.

Chelae almost symmetrical in size and
shape, not showing more than 10 per cent
difference in size; rounded and without
grooves. Fingers compressed, with slight, ir-
regularities on opposing faces; usually with
stiff recurving hairs. Carpus elongate with
outstanding flange into which proximal end
of propodus fits, with slight constriction be-
fore this flange. Merus with upstanding collar
on outer distal side into which proximal end
of carpus fits, with rounded edges, with vari-
able set of long stiff bristles on inner lateral
and superior margins. Ischium also with bris-
tles on superior distal face.

Carpal articles of second thoracic leg with
ratio 10 : 6 : 4 : 4 : 5.

Third legs relatively long and slender, with
ischium 0.5 as long as merus; merus 6.5 times
as long as wide; carpus slightly less than 0.5
as long as merus; propodus slightly longer
than merus; dactylus about 0.3 length of pro-
podus. Merus without spines or teeth; carpus
with long slender spine on distal portion of
inferior margin; propodus bearing 4 or 5
spines on inner margin. Appendage glabrous
except several setae on distal end of merus,
distal end of propodus, and middle of dac-
tylus. "Brush” of fifth thoracic legs of 10
rows of weak bristles.

Species small, maximum length about 12
mm.; color in life not noted.

DISCUSSION: In the collections available
from Hawaii there are 13 specimens of this
species, some broken and some entire. These
specimens vary in a series of points, yet plain-
ly all are members of the same species for they

16

PACIFIC SCIENCE, Vol. VII, January, 1953

show essentially the same structure. The
points of variation are as follows:

1. The nature of the tip of the rostrum (Fig.
Ah, c, d) . In his three specimens Coutiere was
able to distinguish between the species A.
equalis, which had a rostrum with a uniform
taper, and the variety, A. equalis var. truncatus,
which had an abruptly truncate rostrum.
Specimens approaching both conditions are
represented in the Hawaiian collections; how-
ever, in those with the pointed rostrum it is a
little less pointed than in those described by
CoutiHe, and in those in which the rostrum is
truncate it is a little less abruptly so than in
his variety. In addition to the two extremes,
there are a series of intergrading individuals
which show various degrees of truncation. A
number of these specimens have a very slight
tooth at the lower side of the terminal trun-
cation. I conclude, therefore, that the variety
described by Coutiere is merely an individual
variation.

2. The presence of a tooth on the pterygo-
stomial angle of the carapace (Fig. 4c, d) . In
most specimens in the Hawaiian collections
the carapace below the base of the antennae is
rounded, as Coutiere shows for A. equalis; in a
few it has a tooth somewhat similar to that
shown for A. equalis var. truncatus. However,
among the specimens possessing the tooth, it
is not always at the same stage of develop-
ment, and in one specimen there is a tooth on
one side but none on the other. Therefore this
too must be considered an individual varia-
tion.

3. The coverage of the eyes by the carapace
(Fig. Ah, d) . In lateral view, the eyes of most
specimens are completely obscured by the
carapace which may project beyond the cor-
neas for a varying distance; in several, how-
ever, the anterior portion of the corneas of the
eyes projects beyond the margin of the cara-
pace.

4. The size and the armature of the chelae
(Fig. Ae-U). In most of the specimens with

both chelae attached, the larger is only about
10 per cent longer than its mate, but in sev-
eral the larger chela was at least twice the size
of the smaller. It is presumed that in these
asymmetrical specimens the smaller chela was
in the process of regeneration. Some of the
specimens, moreover, had their fingers armed
with a dense covering of recurving bristles; in
others the bristles, while present, were neither
dense nor conspicuous. Finally, all of the meri
of the chelipeds had spine-like bristles, but in
some they were conspicuous and strong and
in others they were inconspicuous and weak.
Between the specimens showing the extremes
of variation of the chelipeds were intergrading
individuals.

If these variations were constant, so the
truncated rostrum was always found with a
pterygostomial spine and a spiny merus, etc.,
and if there were no intergrading individuals,
the differences would be sufficient for the erec-
tion of a new species; as it is, the species must
be considered merely as one in which rather
great variation occurs.

DISTRIBUTION: The species has been re-
ported from the Hawaiian Islands by Ed-
mondson (1946). Specimens in the present
collection have been collected on Oahu as fol-
lows: on algal holdfasts in the intertidal zone
near Koko Head and Kawela Bay; in water
from 10 to 20 feet deep at Hanauma Bay,
Waikiki, and Nanakuli. On Maui it was col-
lected at three localities between Lahaina and
Kalama Park in 3-8 feet of water. On Hawaii
it was collected by Hiatt from a tidal pool on
Keaoi Island, Kau Coast.

Three specimens were collected by the "Al-
batross” in Auau Channel, in water from 32
to 43 fathoms deep (U. S. N. M. 63454, 63455,
63456, "Albatross” Stations 3873, 3876, 3872).

Either the species proper or its variety has
been reported from Djibouti and Permin at
the mouth of the Red Sea, from the Maldives
in the Indian Ocean, and from New Caledonia
and Samoa.

Hawaiian Crangonidae — BANNER

17

Fig. 4. Alpheopsis equalis Coutiere. a, b. Anterior region, dorsal and lateral aspects; c, d, variations in anterior
carapace; large chela, lateral and medial aspects; g, h, merus, large cheliped, lateral and medial aspects;
variations in large chela; k, small chela of same specimen as chela t\ /, second leg; m, third leg; n, fifth leg;
0, telson and uropods of specimen shown in c. {a, b, I, m, scale A; c, <7, i-k, scale B; e-h, scale C; n, o, scale D.)

0.5 mm

18

PACIFIC SCIENCE, Vol. VII, January, 1953

(?) Alpheopsis hiunguiculatus sp. nov.

Fig. 5 a-h

TYPE specimen: a male 8.4 mm. long col-
lected from a head of Pocillopora meandrina
Verrill from the intertidal zone at Halape, Kau
Coast, Hawaii, by R. W. Hiatt. This is the
only specimen of this species in the collection
(U. S. N. M. 93451).

DESCRIPTION: Front of carapace projecting
over eyes but abruptly truncate and flattened
dorsally; without keels, ridges, or grooves.
Rostrum, in dorsal view, very small, narrow
and acute, reaching only 0.7 of length of first
article of antennular peduncle; ventral keel
and abrupt truncation in lateral view. Hoods
over eyes slightly inflated, without teeth or
other projections. Eyes extending slightly be-
yond margins of hoods in lateral view. Ptery-
gostomial angle rounded.

Antennular peduncle short and heavy, with
visible portion of first and second article each
0.2 mm. long, with third article 0.25 mm.
long; second article 0.23 mm. wide. Styloce-
rite short, rounded, with short acute spine
arising abruptly from lower margin, tip not
reaching to end of first antennular article. Sec-
ondary bifurcation of outer flagellum well de-
veloped. All flagella broken before tip.

Lateral spine of basicerite acute, short, not
reaching to end of broad portion of styloce-
rite. Scaphocerite broad, with lateral spine
poorly developed, about as long as squamous
portion. Scaphocerite and antennular pedun-
cle equal in length, both considerably longer
than carpocerite. Antennular flagella lost.

Third maxillipeds with basal article (ischio-
merus) 0.68 mm. long, 0.17 mm. broad; car-
pus 0.30 mm. long, 0.12 mm. broad; third
article (propo-dactylus) 0.30 mm. long, O.IO
mm. broad. Appendages with scattered hairs
along basal joints, 8 rows of short stiff bristles
on terminal article, 2 spiniform bristles on end
of first article, and 4 on second article. Exopod
reaching to middle of carpus.

Large chelipeds lost.

Second thoracic legs 3.2 mm. long. Carpal
articles with ratio 10:8:6:6:10. Length of
first secondary article 5.8 times breadth.

Third legs moderately heavy, 3.1 mm. long.
Ischium 0.34 mm. long, with strong movable
spine. Merus 0.95 mm. long, 0.23 mm. broad
(4.1 times as broad as long), with 2 movable
spines. Carpus 0.45 mm. long, 0.16 mm.
broad, unarmed. Propodus 1.04 mm. long,
0.14 mm. broad, with 6 small movable spines,
2 on distal end. Dactylus 0.29 mm. long,
0.09 mm. broad, with strong secondary un-
guis.

Fifth legs similar to third with slight differ-
ences in proportions. Propodus with no sem-
blance of "brush,” but with 4 feeble spines.
Secondary unguis of dactylus poorly devel-
oped.

Basiopodites of first 4 pairs of pereiopods
with rudimentary exopodites(?) (Fig. 5/^).

Pleura of sixth abdominal segment not ar-
ticulated.

Telson 1. 18 mm. long, 0.55 mm. broad at
base (2.1 times as long as broad), with tip
0.25 mm. broad. Lateral margins slightly con-
vex; terminal margin more markedly convex.
Four pairs of long setiferous bristles between
2 terminal spines. Dorsal spines 0.5 and 0.8
mm. from anterior end. Anal tubercles absent.

Branchial formula: 5 pleurobranchs, I ar-
throbranch (rudimentary), 7 epipodites (mas-
tigobranch on seventh leg poorly formed;
setobranch lacking on eighth).

DISCUSSION: Unfortunately, this most per-
plexing species is represented by a single
broken specimen, so any conclusions about it
must be tentative. It was placed in this genus
because in most characteristics it is very much
like A', equalts Coutiere. However, the lack of
articulation of the pleura of the sixth abdom-
inal segment and the presence of exopodites
on the periopods raise serious doubts as to its
relationship to this genus. The second char-
acteristic may not be of any worth, possibly
being merely a persistent larval characteristic
or an atavism. Because the specimen is incom-

Hawaiian Crangonidae — BANNER

19

Fig. 5. Q)Alpheopsis biunguiculatus sp. nov. a, b, Anterior region, dorsal and lateral aspects; c, third maxilliped;
d, second leg; e, third leg; /, fifth leg; g, telson; h, lateral view of the branchial cavity, the two anterior pleuro-
branchs shown only in outline, {a-g, scale A; h, scale B.)

20

PACIFIC SCIENCE, Vol. VII, January, 1953

plete and because of the dubious worth of the
exopodites as a diagnostic character, it has
seemed advisable not to create a new genus
for this specimen. When another specimen,
complete with the large chelipeds, is found it
may be necessary to remove this species from
this genus.

relationship: If A. hiunguiculatus belongs
to this genus, it can be separated from all
other species by the lack of articulated pleura
on the sixth abdominal segment; it can be
separated from all species save A. fissipes Cou-
tiere by the presence of the biunguiculate dac-
tyli on the third to fifth legs, and from A. fis-
sipes (Coutiere, 1908, and more completely,
1921), by the characteristics presented in the
following tabulation:

A. hiunguiculatus
Rostral tip without hairs
Stylocerite shorter than
first antennular article

Carpocerite shorter than
antennular peduncle
Terminal article of third
maxillipeds with bristles

Ratio of carpal articles of
second legs 10 : 8 : 6 :
6 ; 10

Ratio of articles of third
legs: ischium, 0.75; me-
rus, 2.1; carpus, 1.0; pro-
podus, 2.3

A. fissipes

Rostral tip with hairs

Stylocerite almost as long
as second antennular
article

Carpocerite longer than
antennular peduncle

Terminal article of third
maxillipeds without
bristles

Ratio of carpal articles of
second legs 10 : 5 : 5 :
4 ; 6

Ratio of articles of third
legs: ischium, 0.8; me-
rus, 1.4; carpus, 1.0;
propodus, 1.6

Neoalpheopsis gen. nov.

Carapace projecting to form rostrum and
orbital teeth which obscure eyes, in dorsal
view; corneas may or may not extend slightly
beyond edge of carapace in lateral view. No
pterygostomial tooth or even acute angle.

Antennule with heavy peduncle, basal ar-
ticle nearly as long as both distal articles. Sty-
locerite broad, acute tip reaching distal por-
tion of second antennular article. Inner flagel-
lum as long as carapace, outer somewhat
shorter and with setiferous bifurcation on ba-

sal portion.

Antenna with spine of basicerite broad, but
apically acute. Scaphocerite broad, rounded
with lateral spine not longer than squamous

portion. Carpocerite normal. Flagellum al-
most as long as body.

Last article of third maxilliped with trans-
verse rows of short setae on inner side, similar
to that of Crangon.

First pair of chelae well developed, large,
approaching bilateral symmetry, without sex-
ual dimorphism. Chela can be folded back
against merus, merus triangular, somewhat
excavate and twisted to accommodate chela;
merus without spines or teeth. Carpus cyan-
thiform, more elongate than in Crangon. Chela
proper rounded, smooth, tapering, without
crests or lobes, rotated so dactylus rests
against merus when flexed. Cutting surface of
dactylus with serrations that meet exactly
with similar serrations on fixed finger, or de-
veloped as knife-like ridge. Palmar and digital
adhesive plaques poorly developed.

Second legs with carpus of 5 articles, first
secondary article slightly shorter than remain-
ing 4 together.

Third legs with proximal articles with mov-
able spinules; propodus with few weak spines;
dactylus simple. Fifth legs with usual brush
oti inner side of propodus poorly developed.

Abdomen of usual form. Pleura of sixth ab-
dominal segment articulated. Uropods of
usual form. Telson tapering, convex dorsally.
Tip of telson with 2 pairs of spines, medial-
ly to them margin projects markedly as acute
triangle; triangle with 3 or 4 pairs of setiferous
bristles. No anal tubercles.

See Table 1 for branchial formula.

This genus is distinguished from almost
all other genera of crangonids by a series of
characteristics, the more conspicuous of
which are given here.

It is distinguished from Athanas Leach,
Athampsis Coutiere, Arete Stimpson, Areteopsis
de Man, Automate de Man, Pterocaris Heller,
and Ogyris Stimpson by the covering of the
eyes, in dorsal view, by the projecting cara-
pace; from Betaeus Dana and Parabetaeus Cou-
tiere by the presence of a projecting rostrum;
from Synalpheus Bate and Crangon Weber by

Hawaiian Crangonidae — BANNER

21

TABLE 1

Branchial Formula of the Genus Neoalpheopsh

PLEURO-

BRANCHS

ARTHRO-

BRANCHS

EPI-

PODITES*

Maxillipeds

1

1

2

1

3 ”

a

Thoracic legs

1

1

1

a, 13

2

1

a, 13

3

1

a, 13

4

1

at, /3

5

1

* The designations a and jS are used here as they
are applied by Coutiere (1899: 268-286), where a re-
fers to the arm-like mastigobranch, and ^ refers to the
small setiferous lobe-like setobranch.

t This epipodite reduced to a small protuberance.

the lack of the well- developed plunger and
adhesive plaques on the dactylus of the large
chela.

Neoalpheopsis bears a superficial resemblance
to Jousseaumea Coutiere as both the frontal
border of the carapace and the teeth on large
chela are similar and there is a general simi-
larity of form. However, they can be distin-
guished easily by the modification for the
flexure of the large chela, as the merus is
grooved in Neoalpheopsis and the chela is
grooved in Jousseaumea\ by the pleura of the
sixth abdominal segment which are articu-
lated in Neoalpheopsis and not in Jousseaumea;
by the telson, the posterior margin of which
projects as a triangle in Neoalpheopsis but which
is posteriorly cleft in Jousseaumea. The two
can be distinguished further by the branchial
formula which is 5-1-7 in Neoalpheopsis and
5-1-8 in Jousseaumea.

In all ways Neoalpheopsis shows a close rela-
tionship to Alpheopsis Coutiere. The form of
the carapace is similar in the two genera; the
cephalic appendages are similar; the walking
legs of the thorax are similar; the pleura of the
sixth abdominal segment are similar. The che-
lae and the telson differ, however. The chelae
in Neoalpheopsis are carried folded back against

the merus, the fingers are serrate or toothed,
and the chelipeds are almost symmetrical;
whereas in Alpheopsis the chelae are carried ex-
tended, the fingers are usually without serra-
tions, and the chelipeds usually have marked
asymmetry. The telson in Neoalpheopsis pro-
jects terminally to form a large and acute tri-
angle, whereas in Alpheopsis the terminal mar-
gin is either straight or slightly arcuate.

TYPE SPECIES: Neoalpheopsis hiatti Banner.

SPECIES OF NEOALPHEOPSIS

1, Rostrum reaching to near middle of sec-
ond antennular article; merus of third

legs 4.5 times as long as broad

N. hiatti (p. 21)

Rostrum reaching only to distal third of
first antennular article; merus of third

legs 7.5 times as long as broad

N. euryone (p. 25)

Neoalpheopsis hiatti sp. nov.

Fig. 6 a~l

TYPE SPECIMEN: A male 13.5 mm. long
collected from a head of coral in 20 feet of
water, Hanauma Bay, Oahu. (Sizes and places
of collection of paratypes listed below.) (U. S.
N. M. 93452.)

DESCRIPTION: Rostrum broadly triangular
with tip reaching to initial third of second ar-
ticle of antennular peduncle; rostrum slightly
convex dorsally, with ventral keel. Orbital
teeth broadly triangular with rounded tips;
length 0.3 of rostrum. Carapace slightly in-
flated over eyes. No pterygostomial angle.

Eyes concealed in dorsal view; in lateral
view slightly visible ventrad and anterior to
margin of orbital hood.

First antennular article 0.85 mm. long from
base, with visible portion 0.30 mm. long; sec-
ond article 0.30 mm. long; third article 0.40
mm. long; all about 0.35 mm. in diameter.
Stylocerite broadly acute, not spiniform, ex-
tending to end of second antennular article.
Internal crest of first article well developed,
with acute tooth extending to end of article.

22

PACIFIC SCIENCE, VoL VII, January, 1953

Inner flagellum 4.1 mm. long; outer flagellum
3.7 mm. long, with base thickened for 0.30
mm. and produced as secondary lobe 0.35
mm. long.

Lateral margin of basicerite of antennal pe-
duncle with broad, acute tooth extending to
middle of first article of antennular peduncle.
Scaphocerite reaching to end of antennular
peduncle, almost 0.5 as broad as long, with
setae around rounded distal and inner mar-
gins; lateral spine poorly developed, shorter
than squamous portion. Carpocerite not reach-
ing to end of the scaphocerite. Antennal fla-
gellum 8.0 mm. long.

Basal article of endopod of third maxilliped
(ischio-merus) 1.5 mm. long, with basal 0.7
flattened and curved (not shown in aspect
drawn), straight and subcylindrical distally.
Carpus 0.46 mm. long, cylindrical. Third arti-
cle (propodo-dactylus) 0.7 mm. long, circular
in cross section, tapering; with 8 transverse
rows of spinules.

Chelipeds well developed, 7.5 mm. long,
quite symmetrical, usually carried partially or
completely folded back on merus. Ischium
short, delimited from merus by slight groove.
Merus 2.9 mm. long, 0.45 mm. broad; con-
vex-concave distally in cross section, concav-
ity accommodating chela; edges of concavity
produced as low ridges distally, outer ridge
obsolete proximally, inner ridge distinct to
base. Merus without spines or setae, inferior
internal margin produced as small tooth. Car-
pus conical, 0.7 mm. in length and distal
breadth, unarmed. Upon flexion of both car-
pal articulations chela rests against merus.
Propodus 3.6 mm. long, fingers 1.3 mm. Palm
somewhat flattened, curved, without keels or
sharp angles, 1.0 mm. -high. Maximum width
of fixed finger 0.3 mm. ; dactylus slightly thin-
ner (0.25 mm.). Proximal 0.75 of fixed finger
of right chela with 11 large flattened triangu-
lar teeth, larger distally. These alternate with
corresponding smaller teeth on dactylus. Dac-
tylus of left chela with knife-like crest show-
ing minor irregularities homologous to teeth
of right dactylus. Distal 0.25 of fixed finger of

both chelae with ridge, armed with fine serra-
tions, apposable to bare spot on dactylus
opposite; both fingers taper to unarmed,
curved, acute tip that crosses opposite mem-
ber. Palmar and digital adhesive plaques poor-
ly developed.

Second legs 4.3 mm. long. Articles of car-
pus with ratio 10 : 2 : 2 : 2 : 4. Propodus al-
most twice as long as last article of carpus,
fingers 0.5 of length.

Third legs with ischium 0.8 mm. long;
merus, 1.6 mm. long, 0.35 mm. broad; carpus,
0.7 mm. long, 0.2 mm. broad; propodus, 1.3
mm. long, 0.2 mm. broad; dactylus, 0.35 mm.
long, 0.06 mm. broad. Ischium unarmed; me-
rus with strong movable spine on middle of
inferior margin, usually pressed against the
article; carpus with single small movable spine
at articulation of propodus; propodus with 4
spines, 3 proximal less than 0.5 width of pro-
podus, fourth 0.7 width of adjacent propodus.
Dactylus with uniform taper to acute tip,
slightly curved.

Fourth leg similar to third. Fifth leg with
armature of propodus reduced to 2 short
spines, usual "brush” on distal portion of ar-
ticle reduced to 4 small bunches, not rows, of
bristles, each composed of at most 6 or 7 in-
dividual bristles.

Anterior 5 abdominal pleura rourided.
Pleura of sixth abdominal segment acute, tri-
angular, with margins slightly convex, 0.42
mm. long, 0.32 mm. broad at base. Uropods
of normal form, broad, rounded.

Telson convex dorsally, concave ventrally;
sides straight with marked taper. Proportions
as follows: Total length, 1.80 mm.; width at
base, 0.74 mm.; width at tip, 0.32 mm.; thick-
ness at base, 0.42 mm.; width of base of ter-
minal triangle, 0.14 mm.; length (or height)
of terminal triangle, 0.14 mm.; anterior pair of
dorsal spines 0.98 mm. from base; posterior
pair of dorsal spines 1.40 mm. from iDase; an-
terior pair of dorsal spines 0.10 mm. long;
posterior pair of dorsal spines 0.14 mm. long;
lateral terminal spines 0.09 mm. long; medial
pair of terminal spines 0.50 mm. long. Tip of

Hawaiian Crangonidae — BANNER

23

Fig. 6, Neoalpheopsis hiatti gen. et sp. nov. a, h. Anterior region, dorsal and lateral aspects; <r, anterior region
of specimen 4; d, third maxilliped; e, large cheliped; /, merus; g, left chela; h, right chela; i, large chela of speci-
men 3;y, third leg; k, fifth leg; /, telson, uropods, and articulated pleura of the sixth abdominal segment (drawing
foreshortened), {a, b, d-f,j-l, type specimen; c, g, h, specimen 4; i, specimen 3.) {a, b, d-f, scale A; scale

B; g-t, scale C.)

24

PACIFIC SCIENCE, Vol. VII, January, 1953

telson with 3 additional pairs of long setae.
Anus at anterior end of ventral side of telson;
no anal tubercles.

Color variable, with red and brown chroma-
tophores.

discussion: In addition to the type speci-
men, this series is represented in the collec-
tions by four other specimens:

1. An ovigerous female, 27 mm. long, col-
lected olf Waikiki Reef, Oahu, from a coral
head in water about 20 feet deep.

2. A male, 21 mm. long, collected in Hana-
uma Bay, Oahu, from a coral head in about
10-15 feet of water (chelae lacking).

3. An ovigerous female, 20 mm. long, col-
lected offshore at Halape, Kau Coast, Hawaii,
from a head of Porites in 8-9 feet of water by
R. W. Hiatt. The eggs borne by this specimen
were near hatching and contained larvae in the
protozoal stage of development; the eggs
were 0.25 mm. by 0.72 mm. in diameter.

4. A male, 17.2 mm. long, collected at the
same place as specimen 3 by R. W. Hiatt.

The four specimens agree well with the type
in almost all characteristics except for minor
variations in proportions, e.g., the chela varies
from a little over 3 to a little less than 4 times
as long as broad, and the length-breadth rela-
tionship of the telson varies by 10-15 per cent.
These differences are well within the range of
normal and expected variation.

Specimen 4 shows greater differences from
the type than do 1 and 2. In 4 the anterior re-
gion of the carapace, while bearing the same
proportions as that of the type in respect to
the length tf the rostrum, etc., is more closely
pressed to the bases of the antennae; in con-
sequence, the eyes are almost entirely con-
cealed in lateral view, so that at first glance the
rostrum appears to be longer than that of the
type (Fig. 6c). The chelae also show a series
of finer teeth and generally longer and thinner
proportions (Fig. 6g-h).

However, it is specimen 3 that shows the
greatest differences. In this specimen the chela
(only one is present) bears few and conspicu-

ous teeth on both fingers instead of fine teeth
on the fixed finger in opposition to the blade
of the movable finger (Fig. 6i) . Some of the
teeth appear to be blunted or broken.

It may well be that this last specimen repre-
sents another species. However, inasmuch as
all other characteristics are well within the
range of expected variation, it must be as-
sumed, until other specimens are studied, that
this is an extreme variation from the normal.
Another reason for this conclusion is the fact
that this specimen and specimen 4 were col-
lected from the same habitat and are of differ-
ent sexes. It is possible that these two speci-
mens were mates living in the same burrow,
as is common among the crangonids.

The species is named in honor of Robert H.
Hiatt, the collector of some of the specimens
of this species and of other specimens valu-
able to the study.

RELATIONSHIP: There is only one other spe-
cies in this genus, N. euryone (de Man). If the
specimens discussed above all belong to N.
hiatti, then apparent differences between this
species and N. euryone in the chelae, second
legs, etc., are within the range of variation of
the species. However, there are four sets of
characteristics that exceed the range of vari-
ations noted and which will serve for the
separation of the two species. In N. hiatti
the rostrum reaches beyond the end of the
first antennular article; the stylocerite reaches
to the end of the second antennular article;
the merus of the third legs is 4.5 times as
long as broad, 2.3 times as long as the car-
pus, and 4.5 to 5.5 times as long as the dac-
tylus; the telson is 13 times as long as the
terminal triangle. On the other hand, in N.
euryone the rostrum does not reach past the
middle third of the first antennular article; the
stylocerite reaches only to the middle of the
second antennular article; the merus of the
third legs is 7.5 times as long as broad, 1.6
times as long as the carpus, and only 3.0 times
as long as the dactylus; the telson is 10 times
as long as the terminal triangle.

Hawaiian Crangonidae — Banner

25

Neoalpheopsis euryone (de Man)

Fig. 7 a-g

Alpheopsis ? euryone de Man, Ned. Dierk. Ver.,
Tijdschr. (2), 11(4): 308, 1910.

Alpheopsis ? euryone de Man, Siboga Exped.
39aH2): 184, pi. 5, fig. 19, 1911. [This de-
scription is more available, more complete,
and is provided with plates.]

Rostrum triangular, acute, tip extending
over 0.7 of length of visible portion of first
article of antennular peduncle. Rostral base
and rostrum without dorsal keel, not distinct
from gradual curvature of carapace; rostrum
with ventral keel. Orbital teeth strongly re-
duced, hence frontal border of carapace lateral
to rostrum almost straight in dorsal view; apex
of slight teeth rounded. No pterygostomial
atigle.

Eyes concealed in dorsal view, partially vis-
ible in lateral view.

First article of antennular peduncle 0.74
mm. long; second 0.44 mm. long and as broad
as long; the third 0.37 mm. long. Stylocerite
broad, tip acute, reaching more than 0.7 length
of second article. Inner flagellum 8.5 mm.
long; outer 7.3 mm. long, with secondary fla-
gellum reaching over 1 mm. from base of fla-
gellum.

Lateral angle of basicerite produced and
truncate [however, as in the specimen avail-
able, it ends in such abrupt truncation on both
sides that it may have had a strong and acute
tip broken off.] Scaphocerite broad, flat, al-
most as long as antennular peduncle; lateral
spine shorter than rounded portion. Carpoc-
erite of antenna extending only to middle of
last article of antennular peduncle; flagellum
15.3 mm. long, extending to fifth abdominal
segment.

Third maxilliped lacking,

Chelipeds quite symmetrical in develop-
ment, over 11 mm. long from base, capable of
being folded back against merus. Ischium
similar to N. hiatti. Merus 3.4 mm. long, 0.6
mm. broad, with diagonal longitudinal exca-
vation accommodating propodus when ap-

pendage is flexed; inner and outer inferior
distal corners of merus project slightly as
rounded teeth. Chela proper 5.4 mm. long,
1.3 mm. broad (3 times as long as broad), 1.1
mm. thick, oval in cross section. Palm smooth
without ridges or depressions. Fingers 2.0 mm.
long; fixed finger with narrow crest along in-
ner face, with 11 teeth; ridge depressed distad
of teeth, then produced as small crest with
poorly defined teeth; apex curved, strong,
acute. Dactylus similarly with narrow crest,
but knife-like except for small portion in mid-
dle with feeble teeth, bidentate crest opposite
depressed area on fixed finger, knife edge
diminishing to acute, curved tip. Cheliped
glabrous except for several setae near end and
sparse row of short stiff bristles along knife
ridge of dactylus. Palmar and digital adhesive
plaques poorly developed.

Second legs 6.8 mm. in total length. Carpal
articles with ratio 10 : 1.9 : 1.7 : 1.7 : 3.8.

Third legs slender and elongate, 7.8 mm.
long. Ischium 1.5 mm. long, with 2 movable
spines on inferior surface. Merus 2.2 mm.
long, 0.29 mm. broad (7.6 times as long as
broad), with 2 movable spines on inferior
edge. Carpus 1.4 mm. long, without spines or
teeth. Propodus 2.0 mm. long, with 4 mov-
able spines. Dactylus long and slightly curved,
0.7 mm. long, 0.09 mm. broad, with 2 tufts
of bristles on convex surface.

Articulated pleura of sixth abdominal seg-
ment well developed, triangular, with free cor-
ner acute.

Telson 2.95 mm. long, 1.17 mm. broad at
base, 0.48 mm. broad at tip. Anterior portion
strongly convex dorsally, but posteriorly al-
most flat; lateral margins almost straight. Pos-
terior margin produced as large triangle, 0.25
mm. broad at base and 0.29 mm. long, with
straight sides and acute tip. Two pairs of ter-
minal bristles, inner slightly shorter than tri-
angle (0.28 mm. long), outer about 0.3 as long
as inner. Triangle with 4 pairs of setose bris-
tles. First pair of dorsal spinules of telson 0.4
distance from base (1.1 mm.); second pair
about 0.7 of distance from base (1.9 mm.).

26

PACIFIC SCIENCE, VoL VII, January, 1953

DISCUSSION: De Man’s specimen came from
the Netherlands Indies; because it was lacking
in chelipeds, he referred it to the genus Alphe-
opsis "with some doubt."

The specimen here described agrees very
well with de Man’s description and plates ex-
cept in a few small details. In his specimen
the rostrum was shorter, only one third the
length of the visible portion of the first anten-
nular article instead of two thirds. He also
states that the carapace on his specimen had
its upper surface finely punctate, whereas
that on this specimen was smooth. The inner
pair of spines on the tip of the telson are
longer than the projecting triangle in de
Man’s specimen, whereas in this they are
shorter; however, the ones in this specimen
appear to be broken. The lateral tooth of the
basicerite in his specimen is described as
strong and acute, whereas in this it is short
and truncate, but it probably has been broken.
Finally, the merus of the third leg in the type
is 6 times as long as broad instead of 7.5
times.

Most, if not all, of these differences could
easily be individual variation, or they could
be subspecific differences due to geographic
isolation. The only difference that could be of
specific importance is the sculpturing of the
carapace in the original specimen, and this
difference, also, is too slight to warrant spe-
cific separation without any other strong
differences in nonvarying characteristics, es-
pecially as only two specimens of the species
are known.

For a comparison of this species and N.
hiatti Banner see the discussion under the
latter.

DISTRIBUTION: De Man’s species came
from the Karkaralong group, south of Min-
danao (about 5°N, 125°30'E); the specimen
described here is a male 20 mm. long collected
by R. W. Hiatt at Halape, Kau Coast, Ha-
waii, from a head of Fortes in the intertidal
zone, west end of Keaoi Island.

Synalpheus Bate

Synalpheus Bate, Challenger Rpts. 24: 572,

1888.

Synalpheus Coutiere, Les Alpheidae, p. 334,

1899 [redefined].

Anterior portion of carapace with rostrum
and orbital hoods which completely envelop
eyes except on anteroventral side. Anterolat-
eral margin of carapace produced as distinct
pterygostomial angle or tooth.

Antennules with basal articles preponder-
ant, stylocerites large and well formed. Anten-
nular flagellum with weak bifurcation.
Squamous portion of scaphocerite shorter
than lateral spine. Basicerite bearing one or
more teeth.

Large chela carried extended, entire and
smooth, ovaloid, much larger than small
chela; dactylus very short, with cylindrical
process that penetrates into corresponding
cavity on fixed finger. Small chela simple, with
fingers joining exactly, palm entire, carpus
frequently elongate.

Second pereiopods with carpus composed
of 4 or 5 articles. Following legs short and
compressed. In all Hawaiian species merus
unarmed, propodus armed with weak spines,
dactylus biunguiculate. No anal tubercles.

Branchial formula: 5 pleurobranchs, 1 ar-
throbranch, 2 epipodites.

The generic description has been altered to
accommodate Synalpheus redactocarpus Banner
described below, which has only four instead
of the usual five articles in the carpus of the
second legs.

This genus, represented by over 60 species
in the Indo-Pacific and more than 25 from the
coasts of the Americas, is represented in the
Hawaiian Archipelago by only seven species.
As most of the Indo-Pacific forms have been
collected from off-shore waters up to about
100 meters deep but not from the shallow
reef flats, it is possible that more extensive
dredging around the Hawaiian Islands may
produce more species. However, considering

Hawaiian Crangonidae — BANNER

27

mm

2 mm

Z mm

Fig. 7. Neoalpheopsis euryone (de Man), a, b, Anterior region, dorsal and lateral aspects; c, large cheliped; d,
distal end of large chela; e, second leg; /, third leg; g, telson. {a, b, d, g, scale A; c, scale B; e,f, scale C.)

28

PACIFIC SCIENCE, Vol. VII, January, 1953

that the collections made by the "Albatross”
with its extensive dredging and that the col-
lections made by diving have produced only
two new species, it is likely that the present
list comprises almost all the species found in
these waters.

At least one species of this genus, S. para-
neomeris Coutiere, manifests a remarkable vari-
ation, If this condition has been correctly
interpreted, it casts doubts upon many species
of this genus that are separated by relatively
fine distinctions in the proportions of the
various parts. In fact, the perusal of the de-
scriptions of Coutiere in particular (as for
example in his 1909 publication) would make
one wonder about the validity of some of the
described species and varieties, even without
such an example as that of 5. paraneomeris in
mind. Coutiere, too, was doubtful of the va-
lidity of some of the species he described and
suggested that some of them be raised in
"pure cultures” to determine the amount of
variation found in a homogenous group. Cer-
tainly many of the described species have been
established on criteria so fine that almost no
variation could exist without destroying the
validity of the species.

Coutiere (1909) divided the genus into six
groups (not subgenera, but groups which are
without standing under the International
Rules of Zoological Nomenclature). With
only the small group of species studied for
this paper it is impossible to decide how ade-
quate and helpful these groups are. However,
it appears that there are so many exceptions
to the definitions as they were drawn up that
the groups are of scant value. For example, S.
albatrossi Coutiere, which was placed by that
author in the Comatularum Group, violates
the following parts of his definition of the
Group:

"Supraorbital spines insignificant compared
to the rostrum . . .’’—the supraorbital spines
and the rostrum are of equal size.

"... spines of the basicerite almost equal
. . .” — even in S. albatrossi, where the spines
approach equality more closely than do the

spines of most other species of the group, the
spines are not even subequaL

"... first segment of the carpus of the sec-
ond pair of feet very long . . .’’—the first arti-
cle of the carpus is about 1.2 times as long as
the sum of the following, when compared to
the usual 1.0 times as long found in both 5*.
paraneomeris and S. charon and 1.7 times as
long as in S. prolificus, all of which belong to
other groups which are defined as having a
short carpus.

"... following feet cylindrical . . .”—1 do
not know what is meant here, but at least the
articles of the legs of S. albatrossi are as flat-
tened as were the corresponding articles of
other synalpheids examined.

"... ventral hook of the dactyl obsolete
. . .’’—the ventral hook of the dactylus is no
more obsolete in S. albatrossi than is the sup-
posedly well- developed hook of 5. towsendi
brevispinis Coutiere of the Paulsoni Group {op.
cit., p. 35).

"... telson with an oval median lobe ...”
—the median lobe of the telson is no more
oval than any other Hawaiian species of this
genus.

A similar critique could be applied, I be-
lieve, to most of the other characteristics listed
in the key to the groups given by Coutiere.
Even Coutiere did not seem to distinguish
between the groups, let alone the species, for
a specimen from Hawaii (fragmentary, it is
true) in the United States National Museum
which he determined as S. townsendi in the
Paulsoni Group appears to me to be well
within the range of variation found in S. para-
neomeris of the Neomeris Group. In other
words, the groups that were established to
show relationships and to be of aid to the
classification of the genus appear to show
only dubious relationships and are of almost
no aid in classification. As a consequence, in
this paper the groups are not accepted.

If later workers wish to use these groups,
the Hawaiian species may be divided as fol-
lows:

Hawaiian Crangonidae — Banner

29

Comatularum Group
S. alhatrossi Coutiere
S. hiunguiculatus (Stimpson)

Neomeris Group
S. charon (Heller)

5. prolificus (Bate)

S. paraneomeris Coutiere
5. macromanus Edmondson

For the other species described in this paper,
S. redactocarpus Banner, a new group should
be established. With the shifting of S. biungui-
culatus on the basis of its new description
below from the Biunguiculatus Group to the
Comatularum Group, it would be advisable
to change the name of the former group.

t

HAWAIIAN SPECIES OF SYNALPHEUS

1. Carpus of second legs with 4 articles . .
S. redactocarpus (p. 29)

Carpus of second legs with 5 articles . . 2
2. Inferior hook of dactylus of third legs
reduced in size, shorter and smaller in

diameter than superior hook 3

Inferior hook of dactylus of third legs
larger than superior hook in basal di-
ameter and usually longer 4

3. Stylocerite reaching only to end of first
antennular article. . .5. alhatrossi (p. 30)
Stylocerite reaching to end of second

antennular article

5. hiunguiculatus (p. 32)

4. Inferior hook of dactylus of third legs 3

to 5 times diameter of distal hook,

broad and blunt at tip 5

Inferior hook of dactylus of third legs
twice diameter of distal hook, acute
at tip 6

5. First article of carpus of second legs ap-

proximately as long as rest combined

S. charon (p. 37)

First article of carpus of second legs 1.7

times length of rest combined

5. prolificus (p. 38)

6. Palm of large chela tapering only slight-
ly to fingers; fingers equal to 0.3
length of palm. .S. paraneomeris (p. 40)

Palm of large chela inflated, with diam-
eter at fingers 0.5 maximum diameter;
fingers less than 0.25 length of palm
5. macromanus (p. 45)

Synalpheus redactocarpus sp. nov.

Fig. 8 a-i

TYPE SPECIMEN: A male 5.1 mm. long, col-
lected at Halape, Kau Coast, Hawaii, from
coral heads in the intertidal zone on Keaoi
Island by R. W. Hiatt (U. S. N. M. 93454).

DESCRIPTION: Rostrum narrow and acute,
reaching to near middle of second antennular
article; in lateral view tip slightly depressed
but higher than orbital teeth. Orbital hoods
slightly inflated; orbital teeth acute, 0.7 as
long as rostrum.

Antennular peduncle short and heavy with
visible portion of first article subequal to both
second and third. Second article slightly
broader than long. Flagella also short and
heavy, outer slightly longer than peduncle,
inner twice length of outer. Stylocerite reach-
ing nearly to middle of second antennular
article.

Lateral spine of basicerite feebly developed,
reaching only to base of spine of stylocerite;
basicerite otherwise inermous. Scaphocerite
with lateral spine heavy and broad at base, tip
exceeding antennular peduncle and carpocer-
ite; squamous portion broad, about 0.9 length
of lateral spine. Carpocerite as long as anten-
nular peduncle. Antennal flagella lost.

Distal article of third maxilliped rounded
in section, heavily spinose.

Large chela showing compression and some
torsion, without grooves, ridges, or other
sculpture. Chela almost 3 times as long as
broad, with arcuate finger occupying distal
0.23. Carpus short, broad, expanded. Merus
without spines but with superior distal angle
produced into small tooth; article 2.2 times as
long as broad, less than 0.3 length of chela.

Small chela less than 2.5 times as long as
broad with fingers subequal to palm. Fingers

30

PACIFIC SCIENCE, VoL VII, January, 1953

with strong tufts of setae. Carpus as large as
propodus, expanded, on some sides almost
forming collar for base of palm; article twice
as broad as long. Merus without spines, twice
as long as broad, and equal in length to chela.

Second legs very heavy and short, 2.3 mm,
long, merus 0.16 mm. in maximum width.
Ischium 0.48 mm. long, 0.13 mm. broad.
Merus 0.71 mm. long, 0.16 mm. broad. Only
4 secondary carpal articles with ratio 10 : 3.6
: 2.8 : 7.6; fourth secondary article 1.5 times
as long as broad. Chela 0.6 as long as carpus,
slightly broader than carpus; fingers 0.5
length of chela.

Third legs also relatively heavy, 2.1 mm.
long. Ischium 0.33 mm, long, inermous. Me-
rus 0.76 mm. long, 0.22 mm. broad, unarmed.
Carpus 0.29 mm. long, with superior distal
margin produced as rounded tooth. Propodus
slightly curved, 0.77 mm. long, 0.14 mm.
broad, bearing 4 feeble spines on inferior
margin. Dactylus 0.20 mm. long, 0.07 mm.
broad at base, biunguiculate, with superior or
distal claw not as broad at base as inferior,
about 1.5 times as long as inferior; superior
claw more acute than inferior. Fourth legs
similar.

Fifth legs with "brush” feebly developed,
consisting of but 3 tufts of 3 bristles each.
Dactylus with superior unguis thinner than
on preceding legs,

Telson 1.2 times as long as broad at base,
with tip 0.5 as broad as base. Lateral margins
convex distally; distal margin strongly arcuate
and bearing 4 plumose setae. Large pair of
terminal spines approximately 0.5 as long as
breadth of tip of telson. Dorsal surface slight-
ly convex, without any trace of usual 2 pairs
of dorsal spinules.

DISCUSSION: This species, in spite of the
four-jointed carpus of the second legs, plainly
belongs to the genus Synalpheus. The shape
and development of the anterior region of the
carapace, including the pterygostomial angle,
are similar to other species of the genus; ex-
cept for the second legs, the appendages in
their development parallel the development

found in Synalpheus. The important branchial
formula is the same as that of other members
of the genus.

The unique four-jointed carpus will separ-
ate this species from all other members of the
genus, however. Other genera of crangonids
have fewer articles than the usual five, e.g.,
Fterocaris Heller, Ogyris Stimpson, Arete Stimp-
son, and Alpheopsis Coutiere. Moreover, in
Alpheopsis there are species with both three and
five articles to the carpus. However, the other
characteristics of this species do not fit these
genera.

To my knowledge the lack of dorsal spi-
nules on the telson is a characteristic that is
unique among the Pacific synalpheids, al-
though Coutike states (1899: 316) "Quelques
especes [of the genus Synalpheus] . . . montrent
une reduction tres grande des epines, ou
meme leur disparition totale.”

This species does not fit the subgeneric
groups created by Coutiere very satisfactorily.
It apparently belongs to either the Neomeris
or Paulsoni group. From the former it differs
in that the two hooks of the third and fourth
dactyli are subequal; from the latter in that
the rostrum has no ventral prolongation that
"embraces the ocellary beak” (Coutiere 1909:
4) . Should other species with similar develop-
ment of the carpus be described, they should
be included with this species in a special
group; however, the creation of a new group
does not seem practical for this single species.

Of the Hawaiian specimens studied, S. re-
dactocarpus appears to be most closely related
to S. hiunguiculatus (Stimpson), but it differs
from this species in the length of the spines of
the stylocerite and basicerite, in the large and
small chela, and in the dactyli of the third to
fifth legs.

Synalpheus albatrossi CoutiHe
Fig. 9 a-i

Synalpheus albatrossi Coutiere, U. S. Natl. Mus.,

Proc. 36: 89, fig. 54a-k, 1909-

DESCRIPTION: Rostrum acute, narrow, pro-
jecting to 0.3 of second article of antennular

Hawaiian Crangonidae — Banner

31

Fig. 8. Synalpheus redactocarpus sp. nov. a, b. Anterior region, dorsal and lateral aspects; c, large cheliped, lateral
aspect; d, large chela, dactylus flat; e, small cheliped; /, second leg; g, fourth leg; h, fourth leg, dactylus; /, telson.
{a, b, e-g, i, scale A\ c, d, scale B; h, scale C.)

32

peduncles. Orbital teeth acute, elongate, be-
tween 0.4 and 0.5 as long as rostrum. Margin
between orbital teeth and rostrum U-shaped.

Antennule with visible portion of basal ar-
ticle 1.2 times length of second and 1.3 times
length of third. Second article about as long
as broad. Distal ends of articles without con-
spicuous setae. Stylocerite reaching end of
first antennular article.

Basicerite with both upper and lower outer
spines developed and acute, lower consider-
ably longer than upper, reaching beyond base
of spine of stylocerite. Outer spine of scaph-
ocerite acute, reaching to middle or near end
of last article of antennular peduncle; squa-
mous portion relatively broad and well devel-
oped. Carpocerite reaching past base of an-
tennular peduncle.

Large chela only slightly compressed, mar-
gins rounded. Palm twice as long as wide,
with strong tooth above articulation of dac-
tylus. Dactylus 0.3 as long as palm, strongly
arcuate. Carpus of cheliped short. Merus
slightly more than 0.4 as long as chela, twice
as long as broad; superior distal margin ex-
tended as slight rounded projection. Small
chela with fingers 0.5 as long as palm, narrow,
acute; palm slightly over twice as long as
broad. Carpus somewhat smaller than chela;
merus almost as long as chela, 3 times as long
as broad.

Carpal articles of second legs with ratio
10 : 1.3 : 1.3 : 1.5 : 3.0.

Third legs with ischium unarmed, slightly
over 0.3 as long as merus. Merus also un-
armed, 4.3 times as long as broad. Carpus
slightly over 0.5 as long as merus, with tooth
on superior margin and 2 movable spines
on inferior margin. Propodus 1.1 times as
long as merus, with 9 single and double spines
along inferior margin. Dactylus 0.2 as long as
propodus, 3 times as long as wide, inferior
claw much smaller in diameter at base and
much shorter than superior or distal claw.

Telson less than 1.1 times as long as broad
at base, tip 0.3 breadth of base. Tip strongly
arcuate between lateral spines, with 8 setifer-

PACIFIC SCIENCE, Vol. VII, January, 1953

ous bristles; inner pair of lateral spines almost
as long as width of tip. Dorsal spinules lo-
cated near margins, at 0.5 and 0.75 length of
telson posterior to articulation.

Colors in life not observed.

DISCUSSION: The two specimens studied
were both males 8 mm. long. These speci-
mens agree well with Coutiere’s description
and plates except for two characteristics: First,
the rostrum is described as equal in length to
the orbital spines and shorter than the first ar-
ticle of the antennular peduncle, whereas in
these specimens it is longer than both. Sec-
ond, the figures of the type specimen show
the articles of the antennular peduncle to be
relatively more broad than the articles in these
specimens. However, as these characteristics
were found to be very variable in S. paraneo-
merts Coutiere, and as the specimens agree on
other points, they are referred to this species.

For a discussion of the points of difference
between this species and the related S. hiungui-
culatus (Stimpson) see the discussion under
the latter.

DISTRIBUTION: This species has been col-
lected only twice, once by the "Albatross”
off Laysan in 10-18 fathoms (the type local-
ity), and by myself (the two specimens de-
scribed here) from off Waikiki Reef, Oahu,
from coral in 20 feet of water.

Synalpheus biunguiculatus (Stimpson)
Fig. 10 a-j

Alpheus biunguiculatus Stimpson, Acad. Nat.

Sci. Phila., Proc. I860: 31.

(Nec) Alpheus biunguiculatus de Man, Arch. f.

Naturgesch. 53: 502, figs. 6, 6a, 1887.
(Nec) Alpheus biunguiculatus Bate, Challenger
Rpt. 24: 562, pi. 51, figs. 4, 4o, 4z, 1888.
(Nec) Synalpheus biunguiculatus Coutiere, Soc.
Ent. de France, Bui. 11: 232, figs. 1-4,

1898b.

(Nec) Synalpheus biunguiculatus Coutiere, Fau-
na and Geog. Maid, and Laccad. 2(4): 873,
pi. 71, fig. 8, 1905.

Hawaiian Crangonidae — BANNER

).

Fig. 9- Synalpheus albatrossi Coutiere. a, Anterior region, dorsal aspect (note: specimen lacking bilateral sym-
metry in the scaphocerite) ; b, anterior region, lateral aspect; c, large cheliped, lateral aspect; d, large chela, dactylus
flat; e, small chela; /, second leg; g, third leg; h, third leg, dactylus; /, telson. {a, b, i, scale A; c, d, scale B; e-g,
scale C; h, scale D.)

34

PACIFIC SCIENCE, Vol. VII, January, 1953

neotype; a female 10.5 mm. long, col-
lected from coral in about 10 feet of water in
Hanauma Bay, Oahu (U. S. N. M. 93453).

description: Rostrum acute, narrow,
about 1.5 times as long as broad at base, tip
reaching slightly beyond end of first anten-
nular article; lateral margins concave. Orbital
teeth reaching almost to end of rostrum, con-
siderably broader than rostrum at base; with
concave lateral margins. Dorsal surface of
carapace and orbital hoods uniformly rounded;
both rostrum and orbital teeth somewhat de-
pressed toward tip, in lateral view.

Antennular peduncle relatively broad and
heavy, with second and third articles of ap-
proximately equal size, slightly broader than
long; visible portion of first article slightly
longer than others. Outer flagellum with
thickened basal portion about as long as an-
tennular base. Stylocerite strong, lateral spine
reaching end of second antennular article.

Scaphocerite with lateral margin slightly
concave, lateral spine strong, tip reaching be-
yond ends of antennular peduncle and car-
pocerite. Squamous portion well developed,
reaching end of antennular peduncle. Upper
teeth of basicerite entirely lacking, lower
tooth acute but not reaching beyond end of
first antennular article.

Terminal article of third maxillipeds round-
ed, with usual strong spines.

Large chela compressed but with rounded
and entire margins. Palm slightly less than
twice as long as high, 0.7 as thick as high.
Fingers slightly less than 0.3 length of palm;
dactylus strongly arcuate and closing at 90°
to plane of chela. Merus of cheliped 3 times
as long as broad, with small terminal tooth on
superior distal margin. Small chela less than
0.5 as long as large chela, with palm about
1.3 times length of fingers, twice as long as
broad.

Carpal articles of second legs with ratio
10 : 2.0 : 2.0 : 1.7 : 4.2.

Merus of third legs 4.3 times as long as
broad, twice length of ischium; both ischium
and merus unarmed. Carpus slightly over 0.5

as long as merus, with terminal rounded tooth
on superior margin, with terminal movable
spine on inferior margin. Propodus as long as
merus, 7 times as long as broad, with 6 mov-
able spines. Dactylus relatively long and thin
for genus, almost 4 times as long as wide at
point of maximum breadth, not strongly
curved. Lower unguis only slightly divergent
from axis of dactylus, conical, acute, marked-
ly smaller than superior unguis, length only
0.3 and basal diameter 0.7 that of superior
unguis. Superior unguis acute and lying on
axis of basal portion of dactylus; about 0.3 as
long as entire dactylus. Fourth legs similar to
third. Fifth legs similar to but thinner than
anterior legs, with no trace of ''brush.”

Telson about 1.3 times as long as broad at
base, tip 0.5 as broad as base. Posterior mar-
gins with usual 2 pairs of spines, length of
longer over 0.5 width of posterior margin.
Margin strongly arcuate between spines. An-
terior pair of dorsal spinules over 0.3 distance
from articulation, posterior pair slightly over
0.7.

DISCUSSION: The name Alpheus (now Synal-
pheus) hiungukulatus was applied by Stimpson
to a specimen from an unspecified locality in
the Hawaiian Islands. He described it as fol-
lows (I860 — translated from the Latin):

Closely related to Alpheus neptuna, yet having front
short-toothed; spine at the base of the dactylus of large
chela missing; posterior foot small, dactylus two-
clawed, secondary claw ventral. The merus of the third
and fourth legs without spines on the lower side.
Length .5 inch. Habitat; Hawaiian Islands; in coral.

This description was without plates. Evi-
dently Stimpson planned to publish more
than this preliminary account, as he did for
the other groups (the descriptions of the
Brachyura and Anomura of the expedition
were published posthumously in 1907 by the
Smithsonian Institution). However, nothing
further on the Macrura was ever published.

The type specimen is now lost — whether or
not it was destroyed in the fire of the Phila-
delphia Academy of Science could not be
ascertained— so there is nothing left but the
description, which will fit many synalpheids

Hawaiian Crangonidae — Banner

35>

1 mm

2 m rr

0.2

Fig. 10. Synalpheus hiunguiculatus (Stimpson). a, b, Anterior region, dorsal and lateral aspects; c, large cheliped,
lateral aspect; large chela, dactylus flat; e, small cheliped, lateral aspect; /, second leg; g, third leg; h, third leg,
dactylus; /, fifth leg, distal end; telson. {a, b, j, scale A; c-e, scale B; /, g, /, scale C; h, scale D.)

36

PACIFIC SCIENCE, Vol. VII, January, 1953

with equal ease. In 1898 and 1905 Coutiere
redescribed the species on the basis of speci-
mens from the Indian Ocean. Coutiere evi-
dently did not establish a neotype.

Unfortunately the species Coutiere de-
scribes has not been found in the Hawaiian
Islands. This confirms de Man’s doubts about
the validity of redescribing a species from
such a distant locality (1911: 202-203). Cou-
tiUe’s specimens do not, then, meet the one
definite characteristic given in the original de-
scription: "Habitat: Hawaiian Islands."
Moreover, the original description states that
the tooth at the base of the dactylus of the
large chela is missing, but CoutiUe’s plates
clearly show his specimens to have this tooth
or projection. Einally, the statement in the
original description that the dactyli of the
third and fourth legs had their "secondary
claws ventral" would imply that the ventral
claw is not as heavy as the distal or superior
claw, a condition contrary to that found in
Coutite’s specimens. For these reasons Cou-
tiere’s description of Synalpheus hiungukulatus
must be discarded and a neotype selected
from the fauna of the Hawaiian Islands.

The name hiunguiculaUis has priority over
all names of synalpheids from the Hawaiian
area, and Sampson’s description can be inter-
preted to apply to any local species with
greater or less precision. For example, it could
be applied to 5. paraneomeris Coutiere with
ease. However, if it were applied to that spe-
cies, the entire literature would be thrown into
an even greater snarl. To avoid such further
confusion, the name has been applied to this,
the third most common synalpheid in these
waters, a species that has not been named pre-
viously. It agrees with the few characteristics
given in the original description, especially
with the portions discussed above. The only
difference is in the indefinite statement of the
original description: "... having front short-
toothed . . .’’. It would appear that this de-
scription possibly was of a different species,
but one cannot tell what Stimpson meant by
"short." Perhaps, having the rostrum reach-

ing only to the end of the first antennular ar-
ticle was short in Stimpson’s opinion. More-
over, when more specimens are examined,
this characteristic will perhaps be found to be
variable, as it is in 5. paraneomeris.

This species is most closely related to S. al-
batrossi CoutiUe of those known from the Ha-
waiian Islands. They can be separated by the
characteristics shown in the following tabu-
lation:

S. hiungukulatus

Rostral base narrower than
base of orbital teeth

Third antennular article as
long as second antennu-
lar article

Stylocerite reaching to end
of second antennular ar-
ticle

Basicerite without upper
spine, lower spine reach-
ing end of first antennu-
lar article

Scaphocerite longer than
antennular peduncle

Large chela without tooth
over dactylar articulation
or on superior margin of
merus

Dactylus of third legs with
lower unguis almost par-
allel to upper unguis

Telson with posterior mar-
gin strongly convex

5. alhatrossi

Rostral base broader
than base of orbital
teeth

Third antennular article
shorter than second
antennular article
Stylocerite reaching
slightly beyond end of
first antennular article
Basicerite with acute up-
per spine, lower spine
0.5 length of first an-
tennular article
Scaphocerite shorter than
antennular peduncle
Large chela with tooth
over dactylar articula-
tion and small tooth
on distal end of su-
perior margin of merus
Dactylus of third legs
with upper and lower
ungui divergent
Telson with posterior
margin "very little
convex’’ (Coutiere,
1909)

In addition, there are differences in propor-
tions of several of the appendages, especially
in the merus of the large cheliped and of the
small chela. Of these characteristics, probably
the most important are the relative lengths of
the stylocerites and the presence or absence of
the upper spine of the basicerite. It will be
noted that the characteristics of S. alhatrossi in
the tabulation are derived from Coutiere’s

original description, and that the first, second,
and eighth points do not apply well to the
specimens placed in 5. alhatrossi in this paper.

It is necessary to assign a new name to Cou-
tiere’s species. I propose, therefore, Synalpheus
coutierei nom. nov. to replace Synalpheus hiun-

Hawaiian Crangonidae • — BANNER

37

guiculatus (1898^: 232, figs. 1-4), non
S. hiungukulatus (Stimpson) (I860: 31). All
previous records under the names Alpheus, or
Synalpheus, hiungukulatus other than the orig-
inal description should be attributed to S. cou-
tierei, with the possible exception of the record
made without description by Lenz in 1901
wherein he reported the species from Laysan.
However, it is impossible to know exactly
upon which species he was reporting.

DISTRIBUTION: As Stated above, the type
locality was not designated. The neotype
came from the deeper water along the west
side of Hanauma Bay, Oahu; other specimens
have been collected on Oahu off Waikiki in
18 feet of water and off Nanakuli in 15 feet of
water; at Maui off Kalama Park in 8 feet of
water; and by R. W. Hiatt off Hawaii at Ha-
lape, Kau Coast, near Keaoi Island, from Poc-
tllopora meandrina Verrill in 25-30 feet and
40-45 feet of water.

Synalpheus charon (Heller)

Fig. 11 a-k

Alpheus charon Heller, K. Akad. Wiss. Wien,

Sitzungsber. 44: 272, 1861.

DESCRIPTION: Rostrum acute, narrow tri-
angle with straight margins, tip reaching near-
ly to end of first antennular article. Acute tip
of orbital hoods reaching to near tip of ros-
trum; outer margins of orbital hoods convex,
inner margins slightly concave and separated
from base of rostrum by shallow depression.

Antennular peduncle with visible portion
of first article longest, third slightly shorter
than first, second correspondingly shorter
than third; each article about as broad as long.
Stylocerite well developed, with tip of lateral
spine reaching almost to end of second an-
tennular article.

Lateral spine of scaphocerite slightly longer
than squamous portion, approximately as
long as antennular peduncle, and somewhat
shorter than carpocerite. Superior angle of
basicerite rounded, but lateral angle contin-

ued into strong spine, almost equal in length
to stylocerite.

Large chela slightly compressed and twist-
ed, with margins rounded and entire; shape
somewhat inflated, 2.5 times as long as broad.
Superiolateral edge of face of chela behind ar-
ticulation of dactylus projecting over articu-
lation as small rounded tubercle. Dactylus
strongly arcuate, between 0.25 and 0.3 length
of entire chela. Merus with superior distal
margin projecting as acute tooth. Small chela
likewise rounded and entire, dactylus slightly
over 0.3 length of entire chela. Greatest diam-
eter of carpus slightly larger than that of chela
proper. Merus similar in form to large chela.

Carpal articles of second legs with ratio
10 : 3 : 3 : 3 : 5.

Third and fourth legs with merus unarmed
and 3 times as long as broad. Carpus pro-
longed as rounded lobe on superior distal
margin, armed with spine on inferior distal
margin. Propodus curved and armed with 5
or 6 slender spines. Inferior hook of dactylus
thick and obtuse, with broad spoon-like ex-
cavation on inferior margin. Distal hook,
when seen in profile, longer than inferior
hook, less than 0.3 as broad at base as ventral
hook, tapering to acute tooth. All aspects of
distal hook except straight profile showing
basal half as broad and flat, paddle-like, with
sudden constriction to narrow acute tip (Figs.
\\h, i, the last is in such rotation that the
acute tip is somewhat obscured) .

Telson about as long as broad at base, with
tip 0.5 width of base. Lateral margins anteri-
orly concave, then posteriorly convex; tip
arcuate.

The largest specimen in the collection is a
female 22 mm. long; most specimens are
much smaller. The color is often a brilliant
orange red.

DISCUSSION: Only slight variation was
noted in the specimens of this species in the
collections at hand, possibly in part because
of the small number of specimens available.
Only one specimen showed a notable varia-

38

PACIFIC SCIENCE, Voi. VII, January, 1953

don from the specimens described; in this
specimen the merus of the third legs was only
2.5 times as long as broad and the basal por-
tion of the superor unguis was only slightly
broadened. This specimen had all the other
characteristics of the species, including the
excavated ventral hook of the dactylus, so it
was assigned to this species.

These Hawaiian specimens agree well with
Heller’s original description and with de
Man’s redescription of the same species from
the East Indies (de Man, 1911). Heller did
not remark upon the peculiar superior unguis
of the dactylus of the third legs, but it was
described fully by de Man. All differences
noted were minor ones of proportions, such
as the length of the stylocerite, which was
found to be quite variable in the closely re-
lated S. paraneomeris.

This species is very similar to S. paraneo-
meris Coutiere in general body form and pro-
portion, especially when the variation of S,
paraneomeris is considered. However, the na-
ture of the dactyli of the third and fourth legs
affords a rapid and certain method of distin-
guishing the two. To separate this species
from S. prolificus (Bate) see the discussion
under that species. Ramadan (1938) considers
S. helleri de Man (1911: 194, 245) to be prob-
ably a synonym of S. charon, but reaches no
definite conclusion.

DISTRIBUTION: This species appears to oc-
cur almost exclusively in living heads of Pocil-
lopora meandrina Verrill, which it inhabits
with Crangon ventrosa (Milne-Edwards) and
crabs of the genus Trapezia. It therefore could
be expected from all reefs and other areas
where this coral grows. It has been collected
at Mokulua, Waimanalo, Hanauma Bay,
Black Point, Waikiki, Nanakuli, Mokuleia,
and Kawela Bay on Oahu, and at Lahaina on
Maui, in water ranging from a few feet to
about 20 feet deep. As reported by Coutiere
(1909), the ''Albatross” collected the species
at two stations off Laysan in water from 10 to
30 fathoms deep and off the northeast coast

of Maui in 13 fathoms (Stations 3962, 3955,
4073).

S. charon has been reported from many
places between the Red Sea and Hawaii, in-
cluding several localities in the Indian Ocean,
South China and southern Japan, Australia,
and Thursday Island in the Pacific.

Synalpheus prolificus (Bate)

Pig. 12 a-c

Alpheus prolificus Bate, Challenger Rpt. 24:
556-557, pL 99, figs. 4, 4c, 4m, 1888.

As no specimen of this species was avail-
able the original description is given:

Carapace two-thirds the length of the pleon. The
rostrum narrow. The orbital lobes pointed, broad and
nearly equal in length to the rostrum.

First pair of antennae have the second joint of the
peduncle shorter than the first, the third subequal with
the second. Stylocerite a little longer than the first joint,
flagella subequal and about as long as the carapace.

Second pair of antennae as long as the animal, pe-
duncle longer than that of the first pair, basal tooth as
long as the stylocerite and nearly half as long as the
scaphocerite, which is subequal with the peduncle.

First pair of pereiopoda lost. Second slender, having
the carpos five-articulate, the first articlus being longer
than the four following ones. Third and fourth pairs
robust, meros broad, compressed, without distal tooth;
carpos anteriorly produced on the upper distal margin;
propods curved, furnished with spines on the posterior
margin; dactylos stout, bluntly pointed, with a small
secondary unguis on the outer surface (fig. 4m).

The pleopods are broad and foliaceous.

Telson broad, smooth, furnished with two spinules
on the dorsal lateral surface, tapering and rounded at
the extremity.

Length, entire 20 mm. (0.8 in.)

" of carapace 8

Depth of carapace 5.5 "

Length of pleon 12

Habitat~OE Yionolnln, Sandwich Island; depth 18
fathoms. One specimen; female.

Observations — This species is one of those that ap-
proach Alpheus neptunus Dana (PI. Cl, fig. 2) and Al-
pheus biunguiculatus Stimpson. It differs from the former
in having the carpos of the third and fourth pairs of
legs anteriorly produced at the upper distal extremity
and in having the dactylos biunguiculate, and from the
latter (PI, Cl, fig. 4) in being generally more robust, in
having the rostrum and supraorbital teeth less prom-
inent, the carpos of the third pair of pereiopoda pro-
duced on the upper margin, and the dactylos short and
thick, the second unguis being rudimentary and situ-
ated behind the larger on the outer margin.

Our specimen is a female and carries a very large
mass of ova; hence the name.

Hawaiian Crangonidae — BANNER 39

Fig, 11. Synalpheus charon (Heller), a, h, Anterior region, dorsal and lateral aspects; c, large cheliped, lateral
aspect; d, large chela, dactylus flat; e, small chela;/, second leg; g, third leg; h, i,j, dactylus of third leg, anterior,
posterior, and posteroventral sides; k, telson. {a, b, scale c, d, scale B; e-g, scale C; h-j, scale D; k, scale E.)

40

PACIFIC SCIENCE, Vol. VII, January, 1953

DISCUSSION: This species appears to be most
closely related to S. charon (Heller) of all the
Hawaiian species. However, to judge from
Bate’s description and figures, the two spe-
cies can be separated by a series of character-
istics. Probably the most important are in the
carpus of the second leg, which has the first
article 1.7 times as long as the following four
together in S. prolificus (according to the fig-
ure) and only equal to the sum of the follow-
ing articles in S. charon, and in the biunguicu-
late dactylus of the third legs, where the base
of the superior claw is one sixth and the base
of the inferior claw five sixths the diameter of
the dactylus in Bate’s figures instead of one
fourth and three fourths as in 5. charon. Final-
ly, in the antennules the visible portion of the
first antennular article is longer than the
second and third, which are subequal, and the
tip of the stylocerite reaches only slightly be-
yond the end of the first article in S. prolificus,
whereas in S. charon the first and second an-
tennular articles are subequal, the third is
shorter, and the tip of the stylocerite reaches
to the middle of the second antennular article.

Bate makes no mention of, nor shows any
detail of, the dactylus of the third leg. If it
does show the concavity of the inferior ungis
and the basal expansion of the superior unguis
characteristic of S. charon (and such a small
character could easily have been overlooked),
the two species are certainly most closely re-
lated. However, the nature of the second legs
and of the stylocerite would be valid charac-
teristics for the separation of the two species
even if the dactyli were similar.

De Man (1911) suggests that 5. prolificus
may be a synonym of S. hiunguiculatus (Stimp-
son) ; the redescription of S. hiunguiculatus in
this paper removes that possibility.

There are no specimens of S. prolificus in the
present collection; it has not been reported in
the literature since its original description ex-
cept for two specimens recorded by Ortmann
which were later re-examined and placed in
S. gravieri Coutiere (de Man, 1911). This lack
of distributional records indicates that the

Fig. 12. Synalpheus prolificus (Bate), a. Lateral aspect
(slightly over 3X); b, anterior region, dorsal aspect; c,
third leg, propodus and dactylus. (Redrawn from Bate,
Challenger Rpts., Vol. 24; Plates, 1888, pi. 99, fig. 4.)

species is probably restricted to the deeper
zones and does not reach the reefs; it may also
indicate that the species is endemic to the Ha-
waiian Islands, but insufficient sampling of
deeper water in other areas would make such
a conclusion questionable.

Synalpheus pafaneomeris Coutiere
Figs. 13 a-l, 14 a-h

Synalpheus paraneomeris Coutiere, Fauna and
Geog. Maid, and Laccad. 2: 872, 1905.
Synalpheus townsendi Coutiere {partim), U. S.
Natl. Mus., Proc. 36: 35, 1909.

DESCRIPTION: Rostrum narrow and acute,
not quite reaching to end of first antennular
article. Orbital hoods produced into teeth
that reach almost to end of rostrum. Depres-
sion between rostrum and orbital hoods not
extensive, but lateral margins of rostral base
so abrupt that rostrum appears separated from
anterior carapace in dorsal view.

Hawaiian Crangonidae — BANNER

41

Antennular peduncles broad and heavy;
second article slightly broader than long. Sty-
locerite reaching almost to end of second an-
tennular article. Scaphocerite slightly longer
than antennular peduncle, not reaching to end
of carpocerite. Squamous portion of scapho-
cerite rounded anteriorly, comparatively
broad. External tooth of basicerite only slight-
ly shorter than stylocerite; upper angle not
produced into tooth.

Large chela rounded in outline, only slight-
ly compressed, without tubercle at base of
articulation of dactylus. Length of fingers
somewhat more than 0.3 length of palm; chela
more than 3 times as long as broad. Merus
curved, without spines, with superior margin
obtuse distally.

Small chela 0.3 length of large chela. Car-
pus greater in diameter than propodus. Merus
without spines but produced on upper distal
margin as broad dentate process.

Carpal articles of second legs with ratio
10 • 2.5 : 2.5 : 2.5 : 5.5.

Merus of third legs broad, over 3 times as
long as wide, inermous. Carpus with movable
spine on distal inferior margin, with superior
margin produced distally as sharp tooth. Pro-
podus with 6 spines along inferior margin.
Dactylus biunguiculate, with inferior tooth
solid, conical, and with slightly protuberant
tip; distal or superior tooth longer than in-
ferior tooth, 0.5 as broad at base as inferior
tooth.

Length of largest specimens 15 mm.; usual-
ly 10 mm. or less. Color typically olive brown
to gray, but those living in heads of Pocillopora
meandrina reddish.

DISCUSSION: According to Coutiere’s de-
scription, this species manifests a rather wide
variation in the size and shape of the rostral
front, in the antennules and antennae with
their spines, and in the proportions of the
chelae. An examination of almost 100 speci-
mens available showed all of these and other
variations. The more important variations ob-
served were in the following characteristics:

1. The length of the rostrum and of the orbital
teeth. The rostrum itself varied from two to
three times as long as broad, and its tip reached
variously from two thirds the length of the
visible portion of the first antennular article to
beyond the end of the same article. In some
cases the rostral tip was depressed when seen
in side view, with the curve paralleling that of
the orbital teeth; in others the rostrum con-
tinued straight in the dorsal line of the cara-
pace while the orbital teeth alone were de-
pressed anteriorly. In most specimens the or-
bital teeth were of the same length as the ros-
trum; in some they were somewhat shorter,
and in a few they were only two thirds the
length of the rostrum, a condition that was
approached by the specimen Coutiere de-
scribed.

2. The antennules. These were depicted by
Coutiere as being variable in the proportions
of the articles. The local specimens were found
to parallel this described variability. In addi-
tion, the stylocerite also varied in length, but
its tip always reached beyond the end of the
first antennular article.

3. The spine of the basicerite. When the tip of
this spine was compared to the tip of the sty-
locerite, the two were usually found to reach
to the same level, but in a few cases the tooth
of the basicerite was much shorter.

4. The breadth of the squamous portion of the
scaphocerite. As shown by Coutiere, the squa-
mous portion was two thirds the length of the
lateral spine. In these specimens it varied from
a narrow blade two thirds the length of the
lateral spine to a broad squamous portion five
sixths the length of the spine.

5. The large cheliped. The variation in the
proportions of the large chela were like those
noted by Coutiere, but, in addition, the shape
of the superior distal end of the merus was also
found to be variable. In about half the speci-
mens it was rounded, as specified and shown
by Coutiere; in the rest of the specimens this
corner was produced into a somewhat variable
but small, acute, triangular tooth.

42

PACIFIC SCIENCE, Vol. VII, January, 1953

6. The carpal articles of the second legs. These
were described as having the first article slight-
ly longer than the sum of the remaining four;
however, this ratio was found to vary so that
in some the first article was shorter than the
sum of the distal articles (as is the case for the
specimen described above) ; on the other
hand, the longest article measured was 1.3
times the length of the distal articles.

7. The dactylus of the third and fourth legs.
This was almost always as described and de-
picted by Coutiere, but some variation was
noted. Instead of having the lower unguis
markedly larger than the distal unguis, oen
specimen with three normal dactyli had a dac-
tylus on the fourth legs which had the inferior
hook much shorter and only slightly larger
in basal diameter than the distal tooth (Fig.
ll)k). Others showed conditions approaching
this extreme on one or more legs. The dactyli
also varied in the angle between the two
hooks; in most the spread was 45 degrees or
less (when the main axis of the hooks was
projected from the point of meeting) ; in one
specimen the spread was about 60 degrees.

The series of measurements necessary to
study the variation in proportions of the ap-
pendages and telson were not made, but a
simple visual comparison showed these pro-
portions to exhibit a range comparable to that
discussed above.

This variation is a mere extension of the
variability already noted by Coutiere, and
these Hawaiian forms agree rather well with
the characteristics noted by Coutiere, except
on one point. In his description of the dactyli
of the third and fourth legs Coutiere states
that the ventral hooks are, "... courbe, ob-
tuse, suivie d’une concavite du bord inferieur
parfois assez marquee pour indiquer une
ebauche de triunguiculation.” Yet his draw-
ing of the dactylus shows the ventral hook
far from obtuse and gives no indication of the
location of a concavity. The Hawaiian speci-
mens agree with the figure, not the descrip-
tion.

An attempt was made to determine if any
of these variable characteristics would be con-
stant and discrete enough to separate the
population into either distinct species or va-
rieties. However, all the characteristics seemed
to vary within the limits given above. For ex-
ample, although most of the specimens seemed
to be either the form with a long rostrum or
the form with the short depressed rostrum,
some individuals were intermediate between
the extremes. Again, careful measurements
were made on a series of specimens from the
same location to determine if the length of
the squamous portion of the scaphocerite
would be a good criterion for separation; in
the specimens examined the shortest squa-
mous portion was slightly less than two thirds
the length of the lateral spine, the longest was
more than five sixths the length, but many
were intermediate between the two. It was
concluded that neither these nor the other
characteristics discussed above were constant
enough to be the basis of separation of any
subspecific forms.

However, the species has been separated
into five varieties by Coutiere (1905, 1909)
and de Man (1911): S. paraneomeris var. pro-
latus de Man, S. paraneomeris var. halmaherensis
de Man, S. paraneomeris var. praedahundus de
Man, S. paraneomeris var. prasalini Coutiere,
and S. paraneomeris var. seychellensis Coutiere.

There is confusion about the varietal name
S. paraneomeris var. prolatus, which first ap-
pears as a name in Coutiere (1909: 9), where
he states "5. paraneomeris prolatus, new name
( = 5. paraneomeris oxyceros Coutiere)," yet
there are no references in the Zoological Rec-
ord or in Coutiere’s bibliographies to an S.
paraneomeris oxyceros. A possible clue to the
name appears on page 12 of the 1909 work:

S. lockingtoni differs from 5". paulsoni almost solely by
the spine of the scaphocerite being longer in the latter
and surpassing the carpocerite; with the exception of
the place of origin, the second species would corre-
spond to the ’‘'‘oxyceros" form so often met with that
it appears to be almost a constant variation among the
sub-species of a given species. “

[Footnote]^ ... in deference to the accepted rule of
nomenclature which forbids duplication of names with-

Hawaiian Crangonidae — Banner

43

Fig. 13. Synalpheus paraneomeris Coutiere. a, b, Anterior region, dorsal and lateral aspects; c, d, anterior region,
showing variations; e, large cheliped, lateral aspect; /, large chela, dactylus flat; g, small chela; h, second leg; /,
third leg;/, third leg, dactylus; k, third leg, aberrant dactylus; /, telson. {a, d, /, scale A; c, scale B; b, e, /, scale C;
g-i, scale D;/, k, scale E.)

-- 1 mm

'Imm

2mm
'2mm

0.2mm

44

in a single genus, I have in this paper used different
names for oxyceros having a similar meaning, as . . . pro-
latus. . . .

Later, de Man describes and depicts two
specimens similar to what Coutiere meant to
be S. paraneomeris prolatus (or oxyceros) . There-
fore, it appears to me that if the International
Rules were to be applied to these unrecog-
nized names (which are varietal names, not
names of subspecies), Coutiere merely created
a nomen nudum and de Man is the true author
of the variety.

Only a small number of specimens of each
of these varieties was examined: Coutiere’s
original description was based on 14 speci-
mens—one variety upon four specimens and
the other four varieties upon only two speci-
mens each. It is evident that such a small
number of specimens is not an adequate basis
for the separation of varieties. For example, if,
of the 100 specimens examined for this study,
10 or 20 were selected and studied carefully, it
is likely that at least three seemingly separate
and distinct subspecies or even species could
be distinguished; it is only the large number
of specimens that permits the range of varia-
tion to be understood. Consequently in my
opinion the varieties described merely fall in
the line of normal variation of a large popula-
tion and are not of taxonomic worth; for this
reason the previously named varieties have
not been accepted nor have new varieties been
proposed in this paper.

Coutiere (1909) listed S. townsendi as occur-
ring in the Hawaiian Islands; the record was
based on a single specimen taken by the ’'Al-
batross” on French Frigate Shoal. I have ex-
amined the specimen which is in the United
States National Museum. It has none of the
thoracic legs, and these evidently (from Cou-
tiere’s notes in the vial) were lacking when it
was examined by him. Without the important
dactyli and chelipeds it is impossible to identi-
fy the specimen positively. However, in view
of the facts that the anterior body regions and
its appendages and the telson were within the
range of variation for the local S. paraneomeris,

PACIFIC SCIENCE, Vol. VII, January, 1953

Fig. 14, Synalpheus paraneomeris Coutiere. Specimen
from "Albatross” Station 3969, identified by Coutiere
as S. townsendi Coutike. a. Anterior region, dorsal as-
pect; b, telson.

that S. townsendi is known only from the
Americas, and that it has never been reported
from these islands nor is it represented in the
present large collection of synalpheids, the
specimen has been assigned to 5. paraneomeris,

DISTRIBUTION: This species is one of the
most common in the Hawaiian Islands. In
addition to the usual habitat in dead coral
heads, it is found in heads of living Pocillopora
meandrina Verrill. On Oahu it has been col-

Hawaiian Crangonidae — Banner

45

lected from the reef flats and in water to about
20 feet deep at the following localities: Off
Mokulua, Waimanalo, Hanauma Bay, Black
Point, Waikiki, Nanakuli, Kawela Bay, and
Kaaawa. On Maui it has been collected at
Makena and at five places along the coast
north and south of Lahaina. R. W. Hiatt took
some specimens from the intertidal zone at
Halape, Kau Coast, Hawaii. C. H. Edmond-
son (1925) reported the species from Laysan
Island, Lisianski Island, and Pearl and Hermes
Reef. Collections by the "Albatross” reported
by Coutike included specimens from off
southwest Oahu in 18-69 fathoms, off Laysan
in 19-60 fathoms, and the specimen which he
referred to as S. townsendi from French Frigate
Shoal in 15-16 fathoms.

The species and its varieties have been re-
ported from the Red Sea, the Indian Ocean,

and the Netherlands Indies, and in the central
Pacific from Wake and Palmyra Islands.

Synalpheus macromanus Edmondson

Fig. 15 a-g

Synalpheus macromanus Edmondson, Bernice

P. Bishop Mus., BuL 27: 9-11, figs, la-j,

1925.

DESCRIPTION: Rostrum apparently reaching
to near end of first antennular article, orbital
teeth acute, about 0.5 length of rostrum.

Antennular peduncle short and heavy with
second and third articles about 1.7 times as
broad as long. Stylocerite reaching to near end
of second article. Lateral spine of basicerite
strong, almost as long as stylocerite; dorso-
lateral angle rounded. Lateral spine of scapho-
cerite 1.2 times as long as squamous portion;

j 0.5 mm

, 2 mm

m m

Fig. 15. Synalpheus macromanus Edmondson, a, Anterior region, dorsal aspect, left side not shown; b, anten-
nules and antennae, lateral aspect; c, large cheliped, lateral aspect; d, large chela, dactylus flat; e, second leg;
/, third leg; g, third leg, dactylus. {a, b, g, scale A; c, d, scale B; e, f, scale C.)

46

PACIFIC SCIENCE, Vol. VII, January, 1953

tip reaching beyond end of antennular pe-
duncle. Carpocerite dorsoventrally com-
pressed, as long as scaphocerite.

Large chela very large, inflated, almost cir-
cular in section, only twice as long as broad
at maximum diameter, with fingers much
thinner and lower than palm, occupying dis-
tal 0.25 of chela. Carpus of usual form. Merus
0.25 length of chela, twice as long as broad,
with subacute tooth terminating superior dis-
tal margin.

Small chela of usual configuration, with
chela proper compressed and fingers occupy-
ing distal 0.3.

Carpal articles of second legs with ratio
10 : 1.7 : 1.7 : 1.9 : 6.5.

Ischium of third legs 0.3 length of merus;
merus about 3.5 times as long as broad, un-
armed; carpus 0.45 length of merus, superior
distal margin projecting as rounded tooth, in-
ferior distal margin not projecting but with
slender spine; propodus 0.8 length of merus,
somewhat curved, with 5 movable spines on
inferior margin; dactylus 0.25 as long as me-
rus, biunguiculate, conical inferior unguis 1.3
times as broad at base and 0.75 as long as
curved superior or distal unguis.

DISCUSSION: The description and figures are
of the type specimen previously described and
figured by Edmondson. In this specimen, the
only one known, the frontal regions of the
carapace are so distorted as to render accurate
determination of the characteristics impos-
sible.

This species approaches S. paraneomeris
Coutiere in almost all characteristics. It is pos-
sible that the rostrum and orbital hoods are
quite different in the two species, but, as men-
tioned, the exact condition of the frontal re-
gion of the carapace of S. macromanus cannot
be determined. The articles of the antennules
may be broader in this species; the stylocerite
may be longer; the relative sizes of the claws
of the dactylus of the third legs may be differ-
ent. As these characteristics are among the
most variable in what was interpreted to be
S. paraneomeris, I believe the range of variation

of that species would easily encompass these
differences. However, no specimen of S. para-
neomeris has been seen with a large chela ap-
proaching the inflated condition found in that
of S. macromanus, and it appears that this is a
valid criterion for the recognition of two spe-
cies. Of course, it may be found that the huge
chela of S. macromanus is merely a monstrous
variation of 5. paraneomeris, but no specimens
in the collection studied would indicate it; it
may also be that the gross chela is merely a
growth anomaly, for no other specimens have
been collected with a similar chela. At the
present, in any case, the species appears to be
distinct and valid.

DISTRIBUTION: The type specimen, a female
13.5 mm. long, is from Lisianski Island.

Crangon Weber

Crangon Weber, Nomen. Ent. Sec. Ent. Syst.,

1795.

Alpheus Fabricius, Sup. Ent. Syst., 1798.

Carapace developed as orbital hoods which
completely enclose eyes except on ventral
side. Orbital hoods frequently with spiniform
apex and usually demarked from rostral base
by more or less pronounced depression. Ros-
trum often present and carapace usually with
at least slight rostral carina. Pterygostomial
margin of carapace rounded.

Antennules usually short, frequently with
basal peduncular article and stylocerite re-
duced. Scaphocerite frequently reduced; ba-
sicerite either armed with inferolateral spioe
or rounded; carpocerite usually reaching tn
or beyond end of scaphocerite.

Chela of first legs very asymmetrically de-
veloped. Large chela of variable form, from
subconical to compressed and twisted, sur-
faces from smooth and entire to deeply sculp-
tured with grooves and pronounced "alphe-
opsidean” lobes. Dactylus usually with pis-
ton-like process that fits into cavity on fixed
finger and always with digital and palmar ad-
hesive plaques. Chela always carried extended
from body. Large cheliped with hemispherical

Hawaiian Crangonidae — Banner

47

carpus, with merus at least somewhat triangu-
lar in section. Small chela of simple form, at
times showing marked sexual dimorphism.

Carpus of second legs of five articles of
variable proportions.

Third and following legs robust, com-
pressed; armature and proportions of merus,
carpus, and propodus various, dactylus sim-
ple or biunguiculate. Fifth legs with "brush”
on propodus.

Abdomen usually without lateral compres-
sion, with pleura in females larger than those
of males and usually without acute projections
on margins. Pleura of sixth abdominal seg-
ment not jointed. Telson usually with pos-
terior margin convex and lateral angles dis-
tinct but not acute. Anal tubercles well de-
veloped.

Branchial formula: 5 pleurobranchs, 1 ar-
throbranch, 8 epipodites, and sometimes a
supplementary arthrobranch on first thoracic
legs (except in C. paragracilis Coutiere and C.
nanus Banner) .

This genus, previously known to American
systematists and still known to European sys-
tematists as Alpheus, is by far the largest in the
family, not only in Hawaiian waters, but also
in the entire Pacific. In 1911 de Man listed
128 species of Crangon in comparison with 62
species of Synalpheus, the next largest genus.
In this paper 31 species from Hawaii are dis-
cussed; in addition there are two species re-
ported from Hawaii, the records of which are
doubtful.

To facilitate the classification of the genus,
Coutiere (1899) divided it into five groups,
principally on the basis of evolutionary devel-
opment of the large chela and to a lesser
extent on other characters, such as the charac-
ter of the frontal margins of the carapace, the
character of the dactylus of the third legs, etc.
Later he divided the third group, Crinitus, into
three subgroups (1905). These groups as orig-
inally defined were sharply distinguished one
from another and constituted a valuable aid
to the separation of the species.

Many new species were described after the

groups were defined, for example, Coutiere
(1905) described 23 new species from the
Maldives and Laccadives, and de Man (1911)
described some 20 new species from the col-
lections of the Siboga Expedition. Since these
did not exactly conform to the original de-
scriptions of the groups and subgroups, mod-
ifications of the definitions were made.

On the basis of the Hawaiian species it ap-
pears that these groups, unlike those of Syn-
alpheus, probably are still quite valid in deter-
mining the relationships of the species. How-
ever, should some future worker have avail-
able to him a collection of a large number of
species, these groups should certainly be re-
evaluated and some of the species be assigned
either to others of the present groups or to
new groups; by doing this some of the am-
biguity of the definitions of the groups could
be avoided. If that were done it would be de-
sirable to raise the groups to subgenera.

While the groups in general show close re-
lationships of the species within them, the
utility of the groups as redefined by de Man
is definitely questionable. Some species could
fall into two or three groups, as defined, with
equal ease, as the revised definitions often are
equivocal and ambiguous. For example, de
Man’s definition of the Macrochirus Group
(191 1: 307; italics mine):

Lateral (extracorneal) spines usually present. Inferior
spine of basicerite sometimes large. Larger chela usually
longitudinally grooved or emarginate anteriorly. Merus
of third legs usually unarmed, dactyli of three posterior
legs biunguiculate or simple. External spine of exopod of
caudal fan often black.

Because the groups do seem to follow the
natural relationships, they have been used in
this paper. However, because the definitions
of the groups are not clear and definitive, it
has been impossible for the major divisions
of the key to the species to follow the group-
ings. Instead, the key keeps all members of
one group in a single series and in the se-
quence in which they are discussed in the
text. For ready reference the Hawaiian species
are listed here by groups.

48

PACIFIC SCIENCE, Vol. VII, January, 1953

Megacheles Group

1. C. hailstonei (Coutiere) var. paucispi-

nata Banner

2. C. hrachymerus Banner

3. C. albatrossae Banner

4. C. tuthilli Banner

5. C. oahtiensis Banner

6. C. collumiana (Stimpson)

7. C. deuteroptis (Hilgendorf)

8. C. paradentipes (Coutiere)
Macrochirus Group

9. C. gracilis (Heller) var. simplex Banner

10. C. edmondsoni Banner

11. C. latipes Banner

12. C. ventrosa (Milne-Edwards)

13. C. amir ante i (Coutiere)

14. C. nanus Banner

15. C. hawaiiensis

16. C. paragracilis (Coutiere)

Crinita Group, Obesomanus Subgroup:

Not represented.

Crinita Group, Crinita Subgroup

17. C. paralcyone (Coutiere)

18. C. brevipes (Stimpson)

19. C. clypeata (Coutiere)

Crinita Group, Diadema Subgroup*

20. C. paracrinita (Miers)

21. C. paracrinita (Miers) var. bengalensis

(Coutiere)

22. C. gracilipes (Stimpson)

23. C. pugnax (Ddidd)

24. C. diadema (Dana)

25. C. pseudopugnax Banner

26. C. percyi (Coutiere)

Brevirostris Group

27. C. rapax (Eabr.)

28. C. platyunguiculata Banner

* This name has been changed from Insignis to
reflect the placement of C. insignis (Heller, 1861) as a
synonym to C. diadema (Dana, 1852); although the
rules of zoological nomenclature do not apply to this
subgeneric category, it was thought that the change
would reduce confusion.

t Two species of this group, C. audonini (Coutiere)
and C. strenua , which have been reported from

the Hawaiian Islands, are not included in this list nor
in the key because the records are doubtful (see pp. I4l
and 142).

Edwardsi Group!

29. C. leptochirus (Coutiere)

30. C crassimanus (Heller)

31. C. pacifica (Dd-ndi)

With the change of the generic name from
Alpheus, a masculine noun, to Crangon (from
Gr. Kpayyov) a feminine noun, it has been
necessary to change the endings of some of
the specific names.

It was hoped that in the key to the species
it would be possible to avoid the use of all
sexually dimorphic characteristics and the
large and small chelae which are so often lost.
However, it was soon found that such a key
would be difficult to use and would throw the
species out of their normal sequence. Wher-
ever possible the substitution of other char-
acteristics for these was made, especially for
sexually dimorphic characteristics. Einally, ac-
cessory characters are given where it was
thought that they would be useful.

HAWAIIAN SPECIES OF CRANGON

1. Anterior margins of orbital hoods

with acute teeth 2

Anterior margins of orbital hoods
rounded, flattened, or keeled but

never bearing acute teeth 13

2(1). Outer face of large chela with sev-
eral longitudinal grooves and
crests; dactylar articulation of
chela flanked with strong teeth . 3
Large chela rounded, no grooves,

crests, or teeth 10

3(2). Inferior distal margin of merus of
third leg rounded. (Some aber-
rant specimens of some species
in this group lack acute teeth
on the orbital hoods; they can be
recognized easily, however, by
the twisted and crested nature of

the large chela.) 4

Inferior distal margin of the merus
of the third leg produced into an

acute or obtuse tooth 8

4(3). Crest leading to the palmar adhe-
sive plaque on large chela sharp;

Hawaiian Crangonidae — BANNER

49

dactylus of large chela with small
lobes on inner distal face but not

bulbous at tip 5

Crest leading to palmar adhesive
plaque on large chela rounded;
dactylus of large chela with tip

bulbous 6

5 (4) . Second carpal article of second leg

0.5 the length of the first

C. hailstonei Ydit. paucispinata (p. 51)
Second carpal article of second leg
almost 1.5 times the length of
the first. . . . C. hrachymerus (p. 58)
6(4). Dactylus of third leg with one or
more superior serrations but with
no trace of an inferior unguis . . 7
Dactylus of third leg with no supe-
rior serrations but with a strong
inferior unguis . C. oahuensts (p. 64)
7 (6) . Crest leading to the superior tooth
at the dactylar articulation of the
large chela interrupted by a trans-
verse notch . . C. albatrossae (p. 60)
Crest leading to the superior tooth
at the dactylar articulation of
the large chela continuous from
the superior margin of the

palm C. tuthilli (p. 63)

8(3). Tips of the orbital teeth reaching
about half the length of ros-
trum. Dactylus of third leg bi-
unguiculate; inner face of large
chela with a moderate growth of
setae; lateral spine of basicerite

longer than stylocerite

C. collumiana (p. 67)

Tips of the orbital teeth reaching
at least near the end of ros-
trum, or beyond 9

9(8). Tips of orbital teeth not reaching
to tip of rostrum. Large and small
chelae densely hirsute on inner
faces; small chela heavy and with
straight fingers; dactylus of third
leg simple. . . . C. deuteropus (p. 70)
Tips of orbital teeth extending be-
yond rostrum. Inner faces of

large and small chelae almost
glabrous; small chela slender
with fingers bent at right angles
and crossing; dactylus of third

leg biunguiculate

C. paradentipes (p. 72)

10(2). Dactylus of third leg simple,

curved, conical, with acute tip . .11
Dactylus of third leg with tip either
obtuse or strongly hooked. ... 12
11(10). Large chela about 2.5 times as long
as high, with depressions on su-
perior and inferior margins, Me-
rus of third leg 3-4 times as long
as broad; external spine of uro-

pod usually black

C. gracilis var. simplex (p. 75)

Large chela 3. 0-3. 5 times as long
as high, without depressions on
margins. Merus of third leg 6-7
times as long as broad; external
spine of uropod never black . . .

C. edmondsoni (p. 78)

12(10). Dactylus of third leg with abrupt
taper and abrupt hook in distal
0.3; tip acute. Second article of
carpus of second leg broader
than long; mature specimens 8.5

mm. long C. latipes (p. 82)

Dactylus of third leg almost
straight and very blunt. Second
article of carpus of second leg
about twice as long as broad;
mature specimens up to 42 mm.

long C ventrosa (p. 84)

13(1). Large chela with only 1 or 2 simple
longitudinal grooves on outer
face extending proximally from

articulation of dactylus 14

Large chela with or without
grooves, but never with only 1

or 2 longitudinal grooves 17

14(13). Second article of carpus of second
leg over twice length of first.

Large chela subcylindrical

C. amirantei (p. 87)

Second article of carpus of second

50

PACIFIC SCIENCE, VoL VII, January, 1953

leg shorter than first. Large chela

compressed 15

15(14). Scaphocerite reduced, not reaching
to end of antennular peduncle;
merus of third leg unarmed.
Margins of large chela entire . . .

C nanus (p. 90)

Scaphocerite longer than the an-
tennular peduncle; merus of

third leg armed 16

16(15). Depression on lower margin of
large chela with both sides grad-
ual; second carpal article of sec-
ond leg broader than long

C. hawaiiensis (p. 95)

Depression on lower margin of
large chela proximally delimited
by a lobe; second carpal article
of second leg longer than broad . .

C. paragracilis (p. 96)

17(13). Dactylus of third leg biunguicu-

late 18

Dactylus of third leg simple 19

18(17). First carpal article of second leg
about twice as long as second . .

C. hawaiiensis (p. 95)

Second carpal article of second leg
about 3 times as long as first.
Large chela subcylindrical, taper-
ing; margins entire

C. paralcyone (p. 99)

19(17). Large chela without notches,
grooves, or crests, usually quite

cylindrical 20

Large chela with either transverse
groove, proximal to the dactylus,
or pronounced lobes and depres-
sions, especially along upper and
lower margins; chela sometimes

cylindrical 23

20(19). Rostrum very small or absent; dor-
sal Carina of carapace not extend-
ing to tip of rostrum when pres-
ent; orbital hoods with narrow
to broad vertical keel anteriorly;

chela subcylindrical 21

Rostrum reaching about to middle

of visible portion of first anten-
nular article, with interorbital Ca-
rina extending to tip; orbital
hoods evenly rounded anterior-
ly; chela compressed 22

21(20). Second carpal article of second leg
about 1. 5-2.0 times length of
first; without tooth on basicer-
ite; inferior internal margin of
merus of large chela unarmed;
no sexual dimorphism in small

chela C brevipes (p. 103)

Second carpal article of second leg
about equal to first, not over 1.3
times length of first; basicerite
with tooth; tooth distally on in-
ferior internal margin of liierus
of large chela; dactylus of small
chela of female broadened, twice

as long as broad

C clypeata (p. 107)

22(20). First carpal article of second leg
1.75-1.85 times length of sec-
ond; merus of both large and
small chelipeds with strong

tooth C. paracrinita (p. 110)

First carpal article of second leg
about equal to length of second;
tooth on inferior internal margin
of merus of large chela reduced,
none present on small chela . . .

C. paracrinita y 2.1. hengalensis (p. 110)
23(19). Large chela with only transverse
groove proximal to dactylus,
without lobes on either margin;
chela subcylindrical to com-
pressed 24

Large chela with transverse groove
and also marked lobes on both
superior and inferior margins and
depressed areas on faces; chela

always compressed 30

24(23). Dactylus of third leg conical, large

chela subcylindrical 25

Dactylus of third leg triangular in
section, inferior surface broad
and flat, somewhat twisted, large

Hawaiian Crangonidae — Banner

51

chela strongly compressed ..... 29

25(24). Merus of third leg without tooth
on inferior distal margin. All ap-
pendages long and slender; ros-
tral base narrow and flattened
dorsally; merus of large cheliped
with few movable spines ......

............ C gradlipes (p. 115)

Merus of third leg with strong
tooth on inferior distal margin . 26
26(25). Second carpal article of second leg
twice as long as first. Merus of
large cheliped with series of
- spines, subterminal tooth on in-
ferior internal margin; rostral
base very narrow; rostrum not
depressed.. . . . .C. pugnax (p. 116)
Second carpal article of second leg

about as long as first 27

27(26). Rostral base broad, at least 0.7 as
broad as long. Merus of large
cheliped without spines; rostrum

somewhat depressed

............ C diadema (p. 118)

Rostral base narrow, less than 0.5

as broad as long 28

28(27). Anterior portion of base of ros-
trum sharply carinate with slop-
ing margins; margins overhang-
ing orbital-rostral grooves only
in posterior 0.3. Large chela with
depressed areas on superior por-
tion of inner and outer faces;
merus of cheliped with few
spines but numerous serrations.

C pseudopugnax (p. 122)

Anterior portion of base of ros-
trum flattened, with only slight
Carina; margins of rostral base
overhanging orbital-rostral
groove for entire length. Large
chela without depressed areas as
in C. pseudopugnax, merus with
similar serrations . C (p. 124)
29(24). Second carpal article of second leg
shorter than first. Propodus of
third leg bearing only light setae
C rapax (p. 126)

Second carpal article of second leg
over twice as long as first. Pro-
podus of third leg with numer-
ous long setae

.C. platyunguiculata (p. 130)

30(23). Length of dactylus of third legs
about twice breadth of merus.
Large chela with transverse
groove across top, _ connecting
triangular depressed area on each
face; lower margin with strong
lobe; small chela of male prob-
ably "balaeniceps-shaped” ....

C leptochirus (p. 133)

Length of dactylus of third legs
about equal to breadth of merus . 31
31(30). Posterior margin of groove behind
dactylus of large chela sloping
back gradually (best seen in pro-
file), not projecting over floor of
groove; with more or less con-
spicuous tooth distally on inferi-
or internal margin of merus of
large cheliped; small chela "ba-
laeniceps-shaped” in male, slen-
der in female; usually with spine
on ischium of third legs ......

C crassimanus (p. 134)

Posterior margin of groove behind
dactylus of large chela overhang-
ing floor of groove (best seen in
profile) ; no tooth on inferior in-
ternal margin of merus of large
cheliped; small chelae of both
sexes with slender conical fin-
gers; usually without ischial
spine on third legs ...........

C, pad fica (p. 138)

Megacheles Group

Crangon hailstonei (Coutiere) pauci-
spinata var. nov.

Fig. 16 a-j

Alpheus hailstonei Coutiere, Fauna and Geog.
Maid, and Laccad. 2(4): 879, pL 74, fig. 18,
1905.

52

PACIFIC SCIENCE, Vol. VII, January, 1953

TYPE SPECIMEN: A 16 mm. ovigerous fe-
male, "Albatross” Station 4128, off Kauai,
68-90 fathoms, U. S. N. M. 91602.

DESCRIPTION: Rostrum well developed,
reaching almost to end of first antennular ar-
ticle and almost twice as long as broad at
base; top rounded, not carinate. Orbital hoods
somewhat inflated, with acute orbital teeth
reaching beyond middle of rostrum. Orbito-
rostral furrows slight, rounded.

Visible portion of first antennular article
more than 0.7 length of second article; second
article twice length of third article, and 3
times as long as broad at base. Stylocerite
reaching to end of first article. Lateral spine
of basicerite nearly reaching to level of tip of
rostrum. Squamous portion of scaphocerite
narrow and almost 0.9 as long as lateral spine;
tip of lateral spine reaching to end of anten-
nular peduncle. Carpocerite reaching slightly
past middle of third antennular article.

Large chela slightly over 3 times as long as
broad, strongly compressed, with pronounced
crests and grooves. [For terminology of crests
and grooves, see Fig. 17^.] Superior crest
strong, terminating distally as strong acute
tooth above articulation of dactylus. Trans-
verse groove deep, proximally overhanging,
distally rounded. Superior groove only mod-
erately deep. Plaque crest strong, with sharp
ridge, proximally merging with superior mar-
gin of palm. Palmar groove well defined. In-
ferior crest with lower margin better defined
than upper; ending at dactylar articulation as
tooth smaller than that of superior crest. In-
ferior depression shallow but distinct, shoul-
der rounded. Dactylus with high crest, over
0.2 length of entire chela, thin, flat on outer
face, rounded inner face with several distinct
rounded lobes distally, extending over fixed
finger. Carpus of usual form. Merus 3 times
as long as breadth of outer surface and twice
as long as maximum breadth, superior and in-
ferior external margins rounded; inferior in-
ternal margin with strong, acute tooth and
3 small spinules; superior margin rounded
distally. Ischium with single spine.

Total length of small cheliped 0.7 length
of large cheliped, chela about 0.6 length of
larger. Chela almost 5 times as long as broad
with fingers 0.4 length of entire chela. Palm
ovoid, articulation of ' dactylus with strong
teeth on each side. Dactylus narrow, crested.
Carpus rather small. Merus 0.7 as long as
chela, 1.3 times as long as corresponding ar-
ticle on large chela; slender, about 5.5 times
as long as broad; inferior internal margin
armed with 4 small spines and small tooth
distally. Ischium with single inferior spine.

Carpal articles of secorrd leg with ratio 10 :
5 : 3 : 5.5 : 4.

Ischium of third legs about 0.5 as long as
merus, with strong movable spine. Merus 6.5
times as long as broad, unarmed. Carpus 0.6
as long as merus, with acute tooth distally on
superior margin. Propodus 0.9 as long as me-
rus, with 10 strong spines along inferior and
distal margin; last spine bent distally, with
minute serrations on convex portion. Dac-
tylus 0.23 as long as merus, only slightly ta-
pering in proximal 0.7; basal portion de-
marked from distal 0.25 by slight ridge pro-
longed inferiorly as small projecting tooth;
tip tapering, acute.

DISCUSSION: This particular form is repre-
sented in the collections studied by only two
semi-intact specimens collected by the "Al-
batross” in Hawaiian waters. There is a series
of additional individuals, from the same and
other stations, which are so fragmentary that
positive identifications cannot be made.

The other specimen sufficiently intact for
positive identification, a male 19 mm. long,
varies from the type in the following charac-
teristics: rostrum reaches end of first anten-
nular article; large chela 2.9 times as long as
broad; merus of large chela 1.4 times as long
as maximum breadth, with 5 spines, plus 2
on ischium; small chela lacking; carpal arti-
cle with ratio 10: 5:2. 5:6:4; articles of
third leg with ratio 2.5 : 10 : 6 : 9 : 1.9; pro-
podus with 14 spines; dactylus with short
bristles on superior margin. These differences

Hawaiian Crangonidae ■ — Banner

53

I mm
_i2mm

0.5mm

Fig. 16. Crangon hailstonei (Coutiere) n2X. paucispinata var. nov. a, b, Anterior region, dorsal and lateral aspects;
c, large cheliped, lateral aspect; d, large cheliped, medial aspect; e, large chela, distal end, dorsal aspect; /, small
cheliped; g. second leg; h, third leg; /, /, third leg, dactylus. {a, b, scale A; c-h, scale B; i,j, scale C.)

54

PACIFIC SCIENCE, Vol. VII, January, 1953

are minor and to be expected in the normal
range of variation.

The varietal name refers to the few spines
found on the merus of the large cheliped.

relationship: This form and the next four
described are apparently closely related to
each other, to the species C. hailstonei (Cou-
tiere), and to its two tentative varieties de-
scribed by de Man, C. hailstonei assimulans and
C. hailstonei laetahilis. De Man was not sure of
the separation of his forms from the parent
form described by Coutiere, as the differences
between them, although distinct, were rather
minor. Similarly, these five forms differ from
the previously described forms and from each
other on minor but distinct points.

It is likely, therefore, that when sufficient
intact specimens of these and the previous
varieties are available and when the variation
in a large population is studied it will be nec-
essary to consolidate this and de Man’s varie-
ties, and perhaps the following species as well,
into the species described by Coutiere. But,
on the other hand, it may be necessary to
raise this and the previous varieties to specific
rank. However, with only a few specimens
available, it appears to me that the separation
given in this paper is for the present the most
wise, as this form is separated from C. hail-
stonei and the other varieties by only more
minor characteristics, and the other forms, de-
scribed as new species, by greater differences
in more important characteristics.

Table 2 facilitates the separation of these
eight species and varieties. The characteristics
given are not all of equal worth for the sep-
aration of the species. Eor example, little if
any faith can be placed upon the minor vari-
ations found in the relative lengths of the
carpal articles, except in C. hrachymerus, or in
the length of the rostrum, except possibly in
C. tuthilli.

As can be seen from the table, this variety
differs from C. hailstonei hailstonei and from C.
hailstonei laetahilis chiefly in the number of
spines on the margins of the merus of the
large chela, and from C. hailstonei hailstonei and

from C. hailstonei assimulans in the nature of
the dactylus of the third legs. Other differ-
ences noted in the table are of more variable
characteristics and therefore are of lesser
worth.

DISTRIBUTION: One of the positively iden-
tified specimens has its collection data given
above; the other was collected by the "Alba-
tross” at Station 3868 in Pailolo Channel be-
tween Molokai and Maui at 294-684 fathoms
(U. S. N. M. 63535). Other specimens, with
only one or several appendages attached or
with the appendages loose in the bottle so
that identification was more uncertain, were
collected as follows:

"Albatross” Station 3847 (U. S. N. M.
63528) : South coast of Molokai, 23-24
fathoms; 17 specimens, 14 of which are
completely lacking thoracic legs.

"Albatross” Station 3872 (U. S. N. M.
63532): Auau Channel between Maui
and Lanai, 32-43 fathoms; 1 specimen,
lacking large chelae, second legs, and
some of posterior legs.

"Albatross” Station 3875 (U. S. N. M.
63527): Auau Channel between Maui
and Lanai, 65-34 fathoms; 2 specimens.

"Albatross” Station 4128 (U. S. N. M.
63531): Data as given for type; 3 other
fragmentary specimens.

Other specimens from the "Albatross” col-
lection which undoubtedly belong to this
complex and were identified by Coutiere as
C. hailstonei (MS.), but which are too badly
broken to permit me to even guess as to their
identity, were collected as follows:

"Albatross” Station 3809 (U. S. N. M.
63533): South coast of Oahu, 51-125
fathoms; 1 specimen.

"Albatross” Station 3982 (U. S. N. M.
63534): Near Kauai, 40-233 fathoms; 1
specimen.

"Albatross” Station 4055 (U. S. N. M,
63530): Northeast coast of Hawaii, 50-
62 fathoms; 1 specimen.

TABLE 2

Characteristics Differentiating Crangon hailstonei and Some Related Species and Varieties

Hawaiian Crangonidae — BANNER

55

Ui

'O

pq

<

cq

ci

(M

o

d

o

0

I o

►o

^ s

a

Cm

G A

4->

U(

: a

g

o

<u /-»

o

rO

vr\

G ’a

ctj rt

a

1

1

vq

o

rsj

d

T— <

cTo

d

(N

!>0

I (>0

(N

r—i

I

•«<k

tlH ^

1

a

a

m

c<^

■M

to

l-l

"-G

u-l

o

U4

G A
(U u

W 'Ti

o

O

^ rTi

a

■a §

(U

CO

S-l

<u

>

CO

G

cti

u

: 1 : 0.5

4_, S

52 3

2 cA

o .a

w

o

G 2

2 'ul

-G ^ ^

04

04

00

d

r ^

O

r--

G t:

ctS cti

o

0° S S

a

o u

o

rL

H

d

(N

C/0

d

fTi

o

oo

cTo

oi

t— H

d

X ^

P4

Ph

2 G y

W ^

PQ ^

o

O

d

S ‘a

2 G s

O

^ 3

o ^

^ 3

a

c^S

1 — (

o

^ a

a

S.'S

g3

(N

— < M

* ^ M

a

ir\

a

1

43 52 G

G 43 G

’§1

r-i

d

CO

CO

3

s

w

2

o

(u a .y
o ^ t
tll G

to ci ui
O ^

a 52 3

T3 ^

Jii
u
G
G

u G
^ ^

^ G

ll

a

a

Os

1-H

o

H

^ .a

<U w

' M G

<-G G

O (N

(N

I

o

<

<

<

PQ

<

PQ

« -a
•S

a

o

o

o

o

o

o

42 ~5s

a

^ (U

a a

U4

•UJ

Ui

Ul

Ui

A

xr\

o

oi

d

CJ

ctS

-Si ^

rH

.S?-G

rzn

r=J

rz3

o

o

to

CO

r^.

a

a

a

a

a

a

H

'

<N

3

3

3

3

3

3

a

<

d

G u.

^ ,s

a

c\

CO

(U

6

<u

>

ctj

u

G

CO G

a

G C

d

o

\r\

O

r--

lOi

cr £i

H

CN

U

d

3

W

J2 ^

O O

iz; iz;

-S

o ^

G

<D

>

‘Sib

•M

o

:z;

52

^-G 'g

'g

S ii -y

o c t:
O

CO

iS <u
-G >
bo 52

G
g <D
"o § ^

G t:
. c^i
O

G .. A!
u

CO G a

§•§ S.

w

(U

c .*2

rt «-i
<U
U

U( ^

(U

ti ^

2 ^

CO

CO

a 43

2 So

'T3

ci

<u

«-l

43

So

G

<U

(U to

T3 A

c! .a
^ S

G ’tJ
G bQ
C G

<U qj

G

<

u

G

G

T3 OJ

(u .a

U

S o

a 43

G Q.

a ^

° C/O

OJ

u

o

a

S

U

.tG 4^

^ -5

U bO

<u 2
bo^ a

c^ <i

u4

43

.5?

‘a

43

43

So

TABLE 2 — Continued

Characteristics Differentiating Crangon hailstonei and Some Related Species and Varieties

56

PACIFIC SCIENCE, Vol. VII, January, 1953

TABLE 2 — Continued

Characteristics Differentiating Crangon haihtonei and Some Related Species and Varieties

Hawaiian Crangonidae ■ — Banner

57

o

<u O

s'?

O

-•3 S

^ c . 6
o § B oj

S3 .S

CO

o

so

(N

3

3

<0

d

£L>

d

3

d

<u

CO

(U

CO

0

<U

U

43

3

c

c

1

O

T

C3 >

'O
O itn

(U G

*-> (U

3 CO

U _D

C C

<u

P CO

CO

U)

•S

Cj

I

-13

^ P

n3 *->

S S

CQ ^

m

ir\ ^
(N'Vp

f}

O yr\

m
O (N

ck

d

lU

00

<u

d

<u

CO

3

ON

(U

ir\

u

3

CL

d

<

<

(N

d

d

d

lU

”Sb

d

(U

'So

d

w

1.9

d

0

00

d

OJ

CO

43

43

acid
G d

ri:

VO

3

<

00

CO

C/3

<

0

ci

1

3

t ^

S CL

CO

OJ ^

W) W) c

G O -G

<u t! 3

^ d o

o

d

3 u
CL >,

CO

d CO

o 5 -d

U O (U

d

OJ

>

‘Sib

4->

O

iz;

uT <u

'it

(U •>-i
^ CO

CO

•dt

(U

3

-d

4 f

6^

-d

So

d

43

■i-i

too

d

3

u

Q

bO.'-'

(U C4-1

— ' o

"S-S

S|

<u W

CO

=S.2

• in

^ csi

H ^

-d

<7j

<u

>-<

43

-d

So

d

(U

O >,

2 I

PL Q

The data for C. haihtonei haihtonei taken from Coutiere ( 1905: 879—880, figs. 18— 18c); for C. haihtonei assimulans from de Man (I9II: 331—333, pi. 14, figs. 64a— 64e) ;
for C. haihtonei laetabilis from de Man (I9II: 333—334, pi. 14, figs. 64d— 64e) . Where necessary, the individual characteristics or proportions were derived from the figures.
C. brachymems, C. albatrossae, C. tuthilli, and C. oahttensis are all new species described on the following pages.

58

PACIFIC SCIENCE, Vol. VII, January, 1953

"Albatross” Station undesignated (U. S. N.
M. 84243): Hawaiian Islands; 1 speci-
men.

Crangon brachymerus sp. nov.

Fig. 17 a-j

TYPE SPECIMEN: The type and only known
specimen is an 18 mm. male, from "Alba-
tross” Station 3874 (U. S. N. M. 63536) be-
tween Maui and Lanai Islands, 21-28 fathoms.

DESCRIPTION: Frontal region of carapace
abnormal, distorted and strongly asymmetri-
cal, with right orbital tooth rather poorly de-
veloped and left entirely lacking; rostrum dis-
placed and bent sharply ventrad. Second an-
tennular article about 0.1 longer than visible
portion of first, 1.7 times length of third, al-
most 3 times as long as broad. Stylocerite
reaching to end of first antennular article.

Lateral spine of basicerite about 0.8 as long
as stylocerite. Lateral spine of scaphocerite
reaching slightly beyond end of antennular
peduncle; spine moderately strong, squamous
portion narrow. Carpocerite also reaching
slightly beyond end of antennular peduncle,
but slightly shorter than scaphocerite.

Large chela similar to C. hailstonei var. pau-
cispinata Banner (p. 51), compressed, 2.7
times as long as high, with fingers about 0.25
as long as entire chela. [For terminology of
crests and grooves see Fig. 17^.] Superior crest
acute distally, terminating in strong, acute
tooth above dactylar articulation, merging
proximally with superior edge of chela. Trans-
verse groove deep with proximal margin more
or less overhanging, distally rounded. Superior
groove deep. Plaque crest pronounced, with
sharp ridge, proximally merging with superior
margin of palm. Palmar groove shallower and
less well defined than superior groove. Inferi-
or crest arising from middle of palm, termi-
nating distally in tooth below dactylar articu-
lation, smaller than superior tooth. Inferior
depression marked, shoulder strong but
rounded. Dactylus strongly twisted, with high
crest, narrow blade; inner surface of dactylus

with 2 large rounded lobes distally that meet
cutting surface of immovable finger; distal
lobe with several groups of short strong bris-
tles. Carpus small. Merus very short, only 0.6
as long as height of palm, broad, 3.0 times as
long as breadth of outer face, 1.2 times as long
as maximum breadth at distal end, with 3-4
small spines along superior margin and 5
stronger spines on inferior internal margin.
Superior margin of merus distally produced
but rounded; inferior internal margin greatly
produced distally as broad, flat process termi-
nating as strong tooth. Ischium of usual form,
without spines.

Total length of small cheliped 0.8 length
of large cheliped, with chela 0.6 as long as
large chela and merus 1.3 times as long. Chela
0.25 as broad as long, somewhat compressed,
with teeth, crests, and grooves similar to large
chela but much more poorly developed. Su-
perior crest developed but rounded; trans-
verse groove very shallow; plaque crest con-
tinued as rounded ridge across palm; inferior
crest also rounded; superior and inferior crests
terminating distally as small but acute teeth
at dactylar articulation. Fingers 0.8 length of
palm; immovable finger thin and slightly
twisted; dactylus with high, blade-like crest
continuing almost to end, cutting surface
knife-like, whole article twisted into shallow
S-shape. Carpus slightly larger than that of
large chela. Merus 4 times as long as broad,
with superior margin bearing 4 short spines
and scarcely projecting, rounded, distal ter-
mination; inferior internal margin with 7
small spines and simple terminal tooth. In-
ferior surface of ischium with single spine.

Carpal articles of second legs with ratio
10 : 14.5 : 5.5 : 6 : 6. Chela 0.8 length of first
carpal article.

Third legs in specimen broken. Fourth
legs with ischium 0.5 length of merus with
strong movable spine. Merus 9 times as long
as broad, distally unarmed. Carpus almost 0.6
length of merus; superior margin projecting
as short, acute tooth. Propodus 0.7 length of
merus, with 13 moderately strong spines on

Hawaiian Crangonidae — Banner

j 2 m m

2mm
j 2 mm

j 0.5 mm

Fig. 17. Crangon brachymerus sp. nov. a, b. Anterior region, dorsal and lateral aspects; c, large cheliped, lateral
aspect; d, e, large cheliped from "Albatross” Station 3847, medial and lateral aspects;/, small cheliped; g, small
chela, distal end, superior surface; h, second leg; /, third leg; / third leg, dactylus. Abbreviations on e: ic, inferior
crest; id, inferior depression; PC, plaque crest; PG, palmar groove; s, shoulder; sc, superior crest; SG, superior
groove; TG, transverse groove, {a, b, scale A; scale B; d, e, scale C\j, scale D.)

60

PACIFIC SCIENCE, VoL VII, January, 1953

inferior margin, distal 10 paired. Dactylus 0.25
as long as merus, curved, tapering to acute
tip, with very slight accessory tooth about
0.7 distance to tip; small tuft of short thick
spines or bristles opposite biunguiculation.
Parts of third legs present similar to fourth
but larger.

DISCUSSION: A single chela from the "Al-
batross” Station 3847 (U. S. N. M. 63571),
off Laysan Island, 97-199 fathoms, undoubt-
edly belongs to this species. The chela, shown
in Figure 17^, is almost identical to the form
described above. The palmar grooves of the
outer face are deeper and the inner face has
irregular depressed areas, but a closer exam-
ination shows that without question these are
the result of shrinking during a previous des-
iccation. The only difference is the armature
of the superior margin of the merus where the
small spines are so closely pressed against the
rounded edge that only two can be discerned.
Careful examination did not disclose any
more spines; but possibly in a better pre-
served specimen more would be visible.

The specific name refers to the short, broad
merus of the large cheliped.

relationship: The separation of this spe-
cies from the related species and varieties in
the C. hailstonei complex is given in Table 2.
It is unfortunate that the only specimen of
this apparent species available has the frontal
region of the carapace distorted and abnormal
but other characteristics are sufficient to sep-
arate it from the related forms. It may be pre-
sumed, however, as none of the seven other
species and varieties differ markedly in the
nature of the rostrum and the orbital hoods,
that in this species too the rostrum is rather
elongate and the orbital teeth are well devel-
oped, probably quite like the development of
C hailstonei paucispinata Banner (Fig. 16 a).

The decisive differences between this spe-
cies and the variety C hailstonei var. paucispi-
nata are three: First, the merus of the large
chela is very short and broad, markedly short-
er than the palm of the chela is high; second,
the merus of the small cheliped is less than

0.5 the length of the chela; third and prob-
ably most important, the second article of the
carpus of the second leg is 1.5 times as long
as the first instead of only 0.5 as long. In my
opinion, these three pronounced differences
are great enough to separate this form as a
distinct species.

Crangon albatrossae sp. nov.

Fig. 18 a-m

TYPE specimen: An ovigerous female 33
mm. long, "Albatross” Station 4031 (U. S. N.
M. 63588), Penguin Bank, west of Molokai,
in 27 fathoms. Paratypes listed below.

DESCRIPTION: Rostrum 0.7 as long as visi-
ble portion of first article of antennular pe-
duncle, somewhat rounded dorsally, twice as
long as broad at base; upper surface some-
what depressed toward tip. Orbitorostral
grooves broad and shallow. Orbital hoods
slightly inflated, with teeth reaching 0.5 length
of rostrum. Margin of carapace between base
of rostrum and orbital teeth straight but at
angle of about 30° to transverse line of body.

Second antennular article 3 times as long
as broad, 1.5 times length of visible portion
of first article, over twice length of third arti-
cle. Tip of stylocerite reaching to end of first
article.

Lateral spine of basicerite reaching approxi-
mately to end of rostrum. Lateral spine of
scaphocerite strong, lateral margin somewhat
concave; tip reaching slightly beyond end of
antennular peduncle; squamous portion nar-
row, 0.9 length of lateral spine. Carpocerite
slightly longer than scaphocerite.

Large chela heavy, 2.4 times as long as
broad, compressed, 1.7 times as broad as
thick, tapering and distally sculptured. [For
terminology of crests and grooves, see Fig.
17^.] Superior crest rounded, terminating dis-
tally as subacute tooth above dactylar articu-
lation; merging soon with superior margin of
chela proximal to transverse groove. Trans-
verse groove deep and abrupt, proximally
overhanging. Superior groove deep. Plaque

Fig. 18. Crangon albatrossae sp. nov. a, b, Anterior region, dorsal and lateral aspects (one pair of antennular
flagella lost); c, large cheliped, lateral aspect; d, large cheliped, medial aspect; e, large chela, superior face; /, g,
small chelipeds; h, second leg; /, third leg;/, k, /, third leg, dactylus; m, uropods and telson. {a-d, f, i,j, m, from
type specimen; e, g, h, k, /, from specimens collected off southwest Oahu.) {a, b, m, scale A; c-/, scale B; k, /, scale
C; y, scale D.)

62

PACIFIC SCIENCE, Vol. VII, January, 1953

crest strong but rounded, extending to su-
perior margin of chela. Palmar groove short,
well defined on lower side. Inferior crest
strong but rounded, terminating distally in
small, subacute tooth. Inferior depression
deep, shoulder abrupt but rounded. Dactylus
less than 0.3 length of chela, closing almost
across end; with high crest proximally, dis-
tally expanded to appear bulbous. Chela with
sparse, scattered setae distally, especially near
fingers. Merus 2.2 times as long as broad, 1.7
times as long as broad across maximum
breadth, 0.8 height of palm; superior distal
margin produced but rounded, inferior inter-
nal margin with 7 short movable spines and
poorly developed distal tooth. Inferior sur-
face of ischium with 4 spines.

Small cheliped almost equal in total length
to large cheliped, small chela about 0.75 as
long as large chela. Chela 4 times as long as
broad in female, 3.5 times as long as broad in
male, fingers occupying 0.6 of length. Palm
subcylindrical without pronounced sculptur-
ing; articulation of dactylus flanked by acute
teeth on each side; plaque crest slight,
rounded. Fingers slightly broadened with tips
crossing. Carpus with short, broad, rounded
tooth on upper distal margin. Merus 3 times
as long as broad in female, 2.2 times as long
as broad in male, with 9 irregularly placed
movable spines and small distal tooth on in-
ferior internal margin.

Second legs lacking in type specimen. On
paratypes, carpal articles with ratio 10 : 5-6 :
3-4 : 5-6 : 5-6. Chela about 0.8 length of first
carpal article.

Ischium of third legs with strong movable
spine. Merus 5 times as long as broad, un-
armed. Carpus 0.6 length of merus, with su-
perior margin projecting distad as acute tooth.
Propodus almost 0.8 as long as merus, with 11
spines on inferior and distal margins. Dac-
tylus 0.2 as long as merus, about 3 times as
long as broad at base, curved, tip simple, su-
perior surface with 2 notches.

Telson short and broad, length only 1.2
times breadth at base, breadth at tip 0.6 that

of base. Lateral margins slightly convex; pos-
terior margin very slightly arcuate. Postero-
laterally with only one pair of short spines;
larger dorsal spines about 0.55 and 0.8 dis-
tance from base.

Maximum size: 33 mm. (type specimen).

DISCUSSION: Only slight variation was noted
in seven specimens of this species available.
There was the usual and expected variation in
the proportions and lengths of appendages:
in some the stylocerite was slightly shorter
than the first antennular article, and in some
it was as long; the length of the lateral spine
of the basicerite also varied within narrow
limits; the proportions of the large chela it-
self were quite constant, but the merus of the
large cheliped was somewhat more variable in
length, breadth, and armature, with the in-
ferior internal tooth more highly developed
in some than in the type; the small cheliped
appeared to show the slight sexual dimorph-
ism described above; the limits of the ob-
served variation in the carpal articles is given
above; the merus of the third legs ranged to
almost six times as long as broad in some
specimens; the number of spines on the pro-
podus was as high as 11; the dactylus may
have one to three superior serrations, and in
some cases one of these serrations was devel-
oped into a lobe-like extension on the anterior
surface; the telson was considerably more nar-
row and elongate, but still preserving the basic
form.

The species is named in honor of the
steamer "Albatross” of the United States Fish
Commission which collected the type speci-
mens in 1902.

relationship: This species plainly is close-
ly related to C. hailstonei and its varieties;
therefore it has been included in Table 2.
There are a series of minor characteristics that
would aid in the separation of this species
from the varieties of C. hailstonei and C. hrachy-
merus, but the most important ones are found
in the large chela, in which the crest leading
to the plaque is rounded instead of sharp and
the dactylus has a bulbous end instead of a

Hawaiian Crangonidae — Banner

63

narrow lobe-bearing end, and in the dactylus
of the third legs which has marked superior
serrations and no trace of inferior teeth.

This species is also related to C. staphylinus
(Coutite). From the short description and
plates of Coutiere (1921: 418, pi. 62, fig. 13;
note that this later description is exactly the
same as the original description [1908: 14] and
in addition has figures of the specimen) only
small differences appear. The frontal region
of the carapace is straighter and the orbital
teeth are shorter in C. staphylinus\ the large
chela may be different although with the in-
adequate description and the drawing of the
poorly rotated chela, it is difficult to tell; the
small chela is more slender; and, most im-
portant, the dactylus of the third leg is longer
and without the superior notches which are
found in this species.

DISTRIBUTION: In addition to the type loca-
tion given above, the species has been col-
lected as follows:

"Albatross" Station 3873 (U.S.N.M.
63589), in Auau Channel between Maui
and Lanai Islands, 32-37 fathoms; a male
23 mm. long.

Off southwest Oahu, 40-350 feet, collected
by the "Makua” (specimens in the Ber-
nice P. Bishop Museum) ; five specimens,
male and female, from 20 to 29 mm.

long.

Crangon tuthilU sp. nov.

Fig. 19 a-g

TYPE SPECIMEN: A 23 mm. ovigerous fe;
male, collected off southwest Oahu, 40-350
feet deep, by the "Makua." Sole specimen
known. Specimen in the Bernice P. Bishop
Museum.

DESCRIPTION : Rostrum small, reaching only
to middle of visible portion of first antennular
article, as broad as long, upper surface strong-
ly depressed toward tip. Anterior carapace

__i 2. mm
_j 4 mm
j 0. 5 m m

Fig. 19. Crangon tuthilli sp. nov, a, b. Anterior region, dorsal and lateral aspects; c, large cheliped, lateral aspect;
d, large chela, superior face; e, second leg, distal end; /, third leg; g, third leg, dactylus. {a, b, scale A; c-f, scale
B; g, scale C.)

64

smoothly rounded, without trace of orbito-
rostral grooves. Orbital teeth short and nar-
row, reaching 0.7 length of rostrum.

Second antennular article 1.3 times as long
as visible portion of first, 1.5 times as long as
third, twice as long as broad. Stylocerite
reaching end of first antennular article.

Lateral spine of basicerite strong, longer
than rostrum, extending over 0.7 of length of
first antennular article. Lateral margin of sca-
phocerite almost straight, tip of lateral spine
reaching end of antennular peduncle; squa-
mous portion only 0.05 shorter than spine.
Carpocerite considerably longer than anten-
nular peduncle.

Large chela only moderately heavy, 2.8
times as long as broad, compressed and with
distal sculpturing reduced. [For terminology
of crests and grooves see Fig. lie.] Superior
crest low, rounded, terminating as relatively
small acute tooth above dactylar articulation.
Transverse groove lacking. Superior groove
shallow and broad. Plaque crest low, rounded,
but merging with superior margin of palm.
Palmar groove rounded, narrower than su-
perior groove. Inferior ridge slight, rounded,
distally ending as rounded protuberance be-
low dactylar articulation. Inferior depression
slight, shoulder low, gradual, and rounded.
Dactylus similar to that of C. albatrossae Ban-
ner, with proximal portion narrow and crested,
distal portion expanded and bulbous. Carpus
of usual form. Merus equal in length to height
of palm, 2.7 times as long as breadth of outer
face, 1.6 times as long as maximum breadth;
superior margin rounded, unarmed; superior
distal angle slightly projecting but rounded;
inferior internal margin with 8 small spines
and large terminal tooth. Ischium with 2
spines on superior margin, 5 on inferior
margin.

Small cheliped lost.

Carpal articles of second legs with ratio
10 : 6 : 4 : 4 : 6.

Third legs with ischium 0.4 length of me-
rus, armed with strong spine. Merus 7 times
as long as broad, unarmed. Carpus 0.6 as long

PACIFIC SCIENCE, Vol. VII, January, 1953

as merus, with acute distal projection of su-
perior margin. Propodus 0.8 as long as merus,
with 8 inferior and distal spines. Dactylus 0.25
as long as merus, curved, acute, with 2 su-
perior serrations, no trace of inferior teeth.

DISCUSSION: This single specimen came
from the same dredge haul that produced the
paratypes of C. albatrossae Banner and super-
ficially is very similar to them. The basis for
separation of the species from the other spe-
cies of the C. hailstonei complex is given in
Table 2. Probably the most valid and least
variable characteristics that will serve for the
identification of this species are the nature of
the anterior region of the carapace, the ros-
trum being much shorter and broader than
that of the other species, and the lack of the
transverse groove on the large chela.

Unfortunately, with only one specimen
available, it cannot be determined whether
this specimen represents a heretofore uncol-
lected species or merely an aberrant individ-
ual, probably of C. albatrossae. Minor genetic
mutations could account for the differences
between this and the typical C. albatrossae.
The fact that the specimen was collected with
a series of specimens of C. albatrossae would
incline one toward that belief, but of course
in a half-hour dredge haul there is no reason
to suppose that all the specimens came from
exactly the same habitat. However, the char-
acteristics mentioned above and some of the
more minor characteristics, such as the thick-
ness of the second antennular article, appear
to be rather constant within a species of this
group. For these reasons it was decided to
designate this specimen a species.

The species is named in honor of Dr. Leon-
ard D. Tuthill, Department of Zoology and
Entomology, University of Hawaii.

Crangon oahuensis sp. nov.

Fig. 20 a-m

TYPE SPECIMEN: A male 27 mm. long col-
lected from coral in water about 20 feet deep
off Waikiki Reef, Oahu (U.S.N.M. 93455).

Hawaiian Crangonidae — Banner

65

4mm

2mm

0.5mm

Fig. 20. Crangon oahuensis sp. nov. a, b, Anterior region, dorsal and lateral aspects; c, anterior carapace, dorsal
aspect, showing asymmetrical development of orbital hoods; d, large cheliped, lateral aspect; e, large chela,
superior face; /, large cheliped, merus, internal face; g, small cheliped, lateral aspect; h, small chela, superior face;
i, small chela, section of dactylus showing ridge; y, second leg; k, third leg; /, third leg, dactylus; m, telson.
{a-c, m, scale A; d-h, j, k, scale B; /, /, scale C.)

66

PACIFIC SCIENCE, Vol. VII, January, 1953

DESCRIPTION: Rostrum almost 1.5 times as
long as broad, with tip reaching 0.7 of length
of first antennular article; rostrum and rostral
base slightly rounded dorsally, not carinate.
Orbital hoods not inflated. Orbital teeth
reaching about 0.7 of length of rostrum. Or-
bitorostral furrows slight, shallow, and broad;
anterior margin between base of rostrum and
base of orbital teeth very slightly convex, but
at angle of about 30° to transverse line of
body.

Second article of antennular peduncle twice
as long as broad, 1.4 times length of first ar-
ticle, 1.5 times length of third article, Stlyo-
cerite reaching slightly beyond end of first
antennular article. Lateral spine of basicerite
acute, tip reaching to end of first antennular
article. [In the type specimen, the spines are
developed asymmetrically, with that of one
side as described and that of the other
rounded anteriorly, shorter than the rostrum.]
Lateral margin of scaphocerite only slightly
concave; spine 0.8 longer than squamous por-
tion; squamous portion reaching end of an-
tennular peduncle. Carpocerite equal in length
to spine of scaphocerite.

Large chela heavy, 2.5 times as long as high,
compressed, 1.6 times as high as thick, dis-
tally sculptured. [For terminology of crests
and grooves, see Fig. 17^“.] Superior crest well
developed, terminating distally as strong acute
tooth above dactylar articulation, proximally
merging with superior margin. Transverse
groove deep, abrupt. Superior groove deep
and well defined. Plaque crest rounded but
heavy, merging proximally with superior mar-
gin of palm. Palmar groove with upper mar-
gin poorly defined, lower margin abrupt. In-
ferior crest rounded, strong, terminating as
strong, acute tooth below dactylar articula-
tion. Inferior depression shallow; shoulder
low, gradual and rounded. Dactylus, occupy-
ing distal 0.25 of chela, distorted, hammer-
shaped, end bulbous, middle thin, crested,
and twisted; outer surface of dactylus with 3
grooves .armed with short stiff setae. Chela
with numerous short to medium bristles on

distal 0.5 of inner face. Merus twice as long
as broad, with superior margin terminated by
short obtuse tooth; inferior internal margin
with 9 small movable spines and strong sub-
terminal tooth. Ischium with rounded pro-
jection on superior margin, inferior margin
with 5 spinules similar to those on merus.

Small chela in both sexes of peculiar form,
with palm slightly over twice as long as broad,
but with dactylus long, slender, and curved,

1.4 times length of palm, when seen in lateral
view (as in Fig. 20g) over 11 times as long as
broad in middle; fingers curved in both planes,
tips Cfoss slightly. Superior side of opposing
face of fingers with thin ridge except where
tips curve and cross; ridge with close series of
fine rugae (Fig. 20/) ; on interior side of ridge
opposing faces with orderly series of short
stiff setae. Articulation of dactylus flanked on
each side by strong acute teeth. Merus slightly
over twice as long as broad, slightly heavier
than that of large chela, with similar armature
except spinules more numerous and subter-
minal tooth smaller. Small chela 0.8 as long
as large chela.

Carpal articles of second legs with ratio
10 : 9 : 4 : 4 : 6.

Third legs with ischium unarmed. Merus

3.4 times as long as broad, unarmed. Carpus
0.5 length of merus, superior distal angle pro-
jecting but rounded; inferior margin with 2
nearly spine-like bristles in type specimen,
with distal angle rounded, not projecting.
Propodus 0.75 as long as merus, with 10 long
slender spines on inferior and inferodistal
margins. Dactylus 0.2 as long as merus, curved,
with small secondary unguis on inferior
margin.

Telson 1.5 times as long as broad, base 1.7
times as broad as tip; lateral margins some-
what convex; distal margin arcuate.

DISCUSSION: The 12 specimens of this spe-
cies at hand show some variation, especially
those of smaller si2e when compared to the
large specimen designated as the type. The
general form and appearance was the same in
all cases, but in some of the smaller specimens

Hawaiian Crangonidae — BANNER

67

the articles of the antennular peduncle were
shorter and broader (second article ranges to
1.7 times as long as broad); the stylocerite
was shorter than the first antennular article;
the relative lengths of the antennular pedun-
cle, the scaphocerite, and the carpocerite were
not constant; the number of spines on the
merus of the large and small chelae varied, but
in all cases were numerous; the second carpal
article was at times 0.7 or 0.8 the length of the
first, with the other articles varying to a simi-
lar degree; the merus of the third legs was
more slender, up to 4.0 times as long as broad;
and finally, the number of spines on the pro-
podus of the third legs usually was less than
the 10 spines of the type specimen, often
around six or seven.

The most remarkable variation was ob-
served in the orbital teeth of three specimens.
Two specimens, one shown in Figure 20c, had
the orbital teeth asymmetrically developed,
with a tooth of the usual type on one side and
the other side terminating in an obtuse non-
projecting angle. The third specimen was
lacking in the orbital teeth on both sides.

This species has been named after the island
from which it was collected.

relationship: This species apparently be-
longs to the group of species designated in
this paper as the C. hailstonei complex, and for
that reason its characteristics have been set
forth in Table 2. It can easily be separated
from all other members of this complex and
from C. seurati (Coutiere) (which probably
also belongs in the complex) by the extraor-
dinary elongation and great thinness of the
fingers of the small chela and by the strong
biunguiculation of the dactylus of the third
legs. Other characteristics which will separate
the species from one or several of the group
are the size of the rostrum, the heaviness of
the antennular peduncle, the rounded condi-
tion of the palmar crest and the bulbous dac-
tylus of the large chela, the relatively longer
second carpal article of the second legs, the
breadth of the merus of the third legs, and the
lack of the ischial spine on these legs.

DISTRIBUTION: This species is represented
by 13 specimens which were collected on
Oahu at Hanauma Bay, Kahala, Waikiki, and
Nanakuli (all except the Kahala specimen
from water 10 to 20 feet deep) and on Hawaii
at Halape on the Kau Coast, one specimen
from 10 feet, one from 40 feet deep, the last
two collected by R. W. Hiatt.

Crangon collumiana (Stimpson)

Fig. 21 a-h

Alpheus collumianus Stimpson, Acad. Nat. Sci.

Phila., Proc. 12: 30, I860.

DESCRIPTION: Rostrum acute, short, reach-
ing to middle of visible portion of first anten-
nular articles, demarked laterally from orbital
hoods by abrupt depressions, with small ros-
tral Carina that extends posteriorly between
orbital hoods. Orbital hoods rounded with
anteriolateral margins produced into acute
teeth as long as rostrum. Margin between
base of rostrum and orbital teeth convex.

Lateral spine of stylocerite acute, reaching
beyond rostrum nearly to end of first anten-
nular article. Second article of antennular
peduncle usually 1.8 times length of visible
portion of first article, 1.7 times length of
third article. Lateral spine of basicerite long,
acute, reaching beyond end of first antennular
article. Scaphocerite with strong lateral spine
that exceeds squamous portion by about 0.25,
usually reaching end of antennular peduncle.
Tip of carpocerite extending beyond end of
scaphocerite.

Large chela 2.3 times as long as broad, lat-
erally compressed and distally sculptured. [For
terminology of crests and grooves, see Fig.
17^.] Superior crest heavy, rounded, terminat-
ing distally as strong subacute tooth above
dactylar articulation; merging with superior
margin of palm almost immediately proximal
to transverse groove. Transverse groove deep,
abrupt, and merging with superior groove.
Superior groove deep, broad, but terminating
immediately proximad of transverse groove.
Plaque crest rounded, heavy, and well defined

68

PACIFIC SCIENCE, VoL VII, January, 1953

but not merging with superior margin of
palm. Palmar groove narrow and deep, reach-
ing only about 0.3 length of palm. Inferior
crest' heavy, rounded, terminating distally as
strong, acute tooth below dactylar articula-
tion. Inferior depression moderately devel-
oped; shoulder heavy but rounded. Dactylus
short, about 0.25 length of entire chela, ro-
tated to close almost distally across end of
chela. Chela moderately setose, with numer-
ous groups of setae on inner face, superior
and inferior margins, arid about edge of fin-
gers. Merus less than twice as long as broad,
without spines, with superior and inferior in-
ternal margins slightly produced and rounded.

Small chela somewhat heavy. Palm 1.5
times as long as wide, distally with superior
crest well developed and ending as projecting
tooth which flanks articulation of dactylus.
Fingers about equal in length to palm, dis-
tally strongly hooked. Carpus with flat tooth
on superior distal margin. Merus over twice
as long as wide, without spines.

Carpal articles of second legs with ratio
10 : 9 : 4 : 4 : 7.

Ischium of third legs with movable spine.
Merus 3.3 times as long as wide, armed with
strong but rounded tooth on inferior distal
margin. Carpus with superior margin ending
distally in acute dentate projection. Propodus
with 5-6 sets of strong movable spines along
inferior margin. Dactylus biunguiculate, with
inferior tooth small and located in distal 0.25
of article.

Largest specimens in collection 20 mm.
long. In life body mostly transparent with red
and yellow chromatophores, legs with yellow
and green chromatophores; superior surface
of chelae olive drab.

discussion: This species does not seem to
exhibit the wide range of variation found in
some. However, the following variations were
noted:

1. The length of the second antennular ar-
ticle varied from 1.5 to 2.0 times the length of
the visible portion of the first article; in most
specimens about 1.7 or 1.8 times as long.

2. In some specimens the scaphocerite was
equal in length or slightly shorter than the
antennular peduncle; in others it was slightly
longer than the carpocerite. The carpocerite,
usually markedly longer than the antennular
peduncle, in a few cases was equal to it.

3. In most specimens the small chela was
markedly smaller than the large chela, as in
the Qne drawn; in a few of the larger ones it
was almost 0.8 the length of the large chela
and similar in form to the small chela of C.
deuteropus (Hilgendorf) (Eig. 22^).

4. The second carpal article of the second
legs was usually about 0.9 the length of the
first but varied from 0.8 to 1.0 times as long.

5. The merus of the third legs in a few
specimens carried one or two weak spines,
and the inferior tooth, usually acute with an
angle of 60 or 70 degrees, was in some cases
obtuse with an angle of more than 100 de-
grees.

The Hawaiian form of this species differs
from Stimpson’s original description chiefly
in the nature of the third and fourth legs,
which he describes as, "Pedes tertii quartique
. . . mero lato, inferne spinuloso et apicem
unidentato. ..." In the Hawaiian forms, the
few specimens with feeble spinules on the
merus would hardly be called spinose. The
specimen described by de Man (1911: 334)
had one or two small movable spines on the
carpus of the third legs which are lacking in
the Hawaiian specimens. Finally, in de Man’s
plates (pi. 14, figs. 65, 65a) the superior ridge
of the large chela merges with the surface of
the palm without any abrupt groove demark-
ing it as in the Hawaiian form. These two
characteristics of de Man’s specimen were not
remarked upon in the original description.
These differences are minor and probably in-
significant.

The three specimens listed below, which
were collected by the "Albatross,” had been
examined but not published upon by Cou-
tiere. The Laysan specimen ("Albatross" Sta-
tion 3962) was given a manuscript name for
a new species. The specimen had recently

Hawaiian Crangonidae — BANNER

69

Fig. 21. Crangon collumiana (Stimpson). a, b, Anterior region, dorsal and lateral aspects; c, large chela, lateral
aspect; d, large chela, medial aspect; e, small chela; /, second leg; g, third leg; h, third leg, dactylus. i^a-g,
scale A; scale B.)

70

PACIFIC SCIENCE, VoL VII, January, 1953

molted, and its chitin was soft and distorted;
however, close examination showed only one
difference from the specimens described
above: there were two spines on the carpus of
the third legs (therefore more like the speci-
men described by de Man). The other two
specimens, which Coutiere had identified as
C. dentipes (Guerin)— a species never reported
in the Indo-Pacific— showed no characteris-
tics that would distinguish them from C. col-
lumiana as identified in this paper.

DISTRIBUTION: This rather common species
has been collected from the reef flats and from
waters ranging to 20 feet deep at the following
localities around Oahu: Waimanalo, Hana-
uma Bay, Black Point, Waikiki, Mokuleia,
Haleiwa, Kawela Bay. It was dredged from
100-200 feet off Diamond Head and from 180
feet off Kaena Point, where it was found liv-
ing in sponges. Off Maui it was collected at
Makena, Kalama Park, Lahaina, and two lo-
calities south of Lahaina. The "Albatross”
specimens came from off Laysan, 16 fathoms
(Station 3962, U.S.N.M. 63596), off French
Frigate Shoal in 1414-1 61/2 fathoms (Station
3968, U.S.N.M. 63509), and off Nihoa, 24-40
fathoms (Station 4163, U.S.N.M. 63510). Ed-
mondson (1925) has reported the species from
Pearl and Hermes Reef, Laysan, Lisianski, and
Ocean Islands.

Edmondson (1925) has also reported the
species from Johnston, Wake, and Palmyra
Islands in the central Pacific. The species has
been otherwise reported from Djibouti, at the
mouth of the Red Sea, through the Indian
Ocean and western Pacific to Japan. The type
locality is the Bonin Islands.

Crangon deuteropus (Hilgendorf)

Fig. 22 a-g

Alpheus deuteropus YUlgenAod, K. Akad. Wiss.

Berlin, Monats. 1878: 834.

DESCRIPTION: Rostrum acute, curved
abruptly upward near tip; tip reaching about
to middle of visible portion of first antennular
article, posteriorly continued as distinct ros-

tral Carina; base separated from orbital hoods
by depression. Orbital hoods rounded except
for acute orbital teeth which equal rostrum in
length. Margin between orbital teeth and ros-
trum straight to slightly arcuate.

Antennular peduncle long and slender,
with second article slightly less than 1.5 times
as long as visible portion of first, about twice
as long as third. Lateral spine of stylocerite
from 0.7 as long to as long as first antennular
article. Entire antennular peduncle densely
hirsute. Lateral spine of scaphocerite strong,
1.25 times as long as narrow squamous por-
tion; outer margin strongly concave. Lateral
spine of basicerite strong, extending beyond
end of rostrum. Carpocerite hirsute. Scapho-
cerite usually not reaching end of last anten-
nular article; carpocerite almost always longer
than both antennular peduncle and scaphocer-
ite. [The specimen drawn in Figure 22a is ab-
normal in these relationships.]

Chelae very densely hirsute on inner face
and superior portion of outer face; structure
obscured by setae. Hirsute surfaces papillose
where bundles of setae are attached. Large
chela strongly compressed, ovate in shape,
about twice as long as high. Fingers approxi-
mately 0.2 as long as entire chela. [For termi-
nology of crests and grooves, see Fig. 17^.]
Superior crest strong, rounded, distally pro-
jecting as long, subacute tooth, proximally
ending in transverse groove. Transverse
groove deep. Superior groove deep but short.
Plaque crest heavy, soon merging with face of
palm. Inferior crest heavy, well defined but
not extending proximad to shoulder of in-
ferior depression, distally ending in strong,
acute tooth. Inferior depression shallow;
shoulder slight and rounded. Short immov-
able finger emarginate at tip to receive dac-
tylus. Dactylus rotated to close laterally and
distally across end of chela, compressed and
sharply carinate. Carpus relatively large. Me-
rus heavy, without teeth, with superior distal
angle subacute.

Small chela large, 0.8 as long as large chela,
ovate, densely hirsute on inner face. Fingers

Hawaiian Crangonidae — Banner 7 1

Fig. 22. Crangon deuteropus (Hilgendorf). a, b. Anterior region, dorsal and lateral aspects; c, large cheliped,
lateral aspect, setae not shown; d, large cheliped, medial aspect, only bases of setae shown, length indicated by
dotted line; e, small cheliped, medial aspect, setae not shown; /, second leg; g, third leg.

72

PACIFIC SCIENCE, VoL VII, January, 1953

slightly over 0.5 length of palm. Distal por-
tion of superior crest well developed, ending
distally in very large, acute, projecting tooth,
ending proximally in transverse groove. Other
grooves and crests not developed. Fingers
laterally compressed, thin at cutting edge.
Carpus heavy, with rounded to acute flat tooth
superior to point of articulation of propodus.
Merus heavy, only twice as long as wide,
without teeth.

Second legs only lightly hirsute. Carpal ar-
ticles with ratio 10 : 5 : 2 : 3 : 4 (in specimen
drawn) .

Third legs compressed and wide. Ischium
with long spine. Merus 3.5 times as long as
wide, strong tooth on inferior distal edge,
lined by long setae along superior margin.
Carpus also lined by setae on both edges; each
distal margin projecting as acute tooth. Pro-
podus with superior edge lined by setae, in-
ferior edge with 5 pairs of movable spines and
unpaired proximal spine. Dactylus short, sim-
ple, acute, strongly curved.

Specimens range to 35 mm. long. In life
mostly colorless with scattered small red
chromatophores on body and greenish-brown
tracing of margins of anterior body regions
and appendages; chelae with dark greenish
cast.

DISCUSSION: The 50 specimens of this spe-
cies from Oahu were examined in detail and
the following variations were noted:

1. The rostrum varied from slightly shorter
to slightly longer than the orbital teeth.

2. The frontal margin of the carapace be-
tween the rostrum and the orbital teeth varied
from almost straight to definitely arcuate.

3. The length of the stylocerite and the rel-
ative lengths of the scaphocerite, carpocerite,
and antennular peduncle varied as noted in
the description. Usually the scaphocerite
reached to somewhat past the middle of the
third antennular article.

4. The second carpal article of the second
legs varied from 0;55 to 0.85 times the length
of the first article, usually being about 0.6 to
0.7 its length. The distal articles appeared to

bear a more fixed relationship to the length
of the second article than to the length of the
first article.

5. The number of spines of the propodus
of the third legs varied from five to seven
pairs.

The smallest specimens, 8 millimeters long,
as well as the largest, 35 millimeters long,
were within this range of variation, and no
particular variation was correlated with the
growth differences.

If one considers that Hilgendorf described
the small chela as if it were the large chela —
if he did not, his was a most aberrant cran-
gonid—the Hawaiian form of this species
does not differ from the original description
in any way that is not covered by the observed
variation.

DISTRIBUTION: The species has been col-
lected, mostly in water from 6 to 20 feet deep,
at the following localities about Oahu: Ka-
neohe Bay, Waimanalo, Hanauma Bay, Wai-
kiki, Nanakuli, Kawela Bay. One specimen
was dredged from water 100 feet deep off
Waimanalo. Off Maui it was collected at five
localities north and south of Lahaina. Ed-
mondson (1925) reported the species from
French Frigate Shoal; the Bernice P. Bishop
Museum has two unreported specimens from
Pearl and Hermes Reef.

Elsewhere the species has been reported in
the Pacific only from the Netherlands Indies,
but within the Indian Ocean it has been col-
lected at various points, including the Mal-
dive Archipelago and the east coast of Africa,

Crangon paradentipes (Coutiere)

Figs. 23 a-i, 24 a-h

Alpheus paradentipes Coutiere, Fauna and

Geog. Maid, and Laccad. 2(4): 880, pi. 74,

fig. 17, 1905.

DESCRIPTION: Rostrum short, reaching only
to near middle of first antennular article, nar-
row, rather flattened on top, with carina ex-
tending only short distance on anterior cara-
pace. Orbital hoods not inflated, anterior

Hawaiian Crangonidae — Banner

73

I I m m

i 2. mm

j 0 . 5 m m

C

Fig. 23. Crangon paradentipes (Coutiere). a, b. Anterior region, dorsal and lateral aspects; c, large cheliped,
lateral aspect; d, large chela, ventral aspect; e, /, small chela, female, lateral and ventral aspects; g, second leg;
h, third leg; /, third leg, propodus and dactylus. {a, b, scale A; c-h, scale B; /, scale C.)

74

PACIFIC SCIENCE, Vol. VII, January, 1953

margin not continuous curve, with acute or-
bital teeth that extend beyond tip of rostrum.
Anterodorsal surface of carapace somewhat
depressed; orbitorostral furrows very broad
and shallow, not reaching as far posteriorly
as posterior edge of eyes; anterior margin be-
tween rostral base and base of orbital spines
almost straight, forming angle of about 30°
to transverse line of body, base of orbital
spines advanced.

Second article of antennular peduncle twice
as long as broad, equal in length to visible por-
tion of basal article, 1.3 times as long as distal
article. Spine of stylocerite reaching to end of
first article. Lateral spine of basicerite poorly
developed but reaching to end of rostrum.
Scaphocerite equal in length to antennular
peduncle, slightly exceeded by carpocerite.

Large chela heavy, 2.5 times as long as
broad, compressed, 1.7 times as high as thick,
with pronounced sculpturing distally. [Eor
terminology of crests and grooves, see Fig.
17^’.] Superior crest high and thin, continuing
distally as strong acute tooth lying above dac-
tylar articulation; soon merging with superior
margin of palm proximal to transverse groove.
Transverse groove broad and deep. Superior
groove rather deep. Plaque crest heavy,
rounded. Palmar groove deep. Inferior crest
flattened with margins rounded, distally pro-
jecting as strong, acute tooth below dactylar
articulation. Inferior depression extensive and
deep; shoulder slightly overhanging but with-
out sharp margins. Dactylus strongly com-
pressed, curved, distorted; only 0.2 length of
entire chela because of distortion; chela ro-
tated so finger closes horizontally almost
across end of palm. Merus 3 times as long as
broad, with superior margin terminated by
acute tooth and inferior internal margin with
2 spines and subterminal tooth. Ischium also
with small spine.

Small chela of female 3.5 times as long as
broad, with fingers 0.5 of length. Articulation
of dactylus flanked on each side by strong
teeth. Fingers with tips bent at 90° and cross-
ing. Merus with 3 spines and small distal

tooth on inferior internal margin; ischium also
with spine. [No male in the collection with
the small chela intact, but according to Cou-
tiere the male chela has the fingers broadened
but not hooked.]

Carpal articles of second legs with ratio
10 : 7 : 3 : 4 : 5.

Merus of third legs almost 6 times as long
as broad, armed with strong subterminal
tooth. Carpus 0.6 as long as merus, with small
acute projection on superior distal margin; in-
ferior margin with 4 movable spines, most
distal one on inferior distal angle. Propodus
0.7 as long as merus, with 7 spines on inferior
margin but none distally. Dactylus 0.3 as long
as propodus, biunguiculate with inferior un-
guis shorter than superior.

Telson twice as wide at base as at tip, 1.8
times as long as broad at tip; lateral margins
slightly convex; posterior margin arcuate.

Largest specimen, a female, 10 mm. long;
color in life not noted.

DISCUSSION: The few specimens in the pres-
ent collection show but little variation; in
some the orbital teeth are slightly longer than
in the specimen described; in some the lateral
spine of the basicerite is better developed; the
number of movable spines on the merus of
the large chela is as great as five. The expected
variation in the number of spines on the car-
pus and propodus of the third legs was ob-
served. One interesting specimen from the
' 'Albatross” collection had its sole remaining
second leg with the carpus divided into six
rather than five articles; it appeared as if an
extra article was inserted between the normal
first and second articles.

Allowing for the variation observed, these
specimens agree well with the descriptions
and figures of Coutiere. I believe that the dif-
ference in the merus of the large chela, which
was described as bearing on "son bord supero-
interne [sic] . . . 7-8 dente ...” and which was
shown as being serrate on the inferior internal
margin, probably normally carried movable
spinules as do the specimens described above.
Otherwise there are no marked differences.

Hawaiian Crangonidae — BANNER

75

Fig. 24. Crangon paradentipes (Coutiere). a, Normal
second leg from specimen from "Albatross” Station
4063; b, second leg with a carpus of six articles from
specimen from an undesignated "Albatross” Station
(U.S.N.M. 63600).

DISTRIBUTION: This species is one of the
few that have been found in sponges in
Hawaii, a common habitat for some species
in other parts of the world. However, of the
four separate collections of the individuals
noted below, only one was noted as coming
from a sponge; whether the others did or not
is not known.

All records of the species probably are from
deep water beyond the reefs; two were from
Oahu, off Kaena Point, 180 feet (in sponge),
and off Diamond Head, 100-200 feet; one
taken by the "Albatross” was from north-
east of Hawaii in 61-63 fathoms (Station
4063, U.S.N.M. 63601) ; the fourth, also taken
by the "Albatross,” merely states "Hawaiian
Islands” (U.S.N.M. 63600).

The species is also known from the Lacca-
dive Archipelago, its type locality, from other

localities in the Indian Ocean, and from the
Netherlands Indies.

Macrochirus Group

Crangon gracilis (Heller) var. simplex
var. nov.

Fig. 25 a-k

Alpheus gracilis Heller, K. Akad. Wiss., Wien,

Sitzungber. 44: 271, Taf. 3, figs. 19-20,

1861.

TYPE specimen: a male 12 mm. long, col-
lected on Waikiki Reef, Oahu (U.S.N.M.
93456).

DESCRIPTION: Rostrum acute, rounded dor-
sally, without Carina; tip reaching from slight-
ly past middle of visible portion to end of
first antennular article. Rostrum separated
from orbital hoods by rounded shallow de-
pression. Orbital hoods rounded, with small
acute orbital teeth, well demarked from curva-
ture of hoods; tips reaching almost to end of
rostrum.

Antennular peduncles with second article
somewhat longer than broad and shorter than
visible portion of first or third. Stylocerite
large, lateral spine reaching past middle of
second antennular article. Lateral spine of
basicerite from shorter to longer than ros-
trum. Scaphocerite with strong lateral spine
slightly longer than antennular peduncle and
shorter than carpocerite.

Large chela strongly compressed, slightly
over 0.5 as thick as high, 2.5 times as long as
high, without lobes or grooves. Shallow
rounded depression on superior internal face
proximal to articulation of dactylus, similar
concavity on inferior internal face inferior to
point of articulation. Fingers 0.3 length of
chela. Chela almost glabrous except for usual
bristles near tip of fingers and few scattered
setae along superior and inferior margins.
Merus without teeth, with 4-6 short spines
along inferior internal margin. Small chela
with fingers longer than palm, with tooth at
point of articulation of dactylus; merus with
armature similar to that of large chela.

76

PACIFIC SCIENCE, Vol. VII, January, 1953

Carpal articles of second legs with ratio
10 : 6 : 3 : 3 : 6.

Ischium and merus unarmed; merus 4-5
times as long as broad; propodus tapering
distally, with proximal end about 1.5 times
width of distal; inferior margin with 6-8 long
slender spines; dactylus slender, 0.25 length
of merus, simple, with slight convexity on
inferior margin at point where lower unguis
would be expected.

Telson with sides rather straight and with
pronounced taper, 1.8 times as wide as long
at posterior margin. External spine of uro-
pods almost always brown to black.

Specimens in collection, range to 18 mm. in
length. In life, body and chelae transversely
banded with broad gray bands of red and blue
chromatophores; eggs brilliant yellow. Trans-
verse bands red in preserved specimens.

DISCUSSION: This species varies in a num-
ber of characteristics. The rostrum, which in
most extends only to the middle of the visible
portion of the first antennular article, in some
specimens reaches to the end of that article
and in one specimen extends to one third of
the second article. In some specimens there is
a faint trace of an obtusely angular dorsal Ca-
rina on the rostral base. The orbital teeth vary
in breadth and length, sometimes being al-
most as long as the rostrum and sometimes
only half as long (contrast Fig. 2^a and c); in
all cases, however, the teeth arise abruptly
from a uniformly curved anterior margin of
the orbital hood. There are variations, as
would be expected, in the lengths of the sty-
locerite and of the lateral spine of the basicer-
ite, the latter in a few cases exceeds the length
of the rostrum. The small and large chelae
vary somewhat in their proportions but their
general form and armature remain constant as
shown. The second carpal article of the sec-
ond legs varies from 0.5 to 0.7 the length of
the first, with a similar variation in the other
articles as well. On the third legs the variation
of the breadth of the merus and the number
of spines on the propodus is noted above.
Variations were also noticed in the propor-

tions of the telson where a series of six speci-
mens gave differences in the ratio of the width
of the posterior border to the greatest width
of 1. 6-2.0, and in the ratio of the width of the
posterior border to the length of 2. 4-3.0. A
most peculiar variation was observed in the
external spine of the uropods: in 40 specimens
all but 5 have brown to black spines, 2 have
spines with faint color, and 3 have colorless
spines. This change of color is not the result
of the solvent action of the preserving fluid
(formaldehyde or alcohol), as some specimens
preserved over 14 years still have the spines as
dark as the newly preserved ones. Perhaps the
variation may be the result of moulting, for
several of the specimens with slightly colored
or colorless spines have what appears to be
softer chitin.

The varietal name refers to the lack of a sec-
ondary unguis on the dactylus of the third leg.

RELATIONSHIP: There have been four de-
scriptions published for this species and its
varieties. The type specimen first described by
Heller was subsequently redescribed by de
Man (1911), who corrected some errors found
in Heller’s description and also extended the
original description. Coupled with this rede-
scription, de Man also described a tentative
variety, C. gracilis var. luciparensis which he
distinguished from Heller’s type specimen
chiefly by minor differences in the proportions
of the various appendages and by the external
spine of the uropods being black instead of
brown. It is interesting that the variations
noted above are sufficient to bridge these dif-
ferences. Coutiere (1905) described C. gracilis
var. alluaudi from Mahe which was distin-
guished from the main species by the lack of
a secondary unguis, and on all other points
”ces specimens me paraissent impossibles a
distinguer de V A. gracilis!' One is at a loss to
know whether the specimens were exactly
like Heller’s erroneous description, therefore
different from the type specimen, or exactly
like the type specimen.

The Hawaiian form here described is dis-
tinguished from the original species and from

Hawaiian Crangonidae — BANNER

77

_) I mm
_.2mm
1mm
i0.5mm

Fig. 25. Crangon gracilis (Heller) var. simplex var. nov. a, h. Anterior region, dorsal and lateral aspects; c, ,
anterior carapace, dorsal aspect, showing a more typical development of the orbital teeth; d, e, large chela, lateral
and medial aspects; /, g, large cheliped, merus and carpus, lateral and medial aspects; h, small cheliped; i, second
leg; /, third leg; k, third leg, dactylus. {a-c, scale A; d-g, scale B; h-j, scale C; k, scale D.)

78

PACIFIC SCIENCE, Vol. VII, January, 1953

the two varieties by the following character-
istics: First, the other forms have a distinct
but subacute dorsal carina, while the Hawai-
ian variety is usually evenly rounded dorsally.
Second, the base of the rostrum is separated
from the orbital hoods by rather shallow
rounded depressions instead of deep narrow
grooves as de Man describes them. Third, the
inferior internal margin of the merus of the
large cheliped is without a small obtuse tooth
at its distal extremity in the local specimens.
From the original type for the species and
from de Man’s variety, this variety is distin-
guished, as is C gracilis var. alluaudi, by the
lack of the secondary unguis on the dactylus.

Future workers may decide that the lack of
the secondary unguis is sufficient to separate
this form, together with Coutiere’s, as a spe-
cies distinct from that of Heller. Except for
the fact that this is not supported by other
differences, I would be inclined to make this
separation myself. However, as Coutiere did
not think of this as a specific difference, as
there are no supporting differences, and as
there is the swelling on the dactyl which may
be the remnant of the secondary unguis, I
have decided to leave the form within this
species. The new variety for the Hawaiian
specimens was created to emphasize this dif-
ference and the fact that these specimens do
not have the deep grooves between the orbital
hoods and the rostrum.

C. gracilis var. simplex is most closely related
to C. edmondsoni Banner and C. latipes Banner
of the other local species. The differences be-
tween them are set forth in Table 3.

DISTRIBUTION: This species has been col-
lected both from the shallow reefs and in
waters up to 20 feet deep at the following lo-
cations on Oahu : Waimanalo, Hanauma Bay,
Black Point, Waikiki, Nanakuli, Mokuleia,
Kawela Bay, and Punaluu. On Maui it was
collected at Makena, Kalama Park, Lahaina,
and four localities north and south of Lahaina.
Edmondson (1925) reported it (as C. gracilis')
from Laysan and Lisianski Islands. No speci-
mens were collected by the "Albatross.”

Edmondson (1925) also recorded the spe-
cies from Johnston Island in the central Pa-
cific. It and its two varieties have been re-
ported from the Netherlands Indies, the Indian
Ocean, and the Red Sea.

Crangon edmondsoni sp. nov.

Fig. 26 a-p

TYPE specimen: a female 24 mm. long col-
lected at Kawela Bay, Oahu, by C. H. Ed-
mondson (Bernice P. Bishop Mus. 4059).

DESCRIPTION : Rostrum narrow, longer than
breadth at base; concave sides with 3 or 4
pairs of short bristles; produced as acute tip,
reaching to or slightly beyond end of first an-
tennular article, distally depressed; slightly
rounded dorsally, without carina. Orbital
hoods continuous with curvature of carapace
posteriorly, not sharply delimited from sur-
rounding carapace and separated from base of
rostrum by only slight shallow depressions.
Anterior margin of orbital hoods produced
into acute, dorsally flattened teeth, reaching
almost to mid-point of rostrum. Margin of
orbital hoods between rostral base and apex
of hoods straight to concave.

Visible portion of first article of antennule
slightly larger than second and 0.25 longer
than third. Stylocerite strong, reaching to
middle of second antennular article. Entire
dorsal surface of antennular peduncle with oc-
casional short setae, with denser groupings of
longer setae at points of articulation. Inner
flagellum about 7 times as long as peduncle;
outer flagellum 4 times as long as peduncle.

Lateral spine of basicerite of antenna acute,
tip not reaching to end of first antennular ar-
ticle. Scaphocerite as long as antennular pe-
duncle with lateral spines only slightly longer
than squamous portion, squamous portion ta-
pering toward tip. End of carpocerite reach-
ing slightly beyond scaphocerite. Antennal
flagellum longer than body, about twice as
long as inner antennular flagellum.

Large chela elongate, tapering, margins
rounded, laterally compressed, 3.4 times as

> g

Hawaiian Crangonidae — BANNER

79

Fig. 26. Crangon edmondsoni sp. nov. a, b, Anterior region, dorsal and lateral aspects; c, d, e, large cheliped,
lateral, ventral, and medial aspects;/, g, large cheliped, merus, lateral and medial aspects; h, small cheliped, lateral
aspect; i, j, small chela, ventral and ventrolateral aspects; k, small cheliped, merus, medial aspect; /, second leg;
, third leg; n, third leg, dactylus; o, fifth leg, propodus and dactylus; p, uropod and telson. {a, b,f, g, o, p, scale
; c-e, h-k, scale 'Q\ I, m, scale C; n, scale D.)

80

PACIFIC SCIENCE, Vol. VII, January, 1953

long as high, 5.0 times as long as thick. Mar-
gins entire, with slight trace of longitudinal
groove proximal to dactylus. Fingers com-
pressed, slightly less than 0.5 as long as palm,
slightly depressed from axis of palm, rotated
so dactylus opens and closes on lateral plane.
Dactylus 1.8 times as long as broad, 2.5 times
as long as thick near base, with the upper (or
lateral) margin slightly rounded, distal mar-
gin strongly rounded. Chela with occasional
setae on superior and inner face, with usual
grouping of short bristles on opposing faces
of fingers. Carpus of usual form. Merus with
inferior margins acute, superior margin
rounded. Superior margin with 6 feeble spi-
nules on basal 0.7; distal angle with obtuse,
projecting tooth. Inferior lateral edge of me-
rus with 13 feeble spinules, slightly larger than
those on superior edge, margin produced as
acute tooth. Inferior internal margin of merus
with 7 movable spinules, with bristles be-
tween them. Merus 2.7 times as long as wide
distally. Ischium of usual form, with 3 strong
movable spines on inferior internal face, and
on adjacent margin of merus.

Small chela almost as long as large chela
and with same rotation. Chela almost as high
as thick, 5.0 times as long as btoad, with
slight taper toward fingers. Inner margin of
palm continued as obtuse projection at point
of articulation of dactylus. Fingers almost as
long as palm, tapering, definite hook on tip.
Merus somewhat similar to that of large chela,
but without spines or bristles on superior or
inferior external margins; 5 movable spines
and several bristles on inferior internal mar-
gin. Ischium with 5 smaller movable spines.
Lateral face of merus slightly more than twice
as long as wide.

Second thoracic legs about 0.7 as long as
entire animal. Ischium 1.2 times as long as
merus, 0.7 as long as entire carpus. Carpal ar-
ticles with ratio 10 : 7 : 4 : 4 : 6; fifth article
twice as broad as proximal end of first. Chela
1.5 times as long as first carpal article.

Ischium of third legs 0.4 as long as merus,
with proximal movable spine. Merus 6 times

as long as broad, with rather straight margins,
without distal teeth. Carpus 0.6 as long as
merus, with inferior and superior margins pro-
jecting distally as small rounded teeth. Pro-
podus 0.7 as long as merus, with 6 movable
spines along inferior margin. Dactylus almost
0.3 as long as propodus, curved, simple; su-
perior margin with indentation or tooth 0.7
of distance from base, curve then continuing
to tip, with tuft of short setae on each side of
dactylus at this break. Entire legs with occa-
sional setae. Fourth legs similar. Propodus of
fifth legs with 7 movable spines, and with
'’brush” feebly developed, of but 6 rows of
setae.

Telson 3.0 times as long as width of pos-
terior margin. Sides tapering, with angle of
taper greater in distal 0.2. Anterior pair of
dorsal spines located in middle of dorsal sur-
face, posterior pair 0.7 of distance toward
posterior edge. Posterior margin somewhat
convex.

Specimens range to 27 mm. long. Color in
life not noted; some of preserved specimens
with 7 distinct longitudinal bands reaching
from antennular peduncles to end of abdo-
men; in preservation all bands red. Middle
band very narrow; dorsolateral bands quite
broad; lateral and ventrolateral bands, latter
along margins of abdominal pleura, each
about 0.5 width of dorsolateral band. Distal
0.25 of large and small chelae violet in pre-
servative. No specimens with any dark pig-
mentation on external spine of uropod.

DISCUSSION : Even with the small number of
specimens of this species at hand, 19 in all,
some interesting variation was noted. As
usual, the relative length of the rostrum varied
in some specimens. This may be due more to
the degree of flexure of the antennular pe-
duncle than to the variation in length of the
rostrum. In all cases the inner anterior margin
of the orbital hoods were concave. Slight dif-
ferences were noted in the relationships of the
parts of the antennular and antennal pedun-
cles. The most conspicuous differences were
in the large chela, where the length of the fin-

Hawaiian Crangonidae — Banner

81

gers varied from 0.3 to 0.5 the length of the
palm, and the whole chela from less than 3.0
to more than 3.5 times as long as high. Both
large and small chelae may be glabrous. In all
specimens the chela was without the depres-
sions of the upper and lower margins that
were found in C. gracilis var. simplex Banner.
The teeth on the superior margin of the me-
rus, not conspicuous even in large specimens,
were scarcely visible in small specimens. The
ratio of the first to the second article of the
carpus of the second legs varied from 10 : 7
to 10 : 8. The merus of the third leg varied
from six to seven times as long as broad.

The species is named in honor of Dr. C. H.
Edmondson, Professor Emeritus of Zoology
at the University of Hawaii and Curator of In-
vertebrates at the Bernice P. Bishop Museum.

relationship: This species appears to be
very closely related to C gracilis and, there-
fore, should belong to the Macrochirus
Group. According to de Man’s key (1911),
the characteristics of this species would leave
it close to C. shearmei (Alcock and Anderson) ,
but obviously they are not closely related, be-
cause C. shearmei^ a deep-sea species, has a
minute rostrum, and the large chela lacks en-
tirely the piston on the dactylus.

This species shows many characters in com-
mon with C. gracilis var. simplex Banner and C.
latipes Banner; the three species are distin-
guished in Table 3.

DISTRIBUTION: Besides the type specimen
from Kawela Bay, Oahu, the species has been
collected on Oahu from Hanauma Bay at
15 feet, off Kahala at 150 feet, and off Nana-

TABLE 3

Characteristics Differentiating Crangon latipes, C. gracilis var. simplex, and C. edmondsoni

C. latipes

C. gracilis var. simplex

C. edmondsoni

Maximum observed size 8.8 mm.

Orbital teeth 0.3-0. 5 length of ros-
trum; abruptly set off from orbital
hoods; rostrum reaching end of
first antennular article

Large chela 2.2 times as long as
broad; 2.2 times as broad as thick;
without depressions on margins

Merus of large cheliped with 2 or
less spines on inferior internal
margin; none elsewhere

Carpal articles of second legs with
ratio 10 : 8 : 3 : 3 : 14; second ar-
article as broad as long

Third legs

Ischium with spine
Merus 3-3.5 times as long as
broad

Propodus with 4-5 spines
Dactylus 2.3 times as long as
broad, distally strongly curved
and abruptly tapering

External spine of uropod never dark

Coloration not noted

Maximum observed size 18 mm.

Orbital teeth usually 0.7 length of
rostrum; abruptly set off from or-
bital hoods; rostrum not reaching
end of first antennular article

Large chela 2.5 times as long as
broad; 2.0 times as broad as thick;
with depressions on upper and
lower margins

Merus of large cheliped with 4-6
spines on inferior internal mar-
gin; none elsewhere

Carpal articles of second legs with
ratio 10 : 6 : 3 : 3 : 6; second ar-
ticle twice as long as broad

Third legs

Ischium unarmed

Merus 4-5 times as long as broad

Propodus with 6-8 spines
Dactylus almost 4 times as long
as broad, slightly curved and
gradually tapering throughout
length

External spine of uropod usually
brown to black

Body with longitudinal stripes

Maximum observed size 27 mm.

Orbital teeth 0.5 length of rostrum;
merging with orbital hoods on
inner margin; rostrum reaching to
or beyond end of first antennular
article

Large chela 3.5 times as long as
broad; almost as broad as thick;
without depressions on margins

Merus of large cheliped with 7 spines
on inferior internal margin; su-
perior and inferior external mar-
gins also spinose

Carpal articles of second legs with
ratio 10 : 7 : 4 : 4 : 6; second arti-
cle over twice as long as broad

Third legs

Ischium with spine

Merus 6-7 times as long as broad

Propodus with 6 spines
Dactylus 3 times as long as broad,
slightly curved and gradually
tapering throughout length,
with notch on superior margin

External spine of uropod never dark

Body with transverse stripes

82

PACIFIC SCIENCE, VoL VII, January, 1953

kuli at 15-20 feet; two other specimens came
from unspecified localities on Oahu, presum-
ably from shallow water. One specimen in
Bernice P. Bishop Museum came from Ma-
kena, Maui, and one was also collected in 3
feet of water at Kalama Park on that island.
R. W. Hiatt collected two specimens at Ha-
lape, Kau Coast, Hawaii, from coral heads in
the intertidal zone.

Crangon latipes sp. nov.

Fig. 27 a-j

Alpheus sp. Coutiere, Fauna and Geog. Maid.

and Laccad. 2: 882, pi. 74, fig. 19, 1905.

TYPE specimen: a male 8.5 mm. long col-
lected off Waikiki Reef, Oahu, in 20 feet of
water (U.S.N.M. 93457).

DESCRIPTION: Anterior carapace produced
as well-developed rostrum, tip reaching slight-
ly beyond end of first antennular article; ros-
trum about 0.3 longer than width at base.
Rostral base laterally demarked from orbital
hoods by shallow depressions; depressions
deeper and more abrupt posteriorly; rostrum
and base without definite carina, rounded dor-
sally. Orbital hoods inflated, anterior margin
rounded in both dorsal and lateral aspects.
Orbital teeth small, acute, arising abruptly
from curvature of orbital hoods; tips reach
about 0.3 length of rostrum.

Antennular peduncle in dorsal view with
second article about 0.2 longer than visible
portion of first article and 0.3 longer than
third; second article 1.3 times as long as broad.
Acute tip of stylocerite reaching to middle of
second antennular article.

Lateral spine of basicerite reaching nearly
to end of first antennular article. Scaphocerite
with lateral spine strong, lateral margin al-
most straight, squamous portion well devel-
oped; tip reaches well beyond end of anten-
nular peduncle. Carpocerite reaching almost
to end of scaphocerite.

Large chela smooth in contour, strongly
compressed. Chela 1.8 mm. high, 0.8 mm.
thick, 4.0 mm. long to end of dactylus; dac-
tylus occupying distal 1.1 mm. Only indenta-

tions 2 very shallow grooves reaching proxi-
mally from articulation of dactylus; grooves
very faint, visible only by rotating cheliped.
Merus 1.3 mm. long, about twice as long as
breadth of inner face; inferior internal margin
with 2 movable spines, terminating distally
as small, acute tooth; termination of superior
margin rounded.

Small chela 1.0 mm. broad, 2.0 mm. long;
fingers 0.5 length of chela, hooked distally.
Merus similar in size and proportions to that
of large chela but with 3 short spines on dis-
tal portion of inferior internal margin.

Second legs with ischium 0.7 mm. long;
merus 0.9 mm. long, 0.24 mm. broad; articles
of carpus with total length of 1.0 mm., with
ratio 10 : 8 : 3 : 3 : 14; second secondary ar-
ticle as long as broad; chela 0.6 mm. long,
fingers 0.5 of length.

Ischium of third legs 0.4 mm. long, with
movable spine; merus 1.36 mm. long, 0.44
mm. broad, inermous; carpus 0.6 mm. long;
propodus 1.1 mm. long, with 5 movable
spines; dactylus 0.35 mm. long, 0.13 mm.
broad in basal half, abruptly tapering to sharp-
ly curved acute tip.

Fourth legs similar in structure. Fifth legs
generally similar but more slender, propodus
with only several distal spines and without
well-defined "brush.”

Telson 1.4 mm. long, 0.75 mm. broad at
base, and 0.36 mm. broad at tip, sides with
uniform taper. Outer distal spines 0.07 mm.
long, inner spines 0.18 mm. long; anterodor-
sal spines about 0.5 distance from tip, pos-
terior spines 0.32 mm. from tip.

Three additional specimens: male 6.5 mm.
long; 2 females 8.8 and 8.0 mm. long. Color
of specimens in life not noted.

DISCUSSION: Two of the paratypes agree
quite well with the type specimen. The female
from Nanakuli differs chiefly in that the ros-
trum is asymmetrically bifurcate at the tip
and one of the orbital teeth points toward the
rostrum. This, obviously, is the result of an
accident or of anomalous growth. The female
from Lahaina also agrees with the type quite

Hawaiian Crangonidae — BANNER

83

I mm

— 1 0.4mm

Fig. 27. Crangon latipes sp. nov. b. Anterior region, dorsal and lateral aspects; c, d, large cheliped, lateral and
medial aspects; e, small cheliped, lateral aspect; /, second leg; g, third leg; h, third leg, dactylus; /, fifth leg; j, tel-
son. {a, b, /, g, i, j, scale A; c-e, scale B; h, scale C.)

84

PACIFIC SCIENCE, Vol. VII, January, 1953

well, except that the large chela is somewhat
more slender and its merus does not bear the
two movable spines found in the type.

However, the second male, from the island
of Hawaii, shows a few notable differences.
The rostrum, the stylocerite, and the lateral
spine of the basicerite are all shorter, with the
rostrum and the stylocerite reaching only to
near the end of the first third of the second
antennular article, and the lateral spine of the
basicerite reaching only to the middle of the
visible portion of the first article. The orbital
teeth are half the length of the rostrum, and
the orbital hoods are slightly less inflated.
The large chela is 2.7 times as long as broad
instead of 2.2, and the fingers occupy slightly
over 0.5 of the length of the chela instead of
slightly more than 0.25. The ratio of the car-
pal articles of the second legs is slightly differ-
ent— 10 : 10 : 8 : 6 : 16, with the second arti-
cle slightly broader than long. The merus of
the third legs is 3.5 times as long as broad in-
stead of 3.0 times, and the propodus of the
same legs bears four instead of five spines.
Other parts of the two specimens showed
close similarity.

This male appears to be either within the
normal range of variation or to be a closely
related variety. It is likely that all the differ-
ences may be due to simple growth differences
combined with the inherent variation as shown
in so many other species; in any case, the dif-
ferences are not great enough, especially with
only one specimen showing them, to warrant
the erection of a new species or variety.

RELATIONSHIP: This species is closely re-
lated to two other Hawaiian species, C. gra-
cilis var. simplex Banner and C. edmondsoni Ban-
ner. The more conspicuous differences are
given in Table 3.

Of these characteristics, the most readily
observed and decisive for rapid separation of
this species are the carpus of the second legs
and the dactylus of the third to fifth legs. The
anterior region of the carapace also affords
good characteristics, but it is more variable
and therefore less reliable.

According to de Man’s key (1911) in the
Macrochirus Group, this species keys out to
the two species C faceta (de Man) and C.
spkndida (Coutiere) . It can be separated easily
from these two species on the basis of the
breadth of the articles of the second and third
legs; by the greater breadth of the large chela;
and especially by the lack of the longitudinal
groove on the palm of the chela (characteris-
tics taken mostly from de Man’s key; Cou-
tiere’s original description, without figures, of
C. spkndida was too brief to be of any value) .

This species is apparently identical to Al-
pheus sp. of Coutiere {loc. cit.) which was left
unnamed because his three specimens each
lacked the large chela. The form and propor-
tions of the anterior body region and the pro-
portions of the second legs, both shown in
the plates, are identical to this species; the
unique dactylus of the third legs was neither
mentioned nor drawn. If the dactyli are the
same, the specimens undoubtedly belong to
this species. Incidentally, Coutiere believed
that the species would belong to the Mega-
cheles Group; the large chela shows its affini-
ties to be with the Macrochirus Group.

DISTRIBUTION: This rare species is evidently
widespread among the larger Hawaiian Is-
lands at least, for the four specimens are from
three islands. The type came from water 20
feet deep off Waikiki Reef, Oahu; an 8.8 mm.
female came from 15-20 feet of water off
Nanakuli, Oahu; an 8.0 mm. female came
from 4 feet of water at Lahaina, Maui; and the
aberrant 6.5 mm. male was collected by R. W.
Hiatt from a head of Pocillopora meandrina
Verrill in 5 feet of water at Halape, Kau Coast,
Hawaii. Coutiere’s specimen came from Hu-
lule Male Atoll, Indian Ocean.

Crangon ventrosa (Milne-Edwards)

Fig. 28 a-i

Alpheus ventrosus Milne-Edwards, Histoire Na-
turelle des Crustaces, 2: 352, 1837.

Alpheus laevis Randall, Acad. Nat. Sci. Phila.,
Jour. 8(1): 141, 1839-

Hawaiian Crangonidae — BANNER

Fig. 28. Crangon ventrosa (Milne-Edwards). a, b, Anterior region, dorsal and lateral aspects; c, large chela
female, lateral aspect; d, large cheliped of female, medial aspect; e, small cheliped, lateral aspect; /, second leg;
g, third leg; h, /, third leg, dactylus, inferior and posteroinferior surfaces. (In drawings h and /, the portions shaded
with dashed lines are of thin chitin. Appendages drawn from several specimens, so relative size is not indicated.)
{a-g, scale A; h, i, scale B.)

2mm
1 1 mm

86

PACIFIC SCIENCE, VoL VII, January, 1953

Alphem ohesomanus Boone, Vanderbilt Mar.

Mus., Bui. 6: 135, 1935. [Part of distribution

records, only.]

DESCRIPTION: Rostrum slender, acute,
reaching almost to end of first article of an-
tennular peduncle, straight to depressed at
tip; small groups of short bristles along lower
margin. Rostral base broad, flattened, not
carinate dorsally, separated from carapace by
deep and narrow sulci on each side. Orbital
hoods rounded laterally, extending anteriorly
as acute teeth from inner portion of hood,
0.3-0. 5 length of rostrum.

Antennular peduncle slender, with second
article variable but usually only slightly longer
than visible portion of first or third. Stylocer-
ite well developed, reaching past middle of
second antennular article. Lateral spine on
basicerite acute, equal to or exceeding length
of stylocerite. Scaphocerite long, distally nar-
row, with lateral spine only slightly longer
than squamous portion and reaching slightly
beyond antennular peduncle and tip of car-
pocerite.

Large chela usually less than 2.5 times as
long as broad, compressed, with margins
rounded, without grooves or crests. Dactylus
elongate, not strongly arcuate, 0.5 length of
palm, tip usually acute in males, rounded in
females. Merus with both superior and in-
ferior internal angles obtuse, although pro-
jecting; inferior internal margin with about
5 spines.

Small chela almost as long as larger but
more slender, 2.5-3 times as long as broad.
Dactylus strongly curved at tip, crossing fixed
finger when closed. Inner margin of cutting
face of fixed finger with distinct lamellar ridge
along entire length, series of short strong bris-
tles along outer edge. Fingers about equal in
length to palm. Merus with superior distal
angle subacute, often with several movable
spines on inferior internal edge.

Carpal articles of second legs totaling 1.7
times length of merus and broad in propor-
tion to width; second article only 1.5 times as

long as broad; articles with ratio 10 ; 8 : 6 :
6 : 12.

Ischium of third legs inermous. Merus in-
ermous, over 3 times as long as wide, widest
at 0.7 of length then tapering sharply to tip.
Carpus with inferior margin continued as
small tooth. Propodus with 5 to 7 movable
spines, each with broadened and flattened tip.
Dactylus heavy, blunt, laterally compressed,
with thick longitudinal ridge on anterior face
continuing around tip as transparent flange,
blunt tip carrying ''fingernail” of hard chitin,
tip reinforced with another ridge of chitin in
form of horse’s hoof. Inferior and lower pos-
terior portion of face of dactylus proximal to
"nail” of soft and flexible chitin (see Fig.
28^, /■).

Body in general large and strongly com-
pressed. Large females in collection range to
42 mm. in length. Usually orange-red on up-
per surface, sometimes longitudinally striped
with deeper red along dorsal surface of cara-
pace and abdomen; mottled with spots of
deeper red along superior and superolateral
portions of both large and small chela.

DISCUSSION: The usual variation was noted
in this species; much of it has been remarked
upon in the description. Additional variation
was found in the proportions of the small
chela; in the ratio of the lengths of the carpal
articles of the second legs, especially in the
fifth article which varied from noticeably
shorter than the first to somewhat longer; and
in the number of spines on the propodus of
the third legs. The fingers of the large chela
show sexual dimorphism, but a few specimens
seemed to intergrade. One specimen in the
collection had an orbital tooth on the right
but not on the left side.

These specimens agree well with the brief
description given by Milne-Edwards. De Man
(1911) remarked on the variation manifested
by the few specimens of this species in his col-
lection and noted that the second article of
the antennular peduncle varied from slightly
shorter than the first article to over 1.5 times
as long, the rostrum from shorter than to

Hawaiian Crangonidae — BANNER

87

equal to the first peduncular article, and the
scaphocerite from ' shorter than to as long as
the antennular peduncle. De Man also sug-
gests that the two forms of the large chela
(one with a rather obtuse dactylus, the other
with an acute dactylus) might be a sexually
dimorphic trait; in general this was true in the
Hawaiian specimens.

distribution: C. ventrosa, in Hawaii at
least, has always been found in living heads
of corals of the genus Pocillopora, usually
either P. meandrina var. nohilis Verrill or P. //-
gulata Dana, and occasionally in P. damicornis
(Linnaeus); in these corals it associates with
Synalpheus charon (Heller) and crabs of the
genus Trapezia.

This, although not the most common Ha-
waiian species, is certainly the most spectacu-
lar, both in color and in size, and consequent-
ly there have been many records of it through-
out the Islands. The first record was under the
name of A. laevis made by Randall in 1839; it
was also reported by Dana (1852), Stimpson
(I860), and Bate (1888). In the Islands it
probably occurs wherever Pocillopora is found;
specimens studied have come from the fol-
lowing localities: on Oahu, both in shallow
water and in water up to 20 feet or more deep,
at Waimanalo, Hanauma Bay, Waikiki, Nana-
kuli, and Kawela Bay; on Maui, at Makena
and Lahaina. Previously unreported specimens
collected by the "Albatross” came from the
following locations: Honolulu (further data
unrecorded); south of Molokai, 23-24 fath-
oms; two dredge hauls on Penguin Bank
(west of Molokai), 27-29 fathoms; and off
Kauai, 18-41 fathoms. "Albatross” Stations:
(not recorded), 3847, 4031, 4032, and 4023,
respectively; U.S.N.M.: 63633, 63634, 63636
(63637, 63638 — lot divided into two vials),
and 63635, respectively. Edmondson (1925)
has reported the species from French Frigate
Shoal and Laysan Island; there are 15 speci-
mens in Bernice P. Bishop Museum jfrom
Pearl and Hermes Reef.

Crangon ventrosa has been reported through-
out the entire tropical Indo-Pacific region.

from the Gulf of Akabah at the western tip of
the Red Sea to the Gulf of California in Mex-
ico. Edmondson has reported the species from
Johnston, Wake, and Palmyra Islands, and
others have reported it from Tahiti.

As C ohesomanus (Dana), Boone (1935: 135)
reported the following previous distribution-
al records (in part) : "Gulf of California (Cou-
tiere) ; Sandwich Islands (Randall) ; Honolulu
(Bate) ; . . . Sandwich Islands and Fiji Islands
(Dana) ; . . . Gulf of Aykab [sic] and through
the Red Sea (de Man) . . . . ” She mentions none
of the records noted above for C. ventrosa
(1935: 126). Inasmuch as the records here
quoted from Boone were made by the authors
to which she attributes them not for C. obeso-
manus but for C. ventrosa (A. laevis), it would
appear that these records have been inter-
changed. However, other records listed by her
under C. ohesomanus are correct.

Crangon amirantei (Coutiere)

Fig. 29 a-j

Alpheus amirantei Coutiere, Soc. Philomath.

Paris, Bui. IX, 11(5): 15-16, 1908.

Alpheus amirantei CouXAhtQ, Linn. Soc. London,
Trans. II (Zoology), 14(4): 421, pi. 63, fig.
16, 1931. [Text exactly the same as 1908
reference; plates not previously published.]

DESCRIPTION: Carapace with swollen, al-
most hemispherical orbital hoods, anterior
margin of hoods produced as rounded, sub-
vertical keel slanting mesad ventrally, area be-
tween keel and rostrum flat. Frontal margin
anterior to orbital hoods almost straight; small
notch near base of rostrum. Rostrum small,
extending less than 0.5 length of visible por-
tion of first antennular article, with high ros-
tral Carina only slightly lower than orbital
hoods. Deep depression between rostral base
and orbital hoods. Side of carapace between
base of antenna and orbital hood depressed.

Visible portion of first antennular article as
long as second; third article about 0.75 length
of second; second article about 1.5 times as
long as broad. Stylocerite very short, reaching

88

PACIFIC SCIENCE, VoL VII, January, 1953

only slightly beyond 0.7 of length of visible
portion of first article, and only slightly ex-
ceeding broadened portion. Outer flagellum
heavy, short, not over 1.5 times length of
peduncle.

Basicerite with strong, acute spine. Scapho-
cerite strong, outer margin curved, spine dom-
inant over reduced squamous portion; tip
reaching beyond end of antennular peduncle
and carpocerite. Carpocerite as long as anten-
nular peduncle.

Proximal portion of palm of large chela sub-
cylindrical in section, as thick as high, entire
chela less than 3 times as long as thick; distal-
ly tapering, more flattened. Chela proximal to
articulation of dactylus slightly inflated; lat-
eral face proximad of articulation with poorly
defined but noticeable longitudinal groove.
Dactylus of usual form, about 0.3 length of
entire chela. Merus twice as long as broad,
superior distal margin rounded, inferior inter-
nal margin with strong subterminal tooth,
inermous. Large chela sexually dimorphic in
size, that of female 0.8 and that of male 1.4
times as long as carapace.

Small chela sexually dimorphic in size and
proportions. Appendage of female with fin-
gers 0.4 as long as chela; chela 3 times as long
as broad. Carpus elongate, equal in length to
fingers. Merus twice as long as broad, 0.8 as
long as chela; superior margin irregular; in-
ferior internal margin without tooth. Append-
age of male about 0.3 larger proportionately
than that of female, with fingers 0.6 length of
chela; chela 3 times as long as broad. Carpus
0.5 as long as fingers, Merus similar to that of
female, but 0.7 as long as chela and with su-
perior margin regular.

Carpal articles of second legs with ratio
10 : 23 : 5 : 5 : 10.

Third legs with strong movable spine on
ischium. Merus 2.6 times as long as broad,
with very strong subterminal tooth. Carpus
0.5 as long as merus, with inferior distal mar-
gin projecting as acute tooth; article without
movable spines. Propodus only slightly long-
er than carpus, heavy, with about 10 strong

movable spines. Dactylus simple, strong, and
slightly curved, about 0.25 as long as merus.

Telson 1.5 times as long as broad at base,
anterior breadth 1.7 times breadth at posterior
margin. Lateral margins slightly convex an-
teriorly and slightly concave posteriorly; pos-
terior margin arcuate. Length of larger pair of
posterior spines approximately equal to
breadth of tip; dorsal spines as long as larger
posterior spines, located approximately 0.3
and 0.7 of distance from anterior end of tel-
son; distal border without spinules. Inner uro-
pod with numerous strong spines on distal
and distolateral margins.

Largest female 10.0 mm. long; color in life
not noted.

DISCUSSION: In the four specimens avail-
able, no marked variation was noticed. Unfor-
tunately, a second female did not have a small
chela, so it was impossible to determine
whether the irregularity of the superior mar-
gin of the merus noted above was actually a
sexually dimorphic characteristic or merely an
individual variation.

This form differs in several respects from
the type as described by CoutiUe. In the first
place, in CoutiUe’s specimens the scaphocer-
ite was not longer than the carpocerite, and
both were longer than the antennular pedun-
cle instead of as described here. The basicerite
is described as, "a peu pres inerme,” w’hereas
in this form the basicerite has a relatively
strong spine. The greatest differences are in
the small chelae; the differences in propor-
tions of the finger and the palm between the
specimen drawn by Coutiere and the one
drawn here are merely sexual dimorphism;
however, Coutiere shows his specimens as
carrying a continuous band of setae which are
lacking in all Hawaiian specimens, and he
states that the merus bears a strong tooth
("epine”) which is lacking in the Hawaiian
specimens. The second article of the carpus
of the second legs is over three times as long
as the first, and in the local specimens it is
only 2.3 times as long. Finally, the carpus and
propodus in Coutiere's specimens were slight-

Hawaiian Crangonidae — BANNER

89

. i I i0.5mm

■ A

. 'I I 1 1 1 mm

e ®

Fig. 29. Crangon amirantei (Coutiere). a, h, Anterior region, dorsal and lateral aspects; c, large cheliped, lateral
aspect; d, large chela, ventrolateral aspect; e, large cheliped, merus, medial aspect; /, small cheliped, lateral aspect;
g, second leg; h, third leg; /, third leg, dactylus; 7, telson and uropod. (Drawn from a female.) {a, b, i, j, scale A;
c-h, scale B.)

90

PACIFIC SCIENCE, Vol. VII, January, 1953

ly longer in relation to the merus than they
were in these specimens.

These differences cannot be considered as
of major magnitude because in related species
(see C. paragracilis, C. brevipes, and C. clypeata)
all of these characteristics were found to be
quite variable. Possibly the most important
difference is the lack of the tooth on the merus
of the small cheliped, but, even if this were
constant, it would not be sufficient for the
erection of a new species.

DISTRIBUTION: All five specimens in the
collection came from Oahu: two from Hana-
uma Bay, 20 feet deep, one off Waikiki, 20
feet deep, and two from Nanakuli, 15 feet
deep.

Coutiere’s two specimens, both males, came
from Amirante, 25-80 fathoms, in the Indian
Ocean.

Crangon nanus sp. nov.

Figs. 30 a-n, 31 a-h

Alpheus sp..^, Metalpheus, n. gen., Coutiere,
Soc. Philomath. Paris, Bui. IX, 9(5): 23-26,
1908.

Alpheus sp.?, Metalpheus, n. gen., Coutiere,
Linn. Soc. London, Trans. II (Zoology)
17(4): 419-421, pi. 62, fig. 15, 1921. [The
same description as above, but with fig-
ures.]

TYPE specimen: a female 9.0 mm. long,
collected from coral in water about 10 feet
deep at Hanauma Bay, Oahu (U.S.N.M.
93458).

DESCRIPTION: Rostrum short, acute, barely
reaching end of first antennular article, about
twice as long as wide at base, without distinct
Carina, separated from orbital border by shal-
low rounded concavities. Orbital border pro-
jecting forward, rounded tips 0.7 as long as
rostrum. Orbital hoods inflated.

Antennular peduncle short and thick, with
second article about 0.7 length of visible por-
tion of first and slightly more than 0.7 length
of third; second article about 1.3 times as
broad as long. Stylocerite with tip of spine

reaching beyond middle of second article.
Outer flagellum not bifurcate.

Basicerite of antenna very heavy, reaching
beyond end of first article of antennular pe-
duncle, 0.3 thicker than long. Lateral spine of
basicerite acute, extending to tip of stylocer-
ite. Scaphocerite with squamous portion
broad and rounded, 0.8 as long as lateral
spine; tip of lateral spine not reaching end of
third article of antennular peduncle. Carpocer-
ite heavy, subconical, almost 0.3 longer than
antennular peduncle.

Labrum massive. Incisor process of mandi-
ble expanded, covering inferior portion of
labrum and bearing numerous strong teeth;
mandibular palp also expanded, with strong
bristles in addition to teeth similar to those on
incisor process. Maxillule and maxilla normal.
First maxilliped with endite of basipodite ex-
tended and contours rounded; no visible ar-
ticulations in endopod. Epipodite of second
maxilliped enlarged. Third maxilliped as
shown in Figure 30c.

Large chela strongly compressed laterally,
twice as long as broad. Margins entire except
for several small rounded eminences on in-
ferior edge where bristles arise. Outer face
with 2 shallow grooves extending 0.25 of
length proximally, superior starting at point
of articulation of dactylus, inferior near socket
for piston of dactylus. Palm almost 4 times as
long as fingers. Dactylus laterally disposed,
upper margin strongly arcuate and closing
across end of chela. Fixed finger very short,
with terminal indentation to accommodate
dactylus. Chela with scattered setae and bris-
tles distally on inner face and on proximal
portion of dactylus. Merus over 0.3 as long as
chela, 1.5 times as long as broad, inermous,
triangular in section, with inferior margins
rounded.

Small chela more than 0.7 length of larger,
fingers as long as palm. Palm 1.5 times as long
as high. Fingers curved, hooked tips crossing,
immovable finger with thin knife-like ridge
on inner edge. Distal portions of palmar edges
and fingers with scattered groups of short

Hawaiian Crangonidae — Banner

91

Fig. 30. Crangon nanus sp. nov. a, b. Anterior region, dorsal and lateral aspects; c, third maxilliped; d, e, large
cheliped, lateral and medial aspects; /, g, small cheliped, lateral and medial aspects; h, second leg; /, third leg;
y, third leg, dactylus; k, fifth leg; /, fifth leg, dactylus; m, telson and uropods {c-g from a paratype). {a, b, j, /, m,
scale A\ c, h, i, k, scale B; d-g, scale C.)

92

PACIFIC SCIENCE, Vol. VII, January, 1953

setae and bristles. Carpus slightly longer than
that of large cheliped and with rounded pro-
jection on inner distal edge. Merus similar in
form and proportions to that of large
cheliped.

Ischium of second legs almost as long as
merus. Carpal articles together 1.2 times
length of merus, with ratio 10 : 6 : 4 : 3 : 8.
Fingers of chela as long as palm. Entire ap-
pendage shorter than third legs.

Third legs without spines on ischium. Me-
rus tapering, inermous, 3 times as long as
wide. Carpus 0.4 as long as merus, without
spines but with both margins continued dis-
tally to form rounded tooth. Propodus arcu-
ate, 0.7 as long as merus, with 6 movable
spines. Dactylus biunguiculate with inferior
hook larger in diameter but slightly shorter in
length than distal hook. Fourth legs similar
but larger. Fifth legs with proximal 3 articles
shorter and of slightly different proportions;
propodus and dactylus equal to those of third
legs; brush of propodus of but 2 rows of bris-

tles; inferior tooth of dactylus feebly devel-
oped.

First pleopod of female reduced and with-
out articulations in both rami; endopod lack-
ing appendix interna; second and following
pleopods with endopods bearing separate
broad lobes for appendix interna. Second pleo-
pods of male of peculiar form (see Fig. 31^).

Telson 1.7 times as long as broad, tip about
0.5 as broad as ba*se; margins shghtly convex;
dorsal spinules at 0.4 and 0.7 of distance from
anterior articulation.

Branchial formula without mastigobranch
on base of seventh thoracic appendage and
setobranch on eighth; pleurobranch of third
also lacking.

Mature individuals of both sexes ranged
from 6 to 10 mm. in length. Color in life not
recorded.

DISCUSSION: The 20 specimens of this spe-
cies show little variation. One peculiar tend-
ency, similar to that observed in C. paragra-
cilis, is for the anterior portion of the body to

Fig. 31. Crangon nanus sp. nov. a, Second pleopod of male; b, first pleopod of female; c, d, e, endopods of
second to fourth pleopods of female;/, anterior region of specimen showing slight protrusion of eyes; g, h, anterior
region of specimen showing extreme protrusion of eyes, {a-e, scale K\ f-h, scale B.)

Hawaiian Crangonidae — BANNER

93

be displaced, exposing the eyes as is shown in
Figure Five specimens of this species

show this condition to some degree. The pos-
sible causes of the condition are discussed
under C. paragracilis.

Normally occurring variations were slight.
In one specimen the orbital hoods are equal
to the rostrum in length. Slight differences in
the proportions of the carpal articles of the
second legs and in the proportions of the ar-
ticles of the third legs were observed but all
articles, exhibited a generally consistent if not
exact relationship one to another. Finally, the
articles of the antennal peduncle between the
basicerite and the carpocerite are or are not
visible in lateral view, the differences evident-
ly caused by the degree of rotation of the car-
pocerite.

relationship: Because of the close rela-
tionship this species bears to C. paragracilis
(Coutiere) it has been placed in the Macro-
chirus Group. According to de Man’s key
(1911) this species is a relative of C. macro-
chirus (Richters) . However, it does not appear
to be closely related when the characteristics
of the latter species are considered, for macro-
chirus has deep orbitorostral grooves, the sec-
ond antennular article is three times the length
of the third antennular article, and the fingers
of the large chelae are proportionally larger.

Certainly, of all Hawaiian species this spe-
cies is most closely related to C. paragracilis
(Coutiere) and to the close relative of C. para-
gracilis, C. hawaiiensis Edmondson. As C. ha-
waiiensis is distinguished from C. paragracilis
principally in the form of the large chela, as
C. paragracilis is better known, and as C. para-
gracilis is probably the parent species to both
this species and C. hawaiiensis, C nanus is com-
pared chiefly to C, paragracilis. In the two spe-
cies the form of the anterior carapace is almost
exactly the same; the compact antennular pe-
duncle and the styloceiite are similar; the long
and heavy carpocerite is similar; the peculiar
mouthparts are similar; the general shape of
both the large and small chela are similar
(specific differences are noted below) ; the re-

lationship of the lengths of the carpal articles
of the second legs are roughly the same; even
the curvature of the propodus is similar in
both species. The branchial formula is similar
and evidently different from all other mem-
bers of this genus.

In fact, there is sufficient similarity between
the two species so that they could easily be
confused unless they were closely examined.
The chief differences lie in the following char-
acteristics: the large cheliped, in which the
chela has the inferior margin entire and the
merus unarmed in C. nanus, but with the chela
bearing a shoulder on its inferior margin and
the merus with spines on the inferior internal
margin and a spine on the superior distal
angle in C. paragracilis; the pleopods, which
are of the usual form in C. paragracilis and as
shown in Figure 31 in C. nanus; the merus of
the third leg which is unarmed in C nanus
and bears an acute tooth in C paragracilis.
Furthermore, in C. nanus the scaphocerite is
shorter than the antennular peduncle, the me-
rus of the second leg is equal in length to the
first four carpal articles, and the inferior un-
guis of the dactylus of the third leg is normal-
ly about 0.75 as long as the distal unguis,
whereas in C. paragracilis the scaphocerite is
longer than the antennular peduncle, the me-
rus of the second leg is equal in length to the
first three carpal articles, and the inferior
unguis of the third legs is 0.25-0.3 the length
of the distal claw. The differences between
the two species appear to be constant.

There is one other description in the litera-
ture that appears to be of the same or a closely
related species, that of " Alpheus sp.? Metal-
pheus n. gen.?" of Coutiere {loc. cit.). There
seems to be some confusion about this spe-
cies, for while Coutiere states it shows rela-
tionship to C. paragracilis and considers it as
possibly identical to the unfigured C. rostra-
tipes (Pocock, 1890), he also considers it as a
representative of a new genus.

C nanus is similar to " Alpheus sp." in many
important respects: the form of the orbital
hoods, anterior carapace, antennules and an-

94

PACIFIC SCIENCE, Vol. VII, January, 1953

tennae, mouthparts (with the exceptions
noted below); small chela (the large chelae
were lacking in Coutiere’s specimens), second
and following legs, including the important
dactyli; and the peculiar pleopods of the
male and female. Even the tendency toward
protuberant eyes, noted above, was found in
Coutiere’s specimens, as in the plate the me-
dial portion of the eyestalks appears beyond
the anterior margin of the carapace.

C. nanus differs from Coutiere’s specimen in
the rostrum, which is poorly demarked lat-
erally even at the edge of the carapace and
which has its tip depressed in lateral view;
in the endopod of the third maxilliped, in
which the basal article is narrower, the second
article broader, and the third article bearing a
distal tuft of setae; and in the small chela,
which lacks spines on the lower margin of the
palm. In Alpheus sp. the rostral ridge is still
discernible between the orbital hoods, the
rostral tip is not depressed, and the palm of
the small chela bears five {}) spines on its in-
ferior margin. When the wide range of varia-
tion in the closely related C. paragracilis is
considered, these minor differences certainly
do not appear to be of specific worth.

The possibility suggested by Coutiere that
C. nanus is a synonym of C rostratipes (Po-
cock) can be ruled out. In the first place, Po-
cock’s specimen came from Eernando Noron-
ha in the Atlantic Ocean off Brazil; while this
does not exclude the possibility that the two
species are identical, it certainly reduces the
probability, for almost no species of this
genus found off the Atlantic coast of the
Americas is also found in the Indo-Pacific
area. Anatomically, several characters separate
the species in spite of the fact that Pocock
published only a short description without
plates : C. rostratipes is described as having the
rostrum "springing from center of a depres-
sion,’’ which certainly could not be applied to
the Hawaiian form; the articles of the anten-
nular peduncle are described as being equal
in length, whereas in C. nanus the third article
is the longest; the merus of the small chela is

described as having the superior margin pro-
duced into a "conspicuous tooth,’’ a descrip-
tion that would not suit the subacute projec-
tion of this species; finally, the articles of the
carpus of the second legs in C rostratipes have
the ratio 10 : 5 : 2 : 3 : 4, instead of 10 : 6 :
4:3: 8.

Returning to the contention of Coutiere
that his specimen may represent a distinct new
genus, he states:

Volume des fouets antennaires et de I’antenne, incom-
plete protection des yeux, volume du labre, forme tres
speciale des mandibules, du 2® maxillipede, du 3® max-
illipede, des pinces de la 1® paire (?), brievete de la
2® paire, reduction du nombre des epipodites, forme
tres speciale des pleopodes, tels sont les caracteres que
Ton pourrait invoquer pour la separation de I’A. rostra-
tipes et des formes affines. Le nouveau genre pourrait
recevoir le nom de Metalpheus s’il etait conserve.

The prime basis for separation, not empha-
sized in this paragraph, was the supposition
of Coutiere that the large chela (lost in his
specimens) was similar in form to the small
chela. This was an erroneous assumption.

Of these characteristics, all save the short-
ness of the second thoracic legs and the modi-
fied pleopods are found in C. paragracilis as
well as C. nanus. The similarity of the two
species, as pointed out above, is especially
marked in the peculiar mouthparts and bran-
chial formula. This relationship is so close
that C. paragracilis and C. nanus must be in-
cluded in the same genus. However, as C.
paragracilis shows definite affinities to other
members of the genus Crangon, I do not be-
lieve that the erection of the proposed genus
would be justified.

distribution: Most of the specimens of
this species came from coral heads in water
10-20 feet deep immediately outside the shal-
low portions of the reef. The few specimens
from near the intertidal zone were collected
at the following places: from algal holdfasts
at Kawela Bay and on the open coast east of
Koko Head, on Oahu; from coral at Kalama
Park and at Lahaina, on Maui. The specimens
from deeper water came from the following
localities: off Mokulua Islands, Hanauma
Bay, Waikiki Reef, and Nanakuli, on Oahu;

Hawaiian Crangonidae — BANNER

95

in the deeper water off Kalama Park, on Maui.

Coutiere did not record the locality in the
Indian Ocean where his specimens were col-
lected.

Crangon hawaiiensis Edmondson
Fig. 32 a-h

Crangon hawaiiensis Edmondson, Bernice P.

Bishop Mus., Bui. 27: 14, figs. 2a-i, 1925.

DESCRIPTION : Rostrum triangular with nar-
row acute tip, about 1.5 times as long as broad
at base, tip probably reaching to near end of
first antennular article. Orbital hoods prob-
ably subacute, pointed, and somewhat shorter
than rostrum. Orbitorostral grooves shallow.

First antennular article heavy, probably
with visible portion slightly longer than sec-
ond article. Second article about 1.3 times as
broad as long, somewhat shorter than third
article. Stylocerite reaching to or slightly be-
yond end of first antennular article. Lateral
spine of basicerite longer than stylocerite.
Lateral margin of scaphocerite slightly con-
cave; lateral spine 0.2 longer than squamous
portion; tip reaching beyond end of anten-
nular peduncle. Carpocerite heavy, reaching
about to end of scaphocerite.

Mouthparts similar to C. paragracilis and
C nanus (see pp. 96 and 90).

Large chela almost 2.5 times as long as
broad, strongly compressed but not twisted,
fingers occupying distal 0.3. Palmar surfaces
smooth, rounded; slight concavity in inferior
margin below and proximal to articulation of
dactylus; faint groove on outer face of palm,
extending short distance proximad from
upper margin of dactylar articulation. Merus
1.8 times as long as maximum breadth; in-
ferior external margin rounded and not def-
inite; superior margin terminating in acute
tooth. Scattered setae on inner face of chela
and on inferior internal margin of merus.

Small chela lost.

Carpal articles of second legs with ratio
10 : 5 : 5 : 5 : 7; second article 1.1 times as
broad as long.

Ischium of third legs 0.5 length of merus,
unarmed; merus 3.3 times as long as broad,
with small acute tooth or smaller obtuse pro-
jection; propodus 0.3 as long as merus, almost
twice as wide distally as proximally, with
neither inferior nor superior margins project-
ing but terminating at approximately right
angles without spines or teeth; propodus 0.9
length of merus, curved, with 6 small spines;
dactylus biunguiculate, curved and heavy, less
than 0.4 length of merus, 2.2 times as long as
broad at base, with distal unguis longer but
less broad at base than inferior unguis.

Sole specimen known female 12 mm. long.

DISCUSSION: The type specimen was re-ex-
amined and redrawn for this study. Unfor-

Fig. 32. Crangon hawaiiensis 'E.dmondson. a, Anterior
carapace, dorsal aspect (the right orbital hood is folded;
the left is badly crumpled and its restored outline is
indicated by the dashed line); b, c, antennule and an-
tenna, dorsal and lateral aspects; d, large cheliped; e,
second leg; /, third leg; g, third leg, dactylus; h, fifth
leg, propodus and dactylus. {a-c, g, scale A; d-f,
scale B.)

96

PACIFIC SCIENCE, Vol. VII, January, 1953

tunately the type not only lacks the small
chela, but the chitin of the anterior thoracic
regions is so soft and distorted that the true
nature of the orbital hoods could not be as-
certained.

Careful comparison of the specimen to the
original description and plate of Edmondson
showed only two minor differences: first,
there was a faint longitudinal groove on the
palm of the large chela that was not previous-
ly noticed; second, the inconsistency of the
specimen as to the armature of the merus of
the third legs was not remarked upon. On this
last point the omission is probably of no great
moment, for in other species it has been ob-
served that occasional specimens may lack a
tooth on one or more legs, as this one does,
yet the majority of the specimens will have it.

Of the Hawaiian species this stands mid-
way between C. paragracilis (Coutiere) and C.
nanus Banner. It is like both species in the
form of the rostrum and orbital hoods (prob-
ably), in the mouthparts, in the general form
of the antennular and antennal bases, in the
general form (but not the specific sculptur-
ing) of the large chela, in the general form of
the third legs, especially in the curved pro-
podus and the shape of the biunguiculate dac-
tylus. It is like C. paragracilis in the length of
the scaphocerite and in the tooth on the merus
of the third legs, although in this species it is
more poorly developed. It is like C nanus in
the heavy form of the basicerite although not
as heavy, in the breadth of the articles of the
carpus of the second legs, and in the lack of
the deep sculpturing of the large cheliped.

However, it can be easily distinguished
from these two species, as the lower concavity
of the chela of C. hawaiiensis is slight and
gradual, whereas in C. paragracilis it is delim-
ited posteriorly by a lobe, and in C. nanus the
entire depression is lacking; the second article
of the carpus of the second legs is broader
than long instead of slightly or markedly
longer than broad, as it is in the other two
species. From C. paragracilis it may be dis-
tinguished also by the lack of spines on the

inferior internal margin of the merus, by the
shorter spine of the stylocerite, and by the
ratio of the articles of the carpus of the second
legs. It may also be distinguished from C.
nanus by the shorter stylocerite and the longer
scaphocerite.

Schmitt (1939) has described a new subspe-
cies, C. hawaiiensis clippertoni from Clipperton
Island in the eastern Pacific. It was distin-
guished from the type by two characteristics;
the supposed lack on the type of longitudinal
grooves on the palm and by the rostrum
which in Schmitt’s specimen did not reach
beyond the orbital hoods. The first of these
differences is already removed, and it is likely
that, if this species shows variation similar to
that of C. paragracilis, the other difference will
be within the range of variation for the spe-
cies. However, until such a time as more speci-
mens are examined, it would be best to leave
the subspecies as it now stands.

DISTRIBUTION: The type locality for this
species is Lisianski Island, in shallow water.

Crangon paragracilis (Coutiere)

Fig. 33 a-h

Alpheus paragracilis Coutiere, Soc. Ent. de
Erance, Bui. (7): 149-152, 1898a.

Alpheus paragracilis Coutiere, Fauna and Geog.
Maid, and Laccad. 2(4): 883, pi. 76, fig. 22,
1905. [Description same as original; figures
added.]

DESCRIPTION: Carapace produced into short
acute rostrum almost reaching end of first ar-
ticle of antennular peduncle.- Rostrum and
carapace rounded dorsally, without carina.
Shallow, rounded depression between base of
rostrum and orbital hoods. Orbital hoods
rounded, without processes.

Stylocerite acute, extending to middle of
second article of antennular peduncle. Pedun-
cle proper short and heavy, with second arti-
cle shorter than visible portion of first or third.
Lateral spine of scaphocerite not over 0.2
longer than squamous portion, as long as car-
pocerite, definitely longer than antennular pe-

Im m

1 Zmm

j I mm
0*5mm

Fig. 33. Crangon paragracilis (Coutiere). a, b. Anterior region, dorsal and lateral aspects; c, d, large cheliped,
lateral and medial aspects; e, small cheliped; /, second leg; g, third leg; h, third leg, dactylus. {a, b, scale A; c-e,
scale B; /, g, scale C; h, scale D.)

Hawaiian Crangonidae — BANNER

98

PACIFIC SCIENCE, VoL VII, January, 1953

duncle. Outer margin of scaphocerite some-
what concave. Lateral spine of basicerite ex-
tending beyond rostral tip.

Large chela compressed, slightly over twice
as long as high. Palm with 2 indentations on
outer face, longitudinal groove behind articu-
lation of dactylus and vertical shoulder on in-
ferior side slightly proximal to articulation.
Distal end of chela rotated almost 45° to
proximal end, dactylus opening somewhat
laterally. Fingers slightly less than 0.3 length
of palm. In addition to usual bristles on im-
movable finger and dactylus, row of several
groups of bristles on inner face near superior
margin, occasional bristles on inner face of
chela. Merus with superior edge produced as
acute tooth, with series of 8-10 very short,
conical spines along inferior internal margin.

Small chela about 0.75 as long as large
chela, slender; fingers curved, about as long
as palm. Propodus with acute tooth at point
of articulation of dactylus. Merus similar in
form to that of large chela.

Carpal articles of second legs with ratio
10 : 7 : 4 : 5 : 7.

Third legs with merus over 3 times as long
as broad, with small acute tooth distally on
inferior margin; carpus with distal projections
of superior and inferior margins poorly devel-
oped, rounded; propodus shorter than merus,
strongly arched, with 6-10 strong movable
spines along inner edge; dactylus biunguicu-
late, with both claws strong and acute, distal
claw twice as long as and broader at base than
inferior.

Branchial formula without mastigobranch
on seventh thoracic appendage, without seto-
branch on eighth appendage, without auxil-
iary pleurobranch at base of third maxilliped.

Largest specimens reaching 17 mm. in
length; body with bright blue and smaller red
chromatophores, blue predominating on
body, especially on antennular and antennal
bases, chelae, and caudal fan; tips of chelae
yellow- green; eggs green.

DISCUSSION: This species shows some vari-
ations but they are not as marked as they are

in some other members of the genus. Like
most species studied, there is some variation
in the proportions of the appendages, for ex-
ample, the first 2 carpal articles of the second
legs which vary in ratio from 10 : 5 to 10 : 7.
The tooth on the merus of the third legs varies
from a strong and acute tooth to a less well-
developed tooth with the point approaching
an obtuse angle. The squamous portion of the
scaphocerite varies from broadly rounded as
shown in Figure 33^ to narrow as shown by
Coutiere for the type (1905, fig. 22).

When this variation is considered, there is
only one minor difference apparent between
these specimens and those described by Cou-
tiere, the difference being that the propodus
of the third legs in the type bears only four
weak spines instead of the 6-10 in the Ha-
waiian form.

A peculiarity of this and the related species,

C. nanus Banner, is that these species, seem-
ingly alone among the local crangonids, can
have their eyes forced out from under their
carapace. In the collection of both species
available about 5 per cent of the specimens
showed this condition, which is evidently
caused by rough handling in collecting, a
thing very easily done with specimens so
small. Either too much pressure on the thorax
or too much pull on the antennules and an-
tennae will displace the anterior portion of the
body so the eyes will be exposed partially or
completely (see Fig. ^if-h for C. nanus). As
there is usually no distortion of the body or
the appendages accompanying this protru-
sion, the condition could be confusing to an
inexperienced worker.

DISTRIBUTION : This is one of the most com-
mon species about the Islands and has been
collected from most of the seaward reefs of
Oahu. It has been found in heads of coral, in
coralline algae, and in the holdfasts of algae.
Specific localities on Oahu where it has been
collected, from water up to 20 feet deep, in-
clude: Kaneohe Bay, Waimanalo, Hanauma i
Bay, Black Point, Waikiki, Nanakuli, Moku- I
leia, Kawela Bay. On Maui, at a similar depth i

Hawaiian Crangonidae — BANNER

99

rangCj it was collected at Makena, Kalama
Park, Lahaina, and four localities ranging
from 12 miles south of Lahaina to 10 miles
north of Lahaina. The only record of the spe-
cies from deeper water was a collection from
180 feet off Kaena Point, Oahu, where it was
living in sponges. Edmondson (1925) has re-
ported it from French Frigate Shoal, Pearl and
Hermes Reef, Laysan, Lisianski and Ocean Is-
lands. The "Albatross” took only one group
of specimens, those from Laysan Island, evi-
dently from shallow water.

The species has been reported from Johns-
ton and Wake Islands (Edmondson, 1925)
and from Tahiti in the central Pacific, and
from the Indian Ocean. It was not reported
by de Man from the Siboga Expedition ma-
terial, although the range of the species ex-
tends to either side of the area investigated by
that expedition.

Crinita Group, Crinita Subgroup

' Crangon paralcyone (Coutiere)

Fig. 34 a-i

Alpheus paralcyone Coutiere, Fauna and Geog.
Maid, and Laccad. 2: 895, pi. 80-81, figs.
34a-h, 1905.

Crangon laysani Edmondson, Bernice P. Bish-
op Mus., Bui. 27: 17, figs. 3a-j, 1925.
Crangon hucephalus (Coutiere), Edmondson,
Bernice P. Bishop Mus., Bui. 27: 14, 1925
[at least in part].

description: Rostrum not reaching be-
yond middle of visible portion of first anten-
nular article, broad yet acute, with dorsal Ca-
rina continuous from tip to behind orbital
hoods. Orbital hoods with no trace of teeth,
depressed anteriorly as flattened shelf expand-
ing as broad area between base of rostrum and
anterior portion of orbital hoods; frontal mar-
gin of this area varying from arcuate to S-
shaped. Margin of carapace above base of an-
tenna higher than at base of the orbital hoods,
forming slight but distinct depression.

Antennular peduncle short, with second
article somewhat longer than visible portion

of first, about twice as long as third (note that
the elongation of the second article shown in
Figure 34^ is atypical) . Stylocerite short and
variable with lateral spine not reaching end of
first antennular article. Basicerite with acute
lateral spine of varying size, usually quite
small. Scaphocerite relatively narrow, with
strong lateral spine 1.3 times as long as squa-
mous portion; lateral margins concave. Lat-
eral spine of scaphocerite usually longer than
antennular peduncle, usually greatly exceeded
by carpocerite.

Large chela subcylindrical, tapering, with-
out grooves or lobes; distal portion slightly
bent, thus lower surface concave in profile.
Fingers about 0.3 length of entire chela. Dac-
tylus laterally disposed, with strongly arcuate
margin. Distal portion of inner face of chela
sometimes sparsely covered with setae. Carpus
of usual form. Merus short, triangular, with
superior margin ending distally as rounded to
subacute projection, inferior internal margin
with acute tooth distally.

Small chela short, approximately 3 times as
long as broad. Fingers 0.6 as long as palm,
broad, flattened, lined with dense setae in
male, narrower and conical in female. Palm
somewhat triangular in section. Merus similar
to that of large chela.

Carpus of second legs variable, but usually
with ratio 10 : 30 : 9 : 9 : H.

Ischium of third legs with small blunt spine
(not shown in Fig. 34/?). Merus about 3.5
times as long as broad, with strong acute
tooth on distal portion of inferior border.
Carpus with 1-5 movable spines on inferior
border, continued distally as acute tooth; su-
perior margin with numerous long setae. Pro-
podus with varying number, usually about 12,
movable spines along inferior margin, most of
them in pairs, with many long setae on su-
perior margin. Dactylus short, 0.2 as long as
propodus, with small secondary unguis ro-
tated so as to be almost invisible, except when
legs are turned slightly. Fifth legs with
"brush” well developed.

100

Pleura of first abdominal segment showing
sexual dimorphism, in male ending posteri-
orly as dentate process that points slightly
posteriorly, in female rounded. Following
pleura of usual shape.

Telson 1.7 times as long as broad, tip 0.7 as
wide as base; sides straight and posterior mar-
gin somewhat arcuate; dorsal surface slightly
depressed medially. Dorsal spinules large, pos-
terior pair located near middle; posterior mar-
gin usually with series of about 10 very feeble
spinules. Inner branch of uropod with strong
spines on posterolateral margin.

Most specimens were 10 mm, long or less;
however, those specimens from Kaneohe Bay,
Oahu, listed below reach 17.6 mm. in length.
Color in life not observed.

DISCUSSION: This species, as well as the two
other members of this subgroup from Hawaii,
has been found to vary considerably in char-
acteristics commonly accepted as of taxonom-
ic importance.

The ratio of the lengths of the articles of the
antennular peduncle varies considerably, with
the second article varying from 1.4 to 2.0
times the length of the visible portion of the
first article. The stylocerite varies from mark-
edly shorter than the first antennular article to
almost equal in length.

The relative lengths of the antennular pe-
duncle, the scaphocerite, and the carpocerite
also vary. In some specimens the scaphocerite
is scarcely longer than the antennular pedun-
cle, and in turn is greatly exceeded by the car-
pocerite; in others the scaphocerite is almost
equal to the carpocerite, and both extend far
beyond the end of the antennular peduncle;
although the carpocerite is always longer than
the antennular peduncle, the amount by which
it exceeds is variable. The lateral spine of the
basicerite also varies in size and in some speci-
mens is so reduced that it is difficult to dis-
cern.

The large cheliped is constant in its general
form in all specimens but varies somewhat in
its proportions; in one specimen the tooth of

PACIFIC SCIENCE, Vol. VII, January, 1953

the inferior internal margin is reduced and

rounded (Fig. 34^).

As noted by Coutiere, the small chela is
sexually dimorphic, but the breadth of the
dactylus of the male is greater in proportion
in larger specimens, and in the largest speci-
mens it is broader than that described by Cou-
tiere.

In the second legs the second carpal article
is usually nearly three times the length of the
first, but it varies more or less continuously
from 2.2 to 4.0 times as long. The proportions
of the other articles also vary but not as much.

The spines on the carpus of the third legs
usually are two in number but vary from one
to five. The spines along the inferior margin
of the propodus vary from 8 to 13. Studies
were not made on the size of the secondary
unguis of these legs because it was so difficult
to rotate the leg to where the full unguis could
be seen; however, they, too, appear to vary,
being quite reduced in some specimens.

In none of the specimens examined does
the sixth abdominal somite bear spines. The
telson was not measured over a series of in-
dividuals, but examination did not show that
it varies greatly in general shape. How vari-
able the spinules on the tip of the telson are
could not be determined, as at best they are
exceedingly difficult to discern. In some speci-
mens they may have been missing.

This form evidently has been described
under two names: as C. paralcy one (Coutiere)
from the Maldives and Laccadives in the In-
dian Ocean and as C laysani Edmondson from
the Hawaiian Archipelago. The two species
appeared to be separate and valid on the basis
of the original descriptions, but when the
variation within the form is considered it ap-
pears that there are no important differences.

The original descriptions of the two species
showed the following characteristics in com-
mon: the length of the stylocerite; the shape
of the front of the carapace except for the ros-
trum; the shape of the large and small cheli-
peds; the shape and armature of the third legs;
and the shape of the telson. On the basis of

Hawaiian Crangonidae — Banner

101

Fig. 34. Crangon paralcyone (Coutiere). a, b. Anterior region, dorsal and lateral aspects; c, basicerite, showing
tooth (same specimen); d, large cheliped, lateral aspect; e, large chela, ventral aspect; /, small chela; g, second
leg; h, third leg; /, third leg, dactylus. (a-c, scale A; d, scale f-h, scale C; /, scale D.)

102

PACIFIC SCIENCE, Vol. VII, January, 1953

other characteristics, it appeared that the two
forms might be separated. For example, the
relative lengths of the scaphocerite to the an-
tennular peduncle and carpocerite; the relative
length of the second article of the second
thoracic legs, described by Coutiere as 2.43
times the length of the first and by Edmond-
son as 3.0 times the length of the first; the
armature of the carpus of the third legs; and
the presence of spinules on the posterior bor-
der of the telson. However, all of these char-
acters are found to be variable and the differ-
ences between the Indian and the Pacific forms
removed by the variation within the Hawaiian
population.

There are several differences that seem to
remain between this form and the form de-
scribed by Coutiere: first, in the Indian Ocean
form, the rostrum is but slightly produced,
whereas in this form it usually reaches the end
of the first article of the antennular peduncle;
however, Coutiere has shown in his plates that
this varies in his specimens, and similar varia-
tion has been seen in the local specimens. Sec-
ond, the lateral spine of the basicerite is longer
in Coutiere’s specimens than in the local speci-
mens; however, in the latter it is quite vari-
able. Third, the merus of the large chela was
shown by Coutiere without spines or teeth
(although he described it as bearing ”une forte
epine a son apex infero-interne”) ; in the Ha-
waiian form there is a pronounced and almost
always acute tooth. The external spine of the
uropod is shown to be bent in Coutiere’s
plates, whereas in this form it is straight. Fi-
nally, de Man in his key (1911: 314) puts em-
phasis on the presence of two teeth on the
posterior dorsal margin of the sixth abdom-
inal segment, teeth that are not found in the
Hawaiian specimens; however, Coutiere re-
ported that, "... 2 prolongments epineux . . .
pouvant parfois manquer ou etre tres peu
marques."

Therefore, as all the differences between the
Hawaiian forms and the type described by
Coutiere are either bridged by the variations
noted or are both minor and variable so the

differences can be anticipated to be bridged
by the study of additional specimens, it is ad-
visable to place the species C. laysani as a syn-
onym of C. paralcyone.

The specimen from Laysan identified by
Edmondson (1925) as C. hucephalus (Coutiere)
was re-examined and found to be a member
of this species.

DISTRIBUTION: This species appears to be
more abundant in deeper water than in shal-
low water. It has been collected at the follow-
ing locations in water up to 20 feet deep: at
Kaneohe Bay, Hanauma Bay, Kahala Beach,
Waikiki Beach, off Nanakuli, on Oahu; at
Makena, Lahaina, and 10 miles south of La-
haina, on Maui; at Halape, Kau Coast, Ha-
waii, from coral at 35-40 feet deep. It was re-

TABLE 4

Deep-water Collections of Crangon paralcyone

LOCATION

DEPTH

U.S.N.M.

STATION CATALOGUE
NUMBER* NUMBER

NE of Hawaii

fathoms

29-26

4053

63623

26-50

4054

63619

24-83

4061

63608

Auau Channel

32-43

3872

63607

34-65

3875

63617

28-43

3876

63609, 63612,

Off Puaaloa, Maui

26-27

_

63625

N of Maui

45-52

4070

63626

49-57

4075

63611

S of Molokai

23-24

3847

63618

44-73

3848

63613

43-73

3849

63621, 63624

Penguin Bank

27-29

4032

63616

Oahu

Off Mokulua Island

20

_

_

1 mile S of Kahala

25

—

—

Off Diamond Head

35

—

—

Off Kaena Point

25

—

—

Off SW Oahu

6-60

—

—

Off Kauai

24-43

4024

63610

Off Kaula Rock,

SW of Niihau

35

_

_

Off Nihoa

26

4147

63622

27-29

4158

63614

Off French Frigate Shoal 14-17

3968

63615

17

3971

63620

* Station numbers are those of the "Albatross”;
collections without station numbers were made by the
"Makua” and "Salpa.”

Hawaiian Crangonidae — BANNER

103

ported from shallow water at Laysan as C lay-
sani and C hucephalus by Edmondson (1925).
It has been dredged at the locations given in
Table 4.

The species has been reported from various
locations in the Indian Ocean and from the
Netherlands Indies.

(?) Crangon paralcyone (Coutiere)

A single 9 millimeter specimen, collected at
Hanauma Bay, Oahu, in 20 feet of water, has
been referred to this species with doubts. It
agrees with C. paralcyone in the form of the
rostrum and the orbital hoods, the stylocer-
ites, the spine on the basicerite (on one side
only; the spine is lacking on the other side),
the relative lengths of the scaphocerite, car-
pocerite, and antennular peduncle; somewhat
in the form of the large and small chelipeds;
in the relative lengths of the articles of the
carpus of the second legs; and somewhat in
the form and armature of the third legs. It dif-
fers in the following points : the articles of the
antennular peduncle are broader; the dactylus
of the large chela is relatively shorter; the me-
ral tooth of the third legs is reduced to a very
short rounded protuberance, the spines of the
propodus are much longer, almost comb-like
in form.

Most of these differences could be individ-
ual differences in a variable species, the most
notable being the dactylus of the large chela
and the armature of the third legs which has
the reduced tooth on the merus and the longer
propodal spines. It is possible that these too
may be individual differences, or they may be
constant and adequate criteria to separate this
specimen as a species. However, with only
one such specimen, and with the parent spe-
cies showing such marked variation, it is con-
sidered advisable to place the specimen in C.
paralcyone, with reservations.

Crangon brevipes (Stimpson)

Figs. 35 a-j, 36 a-e, yi
Alpheus brevipes Stimpson, Acad, Nat. Sci.

Phila., Proc. 12: 30, I860.

NEOTYPE: A female 13.4 mm. long, col-

lected from a head of coral in 10 feet of water
off Nanakuli, Oahu. A male paraneotype 9-6
mm. long has also been designated from the
same locality (U.S.N.M. 93459).

DESCRIPTION: Anterior portion of carapace
with dorsal carina high and thin, ending
abruptly at base of rostrum. Rostrum short,
triangular, seldom reaching past anterior mar-
gin of orbital hoods. Orbital hoods hemi-
spherical, continued forward as short, sub-
acute, subvertical keel that slants mesally
toward ventral side; anterior margin of hoods
almost vertical when seen in profile, reaching
higher than rostral carina. Area between or-
bital hoods and base of carina flattened, with
anterior margin varying from concave to
straight. Lateral margins of carapace with in-
dentation between lateral margin of orbital
hoods and covering of base of antennal pe-
duncles.

Second article of antennular peduncle from
1.5 to 2.0 times as long as visible portion of
first article; third article about as long as first.
Stylocerite short, broad, lateral spine reaching
only 0.8 length of visible portion of first an-
tennular article. Basicerite unarmed. Scapho-
cerite narrow, tapering, squamous portion
about I.I-I.2 times as long as lateral spine;
lateral margin slightly concave. Scaphocerite
usually slightly longer than antennular pedun-
cles and slightly shorter than carpocerite.

Large chela without notches or grooves,
subcylindrical, tapering. Dactylus heavy, be-
tween 0.25 and 0.3 length of entire chela,
strongly arched dorsally. Merus about 1.5
times as long as broad, unarmed, distal end of
superior margin rounded. Chela sexually di-
morphic in both size and proportions, in fe-
male noticeably shorter than carapace, in male
longer than carapace, with relatively greater
thickness and shorter fingers than in female.

Small chela about 3 times as long as fingers,
latter narrow and conical in both sexes; in
male fingers about twice as long as broad, in
female about 3 times as long as broad. Carpus
0.3 length of chela. Merus 3 times as long as
broad but otherwise similar to large chela.

104

PACIFIC SCIENCE, VoL VII, January, 1953

Carpal articles of second legs variable, but
ratio usually within following range: 10 : 15-
21 : 5-7 : 5-7 : 9-12.

Third legs with spine on ischium. Merus
2. 7-3.1 times as long as wide, with strong sub-
apical tooth on inferior margin. Carpus with
acute tooth on distal end of inferior margin,
without movable spines. Propodus with 10
movable spines along inner margin. Dactylus
simple, curved, acute.

Pleura of abdomen rounded in both sexes.
Telson broad, flat, slightly tapering distally,
point of greatest width 1.3-1. 8 times as wide
as posterior margin, length 2. 7-2. 9 times
breadth of posterior margin; both pairs of
dorsal spines heavy, 0.3 and 0.6 of distance
from point of articulation.

In addition to usual setiferous bristles pos-
terior margin of telson with about 10 small
spinules, these difficult to discern. Inner
branch of uropod also with series of strong
movable spinules, about 20, along postero-
lateral margins.

Specimens reaching 16.0 mm. in length;
color in life not observed.

DISCUSSION: Inquiry at the major American
museums revealed no records of Stimpson’s
type specimen, so it is presumed to have been
lost. Unfortunately from Stimpson’s descrip-
tion it is not possible to determine the exact
type locality, for all that he specified was, "in-
sulas Hawaienses; inter ramos madreporar-
um." The neotype meets those specifications.

All the specimens available were examined
carefully, but a group of specimens from Wai-
kiki Reef which exhibited the usual range of
variation were studied in greater detail. In this
group, the variable characteristics were meas-
ured as accurately as possible and the results
are here summarized.

Anterior region of the carapace: The dorsal
Carina varied from broad and dorsally rounded
to narrow and acute; in no specimen did it
reach the anterior tip of the rostrum. The vari-
ation in thickness of the anterior ridge of the
orbital hoods may be seen in the drawings.
One of the most variable characteristics was

that of the anterior margin of the carapace be-
tween the ridges of the orbital hoods and the
rostrum; this varied from a concave margin in
which the tip of the short rostrum did not
reach the anterior margins of the orbital
hoods, to a straight margin with the rostrum
reaching beyond the hoods; in some speci-
mens the frontal region was asymmetrical.

Antennules and antennae: The observed range
in proportions of the articles of antennular
peduncle is reported in the description. The
relative length of the scaphocerite also varied,
but without exception it was no shorter than
the antennular peduncle and no longer than
the carpocerite. In all specimens the basicerite
lacked a lateral spine.

Chelipeds: Some of the variation of this sex-
ually dimorphic appendage is indicated in the
description. It was also found to differ in the
degree of compression, with the breadth al-
most equaling the height in some specimens
while in others the breadth was only about 0.7
of the height. The palm was found to vary
from 1.5 to almost 3.0 times the length of the
merus. In none of the specimens did the merus
have an acute tooth on the end of the inferior
internal margin, but although the margin was
straight in most cases, a few specimens were
found to bear a small obtuse and rounded pro-
jection.

The small cheliped was quite uniform ex-
cept for the slight sexual dimorphism de-
scribed above.

Second legs: The range of variation of the
carpal articles has been given; a further dis-
cussion of this variation is given below where
C. hrevipes is contrasted to C. clypeata (Cou-
tiere) .

Third legs: These appendages were quite
constant in armature and proportions, with
the ischium always bearing a spine; the merus
varying from 2.7 to 3.0 times as long as broad,
that of the smaller specimens being more nar-
row than the larger; the merus always with a
strong distal tooth; the propodus always bear-
ing strong spines; the dactylus always simple.
However, in the smallest specimens (those

Hawaiian Crangonidae — Banner

105

-> I mm
I mm
0.5mm

Fig. 35. Crangon brevipes (Stimpson). a. Anterior region, dorsal aspect (asymmetry of rostrum and scaphocerite
found in specimen); b, anterior region, lateral aspect; c, large cheliped, lateral aspect; d, large chela, ventral aspect;
e, large cheliped, merus and carpus, medial aspect; /, small cheliped, lateral aspect; g, small cheliped, merus and
carpus, medial aspect; h, second leg; i, third leg;/, third leg, dactylus. {a, b, scale A; c-i, scale B;/, scale C.)

106

PACIFIC SCIENCE, Vol. VII, January, 1953

Fig. 36. Crangon brevipes (Stimpson). Variation in
the frontal region of the carapace, antennules, and an-
tennae; specimens drawn in a, b, and c came from Wai-
kiki Beach, Oahu; the specimen in d and e came from
Mokuleia, Oahu, {a, b, scale A; c, scale B; d, e, scale C.)

about 5 millimeters long) the propodus bears
only 4-5 well- developed spines instead of the
usual 8-11 spines.

Telson: The measured range of variation of
the telson has been given. No attempt was
made to determine whether the small spinules
on the posterior border of the telson or on the
posterolateral margin of the inner uropod
varied in number, as it was too difficult to
count them accurately. In all specimens that
were studied carefully the spinules were pres-
ent.

These specimens agree perfectly with my
interpretation of the short description given,
without figures, by Stimpson; the description
is adequate to separate this species from all
other species known to occur in the Hawaiian
Archipelago.

Until this species was thoroughly studied,
it was felt that probably it was identical with

C. clypeata (CoutiUe), a species found in the
same habitat; if that had been so, C. clypeata
would have been relegated to synonymy. The
two species agree in most characteristics: in
the general shape and configuration of the
anterior portion of the carapace; the propor-
tions of the antennular peduncle and antennal
peduncle; in the large chela; in the third legs;
in the telson.

When a large number of specimens was
studied, however, the following characteris-
tics were found to be reliable for their separa-
tion:

1. Sexual dimorphism of the small chela of
males. In C. clypeata the dactylus of the small
chela of males, although somewhat variable,
is always noticeably broadened or subspatu-
late and surrounded by dense setae; in C. hrev-
ipes it is never subspatulate.

2. Tooth on hasicerite. This tooth, or spine,
is almost always present in C. clypeata and is
always lacking in C. brevipes\ however, in a few
of the specimens of C. clypeata it is very re-
duced, and sometimes it is entirely lacking.

3. Tooth on merus of large cheliped. This
tooth is usually large and acute in C. clypeata
and entirely lacking in C brevipes\ however,
this characteristic alone is not adequate to
separate the species, as sometimes in C.
clypeata it may be reduced and obtuse or
rounded, or even entirely lacking, whereas
in C. brevipes the margin may bear an obtuse
projection.

4. The relative length of the second article of
carpus of the second legs. In an attempt to find
the extent of variation in the relative lengths
of the first and second carpal articles, they
were measured in 103 specimens of both spe-
cies; the result of this study is shown in Figure
37. This shows that most of the specimens of
C. clypeata have the ratio of the second to the
first articles of 1.0 and that most of the speci-
mens of C. brevipes have the ratio of the same
articles between 1.7 and 2.0, However, speci-
mens that on the basis of the other character-
istics were definitely C. clypeata did reach the
maximal ratio of 1.3, and similarly specimens

Hawaiian Crangonidae — BANNER

107

that were definitely C. brevipes did reach the
minimal ratio of 1.4. Therefore, it is likely
that, if enough specimens were measured, the
edges of the normal distributional curve would
overlap. Another graph, not reproduced here,
was made to ascertain whether the ratio be-
tween the articles varied with the size of the
specimens; in this the ratio of the lengths of
the first two carpal articles was plotted against
the length of the carapace. This study showed
no correlation whatsoever, with both species
having great variation in ratios for all size
ranges.

On the basis of these four variable charac-
teristics, satisfactory separations of the two
species may be made. If, for example, the ratio
of the first two articles of the carpus of the
second legs was between 1.3 and 1.5, in cases
where it was undecisive, then the specimen
could be assigned to its correct species on the
basis of the teeth of the merus of the large
chela or of the basicerite. In none of the well
over 200 specimens examined was any doubt
encountered as to the identity of an individual
specimen.

The belief that these are indeed separate
species is confirmed by the study of paired
specimens. Both of these species live in algae-
covered tubes on old coral; in each tube there
is a single pair of specimens, a male and a
somewhat larger female which are evidently
mates. During routine collecting, whenever it
was possible to get both members of this pair,
they were removed and preserved for special
study. In the small number of these pairs
available, none consisted of mixed species—
either both were plainly C. brevipes or both
were plainly C. clypeata.

This endemic species apparently has been
derived from the Indo-Pacific C. clypeata. As
the differences between the species are neither
constant nor of great magnitude, as the hab-
itat remains the same, the separation of the
two species can be presumed to have been in
relatively recent time.

Coutiere described a species, C. parabrevipes
(1898^: 151), which he thought might be iden-

tical with this species. It was plainly a differ-
ent species as C parabrevipes had grooves and
ridges on its chela which are entirely lacking
in C. brevipes. C. parabrevipes (Coutiere) was
considered a synonym of C. acuto-femorata
(Dana) by de Man (1911).

DISTRIBUTION: The species appears to be
moderately plentiful on Oahu, where it was
collected both in shallow water of the reef flat
and outside the reefs in water to about 20 feet
deep at the following locations: off Mokulua
Islands, Waimanalo, Hanauma Bay, Waikiki,
Nanakuli, Mokuleia, Kawela Bay. It was col-
lected at Halape, Kau Coast, Hawaii, by R.
W. Hiatt, in coral at a depth of 30-40 feet.
The only record of the species from deep water
is two specimens collected in 30 fathoms off
Waikiki Beach.

Crangon clypeata (Coutiere)

Figs. 37, 38 a-k, 39 a-u

Alpheus clypeatus Coutiere, Fauna and Geog.
Maid, and Laccad. 2: 897-898, pis. 81-82,
figs. 36-36g, 1905.

DESCRIPTION: Frontal border, seen from
above, truncate and projecting, with margin
from slightly concave to slightly convex. Ros-
trum either small or lacking. Dorsal carina
high and definite, sharp or rounded, anteriorly
terminated abruptly before reaching edge of
frontal area. Area between orbital hoods and

■Ratio of second to first article
of carpus of second legs

Fig. 37. Comparison of carpal ratio of Crangon cly-
peata (Coutiere) and Crangon brevipes (Stimpson).

108

PACIFIC SCIENCE, VoL VII, January, 1953

rostrum convex dorsally. Orbital hoods hemi-
spherical. Depression between lateral margins
of orbital hoods and portion of carapace
above base of antennae.

Antennular peduncle with second article 1-
2 times as long as visible portion of first arti-
cle; third article equal to or somewhat longer
than visible portion of first; second article 1.5-

2.0 times as long as broad. Short lateral spine
of stylocerite reaching nearly 0.7 length of vis-
ible portion of first article. Basicerite almost
always bearing thin, acute lateral spine. Sca-
phocerite, antennular peduncles, and carpo-
cerite subequal in length, all variable, but car-
pocerite almost always slightly longer.

Large chela of male subcylindrical, usually
slightly compressed, entire and smooth, ta-
pering toward fingers. Entire chela about 4
times length of fingers and about 2.5 times
its greatest height. Chela rotated, with dac-
tylus lateral in position. Merus about 1.5
times as long as wide, with superior distal
angle not greatly produced; inferior internal
margin usually with strong acute tooth dis-
tally. Chela relatively shorter and thinner in
female than in male.

Small chela of male with dactylus expanded,
about twice as long as broad, with dense
fringe of short setae on margin; fingers slight-
ly shorter than palm; height of palm slightly
over 0.3 length of chela. Merus twice as
long as wide, about 0.7 as long as chela, with
all distal angles rounded. In female dactylus
narrow, 3 times as long as broad, uniformly
tapering; palm about 1.5 times as long as fin-
gers; chela slightly over 3 times as long as
high; merus thinner than in male.

Ratio of carpal articles of second legs vary-
ing thus: 10 : 9-13 : 2.9-4.7 : 2.9-4.7 : 6.3-

10.0 (see Fig. 37).

Ischium of third legs armed with short
strong movable spine. Merus 3. 1-3. 5 times as
long as wide, with strong tooth distally on
inferior margin. Carpus prolonged distally on
inferior border as rounded tooth, otherwise
unarmed. Propodus broad and flattened, with
9-11 movable spines along inferior margin.

Dactylus simple, curved. Fourth legs similar
to third legs except meral tooth less pro-
nounced.

Telson with sides of uniform taper 2. 2-2. 9
times as long as width of posterior margin,
greatest breadth 1.3-2. 2 times as wide as pos-
terior margin. Pairs of dorsal spines about 0.4
and 0.7 distance from articulation. Posterior
margin between posterolateral spines arcuate,
with about 10 small spinules; inner branch of
uropod similarly with spinules along distolat-
eral margin.

Mature specimens usually reaching maxi-
mum length of 18 mm.; one female, however,
23 mm. long. Color of live specimens variable,
body transparent, covered dorsally with red,
yellow, and blue chromatophores in reticu"
lated pattern; abdomen with reticulations in
transverse bands; legs usually transparent;
large chela with more red and blue chromato-
phores than body; eggs bright green.

DISCUSSION: Like C. parakyone and C. brevP
pes, this species shows great variation in many
characteristics. Some 10 specimens were
studied intensively to determine the range of
variation; unfortunately, time did not permit
the making of the complete series of measure-
ments for the almost 100 specimens available.
The specimens not measured, however, were
all examined to determine any marked varia-
tion.

The ranges of variation of the frontal region
of the carapace, of the antennular and antennal
peduncles, of both chelipeds, of the third legs,
and of the telson either have been given in the
description or are shown in Figures 38 and 39-
The range of variation of the second legs is
given in the description and shown graphical-
ly in Figure 37. A few additional variations are'
noteworthy: The lateral spine of the basicer-
ite, usually well developed, is sometimes re-
duced or absent. The large chela, already
noted as sexually dimorphic, also varies from
almost circular in cross section to distinctly
compressed. The merus of the large cheliped
almost always carries a well- developed tooth
on the inferior internal margin, but in some

Hawaiian Crangonidae — BANNER

1 2m

1 2mm

Fig. 38. Crangon clypeata (Coutiere). a, b, Anterior region, dorsal and lateral aspects; c, large cheliped, male,
lateral aspect; d, large cheliped, merus and carpus, medial aspect; e, small cheliped, male; /, dactylus of e showing
maximum breadth; g, small cheliped, female; h, second leg; /, third leg; j, third leg, dactylus; k, telson and uro-
pods. {a, b, e-i, k, scale A; c, d, scale B;/, scale C.)

no

specimens it is reduced and rounded and in a
very few completely absent. On the third legs
of very small specimens (less than 5.0 mm.
long) there are only 4-5 spines on the pro-
podus instead of the usual 9-11-

The observed range of variation of this spe-
cies is great, and if a few specimens on the
fringes of the normal curve of variation were
examined alone they probably would be inter-
preted as separate but related species. How-
ever, in the large number of specimens exam-
ined, the gaps between the extremes were
closed so the collection presented an almost
continuous spectrum of variation from one
extreme to the other.

The description of the type specimen of
Coutiere easily falls within the range of varia-
tion noted above; there should be no doubts
as to the specific identification of this species.

Locally this species can easily be confused
with C. hrevipes\ the separation of the two spe-
cies is discussed in detail on page 106.

DISTRIBUTION: This species is usually found
in the same habitat as C. hrevipes\ like C. hrevi-
pes, it is moderately common around Oahu.
In the shallow water of the reef surface and on
the outer sides of reefs to the depth of about
20 feet it has been collected at the following
localities: off Mokulua Islands, Waimanalo,
Hanauma Bay, Waikiki, Nanakuli, Mokuleia,
on Oahu; at Lahaina and at 10 miles north of
Lahaina, on Maui. Edmondson (1925) has re-
ported this species from French Frigate Shoal,
Pearl and Hermes Reef, and Ocean Island.
One chela, collected by the ’’Albatross” at
Honolulu — without further designation —
probably is this species (U.S.N.M. 63633).

Edmondson has also reported the species
from Johnston and Wake Islands. Its original
description was based on specimens from the
Indian Ocean, but it was not collected by the
Siboga Expedition.

Crinita Group, Diadema Subgroup
Crangon paracrinita (Miers)

Alphetis paracrinitus . Miers, Ann. and Mag.

Nat. Hist. V, 8: 365, pi. 16, fig. 6, 1881.

PACIFIC SCIENCE, Vol. VII, January, 1953

As I have seen no specimen of this species,
the original description is given.

Rostrum triangular, acute, arising from the frontal
margin of the carapace (which is slightly concave on
each side of its base), but not prolonged backward as
a dorsal carina. Orbital arches entire, arcuated, without
spinules; anterior margin of the carapace sinuated on
the sides, without spines. Postabdominal segments
smooth, entire, with the lateral margins broadly
rounded; terminal segment not three times as long as
broad at the base, with its distal end subtruncated.
Eyes completely concealed beneath the carapace. An-
tennules with three joints of the peduncle exposed, of
which the middle one is slightly the longest, with a
small spine-like scale at base, reaching nearly to the end
of the basal joint. Basal scale of antennae about reach-
ing to the end of the antennal peduncle, with the outer
margin convergent towards it and clothed with long
hairs. Anterior legs or chelipeds having the merus and
carpus slender; merus with a small tooth or spine at.the
distal end of its under margin; palm of larger cheliped
rather more than twice as long as broad, smooth, with-
out notches or ridges, largest at its rounded basal end,
with an impressed curved line on its upper and proxi-
mal end; fingers nearly half as long as the palm; the
upper with its superior margin arcuated. Smaller cheli-
ped with the carpus rather longer, and chela very slen-
der, fingers hairy. Second legs with the first joint of the
carpus longer than the second, the last three joints of
nearly equal length, the last a little the longest, the
joint preceding these somewhat longer. Ambulatory
legs somewhat hairy. Distal ends of the rami of the
uropods clothed with long hairs. Color light yellowish
(in spirit). Fingers of larger cheliped pinkish. Length
7 lines (nearly 15 millim.).

Only one specimen has been reported from
the Hawaiian Archipelago, that by Edmond-
son (1925) from Laysan Island.

Elsewhere in the Pacific the species has been
recorded from Johnston and Palmyra Islands
by Edmondson. It has also been reported from
Djibouti in the Indian Ocean. The type local-
ity is Senegambia, West Africa.

Crangon paracrinita (Miers) var.
bengalensis (Coutiere)

Fig. 40 a-k

Alpheus paracrinitus var. bengalensis Coutiere,

Fauna and Geog. Maid and Laccad. 2: 901,

pi. 32, figs. 37-37e, 1905.

DESCRIPTION: Rostrum triangular, acute,
longer than width at base, tip reaching ap-
proximately to middle of visible portion of
first antennular article, rounded dorsally, not

Hawaiian Crangonidae — Banner

111

Fig. 39. Crangon clypeata (Coutiere). Showing variation in front of carapace, antennules, antennae, and cheli-
peds. {a, b, v, are of one specimen; c, d,j-l, of another; e, m-o, of a third; /, g, p-r, of a fourth; h, s-u, of a fifth.)
{a, b, e, i-l, v, scale A.\ c, d, /, g, scale B; h, scale D; m-u, scale C.)

112

carinate, separated from anterior orbital hoods
by short shallow round-edged depressions.
Orbital hoods not conspicuously inflated,
rounded anteriorly. Frontal margin between
orbital hoods and base of rostrum concave.

Visible portion of first antennular article
subequal to second; third article shortest. Tip
of stylocerite reaching beyond tip of rostrum,
almost to end of first antennular article. Lat-
eral spine of scaphocerite about 1.1 times as
long as squamous portion. Basicerite with
acute spine. End of carpocerite greatly exceed-
ing both scaphocerite and antennular pedun-
cle; scaphocerite equal to or slightly longer
than antennular peduncle.

Large chela compressed, with smooth regu-
lar margins and faces except for very slight
concavity in upper margin proximal to articu-
lation of dactylus and longer, more gradual
concavity in lower margin below articulation
of dactylus. Chela 3 times as long as broad,
fingers about 0.3 length of chela. Finger 1.5
times as long as greatest height. Merus of
large cheliped 2.0 times as long as broad, su-
perior edge distally rounded, inferior external
edge rounded in basal portion, inferior inter-
nal edge with small acute tooth distally.

Small chela subcylindrical, almost 5 times
as long as broad, fingers 0.5 length; shoulders
at articulation of dactylus rounded. Carpus
elongate, slightly over 0.3 length of chela.
Merus 3.4 times as long as broad, similar in
form to that of large cheliped but with inferior
internal spine either poorly developed or
lacking.

Both large and small chelae sexually dimor-
phic in size, large chela of male 1.3 times
length of carapace, equal in female. Small
chela with similar range; in addition, dactylus
of male broader than that of female and bear-
ing heavy fringe of bristles.

Carpal articles of second legs with ratio
10 : 10 : 5 : 5 : 8.

Ischium of third legs with movable spine.
Merus 5 times as long as wide, rounded dis-
tally (in specimen drawn only left leg rounded
distally, that of right leg with small acute

PACIFIC SCIENCE, Vol. VII, January, 1953

process distally). Carpus slightly over 0.5 as
long as merus with neither distal angle greatly
projecting or acute. Propodus about 0.8 as
long as merus, with 6-8 spines. Dactylus
simple, curved and elongate, 0.3 as long as
propodus.

Abdominal pleura of both sexes rounded.
Telson 1.7 times as long as broad at base, 1.7
times as broad near base as at tip. Proximal
0.6 of telson broad, only slightly tapered; dis-
tal 0.4 abruptly tapered anteriorly, gradually
tapered posteriorly. Posterior margin almost
straight. Paired dorsal spines 0.3 and 0.6
of length from base.

Length up to 14 mm. Body usually trans-
lucent white with definite narrow transverse
bands of red on abdomen and somewhat simi-
lar bands, poorly defined, on thorax; bands
across bases of antenna and antennules more
definite; chelae with red mottling; eggs dark
yellow in early stages, more greenish in ad-
vanced stages.

DISCUSSION: No marked variation in the
specimens in the collection was noted, al-
though a detailed study of the ratios of the
parts of the appendages was not undertaken.
Slight differences were noted, as would be ex-
pected, in the proportions of the antennules,
antennae, large and small cheliped (especially
in the length-width relations of the merus,
which varied from 1.8 to 2.7 times as long as
broad), in the merus of the third legs, and in
the telson. It should be noted that in the 26
specimens of this variety in the collections the
second article of the carpus of the second legs
varied from 0.85 to 1.3 times as long as the
first. The merus of the large cheliped was
armed with teeth of varying size on the in-
ferior internal margin, and the corresponding
margin of the small cheliped often had a
feebly developed tooth. However, in general
the specimens were quite similar.

This variety was separated from the parent
species by Coutiere on the basis of three char-
acteristics : first, the parent species has a small
tooth at the articulation of the dactylus of the
small cheliped, whereas the variety has none.

Hawaiian Crangonidae — BANNER

Fig. 40. Crangon paracrinita (Miers) var. hengalensis (Coutiere). a, b, Anterior region, dorsal and lateral aspects;
c, large cheliped, lateral aspect; d, large chela, medial aspect; e, large cheliped, merus and carpus, medial aspect;
/, small cheliped, lateral aspect; g, small chela, ventral aspect; h, second leg; /, third leg (meral tooth abnormal);
y, third leg, meral-carpal articulation normal, from same specimen as i; k, telson. {a, b, scale A; c-e, h-j, scale B;
/, g, scale C; k, scale D.)

j I m m

I 2 mm

j 2 mm

1 1 m m

114

PACIFIC SCIENCE, Vol. VII, January, 1953

Second, the tooth on the inferior internal mar-
gin of the merus of the large cheliped is re-
duced, and the one at the corresponding loca-
tion on the small chela is lacking in the va-
riety, whereas both are present and well devel-
oped in the type described by Miers. Finally,
the first carpal article of the second legs is al-
most twice as long as the second in C. para~
crinita (Coutiere: . rapport de 1.75 a

1.85 . . loc. cit.), whereas in the variety the
two articles are of almost equal length.

The differences between the Hawaiian
specimens and those described by Coutiere
are slight except for the variations noted.

All specimens collected in the windward
Hawaiian Islands have been assigned to this i
variety. It is true that the armature of the me- j
rus of the chelipeds has been overlooked in
making this separation, but first, the variation ,
of the armature indicates that too much faith
should not be placed upon it; second, the Ha-
waiian forms agree with the variety in the ratio
of the articles of the carpus of the second legs
— at least, they lie within the expected range
of variation— and none had the tooth flanking
the articulation of the dactylus of the small
chela. If the variation in these two forms is
similar to that found in C. brevtpes (Stimpson)

TABLE 5

Characteristics Differentiating the Species of the Diadema Subgroup

CHARAC-

TERISTIC

C. pugnax

C. diadema

C. pseudopugnax

C. percyi

C. gracilipes

C. paracrinita
var.

hengalensis

Rostral base

Very narrow
triangle
without Ca-
rina; not
depressed

Broadly trian-
gular with Ca-
rina; some-
what de-
pressed

Simple ridge
anteriorly,
triangular
posteriorly;
strongly de-
pressed

Slightly broad-
er than in
pugnax; with-
out Carina
anteriorly;
strongly de-
pressed

Similar to

pugnax

Rounded,
not well
marked;
slightly
depressed

Orbitoros-
tral groove

Narrow, deep

Broad and
shallow
anteriorly

Broad and
deep

Narrow, deep

Narrow, deep

Short, shal-
low, round-
ed

Large chela

Transverse

groove*

Transverse

groove

Transverse
groove and
depressed
areas

Transverse

groove

Transverse

groove

Without

groove

Large cheli-
ped, merus

Strong tooth
on superior
margin; in-
ferior mar-
gin with
tooth and
series of
spines (?)

Similar teeth,
lacking meral
spines

Similar teeth,
with serra-
tions on all
margins

Same as C.
pseudopugnax

Similar teeth,
2-4 spines
on inferior
margin

Without su-
perior tooth,
inferior
tooth weak;
no spines

Carpal arti-
cles of sec-
ond legs,
ratio of first
to second

10 : 25

10 : 10

10 : 11

10 : 10

10 : 10

10 : 10

Third legs,
merus

Strong tooth

Strong tooth

Strong tooth

Strong tooth

Without

tooth

Without

tooth

*De Man states in his key ( 1911 ) "Chela without a transverse groove near the articulation of the dactylus.” However, Dana
(loc. cit.) states "narrow emarginate above, near articulation,” and shows in his figure what might be the groove.

Hawaiian Crangonidae — Banner

115

and C clypeata (Coutiere), the diagnostic char-
acteristic afforded by the ratio of the lengths
of carpal articles may prove to be of some-
what questionable worth.

To separate this from the related Hawaiian
species, see Table 5.

DISTRIBUTION: On Oahu four specimens
were from unrecorded localities, presumably
from the shallow water at the reef surfaces;
specific localities were Kaneohe Bay, under
coral heads on sand flats about a foot deep at
low tide, where the species was common;
Waimanalo, 6 feet deep; Hanauma Bay, 20
feet deep; Black Point, 2 feet deep; and Wai-
kiki, 2 feet deep. On Maui, specimens were
collected at Kalama Park, 3 and 8 feet deep;
10 and 12 miles south of Lahaina, both at 6
feet; and at Lahaina, 4 and 6 feet deep. There
are no other records of the variety from the
Hawaiian Islands.

Crangon gracilipes (Stimpson)

Fig. 41 a-i

Alpheus gracilipes Stimpson, Acad. Nat. Sci.

Phila., Proc. 12: 31, I860.

DESCRIPTION: Rostrum narrow, triangular,
acute, flattened dorsally, without carina, tip
slightly depressed, reaching 0.7 of length of
visible portion of first antennular article. Ros-
trum separated from posterior portion of or-
bital hoods by deep narrow depressions, more
anteriorly by wide flattened areas with abrupt
sides against both orbital hoods and rostrum,
latter overhanging groove. Orbital hoods in-
flated, large, with abrupt medial margin, pos-
teriorly merging with carapace, anteriorly with
dorsoventral keel. Anterior margin of orbito-
rostral area concave.When seen in lateral view,
anterior carapace without definite angle along
dorsal surface at base of rostrum [as there is in
C. percyi (Coutiere)], and without tubercle at
this point.

Antennular peduncle long and slender with
second article over twice as long as broad,
longer than visible portion of first article, over
twice as long as third article. Stylocerite reach-

ing to or slightly beyond end of first antennu-
lar article. Scaphocerite long, with strong lat-
eral spine about 1.1 times as long as squamous
portion; outer margin somewhat concave;
squamous portion somewhat reduced. Strong
lateral spine of basicerite reaching tip of ros-
trum. Scaphocerite slightly longer than anten-
nular peduncle; tip of carpocerite reaching
slightly beyond end of second antennular ar-
ticle. Antennal flagellum about 1.5 times body
length.

Large chela subcylindrical, 1.3 times as high
as broad, 3.7 times as long as high, tapering
toward distal end. Palm with deep transverse
groove proximal to articulation of dactylus.
Palm, at articulation of dactylus, with narrow
depression to accommodate high crest of dac-
tylus when flexed. Dactylus 0.3 length of
chela, compressed, strongly arcuate, longer
than fixed finger. Merus about 2.5 times as
long as broad; superior margin somewhat
rounded but usually projecting as acute tooth
(specimen drawn also with secondary tooth) ;
inferior internal margin acute, with 2-4 spines
and acute tooth distally.

Small chela not sexually dimorphic, long
and narrow, over 5 times as long as broad,
fingers somewhat broadened, 0.7 length of
palm. Dactylus with fringe of long setae run-
ning longitudinally from near point of articu-
lation over crown of article before apex. Small
subacute tooth on propodus above articula-
tion of dactylus. Merus usually with acute
teeth distally on inferior internal and superior
margins ; inferior internal margin usually with
one to several spinules.

Carpal articles of second legs with ratio
10 : 10 : 4 : 5 : 6.

Ischium of third legs with small movable
spine; merus 8 times as long as broad, iner-
mous; carpus 0.5 as long as merus, with tooth
at inferior distal margin poorly developed;
propodus almost as long as merus, with 9-11
slender spines; dactylus simple, curved, 0.2
length of merus.

Telson 2.1 times as long as broad at base,
tip 0.6 width of base. Lateral margins almost

116

PACIFIC SCIENCE, Vol. VII, January, 1953

straight, posterior margin arcuate. Dorsal
spines moderately developed, 0.4 and 0.7 of
distance from articulation to tip. Length of
larger pair of terminal spines less than 0.5
breadth of tip. Tip of telson and distolateral
margin of inner uropod with series of small
spinules.

Length up to 22 mm. One specimen ob-
served alive; dark, almost black, to unaided
vision; color from greatly expanded blue and
red chromatophores. Under microscope body
mottled, with clear areas over eyes and on
branchiostegites over third maxillipeds and
elsewhere, and with darker areas like two "eye
spots" on sides of abdomen. Antennules and
antennae, second to fourth legs, and caudal
fan bluish; fifth legs reddish; chelae slightly
darker than body with pink cast at dactylar
articulations and on tips of fingers.

DISCUSSION: In a few specimens available
for comparison no great variation was noted.
The rostral length varied, with the tip reach-
ing from 0.7 of the length of the first article
of the antennular peduncle to beyond the end
of the same article. In some cases the second
antennular article was over twice the length of
the first. The merus of the large chela varied
as noted above; almost always the distal teeth
were well developed; the same is true of the
small chela. In some of the specimens the sec-
ond carpal article of the second legs was
slightly shorter than the first. The number of
spines on the propodus of the third legs varied
slightly.

The only possible difference between this
and the original description of Stimpson lies
in the lateral spine of the basicerite which he
specifies to be minute, whereas on all of these
specimens it is large and well formed; how-
ever, exactly what Stimpson meant by "mi-
nute” is not known. The specimens also agree
well with the specimens described by de Man
(1911) from the southwest Pacific.

To separate this from the related Hawaiian
species, see Table 5.

DISTRIBUTION: The few specimens available
were collected from Kaneohe Bay, Kaaawa,

and Waikiki, Oahu, and one from Maalaea,
Maui; these were all without records as to
depth, but presumably they were from shal-
low water. Edmondson (1925) reported two
specimens from Lisianski Island. The species
has also been reported from the Hawaiian Is-
lands by Coutiere.

C. gracilipes occurs from the Red Sea through
the Indian Ocean, the East Indies, and Japan,
to Samoa, Tahiti, and Hawaii. The type local-
ity is Tahiti.

Crangon pugnax (Dana)

Fig. 42 a-h

Alpheus pugnax Dana, U. S. Explor. Exped.

13: 554, pi. 32, figs. 6a-6h, 1852.

Alpheus pugnax Dana, Acad. Nat. Sci. Phila.,

Proc. 6: 27, 1854. [Description without fig-
ures.]

As no specimens are available, the original
description is given:

Beak [rostrum] acute, narrow triangular, flat above,
arising from between the bases of the eyes. Basal spine
of outer antennae [of basicerite] small, basal scale
[scaphocerite] longer than base [carpocerite]. Base [pe-
duncle] of inner antennae shorter than scale of outer;
basal spine [stylocerite] of inner antennae not shorter
than the first joint, second joint short. Anterior feet
unequal; larger hand long, smooth with rounded mar-
gins, narrow emarginate above near articulation, fingers
short (about one-third the length of hand), arm having
a spine at both inner and outer apex. Feet of second
pair long, first joint short, hardly half the second in
length. Third and fourth pairs slender, third joint with
an acute tooth at lower apex.

At Lahaina, Island of Maui, Hawaiian group.

Length, twelve lines. The beak from its base is very
narrow, triangular, with straight sides, and there is no
Carina prolonged down the back. Second joint of inner
antennae hardly longer than first. Larger hand partly
pubescent, somewhat fusiform, but little compressed,
the finger turned out of plane of the hand; movable
finger short, thin above, with arcuate dorsal margin. !
First joint of carpus of second pair of feet less than half
the second; third shorter than fourth; fifth longer than
fourth; hand as long as fourth and fifth. The fifth joint
of third pair of legs long, very much longer than fourth,
about eight set of spinules on its lower side, rather long ■
hairy above.

Dana further states in his key: "Base of ^
rostrum rising between the eyes, deep sulci in \
carapace either side of rostrum. Orbital mar- :

Hawaiian Crangonidae — BANNER

117

Fig. 41. Crangon gracilipes (Stimpson). a, b. Anterior region, dorsal and lateral aspects; c, d, large cheliped,
lateral and medial aspects; e, large chela, distal end, ventral aspect; /, small cheliped, male, lateral aspect; g, small
chela, male (different specimen), ventral aspect; h, second leg; /, third leg. {a, b, g, h, scale A; c-f, i, scale B.)

118

PACIFIC SCIENCE, Vol. VII, January, 1953

Fig. 42. Crangon pugnax (Dana), "a, Front, side view,
enlarged; b, upper view; c, part of outer maxilliped; d,
e, larger hand, in different positions; /, smaller hand;
g, part of leg of second pair; h, part of leg of third pair.”
(Figures and legend from Dana; figures redrawn from
U. S. Exploring Expedition, Folio Atlas to Vol. 13,
1855, plate 35, figures 6a-h.)

gin inermous.” [Translated from Latin; Dana,
1854.]

DISCUSSION: As far as I have been able to
determine, this species has not been reported
since its original description. Moreover, the
type specimen has been lost. As no specimens
in the collections available agreed with the de-
scription on all characteristics, a special field
trip was made to the type locality, Lahaina,
Maui, to collect neotypes of this species and
of C. diadema (Dana). Unfortunately, the con-
ditions at Lahaina are not the same as they
were 100 years ago when the collections were
made by Dana; now the near-by sugar mill is
discharging fresh water charged with large
quantities of red mud onto the reef area, kill-
ing most of the coral and the other life on the
reef. It was possible to collect quite a few
specimens in spite of the pollution, but none
had the characteristics of C. pugnax. In a fur-

ther attempt to obtain specimens of the spe-
cies, the reefs 5, 10, and 30 miles away from
Lahaina were visited; none of these reefs du-
plicated the original conditions of the La-
haina reef, for they were either deeper, or more
sandy, or less well developed, etc., and no
specimens of C. pugnax were obtained.

However, on the basis of the original de-
scription and figures alone, it is possible to
separate clearly and decisively C pugnax from
all other species of Hawaii. Table 5 gives
the criteria for its separation from the obvi-
ously closely related Hawaiian species of the
subgroup Diadema.

If this species is not extinct, and there is no
good reason to presume it is until far more
extensive collections are made about the Ha-
waiian Islands, it is valid and recognizable.

Crangon diadema (Dana)

Frontispiece, Fig. 43 a-k

Alpheus diadema Dana, U. S. Explor. Exped.

13: 555, pL 25, figs. 7a-7e, 1852.

Alpheus diadema Dana, Acad. Nat. Sci. Phila.,
Proc. 6: 23, 1854. [Description without fig-
ures.]

Alpheus insignis Heller, K. Akad. Wiss. Wien,
Sitzungber. 44: 269, Taf. 2, figs. 17-18, 1861.

neotype: a male, 22 mm. long, collected
at Lahaina, Maui, from a head of coral (Po-
rites) in about 4 feet of water (U. S. N. M.
93460).

DESCRIPTION: Rostrum short, reaching to
middle of visible portion of first antennular
article; rostral base broad, flattened, extending
posteriorly between eyes; margins of rostrum
and base anteriorly slightly concave and taper-
ing, almost parallel in middle, posteriorly
slightly concave and spreading; margins lat-
erally overhanging orbitorostral groove, with
opposing face of groove almost touching ros-
tral portion ; carina prominent only in middle
section; posteriorly marked by slight protu-
berance. Orbital hoods high, rounded, ex-
ceedingly clear, conspicuously demarked on
all sides, anteriorly projecting as slight

Hawaiian Crangonidae — BANNER

119

rounded vertical keel. Area between orbital
hoods and narrow orbitorostral furrow flat-
tened; anterior margin arcuate.

Antennular peduncle with first article
slightly shorter than second, correspondingly
longer than third; second article almost twice
as long as broad. Stylocerite short and broad,
lateral spine reaching end of first antennular
article. Lateral spine of basicerite well devel-
oped, shorter than stylocerite. Carpocerite
reaching end of antennular peduncle, ex-
ceeded by scaphocerite. Scaphocerite with lat-
eral spine dominant, squamous portion re-
duced; lateral margins concave.

Large chela subcylindrical, almost as broad
as high, about 3 times as long as broad, taper-
ing distally; surface without sculpture except
for transverse groove proximad of articulation
of dactylus; chela sparsely hirsute on upper
and inner surfaces. Dactylus about 0.4 length
of chela, strong, with margin arcuate. Merus
about twice as long as broad, about 0.25 as
long as chela; superior distal margin project-
ing as acute tooth, inferior internal margin
with tooth distally. Large chela of female sim-
ilar in form but relatively much smaller.

Small chela of male 0.8 as long as carapace,
subcylindrical, tapering, 3 times as long as
broad, without sculpturing. Fingers slightly
over 0.3 length of whole chela, expanded,
about twice as long as broad, with dense fringe
of setiferous bristles along both sides and over
upper distal surface of finger ("balaeniceps-
shaped,” see Fig. 43/^g). Carpus slightly
elongate, 0.25 length of chela. Merus similar
to that of large chela but without inferior in-
ternal tooth. Small chela of female quite simi-
lar in form but much smaller, without balae-
niceps-shaped dactylus and with carpus rela-
tively more elongate.

Carpal articles of second legs with ratio
10 : 10 : 3 : 4 : 5.

Third legs with strong spine on ischium.
Merus 3.5 times as long as broad, strong,
curved, acute tooth subterminally on inferior
margin. Carpus 0.5 as long as merus, inferior
margin projecting as strong tooth. Propodus

about 0.8 as long as merus, tapering, with
about 20 movable spines of varying sizes.
Dactylus simple, slightly curved, acute, as
long as merus.

Telson almost twice as long as broad, pos-
terior margin 0.75 as broad as anterior section;
lateral margins slightly concave. Posterior
margin arcuate. Dorsal spines large, located
about 0.3 and 0.7 of distance from articulation
to tip. Length of larger pair of posterolateral
spines 0.3 width of tip; smaller pair less than
0.5 as long as longer pair. Margin between
posterolateral spines with 9 somewhat irreg-
ularly placed spinules. Distolateral margin of
inner uropod with 4 large spines and 9 spi-
nules.

Large females up to 26 mm. long; no males
exceeding 22 mm. of neotype. Color in life
variable, but usually dark, olive green, reddish
brown, etc., with pronounced irregular mot-
tling of lighter color; specimens at times al-
most transparent.

DISCUSSION: All the specimens at hand were
examined, but detailed studies on the varia-
tion in proportions of the appendages were
not made. However, variation was noted in
three parts. First was in the anterior region of
the carapace. Small specimens, 8 millimeters
long or less, had a markedly different rostrum
than the adults. In these specimens the ros-
trum was narrow and of almost uniform taper,
with its tip reaching almost to the end of the
first antennular article; the rostral carina was
well developed along the complete length;
moreover, in these specimens the area between
the rostrum and the orbital hoods extended
relatively further forward, markedly surpass-
ing the anterior margin of the orbital hoods.
In specimens of increasing size, the condition
of the rostrum approached that of adults. In
mature specimens, variation was noticed es-
pecially in the shape of the base of the ros-
trum between the orbital hoods; in some it
was as described above with the margins of
the middle portion straight and almost paral-
lel, whereas in others the curve from the edge
of the carapace to the posterior limits was uni-

120

PACIFIC SCIENCE, VoL VII, January, 1953

form and gradual. Furthermore, although in
most large specimens the tip of the rostrum
was depressed as shown in Figure 43^, in a
few of intermediate size it was found to con-
tinue almost ' ’level,” and in one it was slight-
ly elevated.

A second field of variation was in the carpal
articles of the second legs; some had the ratio
given above, but in some the ratio of the first
two articles approached 10 : 13. None were
noted where the second article was shorter
than the first.

The third variation was in the propodus of
the third legs. 'Small specimens had only four,
five, or six spines well developed; among the
large specimens some had a few more, some
had a few less, than the 20 described.

Since the original description of this spe-
cies from the Hawaiian Islands by Dana, no
specimens have been reported. In 1861 Heller
described from the Red Sea what was appar-
ently a closely related form, Alpheus insignis,
and all later workers, evidently because they
were not certain of the characteristics of
Dana’s species, used Heller’s name. As C. in-
signis the species was later reported from the
Hawaiian Islands. Coutiere in 1899 (p. 500)
listed C. insignis as a synonym of C. diadema
but in 1909 (p. 899) reviewed the situation
and decided, like de Man, to reserve judg-
ment until Dana’s type was redescribed. Un-
fortunately, the type of Dana’s species was
lost, and it was impossible to determine the
exact characteristics of the species.

To resolve this uncertainty, a special trip
was made to Lahaina, Maui, the type locality,
to collect replacement type specimens for
Dana’s description. This effort was successful
as the reef at Lahaina had a larger proportion
of this species than any other locality visited,
and the specimens also were larger and more
robust. One of them is described above.

This neotype differs in five minor ways
from Dana’s drawings and description. First,
the margins of the base of the rostrum were
shown as definitely and uniformly concave
throughout their length between the orbital

hoods, instead of straight and parallel as they
are in this specimen; however, as noted above,
this is a variable character. Second, his profile
drawing shows the rostrum as turning up near
its apex, instead of being depressed as in this
specimen; this, too, is a variable characteristic.
Third, the margin between the base of the ros-
trum and the anterior orbital hoods is shown
as concave instead of convex; this may be due
to faulty delineation in the drawing. Fourth,
the second carpal article of the second legs
was described as being much shorter than the
first and was shown to have the ratio (approxi-
mately) of 10 : 7 instead of 10 : 10 or 10 : 12,
as found in all the specimens in the collection.
Finally, the propodus was described as bear-
ing "six sets of spinules on inner side,” in-
stead of approximately 20 found in the neo-
type. This last difference could be attributed
either to the immaturity of Dana’s specimens
or to inadequate observation.

Thus, the only significant difference be-
tween Dana’s description and this neotype is
in the ratio of the carpal articles of the second
legs. This may be due to a chance variation in
the specimens or to faulty observation; but,
even if an actual difference, it would not be a
sufficient difference to warrant calling the lo-
cal specimens other than diadema.

I can find no significant differences between
this species and the form described by Heller.
Heller shows the margin between the rostrum
and the anterior margin of the orbital hoods
to be concave instead of convex (as did Dana)
and somewhat more narrow; he describes the
propodus of the third legs as bearing seven or
eight spines. As these differences, if correct,
would be at most of only subspecific value,
Heller’s species is placed in synonymy.

Table 5 contrasts this with related Hawaiian
species.

distribution: This species is common in
the shallow waters of the Hawaiian Islands.
On the reef flats or in water to 20 feet deep it
has been collected from the following locali-
ties: on Oahu, Kaneohe Bay, Mokulua Is-
land, Waimanalo, the shore near Koko Head,

Hawaiian Crangonidae — Banner

121

Fig, 43. Crangon diadema (Dana), a, h. Anterior region, dorsal and lateral aspects; c, large cheliped, lateral
aspect; d, large chela, ventral aspect; e, large cheliped, merus, medial aspect;/, small cheliped, male, lateral aspect;
g, small chela, distal end, male, ventral aspect; h, small cheliped, female, lateral aspect; /, second leg; / third leg;
k', telson and uropods, (Drawings of neotype.) {a, h, k, scale A; c-j, scale B.)

122

PACIFIC SCIENCE, Vol. VII, January, 1953

Hanauma Bay, Black Point, Waikiki, Nana-
kuli, Mokuleia, Kaaawa, and Kawela Bay; on
Maui, Makena, Kalama Park, Lahaina, and
12 miles south of Lahaina. The ” Albatross”
collected five specimens from "Honolulu
Reef” (U.S.N.M. 63552, 63553). There is only
one dredged specimen, taken between 40 and
350 feet deep off the southwest coast of Oahu.
Edmondson (1925) has reported the species
(as C. insignis) from Laysan and Lisianski Is-
lands and Pearl and Hermes Reef.

As C. insignis the species has been reported
throughout the tropical Indo-Pacific region
from the Red Sea to Samoa; Edmondson
(1925) has reported it from Johnston Island
in the central Pacific.

Crangon pseudopugnax sp. nov.

Fig. 44 a-i

TYPE SPECIMEN: A female 15.6 mm. long,
collected at Kalama Park, southeast Maalaea
Bay, Maui; it was collected from old coral at
a depth of about 8 feet. Five paratypes, one
from shallow water at Makena, Maui, one
from shallow water at Waikiki, Oahu, and
three from about 18 feet of water off Waikiki
Reef, Oahu (U.S.N.M. 93513).

DESCRIPTION: Rostrum acute, twice as long
as broad at anterior margin of base, tip reach-
ing to end of first antennular article. Rostral
Carina well- developed crest continuing to mid-
dle of orbital hoods, then broadening into
dorsally flattened triangular area. Orbital
hoods inflated, rounded anteriorly, well de-
marked from surrounding carapace, except on
posterior margins merging with carapace; or-
bital hoods very clear. Area between orbital
hoods and rostrum flattened; anterior margin
arcuate. Anterior portion of carapace de-
pressed, seen in profile, with small medial
protuberance or lobe.

Second and third articles of antennular pe-
duncle equal in length but both slightly
shorter than visible portion of first article; sec-
ond article about 1.3 times as long as broad.
Stylocerite well developed, tip reaching be-
yond end of first antennular article.

Lateral spine of basicerite well developed
but not reaching end of first antennular arti-
cle. Scaphocerite with lateral spine dominant
and squamous portion reduced; lateral mar-
gin strongly concave. Carpocerite as long as
antennular peduncle; both exceeded by sca-
phocerite.

Large chela subcylindrical, 1.3 times as
broad as high, 2.7 times as long as broad
[note: Figs. 44<; and 44^ show neither maxi-
mal nor minimal diameters] ; margins rounded;
tapering toward fingers. Palm marked by deep
transverse groove proximal to articulation of
finger, which spreads and soon disappears on
superior face, but on inferior face continues
into shallow, poorly defined depressed area
extending distally to articulation of dactylus
and proximally an equal distance. Adhesive
plaque on independent lobe, demarked by
palmar depression on inferior side and by
deep, short depression on superior side that
accommodates ridge of dactylus upon flexure.
Dactylus about 0.3 length of entire chela;
high, narrow, arcuate, with pronounced ridge
along free margin; tip rounded. Fixed finger
blunt at tip. Chela with occasional setae and
usual tufts on fingers. Merus triangular, sharp-
ly angled; superior margin terminating as
acute dentate projection, with 5 small inden-
tations proximally from which very small
bristles arise; inferior external margin with 7
similar indentations; inferior internal margin
with 9 indentations and strong subacute tooth
subterminally. Ischium with 3 indentations
for bristles.

Small chela cylindrical, 5 times as long as
broad, fingers about 0.7 as long as palm. Fin-
gers narrow. Chela with usual setae. Carpus of
usual form. Merus about 0.7 as long as chela,
triangular, with indentations similar to merus
of large chela but fewer; inferior internal mar-
gin unarmed.

Carpal articles of second legs with ratio
10 : 11 : 4 : 5 : 7.

Ischium of third legs with movable spine
on inferior margin (difficult to see) . Merus 5
times as long as broad, with strong, acute

Hawaiian Crangonidae — Banner

123

-I 1 2 mm

A

-I • 2mm

B

Fig. 44. Crangon pseudopugnax sp. nov. a, b, Anterior region, dorsal and lateral aspects; c, large cheliped, lateral
aspect; d, large chela, ventral aspect; e, large cheliped, merus and carpus, medial aspect; /, small cheliped, lateral
aspect; g, second leg; h, third leg; i, telson and uropod. (Setae are not shown on the chelipeds or the second
legs.) {a, b, i, scale A; c-h, scale B.)

124

PACIFIC SCIENCE, VoL VII, January, 1953

tooth on distal inferior margin. Carpus 0.5
as long as merus, unarmed except for tooth on
inferior distal angle. Propodus 0.8 length of
merus, with 10 movable spines in addition to
usual setae. Dactylus simple, acute, somewhat
curved.

Telson almost twice as long as broad, tip
slightly more than 0.5 as broad as base; lateral
margins slightly convex anteriorly, slightly
concave posteriorly; posterior tip strongly
convex. Dorsal spines moderately developed,
anterior approximately 0.4 and posterior 0.7
of length of telson from base; medial pair of
terminal spines narrow, acute, over 0.5 as long
as breadth of tip of telson. Medial margin of
tip with 3 irregularly placed, feeble spinules.
Inner uropod with 5 spines of graduated size
along distolateral margin; spine of outer uro-
pod long, rather slender.

In life, type specimen with broad, red, trans-
verse bands.

DISCUSSION: Only one of the paratypes had
all its appendages; all were smaller than the
type specimen. In these few specimens not
much variation was noted. The most conspic-
uous difference was in the rostral base between
the eyes. Most of the specimens showed a
narrow but carinate base that was set off lat-
erally by shallow grooves, while the type
lacked these grooves. One specimen had a
slight spine on the merus of the small chela,
and a specimen half the size of the type did
not have the depressed areas on either side of
the groove of the large chela. Aside from these
differences and slight differences in the pro-
portions of the articles, all the specimens were
similar. Unfortunately there were no intact
male specimens in the collection, but it is pre-
sumed that the small chela of the male is ba-
laeniceps-shaped as it is in the related species.

RELATIONSHIP: This species plainly belongs
to the Diadema Subgroup, and shows rela-
tionship to C. diadema (Dana) in the nature of
the orbital hoods and the transverse groove on
the palm of the large chela; the rostrum is
somewhat like that of C gracilipes (Stimpson)
and the general configuration is sufficiently

close to C. pugnax (Dana) that I tentatively
identified it as that species in the field.

C.pseudopugnax can be separated easily from
these related species by a combination of
characteristics. From C. diadema the species
may be separated most easily by the nature of
the base of the rostrum; from C gracilipes by
the presence of a groove on the large chela
and by the tooth on the merus of the third
legs; from C. pugnax by the lengths of the first
two articles of the second legs. (See Table 5.)

This species shows a close affinity to an-
other related Hawaiian species, C.percyi (Cou-
tiere). They can be distinguished by the na-
ture of the rostrum, which in C. pseudopugnax
bears an acute carina on the anterior portion
of the rostral base and does not overhang the
orbitorostral groove except at the posterior
end, while in C. percyi the anterior surface is
broad and flat, and the margins overhang the
orbitorostral grooves for their entire length;
by the large chela, which bears a depression
on the lower face in C. pseudopugnax that is
lacking in C.percyi\ and by the small cheliped,
which lacks teeth at the dactylar articulation
and a tooth on the inferior internal margin of
the merus in C. pseudopugnax^ whereas the dac-
tylar articulation is flanked by two teeth and
the inferior internal margin of the merus bears
a distal tooth in C. percyi.

This species is also related to C. philoctetes
(de Man), but the latter has a series of mov-
able spinules on the merus of the small chela
and the merus of the third legs, which are ab-
sent in C. pseudopugnax. Unfortunately the
large chela was absent in de Man’s type speci-
men.

DISTRIBUTION: All Specimens known are re-
corded above.

Crangon percyi (Coutiere)

Fig. 45 a-g

Alpheus percyi Conxihs^, Soc. Philomath. Paris,
Bui. IX, 9(5): 21, 1908.

Alpheus percyi Coutiere, Linn. Soc. London,
Trans. II (Zool.) 17: 426, pi. 64, figs. 22-

Hawaiian Crangonidae — Banner

22f, 1921. [Same description as original, but

with figures.]

description: Rostrum acute, reaching to
end of first article of antennular peduncle;
posteriorly continued as broad base between
eyes; lateral margins almost straight, over-
hanging orbitorostral furrow; base with low,
indistinct dorsal carina. Orbitorostral furrows
deep and narrow, flattening anteriorly, with
anterior margin somewhat sinuate. Orbital
hoods hemispherical anteriorly, high and of
clear chitin; merging with carapace posterior-
ly. Anterior region of carapace, seen in pro-
file, depressed; beginning of depressed region
with small medial protuberance.

First and second articles of antennular pe-
duncle subequal; third article somewhat
shorter. Stylocerite strong, slightly exceeding
first article. Lateral spine of basicerite some-
what shorter than stylocerite. Scaphocerite
with strong lateral spine, squamous area re-
duced; lateral margins strongly concave. Ba-
sicerite as long as antennular peduncle, both
exceeded slightly by scaphocerite.

Large chela rounded, subcylindrical with
maximum diameter 1.2 times minimum diam-
eter, chela 2.7 times as long as maximum di-
ameter. Palm with deep transverse groove
proximal to dactylus but without other sculp-
turing. Dactylus about 0.3 length of entire
chela, arcuate, high and narrow. Chela with
only scattered setae except for tufts about fin-
gers. Merus over twice as long as broad, su-
perior and inferior internal margins with
strong acute teeth; all margins with series of
small indentations from which fine setae arise.

Small chela subcylindrical with tapering fin-
gers, 2 poorly developed teeth flanking articu-
lation of dactylus; fingers in female conical,
in male balaeniceps-shaped, or bearing a dense
fringe of setae over broadened dactylus (ac-
cording to Coutiere; no males collected from
Hawaii) . Merus similar to that of large chela
except more slender and superior and inferior-
internal teeth more poorly developed.

Carpal articles of second legs with ratio
10 : 10 : 4 : 5 : 6.

125

Ischium of third legs with moderately
strong movable spine. Merus 5 times as long
as broad, with strong tooth distally on inferior
margin. Carpus 0.5 as long as merus, with
usual inferior distal tooth. Propodus almost
0.8 as long as merus, with 11 relatively long
spines on inferior and distal margins. Dactylus
0.2 as long as merus, somewhat curved, acute,
simple.

Telson of usual form, similar to that of C.
pseudopugnax, with series of small spinules on
posterior margin. Distolateral margin of inner
uropod with about 12 small spinules.

Specimens in collection 3 females, one 25
mm., others about 20 mm. long.

DISCUSSION: The other specimens agreed
with the one described and illustrated on most
characteristics. The points of difference be-
tween one or the other of the two undescribed
specimens and the one described above were:
First, the "break,” or angle, the anterior cara-
pace and rostrum made to the dorsal surface
of the carapace was less pronounced, and the
tubercle was less noticeable. Second, the ros-
trum was shorter, reaching only 0.7 of the
length of the visible portion of the first anten-
nular article. Third, the second antennular ar-
ticle was relatively longer, being 1.3 times the
length of the visible portion of the first article.
Finally, on the large chela there was a poorly
marked shallow depression that ran from the
transverse groove toward the articulation of
the dactylus. These differences probably are
individual variations.

The specimens of this species, which has
not previously been reported from the Ha-
waiian Islands, differ in only a few details
from the original description and figures by
Coutiere. It is uncertain from Coutiere’ s draw-
ings whether the configuration of the base of
the rostrum and the orbitorostral furrows are
exactly the same in the two forms, but in any
case they are very similar. Coutiere does not
show as much depression of the anterior mar-
gin of the carapace as is shown in these speci-
mens, nor does he show the median tubercle.
The carpocerite may be somewhat shorter in

126

PACIFIC SCIENCE, Vol. VII, January, 1953

Coutiere’s specimens. It should be noted that
on the left side of the described specimen
(Fig. 45^) the spine of the basicerite was [like
that shown by Coutiere, but the one on the
right was entirely absent. The large and small
chelae are similar, but the merus of these ap-
pendages in the type may or may not have
the series of fine indentations found in the lo-
cal specimens, for Coutiere’s plates are not
clear at the point. The third legs are similar
except that Coutiere does not indicate the
ischial spine, although he does state that the
appendage "est tres semblable [to C dasy-
cheles (Cout.)] comme forme ...” and that C.
dasycheles has the spine. Finally, on the tip of
the telson and on the inner uropod he indi-
cated no spinules; these, however, are very
difficult to discern on the local specimens be-
cause they are confused with the bases of the
setiferous bristles, and it is likely that Cou-
tiere merely overlooked them.

In my opinion none of these differences are
of sufficient magnitude, constancy, or general
reliability to warrant the erection of a new spe-
cies for the local form.

The characteristics which will distinguish
this from closely related local species are
given in Table 5.

DISTRIBUTION: The three specimens col-
lected locally came from Oahu in 20 feet of
water at Hanauma Bay, in 15 feet of water off
Nanakuli, and at Halape, Kau Coast, Hawaii,
in water 35-40 feet deep (last collected by R.
W. Hiatt).

Brevirostris Group

' Crangon rapax (Fabricius)

Fig. 46 a-i

Alpheus rapax Fabricius, Sup. Ent. Syst. p.
405, 1798.

Alpheus malaharkus Hilgendorf, K. Akad.

Wiss., Berlin, Monats. p. 832, 1878.

Alpheus brevirostris de Man, Linn. Soc. Lon-
don, Jour. 22: 261, 1888.

Alpheus rapax de Man, Soc. Zool. France,
Mem. 22: 147-155, 1909. [Species rede-

scribed and illustrated; see these and the
following pages in this reference for a com-
plete synonymy.]

DESCRIPTION: Rostrum small, acute, equi-
lateral, tip reaching to end of first 0.3 of
visible portion of first antennular article; ros-
tral Carina anteriorly sharp, posteriorly some-
what rounded, extending to end of orbital
hoods. Orbital hoods large, inflated, rounded
anteriorly, posteriorly merging with the cara-
pace; higher in middle than rostral carina;
transparent, hence from side rostral carina
visible through hood. Anterior margin of
carapace from orbital hoods to rostrum al-
most straight.

Antennular peduncles elongate and thin,
second article 3. 5-4.0 times as long as broad,
2.5 times length of first article or of third
article. Stylocerite rounded, leaf-like, anterior
spine represented by very small tooth reaching
about 0.75 of length of visible portion of first
antennular article. Antennular articles with
few short bristles near points of articulation;
stylocerite with fringe of short bristles on
margins. Basicerite with distinct but short
tooth about same length as rostrum. Scapho-
cerite with strong lateral spine, squamous
portion narrow; lateral margins concave. Sca-
phocerite slightly longer than carpocerite
which reaches to end of antennular peduncle.

Distal articles of third maxillipeds large, of
usual form, densely bristled on inner face.

Large chela high, compressed, margins
rounded, 2.7 times as long as high, about
twice as high as wide. Upper surface with
rounded transverse groove extending short
distance down each face. Dactylus strong,
high, compressed, 0.35 as long as entire chela.
Carpus of usual form. Merus 2.6 times as long
as broad, with superior distal angle rounded,
inferior internal margin with 5 movable spines
and acute terminal tooth of moderate size.
Dactylus, margins of palm, superior and in-
ferior internal margins of merus with scattered
long setae.

Hawaiian Crangonidae — Banner

127

Fig. 45. Crangon percyi (Coutiere). a, b, Anterior region, dorsal and lateral aspects; c, d, large cheliped, lateral
and ventral aspects; e, small cheliped; /, second leg; g, third leg. {a, b, scale A; c-g, scale B.)

128

Small chela of male with palm almost 2.5
times as long as wide, fingers 1.5 times length
of palm; margins without grooves. Dactylus
broad with opposing faces flattened and
fringed with dense setiferous bristles, or ba-
laeniceps-shaped; both dactylus and fixed fin-
ger strongly hooked at tip. Carpus somewhat
longer than broad. Merus narrower proxi-
mally than distally, maximum breadth 1.5
times minimum, article twice as long as broad
at maximum breadth; armature like that of
large cheliped except tooth on inferior inter-
nal margin lacking. Small chela of female
reported (de Man, 1909) as similar in form to
that of male except lacking rows of setae on
dactylus (therefore not of balaeniceps-shape)
and slightly smaller in proportions.

Carpal articles of second legs with ratio
10 : 7 : 3 : 3 : 5.

Ischium of third legs with strong spine.
Merus 4.4 times as long as broad, with few
setae along margins. Propodus slightly less
than 0.5 length of merus, superior distal mar-
gin with sparse long setae, superior distal
angle projecting as tooth. Propodus 0.7
length of merus with 7 spines on inferior and
terminal margins, scattered long setae on in-
ferior and superior margins. Dactylus 0.5
length of propodus (or 0.3 length of merus),
narrow, almost straight, tip acute, inferior
surface flattened, superior surface rounded;
rounded surface with 2 short bristles.

Telson similar in form to that described for
C. platyunguiculata (p. 130), 1.8 times as long
as wide at point of maximum breadth; tip
0.7 as wide as maximum breadth; postero-
lateral spines slightly shorter and weaker than
those of C. platyunguiculata. Tip of telson and
distal lateral margins of inner uropod with
weak spines similar to those of platyunguku-
lata.

Sole specimen in collection, male 28 mm.
long; specimens reported by de Man {loc.
cit.) to reach 51 mm. Color in life not noted.

. DISCUSSION: Crangon rapax (Fabr.) was re-
described and illustrated by de Man in 1909.
The specimen from Hawaii differs from this

PACIFIC SCIENCE, Vol. VII, January, 1953

description in a few characteristics. The speci-
mens are similar in general configuration, and
the general relationship of the size, shape, and
armature of the appendages. Some minor
differences were noted in a comparison of
this specimen with de Man’s description, for
example: the rostrum reaches to the end of
the first third of the visible portion of the
first antennular article in this specimen, and
from one half to two thirds of the length in
de Man’s; the visible portion of the first
antennular article is slightly longer and the
third article is slightly shorter, when compared
to the second article, than those described by
de Man; the merus of the large chela is 2.6
times as long as broad in this specimen and
bears five spinules, while it is 3. 0-3 .5 times
as long as broad and bears six to seven spinules
in the described specimens; the ratio of the
carpal articles is 10 : 7 : 3 : 3 : 5 instead of
10 : 8-9 : 3 : 3 : 4; and so on. Two other
differences may be of greater importance: the
merus of the small cheliped of the male is
broader and lacking the small terminal tooth
on the inferior internal margin found in the
specimens described by de Man; the carpus
and propodus of the third leg in this specimen
bears only scattered long setae, while de Man
describes and depicts his specimens as being
densely hirsute on these articles.

When considering these differences it should
be remembered that de Man’s description is
based upon two specimens, a male 51 mm.
long and a female 46 mm. long, while the
Hawaiian specimen is only 28 mm. long.
Therefore, it appears that all these dif-
ferences are of minor importance and may be
growth differences, or differences normally
occurring in a population, or, possibly, they
may be differences indicative of a geograph-
ical subspecies of the parent C. rapax. How-
ever, in my opinion, the description of this
Hawaiian form as a new species or a new
subspecies is not indicated on the basis of the
present specimen.

This species can be separated from C. platy-
unguiculata, the only other Hawaiian species

Hawaiian Crangonidae — BANNER

129

Fig. 46. Crangon rapax (Fabricius). a, b. Anterior region, dorsal and lateral aspects; c, large cheliped, lateral
aspect; d, large chela, dorsal aspect; e, small cheliped, lateral aspect; /, small cheliped, merus and carpus, medial
aspect; g, second leg; h, third leg; /, third leg, dactylus. {a, h, g, h, scale A; c-f, scale B; /, scale C.)

130

PACIFIC SCIENCE, Vol. VII, January, 1953

of the Brevirostris Group and the only other
Hawaiian species with which it could be con-
fused, by the relative lengths of the second
carpal articles of the second legs, almost 3.0
times the length of the first article in C.
platyunguiculata and only 0.7 its length in this
species; by the excessive setae on the third
legs in C. platyunguiculata and by the thicker
chela in that species. Other characteristics for
the separation of the two species are found in
the frontal region of the carapace, the relative
lengths of the scaphocerite and the anten-
nular and antennal peduncles, and the small
chelae of the males and females.

DISTRIBUTION: This specimen was collected
on the sand flats of Kaneohe Bay, Oahu,
where the water is only a few inches deep at
low tide. For the possible habitat of this
species, see the discussion under C platyun-
guiculata.

It has been reported from Zanzibar, Dji-
bouti, and the Mergui Archipelago (Andaman
Sea) in the Indian Ocean, from the Nether-
lands Indies, Japan, and near Queensland in
the Pacific, but not previously from the cen-
tral Pacific region.

Crangon platyunguiculata sp. nov.

Fig. 47 a-k

TYPE SPECIMEN: A female 25 mm. long,
collected at Waialua Bay, Oahu, from old
coral in about 6 feet of water. Paratypes, two
males, one 20 mm., the other 10 mm. long,
from the same location (U. S. N. M. 93461).

DESCRIPTION: Rostrum small, triangular, as
long as broad at base, tip reaching less than
0.5 length of visible portion of first anten-
nular article. Rostral carina distinct and con-
tinuous from tip of rostrum to between orbital
hoods, thereafter more rounded and less well
defined but continuing to posterior margin
of hoods. Orbital hoods large, rounded an-
teriorly, frontal margin reaching to middle of
rostrum; posteriorly merging with convexity
of carapace; in lateral view higher than rostral
base; transparent, hence outlines of rostral

base visible through hoods in lateral view;
much larger than eyes.

Antennal peduncle long, rather slender.
Second article 3 times as long as broad, 1.3
times as long as first article, twice as long as
third article. Stylocerite large, with lateral
spine poorly developed, reaching only slight-
ly beyond curvature of scale; tip reaching
almost to end of first antennular article. Later-
al spine of basicerite well developed, slightly
shorter than stylocerite. Scaphocerite large,
with heavy outer spine; lateral margins slight-
ly concave, tip relatively heavy and somewhat
incurved; squamous portion narrow. Carpo-
cerite reaching beyond end of scaphocerite,
which slightly exceeds end of antennular
peduncle.

Third maxillipeds of usual form, but inner
faces of terminal and subterminal articles very
densely clothed with setae and short bristles.

Large chela compressed, 2.3 times as high
as thick, 2.3 times as long as high, margins
rounded, broadest near middle, slightly taper-
ing proximally, somewhat more distally. Up-
per margin proximal to dactylus with deep
transverse furrow expanding into shallow de-
pressed area on each face. Margin above
articulation of dactylus with shallow excava-
tion adjacent to palmar adhesive plaque for
accommodation of crest of dactylus when
flexed. Dactylus 0.3 length of chela, crested,
strongly arcuate, heavy, 2.4 times as long as
wide, compressed like chela. Chela with nu-
merous long setae on margins. Carpus of
usual form. Merus 2.5 times as long as broad,
superior distal margin rounded, inferior in-
ternal margin with 2 strong movable spines,
one about middle, other more distal, weak
but acute subterminal tooth; long setae found
only on distal portions of article.

Small chela of female somewhat com-
pressed, 3.5 times as long as broad, fingers
between 0.5 and 0.6 of length; surfaces with-
out grooves or depressions; articulation of
dactylus flanked by small subacute projection;
fingers quite rounded. Entire chela heavily
covered with long setae, especially along up-

Hawaiian Crangonidae — BANNER

131

-<2mm

j3mm

Fig. 47. Crangon platyunguiculatus sp. nov. a, b, Anterior region, dorsal and lateral aspects; c, large cheliped,
lateral aspect (setae not shown); d, large chela, dorsal aspect; e, large cheliped, merus, medial aspect (setae not
shown); /, small cheliped, medial aspect; g, small chela, male, lateral aspect; h, second leg; /, third leg;/, third
leg, dactylus; k, telson and uropods. {a, b,j, k, scale A; c-i, scale B.)

132

PACIFIC SCIENCE, VoL VII, January, 1953

per and lower margins and margins of op-
posing faces of fingers. Carpus O'. 2 5 as long
as chela, 1.2 times as long as broad. Merus
similar in form to that of large chela but
without terminal tooth on inferior internal
margin, with 4 small movable spines instead.
Small chela of male 3.7 times as long as
broad, fingers occupying slightly more than
0.6 length of chela. Dactylus not rounded,
but with opposing face flattened, with short
dense fringe of stiff setae along margins of
face in middle 0.7; fixed finger also flattened,
with corresponding fringe of setae in opposi-
tion; other setae similar to those of female.
Carpus and merus in male similar to those of
female.

Carpal articles of second legs with ratio
10 : 23 : 7 : 10 : 11.

Ischium of third legs with strong movable
spine. Merus 4.0 times as long as broad,
inermous with occasional short setae. Carpus
0.5 as long as merus, distal angles not pro-
jecting but rounded; superior margin with
series of long setae. Propodus 0.6 as long as
merus, tapering toward tip, straight, with 7
movable spines of moderate size, with nu-
merous long setae on both margins, some of
setae 0.5 as long as article. Dactylus long,
simple, almost 0.3 as long as merus, superior
margin rounded, inferior margin broad and
flattened, somewhat curved; tip acute; lateral
margins with short movable spine and short
curved bristle.

Telson with anterolateral margin convex,
broadest at about 0.3 of length, posterolateral
margin concave and narrowing; 1.8 times as
long as broad, tip 0.7 as broad as maximum
breadth. Posterior margin strongly arcuate,
reaching beyond tip of posterolateral spines.
Posterolateral spines about 0.3 as long as
breadth of tip; dorsal spines poorly developed,
located 0.4 and 0.6 of distance from articula-
tion to tip. Tip of telson and distal lateral
margin of inner uropod with weak spinules.

Body translucent white to gray base color,
with red and blue chromatophores arranged
in poorly defined transverse bands on abdo-

men and in narrow band across eyes; similar
poorly defined bands on chelae. In one live
specimen blue chromatophores expanded,
giving over-all blue-gray appearance; in other,
red chromatophores expanded, giving reddish
cast.

DISCUSSION: No significant variation was
noted in the three specimens.

RELATIONSHIP: This species plainly belongs
to the Brevirostris Group and can be separated
easily from all other Hawaiian species save
C. rapax (Fabr.) by the appearance of the
anterior margin of the carapace, the com-
pressed chela with a transverse groove, and
the dactylus of the third legs.

Within the Brevirostris Group this species
and C. sauvensis (de Man) can be distinguished
by the characteristics given in de Man’s key
(1911: 322-325). From most of the other

species in the group these two can be dis-
tinguished by the following characters: the
presence of a transverse groove on the palm
of the large chela; the balaeniceps-shaped
dactylus of the small chela of the males; the
lack of a tooth on the merus of the third leg.
Three more closely related species, C. djedde fi-
sts (Coutiere), C. djihoutensis (de Man), and
C. pubescens (de Man), all have a longer and
thinner merus on the third leg; in addition
C. djeddensis has a transverse groove on the
third chela; in C. djihoutensis the terminal spine
of the scaphocerite is much longer relative to
the squamous portion; and in C. pubescens the
rostral carina is longer.

C. platyunguiculata can be separated from
C. sauvensis by the characteristics given in the
following tabulation:

C. platyunguiculatus

Rostrum reaching beyond
orbital hoods by 0.5 its
length

Stylocerite with tip
straight

Scaphocerite shorter than
carpocerite

C. sauvensis

Rostrum reaching beyond
orbital hoods by 0.8 its
length

Stylocerite with tip in-
curved

Scaphocerite as long as
carpocerite

Large chela with palm 2.0
times as long as fingers
Carpal articles of second
legs with ratio 10 : 23 :
7 : 10 : 11

Large chela with palm 1.5
times as long as fingers
Carpal articles of second
legs with ratio 10 : 15 :
6:6:6

V

Hawaiian Crangonidae ■ — BANNER

133

Propodus of third legs 1.2 Propodus of third legs 1.5
times length of carpus times length of carpus
Dactylus of third legs not Dactylus of third legs spi-
spiniform, flattened on niform, not flattened
inferior face

On almost ail other points the two species are
similar , so similar in fact that, with only one
or two of the differences listed above, the
Hawaiian form could be considered at most
a variety. However, the great difference in the
dactylus of the third leg reinforced with the
other more minor differences leaves little
doubt but that this species is valid.

The differences between this species and
C rapax, its closest relative locally, are dis-
cussed under that species.

distribution: In addition to the type
series listed above, two specimens were col-
lected from Kaneohe Bay, Oahu, where they
v/ere found underneath coral heads on shallow
sand flats, less than a foot deep at low tide.
In many of the areas of silty sand in Kaneohe
Bay there are numerous burrows evidently
made by this species or by C. rapax (Fabr.).
Some of these burrows are also inhabited by
specimens of a small fish. Unfortunately, I
was unable to collect either the crangonids or
the fish, as the burrows penetrated through
the veneer of sand into the underlying con-
solidated coral of the reef. It was also im-
practical to seize the animals in their burrows,
to entice them out, or to drive them out. The
two that were collected evidently had burrows
that led under large loose coral heads lying
on the surface of the sand; when the heads
were moved the shrimp were caught in a dip
net.

Edwardsi Group

Crangon leptochims (CoutiUe)

Fig. 48 a-h

Alpheus leptochims Coutiere, Fauna and Geog.
Maid, and Laccad. 2(4): 914-916, pi. 87,
figs. 54-54e, 1906.

DESCRIPTION: Rostrum triangular, about
1.5 times as long as broad, tip reaching almost
to end of first antennular article. Rostral carina

rounded, extending slightly posterior of eyes.
Orbital hoods somewhat inflated, rounded
anteriorly, demarked from base of rostrum by
shallow rounded grooves. Anterior margin of
carapace somewhat emarginate between orbi-
tal hoods and rostral base.

Visible portion of first antennular article
as long as second article; second article about
1.8 times as long as broad; third article about
0.7 length of second. Tip of spine of stylo-
cerite reaching to end of first article.

Lateral spine of basicerite reaching to tip
of rostrum. Scaphocerite strong, lateral mar-
gin slightly concave, tip reaching consider-
ably beyond end of antennular peduncle.
Carpocerite longer than antennular peduncle,
reaching to end of squamous portion of
scaphocerite.

Large chela rather slender, 3 times as long
as broad, compressed, with grooves and de-
pressions. Transverse groove on superior
margin, with distal edge rounded and prox-
imal edge overhanging floor of groove;
groove expanding on external face to elon-
gate, narrow, triangular depression with poor-
ly demarked margins; on internal face con-
tinuous with similar but less extensive
depressed area. On inferior margin, opposite
groove on superior margin, notch with abrupt
proximal edge and gradual distal edge; on
external face notch continues, well-defined,
for 0.3 height of face; on internal face notch
poorly defined and continuing for only 0.25
of height. Finger of usual form, less than 0.3
length of entire chela. Merus 3 times as long
as broad, with superior margin ending in
obtuse angle; inferior internal margin with 2
movable spines and distal tooth.

Small chela of female almost 5 times as long
as broad, without sculpturing, fingers occupy-
ing almost 0.5 length of chela. Articulation
of dactylus flanked by small obtuse projection
on inner side. Carpus with small tooth pro-
jecting over propodus. Merus 4.5 times as
long as broad, almost 0.3 longer than merus
of large chela, with 3 movable spines and sub-
terminal tooth on inferior internal margin.

134

PACIFIC SCIENCE, VoL VII, January, 1953

Carpal articles of second legs with ratio
10 : 11 : 5 : 6 : 7.

discussion: Unfortunately all of the three
specimens in the collection were dredged from
deeper water and only one is reasonably in-
tact, the others having lost most of their ap-
pendages in the rough handling in the dredge
net. In the Edwardsi Group, the largest of the
genus, positive identification cannot be made
without reference to the small chela of the

male; and, as the small chela of the sole male
was lacking in these specimens, they are re-
ferred to this species with some doubts.

The specimens compare quite well with the
description and plates of Coutiere except on

the following points:

C. leptochirus

Second antennular article
1.4 times length of visi-
ble portion of first
Scaphocerite exceeding
antennular peduncle by
0.1 its length
Large chela with parallel
sides; proximal edge of
superior groove round-
ed; depressed areas on
upper portions of faces
very narrow

Meri of large and small
chelae each with 4 spines
Second carpal article of
second legs 0.7 length of
first

Third legs without ischial
spine, merus 7.5 times as
long as broad

Hawaiian Form
Second antennular article
as long as visible portion
of first

Scaphocerite exceeding
antennular peduncle by
0.2 its length
Large chela with slight ta-
per; proximal edge of
superior groove abrupt,
overhanging; depressed
areas on faces broader

Meri of large and small
chelae with 2-3 spines
Second carpal article of
second legs 1.1 length of
first

Third legs with ischial
spine, merus 5.5 times as
long as broad

These differences appear to be decisive
when listed out in this fashion; however,
studies on other species in this group and in
other groups of the genus have shown that
each of these characteristics are variable at

least to the extent here observed. Therefore no
dependence can be placed upon any one of
the differences listed; possibly in aggregate
they are significant, but without the important
small chela of the male, and with but a single
intact specimen to study, I do not believe
that the designation of this form as a distinct
species is warranted.

DISTRIBUTION: The three specimens were
taken in a dredge haul off Diamond Head,

Oahu, in 100-200 feet of water. Coutiere’s
specimens came from the Indian Ocean.

Crangon crassimanus (Heller)

Fig. 49 a-k

Alpheus crassimanus Heller, Reise der . . .No-
vara. . . . Zook Theil 2(8): 107, pi. 10, fig.
2, 1865.

DESCRIPTION: Rostrum acute, reaching al-
most to end of first antennular segment, with
margins gradually curving out to merge with
frontal portion of orbital hoods and carapace,
demarked from orbital hoods by broad, shal-
low depression. Subacute rostral carina arising
between anterior portion of orbital hoods.
Orbital hood hemispherical, without keels or
ridges.

Lateral spine of stylocerite slightly longer
than first antennular article. Antennular pe-
duncle slim, with second article equal to or
longer than visible portion of first, from 1.5
to 3.0 times as long as third. Lateral spine of
basicerite not as long as stylocerite. Scapho-
cerite with heavy lateral spine; lateral margin
concave; squamous portion narrow, with an-
terior margin not rounded but joining spine
at acute angle. Scaphocerite and carpocerite
almost equal, slightly longer than antennular
peduncle.

Large chela heavy, 2.2 times as long as
broad, laterally compressed, 1.7 times as high
as thick, with grooves and depressions.
Rounded transverse groove on superior mar-
gin proximal to articulation of dactylus, con-
tinuous with irregular depression with
rounded edges on both inner and outer face.
Proximal margin of transverse groove, in pro-
file, rounded and sloping back to superior
margin of chela. Another groove on inferior
margin of chela, directly below superior
groove, rounded only distally and demarked
by abrupt shoulder proximally, ending
abruptly on outer face about 0.2 distance from
lower edge, and on inner face gradually di-
minishing but still visible beyond mid-line.
Dactylus 0. 3-0.4 length of chela, heavy and

Hawaiian Crangonidae — BANNER

135

mm

jO.5

Fig. 48. Crangon leptochirus (Coutiere). a, b. Anterior region, dorsal and lateral aspects; c, large cheliped, lateral
aspect; d, large chela, medial aspect; e, small chela, female; /, second leg; g, third leg; h, third leg, dactylus. (a, b,
scale A; c-g, scale B; h, scale C.)

136

PACIFIC SCIENCE, Vol. VII, January, 1953

curved. Merus twice as long as broad; superi-
or margin unarmed; inferior internal margin
with tooth distally.

Small chela of usual form in female, sub-
spatulate in male. In female 3.4 times as long
as broad, without sculpture; fingers slender,
tapering, about as long as palm; short acute
tooth at point of articulation of dactylus
usually present, as illustrated, but sometimes
reduced or completely absent. In male small
chela relatively larger and heavier, 2.9 times
as long as broad, without sculpturing except
for slight depressed area on upper surface
proximal to articulation of dactylus. Dactylus
0.5 total length of chela, broad, flat, 2.7 times
as long as broad, with dense fringe of setifer-
ous bristles along margins, crossing top of
article proximal to tip; tip compressed and
hooked (dactylus balaeniceps-shaped). In
both sexes carpus projecting over propodus
as broad flat tooth, merus without tooth on
inferior internal margin, otherwise similar to
that of large chela.

Carpal articles of second legs with ratio
10 : 10 : 5 : 5 : 8.

Ischium of third and fourth legs usually
with movable spine; merus inermous; distal
margin of carpus not produced into acute
processes; propodus usually with about 10
movable spines; dactylus simple, slightly
curved.

Largest specimen, female from Kaneohe
Bay 26 mm. long; color usually transparent
with reddish to greenish transverse bands.

DISCUSSION: Marked variation was noted in
C. crassmanus, as in several other Hawaiian
species of which numerous specimens were
collected. In this species the careful append-
age-by-appendage study of size ratios was not
made, but the following variations were noted.

Antennular peduncles: The second article
varies' in relative length from as long as the
visible portion of the first and 1.5 times as
long as the third to twice as long as the first
and three times as long as the third.

Large cheliped: The amount of sculpturing
on this appendage varies markedly as is shown

in Figure 49c, d, e, f. In some specimens, es-
pecially the smaller ones, the sculpturing is
very much reduced and noticeable only in the
area adjacent to the margins; in larger speci-
mens the shallow depression may be quite ex-
tensive over both faces, usually with rounded
and ill- defined margins but at times with def-
inite limits. The tooth on the inferior internal
margin of the merus of the cheliped is usually
present although it is not as large as in many
other species; however, in some specimens it
may be reduced or entirely absent. In no speci-
men seen was the upper groove other than
gradually rounded on both margins.

Small cheliped: In some of the larger females
the dactylus, although it never approaches the
extreme condition found in males, is often
proximally broadened, not slender and taper-
ing as in younger females. The tooth on the
inner side of the articulation of the dactylus is
almost always present, although sometimes re-
duced; occasionally it is reduced to a rounded
shoulder. No specimens were observed with
a tooth on the merus like that of the large
chela. No males were found without the ba-
laeniceps-shaped dactylus.

Second legs: The second article of the carpus
varies from 0.75 to 1.3 times the length of the
first, and the fifth article from 0.5 to 1.0 times
the length of the first.

It has been observed that those members of
this species collected from Kaneohe Bay and
especially those collected on the inner arms of
Pearl Harbor, where the water is very warm,
quiet, and filled with silt, are larger, averaging
about twice the length of, and have a redder
color than, specimens collected from coral
boulders of the open reefs. No other charac-
ters distinctive of the two groups of specimens
could be found.

The local specimens agree well with the
original description of Heller, differing only
in one possible point. Heller states, "Anten-
narum spina externa basalis minima,” and, in
his plate, does not show a spine at all, while
in the Hawaiian specimens it is well formed.
The ratio of the articles of the carpus of the

Hawaiian Crangonidae — BANNER

137

j I min
Zmm

Fig, 49. Crangon crassimanus (Heller), a, b, Anterior region, dorsal and lateral aspects; c, d, large cheliped,
lateral and medial aspects; e,f, large chela from another specimen, setae not shown, lateral and medial aspects;
g, small cheliped, female; h, small chela, male, long setae not shown, lateral aspect; /, second leg; /, third leg;
k, third leg, propodus and dactylus. {a, h, k, scale A; c-j, scale B.)

138

PACIFIC SCIENCE, VoL VII, January, 1953

second leg (10 : 6 : 2 : 2 : 4, as taken from the
drawings) are almost within the range of vari-
ation of the Hawaiian form.

De Man (1911) suggests that possibly C
crassimanus is a synonym for C. lohides (de
Haan) , but a study of the literature following
the publication of the note reveals that the
question has not yet been settled and that C
crassimanus is still accepted as a valid species.

One specimen of this species in my collec-
tion and two in the collections of the Bernice
P. Bishop Museum (No. 5057) bear an un-
usual parasite attached to the posterior ven-
tral portion of the thorax. It is in the form of
a more or less bean-shaped body, whitish in
color and about 5 mm. long, which is attached
by a short stalk. The specimens from the Ber-
nice P. Bishop Museum were described as
new by Nierstrasz and Brandis (1930) as Faba
glabra^ while my specimen was identified as
this species by Dr. Edward C. Reinhard. The
parasite is of unknown systematic position,
but the original authors suggest it may be an
epicaridan crustacean. It is interesting to note
that, of all the crangonids from Hawaii, the
parasite has been found only on this one spe-
cies and that on that species it is extremely
rare.

DISTRIBUTION: This species is one of the
most common in the shallow inshore waters
of the reefs. There is, in Hawaii, no record of
its collection from water over 5 or 6 feet deep.
On Oahu it has been collected from the fol-
lowing localities: Kaneohe Bay, Koko Head,
Black Point, Waikiki, Pearl Harbor, Moku-
leia, Haleiwa, Kaaawa, Punaluu; on Maui it
was collected at Kalama Park and 12 miles
south of Lahaina; on Molokai at Kainalu.
Edmondson (1925) reported the species from
Laysan and Lisianski Islands. The "Albatross”
collected specimens in Pearl Harbor (U.S.N.
M. 63559) and Honolulu Reef (U.S.N.M.
63560, 63562).

In the central Pacific the species was re-
ported by Edmondson (1925) from Johnston
Island. It ranges through the Indo-Pacific re-
gion and has been reported from places as far

distant from Hawaii as Djibouti, at the mouth
of the Red Sea, and Cape York, Australia.

Crangon pacifica (Dana)

Fig. 50 a-i

Alpheus pacificus Dana, U. S. Explor. Exped.

13 (1): 544, pi. 34, fig. 5, 1852.

Alpheus pacificus Dana, Acad. Nat. Sci. Phila.,

Proc. 6: 21, 1854. [Diagnosis without fig-
ures.]

NEOTYPE: A male 32 mm. long, collected
at Kalama Park, about 18 miles southeast of
Lahaina, Maui, from a coral head in about 3
feet of water (U.S.N.M. 93462).

DESCRIPTION : Rostrum acute, triangular, al-
most twice as long as broad at base, reaching
nearly to 0.7 of visible portion of antennular
peduncle; margins straight, with few short
setae. Rostral carina rounded, arising slightly
posterior to eyes. Orbital hoods not greatly
inflated, rounded anteriorly. Orbitorostral
groove moderately shallow with gradually
sloping margins ; anterior portion of carapace
between rostral base and front of orbital
hoods concave.

Second article of antennular peduncle about
1.5 times as long as broad, 1.2 times as long
as visible portion of first or third articles.
Spine of stylocerite reaching slightly past end
of first antennular article.

Lateral spine of basicerite reaching almost
as far as spine of stylocerite. Scaphocerite with
squamous portion well developed, more than
0.9 length of slender lateral spine; lateral mar-
gins only slightly curved. Carpocerite slightly
longer than scaphocerite; scaphocerite 0.1
longer than antennular peduncle. Antennal
flagellum longer than body.

Large chela massive, 2.2 times as long as
broad, compressed, twice as broad as thick,
with lobes and crests. Superior margin cut by
deep transverse groove proximal to dactylus,
proximal edge of groove overhanging, distal
edge rounded; groove continued on outer
face as shallow, poorly defined, depressed
area extending downward about 0.3 breadth

Hawaiian Crangonidae — BANNER

139

Fig. 50. Crangon pacifica (Dana), a, h. Anterior region, dorsal and lateral aspects; c, d, large cheliped, lateral
and medial aspects; e, small cheliped, male, lateral aspect; /, small chela, female, lateral aspect; g, second leg; h,
third leg; /, telson. {a, b, i, scale A; c-f, scale B; g, scale C.)

m m

5 mm

3 mm

140

PACIFIC SCIENCE, VoL VII, January, 1953

of face and posteriorly beyond middle of
palm, continued on inner face as much smaller
triangular depressed area immediately below
groove. Inferior margin with much deeper
notch below upper groove, the proximal edge
overhanging, the distal edge abrupt but
rounded; notch continues more than 0.3 dis-
tance up outer face with proximal edge well
defined throughout but distal edge fading;
depression on inner face similar to that on
outer face but with both edges gradual. Dac-
tylus 0.4 length of chela, heavy, with distal
margin strongly curved. Scattered setae on
superior margin and inner distal face of chela
and on fingers. Merus about 0.25 as long as
chela, twice as long as broad, superior distal
margin obtuse and lacking inner distal tooth;
with occasional setae along margins. Large
chela of female similar.

Small chela of male 3.4 times as long as
broad, fingers 0.6 of total length. Palm com-
pressed, without grooves, ridges, or depres-
sions except for slight indentation at base of
fixed finger. Both fingers curved distally so
that points cross; both fingers with dense
rows of marginal setae proximally, becoming
more scattered distally; oppositional faces of
both fingers somewhat flattened. Articulation
of dactylus not flanked by tooth. Carpus with
tooth extending over base of propodus. Me-
rus similar in form to that of large chela. Small
chela of female of same proportions and arma-
ture as that of male except perhaps with few
less setae proximally on fingers; size relatively
less than that of male.

Carpal articles of second legs with ratio
10:8:3:3:5.5; second article of carpus 4.4
times as long as broad; chela as long as first
carpal article.

Ischium of third legs with strong movable
spine; merus 4 times as long as broad, taper-
ing slightly distally, unarmed; carpus about
0.5 as long as merus, neither superior nor in-
ferior distal margin projecting distally as acute
tooth; propodus 0.75 as long as merus, with
II moderately large spines on inferior and dis-
tal margins; dactylus acute, curved, simple.

slightly shorter than width of merus. Merus
with several short setae; superior margin of
carpus and propodus with scattered, long
setae. Brush on fifth legs well developed.

Telson tapering only slightly in posterior
third, anterior margins convex; 2.2 times as
long as broad, maximum breadth 1.4 times
breadth of tip. Margin of tip only slightly ar-
cuate. Posterolateral spines feeble, scarcely
reaching beyond rounded portion of tip; dor-
sal spines heavy, 0.4 and 0.6 of distance from
articulation to tip. Middle of posterior border
and distolateral margin of inner uropod with
small, irregularly placed spinules.

Neotype about maximal size observed.
Color in life, banded red and transparent, with
blue-violet markings on appendages, especial-
ly large chela.

DISCUSSION: Although not as much varia-
tion was noted in this species as in the closely
related C crassimanus (Heller) the following
points are noteworthy: The length of the sec-
ond antennular article was as much as twice
the length of the visible portion of the first in
some smaller specimens; the sculpturing on
the upper parts of both faces of the large chela
varied in extent; the number of setae on the
opposing faces of the fingers of the small
chela also varied from the neotype, some hav-
ing more, some less; the second article of the
carpus of the second legs was almost as long
as th^ first article in some specimens; and the
number of spines on the propodus of the third
legs varied slightly from the II found in the
type specimen. In none of the specimens,
however, was there a tooth on the merus of
the large chela, nor at the articulation of the
dactylus of the small chela; in no case was the
proximal margin of the superior groove of the
large chela other than overhanging the floor
of the groove nor was the small chela of the
males approaching the condition found in C.
crassimanus (balaeniceps-shaped). ^

Dana’s original specimen, now lost, was
very large, IM inches (or about 45 mm.), and
was collected at some undesignated spot in
the Hawaiian Islands. The neotype, the larg-

Hawaiian Crangonidae — BANNER

141

est in the present collection, unfortunately is
somewhat smaller (32 mm.), and was collected
on Maui near where Dana had collected other
Hawaiian types (see under C diadema and C
pugnax) . It agrees well with Dana’s description
and plates except in the ratio of the articles of
the carpus of the second leg which according
to his plate had a ratio of 10 : 5 : 3 : 2.5 : 4,
which is 10 : 8 : 3 : 3 : 5.5 in this specimen.
While this character was quite variable in the
specimens in the present collection, in no case
did it approach the extreme given by Dana;
however, it is likely that this difference still
can be individual variation or possibly a
growth difference. A second more slight dif-
ference is that Dana’s plates do not show the
proximal margin of the superior groove of the
large chela overhanging the groove; this char-
acter, too, is somewhat variable in these speci-
mens, and may well be either an individual or
a growth difference.

This species resembles C crassimanus (Hel-
ler) very much, and there is no positive way to
distinguish between them except by the char-
acter of the first thoracic legs. The small chela
in the male is subspatulate in C crassimanus
and of the usual form in both the male and
female in C. pacifica. On the large chela the
most useful character is the presence of a
tooth on the internal distal angle of the merus
in C. crassimanus, with none in C. pacifica;
however, this tooth is variable in size and
often very much reduced. The proximal edge
of the upper depression is rounded in C. cras-
simanus and usually overhangs in C. pacifica,
but some specimens of both species approach
the perpendicular. The superior depression on
the internal face usually extends posteriorly
past the mid-line of the palm in C. crassimanus
and scarcely back of the upper edge of the de-
pression in C pacifica.

distribution: This species, like C. crassi-
manus, is common in shallow waters. It does
not seem to penetrate the inshore portions of
the reefs nor the quieter waters of enclosed
bays as much as C. crassimanus (although it

does penetrate as far into Kaneohe Bay) ; it also
reaches a bit further beyond the tidal zone
having been collected from water l5 feet deep
at Nanakuli, Oahu. It has been collected from
the following localities on Oahu: Kaneohe
Bay, Waimanalo, Hanauma Bay, Black Point,
Waikiki, Nanakuli, Mokuleia, Kawela Bay,
and Kahana Bay; on Maui it has been col-
lected at Makena, Kalama Park, Lahaina, and
12 miles south and 10 miles north of Lahaina.
The ’’Albatross” collected specimens on Ho-
nolulu Reef and at Waialua, Oahu (U.S.N.M.
63590 and 63593) and Napili Harbor Reef,
Maui (Station No. 3881, U.S.N.M. 63592).
Edmondson (1925) has reported the species
from Laysan and Ocean Islands.

It is widely spread through the Indo-Pacific
region, from the Red Sea and Madagascar to
New South Wales and Wake Island.

Species of Doubtful Record
Crangon strenua (Dana)

AlphcnS strenuus Dana, U. S. Explor. Exped.

13(1): 543, pk 34, fig. 4, 1852.

Originally described from Tongatabu,
Tonga, this species is of wide distribution in
the south Indo-Pacific, from the Galapagos
on the east to Djibouti at the mouth of the
Red Sea on the west. It was reported from an
unspecified locality in Hawaii by Stimpson
(1861) as A. avarus Fabr. (he regarded A.
strenuus as a synonym for A. avarus). How-
ever, as the species has not been reported
from this archipelago since that time, and as
it can be confused most easily with the com-
mon C. crassimanus (Heller) (the two species
can be separated with certainty by the pres-
ence of a balaeniceps-shaped small chela in
both the male and female in C. strenua in
contrast to having the condition only in the
male in C. crassimanus), it is likely that Stimp-
son made a mistake in identification.

142

PACIFIC SCIENCE, Vol. VII, January, 1953

C. audouini (Coutiere)

Alpheus audouini Coutiere, Fauna and Geog.
Maid, and Laccad. 2: 911-9135 pi. 87, lig.
52, 1905.

This species was reported from Hawaii by
Coutiere in his original description: "L’espece
se trouve dans toute la Mer Rouge, I’Ocean
Indien, la Malaisie, et de la ZHande aux
lies Sandwich.” Unless this species is con-
fused with the closely related C. pacifica (in
which case the name C pacifica would take
priority), it has not been reported from the
Islands since, nor was it found in the present
collections. Therefore it seems likely that
Coutiere’s sweeping distributional list was a
little too inclusive.

REFERENCES

Anonymous. 1904. Records of the dredging
and other collecting stations of the U. S.
Fish Commission Steamer Albatross in 1901
and 1902. Report ofi the Commissioner for the
Year Ending June 30, 1902, U. S. Commission
of Fish and Fisheries 1902: 399-432.

Balss, Heinrick. 1927. Decapoda. In: Hand-
buch derZoologie [Willy Kiikenthal, ed.] Bd.
Ill, Pt. 1: 841-1082. W. de Gruyter u. Co.,
Berlin u. Leipzig.

Bate, C. Spence. 1888. Report on the Crus-
tacea Macrura dredged by H. M. S. Chal-
lenger during the years 1873-76. In: The
Voyage ofH. M. S. Challenger, Zoology 2Y xc
d- 942 [157 pis. in separate vol.]. Eyre &
Spottiswoode, London.

Boone, Lee, 1935. Scientific results of the
world cruise of the yacht "Alva” 1931, Wil-
liam K. Vanderbilt, Commanding. Crusta-
cea. . . . Vanderbilt Mar. Mus., Bui. 6: 1-
264, 96 pis.

Coutiere, Henri. 1896. Note sur quelques
genres nouveaux ou peu connus d’Alphe-
ides, formant la sous-famille des Alpheops-
ides. Paris Mus. d'Hist. Nat., Bui. 2(8): 380-
386.

1898^. Note sur quelques Alpheides

nouveaux de la collection du British Mu-

seum. Soc. Fnt. de Prance, Bui. 1898 (6) : 149-
152; (7): I66.

1898^. Note sur Synalpheus biunguicu-

latus Stimpson.^ deMan [Crust.]. Soc. Fnt. de
France, Bui. 1898 (11): 232-233.

1899. Les "Alpheidae,” morphologie

externe et interne, formes larvaires, biono-
mie. Theses presentees a la Faculte des Sci-
ences de Paris. . . . Ser. A, No. 321 [No.
d’Ordre 980]. 599 pp., 409 text figs., 6 pis.
Masson et Cie., Paris. [Also in: Ann. Sci.
Nat., VIII, Zook 9: 1-560.]

1903. Note sur quelques Alpheidae

des Maldives et Laquedives. Soc. Philomath.
Paris IX, 5(2): 72-91.

1906. Marine Crustacea, XV. Les Al-
pheidae. In: Fauna and Geography of the Mal-
dive and Laccadive Archipelagoes. [J. Stanley
Gardiner, ed.] Vol. 2, Rpt. 21, pp. 852-921,
pis. 70-87, text figs. 127-139. University
Press, Cambridge.

I9O8. Sur quelques nouvelles especes

d’Alpheidae. Soc. Philomath. Paris, Bui. IX,
11(5): 1-26.

1909. The American species of snap-
ping shrimps of the genus Synalpheus. U. S.
Natl. Mus., Proc. 36: 1-93, text figs. 1-54.

— 1921. Les especes d’Alpheidae rap-

portees de I’Ocean Indien par M. J. Stan-
ley Gardiner. Linn. Soc. London, Trans. II
(Zool.) 17(4): 413-428, pis. 60-64. [Also
titled: Reports of the Percy Sladen Trust Expe-
dition to the Indian Ocean in 1903. . . . 6(10):
413-428, etc.]

Dana, James D. 1852. Crustacea. In: United
States Exploring Expedition, During the Years
1838, 1839, 1840, 1841, 1842, under the
Command of Charles Wilkes, U. S. N. Vol. 13,
viii + 685. [Folio Atlas of 96 plates pub-
lished in 1855.]

— 1854. Conspectus Crustaceorum, 'etc.

Conspectus of the Crustacea of the Explor-
ing Expedition under Capt. Wilkes, U. S. N.
Acad. Nat. Sci. Phila., Proc. 6: 6-28 [Proc.
for 1852].

DE Man, Johannes Govertus. 1909. Note
sur quelques especes du genre Alpheus

Hawaiian Crangonidae — Banner

143

Fabr. appartenant au Group Brevirostris.
Soc. Zool. de France, Mem. 22: 146-164, pis.
7-8.

1910. Diagnoses of new species of

macrurous decapod Crustacea from the "Si-
boga Expedition,” V. Ned. Dierk. Ver.,
Tijdschr. II, 11(4): 287-319.

1911. The Decapoda of the Siboga Ex-
pedition. Part II. Family Alpheidae. In:
Siboga-Expeditie 39a ^ (2): 133-327 [Livre
60]; 1915, Supplement [Plates and explana-
tions] 39a^ (2): 23 pis. [Livre 74].

1922. The Decapoda of the Siboga Ex-
pedition. On a collection of macrurous dec-
apod Crustacea of the Siboga Expedition,
chiefly Penaeidae and Alpheidae. In: Siboga-
Expeditie 39a^ (5): 1-51, pis. 1-4 [Livre 93].

Edmondson, Charles Howard. 1923. Crus-
tacea from Palmyra and Fanning Islands.
Bernice P. Bishop Mus., Bui. 5: 3-43.

1925. Crustacea. Marine zoology of

the tropical Central Pacific. Bernice P. Bishop
Mus., Bui. 27: 3-22.

1930. New Hawaiian Crustacea. Ber-
nice P. Bishop Mus., Occas. Papers 9(10):
3-18.

1933. Reef and Shore Fauna of Hawaii.

295 pp. Bernice P. Bishop Museum, Special
Publication 22, Honolulu. [Revised 1946.]

Fabricius, Johann C. 1898. Supplementum En-
tomologia systematica emendata et aucta ....
572 pp. Hafniae [Copenhagen].

Gurney, Robert. 1938. The larvae of the
decapod Crustacea. In: Great Barrier Reef
Expedition 1928-1929, Scientific Reports. 6(1):
1-60. British Museum (Natural History),
London.

Haskell, Daniel Carl. 1942. The United
States Exploring Expedition 1838-1842 ....
xii + 188 pp. The New York Public Li-
brary, New York.

Heller, Camil. 1861. Beitrage zur Crustace-
en-Fauna des Rothen Meeres. K. Acad.
Wiss. Wien, Sitzungber. 44: 241-295, 3 Taf.

1865. Crustaceen. [Abth. 8] In: Reise

der Osterreichischen Fregatte Novara um die
Erde in den Jahren 1857-38-39 unter den Be-

fehlen des Commodores B. vonWiillerstorf-Ur-
bair. Zool. Theil. Bd. 11. 280 pp., 25 Taf.

Hilgendorf, Franz Martin. 1878. Die von
Herrn W. Peters in Mozambique gesam-
melten Crustaceen. K. Akad. Wiss. Berlin,
Monatsber. 1878: 782-851, 4 Taf.

Hult, Joran. 1938. Crustacea Decapoda
from the Galapagos Islands collected by
Mr. Rolf Blomberg. Arch, for Zool. 30(5) :
1-18.

International Commission on Zoologi-
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25] rendered by the International Commis-
sion on Zoological Nomenclature. Smithsn.
Inst., Pub. 1938: 1-61. [Opinion 17, pp.
40-42.]

1922. Opinions rendered by the In-
ternational Commission on Zoological No-
menclature, Opinions 68 to 77. Smithsn.
Inst., Misc. Pub. 73(1): 1-73. [Opinion 74,
pp. 32-34.]

Lenz, Heinrich. I90I. Ergebnisse einer Reise
nach dem Pacific. Crustaceen. Zool. Jahrb.,
Abt. f. System., Geog. u. Biol. Tiere 14(5):
429-482.

Miers, Edward John. 1875. On some new
or undescribed species of Crustacea from
the Samoan Islands. Ann. and Mag. Nat.
Hist. IV, 9: 341-344.

1881. On a collection of Crustacea

made by Baron Hermann Maltzan at Goree
Island, Senegambia. Ann. and Mag. Nat.
Hist. V, 8: 204-377, 4 pis.

1884. Crustacea. In: Report on the Zoo-
logical Collections made in the Indo-Pacific
Ocean during the Voyage ofH. M. S. '' Alert
1881-2, XXV + 684 pp., 54 pis. British Mu-
seum (Natural History), London.

Milne -Edwards, Henri. 1837. Histoire natu-
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physiologie et la classification de ces animaux.
Tome 11. 532 pp. Roret, Paris.

Nierstrasz, H. F., and G. A. Brender a
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new species of parasitic Crustacea. U. S.
Natl. Mus., Proc. 11 {9)'- 1-9-

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POCOCK, R. L 1890. Crustacea of Fernando
Noronha. Linn. Soc. London, Jour., Zool.
1890: 506-529, pi. 20.

Ramadan, M. M. 1936. Report on a collec-
tion of Stomatopoda and Decapoda from
Ghardaqa, Red Sea. Egypt. Univ., E acuity
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Randall, John W. 1839. Catalog of the
Crustacea brought by Thomas Nuttall and
J. K. Townsend, from the west coast of
North America and the Sandwich Islands
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147.

Rathbun, Mary J. 1904^^?. Some changes in
crustacean nomenclature. Wash. Biol. Soc.,
Proc. 17: 169-172.

1904^ and 1910. Decapod crustaceans

of the northwest coast of North America.
Smithsn. Inst., Harriman Alaska Series 10:
1-210. [First privately published in New

York under the earlier date as Vol. X of
Alaska.]

Richters, F. 1880, Crustacea Decapoda of
Mauritius and the Seychelles. In: Beitrdge
zur Meeresfauna der Insel Mauritius und der
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pp,, 25' pis. Gurmann, Berlin.

Schmitt, Waldo L. 1939. Decapoda and
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Stimpson, W. 1861. Prodromus descriptionis
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Hamburgi.

I

News Notes

The Congress of the Republic of the Philip-
pines, in a special session, has made funds avail-
able for expenses in connection with the Eighth
Pacific Science Congress. The Eighth Congress
is to be held in Quezon City, The Philippines,
from November 16 to 28, 1953. Dr. Vidal A.
Tan is President of the Congress, and Dr. Patro-
cinio Valenzuela is Secretary-General. (From
the Pacific Science Association Information Bul-
letin. )

According to Dr. Gilbert Archey, Secretary-
General of the Seventh Pacific Science Congress,
the final volume of the Proceedings of the Sev-
enth Congress is now in the hands of the print-
ers. All volumes should be issued by February
or March of 1953. The Meteorology and
Oceanography volume is being distributed at
the present time. Enquiries and orders for the
set of the Proceedings or for the separate vol-
umes are being handled by the Secretary, Royal
Society of New Zealand, Victoria University
College Buildings, Wellington. (From the Pa-
cific Science Association Information Bulletin.)

Dr. Alexander Spoehr has been appointed
Director of the Bernice P. Bishop Museum,
Honolulu, succeeding the late Sir Peter Buck.
Dr. Spoehr, who is Curator of Oceanic Eth-
nology at the Chicago Natural History Museum,
will assume the directorship in January, 1953.
He is an authority on native kinship and social
organization, and is recognized as an expert on
field and museum technique. (From the Pacific
Science Association Information Bulletin.)

The South Pacific Commission has appointed
as its Executive Officer for Economic Develop-
ment, Dr. A. H. J. Kroon, a Dutch agricultural

scientist with many years’ experience in agri-
cultural development in Indonesia.

In association with Dr. E. M. Ojala, Deputy
Chairman of the Research Council, Dr. Kroon
will be responsible for carrying out the Com-
mission’s projects for fostering the economic
development of Pacific territories. He will work
in collaboration with the Executive Officers in
charge of the two other main fields of Com-
mission activity — health (Dr. Emile Massal)
and social development (H. E. Maude).

From the summer number of the American
Council of Learned Societies Newsletter we re-
print, by permission, this delightful poetic
comment by Kit Berry:

O SYLLABUS, O SYLLABI

Servitude Scholastica Grandifiora

"The Oxford don, with brow so high,
Pronounces it gladioli’ ”,

And Cambridge, Mass., aspiring higher.
Builds Babel’s tower the heavens nigher.

O syllabus, O syllabi.

O octopus, O octopi.

The antique hybrids still appear,

With Grecian front and Latin rear,
Embodied in our English tongue
By no true graft, but stiffly hung,

Mere fossils from an ancient lore.

Which learning with less effort wore,

And, when it grafted, grafted true.

And made a hybrid live and new.

O syllabus, O syllabi.

O omnibus, O omnibi? ?

Yet it has been said by a liberal scholar that
"Culture is forgotten learning,” and by a
prophet that "The letter killeth, but the spirit
giveth life.”

■ [145]

p

Index to Crangonids

Names marked with an asterisk ( * ) are either new or redefined, those in
italics are synonyms. Page numbers in bold type indicate descriptions.

Alpheus (see Crangon or Synalpheus for species), *pacifica, 48, 51, 138

9, 46

Alpheopsis, 10, 14, 21, 30
key, 15
aequalis, 1 5
*biunguiculatus, 15, 18
chilensis, 15
equalis, 15
fissipes, 20
idiocarpus, 14
Arete, 20, 30
Areteopsis, 20
Athanas, 20
Athanopsis, 20
Automate, 20

Betaeus, 20
harrimani, 5
truncatus, 5

Crangon, 9, 10, 20, 46
key, 48

acuto-femorata, 107
*albatrossae, 48, 49, 55, 60, 64
amirantei, 48, 49, 87
audouini, 48, 142
avarus, 14 1

*brachymerus, 48, 49, 55, 58
*brevipes, 48, 50, 103
hrevirostris, 126
bucephalus, 99, 102
clypeata, 48, 50, 106, 107
collumiana, 48, 49, 67
crassimanus, 48, 51, 134, 14 1
dasycheles, 126
dentipes, 70
deuteropus, 48, 49, 70
*diadema, 48, 51, 114, 118
djeddensis, 132
djiboutensis, 132
*edmondsoni, 48, 49, 78
faceta, 84

gracilipes, 48, 51, 115
gracilis, 75, 76
alluaudi, 76
luciparensis, 76
^simplex, 48, 49, 75, 77, 81
hailstonei, 51, 54
assimulans, 54, 55
hailstonei, 54, 55
laetabilis, 54, 55
paucispinata, 48, 49, 51, 60
hawaiiensis, 48, 50, 92, 95
clippertoni, 96
insignis, 118
laevis, 84

*latipes, 48, 49, 81, 82
lays ant, 99

leptochirus, 48, 51, 133
lobides, 1.38
malabaricus, 126
* nanus, 47, 48, 50, 90
obesomanus, 86
*oahuensis, 48, 49, 55, 64

parabrevipes, 107
paracrinita, 48, 50, 110

bengalensis, 48, 50, 110, 114
paradentipes, 48, 49, 72
paragracilis, 47, 48, 50, 92, 96
paralcyone, 48, 50, 99
(?) paralcyone, 103
percyi, 48, 51, 114, 124
philoctetes, 124

*platyunguiculata, 48, 51, 128, 130
*pseudopugnax, 48, 51, 114, 122
pubescens, 132
pugnax, 48, 51, 114, 116
rapax, 48, 51, 126, 133
rostratipes, 93, 94
sauvensis, 132
seurati, 67
shearmei, 81
splendida, 85
staphylinus, 63
strenua, 48, 14 1
*tuthilli, 48, 49, 55, 63
ventrosa, 48, 49; 84

Crangonidae, 8

key to Hawaiian genera, 10

Jousseaumea, 10, 21
key, 10

brevirostris, 10, 12
mauiensis, 10, 12

Metalpheus, 90, 93

*Neoalpheopsis, 10, 20
key, 21

euryone, 21, 25
*hiatti, 21

Ogyris, 20, 30

Parabetaeus, 20

Pterocaris, 20, 30

Synalpheus, 10, 20, 26
key, 29

albatrossi, 28, 29, 30, 36
*biunguiculatus, 29, 32, 36
charon, 28, 29, 37, 40
*coutierei, 37
gravieri, 40
helleri, 38
lockingtoni, 42
macromanus, 29, 45
paraneomeris, 28, 29, 36, 40, 46
halmaherensis, 42
oxyceros, 42
praedabundus, 42
prasalini, 42
prolatus, 42
seychellensis, 42
prolificus, 28, 29, 38
* redactocarpus, 26, 29
townsendi, 28, 40, 44
brevispinis, 28

{147]

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Batzo, Roderick L., and J. K. Ripkin. 1849.
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Crawford, David L. 1920^i. New or interesting
Psyllidae of the Pacific Coast {Uomop.) . Ent.
News 51 (1): 12-14.

1920^. Cerotrioza (Psyllidae, Homop-

tera). Hawaii. Ent. Soc., Proc. 4 (2): 374-
375.

Rock, Joseph F. 1916. The sandalwoods of Ha-
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APRIL, 1953

NO. 2

' ’ . -F ! i 1

►L. VII

PACIFIC

SCIENCE

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

IN THIS ISSUE: Yamamoto — Rotatorian Fauna of Korea

• St. John and Mason — Vernacular Names of Plants of
Bikini • Landa — Body Length to Scale Length in the
Pacific Pilchard • Vandel — A New Isopod from Oregon

• Li — Plant Interchanges between Formosa and the Phil-
ippines • Levi-Castillo — Culex (Phalangomyia) in Ecua-
dor • Parrott — Catalogue of Australian Braconidae •
Harry — Bathypelagic Fishes^ Paralepididae % NOTES

Published by

THE UNIVERSITY OF HAWAII PRESS
HONOLULU, HAWAII

BOARD OF EDITORS

Leonard D. Tuthill, EMtor^n-Chief
Department of Zoology and Entomology, University of Hawaii

O. A. Bushnell, Associate Editor
Department of Bacteriology, University of Hawaii

Marjorie Milnes, Assistant to the Editors
Office of Publications and Information, University of Hawaii

Earl J. Anderson

Pineapple Research Institute Pathologist
Pineapple Research Institute
Honolulu 14, Hawaii

Townsend Cromwell
Pacific Oceanic Fishery Investigations
Oceanographer

Pacific Oceanic Fishery Investigations
Honolulu 14, Hawaii

Maxwell S. Doty

Department of Botany, University of Hawaii

R. A. Falla

Dominion Museum, Wellington, New Zealand

Christopher Gregory

Department of Mathematics
University of Hawaii

D. Elmo Hardy

Associate Entomologist, University of Hawaii
Agricultural Experiment Station

Wendell A. Mordy

Department of Meteorology
Pineapple Research Institute
Honolulu 14, Hawaii

Norman S. Noble
Commonwealth Scientific and Industrial
Research Organization
314 Albert Street

East Melbourne, C2, Victoria, Australia
Harold St. John

Department of Botany, University of Hawaii

G. Donald Sherman
Chemist, University of Hawaii
Agricultural Experiment Station

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University of Hawaii

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Scripps Institution of Oceanography
La Jolla, California

Thomas Nickerson, Managing Editor
Office of Publications and Information, University of Hawaii

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( Continued on inside hack cover)

PACIFIC SCIENCE

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

VOL. VII APRIL, 1953 NO. 2

Previous issue published January 2, 1953

CONTENTS

PAGE

Pfelifninary Studies on the Rotatorian Fauna of Korea.

Kokichi Yamamoto 151

Vernacular Names of the Plants of Bikini, Marshall Islands. Pacific Plant

Studies 12. Harold St. John and Leonard E. Mason 165

The Relationship between Body Length and Scale Length in Live Y ear-Classes of

the Pacific Pilchard or Sardine, Sardinops caerulea (Girard, 1834).-
Antonio Landa 169

A New Terrestrial Isopod from Oregon, Caucasonethes Rothi n sp.

A. Vandel 175

Floristic Interchanges between Formosa and the Philippines.

Hui-Lin Li ... 179

Observations on the Subgenus Phalangomyia of the Genus Culex in Ecuador
with Description of a New Species (Diptera: Culicidae) .

Roberto Levi-Castillo 187

A Systematic Catalogue of Australian Braconidae.

Arthur W. Parrott . 193

Studies on the Bathypelagic Fishes of the Family Paralepididae. 1. A Survey

of the Genera. Robert R. Harry 219

NOTES:

Faunal Speciation in Netv Georgia, Solomon Islands.

R. J. A. W. Lever 250

News Notes 251

Pacific Science is published quarterly by the University of Hawaii Press, in January,
April, July, and October. Subscription price is three dollars a year; single copies are
.one dollar. Check or money order payable to University of Hawaii should be sent to
University of Hawaii Press, Honolulu 14, Hawaii, U.S.A.

Preliminary Studies on the Rotatorian Fauna of Korea^

Kokichi Yamamoto^

INTRODUCTION

Thorough limnological surveys in Korea
have not been carried out up to the present
time; only a few brief studies on the plankton,
especially on the rotatorian fauna, have been
made in several districts. There are papers
relating to the Korean rotatorians published
by five authors, namely, Kawamura (1917),
Hada (1936), Ueno (1941), Sato (1939, 1941),
and Yamamoto (1941). One of the reasons
why such studies have been neglected is the
paucity of lakes and ponds suitable for col-
lections, except for a series of brackishrwater
lakes on the coast of the Japan Sea and reser-
voirs recently built for hydroelectric plants.
The studies of the fresh-water fauna of Korea
are quite incomplete when compared with
those made on the fauna of Manchuria, which
has been clarified by several researches under
the direction of Professor T. Kawamura and
summarized by Yamasaki (1940).

The author, accompanied by M. Ueno,
made a trip for limnological work to the mid-
dle, eastern, and western districts of Korea
from November 19 to 26, 1941. Since return-
ing to the laboratory the author has engaged
in a microscopical study of the rotatorians in
the plankton obtained. At the same rime,
collections made by us in the preceding year
in the northern part of Korea, as well as ma-
terials preserved in our laboratory, were ex-
amined. These latter consist of specimens
which were collected by T. Kawamura during

1 Contribution No. 133 from the Otsu Hydrobio-
logical Station of the University of Kyoto.

^ Otsu Hydrobiological Station, University of Kyo-
to. Manuscript received March 23, 1949.

June, 1936, and by A. Tanaka in Lake Puchon-
ho in 1931.

Through the courtesy of T. Kamita of Kei-
jo Normal School, some plankton samples
collected by T. Sato in the City of Keijo
(Seoul) and the neighborhood were placed at
my disposal for study. The rotatorians found
in all these collections are reported upon in
the following pages. Previous records are in-
cluded in the list.

As the specimens treated here have been
collected not only in a limited season but also
from comparatively confined localities, they
are not sufficient basis for a full discussion of
the rotatorian fauna of Korea. The survey was
to include different seasons and additional
various localities, but the program was of
necessity changed by the outbreak of war.
Difficulties of travel had increased day by day,
and finally the war made continuation of our
survey in this peninsula impossible. There-
fore, this report is based on limited material.

I wish to express my thanks to Doctor
Ueno for his kind advice during the journey
and in the research room.

LOCALITIES

The specimens examined were obtained
from 16 bodies of water, which included all
localities from which there are previous rec-
ords. They are placed in four regions —
northern, middle, eastern, and western — no
collections having been made in the southern
portion of the peninsula. The stations are
shown on the sketch map in Figure 1. Table
1 shows some ecological data for the stations
examined by Ueno and Yamamoto; such data.

151

152

PACIFIC SCIENCE, Vol. VII, April, 1953

Fig. 1. Map of Korean Peninsula showing the five
main regions and the collection localities (1-16).
(Heavy broken line indicates railways.)

are unavailable for the earlier collections by
Tanaka, Kawamura, and Sato.

1. Daitaku (cf. Ueno, 1941) (Tai-taik).

This is a bog lake situated in the moorland

of the Paik-mo high plateau, Ham-Gyon Puk
Do. Pelagic (station \d) and littoral {ih)
plankton were collected. Some microorgan-
isms were also obtained among the mosses of
the surrounding swamp.

2. Lake Fusenko (cf. Ueno, 1941) (Puchon-
ho).

This is a gigantic reservoir for the hydro-
electric plant which has dammed the river
Puchon-gang, an upper tributary of the river
Yalu. The collections by A. Tanaka {2h) from
this lake were obtained during its early stages
as a lake.

3. A small pond at Shochudan in Seoul.

4. A small pool at Shindocho in Seoul.

5. Eitoho, southwestern suburb of Seoul.

6. River Kan-ko (the Hang-gang).

The potamoplankton was collected on the
river in cross section at Tokuson, southeastern
suburb of Seoul.

7. Seiko or West Lake, south of Seoul.

One of the collections examined was made

by T. Kawamura (Jo) 5 years before our visit
{ih). This lake was built artificially about 300
years ago at the foot of Mt. Reiki, imitating
the site of Si-hu in Hangchow, China.

8. Kanggyong.

Station Sa is a pond with luxuriant growth
of water weeds among rice fields. Its surface
area is about 100 square meters.

Station is an oxbow lake on the mean-
dering lower reaches of the river Kum-gang.

9. Lake Kodoho (Kangdong-po).

This brackish lake of medium area is situ-
ated near the coast of the Japan Sea. The
collections were done at three stations, one
being the center of the lake {9a), the next
{9h) near "Jichudai,” an island situated south
of the center, and the last (9c) in the southern
basin.

10. A pond near the Sanjippo railway station.
On the east of the railway station there is

a small pond with clear water in an area sur-
rounded by granite rocks.

11. Lake Sanjippo (Sam-il-po).

This lake is also situated in the granite area
and has very clear water. The collection was
made by using a boat at the southern ex-
tremity of the lake.

12. Lake Kanko (Kam-ho).

This is a small pond which is situated among
pine forests on the coast of the Japan Sea.

13. Inner Diamond Mountain (Kumgang
san).

The plankton was collected by Kawamura
from the small pond of the Choanji Temple.

14. Gan-o-chi.

This small pond of badly contaminated
water is historically famous. It is said that
this pond was built nearly 2,000 years ago in

Rotatoria of Korea — Yamamoto

153

TABLE 1

Ecological Data at Various Collection Stations in Korea

STATION

NUMBER

REGION

HOUR

DATE

AIR

TEMP.

WATER

TEMP.

pH

PER CENT

. CL

°C.

°C.

l{ap)

n

14

15

VII 1940

21.0

19.5

6.3

6.1

2a

N

10

11

VII 1940

21.0

6.8

6

M

10

22

XI 1941

11.2

9.8

lA

7.3

lb

M

14

9

XI 1941

17.6

12.7

6.6

6.7

%a

M

13

24

XI 1941

13.0

11.4

8.6

8.2

Qb

M

14

24

XI 1941

11.9

11.8

7.8

7.8

0.08

9a

E

14

14

XI 1941

15.8

12.0

7.7

7.3

9b

E

14

XI 1941

7.9

7.6

9c

E

16

14

XI 1941

12.4

8.6

8.6

0.97

10

E

9

12

XI 1941

9.8

9.2

6.3

6.3

11

E

10

12

XI 1941 •

15.0

11.5

6.5

6.4

12

E

15

12

XI 1941

15.5

15.8

7.0

7.2

14

E

10

27

XI 1941

9.8

8.7

6.8

6.9

15

E

9

26

XI 1941

5.9

9.2

6.9

7.0

16

W

14

17

XI 1941

18.5

11.7

7.7

7.4

0.02

the garden of the Royal Palace in Keishu
(Kyongju).

15. A pond west of the Bukkokuji railway
Station.

Two kinds of collections were made, one
from near the surface (15^) and the other
from the bottom (15/;^).

16. River Taidong-gang at Phongyang.

The potamoplankton was collected on the

river near Hangetsu-to (Halfmoon Island) in
5 minutes’ upstream sailing.

SPECIES FOUND IN PRESENT MATERIAL

As the result of research on the materials
from the stations mentioned above, 73 spe-
cies (including 8 varieties and 7 forms) be-
longing to 10 families and 26 genera have
been identified. Besides these there were sev-
eral species impossible to identify because of
the strong contraction of the body. Seven
species, which appeared in the previous rec-
ords, could not be found in the present exam-
ination. Thus, at present 83 species are known
to occur in Korea. Although this is not a
large number of species, it compares well with
the number recorded from Japan.

In the following list the distribution within
Korea is indicated by region and station. The
known distribution in other parts of the Far

East is also given. Species marked with an
asterisk are not present in the material studied.

Order FLOSCULARIACEA
Family CONOCHILIDAE

1. Conochiloides dossuarius (Hudson) (M-8(^)
Japan, Manchuria, Sumatra, Bali.

2. Conochilus hippocrepis {Schmnk) (N-1^, lb,
2h\ M-4) Japan, China, Manchuria.

3. Conochilus imicornis Rousselet (M-7^, lb
[Hada, 1936]) Japan, China, Manchuria.

Order PLOIMA
Family Notommatidae

4. Monomfnata grandis Tessin (N-H) Japan,
Manchuria, Sumatra.

‘b.'^Notommata sp. (N-1^).

6. Scandium eudactylotum Gosse (E-15^) Ja-
pan, China, Manchuria.

7. Diaschizza gtbba (Ehrenberg) (N-1^) Ja-
pan, China, Sumatra, Java.

Family Synchaetidae

8. Synchaeta longipes Gosse (M-7^).

9. Synchaeta oblonga Ehrenberg {N-2a; M-lb;
E-9^, 9b, 9c, I5a) Japan.

10. Synchaeta tremula (O. F: Muller) (N-7^;
E-9c) Japan, China, Java.

^Synchaeta spp. (N-2c; E-12, 14).

154

PACIFIC SCIENCE, Vol. VII, April, 1953

W.'^Floesoma hudsoni\T£>^o{ (M- [Hada, 1936])
Japan.

12. Poesomla triacanthum Bergendal (N-l;^)
Japan.

13. Ploesoma truncatum (Levander) (N-2^, 2c;
M-7^, 8^) Japan, Manchuria.

14. Polyarthra euryptera (Wierzejski) (M-7^,
ih, 8^, 8^; W-16) Japan, China, Man-
churia.

15. Polyarthra trigla (Ehrenberg) (N-1^, \h,
2a, 2c; M-3, 5, la, lb [Hada, 1936], 8^;
E-9^, 9h, 9c, 10, 11, 12, 14, 15) Japan,
China, Manchuria, Java.

16. Polyarthra trigla var. major Burckhardt
(M-7^, ih) China.

17. Polyarthra trigla var. remata Skorikov (M-
la, lb) China.

Family Trichocercidae

18. Trichocerca capucina Wierzejski et Zacha-
rias (M-5, la, lb [Hada, 1936]) Japan,
China, Manchuria.

19. Trichocerca cristata Harring (E-9c) Japan,
China, Manchuria.

20. Trichocerca cylindrica (Imhof) (M-7^, lb\
E-9c) Japan, Manchuria.

2\. '^Trichocerca elongata (Gosse) (M-[Hada,
1936; Kawamura, 1916]) Japan, China,
Manchuria.

22. Trichocerca longiseta (Schrank) (N-l^^;
M-8^, 8*^; E-12, 14) Japan, China, Man-
churia.

23. Trichocerca pusilla jtnnmgs {Idl-lb [Hada,
1936]) Japan, China, Manchuria.

24. Trichocerca stylata (Gosse) (M-7/^) Japan,
Manchuria.

25. Diurella brachyura {Gossq) (E-15^) Japan,
China, Manchuria, Sumatra, Java.

26. Diurella porcellus (Gosse) (E-15^) Japan,
China, Manchuria.

27. Diurella stylata Eyferth (M-5, lb-, E-14)
Japan, China, Manchuria, Sumatra, Java.

28. Diurella tigris (O. F. Muller) (N-1^; M-
[Hada, 1936]; E-15^, 16^) Japan, Man-
churia.

29. Diurella weberi jtnmngs (E-15^) Sumatra,
Java.

Family Asplanchnidae

30. Asplanchna priodonta Gosse (N-2^, 2b, 2c;
M-3, 5 [Sato, 1941], la, lb [Hada, 1936],
8^, ^b; E- [Sato, 1941], 10, 13, 15^) Japan,
China, Manchuria. Java.

Family Brachionidae

blABrachionus angularis Gosse (E- [Sato,
1941]) Japan, China, Manchuria, Java.

32. Brachionus angularis var. bidens (Plate)
(M-3, 8^, 8^) Japan.

33. Brachionus budapestinensis Daday (M-3)
Japan, China, Manchuria, Eormosa.

34. Brachionus calyciflorus Pallas (M-8^; E-14)
Japan, China, Manchuria, Java.

35. Brachionus calyciflorus f. amphiceros Ehren-
berg (M-3, 8^) Japan, China, Manchuria.

36. Brachionus calyciflorus f. anuraeiformis
Brehm (M-3, ^a) Japan, China, Manchu-
ria.

37. Brachionus calyciflorus var. dorcas Gosse
(M-3, 8^) Japan, China, Manchuria.

38. Brachionus calyciflorus var. dorcas f. spinosus
Wierzejski (M-3, ^a) Japan, China, Man-
churia.

39. Brachionus forflcula Wierzejski (M-5) Ja-
pan, China, Manchuria.

40. Brachionus quadridentatus (M-3;

E- [Sato, 1941]) Japan, China, Manchuria,
Java.

41. Brachionus quadridentatus var. brevispinus
(Ehrenberg) (E-9c) Japan, China, Man-
churia.

42. Brachionus urceolaris O. E. Muller (E-
[Sato, 1941], 14) Japan, China, Manchu-
ria.

A'bAPlatyias patulus (O. F. Muller) (M-[Hada,
1936]) Japan, China, Manchuriap Java,
Bali.

44. Keratella cochlearis (Gosse) (N-2^, 2b; M-3,
5, 6, la, 7^ [Hada, 1936], 8^, ^b; E-10,
13, 14, I5a, 15b) Japan, China, Manchu-
ria, Java.

45. Keratella cochlearis var. macracantha (Lau-
terborn) (N-la, lb) Japan.

46. Keratella cochlearis tecta (Gosse) (N-2^;

Rotatoria of Korea — Yamamoto

155

M-5, la, Ih [Hada, 1936], 8^, 8^; E-9^,
12, 14) Japan, China, Manchuria.

47. Keratella cochlearis var. tecta f. micracantha
(M-5, la, lb).

48. Keratella cruel formis (Thompson) var. eich-
waldi (Levander) (E-9^, 9h, 9c) Japan.

49. Keratella quadrata (O. F. Muller) (N-1^,
la, 2c; W-16) Japan, China, Manchuria.

50. Keratella serrulata (Ehrenberg) (N-1^)
Japan.

51. Keratella valga (Ehrenberg) f. asymmetrka
Barrois et Daday (M-3, 8^; E- [Sato,
1941]) Japan, China, Manchuria.

52. Keratella valga (Ehrenberg) f. monstrosa
Barrois et Daday (M-3, 5) Japan, China,
Manchuria.

53. Keratella valga (Ehrenberg) f. tropica Ap-
stein (E-9^, 9b, 9c, 12) Japan, China,
Manchuria.

54. Notbolca acuminata Ehrenberg (E-9^, 9c,
12) Japan, Manchuria.

55. Notholca labis Gosse (E-9^, 12) Japan.

%.'^Anuraeopsis^^. (M- [Hada, 1936]).

Family Euchlanidae

57. Euchlanis alata Voronkov (N-l*^) Japan.

58. Euchlanis dilatatu Ehrenberg (E-9^^, 9b, 9c,
12, 14; W-16) Japan, China, Manchuria,
Sumatra.

59. Euchlanis triquetra Ehrenberg (N-1^) Ja-
pan.

60. Lecane brachydactyla (Stenroos) (N-1^, Ic)
Japan.

61. Lecane ludwigii (Eckstein) (E-15^) Japan,
China, Manchuria, Sumatra, Java.

62. Lecane luna (O. F. Muller) (M- [Hada,
1936; Kawamura, 1917]; E-9^) Japan,
China, Manchuria, Sumatra, Java.

63. Monostyla arcuata Bryce (M-Sa) Japan,
China, Manchuria, Sumatra.

64. Monostyla crenata Harring (N-1^) Japan,
China, Manchuria, Java.

65. Monostyla hamata Stokes (N-lc) Japan,
China, Manchuria, Sumatra, Java, Bali.

66. Monostyla lunaris (Ehrenberg) (N-1^, 1^;
M- [Hada, 1936]; W-l6) Japan, China,
Manchuria, Sumatra, Java.

67. Monostyla pyriformis Daday (E-11) Japan,
Sumatra, Java.

68. Lepadella acuminata (Ehrenberg) (E-15^)
Japan, Sumatra.

G9 A Lepadella borealis Harring (M- [Hada,
1936]).

70. Lepadella ovalis (O. F. Muller) (N-1^;
E-12) Japan, China, Manchuria, Sumatra.

71. Colurella adriatica (Ehrenberg) (M-8^;
E-9^) Japan, China.

72. Colurella lepta (Gosse) (E-9^, 9c) Japan,
China.

73. Colurella obtusa (Gosse) (E-9^, 9b, 9c)
Japan, China, Sumatra.

74. Mytilina trigona (Gosse) (N-1^) Japan.

75. Mytilina ventralis (Ehrenberg) (N-1^) Ja-
pan, China, Manchuria.

IG.'^ Mytilina ventralis brevispina (Ehrenberg)
(M- [Hada, 1936]) Japan, China, Man-
churia, Java.

77. Trichotria tetractis (Ehrenberg) (N-1^) Ja-
pan, China, Manchuria, Sumatra, Java.

78. Trichotria truncata (Whitelegge) (M- [Ha-
da, 1936], 8^; E-14, I5b) Japan, Man-
churia.

Family Filiniidae

79. Eilinia longiseta (Ehrenberg) (N-2^, lb\
M-3, 5, 8^, 8^; E-9c, l^a) Japan, China,
Manchuria, Sumatra, Java.

80. * Pedalia mira (Hudson) (E- [Sato, 1941])
Japan, China, Manchuria, Sumatra.

Family Testudinellidae

81. Testudinella bidentata (Ternitz) (N-1^)
Japan.

82. Testudinella patina (Hermann) (E-9^, 9b,
9c, 15^) Japan, China, Manchuria, Suma-
tra, Java.

Order BDELLOIDA

Family Philodinidae

85.*Rotaria sp. (N-lc; E-9^, 10, 15^).

NOTEWORTHY SPECIES

Brief descriptions of some unusual species

which have been observed are given here.

156

PACIFIC SCIENCE, Vol. VII, April, 1953

Euchlanis alata Voronkov, 1912
PL 2, Figs. 26, 26^, 26^, 27

1912 Euchlanis alata Voronkov, Mus. Zook

[St. Petersburg], Ann. 16: 210.

Dorsal plate: length, 335 ix\ width, 315 \x\
frontal width, 160 \x (winged form). Foot,
110 ji\ toe, 18 jj,.

The body consists of two plates, dorsal and
ventral, the dorsal one being much the larger,
about twice as wide as the ventral. No keel
is present on the back. In the winged form,
both sides of the dorsal plate are expanded to
at least two thirds the length. The posterior
tip of the wing is pointed, turned slightly
upward, making the greatest width. The cen-
tral posterior part gradually becomes low and
has no notch. At the anterior extremity a large
V-shaped notch is present. The wingless form
is elliptical.

The ventral plate is slightly convex, elon-
gate elliptical, its posterior border parallel to
that of the dorsal plate.

The foot is large, two-jointed, with one
pair of toes.

A large central, anterior eye-spot is present.

Both the winged and the wingless forms
were collected from the littoral region of Tai-
taik (1/3). They are 20 per cent larger than the
form recorded by Myers (1930). According to
his note, various stages of "wing” develop-
ment were found in this species in the acid
waters of Mt. Desert Island, Maine, but the
forms from Atlantic County, New Jersey, had
no lateral wing-like processes. At that time,
this species had been found in only four
places in the world, i.e., near Moscow, Rus-
sia, Arc-en Barrois (Haute-Marne), France,
and the two localities mentioned above. The
present finding is probably the first record in
the Far East.

Euchlanis triquetra Ehrenberg, 1838
PI. 2, Figs. 28, 28^

1838 Euchlanis triquetra Ehrenberg, Infusions-

thierchen, p. 461.

Length, 220 /x; width, 240 \x.

The dorsal plate is larger than the ventral
plate, is triradiate in cross section, and has a
median keel. The sides are roundly expanded;
in lateral view the highest portion is at the
center, gradually decreasing in height ante-
riorly and decreasing suddenly near the pos-
terior end. The ventral plate is slightly con-
cave, about two thirds as wide as the dorsal
plate. This species was found in the littoral
region of Tai-taik {ih).

Testudinella bidentata (Ternitz, 1892)

PI. 3, Figs. 34, l)Aa

1892 Pterodina bidentata Ternitz, Rotatorien I
der Umgebung Basels, p. 20. |

This species is nearly circular in shape. The
dorsal plate has a convex frontal edge with a
small central notch. On each side of the hind i

ji

portion a small triangular projection is pres-
ent, the tips of which are 100 micra apart. In
the ventral plate the frontal edge is concave, |
but it swells out slightly centrally. A poly-
gonal foot-opening is situated at the posterior
extremity. In cross section (Fig. 34^), the
ventral side is flat, but the dorsal side forms
a low arc. This species was found in the region
off the shore of Tai-taik (1/3). In Japan there
is only a single record from the Koishikawa
Botanic Garden of Tokyo (T. Kawamura).

Ploesoma triacanthum (Bergendal, 1892)

PL 3, Fig. 36

1892 Gastroschiza triacanthum Bergendal, Univ.
Lundensis, Acta 28: 1.

The anterior end of the head is separated
into three triangular processes of which the
central one is the longest. In general, this
portion bends downward as a shield-like head.
This species was found only in the littoral
region of Tai-taik (1^).

Keratella cruciformis (Thompson, 1892)
var. eichwaldi Levander, 1894
PL 1, Fig. 14, 15

1892 Anuraea cruciformis Thompson, Liver-
pool Biol. Soc., Proc. 6: 77.

Rotatoria of Korea — Yamamoto

157

1894 Anuraea ekhwaldi Levander, Soc. Fauna

Flora Fennica, Acta 12: 62.

Total length, 120 /r.

This brackish-water species was found in
every part of Lake Kandong-po (9^, h, c).
The hind process is absent. Both the punctate
and reticular sculptures are remarkable. In the
dorsal lorica, the frontal plate is very short
c. 10 m), but the first carinal plate is remark-
ably long (c. 75 m)-

Keratella serrulata (Ehrenberg, 1838)

PI. 2, Fig. 20

1838 Anuraea serrulata Ehrenberg, Infusions-

thierchen, p. 508.

Total length: 195 m (from the frontal notch
to the posterior end, 130 \x, length of the
ventral plate, 120 yl). Width: frontal end 98
y, greatest, 112 y, hind end, 54 y\ frontal
processes: central, 65 y, sublateral, 22 y, later-
al, 25 y.

Both the punctate and reticular sculptures
are remarkable in the dorsal plate. 'The last
central plate is hexagonal, and from its pos-
terior corner a central line runs posteriorly.
Both posterior corners project slightly. The
central pair of the frontal processes are long
and stout, bending downward remarkably
near the tip. This is a characteristic species of
dystrophic water. Skadowski (1923) has found
this species in moorland waters, in which the
pA values ranged from 3.2 to 4.5. The pA
value of the present locality (1^) measured as
high as 6.3.

DISTRIBUTIONAL RELATIONSHIPS

As already noted, in the western region
only a collection of potamoplankton was
made, on the river Taidong-gang (16), where
only five rotatorian species were found. This
number is not small for a collection from a
river, but there are very few individuals of the
species, four of which were found in three
other regions. Therefore, the fauna of the
western region should not be considered here.

The species common to the three other re-
gions number eight, or less than 10 per cent.

In general, these species are well known to be
widely distributed animals. They are all pe-
lagic except Diurella tigris. The frequencies of
occurrence are as follows:

Polyarthra trigla

12 stations

Keratella cochlearis

10 stations

Keratella cochlearis var. tecta

7 stations

Asplanchna priodonta

9 stations

Filinia longiseta

6 stations

Synchaeta ohlonga

4 stations

Trichocerca longiseta

4 stations

Diurella tigris

3 stations

The number of species peculiar to any one

region of northern, middle, or eastern Korea
are 17, 25, and 19, respectively. The total pe-
culiar to one region, 61, is three fourths of the
total number. Of the 13 species remaining, 8
are found both in the middle and eastern re-
gions, 3 others are found both in the northern
and middle regions. One species {Lepadella
ovalis) is found in both the northern and the
eastern regions.

The collections in northern Korea were
made chiefly in summer. Seventeen species
obtained from this region are, without excep-
tion, found only from Tai-taik. Only one
from Tai-taik, Keratella quadrata, is also found
in the collection from Puchon-ho, and in the
Taidong-gang of the western region. This
suggests that the conditions of Tai-taik differ
greatly from those of any other waters visited.
The occurrence of Keratella cochlearis var. ma-
cracantha is worthy of note as this collection
was made in midsummer and this has been
considered heretofore to be a winter form.

In Tai-taik 23 species of Rotatoria were
found, i.e., 8 from the pelagic, 18 from the
littoral, and 3 among the mosses of the mar-
ginal region.

In a work on the plankton of Lake Puchon-
ho, the present author (1941) has noted that
at the early stage of the development of the
reservoir many heleoplankters such as Cono-
chilus hippocrepis were found, but after several
years the condition of the water so changed
that Synchaeta spp. and Keratella quadrata ap-

158

Q C.v. tectso O VaJ,^^t-.wo»i5tiro5a.

P C. f .mbrdxacntha^ Q vaLgd,f tropica^

X) c. ^/.m^£.r^y0^^K 0 serrulate

Fig. 2. Occurrences of the genus Keratella in Korea.

peared. In July, 1941, Asplanchna priodonta
was the predominant species, being especially
abundant in the epilimnion. The nine species
reported from this reservoir, excepting Kera-
tella quadrata, are common to the middle and
eastern regions.

In the middle region, three of the samplings
were made by Sato from localities in and
around Seoul. Among them, one from a pool
at Shindo-cho (4) contained only a single
species, Conochilus hippocrepis. The collections
from a pool at Shochudan (3) and from Ei-
toho (5) contained 11 species each. A remark-
able condition common to these two localities

PACIFIC SCIENCE, Voi. VII, April, 1953

is the richness of species belonging to the
family Brachionidae, i.e., eight from the for-
mer and six from the latter. It is particularly
noteworthy that species of a warm-water type
are found here, e.g., Brachionus forficula in the
latter and Keratella valga in both localities.

As potamoplankton of the Hang- gang (6),
only a single species {Keratella cochlearis) was
found, but Sato (1939) has reported two spe-
cies, Asplanchna priodonta and Testudinella pa-
tina, from the same river in winter.

Besides our collection from Seiko (7), those
taken by Kawamura on June 2, 1936 (7^),
were examined. Notwithstanding the fact that
the season of collection was quite different,
11 species found in early summer were all
included among the 18 species collected in

Fig. 3. Occurrences of the genus Polyarthra in Korea.

Rotatoria of Korea — Yamamoto

159

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

Plate I

(Line scales represent 100 micra except those marked otherwise)

1. Polyarthra euryptera Fig. 9. Brachionus forficula

2. Polyarthra trigla Fig. 10. Brachionus calyciflorus

3. Polyarthra trigla v2lT. remata Fig. 11. Brachionus quadridentatus

4. Polyarthra trigla var. major

5. Conochiloides dossuarius

6. Brachionus angularis

7. Brachionus angularis (type)

8. Brachionus budapestinensis

Fig. 12. Keratella quadrata

Fig. 13. Keratella valga f. tropica

Fig. 14. Keratella cruciformis var. eichwaldi

Fig. 15. K. cruciformis var. eichwaldi (from Sakhalin,

after Yamaguchi)

Fig. 16. Keratella cochlearis var. macracantha

160

PACIFIC SCIENCE, VoL VII, April, 1953

late autumn. In both cases Asplanchna priodon-
ta and Keratella cochlearis were found abun-
dantly. The length of the posterior process of
the latter species varies greatly, showing stages
almost continuous from to typica-

type (Fig. 2). Polyarthra trigla shows a very
wide distribution (Fig. 3). In the specimens
from Seiko, two varieties were included (var.
major and var. remata).

Of 17 species reported by Hada (1936)
from this lake, 7 were also obtained in the
present survey. They belong chiefly to the pe-
lagic species, while the remainder are those
of the creeping type. The occurrence of such
bottom-inhabiting forms may be attributed
to vertical hauls. The total number of species
recorded from this lake is 28. This is the
greatest number of species found in any single
locality in Korea.

Seventeen species were found in two collec-
tions from Kanggyong. One collection from
a small pond (8^) contained 13 species. Of
these, Brachionus calyciflorus shows great vari-
ability, many types such as typica, amphiceros,
amiraeiformis, dorcas, and spinosus being iden-
tified. Another collection from a pond (8i^)
contained 10 species, of which 7 are also
found in pond 8^?. No remarkable morpholog-
ical characteristics can be found.

In the eastern region, the greatest number
of species was found from Kangdong-po
(9^, b, c). The total here is 23, if Sato’s (1941)
record is added. The present collections were
made from a small boat at three stations on
the lake. Notholca occur in all three stations,
Notholca acuminata in the first and third sta-
tions, and Notholca lahis in the second. These
species have been known as brackish-water
forms and were also found in Lake Kam-ho
(12).

Of eight species found in Kam-ho, six are
also found in Kangdong-po.

The remaining two localities are located in
the far southern region of eastern Korea, near
the coast of the Japan Sea. An unnamed pond
is situated to the west of the Bukkokuji rail-
way station (14). Its area is considerable and

it receives several brooks from the rice fields
around it. Eight species were found both in
the collections from the surface and in those
from the bottom, the total being 14. There
are two common species in these two places,
one pelagic and the other a creeping species
— Keratella cochlearis and Diurella tigris.

Throughout all the regions surveyed, the
species belonging to the genus Keratella occur
most frequently. The features of its distribu-
tion seem to be characteristic. Keratella coch-
learis in both forma typica and var. tecta is
distributed in the northern, middle, and east-
ern regions, whereas the remaining forms
seem to be limited to a single region. K,
quadrata, K. serrulata, and K. cochlearis var.
macracantha are distributed chiefly in the
northern region, K. cochlearis f. micracantha,
K. valga f. asymmetrica, and f. monstrosa in the
middle, and K. cruciformis var. eichwaldi and
K. valga f. tropica in the eastern region. K.
quadrata was, however, found also in the
western region, and its forma asymmetrica has
been recorded by Sato (1941) as a summer
plankton animal of Kangdong-po.

It is remarkable that the distribution of spe-
cies of southern origin, such as K. valga f.
tropica, have shown a late autumn occurrence
in the district of the Japan Sea coast. This
suggests that the influence of the northward-
flowing warm current along the coast of
North Korea upon the climate of this district
may be one of the factors acting in the distri-
bution of this species. It is also interesting
that, in spite of the collections having been
made in the northern region in summer and
in the southern region in late autumn, K.
quadrata appears to be distributed strictly in
the northern part and K. valga only from the
southern region (Fig. 2).

Species belonging to the genus Brachionus
occurred in comparatively few places. Two of
theso localities were the stations in and about
the city of Seoul where the collections were
made by Sato. In the present survey, Brachi-
onus species were found in only three locali-
ties, i.e., Kanggyong, the Kangdong-po, and

Rotatoria of Korea — Yamamoto

161

Fig.

17.

Keratella cochlearis var. tecta

Fig. 23.

Colurella lepta

Fig.

l^a-c.

Keratella cochlearis f. micracantha

Fig. 24.

Colurella obtusa

Fig.

l^d.

Keratella cochlearis s. str.

Fig. 25.

Lepadella acuminata {a; side view)

Fig.

19.

Keratella valga f. monstrosa

Fig. 26.

Euchlanis alata {a: side view; b: frontal

Fig.

20.

Keratella serrulata

view)

Fig.

21.

Notholca acuminata

Fig. 27.

Euchlanis alata (wingless form)

Fig.

22.

Notholca labis

Fig. 28.

Euchlanis triquetra {a: cross section)

162

PACIFIC SCIENCE, Vol, VII, April, 1953

(3 buJApestLnensls ^ (^uAdrldentatus
cadyclfloTU-s w-TceolstTls

Fig. 4. Occurrences of the genus Brachionus in Korea.

the Gan-o-chi. In Gan-o-chi, two species of
Brachionus {B. urceolaris and calyciflorus) oc-
curred, whereas in Kangdong-po only a sin-
gle species {B. quadridentatus var. hrevispinus)
was found. At Kanggyong two species {B.
angularis var. bidens and B. calyciflorus) oc-
curred; five forms of B. calyciflorus were ob-
served in the pond along the railroad (8^) as
previously mentioned.

The values of the hydrogen ion concentra-
tion of water in these places indicate that,
although the range tolerated by this genus
would be rather wide, the organisms prefer
alkaline waters. This seems also to correspond
to conditions in Korean waters, though the

occurrences of this genus are limited to fewer
localities. As is mentioned above, none of
them was found in the northern region. As
far as our collections are concerned, one spe-
cies was found from the middle and two from
the eastern region, a fourth was common to
both. Besides these, two {hudapestinensis and
forflcula) were found in the collections made
by Sato. They are warm-water forms. Thus, a
total of six species of Brachionus is known
from the middle and eastern regions (Fig. 4) .

SUMMARY

1. Plankton collections were made in Ko-
rea at 16 places, which may be classified
roughly into four regions — the northern,
western, middle, and eastern.

2. Fifty-eight species of Rotatoria were
identified in these collections. In addition,
seven species can be added from previous
records. With the addition of eight varieties
and seven forms, plus three undeterminable
species, the total number becomes 83.

3. The species of wide distribution are Ke-
ratella cochlearis and Polyarthra trigla, both in-
cluding several varieties or forms.

4. Species of Brachionus were found only
in a few stations.

REFERENCES

Ahlstrom, E. H. 1940. A revision of the
rotatorian genera Brachionus and Platyias
with descriptions of one new species and
two new varieties. Amer. Mus. Nat. Hist.,
Bui. 77: 143-184.

Carlin, B. 1943. Die Planktonrotatorien des
Motalastrom. Lund Univ., Limnol. Inst.,
Meddel. 5: 1-255.

Collin, A., H. Dieffenbach, R. Sachse, and
M. Voigt. 1912. Rotatoria und Gastrotri-
cha. In Brauer: Die Susswasser fauna Deutsch- ,
lands. Vol. 14. 274 pp. Gustav Eischer, Jena.
Edmondson, W. T., and G. E. Hutchinson.
1934. Report on Rotatoria. Yale North In-
dia Expedition. Conn. Acad., Mem. 10: 153-
186.

Rotatoria of Korea — YAMAMOTO

163

Fig. 29. Lepadella ovalis

Fig. 30. Lecane brachydactyla

Fig. 31. Monostyla lunarh

Fig. 32. Monostyla furcata

Fig. 33. Monostyla crenata

Fig. 34. Testudinella bidentata {a: cross section)

Fig. 35. Testudinella patina

Fig. 36. The soma triacanthum

Fig. 37. Diurella tigris {a: head; b: toes)

Fig. 38. Diurella porcellus
Fig. 39. Trichocerca cristata
Fig. 40. Trichocerca capucina
Fig. 41. Trichocerca pusilla
Fig. 42. Trichocerca longiseta
Fig. 43. Trichocerca cylindrica (head)
Fig. 44. Scaridium eudactylotum
Fig. 45. Monommata grandis

164

PACIFIC SCIENCE, VoL VII, April, 1953

Hada, Y. 1936. (The plankton of Lake Seiko,
Suigen in autumn.) Chosen Nat. Hist. Soc.,
Jour. 21: 1-11. [In Japanese.]

1938. Rotatorian fauna of Manchou-

kou. Sapporo Nat. Hist. Soc., Trans. 15:
171-186.

Harking, H. K. 1913. Synopses of Rotatoria.

U. S. Natl. Mus., Bui. 81: 1-226.

Hauer, J. 1938. Die Rotatorien von Sumatra,
Java und Bali nach den Ergebnissen der
Deutschen Sunda-Expedition. Arch. f. Hy-
drohiol., Suppl. 15: 297-384, 507-602.
Hudson, C., and P. Gosse. 1886-89. The
Rotifera or wheel- animalcules . 312 pp. Long-
mans, Green & Co., London.

Kawamura, T. 1917. Freshwater biology of
Japan. 2 vols., 579 pp. Shokabo, Tokyo.
[In Japanese.]

Myers, E. J. 1930. The rotifer fauna of Wis-
consin. V. The genera Euchlanis and Mo-
nommata. Wis. Acad. Sci., Arts, Letters, Trans.
25: 353-413.

Rylov, W. M. 1935. Das Zooplankton der
Binnengewasser. Die Binnengewdsser. Vol.
15. 272 pp. Schweizerbart, Stuttgart.

Sato, T. 1939. Potamoplankton of the River

Hang-gang in early winter. Jap. Jour. Lim-
nol. 10: 128-130. [In Japanese.]

1941. Biological notes on the brackish

water lake Kodoko on the Japan Sea coast
of Korea. Jap. Jour. Limnol. 11: 85-88.
[In Japanese.]

SCHONICHEN, W. 1917. Einfachste Lehensfor-
men des Tier und Pflanzenreiches. Vol. 2.
Urtiere, Radertiere. 522 pp. Hugo Ber-
muehler, Berlin.

Ueno, M. 1941. Introductory account of the
biological survey of inland waters of north-
ern Chosen [Korea]. Jap. Jour. Limnol. 11:
96-107. [In Japanese.]

Yamamoto, K. 1941. The plankton of lake
Fusenko of northern Chosen [Korea]. Jap.
Jour. Limnol. 11: 108-117. [In Japanese.]

— ■ 1944. The Rotatoria from the fresh

waters of the Yangtse delta. Reports on the
limnological survey of central China. XVII.
Shanghai Sci. Inst., Jour. 14: 90-117. [In
Japanese.]

Yamasaki, M. 1940. Freshwater Rotatoria of
Manchoukuo. Report of the limnological sur-
vey of Kwantung and Manchoukuo. T. Kawa-
mura, ed. 573 pp. Government of Kwan-
tung Prov., Dairen. [In Japanese.]

A

Vernacular Names of the Plants of Bikini, Marshall Islands
Pacific Plant Studies 12^

Harold St. John and Leonard E. Mason ^

In 1946, in connection with the atomic bomb-
ing at Bikini, "Operation Crossroads,” gen-
eral scientific studies were made upon this
coral atoll in the northern Marshall Islands,
in the central Pacific Ocean. In the plant
kingdom general collections were made, even
of the bacteria and the phytoplankton. The
remaining plant groups were collected in 1946
by Dr. W. R. Taylor, a specialist on marine
algae, and he studied and reported on most
of them. His general account of the flora has
now appeared in book form (Taylor, 1950).
In this book he includes 41 species of angio-
sperms as composing the terrestrial flora of
Bikini. Taylor did not obtain a record of the
native names of the plants in use by the
people of Bikini, who had already been re-
moved to Rongerik atoll, since their home
island was to become dangerously radioactive.
They went first to Rongerik which, after 2
years of residence, proved inadequate, then
temporarily to Kwajalein until a final reloca-
tion was made in November, 1948, at Kili
Island in the southern Marshalls.

Appreciating the value of the Marshallese
ethnobotany, St.John urged Mason, who was

^ This is the twelfth in a series of papers designed
to present descriptions, revisions, and records of Pacific
island plants. The preceding papers were published as:
Bernice P. Bishop Mus., Occas. Papers 17 (7), 1942; 17
(13), 1943; 18 (5), 1945; Amer. Fern Jour. 35: 87-89,
1945; Torrey Bot. Club, Bui. 73: 588, 1946; Pacific Sci.
1(2): 116, 1947; 2(2): 96-113, 1948; 2(4): 272-273,
1948; 5(3): 279-286, 1951; 6(2); 145-150, 1952; Web-
bia 8: 225-228, 1951.

2 Chairman, Department of Botany and Chairman,
Department of Anthropology and Sociology, Univer-
sity of Hawaii, respectively. Manuscript received May
10, 1951.

about to visit Kili and the Bikini people on an
inspection trip, to attempt to compile it from
native informants. On August 14, 1949, Ma-
son spent 2 hours collecting plants, beginning
at the village on the north shore, following
the beach westward to the end of the island,
and then cutting back to the village through
the interior of the island. The plants collected
and here listed are those recognized as char-
acteristic Bikini plants by Jibaj, Ejkel, and
Joaj, male adults and former residents of
Bikini, who were selected because of their
knowledge of Bikini’s plant life. Seventeen
plants common to Kili and Bikini were col-
lected, and they have been identified by St.
John and deposited in the Bishop Museum.
The informants were in unanimous agreement
about the naming of 16 of them, but one man
disagreed on the spelling of a single vernacu-
lar name, that for Eleusme indica, although the
difference in their spelling and pronunciation
was slight and was due to the inadequacies of
the orthography used by the Marshallese in
spelling their language.

The orthography here used is that estab-
lished by the Protestant Mission and pub-
lished in their Bible in Marshallese, printed
in 1885. Vowel pronunciation is as follows:
a, e, o, and u are like those in Spanish; a, as
in English hat; i, as in bit; o is a central vowel
somewhat like the u in English cut but higher
in the mouth. Frequently certain vowels are
preceded by a definite w sound. The conso-
nants t and d are spelled as t, but when placed
medially in a word are pronounced more like
d; the same applies to the pair k and g, k

165

166

PACIFIC SCIENCE, VoL VII, April, 1953

being written to stand for both, though pro-
nounced more like g when placed medially.
The letter g is used only with the sound ng
which the Marshallese spell as h.

VERNACULAR NAMES USED BY NATIVES
OF BIKINI

'Pandanus sp. ’’Bob anbwiri”; a second variety
"bob erwan." (Because the specimens of
the fruits were lost in transit, specific identi-
fication could not be made.)

Eleusine indica (L.) Gaertn. "Ujuj” (or "ujoj").
Although this plant was not collected by
Taylor on Bikini, it is common on many of
the Marshall Islands, and this report for
Bikini is trustworthy. "Ujuj" is a general
name for a number of weeds, mostly grasses.
This one may be good in treatment of
diarrhea.

Cocos nucifera L. "Ni." Very generally useful
to the Marshallese.

Tacca Leontopetaloides (L.) Ktze. "Mokmok."
It is good in the treatment of diarrhea when
eaten by itself, or as food when mixed with
coconut cream, Pandanus fruit pulp, sugar,
or toddy. The seed (fruit) is wrapped in a
piece of coconut cloth, "inubil," and used
as a scent-stimulant by pregnant women
experiencing morning-sickness.

Fleurya ruderalis (Forst. f.) Gaud. "Nen kot-
k5t.” It is used only for medicine. The
leaves, root, and stem are crushed, the juice
thus obtained being mixed with water and
coconut milk. The potion is given to a
person who is distressed and yelling or cry-
ing, in order to make the evil spirit go away.
This is a new record for Bikini, as Taylor
did not find it there. However, it is not
unexpected as it is known on Eniwetok and
is widespread in the Marshalls.

Pisonia grandis R. Br. "Kangdi." A tree useful
for house timbers, especially posts, because
the wood lasts a long time.

Triumfetta procumhens Forst f. ' ' Atat. ’ ’ A ground
creeper, said to be the most valuable vine
in Bikini. The bark is stripped off, then, by
fretting in the hands, the easily broken

outer bark is removed. The smooth, fibrous
band of inner bark is dried in the sun to an
orange-brown color of varying shades and
used as a fiber to provide ornamentation in
finely woven Pandanus matting. The leaves
are crushed and placed as medicine on open
wounds. Sometimes the leaves, yellow and
brown, and the woody part of the stem are
mashed, giving a liquid used as a potion
for treatment of gonorrhea.

Hibiscus tiliaceus L. "Lwo” (pronounced like
the English "law"). A tree, the wood of
which is light, for which reason it is used
for masts and booms of sailing craft and
for pieces of the outrigger frame of canoes.
The inner bark is used for handicraft fiber,
for cord, or for string.

Pemphis acidida Forst. "Konge." A tree, the
hard wood of which is used for the canoe
keel, for the apex of the canoe mast, and j
for the pieces fitted into the canoe bow and |
stern where the sail booms are lashed. It is |
also used for the husking stake ("ton"), I
for the digging stick to uproot Tacca, and |
for a beater for pounding soaked coconut i
husks when making sennit ("eokwal").

Terminalia litoralis Seem. "Ekong," A large
tree, the wood of which is used for making
the elongate bowls in which the "beru"
(fruits of Pandanus and tubers of Tacca) and
the "bwiru" (preserved Artocarpus fruits) I
are mixed. The leaves and root are used for
medicine.

Ipomoea tuba (Schlecht.) Don. "Mar b5le." I
A vine which is regarded as harmful, since ||
it kills any tree on which it grows. It has
small roots (tubers as small as marbles)
which long ago were used for food. After
being cooked overnight in the "umum"
(underground oven), they were eaten like
a potato. Previous records from the Mar-
shalls have been called 1. grandiflora or 1.
alba, but these were misdeterminations.

Cordia subcordata Lam. "Kono." A tree with
red flowers that are used in making leis.
The wood is used for many purposes in
Bikini where the tree occurs in great abun-

Plant Names of Bikini — St. John AND MASON

167

dance, more so than elsewhere in the Mar-
shalls. The wood is used for making the
block on which Pandanus or Cocos fiber hats
are plaited, for the hull, bow and stern
pieces, and interior hull braces of a canoe,
and for paddles. Recently it has been used
for many tourist souvenirs, such as out-
rigger canoe models, bowls, cups, and
clogs.

Messerschmtdia argentea (L. f.) I. M. Johnston.
”Kiren.” A tree which, if large, is used for
canoe hulls, if small, for the curving booms
of the outrigger. The bark is useful. First
the dry outer bark is scraped off and re-
moved, then the inner bark is scraped,
making wet shavings that are good for
placing on a cut. The young leaves are
pounded to obtain a juice which is good
to drink for a stomach-ache or for any
general ache in the body. The seeds are
pounded to obtain a juice which is applied
to gum boils or canker sores in children’s
mouths; it is also good for treatment of
diarrhea.

Guettarda speciosa L. '’Wut.” A tree, the wood
of which is useful for house frames, house
posts, outrigger booms, and the diagonal
braces between the outrigger and the canoe
hull. When large, it is also used for canoe
hulls. The leaves are medicinal.

Morinda citri folia L. "Nen.” A tree, the wood
of which is used for the framework of a
house. The leaf, root, and fruit are strongly
medicinal. The seeds are pulverized and the
pulp applied to any tooth which is causing
pain. The root is scraped, then the scrap-
ings, together with young coconut husk,
are boiled to make a light-brown dye for
handicraft fibers, such as Triumfetta, Hibis-
cus, and young coconut leaf, but not for
Pandanus.

Scaevola frutescens (Mill.) Krause. "Konnat.”
A shrub or small tree, the wood of which
is used for the curving booms of the out-
rigger; also for the roof frame of a house.
The leaf is used to make a bitter drink
which is given to a pregnant woman just

before and just after childbirth (''to clean
up the belly inside”) ; it is also used to wash
the mother’s body.

Mason also obtained notes on other species
not collected on the excursion but known to
be common to Kili and Bikini. Since they are
known by the same vernacular names on the
other Marshall Islands, it is possible to iden-
tify them with surety. These are:

Artocarpus incisus L. f. "Ma.” Two varieties
exist: "betaktok,” the fruits of which are
large and seedless and make excellent food;
and "mejwan,” the fruits of which are seed-
bearing.

Hernandia ovigera L. ’'Bing bing.” A tree, used
for canoe hulls.

The three Bikinian informants also told of
eight other plants characteristic of Bikini but
not occurring on Kili. For each they furnished
the vernacular name and a description. By use
of these we have identified six of them. They
are:

Portulaca samoensis v. Poelln. "Bujang.” ''A
low bush.” In Taylor’s book this is listed
as P. quadrifida L., the valid name of an
African species, but we consider it a mis-
identification of the Pacific species.

Cassytha frliformis\^. "Kanon.” "A leafless vine
which kills the trees on which it grows.”
Sophora tomentosa L. "Kille.” "A tall tree,
about 15 to 20 feet high.”

Suriana maritima L. "Kalange.” ''Somewhat
like 'konge’ {Pemphis acidula) but smaller.”
Euphorbia Atoto Forst. f. "Berrol.” "A low
bush, with small leaves, which when cut
exudes a milky juice.” Taylor (1950: 3)
stated that members of the Euphorbiaceae
do not occur on Bikini or the northern
Marshalls, but this species and three other
members of the family are now known to
occur on Eniwetok, so this report is credible.
Sida fallax Walp. "Kio.” ''A bush with a
yellow flower.”

Of the list of eight species named and de-
scribed, there remain two which we have not
identified:

168

"Kora.” "A bush about two feet high, with
leaves like ’nen kotkdt’ (Fleurya ruderalis),
but larger.”

"Kaliklik.” "A thorny bush.” The name "kai-
liklik” is regularly used in the Marshalls for
a cultivated variety of Cyrtosperma Chamis-
sonis that has prickly petioles, but that does
not seem to be the right application of it
here. The vernacular name means thorny or
prickly. On Bikini this name might be ap-
plied to Achyranthes canescens, which is prick-

PACIFIC SCIENCE, Vol. VII, April, 1953
ly enough to deserve the description.

REFERENCES

Anonymous, 1885. Bible: New Testament: ak
Kalimur Ekdl an ar Iroij im Dri Lomor Jisds
Kraist. American Bible Society, New York.

Taylor, W. R. 1950. Plants of Bikini and other
northern Marshall Islands. University of
Michigan Studies, Sci. Ser. 18. xv + 227
pp., 79 figs. Ann Arbor.

I

The Relationship Between Body Length and Scale Length in Five
Year-Classes of the Pacific Pilchard or Sardine,
Sardinops caerulea (Girard, 1854).

Antonio Landa’

INTRODUCTION

By correlating the marginal growth of
scales (the amount of growth between the
last ring or annulus and the margin of the
scale) with the season of the year in which
the scales were collected, Walford and Mosher
(1943^: 9 and 1943^: 12) have shown for the
Pacific pilchard or sardine, Sardinops caerulea
(Girard) 1854, that these rings are formed
annually and consequently can be used for
age determination as well as for back-calcula-
tion of the length of the fish at a given earlier
age. The application of the scale rings to the
problem of age determination in the pilchard
has been thoroughly discussed in the papers
cited above. The application of the same phe-
nomenon to back-calculation of lengths,
which lengths are used in several different
types of problems, requires further considera-
tion. The literature on this subject is volumi-
nous and of considerable interest. Reference
should be made to the works of Blackburn
(1949), Buchanan-Wollaston (1934), Lea
(1910, 1913), Lee (1912), Ottestad (1938),
and Schuck (1949) for further information.

The estimation of body length at a previous
age, using length of the scale from its center
to the ring corresponding to that age, in-
volves the regression of body length on scale
length. The regression of scale length on body
length, sometimes used, will give the mean
scale length corresponding to a given body
length, information which is not pertinent to

1 Biologist, Cia. Admora del Guano, Lima, Peru.
Manuscript received December 4, 1951.

the problem. It should be noted at once that
different regression lines could be obtained
from a given set of material depending on the
manner of sampling and that this manner, in
turn, depends on the use for which the re-
gression values are intended. Thus, if the goal
is to know at which body length the scales
start growing, it should be emphasized that
the sampling be comprehensive of the very
small body lengths. Moreover, such sampling
should be equally representative of all the
body lengths if a further goal is desired, i.e.,
to know whether or not the regression of body
length on scale length is the same throughout
all different body lengths.

In the particular case of the Pacific pilchard
investigations, and, presumably, in others of
similar commercial fisheries, the goal is to
know the body lengths that the fish caught
commercially had in previous years. It follows
that the sampling should, before anything
else, be representative of the commercial catch
even if that requirement makes the data un-
suitable for the attainment of goals of the
type indicated above. Once the scope of the
problem is thus limited, the following are
some of the questions that arise in the back-
calculation and use of estimated lenghts:

(1) Are the regression constants (regres-
sion coefficient and body intercept) of
body length on scale length the same
for all year-classes regardless of region
of capture?

(2) Are the regression constants for a given
year-class the same regardless of region
of capture?

169

170

PACIFIC SCIENCE, Vol. VII, April, 1953

(3) Can the regression of body length on
scale length be expressed by a straight
line when fish of successive ages are
considered?

(4) If the regression is linear, does it have
its origin at 0.0? (If so, the actual pro-
cess of calculating lengths at previous
ages may be greatly simplified.)

Answers to these four questions are neces-
sary in order to decide if, for back-calculation
of lengths, a given body of data should be
treated as a unit or grouped by year-classes,
by regions in which the fish were captured,
by age groups, or by groups combining two
or three of these categories.

Acknowledgments

I wish to thank Mr. John C. Marr, Dr.
Frances E. Felin, and Mr. Theodore M. Wid-
rig of the U. S. Fish and Wildlife Service for
making the material used in this study avail-
able to me and for other assistance.

SOURCE OF DATA

Material with which to investigate these
questions consists of records of scale readings
for the Pacific sardine on file at the laboratory
of the South Pacific Fishery Investigations,
U. S. Fish and Wildlife Service, at Stanford
University. The material was obtained as a
result of a co-operative program between the
U. S. Fish and Wildlife Service, the California
Department of Fish and Game, the Fisheries
Research Board of Canada, the Washington
State Department of Fisheries, and the Fish
Commission of Oregon. The methods of col-
lecting samples, examining scales, and record-
ing data are explained by Felin and Phillips
(1948). In brief, each fish is represented by a
card which contains information on fish
length, sex, date and locality of capture, and,
on a printed scale, the relative position, of the
focus, annuli, and margin of the fish scale.
The data are, in general, representative of the
populations commercially fished along the
Pacific Coast of the United States and Canada
(with the exception of bait fisheries in south-
ern California).

From the material available (about 60,000
cards), the data for the 1938 through 1942
classes for fish caught off Canada, Washing-
ton and Oregon, San Francisco, Monterey,
and San Pedro during the fishing seasons of
1939-40 through 1947-48 were selected in
order to have (1) a series of year-classes repre-
sented, (2) all the commercial ports repre-
sented (3) for each year-class, fish with one to
five annuli represented.

METHODS

To reduce this material (about 40,000 cards)
to workable proportions and still have a repre-
sentation of the fish taken by the commercial
fishery, a sample was taken in the following
manner:

(1) All the cards belonging to the material
selected were considered as a single
"population” without regard to year-
class, port of landing, age, or season
of the year in which the fish were
caught.

(2) One thousand one hundred and ninety-
nine bibliographic cards (about 3 per
cent of the above population) were
marked with numbers taken from a ran-
dom nurhbers table (Table 1 in Dixon
and Massey, 1949) using five-digit num-
bers. The cards were arranged in as-
cending order. Three percent was cho-
sen because, after some trials, it was
seen to be the minimum number of
cards likely to include every category
in the sample.

(3) The scale reading cards were counted,
and, when the count coincided with the
number on the first bibliographic card,
the latter was removed, and the corres-
ponding scale card was taken as part
of the sample. This process was re-
peated, without interrupting the orig-
inal count, until all the bibliographic
cards were removed.

(4) The sample so obtained was then sub:
divided into all possible year-class and
port combinations. By chance, the

Body Length and Scale Length in Pacific Pilchard — Landa

171

TABLE 1

Coded Values Taken from Scatter Diagrams
(Y = Body Length; X = Scale Length)

YEAR-CLASS

AND PORT

COMBINATION

nj*

Y't

Y'2

XT

X'2

X'Y'

1938 in:

Pacific

Northwest. . .

10

-29

1015

14

88

91

San Francisco . .

91

-16

4514

79

843

1138

Monterey

76

269

4371

26

640

938

San Pedro

60

-26

1692

-32

374

549

1939 in:

Pacific

Northwest. . .

43

138

5472

-33

349

807

San Francisco . .

150

1079

27721

16

2302

5388

Monterey

166

640

15716

-63

1541

2620

San Pedro

124

-128

4926

54

716

1065

1940 in:

Pacific

Northwest . . .

14

25

1681

22

45

96

San Francisco . .

50

-12

2120

-4

368

430

Monterey

75

487

12303

93

1111

2639

San Pedro

70

-109

3823

-23

535

1091

1941 in:

San Francisco . .

5

-4

102

3

37

15

Monterey

26

125

3171

42

274

726

San Pedro

90

153

2923

90

484

809

1942 in:

Pacific

Northwest. . .

7

61

1415

6

84

270

San Francisco . .

23

16

1531

6

192

204

Monterey

68

491

12327

159

1283

3464

San Pedro

51

-6

4486

10

502

1308

*nj = number of items in jth group,
fY' = sum of coded body lengths.

J X'= sum of coded scale lengths.

1941 year-class in the Pacific North-
west did not appear in the sample.

For* each year-class and port combination
a two-variable frequency table was made, plot-
ting standard body lengths against scale
lengths. Body lengths were grouped by 2-
millimeter intervals, scale lengths by 6-milli-
meter intervals. In order to code, arbitrary
means were independently chosen for each of
the variables in every frequency table. Owing
to the magnification used in reading the
scales, the scale-length values are 30 times
larger than the actual scale lengths (Felin and
Phillips, 1948).

The values obtained from these frequency
tables are given in Table 1. It is possible to
pool these values in such a way as to obtain
combinations other than port and year-class
combinations. For example, to obtain infor-
mation about the 1938 class as a unit, the
sums of X' for the 1938 class from all ports
are pooled in a single sum, the sums of Y'
are similarly pooled, etc.

TABLE 2

Values Used in the Covariance Tests

TEST

Si

S2

S3

S4

N*

Pf

1

53941

1046

646

125

1199

5

2

6631

26

882

1162

237

4

3

21828

292

1522

18

483

4

4

9154

392

421

2351

209

4

5

2899

175

30

405

121

3

6

5542

401

279

322

149

4

* N = total number of observations,
t p = number of groups.

Pooled data obtained in this manner were
used to get the Si, So, S3, and S4 values (see
Table 2) as defined in Kendall (1946: 238 ff.)
to make the tests indicated in Table 3. Tests
were made after Kendall {op. cit.), at the .05
level of significance and each consisted of
testing the hypotheses (1) that the regression
coefficients of the subclasses considered could
have been drawn from the same populations
and (2) that the regression of body length on
scale length was a straight line (see footnote
to Table 3). They were performed with coded
data; later the data were uncoded to calculate
the means, deviations, regression coefficients
of body length on scale length, and body-
length intercepts for the groups that were
shown, by the tests, to be significantly dif-
ferent. Test 1 indicated that the hypothesis
(1) above should be rejected, and therefore
it was necessary to consider each year-class
separately.

RESULTS

From the results of Tests 1 to 6 (Table 3)
and from the values found (Table 4), the
questions posed can be answered, as far as the
material treated is concerned, as follows:

172

PACIFIC SCIENCE, Vol. VII, April, 1953

TABLE 3

Covariance Tests for the Regression of Body Length on Scale Length

F. RATIO*

TEST

DATA COMPARED

HYPOTHESES

STATISTIC*

(.05 1. of S.)

1

1938, 1939, 1940, 1941,

For all the data, the relationship can be expressed by

and 1942 year-classes,
ages 1-5, data pooled

a straight line.

.19

2.93

from all ports.

No difference between the regression coefficients of
each year-class.

5.76

2.37

2

1938 class in:

For all the data, the relationship can be expressed by

Pacific Northwest

San Francisco

a straight line.

.84

3.67

Monterey

No difference between the regression coefficients of

San Pedro

the 1938 class in each port.

.30

2.60

3

1939 class in:

For all the data, the relationship can be expressed by

Pacific Northwest

San Francisco

a straight line.

.15

3.67

Monterey

No difference between the regression coefficients of

San Pedro

the 1939 class in each port.

2.12

2.60

4

1940 class in:

For all the data, the relationship can be expressed by

Pacific Northwest

San Francisco

a straight line.

.09

3.67

Monterey

No difference between the regression coefficients of

San Pedro

the 1940 class in each port.

2.87

2.60

5

1941 class in:

For all the data, the relationship can be expressed by

San Francisco

a straight line.

.16

5.66

Monterey

No difference between the regression coefficients of

San Pedro

the 1941 class in each port.

3.47

3.07

6

1942 class in:

For all the data, the relationship can be expressed by

Pacific Northwest

San Francisco

a straight line.

.23

3.67

Monterey

No difference between the regression coefficients of

San Pedro

the 1942 class in each port.

3.40

2.60

* Statistic to test if the relationship can be expressed by a straight line:

s, 2p-2 ^ J . N-2p

. to be compared with F. ratio:

N — 2p S2H“S3+S4 2p — 2

Statistic to test if there is difference between the regression coefficients of the subclasses involved:

S2 N-2p . p-1

♦ to be compared with F. ratio:

p-1 Si N-2p

(1) The coefficient of regression of body
length on scale length and the body
intercept are significantly different for
the 1938, 1939, 1940, 1941, and 1942
classes (Test 1).

(2) There is no significant difference among
the regression constants for the whole
1938 class regardless of area of capture
(Test 2). The same is true for the 1939

class (Test 3). The regression constants
for the 1940 class are significantly dif-
ferent for fish sampled in the Pacific
Northwest, San Francisco, Monterey,
and San Pedro (Test 4). This also is
true of the 1941 class (Test 5) and the
1942 class (Test 6).

(3) The relationship between body length
and scale length for 1- to 5-ring fish
of the year-classes involved does not

Body Length and Scale Length in Pacific Pilchard — Landa

173

TABLE 4

Regression Values Found for Homogeneous Groups

GROUPS

N*

by.xt

Sxt

Sy§

r||

Sy.xl

Sb **
y.x

Y.tt

x.«

a§§

(Y)llll

(X)1I1[

1938 year-class

237

15.90

.58

14.42

.64

11.1

1.24

210

5.20

127

202

124

1939 year-class

483

21.90

.70

19.89

.77

12.7

.85

211

5.17

98

204

154

1940 year-class in;

Pacific Northwest.

14

57.00

.17

22.43

.43

20.3

32.54

226

5.76

-102

222

164

San Francisco

50

11.66

.54

13.14

.479

11.5

3.01

227

5.65

161

228

170

Monterey

75

20.45

.73

22.23

.67

16.4

2.60

213

5.30

105

200

152

San Pedro

70

21.38

.55

14.56

.81

8.5

1.86

201

5.00

94

204

152

1941 year-class in:

San Francisco

5

5.00

.55

10.00

.28

9.6

8.10

220

5.80

191

222

170

Monterey

26

25.36

.57

20.28

.71

14.3

4.99

210

5.40

73

200

152

San Pedro

90

16.64

.42

10.94

.64

8.4

2.10

199

5.06

115

196

146

1942 year-class in:

Pacific Northwest.

7

28.00

.72

24.20

.83

13.5

7.63

233

5.63

75

216

164

San Francisco

23

10.50

.58

16.62

.37

15.4

5.65

217

5.53

159

216

164

Monterey

68

25.40

.73

22.89

.81

13.3

2.20

200

5.13

70

186

140

San Pedro

51

26.20

.60

18.94

.83

10.5

2.35

202

5.10

68

202

152

* N = number of items in the group.

NSX'Y'-(2X0(2Y0

t hy.x = regression coefficients of y on x = NSX'^— (SX')^

[For actual calculation the uncoded values were used: X' = (X')

30X; Y'

{ Sx = Standard deviation of mean scale length =

V

'2X'2(2X')2

N

N-1

§ Sy = Standard deviation of mean body length

^x

II r = correlation coefficient =-^ — by.x

y

1 Sy.x = Standard error of estimate = Sy(l— r)

1

2

/2Y'2(SY')2
/ N

^ N-1

(Y)-Y]

** Sb = variance of regression coefficient = -

SxVn-1

2Y'

ft Y. = mean body length = + assumed mean of Y

N

SX^

X. = mean scale length = h assumed mean of X

N

§§ a = body intercept = Y. — bX.
III! (Y) = assumed mean of Y
Ifl (X) = assumed mean of X

30

deviate significantly from a straight line
(Tests 1 to 6 and Table 4).

(4) The values found for the y-intercepts
are so great that it seems likely that the
regressions do not pass through the
origin (0.0) although this was not
tested.

These answers imply that, in order to back-
calculate lengths, the formula ln = a+bsn (in

which In = body length at age n and sn = scale
length at age n) should be used in which, for
each homogeneous group, the values of ”a”
and "b” are different (see Table 4). These
values are used in a separate work for a study
of the rate of deceleration of growth in the
Pacific sardine. It should be noted that the
formula given above, which applies to aver-
ages, suits our purpose quite well as long as

174

PACIFIC SCIENCE, VoL VII, April, 1953

our sample is reasonably representative of the
commercial catch; also, that its use avoids an
error likely to be introduced when only the
individual relationship, body length-scale
length, is used, i.e., the error caused by the
use of scales of different parts of the body.

SUMMARY AND CONCLUSIONS

To determine what method will most ac-
curately serve to back-calculate lengths of
pilchard at earlier ages, covariance analysis of
observed scale and body-length regressions
is useful. Emphasis is given to the most ap-
propriate sampling criteria to be used in order
to get the regression values, as well as to the
actual method of sampling.

REFERENCES

Blackburn, M. 1949. Age, rate of growth,
and general life-history of the Australian
pilchard {Sardinops neopilchardus) in New
South Wales waters. Commonwealth Scienti-
fic and Industrial Research Organization Bui.
242, Div. Fisheries Rpt. 12; 1-86, figs. 1-8,
pis. 1-8.

Buchanan-Wollaston, H. J. 1934. The
theory of variation, correlation and regres-
sion. Its relevance in researches on propor-
tional growth. Cons. Berm. Internat. L'Ex-
plor. Mer, Rapp. Proc.-Verh. 89 (4): 33-44,
figs. 1-3.

Dixon, W. J., and F. G. Massey. 1949. In-
troduction to statistical analysis. Statistical Lab-
oratory, Department of Mathematics, Uni-
versity of Oregon, Eugene, viii+217 pp.,
14 tables.

Felin, F. E., and J. B. Phillips. 1948. Age
and length composition of the sardine
catch off the Pacific Coast of the United

States and Canada. Califi. Div. Pish and
Game, Fish Bui. 69: 1-122, 1 fig.

Kendall, M. G. 1946. The advanced theory oj
statistics. Vol. IF Chas. Griffin & Co., Ltd.
vii+521 pp., figs. 19.1-30.10.

Lea, Einar. 1910. On the methods used in
the herring-investigations. Cons. Perm. In-
ternat. L’Explor. Mer, Puhl. Cir Constance 53:
7-174, 10 figs., 1 pi.

1913. Further studies concerning the

methods of calculating the growth of her-
rings. Cons. Perm. Internat. UExplor. Mer,
Puhl. Cir Constance 66: 3-36, 6 figs.

Lee, R. M. I912. An investigation into the
methods of growth determination in fishes.
Cons. Perm. Internat. h'Explor. Mer, Puhl.
Circonstance 63: 1-34, 3 figs.

Ottestad, P. 1938. On the relation between
the growth of the fish and the growth of
the scales. Cons. Perm. Internat. h'Explor.
Mer, Rapp. Proc.-Verh. 108 (4): 24-31, 3
figs.

SCHUCK, H. A. 1949. Problems in calculating
size of fish at various ages from propor-
tional measurements of fish and scale sizes.
Jour. Wildlife Management 13 (3): 298-303,
1 fig., 11 pis.

Walford, L., and K. Mosher. 1943^. Stud-
ies on the Pacific pilchard or sardine {Sar-
dinops caeruled) 2. Determination of the age
of juveniles by scales and otoliths. U. S.
Fish and Wildlife Serv., Spec. Sci. Rpt. 20:

1-19, figs. 1-32.

and 1943^. Studies on the Paci-
fic pilchard or sardine (Sardinops caerulea)
3. Determination of age of adults by scales,
and effect of environment on first year’s
growth as it bears on age determination.
FJ. S. Fish and Wildlife Serv., Spec. Sci. Rpt.
21: 1-22, figs. 1-6. ^

A New Terrestrial Isopod from Oregon, Caucasonethes rothi n. sp.

A. Vandel^

Mr. Vincent D. Roth, Technician and Cu-
rator, Entomology Department, Oregon State
College, captured a specimen of a remarkable
terrestrial isopod 8 miles east of Gold Beach,
Oregon. Upon the suggestion of Dr. Robert
J. Menzies, San Diego, California, Roth sent
me this specimen for determination.

This specimen belongs to the genus Cau-
casonethes Verhoeff, and to a new species,
named C. rothi n. sp. in honor of the dis-
coverer.

The genus Caucasonethes belongs to the fam-
ily Trichoniscidae and to a group of genera
which includes Protrichoniscus Arc., Caucaso-
nethes Verb., Escualdoniscus Vand., and Fina-
loniscus Brian (and, perhaps, Brackenridgia Ul-
rich, still imperfectly known).

GENERIC CHARACTERS OF CAUCASONETHES
VERHOEFF 1932

I consider today that Amerigoniscus Vandel
1950 is a synonym of Caucasonethes Verhoeff
1932. The generic characters of Caucasonethes
are the following:

(1) Pleonites 4 and 5 destitute of glandular
pores.

(2) Peraeopod VII without secondary sex-
ual characters.

(3) Endopodite of pleopod 1 male uniseg-
mented, likened to a lengthened tri-
angular blade, destitute of terminal
bristles.

(4) Exopodite of pleopod 1 male with pos-
terior end split into two lobes.

(5) Endopodite of pleopod 2 male biseg-

1 Professor of Zoology, Faculte des Sciences, Uni-
versity of Toulouse, Toulouse, France. Manuscript
received March 20, 1952.

mented; distal article terminating in an
acute and short point.

SPECIES OF THE GENUS CAUCASONETHES

The genus Caucasonethes includes three spe-
cies: horutzkii N QxhoQFi 1932 (caves of Kutais
and Zchal-Ziteli, in Transcaucasus; Birstein,
1950: 356); henroti (Vandel, 1950) {Amerigo-
niscus) (Gilly’s Cave, Pennington Gap, Vir-
ginia); and rothi n. sp.

The three species are very near one another
but are easily distinguishable thus:

1. Endopodite of pleopod 1 male 8 times

longer than broad, rounded at tip

henroti (Vandel)

Endopodite of pleopod 1 male 10 times
longer than broad, acute at tip 2

2. Exopodite of pleopod 1 male with 2
acute terminal lobes, widely separated

from each other horutzkii Verhoeff

Exopodite of pleopod 1 male with 2
terminal approximated lobes, one quad-
rangular, the other triangular, .rothi n. sp.

Caucasonethes rothi n. sp.

TYPE LOCALITY: Eight miles east of Gold
Beach, Oregon, on the north bank of the
Rogue River, under brocken rock and moss
in a dense forest, April 29, 1951, V. Roth
collector. 1 cf . The type is deposited in the
collection of the author.
size: (T, 5.5 X 2 mm.

COLORATION: White; without pigment.
EYES: absent.

TEGUMENTAL CHARACTERS: Surface of head
with 4 rows of granulations, the last one con-
sisting of strong, projecting tubercles. Tergite
I with 3, tergites II-VII with 2 rows of granu-

175

176

PACIFIC SCIENCE, VoL VII, April, 1953

lations, last one strongest. Granulations of
last segments much weaker than those of
anterior segments. Some weak granulations
upon pleonites 1-3. Every granulation covered
with a bristle-scale {Schuppenborste of German
authors) surrounded with numerous accessory
scales (Fig. \d).

Carapace covered with laciniate scales.

Pleonites 4-5 without glandular pores.

SOMATIC characters: Antennary tubercles
of cephalon, large triangular, projecting for-
ward.

Tip of telson rounded, with 4 bristles (Fig.

!.?)•

APPENDAGES: Afitemula (Fig. Ic)— 3-seg-
mented, last article with 10 sensory hairs
{aesthetascs) .

Antenna (Fig. 1^/)— long; articles 2, 3, and
4 grooved; articles 4 and 5 with 3 rows of
scaly tubercles. Flagellum 6-segmented; arti-
cles 2, 3, and 4 with sensory hairs (aesthetascs^.

Right mandible (Fig. l/) — 2 median peni-
cills and 1 molar penicill.

Left mandible — (.^), crushed.

Maxillula—outQt endite (Fig. 2b) termi-
nated with 11 teeth and 2 stalks, one long and
cylindrical, the other short and feathered.

Maxilliped (Fig. 2^)— palp 4-segmented;
only first article individualized. Tip of endite
terminated with 3 strong bristles and conical,
ciliated spindle.

Peraeopods—nosmA, long and slender.

Pleopod 3 — exopodite quadrangular.

Pleopods 4 and 5— exopodite triangular.

Uropod (Fig. 1^)— endopodite a little short-
er and very much narrower than exopodite,
terminated with a very long stalk; exopodite
with cluster of short stalks.

SECONDARY SEXUAL MALE CHARACTERS:

Peraeopod 7— meros and carpos fringed with
hyaline scales.

Peraeopod VII — destitute of sexual differ-
ences.

Genital duct (Genitalkegel of German au-
thors) (Fig. 2^)— very long, narrow, conical
at tip.

Fig. 1. Caucasonethes rothi n. sp.; a. Bristle-scale of
tergite I; b, laciniate scale of carapace; c, antennula;
d, antenna; e, uropod; /, right mandible; g, telson.

New Isopod — Vandel

177

Fig. 2. Caucasonethes rothi n. sp.: Maxilliped; h, maxillula, end of the outer endite; c, first pleopod of male,
en — endopodite,^Ar — exopodite; d, posterior end of exopodite of first pleopod of male; e, genital duct; /, second
pleopod of male. '

Pleopod 1 (Figs. 2c, <7)— endopodite blade-
like, triangular, very narrow and lengthened,
destitute of terminal stalk. Exopodite large,
quadrangular, terminated with 2 lobes, one
quadrangular and ciliated, the other triangular
and pointed.

Pleopod 2 (Fig. 2/)— endopodite bisegment-
ed, sharpened at end; exopodite rectangular.

RELATIONSHIP OF CAUCASONETHES ROTHI
N. SP. AND OREGONISCUS NEARCTICUS

(arcangeli)

Arcangeli (1932: 137) gave the name of
Trlchonlscus nearctkus to a single female from
MacLeay Park, Portland, Oregon. Hatch
(1947: 190) proposed the generic name Ore-
goniscus for this species.

178

PACIFIC SCIENCE, Vol. VII, April, 1953

It is difficult to recognize the species of
terrestrial isopods based only upon females,
but, I think that Oregoniscus nearcticus (Arc-
angeli) and Caucasonethes rothi n. sp. are dis-
tinct. They differ in many points:

(1) The size is very different: 0. nearcticus,
2.73 mm. long; C. rothi, 5.5 mm. long.

(2) The number and disposition of sensory
hairs {aesthetascs) of antennula is very
different in the two species.

(3) Antennae very long in C. rothi, short in
0. nearcticus "caratteristica e la incava-
tura che dorsalmente presenta nelle
meta distale esterna il 5° articolo dello
scapo” (Arcangeli, 1932). This charac-
ter is missing in C. rothi. The flagellum
of the antenna is six-segmented in C.
rothi, four-segmented in 0. nearcticus.

(4) The legs of C. rothi are long and slend-
er, whereas in 0. nearcticus "e da notarsi
la larghezza e relativamente piccola
lunghezza specialmente del meropodi-
te, carpopedite, e propodite, che con-
feriscono al membro un aspetto mas-
siccio” (loc. cit.).

(5) In C. rothi the endopodite of the uro-
pod is only a little shorter (about one
tenth) than the exopodite. In 0. nearc-
ticus, the endopodite is about one half
as long as the exopodite.

REFERENCES

Arcangeli, A. 1932. Isopodi terrestri raccolti
dal Prof. Silvestri nel Nord- Am erica. Por-
tici R. Scuola Super, di Agr., Lab. Zool. Gen.
e Agr., Bo/. 26: 121-141, 7 figs.

Birstein, J. a. 1950. The Fauna of the west-
ern Transcaucasus caves. Zoologitchesky Jour.
29: 354-366.

Hatch, M. H. 1947. The Chelifera and Iso-
poda of Washington and adjacent regions.
Wash, [state] Univ., Pubs. Biol. 10: 155-274,
18 pis.

Vandel, a. 1950. Isopodes terrestres recueil-
lis par C. Bolivar et R. Jeannel (1928) et le
Dr. Henrot (1946). Arch, de Zool. Expt. et
Gen. 87: 183-210, 20 figs.

Verhoeff, K. W. 1932. Cavernicole Onis-
coideen. 44. Isopoden-Aufsatz. Hohlen-u,
Karstf. Berlin, Mitt. 1932: 3-15, 21 figs.

I

Floristic Interchanges Between Formosa and the Philippines

Hui-Lin Lii

INTRODUCTION

The floras of Formosa and the Philippines
are, in spite of their geographic proximity,
distinctive in their general nature. The two
regions were separated by the Formosan rift
(the Bashi Channel), probably in the early
Tertiary, long before their connections with
other respective neighboring areas were sev-
ered. Thus the bulk of the Formosan flora was
derived from China or from other floras that
spread to Formosa through the Chinese main-
land, while the bulk of the Philippine flora
was derived from Malaysia and other lands
south of the Asiatic continent.

The distinctiveness of the two floras has
been amply demonstrated by Merrill (1923).
It will suffice to mention here the many tropi-
cal and southern families that are present in
the Philippines but absent from Formosa,
such as the Dipterocarpaceae, Centrolepida-
ceae, Monimiaceae, Nepenthaceae, Cunonia-
ceae, Erythroxylaceae, Dichapetalaceae, Da-
tiscaceae, Clethraceae, Epacridaceae, and Sal-
vadoraceae. On the other hand, many other
families, clearly of northern origin, occur in
Formosa but are entirely absent from the
Philippines, such as Cephalotaxaceae, Taxo-
diaceae, Cupressaceae, Betulaceae, Lardiza-
balaceae, Trochodendraceae, Papaveraceae,
Geraniaceae, Callitrichaceae, Pyrolaceae, Dia-
pensiaceae, Valerianaceae, and Dipsaceae.

^ Research Associate, Morris Arboretum, University
of Pennsylvania, Philadelphia. Manuscript received July
1, 1952. The author is indebted to Dr. A. C. Smith of
the Smithsonian Institution for his kindness in reading
the manuscript and offering suggestions for its im-
provement.

However, though the basic constituents of
the two floras are quite different, indicating
their separate origins, their close geographic
proximity allows certain floristic interchanges,
apparently of relatively recent times. The dis-
tance between the main islands, Formosa and
Luzon, is about 350 kilometers. Stretching in
the sea between the two is a chain of small
islands, parts of the same volcanic line. The
first group consists of three islands off the
southeastern coast of Eormosa, Hoshaotao
(Kotosho in Japanese), now called Lutao,
Hungtauyu (Katosho in Japanese), now called
Lanyu and commonly known as Botel Toba-
go, and the tiny island. Little Botel Tobago.
These islands are separated from the Batan
(or Batanes or Bashi) Islands by the Bashi
Channel at a distance of about 100 kilo-
meters. The Batan Islands are separated from
the Babuyan Islands, which are close to the
northern coast of Luzon, by the Balintang
Channel, about 70 kilometers wide. The dis-
tance is nowhere great enough to make plant
migration difficult. At South Cape, the south-
ernmost tip of Eormosa, the small island
Y’Ami of the Batan group, the northernmost
of the Philippine Islands, is actually within
sight.

These small islands show remarkable affini-
ties with the Philippine flora, even the north-
ernmost ones, Lutao and Lanyu, which are
closest to Eormosa. The conspicuous rela-
tionship of the floras and faunas of the two
latter islands to the Philippines has led some
to suggest extending the Neo-Wallace Line
northward from the Philippines, passing it
between the main island of Formosa and these

179

180

PACIFIC SCIENCE, Vol. VII, April, 1953

two small islands (Kano, 1931). However, it
has been shown that the southern tip of For-
mosa, commonly known as the Hunchuen
(Koshun) Peninsula, has a flora quite dis-
tinct from that of the Formosan mainland but
very close to that of these two small islands,
showing similarly distinctive relationships
with the Philippine flora. There is a marked
infiltration of austral elements from the Phi-
lippines to Formosa, particularly in the ex-
treme southern part of the island and in these
two small islands (Li and Keng, 1950).

SOUTHWARD MOVEMENT OF
TEMPERATE ELEMENTS

Before the inception of this northward mi-
gration of relatively recent occurrence, as will
be discussed later, there occurred another ex-
pansion of floristic elements in the opposite
direction, that is, from Formosa to the Phi-
lippines. These elements, in contrast to the
southern tropical elements, mostly of the low-
lands, are cold-temperate in nature and lim-
ited to montane or alpine regions. In Formosa,
because of the massive and lofty mountain
ranges which occupy most of the central por-
tion of the island, there developed an exten-
sive alpine-montane flora rich in number of
species. These plants show distinctively close
relationships with those of western China and
the eastern Himalayas and also to some ex-
tent with the montane flora of Japan in the
north.

There are over 40 peaks in the Formosan
mountain systems that exceed 1,000 meters
in height. The highest peak is Yu Shan, or
Mt. Morrison, towering to a height of 1,950
meters, the highest of all eastern Asia. Such
an extensive, high mountain chain permits the
existence of an alpine flora that otherwise
exists only in the Himalayas, western China,
and other mountain regions far north of For-
mosa.

From Formosa, a large number of tem-
perate elements, mostly of mountain regions,
extend southward to the Philippines. In the
Philippine Islands, these Asiatic elements are

largely confined to medium and high altitudes
in northern Luzon. Such characteristically
northern or Asiatic elements, mostly common
genera in the mountains of Formosa, such as
Ltlium^ Liriope, Saururus, Tbesmm, Arenaria,
Sedum, Duetzia, Rosa, Skimmia, Buxus, Pistacia,
Androsace, Hoppea, Salvia, Ellisiophyllum^ Hemi-
phragma, Peracarpa, Aster, Solidago, Anisopap-
pus, and Artemisia, occur nowhere in the Ma-
layan region outside of northern Luzon (Mer-
rill, 1923-26). It is apparent that Formosa is
an important route of migration of these
Asiatic types to the regions in the south (Van
Steenis, 1946).

We may picture that during the Pleistocene
or earlier, when the general temperature was
much lower than now, these montane species
would have inhabited lower elevations and
thus had greater ranges. Their ranges might
have been more or less continuous, extending
from the Himalayas and western China through
the mountains of southern China to Formosa
and Luzon. Subsequent rising of tempera-
tures forced these plants to higher elevations
until finally they occupied the present iso-
lated mountain regions suitable for their exist-
ence. Thus, species like Hemiphragma hetero-
phyllum Wall, and Ellisiophyllum pinnatum
(Wall.) Makino now have widely disjunct
ranges on the high mountains of Luzon, For-
mosa, western China, and the eastern Hima-
layas, etc.

These Philippine plants of northern origins
are sometimes specifically or subspecifically
distinct from their congeners in Formosa,
while others may be specifically identical.
That some of these show morphological dif-
ferentiation indicates that their separation
must have been for a considerable length of
time. As such plants are now confined to
isolated and distant mountain regions and as
there were drastic changes in climate during
the geologically recent past, such a south-
ward migration of the montane flora may be
considered largely a matter of the past, there
being little possibility that the same process
is going on extensively at the present.

Floristic Interchanges — Li

181

NORTHWARD MOVEMENT OF
TROPICAL ELEMENTS

In contrast to the southward migration of
these temperate elements, the northward move-
ment of tropical elements from the Philippines
to Formosa is limited practically to the low-
lands and particularly to the southern ex-
treme of the island of Formosa. Favored by
the gradual rising of temperature since the
Pleistocene, these tropical, or southern, plants
that find their way to Formosa are able to
establish themselves there.

Apparently many strand plants migrate by
the help of ocean currents. In the case of
Formosa, the main current is supplied by the
Japan Stream from the south which passes
along the island chain connecting the Philip-
pines and Formosa. This chain of small islands
also acts as a series of stepping stones for the
migration of plants, especially from south to
north. This northward migration of plants is
also aided by wind. The prevalent monsoon
during the summer and fall and the general
track of typhoons rampant in the eastern Paci-
fic during this same period of the year are also
in a northeastern direction from the tropics,
often leading from the Philippines to For-
mosa.

That these islands serve as an efficient mi-
gration route of the northward movement of
southern plants can be demonstrated by the
presence of the following floral elements in
Formosa: (1) species of wide tropical distri-
bution, (2) species common to the Philip-
pines and Formosa, (3) species common to
the Philippines and the southernmost part of
Formosa, and (4) species common to the
Philippines and the islands of Lutao and
Lanyu.

In the first group are many paleotropic
species that extend to the whole island of
Formosa but more commonly only to the
southern part of the island. Such common
species are Dodanea viscosa (L.) Jacq., Croton
cumingia Muell.-Arg., Litsea cuheha Pers., and
Murrya paniculata (L.) Jack. Those that also
occur in the northern part are mostly along

the coastal lowlands. Their wide ranges are
due apparently to their more aggressive na-
ture and especially to their wider tolerance of
temperature extremes.

In the second group are species like Acacia
confusa Merr., Ipomoea polymorpha R. & S., and
Oreocnide trinervis Wedd., which occur widely
in the Philippines and extend also to Formosa,
especially along the coasts and lowlands, from
south to north. The number of such species
with wide ranges in Formosa is few, indicating
the insignificance of such elements and pos-
sibly also their recent and scattered arrival.

There are many more species common to
the Philippines and Formosa but confined in
the latter island to the southern part. These
species are sometimes limited only to the
southernmost tip of the island — the Hun-
chuen Peninsula and the small islands of Lu-
tao and Lanyu. Such species as llligera luzon-
ensis Merr., Schizostachyum diffusum Merr.,
Goniothalamus amuyon Merr., Aglaia elliptici-
folia Merr., Maha huxi folia Pers., Guettarda
speciosa L., Morinda citri folia L., Pemphis acidula
Forst., etc., are often important elements in
the local vegetation. That their occurrence is
often limited to the seashore and coastal areas
in Formosa distinctly indicates that they are
relatively recent arrivals.

The flora of the two small islands, Lutao
and Lanyu, off the southeast coast of Formosa
is much more intimately related to the south-
ernmost part of Formosa than is generally
understood (Li and Keng, 1950). The south-
ernmost tip of Formosa manifests close floris-
tic relationships with the small islands as well
as with the Philippines in general. Its flora is
quite distinct from that of northern Formosa.
The fact that some species are present on these
two small islands but absent in the southern-
most part of Formosa is due apparently to
human factors, as the mainland of Formosa
has been more severely exploited. As a result,
there are a number of Philippine species that
are present on these islands but do not occur
on the mainland of Formosa, such as Myris-
tica cagayanensis Merr., Boerlagiodendron pecti-

182

PACIFIC SCIENCE, Vol. VII, April, 1953

natiim Merr., Dysoxylum ctmingiana DC.,
Styrax kotoensis Hay., Turpinia lucida Nakai,
Macaranga dipterocarpifolia Merr., etc.

The importance in floristic migration played
by the chain of small islands between the
Philippines and Formosa can be illustrated
more precisely by a few cases. Among the
wide paleotropic species, chiefly along or near
the coast and common throughout the Philip-
pine Islands, are Morinda citrifolia L. and
Maha huxifoUa (Rott.) Pers. ( == Diospyros ferrea
Bakh.). These species extend through the
Batan and Babuyan Islands to Formosa, lim-
ited in the latter case only to Lutao and Lanyu
and the southernmost Hunchuen Peninsula.
Among the less wide species are Aglaia for-
mosana Hay. and A. elliptici folia Merr., both
of the northern part of Luzon. These species
also extend northward, following the same
route, through the Batan and Babuyan Is-
lands to Hunchuen Peninsula in Formosa and
to Lutao and Lanyu.

That these southern tropical elements are
relatively recent arrivals in Formosa can be
attested by their occurrence in coastal regions
and in secondary forests as well as by their
specific identity with Philippine plants. In
most cases, the species in Formosa are exactly
the same as in the Philippines. These species
have not been isolated long enough to have
undergone any morphological differentiation.
It may be presumed that this northward mi-
gration is not only very recent but is still
continuing at the present. Success of estab-
lishment of these southern species in Formosa
is controlled mainly by climate factors. With
probable continued rise of temperature in the
future, there may be more southern species
entering into the flora of Formosa. At the
same time, many of the species will be able
to extend their range to the northern part of
the island.

SOME FLORISTIC NOTES

In the past, this southern affinity of the
Formosan flora has not been properly empha-
sized because of our inadequate information.

The floras of Formosa and the Philippines
have been studied, in most cases, independ-
ently of one another. Many of the species
proposed from Formosa are later found to be
conspecific with earlier-named Philippine spe-
cies. Such scattered findings are increasing in
number. More recently, with ample reference
collections from the Philippine Islands at
hand, the present writer restudied many For-
mosan plants in connection with related spe-
cies from neighboring regions. It was dis-
covered that many species, especially those
considered endemic to southern Formosa or
Lanyu (Botel Tobago), are synonymous with
certain generally widespread species of the
Philippine Islands, especially of Luzon and
the northern small islands. Some of these
notes have been published in a few scattered j
papers (Li, 1950, 1952). A number of recent |
findings are enumerated below. Cited speci- j
mens are selected from the herbarium of the i
National Taiwan University, Formosa (NTU),
and the U.S. National Herbarium, Smith-
sonian Institution (US).

Urticaceae

1. Laportea batanensis C. B. Robinson in

Philip. Jour. Sci. 5. Bot.: 481, 1910.

Laportea kotoensis Hay., Gen. Ind. FI. For-
mos. 70, 1916, nomen; Kanehira, For-
mos. Trees 529, 1917; ibid., rev. ed. 170,
1936. Syn. nov.

Batan Islands, Botel Tobago.

Neither Hayata nor Kanehira cited speci-
mens, but Kanehira’s very brief description
shows that the plant is identical to the Philip-
pine species, L. batanensis, known from the
Batan Islands only.

Myristicaceae

1. Myristica cagayanensis Merr. in Philip.

Jour. Sci. 17: 255, 1920; Kanehira, Formos.

Trees rev. ed. 193, f. 141, 19^6.

Myristica glomerata Kudo & Masamune in
Ann. Taihoku Bot. Card. 2: 89, 1932.
Syn. nov.

Floristic Interchanges — Ll

183

Luzon; Formosa, in southern part, and in
Botel Tobago.

Formosa: Cuntin, Hunchuen Peninsula, H.
Keng, Oct. 31, 1950 (US).

Myristica glomerata Kudo & Masamune is a
name based on Sterculia glomerata Blanco, FI.
Filip, ed. 1. 764. 1837, which had earlier been
made into the combination Knema glomerata
(Blanco) Merr. in Jour. Str. Branch. Roy. As.
Soc. 76: 81. 1917, Sp. Blanco. 151. 1918, a
species of Borneo and the Philippines but not
of Formosa. Kudo and Masamune were mis-
led by the previous misidentification of the
Formosan plant in question as Myristica heter-
ophylla sensu Hay., Gen. Ind. PL Formos. 61.
1917, etc., non F.-Vill. Myristica heterophylla
F.-Vill. is a synonym of Knema glomerata
(Blanco) Merr. The two plants in question
differ greatly and are now placed in two dif-
ferent genera.

Rutaceae

1. Evodia confusa Merr. in Philip. Jour. Sci.
20: 391, 1922.

Evodia merrillii Kanehira & Sasaki in Kane-
hira, Formos. Trees rev. ed. 313, f. 267,
1936.

Evodia roxhurghiana sensu Matsum & Hay.
in Jour. Coll. Sci. Tokyo 22: 70, 1906
(Enum. PL Formos.); Hay. Icon. PI. For-
mos. 1: 118, 1911, non Benth.

Celebes and Philippines, widely distributed
and common in forests at low and medium
altitudes; Formosa, mostly in northern part.

Formosa: Shirin, Taihoku, T. Tanaka &
Y. Shimada 11163 (US) ; Kangu, H. Keng, Oct.
26. 1950 (US); Sinten, H. Keng 1811 (US);
Tykutan, Taihoku Sou-Gen Lin, Aug. 30,
1932 (NTU).

Kanehira and Sasaki cited no specimen for
their species. The description and illustration
are distinctive. This characteristic plant with
large leaves is in all respects referable to the
common Philippine species Evodia confusa
Merr., known earlier as glabra T

2. Glycosmis cochinchinensis (Lour.)
Pierre ex Engl. & Prantl, Nat. Pflanzenfam.

III. 4: 185, f. 106, 1895; Kanehira, Formos.

Trees rev. ed. 314, f. 269, 1936.

Toluifera cochinchinensis Lour. El. Cochinch.
262, 1790.

Glycosmis citri folia Lindley in Trans. Hort.
Soc. 6: 72, 1826.

Glycosmis pentaphylla Correa in Ann. Mus.
Nat. Hist. Paris 8: 386, 1806; Matsum.
& Hay. in Jour. Coll. Sci. Tokyo 22: 73,
1906; Hay. Icon. PL Formos. 1: 121,

1911.

Citrus erythrocarpa Hay. Icon. PL Formos.
6: 13, 1916. Syn. nov.

Glycosmis erythrocarpa Hay. Icon. PL For-
mos. 8: 14, 1919. Syn. nov.

India to Malaysia, Philippine Islands, Hai-
nan; Formosa, common in forests at low
altitudes.

Formosa: Taipei, H. Keng 1044 (US), K.
Odashima 13607 (US), Kei-Dai Lin, Nov. 12,
1930 (NTU); Tamali, Taitung, H. Keng 1376
(US); Bankinsing, A. Henry 1487 (US), 1387
(US); Kuraru, Koshun, E. H. Wilson 11038
(US)._

This is a polymorphous tree, very variable
in its features. Hayata’s plant is described with
smaller leaves which are usually ternate, but
the characters are within the range of varia-
tion of the species.

Anacardiaceae

1. Semecarpus gigantifolia Vidal, Sinopsis

Atlas 22, t. 36, f. A. 1883.

Semecarpus vernicifera Hay. & Kawakami in
Hay. Icon. PL Formos. 2: 108, 1912;
Kanehira, Formos. Trees rev. ed. 367^
f. 322, 1936. Syn. nov.

Widely distributed in the Philippines, in
forests at low altitudes; Formosa, along sea-
shore of the south and east coasts, and on
Botel Tobago Island; cultivated in other
places.

Formosa: Taihoku, cult., E. H. Wilson 9910
(US), Kei-Dai Lin, Nov. 5. 1936 (NTU).

The Formosan plant proves to be identical
with this common Philippine species, many
specimens of which are available for compar-
ison.

I

1

184

PACIFIC SCIENCE, Vol. VII, April, 1953

Aquifoliaceae

1. Ilex asprella (Hook. & Arn.) Champ, in
Kew Jour. 4: 329, 1852; Kanehira, Formos.
Trees rev. ed. 370, f. 324, 1936; S. Y. Hu
in Jour. Arnold Arb. 30: 269, 1950.

Prims asprella Hook. & Arn. Bot. Beech.

Voy. 176, pi. 36, f. 1, 2, 1833.

Ilex arisanensisY2.m2SS\oio, Suppl. Icon. PI.
Formos. 1: 30. f. 10, 1952; Kanehira,
Formos. Trees rev. ed. 370, 1936. Syn.
nov.

Luzon to southeast China; Formosa, in the
thickets from low to medium altitudes
throughout the island.

Formosa: Sozan, Taihoku-shu, T. Tanaka
& Y. Shimada 11009 (US), Y. Kudo, S. Suzuki
& K. Mori, April 21, 1929 (NTU); Mt. Kwan-
non, Taihoku-shu, T. Tanaka & Y. Shimada
11094 (US); Taipei, H. Keng 1024 (US), T.
Tanaka 76 (US) ; Shirin to Sozan, E. H. Wilson
10293 (US); Musha, Nanto, E. H. Wilson
10063 (US); South Cape, A. Henry 1334 (US);
Arisan, Faurie 186 (cotype of /. arisanensis
Yamamoto, photo US).

Photograph of Yamamoto’s type together
with his original description prove that L
arisanensis Yamamoto is clearly conspecific
with /. asprella. This species has characteristi-
cally thin leaves with caudate tips and long
slender pedicels. Faurie’s plant, a mature fruit-
ing specimen, has relatively larger leaves than
others and also was collected at higher alti-
tudes than most others. Ilex arisanensis Yama-
moto was not considered in S. Y. Hu’s study
of the genus, and it was maintained as a dis-
tinct species by Kanehira.

Icacinaceae

1. Gonocaryum calleryanum (Baill.) Becc.
in Malesia 1: 123, 1877.

Phlehocalymna calleryanum Baill, in Adan-
sonia 9: 147, 1896.

Gonocaryum diospyrosifolium Hay. Icon. PI.
Formos. 2: 106, 1912; Kanehira, For-
mos. Trees rev. ed. 400 f. 359, 1936.
Syn. nov.

Luzon, Batan Islands; Formosa, in forests
along the coast, rare in Hunchuen Peninsula
only.

Formosa: Koshun, Kuraru, Y. Kudo & S.
Suzuki 13889 (NTU); Kuraru, B. Hayata &
Sasaki 13621 (isotype of G. diospyrosifolium
Hay., photo US).

Gonocaryum diospyrosifolium Hay. proves to
be the same as the Philippine G. calleryanum
(Baill.) Becc. {G. tarlacense Vidal). The plant
is a common tree in Luzon and is also found
on the Batan Islands between Luzon and For-
mosa. In Formosa, it is a rare plant grown
only in the shore forests of Kuraru, Hun-
chuen f^eninsula, sometimes on coral rocks.

Melastomataceae

1. Astronia ferruginea Elm. Leafl, Philip.

Bot. 4: 1205, 1911; Merr. in Philip. Jour.

Sci. 8: Bot. 342, 1913.

Astronia formosana Kanehira, Formos. Trees
259, 1917. Syn. nov.

Astronia pulchra sensu Hay. in Jour. Coll.
Sci. Tokyo 30(1): 114, 1911; (Mat. FI.
Formos.), Icon. PI. Formos. 2: 25, 1912;
non Vidal.

Astronia cumingiana sensu Kanehira, For-
mos. Trees rev. ed. 507. f. 469, 1939; non
Vidal.

Philippine Islands, from Mindanao to Lu-
zon; Formosa, in forests, Hunchuen Penin-
sula, Lutao and Lanyu (Botel Tobago).

Formosa: South Cape, A. Henry 638 (US);
Kuskus, Y. Kudo & S. Suzuki 16066 (NTU).

The Formosan plant is neither a distinct
species nor referable to either A. pulchra or
A. cumingiana of the Philippines. The species
is actually identical with A. ferruginea Elm.,
a more or less common species of the Philip-
pines. The ferrugineous lepidote inflorescence
and undersurface of the leaves are very dis-
tinctive and characteristic of this species.

Boraginaceae

1. Cordia cumingiana Vidal, Phan. Cum-
ing. Philip. 187, 1885, Rev. PI. Vase. Filip.

192, 1886.

Floristic Interchanges — Li

185

Cordia kanehirai Hay. Icon. PI. Formos.
6: 31, 1916; Hou in Taiwania 1; 207,
1950. Syn. nov.

Luzon; Formosa, in the thickets in southern
part only, scarce.

Formosa: Koshun, E. Matuda 1333 (NTU);
Kuraru, Koshun, R. Kanehlra 7 (isotype of
C kanehirai, photo US).

Hayata originally placed this plant as a
close ally of C. cumingiana. In comparing spe-
cimens from the two regions, it is found that
the Formosan plant cannot be differentiated
from the Luzon plant and must be considered
as conspecific.

2. Cynoglossum lanceolatum Forsk, FI.

Aegypt. Arab. 41, 1775; Hou in Taiwania

1: 217, 1950.

Cynoglossum alpestre Ohwi in Acta Phytotax.
Geobot. 2: 150, 1933; Hou in Taiwania
1: 218, 1950. Syn nov.

India, Southern China to the Philippines;
Formosa, mountain regions.

Formosa: No precise locality, A. Henry
1013 (US); Arisan, T. Tanaka 320 (US); S.
Suzuki, ]\Ay 17, 1937 (NTU).

No isotypes of Ohwi’s species are available,
but, among the cited specimens, Suzuki j. n.
most closely approaches the original descrip-
tion of C. alpestre in having longer, denser
hairs and smaller, thicker leaves than other
specimens of C. lanceolatum. These characters
are, however, evidently due to higher eleva-
tions, whereas there are no structural differ-
ences between plants of higher and lower
altitudes. Cynoglossum alpestre Ohwi can at most
be considered an alpine form of the species
C. lanceolatum, but, in view of the wide dis-
tribution and variable nature of the species,
it seems not desirable to recognize it taxo-
nomically.

3. Ehretia navesii Vidal, Rev. PL Vase. Filip.

194, 1886.

Ehretia resinosa Hance in Jour. Bot. 299,
1880; Hou in Taiwania 1: 203, 1950.
Syn. nov.

Ehretia formosana Hemsl. in Jour. Linn.
Soc. Bot. 26: 144, 1896. Syn. nov.

Philippine Islands; Formosa, in southern
part, along the seashore.

Formosa: Takao, A. Henry j. n. (US), K.
Moritani 2308 (NTU); Kuraru, E. H. Wilson
10967 (US).

The Formosan plant proves to be the same
as the Philippine E. navesii Vidal, a charac-
teristic plant readily distinguished by the
long-linear and persistent calyx-lobes which
enclose the globose fruit at maturity.

Verbenaceae

1. Premna nauseosa Blanco, FI. Filip. 489,

1837.

Premna integri folia Blanco, FI. Filip, ed. 2.
342, 1845; Matsum. & Hay. in Jour.
Coll. Sci. Tokyo 22: 299, 1906 (Enum.
PI. Formos.); non Linn.

Premna ohtusifolia sensu Sasaki, List. PL
Formos. 353, 1938; Mori in Masamune,
Short FI. Formos. 181, 1936; non R. Br.

Premna odorata sensu Yamamoto in Jour.
Soc. Trop. Agr. 6: 554, 1934, p. p.; Ka-
nehira, Formos. Trees rev. ed. 654, 1936,
p. p.; non Blanco.

Philippine Islands; Formosa, in southern
part only.

Formosa: No precise locality, A. Henry 613^
791 (US).

This is an exceedingly common species in
the Philippine Islands, generally known as
"P. integri folia." This species was confused by
many authors, including Kanehira, with P.
odorata, another Philippine species that occurs
also in southern Formosa. In P. odorata the
leaves are densely tomentose beneath. In P.
nauseosa the leaves are more or less glabrous
beneath.

2. Premna odorata Blanco, FI. Filip. 488.

1837; Yamamoto in Jour. Soc. Trop. Agr.

6: 554, 1934, p. p.; Kanehira, Formos.

Trees rev. ed. 654, f. 6IO, 1936, p. p.

Premna vestita Schauer in DC. Prodr. 11:

631, 1847; Henry in Trans. As. Sci. Jap.
24. SuppL: 70, 1896.

Widely distributed in Luzon; Formosa, in
forests near seashore in northern and southern
parts.

186

SUMMARY

The floras of Formosa and the Philippines
are distinctive in their general nature. The
former is related especially to the Chinese
mainland and the latter to Malaysia. Their
close geographic proximity, however, has al-
lowed certain floristic interchanges in relative-
ly recent times. An earlier movement was the
southward migration of temperate elements
from Formosa to the Philippines. These are
mostly montane and alpine elements and, in
the Philippines, largely confined to the moun-
tains of northern Luzon. These elements are
now of wide disjunct distribution and are
related to the alpine floras of western China
and the Himalayas. A more recent movement
is the northward expansion of tropical ele-
ments from the Philippines to Formosa. These
are usually lowland plants and are found in
Formosa limited mostly to the southern ex-
treme of the island and the adjacent small
islands Lanyu and Lutao. Floristic notes on
certain species common to the Philippine and
Formosan floras but considered by former
authors as distinct are given.

PACIFIC SCIENCE, Vol. VII, April, 1953
REFERENCES

Kano, T. 1931. Notes on Pachyrrhynchides
from Kotosyo and their zoogeographical
significance. Biogeogr. Soc. Jap., Bui. 2;
193-208. [in Japanese.]

Li, H. L. 1950. Studies in the Scrophulariaceae
of Taiwan. Quart. Jour. Taiwan Mus. 3:
55-80.

1952. Notes on some families of For-
mosan phanerogams. Wasl^. Acad. Sci., Jour.
42; 39-44.

and H. Keng. 1950. Phytogeographi-

cal affinities of southern Taiwan. Taiwania
1: 103-128.

Merrill, E. D. 1923. Die pflanzengeographi-
sche Scheidung von Formosa und den Phi-
lippinen. Bot. Jahrh. 58: 599-604.

1923-26. An enumeration of Philippine

flowering plants. 4 vols. Bureau of Printing,
Manila.

Van Steenis, C. G. G. J. 1946. Preliminary
revision of the genus Lonicera in Malaysia.
Arnold Arboretum, Jour. 27: 442-452.

Observations on the Subgenus Fhalangomyia of the Genus Culex in Ecuador with
Description of a New Species (Diptera: Culicidae)

Roberto Levi-Castillo^

The subgenus Fhalangomyia has rather gen-
erally been considered as a synonym of the
subgenus Culex although, especially in the
species found in Ecuador and described here-
in, it has characters which differentiate it very
clearly. The terminalia are very characteristic.
The strongly chitinized structure of the clasp-
er and the presence of strong hairs on its
outer surface constitute a great difference
from the condition found in members of the
subgenus Culex. Fhalangomyia seems to repre-
sent one of the oldest branches of the genus
Culex, as indicated by its adaptation to a high-
altitude environment.

All the members of this subgenus are found
only in the Andean countries of South Ameri-
ca. They are large mosquitoes with very well-
developed bodies and strong, long legs. They
are adapted for high-altitude flying and for
low oxygen consumption and, therefore, are
always found at altitudes of more than 2,000
meters above sea level, where no other mos-
' quitoes and few other insects of any sort can
live. One of the two species considered here
was found at an altitude of 2,900 meters near
the city of Quito and the other near the town
of Cuenca at an altitude of 2,500 meters. In
the Andean region of Ecuador the median
temperature is ordinarily between 14.5° and
20°C. (58°— 68°F.), and wind velocities of 5 to
25 miles per hour prevail. Thus, in order to
live, mosquitoes must be adapted to this en-
vironment.

^ Director, Ecuadorian Center for Entomological Re-
search, Guayaquil, Ecuador. Manuscript received April
14, 1952.

The most interesting characteristic of these
mosquitoes is the fact that larvae, pupae, or
adults die very shortly if they are taken to sea
level, apparently because their metabolism has
been adapted to the oxygen shortage of the
high altitudes of the Andes. Thus the author
was unable to breed larvae of Fhalangomyia at
sea level, even when water from the breeding
places, with all the algae present, was brought
to the laboratory. Apparently, then, compres-
sion is a main factor in killing the larvae,
although they sometimes survive 10 to 12
hours of being at sea level. Also temperature
changes will kill the larvae, and at sea level
both the temperature and humidity are higher
than those at over 2,000 meters above sea
level. However, even placing the larvae
brought to sea level in refrigerators held at
the same temperatures as the highlands does
not prevent their dying in a few hours. This
proves that the subgenus Fhalangomyia is only
adapted to live at high altitudes, in the An-
dean region of Ecuador. The problem of the
effects of oxygen, basal metabolism, and en-
vironment on highland mosquitoes has not
been studied sufficiently, and it is one of the
research projects of the Ecuadorian Center for
Entomological Research.

Two species of Fhalangomyia have been
taken in the highlands of Ecuador, one pre-
viously known and one new to science. They
are both described here.

Culex (Fhalangomyia) archegus

Dyar, 1929

LARVA (Fig. \a, h)\ Head rounded, wider
than long. Antennae long, tapering, with a sub-

187

188

Fig. 1. Larval structures of C. {Ph.) archegus. a.
Head; h, cauda.

apical hair tuft, spiculate body, and 2 pairs
of long apical hairs. Freantennal hairs 6 to 8,
finely spinulate, longer than the antennae. In-
terior dorsal hairs 4 to 6, finely spinulate. Ex-
terior dorsal hairs 3 to 4, spinulate.

Comb of the eighth abdominal segment (Fig.
lb) formed by a patch of elongate teeth with
apex hairy. Air tube long, 4.5 times as long
as wide. Pecten of 13 to 15 triangular scales,
sharp and with 3 to 5 teeth each. Tuft of the
pecten double and long, other tufts double
and long, located at different places along the
air tube. Anal segment longer than wide, finely
spinulate on most of its surface. Lateral hair
tuft double and large. Dorsal brush formed by
3 long, strong elements. Ventral brush formed

PACIFIC SCIENCE, Vol. VII, April, 1953

by 8 hairs, 4 elements each, very well devel-
oped and long.

PUPA (Fig. 2a, b) (Chaetotaxy according to
Knight and Chamberlain, 1948) : Segment I,
hair 10 well developed and long, hair 2 with
strong base, very ramified; Segment II, hair 8
developed, long, hair 2 long and thin; Seg-
ment III, hairs 2 and 5 long and ramified, hair
7 long; Segment IV, hairs 2 and 5 ramified and
long, hairs 4, 6, 7 long; Segment V, hair 2
ramified and long, hair 5 very long and di-
ramified; Segment VI, hair 2 diramified and
very long, hair 5 ramified and long, hair 7
long and simple; Segment VII, hair 2 medium

Fig. 2. Pupal structures of C. {Ph.) archegus Dyar
and C {Pht) quitensis n.sp. a. Trumpet of C. archegus;
h, dorsal aspect of abdomen of C. archegus; c, trumpet
of C. quitensis; d, dorsal aspect of abdomen of C.
quitensis.

Subgenus Phalangomyia — Levi-Castillo

and diramified, hair 6 long and simple, hair 5
medium and diramified, hair 8 ramified and
very well developed; Segment VIII, hair 5 me-
dium and simple, hair 8 multiramified, very
well developed and spiculated. Pupal trum-
pets longer than wide, with a triangular open-
ing, rounded at the angular borders, tapering
and very sinuous. Pupal paddles rounded with
strong medium rib, apical hairs small.

ADULT (both male and female): Proboscis
medium, slightly enlarged near the tip, olive
green. Palpi thin and long in the males, short
and lobulous in the females, with many setae,
olive green and black. Vertex with dark in-
tegument revetted with plumous white scales,
which form an overlapping tuft toward the
clypeus, and bordering the eyes. Occiput dark
with plumous scales as on the vertex. Pro-
thoracic lobes revetted with golden scales and
dark hairs. Mesonotum with 3 strips of small,
compact brown scales forming circular spots
at the wing base, the disc revetted with small
golden hairs, with clear spaces among the
compact scales. Scutellum trilobate with the
median lobe larger than the others, with gold-
en scales as on the mesonotum, with long
dark hairs on all the lobes. Postnotum bare
of scales, light brown. Abdomen olive brown,
segments with basal white bands that become
larger, forming wide white bands that cover
lateral portion of abdomen, white scales cov-
ering nearly all of preapical segment and api-
cal segment except for small black spot. Ven-
ter with central dark portion toward which
lateral white bands converge, forming latero-
ventral white spots. Legs long, second and
third pairs successively larger. All legs black,
with basal and apical white spots on femora
and tibiae. Segments of tarsi with small white
spots at base and tip. Wings revetted with
small dark scales.

Male Terminalia: Coxite (Fig. 3^) triangu-
lar, tapering, external surface bearing many
long and strong setae, basal portion with mi-
cropili and sinuosities. Subterminal lobe prom-
inent, short, with 3 sinuous rods, median one
slightly curved apically, all 3 arising from

189

Fig. 3. Male terminalia of C. {Ph.) archegus. a,
Coxite and clasper; h, mesosomal plate, tenth sternites,
and ninth tergites.

tubercular bases, lobe also bearing very thin
hair-like filament slightly curved at tip and
shorter than rods. In addition, subterminal
lobe with very large ovate leaf and long rod-
like seta, almost as stout as rods. Clasper
curved, broadened centrally, tapered distally.

190

with triangular spine nearly terminal. Meso-
somal plate (Fig. 3^) with upper limb short,
lower recurved, thumb-shaped, with several
curved teeth between the lower and a very
large tooth which arises from base and which
tapers sharply distally and projects beyond
plate. Tenth sternites with tips tufted with
spines, with wide, very long curved arm from
base. Ninth tergites forming elongate lobes
with 6 to 8 thin long hairs, lobes very hairy
and well sclerotized.

Culex {Phalangomyia) quitensis n.sp.

LARVA (Fig. Aa, h ) : Head round. Antennae
long, tapering, with subapical hair tuft, spi-
culate body, 1 pair of long tapering hairs, 1
pair of short tapering hairs. Preantennal hairs
4 to 6 finely spinulate hairs. Interior dorsal
hairs a bunch of 4 to 6 finely spinulate hairs.
Exterior dorsal hairs 4 to 6 finely spinulate
hairs.

Comb of eighth abdominal segment (Fig. Ab)
formed by patch of oval teeth with apex
hairy. Air tube long, 3.5 times as long as wide.
Eecten of 14 to 16 conical scales with 4 to 6
teeth each. Tuft of pecten simple or double,
finely spinulate, other tufts formed of 1 to 3
spinulated hairs located at different places
along air tube. Anal segment wider than long,
spinulate, with small and medium spinules on
most of surface. Lateral hair tuft double, long
and spinulate. Dorsal brush formed of 3 long,
strong, spinulate hairs. Ventral brush formed
of 6 to 8 hair tufts of 4 elements each, very
well developed and long.

PUPA (Fig. 2c, d) (Chaetotaxy according to
Knight and Chamberlain, 1948): Segment I,
hair 10 well developed and long, hair 2 with
strong base, very ramified; Segment II, hairs
2, 4, 8 long and simple; Segment III, hairs 2,
5, 8 long and simple; Segment IV, hairs 2, 8
long and simple; Segment V, hairs 2, 5, 6 long
and simple; Segment VI, hairs 2, 5, 7 long and
simple; Segment VII, hairs 2, 5, 7 long and
simple, hair 8 long and diramified; Segment
VIII, hair 5 long and simple, 8 long and mul-
tiramified. Pupal trumpets longer than wide.

PACIFIC SCIENCE, Vol. VII, April, 1953

Fig. 4. Larval structures of C. iPh.) quitensis. a. Head;
b, cauda.

with wide pyramidal opening, rounded on one
angle, tapering and very sinuous. Pupal pad-
dles rounded, with strong median rib, apical
hairs medium.

ADULT (both male and female): Proboscis
black with white tip. Palpi black with white
tips, small and hairy in female, long and thin
in males. Vertex with white, truncate, triangu-
lar scales, dark triangular scales, and black
setae, forming tufts that overhang clypeus.
Occiput with white and black triangular scales.
Borders of eyes with dark setae. Prothoracic
lobes with integument dark brown, with long
and strong hairs. Mesonotum with two bare
lines, forming a lyre-like structure, with white
scales, very compressed, forming tufts at bases
of wings. Disc covered with dark-brown com- I
pressed scales and dark setae, posteriorly re- .
vetted with white scales and dark long setae.

Subgenus Phalangomyia — Levi-Castillo

191

Scutellum trilobed, revetted with white scales
and dark hairs. Postnotum bare with integu-
ment dark brown. Abdomen covered dorsally

Fig. 5. Male terminalia of C. (Ph.) quitensis. a,
Coxite and clasper; b, mesosomal plate, tenth stern-
ites,,and ninth tergkes.

by brown scales and with basal spots of sil-
very white scales in each segment which
become lateral bands. Venter with dark-
brown scales in straight line up to tip, bands
of white scales diverging from this line to
form lateral spots of white scales. Legs very
long, posterior pair longest. Pleurae and cox-
ae revetted with white scales forming tufts
and spots. Legs dark brown with femora and
tibiae white ventrally and on dorsum basally
and apically, segments of tarsi dark brown
with white spots on knees. Wings large with
small dark scales.

Male Terminalia: Coxite (Fig. 5^) triangu-
lar, tapering, external surface bearing many
short, strong setae, and very small hairs. Sub-
terminal lobe prominent, short, with 4 rods,
one smaller and with tip curved like question
mark, other 3 sinuous, slightly curved sub-
apically, lobe also bearing medium-sized ovate
leaf and long hair-like seta, all arising from
tubercular bases. Clasper curved, very broad
basally, tapering apically, with two small sub-
apical setae and very small terminal spine.
Mesosomal plate (Fig. ‘bh) with upper limb
broad, with several spine-like teeth, very large
formation of teeth from base curving sharply
outward and projecting as a point beyond
plate. Tenth sternites with tips tufted with
spines, with wide, curved, very long arm from
base. Ninth tergites forming elongate^ lobes
with 6 to 10 long, strong, terminally curved
hairs, lobes very hairy and well sclerotized.

TYPES: 2 males, 1 female, and their larval
and pupal pelts in the collection of the Ecua-
dorian Center for Entomological Research.

TYPE locality: Quito, Ecuador (South
America) .

REFERENCES

Dyar, H. G. 1928. The mosquitoes of the

Americas. Carnegie Inst. Wash., Tub. 387:

358-360.

1929. Remarks on the subgenus Fha-

langomyia of Culex Linn. Amer. Jour. Hyg.

9(2): 511.

192

PACIFIC SCIENCE, Vol. VII, April, 1953

and F. Knab. 1914. New mosquitoes

from Peru. Insecutor Inscitiae Menstruus 2(4) ;
58-61.

Edwards, F. W. 1932. Diptera family Culi-
cidae in Wytsman’s Genera Insectorum. Fasc.
194. 258 pp. Brussells.

Knight, K. L., and R. W. Chamberlain.

1948. A new nomenclature for the chaeto-
taxy of the mosquito pupa, based on a
comparative study of the genera. Helmin-
thoL Soc. Wash., Proc. 15(1): 1-18.

Lane, J. 1939. Catalogo dos mosquitos neo-
tropicos. BoL Biol. Sao Paulo, Ser. Monog.
No. 1: 46.

A Systematic Catalogue of Australian Braconidae

Arthur W. Parrott^

INTRODUCTION

It has been 60 years since a list of the
Braconidae of Australia was published, ex-
cepting the world lists compiled by Dalla
Torre (1898) and Szepligeti (1904). In 1891,
Froggatt published his "Catalogue of De-
scribed Hymenoptera of Australia” in which
he lists eight species of braconids. Nine years
later Dalla Torre’s world catalogue appeared
in which he records 28 species from Austra-
lia and Tasmania. As far as the Australian
fauna is concerned, there are errors and om-
missions which diminish the value of this
great work, and in the present catalogue it is
referred to only when the nomenclature of a
species is affected. For example, Dalla Torre
does not mention Guerin’s genus Trachypectus
and records Bracon hicolor Brulle from Africa
and Myosoma mutator Fabr. from America. The
next list was published in Szepligeti’s "Genera
Insectorum,” Parts I and II, which appeared
in 1904. While Szepligeti was preparing this
work, he was actively engaged in a study of
the Australian fauna and thus added greatly
to our knowledge of this group. He lists 37
species from Australia and Tasmania. The
present list records 224 species from Australia
and Tasmania.

Unquestionably the taxonomy of the Bra-
conidae is rather confused. Further studies
will undoubtedly result in much more syno-
nymy and in the resurrection of some species
and genera now suppressed. The great divers-
ity of species occurring in Australia and neigh-
bouring regions necessitates an enormous

1 Entomologist, Cawthron Institute, Nelson, New
Zealand. Manuscript received March 1, 1952.

amount of work before a proper arrangement
of species and their synonymy can be formu-
lated. The arrangement of the genera and
higher groups, in the present catalogue, is
substantially that used by Muesebeck (1951:
90-184).

For the majority of species listed, I have
had access to the original works, but those
that were not available to me I have checked
with other authors and in the Zoological Rec-
ord. In compiling the synonymy of the older
species, especially those of Fabricius and
Brulle, and of the several more or less cos-
mopolitan species, a complete bibliography
is not given, but invariably the original ref-
erence, together with all subsequent references
where nomenclature is affected, is included.
All references are included when a fuller de-
scription or figures are given, or when ref-
erence is made to the host or distribution of
the species concerned. At the end of the cata-
logue is a list of beneficial species introduced
and liberated for purposes of biological con-
trol. Established aliens, accidentally introduced
species, and those occurring naturally outside
the Australian area are given in the body of
the catalogue with an indication of their dis-
tribution outside Australia. A host index is
included as well as a general index, listing all
subfamily, generic, specific, and subspecific
names included in the catalogue.

Every endeavour has been made to make
this catalogue as complete as possible; never-
theless, there may be ommissions, and perpet-
uations of errors occurring in early records,
and subsequent authors’ corrections may have
been overlooked, but it is hoped that such
errors have been reduced to a minimum.

193

194

PACIFIC SCIENCE, Vol. VII, April, 1953

HISTORICAL

The history of Australian braconid system-
atics may be divided conveniently into four
major periods of activity, namely, 1777 to
1890, 1891 to 1913, 1914 to 1927, and 1928
to 1951.

First Period 1777-1890.

Australian braconid systematics com-
menced with the publication of the descrip-
tion of new species by Fabricius in 1777, and
was ably carried on by Brulle in 1846. Other
workers who described Australian species dur-
ing this period were Erichson, Guerin-Mene-
ville, and Holmgren. During this period, 15
species were described from Australia and
Tasmania. Fabricius worked on the collec-
tions made by Banks and Solander, naturalists
on the "Endeavour” during Cook’s voyages.
These early Australian species were collected
by Banks and Solander on the return journey
of the "Endeavour” to England after observ-
ing the transit of Venus at Tahiti. The "En-
deavour” was in Australian waters from April
to August, 1770, and, according to Musgrave
(1930; 190), the insects were mostly captured
around Botany Bay and the east coast of
Australia. From this material, four species of
Braconidae were described, and the types are
in the Banksian Collection in the British
Museum. It was not until 1824 that the next
braconids to be described from Australia were
obtained during the voyage of the "Coquille”,
commanded by L. I. Duperrey. Lessen and
Garnot were the naturalists, and the insects
were later described by Guerin-Meneville. A
footnote (a) by Musgrave (1930; 194) states
"In Duperrey’s 'Voyage autour du Monde
. . . sur ... La Coquille, pendant les Annees
1822-1825.’ The plates, with the scientific
names of the insects appeared between 1827-
1832, but the text did not appear until 1838.”

Erichson described a species from Tasmania
in 1841. Five years later, Brulle (1846) de-
scribed eight species from Australia and Tas-
mania in his great work "Histoire Naturelle
des Insects.” A specimen collected at Sydney,

New South Wales, was described by the well-
known European hymenopterist Holmgren in
1868. No further species were described dur-
ing this period. Thus, 15 species of braconids
were known from Australia and Tasmania to
1890.

Second Period 1891-1913

This period may conveniently begin with
the publication of Froggatt’s "Catalogue of
Described Hymenoptera of Australia,” in
which he lists eight species of braconids from
Australia (actually 15 species were known at
this time). During the 22 years covered by
this period, Ashmead, Froggatt, Cameron,
Szepligeti, Bingham, and Kokujew contribut-
ed to the knowledge of the Australian mem-
bers of this family. Szepligeti, between the
years 1901 and 1908, played the most promi-
nent part.

A German expedition collected a number
of insects in southwest Australia in 1905
which later were described by specialists in
"Die Fauna Siidwest-Australiens. 1907-1910,”
in which Szepligeti (1908) described the Bra-
conidae.

Third Period 1914-1927

The works of R. E. Turner (1917-27) dom-
inate this period, although those of Froggatt,
Fullaway, and Baker must be mentioned. The
period closes with the publication in 1926 of
Baker’s work on the Australian and Philip-
pine Cheloninae.

An expedition, headed by Dr. E. Mjoberg,
traveled into Queensland and northwest Aus-
tralia. The Braconidae were described by
Roman (1915).

Fourth Period 1928-1931

During this period a great amount of im-
portant revisional work was undertaken, prin-
cipally by Wilkinson, on the Microgaster-
inae; this work was ably continued by Nixon.
These authors dealt with the faunas of large
regions and placed the systematics of Aus-
tralian Braconidae on a sound basis in indi-
cating their relationship with other faunas.

Australian Braconidae — PARROTT

195

The works of Wilkinson and Nixon in
England and Muesebeck in the United States
have undoubtedly contributed toward a
sounder knowledge of braconid systematics
generally, and their work forms a basis for a
clearer interpretation of the Australian forms
in particular.

Other workers have contributed to the
knowledge of the Australian braconid fauna,
but those mentioned have made the principal
contributions.

ACKNOWLEDGMENTS

To Dr. Leonard Tuthill, Professor of En-
tomology, University of Hawaii, Honolulu,
I want to convey my very sincere thanks for
suggestions on the layout of this catalogue;
without his kindly co-operation this work
would not have been completed.

To Dr. David Miller, Director, and Mr.
L. J. Dumbleton, Senior Entomologist, En-
tomological Research Station, Nelson, New
Zealand, my thanks are due for helpful sug-
gestions during the final stages of preparation
of this catalogue.

To Miss Shirley Armstrong, Librarian, En-
tomological Library, Cawthron Institute, Nel-
son, New Zealand, I am indebted for her
untiring efforts in tracing references and ob-
taining publications.

To Mr. G. E. J. Nixon, British Museum
(Natural History), London, Mr. Anthony
Musgrave, Entomologist, and Mr. G. Gross,
Assistant Entomologist, Australian Museum,
Sydney, go my thanks for the literature which
they supplied.

To Mr. Erank Wilson, Principal Research
Officer, C.S.I.R.O., Canberra, Australia, I am
indebted for the list of introduced Braconidae.

To Dr. Rene Malaise, Natural History Mu-
seum, Stockholm, my thanks are due for a
list of Roman’s types and other Australian
species in the collections of the Museum.

ABBREVIATIONS OF MUSEUMS

The following abbreviations are used in
giving the location of types of Australian
Braconidae.

B.M.

H.D.O.M.

H.E.S.

N.C.A.C.

N.H.M.

N.H.M.S.

N.H.M.V.

N.K.

S.C.M.G.

U.S.N.M.

British Museum of Natural
History, London

Hope Department, Oxford
Museum, Oxford.

Hawaii Entomological Society
Collections, Honolulu.

National Collections of Aus-
tralia, Canberra.

National Hungarian Museum,
Budapest.

Natural History Museum,
Sweden.

Natural History Museum,
Vienna.

Location not known.

Saussure Collection, in Mu-
seum, Geneva.

United States National Mu-
seum, Washington.

Order HYMENOPTERA

Suborder apocrita ( = clistogastra)
Superfamily ichneumonoidea

Eamily BRACONIDAE

Subfamily APHIDIINAE

All members of this subfamily are internal
parasites of aphids.

Genus Diaeretus Eoerster, 1862

rapae (Curtis)

Curtis [Aphidius] (1855, 2: 194) 9 descr.
Ashmead [Lipolexis] (1900: 353) Austr. rec-
ord.

Eroggatt [Lepolexis] (1907; 87) noted.
Cameron [Lipolexis] (1912; 197) Austr. rec-
ord.

Smith (1944: 101) syn., descr., fig.

Loc: Victoria; New South Wales.

Hosts; Rhopalosiphum pseudobrassicae
(Davis).

Mysus persicae (Sulz) .

Brevicoryne hrassicae (Linn.).

Type: N.K.

Note: This European species is now well
established in at least the southern

196

PACIFIC SCIENCE, Vol. VII, April, 1953

states of Australia and also in New
Zealand.

Genus Ephedrus Haliday, 1833

persicae Froggatt

Froggatt (1904: 611) 9 descr.

Froggatt (1907: 86) note, fig.

Froggatt (1910: 342) host, fig.

Loc: New South Wales.

Host: Peach aphis.

Type: N.K.

Note: This species is probably a European
species that has become established
in Australia.

Subfamily EUPHORINAE
(= Meteorinae)

The members of this subfamily attack a
wide range of hosts, but all species are in-
ternal parasites.

Genus Aridelus Marshall, 1887

Helorimorpha Schmied., 1907)
exiles (Turner)

Turner [Helormorpha] (1927: 558) cf descr.
Loc: Mackay, Queensland.

Type: B.M.

Genus Meteorus Haliday, 1835
(== P rote his EoQVSttr:, 1862)

Most species of this genus are parasitic on
lepidopterous larvae, but some are known to
parasitise Coleoptera.
antipodalis Ashmead
Ashmead (1900: 353) 9 descr.

Loc.: New South Wales.

Type: U.S.N.M. (No. 4908).
bicolor Szepligeti
Szepligeti (1905: 53) 9 descr.

Loc: New South Wales.

Type: N.H.M.
dumbletoni Muesebeck

Muesebeck (1939: 172) cT* 9 descr.
Dumbleton (1940: 325a) host, bioL, distr.
Loc: Hobart, Tasmania.

Host: Tortrix postvittana Walk.

Type: U.S.N.M. (No. 53341).

lutens (Cameron)

Cameron [Protelus] (1911: 341) 9 descr.
Loc: Sydney, New South Wales.

Host: Teara sp. [Notodontid.].

Type: B.M.

Genus Perilitus Nees, 1818
(= Dinocampus PoQtstQT, 1862)

The species of this genus are internal para-
sites of adult Coleoptera.
coccinellae (Schrank)

Schrank [Ichneumon] (1802, 2: 310) 9 descr.
'^QQs[Braconterminatus]{lSll\26) 9 descr.
Nees [Perilitus terminatus] (1834: 30) 9

descr.

Dalla Torre (1898: 122) listed.

Szepligeti [Dinocampus terminatus] (1904:
175) listed.

Timberlake [D. terminatus] (1918: 401)
Austr. record.

Cushman [Dinocampus] (1922: 242) syn.
Muesebeck (1951: 101) syn., listed.

Loc: Queensland. This species is nearly

cosmopolitan.

Hosts: Hippodamia convergens Guer.

H. 3-signata (Kby.).

Coccinella californica Mann.
Coleomegilla maculata (Deg.),
Cycloneda sanguinea (L.).

C. trifasciata juliana Moils.

Type: N.K.

Subfamily MACROCENTRINAE

Internal parasites of lepidopterous larvae.
The gregarious forms seem to be polyembry-
onic.

Genus Macrocentrus Curtis, 1833
(= Amicroplus Eoetster, 1862)

( = Ehogra Cameron, 1901^)

ancylivorus Rohwer

Rohwer (1923: 56) 9 descr., N. America.
Loc: Liberated 'in Goulburn Valley, Vic-

toria.

Host: Grapholitha molesta, Busck. [Ole-
threutid.].

Type: U.S.N.M.

Australian Braconidae — PARROTT

197

rubromaculatus (Cameron)

Cameron \Fhogra\ (1901^/ 104) descr., New
Zealand.

Wilkinson [Amicroplus tasmanicm] (1928^;

265) descr. [New synomymy].

Loc: New Zealand; Longford, Tasmania,

Host: Argotis sp. (or allied Noctuid).
Type: B.M.

Subfamily HELCONINAE
The species of this subfamily, with few
exceptions, appear to be parasitic on wood-
boring coleopterous larvae.

Tribe helconini

Genus Austrohelcon Turner, 1918
australiansis (Kokujew)

Kokujew [Helcon] (1901: 15) & descr.
Szepligeti [Aspidocolpus] (1904: 152) listed.
Turner (1918^/ 165) nomencl., note.

Loc: Tasmania (Kokujew); New South

Wales (Turner).

Type: N.K.
erythrocephalus Turner
Turner {lS>lSd: 167) cf descr.

Loc: Victoria.

Type: B.M.
indultor (Erichson)

Erichson [Helcon] (1842: 258) cT’ descr.
Dalla Torre [Helcon] (1898: 84) listed.
Szepligeti [Helcon] (1904: 151) listed.
Turner (1918V.' 165) nomencl., note.

Loc: Tasmania.

Type: N.K.
inornatus (Kokujew)

Kokujew [Helcon] (1902: 15) cT descr.
Szepligeti [Aspidocolpus] (1904: 152) listed.
Turner (1918V.' 165) nomencl., note.

Loc: Australia.

Type: N.K.
meridionalis Turner

Turner (1918V.' 167) descr.

Loc: Victoria.

Type: B.M.

Genus Calohelcon Turner, 1918
obscuripennis Turner
Turner (1918V.' 165) descr.

Loc: Victoria.

Type: B.M.

Genus Helcon Nees, 1814 (1812)

(= GymnoscelusYo^t^t^s, 1862)

rufithorax Turner

Turner [Gymnoscelus] (1918V.' 170) descr.
Loc: Victoria.

Type: B.M.
rufoniges Turner

Turner [Gymnoscelus] (1918V,' 169) descr.
Loc: Tasmania.

Type: B.M.

Genus Parahelcon Kokujew, 1901
konowi Kokujew

Kokujew (1901: 15) 9 descr., n. gen.
Cameron [Opius euthyrrhini] (1912: 197) 9
descr., host.

Loc: New South Wales, Australia.

Host: Euthyrrhinus meditahundas Eabr.
(palm weevil).

Type: N.K.; Cameron’s type, B.M.

Genus Trichiohelcon Turner, 1818

phoracanthae (Eroggatt)

Froggatt [Iphiaulax] (1916: 565) 9 descr.,

fig-

Froggatt [Iphiaulax] (1923: 42) descr., hg.,
host.

Turner (1918V,' 168) nomencl., note.

Loc: S. E. Australia; Tasmania.

Host: Trogodendron fasciculatum Schreiber
(Clerid.).

Genus Trachypetus Guerin, 1838

The true position of this genus is doubtful;
it was placed in the subfamily Cheloninae by
Szepligeti (1904: 99), but Brues transferred it
to the Helconinae.
clavatus Guerin

Guerin (1830: 201) cT* descr., fig.

Schulz (1911: 85) noted.

Brues (1920: 59) nomencl.

Loc: New South Wales, Queensland.
Type: N.K.

198

PACIFIC SCIENCE, Vol. VII, April, 1953

Tribe diospilini

Genus Diospilus Haliday, 1833

ruficeps Szepligeti

Szepligeti (1905: 14) descr.

Loc: New South Wales.

Type: N.H.M.

Genus Westwoodiella Szepligeti, 1904

bicolor Szepligeti

Szepligeti (1904: 155) 9 descr.

Loc: Sydney, New South Wales,

ruficeps (Brulle)

Brulle [Westwoodia\ (1846: 127) descr., fig.
Loc: Tasmania.

Type: N.K.

Tribe zelini

This tribe has previously been placed in the
Macrocentrinae, but I have followed Muese-
beck (1951: 111) in placing it in the

Genus Zele Curtis, 1832

australiensis Nixon
Nixon (1938: 419) 9 descr.

Loc: Mackay, Queensland.

Type: B.M.

Subfamily BLACINAE
Genus Orgiloneura Ashmead, 1900

antipoda Ashmead

Ashmead (1900: 355) 9 descr.

Loc: Australia.

Type: U.S.N.M. (No. 4910).

Genus Orgilus Haliday, 1833
( = Ischius Wesmael, 1837)

leucogaster (Holmgren)

Holmgren {lschius\ (1868: 429) cf descr.
Froggatt \lschius\ (1890: 713) listed.

Dalla Torre (1898: 128) listed.

Szepligeti (1904: 120) listed.

Loc: Sydney, New South Wales.

Type: N.H.M.S.

Subfamily AGATHIDINAE

As far as is known, all species of this sub-
family are internal parasites of lepidopterous
larvae.

Genus Agathiella Szepligeti, 1902
festinata Turner

Turner (1918^,' 111) 9 descr., key.

Loc: South Australia.

Type: B.M.
latibalteata (Cameron)

Cameron {Agathis\ (1906: 26) cT’ descr.
Turner (1918^?.’ 110) note., nomencl., key.
Loc: Australia.

Type: B.M.
maligna Turner

Turner (1918^.' 112) 9 descr., key.

Loc: S.W. Australia.

Type: B.M.
meridionalis Turner

Turner (1918^; 110) 9 descr., key.
Dumbleton (1940: 325a) host.

Loc: Mt. Wellington, Tasmania.

Host: Tortrix postvittana Walk. (Tortri-
cid.).

. Type: B.M.
minima Turner

Turner (1918^; 113) 9 descr., key.

Loc: Kuranda, Queensland.

Type: B.M.

Note: 'Ttis possible that Ashmead’s genus
Orgiloneura may be founded on a
species of this genus, with some-
what reduced neuration, but his de-
scription is too short for any con-
clusions to be drawn,” (Turner
1918^; 113).
ruficeps Szepligeti

Szepligeti (1905: 52) cT descr.

Turner (1918^.' 110) note, key.

Loc: Sydney, New South Wales.

Type: N.H.M.
rugosa Turner

Turner (1918^; 112) 6^ 9 descr., key.

Loc: Tasmania.

Type: B.M.
tenuissima Turner

Turner (1918^.' Ill) 9 descr., key.

Loc: Victoria. \

Type: B.M.

Note: Possibly a female of A. ruficeps
Szepl. (Turner).

Australian Braconidae — PARROTT

199

tricolor Szepligeti

Szepligeti (1905: 52) 9 descr.

Turner (1918^/ 110) note, key.

Loc: Sydney, New South Wales.

Type: N.H.M.
unimaculata Turner

Turner {I9l8a: 111) d' descr., key.

Loc: Kuranda, Queensland; Sydney,

New South Wales.

Genus Agathis Latreille, 1804

( = Bassus Fabr., 1804)

(= Mkrodus Nees, 1814)

The two species described by Brulle in 1849
are listed under this genus, but it is doubtful
whether they are typical Agathis. In 1918
Turner (p. 110) stated "as far as I am aware
typical Agathis does not occur in Australia."
Brulle’s species may probably belong to Aga-
thiella Szebk Muesebeck (1951: 117) syno-
nymises Microdus Nees, 1814 (1812) with
Agathis Latr., 1804, and the four species de-
scribed by Turner under Microdus are listed
here under Latreille ’s genus,
bicolor Brulle

Brulle (1846:483) descr.

Froggatt (1890: 710) listed.

Loc: Australia.

Type: N.K.
dimidiata Brulle

Brulle (1846: 487) descr.

Froggatt (1890: 710) listed.

Loc: Tasmania.

Type: N.K.
martialis (Turner)

Turner \Microdus\ (1918^.* 108) d 9 descr.,
key.

Loc: Kuranda, Queensland.

Type: B.M.
pedunculatus Szepligeti

Szepligeti (1905: 51) 9 descr.

Turner (1918^.* 106) note.

Loc: Sydney, New South Wales.

Type: N.H.M.
rufithorax (Turner)

Turner \Microdus\ (1918^,' 106) 9 descr.,
key.

Loc: Kalamunda, Yallingup, S.W. Aus-

tralia.

Type: B.M.
rufobrunneus (Turner)

Turner \Microdus\ (1918^.' 106) 9 descr.,
key.

Loc: Townsville, Queensland.

Type: B.M.
xanthopsis (Turner)

Turner \Microdus\ (1918^.' 107) d 9 descr.,
key.

Loc: Yallingup, S.W. Australia.

Type: B.M.

Genus Biroia Szepligeti, 1900

solitaria Turner
Turner (1918c.* 229) d 9 descr.

Loc: Mackay, Queensland.

Type: B.M.

Genus Braunsia Kriechbaumer, 1894

bicolor Brulle

Brulle (1846: 483) 9 descr.

Dalla Torre \Agathis\ (1898: 138) listed.
Loc: Australia.

Type: N.K.
diversipennis Turner

Turner (1918c,‘ 224) d descr.

Loc: Mackay, Queensland.

Type: B.M.

Genus Cremnops Foerster, 1862

(= Bracon Auct. not of Fabricius. See Opin-
ion 162 of the Internatl. Comm, on Zool.

Nomencl. 1945.)
commutator Turner

Turner (1918c,’ 223) 9 descr., key.

Loc: Mackay, Queensland (Type), Ku-

randa, Townsville, Queensland;
Port Darwin, Northern Territory.
Type: B.M.
dissimilis Turner

Turner (1918c,* 222) d 9 descr., key.

Loc: Mackay, Queensland.

Type: B.M.
marginipennis Turner

Turner (1918c,* 222) d 9 descr., key.

200

PACIFIC SCIENCE, Vol. VII, April, 1953

Loc: Kuranda, Queensland.

Type: B.M.
xanthostigma Szepligeti

Szepligeti (1900: 61) 9 descr.

Turner (1918^/ 221) note.

Loc: Kuranda, Mackay, Queensland; al-

so New Guinea.

Type: N.H.M.

Genus Disophrys Foerster, 1862
diluta Turner

Turner (I9I80* 228) 9 descr., key.

Loc: Mackay (type), Kuranda, Queens-

land.

Type: B.M.
diversipes Turner

Turner (1918c.” 227) 9 descr., key.

Loc: Kuranda (type), Mackay, Queens-

land.

Type: B.M.
exornata Turner

Turner (1918c.' 227) 9 descr., key.

Loc: Kuranda, Queensland.

Type: B.M.
nigropectus Turner
Turner (1918c.‘ 229) 9 descr., key.

Loc: Kuranda, Queensland.

Type: B.M.
ruberrima Turner

Turner (1918c.' 226) 9 descr., key.

Loc: Mackay (type), Townsville,

Queensland. . ^

Type: B.M.
rufifrons Turner

Turner (1918c.' 226) 9 descr., key.

Loc: Port Essington, Northern Territory.

Type: B.M.
signatipennis Turner

Turner (1918c.' 225) & descr.

Loc: Kuranda, Queensland.

Type: B.M.
similipicta Turner

Turner (1918c.' 228) 9 descr., key.

Loc: Mackay (type) Townsville, Queens-

land.

Type: B.M.

Genus Platyagathis Turner, 19 18
leaena Turner

Turner (1918^3^.’ 114) cT 9 descr., new
genus.

Loc: Yallingup, S.W. Australia.

Type: B.M.

Subfamily MICROGASTERINAE

All the species of this subfamily appear to
be internal parasites of lepidopterous larvae.

Genus Adelius Haliday, 1833

australiensis (Ashmead)

Ashmead \Acoelius\ (I9OO: 358) 9 descr.
Loc: Australia.

Type: U.S.N.M. (No. 4911).

Genus Apanteles Foerster, 1862
anthelae Wilkinson

Wilkinson (1932^?.' 335) cP 9 descr., key.
Loc: Victoria.

Host: Anthela ocellata Walker (Anthelid.).
Type: B.M.
australiensis Ashmead
Ashmead (1900: 356) 9 descr.

Wilkinson (1930: 485) host. ‘

Wilkinson {I9^2a: 334) key.

Loc: New South Wales, Victoria.

Host: Antheraea eucalypti Scott (Satur-
niid.).

Type: U.S.N.M. (No. 4931).
biroi Szepligeti
Szepligeti (1905: 49) descr.

Wilkinson (1928: 121) cf 9 descr., key.
Wilkinson (1932^.' 337) key.

Loc: Sydney, New South Wales.

Type: N.H.M.
deliadis Bingham

Bingham (I906: 125) & 9 descr., host. j
Wilkinson (1932^; 334) 9 descr., key. j

Loc: Townsville, Queensland. |

Host: Delias argenthonaE2}oi. \

Type: H.D.O.M. !

flavipes (Cameron) j

Cameron \Cotesia\ (1891: 185) descr.

Olliff [Apanteles nonagriae] (1893: 381) I

descr., fig. I

Australian Braconidae — PARROTT

201

Wilkinson (1928^/ 93) & 9 descr.
Wilkinson [A. nonagriae] (1928^; 136)
descr.

Wilkinson (1929^.' 108) descr.

Wilkinson (1932: 337) key.

Wilkinson (1934A' 156) key.

Loc: Australia; also India and Formosa.

Host: Phragmatiphila truncata Walk.

(Noctuid.).

Type: B.M.
glomeratus (Linnaeus)

Linnaeus [Ichneumon} (1758: 568) descr.
Wilkinson (1932^.' 334) key.

Loc: Nearly cosmopolitan; introduced

into Australia.

Hosts: Pieris rapae (Linn.) (Pierid.).

P. hrassicae (Linn.) (Pierid.).

Type: N.K.
guyanensis Cameron
Cameron (1911: 327) cf 9 descr.
Wilkinson (1930: 483) host., key.

Loc: Western Australia; also British Gui-

ana.

Host: Probably Utsthesia pulchella Lis.

(Arctiid.).

Type: B.M.
philoeampus Cameron

Cameron (1911: 342) 9 descr.

Wilkinson (1928r,' 96) descr.

Wilkinson (1932^.* 333) key.

Loc: Wattle Flat, New South Wales.

Type: B.M.
radiantis Wilkinson

Wilkinson (1929^.* 110) cT’ 9 descr., host.
Wilkinson (1932^.- 334) key.

Loc: Gatton, Queensland.

Host: Euxoa radians Guen. (Noctuid.).
Type: B.M.
rubecula Marshall

Marshall (1885: 175) 9 descr.

Marshall (1889: 420) 9 descr.

Wilkinson (1932^,‘ 334) key, Austr. record.
Loc: Europe; Australia; New Zealand.

Host: Pieris rapae lAnn. (Pierid.).

Type: N.K.
ruficrus (Haliday)

Haliday [Microgaster] (1835: 253) cT 9

descr.

Ashmead [Apanteles antipoda] (1900: 355)
cf 9 descr., host.

Tryon (1900: 142) descr., host hg.
Cameron [A. sydneyensis] (1911: 342) 9
descr., host.

Viereck [A. narangae] (1913: 642) descr.
Wilkinson [A. antipoda} (1928: 95) descr.
Gahan (1928: 256) syn.

Wilkinson (1929^.* 108) note, syn.
Wilkinson (1932: 333) syn., key.

Loc: Nearly cosmopolitan; Queensland;

New South Wales.

Hosts: Heliothis armigera ■ Hubn. (Noc-
tuid.).

Agrotis sp. (Ashmead) (Noctuid.).
Plusia sp..^ (Cameron) (Noctuid.).
Leucania unipuncta Haworth (Noc-
tuid.).

Type: N.K. (type A. antipoda Ash.,
U.S.N.M. No. 4912).
rufiventris (Bingham)

Bingham [Protapanteles} (1906: 127) 9

descr., host.

Wilkinson (1932^.' 335) key, host.

Loc: Townsville, Queensland.

Type: H.D.O.M.

Host: Ogyris genoveve Wo'w. (Lycaenid.).
sicarius Marshall

Marshall (1885: 209) d" 9 descr.
Wilkinson {Vy^2a: 337) key.
tasmanica Cameron

Cameron (1912: 196) 9 descr.

Wilkinson (1932^.' 337) cf 9 key.
Dumbleton (1940: 325a) host, N. Z. record.
Dumbleton (1935: 572) host, bioL, fig.
Loc: Tasmania; New Zealand.

Type: B.M.

Host: Tortrix postvittana Walk.
(Tortricid.).

Genus Microgaster Latreille, 1804

magnifica Wilkinson

Wilkinson (1929^.* 120) cT 9 descr., key.
Loc: Queensland.

Type: B.M.
morata Wilkinson

202

PACIFIC SCIENCE, Vol. VII, April, 1953

Wilkinson (1929^/ 106) d' 9 descr., key.
Loc: Victoria.

Host: Thyridopteryx herrkhi Westr. (Psy-
chid.).

Type: B.M.
perniciosa Wilkinson

Wilkinson (1929^.' 112) 9 descr., key.
Loc: Victoria.

Type: B.M.
petiolaris Szepligeti

Szepligeti (1905: 48) d 9 descr.
Wilkinson (1929^." 104) d 9 descr., key.
Loc: New South Wales.

Type: N.H.M.
resplendens Wilkinson

Wilkinson (1929^.' 106) 9 descr., key.

Loc : Mt. Wellington, Tasmania (2,000 ft.)

Type: B.M.
rixosa Wilkinson

Wilkinson (1929^.' 108) d 9 descr., key.
Loc: New South Wales.

Type: B.M.
sons Wilkinson

Wilkinson (1932A' 87) descr., fig.

Loc: Brindabella, A.C.T.

Type: N.C.A.
tearae Wilkinson

Wilkinson (1929^.' 107) 9 descr., key.,
host.

Loc: Australia.

Host: Teara tristis Lewin (Notodontid.).
Type: B.M.
tegularis Szepligeti

Szepligeti (1905: 49) d descr.

Wilkinson (1928^.* 105) d descr., key.
Loc: New South Wales.

Type: N.H.M.
vulpina Wilkinson

Wilkinson (1929^.* 109) d 9 descr., key.
Loc: Victoria.

Type: B.M.

Genus Microplitis Foerster, 1862 .

basalis (Bingham)

Bingham {M.krogaster\ (1906: 123) d descr.
Dodd {MkYogaster\ (1906: 120) host.
Wilkinson (1929^.' 122) d 9 descr.

Wilkinson (1930^.' 24) host.

Loc: Queensland.

Host: Theratra oldenlandiae Walk.

Type: H.D.O.M.
demolitor Wilkinson

Wilkinson {193^a: 119) cT" 9 descr., host.
Loc: Stanthorpe, Queensland.

Hosts: Heliothk ohsoleta^koi. (Noctuid.).

H. armigera Hubn. (Noctuid.).
Type: B.M.
perelegans (Bingham)

Bingham [Mkrogaster] (1906: 126) 9

descr.

Dodd [Mkrogaster] (1906: 120) host.
Wilkinson (1929^/ 122).

Wilkinson (1930^.' 24) host.

Loc: North Queensland.

Host: Pheressaces cycnoptera Lowes (Noto-
dontid.).

Type: H.D.O.M.

Subfamily CARDIOCHILINAE

There are no host records from Australia,
but, from the few host records available from
other countries, the species of this family
apparently parasites lepidopterous larvae.

Genus Cardiochiles Nees, 1818
assimilator Turner

Turner (1918A 49) d 9 descr.

Loc: Kuranda, Mackay, Queensland.

Type: B.M.
dissimulator Turner

Turner (1918A' 50) 9 descr., key.

Loc: Thursday Island, Queensland.

Type: B.M.
rufator Romans

Romans (1915: 17) 9 descr.

Loc: Kimberley, N.W. Australia.

Type: N.H.M.S.
uniformis Turner

Turner (1918A 31) d 9 descr., key.

Loc: Mackay, Queensland.

Type: B.M.
verticalis Turner

Turner (1918^* 51) 9 descr.

Loc: Mackay, Queensland. ^

Australian Braconidae — PARROTT

203

Type: B.M.

Subfamily CHELONINAE

Parasites of lepidopterous larvae. Eggs are
laid in eggs of the hosts but the larvae of the
parasite do not become mature until the hosts
have reached maturity.

Genus Ascogaster Wesmael, 1835

abdominalis Szepligeti

Szepligeti (I9O8A 319) 9 descr.

Loc; S.W. Australia.

Type: N.H.M.
antennalis Szepligeti

Szepligeti (I9O8A 319) descr.

Loc: S.W. Australia.

Type: N.H.M.
australiensis Szepligeti

Szepligeti (1905: 47) descr.

Loc: S.W. Australia.

Type: N.H.M.
caudata Szepligeti

Szepligeti (1905: 46) descr.

Loc: Australia.

Type: N.H.M.
cinctus Baker

Baker (1926: 477) 9 descr., key.

Loc: New South Wales.

Type: U.S.N.M.
detectus Baker

Baker (1926: 478) cf descr., key.

Loc: New South Wales.

Type: U.S.N.M.
distinctus Baker

Baker (1926: 481) 9 descr., key.

Loc: New South Wales.

Type: U.S.N.M.
erythropus Cameron

Cameron [Areogaster] (1911: 340) 9 descr.

Loc: Mittagong, New South Wales.

Type: B.M.

Note: Areogaster, which occurs in Camer-
on, 1911, is an evident typographi-
cal error for Ascogaster.
intensus Baker

Baker (1926: 473) cf descr., key.

Loc: New South Wales.

Type: U.S.N.M.
laeviventris Baker

Baker (1926: 475) 9 descr., key.

Loc: New South Wales.

Type: U.S.N.M.
maculaticeps Baker

Baker (1926: 482) <T descr., key.

Loc: New South Wales.

Type: U.S.N.M.
modestus Baker

Baker (1926: 474) 9 descr., key.

Loc: New South Wales.

Type: U.S.N.M.
palpalis Szepligeti

Szepligeti (1905: 46) descr.

Loc: ' Australia.

Type: N.H.M.
tegularis Szepligeti

Szepligeti (1905: 46) descr.

Loc: Australia.

Type: N.H.M.
tricolor Szepligeti

Szepligeti (1905: 46) descr.

Loc: Mt. Victoria, New South Wales.

Type: N.H.M.
vividus Baker

Baker (1926: 479) cT’ descr., key.

Loc: New South Wales.

Type: U.S.N.M.

Genus Chelonus Panzer, 1806

australiensis Szepligeti
Szepligeti (1905: 46) descr.

Loc: New South Wales.

Type: N.H.M.
coriaceus Szepligeti

Szepligeti (1905: 45) descr.

Loc: New South Wales.

Type: N.H.M.
megaspilus Cameron

Cameron (1911: 341) 9 descr.

Loc: Tamworth, New South Wales.

Type: B.M.
similis Szepligeti

Szepligeti (1905: 44) descr.

Loc: New South Wales.

Type: N.H.M.

204

unimaculatus Szepligeti
Szepligeti (1905: 45) descr.

Loc: New South Wales.

Type: N.H.M.

Genus Phanerotoma Wesmael, 1838

australiensis Ashmead

Ashmead (1900: 354) 9 descr.

Roman (1915: 17) d' descr., noted.

Loc: Broome, W. Australia (Roman).

Type: U.S.N.M. (No. 4904).
leeuwinensis Turner
Turner (1917: 247) descr.

Loc: Australia.

Type: B.M.

Genus Sigalphus Latreille, 1802

conjugator (Turner)

Turner [Sphaeropyx] (1917: 245) descr.

Loc: Australia.

Type: B.M.
tripartitus Szepligeti

Szepligeti (1905: 43) descr.

Loc: New South Wales.

Type: N.H.M.

Subfamily OPIINAE

The species of this subfamily are, with few
if any exceptions, parasitic on Diptera. Some
species are important in the biological control
of fruit-flies.

Genus Opius Wesmael, 1835

Refer to Muesebeck (1951: 153) for the
generic synonymy,
albimanus Szepligeti

Szepligeti (1905: 54) descr.

Loc: New South Wales.

Type: N.H.M.
brevicaudis Szepligeti
Szepligeti (1905: 25) descr.

Loc: New South Wales.

Type: N.H.M.
carpocapsae (Ashmead)

Ashmead [Dmcbasma] (1900: 357) descr.
Szepligeti [Dmchasma] (1904: 162) listed.
Loc: New South Wales; New Zealand.

PACIFIC SCIENCE, Vol. VII, April, 1953

Host: Probably Darus tryoni (Froggatt)
(Trypetid.).

Type: U.S.N.M. (No. 4914).
deeralensis Fullaway

Fullaway (1950: 65) descr., host.

Fullaway (1951: 244) note.

Loc: Deeral, Queensland.

Host: Dacus latkaudus Hardy.

Type: H.E.S.
fijiensis Fullaway

Fullaway (1950; 67) cf 9 descr., host,
recorded Australia.

Fullaway (1951: 244) descr., host.

Loc: Australia; Fiji; New Caledonia.

Host: Dacus sp.

Type: H.E.S.
froggatti Fullaway

Fullaway (1950: 67) descr., host.

Fullaway (1951: 250) note.

Loc: Deeral, Queensland; Atherton Ta-

bleland.

Host: Dacus sp.

Type: H.E.S.
kraussi Fullaway

Fullaway (1951: 249) cf 9 descr., host.
Loc: Deeral, Queensland.

Host: Dacus latkaudus Hardy.

Type: H.E.S.

Note: Reared in Hawaii from fruit-fly pu-
pae collected in Australia,
perkinsi Fullaway

Fullaway (1950: 66) descr., host.

Fullaway (1951: 244) listed, host.

Loc: Deeral, Queensland.

Host: Dacus latkaudtis Hardy.

Type: H.E.S.
persulcatus (Silvestri)

Silvestri \Biostem\ (1916: 167) descr.
Fullaway (1950: 67) note, Australian record.
Loc: Australia.

Host: Dacus sp.

Type: N.K.
tryoni Cameron

Cameron (1911: 343) d descr.

Gurney and Gallard (1910: 428) noted,, fig
Fullaway (1951: 243) noted, host.

Loc: New South Wales.

Australian Braconidae — PARROTT

205

Hosts: Dacus la^kaudus Hs-rdy (Trypetid.).

Dacus ferrugineus Fabr. (Trypetid.).

Ceratitis capitata Wied. (Trypetid.).

Type: B.M.

Subfamily BRACONINAE
{= V^pwnmae Yier.)

The members of this subfamily have a wide
variety of hosts, many of which are important
pests. Although Thomson’s genus Ipobracon
is closely related to, and considered by some
authors to be synonymous with, Foerster’s
genus Iphiaulax. I am at present retaining the
identity of Ipohracon.

Genus Atanycolus Foerster, 1862

tomentosus Szepligeti

Szepligeti (1905: 25) descr.

Loc: Queensland.

Type: N.H.M.

Genus Bracon Fabricius, 1804-05

Froggatt (1916: 561) describes a species of
Bracon reared from Lyctus hrunneiis infesting
tea chests that were imported from India,
australasicus Cameron

Cameron (1912: 193) 9 descr.

Loc: Australia.

Type: B.M.
australiensis Szepligeti

Szepligeti (1905: 38) descr.

Loc: Australia.

Type: N.H.M.
bimaculatus Szepligeti

Szepligeti (1905: 38) descr.

Loc: Australia.

Type: B.H.M.
bimaris Turner

Turner (1918^/ 93) 9 descr.

Loc: Tasmania.

Type: B.M.
capitator (Fabricius)

Fabricius [Ichneumon] (1775: 335) 9 descr.

Fabricius (1804: 108) descr.

Dalla Torre (1898: 261) listed.

Szepligeti (1904: 37) listed.

Loc: Australia.

Type: B.M.
defensor (Fabricius)

Fabricius [Ichneumon] (1775: 335) 9 descr.
Fabricius (1804: 108) descr.

Dalla Torre (1898: 262) listed.

Szepligeti (1904: 37) listed.

Loc: Australia.

Type: B.M.
eucalypti Cameron

Cameron (1911: 340) cA 9 descr.

Loc: Sydney, New South Wales.

Type: B.M,

Note: Bred from reddish galls on leaves
of eucalyptus,
froggatti Cameron

Cameron (1911: 339) 9 descr.

Loc: Reedy Creek, near Inverell, New

South Wales.

Type: B.M.
hartmeyeri Szepligeti

Szepligeti (1908: 317) descr.

Loc: S.W. Australia.

Type: N.H.M.
hebetor Say

Say (1836: 252) cf 9 descr.

Kirby [as hrevicornis] (1884: 31) descr.
Marshall [as hrevicornis] (1885: 24) descr.
Ashmead [diS juglandis] (1888: 621) descr.
Johnston [Macrohracon] (1895: 324)
Froggatt (1912: 310) descr., fig., host.
Myers [Microhracon] (1929: 425) recorded,
host.

Loc: Nearly cosmopolitan; Australia.

Hosts: Plodia interpunctella Huebner.
Ephestia kuehniella Zeller.

Ephestia eluttella Huebner.

Galleria mellonella Linnaeus.

Type: hehetor, \ost\ juglandis , U.S.N.M.
hospitator Fabricius

Fabricius [Ichneumon] (1775: 335) descr.
Fabricius (1804: 106) descr.

Dalla Torre [B. arvensis] (1898: 37) listed.
Szepligeti (1904: 37) listed.

Loc: Australia.

Type: B.M.
levisulcatus Cameron

Cameron (1912: 194) descr.

206

Loc: Reedy Creek, Inverell, New South

Wales.

Type: B.M.
pilitarsis Cameron

Cameron (1912: 193) 9 descr.

Loc: Reedy Creek, Inverell, New South

Wales.

Type: B.M.
pulchellus Brulle

Brulle (1846: 432) descr.

Froggatt (1890: 713) listed.

Dalla Torre (1898: 285) listed.

Szepligeti (1904: 37) listed.

Loc: Australia.

Type: N.K.
thalpocharis Ashmead
Ashmead (1900: 359) d' 9 descr.
Szepligeti [B. talpocharis] (1904: 46) listed
[error in spelling.]

Froggatt [Microhracon thalpocaris] (1907: 87)
descr.

Loc: Australia.

Host: Thalpocharis coccophaga Walker.
Type: U.S.N.M. (No. 4915).
tricolor Ashmead

Ashmead (1900: 359) 9 descr.

Szepligeti (1904: 46) listed.

Loc: Australia. ,

Type: U.S.N.M. (No. 4916).
turner! Cameron

Cameron (1906: 26) d descr.

Loc: Australia.

Type: N.K.
walkeri Turner

Turner (1918^.' 93) 9 descr.

Loc: Queensland; Northern Territory.

Type: B.M.

Genus Campyloneurus Szepligeti, 1906

australiensis (Szepligeti)

Szepligeti \lphiaulax\ 369) descr.

Szepligeti (1906: 561) n. gen.

Turner (1918^^; 100) key.

Loc: Australia.

Type: N.H.M.
biplicatus Roman

Roman (1915: 12) 9 d descr.

PACIFIC SCIENCE, Vol. VII, April, 1953

Loc: Kimberley District, N.W. Australia.

Type: N.H.M.S.
mutator (Fabricius)

Fabricius [Ichneumon] (1775: 335) descr.
Fabricius [Bracon] (1804: 109) descr.

Brulle \M.yosoma\ (1846: 453) descr.
Froggatt [M.yosoma\ (1890: 710) listed.
Dalla Torre [Myosoma\ (1898: 255) listed,
loc. error.

Turner (1918^?; 100) nomenclature.

Loc: Queensland.

Type: B.M.
praeclarus Turner

Turner (1918^.* 101) 9 descr., key.

Loc: Queensland.

Type: B.M.

Note: Very near C profugus Turner, of
which it may be a variety (Turner
1918^.* 101).
praepotens Turner

Turner (I9l8a: 101) 9 descr., key.

Loc: Queensland.

Type: B.M.
profugus Turner

Turner {I9l^a: 100) 9 descr., key.

Loc: Queensland.

Type: B.M.

Genus Cyanopterus Haliday, 1835

calligaster Roman

Roman (1915: 16) 9 descr.

Loc: Fremantle, W. Australia.

Type: N.H.M.S.
crassicaudis Szepligeti

Szepligeti [Bracon] (1901: 397) descr.
Szepligeti [Iphiaulax] (1904: 21) listed.
Turner (1918^; 94).

Loc: Doubtful, probably Australia.

Type: ^N.H.M.

Note: This may be synonymous with C.
profiscator (Fabr.) (Turner 1918^;
94).

innotatus Turner

Turner (1918^?; 95) 9 descr., key.

Loc: Queensland.

Type: B.M.

Australian Braconidae — PARROTT

207

proficiscator (Fabricius)

Fabridus [Ichnetmon] (1775: 335) descr.
Fabridus [Bracon] (1804: 105) descr.
Szepligeti [Bracon] (1904: 37) listed.

Turner (1918^.' 94) noted, synonymy, key.
Loc: Australia.

Type: B.M.
rufus Szepligeti

Szepligeti [Iphiaulax] (1901: 397) descr.
Szepligeti [Iphiaulax] (1904: 23) listed.
Szepligeti (1906: 586) genus changed.
Turner (1918^; 95) noted, key.

Loc: New South Wales.

Type: N.H.M.

Genus FIybogaster Szepligeti, 1906

australiensis (Ashmead)

Ashmead [Iphiaulax] (1900: 360) cf descr.
Szepligeti (1904: 23) listed.

Roman (1915: 15) d’ 9 noted, nomen-
clature.

Loc: Australia, Kimberley (Roman).

Type: U.S.N.M. (No. 4918).

Genus Iphiaulax Foerster, 1862
(= Ipohracon Dalla Torre, 1898)

afer Szepligeti

Szepligeti (1905: 29) descr.

Loc: Australia.

Type: N.H.M.
australis Szepligeti

Szepligeti (1901: 369) cT descr.

Szepligeti (1904: 23) listed.

Loc: Australia.

Type: N.H.M.
bifasciatus Szepligeti

Szepligeti (1905: 28) descr.

Loc: Australia.

Type: N.H.M.
biparti tus Szepligeti

Szepligeti (1905: 35) descr.

Loc: Australia.

Type: N.H.M.
bisignatus Szepligeti

Szepligeti (1905: 29) descr.

Loc: Australia.

Type: N.H.M.

camerunensis Szepligeti
Szepligeti (1905: 29) descr.

Loc: Australia.

Type: N.H.M.
crenulatus Szepligeti

Szepligeti (1905: 34) descr.

Loc: Australia.

Type: N.H.M.
cristatus Szepligeti

Szepligeti (1905: 30) descr.

Loc: Australia.

Type: N.H.M.
limbatus (Brulle)

Brulle [Bracon] (1846: 433) 9 descr.
Froggatt [Bracon] (1891: 713) listed.
Ashmead [Callihracon] (1900: 358) descr.
n. gen.

Szepligeti (1904: 23) listed.

Froggatt [Bracon] (1907: 86) descr.

Loc: Australia.

Type: N.K.
longicornis Szepligeti
Szepligeti (1905: 32) descr.

Loc: Australia.

Type: N.H.M.
longiseta Szepligeti

Szepligeti (1905: 28) descr.

Loc: Australia.

Type: N.H.M.
micans Szepligeti

Szepligeti (1905: 35) descr.

Loc: Australia.

Type: N.H.M.
perspicax Szepligeti

Szepligeti (1905: 32) descr.

Loc: Australia.

Type: N.H.M.
pretiosus Szepligeti

Szepligeti (1905: 36) descr.

Loc: Australia.

Type: N.H.M.
pulchricaudis Szepligeti
Szepligeti (1905: 30) descr.

Loc: Australia.

Type: N.H.M.
rubriceps Froggatt

Froggatt (1916: 564) 9 descr., fig.

208

PACIFIC SCIENCE, Vol. VII, April, 1953

Eroggatt (1923: 40) descr., fig., host.

Loc: New South Wales.

Host: Trogodendron fasciculatum Schreibers
(yellow-horned clerid.).

Type: N.K.
ruficeps Szepligeti

Szepligeti (1905: 33) descr.

Loc: Australia.

Type: N.H.M.
scoparius Szepligeti

Szepligeti (1905: 31) descr.

Loc: Australia.

' Type: N.H.M.
semirufus Szepligeti

Szepligeti (1905: 33) descr.

Loc: Australia.

Type: N.H.M;
speciosissimus Szepligeti
Szepligeti (1905: 31) descr.

Loc: Australia.

Type: N.H.M.
transiens Turner

Turner (1918^; 95) cf’ 9 descr.

Loc: Queensland; Northern Territory;

N.W. Australia.

Type: B.M.
trifasciatus Szepligeti

Szepligeti (1905: 31) descr.

Loc: Australia.

Type: N.H.M.
trinotata Ashmead
Ashmead (1900: 359) (T descr.

Szepligeti (1904: 23) listed.

Loc: Australia.

Type: U.S.N.M. (No. 4917).

Genus Ipobracon Thomson, 1892

emeraldensis Eahringer
Eahringer (1942: 40) descr.

Loc: Australia.

Type; N.H.M.
flaviceps (Cameron)

Cameron [Poecilobracon] (1901a: 122) cf
descr.

Szepligeti [Jphiatdax\ (1904: 23) listed.
Turner (1918^.‘ 105) descr., variety.

Loc: Australia.

Type: B.M.

flaviceps Roman mackayensis Turner
Turner (1918^.' 105) descr.

Loc: Queensland.

Type: B.M.
frater cuius Turner

Turner (1918^; 105) 9 descr.

Loc: S.W. Australia.

Type: B.M.
gilberti Turner
Turner (1918^.- 104) 9 descr.

Loc: Queensland.

Type: B.M.
ingressor Turner

Turner (1918^; 102) 9 descr.

Loc: Queensland.

Type: B.M.
pallidicolor Turner

Turner (1918^; 102) d' 9 descr.

Loc: Queensland.

Type: B.M.
punctistigma Roman

Roman (1915: 10) 9 descr.

Loc: Kimberley District, N.W. Australia.

Type: N.H.M.S.
quadricolor Turner

Turner (1918a: 103) 9 descr.

Loc: Queensland.

Type: B.M.
torridus Turner

Turner (1918^.* 104) 9 descr.

Loc: Queensland.

Type: B.M.

Genus Macrobracon Szepligeti, 1902
nobilis Turner

Turner (1918^; 96) d 9 descr.

Loc: Queensland.

Type: B.M.

Genus Megalommum Szepligeti, 1900

annulatum Turner

Turner (1918^/ 97) d 9 descr.

Loc: S.W. Australia; Tasmania.

Type: B.M.

Australian Braconidae — PARROTT

209

Genus Merinotus Szepligeti, 1906
xanthocephalus Turner

Turner (1918^?.' 98) & 9 descr.

Loc; Queensland.

Type: B.M.

Note: This genus is very near, if not iden-
tical with, Sigalphogaster Cameron.

Genus Monocoila Roman, 1910
parva Roman

Roman (1915: 13) 9 descr.

Loc: Kimberley, N.W. Australia.

Type: N.H.M.S. [Not marked as type.]

Genus Platybracon Szepligeti, 1900
nigriceps Cameron
Cameron (1911: 338) 9 descr.

Loc: Gin Gin, Queensland.

Type: B.M.

Genus Stigmatobracon Turner, 1918
basipennis Turner
Turner (1918a: 92) 9 descr., key.

Loc: Queensland.

Type: B.M.
diversipennis Turner
Turner (1918^.' 92) 9 descr., key.

Loc: Victoria.

Type: B.M.
torresensis Turner

Turner (1918<^.' 93) cf descr., key.

Loc: Cape York, Queensland.

Type: B.M.
xanthostigma Turner

Turner (1918^.- 92) descr., key.

Loc: Queensland.

Type: B.M.

Genus Vipio Latrielle, 1905
gestroi Mantero
Mantero (1897: 119) 9 descr.

Dalla Torre (1898: 305) listed.

Szepligeti (1904: 14) listed.

Loc: Australia.

Type: N.K.

Subgenus Vipiellus Roman, 1915
mjobergi Roman

Roman (1915: 8) 9 descr.

Loc: Kimberley, N.W. Australia.

Type: N.H.M.S.

Subfamily SPATHIINAE

As far as is known at present, the species
of this family are external parasites on coleop-
terous larvae.

Genus Leptospathius Szepligeti, 1902
formosus Szepligeti

Szepligeti (1902: 49) cf 9 descr., genotype.
Szepligeti (1904: 54) listed.

Loc: Australia.

Type: N.H.M.

Genus Spathius Nees, 1818

apicalis Westwood
Westwood (1882: 43) 9 descr.

Wilkinson [Y festinans] (1931^/ 261) cf 9
descr.

Nixon (1943^.' 378) descr., key, Austr.
record.

Loc: Kuranda, Queensland; India.

Type: N.K.
iridescens (Schletterer)

Schletterer [Stenophasmus] (1889: 203) cf
descr.

Szepligeti (1904: 53) listed, nomenclature.
Wilkinson (I9?)lh: 526) noted.

Loc: Cape York, Queensland; Sydney,

New South Wales.

Types: N.H.M.V., S.C.G.M.
kurandaensis Nixon

Nixon (1943^/ 393) descr., key.

Loc: Kuranda, Queensland.

Type: B.M.
turner! Nixon

Nixon (1943^.' 370) descr.

Loc: Tambourine Mountains, S.E.

Queensland.

Type: B.M.
sp. ?

Froggatt (1927: 55-56) recorded and fig-
ured.

Loc: New South Wales.

Note: Collected from imported tea chests
infested by a hostrychid beetle.

210

PACIFIC SCIENCE, Vol. VII, April, 1953

Subfamily HECABOLINAE
Genus Heterospilus Haliday, 1862
asion Nixon

Nixon (1943^.' 260) d' 9 descr., fig.

Loc: Maryborough and Imbil, Queens-

land.

Host; X.ylopsocus gihhicollis (Bostrychid.).

Type; B.M.

Genus Monolexis Foerster, 1862
ads Nixon

Nixon (1943<^.' 261) d 9 descr., fig.

Loc; Atherton, Palm Is., Queensland.
Host; Lyctus hrunneus {Lyciidi).

Type; B.M.

Note; "It is possible that my new species
is a European insect which has been
introduced with Lyctus hrunneus for
I have examined a single specimen,
a female (Italy, bred from Bostrychid
in figs) which appears to be identi-
cal with atisd (Nixon, 1943^?.’ 262.)

Subfamily EXOTHECINAE
Genus Opiopterus Szepligeti, 1908
parvus Szepligeti

Szepligeti (1908; 318) descr., genotype.
Loc; S.W. Australia.

Type; N.H.M.

Genus Spin aria Brulle, 1846

aliciae Turner

Turner (1917; 244) descr.

Loc; Australia.

Type; B.M.

Genus Xenarcha Foerster, 1862
tricolor Szepligeti

Szepligeti (1908; 318) descr.

Loc; S.W. Australia.

Type; N.H.M.

Subfamily ROGADINAE

The majority of the species of this sub-
family are parasitic on lepidopterous larvae

Genus Rogas Nees, 1818
( = Rhogas Agassiz 1846)

politus Szepligeti

Szepligeti (I906; 617) descr.

Loc; Australia.

Type; N.H.M.
tricolor Haliday

Haliday (1836; 98) descr.

Szepligeti (1904; 87) listed.

Loc; Australia.

Type; N.K.

Subfamily DOi^TCT/NHE

Most species of this family are external
parasites of coleopterous larvae.

Genus Acanthodoryctes Turner, 1918
gilberti Turner

Turner (I9I8A' 56) 9 descr., key.

Loc; Mackay, Queensland.

Type; B.M.
morleyi (Froggatt)

Froggatt [Iphiaulax] (1916; 566) 9 descr.,
host.

Froggatt \lphiaulax\ (1923: 42) descr., host.
Turner (I9I8A' 56) noted, genotype.

Loc; East Coast, Australia.

Host; Trogodendron fasciculatum (Clerid.).
Type; B.M.

Genus Caenopachyella Szepligeti, I9O8
lutea Szepligeti

Szepligeti (I9O8; 399) d 9 descr., geno-
type.

Loc; New South Wales.

Type; N.H.M.

Genus Doryctes Haliday, 1836

laemosacci Nixon

Nixon (1943: 257) 9 descr., host'

Loc; Emu Valley, Queensland.

Host; Laemosaccus sp. (Curculionid.).
Type; B.M.
parvus Muesebeck

Muesebeck (1941; 150) d 9 descr.

Loc; Queensland, Australia. The type
locality is Mayaguez, Puerto Rico.
Type; U.S.N.M. No. 55678.

Host; Dinoderus minutus (Fabr.) (Type)
LycUis sp. (Queensland.)

Australian Braconidae — PARROTT

211

Note: This is rather a common species in
various tropical regions of the
world.

syagrii (Fullaway)

Fullaway [Ischwgoms] (1922: 201) & 9

descr., host.

Loc: New South Wales.

Host: Syagrms fulvitarsis Pasc. (fern weev-
il).

Type: N.K.

Note: Fullaway (1921: 101, 114) records
a species of Doryctes parasitic on the
fern weevil ( Syagrius fulvitarsis) from
New South Wales. This probably is
the same species as recorded above.

Genus Liodoryctes Szepligeti, 1906
atriceps Turner

Turner (1918A' 54) 9 descr., key.

Loc: Mackay, Queensland.

Type: B.M.

australiensis (Szepligeti)

Szepligeti [Acanthohracon] (1902: 48)
descr.

Szepligeti [Neotrimorus] (1904: 64) listed.
Szepligeti (1906: 599) noted, genotype.
Turner (19 18 A 55) noted.

Loc: Cooktown, Kuranda, Mackay,

Queensland.

Type: B.M.
erythrothorax Turner
Turner (1918A' 54) cf 9 descr., key.

Loc: Mackay, Queensland.

Type: B.M.
nigridorsalis Turner
Turner (1918A' 53) 9 descr., key.

Loc: Port Darwin, Australia.

Type: B.M.

Genus Syngaster Brulle, 1846
annulicornis Brulle
Brulle (1846: 454) descr.

Froggatt (1890: 714) listed.

Szepligeti (1904: 67) listed.

Turner (191 8 A 57) noted.

Loc: Australia.

Type: N.K.

Note: Turner (1918A' 57) had not seen
this species, and is doubtful of its

true generic position,
lepidus Brulle

Brulle (1846: 459) descr.

Froggatt (1890: 714) listed.

Szepligeti [S. lepida\ (1904: 67) listed [error
in spelling.]

Cameron (1912: 195).

Turner (1918A 57) noted.

Loc: S.E. Australia; Tasmania.

Type. N.K.

introduced species

I am indebted to Mr. Frank Wilson, Prin-
cipal Research Officer, C.S.I.R.O., Entomolo-
gy Division, for the following list of braconids
introduced into Australia in connection with
biological control. No information is available
on the extent to which they have become
established in the various localities in which
they were liberated.

'^Apanteles ruhecula Marshall.

'^Apanteles glomeratus Linn.

Ascogaster carpocapsae Vier. {A. quadridenta-
tus Wesm.)

Bassus diversus (Mues.) {Agathis diversus)
Chelonus phthorimaeae Gahan
^ Macrocentrus ancylivorus Rohwer
Microhracon gelechiae Ashmead {Bracon ge-
lechiae)

Triaspis thoracicus (Say) (T. sayi Mues. and
Walkley.)

The above species are listed under the names
by which they are generally referred to in the
literature, the correct nomenclature is given
in parentheses where this is different.

Species that have become definitely estab-
lished are marked with an asterisk and in-
cluded in the body of the catalogue.

REFERENCES

Ashmead, W. H. 1888. Descriptions of new
Braconidae in the collections of the U. S.
National Museum. U. S. Natl. Mas., Proc.
2: 611-671.

1900. Notes on some New Zealand

and Australian parasitic Hymenoptera, with
descriptions of new genera and new species.
Linn. Soc. N. S. Wales, Proc. 25: 327-360.

212

PACIFIC SCIENCE, Vol. VII, April, 1953

Baker, C. F. 1926. Braconidae-Cheloninae of
the Philippines, Malaya and Australia. I.
Chelonini (except Chelonus). Philippine
Jour. Sci. 31: 451-499.

Baker, G. C. 1906. Fruitfly parasites. West.
Austral., Dept. Agr., Jour. 16: 27-28.

Bingham, C. T. 1906. New species of Braco-
nidae and Chalcididae from North Queens-
land bred by F. P. Dodd. Ent. Soc. London,
Trans. 1906 (1): 125-130.

Brues, C. T. 1920. The braconid genus Tra-
chypterus Guerin. Psyche 27: 59-62.

Brulle, G. a. 1846. Histoire naturelle des in-
sectes. Hymen. Tom. IV. 680 pp. Atlas, col.
pis. Roret, Paris.

Cameron, P. 1891. Hymenopterological no-
tices. Manchester Phil. Soc., Mem. 4 (4):
182-194.

1901^. Descriptions of three new gen-
era and species of Flymenoptera from East-
ern Asia and Australia. Ann. and Mag. Nat.
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I901A On a Collection of Hymenop-

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1906. Descriptions of two new species

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1911. A collection of parasitic Hy-

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1912. On a collection of parasitic Hy-

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Australian Braconidae — PARROTT

213

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PACIFIC SCIENCE, Vol. VII, April, 1953

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1908^. Braconiden aus der Sammlung

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— — — 1918^. Australian Braconidae in Brit-
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91-114.

1

Australian Braconidae — PARROTT

215

1918^. Notes on Braconidae in Brit-
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1918c. Notes on Braconidae in Brit-
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221-230.

19 18^/. Notes on Braconidae in Brit-
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1927. New parasitic Hymenoptera in

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— • 1928^. New parasitic Hymenoptera

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1929^. New parasitic Hymenoptera

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1930^. A revision of Indo-Australian

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1930^. New species and host records

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1931^. Five new species of Spathius.

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19Mb. On the Indo-Australian and

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1932<^. A revision of the Ethiopian

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Ent. Soc. London, Trans. 80: 301-344.

1932^. Three new braconids. (Hymen.

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\9Ma. On some Microgasterinae.

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1934^. On some Apanteles. (Hymen.

Bracon.). Stylops 3: 145-156.

INDEX TO AUSTRALIAN BRACONIDAE

abdominalis Szepl, (Ascogaster) 203

Acanthobracon see Liodoryctes 211

Acanthodoryctes 210

Acoelius see Adelius 200

Adelius . 200

afer S2epl. (Iphiaulax) 207

Agathiella I98

AGATHIDINAE 198

Agathis (see also Agathiella) 199

albimanus Szepl. (Opius) 204

aliciae Turner (Spinaria) 210

Amicroplus see Macrocentrus 196

ancylivorus Rohw. (Macrocentrus) 196

annulatum Turner (Megalommum) 208

annulicornis Brulle (Syngaster)- 211

antennalis Szepl. (Ascogaster) 203

anthelae Wilkn. (Apanteles) 200

antipoda Ashm. (Orgiloneura) 198

antipoda Ashm. (Apanteles) see ruficrus 201

antipodalis Ashm. (Meteorus) 196

Apanteles 200

Aphidius see Diaeretus 195

Aphidiinae 195

apicalis Westwood (Spathius) 209

Aridelus 196

Areogaster .. 203

Ascogaster 203

asion Nixon (Heterospilus) 210

Aspidocolpus see Austrohelcon 197

assimilator Turner (Cardiochiles) 202

Atanycolus 205

atis Nixon (Monolexis) 210'

atriceps Turner (Liodoryetes) 211

australasicus Cam. (Bracon) 205

australiansis Kokj. (Austrohelcon) 197

australiensis Ashm. (Adelius) 200

australiensis Ashm. (Apanteles) 200

australiensis Szepl. (Ascogaster) 203

australiensis Szepl. (Bracon) 205

australiensis Szepl. (Campyloneurus) 206

australiensis Szepl. (Chelonus) 203

australiensis Ashm. (Hybogaster) 207

australiensis Szepl. (Liodoryctes) 211

australiensis Ashm. (Phanerotoma) 204

australiensis Nixon (Zele) 198

australis Szepl. (Iphiaulax) 207

Austrohelcon 197'

216

PACIFIC SCIENCE, VoL VII, April, 1953

basalis Binghm. (Microplitis)

basipennis Turner (Stigmatobracon)

Bassus see Agathis

bicolor Brulle (Agathis)

bicolor Brulle (Braunsia)

bicolor S2epl. (Meteorus)

bicolor Szepl. (Westwoodiella) ....

bifasciatus Szepl. (Iphiaulax)

bimaculatus Szepl, (Bracon)

bimarus Turner (Bracon)

bipartitus Szepl. (Iphiaulax)

biplicatus Roman (Campyloneurus)

biroi Szepl. (Apanteles)

Biroia

bisignatus Szepl. (Iphiaulax)

BLACINAE

Bracon

BRACONINAE

Braunsia

brevicaudis Szepl. (Opius)

brevicornis (Bracon) see hebetor. . .

Caenopachyella

Callibracon see Iphiaulax

calligaster Roman (Cyanopterus) . . .

Calohelcon

camerunensis Szepl. (Iphiaulax) ....

Campyloneurus

capitator Fabr. (Bracon)

Cardiochiles

CARDIOCHILINAE

carpocapsae Ashm. (Opius)

caudata Szepl. (Ascogaster)

CHELONINAE

Chelonus

cinctus Baker (Ascogaster)

clavatus Guerm. (Trachypterus) . . .

coccinellae Schrank (Perilitus)

commutator Turn. (Cremnops) . . . .

conjugator Turn. (Sigalphus)

coriaceus Szepl. (Chelonus)

Cotesia see Apanteles

crassicaudis Szepl. (Cyanopterus) . .

Cremnops

crenulatus Szepl. (Iphiaulax)

cristatus Szepl. (Iphaiulax)

Cyanopterus

deeralensis Full. (Opius)

defensor Fabr. (Bracon)

deliadis Bingham (Apanteles)

demolitor Wilkn. (Microplitis) . . . .

detectus Baker (Ascogaster)

Diachasma see Opius

Diaeretus

Diluta Turner (Disophrys)

dimidiata Brulle (Agathis)

Dinocampus see Perilitus

Diospilini

Diospilus

Disophrys

dissimilis Turner (Cremnops)

dissimulator Turner (Cardiochiles) .
distinctus Baker (Ascogaster)

diversipennis Turner (Stigmatobracon). . ._ 209

diversipennis Turner (Braunsia) 199

diversipes Turner (Disophrys) 200

Doryctes 210

DORYCTINAE 210

dumbletoni Mues. (Meteorus) 196

emeraldensis Fahr. (Ipobracon) 208

Ephedrus 196

erythocephalus Turner (Austrohelcon) 197

erythropus Cam. (Ascogaster) 203

erythrothorax Turner (Liodoryctes) 211

eucalypti Cam. (Bracon) 205

EUPHORINAE I96

euthyrrhini Cam. see konowi (Parahelcon) 197

exiles Turner (Aridelus) I96

exornata Turner (Disophrys) 200

EXOTHECINAE 210

festinans see apicalis 209

festinata Turner (Agathiella) 198

fijiensis Full. (Opius) 204

flaviceps Cam, (Ipobracon) 208

flavipes Cam. (Apanteles) 200

Fohgra see Macrocentrus 196

formosus Szepl. (Leptospathius) 209

fraterculus Turner (Ipobracon) 208

froggatti Cam. (Bracon) 205

froggatti Full. (Opius) 204

gestroi Mant. (Vipio) 209

gilberti Turner (Acanthodoryctes) 210

gilberti Turner (Ipobracon) 208

glomeratus Linn. (Apanteles) 201

guyanensis Cam. (Apanteles) 201

Gymnoscelus see Helcon 197

hartmeyeri Szepl. (Bracon). 205

hebetor Say. (Bracon) 205

HECABOLINAE 210

Helcon (also see Austrohelcon) 197

HELCONINAE 197

Helconini 197

Helorimorpha see Aridelus 196

Heterospilus 210

hospitator Fabr. (Bracon) 205

Hybogaster 207

indultor Erich. (Austrohelcon) 197

ingressor Turner (Ipobracon) 208

innotatus Turner (Cyanopterus) 206

inornatus Kokj. (Austrohelcon) 197

intensus Baker (Ascogaster) 203

Iphiaulax see also Campyloneurus 206

Iphiaulax see also Trichiohelcon 197

Iphiaulax see also Cyanopterus 206

Iphiaulax 207

Ipobracon 208

Ipobracon see also Iphiaulax 207

iridescens Schl. (Spathius) 209

Ischiogonus see Doryctes 210

Ischius see Orgilus 198

juglandis Ashm. see hebetor Say 205

konowi Kokj. (Parahelcon) 197

kraussi Full. (Opius) 204

kurandaensis Nixon (Spathius) 209

laemosacci Nixon (Doryctes) 210

laeviventris Baker (Ascogaster) 203

I

202

209

199

199

199

196

198

207

205

205

207

206

200

199

207

198

205

205

199

204

205

210

207

206

197

207

206

205

202

202

204

203

203

203

203

197

196

199

204

203

200

206

199

207

207

206

204

205

200

202

203

204

195

200

199

196

198

198

200

199

202

203

Australian Braconidae — PARROTT

latibalteata Cam. (Agathiella)

leaena Cam. (Platyagathis)

leeuwinensis Turner (Phanerotoma) . . .

lepidus Brulle (Syngaster)

Leptospathius

leucogaster Holmgr. (Orgilus)

levisulcatus Cam. (Bracon)

limbatus Brulle (Iphiaulax)

Liodoryctes

Lipolexis see Diaeretus

longicornis Szepl. (Iphiaulax)

longiseta Szepl. (Iphiaulax)

lutens Cam. (Meteorus)

lutea Szepl. (Caenopachyella)

Macrobracon

MACROCENTRINAE

Macrocentrus

maculaticeps Baker (Ascogaster)

magnifica Wilkn. (Microgaster)

maligna Turn. (Agathiella)

marginipennis Turn. (Cremnops)

martialis Turn. (Agathis)

Megalommum

megaspilus Cam. (Chelonus)

meridionalis Turner (Agathiella)

meridionalis Turner (Austrohelcon) . . .

Merinotus

METEORINAE see Euphorinae

Meteorus

micans Szepl. (Iphiaulax)

Microbracon see Bracon

Microdus see Agathis

Microgaster

Microgaster see also Apanteles

MICROGASTERINAE

Microplitis

Minima Turner (Agathiella)

mjobergi Roman (Vipiellus)

modestus Baker (Ascogaster)

Monocoila

Monolexis

morata Wilkn. (Microgaster)

morleyi Frogg. (Acanthodoryctes) ....

mutator Fabr. (Campyloneurus)

Myosoma see Campyloneurus

Neotrimorus see Liodoryctes

narangae Vier. see ruficrus

nigriceps Cam. (Platybracon)

nigridorsalis Turner (Liodoryctes)

nigropectus Turner (Disophrys)

nobilis Turner (Macrobracon)

nonagriae OllifF. see flavipes Cam

obscuripennis Turn. (Calohelcon)

Opiinae

Opiopterus

Opius

Orgiloneura

Orgilus

pallidicolor Turn. (Ipobracon)

palpalis Szepl. (Ascogaster)

Parahelcon

parva Roman (Monocoila)

parvus Mues. (Doryctes)

217

parvus Szepl. (Opiopterus) 210

pedunculatus Szepl. (Microdus) 199

perelegans Bingh. (Microplitis) 202

Perilitus I96

perkinsi Full. (Opius) 204

perniciosa Wilkn. (Microgaster) 202

persicae Frogg. (Ephedrus) 196

perspicax Szepl. (Iphiaulax) 207

persulcatus Silv. (Opius) 204

petiolaris Szepl. (Microgaster) 202

Phanerotoma 204

philoeampus Cam. (Apanteles) 201

phoracanthae Frogg. (Trichiohelcon) 197

pilitarsis Cam. (Bracon) 206

Platyagathis 200

Platybracon 209

Poecilobracon see Ipobracon 208

politus Szepl. (Rogas) 210

praeclarus Turner (Campyloneurus) 206

praepotens Turner (Campyloneurus) 206

pretiosus Szepl. (Iphiaulax) 207

proficisator Fabr. (Cyanopterus) 207

profugus Turner (Campyloneurus) 206

Protapanteles see Apanteles 200

Proteins see Meteorus 196

pulchellus Brulle (Bracon) 206

pulchicaudis Szepl. (Iphiaulax) 207

punctistigma Roman (Ipobracon) 208

qua'dricolor Turner (Ipobracon) 208

radiantis Wilkn. (Apanteles) 201

rapae Curtis (Diaeretus) 195

reficeps Szepl. (Iphiaulax) 208

resplendens Wilkn. (Microgaster) 202

RHOGADINAE see ROGADINAE 210

Rhogas see Rogas 210

rixosa Wilk. (Microgaster) 202

ROGADINAE 210

Rogas 210

rubecula Marsh. (Apanteles) 201, 211

ruberrima Turner (Disophrys) 200

rubriceps Frogg. (Iphiaulax) 207

rubromaculatus Cameron (Macrocentrus) 197

rufator Romans (Cardiochiles) 202

ruficeps Szepl. (Agathiella) 198

ruficeps Szepl. (Iphiaulax) 208

ruficeps Szepl. (Diospilus) 198

ruficeps Brulle (Westwoodiella) 198

ruficrus Hal. (Apanteles) 201

rufifrons Turner (Disophrys) 200

rufithorax Turner (Helcon) 197

rufithorax Turner (Agathis) 199

rufiventris Bingh. (Apanteles) \ . 201

rufobrunneus Turner (Agathis) 199

rufoniges Turner (Helcon) 197

rufus Szepl. (Cyanopterus) 207

rugosa Turner (Agathiella) 198

scoparius Szepl. (Iphiaulax) 208

semirufus Szepl. (Iphiaulax) 208

sicarius (Apanteles) 201

Sigalphus 204

signatipennis Turner (Disophrys) 200

similipicta Turner (Disophrys) 200

similis Szepl. (Chelonus) 203

198

200

204

211

209

198

205

207

211

195

207

207

196

210

208

196

196

203

201

198

199

199

208

203

198

197

209

196

196

207

205

199

201

200

200

202

198

209

203

209

210

201

210

206

206

211

201

209

211

200

208

200

197

204

210

204

198

198

208

203

197

209

210

218

solitaria Turner (Biroia)

sons Wilkn. (Microgaster) .

SPATHIINAE

Spathius

speciosissimus Szepl. (Iphiaulax)

Sphaeropyx see Sigalphus

Spinaria

Stenophasmus see Spathius

Stigmatobracon

syagrii Full. (Doryctes)

sydneyensis Cam. see ruficrus Haliday .

Syngaster

talpocharis see thalpocharis

tasmanica Cam. (Apanteles)

tasmanicus Wilkn. (Macrocentrus) ....

tearae Wilkn. (Microgaster)

tegularis Szepl. (Ascogaster)

tegularis Szepl. (Microgaster) ........

tenuissima Turner (Agathiella)

terminatus see coccinellae

talpocaris see thalpocharis

thalpocharis Ashm. (Bracon)

tomentosus Szepl. (Atanycolus)

torresensis Turner (Stigmatobracon). . .

torridus Turner (Ipobracon)

Trachypterus

transiens Turner (Iphiaulax)

Trichiohelcon

tricolor Szepl. (Agathiella)

tricolor Szepl. (Ascogaster)

tricolor Ashm. (Bracon)

tricolor Hal. (Rogas)

tricolor Szepl. (Xenarcha)

trifasciatus Szepl. (Iphiaulax)

trinotata Ashm. (Iphiaulax)

tripartitus Szepl. (Sigalphus)

turner! Cam. (Bracon)

turner! Nixon (Spathius)

tryoni Cam. (Opius)

uniformis Turner (Cardiochiles)

unimaculata Turner (Agathiella)

unimaculatus Szepl. (Chelonus)

verticalis Turner (Cardiochiles)

Vipiellus (Sub-genus)

Vipio

VIPIONINAE see BRACONINAE

vividus Baker (Ascogaster)

vulpina Wilkn. (Microgaster)

walker! Turner (Bracon)

Westwoodia see Westwoodiella

Westwoodiella

xanthocephalus Turner (Merinotus) . . . .

xanthopsis Turner (Agathis)

xanthostigma Szepl. (Cremnops)

xanthostigma Turner (Stigmatobracon)

Xenarcha

Zele

Zelini

PACIFIC SCIENCE, Vol. VII, April, 1953

HOST INDEX OF AUSTRALIAN BRACONIDAE

Coleoptera

Coccinella California Mann. (Coccinellid.) 196

Coleomegilla maculata (Deg.) (Coccinellid.) 196

Cycloneda trifasciata juliana Muls.

(Coccinellid.) 196

Cycloneda sanguinea (Linn.) (Coccinellid.). I96

Dinoderus minutus (Fabr.) (Bostrychid) 210

Euthyrhinus meditabundas Fabr. (Curculionid.) . . 197
Hippodamia convergens Guer. (Coccinellid.) .... I96

Hippodamia 5-signata (Kby.) (Coccinellid.) I96

Laemosaceus sp. (Curculionid.) 210

Lyctus brunneus (Lyctid.) 210

Syagrius fulvitarsis Pasc. (Curculionid.) 211

Trogodendron fasiculatum Schreibers

(Clerid.) 197, 208, 210

Xylopsocus gibbicollis (Bostrychid.) 210

Diptera

Ceratitis capitata Wied (Trypetid.) 205

Dacus sp. (Trypetid.) 204

Dacus ferrugineus Fabr. (Trypetid.) 205

Dacus laticaudus Hardy. (Trypetid.) 204, 205

Dacus tryoni Cam. (Trypetid.) 204

Homoptera

Brevicoryne brassicae (Aphid.) 195

Mysus persicae (Sulz.) (Aphid.). 195

Rhopalosiphum pseudobrassicae (Davis)

(Aphid.) 195

Lepidoptera

Agrotis sp. (Noctuid.) 201

Anthela ocellata Walk. (Lymantriid.) 200

Antheraea eucalypti Scott (Saturnid.) 200

Delias argenthona Fabr. (Pierid.) 200

Ephestia eluttella Hueb. (Phycitid.) 205

Ephestia kuehniella Zell. (Phycitid.) 205

Galleria mellonella Linn. (Galleriid.) 205

Grapholitha molesta Busck. (Olethreutid.) 196

Heliothis armigera Hubn. (Noctuid.) 201, 202

Leucania unipuncta Haworth (Noctuid.) 201

Ogyris genoveve How. (Lycaenid.) 201

Pheressaces cycnoptera Lowes (Noctuid.) 202

Phagmatiphila truncata Walk. (Noctuid.) 201

Pieris brassicae Linn. (Pierid.) 201

Pieris rapae Linn. (Pierid.) 201

Plodia interpunctella Hubb. (Phycitid.) 205

Plusia sp 201

Teara tristis (Lewin) (Notodontid.) 202

Thalpocharis coccophaga (Noctuid.) 206

Theretra oldenlandiae fumata Walk. (Sphingid.) . 202

Thyridopteryx herrichi West. (Psychid.) 202

Tortrix postvitt'ana Walk. (Tortricid.) 198, 201

Utsthesia pulchella Linn. (Atctiid.) 201

199

202

209

209

208

204

210

209

209

211

201

211

206

201

197

202

203

202

198

196

206

206

205

209

208

197

208

197

199

203

206

210

210

208

208

204

206

209

204

202

199

204

202

209

209

205

203

202

206

198

198

209

199

200

209

210

198

198

Studies on the Bathypelagic Fishes of the Family Paralepididae.
1. Survey of the Genera

Robert R. Harryi

INTRODUCTION

The Paralepididae are large-eyed, elongate
fishes lacking light organs and are members
of the suborder Alepisauroidea (Alepisaurina,
using the recently proposed endings of Sten-
zel, 1950: 94) of the order Iniomi (also called
Scopeliformes by Berg, 1940: 256; Iniomida
by Schultz, 1948: 233; Iniomida- or Mycto-
phida, using Stenzel’s endings). These abun-
dant pelagic and deep-sea teleosts of world-
wide distribution have been termed barracu-
dinas by Fowler (1936). The family appears
to be most closely related to the Anotopteri-
dae and Alepisauridae.

While I was endeavoring to determine the
relationships of a new species of the genus
Lestidium from off the coast of central Califor-
nia, it became evident that the classification
of the Pacific paralepidids was in deplorable
confusion, especially in regard to generic
grouping. Comparison of the form and de-
velopment of various parts of the body in the
different genera has shown, unexpectedly,
that certain structutes not heretofore used are
of phylogenetic or taxonomic importance.
Among the most useful are the structure of
the maxillaries, the form and distribution of
the dentition and gillrakers, modifications of
squamation, and variation of the lateral-line,
tube. These new observations form the basis
for this revision of the Paralepididae.

In addition, these fish form an important
food of tunas and other large oceanic fishes.
A more complete knowledge of them, partic-
ularly in regard to speciation and limits of

1 Academy of Natural Sciences of Philadelphia.
Manuscript received. August 23, 1950.

distribution, would furnish a valuable tool
for obtaining information on the migration
of these larger food fishes by means of stom-
ach analyses.

In the course of the present study I have
been able to examine material totaling several
hundred specimens, , representing all the
known genera {Sudis, Notolepis, Paralepis, Les-
tidium, Macroparalepis, Arctozenus, Bathysudis,
and Lestidiops), and all the generic types. One
genus and one species are described as new.
The revision is divided into parts comprising
(1) a survey of the genera, (2) a review of the
North Pacific species, and (3) a synopsis of
the South Pacific species. The illustrations
are by the author.

Acknowledgments: Many people have been
extraordinarily helpful by giving, loaning, ex-
changing, or examining specimens, and by
offering helpful criticism during the prepara-
tion of this paper. Of these I wish to thank
particularly Dr. Vilh. Ege of the Marinbio-
logisk Laboratorium, Charlottenlund Slot,
Denmark, Dr. George S. Myers of Stanford
University, Dr. Rolf L. Bolin of the Hopkins
Marine Station, Dr. Carl L. Hubbs of the
Scripps Institution of Oceanography, Mr.
John E. Fitch of the California Division of
Fish and Game, Dr. E. H. Ahlstrom of the
United States Fish and Wildlife Service, Dr.
Ethelwyn Trewavas and Mr. N. B. Marshall
of the British Museum, Mr. G. E. Maul of the
Museum Municipal do Funchal, Dr. Arthur
D. Welander and Dr. A. W. Herre of the
University of Washington, Dr. Earl S. Herald
of the California Academy of Sciences, Dr.
Leonard P. Schultz of the United States Na-
tional Museum, Dr. William Beebe of the

219

220

PACIFIC SCIENCE, Vol. VII, April, 1953

New York Zoological Society, Dr. Kiyoma-
tsu Matsubara of Kyoto University, Mr. John
W. Reintjes of the Pacific Oceanic Fishery
Investigations, Dr. Daniel Merriman of the
Bingham Oceanographic Institute, Dr. Enrico
Tortonese of the Universita di Torino, and
Mr. George Vanderbilt of Honolulu. The of-
ficers of the California Academy of Sciences,
Stanford University, New York Zoological
Society, Scripps Institution of Oceanography,
United States National Museum, Academy of
Natural Sciences of Philadelphia, and Pacific
Oceanic Fishery Investigations Laboratory,
Honolulu, have allowed me to examine all
their collections of the family Paralepididae.

The work was done in the Natural History
Museum of Stanford University and in the
Research Laboratory of the George Vander-
bilt Collections of the California Academy of
Sciences.

HISTORY OF GENERIC CLASSIFICATION

The family Paralepididae has been various-
ly expanded and restricted. It has, in the past,
encompassed a wide assortment of remark-
ably divergent groups ‘(r/V.? Regan 1911, Parr
1928, Fowler 1944), but it is generally accept-
ed by recent authors that the family includes
only the genera Paralepis, Lestidium, Macro-
paralepis, and Sudis. The genus Luciosudis
Fraser-Brunner belongs in the suborder Myc-
tophoidea, probably in the family Chloroph-
thalmidae. The genus Notosiidis Waite may
belong in the suborder Alepisauroidea, but,
if it does, it definitely does not belong in the
Paralepididae. It is perhaps near the family
Anotopteridae. The genus Neosudis Castelnau
(1873) has not been rediscovered since the
original account, and its systematic position
is not known. Many characters are attributed
to this genus which do not agree with the
suborder Alepisauroidea as presently under-
stood, and it is not included in the present
study.

The recent workers who have contributed
most toward a better understanding of the
classification of the Paralepididae are Regan,
Parr, Ege, and Maul.

Regan’s arrangement of the order Iniomi
(1911) has been generally accepted as the
first reasonably natural system proposed for
these forms. He relegated Paralepididae (using
the name Sudidae, which he later sometimes
abandoned in favor of Paralepididae) to the
suborder Myctophoidea together with Syno-
dontidae, Aulopidae, and Myctophidae. Un-
fortunately, he placed Sudidae in the wrong
suborder because he based his family descrip-
tion on Chlorophthalmus instead of on Sudis,
Paralepis, or a genus closely related to them.
While Chlorophthalmus is obviously close to
the three families Regan placed in the sub-
order Myctophoidea, its relationships to the
paralepidids are distant. This unfortunate link-
age of Chloropththalmus with the Paralepididae
has contributed to the subsequent confusion
of the subordinal relationships of the Myc-
tophoidea and Alepisauroidea — confusion
which has extended to the present. Actually
it is clear that Sudis and its allies are more
closely related to the Alepisauroidea than to
the Myctophoidea and, therefore, should have
been placed in the former suborder. Further-
more, Regan used Sudidae as a catch-all for
forms that did not fit into his system, and he
divided it into three groups containing nine
genera which previous authors had appor-
tioned to several families. Recent authors re-
gard his third section, exclusive of Parasudis,
as the only one of his three divisions that
really belongs in the Paralepididae.

The most important extensive studies of the
Iniomi have been those of Parr. In his survey
of the Paralepididae (1928), he accepted and
named Regan’s three groups as subfamilies
and added the problematical Notosudis, which
was described subsequent to Regan’s paper,
as a fourth subfamily.

Parr’s discussion of his "Paralepidini” in-
cludes a review of all genera and species
(mostly prepared from the literature, as he
had little material) but his entire classification
was greatly revised, immediately after it ap-
peared, in a series of papers by Ege and him-
self. Parr (1928) recognized only two genera

Genera of Paralepididae — Harry

221

as valid, on the basis of the presence of scales
in one genus and their absence in the other.
Parr and Ege attempted by subsequent in-
vestigation to demonstrate that this type of
division was not justified. The two reasons
why Parr’s classification of 1928 was not ac-
cepted by subsequent authors (including him-
self) are: (1) he did not recognize that Sudis
hyalina was so distinct from all other para-
lepidids that it deserved monotypic generic
status; (2) he based his generic characters for
Sudis and Lestidium solely on the presence or
absence of scales. It was not known at that
time that the species of the family could be
arranged in a uniformly graded series from
(a) those completely scaled on head and body,
through (b) those scaled only on body or
head, through (c) those scaled only late in
development, to (d) those lacking scales at
all stages of development. Furthermore, he
used the generic name Sudis for the scaled
species, but the generic type {hyalina) is al-
most completely naked at all stages. In addi-
tion, each of his genera included both naked
and scaled species.

Parr (1929: 4, 7) in a paper on the osteology
and classification of the Iniomi, pursued fur-
ther the relationships of the families. His in-
vestigations of the Paralepididae are based on
the partial osteology of two representatives of
''Lestidium' {Paralepis hrevirostris and a spe-
cies of Lestidium). It represents the first re-
search of this kind on the family. Parr pro-
posed a phylogenetic line on the basis of
osteological characters, from Chlorophthalmus
through Bathysudis to Omosudis, "thereby in-
directly establishing a relationship between
the Sudidae and Alepisauridae."

The generic relationships of the Paralepi-
didae adopted by Parr (1928, 1929) and others
were subsequently greatly modified on the
basis of an extensive distributional and sys-
tematic research on the genus "Paralepis" of
the North Atlantic by Ege (1930). His rich
material was composed almost exclusively of
larval to juvenile specimens. The identifica-
tions he made of these early stages with pre-

viously known adult forms are particularly
valuable for the elucidation of the taxonomic
and nomenclatural problems which had de-
veloped in the earlier literature. His generic
classification is essentially the same as that
used by most previous European authors. He
retained the genus Sudis as monotypic and
expanded the limits of Paralepis. Although
the relationship of the genus Lestidium to
Paralepis was considered by Ege to be a matter
beyond the scope of his investigation, he
maintained that the recognition of Lestidium
on the premise of its being devoid of scales
was completely untenable. The Mediterranean
and North Atlantic material available to him
was represented by young material complete-
ly devoid of scales; but he did not consider
the lack of scales in the younger individuals
necessarily to indicate their absence in the
fully adult stage. For example, he found an
individual oi" Paralepis rissoi" (which is scaled
when adult) 188 millimeters in length which
completely lacked any trace of scales.

Unfortunately, Ege did not investigate the
generic classification of the Paralepididae as
carefully as he did the specific. He never de-
fined the limits of the genus "Paralepis" as
he accepted it, but apparently used it in the
broad sense of early European authors. Ac-
tually he did not realize the bearing of Lesti-
dium to his work. He claimed that the genus
Lestidium was beyond the scope of his in-
vestigation, but included all species of this
genus of which he had material in "Paralepis"
and excluded almost all the others which he
had not seen.

While he claimed that Lestidium is unten-
able on the basis of the lack of scales, Ege
proved on morphometric grounds that the
naked species can be recognized {vide the
differentiation of groups I and II in his key).
Apparently he never realized that his first
division included all the scaly forms, and the
second division all the forms naked even as
adults. The mention of such discrepancies is
not meant to detract from the value of his
work. He was not concerned particularly with

222

PACIFIC SCIENCE, Vol. VII, April, 1953

the major classification of the family, but was
primarily interested in development and spe-
ciation. In fact, he has made the Paralepididae
the best-known group of the order Iniomi in
this respect.

Ege’s results were interpreted by Parr
(1931^), who assumed that Lestidium was un-
tenable and that the limits of the genus Para-
lepis had to be considerably extended to en-
compass a highly variable group— at least as
long as practicable generic distinctions had
not become re-established on a different mor-
phological basis. Thus, if Parr’s classification
of 1928 were altered to fit this idea, all the
species of "Paralepidini,” tyiQQpt Sudis hyalina,
would be placed in the genus Paralepis.

Ege (1933) preliminarily diagnosed 15 new
species and a new genus, Macroparalepis, from
a part of the paralepidid material collected—
usually at great depths— by the 1928-30 "Da-
na” Expedition around the world. Unfortu-
nately, the specimens are all postlarvae or
juveniles, the descriptions are brief, and the
relationships are not indicated for the species
of Macroparalepis. Lestidium and Paralepis were
compared with Macroparalepis. Dr. Ege is still
studying this material and plans to publish
more complete accounts.

Gregory and Conrad (1936; 28) also at-
tempted to give a natural phylogeny of the
order Iniomi. Their classification, which ap-
pears to have been done primarily by a com-
parison of illustrations of the species and
superficial examination of the literature, adds
little to our knowledge of the family Para-
lepididae. They apparently misinterpreted
some of Parr’s findings of 1929 and did not
include the papers of Ege (1930, 1933) and
Parr (1931).

Jensen (1942), in his critical review of the
paralepidids of Greenland, gives important
discussions of the genus Paralepis. One of the
most important parts of his results is a de-
tailed account of the structure of the lateral-
line and scales. It is important to note that
Jensen confuses the ossified sections of the

lateral-line tube and the overlying lateral-line
scales.

In a monograph of the paralepidid fishes
in the collection of the Municipal Museum
of Funchal, Madeira, Maul (1945) included
material of Sudis, Paralepis, and Macropara-
lepis and confined the limits of the family to
these genera. He presented definitions of the
genera for the first time since Parr (1928).
Maul described more adult material than any
other recent worker and described larger spe-
cimens of Macroparalepis for the first time.

These classifications of Parr, Ege, and Maul
differ essentially from my results, presented
in the next section, in that I place Sudis in a
distinct subfamily (Sudinae) and divide the
Paralepidinae into two basic groups of genera
that can be easily distinguished by osteolo-
gical characters, by gillraker structure, by ex-
tent of squamation in the adults, and by
morphometry. These two basic divisions of
the Paralepidinae correspond to groups I and
II of Ege’s key (1930: 7). An outline of this
classification is presented with the description
of the paralepidids collected by the Bermuda
Oceanographic Expeditions of the New York
Zoological Society (Harry, 1951).

DISCUSSION

Family PARALEPIDIDAE

Paralepidini Bonaparte, 1832-41 (no pagina-
tion), 1846; Bellotti, 1878; Parr, 1928: 17,
32; Parr, 1929: 28.

Paralepidina Gunther, 1864: 418.
Paralepididae Gill, 1874: 16; Gill, 1893: 131;

Goode and Bean, 1895: 118.

Paralepidinae Jordan and Evermann, 1896:
599.

Paralepidae Dollo, I9O8: 59; Jordan, 1923:
154.

Sudidae (in part) Regan, 1911: 125; Parr, 1928:
15; Parr, 1929: 27; Ege, 1933: 223; Fowler,
1944: 438.

Sudini Berg, 1940: 257, 437.

Sudidae 1942: 3; Maul, 1945; 3.

DIAGNOSIS: Body compressed, elongate.
Belly usually constricted, forming a pro-

Genera of Paralepididae — Harry

223

Fig, 1. Front of snout of a paralepidid (illustrated
from the holotype of Lestidium prolixum, n. sp.). The
teeth in the emargination of the uppet jaw are tohse of
the palatines. The premaxillary lacks teeth in this emar-
gination.

nounced mid-ventral carina. Anus moves pos-
teriorly during early ontogeny; finally situ-
ated in region of pelvic fins. Eyes moderately
large, with adipose lids. Mouth with exten-
sive buccal valves. Rictus well before eye;
tip of maxillary not reaching past a vertical
from anterior border of eye except in early
postlarvae. Symphysis of lower jaw more or
less elevated. Corresponding arched, tooth-
less emargination of upper jaw below end of
snout (see Fig. 1). Lower jaw included when
mouth is closed. Mandibles with 1 to 3 rows
of teeth (sometimes atrophied in adults of
scaled genera) ; if present, inner series is com-
posed of depressible canines which recline

backward, the outermost series of short fixed
teeth of similar shape. Premaxillary closely
bound to maxillary and not protractile. Pre-
maxillaries anteriorly with a single row of
depressible canines, situated on the inner face
of this bone, followed by single series of short
fixed teeth on edge of bone. Dorsal process
of premaxillary with wide circular foramen in
all genera except Sudis. Supramaxillary slender,
approximately 0.5 length of maxillary. Vomer
toothless, or with 1 or 2 tiny teeth. Palatine
long and slender, anteriorly attached to sides
of mesethmoid and vomer. Teeth on palatines
in 1 or 2 rows. Teeth on tongue absent or
minute.

Fig. 2. Lateral-line segments of a paralepidid (illus-
trated from the holotype of Lestidium thunnorum, n.
sp.). a. Top of an ossified segment: 3, posterior view
of an ossified segment with adjoining pores stippled;
c, bottom of an ossified segment; d, a series of ossified
segments in normal position along the lateral-line. The
lateral-line scales of the genera Megalosudis, Paralepsis,
and Notolepis overlie these ossified segments.

Scales when present cycloid, generally ca-
ducous. Lateral-line tube large, imbedded in
the skin, enclosed in characteristic partly os-
sified segments (see Fig. 2), which perhaps
are highly modified scales. In the naked gene-
ra, the outer shield-like section of each partly
ossified segment forms an imbedded scale-
like structure. In scaled genera, these struc-
tures overlain by lateral-line scales. Tube va-
riously pierced by pores medially, above, and
below.

Dorsal fin small, with 9-15 rays, situated
near middle of body length. Adipose dorsal
fin present over posterior end of anal fin.
Anal fin far posterior, with long base and
numerous rays (21-50). Caudal fin small, very
short and forked, free from adipose and anal

224

PACIFIC SCIENCE, VoL VII, April, 1953

fins, with 18-20 principal rays. Pectoral hns
moderately low, below midaxis of body, with
9-17 rays. Pelvic fins small, close together,
behind middle of standard length, with 8-12
rays; their position varies from far before dor-
sal hn to behind dorsal base. Vertebrae 65-
102, as far as known. No light organs present.

Secondary sexual dimorphism not known.
Specimens have been taken up to approxi-
mately 20 inches in length.

Sudis Rafinesque (1810) was the first genus
of the barracudinas to be described, but it
was indicated as being related to Sphyraena
and placed in ’’Esocidi.” Cuvier described
Paralepis in 1817 for two species mentioned
by Risso (1810) and also related it to Sphy-
raena. Apparently Bonaparte (1832-46) first
elevated one of the genera to familial status
when he placed ”Paralepidini” as a subfamily
of Sphyraenidae (1832-41) and of Scopelidae
(1846), using, ipso facto, Paralepis as the family
type. He recognized both Sudis and Paralepis
in this group. As far as I can find, all authors
up to the time of Regan (1911) used Para-
lepis as the type genus, with Gill (1874) the
first author to taise the group to the status of
a full family (Paralepididae). Dollo (1908)
and many subsequent authors shortened the
name to Paralepidae, but this is not proper
form. Regan (1911) based the family name on
Siidis because it is the oldest generic name.
According to the Rules of the International
Commission of Zoological Nomenclature as
clarified by Opinions 133 and 141, however,
this is not acceptable, because Paralepididae
was proposed before Sudidae. Unless I have
missed a reference earlier than Bonaparte, in
which a family name was based on Sadis,
Paralepididae is the proper name. It seems
unlikely that there is an earlier reference, since
Bonaparte was apparently the first ichthyolo-
gist to use family endings.

Relationships within the Family

Most authors, except Ege, have examined
only a few of the genera and usually only very
immature material. As a result, few workers
have attempted to analyze the major parale-

pidid relationships. Most current authors have
accepted Ege’s classification of the family
which is based primarily on differences of
postlarval development and recognizes only
a few generic units. My investigations have
primarily stressed the morphological aspects
of as mature material as possible although I
have utilized several complete ontogenetic
series. As there is some important variation
between stages of development, it is believed
that a clearer picture of paralepidid classifica-
tion can be obtained by utilizing more than
one growth stage.

Fig. 3. Top of head of a paralepidid (illustrated from
a paratype of Lestidium Thunnorum n. sp.), showing the
cranial ridges and the foramina in the premaxillaries.

Although our present knowledge of these
fishes is so inadequate as to make very exten-
sive splitting seem premature, certain aggre-
gates of species appear to be so widely sepa-
rated from their nearest relatives on the evi-
dence of prominent characters in stages older
than postlarvae that their inclusion in Para-
lepis seems much too conservative. The reten-
tion of only three genera does not allow us to
depict several important and easily recognized
lines of evolution. In my own investigations,
conclusions have been reached which involve
a splitting of the genus Paralepis, a resurrec-
tion of the previously suppressed genera Les-

MAGNISUDIS
PARALEPIS
PROFUNDISUDIS
NOTOLEPIS
LESTIDIOPS
LESTIDIUM
LESTROLEPIS
MACROPARALEPIS
STEMONOSUDIS

SUDIS

Fig. 4. Diagram of the extent of relationships be-
tween the genera of the family Paralepididae.

Genera of Paralepididae — HARRY

225

Fig. 5. Genotype of Magnisudis {M. harysoma, n. sp., paratype, 510 mm. in standard length, from off Lower
California).

tidium and Notolepis, and the drawing of rela-
tionships along lines somewhat different than
those given by Ege.

There is no doubt that Sudis is unique and
remote from the other barracudinas, and there
may be some question whether this genus
even belongs in the same family with them.
Siidis is distinguished from all other parale-
pidids primarily by lack of a foramen in the
premaxillary and by other osteological pecu-
liarities, by the unique mandibular dentition,
by the reduction of the anterior nostril to a
minute pore in adults, by the presence of
scales only on the preopercles, and by the
different larval development, particularly of
the pectoral fins (Ege, 1930: 6). Despite these
differences, it is believed that family relation-
ships within the suborder Alepisauroidea can
best be shown by retaining Sudis in the Para-
lepididae. The family would then be divided
into the subfamilies Paralepidinae and the
monotypic Sudinae. The Sudinae most closely
resemble the genus Lestidium in gillraker struc-
ture, lack of body squamation, lateral-line
form, and other characters.

The subfamily Paralepidinae falls into two
basic generic subdivisions, ^hich are sepa-
rated in part by the characters given in Ege’s
key (1930: 7) to separate "Groups I and IT’
of Paralepis. The most obvious differences
between the main groupings are that all spe-
cies in division I are scaled as adults and have
numerous teeth in two or more rows on each
gillraker and that those of division II are
scaleless as adults and have a few teeth in one
row on each gillraker. This classification cuts
across Ege’s system. His groups I and II of
Paralepis are apparently only remotely related.

representing groups of genera that might be
designated as tribes (e.g., Paralepidini and
Lestidiini) .

In determining the relationships within the
family, the following characters are probably
primitive and apparently trace back into the
stock now represented by the Chlorophthal-
midae and Aulopidae, the two most general-
ized families in the order Iniomi: short and
massive head; completely scaled body; lack
of nonossified prolongation on the lower jaw
or snout; large eye far forward; tiny fixed
teeth on the jaws, vomer, and palatines; lack
of or slight development of an anterior for-
amen in the premaxillary; hyoid apparatus
extending far forward, with gillrakers strongly
developed on all five arches; simple lateral-
line tube and unmodified lateral-line scales;
low number of dorsal and anal rays.

Such characteristics are approached to a re-
markable degree by a new species in a new
genus, Magnisudis harysoma. The conservative
Magnisudis has a short, broad, massive head
and snout, a moderately short, heavy body.
The eye is large. The teeth on the jaws are
minute; this species is the only paralepidid
with teeth on the vomer in the adult. The
supramaxillary is free at its posterior end. The
gillrakers are reduced to flat bases armed with
4 long filaments (it is of interest that this type
of gillraker is unique to Magnisudis within
the Paralepididae and is known in only one
other family of the Iniomi, the Alepisauridae) .
The gillrakers on the first arches extend far
forward on each side of the tongue into the
anterior part of the mouth. The tip of the
tongue is far forward, near the symphysis of
the lower jaw. The lateral-line structure is the

226

PACIFIC SCIENCE, Vol. VII, April, 1953

simplest in the family. A single row of large,
round, tenaciously adherent scales is present
over the lateral-line tube; each scale is pierced
by a median pore only. The scales over the
remainder of the body are very small, round,
and deciduous. The anal fin has comparative-
ly few rays.

Paralepis, which is restricted to a few species,
is undoubtedly a derivative of the Magnisudis
line of evolution. These fishes resemble Mag-
nisudis and differ from all the remaining para-
lepidids in the short, heavy head and body,
anteriorly free supramaxillary, long anterior
teeth on the gillrakers, and low number of
anal rays. The relationship is, however, not
really close, since Paralepis differs from Mag-
nisudis in several important characters. The
gillrakers are shorter and more spine-like than
in Magnisudis, and there are numerous teeth
on each raker. Only a few anterior teeth on
each gillraker are long, and all the posterior
teeth are very short and needle-like. The
tongue is long, farther back in the mouth than
in Magnisudis, and the gillrakers do not ex-
tend as far forward on each side of the tongue.
The lateral-line structure is more complex.
There are one and one-half or two rows of
caducous scales over the lateral-line tube. In-
stead of having but a single median pore, each
lateral-line segment is often pierced above and
below by another pore. In most species the
upper scale row over the lateral line is pene-
trated near the lower margin by the upper
pore, and the lower scale row receives the
middle and lower pores, although there is
extreme variability in pore structure and a
tendency for the reduction in number of pores
piercing the lateral-line scales. The body scales
are longer than broad, the same size as the
lateral-line scales, and extremely caducous.
There is little variation within the genus Para-
lepis; except in the case of the inadequately
known P. elongata Brauer, the species are very
closely related and somewhat difficult to tell
apart.

The most specialized of the scaled genera
is Notolepis, which approaches Lestidium more

closely than any other scaled form. The snout
is prolonged and pointed, and the body is
more elongate and less deep than in Paralepis.
The teeth in the adults are not lacking or
reduced as in the other scaled genera but are
fairly long and well developed. They are dis-
tinctly arranged in two or three rows, the
outer row of short, fixed teeth, and the inner
series of long, hastate, depressible canines.
The lateral line is essentially similar in struc-
ture to that of Paralepis but has more pores;
the upper scale row is sometimes peculiarly
modified, being notched to receive the large
upper pore instead of being pierced by this
pore series. The body scales are round, over-
lapped for most of their length, and are the
same size as the lateral-line scales. The teeth
on the gillrakers are reduced in length and
consist of numerous tiny teeth in oblique
rows on each raker. The pharyngeal teeth are
reduced in number. It is known from Ege’s
investigations that the scales tend to form
late in life in at least one of the species (N.
rissoi), and it is very probable that there is a
tendency for the suppression of scales in this
group. The three species of Notolepis are wide-
ly separated in distribution: N. rissoi from the
North Atlantic, N. coruscans from the North |
Pacific, and N. coatsi from the Antarctic. The
two northern species are much more conserva-
tive in form than is N. coatsi, which has an
extreme development in dentition, gillrakers,
and lateral line, and these two species are
separated in the new subgenus, Profundisudis.

The next evolutionary step appears to in-
volve a greater development of the elongate
form and dentition, an important reduction
in the number of gillraker teeth, loss of all
scales, further specialization of the partly os- jj
sified lateral-line segments into characteristic !j
scale-like structures, a wide variability in the ii

position of the pelvic fins in relation to the |j

dorsal fin, and a wide range in the number of I
anal rays. The most generalized genus of this
group is Lestidium which corresponds to group
II of Ege (1930: 7).

Lestidium can be distinguished from the

Genera of Paralepididae — HARRY

227

genera Magnisudis, Paralepis, and Notolepis by
the complete absence of scales, by the greater
length of the mandibular teeth, by the ar-
rangement of the teeth on each gillraker into
a single row of short needle-like structures,
and by the reduction of the pharyngobran-
chial teeth to a single patch.

The ancestral stock evidently split rather
early into three developmental lines. One of
these, which can be designated as the sub-
genus Lestidiops, is the most conservative. It
is characterized by a somewhat massive head
and blunt snout, a deep and heavy mandi-
bular symphysis, the absence of nonossified
prolongations on the jaws, a low number of
anal rays (26-28), and the position of the
dorsal fin in the middle third of the space
between pelvic and anal fins.

The second evolutionary line, the subgenus
Lestidium, comprises the majority of the spe-
cies from both the Atlantic and the South
Pacific. It has retained a relatively low number
of anal rays (27-33) but has developed a
slightly less massive head and snout and, in
the most specialized species, a prominent,
nonossified prolongation on the lower jaw
{neks, pacificum, ntidum). In all members of
the subgenus Lestidium the position of the
dorsal fin is over the pelvic fins. The third
evolutionary line of the genus Lestidium,
which is similar to the subgenus Lestidiops in
the position of the dorsal fin in the middle
third of the distance between the pelvic and
anal fins, constitutes a new subgenus, Lestro-
lepis. This clearly delimited group of the genus
Lestidium is distinguished by a high number
of anal rays (33-49), a slender and elongate
lower jaw, a distinctive preorbital pigmenta-
tion, a well-developed, nonossified prolonga-
tion of the lower jaw, and the position of the
dorsal fin.

The remaining Paralepididae have been
placed by Ege in the genus Macroparalepis.
He differentiated this group on the basis of
four characters, presumed to be unique: (1)
the nonossified prolongation on the tip of the
lower jaw; (2) the shape of the lateral-line

segments, which have the greatest height at
or very near the anterior margin; (3) the pres-
ence of two pores above and below on each

FIG. 6. Postlarva of Lestidium sphyraenoides (Risso),
approximately 8 mm. in length, showing the position
of the intestine behind the eye (after Ege, 1930).

lateral-line segment; and (4) the position of
the intestine under the eye in postlarvae (see
Figs. 6 and 7). My investigations show that
at least three of these characters are shared by
other paralepidids, and it may be questionable
that the other is unique to Macroparalepis.
The subgenera Lestidium and Lestrolepis both
have nonossified prolongations of the lower
jaw, and many species of the genus Lestidium
possess characters 2 and 3. The condition
stated for (3) has been found to be the basic
pore pattern for all naked genera and is not
unique to any particular paralepidid group.
In regard to point 4, Ege has stated in corre-
spondence that his groups I and II of ^'Para-
lepsis" both have the same postlarval develop-
ment, with the intestine behind the head, dif-
fering markedly from that of Macroparalepis.
No one, however, has been able to examine
the postlarvae of the species in Lestidium that
most closely approach Macroparalepis, and
there may be some intergradation in these
larval characters.

Fig. 7. Postlarva of Macroparalepis brevis Ege, ap-
proximately 8 mm. in length, showing the position of
the intestine below the eye. (This illustration prepared
from a tracing generously supplied by Ege.)

Ege considers that Macroparalepis is very
distinct and that Lestidium [Paralepis group II]
is much more closely related to Paralepis [Pa-
ralepis group I] than it is to Macroparalepis.
This opinion is based primarily on the dif-
ferences in postlarval development. My con-
clusions are entirely the opposite in regard to
Ege’s Macroparalepis group I. In every adult

228

character, such as in head and body form,
prolongation of the lower jaw, dentition, gill-
rakers, maxillary structure, lateral line, and
lack of squamation, Macroparalepis group I
is very close to Lestidium and can be differ-
entiated from this genus only with difficulty.
While there may be a consistent difference in
the position of the intestine in the postlarvae
of these two genera, I doubt that it is of great
significance.

Ege divided Macroparalepis into two re-
markably distinct groups. The first division,
which was just discussed, contains four spe-
cies which have the anus situated behind the
origin of the dorsal fin, 13-14 dorsal rays, and
24-28 anal rays. Its high number of dorsal
rays surpasses that known for any other form
in the Paralepidinae. The dorsum pigment
pattern is different from that of all other para-
lepidids. This group most closely resembles
the subgenus Lestidiops in general form, anal
ray count, position of dorsal fin, and lateral-
line form and is probably related to the an-
cestral stock of this subgenus. It approaches
the subgenus Lestidium in its nonossified pro-
longation of the upper jaw. The genus Ma-
croparalepis is hereby restricted to Ege’s group
I (since no genotype has been previously
designated).

The second group of Ege’s Macroparalepis,
comprising seven species, is very different
from the first. The anus is situated in front of
the dorsal fin and there are 9-11 dorsal rays
and 37-50 anal rays. The species are very
elongate and some even eel-like, the angle of
the gape extends back almost to the tip of
the maxillary, the tongue is unusually small
and far back, the dentition is considerably
reduced, and the lower jaw is very slender and
pointed with only a slight elevation at its tip.
Also, the shape of the lateral-line segments
is very different from that found in group I
and all other paralepidids, as the length of each
segment is twice the height. Despite the fact
that groups I and II have similar postlarval
development, prolongation of the lower jaw,
and pore pattern, their similarities end with

PACIFIC SCIENCE, Vol. VII, April, 1953

these characters. If these similarities to group
I are put aside for the moment, group II
clearly approaches the subgenus Lestrolepis
with which it shares the prolongation of the
lower jaw, the high number of anal rays, and
the same position of the dorsal fin in relation
to the pelvic fins. In fact, it may very well be
that group II is more closely related to Les-
trolepis than to group I, and that the similari-
ties to the latter group are due to parallel
development or convergence. In any event,
group II is far more different from group I
than group Lis from Lestidium, and it is here
placed in the genus Stemonosudis. A provisional
generic tree showing the degree of relation-
ship of the genera as this now appears to me
is given in Figure 4.

Comparison of specific paralepidid anatom-
ical structures throughout the family shows
some interesting speculative results on the
mode of barracudina evolution. In most char-
acters I have examined there is evident a
straight-line evolution from the primitive con-
dition in the genus Magnisudis to the special-
ized in Stemonosudis through the line of genera
as shown in Figure 4. In almost every instance
this evolutionary pattern involves a loss or
reduction from the form as found in Magni-
sudis. In general, specialization in this family
means a loss or reduction in some pre-existing
character. Such specialization is exemplified
by the gillraker form, which is reduced in the
Paralepididae to a flat bony base. This struc-
ture is most elaborate in the genus Magni-
sudis, being proportionately large and pecu-
liarly notched. Each raker is armed with 4 long
cartilaginous filaments with bony cores ar-
ranged in a square. The genus most closely
approaching this condition is Paralepis. In
this group and all remaining genera the rakers
are flat and simple in outline. Each raker in
Paralepis is armed anteriorly with fairly long
cartilaginous filaments grading posteriorly in-
to short bony spines. The next step is found
in the genus Notolepis where all the armature
of each gillraker consists of short spines.
Thus, we see a trend for the simplification of

Genera of Paralepididae — Harry

229

the form of the gillraker base, the loss of a
cartilaginous sheath from the gillraker spines,
and a progressive reduction in the length of
these spines. There has been, however, an
increase in the number of gillraker teeth. In
Magnisudis there are 3-4, in Paralepis about
5-10, and in Notolepis about 15-30 spines on
each raker. In all three genera the spines are
in two or more rows.

All the rest of the genera in the family
{Lestidium, Macroparalepis, Stemonosudis, and
Sudis) have a singular form of gillraker and
gillteeth which is also characteristic of the
alepisauroid families Scopelarchidae, Everma-
nellidae, Omosudidae, and Anotopteridae. It
approaches most closely the condition in No-
tolepis. The raker base is considerably reduced
in size; the raker spines are very short, similar
in form to those of Notolepis, but in a single
row of 1-7 teeth. There is a tendency for
the loss of gillrakers in Stemonosudis, as indi-
cated by the fact that they appear to develop
later than in other genera.

The degree of similarity of the gillrakers
between the varous genera is the same as the
composite similarity of important characters
and very likely expresses the degree of rela-
tionship. If this hypothesis is true, we could
expect on the basis of gillraker form that
Magnisudis and Paralepis are more closely re-
lated than either is to Notolepis. This state-
ment is corroborated by many other charac-
ters. For instance, the dentition of Magnisudis
and Paralepis is strikingly reduced and tends
to be completely absent in adults. The denti-
tion of Notolepis, however, is well developed,
not atrophied in adults, and approaches more
closely the condition of Lestidium. This same
balance of relationships is found in supra-
maxillary form. The supramaxillary in Para-
lepis and Magnisudis is strongly arched and
widely separated from the maxillary except at
its posterior insertion, but in Notolepis and all
other paralepidid genera it is attached along
its entire border to the maxillary. As a result,
the genus Notolepis is an important link be-
tween the scaled and naked genera; in most

structures it is most closely related to Para-
lepis and Magnisudis, but in some revealing
characteristics it is the same as, or similar to,
the naked genera, particularly Lestidium,
Nevertheless, the genus Notolepis represents
an end point, but in the line of scaled genera
evolution. It has an extreme development of
dentition (particularly in the subgenus Noto-
lepis') and a unique form of lateral-line scales
and tube not equaled elsewhere in the family.
The genus is somewhat intermediate between
Paralepis and Lestidium but the ancestral stock
of the latter genus must have split off rather
early from the Magnisudis-Paralepis-Notolepis
stock. The sharpest break along the evolu-
tionary line of the Paralepidinae as shown in
Figure 4 is definitely between Notolepis and
Lestidium, and the impressive differences of
Magnisudis, Paralepis, and Notolepis (tribe Pa-
ralepidini) from the scaleless genera (tribe
Festidiini) within this subfamily have already
been discussed. The scaleless genera of the
family have very similar gillrakers, and, as
would be expected, all of these genera are
closely related on the basis of a large number
of characters.

Not all characters, however, follow as sim'
pie an evolutionary pattern as the gillrakers.
As has already been demonstrated, the lateral-
line is a structure of considerable phylogenetic
importance but has to be studied carefully
because it can be highly variable even among
individuals of the same species. In the Para-
lepididae the simplest form of the lateral-line
tube and scales is found in Magnisudis, where
each segment has a single pore. In Paralepis
there is usually the addition of a pore above
and below in the tube, and in Notolepis there
are numerous pores in each segment. The
most complex pore patterns and lateral-line
segment shapes are in the genus Lestidium,
and there is finally the stabilization of pore
number in Macroparalepis and Stemonosudis to
two pores above and two below. While com-
parison of lateral-line structure among the
genera will show that the same degree of
relationship exists between the various genera

230

PACIFIC SCIENCE, Vol. VII, April, 1953

as illustrated by the gillrakers, there is not a
progressive reduction in complexity of lateral-
line structure from Aiagnistidis. On the con-
trary, there is a general increase in complexity
of form in the scaled genera from Magnisudis
through Paralepis to the extreme condition
found in Notolepis. Then there is a progressive
reduction in complexity of lateral-line form
in the naked genera from the most complex
in Lestidium through Macroparalepis and Ste-
vionostidts.

Jensen (1942) has presented the most im-
portant discussion on the structure of the
lateral-line tube and its overlying scales. How-
ever, in some species he has confused the
imbedded segments of the tube with overly-
ing lateral-line scales. Therefore, for Paralepis
coregonoides and Paralepis brevis, he figures seg-
ments of the lateral-line tube as lateral-line
scales. As the scales are so caducous in this
genus, the overlying scales were probably
missing (see Fig. 11). Jensen points out that
the lateral-line scales are extremely variable
as to number of pores, which is fully corro-
borated by the present study.

List of Species Currently Recognized

The following list shows all the known
living species of Paralepididae currently recog-
nized in the literature. The synonyms are as
given by Ege (1930), Parr (193H), and Maul
(1945). No attempt is made to put closely
related species together as so many are in-
adequately known.

Sudis squamosa Chapman (1939: 519) from
south of the Aleutian Islands and Lestidium
{Bathysudis) parri Chapman (1939: 519) from
the Gulf of Alaska do not belong in the
famdly Paralepididae. Hubbs and Chapman
(1951) have conclusively identified Sudis squa-
mosa as Mallotus catervarius (Steller) and Les-
tidium parri as Thaleichthys pad ficus (Richard-
son).

Subfamily Paralepidinae Bonaparte, 1846.

Magnisudis, n. gen.

"^harysoma, n. sp. (generic type). Pacific,
off California and Mexico.

Paralepis Cuvier, 1817.

'^coregonoides (Risso, 1810) (generic type).
North Atlantic.

^speciosa Bellotti, 1878. Mediterranean.
elongata (Brauer, 1906). Indian Ocean,
^brevis Zugmayer, 1911. North Atlantic.
*brevirostris (Parr, 1928). North Atlantic.
bronsoni (Parr, 1928). Bahamas.
danae Ege, 1933. Off Cape Verde, West
Africa.

Notolepis Dollo, 1908.

^rissoi (Bonaparte, 1841). North Atlantic.
*coruscans (Jordan and Gilbert, 1881).
North Pacific.

*coatsi Dollo, 1908 (generic type). Ant-
arctic.

Lestidium Gilbert, 1905.
sphyraenoides (Risso, 1810). North Atlan-
tic.

intermedium (Poey, 1867), West Indies.
*ringens (Jordan and Gilbert, 1881). Cal-
ifornia.

jayakari (Boulenger, 1889). Indian
Ocean, off Arabia.

nudum Gilbert, 1905 (generic type). Ha-
waiian Islands, New Zealand (.^), Ja-
pan (.?).

japonicum Tanaka, 1908. Japan.
*sphyraenopsis (Hubbs, 1916). California.
*pseudosphyraenoides (Ege, 1918). North At-
lantic.

atlanticum (Borodin, 1928). North At-
lantic.

thori (Ege, 1930). North Atlantic,
danae (Ege, 1930). North Atlantic.

"^affine (Ege, 1'930). North Atlantic.
*pacificum (Parr, 1931). Pacific off Mexico.
proximum (Ege, 1933). Celebes Sea.
simile (Ege, 1933). Off Cape Verde, West
Africa.

luetkeni (Ege, 1933). Mozambique Chan-
nel.

bellottii (Ege, 1933). New Guinea.
mirabile (Ege, 1933). Celebes Sea.
^philippinum (Fowler, 1934), Philippine Is-
lands.

^bathyopteryx (Fowler, 1944). Pacific, off
Mexico.

Genera of Paralepididae — Harry

231

"^vanderbilti (Fowler, 1944). Caribbean.
^neles, n. sp. California.

"^atrox, n. sp. Hawaiian Islands.
*thunnorum, n. sp. Northeastern Pacific.
^prolixum, n. sp. Japan.

n. sp. Hawaiian Islands.

Macroparalepis Ege, 1933.

"^affine Ege, 1933 (generic type). North
Atlantic.

"^danae Ege, 1933. Fiji Islands, Bermuda

(?).

"^hrevis Ege, 1933. Off St. Helena, North
Atlantic.

egei Maul, 1945. Madeira Island, North
Atlantic.

Stemonosudis Yidsvj , 1951.

macrura (Ege, 1933). Sumatra.
elegans (Ege, 1933). Lord Howe Jsland,
South Pacific.

miscella (Ege, 1933). Sumatra.
elongata (Ege, 1933). Ceylon.
gracile (Ege, 1933). Straits of Macassar,
South Pacific.

"^intermedia (Ege, 1933) (generic type).
Caribbean Sea.

Subfamily Sudinae Regan, 1911.

Sudis Rafinesque, 1810.

"^hyalina Rafinesque, 1810 (generic type).
Mediterranean and Madeira.

Distribution of Genera and Species.

The family Paralepididae is one of the larg-
est in the order Iniomi, consisting of 48
known bathypelagic species. It is world- wide
in distribution and apparently occurs in large
numbers in deep water to a depth of about
4,000 meters. The species are generally wide-
ranging. Adult specimens seldom fall into the
hands of ichthyologists, and the majority of the
adult material has been taken from the stom-
achs of Alepisaurus, tuna, other larger pelagic
fishes, and whales. The general form of para-
lepidids indicates that these are swift-moving
voracious fishes. Larger specimens have very
rarely been taken by net and apparently never
by hook and line. No sexually mature indi-

* Species examined in the present study.

viduals have been reported, and Dr. Bruun
informs me that the "Dana” expeditions were
not able to find or, at least, to determine the
eggs of this family. It is possible that some
or most of the species grow to large size and
that no sexually mature adults have been ob-
tained.

The best-known group, the genus Lesti-
dium, apparently is the only genus whose
members come to the surface where adults
may be taken by light at night. The range and
abundance of the various species of barra-
cudinas are best understood in the North At-
lantic and Mediterranean, mainly because of
the extensive Danish Oceanographical Expe-
ditions, which obtained the large collections
reported on by Ege. No examples have bene
recorded from the South Atlantic (excluding
the Antarctic), but 20 species are recognized
in the North Atlantic. Sudis hyalina is appar-
ently fairly common in the Mediterranean and
is also known from Madeira.

About half of the known species of the
Paralepididae occur in the Pacific Ocean. The
northernmost form is a species of Notolepis
taken off the coast of Washington. Eleven
species of Lestidium and Magnisudis b ary soma
are variously recorded in the North Pacific
from off Japan, the Philippine Islands, the
Hawaiian Islands, Mexico, and California.
These species of Lestidium (except L. nudum,
which is apparently known from the Hawaiian
Islands, Japan, and New Zealand) are known
only from restricted localities. The barracu-
dinas have been collected in the more south-
erly Pacific in the East Indies and off New
Zealand where five species of Lestidium, Ma-
croparalepis danae, and five species of Stemono-
sudis have been recorded. All the East Indian
species are known only from postlarval or
juvenile material, except Lestidium nudum.

Very little material has been collected from
the Indian Ocean, two of the three recorded
species being represented by the holotypes
only. These forms are Lestidium luetkeni from
the Mozambique Channel, L. jayakari from
Muscat, Gulf of Oman and Paralepis elongata
from the Seychelles Islands, Chagos Archi-

232

PACIFIC SCIENCE, VoL VII, April, 1953

pelago, south Ceylon, Bay of Bengal, and
Cocos (Cocos Keeling) Islands. The distinc-
tion between the Indian and Pacific faunas
cannot be sharply defined. P. elongata, the
only species known from the eastern Indian
Ocean, is recorded very near the Pacific. This
species apparently has the greatest latitudinal
range of all barracudinas, but, considering
that Brauer’s types are postlarvae ranging
from 8-30 mm., there may be more than one
species represented. It is expected that more
collecting will reveal various species ranging
through the Indo-Pacific region.

The distribution of the Paralepididae in the
Antarctic is fairly well known. The "Erebus
and Terror," "Challenger," "Pourquoi Pas,"
"Scotia," "Terra Nova," and the B.A.N.Z.
(British, Australian, and New Zealand) Ant-
arctic Research Expeditions collected material
from this region, but it was composed almost
exclusively of larval to juvenile specimens
whose relationships and classification were
generally not understood by the original de-
scribers. Such confusion has resulted in the
subsequent identification of these forms, that
only one abundant species, Notolepis coatsi, of
circumpolar distribution, is definitely recog-
nized from the Antarctic.

No species has been recorded from both
the Atlantic and Pacific Oceans, although this
apparent distinctiveness may be due to a lack
of sufficient material and of proper compari-
son. Dr. Ege is presently examining the rela-
tionships and supposed differences between
the Atlantic and Pacific species and plans to
publish his findings.

Apparently no paralepidids have been re-
corded from the Arctic Ocean.. The distribu-
tion of the three species of Notolepis in the
North Pacific, North Atlantic, and Antarctic
gives a good example of bipolar (pantemper-
ate) distribution.

Explanation of Morphological Figures.

Figures 8, 10, 13, 14, 16, 18, 20, and 22
have been prepared in a standard manner to
facilitate comparison of some of the more
important morphological characters. These

drawings are somewhat diagrammatic but are
accurate for the characters involved. These
figures present the following information:

A: Anterior part of snout. The teeth that are
solid black are depressible; the remainder
are fixed. The buccal valves and supra-
maxillary membranes are stippled. The nos-
trils on the snout and larger pores on the
lower jaw are indicated.
b: Lateral aspect of an enlarged section of the
fixed teeth on the middle of the premaxil-
lary.

C: The anterior lateral-line segments on the
left side. The stippled area represents the
weak membranous part of the tube. The
area with longitudinal parallel lines de-
limits the partly ossified center shield in the
naked genera {Lestidium, Macroparalepis,
and Stemonosudis) and the central row of
scales in the scaled genera (Magnisudis, Pa-
ralepis, and Notolepis) . The dotted lines show
the outline of the internal limits of the
lateral-line tube. The crossed lines indicate
the scales above and below the central
lateral-line row. The characteristic pigmen-
tation in the lateral-line region is presented
for some of the species.
d; a section of the ceratobranchial of the
first arch showing the gillrakers and gill-
teeth. The parallel lines indicate the gill
arch. The light line over the tips of the
teeth shows the mucus line.

E: Dorsal surface of tongue (glossohyal) and
anterior portion of first basibranchial. The
stippled area represents the fleshy tongue.
The glossohyal and basibranchial are indi-
cated by longitudinal parallel lines. The
small hooked circles indicate the teeth.

KEY TO GENERA AND SUBGENERA

1. Teeth on mandible large, triangular,
strongly compressed, with serrate edges;
always present. Anterior process of pre-
maxillary without a foramen. Outer rays
of pelvic fins longer than inner rays.
(Subfamily sudinae, one genus) . . . Sudis
Teeth on mandible slender, basally
round, sometimes hastate, but never

I

Genera of Paralepididae — Harry

serrate; sometimes absent. Anterior pro-
cess of premaxillary with wide circular
foramen. Outer rays of pelvic fins dis-
tinctly shorter than inner rays (Sub-
family paralepidinae) 2

2. Teeth on each gillraker in 2 or more
rows (Fig. IOd). Two large, sep-
arate tooth patches on pharyngobran-
chials on each side. Teeth on lower jaw
comparatively short or (in some adults)
missing, their length up to as much as
5 per cent of snout length. Body scaled

in adults 3

Teeth on gillrakers in one row (Fig.
16d). One tooth patch on pharyngo-
branchials. Teeth on lower jaw long,
their length 5-11 per cent (generally
above 7) of snout length Fig. 16a).
Body naked 6

3. Each gillraker with 4 long filaments in
2 rows (Fig. 8d). Gillrakers extending
far forward into mouth, beyond angle of
gape. Each lateral-line scale tenacious,
much larger than surrounding scales,
bordered above and below by at least 3

scales (Fig. 8c) Magnisudis

Each gillraker with many spine-like
teeth in numerous series (Fig. 13d,
14d). Anterior limb of gillarches naked;
gillrakers not extending forward to angle
of gape. Each lateral-line scale caducous,
same size as surrounding scales, bor-
dered above and below by single scale
(Fig. lOc) 4

4. Teeth on each gillraker not uniform in
size, arranged in a bunch, anterior ones
long, extending over next raker; pos-
terior teeth short, needle-like (Fig. IOd).

' Supramaxillary free from maxillary ex-
cept at posterior insertion (Fig. 10a).
Upper jaw terminating slightly before to
slightly behind a vertical from anterior
border of eye. Nostrils placed one-fourth
or more of upper jaw length anterior to

posterior tip of maxillary Paralepis

Teeth on each gillraker very short, sub-
uniform in size, arranged in oblique

233

rows (Fig. 13d). Supramaxillary closely
bound to maxillary (Fig. 13a). Upper
jaw terminating almost an eye diameter
before orbit. Nostrils over posterior tip
of maxillary Notolepis 5

5. Mandible with 3 rows of teeth, each
series having approximately 35 canines
(Fig. 13a). Palatine with approximately
50 short fixed teeth in single irregular

row Subgenus Notolepis

Mandible with 2 rows of teeth, totaling
approximately 25 canines (Fig. 14a).
Palatine anteriorly with 3 long, widely
spaced, depressible canines accompanied
by short fixed teeth; posteriorly 30 or

fewer fixed canines in single row

Subgenus Profundisudis

6. Lower jaw fairly massive, blunt, tip dis-
tinctly elevated. Nostrils distinctly be-
fore a vertical from posterior tip of
maxillary (except in Lestidium neks).
Angle of gape far before posterior tip
of maxillary. Pattern of dorsum variable,

but without saddle-like blotches 7

Lower jaw very slender, pointed, tip not
elevated (Fig. 20a). Nostrils distinctly
behind posterior tip of maxillary. Angle
of gape near tip of maxillary. Dorsum
not evenly pigmented; posteriorly sev-
eral saddle-like blotches on dorsum, al-
ternating with similar blotches on base
of anal fin (Fig. 19) Stemonosudis

7. Dorsal fin with 9-12 rays. Dorsum light-
ly and evenly pigmented only part way
down to lateral-line (Fig. 15) except in
Lestidium pofi, which is evenly pigmented
all over. Lateral-line segments unpig-
mented or with a few scattered melano-
phores. Anterior lateral-line segments
usually longer than broad. (Fig. I6c)

Lestidium, 8

Dorsal fin with 11-14 rays (usually
13-14). Dorsum heavily pigmented with
large melanophores down onto lateral-
line scales (particularly posteriorly).
Adults with lateral-line segments usually

broader than long (Fig. 18c)

Macroparalepis

I

234

PACIFIC SCIENCE, Vol. VII, April, 1953

8. No prominent markings before eye.

Anal rays 26-33* 9

Prominent round black spot covering
vertical finger-like projection immedi-
ately before eye. Anal rays 33-49

Subgenus Lestrolepis

9. Origin of dorsal fin more than half a
head length behind a vertical of pelvic

fins. Anal rays 26-28

Subgenus Lestidiops

Dorsal fin over or nearly over pelvic fins,
origin less than one third of a head
length behind pelvic base. Anal rays
28-33 Subgenus Lestidium

Subfamily Paralepidinae
Head and lower jaw moderately large. Pair
of large nostrils on each side of snout. An-
terior process of premaxillary with large cir-
cular foramen. Teeth on mandibles, if present,
slender, basally round, sometimes hastate, but
never serriform. Dorsal fin with 9-14 (seldom
14) rays. Pectoral fin moderately developed,
not proportionately larger in younger stages.
Pelvic fins with inner rays distinctly longer
than outer rays. Scales, if present, never just
restricted to preoperculum.

Magnisudis new genus
Figs. 5, 8, 11^

Recently, Dr. Carl L. Hubbs obtained two
adults of scaled paralepidids a foot long from
the stomachs of marlin and the great blue
shark off Lower California. In addition, the
California Division of Fish and Game ob-
tained two adults approximately 20 inches
long and some smaller examples from Califor-
nia. Not only are these specimens the largest
paralepidids known, but they represent the
most generalized of the scaled barracudinas
and are distinct in many important characters
from all known genera.

DIAGNOSIS: Body large and massive, mod-

* Matsubara’s material oi" Lestidium japonicum" with
35 anal rays (1941: 8) is not included, as it appears to
be a new species and the description is not complete
enough to place it in a subgenus. Probably it belongs
to the subgenus Lestrolepsis.

Fig. 8. Genotype of Magnisudis {M. barysoma, n. sp.,
paratype, 510 mm. in standard length, from off Lower
California). A, Anterior part of snout: B, enlarged sec-
tion of teeth on middle of premaxillary; c, anterior
lateral-line scales; D, gillrakers on first arch; e, dorsal
surface of tongue. (See explanation of morphological
figures in text.)

erately compressed. No carinae. Head large,
conical, moderately compressed. Eye larger
Pupil small, crescent or half-moon shaped,
much smaller than lens. Snout massive. Nos-
trils well before end of maxillary. Tip of lowe,
jaw elevated. Upper jaw terminating well be-
fore vertical from anterior border of eye.,;
Supramaxillary long, approximately 0.33-0.5 ;
as long as maxillary, curved and rod-shaped, >
free from maxillary except at posterior inser- ^
tion. Teeth on lower jaw sparse or absent; if)
present very short; all fixed. Teeth on upper ;
jaw very tiny, closely spaced; fixed teeth in y
saw-like row. Vomer toothless or with single
median tooth. Teeth on palatines very short,

Genera of Paralepididae — Harry

235

in single row, fixed. Tongue (glossohyal and
its fleshy expansion) small and short, but
projected far forward near tip of lower jaw.
Gillrakers and entire branchial apparatus ex-
tending far forward into mouth. Gillrakers
well developed on all 5 arches. Each gillraker
with 4 long, stiff, cartilaginous, depressible
filaments, longest filament 1.5 times greatest
pupil diameter. Pharyngobranchial teeth de-
veloped in 2 tooth patches on each side. Gill
membranes free and separate, joined far for-
ward under vertical from nostrils. Left bran-
chial membrane overlaps right.

Sides of head heavily scaled forward onto
preorbital. Occiput covered with scales and
lacking tubes or ridges. Body completely
scaled. Scales small and moderately adherent,
slightly pointed posteriorly and often indent-
ed anteriorly, appearing heart-shaped. Scales
extending on middle of caudal base for ap-
proximately 0.25 its length. Circuli of each
scale not continuous except for inner 8 ridges
or so; remaining circuli run obliquely off
scale. Lateral-line tube fairly small with single
median pore on each segment. One row of
enlarged, strongly adherent, membranous
scales over the tube; each lateral-line scale
bordered by 3 or more scales above and below.
Body scales much smaller than lateral-line
scales.

Pectoral fin with 17 rays. Anal rays 22-25.
Dorsal fin origin slightly in advance of pelvic
fin origin. Anus behind vertical from dorsal
base. Pelvic fins very short, approximately
0.33 length of pectoral fins.

Generic TYPE and only known form, Mag-
nisudis harysoma n. sp.

The name Magnisudis is formed from mag-
nus, L.— large 4- stidis, L.— stake, pile, pike.

The above description will serve to dis-
tinguish and demonstrate the relationships of
both the genus Magnisudis and its single spe-
cies M. harysoma, although the full specific
description will appear only in Part 2 of the
present study. The holotype of M. harysoma,
from off southern California, and the para-
types are deposited at Stanford University.

Genus Paralepis Cuvier
Ligs. 9, 10, \\h

Paralepis Cuvier, 1817: xi, 289 (generic type
by subsequent designation of Jordan and
Evermann, 1917: 104, 120, Paralepis core-
gonoides Risso); Bose, 1818: 520; Cuvier,
1829: 156; Cuvier and Valenciennes, 1829:
556; Risso, 1826: 472; Gunther, 1864: 418;
Goode and Bean, 1895: 118; Jordan and
Evermann, 1896: 602; Regan, 1911: 127;
Jordan and Evermann, 1917: 104; Jordan,
1923: 154; Parr, 1928: 34, 71; Parr, 1929:
29; Ege, 1930: 6; Parr, 1931^: 19; Parr,
193U: 152; Whitley, 1937: 11; Maul, 1945:
4; Harry, 1951: 18.

Arctozenus Gill, 1865: 188 (generic type by
original designation, Paralepis horealis Rein-
hardt); Goode and Bean, 1895: 5 16; Jordan
and Evermann, 1896: 601; Jordan, 1919:
330; Jordan, 1923: 154; Parr, 1928: 33.
Symproptopterus Cocco, 1885 (no generic type,
see Jordan, 1920: 430).

Sudis (in part) Parr, 1928: 34.

Bathysudis Parr, 1928: 41 (generic type by
original designation, Paralepis speciosa Bel-
lotti); Ege, 1930: 53; Parr, 193U: 153;
Gregory, 1933: 207, 209; Gregory and Con-
rad, 1936: 33; Chapman, 1939: 524.

The author of Paralepis has been often given
as Risso. However, Cuvier was the first to use
the name in a generic sense. According to
Whitley, the generic name was introduced
strictly in vernacular form (Les Paralepis Cv.)
by Cuvier, 1817, and Bose was the first to
latinize it, thus becoming the author of the
genus. However, Cuvier listed Paralepis as a
generic name {Paralepis C.) in the "Table
Methodique,” and there appears to be no rea-
son for not accepting Cuvier as the author of
the genus (see Opinion 39 of the International
Commission of Zoological Nomenclature for
a similar case involving Cuvier, 1800).

diagnosis: Body compressed, moderately
short and deep. Ventral carina on belly little
developed. Head and eye large. Pupil round,
larger than lens. Snout short and broad. Nos-
trils well before end of maxillary. Tip of lower

236

PACIFIC SCIENCE, Vol. VII, April, 1953

Fig. 9- Representative of Paralepis (P. brevirostris (Parr), 137 mm. in standard length, from off Medeira).

jaw strongly elevated. Upper jaw reaching to
or slightly beyond vertical from anterior bor-
der of eye. Supramaxillary long, in adults
almost 0.5 as long as maxillary, curved and
rod-shaped, free from maxillary except at pos-
terior insertion. Teeth on lower jaw short and
weak, tending to be absent in adults. Teeth
on palatines short, in 2 irregular rows anterior-
ly, one row depressible; posteriorly in one row
of up to approximately 15 short, fixed teeth.
Tongue large, moderately forward in mouth.
Gillrakers developed on all 5 arches, not ex-

Fig. 10. Genotype of Paralepis (P. coregonoides'BJisso,
from off Funchal, Madeira, 145 mm. in standard length).
A, Anterior part of snout; B, enlarged section of teeth
on middle of premaxillary; c, anterior lateral-line scales;
D, gillrakers on first arch; E, dorsal surface of tongue.
See explanation of morphological figures in text.

tending forward beyond angle of gape. Gill-
rakers spinous, anterior teeth on each raker
long; posterior teeth very short, needle-like,
in numerous rows. Longest rakers as long as
pupil diameter. Pharyngobranchial teeth de-
veloped in 2 tooth patches on each side.
Branchial membranes do not overlap.

Body fully scaled. Scales very deciduous,
moderately pointed posteriorly, lacking any
indentation on posterior margin. Circuli on
scales not continuous except for inner 5 rings
or so; remaining circuli run obliquely off the
scales (see Fig. 11^, and Jensen, 1942: 23,
Fig. 9). Lateral-line tube moderately large; 1.5
or 2 scale rows lying over tube. (In the basic
pore pattern for this genus the upper scale
row over the lateral-line is pierced by the
upper pore, and the scale row over the middle
of the tube is penetrated by the median and
lower pores. There is often reduction in the
number of pores piercing the scales.) Body
scales same size as lateral-line scales. Pectoral
fin rays 14-17. Anal rays 21-26. Vertebrae
67-74.

Disregarding the inadequately known Pa-
ralepis elongata (Brauer), the genus Paralepis
could be divided into two subgenera by seg-
regating the generic type, P. coregonoides, in '
the subgenus Paralepis and recognizing the
remainder of the species in the subgenus
Bathysudis. I have not done this because these
forms are quite similar and closely related..
Paralepis elongata is distinct in several basic
characters from the remaining species of this
genus. If it belongs in Paralepis it could be
readily placed in a new subgenus. '

Species I have examined that are undoubted

Genera of Paralepididae ^ — HARRY

237

Fig. 11. Scales taken from behind and above the
pectoral fins of three genera: a, Magnisudis {M. bary-
soma, n. sp.); h, Paralepis (P. coregomides Risso); c,
Notolepis (N. coatsi Dollo). (Illustrations prepared from
direct projections.)

members of the genus Paralepis are: P. hrevis
Zugmayer, P. hrevirostris (Parr), P. coregonoides
(Risso), and P. speciosa Bellotti. I believe that
P. danae Ege (1933) and perhaps P. elongata
(Brauer) also belong here. P. hronsoni (Parr)
appears to be a synonym of P. hrevirostris
(Parr) (Harry, 1951: 19).

Genus Notolepis Dollo
Figs. 11c, 12, 13, 14

PPrymnothonus Richardson, 1845: 51 (generic
type by monotypy, Prymnothonus hookeri
Richardson); Gunther, 1870: 175; Gunther,

1889: 39; Dollo, 1904: 8; Dollo, 1908: 35;
Page, 1910: 16; Regan, 1911: 127; Regan,
1913: 233; Regan, 1914: 38; Regan, 1916:
125; Jordan, 1920: 527; Jordan, 1923: 154;
Parr, 1928: 33; Norman, 1937: 83.

Notolepis Dollo, 1908: 58 (generic type by
monotypy, Notolepis coatsi Dollo); Regan,
1911: 127; Regan, 1913: 233; Regan, 1914:
38; Regan, 1916: 125; Jordan, 1920: 527;
Jordan, 1923: 154; Parr, 1928: 33; Norman,
1937: 83; Harry, 1951: 26.

Paralepis (in part) Ege, 1930; Maul, 1945: 4.
PPrymnothonoides Whitley and Phillips, 1939:
228 (generic type by original designation,
Prymnothonoides reganiWhkXtY and Phillips) .
Although this genus was very poorly de-
scribed by Dollo and was based on irrelevant
and insignificant characters, it was erected for
one of the most distinctive scaled forms of
barracudinas. Notolepis is resurrected on en-
tirely different characters of both postlarvae
and adults.

DIAGNOSIS: Body compressed and elongate.
Ventral carina on belly moderately developed.
Head and snout long and sharply pointed.
Nostrils behind or over posterior tip of maxil-
lary. Eye moderately large. Pupil round or
elleptical, larger than lens. Tip of lower jaw
moderately elevated. Upper jaw terminating
approximately one orbital diameter before eye.
Supramaxillary short, splinter-like, closely
bound to maxillary. Teeth on lower jaw well
developed, in 2 or 3 rows. Teeth on palatines
short, in 1 or 2 rows anteriorly; posteriorly in
one row of 30-50 teeth. Tongue narrow, well
forward in mouth. Gillrakers not extending
forward beyond angle of gape; gillrakers spin-
ous, entirely reduced to many rows of short
fixed needle-like teeth. Longest rakers much
shorter than pupil diameter. Pharyngobran-

Fig. 12. Representative of Notolepis (N. coruscans ]orda.n and Gilbert, from the Pacific off Washington, 159
mm. in standard length).

238

Fig. 13. Genotype of Notoleph (N. coals/ Dollo, from
the Antarctic, 302 mm. in standard length). A, Anterior
part of snout; B, enlarged section of teeth on meddle
of premaxillary; D, gillrakers on first arch; E, dorsal
surface of tongue. (See explanation of morphological
figures in text.)

chial teeth developed in 2 patches on each
side. Branchial membranes do not overlap.

Body fully scaled. Scales very deciduous,
rounded or pointed posteriorly, occasionally
with indentation on anterior margin. The cir-
culi are continuous for inner 8 rings or more,
outermost rings run obliquely off the scales
(see Fig. 11c, and Jensen, 1942: 22, Fig. 6).
Lateral-line tube large, with at least 1 pore
above, medially, and below on each segment;
tube covered by IJ^ rows of scales. Upper
scale row sometimes notched to receive upper
pore, as shown in Fig. I4c, or may be pierced
by pore series as in Paralepis. Lower scale usu-
ally penetrated by median and lower pores.
Body scales same size as lateral-line scales.
Pectoral fin with 11-13 rays. Anal rays 27-34.
Vertebrae 78-95.

All three known species of the genus f^oto-
lepis have been examined. These can be di-
vided into two subgenera as N. coatsi Dollo
differs markedly from N. rissoi (Bonaparte)
and N. coruscans (Jordan and Gilbert) in sev-
eral characters.

Subgenus Notolepis Dollo
Fig. 11c, 13

DIAGNOSIS: Mandible with 3 series of teeth,
each row with approximately 35 canines. Pa-
latine with approximately 50 short fixed teeth
in single row. Each gillraker on ceratobran-

PACIFIC SCIENCE, Vol. VII, April, 1953 .*^

chial of first arch with approximately 30 short, ‘
fixed, needle-like teeth. Apparently, each
lateral-line segment with large median-pore
at posterior margin and tiny pore above and
below near anterior margin. Anal rays 27-29.

This subgenus is monotypic, containing
only Notolepis coatsi from the Antarctic.

Profundisudis new subgenus
Eigs. 12, 14

Arctozenus (in part) Jordan and Evermann,

1896: 601.

DIAGNOSIS: Mandible with 2 series of teeth,
totaling approximately 25 canines. Palatine
anteriorly with long widely spaced depressible
canines accompanied by short fixed teeth,
posteriorly with 30 or fewer short fixed can-
ines in single row. Each gillraker on cerato-
branchial of first arch with approximately 15
short fixed spine-like teeth. Each lateral-line
segment with 1 large pore above and another
below at posterior margin, with pair of minute

Fig. 14. Representative of the genus Notolepis and
the new subgenus Profundisudis (N. coruscans Jordan
and Gilbert), from the Pacific off Washington, 159
mm. in standard length). A, Anterior part of snout;

B, enlarged section of teeth on middle of premaxillary;

C, anterior lateral-line scales; d, gillrakers on first arch;

E, dorsal surface of tongue. (See explanation of morph-
ological figures in text.) .

Genera of Paralepididae — HARRY 239

Fig. 15. Representative of the genus Lestidium (L. pacificum (Parr), holotype, from the Pacific off Mexico, 164
mm. in standard length). This species is closely related to the genotype.

pores between; another pair of minute pores
near anterior margin of each' section. Anal
rays 30-34.

Subgeneric TYPE, Arctozenus coruscans
and Gilbert, from the North Pacific. This sub-
genus also contains N. rissoi from the North
Atlantic.

The name is formed from profundus, L. —
of the depths + sudis, L.— stake, pile, pike.

Genus Lestidium Gilbert
Figs. 1, 2, 3, 6, 15, 16

Paralepis (in part) Cuvier, 1817: xi, 289; Ege,
1930; Maul, 1945: 4.

Lestidium Gilbert, 1905: 607 (generic type by
original designation, Lestidium nudum Gil-
bert); Regan, 1911: 127; Hubbs, 1916: 154;
Jordan, 1920: 513; Jordan, 1923: 154; Bo-
rodin, 1928: 10; Parr, 1928: 33; Parr, 1929:
29; Parr, 1931^: 19; Parr, 1931^: 153; Greg-
ory, 1933: 209; Gregory and Conrad, 1936:
33; Ege, 1933: 229; Norman, 1937: 83;
Chapman, 1939: 524; Harry, 1951: 26.
Lestidiops Hubbs, 1916: 154 (generic type by
original designation, Lestidiops sphyraenopsis
Hubbs); Jordan, 1920: 559; Parr, 1928: 33;
Parr, 1931^: 19; Parr, 193l/>': 153.

This genus is the largest in the family and
also the most abundant. Generally it has been
taken wherever the family has been recorded.
It is best known in the northern hemisphere
where the majority of the species have been
found. It will probably be found to be abund-
ant also in the Central Pacific, particularly
from the region of the Philippine Islands
northward to Japan. Lestidium is most closely
related to the genera Macroparalepis and Ste-
monosudis and appears to be the least spe-
cialized of these three groups.

Adults are attracted by light at night and
can be most readily taken by this means. In

fact. Dr. Earl S. Herald and Dr. A. W. Herre
have informed me that large examples of Les-
tidium' philippinum are found in the fish mar-
kets of the Philippine Islands and are taken
by the native fishermen with lights.

Gilbert (1905) described Lestidium as hav-
ing "a photophore directed downward and
backward at lower orbital margin”. This was
in error; no paralepidid has any light organ.

DIAGNOSIS: Body compressed, elongate.
Ventral carina on belly well developed. Head
and snout long, pointed. Eye large. Pupil
oval or round, larger than lens. Nostrils before
or over posterior tip of maxillary. Tip of lower
jaw strongly elevated. Upper jaw terminating
at or well before vertical from anterior margin
of orbit. Angle of gape well before tip of
maxillary. Supramaxillary short, splinter-like,
closely bound to maxillary. Teeth on lower
jaw long and well developed, with antrorse
and retrorse canines. Vomer sometimes with
few teeth in young. Teeth on palatines long,
in 2 rows anteriorly, 1 row depressible; pos-
teriorly with approximately 5-15 fixed teeth
in 1 row. Tongue large, moderately forward
in mouth. Gillrakers not extending beyond
angle of gape; spinous, with 1 row of short
fixed teeth on small base. Pharyngobranchial
teeth developed in 1 patch only on each side.

Body and head naked, lacking scales.
Lateral-line tube large. Membrane over lateral-
line modified into scale-like structures which
are highly variable between species. Each
lateral-line segment variously pierced above
and below by 1 to 5 pores but basic pore
pattern of 2 pores above and below. Some
forms also with median pore. Greatest height
of each lateral-line segment variously at front,
middle, and hindmost point. Pectoral fin with
11-13 rays. Dorsal rays 9-12. Anal rays 27-49.

240

PACIFIC SCIENCE, Vol. VII, April, 1953

Fig, 16. Representative of the genus Lestidium (L.
pacificum (Parr), holotype, from the Pacific off Mexico,
164 mm. in standard length). A, Anterior part of snout;

B, enlarged section of teeth on middle of premaxillary;

C, anterior lateral-line segments; D, gillrakers on first
arch; E, dorsal surface of tongue. (See explanation of
morphological figures in text.)

Vertebrae 77-94, so far as known.

As previously stated, the genus Lestidium
can be split into three distinct evolutionary
lines that may be designated as subgenera.

Subgenus Lestidiops Hubbs

DIAGNOSIS: Head massive. Snout deep and
blunt. Tip of lower jaw with 3 vertical non-
ossified ridges on symphysis, but lacking any
prolongations. Eye large, 4. 2-5. 7 into head.
No distinct markings before eye. Dorsal fin
behind pelvic fins; distance between dorsal
and pelvic fins more than 0.5 length of head.
Anal rays 27-28. Each lateral-line segment
with 1 to 5 pores (generally 1 or 2) above and
below.

This subgenus contains L. sphyraenopsis
(Hubbs), L. thunnorum n. sp., and L. apfine
Ege. All three species have been examined.
Subgeneric type, Lestidiops sphyraenopsis

Hubbs.

Subgenus Lestidium Gilbert

DIAGNOSIS; Head slender. Snout moderately
pointed, not deep. Tip of lower jaw with or
without nonossified ridges or prolongations.
Eye large. No distinct markings before eye.
Dorsal fin near or over pelvic fins; distance
between dorsal and pelvic fins less than 0,33
length of head. Anal rays 28-33. Each lateral-
line segment with 3-5 pores above and below.

Among the species that belong in this sub-
genus I have examined L. nudum Gilbert,
L. pseudosphyraenoides (Ege), L. neks n, sp.,
L. pofi n. sp., L. prolixum n. sp., L. pacificum
(Parr). Other species that apparently belong
here: L. proximum (Ege), L. sphyraenoides
(Risso), L. simile (Ege), L. luetkeni (Ege), L.
mirabile (Ege), L. atlanticum Borodin, L.
iayakari (Boulenger) .

Subgeneric type, Lestidium nudum Gilbert,
Lestrolepis new subgenus

DIAGNOSIS: Head slender. Snout pointed,
not deep. Tip of lower jaw with prominent
nonossified prolongation. Eye relatively small,
6. 3-7. 5 into head. Prominent round black
spot immediately before eye. Each lateral-line
segment with 3-5 pores above and below.
Dorsal fin behind pelvic fins; distance be-
tween dorsal and pelvic fins less than 2.5
times into head. Anal rays 35-49.

Subgeneric type, Paralepis philippinus Fow-
ler. This subgenus probably also contains L.
japonicum Tanaka, L. intermedium (Poey), and
L. hellottii (Ege), but only the subgeneric
types, L. vanderhilti (Fowler), and L. atrox n.
sp. have been examined.

The name is formed from lestro, Gr.— pirate
d- lepis, Gr.— scale.

Genus Macroparalepis Ege
Figs. 7, 17, 18

Macroparalepis Ege, 1933: 229 (generic type
not given or subsequently designated; here
designated as Macroparalepis affine Ege);
Maul, 1945: 26; Harry, 1951: 29; Maul,
1952: 51.

This genus is here restricted to group I

Genera of Paralepididae — Harry

241

Fig. 17. Genotype of Macroparalepis {M. affine Ege, from off Madeira, 129 mm. in standard length).

II of Ege’s Macroparalepis is placed in another
genus, Stemonosudis.

DIAGNOSIS: Body compressed and elongate.
Ventral carina on belly well developed. Head
and snout long and pointed. Nostrils before
tip of maxillary. Tip of lower jaw strongly
elevated. Upper jaw terminating slightly be-
fore vertical from anterior margin of orbit.
Angle of gape well before tip of maxillary.
Supramaxillary short, splinter-like, closely
bound to maxillary. Teeth on lower jaw long
and well developed, in 2 rows. Teeth on upper
jaw well developed antrorse and retrorse ca-
nines. Vomer toothless. Teeth on palatines
long, in 2 series anteriorly, 1 row depressible;

Fig. 18. Genotype of Macroparalepis {M. affine Ege,
from off Madeira, 129 mm. in standard length).
A, Anterior part of snout; B, enlarged section of teeth
on middle of premaxillary; c, anterior lateral-line seg-
ments; D, gillrakers on first arch; E, dorsal surface of
tongue. (See explanation of morphological figures in
text.)

few short teeth in single row posteriorly.
Tongue large, moderately forward in mouth.
Gillrakers not extending forward beyond an-
terior border of eye or angle of gape. Gill-
rakers spinous, with 1 row of short fixed
teeth on small base. Pharyngobranchial teeth
developed in 1 patch on each side.

Body and head lacking scales. Lateral-line
tube large. Membrane over lateral-line modi-
fied into large scale-like structures, pierced
above and below by 2 pores on each segment.
Lateral-line segments partly ossified in double
concave form as in Lestidum. Greatest height
of each lateral-line segment variously at front
or hindmost point. Pectoral fin with 10-11
rays. Dorsal rays 11-14. Anal rays 24-28.

I have examined the following species of
Macroparalepis: M. danae Ege, M. brevis Ege,
and M. affine Ege. The genus also contains
Macroparalepis egei Maul. Generic type Ma-
croparalepis affine Ege.

Genus Stemonosudis Harry
Eigs. 19, 20

Macroparalepis (in part) Ege, 1933: 229.
Stemonostidis Harry, 1951: 32 (generic type by
original designation Macroparalepis inter-
media Ege) .

DIAGNOSIS: Body slightly compressed and
very elongate. Ventral carina on belly moder-
ately developed. Head and snout long and
pointed. Nostrils behind posterior tip of max-
illary in Stemonosudis intermedia. (Dr. Ege ex-
amined the types of Stemonosudis elongata, S.
gracile, and S. intermedia [his other species of
this genus are represented by postlarvae only]
and finds them also with the nostrils behind
posterior tip of maxillary.) Tip of lower jaw
not elevated or only slightly so. Upper jaw
terminating approximately 1 orbital diameter

242

PACIFIC SCIENCE, Vol. VII, April, 1953

Fig. 19. Genotype of Stemonosudis (S. intermedia (Ege), from off Bermuda, 125 mm. in standard length).

before anterior margin of eye. Angle of gape
near tip of maxillary. Supramaxillary short,
thread-like, closely bound to maxillary. Teeth
on upper jaw well- developed antrorse and re-
trorse canines. Vomer toothless. Teeth on
palatines short, in 2 rows anteriorly, 1 row de-
pressible; posteriorly few short teeth in single
row. Tongue very small, far back in mouth,
its anterior tip somewhat near angle of gape.
Gillrakers and pharyngobranchial teeth com-
pletely undeveloped on material examined.
(Dr. Ege informs me that the types of Ste-
monosiidis elongata, S. gracile, and S. intermedia
have "gillrakers present on the lower part
of the first gill arch, although very small,
their length equal to about a fourth of the
breadth of the bone of the gill arch.’’)

Body and head scaleless. Lateral-line tube
very large. Membrane over lateral-line tube

Fig. 20. Genotype of Stemonosudis (S. intermedia
(Ege), from off Bermuda, 125 mm. in standard length).
A, Anterior part of snout; B, enlarged section of teeth
on middle of premaxillary; C, anterior lateral-line seg-
ments; D, toothless first gill arch; E, dorsal surface of
tongue. (See explanation of morphological figures in
text.)

modified into large scale-like structures pierced
above and below by 2 pores on each segment.
Greatest height of each lateral-line segment
variously at front or hindmost point. Pectoral
fin with 10-13 rays. Dorsal rays 9-11. Anal
rays 37-50.

Generic type Stemonosudis intermedia (Ege) .
It is presumed that S. macrura (Ege), S. elegans
(Ege), S. elongata (Ege), and S. gracile (Ege)
belong in this genus, although I have been
unable to examine any of them, 5. intermedia
is from the West Indies; all other members
of the genus are East Indian. Further inves-
tigation might show that the East and West
Indian forms belong in separate genera or
subgenera.

Subfamily Sudinae

Head and lower jaw very large and massive.
Nostrils of equal size in young. Adults with
posterior nostril normal, anterior nostril re-
duced to tiny pore in anterior rim of other
nostril. Anterior process of premaxillary with-
out foramen. Teeth on mandible very large,
triangular in shape and strongly compressed;
edges finely serrate, not accompanied by
smaller teeth. Dorsal fin with 14-15 rays.
Pelvic fins with outer rays distinctly longer
than inner rays. Pectoral fin very long and
large, proportionately greatly enlarged in post-
larvae and juveniles. Body naked. Head scaled
in 2 series on preoperculum.

This group contains a single genus and
species, Sudis hyalina, known from the Medi-
terranean and Madeira.

Genus Sudis Rafinesque Schmaltz
Figs. 21, 22

Sudis Rafinesque Schmaltz, 1810: 60 (generic
type by monotypy Sudis hyalina Rafinesque
Schmaltz); Bonaparte, 1846: 35; Canestrini,
1877: 127; Ege, 1930: 6; Maul, 1945: 33;
Harry, 1951: 3.3.

Genera of Paralepididae — HARRY

243

Fig. 21. Genotype of Sudis (S. hyalina Rafinesque, from off Madeira, 315 mm. in standard length).

Paralepis (in part) Cocco, 1839: 49; Cams,

1893: 566; Sanzo, 1917: 1.

Siidis (in part) Parr, 1928: 34.

DIAGNOSIS: Body compressed and elongate.
Ventral carina on belly moderately developed.
Head large, strongly compressed. Snout very
large and deep. No nonossified prolongations
on tip of lower jaw. Nostrils situated almost
0.33 length of maxillary before posterior tip
of maxillary. Eye very large. Pupil vertically
oval, much larger than lens. Supramaxillary
large, more than 0.5 as long as maxillary,
splinter-like, closely bound to maxillary.
Teeth on lower jaw very large, in 1 or 2 series,
of triangular shape and strongly compressed;
edges minutely serrate. Few mandibular teeth
depressible. Teeth on premaxillary minute,
serrate, all fixed. Teeth on palatines moderate-
ly reduced, all fixed; anterior teeth in 2 rows
with long teeth sometimes accompanied by
short ones; anterior double series followed by
single series of short or minute teeth. Tongue
large, moderately far forward in mouth. Gill-
rakers tiny, spinous; each raker composed of
2-4 short teeth in single row; rakers not ex-
tending forward beyond angle of gape. Pha-
ryngobranchial teeth well developed in single
patch on each side.

True scales developed on preoperculum in
2 series. Otherwise body and head scaleless.
The lateral-line tube covered by single row
of distinct, overlapping, scale-like segments
lacking all trace of circuli or annuli. Each
segment pierced by 5-8 pores above and be-
low near upper and lower margins. Dorsal fin
with 14-16 rays. Vertebrae 59, as far as known.

FOSSIL HISTORY

The order Iniomi first appeared in the Cre-
taceous, in which the recent families Aulopo-

didae, Chlorophthalmidae, Myctophidae, and
probably Bathypteroidae are clearly evident.
The suborder Alepisauroidea ( = Paralepidoi-
dea Gill, 1893) appears to be represented in
the Cretaceous by the family Ichthyotringidae
(new family name to replace Rhinellidae Jor-
dan, 1905, because the monotypic genus
Rhinellus Agassiz, 1844, is preoccupied; Ich-
thyotringa Cope, 1878, is the next name avail-
able), and perhaps the Dercetidae also belong
here. Ichthyotringa resembles a Stidis with an
especially prolonged snout. The exact extent
of relationships are obscure, however, because
the head bones are inadequately known. The
Dercetidae, containing perhaps six genera (see
the list by Romer, 1945: 584), look remark-

Fig. 22. Genotype of Sudis (S. hyalina Rafinesque,
from off Madeira, 315 mm. in standard length). A, an-
terior part of snout; B, enlarged section of teeth on
middle of premaxillary; C, anterior lateral-line segments;
D, gillrakers on first arch; E, dorsal surface of tongue.
(See explanation of morphological figures in text.)

244

PACIFIC SCIENCE, Vol. VII, April, 1953

ably similar to an elongate paralepidid or
anotopterid, but again the head bones are
inadequately known, and the resemblance
may be only superficial.

The first definite representative of a recent
alepisauroid family appears in the Eocene on
the basis of the paralepidid genus Holosteus
Agassiz. Scopeloides Wettstein from the Oligo-
cene might be a scopelarchid. A fossil Alepi-
saurus has been discovered in the Miocene of
Europe. The genus Apateodus Woodward of
the Cretaceous family Enchodontidae has a
dentition similar to the Alepisauridae, and
several authors have suggested a close rela-
tionship.

The Paralepididae are the best known ini.
omous group found in the Tertiary, compris-
ing six fossil genera {Paralepis Cuvier, Tros-
!/////! Jordan, Lestkbthys ]ord3.a, DrmysJordd.n,
Holosteus Agassiz, and Parascopelus Sauvage)
and approximately 10 species. The references
to fossil members of the genus Sudis belong
with Paralepis. The paralepidids from the Mio-
cene diatom beds at Lompoc, California, and
Parascopelus from the Miocene of Europe ap-
pear to provide a complete intergradation
between this family and the Chlorophthalmi-
dae. Thus it would appear that the suborder
Alepisauroidea had not become fully differ-
entiated until relatively recently. This is cor-
roborated by the degree of relationships of
the recent forms. The family Paralepididae is
distinctly the most generalized group in the
suborder Alepisauroidea, and particularly the
genus Magnisudis shows a revealing number
of similarities to chlorophthalmids. The other
alepisauroid families (Alepisauridae, Anotop-
teridae, Scopelarchidae, Evermannellidae,
Omosudidae) are much more specialized and
probably were well differentiated by the begin-
ning of the Tertiary. The Dercetidae resemble
Anotopteridae in which the size of the dorsal
fin is between that of the Paralepididae and
the Alepisauridae. If there is a relationship
here, it indicates that these groups were high-
ly evolved even by the Upper Cretaceous.

Despite the fact that the Miocene fossils
seem to show no distinction between the
Chlorophthalmidae and Paralepididae, the
Eocene, Pliocene, and certain Miocene forms
are so similar to recent paralepidid genera
that only minor differences are evident in the
osteology.

Among recent groups the Paralepididae are
most closely related to the Anotopteridae.
They both have the same general proportions,
essentially similar osteology, the same pecu-
liar cartilaginous development of the jaws
which is found in these two families alone
in the order, and a good number of other
similarities. In turn, the Anotopteridae are
very closely related to the highly specialized
Alepisauridae. There are secondary modifica-
tions of the suspensorium that separate the
Anotopteridae from the Alepisauridae, but the
cranial osteology and dentition is very similar,
the skin has exactly the same pore system in
the adults of both families, which is found
only in the adults of these two families in the
order, and the proportions are closely similar.
Therefore, these three families appear to repre-
sent one line of evolution arising from a
chlorophthalmid-like ancestor, with the Ale-
pisauridae representing the most highly spe-
cialized end-point. On the basis of their
postlarval development and morphology the
Scopelarchidae, Evermannellidae, and Omo-
sudidae appear to have branched off from the
alepisaurid line of evolution in various direc-
tions and each represents its own unique end-
point of evolution. Among these three fami-
lies the Scopelarchidae are closest to the
Chlorophthalmidae, and the Omosudidae are
most closely related to the Alepisauridae. !

The Alepisauroidea seems to be a valid
suborder despite the fact that at least one of
its families (Paralepididae) may have become i
separated from the Chlorophthalmidae of the
suborder Myctophoidea (= Aulopodoidea
Gill, 1893) as late as the Miocene or Pliocene, j
In fact, future research may very well reveal ^
intermediate recent forms between the chlo- !
rophthalmid line and the Paralepididae. If this 1

Genera of Paralepididae — Harry

245

happens, the retention of the Alepisauroidea
should be still justifiable because the greatest
phylogenetic break in this suborder is between
the Paralepididae and the other families.

The earliest known paralepidid, Holosteus
esocinus Agassiz (1844: 85, PL 43, Fig. 5) ap-
pears to be very close to the recent genus
Lestidium and may actually belong in it. The
dorsal fin seems to have been pushed back
out of position during fossilization, and it
appears from the illustration that it should
be approximately equidistant between verti-
cals from the ventral and anal fins.

The most primitive fossil paralepidid seems
to \y^Drimys defemor]oidi2.x\ (1925: 14, PI. 4b),
the clearest intermediate stage yet found be-
tween the Chlorophthalmidae and Paralepidi-
dae. Unfortunately the only known Miocene
fossil is badly crushed. There are 45 vertebrae,
a number considerably lower than known for
any recent paralepidid. The presence of large,
coarsely striated scales is more like a chlo-
rophthalmid, whereas the description of the
dentition of the upper jaw is very much like
that of a paralepidid.

T r ossulus exoletus (1921: 250, Pis. 8c,
28a) is another Miocene fossil from the Lom-
poc diatom beds. In general appearance it
seems to be a generalized paralepidid with a
foreshortened body. It shows closest similari-
ties to Magnisudis but actually may be an
intermediate form between a chlorophthalmid
and a paralepidid. The eye is small, the last
dorsal rays are over the anterior anal rays.
Of particular significance is the fact that the
maxillary is narrow and the premaxillary forms
the margin of the upper jaw. There are 50
vertebrae, a number most closely approaching
the genera Paralepis and Magnisudis.

The fossil Lestichthys porteousi ]oM2in (1921:
250, Pis. lOb and 22b) from the Lompoc
Miocene diatom beds appears to be very close
to the genus Lestidium. The vertebral count
of 86, proportions, and osteological notes
seem to place it in the tribe Lestidiini of the
subfamily Paralepidinae.

One of the best known fossils is Paralepis
alhyi Sauvage ( = Paralepis sphekodes) from the
Pliocene. This form is very ably reviewed by
Arambourg (1927), who cites the scattered
literature and gives sketches of the jaw bones
and scapula.

The genus Parascopelus Sauvage appears to
belong in the Paralepididae near the genus
Sudis, but the position of the ventral fins far
forward is unique in the family.

SUMMARY

The family Paralepididae comprises seven
genera {Paralepis, Notolepis, Magnisudis, Lesti-
dium, Macroparalepis, Stemonosudis , Sudis) and
48 living species. It is differentiated from all
other families of the order Iniomi by a charac-
teristic toothless emargination in the upper
jaw below the end of the snout, by a large
foramen through the premaxillary in all gen-
era except Sudis, by the rictus being well be-
fore the eye, and by the position of the dorsal
fin which is near the middle of the body
length far behind the pectoral fins. The Para-
lepididae are regarded as most closely related
to the Anotopteridae and Alepisauridae of the
suborder Alepisauroidea.

The Paralepididae are divided into two sub-
families—the Paralepidinae and Sudinae. The
former is characterized by the presence of a
foramen in the anterior process of the pre-
maxillary, by slender smooth teeth, by the
inner pelvic rays being longer than the outer,
and by the pectoral fins remaining small
throughout ontogeny. In the Sudinae the pre-
maxillary lacks a foramen, the mandibular
teeth are broad and serrate, the outer pelvic
rays are longer than the inner rays, and the
pectoral fins are greatly prolonged during
early postembryonic stages. The subfamily
Sudinae is monotypic, containing the genus
Sudis which is known from the Mediterranean
and Madeira.

Evolution in the family generally appears
to have progressed by losses of pre-existing
characters. This is particularly evident with
such structures as squamation, gillteeth, pig-

246

PACIFIC SCIENCE, Vol. VII, April, 1953

mentation, and massiveness of form. In this
regard the subfamily Paralepidinae has two
basic divisions. The members of one division,
comprising the genera Paralepis, Magnisudis
(described herein as new), and Notolepis, are
scaled as adults and have numerous teeth in
two or more rows on each gillraker; the gene-
ra of the other division {Lestidium, Macropara-
lepis, and Stemonosudis) are completely scaleless
and have few teeth in one row on each gill-
raker. This classification cuts across the sys-
tem currently recognized in the literature.

The new genus Magnisudis appears to be
the most primitive in the family and shows
the closest link back to the Chlorophthalmi-
dae and Aulopidae, the most generalized
families in the order Iniomi. The genus Noto-
lepis has many unique characteristics and rep-
resents the end point in scaled genera evolu-
tion, approaching more closely the scaleless
genera than do either Paralepis or Magnisudis.
The genus Lestidium appears to be the most
generalized of the scaleless genera and clearly
approaches the scaled genera more closely
than either Macroparalepis or Stemonosudis.

The fishes of the family Paralepididae are
world-wide in distribution, except for the
Arctic. The genus Notolepis is bipolar (pan-
temperate) in distribution. One of its sub-
genera {Notolepis) is known from the Ant-
arctic and the other {Profundisudis) occurs in
the North Pacific and North Atlantic. Most
of the other genera are also wide-ranging.

The suborder Alepisauroidea appears to be
represented in the Cretaceous by the family
Ichthyotringidae (new name to replace Rhi-
nellidae Jordan, 1905) and perhaps by the
Dercetidae. Six fossil genera {Paralepis Cuvier,
Trossulus Jordan, Lestkhthys Jordan, Drimys
Jordan, Holosteus Agassiz, and Parascopelus
Sauvage) and approximately 10 species of the
Paralepididae are known from the Tertiary.
Even as recently as the Miocene there seems
to be intergradation between the Chloroph-
thalmidae and Paralepididae, and as a result
the Alepisauroidea became fully differentiated

from the suborder Myctophoidea relatively

recently.

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Berg, Leo Shemenovich. 1940. Classifica-
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PACIFIC SCIENCE, Vol. VII, April, 1953

1893. Families and subfamilies of

fishes. Natl. Acad. Scl, Mem. 6: 125-138.

Goode, George Brown, and Tarleton
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NOTES

Faunal Spedation in New Georgia, Solomon Islands

Some 16 years ago the writer made certain
contributions to the knowledge of the fauna
of San Cristobal, the most eastern of the main
Solomon Islands (Lever, Limn. Soc. London,
Proc. 148(2): 82-83, 1936); this was followed
by more general notes on the whole group
(Lever, Trop. Agr. [Trinidad] 20(2): 40-42,
1943) . The purpose of the present paper is to
draw attention to an extensive local develop-
ment of species formation in the New Geor-
gia group of islands lying in the centre of the
whole Solomon archipelago.

Besides the main island of New Georgia,
which is some 50 miles in length, the group
comprises seven closely associated islands in-
cluding Kolombangara which, though only
15 miles in diameter, rises to a height of over
5,500 feet. All these islands, despite their near-
ness to one another, are separated by very
much deeper water than any other comparable
group in Melanesia (Fig. 1). Thus, although
Kolombangara is only 10 miles from Gizo,
the intervening sea is 245 fathoms deep, and
soundings between Rendova and Wana Wana,
which are only 12 miles apart, reached 552
fathoms.

Recognition of an inter-insular variation in
the animals of these islands was first noted
in the birds by Mayr {Amer. Mus. Novitates
522: 1-22, 1932), who showed that while the
thick-head Pachycephala pectoralis is represent-
ed by a different species in each of the larger
Solomon Islands, in New Georgia alone are
several species found. Later he showed that
specimens of the white eye, Zosterops rendovae,
from Vella Lavella and Bagga were distinct
from those occurring in Ganongga and Gizo
to the south and Kolombangara to the south-
east {Amer. Nat. 74: 249-278, 1940). In fact
the flying time of 3.4 minutes which these

Fig. 1. Map of the New Georgia Group, Solomon
Islands. The outlines of this map have been taken large-
ly from United States Army maps, as the most recent
official British map (No. 17/30 of Directorate of Colo-
nial Surveys) shows Arundel Island as a portion of New
Georgia although in fact a deep-water channel occurs
between the former and the intervening Woodford
Island, with a constriction at the Diamond Narrows.
The D.C.G. map marks, but leaves unnamed, the large
island of Vella Lavella; the spot levels have, however,
been followed here. The most recent edition (1945)
of the British Admiralty chart No. 2392 indicates only
tentative outlines for the coast of western Vella Lavella
and adds that Gananggo may be 2 to 4 miles further
west than its marked position. (Altitude in feet, sea
depth in fathoms.)

small birds take to fly 1.7 kilometers would
have enabled them to bridge the gap which
however, is sufficient to allow of subspecies
occurring on the different islands.

That this same phenomenon occurs in in-
sects has not heretofore been appreciated.
However, well-marked subspecies of the wee-
vil Exophthaimida coerulescens Gunther, taken
by the writer in 1936, were shown to exist by
Marshall {Roy. Ent. Soc. London, Proc., Ser. B.
10(4): 51-55, 1941). The typical form, from
Vella Lavella, is opalescent grey with many
black spots whereas E. c. ganongae is metallic
green, and E. c. kolombangarae is fawn with
small dark spots. A further point of interest

250

NOTES

251

is that, although much more intensive col-
lecting has been done on the more easterly
islands of Guadalcanal, Isabel, San Cristobal,
and Malaita, this conspicuous metallic weevil
was never taken on any of them.

This local endemism, which is probably of
relatively recent origin, is not comparable
with the inter-insular variation recorded by
Marshall (Roy. Ent. Soc. London, Trans. 87(3):
69~101, 1938) for another genus of weevils
{Elytrufus) from Fiji and the New Hebrides or
various other insects found in the Marquesas
(Mumford and Adamson, Congr. Internatl.
Ent., Paris: 431-450, 1933) , as in these cases
the islands are separated by varying but ex-
tensive distances of open sea.

Among the Lepidoptera one extreme case
of aberration of a bird-winged butterfly can
be cited, viz., Troides {Ornithoptera) victoriae
ruhianus Roths. Among a series from Ganong-
ga [published as Ranonga], one male had an
unusually large green area on the forewing,
other males showing a less extensive develop-
ment (Dicksee, Roy. Ent. Soc. London, Proc.
8: 144, 1934). As was remarked when these
specimens were exhibited at a meeting of the
Royal Entomological Society, it is strange
that this Ganongga race should be so very
variable as specimens from other islands near-
by seemed to be constant in wing pattern.
[The distribution of this butterfly is given in

News

Several meetings of general importance were
held in Honolulu during the week of February
16-21.

The American Institute of Biological
Sciences' Advisory Panel on Biology to the
Office of Naval Research held its spring meet-
ing at the University of Hawaii February 18
and 19. In conjunction with this meeting the
Hydrobiology Committee of the A.IB.S. met
at the University on February 20.

The annual meeting of the Invertebrate Con-
sultants Committee for the Pacific, of the
Pacific Science Board of the National Research

Seitz {Macrolepidoptera of the World, vol. 9: 12,
1927) as being Kolombangara Island and Ru-
biana, the latter (also spelled Roviana) being
used both for a village on the west of New
Georgia and an island lying to the south of
the main island.]

Although the bulk of the writer's collec-
tions of anisopterous Odonata were wide-
ranging forms, a handsome new species taken
in 1936, Agriocnemis salamonis Lieftinck (Treu-
bia 20(2): 319-374, 1949), has recently been
described from Ganongga, and it seems likely
that adjacent islands will yield more new
material from this neglected order.

As few classes of animals have been inten-
sively collected in this area, it is probable that
many interesting varieties remain to be dis-
covered although probably not in the Amphi-
bia and Reptilia. These were thoroughly col-
lected and reported on by the Burts {Amer.
Mus. Nat. Hist., Bui. 63(5): 462-597). Only
one snake {Denisonia woodfordii (Boul.)) is
confined to New Georgia, and it shows no
inter-insular variation.

As the writer is no longer stationed in
Melanesia, he has thought it desirable to pre-
sent these few observations and thus draw
attention to the intensive endemism to be
found in this relatively small but very interest-
ing area. — R. J. A. W. Lever, Department of
Agriculture, Kuala Lumpur, Malaya.

Notes

Council, was held at the Experiment Station
of the Sugar Planters’ Experiment Station,
February 18 and 19. L. J. Dumbleton, Plant
and Animal Quarantine Officer of the South
Pacific Commission, stationed in Noumea, was
an observer at the meeting.

Book Notice

Ekman, Sven. 1953. Zoogeography of the Sea.
xiv+417 pp., 121 figs. Sidgwick and Jackson
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York. $6.50. A revised English edition of a
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Crawford, David L. 1920^^. New or interesting
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News 51 (1): 12-14.

1920^. Cerotrioza (Psyllidae, Homop-

tera). Hawaii. Ent. Soc., Proc, 4 (2): 374-
375.

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E4CIFIC

SCIENCE

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

IN THIS ISSUE: Matthews — Development of Pedunculate
Spermatophore of Dardanus asper • McCracken — Prelim-
inary Report on Soils of Saipan • Tseu — Seasonal
Variations in Physical Environment of Marine Ponds •
Glassman — New Plant Records from Eastern Caroline Is-
lands • Belkin — Mosquitoes of Genus Uranotaenia in
Solomon Islands • NOTES

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PACIFIC SCIENCE

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

VOL. VII JULY, 1953 NO. 3

Previous issue published April 30, 1953

CONTENTS

PAGE

The Development of the Pedunculate Spermatophore of a Hermit Crab,

Dardanus asper (DeHaan). Donald C. Matthews 255

A Preliminary Report on the Soils of Saipan, Mariana Islands.

Ralph J. McCracken 267

Seasonal Variations in the Physical Environment of the Ponds at the Hawaii
Marine Laboratory and the Adjacent Waters of Kaneohe Bay, Oahu.

Winifred Shut Lin Tseu 278

New Plant Records from Eastern Caroline Islands, with a Comparative Study

of the Native Plant Names. S. F. Glassman 291

Mosquitoes of the Genus Uranotaenia in the Solomon Islands (Diptera:

Culicidae). John N. Belkin 312

NOTES:

On the Establishment of the Oahu Research Center of the University of

California. Clarence E. Palmer 392

News Note . 395

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The Development of the Pedunculate
Spermatophore of a Hermit Crab,
Dardanus as per (De Haan)^

Donald C. Matthews^

Although in a few pagurian genera {Ana-
pagurus, Clibanarius, Diogenes, Eupagurus, Pa-
gurus) the morphological, physiological, and
mechanical phenomena associated with the
elaboration of the pedunculate spermato-
phores have been thoroughly investigated,
in most pagurian genera these phenomena
have been completely neglected.

This paper extends the knowledge of pe-
dunculate spermatophore elaboration and
compares the process in Dardanus asper (De
Haan) with that of other species.

The literature pertinent to the study of the
development of pedunculate spermatophores
is adequately covered in Mouchet’s (1931)
bibliography. In so far as can be ascertained,
no published work on the development of the
spermatophore of Dardanus asper has been
reported.

METHODS AND TECHNIQUES

Specimens of D. asper (Fig. Ic) obtained
from the Honolulu Aquarium between March,
1949, and November, 1952, were used in this
study. These were transported in sea water
to the laboratory at the University of Hawaii
and used immediately. Cutting the apices of
the covering shells {Tonna perdix) (Fig. lb)
caused the hermit crabs to abandon them
quickly. The crabs were then seized, their
nerve cords severed, and the dorsal surface
of their abdominal wall removed.

The exposed reproductive systems (Fig. 2c,
d) were then freed of the blood vessels and
connective tissues which encompassed them.
Any macroscopic effects of the asymmetrical-
ly placed viscera on the reproductive systems

^ Contribution No. 32, Hawaii Marine Laboratory.

2 Department of Zoology and Entomology, Uni-
versity of Hawaii, Honolulu, Hawaii. Manuscript re-
ceived May 16, 1952.

were recorded. Small portions (about 0.5 cm.)
of the distal vasa deferentia (Fig. Aa) of both
the right and left sides were removed and
placed immediately in fixative. A label indi-
cating specimen number and side was inserted
in each vial so that any dimorphism of the
spermatophores could be correlated with that
side of the reproductive system most affected
by the asymmetric abdomen. The remaining
portions of the right and left vasa deferentia
and testes were then freed, that of one side
was placed in fixative for future sectioning,
and the other was dissected. In this manner
both right and left testes and right and left
vasa deferentia were alternately placed in fix-
ative and alternately dissected.

The testes and vasa deferentia were placed
in Bouin’s fixative, cleared in toluene, em-
bedded in Tissuemat (54-56°C.) and were
serially sectioned at 10 microns. The sections
were stained with standard alum-haematoxy-
lin and counterstained with eosin (0.5 per
cent solution in 90 per cent alcohol to which
4.0 cc. of O.lN HCl was added).

Other living right and left vasa deferentia
were injected with neutral red (Ehrlich) so
that the effects of the muscular contractions
on the sperm mass could be more clearly
discerned.

All figures were drawn by Evan Gillespie
from dissections or from slides prepared by
the author.

My appreciation is extended to Mr. Spencer
Tinker, Director of the Honolulu Aquarium,
who generously supplied the specimens.

DISCUSSION

The abdomen of D. asper (Fig. 2a), like that
of most hermit crabs, exhibits an asymmetri-
cal arrangement of its viscera upon dissection.

255

256

Fig. 1. Dardanus asper (DeHaan) (c) in a shell of Tonna perdix {b) encrusted with anemones, Calliacth armillatas
{a). (0.8 X.)

■Hi

5.

PACIFIC SCIENCE, VoL VII, July, 1953

The large hepatopancreas (Fig. 2b), in follow-
ing the clockwise coil of the abdomen, carries
with it the testes (Fig. 2c). Although micro-
scopic dissection is necessary to free the testes
of the connective tissue and blood vessels
which bind one to the other, they are sep-
arate organs not joined by a transverse bridge.

The dissected testis (Fig. 3) discloses nu-
merous sacculi {a) through whose thin walls
sperm-forming elements {b) are clearly dis-
cernible. Each sacculus contributes its portion
to the continuous sperm mass {c) which
traverses the length of the testis (/) . Although
differences in the size of the testes are ob-
served, the function of the sacculi is not im-
paired. Both right and left testes of all the
specimens examined during the course of this
study were actively producing spermatozoa.
There is no sexual season in the male.

Differences in the size of the testes are
usually accompanied by differences in the
size of the vasa deferentia. However, when
right and left vasa deferentia are carefully dis-
sected, both consistently reveal the same fun-
damental regions (Figs. 4, 5). The distal por-
tion of both right and left vasa deferentia
(Fig. Aa) are always distended with sperma-
tophores. I have found no evidence of para-
sitism or dimorphism of the spermatophores.

Although in tracing the development of
the spermatophore reference is made to speci-
fic cross sections (Figs. 6, 7, 8, 9, 12, 13, 14),
serial sections reveal that the morphological
changes of the vas deferens are gradual and
that these gradual morphological changes are
paralleled by physiological changes.

These changes will be discussed as the
course of spermatophore development is
traced throughout the vas deferens.

Fig. 2. Dorsal view of dissected D. asper. a. Asymmetrical abdomen; b, hepatopancreas; c, testes; d, left vas
deferens; intestine. (0.6X.)

Spermatophore of Dardanus asper — Matthews

Morphology of Living Vas Deferens
as Revealed by Dissection

From the comparative morphological study
of hermit crab vasa deferentia, Mouchet (1931)
concludes that spermatophores with a pedes-
tal and a veil are elaborated in those vasa
deferentia which possess two helices. The
pedunculate spermatophore of D. asper (Fig.
15) which possesses a pedestal {e) and a veil
{d) should, therefore, owe its origin to a vas
deferens with two helices. Mouchet {op. cit.)
also states that the examination of the external
form of the hermit crab vas deferens allows
one to predict the principal characteristics of
the spermatophore that it produces and also
to know the method of fragmentation of the
sperm column at the moment of its trans-

formation into successive ampullae. She fur-
ther states that in all hermit crabs it is at the
exact point of change of curvature of the two
consecutive helices that the continuous flow
of sperm is fragmented either into arches or
into successive rectilinear fragments.

For those hermit crabs whose vasa deferen-
tia offer little complexity, it may be possible
to predict the principal characteristics of the
spermatophore and to determine the method
of fragmentation of the sperm column, but
such is not the case in the highly complex
vasa deferentia of D. asper. The enlarged por-
tion of the vas deferens of D. asper (Fig. 5)
reveals that the curvatures (c, d, e) change
direction three times before the compact left-
handed helix (g) is encountered. These coils
are referred to as right-handed or left-handed

258

Fig. 3. Posterior portion of the testes, a, Sacculi;
h, sperm-forming elements; c, continuous sperm mass;

posterior portion of left testis; e, indeterminate por-
tion of left vas deferens; /, right testis; g, portion of the
right vas deferens. (4.5 X; insert lOX.)

in relation to the course traversed by the sperm
flow; hence the curvatures at c are right-
handed, at d left-handed, at e right-handed,
and at g left-handed. The region c-e, with its
changes in curvature, was at first included in
the indiscriminate region h which emanates
from the testis. This interpretation must be
abandoned as neutral red, injected into the
enlarged, distal portion of the vas deferens
(Fig. Ad), permeates the regions e, d, c (Fig.
5) and reveals a continuous sperm column
already encased in its sheath. The change of
curvature id) should mark the^ exact location
of the fragmentation of the sperm column if
Mouchet’s thesis is correct. As this fragmen-
tation does not occur until the compact, left-
handed coils (g) are encountered, Mouchet’s
assignation of the region of sperm-column
sheath formation solely to the right-handed
coils of the first helix (c) fails to account for
any activity in curvatures (d, e). Obviously,
sheath formation and fragmentation are not
limited to two consecutive helices in D. asper.

PACIFIC SCIENCE, VoL VII, July, 1953

The utilization of the dissecting microscope
serves not only to reveal the nature of the
muscular contractions but also enables one
to correlate the changing morphology of the
lumen with the complicated molding of the
spermatophore.

Cognizant of the arrangement of the ob-
lique muscular fibers in the wall of the vas
deferens, one might expect to observe peris-
taltic waves traversing the tube from the testis
distad to the genital pore. These contractions
would serve to force the homogeneous sperm
mass through a series of ever-changing dies,
and, aided by secretions from the epithelial
lining of the wall, the heterogeneous sperma-
tophore would evolve. Actual observations,
however, prove this supposition to be some-
what inaccurate although correct in principle.

The entire vas deferens, with the possible
exception of the enlarged distal portion (Fig.
Ad), exhibits isolated, intermittent contrac-
tions, which persist for well over an hour,
even though connective tissue, blood vessels,
and nerves are severed. These spasmodic con-
tractions might possibly be attributed to the
austere dissection had not this activity been
observed in superficial regions of the vas def-
erens prior to dissection. These muscular con-
tractions are occasionally simple, sphincter-
like twitches; however, at times they seem to
result in a lateral compression of the wail and
at other times in a dorsoventral flattening.

Fig. 4. Dissection of the right testis and vas deferens.
a, Enlarged distal portion of the vas deferens; b, sinuous
portion of the vas deferens; c, posterior portion of the
testis. (1.5 X.)

Spermatophore of Dardanm asper — MATTHEWS

259

Fig. 5. Dissection of right vas deferens showing exact nature of the coils, a, Posterior region of right testis;
b, (1, 2, 3) indeterminate region; c, first right-handed coil; d, first left-handed coil; e, second right-handed coil;
f, region of fragmentation of the sperm column; g, compact left-handed helix; h, flat spiral; /. enlarged, highly-
contorted region. (6X.)

Even when the internal mass is vitally stained,
the result of these many and varied contrac-
tions on the formation of the spermatophore
is difficult to perceive. However, the internal
mass, regardless of what portion of the vas
deferens is observed, appears to respond to
the ever-changing, restless wall. The com-
bined effect seems not so much to move the
mass along as to mold it to an internal die.
If the activity which is seen in the altered vas
deferens approximates the activity which is
obscure in the unaltered vas deferens, sperma-
tophore formation in D. asper is a slow and
complicated process. During the course of an
hour the internal mass rarely moves more than
a few millimeters. This mechanical activity,
brought about by contraction of the muscular
fibers, must be correlated with the morpho-
logy and physiology of the vas deferens if
the process of spermatophore formation is to
be understood.

Correlation between Morphology,

Physiology of Epithelial Cells, and
Contractions of Muscular Walls
of the Vas Deferens

The region of the vas deferens (Fig. ‘5bl)
which receives the sperm from the testis pre-
sents in cross section (Fig. 6) a tube whose
muscle layer {a), epithelial layer {b), and lu-
men (c) are little specialized. The muscle wall

is thin. The cuboidal epithelium is evenly
distributed around a cylindrical lumen which
contains loose clusters of spermatozoa {d).
The cuboidal epithelium secretes a substance
{e) which enters the lumen and mixes freely
with the spermatozoa. Whether this substance
serves as a nutrient or as a lubricant was not
ascertained. The muscular activity, revealed
by the dissecting microscope, serves not only
to move the sperm mass and to mix it with
the epithelial secretion but also to mold the
mass in compliance with the cylindrical lu-
men.

Gradually the internal morphology of the
vas deferens changes. From the indeterminate
region (Fig. 5^2) a cross section (Fig. 7) re-
veals that the cuboidal epithelium has given
way to columnar epithelium (b) except at two

Fig. 6. Cross section through the indeterminate re-
gion of the vas deferens (Fig. 5^i). a. Muscular layer;
b, cuboidal epithelium; c, cylindrical lumen; d, loose
clusters of spermatozoa; e, secretion produced by epi-
thelial cells. (54 X.)

260

PACIFIC SCIENCE, VoL VII, July, 1953

C

Fig. 7. Cross section through the indeterminate re-
gion of the vas deferens (Fig. 5^2), a. Muscular layer;
b, columnar epithelium; c, d, regions where cuboidal
epithelium still persists; e, sperm column; /, elliptical
lumen; gS secretion produced by epithelial cells.
(54X.)

regions {c, d) which lie directly opposite each
other. This change results in an elliptical lu-
men (/) which in the more distal region (Fig.
5^3) gradually increases both in width and
in height. The columnar epithelial cells {b)
appear to remain unaltered. Their secretion
(gb continues but serves less to mingle with
the spermatozoa than to remain superficially
around the more elliptical and compact sperm
mass {e). The cuboidal cells (c, d) have an
affinity for the nuclear instead of the cyto-
plasmic stain. The activity of the muscle wall
{a) serves to move the sperm mass and to
mold it in compliance with the elliptical lu-
men. There is no evidence as yet of a sperm-
column sheath.

From a region still more distad (Fig. 5c)
a cross section (Fig. 8) reveals that the morph-
ological change in the epithelial layer is even
more pronounced. The cuboidal epithelial
cells (c, d) which take up the nuclear stain,
now seem isolated between diagonally placed
columnar cells {h, i and/, k), which likewise
take up the nuclear stain. These diagonally
placed cells produce a new secretion (g^),
which flows over and covers the distal ends

of the epithelial cells {b) bounding the ellip-
tical lumen (/).

From the region slightly more distad (Fig.
5d) a cross section (Fig. 9) reveals that this
secretion (Fig. Sg^) now lies contiguous with
the sperm column. This is the sperm-column
sheath (Fig. 9/). The phenomena associated
with this change of position are obscure but
may exist because (1) the sperm column {d),
due to an increase in size, now presses against
the sheath-forming material (/) ; (2) the sheath-
forming material (/) is freed from the edge of
the epithelium {b) by a secretion from these
cells; or (3) the contractions of the muscular
wall {a) force the sheath-forming material
(Fig. 8g^) into contact with the more viscous
sperm column (Fig. 9d).

Mouchet {op. cit.), in describing the sperm-
column sheath formation for Eupagurus bern-
hardus, states:

At the two points of maximum depression
of the arch [equivalent to areas c and d of my
figure 8], grooves take shape, and in these

h e t

Fig. 8. Cross section through region c of the vas
deferens (Fig. 5). Muscular layer; b, columnar epi-
thelium; c, d, regions where cuboidal epithelium still
persists; e, sperm column;/, elliptical lumen; g^, sheath
secretion produced by epithelial cells h, i and /, k.
(54X.)

Spermatophore of Dardanus asper — MATTHEWS

i c h

Fig. 9. Cross section through region d of the vas
deferens (Fig. 5). a. Muscular layer; b, columnar epi-
thelium; c, cuboidal epithelium; d, sperm column;
e, lumen; /, sperm-column sheath; g, h, diagonally
placed epithelial cells. (54 X.)

grooves a new secretion accumulates, one which
lines the interior of the canal and which identi-
fies itself with its basic affinities. . . . Then a
continuous cylinder encloses the sperm and it
presents two little lateral swellings which seem
to glide about in the little grooves ... in the
last turn of the first spindle one of the grooves
develops greatly while the other disappears.

It is interesting to note that, although the
method of sperm-column sheath formation
is quite similar to that of D. asper, neither
cross sections nor longitudinal sections
through this region reveal any lateral swell-
ings associated with the grooves. By the time
the sperm column, now enclosed in its sheath,
has advanced as far distad as region e (Fig. 5),
morphological changes of the epithelial cells
result in the formation of a pear-shaped lu-
men. The change in lumen shape, however,
is begun in the preceding region (Fig. "^d)
and can be best explained from a cross section
through this region (Fig. 9). The sperm col-
umn {d), now enclosed in its sheath (/), no
longer occupies the center of the lumen {e)
but is nearer one end. At this end the cu-

261

boidal cells (c) and the diagonally placed
columnar cells (g, h) persist, but those at the
opposite end of the lumen (Fig. ^d, /, k)
disappear. The only significant difference ob-
served between regions d and e (Fig. 5) is the
shape of the lumen.

When the living vas deferens is injected
with neutral red and observed under the dis-
secting microscope, the sperm column with
its sheath appears, in the last right-handed
coils (Fig. 5^), to be continuous, whereas in
the left-handed coils (Fig. 5g) it appears to
be discontinuous. Interest, therefore, was fo-
cused on the region of the vas deferens re-
sponsible for the change (Fig. 5/). Mouchet
{op. cit.) describes this differentiation in Dio-
genes pugilator thus :

Everything happens as if the incurved column
of sperm flow, molded by its passage in the
first spiral, were maintaining its curving until
reaching the second spiral, which is rolled up
inversely. Encountering then the wall of the
canal on its internal concave face, it hits against
it and by successive deflections, describes arcs
whose curve is contrary to the canal which con-
tains them. Each point of deflection becomes
the extremity of an ampulla in which the sperma-
tozoids of two neighboring half-arches come to
accumulate. The slender base of the ampule is
formed by the union two by two of the extremi-
ties of the arches.

This description is not in accord with the
observed process of arch formation in D.
asper. Likewise, Mouchet’s {op. cit.) descrip-
tion of the modifications of the process in
Clibanarius misanthropus, Eupagurus hernhardus,
E. prideauxi, E. cuanensis, and E. hyndmanni
fails to agree with the process in D. asper.

The muscular contractions at this region
(Fig. 5/) appear not unlike those of other
regions. Both dorsoventral flattening and lat-
eral compression of the region are observed.
This region also shortens and lengthens. The
over-all effect of these contractions on the
sperm, column is comparable to that produced
by a tucker attachment of a sewing machine.
The continuous sperm column, enclosed in
its sheath, arrives at this region and, by the

262

PACIFIC SCIENCE, Vol. VII, July, 1953

Fig. 10. Reconstruction of the vas deferens through region g (Fig. 5). a, Ampulla of sperm forming as arch;
b, pear-shaped sperm column; c, connecting sperm-column sheath. (55 X.)

contractions of the muscular wall, undulates
into continuous sinusoidal curves.

Because of the compact coils of the left-
handed helix (Fig. 5g), sections parallel to
the longitudinal axis of the lumen and at the
same time through the plane c-d (Fig. 8) are
difficult to obtain. Sufficient portions are avail-
able, however, to permit a fairly accurate re-
construction (Fig. 10). Although the sperm

column appears to be segmented into sep-
arate and distinct arches, when viewed through
the wall of the vas deferens (Fig 5g), these
arches are in reality joined one to the other.
This obvious discrepancy is easily explained.
Throughout the compact, left-handed coils
of this region (Fig. 5g) the wider portion of
the pear-shaped lumen occupies a position
nearer the outer edges of the coils, whereas

Fig. 11. Reconstruction of the contents of the vas deferens through regions g, h (Fig. 5). a. Continuous thread
of stalk material; b, c, d, e, stalk-forming material accumulating between the closing arches. (55 X.)

Spermatophore of Dardanus asper — Matthews

263

Fig. 12. Cross section through region h of the vas
deferens (Fig. 5). a. Stalk-forming epithelium; b, stalk;
c, deepened groove of lumen; d, pear-shaped ampulla,
(55X.)

the thinner portion of the pear-shaped lumen
occupies a position nearer the inner edges of
the coils. The sperm column is molded in
compliance with the pear-shaped lumen by
muscular contractions of the walls of the vas
deferens. When the sperm column is viewed
through the wall of the vas deferens, only the
thicker portion (Fig. lOb) is visible, hence the
sperm column appears to be segmented. Each
future ampulla of sperm is composed of an
arch (10^^). These are connected by the sperm-
column sheath (10c). The fate of the connect-
ing sheaths is shown later.

As the last compact coils of the left-handed
helix (Fig. 5g) are traversed and the region
of the flattened spiral (5^) is entered, the
groove of the pear-shaped lumen deepens.
The activity of the epithelial cells at the base
of this groove now produces a new secretion.
This at first is thread-like, but in the more
distal portion of the flat spiral (Fig. ^h) thick-
enings appear. The contents of the dissected,
flat spiral which are shown in Figure 11 reveal
that this secretion {a) accumulates at definite
regions {b, c, d, e) between the closing arches.
The thickenings form short, blunt stumps
{d, e) which are the precursors of the stalks
(Fig. 15c). The importance of the connecting
sheath (Fig. 10c) is now apparent. As the
closing arches move distad in the deep groove
of the vas deferens, the stalk-forming material
accumulates only in the regions between the
closing arches; the connecting sheaths be-

Fig. 13. Cross section through region i of the vas
deferens (Fig. 5). a, Stalk-forming epithelium; b, stalk;
c, veil-producing epithelium; d, veil-producing secre-
tion; e, ampulla of sperm. (55 X.)

tween adjacent ampullae prevent the stalk-
forming material entering the lumen. By mus-
cular contraction of the walls of the vas
deferens, the arches finally close, but not until
the stalk-forming material has^ accumulated
between them.

In the region distad to the flattened spiral
(Fig. 5/), a cross section (Fig. 12) reveals that
the stalk (Jb) in the deep groove of the lumen
(c) lengthens both by continued secretion by
the epithelial cells {a) and by the muscular
activity of the walls of the vas deferens.

The lengthening of the stalks carries the
ampullae of sperm ” aloft,” and the connecting
sperm-column sheaths between adjacent am-
pullae become extremely thin and finally ob-
scure.

As this lengthening process continues, a
more distal cross section (Fig. 13) reveals that
the epithelial cells (c), which line the deepened
groove of the lumen, produce still another

264

PACIFIC SCIENCE, Vol. VII, July, 1953

secretion (d). This secretion is the precursor
of the veil (Fig. X’bd). As the vas deferens is
traversed, this secretion, molded by the lateral
contractions of the muscular wall, surrounds
the stalks (Figs. 13^, 15c). The secretion
from the diagonally placed epithelial cells at
the base of the deep groove forms the viscous
pedestal (Figs. 14^, 15c).

The deep groove, so characteristic of the
lumen, is no longer present in the enlarged,
apical portion of the vas deferens (Fig. Ad).
For a considerable distance the vas deferens
presents a cylindrical lumen. Gradually two
folds appear in the epithelium. From these
typhlosole-like folds (Fig. 14c) a new secre-
tion (/) encompasses the completed sperma-
tophores.

Conspicuous longitudinal muscles (Fig.
14^) serve to move the completed sperma-
tophores toward the genital pore. When the
spermatophoric mass is first extruded, it is
difficult to distinguish the pedestal from the
veil. It is the viscous pedestal, however, which
allows the spermatophore to become attached;
the veil is not sticky.

Neither the pedestal nor the veil is seg-
mented, The spermatophore of D. asper is,

Fig. 14, Cross section through the apical region of
the vas deferens (Fig. Ad), a, Longitudinal muscle layer;
b, pedestal of a single stalk; c, "typhlosole”; d, portion
of a stalk; e, ampulla of sperm; /, mucus. (42 X.)

a b

Fig. 15. The completed spermatophore. a, Sheath
of ampulla; b, spermatozoa; c, stalk; d, veil; e, pedestal.
(35X.)

therefore, a continuous ribbon, or two ribbons
if both vasa deferentia expel their contents
simultaneously.

Comparison of Regions of Spermatophoric

Development in D. asper with
Those of Other Species

Mouchet {op. cit.) assigns nine regions of
activity to the vas deferens of Diogenes pugi-
lator Roux. The spermatophoric differences
observed in other species are attributed to
vasa deferentia which lack one or more of
these regions. The following are the nine re-
gions of activity assigned to the vas deferens
of D. pugilator:

1. A rectilinear, irregular part of the canal
coming from the testis.

2. A right-handed helix, where the canal
narrows, in the last turns of which the shell
of the ampulla is secreted.

3. A left-handed helix, where the canal in-
creases in diameter, in the first turns of which
the column of sperm is fragmented into seg-
ments by its being curved into small arches
and by ampullae being formed by the uniting
of their extremities.

4. In the last turns of the second helix each
ampulla acquires a short and thick stalk.

5. The internal canal enlarges into a spindle-
shaped chamber. At the beginning of the spin-
dle, in the thin part, the secretion of the

Spermatophore of Dardanus asper — Matthews

265

TABLE 1

The Presence, Absence, or Modification in Other Species of Crabs of the
Nine Regions of the Vas Deferens Proposed for D. pugilator

SPECIES

region

Diogenes pugilator.

1

2

3

4

5

6

7

8

9

Eupagurus hernhardus

p

P

P

a

P

a

P

P

P

Eupagurus prideauxi.

P

P

P

a

P

a

P

P

P

Eupagurus cuanensis

P

P

P

a

P

a

P

P

P

Anapagurus hyndmanni (left side)

P

P

P

a

P

a

P

P

a

Anapagurus hyndmanni (right side)

P

P

P

a

P

a

P

P

P

Clihanarius misanthropus

P

P

P

a

P

a

a

a

P

Pagurus arrosor.

P

m

m

m

m

a

a

a

P

Dardanus asper. •

P

m

P

P

P

P

a

m

P

p ~ presence; a ~ absence; m = modified.

pedestal is produced in the form of a con-
tinuous ribbon which upholds the individual-
ized stalks.

6. In the thick part of the spindle the stalks
are stretched.

7. At the distal extremity of the spindle,
the segmentation of the ribbon of the pedestal
takes place.

8. In the tube which comes after the spin-
dle, the spermatophores are in single file in
the canal, the ampullae directed proximally.

9. The canal becomes larger, the ampullae
dispose themselves in any way, the pedestals
staying in rank, one after another, until they
reach the genital aperture. At the beginning
of this region the gland cells secrete a mucus
which surrounds the spermatophores.

The presence, absence, or modifications of
these nine regions in the species so far ade-
quately investigated are summarized in Table
1. Dardanus asper, although possessing region
1, does not possess a well-defined region 2
in which the coils of a right-handed helix
secrete the sperm-column sheath. In this re-
spect the vas deferens of D. asper resembles
Pagurus arrosor in which a fine tube of con-
secutive, closed turns rotates now to the right
and now to the left. D. asper resembles Eupa-
gurus hernhardus, E. prideauxi, E. cuanensis,
Anapagurus hyndmanni, Clihanarius misan-
thropus, and Diogenes pugilator in that the
sperm column is segmented between regions
2 arid 3. Dardanus asper resembles Diogenes

pugilator in possessing region 4, where the
stalks appear, region 5, where the pedestal
is developed, and region 6, in which the stalks
are lengthened, but differs from Diogenes pugi-
lator, E. hernhardus, £. prideauxi, E. cuanensis,
and A. hyndmanni in not having region 7,
where the pedestal is segmented. Although
D. asper possesses region 8, in which the
spermatophores are in single file, the ampullae
are not necessarily directed toward the begin-
ing.

In so far as can be determined, D. asper
is the only hermit crab which has been in-
vestigated in which the stalks lengthen, but
the pedestal fails to segment. The sperma-
tophore of D. asper may appear sufficiently
different from those previously described for
other hermit crabs to set it apart in the scheme
of spermatophoric development. However,
too few genera have been investigated to draw
any valid comparisons either on the com-
plexity of their vasa deferentia or the sperma-
tophores which they produce. In all hermit
crabs studied the spermatophore is developed
with great precision. It is always possible to
determine the species by careful examination
of the spermatophore. The effects of a chang-
ing environment on the form of the sperma-
tophore should prove in Coenohita and Birgus
a lucrative field for further investigations.

RESUME

The effect of crowding on the reproductive

266

PACIFIC SCIENCE, Vol. VII, July, 1953

organs is negligible, as spermatophores pro-
duced by right and left vasa deferentia reveal
no dimorphism.

The living vasa deferentia, freed of the con-
cealing testes, exhibit spasmodic contractions.
These serve not only to move the sperm mass
but to mold it in compliance with the gradual-
ly changing internal die of the vasa deferentia,
i.e., first cylindrical, then elliptical, then pear-
shaped. Serial sections reveal that these grad-
ual morphological changes are paralleled by
physiological changes.

A continuous sperm mass emanates from
the testis and enters the undifferentiated por-
tion of the vas deferens. Here the epithelial
cells secrete a substance of unknown function
which mixes with the spermatozoa.

Epithelial cells, isolated at opposite ends
of the elliptical lumen, produce a new secre-
tion which at first covers the distal ends of
the epithelium but later comes to lie con-
tiguous with the sperm column. This is the
sperm-column sheath.

The sperm-column sheath is formed
through several regions of the vas deferens,
not solely in the coils of a right-handed helix.

Longitudinal sections through the region
of the last compact, left-handed helix fail to
show the fragmentation of the sperm column
into separate ampullae. The sperm column re-
mains continuous, but by-muscular contrac-
tions of the wall of the vas deferens it is
forced into continuous sinusoidal curves or
arches.

The groove of the now pear-shaped lumen
deepens, and a new substance is secreted from
the epithelial cells at its base. This is at first
thread-like but later thickens in the regions
between the closing arches. These accumula-
tions are the precursors of the stalks. The
muscular activity which closes the arches to
form the ampullae of sperm is co-ordinated

with that which lengthens the accumulations
of this secretion to form the stalks.

Cross sections through the region of the
flattened spiral reveal the continued lengthen-
ing of the stalks and show the origin of a new
secretion, the precursor of the veil. As the
lumen of the vas deferens increases in dia-
meter, the veil and the stalks become more
elongate, and the ampullae of sperm are borne
on the distal ends of the stalks. The sperm-
column sheaths between adjacent ampullae
are stretched by this process. They become
extremely thin and, finally, are no longer de-
tectable, at least not with the staining tech-
nique employed.

Cross sections through the enlarged, apical
portion of the vas deferens reveal a cylin-
drical lumen distended by fully formed, pe-
dunculate spermatophores. However, two
typhlosole-like folds develop which produce
a mucoid secretion which encompasses the
spermatophores.

Muscular contractions of the wall of the
vas deferens serve now to expel the com-
pleted spermatophore. When first expelled,
it is difficult to distinguish the pedestal from
the veil. If the spermatophore of D. asper is
allowed to stand in tap water, the veil soon
disintegrates, leaving the upright stalks with
their ampullae directly attached to the con-
tinuous, unsegmented pedestal. Dardanus
asper is the only hermit crab so far studied
whose vasa deferentia possess the region for
the lengthening of the stalks but lack the
region for the segmentation of the pedestal.

REFERENCE

Mouchet, S. 1931. Spermatophores des crus-

taces decapodes, anomures et brachyures

et castration parasitaire chez quelques pa-

gures. Sta. Oceanogr. de Salammbo^ Ann. 6:

1-203.

A Preliminary Report on the
Soils of Saipan, Mariana Islands^

Ralph J. McCracken2

INTRODUCTION

The purpose of this paper is to provide a
preliminary description of the nature of Sai-
pan soils and their factors of formation. A
preliminary classification and correlation of
the soils is presented. Arrangements have been
completed for publication, at a later date, of
a complete discussion of Saipan soils in con-
junction with a series of discussions of the
geology of Saipan, under the auspices of the
United States Geological Survey. Laboratory
analyses of representative soil-profile samples
are at present under way. Genesis and classi-
fication of Saipan soils will be discussed more
fully in the later publication, utilizing the
results of the laboratory analyses.

This study demonstrated the applicability,
with minor modifications, of American theo-
ry and methods of soil survey to pioneer in-
vestigations of tropical island groups. How-
ever, the study indicates the desirability of
improving nomenclature and of differentia-
ting criteria of classes of tropical soils in the
higher categories of the natural system of
classification.

The survey reported in the present paper
is presented as an example of the detailed
areal soil studies advocated by students of
tropical soil classification (Kellogg, 1948;

^ A detailed soil survey of Saipan Island, in which
the author participated, was conducted in 1949 under
direction of the U. S. Geological Survey. The present
paper is a report of this work performed while the
author was on loan to Military Geology Branch, U.
S. Geological Survey, and its publication was encour-
aged by that organization. Manuscript received August
19, 1952.

2 Soil Scientist, Division of Soil Survey, Bureau of
Plant Industry, Soils and Agricultural Engineering, U.

S. D. A.

Pendleton, 1948) as one of the next important
steps .in arriving at a true understanding of
tropical soils.

Acknowledgments

The author wishes to express his apprecia-
tion to P. E. Cloud, Jr., and R. G. Schmidt of
the U. S. Geological Survey for providing
information on geologic and petrographic
studies of Saipan currently in progress. Thanks
are due M. G. Cline of Cornell University and
R. W. Simonson of the Division of Soil Sur-
vey, Bureau of Plant Industry, Soils and Agri-
cultural Engineering, United States Depart-
ment of Agriculture, for critical reading of
the manuscript and for helpful suggestions.

GEOGRAPHY OF SAIPAN

Saipan is located at 15° north latitude, about
1,500 miles southeast of Japan proper and
1,200 miles east of the Philippines. It lies in
approximately the center of the Mariana Is-
lands, about 150 miles north of Guam, which
is at the southern end of the chain.

This 48-square-mile island consists essen-
tially of a core of volcanic rock around and
over which limestones have been formed. The
highest elevation, about 1,550 feet above sea
level, is a peak on the island’s central ridge,
which is composed of uplifted limestone. The
island is predominantly rolling and hilly ter-
rain with the exception of limestone plains
at the southern end, on the eastern peninsula,
and at the northern tip.

NATURE OF SOIL-FORMING FACTORS
Parent Materials

Volcanic rocks and volcanically derived
rocks (sandstones and conglomerates) of Eo-

267

268

PACIFIC SCIENCE, Vol. VII, July, 1953

cene age are the oldest outcropping rocks on
Saipan. They underlie less than one third of
the soils.

Dacitic (silica-rich) rocks of the Sankaku-
yama formation are the oldest volcanic rocks.
The dacites are exposed in two small areas
totaling about square mile (included in the
land type of rough stonyland on dacite (Fig.
1). They contain about 80 per cent Si02 (Fig.
5).

Next younger are rocks of the Hagman and
Densinyana formations. The Hagman rocks
are andesitic breccias, conglomerates, tuffs,
and flow rocks. Chemical composition of one
of the andesites is shown in Fig. 5. The Den-
sinyana formation is composed entirely of
reworked, water-deposited conglomerates,
sandstones, and breccias derived from vol-
canic rocks. These two formations underlie
about 8 to 10 square miles.

Limestones underlie about two thirds of
the soils of the island. The Tagpochau lime-
stone of Miocene age and the Mariana lime-
stone of Pliocene or Pleistocene age are the
most extensive and important in relation to
soils.

The Tagpochau limestone makes up the
central backbone of the island and is the most
extensive geological unit. The Chinen soil
and the land type of rough stonyland on
limestone are extensive on this limestone.
Deeper pocket-like areas are the sites of the
Chacha and Saipan soil series. Sandstones and
conglomerates of material derived from vol-
canic rocks are found in the same timeplanes
as this Miocene limestone. Soils developed
from this material have been classified with
the Teo series (a minor soil unit) or with the
Akina or Dago series, according to their
morphology.

The Mariana limestone is found chiefly in
rather level plains in southern and eastern
Saipan. This limestone contains little volcanic
contamination but does include an argilla-
ceous (clayey) facies. The Dandan and Chinen
soil series are developed from Mariana lime-

stone, and some areas of Chacha and Saipan
soils are also underlain by this material.

Relief

The most striking landforms of Saipan are
the highlands of predominantly Miocene
limestone which make a "backbone” along
the center of the island, and bench levels in
Miocene and younger limestone which flank
this ridge (Fig. 1). The central ridge and the
benches are broken in several localities by
exposures of volcanic rocks. The central lime-
stone ridge, most of which has a maximum
elevation of 800-1,000 feet, rises to a point
of 1,550 feet above sea level.This peak, known
as Tagpochau, is the island's highest point.
The wave-cut limestone benches are especi-
ally well developed in the northern and eastern
parts of the island. These bench levels are
generally nearly level or gently sloping and
vary from a few hundred to a few thousand
feet in width. They are ordinarily separated
by steep cliffs which give a giant stair-step
aspect to regions where they are well devel-
oped. Broad, nearly level limestone plains in
the southern portion vary from about V2 to
1 mile in width. Presence of a few small sink-
holes and caverns indicates some solution of
the limestone in these areas.

Soil depth and development in areas un-
derlaid by limestone is closely related to slope.
Shallow stony soils with little profile devel-
opment (the Chinen series) and rough stony-
land with little soil material are found on
steeper slopes where geologic erosion ap-
proximately keeps pace with soil develop-
ment. Deeper soils with some horizon differ-
entiation have developed on the bench levels
and plains where geologic erosion is less
rapid and soil material is received from ad-
jacent steeper areas by colluviation. The deep
soils on undulating and rolling topography
(Saipan series) show subsoil colors of dom-
inantly reddish hues, whereas deep soils of
the nearly level areas (Chacha series) are of
dominantly yellowish hue with some mottles

ROUGH STONYLANO ON OACITE

Fig. 1. Landscape of Saipan and typical sites of occurrence of important soils and land types.

270

PACIFIC SCIENCE, VoL VII, July, 1953

in the subsoil, indicating less favorable oxida-
tion and aeration.

The areas of volcanic rock outcrops are
highly dissected and are at maximum relief.
Ridges are generally sharp and narrow; ra-
vines are deep. Many of these areas are under-
going accelerated erosion with prominent
gullies and erosion scars. As a result, very
little soil material (i.e., horizons of organic
accumulation and modified parent material)
is present, and the weathered volcanic rocks
(the "saprolite” or ^zersatz” of some authors)
are exposed at the surface. Such areas are
classified with the land type of rough broken
land. Shallow soils, such as the Teo soils, are
found where erosion is less severe. Deep soils
underlaid by volcanics have developed only
on broad ridge tops and the less strongly
sloping areas.

OTHER 2%

TOTAL AREA -48 SQUARE MILES

Fig. 2. Percentages of Saipan soil groups and land
types.

Climate

Saipan at 15° north latitute is within the
zone of tropical climates with year-round high
temperatures (Reed, 1941).

Mean monthly temperatures range from 82
to 86°F. Absolute minima and maxima are
within 10 to 15 degrees of the mean tem-
peratures (Fig. 3).

A weak dry season (according to criteria
of Mohr, 1944) occurs in March and April.

Fig. 3. Mean monthly temperatures of Saipan Island,

Average yearly rainfall is 82.5 inches (Fig. 4).
(The foregoing climatic data are from Japan-
ese sources for the years 1928-1937, inclu-
sive.)

Time

Geomorphic and stratigraphic evidence in-
dicates that weathering and soil formation in
the uplands have been proceeding without
interruption since at least the latter part of the
Pleistocene period. Presence of marine sedi-
ments in the Tagpochau limestone indicates
encroachment of the sea upon the land mass
during the Miocene period. However, since
probably late Pleistocene time and the uplift
of the Mariana limestone, no interruptions
in weathering and soil-forming processes on
the uplands of the island have taken place.
It may then be concluded that soil develop-
ment in the upland areas has been continuous
for more than 25,000 years.

Vegetation

The original vegetation of Saipan, inferred
to have consisted of fairly dense forests and
some savanna-like areas of minor extent, was
greatly decimated by extensive clearing for
sugar cane culture during the period of Jap-
anese control.

Fig. 4. Mean monthly rainfall of Saipan Island.

Soils of Saipan^-McCRACKEN

271

Primary, climax forests apparently occupied
the areas of moderate relief which had some
soil mantle. They appear to have been com-
posed of such species as daog {Calophyllum
imphyllum), ifilwood {Intsia hijuga), bread-
fruit {Artocarpus spp.), several species of
Pandanus, and others. Soils derived from a
variety of parent materials were found to
have no significant differences in organic-
matter content and carbon- nitrogen ratios
(Kawamura et al., 1940), indicating no im-
portant differences in vegetation.

Areas of steep relief and of rough stony-
knd in general have a cover of secondary
tree species, such as Leucaena glauca and Acacia
confusa, and several species of shrubs. In the
minor savanna areas, the dominant species is
"swordgrass” {Miscanthus floridula ?), with
some Casuarina present. To what extent such
areas are anthropic (man-caused), was not de-
termined, but they do occur in areas under-
laid by both limestone and volcanic rocks.

PRINCIPAL SOILS AND LAND TYPES AND

THEIR RELATIONS TO SOILS ELSEWHERE

Distribution of the principal soils and land
types is shown in Figure 6. Because of the
intricate pattern of occurrence of the soil se-
ries and the small scale of the map, it has been
necessary to generalize the upland soils both
cartographically and categorically from the
original survey. With categorical generaliza-
tions only a limited number of precise state-
ments can be made about the classes, or units,
shown, as the class interval of each is broad
and includes a wide range of soil conditions.
The homogeneity of the areas shown is re-
duced by the cartographic generalization. Soil
associations or landscapes, rather than dis-
tinctive soils, are shown. However, the map
is useful for obtaining a general picture of
soil distribution on the island.

Miscellaneous Land Types

Rough stonyland on limestone includes
a range of conditions from a few inches (less
than 4 or 5) of stony soil mantle to bare

UPPER BAR OF EACH OF THE PAIRS BELOW REPRESENTS PERCENTAGE
COMPOSITION OF UNALTERED CORE OF BOULDER OF HAGMAN CALCIC
ANDESITE, LOWER BAR REPRESENTS WEATHERED OUTER MANTLE
OF SAME BOULDER. (I)

SiO

2

AIzOs

H 60.95%
54. 39 %

18.06 %
24.03 7o

FezOs

FeO

MgO

CaO

NogO

2.41 %
■ 5.157«

■ 2.617®
0.58 7®

I 2.37%
0.58 7®

■■18.16 %

0.10 7®

■ 3.107®

I 0.44

CHEMICAL COMPOSITION OF DACITE PORPHYRY IN SANKAKU^AMA

F0RMAT10N(2)

SiOg 79 0 %

AlgOa tm 9.94 7®

FegOg 11.28 7®

NagO 13.88%

11.20 7®

(1) ANALYSES BY A.C. VLISIDIS, SAMPLES COLLECTED BY R 6. SCHMIDT,
BOTH OF U.S. GEOLOGICAL SURVEY.

(2) ANALYSES BY F A. GONYER, DEPT. OF MINERALOGY, HARVARD
UNIVERSITY, SAMPLE COLLECTED BY R.S. SCHMIDT.

Fig. 5. Chemical composition of some Saipan vol-
canic rock.

limestone outcrops. The modal condition is
a small amount of mineral soil incorporated
with loose stones, the soil material extending
down into cracks and fissures in the limestone.
The widespread occurrence of this unit (Fig.
2) is due to the steep topography associated
with the uplifted limestones. Normal geologic
erosion keeps pace with soil formation, and
recurring stages of uplift have accelerated soil
removal.

Rough stonyland on dacite describes a
condition of limited extent in northern Saipan
(less than 1 square mile) in which practically
barren dacite or very few inches of gray, acid,
loamy soil are found overlying the dacite on
less steep slopes.

Rough broken land includes those areas
(Figs. 1, 6) in which weathered rock of vol-
canic origin is present at or near the surface,
a zone of organic accumulation being absent
or very thin. Numerous gullies and erosion
scars are evident. The weathered, weakly con-
solidated tuffs and waterlaid sediments from
volcanic rocks are extremely susceptible to
erosion, markedly so on steep slopes.

272

PACIFIC SCIENCE, Vol. VII, July, 1953

CHINEN-SAIPAN SOIL ASSOCIATION (INCLUDES DANDAN AND
CHACHA SOILS AND ALLUVIAL CLAYS)

AKINA-DAGO SOIL ASSOCIATION (INCLUDES TEO SOILS)
SHIOYA SAND

ROUGH STONYLAND- LIMESTONE
ROUGH STONYLAND - DACITE
ROUGH BROKEN LAND
MARSH

COMPILED FROM DETAILED SURVEY BY R J.
McCRACKEN AND R E. ZARZA, 1949

Fig. 6. Generalized soil map of Saipan Island.

Soils of Saipan- — McCracken

273

Shallow and Moderately Deep Soils
over Limestone

Chinen stony clay loam, the most wide-
spread (Fig, 2) of Saipan soil series, is a stony,
dark brown, alkaline clay loam (to light clay)
12 to 18 inches deep over limestone (Fig. 1).
It can be fitted into the American scheme of
classification as a lithosol (Baldwin et aL,
1938).

Dandan clay loam is a friable, brown,
slightly alkaline soil 18 to 42 inches deep over
the younger limestone (Fig. 2). Bodies of
this soil are located on the lower limestone
bench levels, chiefly at southern and northern
ends of the island. On the basis of the above-
described properties, this soil can be classified
as a brown forest soil (Baldwin et aL, 1938).
By inference from observations of tropical
soils elsewhere, this soil can be expected to
become more acid and redder with time, ow-
ing to increasing loss of bases and to ses-
quioxide concentration under the continuous-
ly high temperature and moderately high
annual rainfall prevailing.

Shallow and Moderately Deep Soils over
Volcanks and Volcanically Derived Rocks

T eo soils are firm, reddish brown or brown,
acid, of medium to heavy textures, and aver-
age 12 to 18 inches deep over weathered
volcanic tuffs, sediments from volcanic rocks,
and highly impure limestones (Fig. 1). They
are of limited extent (Fig. 2). In the central
hilly uplands where these soils are found,
parent materials are highly variable within
short distances laterally, therefore these soils
have been mapped as a "complex” rather than
homogeneous soil series. Because of the un-
consolidated nature of the parent materials
and the shallowness of the solum (A and B
horizons), these soils may be classified as re-
gosols (Thorp and Smith, 1949).

Deep Soils over Limestone

Two principal soil series were recognized
in the deeper soils underlaid by limestone.

These two are of essentially similar texture,
consistence, and reaction but differ chiefly in
subsoil color.

Saipan clay has a dark-brown granular clay
surface soil about 6 inches thick which is
neutral in reaction; the subsoil is yellowish
red to red (the former dominant), firm, plas-*
tic, and very slightly add. Depth to limestone
ranges from a little less than 4 to 6 feet (pro-
file diagram, Fig. 1). This soil is found chiefly
on colluvial slopes and in pockets in rolling
and hilly areas of the central uplands (Fig. 6).
This soil series is the most extensive of the
deeper soils of the island (Fig. 2). Tables 1,
2, and 3 show chemical analyses of soil sam-
ples from sites apparently within areas of Sai-
pan clay reported by Kawamura et al. (1940).
They are of uncertain value for our use, owing
to uncertainty of correlation of the sampling
sites described in this paper, uncertainty as
to laboratory procedures used due to trans-
lation difficulties, and lack of complete hori-
zon-by-horizon determinations for a particular
profile. Flowever, the results do give some
idea of developmental trends and bulk chem-
ical composition. A silica content (whole soil)
of about 25 per cent, a cation exchange capa-
city near 7 milliequivalents per 100 grams of
dry soil, and a derived silica-sesquioxide ratio
of slightly less than unity in the subsoil are
indicated. (The significance of these data lies
in their use as rough approximations, within
limits, to the degree of tropical weathering
which has taken place and for comparison to
soils elsewhere.)

Chacha clay has a dark-brown granular
surface, which is about neutral in reaction and
averages 6 inches in thickness; the subsoil is
of strong, brown to yellowish-brown, firm
plastic clay, slightly acid in reaction. Average
depth to limestone is 4 to 6 feet (profile
diagram, Fig. 1). Subsoil colors yellower than
in the Saipan series appear to be correlated
to the less well-drained position of this soil
on the level limestone plains of the eastern
peninsula and south-central part of the island.
However, numerous manganiferous concre-

274'

PACIFIC SCIENCE, VoL VII, July, 1953

TABLE 1.

Chemical Composition of Some Saipan Soil (Analyses of
Whole Soil, after Kawamura et al.)

SOIL TYPE

SAMPLE

DEPTHS

LOSS ON

IGNITION

sio2

AI2O3

Fe203

Cao

MgO

K20

Na20

Inches

Per cent

Per cent

Per cent

Per cent

Per cent

Per cent

Per cent

Per cent

"Red-colored limestone soil"

= Saipan clay

0- 6

22.36

25.62

31.46

14.18

2.96

2.10

0.27

0.64

6-45

13.10

26.38

40.86

15.07

2.63

2.36

0.25

0.63

"Yellow-colored limestone

soil" = Chacha clay

0- 8

18.82

31.99

29.20

15.90

1.67

0.12

0.85

8-27

13.37

32.32

32.28

15.10

0.64

0.22

0.64

"Brown-colored limestone

soil” “ Dandan clay

0-10

18.98

24.81

35.15

11.58

5.86

2.66

0.24

0.92

10-33

14.22

15.38

43.64

15.91

6.85

1.05

1.55

1.03

"Red andesitic soil"

= Dago clay

0-12

14.12

34.24

27.57

21.05

0.76

0.94

0.17

0.25

12-50

10.67

37.95

31.01

17.26

0.45

1.25

0.14

0.50

"Red tuffaceous soil"

= Akina clay.

0- 8

14.02

46.61

19.76

14.52

1.21

0.25

0.33

8-24

10.33

50.37

24.60

12.20

0.95

0.19

0.43

24 +

7.52

53.02

23.76

12.34

0.82

0.15

0.74

Data are abstracted from a number of analyses reported by the Japanese investigators. These analyses were carried out with
the carbonate fusion method.

dons and streaks which are present may be
indicative of parent material differences or
introduction of such material in ground water.
Chemical analyses (Tables 1,2,3) indicate a
silica content (whole soil) of about 30 per
cent, cation exchange capacity near 6.5 milli-
equivalents per 100 grams of dry soil, and a
derived silica-sesquioxide ratio near 1.5.

These soils differ in many respects from
most deep soils over limestone of warmer
climates elsewhere, such as the ”Terra Rossa”
soils, the ”laterite” soils of Cuba and Puerto
Rico, and the "reddish-brown lateritic” soils
of the southern United States, but do seem
to resemble some Puerto Rican soils as de-
scribed by Roberts et al. (1942). They differ
from Terra Rossa, an ill-defined group of soils
especially common over limestones of the
Mediterranean area (Reifenberg, 1938; Joffe,
1949) by reason of higher organic matter
content and lower content of bases. Many
investigators wish to restrict Terra Rossa to
red soils over limestone developed under a
Mediterranean climate-^cool, moist winters
and hot, dry summers. Red soils over lime-

stone in the southern United States, especially
the Dewey and Decatur series, differ from the
Saipan and Chacha series in consistence and
reaction (data on these American soils are
presented by Alexander et ah, 1939). Saipan
clay and Chacha day are more plastic and
firm and have higher pH values, being only ^
very slightly acid in the subsoil. The "lat-
erite” soil over limestone in Cuba and Puerto
Rico (the Matan2as series) is also more friable
and acid than these soils of Saipan (Bennett
and Allison, 1928; Roberts et al., 1942). The
firm soils over limestone in Puerto Rico, es-
pecially the Coto series, appear to resemble
the Saipan and Chacha series.

Since the Saipan and Chacha series do not i
have some of the important characteristics I
described by Kellogg (1949) as criteria of |
latosols, they are provisionally termed "lato-
solic inter grades.” They are not highly porous
or friable but are firm and plastic. They are
nearly neutral in reaction which, together with
available data from chemical analyses, indi-
cates that they do not have low silica-sesqui-
oxide mole ratios, extreme silica depletion,

Soils of Saipan — McCracken

275

TABLE 2.

Some Constants of Representative Saipan Soils
(after Kawamura et al.)

SOIL TYPE

SAMPLE

DEPTHS

CARBON

HUMUS

NITROGEN

c/n

CATION

EXCHANGE

CAPACITY

Inches

Per cent

{Cxl 724)

Per cent

Ratio

m.e./lOO g.

"Red-colored limestone soli”

= Saipan clay

0- 6

5.95

10.26

0.52

11.4

7.2

6-45

7.2

"Yellow-colored limestone

soil” = Chacha clay

0- 8

2.48

4.28

0.24

- 10.3

6.6

8-22

"Brown-colored limestone

soil” = Dandan clay loam. .

0-10

3.98

6.86

0.34

11.7

10-33

9.0

"Red andesitic soil”

= Dago clay

0-12

2.9

5.0

0.23

12.6

12-50

12.1

"Red tuffaceous soil”

“ Akina clay

0- 8

0.28

8-24

35.2

These determinations were completed for a limited number of horizons of a few profiles. Though not sufficiently complete
for basis of conclusions, they are presented here because they do show trends in soil development processes. Carbon was deter-
mined by the wet oxidation method and nitrogen by the Kjeldahl method. Due to some questions concerning translation, method
of determining cation exchange is uncertain.

or extremely high sesquioxide concentrations.
They were observed to crack in drier seasons
and swell during wet periods. However, they
do possess some red color, contain low
amounts of soluble constituents, and have
developed under a climate of year-round
weathering with moderately high rainfall.

Deep Soils over Volcanic Rocks

Akina clay has a reddish-brown, granular
clay surface averaging about 6 inches in thick-
ness and is acid in reaction {pH 6.0). The
subsoil is of yellowish-red, firm plastic clay to
a depth of about 18 inches. Within an approx-
imate depth range of 18 to 30 inches below
the surface, the subsoil is of yellowish-red and
red hues, often displaying light-gray spots
and flecks which appear to be relict colors of
highly weathered minerals of the parent ma-
terial. Below this horizon is a variegated red,
yellowish-red, and light-gray clay, strongly
acid (^H 5.0 to 5.5), with relict textures of
the weathered parent material and feldspar
''ghosts” (outlines of weathered feldspar crys-

tals) commonly preserved (profile diagram.
Fig. 1). Depth to unaltered, moderately basic
volcanic rocks ranges from 15 to 25 feet or
more. The chemical data shown in the tables
for soil samples believed to correlate with this
soil series are somewhat out of line with sam-
ples from other soils of the island. More than
40 per cent silica, a cation exchange capacity
of 35 milliequivalents per 100 grams of dry
soil, and a derived silica-sesquioxide ratio of
greater than two is reported for the subsoil
in samples believed to correlate with the Aki-
na soil. These data are indicative of a less
strongly weathered soil. The Akina subsoil
was observed to be firm and plastic and ex-
hibits cracking and swelling at extremes of
moisture, not indicating extreme sesquioxide
concentration or silica depletion.

Dago clay has a dark reddish-brown, gran-
ular clay surface, slightly acid in reaction. The
subsoil is a dark-red firm acid (^H 6.0) clay
(to an average depth of 30 inches) . Below this
horizon a yellowish-red and dusky red, firm,
plastic, acid {pH 5.5) clay extends to a depth

276

PACIFIC SCIENCE, VoL VII, July, 1953

TABLE 3.

Chemical Analyses of the Fine Fraction ( <0.002 mm.) of
Representative Saipan Soils (after Kawamura et al.)

SOIL TYPE

sample

DEPTHS

Si02

AI2O3

Fe203

si02

AI2O3

si02

R2O3

Inches

Per cent

Per cent

Per cent

MoL Ratio

MoL Ratio

"Red-colored limestone soil”

= Saipan clay

0- 6

23.53

34.79

15.77

1.15

0.89

6-45

25.91

40.56

10.35

1.08

0.93

"Yellow-colored limestone

soil” = Chacha clay

0- 8

33.99

43.21

15.36

1.33

1.09

8-22

37.50

36.37

8.77

1.74

1.51

22 +

40.70

38.78

6.52

1.78

1.58

"Brown-colored limestone

soil” = Dandan clay loam. . .

0-10

15.09

40.05

10.81

0.64

0.54

10-33

12.25

43.30

13.91

0.48

0.40

"Red andesitic soil”

= Dago clay

0-12

33.90

33.24

11.32

1.73

1.42

12-50

36.32

33.39

13.27

1.84

1.47

"Red tuffaceous soil”

= Akina clay

0- 8

49.37

25.96

11.15

3.20

2.51

8-24

45.98

27.92

7.96

2.66

2.25

24 -f

42.59

28.06

11.34

2.58

2.05

Fractionation was obtained by the pipette method.

of 4 feet or more. A highly variegated red,
yellow, and white strongly acid clay of vary-
ing consistence extends to depths of 15 feet
and more, below which are unaltered volcanic
tuffs, flow-rocks, and sediments of volcanic
origin (profile diagram, Fig. 1). Chemical data
of Kawamura (Tables 1, 2, 3) for samples
apparently correlating with this soil series in-
dicate about 35 per cent silica in the fine
fraction of the subsoil. Cation exchange ca-
pacity of 12 milliequivalents per 100 grams of
dry soil and a derived silica-sesquioxide ratio
of about 1.45 for the subsoil are reported.

These deep soils underlaid by volcanic
rocks do not fully qualify as latosols on the
basis of their morphological and chemical
properties, as described above. One would
expect to find latosols, as environmental con-
ditions of year-round high temperatures and
moderately annual rainfall are similar to other
parts of the world where latosols have been
described as developed over volcanic rocks.
Examples are the Flawaiian Islands, where
latosols have been described by Cline (manu-
script) and chemical data presented by Hough

and Gile (1941), and the soils of the Belgian
Congo described by Kellogg and Davol
(1949). However, the Cialitos soils of Puerto
Rico as described by Roberts et aL (1942)
seem to be similar to the Akina and Dago
soils in properties and environment and were
classified as "Reddish Brown Lateritic.” The
tuffaceous shales and volcanic rocks described
as the parent material of this Puerto Rican
soil may, however, have had some extreme
effects on its direction of development.

As the deep soils of Saipan underlaid by
volcanic rocks more closely resemble latosols
than any other class at present defined, it is
proposed also to term them "latosolic inter-
grades.” This signifies that their properties
are in a range between those of a modal latosol
and some other class or classes not at present
defined. More precise definition of classes of
tropical soils in the higher categorical levels
of classification and more laboratory analyses
of Saipan soils are necessary before any fur-
ther statements on their classification and 1
correlation can be made. i

Soils of Saipan — McCracken

111

SUMMARY

Saipan Island, 48 square miles in area, is
located in the center of the Mariana Islands
in the western Pacific Ocean, well within the
zone of tropical climates. Mean annual rain-
fall is about 80 inches with a weak dry season.
Mean monthly temperatures range from 82
to 86°F.

Limestones underlie about two thirds of the
soils of the island, volcanic rocks one third.
Deep, well-drained upland soils are rather mi-
nor in area, as a shallow stony soil and rough
stonyland on limestone make up 29 and
36 per cent, respectively, of the surface area
of Saipan.

Though of limited occurrence, the deep
upland soils pose interesting problems in soil
genesis and classification. They are lacking
in sesquioxide concentrations and silica de-
pletion and possess features, such as plas-
ticity and firmness, which do not conform
to the criteria of latosols. They are provision-
ally termed latosolic intergrades.

Laboratory analyses of samples of Saipan
soils are currently in progress.

REFERENCES

Alexander, L. T., H. G. Byers, and G.
Edgington. 1939- A chemical study of
some soils derived from limestone. U. 5.
Dept. Agr., Tech. Bui. 678: 1-20.

Baldwin, M., C. E. Kellogg, and J. Thorp.
1938. Soil classification. In Soils and men.
U. S. Dept. Agr., Yearbook for 1938: 919-
1001.

Bennett, H, H., and R. V. Allison. 1928.
The soils of Cuba, xxiv + 410 pp., 6 figs.,
1 map. Tropical Plant Research Founda-
tion, Washington.

Cline, M. G. (In Press.) The soils of Haw^aii.
Hough, G. J., and R. S. Gile. 1941. Rock
weathering and soil profile development in

the Hawaiian Islands. U. S. Dept. Agr.,
Tech. Bui. 752: 1-30.

JOFFE, J. G. 1949. Pedology. (2nd ed.) xv +
662 pp., illus., maps. Pedology Publica-
tions, New Brunswick, New Jersey.

Kawamura, J., T. Tanaka, and T. Inagaki.
1940. On the soils of Saipan, Tenian and
Rota Islands. [Translated from Japanese by
R. Endo.] Jour. Set. Soil and Manures 14:
439-484.

Kellogg, C. E. 1948. Preliminary sugges-
tions for classification and nomenclature of
great soil groups in tropical and equatorial
regions. Commonwealth Bur. Soil Sci., Tech.
Commun. 46: 76-85.

— and E. D. Davol. 1949. An explora-
tory study of soil groups in the Belgian
Congo. Inst. Natl, pour C Etude Agron. Congo
Beige, Ser. Sci. 46: 1-61.

Mohr, E. C. J. 1944. The soils of equatorial
regions with special reference to the Neth-
erlands East Indies. [Translated by R. S.
Pendleton.] 776 pp., illus. inch maps, table.
J. W. Edwards Co., Ann Arbor.

Pendleton, R. S. 1948. The classification
and mapping of tropical soils. Common-
wealth Bur. Soil Sci., Tech. Commun. 46:
93-96.

Reed, W. W. 1941. Climates of the world.
In Climate and man. D. S. Dept. Agr., Year-
book for 1941 : 665-684.

Reifenberg, a. 1938. The soils of Palestine.
[Translated by C. S. Whittles.] 131 pp. T.
Murby Co., London.

Roberts, R. C., et al. 1942. Soil survey of
Puerto Rico. 503 pp., 145 figs., map (folded).
U. S. Department of Agriculture, Division
of Soil Survey, series 1936, No. 8.

Sherman, G. D. 1949. Development of lat-
eritic and laterite soils in the Hawaiian Is-
lands. Pacific Sci. 3: 307-314.

Thorp, J., and G. D. Smith. 1949. Higher
categories of soil classification. Soil Sci. 61:
117-126.

Seasonal Variations in the Physical Environment of the
Ponds at the Hawaii Marine Laboratory and the
Adjacent Waters of Kaneohe Bay, Oahu^

Winifred Shui Lin Tseu

INTRODUCTION

Kaneohe Bay, located on the northeastern,
or windward, side of the island of Oahu,
Territory of Hawaii, is approximately 7.3 nau-
tical miles long and 2.5 nautical miles wide,
with 3.8 nautical miles fronting on the Pa-
cific. It covers an area of 14.1 square nautical
miles. The entrance is protected for almost
all its length by coral reefs and has only two
deep channels into the adjacent ocean. Within
the bay are numerous coral reefs with channels
30 to 60 feet deep between them. Adjacent
to the shores the bay has broad shoal areas
of mud or sand that usually terminate in steep
fronts of growing coral. Numerous small
streams flow into the bay, but none of these
carries much water except at times of the
extremely heavy rainfall during southerly, or
"kona,” storms.

Located approximately 1.5 nautical miles
from the southeastern shore of the bay and
0.3 nautical miles from the closest land to the
southwest is a small island, Moku o Loe or
"Coconut Island.” The island is about 18
acres in area and reaches a height of 55 feet.
Originally it was surrounded by broad coral
flats of sand and mud, with growing coral on
the margins. However, a previous owner of
the island had numerous channels dredged
to the edge of the island, the dredged coral-
line material being thrown up to form jetties
protecting the channels. In one area a series
of fish ponds was constructed. The ponds are
separated by the coral fill and are connected

^ Prepared as partial requirement for the degree of
Master of Science, University of Hawaii. Contribution
No. 31, Hawaii Marine Laboratory, Manuscript re-
ceived July 11, 1952.

by concrete spillways into which screens can
be fitted. The sides of these ponds which
are almost vertical were constructed either of
disc-shaped coral heads or of concrete. The
bottom of the ponds is covered with a fine
silt, similar to that covering most of the bot-
tom of Kaneohe Bay.

With the establishment of the Hawaii Ma-
rine Laboratory on the island and the pro-
jected use of these ponds for holding ex-
perimental fish, it became desirable to learn
of the changes in the physical environment
in the ponds as contrasted to those in the
adjacent waters. This study attempts to fill
this need and, in addition, provides informa-
tion of more general interest as no study of
seasonal variation in inshore marine environ-
ments has been made in Hawaii heretofore
or, as far as can be ascertained, in other re-
gions of the tropical Central Pacific. More-
over, no study of seasonal variation of enclosed
waters in Hawaii has been made heretofore,
although ponds somewhat similar to the ones
studied have long been used in Hawaii for
rearing fish commercially.

The study, as carried out, fell into four
sections: first, the survey of the ponds to
determine their dimensions and depths; sec-
ond, the study of currents and tides in the
pond area, the study of the tides being neces-
sary for later sections of the study; third, the
study of the meteorology— rainfall, wind, and
air temperature— and water temperature, and
its correlation with the changes in the ponds
and adjacent waters; and fourth, an investi-
gation principally of changes in the chlorinity
and the oxygen content of the water.

278

Seasonal Variations — Tseu

279

Fig. 1. Map of Moku o Loe, "Coconut Island,” and vicinity showing stations, tide staff, temperature record-
ers, and standard rain gauge.

In the course of the 24 months’ investiga-
tion, 12 stations were established, 11 for the
first portion of the study, with the last station
in front of the Hawaii Marine Laboratory
added for the last 19 months. For convenience
of reference the stations have been numbered
as indicated in Figure 1. Stations 3 to 7 and
11 were located in the ponds; station 2 was
in the open water immediately adjacent to the
ponds; station 10 on the coral flat, exposed
at extreme low water and near the principle
series of ponds; and station 1 in the channel
off the island where the water is about 60
feet deep. This last station was used as typical
of the open water of the bay. These stations
were visited weekly for the first 14 months
of the study and monthly for the next 10
months. The period during which the study
was carried on covered the period from Feb-
ruary 19, 1949, to January 17, 1951.

Three 24-hour cycles of observations were
made to record diurnal changes in tide, tem-
perature, chlorinity, and oxygen content. The
dates of these three periods were selected to

permit comparison of the different seasons.

MENSURATION OF PONDS

The ponds and some adjacent waters were
surveyed to permit the drawing of charts with
depth contours and the estimation of volumes
and volume exchanges in the ponds. The
surveys were made in the following manner:
At regular intervals along the length of a pond
or channel, the distances depending on the
length and the configuration of the body, a
length of line marked off every 6 feet was
stretched across the pond. Then the observer,
either by wading or swimming, took a sound-
ing with a lead line at the marked intervals.
The depths were recorded to the nearest quar-
ter foot. Tidal readings were taken before and
after surveying each pond so depths could
be reduced to the zero tide line (the marine
datum line, 0.8 feet below mean sea level in
Honolulu) . The tidal readings throughout the
entire period varied only about 2 inches be-
cause at the time selected for the survey there
was a 'Vanishing tide” with its long station-

280

PACIFIC SCIENCE, Vol. VII, July, 1953

0

ro

2

g

1

cn

Fig. 3. Contour map of pond 5 showing depth in feet and location of station 3.

Seasonal Variations — TsEU

281

ary phase. These corrected values formed the
basis of the depth contours of the maps (Figs.
2 to 8). Contours were drawn not only from
the location of the soundings but also from
the known configuration of the ponds and
channels.

Areas of the different contours in the ponds
were then obtained by means of a polar plani-
meter (for description and use see Welch,
1948: 79-82). Using these data, the volumes
of the several ponds were then determined,
using the method outlined by Welch (1948:
95). The volumes of the several ponds are
shown in Table 1.

The volumes of average and maximum tidal
exchange as well as the percentage exchanges
in volume at mean and maximum tidal range
were computed on the basis of these area and
volume figures. The volumes of average tidal
exchange are the product of the area at the
zero tide line and the mean range of tide.
The percentage exchanges in volume at mean
tide range were obtained by dividing the vol-

ume of average tidal exchange by the total
pond volume at mean high water. The vol-
umes of maximum tidal exchange are the sum
of the maximum volume above and the mini-
mum volume below the zero tide line; that
is, the result of multiplying the pond area
(at zero tide) by the difference in height be-
tween the lowest and highest tide. The per-
centage exchanges in volume at maximum
tidal range were obtained by dividing the
volume of maximum tidal exchange by the
total pond volume at the maximum high wa-
ter. The values for minimum and maximum
height of tides and the mean range of tide
were calculated from the 1949 Tide Tables
for Waikane, Kaneohe. The value for mean
low water was obtained from the United
States Coast and Geodetic Survey. Inasmuch
as the walls above the zero tide line are almost
vertical, the volumes for the higher tides were
based on the area at the zero tide line. The
values for pond 11 were not computed be-
cause the shoreward margin of the pond is
gradually sloping and irregular.

Fig. 4. Contour map of pond 1 showing depth in feet and location of station 4.

282

PACIFIC SCIENCE, Vol. VII, July, 1953

TABLE 1

Area, Volume, and Volume Exchange of the Ponds and Boat Channel

POND

AREA

VOLUME AT

ZERO TIDE

VOLUME OF

AVERAGE

TIDAL

EXCHANGE

PERCENTAGE

EXCHANGE IN

VOLUME AT

MEAN TIDAL

RANGE

VOLUME OF

MAXIMUM

TIDAL

EXCHANGE

PERCENTAGE

EXCHANGE IN

VOLUME AT

MAXIMUM

TIDAL RANGE

1

Square feet
5,800

Cubic feet
21,000

Cubic feet
8,100

26

19,300

51

2

5,400

8,000

7,500

43

18,100

75

3

4,000

5,100

5,600

46

13,600

80

4

1,700

2,300

2,400

45

5,680

78

5

23,000

97,000

32,000

23

75,100

46

Channel

33,000

200,000

47,000

18

110,000

37

CURRENTS AND TIDES

Tidal current patterns were noted in ponds
1 to 5 of the main pond systems. Observations
were made on two flood and two ebb tides,
with winds of force 3-4 coming from the
north on the flood tides, and winds of force
0-2 coming from the southwest and west on
the ebb tides. To trace the water flow a sat-
urated solution of fluorescein dye in sea water
was poured into the water at various points.
The movements of the fluorescein dye were
noted at intervals, the total time of observa-
tion for each pond depending on the speed
of the dye movements.

The currents on the flood tides reach the
ponds by way of the channels north and west
of the net house (Fig. 1, B), and on the ebb
tides they leave the ponds by the same route;
that is, currents enter and leave ponds 1 and
5 from the adjacent channels, enter and leave
pond 2 mostly through pond 1, and enter
and leave the smaller ponds, 3 and 4, through
pond 5, though some slight water exchange
exists between ponds 2 and 3. On the flood
tides no currents were observed entering pond
5 by way of the tide flats (Fig. 1, 10). Instead,
water coming in from the channels moved
onto the flats.

Short-period reversals of flow several min-
utes in length were noted, especially on the
flood tides and at some of the gates. It was
not in the scope of this study to investigate

the phenomenon, but it is suggested that per-
haps these reversals may be either the result
of long-period waves sweeping over the outer
reef and reaching the island as imperceptible
changes in water level or the cumulative re-
sult of interference patterns of the smaller
off-shore waves.

As it was not possible to establish an auto-
matic tide gauge on the island, approxima-
tions of the tide were made using a tide staff.
The height of the tides was recorded in inches
above an arbitrary 2ero level by means of a
tide staff nailed to the dock (Fig. 1, A). A
series of comparisons, including those of the
three diurnal cycles, was made between the
observed Kaneohe tide and the theoretical
Honolulu tide. The times of high and low
water as well as the amplitude of the tidal
curves were compared.

The regularity of tidal movements is sub-
ject to the influences of wind. For example,
it is generally known that an onshore wind
tends to raise the level and an offshore wind
tends to lower the level of the sea along a
coast. Steady winds did not affect the times
of high and low water but merely their heights .
Variable winds, on the other hand, not only
affect the heights but also the times of high
and low water. In an enclosed bay such as
Kaneohe, with its wide shallow front as en-
trance and two comparatively narrow chan"
nels as exits, a period of high onshore winds
can raise the sea level higher and maintain

Seasonal Variations — TsEU

STATION 5 O °

Fig. 5. Contour map of pond 2 showing depth in feet and location of station 5.

o+ '4

o\%

0

I

G2 '/4

2

02 '4

oz<4 O

02^4

’/4

STATION 6 O

oi '4

02 '/4-

0 2^4

02^/4

0 2%

02 '4

O 1%

o%

I

^ 20'

Fig. 6. Contour map of pond 3 showing depth in feet and location of station 6.

284

PACIFIC SCIENCE, Vol. VII, July, 1953

BEACH

O 0 20 40

' ' '

Fig. 7. Contour map of pond 6 showing depth in feet and location of station 11.

it longer than it can on an open ocean coast.

Because of the small amplitude of the lunar
tide in Hawaii and because of the great in-
fluence of winds upon the tide in Kaneohe
Bay, it is 'doubtful if the present data will
permit the accurate establishment of either
the zero tide line or the time differential with
Honolulu tides.

However, for purposes of comparison an
average zero tide line and a time differential
were determined. Readings in the diurnal se-
ries of observations were taken every 15 min-
utes at the expected times of high and low
water. The average zero tide reading was sub-
tracted from these, and the corrected readings
were then used in plotting and drawing the
diurnal tidal curves for Kaneohe Bay (Fig. 9).
In the first three sets of curves, interest was
directed mainly to comparing the curves—
tidal amplitudes as well as the correspondence
of high and low waters. Hence, the initial
high or low water of Kaneohe Bay and Hono-
lulu were made to correspond with regard to
time. The Kaneohe Bay tidal curve for the
December series fitted the Honolulu curve
poorly, especially with regard to correspond-
ence of high and low waters. This was prob-
ably the result of stormy weather with variable
winds during the period studied. The April
and August series compared a little more fa-

vorably. The last set of curves was drawn
mainly to compare the time differential be-
tween Kaneohe Bay and Honolulu. The April
tidal curves were chosen for this comparison
because they displayed the best fit of the
three observations. The average time differ-
ential in this series was 1 hour 24 minutes,
whereas the average time differential as de-
termined in the study was 1 hour 50 minutes,
and the time differential between the theore-
tical Honolulu tide and that of Waikane,
Kaneohe Bay, the closest reference station,
is 1 hour 35 minutes.

STATION 7 O

Fig. 8. Contour map of pond 4 showing depth in
feet and location of station 7.

Seasonal Variations — TsEU

285

TIME

Fig. 9. Comparison of observed Kaneohe and theo-
retical Honolulu tidal curves. The upper three graphs
compare the heights of the tides with the time adjusted
so that the first tide of the cycle coincides. The bottom
graph shows the actual difference in the time of the
tide. The solid lines represent the Kaneohe data, the
broken lines represent the Honolulu data.

METEOROLOGICAL CONDITIONS

Rainfall was measured by means of a stand-
ard rain gauge installed on Coconut Island
in April, 1949 (Fig. 1, C). Measurements were
read at weekly intervals, and the results were
graphed (Fig. 10) to show seasonal variations.
Values on the graph topped by a heavy hori-
zontal line indicate that the rainfall during
the week was greater than 2.10 inches, the
upper limit of the rain gauge. Rainfall ranged
from 0.00 inches to more than 2.10 inches,
with the heaviest rainfall in both years occur-
ring during the winter months.

Air temperatures were recorded by means
of a continuous temperature recorder installed
on April 3, 1949, in the net house (Fig. 1, B).
Daily minimum and maximum temperatures
were recorded and averaged to obtain weekly
minimum and maximum temperatures, which
were graphed to show seasonal variations.
During the period studied, minimum air
temperatures ranged from 19.3 to 24.3°C, and
maximum air temperatures from 23-9 to
30.7°C. Both of the years studied exhibited
a rapid drop of temperature from the highest
in September and October to the lowest in
November and December.

WATER TEMPERATURES

Water temperatures were obtained by means
of a continuous temperature recorder and by
sampling at intervals.

The water temperature recorder was in-
stalled on April 3, 1949, in the net house near
station 2 (Fig. 1, B). Both surface and bottom
water temperatures were taken. Top and bot-
tom values for each day varied very little,
therefore they were averaged, and the results
were averaged again to obtain a weekly mini-
mum and maximum. These weekly tempera-
tures were then graphed to show seasonal
variations. Minimum water temperatures as
recorded on the continuous temperature re-
corder ranged from 21.5 to 27.0°C and maxi-
mum water temperatures from 21.8 to 27.2°C.

286

PACIFIC SCIENCE, Vol. VII, July, 1953

Fig. 10. Comparison of rainfall and chlorinity curves for a 2-year period from February, 1949, to February,
1951. Chlorinity of surface waters is indicated by broken lines, of bottom waters by solid lines.

In the sampling at intervals, at first both
surface and bottom temperatures were taken
weekly at all the stations where the depth of
the water permitted, and bottom temperatures
were taken at those stations where the water
was too shallow to obtain differential samples
with the modified oxygen sampler. As it was
soon found that there was no great tempera-
ture differential between top and bottom at
the shallow stations, 5, 6, 7, and 11, only
one temperature was later taken at these sta-
tions, and as stations 4 and 6 were like stations
3 and 5, respectively, temperatures were taken
monthly at stations 4 and 6. After March,
1950, temperatures were taken monthly at all
stations.

Water temperatures were taken by means
of a thermometer graduated in 1°C divisions
and attached to the modified oxygen sampler.
Temperatures were estimated to the closest
0.1 °C and were graphed to show seasonal
variations (Fig. 11), averages being used when
the differential between top and bottom was
less than 0.5°C. As expected, there was a close
agreement of changes from station to station.

and all stations showed a definite seasonal
change. Deeper stations 1 and 2 showed the
most gradual seasonal changes, and station
10, the very shallow station on the flats,
showed the greatest extremes of temperature
with a range from 20.5 to 33.3°C.

The extreme alternation between high and
low temperatures on successive weeks, most
noticeable in March and April, 1949, at sta-
tion 10, was characteristic of the shallower
stations and was the result of tidal differences.
One week, at the time of sampling, the tide
would be very low, while on the following
week at the same hour it would be high. In
weeks when the tide was low, the shallow
waters were warmed and were not mixed with
the cooler deep waters until the next flooding
tide. In the following week when the tide
was high at the same hour, the warmed waters
were mixed with the cooler water brought in
from the deeper channels.

Under normal conditions the surface waters
tended to be warmer than bottom waters be-
cause of the warming effects of the sun and
the decreased density of warmer water. How-

Seasonal Variations — TsEU

287

ever, during the winter months from Decem-
ber, 1949^ to January, 1950, and from Decem-
ber, 1950, to January, 1951, the periods of
”kona” weather and heavy rainfall (Fig. 10),
a reverse situation was noted, and surface
waters of lower temperatures were observed
(Fig. 11). This was caused by the colder rain
water floating on top of the warmer, more
saline, and therefore more dense, bay water.

In the three 24-hour periods of observation,
a definite diurnal change in temperature was
found with a minimum temperature in the
morning hours from 0500 to 0800 and a max-
imum temperature in the afternoon hours
from 1200 to 1400 (Fig. 12). An increase of
temperature range was noted from the De-
cember to the August series.

CHEMICAL CONDITIONS

Water samples for determining tempera-
ture, chlorinity, and oxygen content were col-
lected simultaneously by means of the modi-
fied oxygen sampler. Water temperatures were
recorded immediately, chlorinity samples were
transferred from the reservoir bottle into ci-

Fig. 11. Comparison of water temperatures for a
year at stations 1, 5, and lo. Temperature of surface
waters shown by broken line, of bottom waters by
solid line.

trate of magnesia bottles for storage, and
oxygen samples were preserved immediately
in the BOD bottles in which they were col-
lected.

Chlorinity samples were analyzed by the
Mohr technique (Thompson et aL, 1950: see
section, ”The Determination of the Chlorin-
.ity of Sea Water”; Kolthoff and Sandell, 1943:
568-570).

All chlorinity values were graphed to show
seasonal variations and to compare some of
the stations (Fig. 13). Averages were used
when the differential between top and bottom
was less than 0.25 parts per thousand. Chlorin-
ities ranged from 14,94 parts per thousand
to 19.88 parts per thousand. Throughout a
greater part of the year, chlorinities did not
show much variation until the great drop in
chlorinity found in the winter months of Janu-
ary, 1950, and December, 1950, and associ-
ated with the onset of heavy rainstorms (Fig.
10). To show the influence of rainfall on
salinity, data were gathered routinely and
compared with the chlorinity values of rep-
resentative station 2 (Fig. 10).

TIME

Fig, 12. Comparison of water temperatures for three
diurnal observations. Temperature of surface waters
shown by broken line, of bottom waters by solid line.

288

Fig. 13. Comparison of chlorinity for a year at sta-
tions 1, 5, and 10. Chlorinity of surface waters shown
by broken line, of bottom waters by solid line.

The surface layers were especially affected
by these rainstorms. In the shallow stations,
5, 6, 7, 10, and 11, where only one sample
was taken, chlorinities dropped to about the
same degree as the surface waters of the deep-
er stations such as 1 and 2. The surface layer
of brackish water lost its separate identity
after about a month, and deep and shallow
water became almost homogeneous but of a
lowered chlorinity. In the winter season of
1949-50, about 2 months were required for
the chlorinity values to resume a normal level
after the heavy rains.

In the 24-hour cycles, no diurnal changes in
chlorinity were observed which could be cor-
related with either day and night or with tides
(Fig. 14).

Oxygen samples were determined by the
Winkler method for dissolved oxygen (Hol-
lister, 1950: see section, "Determination of
Dissolved Oxygen in Fresh and Sea Water”;
Kolthofif and Sandell, op. cit., pp. 614-619)-
All oxygen values were graphed to show sea-
sonal variations and to compare some of the
stations (Fig. 15). Averages were used when
the differential between top and bottom was

PACIFIC SCIENCE, Vol. VII, July, 1953

Fig. 14. Comparison of chlorinity for three diurnal
observations. Chlorinity of surface waters shown by
broken line, of bottom waters by solid line. Part of
August samples were lost, hence break in graph.

less than 0.25 milliliters of oxygen at normal
temperature and pressure per liter of sea water
at 20°C. From the beginning of sampling to
mid-July, 1949, sampling was done in the
early afternoon with the few exceptions noted.
These afternoon values were, on the whole,
higher than the morning values. Values ranged
from 1.10 milliliters per liter to 7.27 milliliters
per liter, these extreme values being the ex-
ception rather than the rule. On the whole,
no marked seasonal pattern was detected, and
diurnal changes were more noticeable.

Oxygen concentration displayed a definite
diurnal change (Fig. 16) in all three 24-hour
cycles, reaching a minimum in the morning
between 0500 and 0700 and a maximum in
the afternoon between 1400 and 1800. This
diurnal change is presumably related to the
biochemical activity of plants producing an
excess of oxygen in hours of daylight and the
consumption of oxygen by both plants and
animals in the hours of darkness.

Saturation values were determined (com-
putations based on Table 24, Harvey, 1928:

Seasonal Variations — Tseu

289

TABLE 2

Diurnal Changes in Oxygen Content

DATE

TIME

OBSERVED

VALUE

COMPUTED

SATURATION

VALUE

PERCENTAGE

SATURATION

1400 maximum

ot/./L.

5.29

ml.lL.

5.08

104

December 27-28, 1949

1800 median

4.70

5.10

92

0500 minimum

4.10

5.14

80

1600 maximum

5.41

5.04

107

April 28-29, 1950

0200 median

4.82

5.11

94

0700 minimum

4.23

5.10

83

1800 maximum

4.82

4.65

104

August 25-26, 1950

2200 median

4.43

4.71

94

0600 minimum

4.02

4.79

84

60) for the minimum, median, and maximum
oxygen content values in each of the 24-hour
cycles. Results tabulated in Table 2 show that
the minimum morning values represent un-
dersaturated waters and the maximum after-
noon values represent supersaturated water.
All three observations show a similar cycle

Fig. 15. Comparison of oxygen content for a year
at stations 1, 5, and 10. Oxygen content of surface
waters shown by broken line, of bottom waters by
solid line. Unless otherwise indicated, samples for the
first 5V2 months were collected in the afternoon, there-
after in the mornings.

with regard to the amount of oxygen dis-
solved in the water.

At monthly intervals for over a year anal-
yses were attempted for the nutrient salts,
phosphates, nitrites, and silicates. As the ana-
lytical methods were colorimetric, attempts
were first made to measure the color intensi-
ties with a Klett-Sommerson photoelectric
colorimeter, test tube model. Because of the

Fig. 16. Comparison of oxygen content for three
diurnal observations. Oxygen content of surface waters
shown by broken line, of bottom waters by solid line.

290

PACIFIC SCIENCE, Vol. VII, July, 1953

limitations of the instrument, attempts were
then made to measure the color intensities
with a Beckman spectrophotometer, model
B. With neither instrument were the tests or
the measuring apparatus sufficiently accurate
to give reliable results with the low concen-
trations of these salts present.

Inorganic phosphates were analyzed by a
modification of the method of Deniges (as
reported by Robinson and Thompson, 1948^:
33-41). From the attempted analyses it can
be concluded that no samples taken during
the year greatly exceeded the value of 0.50
mygram atoms per liter and that it is likely
that most values were lower.

Nitrites were analyzed by a method origi-
nally developed by Peter Griess and later modi-
fied by Ilosvay (as reported by Robinson and
Thompson, 1948^.' 42-48). None of the sam-
ples taken throughout the year gave measur-
able amounts of nitrite; therefore, the nitrite
was either almost entirely absent from the
waters or at least fell consistently below 0.10
mygram atoms per liter.

Silicates were analyzed by a modification
of a method made practical by Dienert and
Wandenbulcke and first applied to oceano-
graphic investigations by Atkins (as reported
by Robinson and Thompson, 1948c.' 49-55).
With regard to the silicates it can be stated
that no samples taken during the year greatly
exceeded the value of 10 mygram atoms per
liter and that it is likely that most values were
lower.

REFERENCES

Harvey, H. W. 1928. Biological chemistry and
physics of sea water, x + 194 pp. The Mac-
millan Company, New York.

Hollister, H. J. 1950. Determination of dis-
solved oxygen in fresh and sea water. [Sep-

arate pagination.] Manual of oceanographic
methods. Canadian Joint Committee on
Oceanography, Pacific Oceanographic
Group, Nanaimo, B. C., Canada. [Mimeo-
graphed.]

Kolthoff, I. M., and E. B. Sandell. 1943.
Textbook of quantitative inorganic analysis.

xvii + 794 pp. The Macmillan Company,
New York.

Marmer, H. a. 1926. The tide, xi + 282 pp.

D. Appleton and Co., New York.
Robinson, Rex J., and Thomas G. Thomp-
son. 1948^. The determination of phos-
phate in sea water. Jour. Marine Res. 7(1):
33-41.

and 1948^. The determination

of nitrites in sea water. Jour. Marine Res.
7(1): 42-48.

and 1948c. The determination

of silicates in sea water. Jour. Marine Res.
7(1): 49-55.

Sverdrup, H. U., Martin W. Johnson, and
Richard W. Eleming. 1946. The oceans,
their physics, chemistry, and general biology.
X + 1087 pp. Prentice-Hall, Inc., New
York.

Thompson, T. G., et al. 1950. The determina-
tion of the chlorinity of sea water. [Separate
pagination.] Manual of oceanographic meth-
ods. Canadian Joint Committee on Oceano-
graphy, Pacific Oceanographic Group, Na-
naimo, B. C., Canada. [Mimeographed.]
United States Coast and Geodetic Sur-
vey. 1949 (also 1950 and 1951). Tide ta-
bles, West Coast, North and South America
{including the Hawaiian Islands'). 206 pp.
U. S. Department of Commerce, Washing-
ton.

Welch, Paul S. 1948. Limnological methods.

xviii + 381 pp. The Blakiston Co., Phila-
delphia and Toronto.

New Plant Records from the Eastern Caroline Islands, with a
Comparative Study of the Native Plant Names^

S. F. Glassman2

From a botanical standpoint, the Eastern
Caroline Islands (Fig. 1) are poorly known.
The floras of the high islands of this group
— Ponape, Kusaie, and Truk—have been stud-
ied to a reasonable degree. However, the low
coral islands, about 22 in number, have re-
ceived little botanical attention. Only five of
these low islands-— Pingelap, Kapingamaran-
gi, Satawan, Nomwin, and Nukuoro — have
been reported on in any detail. For the re-
maining islands, there are either only a few
scanty records of plant collections or no
known records at all.

During the summer of 1949, the writer
spent 3 months collecting plants on several
islands in the Eastern Carolines. Most of the
time was devoted to Ponape, the flora of
which I have treated in two previous papers
(Glassman, 1952; in press). Approximately 1
day was spent on each of the following low
islands or atolls: Mokil, July 21; Pingelap,
July 22; and Ant, August 10. This study is
based principally on these three islands. Most
of the records listed here are new, as previous
plant collections from these islands have been
so sparse. This paper, therefore, is a list of
new records of species with their accompany-
ing native names, new records of native names,
and changes in nomenclature of species pre-
viously collected from Mokil, Pingelap, and
Ant.

^ The writer is grateful to the Pacific Science Board
of the National Research Council for the fellowship
grant which enabled him to make this trip; to the
Office of Naval Research for financing the trip; to the
University of Oklahoma for supplementary financial
assistance; and to the specialists mentioned below for
determination of specimens.

" Division of Biological Sciences, University of Illi-
nois, Chicago, Illinois. Manuscript received April 14,
1952.

Specimens of vascular plants collected were
deposited in the United States National Her-
barium, Bernice P. Bishop Museum, and
Bebb Herbarium, University of Oklahoma.
Nonvascular plants have been placed in the
Crypto gamic Herbarium of the Chicago Nat-
ural History Museum.

’.Pulusuk.

10^ N-

J^amonuito

JMurili}

East- J^a^u

E^iVfta

'■Cosap

EMatncliJi . ^ ^

Etal

■ Satax

•Kl

. OrohlH

JPafy n . ponape

.fiingekp

'Kusmc

I Mcrthc^

Jlupuoyo

JVAUTICAI MILES
too 0 ,oo

■ . KqpiMgaxxiavaM^i

Fig. 1. Map of Eastern Caroline Islands.

Each island is treated separately with a brief
geographical and historical description fol-
lowed by a catalogue of species for that island.
Each entry in the catalogue of species is based
on specimens collected or observed by the
writer or on names of species published in
taxonomic articles. Synonyms which pertain
to each particular island are also listed. Speci-
mens cited have been either verified person-
ally or determined by specialists in certain
groups as follows: Dr. H. N. Moldenke, New
York Botanical Garden, Verbenaceae; Dr. W.
H. Wagner, Jr., University of Michigan, ferns;
Dr. E. Drouet, Chicago Natural History Mu-
seum, algae; Mr. E. B. Bartram, Bushkill,
Pennsylvania, mosses; and Dr. M. Eulford,
University of Cincinnati, liverworts. Native

291

292

PACIFIC SCIENCE, Vol. VII, July, 1953

names are shown in quotation marks and are
spelled exactly as they sounded to the writer.
Economic uses for some of the species are
also mentioned.

It is interesting to note that each island in
the Eastern Carolines has its own language.
As a matter of fact, on Ponape there are dis-
tinct language differences between different
districts (e.g., Kid and Net) of the island.
Following the catalogue of species for Mokil,
Pingelap, and Ant, native names mentioned
in this paper are tabulated for comparison
with those of the high islands in the Eastern
Carolines — Ponape, Kusaie, and Truk. In-
spection of the table reveals striking differ-
ences as well as similarities in the languages
of these islands.

MOKIL

Mokil Atoll is located approximately 6°40 'N
and 159°47'E. It is about 2 miles long and 1
mile wide and is about 88 miles southeast of
Ponape, 60 miles northwest of Pingelap, and
180 miles northwest of Kusaie. Mokil com-
prises three individual islets— Manton, Kalap,
and Urak. The writer visited only Manton and
Kalap.

Mokil, formerly known as Duperrey Island
and Wellington Island, was discovered by
Duperrey on the corvette 'Ta Coquille” in
1824 (U. S., 1944). In 1838, Lesson wrote
an account of this voyage. Andersson (1854),
botanist with the frigate "Eugenie,” and Skog-
man (1856), also with the same ship, visited
the island in 1852 and subsequently wrote
summaries of the voyage in which the vege-
tation and economic plants are mentioned.
In 1854, Hammet included some notes on the
vegetation in his narrative of the voyage of
the "Serpent.” Moss (1889) gave a brief ac-
count of the vegetation of the island, which
he visited while touring Micronesia in 1886;
and Christian (1899^, b) mentioned Mokil
plants briefly in his accounts of the Caroline
Islands. Thilenius (1927) and Filers (1934)
included some information on the vegetation
and listed a number of plants from a general

survey of the Caroline Islands made by the
Germans between 1908 and 1910.

It appears that the only known previous
collection of plants from Mokil was made by
Ohba in the 1930’s. Several specimens of this
collection were cited by Kanehira (1935).
Murphy, who included Mokil in his geo-
graphical account of the Eastern Carolines
(1949), sent St. John a list of some 25 native
names of plants recorded from the island.
From these names, St. John compiled a ten-
tative list of species (unpublished), most of
which have been corroborated by the present
writer. Both Weckler (1949) and Murphy
(1950) have reported on the agriculture of
Mokil.

A total of 73 species of vascular plants, 5
species of algae, and 2 species of bryophytes
were either collected or observed by the writer.
Numbers 2393 to 2601 inclusive were col-
lected on Manton islet, numbers 2602 to 2633
were taken from Kalap islet. Of the vascular
plants, 34 are indigenous, 12 are crop plants,
16 are ornamentals, and 11 are adventive
weeds. Vernacular names were obtained from
a native guide named Loren.

NONVASCULAR PLANTS
Algae

Collinsiella tuberculata Setch. & Gardn.

2387. On seashore and in shallow water.
Dictyosphaeria favulosa (Ag. ) Dene.

2397^. In shallow sea water.

Halimeda opuntia (L.) Lamx.

2397. In shallow sea water.

Microdictyon okamurai Setch.

2397h. In shallow sea water.

Scytonema figuratum Born. & Flah.

2388. On coral soil, common.

Bryophytes

Microlejeunea bullata (Tayl.) Evans.

2386. On coconut tree, frequent.
Splachnobryum luzonense Broth.

"Lim,” 2612. Terrestrial in coconut grove,
common.

Plant Records from Caroline Islands— Glassman

293

This species was previously thought to be
endemic to the island of Luzon.

VASCULAR PLANTS

'Polypodiaceae

1. Microsorium scolopendria (Burm.)
CopeL, Univ. Calif., Publ. Bot. 16: 112.
1929.

"Kamkam,” 2626. Epiphyte in coconut
grove, common.

2. Nephrolepis sp.

"Boh,” 2611. In coconut grove, terres-
trial and epiphytic, common (sterile).

3. Pteris tripartita Sw., Schrad. Jour. Bot“
67. 1801.

"Papa-ni,” 2624. In coconut grove, ter-
restrial, common.

Cycadaceae

4. Cycas circinalis L., Sp. PL, 1188. 1753.
Observed as an ornamental; introduced
from Ponape.

Fig. 2. Mokil. Cassytha filiformis, a climbing parasitic
plant, on Gmttarda spedosa.

Annonaceae

5. Annona muricata L., ibid., 536.

"Triika shai.” Observed as a cultivated
plant; probably introduced from Ponape.

Lauraceae

6. Cassytha filiformis L., ibid., 35. (Fig. 2.)
"Cossagos,” 2599. Climbing parasite,
along strand, common.

Hernandiaceae

7. Hernandia sonora L., ibid., 981.
"Pingaping.” Observed as a strand plant;
only one medium-size tree was seen.

Crassulaceae

8. Bryophyllum pionatum (Lam.) Kurz,
Jour. As. Soc. Beng. 40: 52. 1871.
"Lamalam.” Observed as a cultivated
plant.

Portulacaceae

9. Portulaca oleracea L., Sp. PL, 445.
1753.

"Ubijon,” 2618. Weed in waste places,
flowers yellow.

10. Portulaca samoensis v. Poelln., Fedde
Repert Sp, Nov. 33: 163. 1933.
"Ubijon,” 2616. Common along strand,
prostrate, flowers yellow,

Amaranthaceae

11. Achyranthes aspera L., Sp. PL, 204.
1753.

"Suga-dugodok,” 2629. Weed in coconut
grove, common.

12. Gomphrena globosa L., ibid., 224.
"Pahwis,” 2613. Planted, bracts pink.

Lythraceae

13. Pemphis acidula Forst., Char. Gen. PL,

68, pi. 34. 1776.

"Kahengy,” Observed as a common
strand plant.

294

PACIFIC SCIENCE, Vol. VII, July, 1953

Onagraceae

14. Jussiaea suffruticosa L., Sp. PL, 388.
1753.

"Kiree.” Observed in Cyrtosperma swamp.
Nyctaginaceae

15. Pisonia sp.

"Mehs.” Observed as tree along the
strand.

Murphy (1949) states that the leaves of
this plant are used as a green manure in
Cyrtosperma swamp.

16. Mirabilis jalapa L., Sp. PL, 177. 1753.
"Four o’clock.” Observed as an orna-
mental plant.

Caricaceae

17. Carica papaya L., ihid., 1036.
"Mamiyap.” Observed as cultivated
plant.

Combretaceae

18. Terminalia litoralis Seem., FI. Vit., 94.
1865.

"Win.” 2602. Along strand, tree 20 feet
high, fruit red, flowers white, common.

Guttiferae

19. Calophyllum inophyllum L., Sp. PL,

513. 1753.

"Isho.” Observed as a tree 50 to 60 feet
high along the seashore.

Tiliaceae

20. Triumfetta procumbens Forst. f..
Prodr., 35. 1786. (Fig. 3.)

"Konup,” 2601. Along strand, procum-
bent, flowers yellow, common.

Malvaceae

21. Hibiscus tiliaceus L., Sp. PL, 694. 1753.
"Pah.” One small tree seen; probably
introduced from Ponape.

22. Sida fallax Walp., Nov. Act. Nat. Cur.
19, suppl. 1: 306. 1843.

"Kau,” 2610. Planted, flowers orange.

Fig. 3. Mokil. Triumfetta procumbens, a decumbent
plant along the beach.

23. Thespesia populnea (L.) Soland. ex
Correa, Ann. Mus. Paris 9: 290. 1807.
"Peneh.” Observed as a frequent tree
along the strand.

Euphorbiaceae

24. Euphorbia heterophylla L., Sp. PL, 453.
1753.

2608. Planted in garden.

25. Phyllanthus niruri L., ibid., 981.
"Limar-mah-pom,” Observed as a weed.

26.

27.

Leguminosae

Bauhinia monandra Kurz, Jour. As. j

Soc. Beng. 42: 73. 1873. |

"Flamboyant.” Observed as an orna- ,|

mental tree with pink flowers. ||

Cassia occidentalis L., Sp. PL, 377. |

1753. ;

"Apschoh,” 2632. Waste places, un-
common. i

Plant Records from Caroline Islands — Glassman

295

28. Poinciana pulcherrima L., ihid., 380.
‘"Shimatada,” 2633. Planted, tree 10 feet
high, flowers yellow.

29. Vigna marina (Burm.) Merr., Inter-
pret. Herb. Amb., 285. 1917.

"Taut-tul,” 2591. Along strand, decum-
bent, flowers yellow, common.

Casuarinaceae

30. Casuarina equisetifolia L., Amoen.
Acad. 4: 143. 1759.

Observed as an ornamental tree; prob-
ably introduced from Ponape.

Moraceae

31. Artocarpus altilis (Parkinson) Fosb.,
Wash. Acad. Sci., Jour. 31: 95. 1941.
"Mai.” "Maipah” is a seeded variety
with serrate leaves, whereas "moshabo-
rok” is seedless with deeply incised
leaves. Breadfruit is very common on
Mokil and is one of the most important
food crops. According to Christian
(1899^), the wood of one variety, "mai-
mat,” is highly prized for house building;
and the wood of other varieties is used
in the construction of canoes.

32. Ficus tinctoria Forst. f.. Prodr., 76. 1786.
"Coain,” 2594. Tree 30 feet high, fre-
quent.

Urticaceae

33. Fleurya ruderalis (Forst.) Gaudich.,
Freyc. Voy. Bot., 497. 1830.
"Nin-kotokot,” 2628. In coconut grove,
frequent.

34. Pilea microphylla (L.) Liebm., Vi-
densk. Selsk. Skr. 5: 302. 1851.

"Reh,” 2619. On stone ledge and in
coral soil, common.

35. Pipturus argenteus (Forst. f. ) Wedd.,
DC. Prodr. 16: 235. 1869.

"Ormuh,” 2620. Tree 20 feet high, in
coconut grove, frequent.

Sapindaceae

36. Allophylus timorensis (DC.) BL, Rum-
phia 3: 130. 1847.

"Kitak,” 2621. Along strand, tree 15 feet
high, flowers white, frequent.

Araliaceae

37. Nothopanax fruticosum (L.) Miq., FI.
Ind. Bat. 1: 765. 1856.

Observed as an ornamental shrub.

Apocynaceae

38. Nerium oleander L., Sp. PL, 209. 1753.
"Uilianter.” Observed as an ornamental
tree.

39. Plumeria rubra L., ihtd., 209.

"Po maria.” Observed as an ornamental
tree.

Asclepiadaceae

40. Asclepias curassavica L., ibid., 215.
"Truka-keree,” 2607. Near native dwell-
ing, uncommon.

Rubiaceae

41. Guettarda speciosa L., ibid., 991. (Fig.

2.)

Fig. 4. Mokil. Flowers of Scaevola frutescens, an abun-
dant shrub along the beach.

296

PACIFIC SCIENCE, VoL VII, July, 1953

Fig. 5. Mokil. Pure stands of Scaevola frutescens with
Cocos nucifera in background.

"Eet,” 2398. Along strand, tree 30 feet
high, flowers white, common.

42. Hedyotis biflora (L.) Lam., Tabl. En-
cycl. 1: 272. 1791.

'’Mussen-buel,” 2622. At base of coco-
nut tree, flowers white, uncommon.

43. Ixora casei Hance, Walp. Ann. Bot.
Syst. 2: 754. 1852.

"Kasaw,” 2603. Cultivated tree, un-
common.

44. Morinda citrifolia L., Sp. PL, 176. 1753.
''Wehmpul.” Observed as a tree along
the strand.

Compositae

45. Vernonia cinerea (L.) Less., Linnaea

4: 291. 1829.

"Mussen-buel,” 2623. Weed in waste
places, flowers blue.

46. Wedelia biflora (L.) DC. ex Wight,
Contrib. Bot. Ind., 18. 1834.
''Morishish,” 2393. Along strand, sub-
scandent, flowers yellow, uncommon.

Campanulaceae

47. Hippobroma longiflora (L.) G. Don,
Gen. Hist. Diehl. PL 3: 717. 1834.

”Ehmah.” Observed as an ornamental
herb.

Goodeniaceae

48. Scaevola frutescens (Mill.) Krause,
Pflanzenr. 4 (277): 125. 1912. (Figs. 4,

5. )

”Romok,’’ 2393. Dominant shrub along
strand, flowers white.

Boraginaceae

49. Cordia subcordata Lam., Tabl. Encycl.
1: 421. 1791.

"Kanaw.” Observed as a small tree along

the strand. The trunk is used in making
canoes.

50. Messerschmidia argentea (L.) John-
ston, Jour. Arn. Arb. 16: 164. 1935. (Fig.

6. )

”Sisin,” 2396. Along strand, tree 30 feet
high, flowers white, frequent.

Solanaceae

51. Capsicum frutescens L., Sp. PL, 189.
1753.

”Chilee.” Observed as an ornamental.

Fig. 6. Mokil. Flowering branch of Messerschmidia

argentea, a frequent beach tree.

Plant Records from Caroline Islands — Glassman

297

Fig. 8. Mokil. Native inhabitant with rhizome and
leaf of Cyrtosperma chamissonis, one of the most import-
ant food plants, banana plants in background.

found scattered along the edges of the
Cyrtosperma swamp and are not a major
source of food on Mokil.

57. Colocasia esculenta (L.) Schott ex
Schott & Endl., ibid., 18.

"Chawa.” Five varieties are recognized
by the natives: "'chawasa,” "peeter,” "ta-
wah,” "pehmeru,” and "chawa-n-jaban.”

58. Cyrtosperma chamissonis (Schott)
Merr., Phil. Jour. Sci., Bot. 9: 65. 1914.
(Fig. 8.)

"Muen.” The following varieties were
recorded: "chaleng welek,” ’'shimeeten-
malou,” ''shumbung-unu,” ’’shigogi,”
and "shirieh.”

This species is probably the most. im-
portant food plant on Mokil.

Convolvulaceae

52. Ipomoea gracilis R. Br., Prodr., 484.
1810.

"Ohlop,” 2606. Trailing along strand,
milky juice, frequent.

Verbenaceae

53. Premna gaudichaudii Schau., DC.
Prodr. 11: 631. 1847.

"Subuk,” 2392. Tree 30 feet high, flowers
white, along strand, frequent.

Hydrocharitaceae

54. Thalassia hemprichii (Ehrb.) Aschers.,
Naturl. Pflzfam. 2 (1): 254. 1889- (Fig.
7.)

"Walap,” 2627. In shallow water, com-
mon.

Musaceae

55. Musa paradisiaca L., Sp. PL, 1043. 1753.
*'Wus.” A few banana trees were seen
near a native dwelling. (Fig. 8.)

Araceae

56. Alocasia macrorrhiza (L.) Schott ex
Schott & FndL, Melet. Bot. 1: 18. 1832.
"Wut”; ’'wut-en-mokil” is a poisonous
variety.

This species and the following one were

Fig. 7. Mokil. Thalassia hemprichii, a common aqua-
tic flowering plant growing in shallow salt water.

298

PACIFIC SCIENCE, VoL VII, July, 1953

Amaryllidaceae
59- Crinum sp.

"'Kiup.” Observed as an ornamental with
purple flowers and leaves.

60. Zephyranthes rosea Lindl., Bot. Reg.,
pL 821. 1824.

"Kiup.” Observed as an ornamental.
Palmae

61. Cocos nucifera L., Sp. PL, 1188. 1753.
(Figs. 5, 10.)

"Ni.” Two varieties were recorded by the
writer: "ni-sikisik” and "atoll.” Murphy
has also recorded the following varieties:
"nerium,” "nikarus,” and "sukabish.”
He also obtained the following names
for various stages of development of the
coconut fruit: "pen,” green nut; "arng,”
ripe nut; and ”par,” sprouted nut. Coco-
nut is undoubtedly the most abundant
and most useful tree on the island, all
parts of the plant being used. Most of
these trees were planted by the early in-
habitants. According to Christian (1899)
the consumption of coconut toddy on
Mokil was prohibited by the American
missionaries.

Pandanaceae

62. Pandanus sp.

Fandamis cylindricus Kanehira, Bot. Mag,
Tokyo 49: 63, fig. 3. 1935.

P. hosinoi Kanehira, ibid., 103, fig. 8.

P. jaluitensis Kanehira, ibid., 103, fig. 9.

P. macrocephalus Kanehira, ibid., 428.

"Kebar.” The natives recognize 19 dif-
ferent varieties which are as follows:
"toboh-tin,” "luaram,” "unmang,”

' ' schwai-pue-ep, ’ ’ "arawan-en-mehluh, ’ ’
"meh-kilikil,” "seepwerik,” "shoni-me-
neyoh,” "pen-bashu,” "musikel,”
‘Tope,” "ninikeh-tahk,” "mokosokos,”
"shaleh- show-wushuh,” "ruboh,” "uhn-
besch,” "enaidah-erewehn,” "shoh-muin-
shong,”and "shee-lahweh.” Pandanus is
a very important plant on Mokil. Nuts of

some of the varieties are edible, and leaves
of most of the varieties are used in thatch-
ing. According to Christian (1899^),
leaves of Pandanus are also used for
canoe sails.

For reasons discussed elsewhere (Glass-
man, 1952), the present writer does not
accept the names of species of Pandanus
described by Kanehira which are listed
in synonymy above.

Taccaceae

63. Tacca leontopetaloides (L.) O. K.,
Rev. Gen. PL, 704. 1891.

"Mokomok.” Observed as a cultivated
and escaped plant in the coconut grove.
The tubers are an important source of
food.

Cyperaceae

64. Cyperus javanicus Houtt., Nat. II Hist.
13: Aanwyz. Plaat. (1), pi. 88, fig. 1. 1782.
"Mordong,” 2609. Along strand and in
coconut grove, common.

65. Fimbristylis atollensis St.John, Pacific
Sci. 6: 145. 1952.

"Puror-en-toge,” 2601. Along strand, fre-
quent.

Gramineae

66. Digitaria pruriens Biise, Miq. PL

Jungh., 379. 1854.

"Muhkarak,” 2627. In coconut grove,
common.

67. Eleusine indica (L.) Gaertn., Fruct.
Sem. PL 1: 8. 1788.

"Puror-en-tuke,” 2611. Weed in waste
places.

68. Eragrostis amabilis (L.) Wight & Arn.,
ex Hook. & Arn., Bot. Beech. Voy., 251.

1841.

"Reh,” 2613. Weed in coconut grove,
common.

69. Lepturus repens (Forst. f.) R. Br.,
Prodr., 207. 1810.

"Limon-gisek,” 2600. Along strand,
common.

2631. Frequent in coconut grove.

Plant Records from Caroline Islands — Glassman

299

70. Oplismenus compositus (L.) Beauv.,

Ess. Agrost., 54. 1812.

"Moh-long,” 2603. In coconut grove,
common.

71. Paspalum vaginatum Sw., Prodr. Veg.
Ind. Occ., 21. 1788.

’'Olee-sakai,” 2630. In coconut grove,
common.

72. Saccharum officinarum L., Sp. PL, 54.
1753.

'Tuh.” Observed as a cultivated plant.

73. Thuarea involuta (Forst. f.) R. Br.,
Prodr., 197. 1810.

"Muhkarak,” 2390. In coconut grove,
common.

Additional species, not collected or ob-
served by the writer, for which tentative
names are given by St. John from native
names recorded by Murphy:
Barringtonia asiatica (L.) Kurz. "We.”
Barringtonia racemosa (L.) BL "Kan-
ge.”

Intsia bijuga (Colebr.) O. K. "Kebuk.”
Ochrosia oppositifolia (Lam.) K.

Schum. "Kacshpar.”

PINGELAP

Pingelap Atoll (Fig. 1) is situated at 6°13'N,
160°E, and is about 60 miles southeast of
Mokil and 144 miles northwest of Kusaie.
It is 2 miles in length and 1.5 miles in width.
The atoll consists of three islets, Tugulo,
Takai, and Pingelap. Only the last islet was
visited by the writer.

Pingelap, also referred to as Musgrave,
MacAskill, Pelelep, and Pingoulap, was dis-
covered by Musgrave with the ship "Sugar
Cane” in 1793 (U. S., 1944). Moss (1889)
gave a brief account of the vegetation of Pin-
gelap which he visited while touring Micro-
nesia in 1886; and in 1899 Christian men-
tioned Pingelap plants in his account of the
Caroline Islands. Thilenius (1927) and Filers
(1934) included some information on the
vegetation and listed a number of plants
based on a general survey of the Caroline
Islands made by the Germans between 1908
and 1910. In 1949 Murphy reported on the
agriculture of the island. The first detailed
paper on the vascular plants of Pingelap was
written by St.John (1948) who listed 57 spe-

Fig. 9. Pingelap. Mangrove swamp showing seedlings of Rhizophora mucronata and surface roots of Sonneratia
caseolaris.

300

cies for the island. In the present treatment,
the writer has added 21 new records of vas-
cular plants, several additional native names,
and some nomenclatural changes. In addi-
tion, one species of algae and two species
of mosses are recorded. Native names were
obtained from Soas, who served as a guide
for both the writer and St. John.

NONVASCULAR PLANTS
Algae

Cladophora membranacea (Ag. ) Harv.
"Lim,” 2638. Along shore and in shallow
water.

Mosses

Ectropothecium monumentorum (Duby)

Jaeg.

2639. On trunk of coconut tree, common.
Leucophanes glauculum C, M.

2643. On coconut tree, common.

VASCULAR PLANTS
Polypodiaceae

1. Pteris tripartita Sw., Schrad. Jour. Bot.,
67. 1801.

"Peypey-eni,” 2631. Coconut grove, ter-
restrial, abundant.

Lauraceae

2. Cassytha filiformis L., Sp. PL, 35. 1753.
"Cossagos,” 2644. Along strand, climb-
ing parasite, common.

Hernandiaceae

3. Hernandia sonora L., ibid., 981.
"Pingapin," 2634. Along strand, tree 15
feet high by 2.5 feet in diameter, com-
mon. Many seedlings were found be-
neath the parent plants. The tree trunks
are used for canoes by the natives.

Rhizophoraceae

4. Bruguiera conjugata (L.) Merr., Phil.
Jour. Sci., Bot. 9: 118. 1914.

PACIFIC SCIENCE, Vol. VII, July, 1953

"Sol,” 2622. Back of mangrove swamp,
tree 40 feet high by 1 foot in diameter.
Only one tree was seen, but the writer was
informed that this species is fairly com-
mon on other parts of the island. The
wood is used in construction of houses,
and a dye from the fruits is utilized for
blackboards. This is the third species of
mangrove recorded for Pingelap, the
others being Khtzophora mucronata Lam.
and Sonneratia caseolaris (L.) Engl. (5.
alba Sm.). (Fig. 9.)

Malvaceae

5. Hibiscus tiliaceus L., Sp. PL, 694. 1753.
"Kalau,” 2633. Along strand, tree 50 feet
high, common. Fibers from the bark are
used in making rope, fish line, hats, and
baskets, and the leaves are often utilized
in washing clothes.

Euphorbiaceae

6. Codiaeum variegatum (L. ) Bl. var. pic-
tum (Lodd.) MuelL-Arg., DC. Prodr.

15: 1119. 1866.

"Kurotong.” Observed as an ornamental
plant; introduced from Kusaie.

Leguminosae

7. Canavalia microcarpa (DC.) Piper,
Biol. Soc. Wash., Proc. 30: 177. 1917.
"Nimelitop,” 2646. Along strand, trail-
ing, flowers pink, common. Plant parts
are used as medicine to aid in childbirth.

8. Inocarpus fagiferus (Parkinson) Fosb., |

Wash. Acad. Sci., Jour. 31: 95. 1941. |

"Marrup,” 2637. Planted, tree 15 feet |
high, introduced from Ponape. j

9. Peltophorum sp. j

"Seh-muatah,” 2638. Planted, tree 8 feet |
high, flowers yellowish orange; intro- ||
duced from Kusaie. )

10. Vigna marina (Burm.) Merr., Inter-
pret. Herb. Amb., 285. 1917. !

"Sau-tul.” St. John recorded the native i
name of this species as "nimelitop,”

Plant Records from Caroline Islands — Glassman

301

which should refer to Canavalia micro -
carpa listed above,

Moraceae

11. Ficus tinctoria Forst. f., Prodr. 76. 1786.
''Kawain.” St. John listed this entry as
Ficus sp.

Urticaceae

12. Fleurya ruderalis (Forst.) Gaudich.,
Freyc. Voy. Bot., 497. 1830.
"Ne-kirrir-ir,” 2640. In coconut grove,
flowers reddish brown, common.

13. Pilea microphylla (L.) Liebm,, Vi-
densk. Skr. 5: 302. 1851.

'’Re” (St.John); "wahpul.”

Rubiaceae

14. Hedyotis biflora (L.) Lam., Tabl. En-
cycl. 1: 272. 1791.

"Musenibuil,” 2655. Base of coconut
tree, flowers white, uncommon.

15. Ixora casei Hance, Walp. Ann. Bot.
Syst. 2: 754. 1852.

Ixora carolinensis (Val.) Hosokawa aff.
var. typica Fosb., Bish. Mus. Occ. Pap.
15: 221. 1940; St. John, Pacific Sci. 2:
112. 1948.

Compositae

16. Ageratum conyzoides L., Sp. PL, 839.
1753.

"Pokaniko,” 2660. Weed in waste places,
frequent. Leaves of this species are com-
bined with those of Ocimum sanctu7n L.
and coconut oil and used as a skin lotion.

17. Vernonia cinerea (L.) Less., Linnaea
4: 291. 1829.

"Musenibuil,” 2656. Weed in coconut
grove and waste places, flowers blue,
common.

Boraginaceae

18. Cordia subcordata Lam., Tabl. Encycl.
1: 421. 1791.

"Ikoh-ik,” 2645. Along strand, tree 50
feet high, flowers orange, frequent.

Convolvulaceae

19. Ipomoea aff. gracilis R. Br., Prodr.,
484. 1810.

"Wahlap,” 2648. Along strand, milky
juice, frequent (sterile).

Verbenaceae

20. Premna gaudichaudii Schau., DC.
Prodr. 11: 631. 1847.

Premna integrifolia L., Mant. PL 2: 252.
1771; St.John, Pacific Sci. 2: 112. 1948.
"Sokuk” (St.John); "sobuk.”

Labiatae

21. Ocimum sanctum L., Mant. PL 1: 85,
1767.

"Teeko,” 2661. In front of dwelling, un-
common.

Musaceae

22. Musa paradisiaca L., Sp. PL, 1043. 1753.
In addition to the native names recorded
by St. John, the following were obtained
by the present writer: "Saipan” and "Ti-
nian.”

Araceae

23. Alocasia macrorrhiza (L. ) Schott ex
Schott & End!., Melet. Bot. 1: 18. 1932.
A number of specimens were seen in the
Cyrtosperma swamp. Of the two varieties
recognized, "wut” is edible and "seh-
buken” is poisonous.

Palmae

24. Cocos nucifera L., Sp. PL, 1188. 1753.
(Fig. 12.)

"Ni” (St.John). Three varieties are rec-
ognized by the natives: "ni-wi-sahsah,”
"ni-mah-uwah,” and "ni-sol.”

Pandanaceae

25. Pandanus sp. (Fig. 10.)

In addition to the native names obtained
by St. John, the following have been re-
corded: "eisesieu-el,” "nanagaisak,” and
"sonomuneyah.” The latter two are prob-

302

PACIFIC SCIENCE, VoL VII, July, 1953

Fig. 10. Pingelap. Native market place showing construction of house. Roof exterior consists of coconut leaves,
posts and cross timbers are from breadfruit wood, lashing is coconut fiber, and thatching of roof interior and
mats in foreground are made from Fandanus leaves.

ably the same as "nanagaisal” and ”so-
numei” of St. John.

Gramineae

26. Centotheca lappacea Desv., Nouv. Bui.
Soc. Philom. Paris 2: 189. 1810.
"Moh-lung,” 26,52. Coconut grove, fre-
quent.

27. Digitaria pruriens Biise, Miq. PI.
Jungh., 379. 1854.

”Reh,” 2641. In coconut grove, common.

28. Eleusine indica (L.) Gaertn., Fruct.

Sem. PL 1: 8. 1788.

"Rosakai,” 2642. Waste places, common.

29. Paspalum vaginatum Sw., Prodr. Veg.
Ind. Occ. 21. 1788.

"Unenekisekik,” 2649- In coconut grove,
common. The leaves of this species and
those of Pisonta are used in the Cyrto-
sperma swamp as a fertilizer.

ANT

Ant Atoll (Fig. 1) is located at approxi-
mately 6°47'N and 158°l'E, and is 6 miles

long by 3.5 miles wide. It is about 8 miles
southwest of Ponape and about 300 miles
east of Truk. Ant comprises 12 individual
islets, only the largest of which, Nikalap, was
visited by the writer.

Ant Atoll, also known as Andema, Fraser,
and Hand, was discovered by Quiros in 1591
(U. S., 1944). Of the few botanical papers
dealing with the island, Meinicke (1876) and
Hambruch (1929) briefly discuss the vegeta-
tion and food plants. There are no known
detailed accounts of the vascular flora of Ant;
however, a few papers concerning the algae
of the island have been published by Yamada
(1944^, b). I

ji

A total of 58 species of vascular plants, 3 !j
species of algae, and 1 species of moss were J
either collected or observed by the writer. Of 1
the vascular plants, 42 are indigenous, 11 are ||
crop plants, 1 is an ornamental, and 4 are
adventive weeds. Vernacular names were ob-
tained from Shokeen, a native guide.

Plant Records from Caroline Islands — GlaSSMAN

303

NONVASCULAR PLANTS
Algae

Enteromorpha prolifera (O. F. MuelL) J.

Ag.

''Muh-so-pweeset/’ 2834. Along beach and
in shallow water.

Laurencia mariannensis Yamada.

’’Moo-nos,” 282 L On shore and in water,
common.

Scytonema hofmannii Born. & Flah.

”Karan-ahl,” 2822. Coconut grove, on
stones and logs, common.

Mosses

Ectropothecium monumentorum (Duby)

Jaeg-

"Limalim,” 2831. On dead coconuts, com-
mon.

VASCULAR PLANTS
Polypodiaceae

1. Microsorium scolopendria (Burm.)
CopeL, Univ. Calif. Publ. Bot. 16: 112.

1929.

”See-see.” Observed as an epiphyte.

2. Nephrolepis exaltata (L.) Schott, Gen.
Fib, pi. 3. 1834.

’’Ahmereh,” 2828. Terrestrial, abundant
in coconut grove.

Lauraceae

3. Cassytha filiformis L., Sp. PL, 35. 1753-
”Wahlee-mah,” 2809. Climbing along
beach, common.

Hernandiaceae

4. Hernandia sonora L., ibid., 981.
’’Ahkaran.” Observed along the beach.

Piperaceae

5. Piper ponapense C. DC., Engl. Bot.
Jahrb. 56: 502. 1921.

’’Ahnuck.” Observed as an epiphyte.

. Capparidaceae

6. Crataeva speciosa Volkens, Engl. Bot.

Jahrb. 31: 463. 1902.

’’Afoosh,” 2824. Tree 30 feet high, flow-
ers cream, fruit edible, introduced from
the Mortlocks.

Lythraceae

7. Pemphis acidula Forst., Char. Gen. PL,
68, pi. 34. 1776.

”Truh-kees,” 2810. Along beach, shrub
10 feet high, flowers white, common.

Onagraceae

8. Jussaiea suffruticosa L., Sp. PL, 388.
1753.

’’Ayah.” Observed in moist places.
Nyctaginaceae

9. Pisonia sp.

”Muk,” 2833. Coconut grove, tree 30
feet high, uncommon (sterile).

Caricaceae

10. Carica papaya L., Sp. PL, 1036. 1753.
’’Mohmiyap.” Observed as a cultivated
plant.

Barringtoniaceae

11. Barringtonia asiatica (L.) Kurz, Jour.

As. Soc. Beng. 45: 70. 1876.

”OoL” A few trees were observed along
the seashore.

Combretaceae

12. Terminalia catappa L., Mant. PL 1:
128. 1767.

’’Uhsass.” Observed along the beach.

13. Terminalia litoralis Seem., FI. Vit., 94.
1865.

”Kin,” 2818. Along beach, tree 20 feet
high, flowers white, fruit red, common.

Guttiferae

14. Calophyllum inophyllum L., Sp. PL,

513. 1753.

’’Ruckiss.” Observed along the strand.
Malvaceae

15. Hibiscus tiliaceus L., ibid., 694.
’’Kileefah.” Observed along the beach.

304

PACIFIC SCIENCE, Vol. VII, July, 1953

16. Thespesia populnea (L.) Soland. ex
Correa, Ann, Mus. Paris 9: 290. 1807.
"Puhneh.” Observed along the strand.

Leguminosae

17. Canavalia microcarpa (DC.) Piper,
Biol. Soc. Wash., Proc. 30: 177. 1917.
"Fin-kalau,” 2812. Along beach, trailing,
flowers purple, frequent.

18. Derris trifoliata Lour., FI, Cochinch.,
433. 1790.

’’Up,” 2819, Liana on beach and in coco-
nut grove, flowers white, abundant.

19. Intsia bijuga (Colebr. ) O. K., Rev. Gen.
PL 1: 192. 1891.

"Choyo,” 2829. Tree 50 feet high, along
beach, uncommon.

20. Vigna marina (Burm.) Merr., Interpret.
Herb. Amb., 285. 1917.

’'Ohloo,” 2804. Trailing along beach,
flowers yellow, common.

Moraceae

21. Artocarpus altilis (Parkinson) Fosb.,
Wash. Acad. Sci., Jour. 31: 95. 1941.
"Mey,” "lukuwol,” "mey-tahit,” "mey-
tee,” "mey-n-patak,” and "mey-os” are
the varieties recognized by the natives.
Breadfruit is a very important food plant
on this island.

22. Ficus carolinensis Warb. apud Schum.
& Lauterb., FI. Deutsch. Schutzg. Siidsee,
242. 1905.

"Kilee-ant,” 2832. In coconut grove, tree
25 feet high by 4 inches in diameter, fre-
quent.

23. Ficus tinctoria Forst. f.. Prodr., 76. 1786.
"Ahwahn,” 2823. Common in coconut
grove, tree 50 feet high, fruit orange,

Urticaceae

24. Fleurya ruderalis (Forst.) Gaudich.,
Freyc. Voy. Bot., 497. 1830.
"Ani-gusgus,” 2813. In coconut grove,
frequent.

25. Pipturus argenteus (Forst. f. ) Wedd.,
DC. Prodr. 16: 235. 1869.

"Orohmah,” 2811. In coconut grove,
tree 40 feet high, flowers green, common.

26. Procris pedunculata (Forst. f. ) Wedd.
ibid., 14: I9I. 1869^.

”Koo-mit.’' Observed as a liana in coco-
nut grove.

Rutaceae

27. Citrus aurantifolia (Christm.) Swingle,
Jour. Wash. Acad. Sci. 3: 465. 1913.
''Kuruhkur.” Observed near a native
dwelling.

Sapindaceae

28. Allophylus timorensis (DC.) BL, Rum-
phia 3: 130. 1847.

"Nguh,” 2813. Shrub along beach and
in coconut grove, flowers white, berries
red, common.

Apocynaceae j

29. Ochrosia oppositifolia (Lam.) K. |

Schum., Natiirl. Pflzfam. 4 (2): 156. j
1895. I

"Oomah.” Observed as a strand plant. |

Rubiaceae

30. Guettarda speciosa L., Sp. PL, 991.
1753.

"Mohsor,” 2807. Along beach, flowers
white, common.

31. Morinda citrifolia L., ibid., 176.

"Nen.” Observed along the beach.

Compositae

32. Vernonia cinerea (L.) Less., Linnaea
4: 291. 1829.

Observed as a weed in waste places.

33. Wedelia biflora (L. ) DC. ex Wight,
Contrib. Bot. Ind., 18. 1834.

"Ahtu-guaht.” Observed as a semi-
scandent shrub along the strand.

Goodeniaceae

34. Scaevola frutescens (Mill.) Krause,
Pflanzenr. 4, 277: 125. 1912.

"Eenut.” Observed as a common shrub
along the strand.

Plant Records from Caroline Islands — Glassman

305

Boraginaceae

35. Cordia subcordata Lam., Tabl. Encycl.
1: 421. 1791.

"Ahlew," 2820. Along beach, tree 50 feet
high by 2 feet in diameter, flowers orange,
common.

36. Messerschmidia argentea (L. ) John-
ston, Jour. Arn. Arb. 16: 164. 1935.
"Amunusut,” 2823. Along beach, tree 10
feet high, flowers white, frequent.

Solanaceae

37. Capsicum frutescens L., Sp. PL, 189.
1753.

’’Mwik,” 2805. Planted, flowers greenish
yellow.

38. Nicotiana tabacum L., ibid., 180.
’’Tamak,” 2817. Planted, flowers pink.

39. Physalis minima L., ibid., 183.

‘'Ti,” 2803. Weed in coconut grove, flow-
ers greenish yellow, common.

Convolvulaceae

40. Ipomoea aff. gracilis R. Br., Prodr., 484.
1810.

"Ahfahmus.” Sterile plants were ob-
served along the beach.

Verbenaceae

41. Clerodendrum inerme (L.) Gaertn.,
Fruct. Sem. PL 1: 271. 1788.

"Oolah.” Observed along the seashore.

42. Premna gaudichaudii Schau., DC.
Prodr. 11: 631. 1847.

’'Orr.” Observed in coconut grove.

Araceae

43. Alocasia macrorrhiza (L.) Schott ex
Schott & EndL, Melet. Bot. 1: 18. 1832.
"Keh.” This species and the following
were of frequent occurrence in the Cyrto-
sperma swamp.

44. Colocasia esculenta (L.) Schott ex
Schott & EndL, ibid., 18.

''Oht.” ”Ohtinyap” and ‘'ohton kusaie”
are varieties distinguished by the natives.

45. Cyrtosperma chamissonis (Schott)

Merr., Phil. Jour. Sci., Bot. 9: 65. 1914.
"Fulah.” The following varieties are rec-
ogni2ed: "onokokung,” "simetun,”
"ponon,” and ’’sinaitah.” This is a very
important starch plant.

Amaryllidaceae

46. Crinum sp.

’'Kiup,” 2835. Along beach, flowers pink-
purple, leaves purple on underside.

Palmae

47. Cocos nucifera L., Sp. PL, 1188. 1753.
"Nu.” The following varieties are dis-
tinguished: ”nu-garaw,” ’'nu-shah,” "nu-
saesael,” and "nu-mow.” The coconut
plantations on this island are operated
by Oliver of Nanpei from Ponape.

Pandanaceae

48. Pandanus sp.

’'Fahss.” ’'Fah-sheh-rah,” 'Tah-tin-
wahl,” "sil-lau,” and "lubush” are some
of the varieties, 2816. Frequent along
beach, tree 30 feet high, trunk branched,
male inflorescence, flowers cream, musky
odor.

Taccaceae

49. Tacca leontopetaloides (L.) O. K.,
Rev. Gen. PL, 704. 1891.

"Mokomok.” This plant is very abun-
dant in the coconut groves and is prob-
ably the most important starch crop on
the island.

Cyperaceae

50. Cyperus sp.

"Pukahret.” Observed along the beach
and in the coconut grove.

51. Cyperus javanicus Houtt., Nat. II Hist.
13: Aanwyz. Plaat. (1), pi. 88, fig. 1.
1782.

”Kusakus,” 2814. Along beach and in
coconut grove, common.

306

PACIFIC SCIENCE, Vol. VII, July, 1953

52. Fimbristylis atollensis St.John, Pacific
Sci. 6: 145. 1952.

"Apuson,” 2830. In coconut grove, com-
mon.

Gramineae

53. Digitaria pruriens Biise, Miq. PL

Jungh., 379. 1854.

’Tahtil-muahn,” 2806. In coconut grove,
common.

54. Eleusine indica (L.) Gaertn., Fruct.
Sem. PL 1: 8. 1788.

"Puhkahr.” Observed as weed in waste
places.

55. Eragrostis amabilis (L. ) Wight & Arn.
ex Hook. & Arn., Bot. Beech. Voy., 251.
1841.

'Tahtil.” Observed as a common weed.

56. Lepturus repens (Forst. f. ) R. Br.,
Prodr., 207. 1810.

"Fahtil-muahn,” 2827. Common in coco-
nut grove.

57. Paspalum conjugatum Berg., Act. Helv.
Phys. Math. 7: 129, pi. 8. 1762.
”Fahtil-rawfut,” 2808. In coconut grove,
frequent.

58. Thuarea involuta (Forst. f. ) R. Br.,
Prodr., 197. 1810.

"Fahtil,” 2826. Common in coconut
grove.

ETHNOBOTANY

Species and native names of plants occur-
ring on Mokil, Pingelap, and Ant are tabu-
lated for comparison with those of the high
islands in the Eastern Carolines— Kusaie, Po-
nape, and Truk (Table 1). The islands are
listed in geographical order with the south-
ernmost island, Kusaie, listed first, and the
northernmost, Truk, listed last. Most of the
vernacular names were recorded by the present

writer during visits to the various islands
(except Truk); however, some names for
Truk, Mokil, and Pingelap were taken from
St. John (1948); some from Truk, Kusaie,
and Ponape were derived from Fosberg (1946) ;
and some others from Truk were extracted
from Elbert (1947), Hosokawa (1937), and
Kramer (1932). Asterisks preceding certain
vernacular names (e.g., ”po maria” and "chi-
lee”) indicate the name is of foreign origin.
Omission of native names in some columns
indicates one of three things : there is no rec-
ord for the species, there is no record for the
native name, or there is no known native name
for the species. An analysis of Table 1 as to
similarity or identicalness of vernacular names
reveals linguistic affinities between the islands.

Table 2 shows the number of similar or
identical native names of the total narries re-
corded for the two islands being compared
in each instance; the percentage of similarity
and the number of native and exotic plants
are also indicated. Ant appears to show the
greatest linguistic affinity with Truk. Of 50
vernacular names recorded, 30 are identical
or similar. Ponape, with 14 similar or iden-
tical names out of 45, shows the next most
important relationship with Ant. Pingelap (9
of 44), Mokil (8 of 42), and Kusaie (4 of 38),
follow in decreasing order of importance.
Most of the present-day inhabitants of Ant
are descendents of people who came from the
Mortlock Islands (Fig. 1) at the beginning of
the present century.

As would be expected, Mokil shows the
greatest linguistic affinity with Pingelap be-
cause of their proximity. Of 56 native names
in common, 39 are the same or modified. For
both Mokil and Pingelap, linguistic relation-
ships for the remaining islands are in the fol-
lowing order: Ponape (24 of 46 and 26 of 48),
Kusaie (12 of 40 and 10 of 44), and Truk
(14 of 50 and 13 of 53).

Plant Records from Caroline Islands — GlassmaN

307

TABLE 1

Vernacular Plant Names from Six Islands in the Eastern Carolines

ISLANDS AND VERNACULAR PLANT NAMES

plant species

KUSAIE

PINGELAP

MOKIL

PONAPE

ANT

TRUK

Microsorium

kaim-kaim,

kiteu

kamakam

kitieu

see-see

chichi,

scolopendria

kemkem

onnum

Nephrolepis spp.

fah, fuah

pues, pue

boh

rah til,

ahmereh

amare, emere.

rawtil

amere

Pteris tripartita

peypey-eni

papa-ni

.

Annona muricata

truka shai

sae

sasaf

Cassytha filiformis

cossagos

cossagos

kohtokot-

shau

wahlee-mah

anau

Hernandia sonora

pung-pung

pingapin

pingaping

pingapin

ahkaran

aguran, agran,
akurang, mosul

Piper ponapense

konok

ahnuck

eresi

Crataeva speciosa

apoot

afoosh

abut, abuts,
afuts, afuch,
apuch

Bryophyllum

lamalam

lamalam

pinnatum

Pemphis acidula

kasugel

kai-i-ni

kahengy

ngi

truh-kees

engi, eegi

Jussiaea sp.

tener-aak

kuri

kiree

telurik

ayah

aunenipuin,

nigaulen,

likeinenpul

Pisonia sp.

mas

mehs

muk

nok, mok,
makku

Carica papaya

hiss, es

kaineap

mamiyap

momiap

mohmiyap

kipau

Barringtonia

bus-bus

wi

we

we

ool

kun, azan,

asiatica

kuun

Barringtonia

kaiengal

kange

winmarr

sun, guon.

racemosa

kun

Terminalia

shufehf,

tepop

tipop

uhsass

as, asas

catappa

Terminalia

sarf

sin

win

kin

sin

litoralis

Bruguiera

shrahl, alol

sol

shorn,

buru bun, on.

conjugata

rhom

ong, oon,
wong

Calophyllum

eetuh, eet

sepang

isho

isyo

ruckiss

rakich, ijau.

inophyllum

mosur, wangu,
legitu, fregits

Triumfetta

konop

konup

kiuin, liodot

procumbens
Hibiscus tiliaceus

lo

kalau

pah

kalau

kileefah

sapuo, silfa,
sinifo, syapo,
shirifa

Sida fallax

kao

kau

sioi le

Thespesia

panu,

penne

peneh

pone

puhneh

pona, okuran,

populnea

pehnuh,

pakeena

'

likokon

Codiaeum

kurotong

kurutun

variegatum
Phyllanthus niruri

limar-mah-

limair-poh

negamaur.

pom

nikammour

Canavalia

shoh-tul-

nimelitop

fin-kalau

chochon.

microcarpa

muet

wonuka

308

PACIFIC SCIENCE, VoL VII, July, 1953

TABLE 1 — continued

ISLANDS AND VERNACULAR PLANT NAMES

PLANT SPECIES

KUSAIE

PINGELAP

MOKIL

PONAPE

ANT

TRUK

Derris trifoliata and

ohsheh,

keh-ohror.

up

pis-uenipot

D. elliptica

op

up

(Ponape, Kusaie)
Inocarpus fagiferus

dark

marrup

marrup

annilla, kurrak

Intsia bijuga

choyo.

choyo

kuren, tuamis.

show

nityanmis

Poinciana

repawtin,

shimatada

sem-tah

simmata

pulcherrima

rapotin

Vigna marina

sratol,

shroh-tulah

sautul

taut-tul

tansilituh

ohloo

wonuka

Artocarpus altilis

mos

mai

mai

mai

mey

mai

Ficus carolinense

she, shra

ayau

kilee-ant

au

Ficus tinctoria

sheh,

kawain

coain

neen

ahwahn

awan, aon,

konyah

apuris, eplis,
mouk

Fleurya ruderalis

ne-kirririr

nin-kotokot

sau mwal

ani-gusgus

ansifichnu,
en ugotnu

Pilea microphylla

re, wahpul

reh

Pipturus argenteus

halkoh

oroma

ormuh

oromah

arome

Procris pedunculata

pahkeh

koo-mit

atatagur,

gumudj,

kimuit,

kemmet,

nitatagul,

adatagul

Allophylus

lah

kitak

kitak

ungeh

nguh

ngo, nga

timorensis

(A. ternatus for Ku-
saie and Ponape)
Citrus sp.

Asclepias

muh

kimeme

truka-keree

karrer

kuruhkur

kurukur

curassavica

Ochrosia

kitee

oomah

uma, uwa

oppositifolia
Plumeria rubra

fohr

*po maria

*po maria

*po maria

sour

Guettarda speciosa

koin-lahk

eles

eet

eet, ith

mohsor

mosor

Hedyotis biflora

musenibuil

mussen-

sing,

•

buel

nisarfonu

Ixora casei

kalsru,

galusa,

kahl-shuh

kalesu

kasaw

kartieu

atiu, achen

Morinda citrifolia

ee

obul

wehmpul

weypul.

nen

nen, nobur,

wumpul

arin

Ageratum

pokaniko

pusen-koh

amshiip,

conyzoides

opolopon,

oponupon

Vernonia cinerea

musenibuil

mussen-

anachuko.

buel

troboasu,

ennetoku

Wedelia biflora

eekeh,

kisuwell

morishish

ingkah.

ahtu-guaht

atuat, eadiat.

agaia

ungkeh

aduduad

Scaevola frutescens

kushosh

ramek

romok

eenut

eenut

not, amoloset,
fremes

Cordia subcordata

eekwahk

ikoh-ik

kanaw

eekoh-eek

ahlew

anau, alau.

anna

Plant Records from Caroline Islands — Glassman

309

TABLE 1 — continued

ISLANDS AND VERNACULAR PLANT NAMES

FLAW 1 arUL-lLS

KUSAIE

PINGELAP

MOKIL

PONAPE

ANT

TRUK

Messerschmidia

shuhshun

sesen

sisin

titin

amunusut

amoneset,

argentea

emoloset,

chen,

amarashet

Capsicum frutescens

*pepper

*chilee

*sele

mwik

muik, moek

Nicotiana tabacum

*tamak

pis-suba

Ipomoea gracilis

oah

wahlap

ohlop

sui-oomp

ahfahmus

ruk, frugrug

Clerodendmm

kyawak

ilau

ilau

oolah

ulo, apuoch,

inerme

etiu, pucherik,
edin, abot

Premna

fienkeek,

sobuk,

subuk

tubuk,

orr

lior, nior,

gaudichaudii

fienket

sokuk

orr

umukau.

(P, corymbosa
for Ponape)
Ocimum saactum

hariin, oarin

teeko

kahterin

umgau, niyol

Thalassia

kahp

walat

walap

olot, ohlot

mut

hemprichii

Musa paradisiaca

ush, oune

wis

wus

ut

uch

Alocasia

unog,

wut

wut

oht

keh

ka, ke, puna

macrorrhiza

wunock

Colocasia

kohtahk,

sawa

chawa

sawah

oht

sawa, onni

esculenta

taka

Cyrtosperma

pashok

muiang

muen

muahng

fulah

pula, pashon,

chamissonis

bura

Crinum sp.

kief-

kiep

kiup

kiup

kiup

kiop, kiaup,

fashfash

kiyop

Zephyranthes rosea

kief-shuck

kiep

kiup

pileep

Cocos oucifera

nu

ni

ni

ni

nu

nu

Pandanus sp.

muen

kipai

kebar

kipar

fahss

kepar, fadj

Tacca

mokmok,

mugamuk

mokomok

mokmok

mokomok

mokomok

leontopetaloides

mukmuk

Cyperus javanicus

mahtok

sapasap

mordong

use

kusakus

amana,

(this is

moirer,

name for
large sedge
on Ponape)

nikaunoun

Fimbristylis sp.

rosakai

puror-en-

apuson

fedil, puker,

toge

fetin, umula

Centotheca

masha-

moh-lung

lappacea

shruck

Digitaria prurieos

reh

muhkarak

fahtil-

muahn

Eleusine indica

rosakai

puror-en-

reh-takai,

puhkahr

toke

rea takai

Eragrostis amabilis

rosakai

reh

shoh-maleh

fahtil

Lepturus repens

rosakai

limon-

fahtil-

gisek

muahn

Paspalum

olee-sakai

uneneki-

timoor

vaginatum

sekik

Saccharum

tuh

seu

tuh

seu

wou

officioarum
Thuarea involuta

mokarak

muhkarak

fahtil

unom

310

PACIFIC SCIENCE, VoL VII, July, 1953

TABLE 2

Similar or Identical Native Names of the Total Names Recorded
BETWEEN Islands and Percentages of Similarty.

ANT

MOKIL

PINGELAP

Kusaie

4 of 38 (11 per cent)

(native 1, food 2, ornamen-
tal 1)

12 of 40 (30 per cent)

(native 6, food 4, ornamen-
tal 2)

10 of 44 (22.7 per cent)

(native 6, food 2, ornamen-
tal 2)

Pingelap

9 of 44 (20.5 per cent)

(native 3, food 4, economic

1, ornamental 1)

39 of 56 (70 per cent)

(native 19, food 8, adventive

6, economic 1, ornamental 5)

Mokil

8 of 42 (19 per cent)

(native 2, food 4, economic

1, ornamental 1)

Ponape

14 of 45 (31 per cent)

(native 7, food 5, economic

1, ornamental 1)

24 of 46 (52 per cent)

(native 11, food 9, adventive

1, ornamental 3)

26 of 48 (54 per cent)

(native 15, food 8, adventive

1, ornamental 2)

Ant

Truk

30 of 50 (60 per cent)

(native 21, food 7, economic

1, ornamental 1)

14 of 50 (28 per cent)

(native 6, food 5, economic

1, ornamental 2)

13 of 53 (24.5 per cent)

(native 6, food 5, economic
1, ornamental 1)

REFERENCES

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mit der Schwedischen Kriegsfregatte Eugenie'
1851-1833. 384 pp. Leipzig.

Christian, F. W. 1899^. The Caroline Islands.
412 pp. Methuen & Co., London.

1899^. Exploration in the Caroline

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Eilers, a. 1934. Inseln um Ponape: In Ergeb-
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8: 1-464. Friederichsen, de Gruyter & Co.,
Hamburg.

Elbert, S. H. 1947. Trukese-English and Eng-
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Fosberg, F. R. Ms. Botanical report on Micro-
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Glassman, S. F. 1952. The flora of Ponape.
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1952. List of nonvascular plants from

Ponape, Caroline Islands. Amer. Midland
Nat. 48: 735-740.

Hambruch, P. [Ed.] 1929. Elf Jahre in Aus-
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Berlin. [A free translation of O’Connell,

J. F. A residence of 11 years in New Hol-
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Hosokawa, T. a. 1937. A preliminary ac-
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cronesian plants. Kyushu Imp. Univ., Dept. \
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Kramer, A. 1932. Truk. In Ergebnisse Siidsee- |
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Lesson, R. P. 1838. Voyage autour du monde j

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Mosquitoes of the Genus Uranotaenia in the
Solomon Islands (Diptera: Culicidae)

John N. Belkin^

INTRODUCTION

In a previous paper (Belkin, 1950) a revision
of the culicine mosquitoes of the Solomon
Islands was initiated, based on the extensive
collections made during the American occu-
pation of these islands from 1942 to 1946.
General information on the background, lo-
calities, collecting and rearing methods, and
location of the material as well as a treatment
of the genus Tripteroides will be found in that
paper.

The present study of the genus Uranotaenia
is based on the examination of 3,967 speci-
mens of seven species, distributed as to stages
in the following manner: 1,459 adults, 415
pupae, and 2,093 larvae. Of this material 107
individuals, including all species, are repre-
sented each by a larval (fourth instar) and a
pupal skin and the corresponding adult. The
importance of such individual rearings cannot
be overemphasized in the genus Uranotaenia
as, in over one half the collections where
association of stages was made only through
mass or lot rearings, at least two species were
involved. The individually reared material
forms the basis of this study. The complete
chaetotaxy of the larva and pupa of 10 indi-
viduals of each species was studied, and in
the associated adults all the external morpho-
logical features were observed in detail, except
as noted under each species. From the study
of the individually reared material, distin-
guishing or diagnostic characters of each spe-
cies were selected, and these in turn were
checked in all the remaining material together

^ Division of Entomology, University of California
at Los Angeles. Manuscript received December 11,
1951.

with a notation of any striking variation in
other characters.

Terminology

In the course of this study it became evi-
dent that the terminologies currently in use
in descriptive culicidology have failed to keep
abreast of, and sometimes have ignored com-
pletely, independent findings in comparative
anatomy. Accordingly I have attempted to
bring up-to-date and to homologize the chae-
totaxy of the mosquito larva (Belkin, 1951:
678-698) and pupa (Belkin, 1952: 115-130).
These revised homologous terminologies are
used in this paper. For the adults, I am using
the same terminologies as in the previous
paper (Belkin, 1950: 208), although it is evi-
dent that these terminologies need to be re-
viewed also. The few changes I have intro-
duced here are self-explanatory.

Descriptions and Illustrations

In deference to the general practice, I have
described the holotypes and allotypes of new
species instead of drawing up composite spe-
cies descriptions from the study of the entire
type material as in the past. The immature
stages are described from the pupal and larval
exuviae of the holotype, thus insuring definite
correlation. While such a procedure prevents
the inclusion of more than one species in each
sex of a new species and is desirable from the
standpoint of definitely associating a nom-
inal species with a specimen, I cannot agree
that it is advisable taxonomically. Such de-
scriptions of types, to be useful, must be
exhaustive and are necessarily lengthy. There-
fore, they almost invariably restrict the author
to much shorter discussion on variation than

312

Uramtaenia in Solomon Islands — Belkin

is needed to characterize the populations de-
scribed and discourage the publication of
much valuable material which could easily
be incorporated in the original composite
description. Furthermore, this practice tends
to the selection of extremes for description
and encourages hasty decisions. In mosqui-
toes it is practically impossible to find a spe-
cimen in a condition to show all the important
features, particularly in reared material. When
males are selected for holotypes, microscopic
preparations of genitalia and appendages de-
stroy some diagnostic characters.

To alleviate in part these disadvantages
without undue repetition in the text, I have
resorted to comparative tables and composite
figures. In the case of the latter the same
arguments may be used in favor of figures
of types as in the case of descriptions. Com-
posite figures based on a study of variation
in the new form are of considerably greater
importance in the identification of other spe-
cimens than a graphic representation of char-
acters already described in detail for the type
specimen, whose departure from the norm
can be determined by comparison with the
composite figure.

I have made the descriptions, tables, and
illustrations as complete as time and the mate-
rial allowed. They may appear unnecessarily
detailed and repetitious. Elsewhere it has been
pointed out (Belkin, 1951: 678) that, in the
immature stages of mosquitoes, culicidolo-
gists have largely restricted themselves to a
few diagnostic characters, particularly in non-
anophelines, and have neglected a multitude
of other morphological features which appear
to be at least as useful. In a revision such as
this, limited both geographically and taxo-
nomically, it is not possible to determine the
important characters and leave out the non-
essential features since it is essentially a pre-
liminary attempt and no comparable pub-
lished study is available at the present. It is
my intent to continue such studies as it is
becoming evident, even in such a limited
field, that in addition to those already in use

313

excellent group characters are to be found in
the thoracic and abdominal chaetotaxy of the
larvae and the cephalothoracic chaetotaxy of
the pupae. As every species treated here be-
longs to a distinct species group and adequate
material of related forms from adjacent areas
was not available for study, little can be said,
as far as specific diagnostic characters are con-
cerned, except that it appears that the larval
and pupal chaetotaxy may be of considerable
value if it is studied quantitatively as well as
qualitatively. For the present I have had to
use a very few disjunctive characters to dis-
tinguish the new species described here be-
cause of the paucity of material and inade-
quate descriptions. Whether the forms de-
scribed here are species or subspecies cannot
be decided without further comparative stud-
ies. I prefer to consider them for the present
as distinct species. In any event, I hope that
it will be possible for other workers to recog-
nize the forms described here and also to test
the usefulness of some of the characters in
the diagnosis of related species.

In the descriptions, strictly generic char-
acters are not repeated since they are indicated
under the discussion of the genus at the be-
ginning of the paper. In the larva the mouth-
parts were not studied at all, and in the female
the genitalia were also neglected. Otherwise
all characteristics previously described were
investigated.

In the tables on the setal branching, the
mean and other statistics were not determined
as the samples were not adequate to provide
significant figures except in cases where the
desired information was evident without com-
putations. It appears that mathematical treat-
ment of multiple characters of chaetotaxy may
prove a valuable tool in diagnosis of related
species. For the present I have confined the
data to an indication of the extremes of varia-
tion and of the "usual number” of branches
in the order of frequency. The latter is the
mode alone or the mode followed by the next
class (or classes) if it is one half or more of
the mode. In general the "usual number(s)”

314

PACIFIC SCIENCE, VoL -VII, July, 1953

represents 75 per cent or more of the total
frequencies observed. In dendritic hairs with
15 or more branches the figures are only ap-
proximate, as the branching varies greatly and
is difficult to observe accurately.

The illustrations are semidiagrammatic in
nature, prepared with the aid of an ocular
reticule, and are composites to show the
"usuaE’ condition, an approximation of the
mode, on the basis of the observations and
measurements of the individually reared mate-
rial. The figures of the female thorax leave
out all detail of the mesonotum, except scale
pattern, and are very inaccurate in the outline
of the head as they were drawn from pinned
specimens. In the male genitalia the anal seg-
ment has been left out completely as it shows
no useful characters, being apparently com-
pletely unsclerotized. In the details of the legs
only pertinent features are indicated. The out-
lines of the unsclerotized portions of the larval
thorax and abdomen are only approximate,
whereas those of the larval head and terminal
abdominal segments and pupal structures are
somewhat more accurate. All elements of
chaetotaxy are shown in as accurate propor-
tions as could be determined.

Acknowledgments

In addition to the individuals mentioned
in the first paper of this series, the following
have been of great assistance to me in the
various phases of this work: F. E. Baisas of
the Department of Health, Manila; Graham
Held, William A. McDonald, and Roy Pence
of the University of California at Los Angeles.

Abbreviations

ADULTS. The standard abbreviations used
by Edwards (1941: 7-9) for the thorax and
those used by Matheson (1944: 12-17) for
the male genitalia are followed. The leg seg-
ments are designated by a combination of a
Roman numeral with an abbreviation of the
segment as follows: I C (prothoracic coxa),
II TR (mesothoracic trochanter), III F (meta-
thoracic femur), I Tib (prothoracic tibia),

II Tar (mesothoracic tarsus), III CL (meta-
thoracic pretarsal claw). In the case of the
tarsus an Arabic numeral is appended to in-
dicate the subsegment as in: III Tar 1 (prox-
imal segment of metathoracic tarsus), II Tar
5 (distal segment of mesothoracic tarsus).
The measurements of the leg segments are
given in comparative figures, using the fore
tibia as the basic unit.

IMMATURES. The following abbreviations
for body regions and special features are used:
A—antenna

C— head (larva), cephalothorax (pupa)
CS— comb scale
G — genital lobe
M — mesothorax

MT — marginal teeth of anal saddle

P— prothorax

PD — paddle

PT—pecten tooth

S — siphon

T--metathorax

I-VIII — segments of abdomen

IX — '*anal flap” of pupa

X— segments X-XII

The individual elements of the chaetotaxy
are indicated by prefixing the number of the
hair to the abbreviation for the segment or
structure as in: 1-C (head hair No. 1), 6-II
(hair 6 on abdominal segment II), 5-IV-VI
(hair 5 on abdominal segments IV, V, and
VI), 3-5-II (hairs 3, 4, and 5 on abdominal
segment II), 2, 4-1 V, VI (hairs 2 and 4 on
abdominal segments IV and VI).

The type of branching is abbreviated thus:
b (branched) when the branches arise at or
near base; f (forked) when the branching takes
place at or beyond the middle of the hair;
d (dendritic) when the branching is secondari-
ly dichotomous or irregularly dendritic.

GENERAL CONSIDERATIONS

Genus Uranotaenia

1891. Uranotaenia Lynch Arribalzaga, La Pla-
ta. Univ. Museo La Plata, Rev. 1: 375,
2: 163-164. Type species: Uranotaenia

Uranotaenia in Solomon Islands — Belkin

pulcherrima Lynch Arribalzaga, 1891;
Las Conchas, Buenos Ayres (the second
of two species, by selection of Neveu-

Lemaire, Soc. Zool. France, Mem. 15:
227. 1902).

1905. Anisocheleomyia Theobald, Entomolo-
gist 38: 52-53. Type species: Anisoche-
leomyia nivipesThtohAd, 1905; Queens-
land (the first of two species, by selec-
tion of Brunetti, Indian Mus. Calcutta,
Rec. 10: 55. 1914).

1905. Pseudouranotaenia Theobald, Jour.
Econ. Biol. 1: 33. Type species: Pseudo-
uranotaenia rowlandi Theobald, 1905;
New Amsterdam, British Guiana
( = Uranotaenia nataliae Lynch Arribal-
zaga, 1891). Monobasic.

1912. Pseudo ficalhia Theobald, Linn. Soc. Lon-
don, Trans., (2) Zool., 15: 89-90. Type
species: Pseudoficalbia pandani Theobald,
1912; Mahe, Seychelles (the first of two
new species, two other species men-
tioned, fide Howard, Dyar, and Knab,
Mosq. North, Central Amer. and West
Indies 4: 899. 1917).

About 130 valid species of Uranotaenia are
recognized at the present time, largely from
the Old World tropics but with representa-
tives in all zoogeographical regions. The
group is so distinctly characterized in the
adult stage that only three additional generic
names have been proposed for this consider-
able assemblage of species. The individual
species offer such striking differences in or-
namentation and special modifications that
only about 35 additional trivial names have
been proposed which are now considered to
be synonyms, some of which may have to be
resurrected in the future. This unusual situa-
tion is partially due to the neglect of this
genus by culicidologists as the few workers
who have studied the group in the tropics
have found it to be represented by a large
number of species even in limited areas. It
appears from the present work that some of
the extremely widespread species are actually
complexes of closely related forms. No mono-

315

graph of any kind has been published on this
genus as yet.

The taxonomic distinctiveness of Uranotae-
nia in the adult stage has led some workers,
notably Dyar (1928: 415), to recognize a sep-
arate tribe for this genus, but more recent
workers have not followed this action. No
subgenera have been recognized since Ed-
wards (1932: 97) synonymized the four gen-
eric names proposed for this group, although
he and subsequent authors have used Pseudo-
ficalhia as a group name for the nonornamented
species of Uranotaenia. The only attempt to
divide the genus into natural groups was made
by Edwards (1941: 43-44) for Ethiopian spe-
cies and was based on adult characters only.
Although these groups have many characters
in common, they are apparently distinct to a
much greater degree in the larval and pupal
stages. The presence of extremely different
types of larvae and pupae in this genus has
not been appreciated by culicidologists in
general, and this has led to such statements
as "Why did Uranotaenia stand still in an
evolutionary sense while Aedes gave rise to
the largest and most successful group of sub-
arctic mosquitoes" (Ross, 1951: 132). There
appear to be several different evolutionary
lines in Uranotaenia, some of which have in-
vaded the same ecological niches, such as
water in living plants, in dead plant material,
in tree holes, etc. The immature stages as well
as the detailed morphology of many such
forms are unknown at the present. Little can
be done in the classification of this group
until these gaps are filled, as such ecological
situations are notorious for the development
of parallel structures in unrelated forms.

In the Solomons there are three distinct
groups of species, two of which are closely
related and very distinct from the third. In
our present state of world knowledge of Ura-
notaenia it is impossible to determine whether
or not these groups are of subgeneric rank,
and rather than add other names to synonymy
I prefer only to bring out the differences and
similarities at this time.

316

PACIFIC SCIENCE, Vol. VII, July, 1953

Adult Characters

In the Solomons, adults of Uranotaenia can
be recognized superficially from all other mos-
quitoes by the following combination of char-
acters: small size; straight proboscis (usually
swollen apically); pulvilli absent; squama
bare; cell R2 always distinctly shorter than
vein R2+3; vein lA strongly curved apically
and ending at level of or before the separation
of Cui and Cu2; wing membrane with minute
microtrichia, not visible under a magnifica-
tion of 50 X; palpus extremely short in both
sexes (stated to be 2-segmented but not seg-
mented in Solomon Island forms).

Edwards (1941: 41-43) gave an excellent
characterization of the genus in the adult
stage. The following features noted in the
Solomon Island species should be added.

Head: Frontal tuft well developed in some
species; orbital bristles (not counting frontal
pair) often conspicuously interrupted, ar-
ranged 1:3; palpus not segmented, composed
of a single piece in both sexes, between 0.10
and 0.05 of proboscis.

Thorax: One to several propleurals; one
posterior pronotal; one spiracular or none;
one pre-alar; upper mesepimeral may be ab-
sent; dorsocentral bristles strong, acrostichals
weak to moderate, supra-alars strong.

Legs: Male claws of mid leg with weaker
claw sometimes almost as long as stronger
claw, sometimes apparently absent, stronger
claw with a tooth in U. atra; in both sexes
apex of fore and mid tibia with specialized
bristles and scales forming tufts or combs.

Wing: No bristles on base of R or Sc.

Male genitalia: Segment VIII long, forming
a broad band at base of genitalia proper, terg-
ite without hairs in all Solomon Islands spe-
cies; mesosome with ventral bridge in some
species.

Edwards (1941: 43-44) recognized four
groups in the Ethiopian region on the basis
of thoracic and wing scaling and characters
of the male genitalia. He did not attempt to
correlate these differences with larval or pupal

specializations. His groups B and D are not
represented in the Solomon Islands. I have
changed his nomenclature from group to sec-
tion to distinguish them from species groups.
Apparently all but one species {JJ. quadrima-
culata) from the Solomons belong to section
A of Edwards on the basis of adult structures.
The section was characterized as follows:
"Scutal scales mostly narrow but a supra-alar
stripe of broad scales; apn scaly; wings usu-
ally with some white scales in lines. Termi-
nalia (where known) with tergite bare, its
lateral corners produced into pointed process-
es or rounded knobs; style short and rather
stout; Ip with strong spines.” Since it appears
that the New World species (Lane 1943:
137-161), including the type species of the
genus, belong to this section although form-
ing a distinct subsection, it may be considered
as representing the typical Uranotaenia. In the
Old World, section A is a complex one con-
sisting of several subsections. In the Solomons
U. wysockii is a member of such a subsection
differentiated from the other species chiefly
by adaptive characters in the larva and pupa
and appearing very similar in the adult stage.
I propose to recognize in the Solomons sub-
section Al for the generalized species breed-
ing in ground water and subsection A2 for
U. wysockii breeding in plants.

U. quadrimaculata superficially fits into sec-
tion C of Edwards which was characterized
as follows: "Scutal scales all narrow, apn de-
void of scales. Terminalia with tergite bare,
more or less produced in middle, but not at
corners; style long and slender; Ip with small
spines.” Scales are present on apn in U. quad-
rimacidata, and as far as I can determine from
descriptions it has a much shorter cell R2 than
any Ethiopian member of this section. In ad-
dition the pre-alar area is not distinctly sep-
arated from the sternopleuron, an unusual
feature in the genus. In the immature stages
U. quadrimaculata and related Papuan forms
appear very distinct from the Ethiopian spe-
cies. The type species of Pseudoficalbia belongs
to section C, but because of these differences

Uranotaenia in Solomon Islands— Belkin

317

I do not feel justified in using it for Aus-
tralasian species. Instead, for the present I
prefer to use subsection C2 for the Papuan
forms related to U. nigerrima and U. quad-
rimaculata.

It is probable that other distinctive groups
of Uramtaenia await characterization among
the Oriental and Australasian species, partic-
ularly the nonornamented forms, some of
which are undoubtedly closely related to or-
namented forms whereas others are very dif-
ferent. Until these are studied, nothing but
speculations can be made about the relation-
ship of the Solomons species.

The different sections present in the Solo-
mons are characterized as follows:

Section Al. —Ninth tergite bare, emarginate
apically, and with lateral lobe either broad
or produced into spine; mesosome with
strong teeth or spines arranged into apical
and median groups. A pre-alar sclerite dis-
tinctly separated from upper sternopleuron.
Cell R2 short or very short, between 0.45
and 0.3 of vein R2+3. Head with at most
a few erect scales on vertex; at least a nar-
row orbital line of light scales; frontal tuft
present or absent. Palpus between 0.05 and
0.08 of proboscis. Scutal vestiture largely
composed of dark, narrow, curved scales
with a patch or line of broad light scales
restricted to front of wing root or extended
cephalad. Pleura with apn light-scaled; ppn
bare or with light scales; stp with transverse
patch of light scales in upper third and
variable number of light or dark scales in
lower two thirds; only one strongly pro-
pleural; one spiracular present; two or more
upper mesepimerals present. Legs without
light-scaled markings on femora. Included
species: U. atra Theobald, £/. barnesi n. sp.,
U. dvinskii n. sp., U. solomonis n. sp., U.
sexaueri n. sp.

Section A2. — As in section Al except for the
following: three or more strong propleu-
rals; spiracular and upper mesepimerals ab-
sent; conspicuous white knee spots on all
femora. Included species: U. wysockii n. sp.

Section C2.— Ninth tergite bare, broadly pro-
duced in middle and without lateral lobe;,
mesosome with numerous weak serrations
not arranged into apical and median groups.
Pre-alar area not distinctly separated from
upper sternopleuron. Cell R2 very short,
less than 0.3 of vein R2-I-3. Head with nu-
merous conspicuous erect scales on vertex,
no distinct orbital light-scaled line; frontal
tuft not developed. Palpus about 0.1 of
proboscis. Scutal vestiture of dark, narrow,
curved scales, lighter near wing root (in
related species broad scales over anterior
end of ppn). Pleura with apn with trans-
lucent scales, no scales on other sclerites;
four strong propleural bristles; one spira-
cular; three or four upper mesepimerals.
Legs entirely dark-scaled. Included species:
U. quadrimaculata Edwards and related Pa-
puan species.

Edwards has shown the importance of the
shape of the ninth tergite of the male geni-
talia in determining relationships in this ge-
nus. Unfortunately, few other workers have
paid much attention to this basic character,
perhaps because of the minuteness of the geni-
talia. The shape of the clasper has been recog-
nized as offering specific differences as well
as group differences. It appears too that the
ornamentation of the mesosome as well as
its shape may be of considerable diagnostic
value. In the Solomons species the shape of
the paramere is distinctive in several species.
Of the other characters used previously, not
enough attention has been given to the pro-
portions of leg segments which are distinctive
in every species I have studied. It appears
that the thoracic scutal and pleural ornamen-
tation is not as important a character as be-
lieved by Edwards as several nonornamented
species in the Oriental region undoubtedly
are closely related to ornamented forms.

Pupal Characters

According to Edwards (1941: 364-365) and
Penn (1949: 28-29) , the pupal stage of JJra-
notaenia is not well defined. Although it is

318

PACIFIC SCIENCE, Vol. VII, July, 1953

true that it is not as clearly distinguished as
the adult stage, the majority of species are
readily recognized. Perhaps the distinctive-
ness of the adults of Uranotaenia has been
overemphasized, as the extremely small size
of the microtrichia is actually the only char-
acter possessed by all species that distin-
guishes them from other mosquitoes, and
this is a relative character whose constancy
has not been checked.

The presence of several distinct types of
pupae hinders the diagnosis of the genus and
indicates its complexity, but the following
characters apply to the majority of species:
abdominal hair 1-IX better developed than
in any other genus, at least one half as long
as segment, usually thickened; palpal case
very short; torus case exceptionally large; hair
4-C placed far caudad and mesad of 5-C; hair
6-C single, usually thickened and very long,
always longer than 7-C; hairs 6, 7-C far re-
moved from 4, 5-C; mesoscutum very strong-
ly arched; paddle with part mesad of midrib
broader than outer part; cereal plates of fe-
male indistinct, usually poorly sclerotized and
never projecting.

The different sections present in the Solo-
mons are differentiated as follows:

Section Al. — Trumpet with a distinct slit into
meatus from pinna, placed closer to mid-
dorsal line than to wing case; paddle with
one weak hair and with indistinct serra-
tions, distinctly longer than wide; cephalo-
thoracic and abdominal hairs conspicuous-
ly branched; hairs 8, 12-11 absent.

Section A2. — Trumpet without slit, placed
closer to wing case than to middorsal line;
paddle without hair; other characters as in
section Al.

Section C2.— Trumpet without slit, placed
closer to wing case than to middorsal line;
paddle with 2 strong terminal hairs and
strong external and internal serrations;
cephalothoracic and abdominal hairs not
conspicuously branched; hairs 8, 12-11 pres-
ent.

Larval Characters

The larval stage of Uranotaenia is by no
means as distinct as generally regarded. Hop-
kins (1936: 40-41), Barraud (1934: 59), and
Edwards (1932: 97) have given good general
diagnoses, but exceptions to the most striking
characters— comb plate and single head hairs
5 and 6 — are becoming more numerous as
more larvae are described.

In the Solomons all the known Uranotaenia
can be distinguished in the larval stage as
follows: Head: Usually slightly longer than
broad but sometimes slightly broader than
long; hairs 8 and 6 spike-like or simple or
even branched; maxillary suture (premaxil-
lary, mental, gular) always completely absent.
Antenna: Short, smooth or minutely spicu-
late; hair 1 short or long, single or branched.
Thorax: Lateral and ventral hairs plumose or
single. Abdomen: Dorsal and ventral hairs slen-
der stellate tufts or single; lateral hair strongly
developed on I, II or I-VI; comb plate always
developed, sometimes united with its mate
dorsally; comb scales and pecten teeth usu-
ally fringed but may be simple or with a few
basal denticles; anal saddle completely ring-
ing segment; acus always present.

The sections present in the Solomons may
be differentiated as follows:

Section Al. — Thorax and abdomen with dor-
sal and ventral stellate tufts; antennal hair
1-A at middle or near base; thoracic hairs |
8, 9-M, 7, 9-T long multiple barbed tufts; I
abdominal hairs 6 strongly developed on j
I and II, weak and stellate on other seg-
ments; hair 8-1 developed, hair ll-I absent;
comb plates separate; comb scales and pec-
ten teeth fringed laterally or apically; ven-
tral valve hair 13-S twisted at base; all valve
hairs single. Antennal hair single, short;
head hairs 5, 6-C spike-like; abdominal hair
4-II stellate; pecten not reaching beyond
0.5 of siphon; siphonal hair 1-S less than
0.5 of siphon length; head hair 14 thick-
ened, spike-like.

Uranotaenia in Solomon Islands — Belkin

Section A2.— Like section Al except: anten-
nal hair 1-A branched, about as long as
antenna; head hairs 5, 6-C slender, not
spike-like; abdominal hair 4-II forked, not
stellate; pecten reaching to 0,8 of siphon;
siphonal hair 1-S about as long as siphon;
head hair 14 dendritic.

Section C2. — Thorax and abdomen without
dorsal or ventral stellate tufts ; antennal hair
1-A near apex of antenna, single; thoracic
hairs 8-M, 7-T short, multiple but not
barbed; hairs 9-M, T long, single; abdom-
inal hairs 6-I-IV subequal, all arising from
tubercles; hair 8-1 absent, hair ll-I present;
comb plates united dorsally into a saddle;
comb scales and pecten teeth simple, with-
out fringes but with or without basal denti-
cles; ventral valve hair 13-S a simple single
hair not twisted at base; some valve hairs
branched; anal hairs 2, 3, 4-X single; head
hair 14 a simple, slender hair.

Species groups within these sections are
separated in the Solomons chiefly on the fol-
lowing characters: head hairs 4, 9, 11-C; an-
tennal hairs 1, 2, 3, 4- A; thoracic hairs 3, 4, 7,
9, 10, 14-P; abdominal hairs 1, 6, 13 on seg-
ments I-VII; pentad hairs; development of
ventral valve and its hairs; hairs 2, 3, 4a, b-X;
size of anal segment. Individual species are
distinguishable apparently on the differential
development and branching of all the larger
hairs, on characters of comb scales and pec-
ten teeth, length of siphon, spiculation of
caudal margin of siphon, and length of anal
gills.

Montschadsky (1930: 580-582; pi. 8, fig.
12) describes and figures for U. ungukulata
Edwards, 1913, a process arising from the
spiracular rim that is similar in form to the
twisted hair 13-S in larvae of sections Al and
hi in the Solomons. Both of these characters
are restricted to Uranotaenia and may prove
to be of value in subdividing the genus.

Biology

The majority of species of Uranotaenia
whose immature stages are known utilize

319

ground waters for breeding, being found in
swamps, marshes, and streams as well as in
temporary pools. The individual species or
groups vary greatly in their requirements for
light, oxygen, and protective cover. As far as
known, all species utilizing such habitats for
breeding lay their eggs in rafts which are
superficially similar to those of Culex and
Mansonia. The larvae of these species rest
parallel to the surface film and immediately
below it because of the shortness of the siphon
and the angle it forms with the abdomen.
They spend most of their time near the sur-
face, apparently feeding just below the sur-
face film. Certain species are easily mistaken
■ for the younger stages of anophelines, partic-
ularly as they are often present in association
with them. The jerky movements of these
larvae are also suggestive of anophelines. All
the species in the Solomons except U. wy-
sockii and U. quadrimaculata belong to this
group. I have collected egg rafts of these
forms but have not reared them.

A number of species have been reported
from specialized ground waters such as rock
holes. It appears that some of these forms are
morphologically distinct from the above-
mentioned group and may lay eggs singly
in some cases. U. stonei Bohart and Ingram,
1946, from Okinawa has a very peculiar larva
with unusual behavior and resting position
but apparently in the adult stage shows re-
semblance to section Al in the shape of the
ninth tergite of the male. On the other hand,
less specialized forms, such as the atra-groxv^
which utilize crab holes for breeding, retain
their similarity to ordinary ground-pool
breeders.

Several species are restricted to breeding in
water collections in living plants such as spe-
cies of Bandanus, Nepenthes, Colocasia, Alocasia,
Curcuma, etc. It is apparent that members of
more than one section of the genus utilize
this type of habitat. In the Solomons U. wy-
sockii belongs to this ecological group. I have
never found mosquito egg rafts in water col-
lections in living plants and consider it likely

320

PACIFIC SCIENCE, VoL VII, July, 1953

that eggs of mosquitoes in such habitats are laid
singly in all genera including Uranotaenia and
Culex. The larval behavior of U. wysockiiis dis-
cussed under that species. It should be noted
that both the behavior and superficial morph-
ology of plant-inhabiting species show mark-
ed convergence in entirely unrelated groups.

A fourth group of larval habitats utilized
by Uranotaenia is found in water collections
in various types of dead plant materials found
on the ground. Tree holes either belong to
this class or may be considered as intermediate
between it and the preceding class. Several
species have been reported breeding in water
collections in opened coconuts, coconut
husks, fallen coconut spathes, fallen bamboo,
and various types of artificial containers. In
the Solomons U. quadrimaculata breeds in
such habitats but in addition is found in taro
(Alocasia, Colocasia sp.). Again it appears that
representatives from several distinct groups
of the genus have invaded this ecological
niche. It is probable that eggs are laid singly
in these species, as reported by Bohart and
Ingram (1946: 58) for U. bimaculata
1908. I have never found egg rafts of U.
quadrmaculata in over 50 collections. The
appearance and behavior of these larvae is
very different from the ordinary ground-water
inhabitants and is discussed in more detail
under the section on the biology of U. quad-
rimaculata.

In the Solomons all the species of Urano-
taenia develop rather slowly in the laboratory,
and I am under the impression that it takes
at least 10 days for the completion of the
aquatic cycle. The pupal stage in the labora-
tory lasts 2 to 3 days and even longer in the
case of U. wysockii.

Adults of Uranotaenia are delicate, shy mos-
quitoes which apparently do not normally
feed on human blood. U. geometrica Theobald,
1901, was reported on the authority of Lutz
(Theobald, 1901: 248) as being a severe biter,
but this has not been confirmed (Shannon,
1931: 24; Lane, 1943: 138). The only definite
records on blood feeding are those of Davis

and Philip (1931: 137, 138) on bird blood and
Remington (1945: 32-37, 64-68) on amphi-
bian blood. In the Solomons all the observed
species are fairly active during the day but
are restricted to shaded situations. They oc-
casionally land on human beings but in my
experience never bite. The majority of species
are found on moist surfaces near the breeding
places, and an occasional specimen shows
blood. Several species come readily to electric
lights, indicating that their activity is not re-
stricted to daylight hours.

Ajjinities of Solomons Fauna

The Solomons species of Uranotaenia as
here understood all belong to species groups
represented in the Australasian and Oriental
regions. All the species are endemic except
U. atra. The specimens determined as the
latter are probably also a distinct species in
the Solomons, but because of the insufficient
knowledge of this group in the other areas
of its occurrence they are not separated no-
menclatorially for the present. All the species
show very strong affinities with Papuan and
Philippine representatives of the same groups
except U. wysockii, a member of the Oriental
alboannulata-grou'^ which has not been recog-
nized previously from these subregions.

There is only one representative of each
species group in the Solomons, and in each
case the separation from related forms is quite
distinct in all known stages although based
on relatively few characters. Probably addi-
tional characters will become apparent when
related species are studied in greater detail.
U. wysockii is an exception for the reason
stated above, and whether it is a form pecu-
liar to the Solomons and without close rela-
tives in adjacent areas is not known. Detailed
discussions of the relationships will be found
under the section on taxonomic discussion
for each species.

Whether the Solomons forms described
here are full species or should be treated as
subspecies only is a matter of opinion. The
differences noted are constant, occur in all

Uranotaenia in Solomon Islands — -Belkin

321

stages, and are more striking than those be-
tween U. quadrmaculata and U. nigerrima,
which Edwards considered distinct species.

To the east of the Solomons only three
species of Uranotaenia have been reported
from the Pacific islands. All of them appear
to be very distinct from any species found in
the Solomons. Perry (1946: 11, 14) reports
U. tibialis Taylor from Espiritu Santo, New
Hebrides. I have seen a single damaged female
thought to be this species from Efate, New
Hebrides (Rentaban Bridge, Sept. 20, 1942,
K. L. Knight Coll. USNM). It appears to be
a distinct species, possibly not related to U.
tibialis at all. The abdomen is broken off, but
the following features are discernible: broad
scales of head all dark, no indication of light
orbital line, one pair of long, erect, vertical
scales in addition to shorter, lighter, erect oc-
cipital scales; scutum with short, broad patch
of bluish scales on front of wing root; apn
and stp each with broad patch of broad, bluish
scales but integument not distinctly light in
line with the patches; wing with white scales
as in tibialis- gtonp, distance between cross-
veins about 1.4 of m-cu; hind tarsus white
from extreme apex of segment 2. Perry (1946:
11) states that in the larva the comb scales
are not fringed or spined and that head hairs
5 and 6 are spike-like. It seems unlikely that
the characteristic leaf-like antennal hairs could
have been overlooked. At any rate, this un-
described species appears to have very little
relationship to the Solomons Uranotaenia.
The other two species from the South Pacific
are both reported from Fiji, Edwards,

1935, is entirely different from any Solomons
species, particularly in the larval stage, and
appears to have no close relative anywhere in
the Papuan subregion. U. colocasiae Edwards,
1928, appears superficially to be related to the
nigerrima-gtow^ but is very unlike U. quad-
rimaculata of the Solomons. Thus, as is shown
also in Tripteroides (Belkin, 1950: 214), there
appear to be no strong affinities between the
mosquito fauna of the Solomons and those
of the South Pacific islands to the east.

Keys to Species

1. ADULTS (MALES AND FEMALES)

1. Posterior pronotum and supra-alar area in
front of wing root each with a dark velvety
integumentary spot; vertex of head with
numerous conspicuous erect scales ......

.U. quadrimaculata Edwards

Posterior pronotum and supra-alar area
without dark spots; vertex of head with
at most a few inconspicuous erect scales . . 2

2. Thorax with broad whitish integumentary

area embracing lower margin of scutum in
front of wing root and upper half of pleura;
vertex of head with broad scales all whitish
U. sexaueri n. sp.

Thorax with scutal integument all dark but
with a narrow patch or a longitudinal line
of whitish or bluish scales on lower margin,
pleura with a median longitudinal line or
patch of light scales; vertex of head with
some of the broad scales dark ......... 3

3. Scutum with a lateral longitudinal line of
white scales extending around the front
margin from wing root to wing root; three
or more strong propleural bristles; spira-
cular and upper mesepimerals absent; con-
spicuous white knee spots on all femora . .

U. wysockii n. sp.

Scutum with a lateral longitudinal line or
patch of white or bluish scales restricted
to area between transverse suture and wing
root on each side; at most one strong pro-
pleural bristle; spiracular and upper mese-
pimerals present; femora without knee
spots, at most indistinctly lighter apically
at articulation 4

4. Abdomen with conspicuous dorsal white

scaling on at least one segment; light scal-
ing of head and thorax white or slightly
bluish, arranged in narrow lines on thorax;
distinct frontal tuft present; males with
front tibia and tarsus without specialized
bristles or scales 5

322

PACIFIC SCIENCE, VoL VII, July, 1953

Abdomen without conspicuous dorsal
white scaling, lateral light patches may be
present but not visible from above; light
scaling of head and thorax strongly azure
blue, arranged in broader patches on tho-
rax; no frontal tuft; males with front tarsus
or tibia and tarsus with specialized tufts
of bristles and scales 6

5. Hind tarsus white-scaled from middle of

segment 3; light scaling of head, thorax,
and wing almost pure white; very narrow
orbital light line; supra-alar line extending
cephalad of spiracle; abdominal tergite 5
with white apical transverse band extend-
ing to sternite U. solomonis n. sp.

Hind tarsus white-scaled from apical fifth
of segment 2; light scaling of head, thorax,
and wing distinctly bluish; broad orbital
light line; supra-alar line not reaching spi-
racle and angled cephalodorsad; abdominal

tergite 5 completely dark-scaled

U. civinskii n. sp.

6. Tarsi all dark-scaled; wing scales all dark

dorsally; abdominal tergites with incon-
spicuous lateral white patches; male with
first segment of front tarsus excavated and
shortened, second segment without tuft
of long bristles and scales, fore tibia with-
out long apical tuft of scales

U. atra Theobald

Hind tarsus white-scaled from apex of seg-
ment 2; wing scales iridescent bluish on
base of R, Cu, and lA; abdominal tergites
completely dark-scaled; male with first seg-
ment of front tarsus not excavated but
with long basal bristles, second segment
with large tuft of long bristles and scales,
fore tibia with long apical tuft of scales . .
U. harnesi n. sp.

2. MALE GENITALIA

1. Gasper slender, gradually narrowed api-
cally; ninth tergite not emarginate apically
and without lateral lobes; mesosome with
numerous short teeth not arranged into

apical and median groups

.U. quadrimaculata Edwards

Gasper broad, parallel-sided or expanded
before apex; ninth tergite emarginate api-
cally, with more or less distinct lateral
lobes; mesosome with a few long spines
or teeth arranged in apical and median
groups 2

2. Mesosome with a complete dorsal sclero-

tized bridge, each plate with two apical
spines and three or more teeth on median
process U. solomonis n. sp.

Mesosome with only a narrow basal dorsal
sclerotized bridge, each plate with only
one apical and one median strong spine . .

3

3. Lobe of ninth tergite produced as a strong-

ly sclerotized blunt spine that is longer
than broad . U. civinskii n. sp.

Lobe of ninth tergite broadly rounded, not
produced as a distinct spine, always broad-
er than long 4

4. Apical spine of mesosome plate short, aris-

ing at extreme apex, median spine arising
close to it 5

Apical spine of mesosome plate long, aris-
ing distinctly before apex, median spine
arising more ventrally 6

5. Ninth tergite broadly emarginate apically

median bridge distinctly shorter than width
of emargination, lateral lobe distinctly pro-
duced U. wysockii n. sp.

Ninth tergite narrowly emarginate apically,
median bridge at least as long as width of
emargination, lateral lobe broadly rounded
. . . f/. sexaueri n. sp.

6. Ninth tergite very narrowly but deeply
emarginate apically, median bridge short,
lateral lobe very broadly rounded; basal
lobe of sidepiece with numerous bristles
U. atra Theobald

Uranotaenia in Solomon Islands — Belkin

323

Ninth tergite broadly and very shallowly
emarginate apically, median bridge long,
lateral lobe slightly but acutely produced;
basal lobe of sidepiece with few bristles
U. harnesi n. sp.

3. PUPAE

1. None of the larger abdominal hairs con-
spicuously branched; trumpet index 3.0 or
less; paddle with two distinct hairs and
strong inner and outer serrations; hairs

1-3-C single; hairs 8, 12-11 present

U. quadrimaculata Edwards

Larger abdominal hairs conspicuously
branched; trumpet index 3.5 or more;
paddle with one indistinct hair or none,
inner serrations always very weak, outer
poorly sclerotized; hairs 1-3-C multiple;
hairs 8, 12-11 absent. 2

2. Trumpet index at least 10; hair 8-C with
long central stem and short basal branches;
hair 2-III laterad or cephalad of l-III. . .3

Trumpet index at most 6.3; hair 8-C with-
out strong central stem, branches subequal;
hair 2-III distinctly mesad of l-III 4

3. Hair 4-1 V, V single, longer than two fol-

lowing tergites together; trumpet uni-
formly dark, tracheoid to pinna

atra Theobald

Hair 4-IV, V 2-5b, shorter than two fol-
lowing tergites together; trumpet dark on
tracheoid and apex, light in between, tra-
cheoid to 0.55 from base. . . U. harnesi n. sp.

4. Trumpet without slit in meatus; paddle

almost as wide as long, without hair; hair
4-IV-VI usually 3b, branches heavy, curved
apically, extending to middle of second
tergite following U. wysockii n. sp.

Trumpet with a distinct slit in meatus;
paddles distinctly longer than wide, hair
present but small; hair 4-IV-VI usually

with more slender straight branches .... 5

5. Hair l-II secondarily branched; hair 4-

V-VII usually 3b, distinctly longer than
following tergite and distinctly longer than
hair 1 on corresponding segments; trum-
pet index 5.0 U. cmnskti n. sp.

Hair l-II primarily branched only; hair
4-V-VII usually 4-6b, slightly longer than
following tergite and about as long as hair
1; trumpet index 4,0 or less 6

6. Trumpet dark basally, golden brown on
apical half or more; hair l-II 6, 7b; paddle
serrations very indistinct, restricted to apex
U. sexaueri n. sp.

Trumpet dark throughout; hair l-II 10,
lib; paddle serrations distinct on apical
half of external margin. . . U. solomonis n. sp.

4. LARVAE (FOURTH INSTAR)

1. Thorax and abdomen without dorsal or
ventral stellate hairs; hair 1-A at apex; hairs
8-M, 7-T short, multiple but not barbed;
hairs 9-M, T single; hair 6-I-IV of ap-
proximately equal size, all arising from
basal tubercles; comb plates united dor-
sally into saddle; comb scales and pecten
teeth not fringed laterally or apically, but

may have a few basal denticles

U. quadrimaculata Edwards

Thorax and abdomen with slender dorsal
and ventral stellate hairs; hair 1-A near
middle or base; hairs 8-M, 7-T long, mul-
tiple, barbed; hairs 9-M, T multiple,
barbed; hair 6-1, II about three times or
more longer than 6-III, IV, the latter with-
out basal tubercles; comb plates separate,
not united dorsally into saddle; comb
scales and pecten teeth with lateral or
apical fringes 2

2. None of head hairs developed into spikes;
hair 1-A long, branched, reaching beyond
apex of antenna; hair 1-S as long as siphon;
pecten extending to 0.8 of siphon. ......

U. wysockii n. sp.

324

Hairs 5, 6-C developed into heavy spikes;
hair 1-A short or minute, never reaching
apex of antenna; hair 1-S distinctly shorter
than siphon; pecten not extending beyond
0.5 of siphon. 3

3. Hairs 2-4-A leaf-like; hair 4-C long, sim-
ple; hair 9-C short, half or less of 8-C; hair

2- X 4, 5b; siphon index 3-5, valves very

long; hairs 1-P, 5-M frayed apically

U. harnesi n. sp.

Hairs 2-4-A simple, acute; hair 4-C short,
branched; hair 9-C about as long as 8-C;
hair 2-X with no more than three branches;
siphon index 4 or more, valves moderate;
hairs 1-P, 5-M simple and acute apically. .4

4. Hair 14-P single; either anal gills short,

rounded, hardly longer than wide or hair
6-1, II 3b 5

Hair 14-P multiple; anal gills long, bluntly
tapered, at least four times as long as wide;
hair 6-1, II 2b 6

5. Hair 6-1, II 2b; hair 11-C 2, 3b; hair 3-P

3- 5b; hair 7, 9-P single; hair 3-VIII 4, 5b;

hair 2-X 2b; anal gills short, rounded,
hardly longer than wide; head poorly pig-
mented, spikes poorly developed

U. atra Theobald

Hair 6-1, II 3b; hair 11-C 4-6b; hair 3-P
6-lOb; hair 7, 9-P 2, 3b; hair 3-VIII 3b;
anal gills about four times as long as wide,
bluntly tapered apically; head heavily pig-
mented, spikes heavy. . . .U. solomonisn. sp.

6. Hair 14-P 12-l6b; hair 1-A less than dia-

meter of antenna in length; hair 9-C 4b;
hair 11-C 8, 9b; hair 4-P 2b; hair 9-P 4,
5b; hair 3-VIII 7, 8b; middle scale of comb
enlarged U. sexaueri n. sp.

Hair 14-P 5-8b; hair 1-A about twice dia-
meter of antenna in length; hair 9-C 2, 3b;
hair 11-C 3-6b; hair 4-P 3b; hair 9-P 2, 3b;
hair 3-VIII 4-6b; middle scale of comb
not enlarged. U. civinskii n. sp.

PACIFIC SCIENCE, Vol. VII, July, 1953

CONSIDERATION OF SPECIES

1. Uranotaenia atra Theobald, 1905
Plates 1, 2

1905. Uranotaenia atra Theobald, Budapest.
Magyar Nemzeti Muz., Ann. Hist. Nat.
3: 114. Type: E; Muina [Muima, near
Madang], New Guinea, 1900 (Biro)
[Budapest. Magyar Nemzeti Muzeum].
1905 . Uranotaenia coeruleocephala lateralis Lud-
low, Canad. Ent. 37: 385-386. Types:
4E; Cottabatto, Mindanao, P. I., 25
Jun. 1905 (E. B. Vedder) [USNM].~
1922. Edwards, Indian Jour. Med. Res.
10: 460 (syn.).

I9O8. Uranotaenia cancer Leicester, Kuala
Lumpur. Inst. Med. Res., Studies 3(3):
215-217. Types: M and E; Port Swet-
tenham and Klang, Malay Penin. (Lei-
cester) [BMNH]. — 1922. Edwards, In-
dian Jour. Med. Res. 10: 460 (syn.),
1910. Uranotaenia ceylonica Theobald, Monog.
Cul. World 5: 503-505. Type: E; Galle,
Ceylon (B. Eletcher) [BMNH]. -1913.
Edwards, Bui. Ent. Res. 4: 238 (syn.).
1914. Uranotaenia propria Taylor, Roy. Ent,
Soc. London, Trans. 61(1913): 704-
705. Type: M; Townsville, Queensland
(H. Priestley) [Sydney Univ..^]. — 1922.
Edwards, Indian Jour. Med. Res. 10:
460 {? syn.), — 1924. Edwards, Bui. Ent.
Res. 14: 357-358 (syn.).

1919. Uranotaenia cairnsensis Taylor, Linn.
Soc. N. S. W., Proc. 43: 839- Type: 2E;
Cairns, Queensland, July 1917 (E. H.
Taylor) [Sydney Univ.?]. — 1924. Ed-
wards, Bui. Ent. Res. 14: 357-358
(syn.).

1934. Uranotaenia atra Theobald. Barraud,
Eauna Brit. India. Diptera v.5 Eamily
Culicidae, Tribes Megarhinini and Cu-
licini pp. 12-1 A: (Eig.: M legs p. 73,
clasper p. 79; larval head, comb plate,
pecten teeth p. 58).

1935. Uranotaenia atra Theobald. Baisas,
Philip. Jour. Sci. 57: 65-66 (Pig.: M

Uranotaenia in Solomon Islands— -Belkin 325

Fig. 1. Uranotaenia atra Theobald, 1905. a-e. Adult; /, g, pupa, a, b. Details of pro- and mesothoracic legs of
male; c, ninth tergite of male; d, male genitalia; e, left lateral aspect of head and thorax of female; /, right lateral
aspect of anterior portion of cephalothorax of pupa; g, metanotum and abdomen of male pupa, left ventral, right
dorsal. Abbreviations as given on page 314.

fore leg, sidepiece, clasper; larval head
and antenna).

1944. Uranotaenia atra Theobald. Knight,
Bohart, and Bohart, Keys Mosq. Aus-
tralasian Reg. pp. 16, 68.

Diagnosis

ADULT.— Head dark centrally; broad orbital
line of azure-blue scales; frontal tuft unde-
veloped. Short supra-alar line of azure-blue
scales; apn and stp each with broad patch of
azure-blue scales, not markedly in line. Tarsi
dark, without definite white on hind leg. Wing
scales all dark. Abdomen with inconspicuous
apicolateral white patches, invisible from
above. Male: I Tar 1 shortened, expanded,
excavated and with tufts of specialized scales;

I Tar 2 with specialized scales; III Tib with
apical and subapical tufts of specialized scales.

PUPA.— Trumpet length 12.0 median width;
tracheoid extending to 0.75; uniformly dark;
slit in meatus. Hair 8-C with central stem and
short basal branches. Hair 2-III laterad or
cephalad of l-III. Hairs 4-IV, V single, dis-
tinctly longer than following two tergites.
Hairs 6-1, II moderate in length. Cephalo-
thorax without strongly contrasting dark ven-
tral pigmentation.

LARVA.— Head about as wide as long; hairs
5, 6-C weak spikes; 4-C short 2, 3b; 7-C 2,
3b; 9"C long, strong, 2, 3b; 1-C short, stout.
Antennal hairs simple; 1-A near base, short.
Thorax and abdomen with well-developed
ventral and dorsal stellate hairs with even

326

branches; hairs moderately long. Thoracic
hairs 9, 10-P well developed; 9-M, T, 8-M,
and 7-T long multiple, barbed; 4-P 2b; 7-P
single; 14- P single; all hairs apically attenu-
ated. Abdominal hairs 6-III-VIII stellate,
much shorter than 6-1, II, without tubercles;
l-I, II 4-6b; 6-1, II 2b, branches uneven;
6-III, IV 5-7b. Comb plates separate; scales
fringed. Siphon index about 4.0; pecten ex-
tending to 0.5; pecten teeth fringed apically
and laterally; hair 1-S moderate, 7-1 lb, un-
even; valves short; hair 9-S a slender hair,
13-S moderate, twisted at base. Anal segment
short; saddle margin with small apical spi-
cules; gills about 0.3 of saddle length, round-
ed; hair 1-X 6-lOb, long; 2, 3-X 2b; 4ab-X
2, 3b.

Description

FEMALE (969-109).— Wing: 1.70 mm. Ab-
domen: 1.20 mm. Proboscis: 1.17 mm. Front
femur: 1.33 mm.

Head: Frontal tuft not developed, a few slight-
ly elongate azure-blue scales projecting an-
teriorly; orbital line of azure-blue scales broad,
slightly expanded toward apn; decumbent
dark scales iridescent bronzy; two pairs of
erect, slender, apically forked, dark vertical
scales; three pairs of slightly shorter, similar
erect occipital scales; occipitals 1:3. Clypeus
brown. Palpus about 0.06 of proboscis; scales
inconspicuous, hairs very conspicuous, over
2.0 length of palpus. Proboscis slightly swol-
len apically; dark-scaled, lighter below; hairs
short and inconspicuous except at apex; label-
la lighter, moderately hairy. Antenna about
1.2 of proboscis; torus creamy except for
brown dorsomesal area, with short hairs and
several minute, outstanding, dark broad scales;
base of first flagellar segment creamy, re-
mainder very dark brown; flagellar whorls
with 6 long bristles; penultimate segment
slightly longer than preceding, apical about
1.2 of penultimate.

Thorax: Scutal integument rather uniformly
brown; dense vestiture of recumbent, narrow,
curved, golden-brown scales; broad patch of

PACIFIC SCIENCE, Vol. VII, July, 1953

semierect azure-blue scales in front of wing
root, scales short anteriorly becoming elon-
gate and lanceolate caudally and forming a
conspicuous tuft; acrostichals moderate, other
bristles strong, all dark. Scutellum brown;
median lobe moderately prominent, with
three strong bristles; lateral lobe with one
weak and two strong bristles; scales dark,
bronzy brown, dense. Pleural integument dark
brown in middle, lighter in line with scaling
and above and below; scaling and chaetotaxy
as figured; apn with large patch of oval azure-
blue scales, not forming a tuft caudally; stp
with a large transverse patch of very broad,
almost circular, azure-blue appressed scales;
small patch of translucent light scales on low-
er stp; bristles dark, propleurals weak. Haltere
light on base and lower part of stem, dark-
scaled on upper part of stem and knob.

Wing: Distance between crossveins about
equal or slightly less than m-cu. Vein R2 about
0.30 of R2-1-3; vein Mi-}-2 about 0.68 of M
beyond m-cu. All scales dark, fringe lighter.
Legs: Coxae and trochanters creamy; scales
light and translucent, with a faint bluish iri-
descence. Femora dark above, lighter ventral-
ly, creamy at base. Tibiae and tarsi dark above,
somewhat lighter below. Leg I : femur 1 ; tibia
1.0; tarsus 0.63, 0.43, 0.30, 0.13, 0.09; claws
very large. Leg II: femur 1.07; tibia 1.43;
tarsus 0.80, 0.40, 0.26, 0.12, 0.10. Leg III:
femur 1.11; tibia 1.25, enlarged apically; tar-
sus 1.05, 0.62, 0.42, 0.25, 0.10, segment 1 with
a small tuft of scales at base ventromesally. !
Abdomen: Tergite 1 with small lateral patch j
of white scales; tergites 2-7 with inconspic- |
uous apicolateral patches of scales, invisible ^ ||
from above, on proximal segments patches ,|
reach to base of segments, on distal restricted ||
to apical margin; remainder of tergites dark- j
scaled. Sternites light cream color, almost j
white. [

MALE (969-102).— Wing: 1.50 mm. Pro- !j

boscis: 1.16 mm. Front femur: 1.25 mm. i

Generally very similar to female. Proboscis }
very strongly swollen apically, conspicuous
ventral light patch on swollen part. Orbital |

Uranotaenia in Solomon Islands — Belkin

327

Fig. 2. Uranotaenia atra Theobald, 1905. Fourth instar larva, a. Head, left dorsal, right ventral; b, left lateral
aspect of distal abdominal segments; c, thorax and proximal abdominal segments, left ventral, right dorsal;
d, dorsal aspect of left antenna. Abbreviations as given on page 314.

328

PACIFIC SCIENCE, Vol. VII, July, 1953

line of light scales narrower than in female.
Torus and flagellum dark brown; flagellar
whorls with about 12-14 bristles, about 0.23
of flagellar length; penultimate segment 1.25
of preceding, apical 1.50 of penultimate. Leg
I: femur 1; tibia 1.0; tarsus 0.28, 0.53, 0.40,
0.20, 0.06, segment 1 shortened, excavated
and with specialized bristles as figured, seg-
ment 2 with specialized bristles as figured;
claws equal, one markedly swollen. Leg II:
femur 1.05; tibia 1.50; tarsus 0.64, 0.33, 0.25,
0.06, 0.18, segments 4 and 5 as illustrated by
Barraud (1934), segment 4 with a long ven-
tral lobe extending to almost middle of 5,
segment 5 with small median ventral lobe;
claws enlarged, larger claw forked. Leg III:
femur 1.13; tibia 1.22, subapical and apical
tufts of specialized scales as illustrated; tarsus
1.02, 0.64, 0.44, 0.17, 0.11; claws equal, not
markedly different.

MALE GENITALIA (969-102) .—As figured.
Ninth tergite widely and deeply emarginate
proximally, sclerotization evanescent in the
middle; apex narrowly but deeply emarginate;
lateral lobe wide, with apical sclerotization
extended ventrolaterally proximad as a strong
curved bar. Proctiger without distinct sclero-
tizations. Basal lobe of sidepiece with dorsal
group of six strong and several small bristles,
and one large and one small bristle ventrally.
Clasper as illustrated. Mesosome with narrow
basal dorsal bridge only; apical tooth long,
weakly curved, three serrations near its base
on apex of mesosome plate; median tooth
broad at base, strongly incurved, one serration
near its base distally on mesosome plate. Para-
mere moderately expanded near base.

PUPA (Exuviae of female, 969-109).— Ab-
domen: 1.80 mm. Trumpet: 0.63 mm. Paddle:
0.47 mm.

Cephalothorax: Dark pigmentation of apical
portion of wing cases and midventral line
including all of leg cases contrasting sharply
with the lighter pigmentation of the remain-
der, except for darker areas at base of wing
cases, middorsally on mesonotum and later-
ally on metanotum. Trumpet dark through-

out; length 12.0 median width; widened in
basal 0.25, slightly flared in apical 0.25; inner
wall indistinct except in reticulate; tracheoid
extending to about 0.75, well developed mes-
ally except on apex, absent from basal 0.08;
reticulate indistinctly ornamented; pinna 0.04;
distinct mesal slit in meatus extending about
0.13 of trumpet length. Hairs moderately pig-
mented; relative position, length, and degree
of development as figured; larger branched
hairs with long slender basal stalk moderately
expanded apically where branches arise, outer
branches often shorter, longer branches with
small but distinct barbs. Hairs: 1(5, 9b, outer
branches of latter shorter), 2(5, 6b), 3 (7b),
4(9b), 5(llb), 6(1), 7(5, 6b), 8(5, 6b, one
branch greatly elongated and thickened),
9(10b), 10(6, 7b), ll(3b), 12(7, 9b).

Abdomen: Moderately pigmented and with
dark areas midlaterally and intersegmentally,
VIII and IX almost completely dark; tergites
II-VII each with a posteromedian patch of
distinct small spicules becoming more exten-
sive caudally; tergite VIII with median patch
of shorter, stronger spines arranged in small
groups; tergal reticulations indistinct; ster-
nites III-VII each with extensive median
patch of small spicules; sternite II without
transverse band; sternite VIII with large an-
teromedian patches of spicules similar to those
of tergite. All hairs moderately pigmented;
relative position, length, and degree of de-
velopment as figured; larger hairs as on ce-
phalothorax. Segment I: hair 1(14 primary
branches, strongly barbed, secondary branch-
ing strong, also with barbs, about 1.1 length
of segment), 2(1), 3(3b), 4(7b), 5(5, 7f),6(4b), j,
7(2f), 10(4b). Segment II: hair 0(1), l(lOb),
2(3b), 3(1), 4(6, 7b), 5(4b), 6(2, 3b), 7(1), |

10(4b). Segment III: hair 0(1), 1(8, lOb), |

2(1), 3(8b), 4(3, 4b), 5(3, 5f), 6(5b), 7(1),
8(3f), 10(4b), 12(3b), 13(1), 14(1). Segment
IV: hair 0(1), l(7b), 2(1), 3(6b), 4(1, minute-
ly barbed distad), 5(3, 4f), 6(5b), 7(1), 8(5b), '

10(2, 3f), 12(3b), 13(1), 14(1). Segment V: j
hair 0(1), l(5b), 2(1), 3(3b), 4(1, minutely
barbed distad), 5(4, 5b), 6(5b), 7(1), 8 (4b), |

Uranotaenia in Solomon Islands — Belkin

10(4b), 12(3b), 13(1), 14(1). Segment VI;
hair 0(1), l(5b), 2(1), 3(3, 4b), 4(2b, minute-
ly barbed distad), 5(5b), 6(4, 5b), 7(1), 8(4f),
10(3b), 12(2b), 13(1), 14(1). Segment VII:
hair 0(1), 1(4, 5b), 2(1), 3(6b), 4(2b, minute-
ly barbed), 5(4b), 6(3, 4b), 7(1), 8(3, 4b),
10(2, 3b), 12(2b), 13(1), 14(1). Segment VIII:
caudal margin of sternite truncate; hair 0(1),
4(5b), 7(4, 5b), 14(1). Segment IX: hair 1(1;
0.6 of segment length). Paddle as figured;
lightly pigmented; midrib strongly sclero-
tized, evanescent apically; external buttress
faintly indicated proximally; basal pigment
bar distinct; external margin with distinct
serrations on apical 0.30; internal margin with
a few scattered sharp spines apically; hair 1
small, distinct. Genital lobe extending to 0.20
of paddle; with ventroapical patch of strong
short spicules. Anal segment indistinct; cereal
sclerites not defined. Male genital lobe (ex-
uviae of 969-102) extending to 0.30 of paddle,
ventral spines arising from weak imbrications;
anal segment indistinct.

LARVA (Fourth instar exuviae of female,
969-109).— Head: 0.60 mm. Siphon: 0.57mm.
Anal saddle: 0.25 mm.

Head: Width about equal to length; ocular
bulge not distinctly defined; pigmentation
light except in antennal and postocular areas;
integumentary imbricate sculpturing faint an-
teriorly, distinct caudally. Labrum long, 0.4
of width at 1-C, anterior margin emarginate
to one half of length. Mental plate moderate;
with six teeth on each side, apical one more
prominent; median apical portion with three
teeth, separated from remainder. Hairs of head
capsule moderately pigmented, well devel-
oped; 5, 6-C developed into thin spikes,
lighter in color than usual, apices sharply
pointed, shafts spiculate basally becoming
barbed apically; 7, 9, H-C minutely barbed,
other hairs simple; relative position, length,
and degree of development as figured. Hair
0(1, short, leaf-like), 1(1, short, stout, bluntly
rounded apically), 3(1, moderate in length),
4(3b), 5(1), 6(1), 7(3b), 8(1, 2f), 9(3b), 10(1,
2f), ll(3b), 12(2, 3f), 13(4b), 14(1, irregular.

329

lightly pigmented spine), 15(2, 3b). Antenna
0.20 of head; shaft distinctly narrowed at
about 0.5; width at middle 0.18 of length;
uniformly moderately pigmented except at
extreme base; shaft without spicules. Anten-
nal hairs well pigmented except 1-A; relative
position, length, and degree of development
as figured; all single; 1-A placed at 0.18-0.27,
length 1.1 of antennal width.

Thorax: Long hairs and tubercles strongly
pigmented; short hairs and stellate hairs light-
ly pigmented; relative position, length, and
degree of development as figured; apices of
long hairs attenuated; barbs when present
long, slender, but conspicuous; hair 5-P about
as long as head. Prothorax: hair 0(8, lOd, b),
1(1), 2(1), 3 (5b, branches strongly barbed,
base elongate, somewhat expanded apically),
4(2b), 5(1), 6(1), 7(1), 8(4b), 9(1), 10(1),
11(3, 4b), 12(2f), 14(1, densely and minutely
spiculate). Mesothorax: hair l(9b), 2(1), 3(3,
4b), 4(4b), 5(1), 6(1), 7(1), 8(4, 5b), 9(4b),
10(1), 11(2, 3b), 12(1), 13(19, 20d), I4(l6d).
Metathorax: hair 1(6, 7b), 2(2f), 3(4b), 4(3b),
5(1), 6(2b), 7(7b), 8(10, 12d), 9(4b), 10(1),
ll(2b), 12(2f), 13(7b).

Abdomen: Tubercles and hairs 6, 7-1, II strong-
ly pigmented, other hairs moderately pig-
mented; relative position, length, and degree
of development as figured; barbs when pres-
ent slender, inconspicuous. Stellate hairs (1,
6, 13) with equal branches; 4-II a stellate
hair. Segment I: hair 1(4, 6b), 2(1), 3(1),
4(7b), 5 (4b), 6(2b, lower branch shorter),
7(1), 8(2, 30,9(4, 6b), 10(3f), 12(3b), 13(2b).
Segment II: hair 0(1), l(6b), 2(1), 3(4, 6b),
4(5b), 5(1, 3f), 6(2b, lower branch shorter),
7(1), 8(2f), 9(3b), 10(2b), ll(2f), 12(1), 13(4,
5b). Segment III: hair 0(1), 1(6, 7b), 2(1),
3(2f), 4(2f), 5(2, 3b), 6(6b), 7(4, 5b), 8(2f),
9(1), 10(2f), ll(2f), 12(1, 2b), 13(4, 5b),
14(1). Segment IV: hair 0(1), l(6b), 2(1),
3(1, 2f), 4(2, 3f), 5(2b), 6(6, 7b), 7(3b), 8(2f),
9(1), 10(2b), 11(1), 12(2b), 13(5, 6b), 14(1).
Segment V: hair 0(1), l(7b), 2(1), 3(3b),
4(2f), 5(2b), 6(6b), 7(5b), 8(2b), 9(1, 2f),
10(1, 2f), 11(1), 12(2b), 13(6b), 14(1). Seg-

330

PACIFIC SCIENCE, VoL VII, July, 1953

merit VI: hair 0(1), l(7b), 2(1), 3(2, 3f),
4(3b), 5(1, 2b), 6(7b), 7(4b), 8(2, 3f), 9(2b),
10(2f), ll(2b), 12(2f), 13(16, 17d), 14(1, 2f).
Segment VII: hair 0(1), 1(6, 7b), 2(1), 3(6,
9b), 4(2f), 5(2, 3b), 6(5b), 7(2f), 8(5, 6b),
9(1, 2f), 10(2b), ll(3b), 12(1), 13(5b), 14(1).
Segment VIII: comb plate strongly sclero-
tized, moderately pigmented, ornamented
with numerous imbrications but lacking spi-
cules except apically; comb scales 7-8, ven-
tral ones smaller, sharply pointed, with short
lateral fringe extending to about 0.75 as fig-
ured; hair 0(1), 1(4, 5b, outer branches short-
er), 2(2, 3f), 3 (4b), 4(2f), 5 (7b, outer branches
shorter), 14(1, unusually strong). Siphon: as
figured; length 4.0 median width; moderately
pigmented, imbrications as on comb plate
but all without spicules; pecten extending to
0.5, teeth 11-12, with lateral and apical fringe
as figured; valves as figured, very darkly pig-
mented, ventral valve short; hair 1(9, lOb,
outer branches shorter, basal expansion short
and asymmetrical, barbs absent), 2(1, unusu-
ally long), 3-5 (only two setal rings visible),
6(1, strong, longer than valve), 7(1, short,
slender), 8(1, moderate, shorter than valve),
9(1, long but very slender and attenuate),
10(hairless setal ring), 11(1, well developed),
12(1, minute), 13(1, strong, twisted at base,
about length of valve). Segment X: saddle
lightly pigmented, darker on the apex dorsal-
ly; median width 0.8 of length; imbrications
much fainter than on comb plate except cau-
dally; caudal margin without long teeth, but
with several rows of small spicules arising
from imbrications; gills short, stubby, round-
ed apically, about 0.3 of saddle length; hair
1(7, lOb, about 0.45 of saddle length), 2 (2b,
about 3.2 of saddle length), 3(2b, about 3.8
of saddle length), 4a(2b, about 1.3 of saddle
length), 4b (2b, about 2.0 of saddle length),
4c(l, about 2.4 of saddle length), 4d(l, about
1.6 of saddle length), 4e(l, about 0.25 of
saddle length).

Variation

. The adults of JJ. atra show considerable
variation in the amount and intensity of blue

scaling on the head and the thorax as do the
other ornamented species, but in all the spec-
imens the azure-blue iridescence is very dis-
tinct. The white scaling of the abdominal
tergites is also subject to considerable indi-
vidual variation but is never absent.

The variation in the chaetotaxy of the im-
mature stages is summarized in Tables 1 and
2. The range of variation of branching in this
species appears to be less than in any other
species known from the Solomons, but this
may be due to the fact that only three collec-
tions were made and from only two localities
on Guadalcanal. One larval specimen shows
a duplication of hair 1-C on the left side,
another has 2-A placed at about the middle
of the antenna instead of the apex.

No information is available on geographical
variation in the Solomon Islands. A single
male from Munda, New Georgia, agrees in
all respects with the material from Guadal-
canal.

Specimens examined: 27M; 28F; 81P; 27L.
Individual rearings: 16 larval, 1 pupal.

Taxonomic Discussion

The application of the name Uranotaenia
atra to the ornate species with the characteristi-
cally modified male front tarsus, entirely dark
legs, and largely dark abdomen rests on Ed-
wards’ identification (1913: 238) of Theo-
bald’s ''mouldy and rubbed and scarcely rec-
ognizable” single female type specimen of
f/. atra as conspecific with IJ. ceylonica Theo-
bald, 1910, another species described from a
single female specimen. Later Edwards (1922:
460) synonymized U. coeruleocephala lateralis
Ludlow, 1905, based on four females from
the Philippines, and JJ. cancer Leicester, 1908,
based on a male and a female from the Malay
Peninsula. Still later (1924: 357-358) Edwards
synonymized JJ. propria Taylor, 1914, based
on a male from Queensland, and U. cairnsensis
Taylor, 1919, based on two females also from
Queensland.

The species as understood by Edwards could
hardly be recognized from Theobald’s de-

Uranotaenia in Solomon Islands — Belkin

scription of U. atra (1905: 114; 1907: 563-
564), but the remaining nominal species are
evidently applicable to the same species or
species complex in different portions of its
range. It is possible that the poor original
condition of the type female of U. atra is
responsible for Theobald’s failure to mention
the characteristic ornamentation of the head,
pleura, and abdomen and the absence of
white scales on the wing. On the other hand
it is also possible that U. atra Theobald rep-
resents a species distinct from Edwards’ con-
cept and one which has not been recognized
since the original description. Theobald’s de-
scription fits U. annandalei Barraud, 1926,
much better than any other species, but U.
annandalei has not been reported to date from
New Guinea although it is known from the
Philippines (Baisas, 1935: 64). For the present
I prefer to follow Edwards’ synonymy and to
retain the name U. atra Theobald, 1905, for
this species.

The reportedly wide geographical distribu-
tion of U, atra in the Oriental and Australasian
regions leads one to suspect that it may break
up into a number of geographical subspecies
or that it may be a complex of related species.
The more or less detailed descriptions of the
adults from India, Ceylon, Malaya, Philip-
pines, and Australia agree quite well in gen-
eral features with the specimens from Guadal-
canal. I find small differences, apparently con-
stant, in the foreleg of the males from the
Solomons when compared with the descrip-
tions and figures of Philippine and Indian
material. The larvae from Guadalcanal are
quite distinct from those described from India
(Barraud, 1934: 74) and Netherlands East
Indies (Bonne-Wepster and Brug, 1939: 1231;
Brug, 1924: 441-442) in the length of the
siphon (Solomons 4:1, others 3:1 or less),
and in addition there appear to be constant
differences in chaetotaxy which cannot be
determined positively without a study of large
series. Baisas (1935: 65) states that the Philip-
pine U. atra breeds in forest streams, an eco-
logical niche different from that occupied by

331

this species in other portions of its range
where apparently it is confined to coastal
swamps and to crab holes. The pupal stage
has not been previously described.

Although it appears likely that U. atra
splits at least into several geographical sub-
species, I consider that our present knowledge
is insufficient to recognize these forms. As
pointed out above, the name U. atra is asso-
ciated with the complex under consideration
on rather inconclusive evidence. As far as I
have been able to determine, neither U. atra
sensu Edwards (other than the single female
of Theobald) nor U. annandalei have been
collected on New Guinea. Until such collec-
tions are made and studied and the types
compared again, it would be unwise to in-
troduce new names in a species complex
which has more names already associated with
it than any other Uranotaenia.

It appears that U. atra is most closely re-
lated to the tibialis- gtou.-^ and to other species
with modified fore tibia and tarsus, as well
as to U. annandalei which shares with this
group the peculiar leaf-like development of
the antennal hairs of the larva. The adults of
these species have very broad scales on the
pleura arranged in broad patches and usually
showing very strong azure-blue iridescence.
In the Oriental and Australasian regions the
known pupae of all these species have ex-
tremely elongate slender respiratory trumpets,
greatly elongated hairs 4-IV, V, and a peculiar
development of hair 8-C. On the other hand
U. atra is quite distinct from all its nearest
relatives in having a larva with a wide head,
poorly developed spike hairs, very short hairs
1-C, unmodified antenna, and very short anal
gills.

Biology

The three larval collections on Guadalcanal
were all made in slightly brackish-water pools
immediately back of the beach in open sunlit
areas. Thorough collections in other Urano-
taenia breeding areas never revealed this spe-
cies. It appears to be restricted on Guadalcanal

332

PACIFIC SCIENCE, VoL VII, July, 1953

to the same ecological niche as Culex sitiens
Wiedemann, 1828, another species with short
hair 1-C and stubby anal gills, characters ap-
parently associated with a brackish-water habi-
tat. The only other species found associated
with U. atra on Guadalcanal was Anopheles
farauti Laveran, 1902.

Living larvae are paler than those of other
species inhabiting ground pools and can be
immediately recognized in the field by the
extremely short anal gills. The pupae are easi-
ly recognized in the field by the entirely dark,
very elongate slender trumpets.

This species has been reported breeding in
crab holes and stagnant pools or swamps with
nipa palms in Malaya (Leicester, 1908:
217), in brackish water on a coral islet in
Java (Brug, 1924; 442), and in forest streams
in the Philippines (Baisas, 1935: 65).

The adults of this species were not collected
on Guadalcanal and there is no published
account of their habits.

Distribution

Solomon Islands, Guadalcanal: mouth of
Lunga River (79-1) Dec. 3, 1943 (M. Cohen);
mouth of Balasuma River (968-3, 969-1) Apr.
30 and May 5, 1945 (JNB et all) [USNM,
CU, JNB]. New Georgia: Munda, 1944, IM
(J. G. Franclemont) [JNB].

Territory of New Guinea, Madang:
Muina (Muima, near Madang) 1900 (Biro)
[Budapest. Magyar Nemzeti Muzeum. (Theo-
bald, 1901)].

Australia, Queensland: Cairns, July, 1917
(F. H. Taylor); Townsville (H. Priestley)
[Sydney Univ..^ (Taylor, 1919)].

Netherlands East Indies, Java: Edam
Islet, Bay of Batavia (S. L. Brug) [Weltevre-
den, Java. Centraal Militair Geneeskundig
Lab. (Brug, 1924)].

Philippine Islands, Mindanao: Cottabato,
June 25, 1905 (E. B. Vedder) [USNM (Lud-
low, 1905)]. 'Palawan: Iwahig (F. E. Baisas).
Luzon: Laguna, Calauan (Santiago); Bulacan,
Tungcong Manga, San Jose (F. E. Baisas)
[Manila. Dept. Health (Baisas, 1935)].

India, Ceylon, Andaman Islands, Ma-
lay Peninsula, Siam

2. IJranotaenia barnesi Belkin n. sp.
Plates 3, 4

1944. IJranotaenia tibialis Taylor. Knight, Bo-
hart, and Bohart, Keys Mosq. Austra-
lasian Reg. p. 15 {partim).

1947. IJranotaenia sp. King and Hoogstraal,
Ent. Soc. Am., Ann. 39: 593.

Diagnosis

ADULT. — Head dark in small triangular cen-
tral area; very broad orbital line of light azure-
blue scales; frontal tuft undeveloped. Mod-
erately long and broad supra-alar line of light
azure-blue scales; apn and stp each with mod-
erately broad patch of light azure-blue scales,
not distinctly in line. Hind tarsus white-scaled
from extreme apex of segment 2, Wing white-
scaled on base of R, Cu, and lA. Abdominal
tergites completely dark-scaled. Male: I Tib
with long apical tuft of scales; I Tar 1 with
long specialized bristles and scales at base;
I Tar 2 with large tuft of specialized scales
beyond middle.

PUPA. — Trumpet length 13.0 median width,
tracheoid extending to 0.55; dark on tracheoid
and apex, light in between; slit in meatus.
Hair 8-C with central stem and short basal
branches. Hair 2-III laterad or cephalad of
l-III. Hairs 4-IV, V 2-5b, shorter than two
following tergites. Hairs 6-1, II very long and
conspicuous. Cephalothorax with strongly
contrasting dark ventral pigmentation.

LARVA. — Head slightly longer than wide;
hairs 5, 6-C strong spikes; 4-C long, simple;
7-C l-3b; 8-C strong, 2, 3f; 9-C short 4-7b.
Antennal hairs 2-4 leaf-like, 2, 3- A removed
from apex; 1-A short, simple. Thorax and
abdomen with very short dorsal and ventral
stellate hairs with even branches; all hairs
short. Thoracic hairs 9, 10-P very short;
9-M, T, 8-M, 7-T long, multiple, barbed;

4- P 2b; 7-P single; 14-P single; 1, 2-P and

5- M with apex frayed or brush-like. Abdomi-

Uranotaenia in Solomon Islands — Belkin

333

Fig. 3. Uranotaenia harnesi Belkin n. sp. a-e, Adult; /, g, pupa, a, b. Details of prothoracic leg of male; c, ninth
tergite of male; d, male genitalia; e, left lateral aspect of head and thorax of female; /, right lateral aspect of anterior
portion of cephalothorax of pupa; g, metanotum and abdomen of male pupa, left ventral, right dorsal. Abbre-
viations as given on page 314.

nal hairs 6-III-VIII stellate, much shorter than
6-1, II, without tubercles; l-I, II 4-6b; 6-1,
II 3b, branches even; 6-III, IV 6-9b. Comb
plates separate; scales fringed. Siphon index
about 3.5; pecten extending to 0.5; pecten
teeth apically fringed; hair 1-S short, 10-1 3b;
valves very long; hair 9-S weak, not hook-
like; hair 13-S very long, strongly twisted at
base. Anal segment very long; saddle margin
with short apical spines; gills about 0.7 of
saddle length; hair 1-X usually 5b; 2-X 4, 5b;
3-X 2b; 4a, b-X 2b.

Description

FEMALE (624-24).— Wing: 1.67 mm. Ab-
domen: 1.00 mm. Proboscis: 1.17 mm. Front
femur: 1.20 mm.

Head: Vertex with decambent scales all broad
and rounded apically, no frontal scale tuft;
a very broad orbital line of light azure-blue

scales to level of apn where it is distinctly
enlarged, leaving a smaller dorsocentral dark-
scaled area; one pair of slender, dark erect
occipital scales; one supernumerary frontal;
occipitals all present, 1:3. Clypeus dark
brown. Palpus about 0.05 of proboscis; with
minute dark scales and numerous dark hairs,
some 1.5 or more length of palpus. Proboscis
distinctly swollen apically; dark-scaled
throughout; swollen portion with long hairs;
labella light brown, moderately hairy. An-
tenna about 1.2 of proboscis; torus brown,
a few short hairs dorsally and longer hairs
mesally; flagellum darker, five or six bristles
in whorls; hairs and scales scanty basally, be-
coming more numerous apically; apical seg-
ment with long light hairs, about 1.4 length
of penultimate.

Thorax: Scutal integument dark brown with
lighter longitudinal areas; sparse vestiture of

334

PACIFIC SCIENCE, Vol. VII, July, 1953

recumbent narrow, elongate golden-brown
scales; supra-alar line of light azure-blue scales
as figured, anterior scales small, almost circu-
lar in outline, becoming more elongate cau-
dally, posterior scales narrow; acrostichals
weak, other scutal bristles strong, all dark,
supra-alars few in number. Scutellum brown,
median lobe very prominent; median lobe
with three strong bristles, scales short, round-
ed, projecting only slightly over base of bris-
tles; lateral lobes each with one weak and
two strong bristles, scales elongate, projecting
strongly over bases of bristles. Postnotum
brown, darker centrally. Pleural integument
light brown; scaling and chaetotaxy as fig-
ured; scales light azure-blue on apn and mid-
dle sternopleuron, patches in line with light-
scaling of head and a light integumentary
area caudad of apn; scales of apn short,
rounded anteriorly, elongate, pointed in small
caudal patch; scales of middle sternopleuron
smaller, almost circular in outline; scales on
lower sternopleuron practically transparent;
larger bristles dark, shorter, golden brown.
Haltere light on base, darker on stem dis-
tally, knob dark-scaled.

Wing: Distance between cross veins 1.7 of
m-cu. Vein R2 about 0.4 of R2+3; vein M1+2
about 0.6 of M beyond m-cu. White scales
dorsally on caudal margin of R to slightly
beyond arculus, on basal 0.3 of Cu, and on
base of lA; remaining dorsal scales dark,
much darker on costal and radial fields, iri-
descent bronzy.

Legs: Coxae light, with few translucent scales
and light hairs and bristles; trochanters light,
with a few hairs; femora dark-scaled above,
lighter below; tibiae entirely dark-scaled; front
and mid tarsi entirely dark-scaled, appearing
lighter ventrally in some lights; hind tarsus
white-scaled on entire segments 3-5 and the
extreme apex of 2. Leg I: femur 1; tibia 1.0,
with small tuft of specialized scales and elon-
gate bristles; tarsus 0.85, 0.50, 0.36, 0.14,
0.07; claws equal, one thickened, other elon-
gate. Leg II: femur 1.14, moderately swollen;
tibia 1.70, with three short stout bristles api-

cally; tarsus O.9O, 0.53, 0.30, 0.09, 0.09; claws
as on I, Leg III: femur 1.30, very slender;
tibia 1.30, with specialized scales on swollen
apex; an inconspicuous light patch subapical-
ly on external face; tarsus 1.20, 0.60, 0.40,
0.20, 0.10; claws minute.

Abdomen: Tergites completely dark, scales
iridescent bronzy; sternites light-scaled, con-
trasting sharply with tergites, especially ba-
sally.

MALE (921-301).— Wing: 1.75 mm. Pro-
boscis: 1.33 mm. Front femur: 1.10 mm.

Generally very similar to female. Vertex of
head more extensively dark-scaled, orbital
light line narrow dorsally, more conspicuous-
ly enlarged laterally to apn. Proboscis more
strongly swollen apically. Flagellar whorls
about 0.28 of flagellum, with about 12 bris-
tles. Light-scaling on wing indistinct, dingy
white. Leg I: femur 1; tibia 0.92, specialized
bristles on apex as figured; tarsus 0.54, 0.40,
1.10, 0.40, 0.12, specializations as figured;
claws equal, one strongly swollen. Leg II:
femur 1.28, strongly swollen; tibia I.9O; tarsus
0.92, 0.66, 0.23, 0.06, 0.17, segment 4 with
long ventral lobe projecting below base of
5; claws enlarged, shorter claw about 0.9 of
larger and more slender, slightly curved; larg-
er claw strongly curved. Leg III: femur 1.38;
tibia 1.52; tarsus 1.42, 0.78, 0.53, 0.28, 0.13;
claws as on foreleg.

MALE GENITALIA (921-301) . —As figured.
Ninth tergite expanded, without bristles;
proximal part strongly emarginate in the mid- |
die and with sclerotization evanescent; apical i
part more shallowly emarginate, poorly de- I
fined, short, broad lateral lobe; a strong ven- ^
trolateral sclerotization from apex of lateral
lobe toward basal arm of sidepiece. Proctiger I
with weak lateral sclerotizations. Basal lobe i
of sidepiece with a dorsal group of three large
and one small specialized bristles arising from
tubercles, ventrally with one specialized long
bristle arising from a tubercle. Clasper as fig-
ured, apex of spine broken off. Mesosome
with a poorly sclerotized ventral bridge in
addition to strongly sclerotized dorsal bridge;

Uranotaenia in Solomon Islands — Belkin

335

Fig. 4. Uranotaenia harnest Belkin n. sp. Fourth instar larva, a. Head, left dorsal, right ventral; h, left lateral
aspect of distal abdominal segments; c, thorax and proximal abdominal segments, left ventral, right dorsal;
d, dorsal aspect of left antenna. Abbreviations as given on page 314.

336

an apical and a median long spine curved
ventrad on each lateral plate, spines subequal.
Paramere moderately expanded in the middle.

PUPA (Exuviae of holotype, 624-24).— Ab-
domen: 1.65 mm. Trumpet: 0.60 mm. Pad-
dle: 0.40 mm.

Cephalothorax: Dark pigmentation of apical
portions of wing cases and mid-ventral line,
including all of leg cases, contrasting very
sharply with the light pigmentation of the
remainder. Trumpet dark on tracheoid and
apex of pinna, light in between and at base;
length 13.0 median width; parallel-sided to
about 0.6, then gradually slightly flared; in-
ner wall indistinct except at apex; tracheoid
extending to 0.55, absent from basal 0.1; reti-
culations absent, replaced by minute, sparse
spicules; pinna about 0.05, opening almost
circular; a distinct slit in meatus extending
about one-half distance of reticulate. All hairs
lightly pigmented and simple; relative posi-
tion, length, and degree of development as
figured; larger branched hairs, except 8-C,
with base elongate proximally and moderate-
ly expanded distally where branches arise.
Hairs: 1(9, lOb), 2(7, 8b), 3(7b), 4(14, 15b),
5(13, I4b), 6(1), 7(5, 7b, branches as distinct
as in other hairs), 8(5, 6b, central stem long
and heavy, basal branches short and slender),
9(6, 8b, branches as distinct as in other hairs,
outer shorter), 10 (6b, outer branches shorter),
11 (5b), 12(8, lib, outer branches shorter).
Abdomen: Lightly pigmented on I and II, pro-
gressively darker distad, except for caudolat-
eral lighter areas on V-VII; tergites III-VII
with a small posteromedian patch of short,
heavy spicules; tergite II with a larger, more
anterior patch; tergite VIII with large an-
teromedian patch of arcuately transverse lines
of spicules; reticulations not visible; sternites
II-VIII with extensive median patch of small-
er spicules, arranged in transverse arcuate lines
on more proximal segments. Hairs lightly
pigmented on basal segments, darker caudal-
ly; relative position, length, and degree of
development as figured; larger branched hairs
with elongate basal stems which are not ex-

PACIFIC SCIENCE, Vol. VII, July, 1953

panded into conspicuous plates distally. Seg- ;
ment I: hair 1(16, 18 primary branches with
occasional apical secondary branches; about
1.2 length of tergite), 2(1), 3(4, 5b), 4(7, 8b),
5(2, 3f, heavy stem), 6(6, 7b), 7(1), 10(3b).
Segment II: hair 0(1), 1(10, lib), 2(4b), 3(11,
13b), 4(1), 5(3, 4f), 6(4, 5b), 7(1), 10(2, 3b).
Segment III: hair 0(1), 1(8, 12b), 2(1), 3(12,
13b), 4(3, 4b), 5(3, 4b), 6(5, 6b), 7(1), 8(4,
5b), 10(2, 4b), 12(3, 4b, well developed), |
13(1), 14(1). Segment IV: hair 0(1), l(lOb), |
2(1), 3(7b), 4(3, 4b), 5(4b), 6(7b), 7(1), 8(3, j
4b), 10(2f), 12(3, 4b), 13(1), 14(1). Segment
V: hair 0(1), l(7b), 2(1), 3(4b), 4(2, 3b), 5(6b),
6(6b), 7(1), 8(2, 3f), 10(5b), 12(3b), 13(1),
14(1). Segment VI: hair 0(1), l(6b), 2(1), !

3(4b), 4(4, 5b), 5(4b), 6(6, 7b), 7(1), 8(3,
4b), 10(3b), 12(2b), 13(1), 14(1). Segment
VII: hair 0(1), 1(5, 6b), 2(1), 3(6, 7b), 4(5b),
5(4, 5b), 6(4, 5b), 7(1), 8(3b), 10(3b), 12(3b),
13(1), 14(1). Segment VIII: caudal margin I
of sternite truncate; hair 0(1), 4(6, 7b), 7(4,
5b), 14(1). Segment IX: hair 1(1, about 0.5
of segment length). Paddle as figured; lightly
pigmented; midrib strongly sderotized, eva-
nescent apically; external buttress indistinct;
basal pigment bar very dark; external margin
with very short, blunt, indistinct serrations;
internal margin with a few faint apical serra-
tions; hair 1 short and indistinct. Genital lobe:
extending to about 0.30 of paddle; ventro-
caudal patch of spicules. Anal segment equal
in length to genital lobe, slightly broader than
the latter apically; cereal sclerites distinct.
Male genital lobe (exuviae of allotype, 921- j
301) extending to 0.30 of paddle; pair of |
large ventral lateral patches of strong spines; |
anal segment with distinct basal sclerotization !
connected to genital lobe. |

LARVA (Fourth instar exuviae of holotype, |
624-24).— Head: 0.55 mm. Siphon: 0.42 mm. I
Anal saddle: 0.36 mm. |

Head: Width 0.9 of length; ocular bulge very .
prominent, preocular area very strongly nar- !
rowed; pigmentation very light except for a
transverse dark band in antennal region and j,
another on posterior part of ocular bulge; |

Uranotaenia in Solomon Islands — Belkin

integumentary imbricate sculpturing distinct
and uniform. Labrum short, 0.25 of width at
1-C, median emargination so deep that la-
brum appears as two prominent tubercles for
1-C. Mental plate small, with about 11 in-
distinct teeth. Dorsal hairs of head capsule
strongly pigmented except for 8, 9, 10-C;
ventral hairs lightly pigmented; 5, 6-C spike-
like, very darkly pigmented, apices sharply
pointed, shaft strongly spiculate; 4, 7-C dark-
ly pigmented, thickened, shaft with a few
minute spicules; other hairs simple; relative
position, length, and degree of development
as figured. Hair 0(1, very large, leaf-like,
strongly flattened and expanded, practically
transparent), 1(1, practically straight, widened
in basal half, sharply pointed), 3(1, well de-
veloped and pigmented), 4(1, very long, ex-
tending beyond apex of 1-C), 5(1), 6(1), 7(1,
2b), 8(2f), 9(6b), 10(3, 4b), ll(5b), 12(3b),
13 (4b), 14(1, long, lightly pigmented, ser-
rated subapically laterad), 15(4, 5b). Antenna
about 0.23 of head; shaft of rather uniform
width but irregular, with subapical notches
for 2, 3-C; width at middle about 0.2 of
length; uniformly very darkly pigmented;
spicules very strong and dark, sparse and of
varying sizes.' Antennal hairs, except 1, 5-A,
strongly pigmented; relative position, length,
and degree of development as figured; all
single; 1-A placed at 0.4, its length 2.5 width
and 0.4 length of shaft.

Thorax: All hairs and tubercles strongly pig-
mented; relative position, length, and degree
of development as figured; apices of long
hairs, except 1, 2-P and 5-M, sharply pointed,
not attenuated; barbs when present, slender
and lightly pigmented; hair 5-P about 0.77
of head length. Prothorax: hair 0(8, 9b), 1(1,
with brush-like frayed apex), 2(1, barbs very
long, brush-like at apex), 3 (7b, base mod-
erately expanded), 4(2b), 5(1), 6(1), 7(1),
8(6, 9b), 9(5b), 10(1), ll(3b), 12(2, 3f), 14(1).
Mesothorax: hair l(5b), 2(1), 3(4b), 4(4b),
5(1, apex brush-like), 6(1), 7(1), 8(4b), 9(5b),
10(1), ll(2b), 12(1), 13(10, 13d), 14(15, l6d).
Metathorax: hair l(4b), 2(1), 3 (6b), 4(4f),

337

5(1), 6(3, 4f), 7(9b), 8(7, 8b), 9(5b), 10(1),
11(1, 2b), 12(2f), 13(7b).

Abdomen: Tubercles on segments I, II and
hairs of all segments strongly pigmented; rel-
ative position, length, and degree of develop-
ment of hairs as figured; barbs when present
slender, lightly pigmented and sparse. Stellate
hairs (1, 6, 13) weak, with equal branches
and sharp points, not attenuated; 4-II a stel-
late hair. Segment I: hair l(4b), 2(2b), 3(1),
4(8, lOb), 5 (2b), 6(3b, equal branches), 7(1),
8(2f), 9(3, 5b), 10(3f), 12(2, 3f), 13(3, 4f).
Segment II: hair 0(1), 1(4, 5b), 2(1), 3(3b),
4(6b), 5 (3b), 6(3b, equal branches), 7(1),
8(2f), 9(1), 10(3b), 11(2, 3f), 12(1, 2f), 13(5,
6b). Segment III: hair 0(1), l(5b), 2(1), 3(3b),
4(3f), 5(2b), 6(7b), 7(3, 4b), 8(2f), 9(1, 2b),
10(2, 3b), ll(2b), 12(1), 13(5, 6b), 14(1).
Segment IV: hair 0(1), 1(4, 5b), 2(1), 3(2, 3f),
4(4b), 5(3b), 6(7, 9b), 7(2, 3b), 8(2b), 9(1),
10(2b), 11(1), 12(3b), 13(5, 6b), 14(1). Seg-
ment V: hair 0(1), 1(4, 5b), 2(1), 3(5b), 4(2,
3f), 5(2b), 6(7, 8b), 7(2, 3b), 8(1, 2f), 9(1),
10(2, 3f), 11(1), 12(1, 2b), 13(7b), 14(1).
Segment VI: hair 0(1), 1(4, 5b), 2(1), 3 (4b),
4(3, 4b), 5(2b), 6(6, 7b), 7(2b), 8(2f), 9(1),
10(2b), ll(3b), 12(2f), 13(16, 21d), 14(1).
Segment VII: hair 0(1), l(4b), 2(1), 3(6, 7b),
4(1, 2f), 5(3b), 6(5, 6b), 7(2f), 8(1, 2f), 9(1),
10(2b), ll(3b), 12(1), 13(4b), 14(1). Segment
VIII: comb plate heavily sclerotized, lightly
pigmented, ornamented with numerous im-
bricated lines with very distinct spicules cau-
dally; comb scales 10-11, median ones longer,
sharply pointed, lateral fringe long but in-
conspicuous, extending to apical third or
more, as figured; hair 0(1), 1(4, 5b), 2(2, 3f),
3(5b, strongly barbed), 4(2f), 5(7, 8b), 14(1).
Siphon: as figured; a deep dorsal emargination
at base; length 3.6 of median width; uniform-
ly lightly pigmented; imbricate ornamenta-
tion without spicules, visible proximally only;
pecten extending to 0.5, teeth 13-13 with
apical fringe and very inconspicuous short
lateral fringe as figured; valves as figured, very
heavily sclerotized and very darkly pigmented,
ventral valve very long; hair 1(11, 12b, small

338

asymmetrical expanded base, no barbs visi-
ble), 2(1, minute), 3(1, distinct seta), 4, 5 (one
setal ring visible), 6(1, heavy at base, sharp-
pointed, stiff, almost reaching to apex of
valve), 7(1, stiff short spine), 8(1, similar to
6, almost reaching to apex of valve), 9(broken
off), 10, ll(not observed), 12(1, short stiff
spine), 13(1, strong, twisted at base, longer
than valve, sharply attenuated) ; tracheal
trunks extremely slender, as figured. Segment
X: saddle lightly pigmented, darker dorsally;
median width 0.53 of length; imbrications
strong dorsally and apically, with faint spi-
cules; caudal margin with a few short, heavy
spines and enlarged spicules on apical im-
brications; gills slender, about 0.7 of saddle
length; hair 1(5, 6b, about 0.2 of saddle
length), 2 (4b, about 1.6 of saddle length,
stiff, heavy, and with nonattenuated, sharp
apex), 3(brokenoff), 4a, b(2b, about 1.1 of
saddle length), 4c, d(l, about 1.1 of saddle
length), 4e(l, about 0.85 of saddle length).

Types

USNM No. 61,417 (holotype, allotype,
paratypes) . Paratypes to be deposited in
BMNH, CU, and CSIR (Canberra); also in
coll. JNB.

HOLOTYPE FLP(624-24) Guadalcanal: To-
girie Swamp, midway between Bonegi and
Poha Rivers, Aug. 23, 1944 (V. R. Roa and
F. B. Wysocki). allotype MLP (921-301)
Guadalcanal: swamp west of Poha River, Mar.
20, 1945 (JNB, M. Cohen, E. Winkler).

PARATYPES (69M, 70F, 37P, 320L; 17 in-
dividual rearings), all collected on Guadal-
canal, as follows: IF, 3L(4-1) Doma Cove,
Oct. 21, 1943 (JNB); 3M, IF, 6L(8) Doma
Cove, Oct. 22, 1943 (JNB, R. J. Schlosser,
L. J. Lipovsky); IMLP (83-33), 2L(83-3)
Matanikau River valley, Dec. 5, 1943 (JNB);
16L(86-1) tributary of White River, Dec. 6,
1943 (S. Civinski); 1M(149) slough, Bonegi
River, Jan. 13, 1944 (JNB) ; 2M, 2F, 31(174-3)
Tassafaronga, Jan. 25, 1944 (S. Civinski); IF,
7L(401-2) Mamara River valley, May 25, 1944
(S. Civinski); 1FLP(455-21), 2L(455-2) Sally

PACIFIC SCIENCE, Vol. VII, July, 1953

Creek, Doma Cove, June 14, 1944 (JNB);
lMLP(547-33) Burns Creek valley, Aug. 1,
1944 (L. J. Lipovsky, M. Cohen, S. Civinski);
1M(564) Poha swamp, Aug. 5, 1944 (V. R.
Roa, F. B. Wysocki); 1MLP(616-11) Poha
swamp, Aug. 19, 1944 (V. R. Roa, F. B.
Wysocki); 4MLP(647-11, 12, 13, 17),
2L(647-1) Poha swamp. Sept. 16, 1944 (V.
R. Roa, F. B. Wysocki); IMLP (654-14)
Poha swamp. Sept. 23, 1944 (V. R. Roa, F.

B. Wysocki); 2FLP (676-31, 32), 1MLP(676-
33), IM, IF, 5L(676-3) West Poha swamp,
Oct. 16, 1944 (JNB, J. Laffoon); IM, IF,
5L(708-4) West Poha swamp, Nov. 1, 1944
(JNB); 2FLP(713-11, 12), 1MLP(713-21),
5M, 6F(713-1), 1F(713-3) Poha swamp, Nov.

3, 1944 (L. J. Lipovsky et al.)\ 2M, 2F(7l4-2),
Poha swamp, Nov. 3, 1944 (J. Laffoon); 5M,
4F, 9L, 11P(734-1) East Burns Creek, Nov.
15, 1944 (C. Calloway); 3M, 2F, 5L(775-3)
Matanikau Village, Dec. 9, 1944 (JNB, J. J.
Cuccio, F. B. Wysocki); 1M,*2F, 4L(796-3)
White River valley, Dec. 27, 1944 (F. B. Wy-
socki, Shaw); 3M, 3F, 26L, 2P(802-3) West
Poha swamp, Jan. 5, 1945 (M. Cohen, C.
Calloway, Shaw); 2M, lF(842-2) Tyler Creek, j
Jan. 24, 1945 (Hawkins); 6M, 6F, 116L(816- |
3) West Poha swamp, Jan. 12, 1945 (M. Co-
hen, F. B. Wysocki); llL(837-3) West Poha
swamp, Jan. 20, 1945 (J. J. Cuccio, F. B.
Wysocki); lF(846-2) Kukum, Jan. 30, 1945 i
(M. Cohen, F. B. Wysocki, J. J. Cuccio);
3F, 59L, 5P(848-2), 4M, 10F(848) Lankford
swamp, Kukum, Eeb. 1, 1945 (J. J. Cuccio, i

C. Calloway, Williams); 2FLP(850-31, 35), '
4M, 2F(850-3), 2M, 1F(850) Lankford |
swamp, Kukum, Feb. 2, 1945 (M. Cohen, J. |
J. Cuccio, F. B. Wysocki); 2F(921-3) same i
data as allotype; 2M, IF, llL(958-2) Poha j!
valley, Apr. 28, 1945 (J. J. Cuccio, E. J. Me- j
Cormick and V. R. Roa) ; 4M, 4F(970-3) West
Poha swamp. May 6, 1945 (JNB); 1F(1140) il
routine night catch, Poha area, Apr. 5, 1944 ||
(S. Civinski); lF(Sta. 5) routine night catch, f
mouth Matanikau River, Mar. 14, 1944; '
2M(1283) flying and resting. West Poha
swamp, Nov. 3, 1944 (L. J. Lipovsky); IF, |

Uranotaenia in Solomon Islands — -Belkin

2L, Tenam, Oct. 18^ 1943 (J.G.Franclemont);
lM(0-24) Tenam, Sept. 10, 1943 (P. W.
Oman); IF, 7L(0-32) Tenam, Sept. 9, 1943
(P. W. Oman); 2M, 1F(K^910) June 21, 1943
(K. L. Knight); 2M(G^40) Dec. 5, 1943 (A.
B. Gurney).

Uranotaenia harnesi is named in honor of
Arthur W. Barnes, Jr., to whom I am greatly
indebted for the painstaking rearing of much
of the material collected on Guadalcanal.

Variation

The adults of U. harnesi show a great deal
of variation in the amount of orbital light
scaling on the head. Usually the light-scaling
is extensive and very conspicuous. The tho-
racic light-scaling shows less variation. The
modified foreleg of the male is extremely vari-
able in the degree of development of the
specialized scales and hairs, but they conform
to the arrangement as illustrated. It is prob-
able that some of the differences noted are
due to the teneral condition of reared speci-
mens. One female has the apical two seg-
ments of the left fore tatsus white-scaled.

The variation in the chaetotaxy of the im-
mature stages is summarized in Tables 1 and
2. There is a rather narrow range of variation
in the branching of the hairs despite the fact
that many collections were made at all seasons
of the year. No unusual variations were ob-
served.

The material from other islands falls within
the range of variation exhibited by specimens
from Guadalcanal.

Specimens examined: 90M; 92F; 5 IP;
402L. Individual rearings: 19 larval.

Taxonomic Discussion

Uranotaenia harnesi is closely related to three
New Guinea species of the tihialis-gtowp de-
scribed by King and Hoogstraal (1947: 585-
596) : U. setosa, U. neotihialis, and U. fimbriata.
It is unfortunate that the two earliest nominal
species in this group, U. tibialis Taylor, 1919,
and U. antennalis Taylor, 1919, are not defi-
nitely recognizable at present. Since they were

339

both proposed for specimens from Cairns,
Queensland, it is unlikely that they would
represent species from the Solomon Islands,
particularly when it appears from the work
of King and Hoogstraal that the tibialis -gtou.^
is composed of a number of closely related
species in New Guinea, none of which can
be definitely associated with Taylor’s species,
and all of which are distinct from U. harnesi.

King and Hoogstraal (1947: 593) examined
specimens of U. harnesi and noted their simi-
larity to U.fimbriata K. and H., 1947. The two
species are very closely related but are distinct
in the following characters of the male of U.
harnesi: light-scaling of head very wide in-
stead of narrow; wing with conspicuous light
scales at base of R, Cu, and lA; fore tibia
longer, 0.9 instead of 0.7 length of femur;
fore tarsus 3 longer, 1.1 instead of 1.0; hind
tarsus white from extreme apex of segment
2 instead of basal third of segment 3; white-
scaling of hind tarsus 2 produced ventrally
from apex, demarcation rather indistinct;
light-scaling of head, scutum, pleura, and
wing a rather intense azure-blue instead of
white. The female of U, fimbriata is unknown
but it appears likely that it would be separable
from that of U, harnesi by the extent of the
light-scaling of the hind tarsus, since this
character does not show marked sexual dif-
ferences in this genus. The same character
will separate U. harnesi females from those of
U. setosa, which has the hind tarsus white
from the apex of the third segment only. The
female of U. tibioclada is unknown, but again
since the male does not show any white-
scaling on the apex of the second segment
it is likely that the female will differ from
U. harnesi in this character also.

The pupal stage of U. fimbriata is not
known, and only the trumpets have been de-
scribed and figured for U, setosa and U. tibio-
clada. The trumpets of U. harnesi agree in
coloration with these two species but appear
to be intermediate in length. The pupa de-
scribed and figured by Penn (1949: 32-33)
as U. albescens is undoubtedly that of a mem-

340

PACIFIC SCIENCE, Vol. VII, July, 1953

her of the tibialis It differs from U.
harnesi in a number of characters.

The larva of U. barnesi is generally similar
to those of U. setosa and U. tibioclada, that of
U. fimbriata being unknown. It differs from
both in having; a much longer siphon, index
about 3.5 against 2.0 or less; siphonal hair
1-S 10, llb(10-13) instead of 8b; hair 5-VIII
8, 7b(6-8) against 4, 5b.

Outside of the Australasian region, three
Philippine species, U. hidlowae Dyar and Shan-
non, 1925, U. clarae Dyar and Shannon, 1925
(= U. delae Baisas, 1935, new synonymy)
and U. reyi Baisas, 1935, appear to me to be
definitely related to U. setosa, U. fimbriata, U.
neotibialis , and U. barnesi on the basis of modi-
fications of the foreleg of the male (where
known), light-scaling of the head, thorax, and
hind tarsus, and in the larva on the basis of
the development of hair 4-C and leaf-like an-
tennal hairs 2, 3, 4-A. As noted by King and
Hoogstraal (1947: 592) U. clarae (as U. delaei
and U. reyi were described from males rather
than females. U. clarae appears to be close)
to U. barnesi than even U. fimbriata in having
the hind tarsus white from the apex of seg-
ment 2 in the adult, and in the definitely
subapical position of antennal hair 2, but dif-
fers in lacking specialized hairs at the base of
segment 1 of the fore tarsus of the male.

It is surprising, in view of the occurrence
of this species complex in the Papuan and
Philippine subregions, that to date no related
species have been described from interme-
diate areas in Borneo or Indonesia. The near-
est approach to this group found in Assam,
Burma, and Hongkong is f/. annandalei
Barraud, 1926, a species also reported from
the Philippines (Baisas, 1935: 64) and pre-
sumably present in intermediate areas. Urano-
taenia annandalei resembles U. barnesi in hav-
ing a modified larval antenna with leaf-like
hairs but differs in the development of hair
4-C in the larva and the absence of modifi-
cations of the foreleg of the male as well as
the coloration of the thorax and hind tarsus
of the adult.

The only other Australasian species (also
occurring in the Oriental region) with modi-
fied foreleg in the male is U. atra Theobald,
1905. In this form the modifications are re-
stricted to the tarsus, do not involve the apex
of the tibia, and the hind tarsus is dark. A
somewhat similar but less conspicuous modi-
fication occurs in the Ethiopian U. pallido-
cephala Theobald, 1908, a species very distinct
in its thoracic ornamentation as well as in the
modifications of the hind tarsus of the male.

Biology

U. barnesi breeds largely in dense fresh-
water jungle swamps having a high organic
content in the water. It prefers shade but will
utilize open situations on occasions. On Gua-
dalcanal it was frequently collected in small
pools, foxholes, and road ruts, and less fre-
quently in rock pools and side pools of small,
densely shaded streams.

The species most frequently associated with
it on Guadalcanal were Bironella hollandi Tay-
lor, 1934, Hodgesia cairnsensis Taylor, 1919,
various species of the subgenera Culex and
Lophoceraomyia, Uranotaenia sexaueri n. sp., U.
civinskii n. sp., IJ. solomonis n. sp., and Ano-
pheles Itingae Belkin and Schlosser, 1944, Ano-
pheles solomonis Belkin, Knight and Rozeboom,
1945, and Anopheles nataliae Belkin, 1946.

Living larvae of U, barnesi are easily recog-
nized from other species of Uranotaenia by
the wide thorax and the narrow, lobed ab-
domen which give it a characteristic slender
appearance. The abdomen and thorax are
usually a dark translucent brown. The short-
ness of the siphon, the characteristic hori-
zontal subsurface resting position, and the
definitely elongate head make this species of
Uranotaenia superficially very similar to the
younger larval stages of anophelines, partic-
ularly Bironella. The pupal stage is easily
separated in the field from all other Urano-
taenia except U. atra by the extremely slender
trumpets and from the latter by the light pig-
mentation of the middle of the trumpets as
well as the strongly contrasting light (dorsal)

Uranotaenia in Solomon Islands — Belkin

341

and dark (ventral) pigmentation of the cepha-
lothorax. The small size and the length of
trumpets will normally separate this species
from all other mosquitoes in the held.

The adults of U. barnesi have been collected
in nature resting on tree buttresses in the
jungle as well as hying in shaded situations.
Both males and females have been collected
occasionally in hand night catches but never
actually biting. They are moderately attracted
to lights at night.

Distribution

Solomon Islands, Guadalcanal: Generally
distributed throughout the year on north-
central and northwest coast (JNB et al., J. G.
Franclemont etal., P. W. Oman, K. L. Knight,

A. B. Gurney, H. E. Milliron et all) [USNM,
CU, JNB]. Florida: Halavo, 6L(K-84l) Dec.
17, 1943 (K. L. Knight) [USNM]. Russell:
Banika, IM, Apr., 1943 (W. G. Downs); Pa-
vuvu, 3L, Mar., 1943 (W. G. Downs); IF,
1944 (R. B. Eads) [USNM]. New Georgia:
Munda, IM, IF, 7L, IP, Jan. 10, 1944 (J. G.
Franclemont) [USNM, JNB]. Bougainville:
Empress Augusta Bay, 16M, 14F, Mar. 1,
1943 (C. R. Bruck); 3L(G-323) Apr. 18, 1944;
5L(G-432) Jul. 8, 1944; IM, 2F('G-423) (A.

B. Gurney) [USNM, JNB].

3. Uranotaenia civinskii Belkin n. sp.
Plates 5, 6

1929. Uranotaenia arg^rotarsis Leicester var.
Edwards in Paine and Edwards, Bui.
Ent. Res. 20: 312-313.

1944. Uranotaenia argyrotarsis Leicester.
Knight, Bohart, and Bohart, Keys
Mosq. Australasian Reg. p. 15, 68
( partim) .

Diagnosis

ADULT. — Head dark centrally; wide bluish-
white orbital line, conspicuously expanded
laterally; four pairs of erect dark vertical
scales; conspicuous frontal tuft of light scales.
Narrow supra-alar line of bluish-white scales
angled dorsad anteriorly; apn and stp each

with narrow line of bluish-white semierect
scales forming streak in line with white-scaling
of head. Hind tarsus white-scaled from apical
fifth of segment 2; apical segments of I, II
Tar cream-colored; small subapical light patch
on III Tib of female. Wing with bluish-white
scales on base of R, Cu, and lA. Abdominal
tergites 2-4 conspicuously white in female;
in male tergite 4 largely white, small white
patches on 2 and 3. Male hind tibia bent at
base and with two long curved, one short
straight bristle.

•PUPA. —Trumpet length about 5.0 median
width; tracheoid extending to about 0.50;
uniformly darkly pigmented; slit in meatus.
Hair l-II secondarily branched; 4-V-VII usu-
ally 3b, longer than following tergite.

LARVA.— Head distinctly longer than wide;
hairs 5, 6-C strong spikes; 4-C moderate,
3b(2-4); 7-C 4, 5b(4-6); 9-C long, 2, 3b.
Antennal hairs simple; 1-A moderate, at basal
third. Thorax and abdomen with strong stel-
late hairs, branches even. Thoracic hairs 9-P
long 2b (2-3), 10-P single, barbed; 9-M, T,
8-M, 7-T long, multiple, barbed; 4-P 3b; 7-P
2b; 14-P moderate, 6, 5b(4-8); apices of long
hairs attenuated. Abdominal hairs 6-III-VIII
stellate, much shorter than 6-1, II, without
tubercles; 7-1, II 5-8b; 6-1, II 2b, branches
uneven; 6-III, IV 6, 7b(5-9). Comb plates
separate; scales fringed, not markedly en-
larged in middle. Siphon index about 5.0;
pecten extending to about 0.5; pecten teeth
with apical and lateral fringe, 16, 15(13-18);
hair 1-S moderate 10, 9b(8-13); valves mod-
erate; hair 9-S moderate, hook-like; 13-S
strong, twisted at base. Anal segment mod-
erate; saddle margin with sparse short apical
spines; gills about 0.6 saddle length; hair 1-X
5, 4b(4-7), moderate; 2-X 3b; 3-X 2b; 4a-X
3b; 4b-X 2b.

Description

FEMALE (504-44).— Wing: 2.30 mm. Ab-
domen: 1.40 mm. Proboscis: 1.50 mm. Front
femur: 1.83 mm.

Head: Vertex with a conspicuous frontal tuft

342

PACIFIC SCIENCE, VoL VII, July, 1953

of elongate bluish-white scales merging into
broad scales; broad orbital line of broad
bluish-white scales, expanding gradually into
a broad patch toward apn; four pairs of dark,
elongate, apically forked erect scales border-
ing the light scales mesally; large triangular
area of dark broad decumbent scales dorso-
centrally; two pairs of shorter slender forked
dark occipital erect scales; inner occipital not
seen, outer three at level of lateral expansion
of light orbital line. Clypeus brown. Palpus
about 0.05 of proboscis; with minute dark
scales and numerous long dark hairs. Pro-
boscis distinctly swollen apically; dark-scaled,
appearing lighter ventrally; short lateral hairs
becoming longer apically on swollen portion;
labella light brown, moderately hairy. Anten-
na about 1.2 of proboscis; torus light brown,
apparently without hairs, appearing flattened
and laterally expanded; flagellum darker,
about six bristles in whorls; hairs and scales
scanty basally, becoming more numerous api-
cally; apical segments gradually lengthened,
apical only slightly longer than penultimate.
Thorax: Scutal integument dark brown with
indistinct lighter longitudinal stripes; mod-
erately dense vestiture of recumbent, narrow,
elongate, golden-brown scales, becoming
broader and darker in prescutellar space;
supra-alar line of bluish-white scales as fig-
ured, anterior scales small, moderately elon-
gate, becoming very slender caudally; acros-
tichals moderate, other bristles strong, all
dark, supra-alars few in number. Scutellum
brown; median lobe moderate with three bris-
tles weaker than those of lateral lobes, scaled
area extensive, scales short, rounded apically,
projecting only slightly over base of bristles;
lateral lobe with two very strong and two
weak bristles, scales elongate, projecting mod-
erately over base of bristles. Postnotum dark
brown, lighter laterally. Pleural integument
brown, light iridescent areas in line with light-
scaling; scaling and chaetotaxy as figured;
scales iridescent bluish-white, patches on apn
and middle sternopleuron in line with light-
scaling of head; scales of apn small and elon-

gate anteriorly, becoming more slender and
longer to form a projecting tuft caudally;
those of middle sternopleuron arranged in
about two rows, individual scales moderately
elongate, larger than on apn, scales semierect;
scales on lower sternopleuron practically trans-
parent, in several irregular small patches; bris-
tles dark, propleurals very short, light colored.
Haltere light on base, darker on stem distally,
knob dark-scaled.

Wmg: Distance between crossveins about
equal to m-cu. Vein R2 about 0.4 of R2-^3;
vein Mi+2 about 0.7 of M beyond m-cu. Iri-
descent bluish-white scales dorsally on caudal
margin of R to slightly beyond arculus and
on anterior margin also at extreme base of
R; larger, almost circular similar scales on
basal 0.5 of Cu; a few white scales on base
of lA; two or three white scales at extreme
base of C; remaining dorsal scales dark, slight-
ly darker on anterior margin, iridescent
bronzy; fringe light, darker on apex.

Legs: Coxae light brown, with translucent
scales and light hairs and bristles; trochanters
light brown, with a few translucent scales and
light hairs; femora dark-scaled above, lighter
below; fore and mid tibiae entirely dark-
scaled; hind tibia with small subapical patch
of light scales on external face; first segment
of front and middle tarsi dark, apical four
segments cream-colored; hind tarsus white-
scaled on 0.8 from base of segment 2 and
entire segments 3-5, remainder dark. Leg I:
femur 1; tibia about 1.0, with small apical
tuft of specialized scales; tarsus 0.90, 0.40, j
0.30, 0.14, 0.08. Leg II: femur 1.11, mod- |
erately swollen basally; tibia 1.60; tarsus 0.93, j
0.41, 0.30, 0.10, 0.09. Leg III: femur 1.20;
tibia 1.22; tarsus 1.10, 0.64, 0.55, 0.22, 0.10. j
Abdomen: Tergites 2, 3, and 4 each with a !
broad median trapezoidal patch of pure white '
scales, patches broader apically; other tergites !
dark-scaled; sternites dull white-scaled. *

MALE (504-45).— Wing: 1.25 mm. Probos- |
cis: 1.40 mm. Front femur: 1.45 mm.

Generally very similar to female. Vertex of
head more extensively dark-scaled. Proboscis !

Vranotaenia in Solomon Islands — Belkin

343

Fig. 5. Uranotaenia civinskii Belkin n. sp. a-d. Adult; e,f, pupa, a, Detail of metathoracic leg of male; b, ninth
tergite of male; c, male genitalia; d, left lateral aspect of head and thorax of female; e, right lateral aspect of anterior
portion of cephalothorax of pupa; /, metanotum and abdomen of male pupa, left ventral, right dorsal. Abbre-
viations as given on page 314,

very strongly swollen apically. Flagellar whorls
about 0.29 of flagellum, about 24 hairs in’
whorl; basal flagellar segments short, basal
part strongly swollen; penultimate segment
slightly less than 2.0 of preceding, apical about
1.7 of penultimate. Leg I: femur 1; tibia 1.0;
tarsus 0.90, 0.33, 0.33, 0.11, O.O6; claws
equal, one broadened, other slender. Leg II:
femur 1.10; tibia 1.40; tarsus 0.90, 0.33, 0.20,
0.06, 0.08, segment 4 with ventral lobe pro-
jecting below 5; claws unequal, enlarged,
smaller 0.6 of larger, both strongly curved.
Leg III: femur 1.16; tibia 1.10, as figured,
straight in basal 0.25, then curved, basad of
curve one short straight and two long curled
bristle; tarsus 1.0, 0.50, 0.44, 0.20, 0.08; claws

equal as on foreleg. Abdomen: tergite 4 with
large patch of dull white scales, broad api-
cally, narrowed triangularly almost to base;
tergites 3 and 2 with smaller median apical
light patches.

MALE GENITALIA (504-45).— As figured.
Ninth tergite long, without bristles; proximal
part very strongly and deeply emarginate;
strong secondary subbasal transverse sclero-
tization; apex widely and deeply emarginate
leaving a strongly projecting, heavily sclero-
tized tooth-like lobe on each side. Proctiger
with a pair of long, weak, ventrolateral sclero-
tizations. Basal lobe of sidepiece with a dorsal
group of eight bristles arranged in three irreg-
ular rows and two ventral bristles. Clasper

344

PACIFIC SCIENCE, VoL VII, July, 1953

as figured. Mesosome with an incompletely
sclerotized ventral bridge in addition to dor-
sal bridge; an apical and a median spine on
each plate, apical spine projecting laterad,
median mesad, spines subequal. Paramere
with a weakly sclerotized median expansion
(not shown in figure).

PUPA (Exuviae of holotype, 504-44).— Ab-
domen: 1.83 mm. Trumpet: 0.32 mm. Pad-
dle: 0.50 mm,

Cephalothorax: Uniformly moderately pig-
mented, slightly darker on leg cases and
metanotum. Trumpet uniformly darkly pig-
mented; length 5.30 of median width; gradu-
ally widened to apex; inner wall indistinct;
tracheoid extending about 0.5 on lateral sur-
face, very poorly developed on mesal surface
except at extreme base; reticulate with im-
brications ending each in one sharp spicule,
more distinct on mesal surface and extending
basad through the tracheoid portion; pinna
about 0.23; distinct mesal slit in meatus ex-
tending about 0.18 of trumpet length. Hairs
moderately pigmented and simple; relative
position, length, and degree of development
as figured; larger branched hairs with slender
base slightly expanded apically where branch-
es arise. Hairs: l(5b), 2(5, 6b), 3(5b), 4(4,
6b), 5(6, 7b), 6(1), 7(4f, base simple), 8(7b),
9(6b, base simple), 10(4b), ll(2b), 12(5, 6b).
Abdomen: Moderately pigmented, somewhat
darker basally, ventral intersegmental sclero-
tizations darker; tergites III-VII each with a
posteromedian patch of distinct small spi-
cules; tergite II with larger patch extending
cephalad; tergite VIII with smaller median
patch; tergal integumentary reticulations in-
distinct; sternites III-VII each with extensive
median patch of small spicules; sternite II
with narrow caudal transverse band of spi-
cules arising from faint arcuate imbrications;
sternite VIII with smaller anteromedian patch
of spicules. All hairs moderately pigmented;
relative position, length, and degree of de-
velopment as figured; larger branched hairs
with inconspicuous short basal part only
slightly enlarged apically where branches

arise. Segment I: hair 1(14, 16 primary branch-
es, dense brush-like secondary branching at
about 0.4-0. 6 from base; about 0.75-0.80
length of tergite), 2(1), 3(3b), 4(5, 6b), 5(3,
4b), 6(2b), 7(2, 3b), 10(2, 3b). Segment II:
hair 0(1), 1(3 primary branches, each with
3-5 secondary apical branches), 2(3, 4b),
3(4b), 4(1), 5(3f), 6(3b), 7(1), 10(2b). Seg-
ment III: hair 0(1), 1(7, 8b), 2(1), 3(5, 6b),
4(3b), 5(2, 3f), 6(4b), 7(1), 8(2f), 10(3, 4f),
12(3b), 13(1), 14(1). Segment IV: hair 0(1),
l(6b), 2(1), 3(5b), 4(4, 5b), 5(2f), 6(3, 4b),
7(1), 8(2, 3b), 10(2f), 12(3b), 13(1), 14(1).
Segment V: hair 0(1), l(4b), 2(1), 3(3f),
4(3b), 5(3b), 6(3, 4b), 7(1), 8(3, 4b), 10(4b),
12(2f), 13(1), 14(1). Segment VI: hair 0(1),
l(4b), 2(1), 3(2f), 4(2, 3b), 5(2, 4f), 6(3b),
7(1), 8(2, 3b), 10(2f), 12(1, 2f), 13(1), 14(1):
Segment VII: hair 0(1), l(3b), 2(1), 3 (4b),
4(3b), 5(1, 2f), 6(3b), 7(1), 8(2, 3b), 10(2f),
12(2, 3f), 13(1), 14(1). Segment VIII: caudal
margin of sternite shallowly emarginate; hair
0(1), 4(1, 3f), 7(2b), 14(1). Segment IX: hair
1(1, about 0.9 of segment length). Paddle as
figured; lightly pigmented; midrib strongly
sclerotized, evanescent apically; external but-
tress distinct proximally; basal pigment bar
distinct; external margin with serrations from
buttress caudad, becoming strongly sclero-
tized on apical half; internal margin with
larger, poorly sclerotized, sparse crenulations;
hair 1 minute, represented largely by socket.
Genital lobe extending to 0.21 of paddle;
with small ventrocaudal apical patch of spi-
cules. Anal segment indistinct, cereal sclerites
not defined. Male genital lobe (exuviae of
allotype, 504-45) extending to 0.24 of paddle;
pair of large ventral lateral basal patches of
strong spicules; anal segment indistinct, ex-
tending slightly beyond apex of IX.

LARVA (Fourth instar exuviae of holotype,
504-44).— Head: 0.70 mm. Siphon: 0.70 mm.
Anal saddle: 0.34 mm.

Head: Width 0.7 of length; ocular bulge prom-
inent; ocular areas lightly pigmented, be-
tween and cephalad of these moderately pig-
mented, caudad very darkly pigmented;

Uranotaenia in Solomon Islands — Belkin

345

Fig. 6. Uranotaenia civinskii Belkin n. sp. Fourth instar larva, a. Head, left dorsal, right ventral; b, left lateral
aspect of distal abdominal segments; c, thorax and proximal abdominal segments, left ventral, right dorsal;
dorsal aspect of left antenna. Abbreviations as given on page 314.

346

PACIFIC SCIENCE, Vol. VII, July, 1953

integumentary imbricate sculpturing very
prominent and uniform. Labrum moderately
long, 0.3 of width at 1-C, anterior margin
very deeply emarginate (twice as deep as fig-
ured). Mental plate (more triangular than
figured) with 21 blunt teeth, faintly indicated.
Hairs of head capsule well pigmented, con-
spicuous; 5, 6-C spike-like, very darkly pig-
mented, apices sharply pointed, shaft very
minutely spiculate, more conspicuously api-
cally; 7, 11-C very minutely barbed, other
hairs simple; relative position, length, and
degree of development as figured. Hair 0(1,
very well developed, reaching to middle of
1-C, very lightly pigmented, flattened and
broadened basally, with external tooth), 1(1,
very slightly curved, short, sharply pointed),
3(1, minute, stiff and heavily pigmented),
4(3b), 5(1), 6(1), 7(4b, short expanded base),
8(1, 2f), 9(2b), 10(2f), ll(5b, slightly ex-
panded short base), 12(2f), 13(5, 6b), l4(bro-
ken off), 15(2f). Antenna 0.23 of head; shaft
distinctly narrowed at 0.6; width at middle
about 0.17 of length; uniformly very darkly
pigmented; spicules small, sparse. Antennal
hairs well pigmented, except apex of 5-A;
relative position, length, and degree of devel-
opment as figured; all single; 1-A placed at
about 0.32, length about 1.4 of antennal
width.

Thorax: All hairs and tubercles strongly pig-
mented; relative position, length, and degree
of development as figured; apices of long
hairs attenuated and sharply pointed; barbs
when present numerous, long and slender and
moderately conspicuous; hair 5-P about as
long as head. Prothorax: hair 0(12, I4d), 1(1),
2(1), 3(8, 9b, large expanded base), 4(3b),
5(1), 6(1), 7(2b), 8(5b), 9(2b), 10(1), ll(3f),
12 (2f), I4(6b, expanded asymmetrical base).
Mesothorax: hair l(6b), 2(1), 3(3, 5b), 4(4,
5b), 5(l),6(l),7(l),8(6b),9(5b), 10(1), 11(1),
12(1), 13(21, 23d), 14(29, 30d). Metathorax:
hair 1(4, 5b), 2(2, 3f), 3(6b), 4(4b), 5(1),
6(2f), 7(8b), 8(llb), 9(5b), 10(1), 11(2, 4b),
12(2f), 13(8b).

Abdomen: Tubercles of segments I, II and

hairs of all segments well pigmented; relative
position, length, and degree of development
as figured; barbs when present slender, lightly
pigmented and sparse. Stellate hairs (1, 6, 13)
with equal branches; 4-II a stellate hair. Seg-
ment I: hair 1(6, 7b), 2(1, 2b), 3(1, 2b), 4(11,
15b), 5(4, 5b), 6(2b), 7(1), 8(2f), 9(9, lib),
10(3, 5b), 12(1, 3b), 13(1, 2f). Segment II:
hair 0(1), 1(4, 5b), 2(1), 3(5b), 4(6b), 5(2,
3b), 6(2b), 7(1), 8(1), 9(1), 10(2b), 11(2, 3f),
12(1, 2f), 13(4, 5b). Segment III: hair 0(1),
1(5, 6b), 2(1), 3(2b), 4(2f), 5(2, 3b), 6(6b),
7(5b), 8(2b), 9(1), 10(2f), ll(3b), 12(2, 3b),
13(5, 7b), 14(1). Segment IV: hair 0(1), 1(6,
7b), 2(1), 3(1, 2f), 4(4, 3f), 5(3, 4b), 6(6, 7b),
7(4, 5b), 8(2, 3f), 9(1), 10(2, 3f), 11(1), 12(1,
2b), 13(6b), 14(1). Segment V: hair 0(1),
l(6b), 2(1), 3(5b), 4(2f), 5(4b), 6(6b), 7(5b),
8(3b), 9(1), 10(2, 3f), 11(1), 12(2, 3b), 13(6,
7b), 14(1). Segment VI: hair 0(1), l(6b), 2(1),
3(2, 4f), 4(2b), 5(3b), 6(6b), 7(3b), 8(2, 3b),
9(1), 10(1, 2b), ll(2f), 12(2f), 13(25, 32d),
14(1). Segment VII: hair 0(1), l(5b), 2(1),
3(8b), 4(2f), 5(3, 4b), 6(6, 7b), 7(4, 5f), 8(7,
9b), 9(2b), 10(2f), 11(2, 3b), 12(1), 13(5, 6b),
14(1). Segment VIII: comb plate heavily
sclerotized, moderately pigmented, ornament-
ed with numerous imbrications with very dis-
tinct spicules caudally; comb scales 8-9, me-
dian ones longer, very sharply pointed, with
conspicuous lateral fringe in basal half as
figured; hair 0(1), 1(3, 5b), 2(2f), 3(6b,
barbed), 4(2f), 5(8, 9b), 14(1). Siphon: as
figured; median width 5.0 of length; uniform-
ly moderately pigmented, ornamented basally
with faint, spiculeless imbrications; pecten
extending to almost 0.5, teeth 16-16 with
lateral and apical fringe as figured; valves as
figured, darker than siphon, ventral valve
moderate; hair l(10b, basal expansion very
short, inconspicuous; barbs extremely small
and sparse), 2(1, moderate, slender), 3-5(only
2 setal rings visible), 6(1, well developed),
7(1, slender, conspicuous), 8(1, slender, about
0.5 of valve), 9(1, moderately strong, hook-
like), 10-12 (not observed), 13(1, strong,
twisted. at base, about length of valve). Seg-

Uranotaenia in Solomon Islands — Belkin

347

ment X: saddle moderately pigmented, darker
dorsad; median width 0.73 of length; imbri-
cations weaker than on comb plate except
caudally; caudal margin with sparse spines of
varying length, proximal ones arising from
imbrications; gills slender, about 0.6 of saddle
length; hair l(5b, about 0.5 of saddle length),
2 (3b, about 3.0 of saddle length), 3 (broken
off), 4a-d (broken off), 4e(l, about 0.2 of
saddle length).

Types

USNM No. 61,418 (holotype, allotype, pa-
ratypes) . Paratypes to be deposited in BMNH,
CU, and CSIR (Canberra); also in collection
JNB.

HOLOTYPE FLP (504-44) and allotype
MLP (504-45) Guadalcanal: Bonegi River
valley, 2 miles from coast, July 18, 1944 (L.
J. Lipovsky, S. Civinski, H. F. Sexauer).

PARATYPES (57M; 56F; 61P; 146L; 9 indi-
vidual rearings), all collected on Guadalcanal,
as follows: llL (14-3) Doma Cove, Still River
area, Oct. 25, 1943 Q'HB et aL) \ 1FLP(64-31),
lM(64-3) Wrights Creek, Nov. 27, 1943
(JNB etaL)\ 2MLP(83-21, 22), 5M, 7L(83-3)
Matanikau River valley, Dec. 5, 1943 (JNB);
1F(86) White River, Dec. 6, 1943 (S. Civin-
ski); 2L(96-2) La Sage Creek, Pt. Cruz, Dec.
13, 1943 (S. Civinski); IM, lF(296-3) Chacon
swamp. Mar. 4, 1944 (L. J. Lipovski, F. B.
Wysocki); 2FLP(397-21, 22), 2L(397-2) Kiwi
Creek, Kukum, May 23, 1943 (V. R. Roa,
F. B. Wysocki); lF(4ll-3) Burns Creek, Lun-
ga. May 27, 1943 (V. R. Roa, F. B. Wysocki);
6L(455) Sally Creek, Doma Cove, June 14,
1944 (JNB); IM, 2F, IL, 1P(472-1) Poha
River valley, Jul. 6, 1944 (JNB et al.)\
IF (490-3) between Poha and Bonegi rivers,
Jul. 14, 1944 (M. Cohen, F. B. Wysocki);
1LP(5 58-21) Matanikau Village, Aug. 4,
1944 (J. J. Cuccio, E. J. McCormick); IMLP
(561-15), 1FLP(561-14), 4M, 6P, 3L, 3P
(561-1) Matanikau River valley, south of
village, Aug. 4, 1944 (L. J. Lipovsky); 4L
(574-3) Matanikau River valley, 2 miles south
of village, Aug. 6, 1944 (JNB, M. Cohen);

1LP(577-31) Matanikau River valley, 3 miles
south of village, Aug. 6, 1944 (JNB, M.
Cohen); 1LP(581-21) Matanikau River val-
ley, 1 mile south of village, Aug. 6, 1944
(JNB, M. Cohen); 1FLP(592-21) Kokum-
bona, Aug. 11, 1944 (J. J. Cuccio, E. J. Mc-
Cormick); 2F, 2L(621-2) Matanikau River
valley, 1 mile south of village, Aug. 22, 1944
(JNB); IM, 1F(654) West Poha swamp. Sept.
23, 1944 (V. R. Roa, F. B. Wysocki); 2M,
lF(66l-2) Poha River, Sept. 23, 1944 (M.
Cohen, J. J. Cuccio, E. J. McCormick);
1L(676) West Poha swamp, Oct. 16, 1944
(JNB, J. Laffoon); 1M(713-1) West Poha
swamp. Sept. 23, 1944 (V. R. Roa, F. B.
Wysocki); 2M, lF(66l-2) Poha River, Sept.
23, 1944 (M. Cohen, J. J. Cuccio, E. J. Mc-
Cormick); 1L(676) West Poha swamp, Oct.
16, 1944 (JNB, J. Laffoon); 1M(713-1) West
Poha swamp, Nov. 3, 1944 (L. J, Lipovsky
et aL)\ 7M, 7F, 15L, 22P(764-3) Matanikau
River valley, 1 to 3 miles south of village,
Dec. 2, 1944 (JNB et al.)\ lF(770-2) Mata-
nikau River valley, Dec. 8, 1944 (J. J. Cuccio,
E. J. McCormick, F. B. Wysocki); 3F(771-2)
Poha River, near mouth, Dec. 8, 1944 (M.
Cohen); 2M, 2F, 9L, 5P(775-3) Matanikau
River valley, 1 mile south of village, Dec. 9,
1944 (JNB et all)\ lF(776-2) same data as
775-3 but in tree hole (JNB et all)\ 2M, 5L,
3P(782-4) Tyler Creek, Matanikau valley,
Dec. 21, 1944 (F. B. Wysocki, Shaw); IM,
2F, 6L, 8P(787-3) Wrights Creek, Matanikau
valley, Dec. 20, 1944 (M. Cohen, F. B. Wy-
socki); IF, 2L(796-3) White River, Dec. 27,
1944 (F. B. Wysocki, Shaw); 8L, 7P(827-2)
Matanikau River valley, 1 mile south of vil-
lage, Jan. 18, 1945 (M. Cohen, F. B. Wy-
socki et al)\ IF, 10L(828-2) Manatikau Vil-
lage (M. Cohen, F. B. Wysocki et al.)\ IM,
2F, 6L(829-2) same as 828-2; lMLP(850-23),
lF(850-2), lM(850-3), 1M(850) Lankford
swamp, Feb. 2, 1945 (Hawkins, Dimard);
6M, 4F(863-4) first tributary south of Poha
Village, Feb. 12, 1945 (JNB et aL)\ IM, IF,
10L(864-3) stream between Poha Villages,
Feb. 12, 1945 (JNB et al.)\ 1M(921) West

348

PACIFIC SCIENCE, VoL VII, July, 1953

Poha swamp, Mar. 20, 1945 (JNB, M. Cohen,
E. Winkler); lE(930-3) Sprague swamp, Do-
ma Cove, Mar. 26, 1945 (JNB, M. Cohen);
2M, 2E(933) Poha River, first tributary north
of village, Apr. 1, 1945 (JNB); lL(958-2)
West Poha swamp, Apr. 28, 1945 (J« J- Ciiccio,
E. J. McCormick, V. R. Roa); lM(970-3)
West Poha swamp. May 6, 1945 (JNB);
3M(1221) Sprague swamp, Doma Cove, rest-
ing on tree buttresses, June 7, 1944 (JNB);
1E(1283) West Poha swamp, flying, Nov. 3,
1944 (L. J. Lipovsky); 2M, 1P(1405) Tassa-
faronga, flying, May 7, 1945 (M. Cohen, E.
Winkler); 3L, Tenaru, Oct. 18, 1943 (J. G.
Eranclemont) ; lE(K-888), Dec. 24, 1943 (K.
L. Knight); 5M, 4E, 8L(K-892) Dec. 26, 1943
(K. L. Knight); 2L(K-897) Dec. 27, 1943
(K. L. Knight); IM, lL(K-949) Aug. 24,
1943 (K. L. Knight); 5L(0-13) Aug. 19, 1943
(P. W. Oman); lE(0-37) June 16, 1944 (P.
W. Oman); 1M(0-251) Sept. 4, 1944 (P. W.
Oman); lL(G-37) Dec. 3, 1943 (A. B. Gur-
ney); 4L(G-74) Nov. 26, 1943 (A. B. Gur-
ney); 4L(G-90) Dec. 5, 1943 (A. B. Gurney);
1L(M-130A) July 22, 1944 (H. E. Milliron).

Uranotaenia civinskii is named in honor of
Stanley Civinski, who contributed greatly to
the knowledge of the mosquitoes of Guadal-
canal through a large number of valuable
collections.

Variation

There is a marked variation in the extent
and intensity of the light-scaling of the head
of the adults, but normally the orbital light
line is wide and a characteristic light cobalt
blue. The light thoracic ornamentation is
somewhat variable in extent and coloration.
The light streak on the stp is normally almost
a pure white but may show a bluish tinge.
The light-scaling of the wing is also quite
variable but is usually extensive and distinctly
cobalt blue. The abdominal light-scaling is
extremely variable. In the female it is rarely
reduced on segments 2 and 3. In some speci-
mens it is more extensive and includes a por-
tion of segment 1 which in the majority of

specimens is a lighter brown than the other
dark portions of the abdomen. In the male,
the light-scaling is usually confined to seg-
ments 3 and 4 and may be even reduced to
segment 4 alone; a few specimens (including
the allotype) show a small light patch on
segment 2 also. This variation in the abdom-
inal light-scaling is apparently an individual
variation for I have not been able to correlate
it with any other variation in the adults, any
characters of the immature stages, or any dif-
ferences in breeding areas. In any given collec-
tion all intergrades may be present, but usually
the majority are either one or the other ex-
treme. The light-scaling of the hind tarsi
always includes the apex of segment 2 but
is quite variable. In some specimens it in-
cludes only the extreme apex of this segment
while in others it may extend basad for as
much as one fourth of its length. The hind
tarsal light-scaling is also more extensive or
at least more conspicuous in the female. The
fore and mid tarsi are extremely variable in
coloration but, as in the holotype, they are
usually distinctly lighter than the rest of the
segments of the leg, and in a few specimens
they are dingy white.

The variation in the chaetotaxy of the im-
mature stages is shown in Tables 1 and 2. The
range of variation is considerable, but no cor-
relation of extremes could be made with any
adult characters or between larva and pupa.

No unusual variations were noted.

The New Georgia specimens agree very '
well with those from Guadalcanal in all stages. !
In specimens from Bougainville the subapical |
light patch on the outer face of the hind tibia
of the female is more conspicuous, more dis- j
tinctly white, and is usually prolonged basad
as a streak. Since no other differences were
noted in the Bougainville material and no
individual rearings are available it appears
best not to recognize this geographical varia- I
tion as a distinct form for the present.

Specimens examined: 102M; 104F; 76P;
211L. Individual rearings: 11 larval. !

Uranotaenia in Solomon Islands — Belkin
Taxonomic Discussion

Uranotaenia civinskii is closely related to IJ.
arg^rotarsis Leicester, 1908, from Malaya, with
which it has been confused in the past. The
only other name proposed to date for mem-
bers of this complex is Tseudouranotaenia pa-
rangensis Ludlow in Theobald, 1910, from the
Philippines, synonymized by Edwards (1922:
460) with JJ. arg^rotarsis. Edwards (1929:
312-313) recognized a single male specimen
from Guadalcanal as being a variety of U.
arg^rotarsis on the basis of the white-scaling
of the hind tarsus embracing the apex of seg-
ment 2, but in his catalog (Edwards, 1932:
98) he no longer listed the Solomons material
as distinct from typical U. arg^rotarsis. Baisas
(1935: 65) found Philippine specimens, pre-
sumably conspecific with P. parangensis, agree-
ing with the description of the Solomons male
in the coloration of the hind tarsus.

Leicester’s original description of U. argiro-
tarsis (1908: 214-215) did not mention the
characteristic, although inconspicuous, mod-
ification of the hind tibia of the male. Ed-
wards (1929: 313), in comparing the Solo-
mons male, noted this characteristic on
Leicester’s type specimen, and Baisas men-
tions it in the Philippine material. It appears,
therefore, that the three geographical forms
belong to the same species complex. No
other species with such a modification of the
hind tibia in the male has been described from
the Australasian or Oriental regions. In the
Ethiopian region, U. pallidocephala Theobald,
1908, has a similar but more complex modi-
fication of the base of the hind tibia, but this
species is markedly different in a multitude
of characters.

U. civinskii differs in the adult stage from
the typical U. argirotarsis from Malaya in the
following characters: hind tarsus white-scaled
from apex of segment 2 in both sexes; ab-
dominal light-scaling usually absent or great-
ly reduced on segment 1 in the female as well
as in the male; hind tibia of female with a
light subapical patch on external face. It
agrees with the Philippine material in the

349

first character but apparently disagrees in the
other two.

The larval stage has not been described
from Malaya, and the description of Philip-
pine material (Baisas, 1935: 65) is not suffi-
ciently detailed to separate U. civinskii. The
characters mentioned by Baisas fall within
the range of variation of IJ. civinskii.

The pupal stage of members of this com-
plex has been figured for Philippine material
by Knight and Chamberlain (1948: 9, fig. 15)
and described and figured for New Guinea
material by Penn (1949: 33-35, fig. 18). Al- ,
though the figures agree in general features,
there is a marked difference in hair l-II and
many minor differences in branching. Penn’s
material agrees with the Solomons specimens
in having hair l-II secondarily branched
whereas it is only primarily branched in the
Philippine specimen. There are differences too
between the New Guinea and Solomons ma-
terial, the principal being the greater number
of branches of hair l-III, the greater length
of hairs 4-V-VII, and the absence of a con-
spicuous external angle on the paddle in the
Solomons form.

On the basis of the above-mentioned dif-
ferences, D. civinskii appears to be quite dis-
tinct from members of the JJ. argirotarsis
complex from Malaya, Philippines, and New
Guinea. Whether or not forms from the latter
two regions are distinct from the typical U.
argirotarsis remains to be determined when
more material is available, but it appears that
they will prove to be distinct.

Biologi

U. civinskii is predominantly a jungle-
stream breeder, utilizing side pools and rock
pools in stream beds in preference to other
habitats. It apparently requires fresh, well-
oxygenated, clear water because, if it is found
in swamps or small pools, these are usually
fed by springs or are in seepage areas. Be-
cause of this requirement it is difficult to rear
under artificial conditions. It is usually found
in densely shaded areas, but when flushed

350

PACIFIC SCIENCE, VoL VII, July, 1953

into open situations it appears to survive
successfully.

Living larvae can usually be recognized in
the field by the long siphon and moderately
pigmented head whereas living pupae can be
distinguished by the moderately long, uni-
formly dark trumpet and long hairs 4-V-VII.

Both males and females are abundantly
found in the vicinity of breeding places and
fly readily in the daytime. They rest most
frequently on shaded, moist rock surfaces and
stream banks but have been found also on
tree buttresses. A small number of males and
females were collected in night hand catches
at lighted quarters.

Distribution

Solomon Islands, Guadalcanal: Tenaru,
IM, IP, Aug. 27, 1928 (R. W. Paine) [Paine
and Edwards, 1929]; generally distributed
throughout the year on north-central and
northwest coast (JNB et al., J. G. Francle-
mont, K. L. Knight, P. W. Oman, A. B.
Gurney, H. E. Milliron et aL) [USNM, CU,
JNB]. Florida: Halavo, 2M, 2F(K-84l), IF
(K-846) Dec. 17, 1943 (K. L. Knight)
[USNM]. Kussell: 4M, 2F, 1944 (R. B. Eads);
Banika, lOL, Mar. 1943 (W. G. Downs)
[USNM]. New Georgia: Segi Pt., 6M, 9F
(B-116) (C. O. Berg) [USNM]; Munda Pt.,
4M, 4F(F-17); 8M, llF(F-20); 6L, Feb. 17,
1944 (J. G. Franclemont) [USNM, JNB],
Bougainville: Empress Augusta Bay, IM, IF.
no date; 5M, 5F, Mar. 1, 1944; 2M, 2F, Mar.
4, 1944 (C. R. Bruck); 1F(G-I4l) Jan. 25,
1944; 2F(G-226) Feb. 19, 1944; 1L(G-310)
no date; IM, lF(G-325) Apr. 14, 1944; IM,
1F(G-341) Apr. 23, 1944; lM(G-349) no
date; lM(G-358) Apr. 27, 1944; lM(G-388)
May 22, 1944; 2L(G-400) June 12, 1944; IM,
2F(G-402) June 14, 1944; 2M, 2F(G-422)
June 27, 1944; 3M, lF(G-436) July 18, 1944
(A. B. Gurney) [USNM, JNBJ.

4. TJranotaenia solomonis Belkin n. sp.

Plates 7, 8

1929. lUranotaenia albescens Taylor. Paine in

Paine and Edwards, Bui. Ent. Res. 20:
304 (misident.).

1944. Uranotaenia albescens Taylor. Knight,
Bohart, and Bohart, Keys Mosq. Aus-
tralasian Reg. p. 15, 68 {partim).

Diagnosis

ADULT.— Head dark centrally; very narrow
white orbital line, not expanded laterally; no
erect occipital scales; conspicuous white front-
al tuft. Narrow supra-alar line of white scales
long, almost reaching scutal angle; apn and
stp each with narrow line of white scales form-
ing streak in line with white-scaling of head.
Hind tarsus cream-white beyond middle of
segment 3. Wing with white scales on base
of R, Cu, and a few on lA. Abdominal ter-
gites 1-3 broadly white-scaled, 4 with apical
median patch, 5 with narrow apical trans-
verse band extending to sternite. Male legs
without striking modifications.

PUPA.— Trumpet length 4.0 median width;
tracheoid extending to about 0.5; uniformly
darkly pigmented; slit in meatus. Hair l-II
usually 10, lib, all primary; 4-V-VIII usu-
ally 4-6b, shorter than following tergite. Pad-
dle serrations distinct on apical half of ex-
ternal margin.

LARVA. — Head longer than wide; hairs 5,
6-C strong spikes; 4-C short, 3, 4b; 7-C 1

4b (3-5); 9-C long, 3b (2-4). Antennal hairs '
simple; hair 1-A short, basad of middle. Tho- |
rax and abdomen with strong dorsal and ven- I
tral stellate tufts. Thoracic hairs 9, 10-P well i
developed; hairs 9-M, T, 8-M, 7-T long, '
multiple, barbed; 4-P 2b; 7-P 2b(2-3); 9-P I
2b(2-3); 14-P single; long hairs with apices
attenuated. Abdominal hairs 6-III-VIII stel-
late, much shorter than 6-1, II, without basal j
tubercles; l-I, II 6-8b(4-ll); 6-1, II 3b, j
branches uneven; 6-II, IV 7-1 lb. Comb
plates separate; scales fringed. Siphon index
about 4.0; pecten extending to 0.45; pecten j
teeth very broad, fringed apically and laterally,
11(11-13); hair 1-S moderate 10, llb(9-13), j
uneven; valves moderate; hair 9-S weak, hook- j
like; 13-S moderate, twisted at base. Anal

Uranotaenia in Solomon Islands-^BELKIN

351

Fig. 7. Uranotaenia sokmonis Belkin n. sp. a-d. Adult; e,f, pupa, a, Detail of mesothoracic leg of male; h, ninth
tergite of male; c, male genitalia; d, left lateral aspect of head and thorax of female; e, right lateral aspect of anterior
portion of cephalothorax of pupa; /, metanotum and abdomen of male pupa, left ventral, right dorsal. Abbre-
viations as given on page 314,

segment long; saddle margin with moderate
apical spines; gills about 0.6 of saddle length;
hair 1-X 5b(5-7), moderate; 2-X 3b; 3-X 2b;
4a-X 3b; 4b-X 2b.

Description

FEMALE (300-212). “Wing: 1.70 mm. Ab-
domen: about 1.10 mm. Proboscis: 1.10 mm.
Front femur: 1.25 mm.

Head: Conspicuous frontal tuft of elongate
white scales partly overlaid by elongate dark
scales; very narrow orbital line of broader
white scales partly overlaid by elongate dark
scales, produced as narrow line toward apn
laterally; recumbent dark scales broader cau-
dad, conspicuously iridescent in a posterior
patch, almost black in front; short erect scales

restricted to extreme caudal part of occiput,
apparently three pairs or more; occipitals
dark, 1:3. Clypeus very dark. Palpus about
0,08 of proboscis; with small dark scales and
numerous long hairs. Proboscis slightly swol-
len apically, dark-scaled, appearing lighter
ventrally; short hairs on shaft and on apex;
labella light brown, moderately hairy. Anten-
na about 1.2 of proboscis; torus very dark,
with a few small hairs mesally; flagellum dark,
about six bristles in whorl; hairs and scales
scanty at base, longer and more numerous
apically; apical segments gradually length-
ened, ultimate only slightly longer than penul-
timate.

Thorax: Scutal integument uniformly dark
brown; dense vestiture of recumbent narrow

352

PACIFIC SCIENCE, Vol. VII, July, 1953

elongate bronzy scales, somewhat broader in
prescutellar space; narrow supra-alar line of
white scales as figured, anterior scales small,
moderately elongate, becoming larger caudal-
ly and forming a small tuft in front of wing
root; acrostichals strong, other bristles strong,
all dark. Scutellum brown; median lobe prom-
inent, with four strong and one weak bristle,
completely covered with short broad scales
projecting slightly over base of bristles; lateral
lobe with one weak and three stong bristles,
completely covered with more elongate scales
projecting strongly over base of bristles. Post-
notum uniformly dark brown. Pleural inte-
gument dark brown, light iridescent areas in
line with white-scaling; scaling and chaeto-
taxy as figured; scales white; line on apn very
narrow, anterior scales short, posterior form-
ing a tuft; sternopleural line with recumbent
scales becoming semierect caudally, three or
four scales in row, scales moderately elongate;
one sternopleural patch restricted to lower
part, scales white; bristles very dark and
strong. Haltere brown at base and stem, dark-
scaled on knob.

Wing: Distance between cross veins about 2.0
of m-cu. Vein R2 about 0.43 of R2+3; vein
M]+2 about 0.8 of M beyond m-cu. Flat white
scales dorsally on caudal margin of R to
slightly beyond arculus and at extreme base
of anterior margin of R; similar scales on
basal 0. 5-0.6 of Cu; a few white scales on
base of lA; two or three white scales on ex-
treme base of C; remaining dorsal scales dark,
strongly iridescent on anterior margin; fringe
light, darker on apex.

Legs: Coxae and trochanters dark brown, with
translucent scales except for flat white scales
proximally on fore and mid coxae; femora
dark-scaled above, light-scaled below; tibiae
dark above, appearing lighter below; fore and
mid tarsi dark basally, lighter apically, partic-
ularly below; hind tarsus distinctly creamy
white beyond middle of segment 3. Leg I:
femur 1; tibia 1.10; tarsus 0.90, 0.42, 0.26,
0.10, 0.07. Leg II: femur 1.10, slightly swollen
basally; tibia 1.44; tarsus 0.90, 0.48, 0.26, 0.13,

0.08. Leg III: femur 1.07; tibia 1.22; tarsus

1.12, 0.64, 0.50, 0.26, O.O9.

Abdomen: Tergite 1 white-scaled except for
small sublateral patch of dark scales; tergites
2 and 3 with broad median white patch ex-
tending from base to apex; tergite 4 with
median apical patch of white scales; tergite
5 with narrow apical transverse band of white
scales extending to sternite; remainder of
tergites dark-scaled; sternites with light
brown to cream-colored scales.

MALE (671-31).— Wing: 1.50 mm. Probos-
cis: 1.20 mm. Front femur: 1.10 mm.

Generally very similar to female. Proboscis
moderately swollen. Flagellar whorls about
0.4 of flagellum, about 30 bristles in whorl;
penultimate segment about 3.0 of preceding,
apical about 0.9 of penultimate. Leg I: femur
1; tibia 1.23; tarsus 0.68, 0.46, 0.25, 0.15,
0.09; claws equal, one broadened. Leg II:
femur 1.15, strongly swollen basally; tibia
1.57; tarsus 0.98, 0.47, 0.24, 0.12, 0.09, seg-
ments 4 and 5 as figured; -smaller claw invisi-
ble. Leg III: femur 1.12; tibia 1.41, apex
slightly enlarged; tarsus 1.23, 0.76, 0.57, 0.30,
0.10, white-scaling on segment 3 less ex-
tensive than in female; claws as on foreleg.

MALE GENITALIA (671-31).— As figured.
Proximal border of ninth tergite deeply and
narrowly emarginate, sclerotization evanes-
cent in emargination; apical border shallowly
emarginate; lateral lobe in form of a long,
slender, heavily sclerotized tooth, prolonged
ventrad basally as a sclerotization; median i
bridge less than length of tooth, with a sub- |
apical sclerotization. Proctiger without dis- !l
tinct sclerotizations. Basal lobe of sidepiece j
with a dorsal group of one very strong, three I

medium, and two small bristles and a ventral jl

group of one long and one short bristle. |
Clasper as figured. Mesosome with a com- '
plete dorsal sclerotization; apex with two :
sharp subequal spines; median part with a ||
ventrolateral lobe bearing three teeth, middle
one longer. Paramere slender at base, expand- .
ed in distal half and with a bifurcation apically. i

PUPA (Exuviae of holotype, 300-212). — |l

Uranotaenia in Solomon Islands — -Belkin

353

Fig. 8. Uranotaenia solomonis Belkin n. sp. Fourth instar larva, a, Head, left dorsal, right ventral; b, left lateral
aspect of distal abdominal segments; c, thorax and proximal abdominal segments, left ventral, right dorsal;
d, dorsal aspect of left antenna. Abbreviations as given on page 314.

354

PACIFIC SCIENCE, Vol. VII, July, 1953

Abdomen: 1.80 mm. Trumpet: 0.28 mm. Pad-
dle: 0.52 mm.

Cephalothorax: Moderately pigmented; ap-
pendage cases, meso- and metanotum darker.
Trumpet dark throughout; length 4.0 median
width; gradually widened from base to about
0.4, then almost parallel-sided; inner wall dis-
tinctly separated except in pinna; tracheoid
extending to about 0.5 on lateral surface, well
developed on mesal surface; reticulate dis-
tinct, without conspicuous spicules; pinna
0.27; distinct mesal slit in meatus extending
about 0.27 of trumpet length. Hairs lightly
to moderately pigmented; relative position,
length, and degree of development as figured;
larger branched hairs with slender basal stalk
only slightly expanded apically where branch-
es arise. Hairs: 1(4, 5b), 2(5b), 3(3, 4b), 4(5,
7b), 5(6, 8b), 6(1), 7(4f), 8(6, 7b, external
branches shorter), 9(4, 5b), 10(3b), 11 (4b),
12(3b).

Abdomen: Moderately and uniformly pigment-
ed, ventral intersegmental scleroti2ations dark-
er; tergites II-VII each with a posteromedian
patch of distinct small spicules; tergite VIII
with anteromedian patch of shorter, stronger
spicules; tergal integumentary reticulations
very faintly visible on I-IV, invisible caudally;
sternites III-VII each with extensive median
patch of small spicules; sternite II with caudal
transverse band of lines of shorter, heavier
spines; sternite VIII with anteromedian patch.
All hairs moderately pigmented; relative po-
sition, length, and degree of development as
figured; larger hairs as on cephalothorax. Seg-
ment I: hair 1(22, 26 primary branches, each
branch densely barbed from near base, some
apical barbs appearing as secondary branches;
about as long as segment), 2(1), 3(4, 5b),
4(6b), 5(4, 5f), 6(2b), 7(2b), 10(4b). Segment
II: hair 0(1), 1(8, lOb), 2(6, 5b), 3(6, 7b),
4(1), 5(4, 5f), 6(2b), 7(1), 10(4, 5b), 12(0).
Segment III: hair 0(1), 1(8, 9b), 2(1), 3(9b),
4(6, 8b), 5(2, 3b), 6(3, 4b), 7(1), 8(2b),
10(3b), 12(1, 2f), 13(1), 14(1). Segment IV:
hair 0(1), 1(7, 8b), 2(1), 3(7, 8b, long stem),
4(7, 9b), 5(2, 3f), 6(4b), 7(1), 8(4b), 10(3b),

12(2f), 13(1), 14(1). Segment V: hair 0(1),
l(6b), 2(1), 3(3, 4f), 4(6, 7b), 5(5, 6b), 6(3b),
7(1), 8(4b), 10(4, 5b), 12(2f), 13(1), 14(1).
Segment VI: hair 0(1), l(5b), 2(1), 3(2f),
4(5b), 5(3, 4f), 6(2, 3b), 7(1), 8(3, 5b), 10(3f),
12(2f), 13(1), 14(1). Segment VII: hair 0(1),
l(5b), 2(1), 3(6b), 4(4, 5b), 5(3f), 6(3b), 7(2,
3b), 8(3b), 10(2f), 12(3f), 13(1), 14(1). Seg-
ment VIII: caudal margin of sternite slightly
emarginate; hair 0(1), 4(3f), 7(3, 4b), 14(1).
Segment IX: hair 1(1, about 0.55 of segment
length). Paddle as figured; lightly pigmented;
midrib strongly sclerotized, evanescent apical-
ly; external buttress distinct proximally; basal
pigment bar distinct; external margin with
distinct serrations on apical half; internal mar-
gin with apical serrations gradually changing
to larger, poorly sclerotized crenulations at
about 0.12 from apex; hair 1 slender, elongate.
Genital lobe extending to 0.22 of paddle,
with large ventrocaudal patch of spicules.
Anal segment indistinct; cereal sclerites well
defined. Male genital lobe (exuviae of allo-
type, 671-31) extending to 0.22 of paddle;
pair of large ventral patches of weak spicules;
anal segment indistinct.

LARVA (Fourth instar exuviae of holotype,
300-212).— Head: 0.70 mm. Siphon: 0.60
mm. Anal saddle: 0.33 mm.

Head: Width 0.8 of length; ocular bulge mod-
erately prominent; pigmentation extremely
dark, except in ocular region; integumentary
imbricate sculpturing very strong, uniform
and distinct. Labrum moderate, 0.30 of width
at 1-C, median emargination deeper than
shown on figure. Mental plate small, with
about 15 teeth. Hairs of head capsule darkly
pigmented, conspicuous; 5, 6-C spike-like, i
deep black, apices frayed, shaft strongly spi-
culate, becoming barbed distally; 7, 9, H-C jj
with minute barbs; other hairs simple; rela-
tive position, length, and degree of develop-
ment as figured. Hair 0(1, curved, sharply
pointed, flattened and expanded laterally at
base, reaching beyond base of 1-C), 1(1, mod-
erately curved, sharply and abruptly pointed),
3(1, well developed and pigmented), 4(3b),

Uranotaenia in Solomon Islands — Belkin

5(1, hair on right side forked at tip), 6(1),
7(3, 4b), 8(2f, strong), 9(3b), 10(2f, strong),
11(4, 5b), 12(2f), 13(5b), 14(1, short, irregu-
lar, flattened and lightly pigmented), 15 (not
seen). Antenna 0.22 of head; shaft wide at
base, gradually and irregularly narrowed to
apex; width at middle about 0.15 of length;
uniformly and extremely darkly pigmented;
spicules small, dark, sparse. Antennal hairs
strongly pigmented, except apex of 5-A; rela-
tive position, length, and degree of develop-
ment as figured; all single; 1-A placed at 0.37
from base, stiff, length 1,1 of antennal width.
Thorax: All hairs and tubercles very strongly
pigmented; relative position, length, and de-
gree of development as figured; apices of long
hairs attenuated; barbs when present numer-
ous and strong; hair 5-P about 0.8 of head
length. Pro thorax: hair 0 (9b), 1(1), 2(1), 3 (8b,
short expanded base), 4(2b), 5(1), 6(1),
7(2b), 8(7b), 9(2b, strong), 10(1), ll(3f),
12(1), 14(1, minutely spiculate). Mesothorax:
hair 1(8, 9b), 2(1), 3(3, 4fb), 4(3f), 5(1), 6(1),
7(1), 8(6b), 9(4, 5b), 10(1), 11(1), 12(1), 13
(34, 35d), 14(37, 39d). Metathorax: hair l(7b),
2(2f), 3(6b), 4(4b), 5(1), 6(2f), 7(7b), 8(15,
17b), 9(5b), 10(1), 11(1), 12(2f), 13(9, 11b).
Abdomen: Tubercles and hairs of all segments
strongly pigmented; relative position, length,
and degree of development as figured; barbs
when present long, conspicuous. Stellate hairs
(1, 6, 13) with equal branches; 4-II a stellate
hair. Segment I: hair l(7b), 2(1), 3(1), 4(9,
lib), 5 (3b), 6(3b, two lower branches short-
er), 7(1), 8(2b), 9(7, 8b), 10(2, 4f), 12(2f),
13(2f). Segment II: hair 0(1), l(6b), 2(1),
3(3, 4b), 4(6, 7b), 5(2, 3b), 6(3b, two lower
branches shorter), 7(1), 8(1), 9(1), 10(3b),
ll(2f), 12(1), 13(6b). Segment III: hair 0(1),
1(7, 8b), 2(1), 3(2, 3f), 4(1), 5(2, 3b), 6(7b),
7(4, 5b), 8(1, 2f), 9(1, 2b), 10(2f), 11(2, 3f),
12(1, 2b), 13(7b), 14(1). Segment IV: hair
0(1), l(8b), 2(1), 3(1), 4(2b), 5(2, 3b), 6(7,
9b), 7(4, 5b), 8(2, 3b), 9(1), 10(2b), 11(1),
12(2b), 13(7b), 14(1). Segment V: hair 0(1),
1(7, 8b), 2(1), 3(2, 3b), 4(1, 2f), 5(2b), 6(9b),
7(6b), 8(2, 3b), 9(2f), 10(2b), 11(1), 12(1,

355

2b), 13(7, 8b), 14(1). Segment VI: hair 0(1),
1(6, 7b), 2(1), 3(2, 3f), 4(2b), 5(2b), 6(8,
10b), 7(2, 3b), 8(2f), 9(1), 10(2f), ll(2f),
12(2, 3f), 13(23, 27d), 14(1). Segment VII:
hair 0(1), 1(6, 7b), 2(1), 3(llb), 4(2f), 5(2,
3b), 6(7b), 7(4, 5f), 8(7b), 9(2b), 10(1),
ll(3b), 12(1), 13(6, 7b), 14(1). Segment VIII:
comb plate heavily sclerotized, moderately
pigmented, ornamented with numerous im-
brications without distinct spicules; comb
scales 7-8, median ones longer, sharply point-
ed, with conspicuous lateral fringe in basal
0.6 as figured; hair 0(1) ,l(5b), 2(2f), 3(6b),
4(2f), 5(8, 9b, branches uneven), 14(1). Si-
phon: as figured; median width 4.0 of length;
heavily pigmented, darker dorsally at base,
ornamented with imbrications as on comb
plate, fainter apically; pecten extending to
0.45, teeth 11-11 with lateral apical and ex-
ternal fringe as figured; valves as figured, very
darkly pigmented, ventral valve moderate;
hair l(llb, basal expansion asymmetrical and
short, no distinct barbs), 2 (broken off), 3-5
(not visible), 6(1, well developed, about as
long as valve), 7(1, slender and conspicuous),
8(1, well developed, about as long as valve),
9(1, very weak, hook-like), 10-12 (not visi-
ble), 13(1, strong, twisted at base, longer than
valve). Segment X: saddle strongly pigment-
ed, darker dorsoapically, imbrications slightly
weaker than on comb plate except distally;
median width 0.54 of length; caudal margin
with sparse spines of varying lengths, prox-
imal ones arising from imbrications; gills
slender, rounded apically, abotit 0.6 of saddle
length; hair l(5b, about 0.5 of saddle length),
2(3b, about 2.6 of saddle length), 3(2b, about
3.3 of saddle length), 4a(3b, about 1.6 of
saddle length), 4b(2b, about 2.0 of saddle
length), 4c(l, about 2.7 of saddle length),
4d(l, about 1.4 of saddle length), 4e(l, about
0.3 of saddle length).

Types

USNM No. 61,419 (holotype, allotype, pa-
ratypes) . Paratypes to be deposited in BMNH,
CU, and CSIR (Canberra); also in coll. JNB.

356

PACIFIC SCIENCE, VoL VII, July, 1953

HOLOTYPE FLP(300-212), Guadalcanal,
Lunga, Apr. 4, 1944 (Brackins), allotype
MLP(671-31), Guadalcanal: Tassi Creek,
mouth of Bonegi River, Oct. 9, 1944 (V. R.
Roa, F. B. Wysocki).

PARATYPES (117M; 138F; 74P; 387L; 34
individual rearings) all collected in various
localities on north and northwest coast of
Guadalcanal as follows: 1M(2), 2M(4), IM
(4-1), 3M, lF(4-2), 1M(6) Doma Cove, Oct.
21, 1943 (JNB); 1F(14-214) Still River, Do-
ma Cove, Oct. 25, 1943 (JNB); 1F(30) Ma-
tanikau River valley, Nov. 11, 1943 (JNB);
1M(52) Burns Creek, Lunga, Nov. 18, 1943
(M. Cohen); lF(64-3) Wrights Creek, Ma-
tanikau valley, Nov. 27, 1943 (JNB et all) \
IM, 3L(83), Matanikau River valley, Dec. 5,
1943 (JNB); lMLP(85-22) La Sage Creek,
Pt. Cruz, Dec. 6, 1943 (S. Civinski); 1F(99)
Lunga, Dec. 13, 1943 (M. Cohen); 1F(100-12)
Burns Creek, Lunga, Dec. 14, 1943 (F. B.
Wysocki); 2MLP(114-211, 213), 1FLP(114-
212), 2L(ll4-2) Butsavu Creek, Lunga valley,
Jan. 3, 1944 (L. J. Lipovsky, F. B. Wysocki);
lF(l44-2) Poha River valley, Jan. 13, 1944
(S. Civinski); 3M, IF, 34L(218-2) Chacon
swamp, Lunga valley, Feb. 21, 1944 (A. W.
Barnes, F. B. Wysocki); 2L(219-2) Lankford
swamp, Kukum, Feb. 21, 1944 (J. J. Cuccio);
2F(289-3) Chacon swamp, Lunga valley, Mar.
29, 1944 (JNB); IM, 1F(296-3) same data
as preceding (L. J. Lipovsky, F. B. Wysocki);
lF(300-2) same data as holotype; 15L(302)
Chacon swamp, Lunga valley, Apr. 7, 1944
(M. Cohen); lF(334-2) Tassafaronga swamp,
Apr. 27, 1944 (S. Civinski); lF(356-2) Belton
Creek, Matanikau, May 6, 1944 (V. R. Roa,
F. B. Wysocki); IF, 2L(374-2) Kukum, May
11, 1944 (Brackins); IL, lP(387-3) Bonegi
River, May 18, 1944 (J. J. Cuccio); 2L(401-3)
Mamara, May 25, 1944 (S. Civinski); ILP
(424-11) Chacon swamp, Lunga valley, June
1, 1944 (JNB); 2L(455) Sally Creek, Doma
Cove, June 14, 1944 (JNB); IF, 1L(472-1)
Poha River valley, July 6, 1944 (JNB, V. R.
Roa, F. B. Wysocki); 3M, lF(506-3) Kukum,
July 18, 1944 (M. Cohen, F. B. Wysocki);

1MLP(564-51), IM, IL, lP(564-5) West Po-
ha swamp, Aug. 5, 1944 (V. R. Roa, F. B.
Wysocki); lF(599-2) same data, Aug. 12,

1944 (V. R. Roa, F. Wysocki); 1MLP(647-
18), 1L(647-1) same data. Sept. 16, 1944 (V.

R. Roa, F. B. Wysocki); 1MLP(654-14),
2FLP(654-15, 16), IM, 1F(654) same data.
Sept. 23, 1944 (V. R. Roa, F. B. Wysocki);
4FLP (671-32, 33, 34, 35), 3M, 6F, 9L, 9P
(671-3) same data as allotype; 1MLP(676-
394), 4FLP(676-34, 38, 39, 392), 7M, 12F,
8L(676-2) West Poha swamp, Oct. 16, 1944
(JNB, J. Laffoon); 2F(681-2) Chacon swamp,
Lunga valley, Oct. 17, 1944 (JNB); 4MLP
(708-105, 106, no, 111), 7FLP(708-101, 102,
103, 104, 108, 109, 112), 20M, lOF, 64L, 21P
(708-1), 1LP(708-107) West Poha swamp,
Nov. 1, 1944 (JNB); 3MLP(713-14, 18, 19),
9M, 20F(713-1) same data, Nov. 3, 1944 (L.

J. Lipovsky et all) \ 2M, 5F, 18L(772-2) Cha-
con swamp, Lunga valley, Dec. 8, 1944 (L.

J. Lipovsky, Shaw, Schultz); 12M, 8F(802-3)
West Poha swamp, Jan. 5, 1945 (M. Cohen,

C. Calloway, Williams); 2lL(8l6-3) same lo-
cality, Jan. 12, 1945 (M. Cohen, F. B. Wy-
socki); IF, l6L(837-3) same locality, Jan. 20,

1945 (J. J. Cuccio. F. B. Wysocki); 2M,
lF(846-2) Kukum, Jan. 30, 1945 (M. Cohen
et all)\ lM(848-2) Kukum, Feb. 1, 1945 (J.

J. Cuccio, C. Calloway, Williams); 1FLP(850-
36), 2M, 3F(850-3) Lankford swamp, Kukum,
Feb, 2, 3, 1945 (M. Cohen, J. J. Cuccio, F.

B. Wysocki); lFLP(921-302), IL, 2P(921-3)
West Poha swamp. Mar. 20, 1945 (JNB, M.
Cohen, E. Winkler); 7M, 6F, 129L(958-2)
same locality, Apr. 28, 1945 (J. J. Cuccio et
all)\ lM(971-2) same locality. May 6, 1945
(JNB); 1F(1020) Lunga, night hand catch,
Jan. 13, 1944 (M. Cohen); 1F(1145) Kukum,
night hand catch, Apr. 5, 1944 (A. W.
Barnes); 2M, 1F(1146) Kukum, night hand 1
catch, Feb. 18, 1944 (J. J. Cuccio, F. B. Wy- |
socki); 1M(1235) West Poha swamp, resting 1
on tree buttress, July 18, 1944 (L. J. Lipovsky, |
F. B. Wysocki); 1F(1338) Lunga valley, night i
hand catch, Jan. 29, 1945 (Tyler); 1F(1373) j
Kukum, night hand catch, Mar. 12, 1945 (C. j

Uranotaenia in Solomon Islands — Belkin

S. Hollingshead, V. R. Roa); lM(Sta. 5)
Matanikau, night hand catch, Mar. 14, 1944
(M. Cohen); 4M, 9F, HL, 3P, Tenaru, Oct.
18, 1943 (J. G. Franclemont) ; 3M(K-949)
Aug. 24, 1943 (K. L. Knight); lM(K-892)
Dec. 26, 1943 (K. L. Knight); lF(K-962)
Dec. 27, 1943 (K. L. Knight); IM, 2F,
4L(NMSS-29-39) Mar. 3, 1943 (Weathersby
and Knapp); lM(0-24) Sept. 10, 1943 (P.
W. Oman); lM(0-33) Sept. 10, 1943 (P. W.
Oman); 2M, 3F(0-34) Sept. 10, 1943 (P. W.
Oman); IM, 2F(0-35) Sept. 13, 1943 (P. W.
Oman); lF(0-38) June 19, 1944 (P. W.
Oman); lL(K-889), 2L(K-908) (K. L.
Knight); IL, lP(K-782) Oct. 14, 1943 (K.
L. Knight).

Variation

The variation in the extent of the light-
scaling of the head and thorax of adults of
U. solomonis is much less than in any other
species of Uranotaenia from the Solomons.
The light scales are almost pure white, show-
ing only rarely a faint bluish tinge, and are
always arranged in narrow lines. The abdom-
inal light-scaling shows a moderate variation
but is always extensive on segments 1,2, and
3. It is frequently greatly reduced on segment
4 and can be almost completely absent from
this segment. The transverse apical white band
on segment 5 is generally very prominent and
reaches the sternite laterally, but in a few spec-
imens it is not complete, perhaps due to a
teneral condition. The white-scaling on seg-
ment 3 of the hind tarsus covers from one to
two thirds of the segment. The light-scaling
of the wing shows the usual variation in ex-
tent but is invariably white.

The variation in the chaetotaxy of immature
stages is shown in Tables 1 and 2. There is
more variation in the branching of the stellate
hairs of the larva than in any other species
studied. Extremes of variation frequently oc-
cur in the same collection and even in the
same individual. The range of variation in the
branching of pupal hairs is moderate. Two
pupae have hair 12-11 well developed, the

357

only instance of the occurrence of this hair
outside of U. quadrimaculata.

The majority of specimens from New
Georgia show a reduction of the light-scaling
on the abdomen with an almost complete
absence of light-scaling on segment 4 and a
corresponding weakening of the apical band
on segment 5, particularly in the female.
There is also in these specimens a more ex-
tensive white-scaling of segment 3 of the hind
tarsus. This is more pronounced in the female,
and in a few specimens there remains only a
faint basal dorsal dark streak on this segment.
I have not been able to find any constant
differences in the immature stages in this ma-
terial and, therefore, am regarding the New
Georgia specimens as a local race for the
present. The Bougainville specimens agree
very well with the material from Guadalcanal.

Specimens examined: 163M; 194F; 89P;
478L. Individual rearings: 36 larval.

Taxonomic Discussion

Uranotaenia solomonis is closely related to U.
albescens Taylor, 1914, described from Towns-
ville, Queensland, and has been treated as a
part of that species in the past (Paine and
Edwards, 1929: 304; Edwards, 1932: 97; Lee,
1944: 19; and Knight, Bohart, and Bohart,
1944: 68). I have not seen specimens of Tay-
lor’s species, but from his description as well
as Lee’s figures of the larva of U. albescens
from New Guinea (1944: 19) it appears that
U. solomonis is at least a distinct subspecies.

The adults of U. solomonis appear to differ
from U. albescens in the more extensive white-
scaling on segment 3 of the hind tarsus as
Taylor states that only its apex is white in
77. albescens. If Taylor’s description is correct,
U. solomonis differs also in that it has a very
narrow line of white scales on apn and less
extensive scaling on abdominal segment 4
and perhaps also 5.

Taylor’s description of the larva is not suf-
ficiently detailed for comparison. It only in-
dicates that the larvae of the two forms belong
to the same group. Lee’s figures (1944: 19)

358

PACIFIC SCIENCE, VoL VII, July, 1953

of a specimen from Milne Bay, Papua, show
striking differences from U. solomonis. In the
latter the anal saddle is almost twice as long
as its width instead of only slightly longer;
hair 1-X is 5-7b instead of 3b; hair 3-X is
2b instead of single; hair 4a-X is 3b instead
of 2b; hair 1-S is 10, llb(9-13) instead of 6b.

Penn’s description and figure (1949: 32-33,
fig. 17) of a single male pupa of U. albescens
from Milne Bay, Papua, indicates an entirely
different species. On the basis of the long
trumpet, peculiar hair 8-C, and elongate hairs
4-V-VII, as well as a number of other char-
acters, I am convinced that Penn described
a member of the tibialis- gtousp as U. albescens.

Uranotaenia solomonis is very sharply sepa-
rated from the other ornamented species of
Uranotaenia from the Solomons in the struc-
ture of the mesosome of the male genitalia.
The "lateral plates" are broadly joined dor-
sally by a heavily sclerotized bridge, and
instead of the two simple teeth on each plate
there are two teeth apically and a ventro-
lateral process bearing three teeth.

U. solomonis and U. albescens appear to form
a species complex peculiar to the Papuan sub-
region. This complex m*ay be related to U.
campestris Leicester, 1908, from Malaya and
U. arguellesi Baisas, 1935, from the Philip-
pines. The latter resemble U. solomonis in ab-
dominal ornamentation but differ in lacking
the white-scaling on the hind tarsus and in
having distinctly bluish scales on the head
and thorax. Another related species may be
U. macfarlanei Edwards, 1914, which has been
reported from Okinawa, China, Hongkong,
Java, Sumatra, Malay Peninsula, and Assam.
This species agrees with U. solomonis in having
the light-scaling white but is distinct in ab-
dominal ornamentation and the absence of
the white-scaling of the hind tarsus.

Biology

Uranotaenia solomonis is the commonest Ura-
notaenia of the Solomons breeding in ground
pools. Usually it is found in rather open situa-
tions, but it will tolerate considerable shade.

It frequently breeds in temporary pools, ruts,
springs, and streams and is common in some-
what open parts of dense jungle swamps.
Stagnant foul water is no deterrent to its
breeding, and it has been collected even in
tin cans. It is interesting to note in connection
with the latter that U. albescens has been also
collected in artificial containers (Taylor, 1914:
706).

This species has such a wide range of breed-
ing habitats that it has been found associated
with practically every species of mosquito
utilizing ground water on Guadalcanal. It is
often found with Anopheles farauti Laveran,
1902, and A. punctulatus Donitz, 1901, but
has a wider range of habitats than even Culex
annulirostris Skuse, 1889-

Living larvae are fairly easily distinguished
in the field from other ground-water Urano-
taenia by their long siphon, black head and
antenna, and normally light-colored bodies.
U. sexaueri has a somewhat similar coloration,
but the pigmentation of the head is not as
strong. Living pupae have a moderately long
trumpet (index 4), dark throughout and quite
like that of U. civinskii, but can be distin-
guished from the latter in the field by the
much shorter abdominal tufts 4-V-VIL

Adults of U. solomonis are commonly seen
in jungle vegetation in proximity to their
breeding places, usually resting close to the
ground but also on tree buttresses. They do
not appear to fly readily until darkness sets in.
In routine night hand catches at lighted quar-
ters they were collected more frequently than
any other Uranotaenia on Guadalcanal, but
this may be due to their greater relative abund-
ance in the populated area.

Distribution

Solomon Islands, Guadalcanal: Generally
distributed throughout the year on north-
central and northwest coast (JNB etal.,]. G.
Eranclemont, K. L. Knight, P. W. Oman,
Weathersby and Knapp) [USNM, CU, JNB].
Russell: Banika, IL, Mar. 1943 (W. G. Downs) ;
IE, 1944 (R. B. Eads) [USNM]. New Georgia:

Uranotaenia in Solomon Islands — Belkin

359

Segi Pt., lM(Be-54a) (C. O. Berg) [USNM];
Munda Pt., lOM, 20F(F-21) Dec. 1943; 4M,
2F, 7L, 2P(F-70) Feb. 14-17, 1944 (J. G.
Franclemont) [USNM, JNB]. Bougainville:
Empress Augusta Bay, IM, 2F (C. R. Bruck) ;
9M, 9F(B-18) Mar. 4, 1944 (C. R. Bruck);
IF, 1944 (W. G. Downs); 2M, 4F(NMSS-
29-1) Dec. 18, 1943 (Weathersby) ; 1L(G-175)
Feb. ■ 3, 1944; lL(G-226) Feb. 19, 1944;
lF(G-247) Feb. 22, 1944; 7M, 2F, IL, 4P
(G-269n) Mar. 6, 1944; 17L(G-306) Mar. 29,
1944; 18L(G-310) Apr. 3, 1944; 1M(G-341);
lF(G-387) ; 2M, IF, 9L(G-388) May 22, 1944;
4M, 4F(G-402) June 14, 1944; lF(G-404a)
July 1, 1944; 1F(G-412) June 19, 1944; 2M,
3F(G-436); lF(G-438) (A. B. Gurney)
[USNM, JNB].

5. Uranotaenia sexaueri Belkin n. sp.

Plates 9, 10

1944. Uranotaenia nivipes (Theobald) . Knight,
Bohart, and Bohart, Keys Mosq. Aus-
tralasian Reg. p. 15 {partim).

Diagnosis

ADULT. — Head entirely white-scaled dor-
sally; small dark patch laterally; few erect
verticals; frontal tuft conspicuous. Lower part
of scutum and upper part of pleura white,
including around the front, with extensive
white-scaling; apn.ppn, and stp each with large
patch of broad white scales; lower half of stp
with dark scales. Hind tarsus white-scaled
from about 0.5 of segment 3; fore and mid
tarsi light on segments 3-5. Wing with con-
spicuous black and white pattern. Abdominal
tergite 1 white-scaled, tergites 2-7 with apical
transverse white bands, broader on basal seg-
ments. Male legs without striking modifica-
tions.

PUPA.— Trumpet length 3.5 median width;
tracheoid extending to about 0.4; dark on
tracheoid, golden brown beyond; slit in mea-
tus. Hair l-II 6-7b, all primary; 4-V-VII
4-6b, slightly longer than following tergite.
Paddle serrations very indistinct, restricted to
extreme apex on external margin.

LARVA. — Head markedly longer than wide;
hairs 5, 6-C strong spikes; 4-C moderate 2b
(2-3); 7-C 4b(3-5); 9-C long 4b(2-5). An-
tennal hairs simple; 1-A very short, near mid-
dle. Thorax and abdomen with well-developed
dorsal and ventral hairs. Thoracic hair 10-P
well developed; 9-P short 4b (4-6); 9-M, T,
8-M, 7-T long, multiple, barbed; 4-P 2b; 7-P
2b(2-4); 14-P very strong (12-l6b); long sin-
gle hairs with apices blunt or frayed. Abdom-
inal hairs 6-III-VIII stellate, much shorter
than 6-1, II, without basal tubercles; l-I, II
7-9b(7-ll); 6-1,11 2b, branches uneven;
6-III, IV 7-8b(7-ll). Comb plates separate;
scales fringed basally only, one or two median
ones enlarged. Siphon index about 5.5; pec-
ten extending to about 0.3; pecten teeth nar-
row, fringed apically and laterally; valves long;
hair 1-S moderate, 7, 8b(6-9); 9-S weak,
straight, inconspicuous; 13-S long, strong,
twisted at base. Anal segment moderate; sad-
dle margin with numerous moderate, narrow,
apical spines; gills about 0.5 of saddle length;
hair 1-X 6, 5b, moderate; 2-X 3b(2-4); 3-X
2b; 4a-X 3b; 4b-X 2b.

Description

FEMALE (713-15).— Wing: 1.55 mm. Ab-
domen: 1.10 mm. Proboscis: 1.15 mm. Front
femur: 1.15 mm.

Head: Vertex with decumbent scales all white;
central and caudal scales with faint iridescence;
scales elongate but rounded apically; frontal
tuft present but largely missing in specimen;
erect occipital scales missing in specimen; in-
ner orbital bristle absent, outer orbitals three.
Postgena with an orbital patch of dark scales
below orbital bristles, followed by grayish-
brown scales. Clypeus dark brown. Palpus
about 0.05 of proboscis, with small scales and
long hairs. Proboscis slightly swollen on ex-
treme apex; dark-scaled, lighter ventrally;
shaft with short hairs laterally; just before
apex a few long hairs, apex with numerous
hairs; labella brown, with vestiture of light
hairs. Antenna about 1.2 of proboscis; torus
light brown, with a few short hairs dorsally

360

PACIFIC SCIENCE, VoL VII, July, 1953

and mesally; flagellum darker; whorls short,
with 6 bristles; hair and scales scanty basally,
more numerous apically; apical segment with
long light hairs, about 1.3 of penultimate.
Thorax: Integument extensively white lateral-
ly as indicated on figure, boundary of white
integument and scales indicated by stippling.
Median portion of scutum with brown inte-
gument and dense vestiture of recumbent nar-
row, elongate, bronzy-brown scales; lateral
portions including a broad anterior border
white-scaled; anterior acrostichal area with a
patch of small, broad, recumbent white scales;
lateral border in front of scutal angle with
scaling denser than shown on figure; supra-
alar scales elongate becoming very slender
and forming a tuft in front of wing root;
acrostichals unusually strong, dorsocentrals
fewer than usual, supra-alars few in number,
all bristles dark. Scutellum dark brown; me-
dian lobe short, partially denuded in speci-
men, scales short, rounded apically, projecting
slightly over base of the three bristles; lateral
lobe with three heavy bristles, longer than
those on median lobe, scaling elongate, pro-
jecting strongly over bases of bristles. Post-
notum dark brown centrally, cream-colored
laterally. Pleural integument extensively white
as indicated on figure, lower part brown; scal-
ing and chaetotaxy as figured; apn entirely
white, with elongate white scales; ppn with
scattered small white scales, almost circular
in outline; stp extensively scaled, scales of
upper part white, dark on lower part, line of
separation very sharp; larger bristles dark,
smaller light, propieurals minute. Haltere
white on base, darker on upper part of stem,
knob dark-scaled.

Wing: Distance between crossveins about 1.7
of m-ciL Vein R2 about 0.4 of R2+3; vein M1+2
about 0.82 of M beyond m<u. Conspicuous
black and white pattern of scaling on dorsal
surface; anterior border dark, median and pos-
terior part white with a subapical transverse
dark band, apex white beyond base of cell
R2; C dark except at base on caudal margin
and at apex; R white at base for 0.3 of wing

length, then dark to 0.42, followed by white
to 0.56 and then dark to level of base of cell
R2; a subapical dark irregular transverse band
including C, R, portion of R2+3 equal to cor-
responding dark area of Ri, five scales on
R4+5 at 0.64-0.76, four scales on M at 0.65-
0.75, five scales on Cui at 0.63-0.74, Cu2 from
0.65 to apex; two dark scales at preapical curv-
ature of lA. Fringe white.

Legs: Coxae and trochanters brown, scales not
seen, larger bristles dark, smaller brown; fe-
mora dark-scaled, light ventrally; tibiae dark,
appearing paler ventrally in some lights; first
two segments of front and middle tarsi dark,
apical three segments lighter; hind tarsus with
apical 0.6 of segment 3 and entire segments
4 and 5 white-scaled, remainder dark above,
lighter below. Leg I: femur 1; tibia about 1.0;
tarsus 0.78, 0.40, 0.23, 0.14, 0.07. Leg II:
femur 1.10, strongly swollen basally; tibia
1.43; tarsus 0.90, 0.42, 0.30, 0.10, 0.08. Leg
III: femur 1.03; tibia 1.20; tarsus I.06, 0.57,
0.43, 0.23, 0.10.

Abdomen: Tergite 1 almost entirely white-
scaled; 2-5 each with apical transverse white
band, broader in center where it occupies
about 0.4 of segment; 6 and 7 with a few
apical white scales; remainder of tergites dark-
scaled; sternites with dull white scales.

MALE (713-12).-— Wing: 1.40 mm. Probos-
cis: 0.98 mm. Front femur: 0.90 mm.

Generally very similar to female. Proboscis
more strongly swollen at extreme apex. Fla-
gellar whorls about 0.45 of flagellum, about
20 bristles in whorl; basal flagellar segments
short, basal part slightly swollen; penultimate
2.0 of preceding, apical equal to penultimate.
Leg I: femur 1; tibia 1.0; tarsus 0.63, 0.36,
0.26, 0,14, 0.08; claws equal, one broadened,
other slender. Leg II: femur 1.13; tibia 1.50;
tarsus 0.90, 0.41, 0.27, 0.10, 0.10, segment 4
with median ventral lobe projecting below
5, segment 5 with dorsoapical lateral lobes;
claws unequal, enlarged, smaller about 0.5
of larger, both strongly curved. Leg III: femur
1.13; tibia 1.30; tarsus 1,15, 0.60, 0.45, 0.25,
0.09; claws as on foreleg. Abdominal tergite

Uranotaenia in Solomon Islands — Belkin 36 1

Fig. 9. Uranotaenia sexaueri Belkin n. sp, a-d^ Adult; e, f, pupa, a. Detail of mesothoracic leg of male; b, ninth
tergite of male; c, male genitalia; d, left lateral aspect of head and thorax of female; e, right lateral aspect of anterior
portion of cephalothorax of pupa; f, metanotum and abdomen of male pupa, left ventral, right dorsal. Abbre-
viations as given on page 314.

1 white-scaled as in female; tergites 2-7 less
extensively white-scaled, apical bands nar-
rower.

MALE GENITALIA (713-12).— As figured.
Ninth tergite long, without bristles; proximal
part shallowly emarginate; apex broadly but
shallowly emarginate; lateral lobes slightly
projecting, with ventrolateral sclerotization.
Proctiger without visible sclerotization. Basal
lobe of sidepiece with a dorsal group of two
strong bristles and three weak ones and one
strong ventral bristle. Clasper as figured. Me-
sosome expanded on each side into a broad,
basal, heavily sclerotized plate on ventral sur-
face; apical spine slightly curved, projecting
laterad; median spine heavy at base, strongly
curved inward and with a basal tooth. Para-
mere expanded near base.

PUPA (Exuviae of holotype, 713-15). — Ab-
domen: 1.62 mm. Trumpet: 0.20 mm. Paddle:
0.42 mm.

Cephalothorax: Uniformly moderately pig-
mented, darker on meso- and metanotum and
bases of wing and appendage cases. Trumpet
dark on tracheoid, golden brown beyond;
length 3.50 median width; gradually widened
from base, almost parallel-sided on apical half;
inner wall distinctly separated only in trache-
oid portion; tracheoid extending about 0.4;
reticulations distinct; pinna about 0.25; a dis-
tinct slit in meatus extending about one-half
distance of reticulate. All hairs moderately
pigmented and simple; relative position,
length, and degree of development as figured;
larger branched hairs with a short stem only
slightly flattened apically where branches arise.

362

P ACIFIC SCIENCE, VoL VII, July, 1953

Hairs: l(6b), 2(5, 6b), 3(5, 6b), 4(7b), 5(8,
9b), 6(1), 7(6f), 8(7, 8b), 9(7, 8b), 10(5, 6b),
11(4, 5b), 12(4, 5b).

Abdomen: Uniformly moderately pigmented
throughout; tergites II-VII each with a large
posteromedian patch of distinct small spic-
ules; tergite VIII with a similar patch antero-
medially; tergites II-VIII with distinct poly-
gonal reticulations, more obvious on caudal
segments; sternites III-VIII each with exten-
sive median patch of small spicules; sternite
II with extensive posteromedian area covered
with numerous short, transverse, arcuate lines
of more heavily sclerotized spicules. All hairs
moderately pigmented; relative position,
length, and degree of development as figured;
larger branched hairs without strong basal
plates, branching taking place short distance
from base. Segment I: hair 1 (about 17 pri-
mary branches, moderate number of strong
secondary branches; about 0.85 length of ter-
gite), 2(1), 3(3b), 4(4b), 5(5, 6b), 6(4b), 7(1,
2f), 10(4, 5f). Segment II: hair 0(1), l(8b),
2(4, 5b), 3(5b), 4(1), 5(3, 4f), 6(2b), 7(1),
10(3f). Segment III: hair 0(1), l(7b), 2(1),
3(6, 7b), 4(4, 5b), 5(3, 4f), 6(2, 3b), 7(1),
8(3f), 10(2f), 12(2, 3f), 13(1), 14(1). Segment
IV: hair 0(1), l(6b), 2(1), 3(5, 6b), 4(6b),
5(3f), 6(2, 3b), 7(1), 8(1, 2b), 10(1, 2f),
12(2f), 13(1), 14(1). Segment V: hair 0(1),
1(5, 6b), 2(1), 3(3, 5f), 4(4, 6b), 5(3, 4b), 6(2,
3b), 7(1), 8(1, 3f), 10(4b), 12(2f), 13(1),
14(1). Segment VI: hair 0(1), 1(4, 5b), 2(1),
3(3, 4f), 4(4, 5b), 5(5f), 6(3b), 7(1), 8(2, 3f),
10(2f), 12(2, 3f), 13(1), 14(1). Segment VII:
hair 0(1), 1(3, 4b), 2(1), 3(4, 5b), 4(4, 5b),
5(2, 4f), 6(2, 3b), 7(3b), 8(3, 4b), 10(2f),
12(2f), 13(1), 14(1). Segment VIII: caudal
margin of sternite truncate; hair 0(1), 4(2, 3f),
7(2, 3b), 14(1). Segment IX: hair 1(1, 0.67 of
segment length). Paddle as figured; lightly
pigmented; midrib strongly sclerotized; ex-
ternal buttress poorly defined; basal pigment
bar distinct; external margin with small teeth
distinct apically only; internal margin with
even shorter teeth; hair 1 extremely weak and
short. Genital lobe extending to about 0.3 of

paddle; with ventrocaudal patch of strong
spicules. Anal segment indistinct. Male gen-
ital lobe (exuviae of allotype, 713-12) extend-
ing to 0.3 of paddle; pair of large ventral
lateral patches of strong spicules; anal seg-
ment indistinct, extending just beyond apex
of IX.

LARVA (Fourth instar exuviae of holotype,
713-15). —Head: 0.65 mm. Siphon: 0.70 mm.
Anal saddle: 0.36 mm.

Head: Width 0.7 of length; ocular bulge
prominent; darkly and uniformly pigmented
except for light ocular areas; integumentary
imbricate sculpturing very prominent and uni-
form. Labrum short, 0.25 of width at 1-C,
anterior margin very deeply emarginate. Men-
tal plate with 15 blunt teeth, conspicuously
defined apically only. Hairs of head capsule
well pigmented, conspicuous; 5, 6-C spike-
like, very darkly pigmented, apices frayed,
shaft minutely spiculate; 7, 11-C minutely
barbed, other hairs simple; relative position,
length, and degree of development as figured.
Hair 0(1, slender, only slightly flattened,
reaching beyond base of 1-C), 1(1, curved,
bluntly pointed), 3(1, minute stiff spine),
4(2b), 5(1), 6(1), 7(4b), 8(3, 4f), 9(4b),
10(2f), ll(8b), 12(3f), 13(6, 7b), 14(1, short,
heavily sclerotized, expanded spine), 15(3,
4b). Antenna 0.23 of head; shaft of rather
uniform width, but irregular; width at middle
about 0.15 of length; uniformly darkly pig-
mented; spicules small and sparse. Antennal
hairs well pigmented except 5- A; relative po-
sition, length, and degree of development as
figured; all single; 1-A placed at middle,
length 0.75 of antennal width.

Thorax: All hairs and tubercles well pigment-
ed; relative position, length, and degree of
development as figured; apices of long hairs
frayed or blunt, appearing as if broken off;
barbs when present numerous, slender, and
lightly pigmented; hair 5-P about 0.8 of head
length. Prothorax: hair 0(15, l6d), 1(1), 2(1),
3(broken off), 4(2b), 5(1), 6(1), 7(2b), 8(8,
9b), 9(4b, equal), 10(1), 11(2, 3b), 12(1),
I4(l4b, large symmetrical semilunar expand-

Uranotaenia in Solomon Islands — Belkin

363

Fig. 10. Uranotaenia sexaueri Belkin n. sp. Fourth instar larva, a. Head, left dorsal, right ventral; b, left lateral
aspect of distal abdominal segments; c, thorax and proximal abdominal segments, left ventral, right dorsal;
dorsal aspect of left antenna. Abbreviations as given on page 314.

364

ed base). Mesothorax: hair 1 (broken off),
2(1), 3(3bf), 4(4f), 5(1), 6(1), 7(2f), 8(bro-
ken off), 9(5b), 10(1), ll(5b), 12(1), 13(28,
30d), 14(24, 30d). Metathorax: hair l(9b),
2(3f), 3(4, 5b), 4(2b), 5(2b), 6(2, 3f), 7(8b),
8(lld), 9(6b), 10(1), 11(3, 4f), 12(1, 2f),
13(13b).

Abdomen: Large hairs and tubercles of seg-
ments I and II well pigmented; hairs of fol-
lowing segments moderately pigmented; rela-
tive position, length, and degree of develop-
ment as figured; barbs when present slender
and lightly pigmented. Stellate hairs (1, 6, 13)
with equal branches; 4-II a stellate hair. Seg-
ment I: hair l(broken off), 2(1), 3(1), 4(10,
13b), 5(2, 3b), 6(2b), 7(1), 8(1, 2f), 9(5b),
10(2, 3f), 12(1, 2f), 13(1, 2f). Segment II:
hair 0(1), l(broken off), 2(1), 3(2, 3b), 4(6b),
5 (3b), 6(2b, one side lower branch 3f), 7(1),
8(1), 9(3b), 10(2, 3b), 11(1, 2f), 12(1), 13(7b).
Segment III: hair 0(1), l(broken off), 2(1),
3(2, 3b), 4(1), 5(2b), 6(7b), 7(4, 5b), 8(1, 3f),
9(2b), 10(1, 3f), 11(2, 3b), 12(1, 2b), 13(7b),
14(1). Segment IV: hair 0(1), 1 (broken off),
2(1), 3(1), 4(2f), 5(2, 3b), 6(8b), 7(4, 5b),
8(2, 3f), 9(1, 2b), 10(1, 3f), 11(1), 12(2b),
13(7, 8b), 14(1). Segment V: hair 0(1), l(9b),
2(1), 3(4b), 4(1), 5(2b), 6(broken off), 7(3b),
8(1, 2f), 9(2b), 10(1), 11(1), 12(1), 13(broken
off), 14(1). Segment VI: hair 0(1), 1 (broken
off), 2(1), 3(2f), 4(4b), 5(1), 6(8b), 7(3b),
8(2f), 9(1), 10(2b), ll(3f), 12(3f), 13(20,
22d), 14(1). Segment VII: hair 0(1), l(broken
off), 2(1), 3(broken off), 4(broken off),
5(3b), 6(6-7b), 7(1, 2f), 8(6b), 9(3b), 10(bro-
ken off), ll(4b), 12(1), 13(9b), 14(1). Seg-
ment VIII: comb plate heavily sclerotized,
well pigmented, ornamented with numerous
imbrications, without visible spicules (at 430
X); comb scales 6-7, second and third from
most ventral greatly enlarged, very sharply
pointed, with very inconspicuous basal lateral
fringe as figured; hair 0(1), l(5b), 2(broken
off), 3 (broken off), 4(broken off), 5 (8b),
14(1, exceptionally long). Siphon: as figured;
length 5.5 median width; uniformly darkly
pigmented, ornamented with imbrications as

PACIFIC SCIENCE, Vol. VII, July, 1953

on comb plate; pecten extending to about
0.3, teeth 12-14 with lateral and apical fringe
as figured; valves as figured, ventral valve
exceptionally long; hair 1(8, 9b, no conspic-
uous basal expansion, barbs absent), 2(1,
short, slender), 3-5 (apparently all present as
setal rings, 3 closer to apex of valve than
usual), 6(1, well developed, longer than valve),

7(1, slender but conspicuous), 8(1, straight,
slender, about as long as valve), 9(1, weak,
straight, inconspicuous), 10-12 (not seen),
13(1, very strong, twisted at base, longer than
valve). Segment X: saddle well pigmented,
darker dorsad; ornamented with imbrications
as on comb plate; median width about 0.5 of
length; caudal margin with numerous sharply
pointed spines, proximally smaller and ap-
pearing to arise from apical imbrications; gills
about 0.5 saddle length, narrow, bluntly point-
ed at apex; hair l(6b, about 0.5 length of sad-
dle), 2(2b), 3(broken off), 4a(2b), 4b-d(bro-
ken off), 4e(l, about 0.3 of saddle length).

Types

USNM No. 61,420 (holotype, allotype, pa-
ratypes). Paratypes to be deposited in BMNH,

CU, and CSIR (Canberra); also incoll.JNB.

HOLOTYPE FLP(713-15) and allotype
MLP(713-12), Guadalcanal: West Poha
swamp, Nov. 3, 1944 (M. Cohen, L. J. Lipov-
sky et all).

PARATYPES (9M; lOF; IIP; 82L; one in-
dividual rearing), all collected on Guadalcanal
as follows: 2L(455) Sally Creek, Doma Cove,
June 14, 1944 (JNB); 2M, IF, 22L, lP(676-3)
West Poha swamp, Oct. 16, 1944 (JNB, J.
Laffoon); 5M, 5F, 4lL, 9P(708-3) West Poha
swamp, Nov. 1, 1944 (JNB); 1MLP(713-16) i
same data as holotype and allotype; 5L(816- |

3) West Poha swamp, Jan. 12, 1945 (M. Co-
hen, F. B. Wysocki); IM, IF, lL(837-3)
West Poha swamp, Jan. 20, 1945 (J. J. Cuccio,

F. B. Wysocki); IM, 91(958-2) West Poha
swamp, Apr. 28, 1945 (J. J. Cuccio, E. J. [
McCormick, V. R. Roa); 2F, lL(K-962) Dec. j
27, 1943 (K. L. Knight).

Uranotaenia sexaueri is named in honor of

Uranotaenia in Solomon Islands — Belkin

Henry F. Sexauer, who, in addition to his
duties in operating a parasitological labora-
tory, found time to contribute to the mos-
quito survey of Guadalcanal.

Variation

Among the small number of adult speci-
mens available there is no marked departure
from the holotype and allotype except for the
extremely variable light-scaling of the fore
and mid tarsi and less extensive variation in
the coloration of the pleural and lateral scutal
integument.

The variation in the chaetotaxy of imma-
ture stages is summarized in Tables 1 and 2.
The range of variation in branching of the
hairs of the larva is considerable but less than
in U. solomonis. In the pupa the branching
shows much less variation.

The six adults from Bougainville show no
striking differences from the Guadalcanal ma-
terial. The Bougainville larvae also fall within
the same range of variation.

Specimens examined: 12M; 15F; 13P; 87L.
Individual rearings: 3 larval.

Taxonomic Discussion

Uranotaenia sexaueri is closely related to U.
nivipes (Theobald, 1905) described from
Queensland. A similar form, U. albofasciata
Taylor, 1920, was described from Northern
Territory, Australia, and has been synonym-
ized with U. nivipes by Edwards (1924: 357).
Neither original description mentioned the
fact that the integument is broadly whitish
on the lower part of the scutum and upper
part of the pleuron and that this band is con-
tinued over the anterior margin of the scutum
but Edwards {loc. cit.), in comparing a para-
type of Taylor’s species with the holotype of
U. nivipes, noted the presence of this charac-
teristic ornamentation in both forms. U. sex-
aueri agrees in this character but is distin-
guished by the extent of the white-scaling of
the hind tarsus, about one half of segment 3
being white whereas only the apex is white in
U. nivipes, and the entire segment is white in

365

U. albofasciata. A further difference may be the
presence of many dark scales in the lower half
of the stp of U. sexaueri, a character not men-
tioned in the other two forms. U. albofasciata
is distinguished from U. nivipes and U. sexaueri
in having the scutellar integument light in-
stead of dark. I have seen a single female
(New Guinea, APO 565, Jul. 5, 1944, E. S.
Ross No. 40) which may represent another
species in the nivipes- group. It differs from U.
sexaueri in having the upper part of the pleural
integument not distinctly white and the lower
part of stp with a few scattered white scales
only.

The immature stages of both U. nivipes and
U. albofasciata have not been described to date.

Related to the above-mentioned Australa-
sian forms are the Oriental U. nivea Leicester,
1908, from Malaya and U. triangulata (Lud-
low, 1908) from the Philippines. Edwards
(1922: 460) synonymized these forms with
U. nivipes but later (1932: 99) considered them
both as representing a single variety of U.
nivipes. Bohart (1945: 36) recognized U. nivea
as a distinct species, retaining U. triangulata
as a synonym. Whether or not these two forms
are distinct from each other cannot be de-
termined here, but they differ from the Aus-
tralasian forms in having the scutal integu-
ment uniformly dark (without the whitish
lateral area), the pleura not distinctly whitish
dorsally, and the sternopleural scales light in
the center and dark above and below. U.
sexaueri agrees with U. nivea and U. triangulata
in the possession of numerous dark scales on
the lower part of the sternopleuron, but ap-
parently these scales are less numerous.

Thus it appears that all the above-men-
tioned forms are members of a complex, the
nivipes- group, occurring in the Oriental and
Australasian regions. U. orientalis Barraud,
1926, from Assam, may be related to this
group, but it has an entirely dark hind tarsus,
and the head scaling is darker. U. sexaueri
shows similarities to both the Oriental and
the Australian forms but appears distinct from
any of the species previously described.

366

PACIFIC SCIENCE, Vol. VII, July, 1953

No attempts have been made in the past
to determine the relationship of the nivipes-
group to other species groups. The ornamen-
tation of the adults is so characteristic and
distinctive that it isolates the group. It is in-
teresting to note that the development of
broad scales on ppn is restricted to just two
species outside of this group, U. nivipleura
Leicester, 1908, and U. hehes Barraud, 1931,
whereas U. unguiculata Edwards, 1913, has
lanceolate scales. These three species appear
entirely unrelated and distinct from the nivipes-
group. The male genitalia of the nivipes-
group have the mesosome similar to- that of
all the other ornamented species found in the
Solomons except U. solomonis and the ninth
tergite much like that of U, wysockii, the
tibialis- gr^on-p, and U. atra. The larva of U.
sexaueri, on the other hand, is most closely
related to that of U. civinskii.

Biolo^

The immature stages of U. sexaueri have
been collected on Guadalcanal only in dense
jungle swamps in association with U. harnesi
and U. solomonis. This species appears to be
rare in the Solomons.

Living larvae are easily confused with U.
solomonis but with some care can be distin-
guished by the lighter pigmentation of the
head capsule and antenna and the consider-
ably longer siphon. The pupae are quite
distinctive as they have a moderately long
trumpet (index about 3.5) that is dark in the
basal half and golden brown apically.

We had little success rearing this species in
the laboratory, although U. harnesi and U'.
solomonis larvae collected at the same time came
through readily.

The adults of this species were not collected
on Guadalcanal, and no information is avail-
able on their habits.

Distribution

Solomon Islands, Guadalcanal: North and
northwest coast (JNB et aL; K. L. Knight)
[USNM, JNB]. Bougainville: Empress Augus-

ta Bay, 2M, 3F(G-103A); IF, 3L(G-388) (A.
B. Gurney) [USNM].

6. IJranotaenia wysockii Belkin n. sp.

Plates 11, 12

Diagnosis

ADULT. — Head dark-scaled centrally; mod-
erate white orbital line; a few dark erect ver-
tical scales; conspicuous white frontal tuft.
Scutal integument entirely dark; narrow line
of white scales on lower margin from wing
root to wing root around front; apn and stp
each with narrow line of white scales forming
streak in line with white-scaling of head. Tarsi
light on all legs from basal third of segment
3, pure white on III, cream-colored on I and
II; conspicuous white knee spots on all fem-
ora. Wing with conspicuous black and
white pattern. Abdominal tergites 1-4 largely
white-scaled, 5 white-scaled on apical third,
6 and 7 with narrow median apical transverse
white bands. Male legs without conspicuous
modifications.

PUPA. — Trumpet length 6.5 miedian width;
tracheoid extending to 0.5; uniformly lightly
pigmented; no slit in meatus. Hairs 4, 5-C
usually 3, 4b; 3-11 2, 3b, close to 2-II; l-III
usually 3b; 4-IV-VI usually 3b, extending to
middle of second segment following; 6-I-V
usually single (1-2). Paddle almost as wide
as long, hair absent.

LARVA.— Head approximately as wide as
long; hairs weakly pigmented, no spikes; 1-A
placed near middle, 3b, almost as long as
antenna; 14-C dendritic. Thorax and abdomen
with slender dorsal and ventral stellate hairs,
usually with four to six branches of uneven
lengths; some very elongate strong single
hairs on thorax. Thoracic hairs 9-M, T, 8-M,
and 7-T multiple, barbed; 4-P 2b; 7-P 2-4b;
9-P 2, 3b, uneven; 14-P single. Abdominal
hairs 6-III-VIII stellate, much shorter than
6-I-II, without tubercles; l-I, II usually 4b,
uneven; 6-1, II usually 2b, uneven; 6-III, IV
usually 4-5b. Comb plates separate; scales
fringed. Siphon index about 3.0; pecten ex-

Vramtaenia in Solomon Islands — Belkin

367

Fig. 11. Vramtaenia wysockti Belkin n. sp. a-d. Adult; e,f, pupa, a. Detail of metathoracic leg of male; b, ninth
tergite of male; c, male genitalia; d, left lateral aspect of head and thorax of female; e, right lateral aspect of anterior
portion of cephalothorax of pupa; /, metanotum and abdomen of male pupa, left ventral, right dorsal. Abbre-
viations as given on page 314.

tending to 0.8; pecten teeth apically and later-
ally fringed; hair 1-S longer than siphon,
usually 4b; hair 9-S hook-like; hair 13-S weak,
slightly twisted at base. Anal segment mod-
erate; saddle margin with long apical spines;
gills 1.6 of saddle length; hairs 1-X 2b; 2-X
3-5b; 3-X 2, 3b; 4a-X 2b.

Description

FEMALE (932-103).— Wing: 1.70 mm. Ab-
domen: 1.20 mm. Proboscis: 1.10 mm. Front
femur: 1.37 mm.

Head: Conspicuous frontal tuft of very long,
narrow, white scales, with shorter scales at
base and a few broad dark scales; orbital line
of white scales moderately broad, angled lat-
erally to apn and slightly enlarged, scales

broad and dense; decumbent dark scales iri-
descent bronzy; one pair of erect, slender,
apically-forked, dark vertical scales; three pairs
of slightly shorter similar erect occipital scales;,
inner occipital bristle absent, outer two out-
side of occipital light line. Clypeus dark
brown. Palpus about 0.05 of proboscis; with
small dark scales and numerous long hairs.
Proboscis slightly swollen apically; dark-
scaled, lighter ventrally; hairs short and in-
conspicuous on shaft, more numerous apical-
ly; labella lighter, moderately hairy. Antenna
about 1.2 of proboscis; torus brown, without
hairs; first flagellar segment creamy, remain-
ing segments dark brown; flagellar whorls
with six bristles; three apical segments sub-
equal.

368

PACIFIC SCIENCE, Vol. VII, July, 1953

Thorax: Scutal integument rather uniformly
brown; moderately dense vestiture of recum-
bent narrow elongate golden-brown scales,
becoming broader and denser in prescutellar
space; line of white scales extending from
wing root to wing root around the lower mar-
gin in front as figured, integument brown;
white scales in patch on anterior acrostichal
area small and narrow; white scales in lateral
portion of line small and narrow at anterior
end becoming longer and broader posteriorly,
not forming a distinct tuft in front of wing
root; acrostichal bristles moderate, dorsocen-
trals very strong, others strong, all dark. Scu-
tellum dark brown; median lobe moderate,
with two strong bristles, scales small, golden
brown, projecting slightly over base of bris-
tles; lateral lobe with two strong bristles,
scales elongate, projecting strongly over base
of bristles. Pleural integument uniformly dark
brown except for light iridescent areas in line
with white-scaling; scaling and chaetotaxy as
figured; scales white; median patches on apn
and stp in line with white-scaling of head;
scales of apn broad anteriorly, more elongate
posteriorly and forming a small projecting
tuft; scales of transverse patch of stp in two
or three rows, individual scales oval, decum-
bent; scales of lower sternopleuron scattered,
translucent dark, broadly oval; bristles dark,
propleurals strong, spiracular and upper me-
sepimerals absent. Haltere light on base and
stem, knob cream-scaled basally, dark-scaled
on extreme apex.

Wing: Distance between cross veins about 1.3
of m-cu. Vein R2 about 0.33 of R2+3; vein
Mi+2 about 0.6 of M beyond m-cu. Conspicu-
ous black and white pattern of scaling on
dorsal and ventral surfaces: anterior border
dark, median and posterior part white, with
indistinct gray subapical transverse band, apex
of wing white beyond base of anterior fork
cell (R2) ; vein C dark except on caudal margin
at base and at apex beyond level of cell R2;
vein Sc dark-scaled ventrally; vein R white
at base to 0.24 of wing length; veins Rs and
R2-f3 dark-scaled below and above; vein R4-1-5

dark-scaled above at extreme base, then for
one third its length starting at 0.17, white-
scaled for remainder of dorsal surface and
entire ventral surface; vein M dark-scaled ven-
trally from m-cu distad; veins M1+2 and M3
dark-scaled at base dorsally; vein Cui dark-
scaled for its apical 0.65, Cu2 for its entire
length; remaining portions of all veins white-
scaled. Fringe white, creamy at apex.

Legs: Coxae and trochanters brown to light
brown; scales translucent except proximally
on coxae I and II where they are white. Fem-
ora dark above, cream-colored below; prom-
inent dorsal knee spots of pure white scales.
Tibiae dark-scaled, cream-colored apically.
First two segments of tarsi dark-scaled, lighter
below; basal third of segment 3 dark above;
remainder of segment 3 and all of segments
4 and 5 white-scaled, pure white on tarsus
III, cream-white on I and II. Leg I: femur 1;
tibia 1.0; tarsus 0.80, 0.45, 0.30, 0.15, 0.10.
Leg II: femur I.06; tibia 1.40; tarsus 0.88,
0.42, 0.30, 0.12, 0.05. Leg III: femur 1.10;
tibia 1.20; tarsus 0.98, 0.60, 0.47, 0.25, 0.06.
Abdomen: Tergite 1 white-scaled except for
cream-colored patch laterally; tergites 2-4
with large median patch of white scales ex-
tending from base to apex, wider apically;
tergite 5 with white-scaled patch occupying
apical two thirds; tergites 6 and 7 with median
apical transverse band of white scales; re-
mainder dark-scaled, Sternites dark-scaled.

MALE (932-102). —Wing: 1.54mm. Probos-
cis: 1.05 mm. Front femur: 1.33 mm.

Generally very similar to female. Proboscis
more distinctly swollen apically. Orbital line
of white scales much narrower than in female.
Flagellar whorls about 0.4 of flagellum, with
about 26 bristles; penultimate segment about
1.65 of preceding, apical about 1.10 of penul-
timate. Leg I: femur 1; tibia 1.0; tarsus 0.75,
0.45, 0.27, 0.13, 0.07; claws equal, one broad-
ened, other slender. Leg II: femur 1.05; tibia
1.41; tarsus 0.88, 0,46, 0.26, 0.12, 0.08, seg-
ment 4 with median ventral lobe projecting
below 5, segment 5 with dorsoapical lateral
lobes; claws enlarged, smaller claw 0.65 of

Uranotaenia in Solomon Islands — Belkin

369

Fig. 12. Uranotaenia wysockii Belkin n. sp. Fourth instar larva, a. Head, left dorsal, right ventral; b, left lateral
aspect of distal abdominal segments; c, dorsal aspect of left antenna; d, thorax and proximal abdominal segments,
left ventral, right dorsal. Abbreviations as given on page 314.

larger. Leg III: femur 1.05; tibia 1.20; tarsus
0.98, 0.62, 0.50, 0.26, 0.10; claws as on foreleg
and as figured. All leg segments paler than
in female.

MALE GENITALIA (932-102) .—Ninth tergite
long; proximal part shallowly emarginate;
apex shallowly emarginate; lateral lobe mod-
erately projecting, 'with ventrolateral sclero-

tization. Proctiger with slight lateral sclero-
tizations. Basal lobe of sidepiece with a dorsal
group of one large and four smaller bristles
and several minute hairs; a ventral group of
several small bristles. Clasper as figured. Me-
sosome with narrow basal dorsal sclerotized
bridge only; apical tooth more slender than
in diagram; median tooth long, directed ba-

370

PACIFIC SCIENCE, VoL VII, July, 1953

sad, only slightly curved mesally. Paramere
slightly expanded near base.

PUPA (Exuviae of holotype, 932-103).—
Abdomen: 1.70 mm. Trumpet: 0.25 mm.
Paddle: 0.32 mm.

Cephalothorax: Uniformly lightly pigmented
except for darker mesonotum and base of
wing pads. Trumpet lightly and uniformly
pigmented; length 6.50 median width; gradu-
ally widened beyond middle to apex; inner
wall separated from base to apex; tracheoid
extending about 0.5; reticulations indistinct;
pinna about 0.2; no slit in meatus. All hairs
lightly pigmented and simple; relative posi-
tion, length, and degree of development as,
figured; base of branched hairs, except 8-C,
without flattened, expanded plate. Hairs:
l(4b), 2(4, 6b), 3(5, 7b), 4(4, 5b), 5(4b), 6(1),
7(2, 3f), 8(9b), 9(2, 3f), 10(2, 3b), ll(4f),
12(2f).

Abdomen: Basal segments unevenly pigment-
ed, with submedian pale areas, connected on
I ; posterior segments uniformly more lightly
pigmented; tergites II-VII with small pos-
teromedian patch of indistinct minute spic-
ules; tergite VIII with anteromedian patch of
imbricate lines of minute spicules; tergal reti-
culations not visible; sternites III-VIII with
extensive median patch of extremely minute
and indistinct spicules. All hairs lightly pig-
mented; relative position, length, and degree
of development as figured; bases of branched
hairs, except l-I, without flattened, expanded
plate. Segment I: hair 1(21, 23 primary branch-
es, moderate number of thin apical secondary
branches; about 0.75 length of tergite), 2(1),
3(3, 4b), 4(3, 5b), 5(3b), 6(1, 2b), 7(1, 3f),
10(2, 3b). Segment II: hair 0(1), l(5b), 2(2bf),
3(2f), 4(1), 5(2b), 6(1), 7(1), 10(3b). Segment
III: hair 0(1), l(3b), 2(1), 3(3b), 4(3b),
5(3b), 6(1), 7(1), 8(2b), 10(2f), 12(2f), 13(1),
l4(not seen). Segment IV: hair'O(l), 1(3, 4b),
2(1), 3 (3b, twice length shown on figure),
4(3b), 5(2f), 6(1), 7(1), 8(3, 4b), 10(3b),
12 (2f), 13(1), l4(not seen). Segment V: hair
0(1), l(3b), 2(1), 3(3, 4f), 4(3b), 5(2, 3bf),
6(2f), 7(1), 8(1, 2f), 10(2b), 12(2f), 13(1), 14

(not seen). Segment VI: hair 0(1), l(3b), 2(1),
3(3, 4f), 4(3b), 5(2, 3f), 6(2b), 7(1), 8(1, 2f),
10(2f), 12(2f), 13(1), l4(not seen). Segment
VII: hair 0(1), 1(3, 4b), 2(1), 3(5, 6b), 4(3b),
5(3b), 6(3b), 7(0, 3b), 8(1, 2b), 10(2f), 12(3b),
13(1), 14(0, 1). Segment VIII: caudal margin
of sternite strongly and evenly emarginate;
hair 0(1), 4(3bf), 7(5, 7b), 14(1). Segment
IX: hair 1(1, very thin, slightly longer than
segment). Paddle as figured; very lightly pig-
mented; midrib strongly sclerotized and dark-
ly pigmented; external buttress poorly defined;
external margin with minute teeth; apex of
inner margin with a few lighter teeth; hair 1
absent. Genital lobe extending to about 0.27
of paddle; patch of strong spicules ventrally.
Anal segment with dorsolateral expansion, no
indication of cereal sclerite. Male genital lobe
(exuviae of allotype, 932-102) extending to
0.4 of paddle, pair of small ventral apico-
lateral patches of weak spicules; anal segment
extending 0.5 from apex of IX to apex of
genital lobe, 0.5 width of latter.

LARVA (Fourth instar exuviae of holotype,
932-103).— Head: 0.60 mm. Siphon: 0.36
mm. Anal saddle: 0.26 mm.

Head: Approximately as long as wide; ocular
bulge near middle, not well defined; very
lightly pigmented, darker dorsocaudad on
midline; integumentary imbricate sculpturing
distinct caudally only. Labrum long, 0.35 of
width at 1-C, anterior margin very shallowly
emarginate. Mental plate with 17 blunt teeth,
conspicuously defined basally on plate. Hairs
of head capsule all very lightly pigmented,
inconspicuous; slender except for 6-C which
is somewhat flattened basally; barbs absent
except minute ones on base of 6-C; relative i
position, length, and degree of development |
as figured. Hair 0(broken off), 1(1, curved, |
sharply pointed), 3(1, simple, thin, flattened |
in basal half, sharply pointed, reaching to base ||
of 1-C), 4(4, 5b), 5(2, 3b), 6(1), 7(4, 5b), ||

8(2f), 9(3, 4b), 10(2f), 11(3, 4b), 12(2f), 13(3,

4b), l4(small flattened tuft of 12, 14 branch- j
es), 15 (3b). Antenna about 0.2 of head, slight- |
ly curved, almost parallel-sided, with slight |

Uranotaenia in Solomon Islands — Belkin

371

swelling near base; width approximately 0.2
of length; very lightly and uniformly pig-
mented; shaft smooth, without spicules. An-
tennal hairs all lightly pigmented and incon--
spicuous; relative position, length, and degree
of development as figured; all single except
1-A which is 3-branched and almost as long
as antenna.

Thorax: All hairs and tubercles very lightly
pigmented, slightly darker than head inte-
gument and hairs; relative position, length,
and degree of development of hairs as fig-
ured; apices of long hairs attenuated and very
elongate; barbs when present very slender and
sparse; hair 5-P about 2.0 of head length.
Prothorax: hair 0(3, 4b, all primary), 1 (broken
off), 2(1), 3(13, 14b, with broad fan-like
asymmetrical base), 4(2b), 5(1), 6(1), 7(2, 3b,
third branch smaller), 8(4, 5b), 9(2b, one
branch very small), 10(1), ll(2f), 12(3, 4f),
14(1, 2f, slightly thickened, with minute
barbs). Mesothorax: hair l(4b), 2(2, 3b), 3(1,
2f), 4(4f), 5(broken off), 6(broken off), 7(1),
8(4b), 9(5b), 10(1), 11(1), 12(1), 13(moder-
ately dendritic tuft of 15-17 branches), 14
(moderately dendritic tuft of 18-19 branches).
Metathorax: hair l(3b), 2(2f), 3(3b), 4(2f),
5 (2b), 6(2f), 7(6b), 8(8, 9b, all primary),
9(5b), 10(1), 11(1), 12(1), 13(5b).

Abdomen: Hairs and tubercles of segments I,
II lightly pigmented as on thorax; hairs of
following segments considerably darker, par-
ticularly stellate tufts; relative position, length,
and degree of development as figured; barbs
when present very slender, short and incon-
spicuous. Stellate hairs (1, 6, 13) with branch-
es of two lengths; 4-II a forked hair, not
stellate. Segment I: hair l(3b), 2(1), 3(1, 2f),
4(7, 9b), 5(2, 3b), 6(2b), 7(1), 8(2b), 9(5, 6b),
10(4b), 12(1), 13(2, 3f). Segment II: hair 0(1),
l(3b), 2(1), 3(3, 4b), 4(2, 3f), 5(3b), 6(bro-
ken off), 7(1), 8(1), 9(1), 10(2f), ll(2f),
12(2f), 13(4b). Segment III: hair 0(1), l(3b),
2(1), 3(3f), 4(2f), 5(2b), 6(3b), 7(6, 7b),
8(2b), 9(1, 2f)., 10(2f), 11(2, 3f), 12(2b), 13(3,
4b), 14(1). Segment IV: hair 0(1), 1(2, 3b),
2(1), 3(1), 4(2, 3f), 5(3, 4b), 6(4b), 7(6b),

8(2f), 9(1, 2b), 10(2, 3f), 11(1, 2b), 12(1, 2b),
13(3, 4b), 14(1). Segment V: hair 0(1), l(3b),
2(1, 2b), 3(3b), 4(2f), 5(2, 3f), 6(4, 5b), 7(4,
5b), 8(2b), 9(2f), 10(2f), 11(1), 12(2f),
13(4b), 14(1). Segment VI: hair 0(1), l(2b),
2(1, 2b), 3(2, 3f), 4(2, 3f), 5(3, 4b), 6(4b),
7(5b), 8(2b), 9(2b), 10(2, 3b), ll(2f), 12(2f),
13(15, I6d), 14(1). Segment VII: hair 0(1),
1(2, 3b), 2(1), 3(3, 4b), 4(1), 5(1), 6(4b),
7(2f), 8(4b), 9(4b), 10(1, 2f), ll(2b), 12(1),
13(4b), 14(1). Segment VIII: comb plate
heavily sclerotized, moderately pigmented,
ornamented with numerous lines of minute
'Spicules; comb scales 8-8, uniform in size,
with inconspicuous fine lateral fringes as fig-
ured; hair 0(1), l(2f), 2(2, 3f), 3(4, 6b, con-
spicuously barbed), 4(2f), 5 (4b, conspicuous-
ly barbed), l4(2f). Siphon: as figured; length
3.0 median width; moderately pigmented, or-
namented with numerous lines of minute spic-
ules; pecten extending to 0.8, teeth 21-20
with lateral and apical fringe as figured; valves
as figured; hair l(4b, two long, two short,
lightly barbed, slightly longer than siphon),
2(1, slender), 3, 4(minute), 6(1, longer than
valve), 7 (not seen), 8(1, weaker than 6, as
long as valve), 9(strong, hook-like), 10-12
(not seen), 13(1, weak, twisted bristle). Seg-
ment X: saddle uniformly and lightly pig-
mented, ornamented with numerous lines of
minute spicules; median width 0.65 of length;
caudal margin with several rows of long,
apically fringed spines as in figure; gills 1.6
length of saddle, narrow, blunt apically; hair
l(2b, about 1.2 length of saddle, uneven),
2 (3b, about 4.0 length of saddle), 3 (broken
off), 4a(2b, about 4.2 length of saddle), 4b
(2b, about 4.0 length of saddle), 4c(l, about
3.8 length of saddle), 4d(l, about 3.8 length
of saddle), 4e(l, about 2.8 length of saddle).
Types

USNM No. 61,421 (holotype, allotype, pa-
ratypes) . Paratypes to be deposited in BMNH,
CU, and CSIR (Canberra); also in coll. JNB.

HOLOTYPE FLP(932-103) and allotype
MLP(932-102), Guadalcanal: Tenaru, Mar.
28, 1945 (JNB, M. Cohen, E. Winkler).

372

PACIFIC SCIENCE, VoL VII, July, 1953

PARATYPES (IM; 7F; 7P; llL; 6 individual
rearings) all collected on Guadalcanal, as fol-
lows: 3L(920-4) Tenaru, Mar. 15, 1945 (JNB,
M. Cohen); 1FP(932-101) same data as ho-
lotype and allotype; IMLP (941-32), 4FLP
(941-31, 33, 34, 35) Kokumbona trail, 8 miles
from north coast, April 8, 1945 (JNB); IF,
lP(0-38) Tenaru, Sept. 28, 1943 (P. W.
Oman); IF, 3L, Tenaru, Oct. 1944 (J. G.
Franclemont) .

Uranotaenia wysockii is named in honor of
F. B. Wysocki in recognition of the many
valuable collections he made on Guadalcanal.

Variation

The small series of adults from Guadal-
canal, although reared from larvae collected
in two widely separated localities in different
plants, show no striking variations. The white-
scaling of the abdomen is quite variable, and
there is often a small basal median triangular
spot on segments 2 to 4 encroaching upon
the white markings. The head, thorax, and
legs show very little variation in coloration.

The variation in the chaetotaxy of immature
stages is shown in Tables 1 and 2. Again
there are no constant differences correlated
with the different origin of the specimens.
Individuals vary considerably in the branch-
ing of the hairs but within the same range as
the other species of Uranotaenia. The follow-
ing larval hairs are unusually variable in
branching: 4-C, 1-A, 0-P, 3-VIII, 2-X. There
is more variation in the number of pecten
teeth than in any other species studied. The
pupal stage is about as uniform as in the
other species.

The material from Bougainville falls within
the range of variation exhibited by the speci-
mens from Guadalcanal without any special
features or concentration at one or other ex-
treme.

Specimens examined: 9M; 25F; 9P; 22L.
Individual rearings: 7 larval, 1 pupal.

Taxonomic Discussion

Uranotaenia wysockii is a very distinct and

isolated species in the alboannulata-gsou^.
This group has not been reported previously
from the Australasian region. The Oriental
species, U. alboannulata (Theobald, 1905)
from South India, U. rutherfordi Edwards,
1922, from Ceylon, and U. trilineata Leicester,
I9O8, from Malaya are highly ornamented
with light spots, lines, or bands on the pro-
boscis (not in U. rutherfordi) and on the fem-
ora, tibiae, and hind tarsus. U. wysockii
agrees with these species in having pictured
wings, a narrow light-scaled line from wing
root around the front margin of the scutum,
and the narrow light-scaled line across stp and
apn connecting with the narrow orbital light-
scaled line on the head. It differs from the
Oriental species in the following combination
of characters: proboscis dark-scaled; all fem-
ora with an apical white knee spot only;
all tibiae dark; hind tarsus white-scaled from
basal third of segment 3; fore and mid tarsi
cream-colored from basal third of segment 3.

The immature stages of the Oriental species
are not known. It is likely that they will be
found in water collections in living plants.

It is possible that U. nivipleura Leicester,
I9O8, from Malaya is related to this group as
it has the characteristic mesonotal scale bor-
der and has been bred from pitcher plants.

It is distinct from the alboannulata- gx.o\x^ in
having dark wings and dark legs. The record
of U. nivipleura by Barraud (1934: 76-77)
from the Western Himalayas does not seem
correct to me, nor Bohart and Ingram’s (1946:
57) record from Okinawa.

It is quite likely that members of the ^Z-
boannulata-gsovc^ have been collected in the
Australasian region and in the Philippines but
have been misidentified as members of the
nivipes- gsoM^. Both groups have the wing
patterned in black and white, and the de-
scriptions of the species of the nivipes- group
are very misleading as to the nature of the
light area around the lateral margin and front '
of the scutum.

Uranotaenia in Solomon Islands — Belkin

373

Biology

Uranotaenia wysockii is known from reared
specimens only. On Guadalcanal larvae were
collected in the water-holding leaf axils of a
short heavy-stemmed, prostrate, stoloniferous
pandanus with broad, almost smooth-mar-
gined leaves {Freycinetia sp..^) growing in a
densely shaded nipa-palm swamp (coll. 920,
932), some 3 miles from the coast. Associated
with it were an undescribed Culex {Lopho-
ceraomyia), Aedes sp. (/^oc/6/- group), and an
undescribed Corethrella. The other collection
came from an elevation of over 3,000 feet,
about 8 miles from the coast, and was made
in leaf axils of a large, erect pandanus with
broad, smooth-margined leaves growing in the
open. Associated with U. wysockii in this hab-
itat were two species of the Aedes kochi-gtou.p
and Culex {Lophoceraomyia) sp.

Superficially, the larvae of U. wysockii bear
little resemblance to those of ground-water
species. The broad head is much like that of
other inhabitants of water collections in plants,
and the simple hairs on the thorax and anterior
portion of the abdomen, as well as those on
the caudal segments, are greatly elongate. The
short, characteristically shaped siphon with
its very long tuft is also very different from
typical Uranotaenia. The presence of a comb
plate and a long siphonal tuft will separate
the larva from all plant breeders on Guadal-
canal. The larvae spend most of the time feed-
ing at the bottom, unlike ground pool breed-
ers. The pupae are of the typical Uranotaenia
type and like the larvae are very lightly pig-
mented. The short, slender, uniformly pig-
mented and widely spaced trumpets are very
characteristic and separate this species from
all other mosquitoes found in similar habitats.

On New Georgia and Bougainville, U. wy-
sockii has been reported as breeding in the leaf
axils of lily-like and aroid plants. I suspect
that the plants were smooth-leaved pandanus
{Pandanus sp. or Freycinetia sp.) as on Guadal-
canal.

Distribution

Solomon Islands, Guadalcanal: Tenaru
and Kokumbona trail (JNB, J. G. Francle-
mont, P. W. Oman) [USNM, JNB]. Bougain-
ville: Empress Augusta Bay, IM, 4F, Mar. 4,
1944 (C. R. Bruck); 1M(G-104A) July 1,
1944; lF(G-270) Mar. 6, 1944; lF(G-34l);
1F(G-344) Apr. 24, 1944; lL(G-356-2), IF
(G-357) Apr. 27, 1944; 6L(G-364) May 3,
1944; lL(G-383) May 17, 1944; 5M, 5F(G-
421) June 27, 1944; 4F(G-431) July 1, 1944
(A. B. Gurney) [USNM, JNB].

7. Uranotaenia quadrimaculata Edwards
in Paine and Edwards, 1929
Plates 13, 14

1929. Uranotaenia quadrimaculata Edwards in

Paine and Edwards, Bui. Ent. Res. 20:
315-6. Types: M, E, L, P; Guadalcanal:
Rere, 19 Aug. 1928; llu, 26 Aug. 1928.
Malaupaina (near San Cristobal) : 4
Sept. 1928 (R. W. Paine) [BMNH
(Paine and Edwards, 1929)]-
1926. Uranotaenia nigerrima Taylor. Edwards,
Bui. Ent. Res. 17: 109 (misident.).
1944. Uranotaenia nigerrima Taylor. Knight,
Bohart, and Bohart, Keys Mosq. Aus-
tralasian Reg. p. 15, 68 {partim, misi-
dent.).

1944. Uranotaenia quadrimaculata Edwards.
Knight, Bohart, and Bohart, Keys
Mosq. Australasian Reg. p. 15, 69
{partim, misident.).

Diagnosis

ADULT. — Head densely covered with long,
dark, erect, vertical scales; decumbent scales
very light purplish-brown. A large velvety
black spot on each side of scutum in front of
wing root and another on ppn, contrasting
sharply with rest of integument. Scutal vesti-
ture of elongate curved decumbent purplish-
brown scales; lighter scales in streak above
and behind dark scutal spot; a few curved,
light scales on scutum above anterior end of
ppn. Pleura light; broad translucent light

374

PACIFIC SCIENCE, VoL VII, July, 1953

scales on apn only. Legs entirely dark. Wings
completely dark-scaled. Abdominal tergites
entirely dark purplish-brown. Male legs with-
out conspicuous specializations.

PUPA.— Trumpet length 3.0 or less of me-
dian width; tracheoid restricted to basal 0.25
or less; uniformly brilliant brown; no slit in
meatus. Cephalothoracic hairs, except 8-C,
single or 2-3b; abdominal hairs, except l-I,
single or 2-3b(rarely 4, 5b); hair 8-II present.
Paddle with distinct serrations on external
and internal margins; hairs 1 and 2 both pres-
ent, strong.

LARVA. — Head longer than wide; hairs
weakly pigmented, no spikes; 1-A placed at
0.7-0. 8 of antenna. Thorax and abdomen
without dorsal or ventral stellate hairs, most
hairs single or 2-4b. Thoracic hairs 9-P, M
single; 8-M, 7-T short, multiple, not barbed.
Abdominal hairs 6-I-IV of approximately
equal size, all with tubercles. Comb plates
united dorsally into saddle; scales not fringed.
Siphon index 2.0 or less; pecten teeth not
fringed but sometimes with basal denticles;
hair 1 very short, inconspicuous; hairs 7, 8-S
usually 2b; hair 13-S simple. Anal segment
very short; gills 3.0 length of saddle; saddle
margin without projecting apical spines; hairs
1-X l-3b; 2, 3-X 1, 2b; 4a-d-X single.

Description

FEMALE (523-23).— Wing: 2.66 mm. Ab-
domen: 1.66 mm. Proboscis: 1.50 mm. Eront
femur: 1.83 mm.

Head: Numerous elongate dark erect vertical
scales, stem very slender, apex expanded, 3-
or 4-pronged; no frontal tuft; recumbent
scales iridescent very light purplish-brown,
lighter laterally toward apn; orbital bristles
rather evenly spaced. Clypeus light brown.
Palpus about 0.09 of proboscis; purplish-
brown scales apically; hairs about 0.5 of shaft
length. Proboscis distinctly swollen apically;
dark-scaled dorsally, lighter ventrally, espe-
cially on swollen part; numerous short hairs
apically; labella light, densely hairy. Antenna
about 1.33 of proboscis; torus light brown.

darker mesally and dorsally, with short hairs
mesally; base of first flagellar segment cream-
colored, remainder of flagellum very dark
brown; first flagellar segment 1.2 of second;
three apical segments subequal.

Thorax: Conspicuous velvety black spots on
posterior pronotum and on scutum in front
of wing root as figured. Scutal integument
uniformly light brown; dense vestiture of
recumbent narrow purplish-brown scales cen-
trally; tuft of erect lighter narrow scales at
extreme anterior end among dorsocentrals and
a few shorter curved recumbent light scales
just external to dorsocentrals above anterior
end of ppn; elongate patch of narrow recum-
bent scales over black spot in front of wing
root, scales lighter than on disc, becoming
whitish in caudal extension over supra-alar
bristles; acrostichals short, ail bristles dark.
Scutellum brown; median lobe moderate, with
one weak and four strong bristles; lateral lobe
with four strong and two weak bristles; vesti-
ture of broad recumbent purplish bronzy-
brown scales projecting moderately over bris-
tles of lateral lobe. Pleural integument cream-
colored, darker along diagonal line from
spiracle to lower mesepimeron and on apn,
velvety black spot on ppn\ scales absent except
broad translucent whitish scales forming large
patch on apn; large bristles dark, smaller light;
propleurals four, upper mesepimerals three or
four. Haltere light at base and lower part of
stem, dark-scaled on upper part of stem and
knob.

Wing: Distance between crossveins 0.4 of
m-cu. Vein R2 about 0.29 of R2-f3; vein M1+2
about 0.72 of M beyond m-cu. Wing scales
all dark. Eringe narrow, dark.

Legs: Coxae and trochanters cream-colored;
scales light and translucent, inconspicuous;
coxa I with about 12 bristles on anterior face;
coxa III with seven bristles on caudal face.
Femora dark-scaled above, lighter below; fe-
mur II swollen basally, cream-colored ven-
trally, with seven conspicuous thin erect hairs
dorsally in basal half and three similar hairs
on external surface of apical third. Tibiae dark

Uranotaenia in Solomon Islands — Belkin

375

Fig. 13. Uranotaenia quadrimaculata Edwards, in Paine and Edwards, 1929. a-f-, Adult; g, h, pupa, a-c, Details
of pro- and mesothoracic legs of male; d, ninth tergite of male; e, male genitalia; /, left lateral aspect of head and
thorax of female; g, right lateral aspect of anterior portion of cephalothorax of pupa; h, metanotum and abdomen
of male pupa, left ventral, right dorsal. Abbreviations as given on page 314.

above, lighter ventrally. Tarsi dark above,
lighter below, apical segments lighter, almost
creamy on II. Leg I: femur 1; tibia 1.09; tarsus
0.87, 0.40, 0.28, 0.14, 0.06. Leg II: femur 0.95;
tibia 1.22; tarsus 0.95, 0.40, 0.28, 0.13, 0.07.
Leg III: femur 0.95; tibia 1.22; tarsus 1.43,
0.73, 0.60, 0.34, 0.10.

Abdomen: Tergites entirely dark iridescent
purplish-brown; sternites entirely cream-
colored.

MALE (939-41).— Wing: 2.30 mm. Probos-
cis: 1.50 mm. Front femur: 1.75 mm.

Generally very similar to female. Proboscis
about the same as in female. Flagellar whorls
with about 24 bristles, 0.4 of flagellar length;
penultimate segment 3.0 of preceding, sub-
equal to apical. Leg I: femur 1; tibia 1.05;

tarsus 0.84, 0.37, 0.28, 0.13, 0.08; claws equal
and similar. Leg II: femur 0.93; tibia 1.16;
tarsus 0.87, 0.37, 0.26, 0.12, 0.07; one claw
enlarged, other short. Leg III: femur 0.90;
tibia 1.15; tarsus 1.40, 0.66, 0.56, 0.36, 0.11;
claws equal, similar.

MALE GENITALIA (939-41) .—Ninth tergite
narrowly and deeply emarginate at base; basal
sclerotization angled caudad before emargi-
nation and evanescent; apex truncate, not pro-
duced into lateral lobes; subapical light
sclerotization extended ventrolaterad prox-
imally as a weak bar. Proctiger with distinct
ventrolateral scoop-shaped sclerotization and
a more apical ventral sclerotization bearing
small spicules. Basal lobe of sidepiece with
one very heavy apical bristle and three rows

376

of progressively shorter bristles basad; one
strong and several short bristles ventrally.
Clasper long, slender; as figured. Mesosome
with fairly broad dorsal basal bridge and with
a very narrow complete ventral bridge; lateral
plate with two rows of ventrolateral short
teeth or serrations. Paramere very broad except
distally.

PUPA (Exuviae of female, 523-23).— Ab-
domen: 2.70 mm. Trumpet: 0.38 mm. Pad-
dle: 0.58.

Cephalothorax: Moderately pigmented; ap-
pendage cases, meso- and metanotum darker.
Trumpet brilliant brown; length 2.8 median
width; gradually widened from base to about
0.5, then almost parallel-sided; inner wall dis-
tinctly separated throughout, distinctly con-
stricted at 0.5 and then flared out, apical
portion with conspicuous elongate imbrica-
tions; tracheoid very poorly developed, re-
stricted to basal 0.25 of anterior face; reticulate
with indistinct small shallow imbrications,
without spicules; pinna 0.2; no slit in meatus.
Hairs moderately to strongly pigmented; rel-
ative position, length, and degree of develop-
ment as figured; branched hairs with simple
base, barbs absent except as noted. Hairs:
1(1, 2f), 2(2, 3b), 3(1, 2b), 4(1, 2b), 5(3b),
6(1), 7(2, 3b), 8(3, 4b), 9(1), 10(1, with con-
spicuous apical and subapical brush of barbs),
11(1, minutely barbed), 12(1), extra hair on
left side caudad of 12, probably homolog of
larval hair 4-T.

Abdomen: Strongly pigmented on proximal
segments with lighter areas around bases of
hairs; distal segments moderately pigmented;
tergites II-VIII with imbrications more dis-
tinct on proximal segments and bearing small
apical spicules which are larger toward mid-
line and form extensive patches; tergite I with
patch of small spicules on median bridge
cephalad of float hairs; sternites II-VIII with
entire surface ornamented with small spicules
arising from faint imbrications. Large hairs
very darkly pigmented, smaller moderately;
relative position, length, and degree of de-
velopment as figured; branched hairs with

PACIFIC SCIENCE, Vol. VII, July, 195 3

simple base, barbs absent except as noted.
Segment I: hair l(dichotomously branched,
with about 10 principal branches, barbs dis-
tinct), 2(1), 3(1), 4(2b),5(l), 6(1), 7(1),
10(2b). Segment II: hair 0(1), 1(2, 3f), 2(1),
3(1), 4(1), 5(3b), 6(1), 7(1), 8(1), 10(2b),
12(0). Segment III: hair 0(1), l(2b), 2(1),
3(1), 4(1), 5(1, 2f), 6(1, 2b), 7(1), 8(1), 10
(2b), ll(setal ring on right side), 12(1, 2b),
13(1), 14(1). Segment IV: hair 0(1), l(2b),
2(1), 3(2, 3b), 4(1), 5(2f), 6(2b), 7(1), 8(2f),
10(2b, f), 11(1), 12(1), 13(1) ,14(1). Segment
V: hair 0(1), 1(2, 3b), 2(1), 3(2b), 4(1), 5(2,
3b), 6(1, 2f), 7(1), 8(2f), 10(2, 3b), 11(1, on
right side only), 12 (2b), 13(1), 14(1). Segment
VI: hair 0(1), l(2b), 2(1), 3(2, 3b), 4(1), 5(2f),
6(1), 7(1), 8(2f), 10(2b), 12(2b), 13(1), 14(1). j
Segment VII: hair 0(1), 1(2, 3b), 2(1), 3(2b), i

4(1), 5(2f), 6(1), 7(1), 8(3b), 10(3b), 12(1, j

2b), 13(1), 14(1). Segment VIII: caudal mar- I
gin of sternite very shallowly emarginate; hair i
0(1), 4(1), 7(1, 2b), 14(1). Segment IX: hair I
1(1, about 0. 7-0.9 of segment length). Pad- |
die as figured; moderately pigmented; midrib |
moderately scleroti2ed, evanescent apically; i
external buttress indistinct; basal pigment bar j
absent; external margin with distinct serra-
tions from basal 0.25, strong apically; internal
margin with even stronger, more sharply
pointed serrations from midrib apex to basal
0.5; hair 1(1, very strong, about 0.25 of pad- i

die), 2(1, strong, about 0.3 of 1). Genital |

lobe extending to 0.33 of paddle, entire ven- j
tral surface with spiculate imbrications. Anal i
segment visible, shorter than genital lobe;
cereal sclerite not defined. Male genital lobe !
(exuviae of 939-41) extending to 0.35 of pad-
dle; ventral surface with faint spiculate im-
brications; anal segment distinct, short, ex-
tending 0.3 from apex of IX to apex of genital
lobe.

LARVA (Fourth instar exuviae of female,
523-23).— Head: 0.80mm. Siphon: 0.52 mm.
Anal saddle: 0.18 mm.

Head: Width 0.9 of length; ocular bulge not
distinctly defined; pigmentation a moderate
bright yellowish brown, darker brown cau-

Uranotaenia in Solomon Islands — Belkin

377

Fig. 14. Uranotaenia quadrimaculata Edwards, in Paine and Edwards, 1929. Fourth instar larva, a, Head, left
dorsal, right ventral; b, dorsal aspect of left antenna; c, left lateral aspect of distal abdominal segments; d, thorax
and proximal abdominal segments, left ventral, right dorsal. Abbreviations as given on page 314.

378

PACIFIC SCIENCE, VoL VII, July, 1953

dally and middorsally; integumentary sculp-
turing indistinct except caudally. Labrum
short, 0.25 of width at 1-C, anterior margin
moderately emarginate. Mental plate large,
acute; with 21 distinct teeth. Hairs of head
capsule weakly pigmented except for 4, 6-C,
all inconspicuous; 5, 6-C slender, not spike-
like; relative position, length, and degree of
development as figured. Hair 0(1, short, leaf-
like) ,1(1, very short, stubby, bluntly pointed) ,
3(1, very short, inconspicuous), 4(1, darkly
pigmented), 5(1), 6(2, 3b), 7(1), 8(1), 9(3b),
10(1), ll(2b), 12(1), 13(2b), 14(1, slender,
simple), 15(1). Antenna 0.25 of head; uniform
in width, curved with concavity lateral; width
at middle 0.16 of length; uniformly moderate-
ly pigmented, darker at base; shaft without
spicules. Antennal hairs moderately pigment-
ed except 1, 2-A; relative position, length, and.
degree of development as figured; all single;
1-A placed at 0.88 from base, length about
1.4 width of antenna.

Thorax: Long hairs and tubercles strongly
pigmented; short hairs lightly pigmented; rel-
ative position, length, and degree of devel-
opment as figured; apices of long hairs
attenuated; long hairs with inconspicuous
barbs or spicules; hair 5-P about 0.6 of head
length. Pro thorax: hair 0(3b), 1(1), 2(1),
3(2b), 4(3b), 5(1), 6(1), 7(2, 4b), 8(3, 4b),
9(1, 2b), 10-12(1), 14(1). Mesothorax: hair
l(3b), 2, 3(1), 4(3b), 5-7(1), 8(3b), 9-12(1),
13(3b), I4(4b). Metathorax: hair l(2b), 2(3b),
3(2b), 4(1), 5(1), 6(1, 2f), 7(3, 5b), 8(3b),
9-12(1), 13(4, 5b).

Abdomen: Tubercles and long hairs very dark-
ly pigmented, other hairs moderately to light-
ly pigmented; relative position, length, and
degree of development as figured; long hairs
with moderate barbs. No true stellate hairs.
Hairs 6-I-V with basal tubercles. Segment I:
hair l(2b), 2(1), 3(3b), 4(2, 3b), 5(1), 6(2b),
7(1), 9(1), 10(2, 3b), 11(3, 4b), 12, 13(1).
Segment II: hair 0(1), l(3b), 2(1), 3(1, 2b),
4(1), 5(2b), 6(2b), 7(1), 8(1), 9(2b), 10(1),
11(2, 3f), 12(2b), 13(4b). Segment III: hair
0(1), 1(2, 3b), 2, 3(1), 4(1, 2b), 5(2b), 6(3b),

7(3b), 8(1), 9(3b), 10-12(1), 13(3b), 14(1).
Segment IV: hair 0(1), 1(2, 3b), 2-4(1), 5 (2b),
6(3b), 7(3, 4b), 8(1), 9(3b), 10-12(1), 13(3b),
14(1). Segment V: hair 0(1), l(2b), 2(1),
3(2b), 4(1, 2b), 5(1, 2b), 6(3, 6b), 7(3b), 8(1),
9(2b), 10-12(1), 13(3b), 14(1). Segment VI:
hair 0(1), l(2b), 2(1), 3(2b), 4(1), 5(2b),
6(3b), 7(not seen), 8(1), 9(3b), 10(2b), 11,
12(1), 13(3b), 14(1), Segment VII: hair 0(1),
l(2b), 2(1), 3(3b), 4, 5(1), remaining hairs not
visible. Segment VIII: comb plates united
dorsally, strongly sclerotized, darkly pigment-
ed, ornamented with strong imbrications from
which minute spicules project; comb scales
6-6, sharply pointed, without lateral fringe
but a few basal spicules as figured; hair 0(1),
l(2b), 2(1), 3(1, 2b), 4(2b), 5(2f, 4b), 14(1).
Siphon: as figured; length slightly less than
2.0 median width; strongly pigmented, im-
brications much heavier than on comb plate,
spicules not distinct; pecten extending to
0.70, teeth 6-7, sharply pointed, heavily
sclerotized and pigmented, without lateral
fringe but with one or two short basal den-
ticles; valves as figured, darker than siphon,
all short; hair l(2b), 2(1, well developed),

3-5 (only 2 setal rings visible), 6(1, well de-
veloped, longer than valve), 7 (2b, minute),

8 (2b, well developed, about 0.5 of valve),

9(1, slender, hook-like, inconspicuous), 10(1,
minute), 11, 12(1, in a conspicuous alveolus),
13(1, simple hair, short). Segment X: saddle j
very short, lightly pigmented; median width j
about 2.0 of length; imbrications faint except |
caudally where two or three rows of short, ;
darkly pigmented spicules arise from their |
apices but do not project from caudal margin ^ !
of saddle; gills extremely long, sausage-
shaped, 3.0 of saddle length; hair 1(2, 3b, j
minutely spiculate, about as long as saddle), ;
2 and 3 arise from large common tubercle, |
2(1, about 3.9 of saddle length), 3(1, about j
4.5 of saddle length), 4a(l, about 3.5 of sad-
dle length), 4b(l, about 4.0 of saddle length), |l
4c(l, about 3.5 of saddle length), 4d(l, about I

2.75 of saddle length), 4e(l, about 0.5 of j

saddle length). j

Urano taenia in Solomon Islands — Belkin

379

Variation

Uranotaenia quadrimaculata adults vary a
great deal in size and to a lesser extent in
coloration. On Guadalcanal all individuals
originating from taro show a conspicuous
dark diagonal band running in line from the
dark spot on ppn across the subspiracular area,
upper middle part of stp, and lower middle
part of mesopleuron just above the lower
mesepimeral bristle. An indication of this dark
integumentary band is present in specimens
from other breeding places, but it is never
conspicuous. I cannot find any structural
characters to differentiate this ecological race
either in the adults or in the immature stages.

The variation in the chaetotaxy of immature
stages is shown in Tables 1 and 2. Although
at first glance the variation does not seem to
be great, actually it is much greater than in
any species of mosquito I have studied, par-
ticularly in the larva. Not only the branching
but also the degree of development of individ-
ual hairs varies a great deal. The position of
smaller hairs is not as constant as it is in other
species. There are cases of duplication of hairs
in the larva. In the pupa there may be two
extra pairs of hairs on the metanotum, prob-
ably representing larval hairs 3, 5-T; on the
abdomen, hair 11 is frequently present on
segments III-V, and there may be a duplica-
tion of hair 12 on these segments as well.
One pupal specimen shows larval hair 9 re-
tained on the left side of segment V. These
anomalies have proved to be useful in estab-
lishing the homologies of the larval and pupal
hairs. They may prove to be of value also
when the potentialities of the mosquito chae-
totaxy and its pattern as tools in genetics are
realized.

The material from the other islands shows
no variations that are not found on Guadal-
canal, and there appear to be no characters
on which geographical subspecies or races
can be recognized within the Solomons.

Specimens examined: 288M; 310F; 96P;
866L. Individual rearings: 15 larval.

Taxonomic Discussion

Uranotaenia quadrimaculata is very closely
related to U. nigerrima Taylor, 1914, from New
Guinea. The two species are distinguished
from all other Uranotaenia by the four dark
integumentary spots, one each on the scutum
in front of the wing root and the ppn. As
pointed out by Edwards (1929: 314), U. quad-
rimaculata differs from U. nigerrima by the
generally lighter thoracic integument, includ-
ing the four dark spots, and the absence of a
patch of broad white scales on the side of the
scutum immediately above the anterior end
of the ppn. Instead of the conspicuous broad
white scales in this area as in U. nigerrima, in
U. quadrimaculata there are a few elongate
light-brown or cream-colored scales. The other
characters mentioned by Edwards also hold
but are difficult to determine because of con-
siderable variation. Knight, Bohart, and Bo-
hart (1944: 15) confused this distinction in
their key by assigning to U. quadrimaculata
the patch of broad, whitish scales which is
characteristic of U. nigerrima. The remaining
characters were properly assigned to the re-
spective species. As a result of this confusion
of characters Knight, Bohart, and Bohart {loc.
cit.) referred specimens from the Solomons,
New Britain, and New Guinea to both species.
I have examined 23 adults of this group from
several localities on New Guinea. They are
all U. nigerrima. All the Solomons specimens
are definitely U. quadrimaculata. The two spe-
cies are allopatric, and there is no overlap
whatever.

The larva of U. quadrimaculata is extremely
close to U. nigerrima. The chief differences
appear to be in the greater development in
U. quadrimaculata of the following thoracic
hairs: 4-P 3b (2-4) instead of 1, 2b; 7-P 2-4b
instead of 1; 7-T 3, 2b(l-5) instead of 1, 2f.
U. nigerrima has the comb scales with distinct
basal spicules whereas these are never strong-
ly developed in U. quadrimaculata. U. niger-
rima also has more prominent basal spicules
on the pecten teeth and strong apicolateral
spicules on the anal saddle.

380

PACIFIC SCIENCE, Vol. VII, July, 1953

The pupa of U. quadrimaculata is very much
like that of U. nigerrima. Penn (1949: 30)
separated the two species on hair lO-II, simple
in U. quadrimaculata^ and 3,4b inf/, nigerrima.
From a study of a series of U. quadrimaculata,
this should be amended to 2b (1-3) for this
species. Additional differences for U. quad-
rimacidata are as follows: 2-C 2b, 1(1-3) in-
stead of 3b; 4-C single (1-2) instead of 2-3f;
5-C 2, 3b instead of 4b; apparently fewer
branches on abdominal hairs 1 and 3.

Two ^ other species from New Guinea are
closely related to U. quadrimaculata. U. papua
Brug, 1924, has a pupa with trumpets and
paddles conforming to the type found in U.
quadrimaculata. U. diagonalis Brug, 1934, has
a larva with the typical features of the head,
antenna, thorax, and abdomen found in U.
quadrimaculata, including the dorsal connec-
tion of the comb plates. The figures of the
male genitalia (Brug, 1934: 511) indicate the
same type of mesosome. It is interesting to
note that the characteristic dark diagonal in-
tegumentary pleural line of this species is
strongly developed in specimens of U. quad-
rmaculata bred from taro.

Uranotaenia obscura Edwards, 1915, from
Borneo and Singapore, has a very similar larva
(Edwards and Given, 1928: 338-339) and un-
doubtedly belongs to the same group. U.
colocasiae Edwards, 1928, from Eiji shows re-
semblance to this group in the larval stage
also and may be related, but in the absence
of specimens I cannot be positive. U. painei
Edwards, 1935, from Fiji according to Ed-
wards (1935: 128) is very much like U. obscura,
U. papua, and U. colocasiae in the adult stage,
but its larva is entirely different from' all these
and shows features of typical Uranotaenia.

The unornamented Uranotaenia of the Ori-
ental region form a complex group of poly-
phyletic origin as shown by the different types
of larvae. As several species are unknown in
the immature stages and the available larval
descriptions are not sufficiently detailed, and
as it appears that superficially similar larvae
are developed in similar habitats, it is im-

possible at the present to determine the rela-
tionship of the several species of this complex
that are found in the Pacific area and to place
U. quadrimaculata and related species in the
natural order.

Biology

Uranotaenia quadrimaculata utilizes for
breeding small water collections in living
plants and dead plant material as well as va-
rious types of artificial containers. It is ex-
tremely common in leaf axils of taro {Colocasia
sp. and Alocasia sp.) where it is preyed upon
by Tripteroides (Rachisoura) mathesoni Belkin,
1950. It has also been found less commonly
in tree holes in association with the usual tree
hole breeders. The greatest number of indi-
viduals is produced from breeding in coconut
shells, coconut spathes, and large leaves fallen
on the ground. A few collections were made
in fallen bamboo and in tree stumps. All sorts
of artificial containers are also very favorable
breeding grounds for this species. It has been
recorded from tin cans, steel helmets, fire
barrels, and cardboard containers. In all such
situations it is preyed upon by Tripteroides
(Rachisoura) stonei Belkin, 1950, which does
not seem to be as effective a check as T.
mathesoni in taro. I have two records of U.
quadrimaculata apparently breeding in ground
pools in a swampy jungle area and in road |
ruts. In both cases the larvae were collected i
following floods so it is very probable that j
they were washed out of their normal breeding j

places, such as coconuts or leaves. '

U, quadrimaculata appears to prefer fairly !|
fresh water in its breeding places. Only oc- j|
casionally has it been found in thick, ferment- '
ing coconut water, such as is preferred by j
Armigeres. A moderate amount of organic
matter is usually present in the majority of ''
its breeding places, except in taro leaf axils. ,

The larvae of U. quadrimaculata bear no j
resemblance whatever to the typical Urano- '
taenia and most closely resemble species of
Armigeres with which they are occasionally
associated in coconuts. They behave like Ar- I

Uranotaenia in Solomon Islands — Belkin

migeres larvae also, spending most of the time
at the bottom, coming to the surface only
rarely, and moving with the same sinuous
undulations of the body. They rest suspended
down from the surface and not parallel to it
as do ground-breeding Uramtaenia. The lar-
vae are extremely large in comparison with
the size of the adults. The extremely long
sausage-shaped gills will separate them in the
field from all forms found in similar habitats,
except for Armigeres. Grossly they can be told
from the latter by the much greater propor-
tionate size of the head and the presence of
long, lateral, abdominal hairs on the first four
segments.

The pupae of U. quadrimaculata are also
very different from those of other Uramtaenia
as the trumpets are very short and relatively
broad. The wide spacing of the trumpets will
aid in recognizing them in the field from
other species in similar habitats.

Adults are often seen in great numbers rest-
ing among coconuts on the ground and in
their other breeding places. They have not
been collected at lights.

Distribution

Solomon Islands, San Cristobal: Malau-
paina. Sept. 4, 1928 (R. W. Paine) [Paine and
Edwards, 1929]. Guadalcanal: Rere, Aug. 19,
1928; Ilu, Aug. 26, 1928 (R. W. Paine) [Paine
and Edwards, 1929]; generally distributed
throughout the year on north-central and
northwest coast (JNB et al., J. G. Francle-
mont, K. L. Knight, A. B. Gurney, H. E.
Milliron et al., P. W. Oman, Lechner, Haage)
[USNM, CU, JNB]. Russell: Banika, 1 adult,
2L, 2P, Mar. 20, 1943 (W. G. Downs)
[USNM]. New Georgia: Segi Pt., 4 adults
(B-3), 5L(B-33), 2L(B-77), 4 adults (B-130)
(C. O. Berg); Munda Pt., 21 adults, 33L,
5P(F-69); 2M, 1F(F-18); 15M, 12F(F-19) (J.
G. Franclemont) [USNM, CU, JNB]. Treas-
ury: 11 adults (P-34) May-June, 1944 (J. H.
Paullus) [USNM]. Bougainville: Empress Au-
gusta Bay, lL(W-4) Dec. 12, 1943; l6L(W-5)
Nov. 17, 1943 (Weathersby and Koch); 9

381

adults, Jan. 28, 1944 (C. R. Bruck); 3 adults,

3L, lP(G-230) Feb. 19, 1944; 6 adults (G-329)

Apr. 16, 1944; 12 adults (G-341) Apr. 23,

1944; 1 adult (G-421) Jun. 27, 1944; 63 adults

(G-433) (A. B. Gurney) [USNM].

REFERENCES

Arribalzaga, Felix Lynch. 1891. Diptero-
logia Argentina. La Plata Univ. Mus, de La
Plata, Rev. 1: 345-377; 2: 131-174.

Baisas, F. E. 1935. Notes on Philippine mos-
quitoes, 11. Uranotaenia group. Philippine
Jour. Sci. 57: 63-80.

Barraud, P. j. 1926. A revision of the culi-
cine mosquitoes of India. Part XVIII . . .
Uranotaenia and Harpagomyia, . . . Indian
Jour. Med. Res. 14: 331-350.

1934. Family CuUcidae. Tribes Megar-

hinini and Culicini. 463 pp. Taylor & Fran-
cis, London [Fauna of British India. Dip-
tera, vol. 5].

Belkin, John N. 1950. Mosquitoes of the
genus Tripteroides in the Solomon Islands.
U. S. Natl. Mus., Proc. 100: 201-274.

[1951.] A revised nomenclature for

the chaetotaxy of the mosquito larva. Amer.
Midland Nat. 44: 678-698.

1952. The homology of the chaeto-
taxy of immature mosquitoes and a revised
nomenclature for the chaetotaxy of the pu-
pa. Fnt. Soc. Wash., Proc. 54: 115-130.

Bohart, Richard M. 1945. A synopsis of the
Philippine mosquitoes. 88 pp. Navy Dept.,
Washington (Nav Med 580).

and R. L. Ingram. 1946. Mosquitoes

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110 pp. Navy Dept., Washington (Nav Med
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Bonne-Wepster, j., and S. L. Brug. 1939.
Larven van Nederlandsch-Indische culici-
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1218-1279.

Brug, S. L. 1924. Notes on Dutch-East-Indian
mosquitos. Bui. Ent. Res. 14: 433-442.

1934. Notes on Dutch East Indian

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Brunetti, E, 1914. Critical review of "gen-
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15-73.

CoQUiLLETT, D. W. 1910. The type species
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Davis, Gordon E., and C. B. Philip. 1931.
The identification of the blood meal in
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Hyg. 14: 130-141.

Dyar, Harrison G. 1928. The mosquitoes of
the Americas. 6I6 pp. Carnegie Institution,
Washington.

and R. C. Shannon. 1925. The types

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Inscitiae Menstruus 13: 66-89.

Edwards, F. W. 1913. New synonymy in
Oriental Culicidae. Bui. Ent. Res. 4: 221-
242.

1922. A synopsis of adult Oriental

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thine) mosquitoes. Part II. Indian Jour.
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1924. A synopsis of the adult mos-
quitos of the Australasian region. Bui. Ent.
Res. 14: 351-401.

1926. Mosquito notes. VI. Bui. Ent.

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1928. Mosquito notes. VII. Bui. Ent.

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1929. Descriptive notes on the mate-
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1932. Diptera Fam. Culicidae. In

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1935. Mosquito notes. XII. Bui. Ent.

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1941. Mosquitoes of the Ethiopian region.

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and D. H. C. Given. 1928. The early

stages of some Singapore mosquitos. Bui.
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Hopkins, G. H. E. 1936. Mosquitoes of the
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Howard, L. O., H. G. Dyar, and F. Knab.
1917. The mosquitoes of North and Central
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King, Willard V., and H. Hoogstraal.
1947. New species of New Guinea Urano-
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Knight, Kenneth L., R. M. Bohart, and
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Knight, Kenneth L., and R. W. Chamber-
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I9O8. Mosquito notes. No. 6. Canad. |

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1910. [Pseudouranotaenia parangen- ||

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Montschadsky, a. 1930. Die Stiegmalplat-
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Uranotaenia in Solomon Islands — Belkin

383

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384

PACIFIC SCIENCE, VoL VII, July, 1953

TABLE 1

LARVA. SETAL BRANCHING

Limits of variation in parentheses, preceded by most usual number(s) observed
in ten specimens (20 hairs) in the order of frequency.

atra

barnesi

ci vinskii

quadrimacul ata

sexaueri

solomoni s

wy sockii

Head 0, 1,

3

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

4

2,3

(2-3)

1

(1)

3

(2-4)

1

(1)

2

(2-3)

3

(3-4)

4, 3

(2-6)

5

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

1

(1)

2

(2-3)

6

1

(1)

1

(1)

1

(1)

1,2

(1-3)

1

(1)

1

(1)

1

(1)

7

3

(2-3)

2

(1-3)

4, 5

(4-6)

1

(1-2)

4

(3-5)

4

(3-5)

4

(3-5)

8

2

(1-2)

2, 3

(2-3)

3,2

(1-3)

1

(1-2)

3

(2-4)

1,2

(1-2)

2

(1-2)

9

3

(2-3)

5,6

(4-7)

2,3

(2-3)

2,3

(2-4)

4

(2-5)

3

(2-4)

4

(4-3)

10

1,2

(1-3)

3

(2-4)

1,2

(1-3)

1

(1)

2, 1

(1-4)

1, 2

(1-2)

2

(1-3)

11

2, 3

(1-3)

6, 5

(4-8)

4, 5

(3-6)

3,2

(2-4)

8,9

(8-10)

5

(4-7)

3

(2-4)

12

2

(1-3)

3,2

(2-4)

2

(1-3)

1

(1)

3,2

(2-3)

2

(2-4)

2,3

(2-4)

13

4

(3-6)

4, 3

(2-6)

5,6

(4-7)

2,3

(1-3)

5,6

(4-7)

4

(3-6)

4, 5

(3-7)

14

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

13

(12-15)

15

3

(2-3)

3

(3-5)

2,3

(2-3)

2, 1

(1-3)

3, 4

(3-5)

2

(1-3)

4, 5

(3-6)

Antenna

1

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

3,2

(2-5)

2

:-6

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

Prothorax

0

8,9

(7-11)

8-10

(5-15)

12

(10-16)

3

(2-5)

15-18(13-21)

9,8

(7-16)

5-7

(3-8)

1

1

(1)

1

(1)

1

(1)

1

(J)

1

(1)

1

(1)

1

(1)

2

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

3

5, 4

(3-5)

9,8

(7-10)

6-8

(5-10)

2

(1-2)

9,8

(7-10)

8,7

(6-10)

12-

14(10-19)

4

2

(2)

2

(2)

3

(3)

3

(2-4)

2

(2)

2

(2)

2

i(2)

5

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

6

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

1

(1)

1

(1-2)

7

1

(1)

1

(1)

2

(2)

4-2

(2-4)

2

(2-4)

2

(2-3)

2,3

(2-4)

8

4

(3-5)

6,7

(5-9)

5

(4-7)

4, 3

(3-6)

7,8

(7-9)

7

( 5- 10 )

5

(4-7)

9

1

(1)

4,5

(3-6)

2

(2-3)

1

(1-2)

4

(4-6)

2

(2-3)

2

(2-3)

10

1

(1)

1

(1-3)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

11

3

(2-4)

3

(2-4)

3

(2-5)

1

(1-3)

3

(2-4)

2,3

(1-4)

2

(1-2)

12

2

(1-3)

4,3

(1-4)

2

(1-3)

1

(1-2)

1.2

(1-3)

1

(1-2)

3, 4

(2-5)

14

1

(1)

1

(1)

6,5

(4-8)

1

(1-2)

14-16(12-16)

1

(1)

1

(1-2)

Meso thorax

1

8

(6-9)

5,6

(4-7)

7,6

(5-9)

3,4

(1-5)

9-11

(8-12)

9,8

(6-9)

4, 5

(4-5)

2

1

(1)

1

(1-2)

1

(1)

1

(1)

1

(1-2)

1

(1-2)

2, 3

(1-4)

3

3,4

(1-4)

3

(2-6)

4, 3

(3-5)

1

(1)

4, 5

(3-6)

3, 4

(2-4)

2-4

(1-4)

4

4

(2-4)

4, 5

(2-6)

5,4

(3-6)

3

(2-4)

4, 5

(2-5)

3,2

(2-4)

4-5

(3-7)

5

1

(1-2)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

6

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

7

1

(1)

1

(1-2)

1

(1)

1

(1)

2

(2-3)

1

(1)

1

(1)

Uranotaenia in Solomon Islands — Belkin

385

TABLE 1 (Continued)

atra

barnesi

ci vinskii

quadrimaculata

sexaueri

Solomon is

wy sockii

8

4,5

(4-5)

4

(4)

5,6

(5-6)

3,2

(2-5)

5

(4-6)

5

(5-6)

4

(4-5)

9

4

(4-5)

5

(4-5)

5

(5-6)

1

(1)

5

(4-6)

5

(4-6)

5

(5)

10

1

(1-2)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

11

1-3

(1-3)

1

(1-3)

1

(1-4)

1

(1)

3

(2-5)

1

(1-3)

1

(1-3)

12

1

(1)

1

(1)

1

(1-3)

1

(1)

1

(1-4)

1

(1)

1

(1)

13

15-

20(8-21)

13-

18(7-20)

20-

25( 16-30)

3. 4

(3-5)

28-

30(28-33)

30-

35(26-36)

15-

20( 10-24)

14

12-

16(10-19)

16-

19(11-21)

30-

35(20-35)

5

(3-6)

28-

32( 18-32)

35-

39(30-39)

15-

20( 10-23)

Metathorax 1

6

(5-7)

4

(4-5)

6

(4-7)

3

(2-4)

9,10 (7-11)

7.8

(5-11)

4,3

(3-5)

2

2

(2-3)

1, 2

(1-3)

3

(2-4)

3-1

(1-3)

2, 1

(1-4)

2,3

(1-3)

2

(2-3)

3

4. 5

(4-6)

6, 5

(4-6)

6

(5-7)

3,2

(2-3)

7,6

(4-9)

4-6

(4-8)

4, 5

(2-5)

4

3,2

(1-4)

2, 3

(1-3)

5, 4

(4-6)

1

(1-2)

4, 3

(2-5)

3-5

(2-6)

3

(2-4)

5

1

(1)

1

(1-2)

1

(1)

1

(1)

3, 2

(1-3)

1,2

(1-2)

2

(1-3)

6

2

(1-2)

3,2

(1-4)

2

(1-3)

1

(1-2)

3, 2

(2-4)

2

(1-3)

2

(2-3)

7

6-8

(6-9)

9,7

(6-9)

8

(7-11)

3, 2

(1-5)

8

(7-8)

7,8

(7-8)

8

(6-10)

8

10-

12(6-12)

7

(5-9)

11-

13(9-15)

3, 4

(2-4)

13-

11(8-15)

15-

18(13-21)

10-

14(8-19)

9

4

(3-5)

5

(4-6)

5

(4-5)

1

(1)

5

(3-6)

5

(4-6)

5

(4-5)

10

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

11

2

(1-4)

1

(1-2)

4, 3

(2-5)

1

(1-2)

3

(3-5)

1

(1-2)

1

(1-3)

12

2, 3

(1-4)

2, 1

(1-3)

2, 3

(1-3)

1

(1-2)

3, 2

(1-4)

2

(1-3)

1

(1-2)

13

7

(6-8)

7

(6-7)

8

(7-9)

5,4

(4-5)

13-

11( 10-14)

9-11 (8-12)

5, 6

(4-6)

Abd. I 1

5,6

(4-6)

4

(4-5)

6, 7

(5-8)

2, 1

(1-4)

8,7

(7-11)

8,7

(4-11)

4

(3-5)

2

1

(1)

2

(1-3)

1

(1-2)

1

(1-3)

1

(1-2)

1

(1)

1

(1)

3

1

(1)

1

(1-2)

1

(1-2)

2, 3

(1-4)

1

(1)

1

(1)

1, 2

(1-2)

4

6,7

(4-8)

8,7

(7-11)

11-

13(10-15)

3, 2

(2-4)

13-

11(9-17)

10, 11(9-15)

8-10 (7-15)

5

3

(3-4)

2,3

(2-4)

4

(3-5)

1

(1-2)

4, 3

(2-5)

3, 4

(2-5)

3,2

(2-4)

6

2

(2)

3

(3)

2

(2)

2, 1

( 1-2)

2

(2)

3

(3)

2

(1-3)

7

1

(1)

1

(1)

1

(1)

1

( 1-2)

1

(1)

1

(1)

1

(1-2)

8

2

(2-3)

2

(1-2)

2

(1-3)

absen t

3, 2

(1-5)

2

(1-2)

2, 3

(2-3)

9

4, 5

(4-7)

3

(2-5)

9,8

(6-11)

1

(1-2)

5-7

(4-9)

8,7

(5-8)

6, 5

(4-7)

10

2, 3

(2-3)

3

(1-4)

5

(3-6)

3,2

(2-5)

3, 4

(2-5)

S

(1-4)

3,4

( 1-4)

11

absen t

absen t

absen t

3

(2-4)

absen t

absen t

absen t

12

2, 3

( 1-3)

2, 1

(1-3)

2

(1-3)

1

(1)

2, 3

(1-3)

2, 3

(2-3)

1,2

(1-2)

13

2

(2)

3,2

(1-4)

2, 3

(1-3)

1, 2

(1-3)

3

(1-5)

1, 2

(1-2)

2, 3

(2-5)

Abd. 11 0

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

5,6

(4-7)

4

(4-6)

6, 5

(5-6)

2, 3

(2-3)

9,8

(7-6)

6,7

(5-10)

4, 3

(3-5)

2

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

3

4, 3

(3-6)

3

(2-5)

6, 5

(4-6)

2

(1-3)

4,3

(1-5)

4, 3

(3-6)

5, 6

(1-8)

4

5,4

(3-5)

6

(6-7)

6, 5

(5-7)

1. 2

(1-3)

7.8

(6-8)

6, 7

(6-9)

3

(2-5)

5

2.3

(1-3)

2, 3

(2-4)

3, 2

(2-4)

2

(1-2)

3

(3-4)

3,2

(1-4)

3

(2-5)

6

2

(2)

3

(3)

2

(2)

2, 1

(1-3)

2

(2)

3

(3)

2

(1-2)

386

PACIFIC SCIENCE, Vol. VII, July, 1953

TABLE 1 (Continued) 3

atra

barnesi

civinskii

quadrimacul ata

sexaueri

so lomoni s

wy sockii

7

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

1

(1)

1

(1)

8

2

(1-2)

2

(1-2)

1

(1-2)

1

(1)

2

(1-3)

2, 1

(1-2)

1

(1-2)

9

2

(2-3)

1

(1)

1

(1)

2

(1-3)

3,4

(3-5)

1

(1)

1

(1-2)

10

2

(2-3)

2, 3

(2-3)

2,3

(1-3)

1

(1-2)

3

(2-4)

2, 3

(2-3)

2

(1-3)

11

1,2

(1-2)

2, 1

(1-3)

2, 3

(1-3)

2, 3

(1-3)

2

(1-4)

2

(1-3)

2, 3

(2-4)

12

1, 2

(1-2)

2

(1-2)

2

(1-3)

2, 3

((1-4)

2

(1-3)

1

(1)

2,3

(1-4)

13

5

(4-5)

6,7

(5-7)

6, 5

(4-7)

4, 3

(3-5)

8,7

(7-12)

6,7

(5-7)

4

(3-5)

Abd. Ill

0

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

6,7

(6-8)

5

(5-6)

7

(5-8)

2

(2-3)

9, 10

(7-11)

10-8

(6-14)

4

(3-5)

2

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

1

(1)

1

(1)

3

2

(1-2)

3

(2-3)

2

(2-3)

1

(1-2)

3, 2

(2-4)

1.2

(1-3)

2, 3

(1-3)

4

2

(1-2)

3

(3)

2, 3

(2-3)

1, 2

(1-3)

2, 3

(1-3)

1

(1-2)

2, 3

(2-3)

5

2

(1-3)

2, 3

(2-3)

3

(2-4)

2

(1-3)

2

(2-3)

2,3

(1-3)

2

(2-4)

6

6

(5-7)

7

(6-7)

6,7

(5-7)

2,3

(2-3)

8,7

(7-9)

7,8

(7-10)

4

(3-4)

7

4, 5

(4-5)

4

(3-4)

5,6

(3-6)

3

(2-3)

5,6

(2-7)

4, 5

(4-7)

6

(6-9)

8

1-3

(1-3)

2

(2)

2,3

(2-3)

1

(1)

3,2

(1-6)

2, 1

(1-3)

2

(2-3)

9

2, 1

(1-2)

1

(1-2)

1

(1)

2, 3

(2-3)

2

(1-3)

1, 2

(1-2)

1

(1-2)

10

2

(1-2)

3, 2

(2-3)

2

(2-3)

1

(1)

2

(1-4)

2

(1-2)

2,3

(2-4)

11

2, 1

(1-3)

2,3

(2-3)

3

(1-4)

1

(1-2)

2,3

(1-5)

2, 1

(1-3)

2, 3

(2-4)

12

2, 1

(1-2)

1

(1-3)

3,2

(2-3)

1

(1)

2, 1

(1-3)

2

(1-2)

2

(2-3)

13

5

(4-6)

5,6

(5-6)

6,7

(5-7)

3,4

(2-5)

9-7

(7-12)

7,8

(6-10)

4

(3-5)

14

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

Abd. IV

0

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

6

(5-7)

5

(4-6)

7

(6-8)

2, 3

(1-3)

10, 11(7-12)

8-10

(5-12)

4

(2-5)

2

1

(1)

1

(1)

1

(1-2)

1

(1-2)

1

(1)

1

(1)

1

(1-2)

3

1

(1-2)

2, 1

( 1-3)

2, 1

(1-2)

1

(1)

1,2

(1-3)

1

(1-2)

1

(1-2)

4

2

(2-3)

4

(3-5)

3

(3-4)

2, 1

(1-3)

1-3

(1-4)

2

(2-3)

2

(2-4)

5

2

(1-2)

2, 3

(2-3)

3,4

(2-4)

2

(1-2)

2, 3

(1-5)

2

(1-3)

2

(2-4)

6

6,7

(6-7)

7

(6-9)

7,6

(6-9)

3,2

(2-3)

8,9

(7-11)

8-10

(7-11)

4, 5

(4-6)

7

4

(2-5)

2, 1

(1-3)

5,4

(4-7)

3,2

(2-4)

5,4

(3-6)

5, 4

(4-7)

5,6

(5-6)

8

2, 3

(1-3)

2

(1-2)

2

(2-3)

1

(1)

2,3

(2-3)

2

(1-3)

2

(2-4)

9

1, 2

(1-2)

1

(1)

1

(1-2)

2, 3

(2-3)

1

(1-2)

1

(1-2)

1, 2

(1-2)

10

2

(1-2)

2

(2-4)

2

(2-3)

1

(1)

2

(1-4)

2

(1-2)

3

(2-4)

11

1

(1-2)

1

(1)

1

(1-3)

1

(1)

1, 2

(1-2)

1

(1-2)

1

(1-2)

2

2, 1

(1-2)

2, 3

(1-3)

2

(1-3)

1

(1)

1,2

(1-3)

2

(2)

2, 1

(1-3)

13

5, 4

(4-6)

5,6

(5-6)

6

(6-7)

3

(3-5)

8

(7-10)

7,6

(5-9)

4

(3-4)

14

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

Abd. V

0

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

6, 5

(5-7)

5

(4-5)

7

(6-8)

2, 3

(2-3)

10-9

(8-12)

10, 11(6-14)

4

(3-5)

2

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

Uranotaenia in Solomon Islands — Belkin

387

TABLE 1 (Continued) 4

atra

barnesi

civinskii

quadrimaculats

» sexaueri

Solomon is

wysockii

3

4,3

(3-4)

5

(4-6)

5,4

(3-6)

2, 3

(2-3)

4,5

(3-6)

3,2

(2-5)

4, 3

(3-5)

4

2

(1-2)

2,3

(2-4)

3,2

(2-3)

2, 1

(1-2)

2,3

(1-4)

2

(1-2)

3,2

(2-4)

5

2

(2)

2

(1-3)

4-2

(2-4)

2

(1-2)

3,2

(1-4)

2

(1-3)

2,3

(2-4)

6

6

(6-7)

7

(6-8)

7,6

(6-9)

3,2

(2-6)

9

(7-10)

9, 10

(8-11)

4,5

(4-5)

7

4,5

(4-6)

3-5

(2-5)

5

(4-6)

3

(2-4)

4,3

(3-8)

5,6

(3-8)

6

(4-6)

8

2,3

(2-3)

2, 1

(1-2)

3,2

(2-4)

1

(1)

2,3

(1-3)

2

(1-3)

2, 3

(2-3)

9

2, 1

(1-2)

1

(1)

1

(1)

2

(2-3)

1

(1-2)

1,2

(1-2)

2,1

(1-2)

10

2, 1

(1-2)

2, 3

(2-3)

2,3

(2-3)

1

(1)

2

(1-3)

2, 1

(1-2)

2, 3

(1-3)

11

1

(1-2)

1

(1)

1

(1)

1

(1)

1,2

(1-2)

1

(1-2)

1

(1)

12

2

(1-2)

2

(1-3)

3

(1-3)

1

(1-2)

1,2

(1-3)

2

(1-2)

2,1

(1.2)

13

6, 5

(4-7)

5-7

(5-7)

6,7

(5-8)

3

(3-5)

8,7

(7-9)

8,7

(6-9)

4

(4-5)

14

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

Abd,

VI

0

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

Cl)

1

6

(5-7)

4

(3-5)

8,7

(5-9)

2,3

(2-3)

9-11

. (8-12)

7-9

(6-14)

3,4

(2-4)

2

1

(1)

1

(1)

1

(1)

1

(1-3)

1

(1)

1

(1)

1

(1-2)

3

2,3

(2-3)

4

(4-6)

3

(2-4)

1,2

(1-2)

2

(2-4)

2,3

(1-3)

3,2

(2-4)

4

2,3

(2-3)

3

(3-4)

3

(2-5)

2, 1

(1-2)

4,3

(3-4)

2

(1-3)

3

(2-4)

5

2

(1-3)

2, 3

(2-3)

3

(2-4)

2

(1-2)

2,3

(1-4)

2

(2-3)

3,4

(2-4)

6

7

(5-8)

7,6

(6-7)

9-7

(5-12)

3,2

(1-3)

9-7

(6-12)

11,10(8-14)

5

(4-7)

7

3, 4

(2-5)

3

(2-4)

3-5

(3-5)

3,2

(2-3)

3

(3-5)

3,2

(2-4)

5,4

(4-6)

8

3,2

(2-3)

1. 2

(1-2)

3,2

(2-4)

1

(1-2)

2,3

(1-3)

2

(1-3)

3,2

(2-3)

9

2

(1-2)

1

(1)

1

(1)

2,3

(2-3)

2, 1

(1-3)

1

(1-4)

2

(1-2)

10

2

(1-3)

2

(1-3)

2, 1

(1-3)

1

(1)

2

(1-3)

2

(1-3)

2, 1

(1-3)

11

2

(1-2)

3

(2-3)

2

(2-3)

1

(1)

3

(2-3)

2, 1

(1-3)

2,3

(1-4)

12

2

(1-3)

2, 1

(1-3)

2

(2-3)

1

(1)

3

(2-4)

1.2

(1-4)

2

(2-3)

13

15-17(8-18)

15-19(12-21)

25- 30(20-33)

2

(1-4)

25-30( 19-32)

22-25(20-31)

15-19(10-20

14

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

Abd.

VII

0

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

5,6

(4-7)

4

(4)

6-8

(5-9)

2,3

(2-3)

8,9

(6-10)

7-9

(5- 10)

4

(2-4)

2

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

3

8,7

(6-9)

6

(5-7)

8-10

(8-10)

3,2

(2-3)

12,13(11-15)

11

(7-15)

4

(3-5)

4

2

(1-3)

1,2

(1-4)

2, 3

(1-3)

1

(1)

2

(1-4)

1.2

(1-3)

2, 1

(1-4)

5

2,3

(2-4)

2

(2-3)

3-5

(2-5)

2

(2)

3

(2-5)

3,2

(1-4)

2. 1

(1-4)

6

4, 5

(3-6)

4,5

(2-6)

6,7

(5- 10)

3,2

(2-4)

7,6

(5-8)

6.7

(5-9)

5,6

(4-7)

7

2, 3

(J-4)

2,3

(1-3)

4

(2-5)

2,3

(2-3)

3,2

(1-4)

4

(3-6)

2

(2-3)

8

5,5

(4-6)

1,2

(1-2)

8-10

(6-11)

2

(2-4)

6-8

(6-9)

7.8

(6-9)

4

(3-6)

9

1,2

(1-3)

1, 2

(1-2)

2

(2-3)

2,3

(2-3)

2

(2-4)

3,2

(2-4)

2

(2-4)

19

2

(1-2)

2

(2)

2

(2-3)

1

(1)

2

(2-3)

2, 1

(1-3)

2

(1-3)

11

2

(1-3)

3

(2-4)

3

(2-3)

1,2

(1-2)

4, 5

(3-6)

3

(2-4)

2

(2-3)

12

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

388

PACIFIC SCIENCE, VoL VII, July, 1953

TABLE 1 (Continued) 5

atra

barnesi

civinskii

quadrimaculata

sexaueri

solomonis

wysockii

13

5

(4-6)

4, 5

(4-6)

5,6

(5-6)

3

(2-5)

8,9

(6-10)

7

(6-8)

5,4 (4-6)

14

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1-2)

1 (1)

Abd. VIII 0

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1 (1)

1

4

(3-5)

3-5

(3-5)

4,5

(3-6)

2

(1-3)

5,6

(4-7)

5

(4-7)

2 (2-3)

2

3

(1-3)

2,3

( 1-4)

3

(1-4)

1

(1-2)

2,3

( 1-5)

1, 2

(1-3)

2 (1-3)

3

5,4

(4-5)

5

(4-5)

6,5

(4-6)

1

(1-2)

7

(7-9)

6,7

(5-8)

5,6 (2-8)

4

2

(1-2)

2

(1-3)

3,2

(1-6)

1,2

(1-3)

2, 1

(1-5)

1,2

(1-2)

2 (1-3)

5

7,8

(5-9)

8,7

(6-8)

8,9

(5-10)

3, 2

(2-4)

10, 11(6-12)

10-8

(7-12)

4 (3-5)

14

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

1 (1-2)

Comb

7,8

(5-8)

10,9

(8-11)

8,9

(6-9)

5,6

(3-7)

6-8

(6-8)

8,7

(7-10)

8,7 (6-9)

Siphon 1

9-11

(7-11)

11, 10(10-13)

10,9

(8-13)

2

(2-3)

8,7

(6-9)

10, 11(9-13)

4 (3-5)

2

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1 (1)

3-6

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1 (1)

7

1

(1)

1

(1)

1

(1)

2

(1-2)

1

(1)

1

(1)

1 (1)

8

1

(1)

1

(1)

1

(1)

2

(2)

1

(1)

1

(1)

1 (1)

9-13

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1 (1)

Pec ten

13, 12( 10-15)

12, 13(11-14)

16, 15(13-18)

7,6

(5-8)

14, 12(12-16)

11

(11-13)

19-21(13-25)

Abd. X 1

7-9

(6-10)

5

(3-6)

5,4

(4-7)

2, 1

(1-3)

6,5

(5-6)

5

(5-7)

2 (2)

2

2

(2)

4,5

(4-5)

3

(3)

1

(1-2)

3

(2-4)

3

(3)

3 (3-5)

3

2

(2)

2

(2)

2

(2)

1

(1-2)

2

(2)

2

(2)

2 (2-3)

4a

2

(2)

2

(2)

3

(3)

1

(1)

3

(3)

3

(3)

2 (2)

4b

2

(2-3)

2

(2)

2

(2)

1

(1)

2

(2)

2

(2)

1 (1-2)

4c

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

1 (1)

4d

1

(1)

1

(]1

1

(1-2)

1

(1)

1

(1)

1

(1)

1 (1)

4e

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

1

(1-2)

1 (1)

Uramtaenia in Solomon Islands' — Belkin

389

TABLE 2.

PUPA. SETAL BRANCHING

Limits of variation in pareKtheses, preceded b'y most usual number(s) observed

in ten specimens (20 hairs) in the order of frequency.
• secondarily branched at apex.

atra

barnesi

civinskii

cpadrimacul ata

sexaueri

solomonis

wysocki i

Cephalo-

1

7

(5-9)

10,9 (8-13)

5

(5-7)

1

(1-2)

6

(4-8)

5

(4-7)

4, 3

(3-4)

thorax

2

6,5

(5-7)

7,8 (7-10)

5

(5-7)

2, 1

(1-3)

6,5

(5-7)

5

(3-5)

4, 5

(3-7)

3

7,6

(5-10)

7,8 (7-10)

6, 5

(5-7)

1, 2

(1-3)

7-5

(5-7)

5

(3-6)

5,6

(4-7)

4

9

(8-10)

14-12(9-16)

6, 5

(4-8)

1

(1-2)

7,6

(6-8)

7,8

(5-8)

4, 3

(3-5)

5

11

(5-11)

12-14(10-17)

7,6

(6-10)

2, 3

(2-3)

8,9

(5-9)

6.7

(5-9)

4, 3

(3-5)

6

1

(1)

1 (1)

1

(.1-2)

1

(1)

1

(1)

1

(1)

1

(1)

7

5

(4-6)

6,7 (4-7)

4. 5

(3-6)

2

(1-3)

6, 5

(3-7)

4

(3-6)

2

( 1-3)

8

6,7

(5-7)

5,4 (4-6)

6.7

(5-11)

4,3

(3-5)

8,9

(7-10)

7,6

(6-8)

7-9

(6-10)

9

9,8

(7-10)

9-11 (6-11)

6, 5

(4-8)

1

(1)

6,7

(5-8)

4, 5

(3-6)

4

(2-5)

10

7-5

(4-8)

5,6 (3-6)

4

(3-6)

2

(1-2)

5

(4-7)

4.3

(2-4)

2

(1-4)

11

4,3

(3-5)

5.4 (4-7)

2, 3

(2-4)

1

(1)

4

(3-5)

5,4

(4-6)

4, 3

(3-6)

12

7,8

(6-9)

10,9 (6-11)

5,6

(4-7)

1

(1)

5,4

(3-6)

3,4

(3-5)

3, 2

(2-4)

Abd. I

1

14-

16(12-17)

16-19(16-25)

18-20( 14-22)

11-

13(9-17)

15-20( 14-25)

20

29( 19-30 )

18-:

22( 15-24)

2

1

(1)

1 (1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

3

3, 4

(3-4)

4 (4-5)

3

(2-3)

1

(1)

5-3

(3-6)

4-6

(4-7)

4,3

(3-4)

4

6-8

(6-8)

7,8 (4-9)

6, 5

(5-7)

2,3

(1-3)

6-4

(3-8)

6

(5-8)

4

(1-5)

5

2, 1

(1-5)

4 (1-6)

3, 4

(1-7)

1

(1)

4

(1-7)

3-5

(3-7)

3.2

(1-4)

6

4,3

(3-6)

7,6 (5-9)

3

(2-4)

1

(1)

3

(3-4)

2

(2-3)

1

(1-2)

7

2

(1-3)

1.2 (1-2)

2,3

(2-3)

1

(1)

2

(1-2)

2

(2-4)

1

(1-4)

10

4

(3-5)

3,4 (3-5)

3-5

(2-5)

2

(2)

5,4

(3-5)

3,4

(2-5)

3,2

(1-3)

Abd. II

0

1

(1)

1 (1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

9-7

(6-10)

10-12(8-13)

4, 5*

(3-8)

1,2

(1-5)

6,7

(5-8)

10, 11(8-13)

4, 5

(3-8)

2

3,4

(2-5)

5,4 (3-5)

3,2

(2-4)

1

(1-2)

4, 5

(2-6)

5-7

(5-9)

2,3

(1-4)

3

7,6

(5-8)

10-14(10-14)

4, 5.

(3-5)

1

(1)

4, 5

(4-5)

5, 6

(4-7)

2, 3

(2-3)

4

1

(1)

1 (1)

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

5

3,4

(3-4)

3,4 (3-5)

4,3

(3-6)

2, 3

(1-4)

3, 4

(1-5)

5-3

(3-7)

2, 3

(2-4)

6

3

(2-3)

5,6 (4-6)

2

(2-3)

1

(1-2)

2

(1-3)

2

(2)

1

(1)

7

1

(1)

1 (1)

1

(1)

1

(1)

1,2

(1-2)

1

(1)

1

(1)

8

absent

absent

absen t

1

(1)

absen t

absent

absen t

10

5,4

(3-5)

4.3 (2-5)

2,3

(2-7)

2

(1-3)

3.4

(1-4)

4. 3

(2-5)

3,2

(2-3)

12

absent

absent

absen t

0,1

(0-2)

absent

0

(0-2)

absen t

Abd. Ill

0

1

(1)

1 (1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

8,9

(7-10)

12-10(8-14)

6.7

(5-8)

1,2

(1-2)

6,7

(6-9)

7.8

(6-10)

3

(2-5)

2

1

(1)

1 (1)

1

(1~2)

1

(1)

1

CD

1

(1)

1

(1)

3

7

(6-10)

12-14(7-19)

6,5

(5-7)

1

(1)

6

(4-7)

8,7

(7-11)

3

(2-4)

4

3

C3-4)

4,3 (2-6)

3

(2-4) .

1

(1)

(3-6)

6,5

(3-8)

3,4

(3-5)

390

PACIFIC SCIENCE, Vol. VII, July, 1953

TABLE 2 (Continued) 2

atra

barnesi

ci

vinski i

quadrimaculata

sexaueri

solomonis

wysockii

5

3,4

(3-5)

3

(2-4)

3

(2-5)

2

(1-2)

3

(2-4)

3,2

(2-4)

3,2

(2-4)

6

4

(3-5)

6

(5-7)

4

(4-5)

1

(1-2)

3

(2-3)

3,2

(2-4)

1

(1-2)

7

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

8

2. 3

(1-3)

2, 3

(2-5)

2, 3

(1-3)

1

(1)

2

(1-3)

2

(2-4)

2

(1-4)

10

4-2

(1-4)

2

(1-5)

3, 4

(3-6)

2

(2-3)

2, 3

(2-5)

3

(2-4)

2-4

(1-4)

12

3

(2,3)

3

(3-4)

3

(2-4)

1,2

(1-3)

3

(1-3)

2

(1-3)

2

(1-4)

13

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

14

1

(1)

1

(1)

1

(1)

1

(0-1)

1

(1)

1

(1)

1

(1)

Abd.

IV

0

h

(1)

1

(1)

1

(1-2)

1

(1)

1

(1)

1

(1)

1

Cl)

1

7-5

(4-8)

10-12(6-12)

5,6

(4-7)

2

(1-2)

6

(5-9)

6,7

(5-9)

3,4

(3- 4)

2

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

3

5,6

(4-6)

7,6

(4-8)

6,5

(4-6)

2

(2-4)

6, 5

(4-7)

6,7

(4-8)

4, 5

(3-6)

4

1

(1)

3,4

(2-5)

5

(4-8)

1

(1)

6

(5-7)

6-8

(5-9)

3

(3)

5

3

(1-4)

3,4

(1-5)

2, 3

(2-4)

2

(1-2)

3, 2

(1-3)

3, 2

(2-3)

2, 3

(2-3)

6

4

(3-5)

7

(4-8)

4

(3-4)

1

(1-2)

3

(2-4)

2. 3

(2-4)

1

(1)

7

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

8

3,2

(2-6)

2-4

(2-4)

4

(2-5)

2

(1-3)

3,4

(1-4)

4, 5

(3-5)

2-4

(1-4)

10

2, 3

(1-3)

2

(2-4)

2, 3

(1-4)

2

(1-2)

2

(1-3)

3

(2-4)

3

(2-4)

12

3

(2-3)

3

(2-4)

2-4

(2-4)

1

(1-2)

2, 3

(1-3)

2

(2-3)

2

(1-3)

13

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

14

1

(1)

1

1

(1)

1

(0-1)

1

(1)

1

(1)

1

(1)

Abd.

V

0

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

5,6

(4-7)

7

(5-9)

4, 5

(4-6)

2

(1-3)

5,6

(4-6)

5

(4-6)

3, 4

(2-4)

2

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

3

3, 4

(3-4)

3, 4

(3-4)

3,4

(2-4)

2

(2)

4, 3

(2-5)

3

(2-4)

3,4

(2-4)

4

1

(1)

3

(2-4)

3

(2-5)

1

(1)

6

(5-7)

5.6

(4-7)

3

(3-4)

5

4.5

(3-5)

5,6

(3-6)

4

(3-6)

2

(1-3)

4, 5

(3-6)

5,6

(4-7)

4, 3

(2-4)

6

4

(3-5)

6,7

(3-9)

3, 4

(3-4)

1

(1-2)

3

(2-4)

3,2

(2-4)

1

(1-2)

7

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

8

2-4

(1-4)

2, 3

(2-3)

4

(3-5)

2, 1

(1-2)

2, 3

(1-3)

4

(2-5)

3

(2-4)

10

4

(2-5)

4, 5

(3-5)

4, 5

(3-7)

2

(2-3)

4

(2-5)

4, 5

(3-6)

3, 2

(1-4)

12

3, 2

(2-4)

2, 3

(2-3)

2

(1-3)

1

(1-2)

3,2

(1-4)

2

(1-3)

O

(2-5)

13

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

14

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

Abd.

VI

0

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

5

(4-7)

5,6

(4-7)

4, 5

(3-6)

2

(1-3)

5.4

(4-6)

5,4

(3-6)

4,3

(3-5)

2

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

3

3

(2-4)

4,3

(2-5)

3

(1-4)

2

(2-3)

4,3

(1-4)

3, 2

(1-3)

3,4

(2-4)

4

2

(1-2)

4

(3-5)

3

(2-4)

1

(1)

5,4

(3-6)

5,4

(4-6)

3

(3)

5

4, 5

(4-5)

5,6

(3-6)

4, 3

(2-4)

2

(1-2)

4,5

(3-6)

3

(3-4)

4-2

(2-5)

Uranotaenia in Solomon Islands — Belkin

391

TABLE 2 (Continued)

atra

barnesi

ci vinskii

quadrimaculata

sexaueri

solomoni s

wysockii

6

-4

(3-5)

5-7

(4-7)

3, 4

(3-4)

1

(1)

3

(3-5)

3,2

(2-4)

2

(2-3)

7

1

(1)

1

(1)

1

Cl)

1

(1)

1

(1)

1

(1)

1

(1)

8

2,3

(2^4)

3, 2

(1-4)

3,2

(1-4)

2

(1-3)

2, 3

(1-3)

4-2

(2-5)

2,3

(1-4)

10

3

(2-3)

3

(1-4)

2

(1-3)

2

(1-2)

2

(1-3)

3,2

(1-4)

2

(1-3)

12

2

(1-3)

2

(2-3)

2

(1-2)

2. 1

(1-2)

2,3

(2-3)

2

(1-3)

2

(1-3)

13

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

14

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

Abd. VII

0

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

5, 4

(3-6)

6,5

(5-7)

3, 4

(3-5)

2, 1

(1-3)

4

(3-5)

4, 5

(3-6)

3,4

(3-5)

2

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

3

4, 5

(3-6)

5,6

(4-7)

5,4

(3-5)

2

(2)

5, 4

(4-6) ■

6,5

(3-6)

7,6

(5-8)

4

2

(2)

5,4

(4-6)

3

(2-5)

1

(1)

5,4

(4-5)

5,4

(3-6)

3

(3-4)

5

4

(3-4)

4-6

(2-6)

3,2

(1-4)

1,2

(1-2)

4

(2-5)

3

(2-3)

2, 3

(2-5)

6

4

(2-4)

5

(3-6)

3,4

(2-4)

1

(1)

3

(2-4)

3,4

(3-4)

3,4

(3-6)

7

1

(1-2)

1

(1-2)

1-3

(1-3)

1

(1)

3

(1-3)

2, 3

(2-3)

2,3

(1-4)

8

4, 3

(2-5)

3-5

(2-5)

4,3

(2-5)

3,2

(2-3)

3, 4

(2-5)

4, 3

(2-5)

3,4

(1-6)

10^

3,2

(2-3)

3

(2-4)

2

(1-3)

2

(1-3)

2

(2)

2, 3

(2-3)

2

(1-3)

;12

2,3

(2-3)

3

(2-3)

2, 3

(1-3)

1

(1-2)

3, 2

(2-3)

3

(1-3)

3,2

(2-4)

is':

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

1

(1)

14

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

Abd. VIII

0

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

4

5,4

(4-5)

5,6

(4-8)

3

(1-5)

1

(1)

3, 4

(1-4)

3

(2-4)

3

(2.-4)

7

4

(3-5)

4

(3-6)

3

(2-5)

1

(1-2)

3,2

(2-5)

3

(2-5)

7-9

(5-16)

1.4

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1-2)

Abd. IX

1

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

(1)

1

CD

Paddle

1

1

Cl)

1

(1)

1

(1)

1

(1-2)

1

(1)

1

(1)

absent

2

absent

absent

absent

1

(1-2)

absent

absent

absent

NOTES

On the Establishment of the Oahu Research Center of the
University of California

In recent years there has been a general
recognition by many national governments
that so-called basic or fundamental scientific
research deserves financial support. Usually
this recognition has been stimulated not so
much by admiration for the scientists or a
desire to increase the cultural heritage of man-
kind as by a hard-headed cost accounting ap-
plied to the history of technological progress
in the past 25 years. In the long run it pays
to support basic research. In the process of
financing apparently useless projects the gov-
ernments have, in spite of waste and some
incompetence, received more than their money s
worth in by-products. In the United States,
naturally, the first example that comes to mind
is the Manhattan Project and its subsequent
development into the complicated investiga-
tions now carried on by the Atomic Energy
Commission. One is apt to forget that the
Federal Government, while making its major
effort in the field of atomic physics, also sup-
ports a multitude of other research projects and
that these are not always developmental in aim
or technological in scope. While some gov-
ernments, operating on the principle that he
who pays the piper calls the tune, have tended
to support basic researches only in their own
laboratories, sometimes to the extent of
starving the universities out of certain fields,
in the United States there has been a consist-
ent effort to avoid centralization by contracting
for basic research with the institutions most
competent to carry it out, namely, the uni-
versities and technological institutes.

The result has been, since the end of World
War II, what is probably the greatest outpour-
ing of the taxpayers’ money into a purely in-
tellectual venture that the world has seen. For
7 years the people of the United States have
paid and have asked no questions. The ad-
ministrators of the program, both civilian and,
surprisingly enough, military, have recognized
that it is almost impossible to assess the value
of a basic research program in a decade. The

results are often intangible, sometimes of a
purely negative kind (as showing, for example,
that a frontal attack on a given problem is
unprofitable) or, in some cases, of value only
in training the technologists and research
workers of the future. Recently, however, there
have been murmurs of discontent. As the tax
bill grows larger and international tensions
increase, three groups have criticized the most
sensitive of the basic research programs: that
supported by the Armed Services. First, the
operating agencies of the Armed Forces, the
men responsible for using weapons and plan-
ning battles, have felt that they have received
practically no benefit from the basic research
program, that progress is too slow in a period
of war such as that in Korea, and that the
money would be better spent on technological
development the results of which can be fore-
seen now. Second, university administrators
have complained that they are slowly being
strangled by the red tape which is inevitably
spun around research projects supported by the
taxpayer and which increases every year. Com-
plicated cost accounting and unimaginative
auditing exasperate them and make them in-
clined to ask research workers in fundamental
science to return to their ivory towers, however
cramped these may now appear. Last, the re-
search workers themselves are complaining.
Some of them have been on contract work for
6 or 7 years, have obtained good results, and
have written voluminous reports, as required
by the university contract system. But these
reports have been buried in official archives;
often the operating arms of the services do not
even know of their existence; sometimes the
solution of a serious military problem already
exists in embryo in the files of the agency most
worried about it. No serious rift has developed
yet, but, if the present tendencies persist, one
can foresee the end of military support for
basic research.

In many of the sciences this rift, no matter
how wide it became, would not be serious for

392

NOTES

393

the universities. Progress would continue at a
slower pace, as it has for 4 centuries. In at
least one field, however, a rift would be dis-
astrous to the universities and, ultimately, to
the Armed Services — the field of geophysics.
In many parts of that science, research cannot
even begin without access to the data, and the
data are beyond the scope of any single uni-
versity or congeries of universities to collect.
Adequate observation depends on costly equip-
ment, the employment of thousands of ob-
servers, and the facilities for collection and
dissemination that only governments possess.
On the other hand, government agencies tend
to see this information in a very practical light.
They may use it for many purposes, but with
few exceptions these amount to attempts to
solve ad hoc and day-to-day operating problems.
Their methods of treatment tend to become
stereotyped, and bureaucratic principles begin
to override scientific principles in the handling
of the data.

Against this background a very small but
significant experiment in co-operation between
the Air Force and a university is being estab-
lished in the Hawaiian Islands. Three insti-
tutions are involved, and the research work
about which the experiment is formed is chiefly
in the field of tropical meteorology. Financial
and physical support is given by Air Force
Cambridge Research Laboratories, the project
being monitored by the Geophysical Research
Directorate of those laboratories. Air Weather
Service, the operating agency of the Air Force
most vitally interested in tropical meteorology,
supplies both physical support and personnel.
The University of California, through its In-
stitute of Geophysics, supplies, under contract,
civilian scientists and technicians and the over-
all direction to the research. The three groups
have begun to work together at Wheeler Field
on the island of Oahu, and full operation is
expected in the very near future. Each agency
has a different name for the project. That used
by the civilian scientists working on the project
and by the University of California is the Oahu
Research Center. The general objectives of the
work are, first, to continue the basic research
work in tropical meteorology that has been
conducted by the Institute of Geophysics at the
University of California at Los Angeles during
the past 5 years; second, to extend those re-
searches to include the effects of mountainous
islands on meteorological phenomena in the
tropics; third, to continue the Institute’s work
on the macrophysics of tropical clouds, especially

as they are affected by high relief; and, finally
and perhaps most important, to translate with
as little delay as possible the results of these
investigations into technological procedures
that can be used by Air Weather Service in any
part of the tropics. The work has been split up
and assigned to three divisions of the project
called, respectively, the basic laboratory, the
experimental laboratory, and the operational
laboratory.

The basic laboratory will continue the in-
vestigations, begun in Los Angeles, of the gen-
eral atmospheric circulation in the Pacific; of
the origin, development, and movement of
typhoons and hurricanes; and of the origin,
development, and movement of upper-level
tropical cyclones. It will extend its work to the
monsoon countries, more particularly those of
southeast Asia, and, in conjunction with the
experimental laboratory, will attempt to develop
an indirect aerology of the tropics.

The experimental laboratory will study the
effects of mountainous islands on the fields of
pressure, temperature, rainfall, and wind. It
will undertake an investigation of orographic
cloud. It will analyze many thousands of' feet
of movie film and stills of tropical clouds over
the Central and South Pacific that have been
accumulated during the past 2 years and will
extend this photographic collection. Investi-
gations of wind structure in the neighborhood
of tropical clouds have been planned, and some
theoretical work has already been done on this
topic.

Both these laboratories will continue to
publish their results in official reports and in
scientific papers. However, the results will be
available to the operational laboratory long
before they reach the press. This laboratory
will be built around a weather station, of the
type that has become standard in the Air Force.
Current data, collected in the station by tele-
type, will be analyzed by both the civilian and
military scientists day by day and under con-
ditions as near those found in the field as
possible. New techniques, suggested by the
general researches of the project, will be
developed, tested, and evaluated in this station.
Any procedure found practicable and useful
will be described in operating manuals suitable
for Air Force use in the field. Finally, the film
produced by the experimental laboratory will
be surveyed and selections made by the oper-
ational laboratory for incorporation in training
movies suitable for the Air Force.

394

PACIFIC SCIENCE, Vol. VII, July, 1953

Each scientist, whether military or civilian,
has been assigned a part of this large field as
a responsibility. To prevent over-specialization,
however, and to ensure the free flow of infor-
mation throughout the project, each scientist
will be assisted by a research committee, whose
task it will be to monitor the work and assist
the responsible investigator in every way. Thus,
each worker will have not only his responsible
task but two committee tasks as well. In ad-
dition, operational problems will be explored
in a daily map discussion, and scientific results
will be reported in formal seminars, held every
2 weeks.

The danger to civilian scientists, inherent in
an off-campus project of this kind, lies in the

lack of outside criticism and stimulation. In
Oahu this stimulation will come from the
Weather Bureau and from meteorologists of
the Pineapple Research Institute and the Air
Weather Service units in the area. As soon as
full operation of the unit is assured, dis-
tinguished scientists from outside Hawaii will
be invited to visit the project and to consult
on its problems.

Military direction of the project is in the
hands of Lieutenant Colonel G. H. Duncan; the
civilian group will be supervised by Professor
C. E. Palmer of the Institute of Geophysics of
the University of California. ■ — Clarence E.
Palmer.

News Note

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Nomenclature Section of the Eighth Inter-
national Botanical Congress to be held in Paris,
July 1954.

Section Nomenclature

Proposals regarding the International Code of
Botanical Nomenclature (1952) must be
submitted to the Rapporteur-General Dr J.
Lanjouw before 1 December 1953. All pro-
posals can be published in Taxon. Botanists
preparing proposals are earnestly requested to
give them the form of the example which will
be published in Taxon voL 2 no. 2 (March
1953).

Office of the Rapporteur-General:

International Bureau for Plant Taxonomy and
Nomenclature, Lange Nieuwstraat 106, Utrecht
— Netherlands.

in any way, and should be mailed flat. Inserts
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Batzo, Roderick L., and J. K. Ripkin. 1849.
A treatise on Pacific gastropods, vii + 326
pp., 8 figs., 1 map. Rice and Shipley, Boston.

Crawford, David L. 1920^i. New or interesting
Psyllidae of the Pacific Coast (Homop.).
News 31 (1): 12-14.

1920^. Cerotrioza (Psyllidae, Homop-

tera). Hawaii. Ent. Soc., Proc. 4 (2): 374-

375.

Rock, Joseph F. 1916. The sandalwoods of Ha-
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VII

OCTOBER, 1953

PACIFIC

SCIENCE

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

IN THIS ISSUE: Townsley — Hawaiian Stomatopod
Crustaceans # Sherman, Kanehiro, Matsusaka — Develop-
ment of Laterite # Hanson — Flukes of Genus Schistorchis

• Knight and Hull — Fart Three of Aedes Mosquitoes
of the Philippine Islands • Morgan — Studies in Electro-
fishing in Sea Water # Bank — Biological Succession in
the Aleutians # Wilson — • Records of Alaskan Eurytemora

• Hardy — Bibionidae of New Zealand # NOTES #
INDEX

i.

p

L

Published by

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HONOLULU, HAWAII

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Leonard D. Tuthill, Editor-m-CMef
Department of Zoology and Entomology, University of Hawaii

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Department of Bacteriology, University of Hawaii

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Office of Publications and Information, University of Hawaii

Earl J. Anderson

Pineapple Research Institute Pathologist
Pineapple Research Institute
Honolulu 14, Hawaii

Townsend Cromwell
Pacific Oceanic Fishery Investigations
Oceanographer

Pacific Oc^nic Fishery Investigations
Honolulu 14, Hawaii

Maxwell S. Doty

Department of Botany, University of Hawaii

R. A. Falla

Dominion Museum, Wellington, New Zealand

Christopher Gregory
Department of Mathematics
University of Hawaii

D. Elmo Hardy

Associate Entomologist, University of Hawaii
Agricultural Experiment Station

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Department of Meteorology
Pineapple Research Institute
Honolulu 14, Hawaii

Norman S. Noble
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Research Organization
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Harold St. John

Department of Botany, University of Hawaii

G. Donald Sherman

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Agricultural Experiment Station

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University of Hawaii

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Scripps Institution of Oceanography
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( Continued on inside back cover)

PACIFIC SCIENCE

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

VOL. VII OCTOBER, 1953 NO. 4

Previous issue published July 3, 1953

CONTENTS

, PAGE

Adult and Larval Stomatopod Crustaceans Occurring in Hawaiian

Waters, Sidney Joseph Townsley 399

The Role of Dehydration in the Development of Laterite, G. Donald

Sherman, Yoshinori Kanehiro, and Yoshito Matsusaka . . . 438

A Discussion of the Trematode Genus Schistorchis (Family Lepocreadiidae)
with Descriptions of Two New Species from Haiuaii. Mary

Louise Hanson . 447

The Aedes Mosquitoes of the Philippine Islands. 111. Suh genera Aedimorphus,
Banksinella, Aedes, and Cancraedes (Diptera, Culicidae). Kenneth L.
Knight and William B. Hull 453

The Response of a Tropical Fish to Direct Current and Its Application to the

Problems of Electrofishing in Sea Water. Morris E. Morgan . . , . 482

Biological Succession in the Aleutians. Theodore P. Bank, II ..... . 493

New Alaskan Records of Eurytemora (Crustacea, Copepoda). Mildred

Stratton Wilson 504

The Bihionidae of New Zealand (Diptera). D. Elmo Hardy 513

NOTES:

Hawaii as a Cloud Physics Laboratory. A. H. Woodcock . . . . . .522
INDEX 525

Pacific Science is published quarterly by the University of Hawaii Press, in January,
April, July, and October. Subscription price is $3.00 a year; single copies are |1.00.
Check or money order payable to University of Hawaii should be sent to University of
Hawaii Press, Honolulu 14, Hawaii, U.S.A.

Adult and Larval Stomatopod Crustaceans
Occurring in Hawaiian Waters^

Sidney Joseph Townsley^

INTRODUCTION

The Stomatopoda constitute the only order
belonging to the division Hoplocarida of the
malacostracan Crustacea. All living forms are
included in a single family, the Squillidae,
whose characteristics are the same as those
of the order. The order, according to Bigelow
(1894), may be defined as Crustacea which
have these characteristics: the stalked eyes and
antennules are borne upon distinct movable
segments; the adult rostrum is separated from
the carapace by a movable joint; the carapace
is small and does not cover the last four tho-
racic somites; the first five pairs of thoracic
appendages are not biramous and serve as
accessory mouth parts, with the second pair
strongly developed into large raptorial limbs ;
the last three pairs of thoracic appendages
are adapted for walking, are biramous and bear
a lateral segment upon the penultimate seg-
ment; the abdomen is well developed; fila-
mentous gills are carried upon the exopod of
the first five abdominal appendages; the sixth
abdominal appendages, the uropods, with the
telson, act as a powerful tail fan.

At the present time the six recognized gen-
era have a world-wide distribution, being
found in the tropical, subtropical, and tem-
perate waters of the Atlantic, Pacific, and In-
dian oceans. All six genera are represented in
the Hawaiian fauna. Edmondson (1921), re-
porting upon the stomatopods in the collec-

^ Contribution No. 37, Hawaii Marine Laboratory,
Prepared as partial requirement for the M.S. degree.
University of Hawaii. Manuscript received November
14, 1951.

^ Graduate assistant, Osborn Zoological Laboratory,
Yale University.

tion of the Bernice P. Bishop Museum, found
six genera and eight species from Hawaii:
Squilla oratoria, 5. alha, Pseudosquilla ciliata^
P. oculata^ Coronida sinuosa, Lysiosquilla macu-
lata, Odontodactylus hanseni, and Gomdactylus
guerini. Since that time no extensive studies
have been made on specimens from Hawaii,
but occasional reports of previously recorded
species from this region have found their way
into the literature. In addition to these eight
species which have been again collected dur-
ing the progress of this study, I have found
two species which have not been recorded
from this region previously, namely, Squilla
hoops and an undescribed species of Squilla.

Stomatopods are usually found in very shal-
low water, but some species occur at greater
depths. I have collected Gomdactylus guerini
in Hawaii only at depths over 100 meters, and
Kemp (1913) reports that Squilla leptosquilla
was dredged by the "'Investigator” at a depth
of 370-419 fathoms. The adults are bottom
dwellers where they may burrow into the sub-
strate or inhabit crevices in coral. They are
seclusive in their habits and during the day
are usually found either in their burrows or
but a short distance away. At times they wan-
der some distance from their burrows, as is
shown by the fact that they are occasionally
captured in crab nets and at submersible lights
used for collecting at night. Inasmuch as the
burrows are quite deep (a foot or more below
the surface), and because the animals retire
to the bottom of them at the least alarm,
stomatopods are quite difficult to collect;
consequently, the group has been studied
very little, the most notable works being those

399

400

PACIFIC SCIENCE, Vol. VII, October, 1953

of Miers (1880), Brocks (1886), Bigelow
(1894), and Kemp (1913). Of these works,
the most useful for the identification of the
Hawaiian species has been Kemp’s mono-
graph on the Indo-Pacific species.

Stomatopods are carnivorous and readily
seize any flesh held in front of their burrows.
This behavior has provided the most satis-
factory means for capturing them, as they can
be lured from their burrows by suspending
fish flesh a short distance away and captured
when they attack the flesh. Considerable dex-
terity is necessary, however, as they withdraw
rapidly into their burrows. It has been im-
possible to keep more than one specimen at
a time in small aquaria as they are highly
cannibalistic.

It is doubtful whether the adults often serve
as food for other animals, although they may
occasionally be found in the stomach con-
tents of fish; but the planktonic larval stages
which compose a substantial proportion of
the neritic plankton form a considerable part
of the diet of pelagic fishes.

Instead of carrying their incubating eggs
attached to the abdominal appendages, as do
most Malacostraca, the stomatopods deposit
them at the bottom of the burrows where they
are aerated by currents of water produced by
the abdominal appendages of the parent.
These appendages are paddle shaped and con-
form to the cylindrical shape of the hole.
The eggs perish when they are deprived of this
constant current of water, and, as the female
will not remain with the eggs when placed
in an aquarium, no larvae have been reared
from the egg in captivity.

The young do not hatch as nauplius larvae.
All known forms hatch in an advanced stage
which bears many adult characteristics, but
it is difficult to assign the early larvae properly
because the diagnostic characters do not ap-
pear until later.

The larvae are delicate, transparent organ-
isms which lead a planktonic existence. The
larval life is long, and they pass through many
molts before reaching the adult stage. Not-

withstanding their great abundance, only two
or three early larval forms have been traced
to their adult counterpart. Their long plank-
tonic existence easily accounts for the world-
wide distribution of the genera and the pres-
ence of species such as Pseudosquilla ciliata
in the Pacific Ocean, Indian Ocean, Red Sea,
and Atlantic Ocean. At present there appear
to be only two species which are endemic in
Hawaii, namely, Coronida sinuosa and Squilla
calumnia n. sp.

Attempts were made to rear larval spec-
imens, collected in plankton tows, in the lab-
oratory, but this proved to be impossible as
their period of survival in captivity was very
short, and none of the larvae ever molted
while in an aquarium. Because rearing of the
larvae was unsuccessful, it was necessary to
study specimens collected in plankton tows
and at night light stations and material from
the stomach contents of pelagic fishes.

The earlier authors such as Miers (1880),
in their work on larval stomatopods, were
content with bestowing separate generic and
trivial names upon the larvae. If the adult
genus to which the larval specimens belonged
was known, they added the suffix -erichthus
to the first syllable of the adult genus plus
a trivial designation (except in the case of
alima of Squilla) . Thus Pseuderichthus communis
Hansen, according to this system of classifi-
cation, represented the larval form of a species
of Pseudosquilla. Because no keys to these
individual larval species were included, future
authors found it difficult to identify specimens
in their possession with those which had been
previously described, and new larval species
were erected to compensate for this. Claus
(1872) and Brooks (1886) have done the most
notable works on the larval forms. Giesbrecht
(1910), Foxon (1932), and Gurney (1942)
have reviewed the works of Claus and Brooks
and expanded their results to include other
unidentified larvae. However, I have found
that the material has not been adequately
simplified below the level of the genus, be-
cause in many cases they have retained the

Hawaiian Stomatopods — Townsley

401

previously assigned generic and trivial names
of the larvae, thus making it difficult to as-
certain the relationship of these species to
those collected in Hawaii. Most of the larvae
belonging to a particular genus resemble one
another so closely that it is virtually impossi-
ble to place them in their proper specific
category without rearing them to maturity.
Because of these difficulties, I have not tried
to relate the Hawaiian larval forms with those
of the previously described species, but where
it has been possible I have attempted to cor-
relate them with their proper adult species.
The specific identification of the larvae was
made in the manner used by Claus (1872)
and Brooks (1886) in which three methods
were employed: (1) a comparison of larval
stages with postlarval forms which show char-
acteristics of both the larvae and adults, in
this manner working from the adult back
through the larval series to the earliest iden-
tifiable stage; this is by far the most accurate
means of identification, except, of course,
where rearing of the larvae to the adult has
been accomplished; (2) a comparison of the
distribution of larval forms and known adults
in which it is possible to make specific diag-
noses with some accuracy in those cases where
only one or two adult species are known to
occur, but cases may arise in which the adult
specimen is unknown; (3) the relative abund-
ance of larval and adult species, although this
is the least accurate method used, has proved
to be useful when one species is known to be
more numerous than others and it is assumed
that the larval forms also retain the same
relative numerical relationship.

Almost all authors who have worked with
larval stomatopods have noted that invariably
there are forms present which in no respects
bear any resemblance to the adults recorded
for the region. This is also true of the material
at hand from Hawaii. This would seem to
indicate that there are numerous species
throughout the world, probably from greater
depths, which are known only in their larval
stages.

The present study was undertaken because
recent investigations on the feeding habits of
pelagic fishes have shown that the larval stages
of the Stomatopoda rank* second in impor-
tance as food for certain species of tuna in
the Hawaiian area, and that they are signifi-
cant in the diet of several other pelagic fishes.
Inasmuch as the adult stages of the pelagic
larvae involved are found on the reefs and
shores of the Islands, it is apparent that the
general economy of the neritic waters in the
vicinity is influenced to some extent by the
productivity of the reef and shore fauna. These
two points suggest a third, namely, that pe-
lagic fishes may be influenced to venture close
to shore because of the increased food supply
there.

In order to assess objectively the impor-
tance of these larval stages in the general
economy of the neritic realm, it is necessary
to be able to identify the species involved,
so that a rapid qualitative and quantitative
analysis of both the plankton and the stomach
contents of pelagic fishes may be made. The
present problem was designed to accomplish
these ends for the Stomatopoda, and it is
hoped that the results obtained will prove of
use to workers in other fields.

Acknowledgments

I wish to express my thanks to Dr. C. H.
Edmondson for permitting me to examine the
specimens in the Bernice P. Bishop Museum,
to Mr. Vernon Brock for making available
specimens collected by the Territorial Divi-
sion of Fish and Game, and to the staff of the
Pacific Oceanic Fishery Investigations for pro-
viding specimens collected by them. I am
indebted to Dr. A. H. Banner for his constant
help and criticism on the collection of mate-
rial and preparation of the manuscript, and
to Dr. R. W. Hiatt for suggesting the problem
and for criticism of the manuscript. In addi-
tion I would like to extend my thanks to Mr.
Kenji Ego, Mr. Daniel Yamashita, and Miss
Winifred Tseu for their help in collecting
specimens.

402

PACIFIC SCIENCE, Vol. VII, October, 1953

ADULT HAWAIIAN SQUILLIDAE

The genera of Squillidae easily fall into two
groups which are distinguished by differences
in the ischio-meral articulation of the raptorial
claw (second maxilliped) . In one group the
articulation between the merus and ischium
is terminal, whereas in the other the ischium
articulates at a point anterior to the proximal
end of the merus. This character is the basis
for the primary division of the family, with
Squilla, Pseudosquilla, Lysiosquilla^ and Coronida
comprising the first group, and with Gono-
dactylus and Odontodactylus comprising the lat-
ter group. Although the phylogenetic sig-
nificance of the ischio-meral articulation of
the raptorial claw is not clearly understood.

the species of each natural group resemble
each other far more closely than they do those
of the other group.

Squilla is separated from the other members
of the first group in that its species all possess
longitudinal carinae on the body segments.
Pseudosquilla, Lystosquilla, and Coronida are sep-
arated from one another by the shape of the
body, the carapace, and the telson, as is in-
dicated in the following key and descriptions.
Odontodactylus and Gonodactylus are readily dis-
tinguished because species of Odontodactylus
possess teeth on the inner margin of the dac-
tylus of the raptorial claw, whereas species of
Gonodactylus lack such teeth.

To avoid confusion in terminology. Figure
1 presents characteristics used in the key and

Fig. 1. Diagrams illustrating morphological details and terminology of stomatopods.

a, Rostrum, carapace, exposed thoracic somites, and first abdominal somite of Squilla. A-i, Structures of carapace.

A, Rostrum; b, anterolateral spine; C, median carina; D, gastric groove; E, intermediate carina; F, lateral carina;
G, marginal carina; H, cervical groove; I, posteriorly reflexed marginal carina. J-N, Structures of thoracic and
abdominal somites. J, Anterolateral spine of fifth thoracic somite; k, submedian carina; L, intermediate carina;
M, lateral carina; N, marginal carina. v-viii, Fifth to eighth thoracic somites, i, First abdominal somite.

3, Second thoracic appendage (raptorial claw). A, Carpus; B, merus; C, propodus; D, dactylus; E, ischium.

c, Telson and last two abdominal somites of Squilla. A-E, Structures of abdominal somites. A, 'Submedian carina;

B, lateral carina; c, marginal carina; D, intermediate carina; E, median carina. F-L, Structures of telson. F, Pre-
lateral denticle; G, lateral tooth; H, lateral denticle; I, intermediate tooth; j, intermediate denticles; K, sub-
median denticles; L, submedian tooth, v-vi. Fifth and sixth abdominal somites.

d, Last abdominal somite and telson of Odontodactylus. A-D, Structures of sixth abdominal somite (vi). A, Sub-
median carina; B, lateral carina; c, second intermediate carina; D, first intermediate carina. E-L, Structures of
telson. E, Marginal carina; F, second lateral carina; G, intermediate carina; H, submedian carina; I, median
carina; J, lateral tooth; K, intermediate tooth; L, submedian tooth.

e, Lateral aspect of head and carapace of Squilla. A, Cornea of eye; b, mandibular palp; C, antennal scale; D, first
thoracic appendage; E, third thoracic appendage; F, second thoracic appendage (raptorial claw).

Hawaiian Stomatopods — Townsley

403

in the descriptions of the species. These have
been partially adapted from Kemp (1913),
whose terminology has been employed
throughout this work.

Key to Adult Hawaiian Squillidae

A. Raptorial claw with articulation of me-
rus and ischium terminal (Fig. 3g), me-
rus grooved ventrally for reception of
propodus throughout its length, pro-
podus finely pectinate or with a series
of fixed spines along the upper margin,
dactylus not inflated at base ........ B

AA. Raptorial claw with ischio-meral arti-
culation at a point anterior to the prox-
imal end of the merus (Fig. 21c), ventral
surface of merus grooved for reception
of propodus for not more than 0.75 of
its length, propodus may or may not be
finely pectinate along the upper margin,
dactylus inflated at base (Odontodac-
tylus and Gonodactylus) .......... I

B. Carapace with well-marked carinae , cer-
vical groove defined across the dorsal
surface; first 5 abdominal somites with
longitudinal carinae (Squilla) ...... C

BB. Carapace without well-marked carinae,
cervical groove not extending over dor-
sal surface; first 5 abdominal somites
without longitudinal carinae ........ F

C. Anterolateral angles of carapace armed
with short, acute spines; cornea of eyes
set obliquely on stalk (Fig. 3^), nearly
1.5 times as wide as greatest breadth
of stalk, distinctly bilobed. ........ .D

CC. Anterolateral angles of carapace round-
ed, without short acute spines (occa-
sionally with spines in S, alhd)\ cornea
of eyes set transversely on eyestalk (Fig.
6), slightly wider than the greatest
breadth of stalk, very slightly bilobed
........................... ......E

D. Dactylus of raptorial claw with 6 teeth
body usually more than 100 mm. in

^ The number given for the teeth on the raptorial
dactylus throughout this study includes the terminal
tooth in all species, except Odontodactylus hanseni.

length from tip of rostrum to tip of

submedian spines of telson

.................. Squilla oratoria

DD. Dactylus of raptorial claw with 5 teeth;
body not reaching over 90 mm. in
length from tip of rostrum to tip of
submedian spines of telson ..........

..................... Squilla boops

E. Submedian teeth on posterior margin of

telson with movable tips; 5 or 6 mov-
able spines along outer margin of exo-
pod of uropod; 2 rounded lobes be-
tween inner and outer spine of basal
prolongation of uropod; body opaque
white throughout . Squilla alba

EE. Submedian teeth on posterior margin
of telson without movable tips; 8 or 9
movable spines along outer margin of
exopod of uropod; 1 rounded lobe
near base of inner spine of basal pro-
longation of uropod; body with definite
brown pigment marking carinae and
margins Squilla calumnia n. sp.

F. Lateral margins of exposed thoracic

somites and abdominal somites parallel
throughout, riot increasing in breadth
posteriorly; telson with sharp median
Carina and with lateral carinae (Pseu-
dosquilla) G

FF. Lateral margins of exposed thoracic and
abdominal somites not parallel through-
out, abdominal somites increasing in
breadth posteriorly; telson with median
Carina low and rounded, and without
lateral carinae (Lysiosquilla and Cor-
onida) H

G. Inner spine of basal prolongation of
uropod longer than outer spine; eyes
small and cylindrical; 6 carinae besides

median carina on telson .

.............. Pseudosquilla ciliata

GG. Inner spine of basal prolongation of
uropod shorter than outer spine; eyes
flattened, club-shaped; 8 carinae be-
sides median carina on telson .

............. Pseudosquilla oculata

H. Alternating bands of light and dark pig-

404

PACIFIC SCIENCE, Vol VII, October, 1953

merit throughout length of body; rapto-
rial dactylus with 9 or 10 teeth on inner
margin; telson with low, rounded, me-
dian Carina .... Lysiosquilla maculata
HH. No alternating bands of light and dark
pigment throughout length of body;
raptorial dactylus with 4 teeth on inner
margin; telson closely studded with
large tubercles in definite patterns .....

Coronida sinuosa

I. Dactylus of raptorial claw with 9 teeth
on inner margin; telson with ordinary
longitudinal carinae; rostrum with even-
ly convex anterior border and evenly
rounded angles; eyes large, breadth of
cornea about equal to 0.5 the length of
the carapace; preserved specimens yel-
lowish white or pink.

Odontodactylus hanseni

II. Dactylus of raptorial claw without teeth
on inner margin; telson with concentric
rings of flesh-tipped spines; rostrum
trispinous; eyes small, breadth of cornea
about equal to 0.125 the length of the
carapace; preserved specimens mottled

red or reddish brown.

Gonodactylus guerini

Squilla oratoria de Haan
Figs. 2, 'ia-k

Squilla oratoria de Haan, 1844: pi. 51, fig. 2
[figures only]; 1849: 223 [description].
Squilla ajjinis Berthold, 1845 : 26, pL 3, figs. 1-2.
Squilla nepa Miers, 1880: 25.

Chloridella ajjinis de Man, 1907 : 439-

description: Carapace considerably nar-
rower anteriorly than posteriorly; nearly twice
as long (including rostrum) as greatest breadth
anteriorly; conspicuous gastric and cervical
grooves, the latter continuous across mid-
dorsal region; seven distinct, longitudinal ca-
rinae, median bifurcated anteriorly for about
0.25 the distance to cervical groove, inter-
mediate extending nearly entire length of ca-
rapace, marginal produced anteriorly into sharp
spine and posteriorly rounded and reflexed

toward gastric grooves. Rostrum longer than
wide, subquadrate, with small median spine,
lateral margins convergent anteriorly, not
covering ophthalmic somite. Eyes large; cor-
nea bilobed, set transversely on stalk; anterior
margin of ophthalmic somite slightly truncate
between base of eyestalks (Fig. '^d), Merus
of raptorial claw articulating terminally with
ischium, ventral surface longitudinally
grooved throughout its length for reception
of propodus; propodus finely pectinate along
upper margin and with four sharp movable
spines; dactylus not inflated at base, with six
long, sharp teeth on inner margin. Propodus
of first, third, and fourth thoracic appendages
nearly circular, that of fifth subcircular, being
slightly longer than wide (Fig. 3/, h, i).
Mandibular palp of three segments (Fig. 3^,
c). Free thoracic and abdominal somites dor-
soventrally flattened, all possessing longitu-
dinal carinae (first five abdominal somites

Fig, 2. Squilla oratoria de Haan (male).

Hawaiian Stomatopods — TowNSLEY

405

Fig. 3. Squilla oratoria de Haan. a. Anterior segments of adult; b, right pars molaris of mandible and mandibular
palp (ventral aspect); c, right pars molaris of mandible (dorsal aspect); d, right pars incisiva of mandible; e, right
maxilla; f,g, right first and second thoracic appendages; h, right third and fourth thoracic appendages; right
fifth thoracic appendage;/, telson and uropods; k, right first abdominal appendage of male.

each with eight carinae) ; fourth through
eighth thoracic somites exposed; intermediate
carinae and lateral carinae of abdominal som-
ites 1-5 end in short, sharp spines; sixth ab-

dominal somite with six spines, two sub-
median, two intermediate, two lateral. Telson
with a single median crest, with radiating
series of pits on each side; distal margin armed

406

PACIFIC SCIENCE, Vol. VII, October, 1953

with three pairs of large immovable teeth,
submedian, intermediate, and lateral; three to
five submedian denticles, seven to nine inter-
mediate denticles, and a single lateral denticle,
also a pair of prelateral denticles. Exopod of
uropods with nine movable spines on outer
margin. Basal portion of uropod elongated
into two spines, inner longer than outer, a
small lobe near the outer edge of the inner
spine (Fig. 3/). Accessory reproductive organ
of male shown in Figure 'bk. The dorsal sur-
face of the specimens is somewhat pitted and
a dull mottled tan in color, the ventral surface
is highly polished and somewhat lighter in
color. Specimens vary from 145 to 200 mm.
in length from the anterior end of the rostrum
to the extremity of the submedian spines of
the telson.

DISCUSSION: Several varieties of this species
are recognized, but the Hawaiian specimens
seem to fit the characteristics given by Kemp
(1913) for those specimens which he had ex-
amined from this region. He found that spec-
imens from Hawaii did not seem to fit any of
the described varieties and did not attempt to
place them in any definite variety.

Three males and three females collected at
Oahu were examined. All were collected in
regions with a muddy bottom suitable for
burrowing. The Bishop Museum has eight
specimens, two females collected at Guam,
and two males and four females bought at a
Honolulu market. All were very similar in
appearance, except for small variations in col-
or and size; characteristics which do not seem
to be of systematic importance. There seems
to be no apparent sexual dimorphism. The
only positive means found for distinguishing
males from females was the penis at the base
of the eighth thoracic appendage, and the ac-
cessory reproductive organ of the male found
on the first abdominal appendage (Fig. 3^).

This species is commonly taken in crab nets
by fishermen at the Ala Wai Canal, Honolulu.
Occasionally they appear in the Honolulu fish
markets, their large size making them desir-
able for food.

DISTRIBUTION: This species appears to be
somewhat more restricted in distribution than
are other stomatopods, being reported only
from the Hawaiian Islands, Guam, Philip-
pine Islands, inland waters of Japan, and the
China Sea.

Squilla boops Kemp

Figs. 4, 5rf-/

Squilla boops Kemp, 191I [May]: 97.

Squilla quadraticauda Fukuda, 19 H [August]:

287, pi. 11, figs. 3-5.

DESCRIPTION: Carapace broad anteriorly,
breadth behind anterolateral spines exceeds ||
0.5 the total length including the rostrum; ij
conspicuous gastric and cervical grooves, the !
latter continuous across the middorsal region; |
seven distinct longitudinal carinae, median j
discontinuous anterior to cervical groove, end- |j
ing in a small pit, marginal produced anterior- ||
ly into a short spine and posteriorly rounded j
and reflexed toward gastric grooves. Rostrum li
broader than long, lateral margins upturned, ||
rounded and slightly convergent anteriorly, i!
a small median carina. Ophthalmic somite
exposed, anterior margin truncate between
eyestalks. Eyes large; cornea distinctly bi-
lobed, set obliquely on stalks, considerably |

more than 0.3 length of carapace; prominent j

lobe on external aspect of eyestalk. Ischio- |

meral articulation of raptorial claw terminal; j

ventral surface of merus longitudinally j

grooved throughout its length for reception |

of propodus; carpus with three carinae; pro- Ij

podus finely pectinate along upper margin |

and with two movable spines proximally; j

dactylus not inflated at base, with a small
ventral tubercle, five teeth on inner margin j

(Fig. 5^). Propodus of third and fourth thor- ij

acic appendages rounded posteriorly, those !

of first and fifth nearly straight and parallel ||

to anterior margin (Fig. '^a.c-e). Mandibular j|

palp composed of three segments, similar to j

that of 5. oratoria (Fig. 3^) . Free thoracic and i|

abdominal somites dorsoventrally depressed, |{

increasing in breadth posteriorly; all possess- ||

Hawaiian Stomatopods — Townsley

407

ing longitudinal carinae; fourth through
eighth thoracic somites exposed, lateral mar-
gin of fifth produced into an anteriorly di-
rected spine and a rounded posterior lobe,
lateral margin of sixth of two rounded lobes,
seventh and eight rectangular; first five ab-
dominal somites with eight carinae, the mar-
ginal and lateral ending in a short acute spine,
all intermediates except that of the first somite
ending in spines, submedian of fifth ending
in spines; sixth abdominal somite with six
carinae, submedian, intermediate, and lateral
all ending in spines. Telson with a single
median crest terminating in a short spine;
radiating series of pits on each side of median
crest; distal margin armed with three pairs of
immovable teeth, submedian, intermediate,
and lateral; three to five submedian denticles,
seven to nine intermediate denticles, one lat-
eral denticle, also a pair of prelateral denticles.
Exopod of uropods with seven movable spines
along outer margin. Basal portion of uropods
elongated into two spines, inner longer than

Fig. 5. Squilla hoops Kemp. a,b,c,d,e, Right first,
second, third, fourth, fifth thoracic appendages; /,
right first abdominal appendage of male.

outer, a small rounded lobe between the two
(Fig. 4). Accessory reproductive organ of
male shown in Figure 5/. Kemp mentions
that the type had the median portion of the
carapace, abdomen, and telson covered with
small, gray chromatophores. These were not
sufficiently abundant to detract from the gen-
eral coloration, which was yellowish in his
type specimen but faintly pink in the spec-
imens at hand. The posterior half of the fifth

408

PACIFIC SCIENCE, Vol. VII, October, 1953

abdominal somite between the lateral and
marginal carinae and the outer angles of the
sixth somite are black in all three of these
specimens, as well as in Kemp’s type spec-
imen. No mention of the color of Fukuda’s
specimen was made. The three local spec-
imens vary from 68 to 75 mm.; Kemp’s was
89 mm., and Fukuda’s 40 mm.

DISCUSSION: One male and two female spec-
imens of this species have been examined.
They were taken from the stomach contents
of black skipjack, Eiithynnus yaito Kishinouye,
caught off Moku Manu [Island] off the north-
east coast of Oahu. One specimen, a male,
was but slightly digested, so the characteris-
tics given by Kemp for this species could be
easily recognized.

5. hoops can be readily distinguished from
S. oratoria by its smaller size and by the pres-
ence of only five teeth on the raptorial dac-
tylus and only seven spines on the outer
margin of the uropod. Early in this study it
was thought that these were small specimens
of S. oratoria, but the differences noted above
have shown them to be the adults of Kemp’s
Squilla hoops.

Kemp and Fukuda’s specimens were both
females; thus previously no description has
been made of the modified accessory repro-
ductive organ of the male found on the first
abdominal appendage. However, Kemp’s
(1913) figures of the female have been useful
in identifying the present specimens, which
I consider conspecific.

DISTRIBUTION: The type specimen, a female,
was taken by the "Investigator” in the Gulf
of Martaban, Burma, at a depth of 67 fathoms.
Another female was collected at Matsuwa,
Sagami Province, Japan. One male and two
females have been obtained from the stomach
contents of a black skipjack {Eiithynnus yaito')
caught off Moku Manu (Bird Island), Oahu.

Squilla alba Bigelow
Figs. 6, la-f

Squilla alhaEigdow, 1893: 103; 1894: 539-541,

pi. 22. [Complete description and figures.]

DESCRIPTION: Carapace longer than great-
est breadth, slightly narrower anteriorly than
posteriorly, with distinct gastric and cervical
grooves, with five longitudinal carinae, an-
terolateral angles rounded and usually armed
with short acute spines, posterolateral angles
broadly rounded. Rostrum longer than broad,
truncate or triangular. Ophthalmic somite par- i
daily exposed, anterior margin straight be- j
tween base of eyestalks, with small rounded
process at base of each eye dorsally. Eyes I
large; cornea set transversely on stalks, only
slightly broader than greatest breadth of stalk,
about 0.3 total length of carapace, including
rostrum. Antennular peduncle about same jl
length as carapace, including rostrum. Ischio- 1|
meral articulation of raptorial claw terminal; j'
ventral surface of merus grooved longitudin- |j
ally throughout its length for reception of
propodus; carpus with a blunt median carina; ||
outer margin of propodus finely pectinate, jj
three movable spines near proximal end (only '
one shown in Fig. ih, other two not visible
in this view); dactylus not inflated at base, |
notched inferiorly, outer margin straight or
slightly concave, inner margin armed with
six teeth. Anterior and posterior margins of j
third and fourth thoracic appendages round-
ed, first and fifth straight and nearly parallel
(Fig. la, c-e). No mandibular palp. Fifth
through eighth thoracic somites exposed; lat-
eral margin of fifth seen in dorsal view com-
posed of two distinct processes, a long, an-
teriorly directed spine and a rounded posterior
lobe; sixth, seventh, and eighth not bilobed, j
but rounded. Longitudinal carinae on all ex- ;
posed somites, eight on first five abdominal^ ||
somites, none ending in spines; posterolateral J
margins acute and rounded; sixth abdominal ||
somite with six carinae ending in spines, two j
submedian, two intermediate, two lateral. Tel- ‘ jj
son with indistinct median carina but a small j
median spine posteriorly; radiating series of
pits on dorsal surface; armed with six mar- [
ginal teeth, two submedian with movable tips, ]
two intermediate, two lateral; five or six min- f
ute submedian spinules, 11 or 12 intermediate, |

Hawaiian Stomatopods — Townsley

409

one lateral. Exopod of uropods with five or
six movable spines along their outer margin.
Basal portion of uropods elongated into two
spines, inner longer than outer, with a dis-
tinct, rounded lobe near base of each spine
(Fig. 6). First abdominal appendage shown
in Figure if. Body of specimens an opaque
white, eyestalks yellow with a very black
cornea. Abdominal and thoracic somites and
carapace with large, white pigment cells dis-
tributed at random over their surface. Telson,
seen from above, nearly transparent so that
tissue projecting into it can be easily seen and
produces the pattern shown in Figure 6. Spec-
imens range in size from 20 to 47 mm. in
length.

DISCUSSION: A small female specimen, 20
mm. long, was collected by Mr. Kenji Ego

Fig. 7. Squilla alba Bigelow. a,b,c,d,e. Right first,
second, third, fourth, fifth thoracic appendages; /,
right first abdominal appendage of female.

at Kawaihae, Hawaii, in a night light station.
This is a very striking species, as the color
of the specimens immediately attracts atten-
tion. Its shape is also somewhat peculiar, the
carapace and exposed portion of the thorax
being equal in length and together making
up nearly 0.44 of the total length of the body.
The entire surface of the body is smooth and
polished, interrupted occasionally by distinct
carinae. The lateral margins of the fifth thor-
acic somite with their long, anteriorly directed
spines and rounded posterior lobes are also
characteristic of this species.

410

PACIFIC SCIENCE, Vol. VII, October, 1953

The type specimen was collected by Bige-
low at Bimini Harbor, Bahamas. Edmondson
(1921) reported that a female specimen was
taken on Waikiki Reef, Oahu, Unfortunately,
I have not been able to locate this specimen
in the Bishop Museum for comparison with
the specimen taken at Kawaihae. Edmondson
states that the specimen which he collected
agreed with Bigelow’s previous description
for the species. The Kawaihae specimen agrees
with Bigelow’s description in color, shape of
the eyes, general body proportions, shape of
the lateral margin of the fifth thoracic somite,
shape of the telson, and the presence of two
lobes between the inner and outer spines of
the uropods. However, there are certain char-
acteristics in which the specimens differ con-
siderably. Bigelow’s type possesses definite
longitudinal carinae on the carapace, the ros-
trum is truncate, the anterolateral angles of
the carapace are rounded and armed with a
short spine, and the outer margin of the uro-
pods is armed with six movable spines. The
Kawaihae specimen, on the other hand, does
not have any carinae on the carapace, the
rostrum is distinctly triangular, the antero-
lateral angles of the carapace are rounded and
unarmed, and the outer margin of the uropods
is armed with only five movable spines. I
believe that these differences are due to the rel-
atively immature condition of the Kawaihae
specimen, and that it is probably the last
postlarval stage of this species. It is possible
that the subsequent molt would reconcile the
differences between this and the Bimini spec-
imen.

Bigelow makes little note of the submedian
spines on the posterior margin of the telson.
His figure (1894: pi. 22) is indistinct, but
there appears to be a line indicating that they
possess movable tips. In the Kawaihae spec-
imen the tips of these spines are unquestion-
ably movable.

No male specimens have been reported.
The largest of Bigelow’s specimens was 41
mm., Edmondson’s was 47 mm., and the Ka-

waihae specimen is considerably smaller, be-
ing only 20 mm. in length.

DISTRIBUTION: The type specimen was col-
lected in calcareous sand at Bimini Harbor,
Bahamas. In the Hawaiian Islands one spec-
imen was taken from dead coral on Waikiki
Reef, Oahu, and another at Kawaihae, Hawaii,
at a night light. These are the only records
known for this species.

SquiUa calumnia n. sp.

Figs. 8, <)a-f

DESCRIPTION: Carapace, excluding rostrum, jl
about as long as greatest breadth; slightly
narrower anteriorly than posteriorly; distinct j;
gastric grooves and cervical groove; median, ij
intermeiate, lateral, and marginal longitu- |!
dinal carinae present, median discontinuous
and bifurcated anterior to cervical groove, ||
lateral posteriorly reflexed toward gastric jl
groove; anterolateral and posterolateral angles i
unarmed and rounded. Rostrum broader than j
long, truncate, small median carina, lateral j!:
margins upturned and rounded anteriorly. I
Ophthalmic somite exposed, anterior margin
concave between base of eyestalks. Eyes large;
cornea set transversely on stalks, only slightly
broader than greatest breadth of stalk, about
0.25 total length of carapace, including ros-
trum. Antennular peduncle about same length
as carapace, excluding rostrum. Ischio-meral
articulation of raptorial claw terminal; ventral
surface of merus longitudinally grooved
throughout its length for reception of pro-
podus; carpus with median and lateral carinae;
propodus finely pectinate along outer surface,
three movable spines near proximal end; dac-
tylus not inflated at base, outer margin slight-
ly concave, armed with six teeth. Anterior
margin of propodus of third, fourth, and
fifth thoracic appendages straight (Fig. 9c-e),
posterior margins rounded; propodus of first
thoracic appendage nearly circular (Fig. ^d).
Mandibular palp composed of two segments.
Fifth through eighth thoracic somites ex-
posed, with longitudinal carinae; lateral mar-

Hawaiian Stomatopods — • Townsley

411

gin of fifth, seen in dorsal view, composed of
two distinct processes, anteriorly directed spine
and nearly straight posterior spine; lateral mar-
gin of sixth nearly straight, angular; seventh
rounded posteriorly. Eight longitudinal cari-
nae on first five abdominal somites, subme-
dian, intermediate, lateral, marginal; all mar-
ginal carinae terminating in a short acute
spine; sixth abdominal somite with six cari-
nae, all ending in spines, two submedian, two
intermediate, two lateral. Telson with median
Carina, ending in a small median spine pos-
teriorly; radiating series of pits on each side

Fig. 9- Squilla calumnia n. sp. a,b,c,d,e. Right first,
second, third, fourth, fifth thoracic appendages; /,
right first abdominal appendage of female.

of median carina; armed with six teeth, two
submedian, two intermediate, two lateral;
four submedian denticles, seven intermediate
denticles, one lateral denticle. Outer margin
of exopod of uropods with eight movable
spines on one side, nine on the other. Basal
portion of uropods elongated into two spines,
inner longer than outer, distinct rounded

412

PACIFIC SCIENCE, VoL VII, October, 1953

lobe near base of inner. First abdominal ap-
pendage shown in Figure 9/ The coloration
of the specimen is sufficient to distinguish it
from others which I have examined. The
ground color is an opaque white, which is
partially obscured by dark-brown chromato-
phores along the posterior border of the thor-
acic and abdominal somites and which also
clearly mark all the longitudinal carinae of
the body. The cornea of the eyes is a deep
black, the eyestalks retaining the opaque
white of the body. Specimen 26 mm, long.

TYPE locality: a single male specimen
26 mm. long was collected by the Territorial
Division of Fish and Game at a night light
station at Hilo Dock, Hilo, Hawaii.

DISCUSSION: Although the penis which is
located at the base of the eighth thoracic
appendage is present, the specimen is ap-
parently immature, as the accessory repro-
ductive organ on the endopod of the first
abdominal appendage is undeveloped. In
spite of this condition I feel that the other
characteristics are sufficiently distinct for the
separation of this specimen from the other
species in this genus.

At first glance this species and Squilla alba
appear to be very similar; but the immovable
submedian spines along the outer margin of
the telson, the more numerous spines along
the outer margin of the uropods, the shape
of the lateral margin of the fifth thoracic som-
ite, and the single lobe at the base of the
inner spine of the uropod will serve to dis-
tinguish this species from 5. alba. The round-
ed anterolateral angles of the carapace, the
large globular eyes, and the six teeth on the
raptorial dactylus are shared by both these
species and will serve to separate them from
the other members of this genus.

The fifth thoracic somite in this specimen
appears to be composed of two distinct proc-
esses when seen from above, but it is different
from others showing this same condition. In
others, e.g., S. laevisHtss (1865), 5. hieroglyph-
ica Kemp (1911), and 5. alba Bigelow (1893),
there is a spine directed anteriorly and a

rounded posterior lobe. Here both processes
are distinctly angulaq the anterior spine
curved forward, the posterior one projecting
laterally. The shape of the sixth thoracic som-
ite is distinctive in this species as well, in
that the others have rounded lateral margins
or are distinctly bilobed.

TYPE SPECIMEN: U.S.N.M. 93097.

Pseudosquilla ciliata (Fabricius)

Figs. 10, Wa-rn

Squilla ciliata Fabricius, 1787: 333.

Cancer ciliatus\Am\2,^\i^, 1790: 2990.

Cancer {mantis) [sic] c/7/^toj- Herbst, 1796: 102.
Squilla stylifera 1818: 189.

Squilla quadrispinosa Eydoux and Souleyet,

1841: 262, pi. 5, fig. 1.

Pseudosquilla stylifera Dana, 1852: 622, pi. 12,

figs. 4a-e.

DESCRIPTION: Carapace longer than great-
est breadth, slightly narrower anteriorly than
posteriorly with conspicuous gastric grooves,
cervical groove wholly absent, lacking cari-
nae, anterolateral and posterolateral angles
unarmed and rounded. Rostrum rounded an-
teriorly, about as wide as long, covering the
ophthalmic somite. Eyes tubular, cornea fol-
lowing long axis of stalk (Fig. 11^). Merus
of raptorial claw articulating terminally with
ischium, ventral surface grooved throughout
its length for reception of propodus; propo-
dus with fine pectinations along outer margin
and with four movable spines near proximal
end; dactylus with three (sometimes four)
long, sharp teeth. Propodus of fifth thoracic
appendage subcircular, slightly longer than^
wide (Fig. Hi). Mandibular palp of three
segments. Free thoracic and abdominal som-
ites compressed, all lacking longitudinal cari-
nae; sixth, seventh, and eighth thoracic som-
ites exposed; posterolateral angles of abdom-
inal somites with small, sharp spines; sixth
abdominal somite with four spines, two sub-
median, two intermediate. Telson with seven
carinae, one median, two submedian, two

Hawaiian Stomatopods — TOWNSLEY

/

413

intermediate, two lateral; armed with six
spines, two submedian with movable tips,
two intermediate, two lateral; no submedian
denticles, two intermediate denticles, single
lateral denticle. Exopod of uropods with nine
(sometimes ten) movable spines on outer
margin. Basal portion of uropod elongated
into two spines, inner longer than outer. Ac-
cessory reproductive organ of male shown in
Figure 11/. Specimens vary from 35 to 85
mm: in length.

DISCUSSION: Fifty-eight specimens of this
well-known species were examined, of which
23 were males and 35 were females. All were
collected from the reefs around Oahu in water
not over 2 fathoms deep. This species repre-
sents the dominant form of the Hawaiian
Islands. It has been collected from both Kauai
and Hawaii where it seems to be as abundant
as it is at Oahu. The .species appears to prefer
the rough substrate of the large exposed reef
flats such as those found in Kaneohe Bay,
Oahu. It has not been found in those regions
having a substrate of coarse shifting sand or
strong wave action.

There appears to be no clearly defined sex-
ual dimorphism, and the only definite means
that I have found for distinguishing males
from females lies in the penis found at the
base of the eighth thoracic leg (Fig. lly) and
the accessory reproductive organ of the male
found on the first abdominal appendage (Fig.
11/). Specimens exhibit all degrees of coloring
from an olive drab through yellow to a light
tan with bright red, blue, and yellow chroma-
tophores. The dactyli of the thoracic append-
ages are, in most cases, light rose.

According to Brooks (1886) the Atlantic
specimens of this species differ in certain small
details from those v/hich are found in the
Indo-Pacific region. Borradaile (1899) named
the Atlantic specimens var. occidentalism but
later workers such as Tattersall (1906) and
Kemp (1913) have shown that the characters
used by Borradaile are to be found in the
Indo-Pacific specimens as well. The speci-
mens from the Hawaiian Islands also appear

Fig. 10. Pseudosquilla ciliata (Fabricius) (male).

to^be quite variable, but this variability is
not sufficient to warrant their being placed
in any other group. I have examined three
specimens from Kaneohe Bay, Oahu, in
which there were ten spines rather than nine
on the exopod of the uropod; and one spec-
imen in which there were nine spines on one
side and ten on the other. Two specimens in
the collection have the outer spine of the basal
prolongation of the uropod longer than the
inner. This is a character which has been con-
sidered to be specific for P. oculata (Brulle),
but all other characters of this specimen are
those of P. ciliata.

Specimens vary in length from 38 to 85
mm. Generally this species is somewhat larger
than its congener, P. oculatam and in the field
they can usually be distinguished in this way.

DISTRIBUTION: There are 12 species of
Pseudosquilla, nine of which are found in the
Indo-Pacific region. P. ciliata and its Hawaiian

Fig. 11. Pseudosquilla ciliata (Fabricius). a, Anterior segments with rostrum removed; b, right antenna with scale;
c, right pars molaris of mandible and mandibular palp; d, right pars incisiva of mandible; e, right maxilla;/,g, right
first and second thoracic appendages; right third and fourth thoracic appendages;/, right fifth thoracic appendage;
y, eighth thoracic appendage of female (top) and male showing penis (bottom) ; kj, right first abdominal append-
ages of female and male; m, telson and uropods.

Hawaiian Stomatopods — ToWNSLEY

415

relative, P. oculata, are both known from the
Pacific, Indian, and Atlantic oceans. Ordina-
rily they are found in the intertidal area in
these regions, but Bigelow (1891) reports
that P. ciliata was collected at a depth of
20-23 fathoms. P. ciliata has been reported
from the Hawaiian Islands, Fiji Islands, So-
lomon Islands, Loyalty Islands, Japan, New
Britain, New Guinea, Red Sea, Madagascar,
Bermuda, Florida Keys, Puerto Rico, and St.
Thomas.

Pseudosquilla oculata (Brulle)

Figs. 12, 13

Squilla oculata Brulle, 1836: 18, fig. 3.

Squilla momdactyla A. Milne-Edwards, 1878:

232.

Pseudosquilla oculata Miers, 1880: 110, pi. 3,

figs. 3, 4.

Pseudosquilla momdactyla Miers, 1880: 110, pi.

3, figs. 1, 2.

DESCRIPTION: Carapace longer than great-
est breadth, slightly narrower anteriorly than
posteriorly, with conspicuous gastric grooves,
cervical groove wholly absent, lacking carinae,
anterolateral and posterolateral angles un-
armed and rounded. Rostrum with small me-
dian spine, about as wide as long, covering
ophthalmic somite. Eyes flattened, appearing
somewhat club-shaped rather than tubular
(compare Figs. \\a and 12), cornea set trans-
versely on stalk. Merus of raptorial claw ar-
ticulating terminally with ischium, ventral
surface grooved longitudinally throughout its
length for reception of propodus; propodus
with fine pectinations along upper surface and
with four movable spines near proximal end;
dactylus with three long, sharp teeth. Propo-
dus of fifth thoracic appendage subcircular,
slightly longer than wide. Mandibular palp
of three segments. Free thoracic and abdom-
inal somites compressed, all lacking longi-
tudinal carinae; fifth, sixth, seventh, and
eighth thoracic somites exposed; postero-
lateral angles of abdominal somites with
small, sharp spines; fifth abdominal somite

with two sublateral spines; sixth abdominal
somite with two submedian and two sub-
lateral spines. Telson with nine carinae, one
median, two submedian, two intermediate,
two sublateral, two lateral; armed with six
spines, two submedian with movable tips,
two intermediate, two lateral; no submedian
denticles, two intermediate denticles, single
lateral denticle. Exopod of uropods with ten
(sometimes 11) movable spines along outer
margin. Elongated basal portion of uropod
with outer spine longer than inner (Fig. 12).
Accessory reproductive organ of male shown
in Figure 13. All specimens were a mottled
tan and all possessed a dark spot on the lateral
edges of the carapace, as noted by Kemp
(1913). The dactyli are tipped with rose. Spec-
imens vary from 35 to 70 mm. in length.

DISCUSSION: This species superficially re-
sembles the more common P. ciliata. In Ha-
waiian waters the two species are commonly
found in the same habitat, and, except through
close scrutiny, P. oculata may often be mis-
taken for P. ciliata. The eyes of this species
are very characteristic in shape and contrast

416

PACIFIC SCIENCE, Vol. VII, October, 1953

Fig. 13. Pseudosquilla oculata (Brulle), right first ab-
dominal appendage of male.

well with those of P. ciliata (cf. Figs. 11^ and
12). The spine-tipped rostrum, the extra pair
of carinae on the telson, and the longer outer
spine of the uropod will all serve to place
specimens in this species.

This species is common in Hawaii: the
Bishop Museum collection has live males and
six females, and in addition I have examined
one male and four females. The largest spec-
imen was a male 70 mm. long, and the small-
est was 35 mm. long. All specimens were
collected at Oahu in shallow water together
with specimens of P. ciliata.

Variability in color and form does not seem
to be pronounced in this species. One spec-
imen has 11 spines on the outer margin of the
exopod of one uropod and 10 on the other.
This variation parallels the condition found
in P. ciliata and indicates that this chracteristic
is quite variable.

There is no clear sexual dimorphism. The
accessory reproductive organ on the first ab-
dominal appendage of the male is shown in
Figure 13. If it is compared with Figure 11/,
it can be seen that both P. oculata and P. ciliata
are very similar with respect to this structure.
Possibly this indicates strong affinities.

DISTRIBUTION: This species and P. ciliata

have been found in the Hawaiian Islands. As
mentioned previously, P. oculata is found in
the same environment as P. ciliata. Specimens
are recorded from Samoa and Hawaii in the
Pacific, from the Macclesfield Bank in the
South China Sea, from Mauritius in the In-
dian Ocean, and from the Cape Verde Islands
and Madeira in the Atlantic.

Lysiosquilla maculata (Fabricius)

Figs. 14, 15^-g !

Squilla arenaria terrestris or locusta Rumphius, I

1705: 4, pi. 3, fig. E. ;

Squilla maculata Fabricius, 1793: 511. ;

Cancer {mantis) [sic] arenarius Herbst, 1796: |

96, pi. 33, fig. 2.

Squilla arenaria Randall, 1839: 146. j

DESCRIPTION: Carapace nearly as broad as |
long, anterior breadth equal to posterior !
breadth, smooth, strongly convex from side I

to side, gastric grooves distinct, cervical jj

groove obsolete or wholly absent, no longi- i
tudinal carinae, anterolateral and postero- jj
lateral angles broadly rounded. Rostrum j
broader than long, smooth, lateral margins I
convergent anteriorly to an acute but blunt
apex, small longitudinal carina on anterior
one third. Ophthalmic somite anteriorly ex- j
posed, dorsally produced into a pair of lobes i
terminating acutely on each side of rostral |i
apex. Eyes large; cornea set obliquely on |!
stalk, breadth about equal to greatest length
of cornea and stalk combined. Antennular I
peduncle with small acute dorsal spine, less I
than one-half length of carapace, excluding |
rostrum. Ischio-meral articulation of raptorial j!
claw terminal; merus grooved ventrally
throughout its length for reception of pro-
podus; carpus with blunt dorsal carina ter-
minating in a sharp spine which overhangs
anterior margin; propodus finely pectinate on
upper margin, four movable spines near prox-
imal end; dactylus slender, 9 to 11 teeth
(usually 10), outer margin straight or slightly

Hawaiian Stomatopods — Townsley

417

concave (Fig. 15^/). Propodus of third and
fourth thoracic appendages orbicular with two
weak spines at proximal end, those of first
and fifth nearly rectangular and lacking spines
(Fig. 15c,/). Fourth through eighth thoracic
somites exposed, lateral margins rounded,
lacking longitudinal carinae. Abdominal so-
mites depressed ; first through fifth lacking lon-
gitudinal carinae, increasing in breadth pos-
teriorly; sixth narrower than preceding so-
mites, with distinct grooves on each side of
mid-line. Telson broader than long, convex
dorsally, concave ventrally, both sides marked
by a feeble median triangular elevation lim-
ited by a pair of posteriorly convergent

Fig. 15. Lysiosquilla maculata (Fabricius). a. Right
pars molaris of mandible and mandibular palp; b, right
pars incisiva of mandible; c,d, right first and second
thoracic appendages; e, right third and fourth thoracic
appendages; /, right fifth thoracic appendage; g, right
first abdominal appendage of male.

418

PACIFIC SCIENCE, VoL VII, October, 1953

grooves; sometimes marked by patches of
pits rather regularly disposed near lateral mar-
gins; posterior margin indistinctly notched
in mid-line with three pairs of blunt lobes,
the two outermost generally sharper and more
conspicuous than the inner. Inner dorsal edge
of peduncular segment of uropods ending in
short spine; exopod of uropods with eight
or nine movable spines; basal portion of uro-
pod elongated into two long spines, inner
nearly twice length of outer, but obscured
dorsally. Modified first abdominal appendage
of male shown in Figure 15g. Animals are
marked by alternating series of light and dark
bands throughout the length of the body.
Specimens vary in length from 69 to 283 mm.

DISCUSSION: This species is very readily
identified. All specimens which have been
hitherto reported, as well as those which I
have examined, have possessed the character-
istic dark transverse bands shown in Figure
14. In large female specimens this appears to
be a secondary sexual characteristic, inasmuch
as the carapace, free thoracic somites, and
sixth abdominal somite are almost entirely
blue-black. Nearly always in males there are
alternating dark and light bands with the tel-
son having a pattern which may vary from
that shown in the figure but which usually
has dark central and lateral areas. Preserved
specimens retain the coloration, but the dark
pigment becomes a brownish black.

There appear to be other characteristics
showing a sexual dimorphism in this species.
Miers (1880) and Kemp (1913) record several
large females in which the carpus and the
basal part of the raptorial propodus bear tufts
of long hairs, and the latter with only two
movable spines at its proximal end instead of
four. The raptorial dactylus in these cases is
also reported by them to have only a series
of eight or nine denticles rather than the long
teeth usually found there; however, none of
the specimens which I have examined has
shown these modifications. Evidently this is
due to the relatively small size of the female
specimens which I have examined compared

with those which they report as having these
modifications.

This species and Squilla oratoria are the two

largest stomatopods found in the Hawaiian
Islands. I have examined six males and three
females ranging from 70 to 185 mm. from
the tip of the rostral spine to the posterior
border of the telson. Other authors have re-
ported specimens ranging from 69 to 283 mm.

Lysiosquilla maculata seems to occur in shal-
low water with a sandy or muddy substrate |
where it may burrow easily. All those spec- |
imens which I have examined had been col- '

lected in the fine silt on the reefs at Oahu and |
Hawaii. Mr. Kenji Ego (personal communi- ]
cation) reports that they occur in abundance '
in the mud flats at Kawaihae, Hawaii. I

DISTRIBUTION: This species is widely dis- '
tributed throughout the Indo-Pacific. In the
Pacific it has been reported from the Hawaiian |
Islands, Marquesas Islands, Samoa, Fiji, Phil- |
ippine Islands, and the Dutch East Indies, j

Fig. 16 Coronida sinuosa Edmondson (female).

Hawaiian Stomatopods — TowNSLEY

419

In the Indian Ocean it has been reported from
Penung, Tuticorin, Madras, Mysore, and Dur-
ban. In the Atlantic Ocean adult specimens
have been collected at Antigua, Dutch West
Indies, and larvae from near the Cape of Good
Hope.

Coronida sinuosa Edmondson
Figs. 16, \la~e

Coronida sinuosa 1921: 295, fig. 2.

DESCRIPTION: Carapace very slightly nar-
rower anteriorly than posteriorly, nearly twice
as long (including rostrum) as breadth at an-
terior border, lacking distinct carinae, cervical
and gastric grooves wholly absent, antero-
lateral and posterolateral angles slightly round-

ed. Rostrum small, rounded anteriorly, breadth
greater than length, not covering ophthalmic
somite. Cornea of eyes set transversely on
eyestalks, slightly bilobed; eyestalks narrow,
elongated, flattened. Merus of raptorial claw
articulating terminally with ischium (Fig. lib),
ventralsurfacelongitudinallygroovedthrough-
out its length for reception of propodus; pro-
podus finely pectinate along upper margin;
dactylus notched and slightly inflated at base,
with four sharp teeth on inner margin. First,
third, fourth, and fifth thoracic appendages
shown in Figure 17^, c, d. Mandibular palp
absent. Free thoracic and abdominal somites
dorsoventrally depressed, all lacking longi-
tudinal carinae; fifth through eighth thoracic
somites exposed, lateral margins of fifth with

420

PACIFIC SCIENCE, Vol. VII, October, 1953

an acute, anteriorly directed process, sixth,
seventh, and eighth narrowed but evenly
rounded (Fig. 16); dorsal surface of sixth
abdominal somite ornamented with a scroll-
like series of tubercles (carinae.^), nearly sym-
metrical in arrangement. Telson with a series
of similar tubercles which may be unsym-
metrical except in the median region (Fig.
16); posterior border medially notched giving
the appearance of two rounded lobes, with a
pair of submedian spines and four to six mar-
ginal spines lateral to each submedian. Exo-
pod of uropods with 8 to 11 movable spines
on outer margin; elongated basal portion of
uropods with two spines, inner longer than
outer. First abdominal appendage of female
shown in Figure 17^. Living animals are trans-
parent white, preserved specimens opaque
white throughout. Specimens range from 9-5
to 17.0 mm. in length.

DISCUSSION; Three female specimens of this
small species have been examined, including
the type specimen in the Bishop Museum.
All had been collected from among dead coral
on Waikiki Reef, Oahu.

The species is apparently closely allied to
Coronida fmdtituherculata (Borradaile) (1898),
but differs considerably in the shape of the
rostrum and in the ornamentation of the sixth
abdominal somite and telson.

Specimens may be easily recognized by the
ornamentation of the last two body segments.
The tubercles (or carinae as they are called by
Edmondson) of the sixth abdominal somite
form a definite pattern which is nearly sym-
metrical on each side of the median line,
suggesting a single median carina, paired sub-
median carinae, paired intermediate carinae,
paired lateral carinae, and paired marginal ca-
rinae. The telson is similarly ornamented, but
is not usually as symmetrical as shown in
Figure 16. Here there is also a suggestion of
a median carina, paired submedian carinae,
and paired intermediate carinae; but all are
somewhat joined to one another forming a
scroll-like pattern. There appears to be con-
siderable variation in the number of spines

found on the posterior margin of the telson,
and I am inclined to believe that there is but
a single pair of submedian spines, the rest
being small, marginal denticles, although Ed-
mondson (1921) recognized two other pairs.

The slightly inflated base of the dactylus
suggests affinities to Gonodactylus and Odonto-
dactylus, but the articulation of the ischium
and merus of the raptorial claw indicates a
close relationship to Squilla, Pseudosquilla,
and Lysiosquilla.

Because of their very small size it can be
seen why this species has not been collected
more often. The three specimens which I have
examined ranged from 9.5 to 17.0 mm. from
the tip of the rostrum to the distal end of the
submedian spines of the telson. Living adults
are also very transparent and difficult to see.

Hawaiian Stomatopods — ToWNSLEY

421

especially so in their native habitat on the
reefs. Preserved specimens become opaque
white or slightly yellowish, with the cornea
of the eyes remaining very dark.

It is difficult to say whether or not the
specimens are mature and whether there is a
distinct sexual dimorphism, because all spec-
imens are females and none were ever collect-
ed with egg masses. The absence of the gastric
grooves and the very elongated antennal so-
mites suggests that possibly they are in an
immature stage. Edmondson (1921) states
that the specimen in the Bishop Museum was
kept alive for a month, during which time it
molted with no apparent increase in size or
change in structure.

DISTRIBUTION: All Specimens collected have
come from dead coral heads on Waikiki Reef,
Oahu.

Odontodactylus hanseni (Pocock)
Figs. 18, 19 a- f

Gonodactylus hansenii Pocock, 1893: 477, pi.

20, figs. 3, 36.

Odontodactylus latirostris hondiddilt, 1907: 212,

pi. 22, figs. 3, 3a.

DESCRIPTION: Carapace longer than great-
est breadth, slightly narrower anteriorly than
posteriorly, with distinct gastric grooves, cer-
vical groove wholly absent, longitudinal cari-
nae lacking, anterolateral and posterolateral
angles unarmed and rounded. Anterior margin
of rostrum evenly rounded, twice as broad as
long. Ophthalmic somite exposed, dorsal
process deeply excavate anteriorly. Eyes large,
globular, corneal diameter 0.25 to 0.5 the
length of carapace, excluding rostrum (Fig.
18). Ischio-meral articulation of raptorial claw
situated at a point in advance of proximal end
of merus (Fig. 19^) ; ventral surface of merus
grooved longitudinally for not more than 0.6
its length for reception of propodus; upper
surface of propodus not pectinate; dactylus
inflated at base, armed with 9-11 teeth on its
inner margin. Propodus of third, fourth, and

Fig. 19. Odontodactylus hanseni (Pocock). a,b,c,d,e,
Right first, second, third, fourth, fifth thoracic append-
ages; /, right first abdominal appendage of male.

fifth thoracic appendages somewhat shield-
shaped (Fig. I9c-e). Mandibular palp of three
segments. Free thoracic and abdominal so-
mites laterally compressed, distinct longitudi-
nal carinae present only on sixth abdominal
somite; sixth, seventh, and eighth thoracic
somites exposed, lateral borders rounded; pos-
terolateral angles of fourth and fifth abdominal
somites ending in a short acute spine; sixth
abdominal somite with four pairs of longitu-
dinal carinae, one submedian pair ending in
spines, one intermediate pair without spines,
one second lateral pair ending in spines, one

422

PACIFIC SCIENCE, VoL VII, October, 1953

marginal pair ending in spines. Telson with
high median carina ending in a short terminal
spine, submedian, intermediate, second lat-
eral, and marginal carinae; armed with three
pairs of strong marginal teeth, submedian
(with movable tips), intermediate, and lateral;
no submedian denticles but a series of minute
spinules, two intermediate denticles, and a
single lateral denticle. Exopod of uropods
with 10 or 11 movable spines on outer margin.
Basal portion of uropods elongated into two
spines, inner shorter than outer. First abdom-
inal appendage of male shown in Figure 19/
Color of the specimens is a uniform light
yellowish pink above with the exposed thor-
acic and first four abdominal somites having
a light pink posterior margin. The large glob-
ular eyes are dark brown and the flagella of
the antennae are pink. The animals are quite
small with the reported range in length from
27 to 80 mm.

DISCUSSION: In dorsal aspect this species
very closely resembles small specimens of
Psuedosquilla, but the ischio-meral articulation
of the raptorial claw and the large globular
eyes are suitable for distinguishing members
of this genus from those of Pseudosquilla. The
inflated dactylus and the ischio-meral articula-
tion of the raptorial claw are features which
members of this genus have in common with
Gonodactylus, but the numerous carinae on the
telson, the nonpectinate upper margin of the
raptorial propodus, and the series of teeth on
the inner margin of the raptorial dactylus will
serve to separate this group from the latter.
As stated by Bigelow (1894) this genus seems
to occupy an intermediate position between
Gonodactylus and Pseudosquilla.

Five female and three male specimens of
this species have been examined, all of which
were taken from stomachs of the skipjack,
Katsuwonus pelamis, and of the yellowfin tuna,
Neothunnus macropterus. The Bishop Museum
has one male and two females which were
dredged off Waikiki, Oahu, at a depth ranging
from 30 to 50 fathoms. Pocock’s (1893) type
specimen was a single female from the Mac-

clesfield Bank, South China Sea, at a depth
of 35 fathoms. Bigelow (1931) reported four
males, one female, and two juveniles taken
in Hawaiian waters by the "Albatross” in
1902 at depths ranging from 24 to 83 fathoms.
The specimens which I have examined are in
agreement with those previously reported.
Some characteristics were relatively difficult
to distinguish, but I attribute this to the fact
that the animals were considerably disfigured
from having been in the stomach. Edmondson
(1921) has illustrated the telson of this species
more clearly than is shown in Figure 18.

There appears to be no distinct sexual di-
morphism. The first abdominal appendage of
the male shown in Figure 19/ is incomplete
because the fixed limb in all three specimens
was indistinguishable.

Fig. 20. Gonodactylus guerini White (female).

Hawaiian Stomatopods — Townsley

423

Fig. 21. Gonodactylus guerini White, a. Right pars molaris of mandible and mandibular palp; b, c, right first and
second thoracic appendages; d, right third and fourth thoracic appendages; e, right fifth thoracic appendage;
/, left first abdominal appendage of male (endopod only).

Evidently this species is quite common in
deeper water because all other specimens have
been taken at considerable depths. Apparently
this species forms a considerable part of the
diet of the tunas, inasmuch as the adults and
larvae are often found in stomach contents.
This indicates that this species is probably
semiplanktonic, because the tunas feed in
mid-water or at the surface. These stomato-
pods certainly spend some time on the bot-
tom because other specimens have been
dredged from a rough substrate.

DISTRIBUTION: Specimens have been re-
ported from the Hawaiian Islands, the South
China Sea, the East Indian Archipelago, and
the Indian Ocean.

Gonodactylus guerini White

Figs. 20, 21^-/

Gonodactylus guerinii White, 1861^.* 43, pi. 7;
1861A 480.

Protosquilla guerinii Brooks, 1886: 75, pi. 16,
figs. 1, 6.

DESCRIPTION: Carapace rectangular, nearly
as wide as long, slightly narrower anteriorly,
anterolateral and posterolateral angles broadly

rounded, lacking distinct carinae, gastric
grooves well defined, cervical almost obso-
lete. Rostrum quadrate, trispinous, covering
ophthalmic somite. Eyes small, tubular; cor-
nea bilobed, set transversely on stalk. Ischio-
meral articulation of raptorial claw situated
at a point in advance of proximal end of merus
so that merus extends backward beyond the
joint (Fig. 21c); ventral surface of merus lon-
gitudinally grooved for not more than 0.75
its entire length for reception of propodus;
upper margin of propodus with a series of
fine acute teeth; dactylus inflated at base,
elongated into a single acute tooth, inner
margin of dactylus serrated. First, third,
fourth, and fifth thoracic appendages shown
in Figure 21^, d, e. Mandibular palp present,
three segments (Fig. 2\d). Free thoracic and
abdominal somites laterally compressed, all
lacking longitudinal carinae (indication of
slight marginal carinae on first five abdominal
somites); fifth through eighth thoracic so-
mites exposed, slightly narrower than posterior
width of carapace; posterior portion of fifth
abdominal somite bears several rows of short
spines, separated from smooth anterior part
by a curved transverse line; sixth abdominal

424

PACIFIC SCIENCE, Vol. VII, October, 1953

somite armed with numerous long spines,
each ending in a blunt, rounded tip from
which protrudes a soft fleshy process. Arti-
culation between telson and preceding seg-
ment barely discernible dorsally. Telson cov-
ered with series of 22 regularly arranged spines
similar to those of preceding segment; distal
margin composed of four spine-like processes
bearing secondary spines on outer edges, two
submedian with single row of secondary
spines, two intermediate with single inner
row of secondary spines and double row of
secondary spines along outer border (Fig. 20).
Exopod of uropods with eight movable spines
on outer margin; endopod with five spines
on dorsal surface; elongated basal portion of
uropods with two spines, outer longer than
the inner. Endopod of first abdominal ap-
pendage of male shown in Figure 21/. Spec-
imens, when alive, are a brilliant red or orange.
Specimens preserved in alcohol immediately
turn a mottled yellow or tan, but those pre-
served in formaldehyde have retained their
red color after several months. Specimens
range in size from 28.5 to nearly 70 mm. from
the tip of the rostrum to the distal margin of
the telson.

DISCUSSION: This is the only species of this
world-wide genus which has been reported
from the Hawaiian Islands. The type spec-
imen was taken by H.M.S. "Herald” at Fiji
Islands. Miers (1880) reported this to be a
female, but Kemp (1913) stated that it was
a male; apparently Kemp was the one who
made the error in identification. Another fe-
male was taken at Honolulu by H.M.S. "Chal-
lenger,” but Brooks (1886) put it in the genus
Protosquilla. The only other record of the spe-
cies is a single male specimen in the Bishop
Museum which was taken off Waikiki, Oahu.
In the spring of 1949 two males and four
females were dredged off Waianae, Oahu, in
water 50-120 fathoms deep. This appears to
be the greatest number of specimens collected
at any one time.

This species is easily recognized by the
radiating series of spines found on the telson

and the last two abdominal somites. The
rectangular carapace with its trispinous ros-
trum is found in several species of this genus.

GomdactylusMiAlx.^ related genus, Odontodac-
tylus, are very similar with regard to the ischio-
meral articulation and the inflated base of the
raptorial dactylus (Fig. 21c); however, Gono-
dactylus is distinguished by the absence of
spines on the raptorial dactylus, which may
be entirely smooth or serrated on its inner
margin (Fig. 21c). Odontodactylus, on the other
hand, has several spines on the inner margin
of the raptorial dactylus.

Brooks (1886) placed this and several allied
species in a new genus, Protosquilla, because
the telson and sixth abdominal somite, al-
though separate, were nearly fused. He also
stated that the small size of the antennary
scales and uropods and the trispinous ros-
trum were sufficiently distinct to warrant the
removal of these species from the genus Gono-
dactylus. Kemp (1913) and Edmondson (1921),
however, did not consider these differences of
a generic nature and retained the original
name applied to the species by White (1861).

Two specimens were kept alive for observa-
tion in an aquarium. Both were females, one
of which was collected with a mass of fertil-
ized eggs; the other laid a mass of eggs in
captivity. Both specimens survived very well
even though they were both dredged from a
depth of over 100 fathoms. The fertilized
eggs developed until the time of hatching but
then began to degenerate. Several eggs from
the mass were removed at intervals and pre-
served in 7 per cent formalin solution with
the hope that eventually they might be used
to describe the embryology of this species.
One female molted once while in captivity
and remained a mottled yellow afterward.
Little has been written about the function of
the elaborate spines on the telson; however,
my observations have shown that the animals
either burrow into the substrate or select holes
in dead coral which are just large enough for
them to crawl into. Ordinarily they are found
partially emerged from their burrows but when

Hawaiian Stomatopods - — ToWNSLEY

425

disturbed they turn around and close the
opening with their telson, which thus serves
as an effective protective mechanism.

distribution: This species has been re-
ported from Fiji and Honolulu, from rela-
tively deep water, the only record being 50-
120 fathoms.

LARVAL HAWAIIAN SQUILLIDAE

Larval stomatopods form a considerable
part of the neritic plankton throughout the
warmer waters of the world. They are trans-
parent organisms which, at hatching, have
many adult characteristics ; however, the diag-
nostic features of the adult do not appear
until late in development, thus making it dif-
ficult to associate the larval forms with the
proper adults. The first attempt to draw up
a key to the larvae was made by Bigelow
(1894); however, he based his key on the
general appearance of the forms, a character-
istic which is often very deceptive. Hansen
(1895) made a careful study of the larvae and
divided them into two major groups, the
alima-type larvae of Squilla and the erichthus-
type larvae of Lysiosquilla, Coronida, Odonto-
dactylus^ Gomdactylus^ and Pseudosquilla^ on the
basis of the shape of the propodus of the
third, fourth, and fifth thoracic appendages.
This division was thought by Hansen to em-
phasize the relationship of the various genera
to one another, but Kemp (1913) and Foxon
(1932) do not believe that this is of phylo-
genetic importance, because none of the larval
types possesses characteristics which can be
considered as an ancestral condition from
which the rest of the genera developed.

Giesbrecht (1910) has noted that in the
Stomatopoda two types of larvae occur at
hatching, but the differences between them
gradually disappear through a succession of
later stages. These two early larval types have
been designated pseudozoea and antizoea.
The antizoea is characterized at the time of
hatching by biramous appendages on the first
five thoracic somites, an abdomen which is
usually unsegmented and lacking pleopods,

and by later development into an erichthus
type of larva. The pseudozoea hatches with
only the first and second thoracic appendages,
the second being in the form of a raptorial
claw as in the adult, a fully segmented ab-
domen bearing four pairs of functional pleo-
pods; later these develop into either an erich-
thus- or an alima-type of larva. Species of
Lysiosquilla and Coronida hatch as antizoea,
later developing into erichthus larvae; species
of Gonodactylus, Odontodactylus, and Pseudo-
squilla all hatch as pseudozoea and later de-
velop as erichthus larvae. Species of Squilla
hatch as pseudozoea but later develop an
alima-type of larva.

Foxon (1932), following the work begun
by Hansen, attempted to bring the classifica-
tion of the larvae into agreement with the
divisions of the adults as presented by Kemp
(1913). In doing this he has set down the
morphological features which serve to identi-
fy the larvae from their earliest stages until
the time they take on the characters observed
in the adults. Brooks (1886), Giesbrecht
(1910), Hansen (1926), and Foxon (1932)
have all noted that the only constant differ-
ence between alima and erichthus larvae is
the number of intermediate denticles between
the submedian and the two lateral denticles
of the telson. In the former there are always
more than four of these denticles, whereas in
the latter there are less than four. If one uses
these denticles as a diagnostic character, it is
then possible to divide the larval forms into
the same major groups as the adults. In using
this system of classification the larval groups
are the same as those of the adults, the like-
ness becoming greater at each successive molt.

This method of classification is unques-
tionably suitable for distinguishing the gen-
era to which the larvae belong, but it is a
much more difficult task to assign them to
their proper species. In no species has an
individual in an early larval stage been reared
to the adult form. Individuals in later stages
have been reared to the postlarval condition,
but they are so delicate that laboratory con-

426

PACIFIC SCIENCE, VoL VII, October, 1953

ditions must be ideal, requiring water which
is well aerated and free of sediment. In con-
sequence, I have found it necessary to assign
the larval species on the basis of the adult
characteristics. If one has identified all adult
species occurring in a particular region, then,
by studying the postlarval forms which show
both adult and larval characteristics, it is pos-
sible to correlate the larval forms occurring
in the same region by working back from the
adult species to the earliest recognizable lar-
vae of those species. Using this method, I
have been able to identify larval stages of
Pseudosquilla as those belonging to adult P.
ciliata. Larvae of Lysiosquilla and Squilla are
likewise assigned to their respective species,
but this identification is not positive. At this
time larvae of Coronida and Odontodactylus can-
not be referred to any definite species. Larval
forms of Gonodactylus have not been found in
these waters.

The following key is designed to facilitate
the identification of the larvae with their re-
spective genera, and is intended to be used
for larvae in all stages of development. There
is no reference to species, but where the adult
species is known it will be mentioned in the
description given for each genus.

Key to the Genera of Larval
Hawaiian Stomatopoda

A. Larvae hatch without raptorial claw (an-
tizoea with biramous appendages on
first 5 thoracic somites) or with rapto-
rial claw (pseudozoea with only first and
second thoracic appendages) and with
the propodus finely pectinate along its
upper margin; ischio-meral articulation
of raptorial claw always terminal in later

stages (Figs. 22^, B

A A. Larvae always hatch with raptorial claw
(pseudozoea) and the propodus never
pectinate along its upper margin; ischio-
meral articulation of raptorial claw at a
point in advance of the proximal end of
the merus in later stages (Fig. 28^). .E

B. Larvae hatch with raptorial claw (Fig.

22d)\ propodus of fourth thoracic ap-
pendage rectangular, generally longer
than broad, and only slightly broader
than that of fifth (Fig. 23/) C

BB. Larvae hatch with biramous thoracic
appendages (Fig. 25^) which undergo
transformation into typical subchelate
form; propodus of fourth thoracic ap-
pendage rounded, broader than long,
more than twice as large as that of fifth
(Fig. 21 h) D

C. Telson with 4 or more intermediate den-
ticles between submedian and lateral

denticles (Fig. 22b)

Squilla (alima larva)

CC. Telson with 3 or fewer intermediate den-
ticles between submedian and lateral

denticles (Fig. 23^) ;

.... Pseudosquilla (pseuderichthus larva)

D. Propodus of raptorial claw broad, dac-
tylus slightly inflated at base (Fig. 2Ge) \
Coronida (coroniderichthus larva)

DD. Propodus of raptorial claw long and
slender, dactylus not inflated at base

(Fig- 25^)

Lysiosquilla (lysioerichthus larva)

E. Spines of telson greatly elongated (Fig.

28^)

. . Odontodactylus (odonterichthus larva)

EE. Spines of telson not greatly elongated
Gonodactylus (gonerichthus larva)

Squilla sp.^, probably S. oratoria |

Fig. 22a-d |

STAGE 1 (Fig. 22a) : This is the earliest stage j
of the alima-type larva which has been found. ’
Larvae bear completely developed first and i
second thoracic appendages, the second being '
in the form of a raptorial claw; propodus of ;
this claw with two small spines near its prox- *|
imal end, upper margin finely pectinate; dac- |
tylus nearly straight, no teeth along inner |
margin; eyes borne on long stalks, this con- ;
dition being retained throughout successive i
stages; antennules fully formed; antennal

Hawaiian Stomatopods — Townsley

427

Fig. 22. Larval forms of Squilla. a, Dorsal vie'w of pseudozoea, first abdominal appendage and telson (probably
S. oratorio)-, b, dorsal aspect of alima larva and first abdominal appendage (probably S. oratorio)-, c, ventral aspect
of alima larva (probably 5. oratorio)-, d, dorsal aspect of alima larva, first abdominal appendage and uropod
{Squilla sp.).

scale present but flagella absent; carapace very
clear, covering nearly 0.5 the entire body, this
relationship remaining the same throughout
successive stages; anterior margin of carapace
trispinous, posterior margin with two pos-
terolateral spines and small zoeal spine be-
tween them; lateral margins of carapace usual-
ly armed with three laterally directed and two
ventrally directed denticles, small denticle at
base of posterolateral spines; abdomen com-
pletely segmented, sixth abdominal somite
indistinct; four pairs of biramous pleopods
without gills, fifth pair represented by small
buds; no uropods; telson longer than wide
with 11 or 12 denticles between submedian

denticles and four between lateral and sub-
median denticles.

The specimens range in size from 5.5 to
8.0 mm. from the tip of the rostral spine to
the posterior margin of the telson. This stage
has been taken in plankton tows in the waters
around Oahu throughout the year, most
abundantly during the summer months.

STAGE 2 (Fig. 22b) : This larval form is sim-
ilar to the preceding stage except for a con-
spicuous change in size, the animals now
measuring 15 to 20 mm. Body proportions
remain the same, carapace still covering nearly
0.5 total length of body; antennal flagella
present; third, fourth, and fifth thoracic ap-

428

PACIFIC SCIENCE, Vol. VII, October, 1953

pendages present; sixth, seventh, and eighth
thoracic appendages represented by small
buds; abdominal appendages all present, with
small gills attached to exopods; uropods be-
ginning to form; sixth abdominal somite with
two small submedian spines; telson well
developed, beginning to assume adult con-
dition with two well-developed spines on
distal margin and distinct submedian and in-
termediate denticles.

As with the first stage, all the specimens
have been collected from regions where S.
oratoria is either known or thought to be
present. Innumerable specimens have been
taken in plankton tows in the waters around
Oahu throughout the year. They may fre-
quently be found in the stomach contents of
the yellowfin tuna, Neothunnus macropterus.

STAGE 3 (Fig. 22c) : By the time the larvae
have reached this stage, they have gained
further adult characteristics. Sixth, seventh,
and eighth thoracic appendages fully devel-
oped; uropods with adult characteristics, basal
portion elongated into two spines and small
rounded lobe between them, exopods with
eight spines, as yet immovable, along their
outer margin; telson more adult-like in ap-
pearance; no longitudinal carinae on body;
carapace still very transparent, no indication
of facies of adult carapace showing through
chitin.

At this time the specimens measure 45 to
60 mm. in length. This stage forms a con-
spicuous part of the plankton and has been
taken in plankton tows from the surface down
to a depth of 200 meters.

There are no specimens beyond this stage
in the collections. It is rather unusual that
no postlarval forms have been found in which
the adult characteristics such as the body cari-
nae, the adult carapace, and the teeth on the
raptorial dactylus are present. I assume from
this that the larvae become bottom dwellers
immediately after they lose the larval carapace.

Squilla sp.^

Fig. 22d

Approximately 200 specimens of this alima
larval stage were taken in night light stations
at Honolulu Yacht Harbor, Oahu. They close-
ly resemble the preceding larval types, but,
as only one stage was collected, it is impossi-
ble to assign the specimens to an adult species.
Carapace covers less than 0.3 total length of
body, very narrow; all appendages fully de-
veloped, gills represented by small buds on
exopod of pleopods; telson very large com-
pared with other body segments, composing
about 0.25 total length of body; fully devel-
oped uropods, bearing five spines along outer
margin of exopod, with basal portion elon-
gated into two spines, inner shorter than
outer.

Pseudosquilla ciliata

Figs. 25a- f, 24a-c

STAGE 1 (Fig. 25a): Brooks (1886) has il-
lustrated a pseudozoea larva similar to this
stage. This represents the earliest larval stage
of members of the genus 'Pseudosquilla. By
observing the relative changes in the other
stages, I have come to the conclusion that
these are the young of P. ciliata. Typical
erichthus larva with only first and second
thoracic appendages present; abdomen with
four pairs of biramous pleopods, without
gills; telson nearly rectangular, with eight
small denticles between submedian denticles;
eyes not borne on long stalks as in alima of
Squilla as articulation is close to ophthalmic
somite; antennules present; antennae appear
as small buds; merus and ischium of raptorial
claw with terminal articulation, propodus fine-
ly pectinate along upper margin, dactylus
composed of single spine; carapace trispinous
anteriorly, posteriorly bearing two long spines
reaching second abdominal somite; may or
may not have small zoeal spine between two
posterior spines; carapace longer than broad,
its lateral margins depressed.

Hawaiian Stomatopods — Townsley

429

f

Fig. 23. Larval forms of Pseudosquilla ciliata (Fabricius). a. Dorsal aspect of pseudozoea and first abdominal
appendage; b, lateral aspect of pseuderichthus larva, dorsal aspect of carapace and first abdominal appendage;
c, dorsal aspect of last abdominal somite, telson, and uropods of larva shown in b; d, e, lateral aspect of larger
pseuderichthus larva and dorsal aspect of last abdominal somite, telson, and uropods;/, second, third, fourth,
and fifth thoracic appendages of larva d.

Larval specimens in this stage of develop-
ment have been taken in great numbers in
plankton tows throughout the year at Kane-
ohe Bay, Oahu. Since adults of P. ciliata and
oculata occur in abundance on the reefs in the
bay, these specimens are believed to be the
first larval stage of a Pseudosquilla. Unfortu-
nately, I find no way to differentiate the early
larvae of species in this genus. The specimens
vary from 3.5 to 5.0 mm. from the tip of the
rostral spine to the posterior margin of the
telson.

STAGE 2 (Fig. 23^, c)\ At this stage in de-
velopment the larvae show characteristics of
both the preceding stage and the fourth stage.
Entire body and larval carapace have increased
in size but retain same relative proportions to
one another; carapace may or may not bear
small zoeal spine; first through fifth thoracic
appendages present; propodus of fourth and
fifth thoracic appendages rectangular in shape

and nearly equal in size (not well shown in
Fig. 23^); no appendages on sixth through
eighth thoracic somites; eyes globular, pro-
truding from anterior margin of carapace; an-
tennae present; pleopods resemble those of
following stage, but have no gills; sixth ab-
dominal somite distinct, bearing two sub-
median spines; uropods composed of short
exopod and endopod, with basal portion elon-
gated into two spines, inner shorter than
outer; telson retains same general appearance
of preceding stage, but somewhat larger.

Specimens measure 9.5 to 12.5 mm. in
length. Approximately 200 individuals in this
stage of development have been examined.
Most of the specimens were taken in plankton
tows from Kaneohe Bay, Oahu, but they have
also been found in the tows from the Molokai
Channel made by the Territorial Division of
Fish and Game.

430

PACIFIC SCIENCE, VoL VII, October, 1953

STAGE 3: This stage has not been found,
but examination of the preceding stage and
that of the next indicates what specimens in
this stage of development should look like.
By this time, all appendages should be pres-
ent, gills should appear on the exopod of the
pleopods, and the telson and uropods should
be similar to the following stage. They should
range from 15 to 20 mm. in length.

STAGE 4 (Fig. 2^d-f): Thoracic appendages
all present, showing adult characteristics, ex-
cept raptorial claw; raptorial claw still very
long and slender, propodus much longer than
ischium and merus combined, dactylus com-
posed of a single long spine with no indica-
tion of teeth along inner margin; facies of
postlarval carapace and rostrum show through
chitin of larval carapace; uropods greatly en-
larged with two immovable spines on outer
margin of exopod, definite pits for others
show through chitin, outer spine of basal
prolongation of uropods longer than innner;
telson armed with two submedian, two inter-
mediate, and two lateral spines, with 15 to 20
denticles between submedian and intermediate
spine; facies of postlarval telson showing
through chitin, with characteristic shape of
adult, being nearly as broad as long.

Specimens of this stage measure 20 to 22.5
mm. in length. Very few of these larvae have
been collected in regions where the previous
stages were most numerous. Occasionally
they are taken in the protected inshore waters,
but they are more often found in the offshore
waters such as the Molokai Channel. This
seems to indicate that these larvae seek less
sheltered waters and return to the shallow
inshore waters only when they lose the larval
carapace.

STAGE 5: There are no great changes from
the preceding stage at this time. Slight in-
crease in size; carapace very loosely attached
to body; mandibular palp present, composed
of three segments; nine or ten immovable
spines along outer margin of exopod of
uropod.

STAGE 6 (Fig. 24^, b)\ This is the first
postlarval form of P. ciliata. No larval cara-
pace, but animals remain planktonic; eyes
resemble adult, nearly tubular in shape, cor-
nea following long axis of stalk; rostrum tri-
angular; carapace distinctly adult in shape,
with distinct gastric grooves, covering only
anterior 0.25 of body; all appendages, except
second thoracic appendage, resemble those
of adult; second thoracic appendage, raptorial
claw, with shortened propodus and dactylus
so that they fit into grooved merus and ischium
when flexed, dactylus still lacks teeth on inner

Fig. 24. Larval forms of Pseudosqutlla ciliata (Fabri-
cius). ajy. Dorsal aspect, second thoracic appendage,
and first abdominal appendage of first postlarval stage;
c, dorsal aspect of anterior segments of last postlarval

Hawaiian Stomatopods — Townsley

431

margin; abdominal somites and appendages
distinctly adult in structure, gills on pleopods
nearly fully developed, sixth abdominal so-
mite with two well- developed submedian and
two lateral spines; uropods with nine or ten
movable spines along outer margin of exopod,
spines of elongated basal portion changing
relationship to one another so that in some
specimens these two spines are nearly equal
in length; telson with three distinct longi-
tudinal carinae, median and two submedians,
posterior margin with pair of movable sub-
median spines bearing series of denticles be-
tween them, pair of immovable intermediate
spines with two denticles between them and
submedian spines, pair of immovable lateral
spines.

The specimens now reach a maximum
length of 20 to 25 mm. This developmental
stage is very common in the plankton during
the spring and summer months and occurs in
both the inshore and offshore waters of Ha-
waii. They may occasionally be found in the
stomach contents of the yellowfin tuna, Neo-
thunnus macropterus, and the skipjack, Katsu-
wonus pelamis.

STAGE 7 (Fig. 24c): There is little change
in this form from the preceding stage. Ros-
trum nearly adult in form, broader than long;
eyes more tubular in shape; raptorial dactylus
composed of single tooth, facies of other teeth
on inner margin can be seen through chitin
in some specimens.

Individuals in this developmental stage
have been taken in the plankton from Ka-
neohe Bay, Oahu, and at Honolulu Yacht
Harbor, Oahu. This indicates that the larvae
are returning to the shallower, more pro-
tected waters and are preparing to settle
down on the bottom to complete their life
cycle.

Lysiosquilla sp.^

Fig. 25a

Brooks (1886) has illustrated this antizoeal
form and states that numerous specimens were
taken by the "Challenger” at Honolulu, Both

Brooks’s specimens and those at hand agree
with those described by Claus (1872) from
Messina. These very small larvae occur in
great numbers in the plankton throughout
the year. Due to their relatively small size
(2.5 to 3.5 mm.) they are very often over-
looked or cast aside as some unknown form.
Although they lack the characteristic sub-
chelate appendages, the eyes and telson have
distinct stomatopod characteristics. These lar-
vae represent the first larval stage of Lysio-
squilla or Coronida, and because of their great
abundance I am inclined to believe that they
belong to Lysiosquilla which is apparently
more abundant in Hawaiian waters than Coro-
nida. There is no diagnostic character suffi-
ciently developed to indicate conclusively the
species to which these specimens belong, and
none have ever been reared in captivity to the
following stage. The carapace completely
covers the animal from head to abdomen
leaving only the eyes and telson exposed.
There are five pairs of biramous thoracic ap-
pendages present, the abdomen is only faintly
segmented, and the pleopods are absent. This
larval form gives rise to an erichthus in later
stages.

Lysiosquilla sp.^

Figs. 25b, c

Nine specimens of this lysioerichthus stage
were collected in a plankton tow in the Molo-
kai Channel. In all respects this larval form
represents the typical lysioerichthus. No spines
between submedian and lateral spines on tel-
son (Fig. 25c); raptorial propodus finely pec-
tinate along upper margin, dactylus composed
of single acute spine, not inflated at base,
ischio-meral articulation terminal; all thoracic
appendages present, propodus of third and
fourth rounded and broader than long, fourth
nearly twice as large as that of fifth; abdomen
increasing in breadth posteriorly; pleopods
all present, gills absent; uropods partially de-
veloped, exopod and endopod both present,
with basal portion elongated into two spines.

Fig. 25. Larval forms of Lysiosquilla. a, Antizoea; b, lateral aspect of lysioerichthus larva and first abdominal'^
appendage; c, dorsal aspect of last abdominal somite, telson, and uropods of same; d, ventral aspect of lysioerich-
thus larva; e, ventral aspect of last larval stage of L. maculata; dorsal aspect of first postlarval stage of L. maculata.

inner longer than outer; carapace nearly as
broad as long from base of anterolateral spines
to base of posterolateral spines, lateral mar-
gins turned downward and toward median
line; most preserved specimens with abdomen
and telson reflexed anteriorly so that ventral

opening of carapace is completely closed.

Specimens measure 14.5 to 28.5 mm. in
length. The 28.5 mm. specimen shows the
facies of ten spines along the inner margin of
the raptorial dactylus much the same as those
seen in the adults of L. maculata.

Hawaiian Stomatopods — ToWNSLEY

433

Lysiosquilla sp?

Fig. 2'bd

This lysioerichthus larval form is represent-
ed by a single specimen taken in a plankton
tow in the Molokai Channel. Its very large
size (45 mm.) immediately draws attention.
All structures are the same as for the previ-
ously described larvae of this genus. It was
temporarily believed to belong to L. maculata,
but its size and relative immaturity (cf. Fig.
25^, which is only 15 mm. long) seem to
indicate that it may be the larval stage of a
species of Lysiosquilla which has not been
found in the adult form in Hawaii.

Lysiosquilla maculata
Fig. 25f, /

Figure 25^ represents one of seven lysio-
erichthus larvae, three of which were collected
in plankton tows at Truk Island and four
from Kawaihae, Hawaii. These are fully de-
veloped larvae, which I believe to be the last
larval stage of L. maculata. This conclusion
is based on an examination of postlarval forms
and the adults of this species. Appendages,
except raptorial claw, telson, and uropods,
resemble those of adult L. maculata; facies
of postlarval telson show through chitin, very
closely resembling telson of postlarval forms
shown in Figure 25/. It is very probable from
these facts that the next molt would produce
the postlarval stage of L. maculata. Specimens
range from 15 to 16 mm. from the tip of the
rostral spine to the posterior margin of the
telson.

The postlarval form (Fig. 25/) undoubtedly
belongs to L. maculata. All anterior append-
ages are the same as those of the adult, and
the raptorial dactylus bears nine teeth on its
inner margin similar to the condition found
in some of the adults of this species. The
telson and uropods have the same general
appearance of the adult as well. These spec-
imens and the adults have eight spines along
the outer margin of the exopod of the uro-

of last larval stage of coroniderichthus larva; h, dorsal
aspect of last abdominal somite, telson, and uropods
of same; c, first abdominal appendage of same; d, dor-
sal aspect of anterior segments of first postlarval stage;
e, second thoracic appendage of same.

pods. A single specimen measuring 16.5 mm.
in length was collected at Kawaihae, Hawaii,
at a night light station, which further sub-
stantiates the identification because the adult
species is very common in that region.

Coronida sp.^

Fig. 2Ga-e

In Figure 2Ga there is represented the last
larval stage (coroniderichthus) of some un-
known species of Coronida. This form is
found in the Ala Wai Canal, Oahu, and at
Kawaihae, Hawaii, in moderate numbers dur-

434

PACIFIC SCIENCE, VoL VII, October, 1953

Fig. 27. Larval form of Coronida sp. a. Dorsal aspect
of coroniderichthus larva; b, second, first, third, fourth,
and fifth thoracic appendages; c, first abdominal ap-
pendage.

ing the summer. Larvae resemble lysioerich-
thus in shape of thoracic appendages, but
with raptorial dactylus slightly inflated at its
base; propodus of fourth thoracic appendage
rounded, nearly twice as large as that of fifth
which is rectangular; all abdominal append-
ages present; no gills; telson rectangular as
in most larval forms, bearing two denticles
between submedian and lateral denticles; uro-
pods present, outer margin of exopod armed
with three spines, basal portion elongated
into two spines, outer longer than inner.

Specimens measure 10.0 to 14.5 mm. from
the tip of the rostral spine to the posterior
margin of the telson.

Figures 26d, e represent the first postlarval
stage of this species which is found in nearly

equal numbers with the previous stage. Only
the carapace and anterior segments and the
raptorial claw have been illustrated as all other
structures are the same as those seen in the
preceding stage. The inner margin of the dac-
tylus is armed with ten teeth. There is no
distinct increase in size over that of the last
larval stage. These forms may be larval stages
of C. sinuosa, but the larval structures bear
no resemblance to diagnostic characteristics
of this species. Apparently the species to which
these larvae belong is quite common, al-
though it has not been found as yet.

Coronida sp.^

Fig. 21a-c

Five coroniderichthus larvae measuring 18
to 20 mm. from the tip of the rostral spine
to the posterior margin of the telson were
collected at Kawaihae, Hawaii. The append-
ages indicate that they belong to some species
of Coronida, but no adult specimens have been
found which have the same characteristics as
those possessed by this larval stage. I believe
that the first postlarval stage would appear
at the next molt because the facies of the
adult carapace, telson, and raptorial dactylus
show through the very clear chitin. These
specimens are unique in that the larval cara-
pace covers all but the last two abdominal
somites and the telson, and the median an-
terior spine and posterolateral spines are
greatly elongated. This condition is not found
in any other larval form, hence it should
serve as a diagnostic character for the separa-
tion of this larval form from others belonging
to this genus.

Odontodactylus sp.^

Fig. 28^-^

Eight odonterichthus larvae, ranging in
length from 18 to 35 mm., were taken in a
plankton tow in the Molokai Channel. They
resemble pseuderichthus larvae very much,
and it is possible to distinguish the two only

Fig. 28. Larval forms of Odontodactylus. a. Lateral aspect of odonterichthus larva, first abdominal appendage,
and dorsal aspect of last abdominal somite, telson, and uropods; h, lateral aspect of odonterichthus larva and
dorsal aspect of last abdominal somite, telson, and uropods; c, first, third, fourth, and fifth thoracic appendages
of same; d, second thoracic appendage of same; e, first abdominal appendage of same.

by comparing the raptorial claws. In pseu-
derichthus larvae this appendage is finely
pectinate along the upper margin of the pro-
podus, and the dactylus is not inflated at the
base. In odonterichthus, however, the rapto-
rial propodus is not pectinate along its upper
margin, and the dactylus is inflated at its
base. In later stages the ischio-meral articula-
tion of the raptorial appendage would be a
useful diagnostic character, but none of the
specimens has attained this condition, which
is present in the adult. This genus is un-
doubtedly closely related to Gonodactylus, but
it has been impossible to compare the larvae
of the two genera because no specimens of
the latter have been collected in Hawaiian
waters. Claus (1872) and Brooks (1886) have
both illustrated gonerichthus larvae which
were collected from other regions, and com-

parisons with their figures show that the Mo-
lokai Channel specimens definitely belong to
the odonterichthus type. Foxon (1932) uses
the character of the long spines found on the
telson of Odontodactylus larvae for separating
them from Gonodacytlus, although he does not
present any figures to illustrate his statements.
Since no Gonodactylus larvae have been col-
lected in the Hawaiian waters, I cannot say
whether or not this is a valid character for the
separation of the larvae of the two genera, but
comparisons with Claus’s and Brooks’s figures
of gonerichthus larvae show that the spines
on the telson of their specimens are consider-
ably shorter than those found in the present
specimens. With this in mind I have placed
the character in the key with the hope that
some future worker who may find the go-
nerichthus larvae will be saved some effort in

436

PACIFIC SCIENCE, Vol. VII, October, 1953

trying to place them in their correct genus.

REFERENCES

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neue oder seltene Reptilien aus Neu-Gra-
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Bigelow, R. P. 1893. The Stomatopoda of
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1931. Stomatopoda of the southern

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1899- On the Stomatopoda and Ma-

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1907. Stomatopoda from the western

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Brooks, W. K. 1886. Report on the Stomatopoda
collected by the H.M.S. "Challenger" during
the years 1873-1876. Vol. 16, part 65, 116
pp., 16 pis. Eyre and Spothswoode, London.

Brulle, M. 1836. In: Histoire naUirelle des
lies Canaries, Zool. Vol. 2, part 2 (Crust.)
[ed. Barker-Webb and S. Berthelot], 152
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Claus, C. 1872. Die Metamorphose der
Squilliden. Gess. Wiss. Gottingen, AbhandL
16: 111-163, 8 pis.

Dana, J. D. 1852. Crustacea. In: United States
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DE Haan, W. 1850. Crustacea. In: Fauna
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DE Man, J. G. 1907. On a collection of Crus-
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from the Inland Sea of Japan; with descrip-
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Edmondson, C. H. 1921. Stomatopoda in the
Bernice P. Bishop Museum. Bernice P.
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Eydoux, F., and L. Souleyet. 1841. Voyage
autour du monde execute pendant les annees
1836 et 1837 sur la corvette la Bonite, com-
mandee par M. Vaillant, Zoologie. Vol. 1,
xxxvii+334 pp. A. Bertrand, Paris.

Fabricius, j. C. 1787. Mantissa insectorum.
Vol. 1. xx-f 330 pp. C. G. Proft, Hafniae.

1793. Entomologia systematica emendata

et aucta. Vol. 2. viii+519 pp. C. G. Proft,
Hafniae.

Foxon, G. E. H. 1932. Report on stomatopod
larvae, Cumacea and Cladocera. Great Bar-
rier Reef Expedition 1928-29, Scientific Reports
[Brit. Mus. (Nat. Hist.)] 4(11): 375-398.

Fukuda, T. 1911. Further report on Japanese
Stomatopoda with descriptions of two new
species. Annot. Zool., Jap. 7: 285-289.

Giesbrecht, W. 1910. Stomatopoden. In:
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der Angrenzenden Meeresabschnitte . Monog.
33(1), 239 pp., 11 pis. R. Friedlander und
Sohn, Berlin.

Gurney, R. 1942. Larvae of decapod Crustacea.
vi+306 pp. Ray Society, London.

Hansen, H. J. 1895. Isopoden, Cumaceen
und Stomatopoden der Plankton Expedi-
tion. Ergebnisse der Plankton Expedition der
Humboldt Stiftung 2: 1-105, 8 pis., 1 chart.
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1926. The Stomatopoda of the Siboga

Expedition. Siboga Expeditie 1899-1900,
Monog. 18, 48 pp., 2 pis. E. J. Brill,
Leiden.

Hawaiian Stomatopods ■ — Townsley

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geschkhte der Krabben und Krebse, VoL 2,
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Hess, W. 1865. Beitrage zur Kennmiss der
Decapoden-Krebse Ost Australien. Arch. f.
Naturgesch. XXXI, 1: 168-170,

Kemp, S. 1911. Preliminary descriptions of
new species and varieties of Crustacea Sto-
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1913. An account of the Crustacea

Stomatopoda of the Indo-Pacific region,
Indian Mm., Mem. 4(1): 1-217.

Lamarck, J. B. 1818. Histoire naturelle des ani-
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Miers, E. J. 1880^. On the Squillidae. Ann.
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Milne-Edwards, a. 1878. Description des
quelques especes nouvelies des crustaces
provenant du voyage aux lies du Cap-Vert
de MM. Bouvier et de Cessac. Soc. Philo-
math., Bui. VII, 2(4): 225-232.

PocoCK, R. 1. 1893. Report upon the sto-
matopod crustaceans obtained by P. W.
Basset Smith, Esq., Surgeon, R.N., during

the cruise in the Australian and China Seas
of H.M.S. "Penguin,” Commander W. U.
Moore. Ann. and Mag. Nat. Hist. VII, 11:

473-479.

Randall, J. W. 1839. Catalogue of Crustacea
brought by T. Nuttall and J. K. Townsend
from the west coast of North America and
the Sandwich Islands, with descriptions of
such species as are apparently new, among
which are included several species of dif-
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Phila., Jour. I, 8: 106-147.

Rumphius, G. E. 1705. D’ Amboinsche Rari-
teitkamer. xliii+340 pp,, 60 pis. Halma,
Amsterdam.

Tattersall, W. M. 1906. Report on the Lep-
tostraca, Schizopoda, and Stoniatopoda,
collected by Professor Herdman at Ceylon
in 1902. In: Herdman, Report to the govt,
of Ceylon on the pearl oyster fisheries of the
Gulf of Manaar. Part 5, pp. 158-188, 3 pis.
Royal Society, London.

White, A. 1861^. Descriptions of two species
of Crustacea belonging to the families Cal-
lianassidae and Squillidae. Zool. Soc. London,
Proc. 1861: 42-44.

1861^. Descriptions of two species of

Crustacea belonging to the families Cal-
lianassidae and Squillidae. Ann. and Mag.
Nat. Hist. Ill, 7: 479-481.

The Role of Dehydration in the Development of Laterite^

G. Donald Sherman, Yoshinori
Kanehiro, and Yoshito Matsusaka^

Soil formation in tropical regions is deter-
mined by the nature and extent of the chemi-
cal weathering of the soil- forming materials.
The intense chemical weathering and the va-
rying length of time of exposure of minerals
to weathering have resulted in the develop-
ment of a broad sequence of soil formation
in the tropical areas of the world. Sherman
(1949) has shown that the soils of the Ha-
waiian Islands exhibit a sequence of soil de-
velopment which can be related to a sequence
of mineral weathering as proposed by Jackson
et aL (1948). This sequence of soil formation
represents a series of successive stages of sec-
ondary mineral development with the follow-
ing order of soils rich in: (1) primary minerals;
(2) montmorillonite; (3) kaolinite; (4) hy-
drated sesquioxides; and (5) dehydrated ses-
quioxides. The end product of this weathering
cycle would be either a ferruginous laterite or
a bauxite laterite, depending on the environ-
mental conditions. These two products rep-
resent two extremes of a wide range of weath-
ered end-products, as the intermediate member
— a ferruginous bauxitic laterite — is quite
common (Harrison, 1934).

Ferruginous laterite was described by Bu-
chanan (1807). As described by Buchanan,
laterite is a soil horizon which has a very high
content of iron oxides. These iron oxides
exist as the minerals, limonite, goethite, hema-

1 Published with the approval of the Director of the
Hawaii Agricultural Experiment Station, Honolulu,
Hawaii, as Technical Paper 273. Manuscript received
December 2, 1952.

2 Chairman and Junior Chemists, respectively. De-
partment of Soils and Agricultural Chemistry, Hawaii
Agricultural Experiment Station.

tite, and magnetite. The profile of the fer-
ruginous laterite must have a horizon rich in
iron oxide which will harden, on exposure, to
an indurate layer with brick-like properties.
The ferruginous ' horizon may contain con-
cretions of iron oxide in which case it is a
pisolithic laterite, or the iron oxide may exist
as a lens in a porous matrix which is known
as a vesicular laterite, according to Pendleton
(1941). Ferruginous laterite may also develop
as a massive horizon near or at the surface
which on exposure will harden and develop
a uniform, structureless crust, or corapace.
Soils of this type have been described by
Aubreville (1947, 1948), Chevalier (1948),
and Sherman (1950).

The mode of origin, or formation, of fer-
ruginous laterite has been a point of contro-
versy. Glinka (1927) believed that laterites
formed under both evergreen and savannah
types of vegetation. Richthofen's views (I860)
differed from Glinka's in that he postulated
the formation of laterite under a former forest
vegetation which had been succeeded by a
shrub and grass vegetation. In other words,
as soils degraded they could not support the
evergreen forest; thus, the forest disappeared^,
and was replaced by a lower level of vegeta-
tion. Harrassovitz (1930), Holmes (1914),
and Campbell (1917) concluded from their
studies that laterite formed only where cli-
matic conditions were conducive to the up-
ward movement of the dissolved soil sesqui-
oxides, which would only occur under alter-
nating wet and dry conditions. On the other
hand, Pendleton (1941, 1942, 1946) believes
that laterites are formed by the oxidation and

438

Dehydration in Laterite Formation — SHERMAN et d.

439

precipitation of the iron oxides in the zone
above the fluctuating water table. This layer
is made up of concretions of iron oxide which
will harden on exposure to give the ironstone
laterite, a material suitable for building mate-
rial. Finally, there are soil scientists who hold
that laterite has developed by residual accu-
mulation as the result of leaching and removal
of silica. Marbut (1932) and Fox (1933) were
the leading advocates of this hypothesis of
the mode of formation of laterite.

In spite of the numerous hypotheses as to
the origin of laterite, there are certain facts
which are included in all proposals. One of
these facts is that in the early stages the
weathering surface is losing silica and bases.
Desilication is taking place with the progres-
sive development of soils having a sequence
of minerals with a lower silica content. Sec-
ondly, as desilication proceeds the sesqui-
oxides accumulate in the soil. These weather-
ing processes are accelerated under high
rainfall and high temperatures which are com-
mon to many tropical regions. Lastly, all
ferruginous laterites harden on exposure. The
hardened laterite is made up of lenses or con-
cretions of iron oxides often cemented to-
gether by a kaolinitic clay.

A number of titaniferous ferruginous lat-
erite crusts have been described in the Ha-
waiian Islands by Sherman (1952). Each
profile of these soils has a massive indurate
surface horizon having both a high bulk den-
sity and a high particle density. This hardened
surface horizon is underlaid by a friable sub-
soil. This subsoil horizon possesses the bulk
density of an average soil. Likewise, the par-
ticle density is lower than that of the surface
horizon. Below the friable subsoil horizon is
an impervious layer which may be the un-
weathered country rock, a plastic clay, or an
unconformity-like relationship which may
have developed by pedogenic processes rather
than being a geological unconformity.

These laterite crusts were either barren or
were covered by scattered bunches of grass
or vines. However, some of the areas which

exhibited the crust-like surface horizon had
a fair amount of vegetation. These areas, also,
exhibited a high bulk density and high par-
ticle density relationship such as is found in
the barren crust areas. From many observa-
tions made during wet and dry weather, it
was noted that the bulk density of the soil
varied according to the soil moisture condi-
tions. In a visit to a barren crust area on
Kauai after a heavy rain, a marked difference
was observed between the bulk density and
hardness of the surface soil of the barren area
and the adjacent identical soil under a vege-
tative cover. It has been observed that the
crust will develop on irrigation ditch banks
where wetting and drying have occurred at
very frequent intervals. The appearance of the
hardened crust can be associated with dry
periods, suggesting that its formation is due
to dehydration. Thus, it is possible that vege-
tation will protect the hydrated minerals from
dehydration and prevent the crust formation.

A study has been conducted to determine
the role of dehydration of the hydrated ses-
quioxides in the development of the laterite
crust. The study also included the determina-
tion of the effect of dehydration on the physi-
cal and chemical properties of the soil. For
this study two sets of soil samples from ad-
jacent profiles were selected. The soil profiles
were located on the slopes of the western rim
of Waimea Canyon on the island of Kauai.
A titaniferous ferruginous laterite crust covers
a rather level bench on this slope. Profiles
were selected at different areas of this laterite
crust. The two main profiles used in this study
were taken at the edge of the barren crust area
so that the profile without a vegetative cover
was taken only 3 feet from the site of the
profile with a vegetative cover. The morphol-
ogy of the two profiles was identical except
for the obvious difference in the physical con-
dition of the surface horizon. The characteris-
tics of the two profiles are shown in Figure 1.

METHODS OF ANALYSIS

Soil samples were collected from the pro-

440

PACIFIC SCIENCE, VoL VII, October, 1953

BARREN SOIL

GRASS AND SHRUB
VEGETATION

% VOLATILE
MATTER

% VOLATILE
MATTER

^///A FERRUGINOUS CRUST \////A HARDENED SOIL ^

FRIABLE SOIL

Fig. 1. The location and relationship of the hardened crust and vegetative cover of two adjacent profiles from
Waimea Canyon, Kauai.

files as shown in Figure 1 and in a like manner
from Other similar areas, including the crust
developed on an irrigation ditch bank. The
bulk density, particle density, differential
thermal analysis, cation exchange capacity,
and content of the major oxides were deter-
mined for each soil sample. The bulk density
of the soil was determined by two methods.
The first was by removing the soil from a
given space and determining its dry weight.
The volume of soil was determined by fitting
a plastic bag to the space occupied by the
soil and ascertaining the amount of water
required to fill the hole. The second method
was similar except that the soil was taken by
a sharp-edged cylinder of a known volume.
The particle density of the soil was deter-
mined by the pycnometer (Wright, 1934).
The cation exchange capacity and the oxide
analysis were made by methods described by
Piper (1944). Differential thermal analysis was
made according to the procedure proposed
by Norton (1939).

EXPERIMENTAL RESULTS

Bulk Density and Particle Density: The most
striking characteristics of the laterite crust are
the high apparent and true specific gravity.
Fujimoto etal. (1948) have described a laterite
crust in which the surface horizon contains

more than 50 per cent heavy minerals. In the
profile there is a tremendous difference be-
tween the indurate surface horizon and the
friable layer in their bulk density and soil
particle density. This same difference was
present in the surface layer of the barren pro-
file but was not apparent in the profile with
a vegetative cover. In Table 1 the bulk densi-
ties and particle densities of the soils taken
from a profile without vegetative cover and
a profile with vegetative cover are given. The
bulk density of the soil samples from the
well- developed laterite crusts was 1.9 or great-
er, and their particle density was higher than
3.8. The bulk density of the soils from the
profile with a vegetative cover was 1.0 or
lower and the particle density ranged from
2.7 to 3.3. The highest bulk<*density and par-
ticle density, 2.5 and 4.1, respectively, were
found in a fragment of the crust from an
irrigation ditch bank wall. The data clearly
demonstrate the obvious difference between
the soils of the barren areas and those with
a vegetative cover. The observations indicate
that the vegetative cover does protect the soil
from hardening to an indurate horizon. The
differences are more clearly demonstrated in
Table 2, which presents the averages of the
different types of soil profiles in Table 1.

Dehydration in Laterite Formation ■ — ShermaN et al.

441

TABLE 1

A Comparison of the Bulk Density and Particle Density of Different Soil Horizons in Barren
' Laterite Crust Areas with Adjacent Areas Having a Vegetative Cover

BARREN SOILS (CRUST)

vegetative COVER (no crust)

location

DEPTH

BULK

DENSITY

particle

DENSITY

location

DEPTH

BULK

DENSITY

PARTICLE

DENSITY

Laterite crust

Waimea Canyon

Site A

Inches

0- 7

2.0

3.84

Waimea Canyon
Site C near
sites A and B

Inches

0- 5

0.8

2.86

Laterite crust

Waimea Canyon

Site A— friable layer

7-14

1.1

2.71

Waimea Canyon
Site C near
sites A and B

5-10

0.9

2.99

Laterite crust

Waimea Canyon

Site B

0- 7

2.2

4.43 ,

Waimea Canyon
Site C near
sites A and B

10-15

0.9

2.82

Laterite crust

Waimea Canyon

Site B-— friable layer

7-14

0.8

2.21

Waimea Canyon
Site C near
sites A and B

15-20

1.0

2.91

Laterite crust

Lower Waimea Canyon
Irrigation ditch bank

Site E

0- 5

1.9

3.98

Laterite crust

Lower Waimea Canyon
■ Irrigation ditch bank

Site E

5-10

2.1

4.16

Waimea Canyon
Site D above
site C

Matted

roots

on

surface

0.9

2.75

Laterite crust

Lower Waimea Canyon
Irrigation ditch bank

Site E

10-15

2.1

4.25

Waimea Canyon
Site D above
site C

0- 5

0.9

3.24

Hardened soil

Lower Waimea Canyon
Irrigation ditch bank

0- 5

1.4

4.00

Waimea Canyon
Site D above
site C

5-10

0.9

3.21

Hardened soil

Lower Waimea Canyon
Irrigation ditch bank

5-10

1.3

3.81

Waimea Canyon
Site D above
site C

10-15

1.0

3.34

Hardened soil

Lower Waimea Canyon
Irrigation ditch bank

10-15

1.3

3.79

Loose clod

Irrigation ditch bottom

2.5

4.14

Chemical Composition: The soil samples from
the two profiles shown in Figure 1 were ana-
lyzed for major oxide constituents, and the
data obtained are given in Table 3. A marked
difference was found in the amounts of cer-
tain constituents and also in the weight loss

on ignition. The soil of the surface horizon
from the area having a vegetative cover showed
a loss of weight amounting to 28 per cent on
ignition, whereas the soil from the same ho-
rizon of the barren profile lost a little over 2
per cent. The same data from the 10-1 5 -inch

442

PACIFIC SCIENCE, VoL VII, October, 1953

TABLE 2

The Average Bulk Density and Particle Density of a Series of Ferruginous Laterites
AT Different Degrees of Hardening

AVERAGE

samples from

SAMPLES FROM

SAMPLES FROM

PROFILES

SAMPLES FROM

CRUST

HARDENED SOIL

HAVING VEGETATION

FRIABLE SUBSOIL

Bulk density

2.13

1.33

0.91

0.95

Particle density

4.13

3.87

3.01

2.46

layer of both profiles showed practically no
difference in their loss of weight on ignition,
being 26 and 28 per cent.

The content of iron and titanium oxides
is the highest in the soil from the 0-5 -inch
and 5-10-inch layers of the profile of the
barren area, while the 10-1 5 -inch layer of both
profiles has a very similar amount of these
oxides. The aluminum oxide content of soil
from the 0-5 -inch and 5-10-inch layers of the
profile with the vegetative cover is higher
than that for the corresponding soil layers of
the profile from the barren area. The soils
from the profile with a vegetative cover are
less acid than the soils from the 'barren area.
In the former the soil reaction ranges from
pH 4.4 to 4.6 and in the latter from pH 3.8
to 4.2.

In Table 4 the data obtained from the ana-
lysis of cation exchange capacities of the soil

are given. The cation exchange capacity of the
soils shows the same relationship as the loss
on ignition. The soil from the surface horizon
of the profile from the area having a vege-
tative cover has a much higher exchange capa-
city than that from the surface horizon of the
profile from the barren area, being 41.0 and
3.7 milliequivalents per 100 grams, respective-
ly. Likewise, a difference existed in the 5-10-
inch layer of the two profiles in which the
cation exchange capacities were 37.0 and 15.6
milliequivalents per 100 grams, respectively.
The cation exchange capacities of the 10-15-
inch layer were very similar, being 38.4 and
34.9 milliequivalents per 100 grams. The or-
ganic matter content of the soil with a
vegetative cover was higher than that found
in the barren area. The organic matter content
is not of sufficient quantity to explain this
change in cation exchange capacity, so de-

TABLE 3

The Chemical Composition of Laterite Profiles; a Comparison of Effect of Dehydration
Due to Exposure on Chemical Composition

LOSS ON

DEPTH

pn

Si02

A1203

Fe203

Ti02

IGNITION

Soil with vegetative cover (no crust)

Inches

Per cent

Per cent

Per cent

Per cent

Per cent

0- 5

4.6

7.50

13.00

41.80

6.03

27.92

5-10

4.6

7.84

15.45

40.99

6.29

26.78

10-15

4.4

14.78

22.60

27.40

4.34

28.53

Barren soil (crust)

0- 5

3.8

6.46

4.13

56.02

31.02

2.06

5-10

4.0

5.68

9.46

60.63

11.87

10.23

10-15

4.2

13.01

22.58

32.10

5.67

25.72

Dehydration in Laterite Formation — Sherman et aL

443

TABLE 4

The Relationship between Loss on Ignition and Cation Exchange Capacity
UNDER Barren Laterite Crust and Adjacent Soil with a Vegetative Cover

barren soil (crust)

SOIL WITH VEGETATION (NO CRUST)

depth

CATION

CATION

PH

organic

LOSS ON

exchange

PH

organic

LOSS ON

exchange

matter

ignition

CAPACITY

matter

IGNITION

CAPACITY

Inches

Per cent

Per cent

m.e.llOOg.

Per cent

Per cent

m.e.llOOg.

0-5

3.8

1.1

2.0

3.7

4.6

8.2

27.9

41.0

5-10.

4.0

10.2

15.6

4.6

26.8

37.0

10-15

4.2

25.7

34.9

4.4

28.5

38.4

hydration is considered to be the cause of its
reduction.

Differential Thermal Analysis: The thermal
curves obtained from the differential thermal
analysis are given in Figure 2 for the soil
profile from the barren area and Figure 3 for
the soil with a vegetative cover. There is a
marked difference in these two sets of curves.
The curve for the surface horizon of the soil
with a vegetative cover showed a strong exo-
thermic peak at 180°C., whereas the surface
soil from the barren area showed no thermal
reaction at this temperature. In fact, the curve
of the latter soil is very similar to that ob-
tained from a sample of hematite. The thermal
curves of the subsoil horizons of both pro-
files exhibit thermal peaks at 180°C. to 190°C.,
but none of the soils exhibited a peak at
710°C. which is characteristic of montmoril-
lonite. The thermal curves of this type are
probably due to the existence of amorphous
hydrated oxides of iron, aluminum, and ti-
tanium. Some of the soils have a small peak
at 350°C. which is probably due to goethite.

As the differential ' thermal analysis was
made on the untreated soils, the question
arises as to whether these curves are influenced
by the difference in organic matter content
of the soil. The surface soil of the profile with
a vegetative cover has a higher content of
organic matter than that of surface soil of
the barren area. It will be noted that the
thermal curves of the 10-1 5 -inch layer of the

profile with a vegetative cover are exactly the
same as that of the surface horizon. As this
horizon has a low organic matter content, the
minerals of the soil must be responsible for
its thermal characteristics. The loss of water
below 300°C. also substantiates the fact that
dehydration of minerals is responsible for
thermal characteristics of the soil, as the loss
of water is the greatest in those soils showing
thermal peaks at 180°C.

The ferrous iron content was determined
in the surface soils from the two profiles. The
soil from the profile with a vegetative cover
contained from two to five times as much
ferrous iron as the soil from the barren area.
As the surface soil from the barren areas is
magnetic and has a low ferrous iron content,
the presence of maghemite is suggested in
these soils. Furthermore, this would suggest
the existence of the lepidocrocite-maghemite
system in these soils as maghemite is the
product of dehydration of lepidocrocite. Fur-
ther evidence is needed to establish the valid-
ity of this suggestion.

DISCUSSION

The results of this study have established
that the indurate laterite crust horizon is de-
veloped by the dehydration of the hydrated
iron and titanium minerals in the soil. The
thermal curves from the differential thermal
analysis and the loss of water under 300°C.
of the two profiles have shown that hardening

444

PACIFIC SCIENCE, VoL VII, October, 1953

of the barren area is due to dehydration. In
the dehydration of the colloidal hydrated iron
oxides, water is lost and the oxides are con-
verted to hematite or related dehydrated iron
oxides. The process will cause an increase in
particle density and, according to Fujimoto
et al. (1948), an increase in particle size. The
question arises as to the length of time re-
quired for a ferruginous horizon to harden
to the laterite crust on exposure. Aubert (1949,
1950) has reported the formation of laterite
crusts under normal conditions in 35 to 60
years. Chevalier (1949) states that, if the forest
is removed by cutting, burning, or loss of
cover due to other reasons, the senile stage
(the laterite crust) will develop very rapidly.
Davis (1940) found that because of the clear
culture required in the culture of teak and
because teak trees lose their leaves during the
dry season, the laterite crust will develop in
a very short period of time [one dry season].
The hardening of Hawaiian soils on exposure
is very rapid, occuring in less than a year.

0 300 500 700 lOOO'C

Fig. 2. The differential thermal curves of the profile
from the barren area of a humic ferruginous latosol
soil.

Fig. 3. The differential thermal curves of a profile
with a vegetative cover of a humic ferruginous latosol
soil.

The dehydration ' of the soil causes very
important changes in the physical and chem-
ical properties of the soil. There is an obvious
change in compaction, bulk density, particle
density, and particle size. The most impor-
tant change is the loss of cation exchange
capacity. It has been observed that these bar-
ren areas are infertile. This infertility very
likely is due to the loss of adsorptive surface
and to the poor physical condition of the soil.
The reforestation of these laterite areas is a
slow process as the conditions for growth are
very poor.

Dehydratioo in Laterite Formation Sherman et aL

445

It has been established by Fieldes et aL
(1952) that the amorphous colloidal hydrated
oxides have a high cation exchange capacity.
In a personal communication from these
workers, they report that they have found
that the hydrated iron oxides lose cation ex-
change capacity on dehydration. These ob-
servations support the findings in this study.

SUMMARY

A study has been conducted to ascertain
the role of dehydration in the development
of the indurate laterite crust. The following
conclusions appeared to be justified:

1. The data obtained from differential ther-
mal analysis, elemental analysis, and physi-
cal measurements have established that the
dehydration of the colloidal hydrated ox-
ides of the soil is responsible for the de-
velopment of the indurate laterite horizon
when exposed to a drying environment.

2. The dehydration of the titaniferous fer-
ruginous horizon will increase the bulk
density and particle density of the soil.

3. Dehydration causes the development of
inert dehydrated minerals as shown by the
data on loss on ignition and cation ex-
change capacity. The colloidal hydrated
iron oxide is converted to hematite or
similar iron oxide minerals.

4. The rates of dehydration of ferruginous
layers will probably vary according to the
mineral content of soil and the vegetative
and climatic environments.

REFERENCES

Aubert, G. 1949. Observations sur le role de
Eerosion dans la formation de la cuirasse
lateritique. Bui. Agr. du Congo Beige 40:
1383-1386.

1950. Observations of the degrada-
tion and lateritic-crust formation in North-
West Dahomey. [Fourth] Internatl. Cong.
Soil Set., Trans. 3: 127-128.

Aubreville, a. 1947. Erosion and "bovali-
zation” in French Black Africa. Agron. Trop.
2: 339-357.

1948. Casamance. Agron. Trop. 3:

25-52.

Buchanan, H. F. 1807. Journey from Madras^
Canara, and Malabar. London. [3 vols.,
reference to vol. 2.]

Campbell, J. M. I917. Laterite: Its origin,
structure, and minerals. Mining Mag. 17:
67-77; 120-128; 171-179; 220-229.

Chevalier, A. 1948. Degradation and con-
servation of soil of tropical Africa: Origin
and extension of laterites and of ferrugin-
ous shell: Struggle against sterilization of
African soil. Rev. Internatl. Bot. Appl. 28:

49-66.

1949. Points de vue nouveaux sur les

sols d'Afrique tropicale sur leur degrada-
tion et leur conservation. BuL Agr. du
Congo Beige 40: 1057-1092.

Davis, P. W. 1940. Preliminary note on Nil-
amber soils with special reference to their
suitability for teak. Indian Forester 66: 658-
671.

Fieldes, M., L. D. Swindale, and J. P,
Richardson. 1952. The relation of col-
loidal hydrous oxides to the high cation
exchange capacity of some tropical soils of
the Cook Islands. Soil Sci. 74: 197-206.

Fox, C. S. 1933. Laterite and laterite soil. In-
dian Forester 59: 630-635.

Fujimoto, Giichi, G. Donald Sherman,
and Ada E. Chang. 1949. The chemical
composition of the separated mineral frac-
tions of a ferruginous humic latosol profile.
Soil Sci. Soc. Amer., Proc. (1948) 13: I66-I69.

Glinka, K. D. 1927. The great soil groups oj
the world. 150 pp. Edward Bros. Ann Arbor,
Michigan.

Harrassowitz, H. 1930. Boden der trop-
ischen Regionen, Blanch’s Handhuch der
Bodenlehre 3: 362-436.

Harrison, J. B. 1933. The katamorphism of
igneous rocks under humid tropical conditions.
79 pp. Imperial Bureau of Soil Science,
Harpenden, England.

Holmes, A. 1914. Investigations of laterites
in Portuguese East Africa. Geol. Mag. 6'

529-537.

446

PACIFIC SCIENCE, Vol. VII, October, 1953

Jackson, M. L., S. A. Tyler, A. L. Willis,
G. A. Bourbeau, and R. P. Pennington.
1948. Weathering sequence of clay-size
minerals in soils and sediments. Jour. Phys.
Colloid. Chem. 52: 1237-1260.

Marbut, C. F. 1932. Morphology of later-
ites. \Second\ Internatl. Cong. Soil Sci., Trans.
5: 72-80.

Norton, F. Id. 1939- Critical study of the
differential thermal method for the identi-
fication of the clay minerals. Amer. Ceramic
Soc., Jour. 22: 54-63.

Pendleton, R. L. 1941. Laterite and its
structural uses in Thailand and Cambodia.
Geog. Rev. 31: 177-202.

and S. Sharasuvanas. 1942. Ana-
lyses and profile notes of some laterites and
soils with iron concretions of Thailand.
Soil Sci. 54: 1-26.

1946. Analysis of some

Siamese laterites. Soil Sci. 62: 423-433.
Piper, C. S. 1944. Soil and plant analysis. 368
pp. Interscience Publishing Co., New York.
Richthofen, F. I860. Bermerkungen fiber
Ceylon. Deut. Geol. GeselL, Ztschr. 12: 531-
545.

Sherman, G. Donald. 1949. Factors influ-
encing the development of lateritic and
laterite soils in the Hawaiian Islands. Pacific

Sci. 3: 307-314.

1950. The genesis and morphology

of Hawaiian ferruginous laterite crusts.
Pacific Sci. 4: 315-322.

1952. The titanium content of Ha-
waiian soils and its significance. Soil Sci.
Soc. Amer., Proc. 16: 15-18.

Wright, C. H. 1934. Soil analysis. 236 pp.
Thomas Murby & Co., London.

A Discussion of the Trematode Genus Schistorchis
(Family Lepocreadiidae) with Descriptions of Two New
Species from Hawaii^

Mary Louise Hanson^

This report is the first to be based on spec-
imens collected during June and July of 1949
at the Marine Laboratory of the University of
Hawaii, Honolulu. To Dr. Robert W. Hiatt,
chairman, and to the members of the Depart-
ment of Zoology and Entomology at the
University of Hawaii I wish to extend my
sincere appreciation for the laboratory space
and the laboratory equipment which were
furnished me. To Dr. William A. Gosline,
ichthyologist in the Department of Zoology
and Entomology at the University of Hawaii,
the author is indebted for his invaluable assist-
ance in identifying the fish hosts. The fishes
were made available through the generosity
and assistance of Mr. Spencer W. Tinker,
Director of the Honolulu Aquarium. From
all these individuals, and from Dr. H. W.
Manter under whose direction the work was
done, the author has received valued encour-
agement. The trip to Hawaii was made pos-
sible by a Robert A. Wolcott Scholarship
granted by the University of Nebraska Foun-
dation.

The trematodes, upon removal from the
hosts, were killed under a cover glass with
F.A.A. fixative (formalin, alcohol, and acetic
acid). Storage for a short time was in 70 per
cent alcohol. The worms were stained in De-
lafield’s haematoxylin, cleared in xylol, and
mounted in Permount.

1 Studies from the Department of Zoology, Univer-
sity of Nebraska, No. 261.

^ University of Nebraska State Museum, Lincoln.
Manuscript received July 22, 1952.

Type specimens are being deposited at the
United States National Museum.

The genus Schistorchis Liihe, 1906, now rec-
ognized to be quite distinct from Pleorchis
Railliet, 1896, with which it was once asso-
ciated, has the following characteristics: me-
dium to large, elongated Homalometroninae;
smooth or spined anteriorly; oral sucker ter-
minal or subterminal; ceca long, with or
without ani; excretory vesicle I-shaped with
long collecting tubules often reaching pha-
ryngeal level; genital pore median or very
slightly to the left, just anterior to the ace-
tabulum; cirrus sac absent; genital sinus pres-
ent; eleven testes intercecal, tandem in a single
row or arranged more or less in a double row;
ovary pretesticular, median or to the right;
seminal receptacle and Laurer’s canal present;
uterus not extending posterior to the ovary;
parasites of marine fishes. Type species: S.
carneus Liihe, 1906.

In addition to the type species the genus
contains four species: 6. callyodontisY dirndigun^
1942; 5. sigani YdiVnd^gMU, 1942; 5. stenosoma;
and S. zancli. The latter two species are here
described. The following key to the genus
Schistorchis is proposed:

1. Testes not in a single median row 3

Testes in a single median row. ........ 2

2. Uterus preovarian .6. zancli

Uterus reaching at least to mid-ovary . . . . ■
. 5. stenosoma

3. Testes not in two lateral, opposite' groups

. .4

Testes in two lateral, opposite groups ....

5. carneus

447

448

PACIFIC SCIENCE, VoL VII, October, 1953

4. Acetabulum near cecal bifurcation

S. callyodontis

Space between acetabulum and cecal bi-
furcation nearly twice length of acetabulum
V sigani

SYSTEMATIC POSITION

Liihe (1906: 102) and subsequent authors
have associated the genus Schistorchis Liihe,
1906, with the genus Pleorchis Railliet, 1896.
Johnston (1913: 383-386) named Pleorchis oli-
gorchis which Odhner (1928: 5) considered
identical with the previously described Schis-
torchis carneus. Stunkard (1931: 713-714)
quoted the latter paper, apparently in agree-
ment. Yamaguti (1942: 362) referred to
'' Schistorchis oligorchis (Johnston, 1914)
[correctly 1913] which is very closely related
to if not identical with, Schistorchis carneus,''
seeming to have reached the conclusion in-
dependently. Manter and Van Cleave (1951:
331) likewise referred to Pleorchis oligorchis
Johnston, 1914 [sic] . . . transferred to Schis-
torchis by Yamaguti (1942).”

Poche (1926: 1-458) placed the genus
Schistorchis with Pleorchis in the family Ple-
orchiidae where it remained until 1942. Cable
and Hunninen (1942: 306) observed that ”the
genera Pleorchis and Schistorchis for which
Poche (1925) [correctly 1926] erected the
family Pleorchiidae, do not seem to be at
all closely related” and considered Pleorchis
among the Acanthocolpidae. Nothing was
suggested for Schistorchis,

At very nearly the same time, Yamaguti
(1942: 364) proposed the separation of Ple-
orchis and Schistorchis and erected the new
family Schistorchiidae to include Schistorchis,
and also Apocreadium Manter, 1937, and Cho-
anodera Manter, 1940, which he considered
to resemble Schistorchis closely. Both Choano-
dera and Apocreadium were considered by
Manter (1940: 344-348) as members of the
family Gyliauchenidae. Cable and Hunninen
(1942: 309) considered the gyliauchenids as
related to Lepocreadiidae, and Manter (1947:
264) stated that '^Apocreadium and Choanodera

are so similar to Homalometron (differing chief-
ly in their lymphatic vessels) that they should
be transferred to the Homalometroninae, fam-
ily Lepocreadiidae.” Thus Schistorchis is the
only remaining genus in the family Schistor-
chiidae.

The genus Schistorchis agrees very well with
the description of Homalometroninae in the
family Lepocreadiidae Nicoll, 1934, as pro-
posed by Cable and Hunninen (1942: 308).
Although this is the first report of multiple
testes among the Homalometroninae, they
are present among the Lepocreadiinae and are
provided for in the description of the family.

The presence or absence of ani is often
difficult to determine in a whole mount. Both
Schistorchis callyodontis and S. sigani were de-
scribed from a single specimen. Both Liihe
and Johnston described S. carneus without ani,
but Odhner (1928: 5) found them to be pres-
ent when he sectioned specimens. Stunkard
(1931 : 723-724) observed that secondary con-
nections between the intestinal ceca and the
exterior have developed independently in sev-
eral different families and that such commu-
nications are not uniformly present in all
members of those families in which they do
occur. The conclusion was drawn that such
connections are relatively recent acquisitions
which developed after the families became
differentiated.

It is proposed that the genus Schistorchis be
classified as Homalometroninae, family Lepo-
creadiidae, and that the family Schistorchiidae
become a synonym of Lepocreadiidae.

Schistorchis stenosoma n. sp.

Fig. la-d

HOST: Cantherines pardalis (Ruppell), file
fish; in all of 7 specimens examined.

location: Intestine.

DESCRIPTION (based on a series of 31 spec-
imens) : Body 7 or 8 times longer than wide,
1.701 to 4.271 mm. long, 0.211 to 0.635 mm.
wide; width about equal along most of body
length, but slightly wider immediately ante-

Genus SchistofcMs — Hanson

449

rior to acetabulum; both ends slightly tapered.
Rows of widely spaced, short, blunt spines
rarely extend as far posterior as acetabulum.
Ventrally spines somewhat heavier and extend
more posteriorly. Traces of eye-spot pig-
ment present. Oral sucker subterminal, 0.161
to 0.300 mm. long, 0.175 to 0.315 mm. wide,
very nearly round. Acetabulum 1/4 to 1/5
body length from anterior end; 0.117 to 0.234
mm. long by 0.119 to 0.234 mm. wide, very
nearly round; sucker ratio 1:0.530 to 1:0.801.
Prepharynx very short, appearing absent in
many specimens; pharynx rounded, 0.080 to
0.146 mm. long by 0.098 to 0.169 mm. wide;
esophagus 0.029 to 0.131 mm. long; bifurca-
tion immediately anterior to acetabulum; area
of bifurcation voluminous; ceca parallel to
body walls, each terminating in anus; ani
terminal, each encircled with cuticular furl.
Genital pore median or slightly to left of
median line, at anterior border of acetabulum.
Testes 11, intercecal, serial and contiguous;
0.044 to 0.190 mm. long by 0.060 to 0.208
mm. wide, anterior testes usually larger than
posterior. Pattern of vasa efferentia varied.
Usually a single vas efferens from each testis
leading to a vas deferens; in some instances
two vasa efferentia arise from a common
origin on a testis (Fig. lb). Two vasa defer-
entia just median to ceca enter posterior por-
tion of seminal vesicle separately. Seminal
vesicle bipartite; posterior portion 0.095 to
0.354 mm. long by 0.085 to 0.248 mm. wide,
rounded, extending posteriorly from level of
mid-acetabulum, never reaching ovary, joined
to anterior portion by a short, narrow tube;
anterior portion much smaller than posterior,
longer than wide, variable in size, opening
into genital sinus with uterus. Prostatic cells
loosely surround anterior portion of seminal
vesicle. Ovary round, 0.073 to 0.162 mm.
long by 0.080 to 0.162 mm. wide, lying to
right of median line just anterior to anterior
testis, but never touching it. Uterus with 3
or 4 intercecal coils; entering genital sinus
dorsal to acetabulum (Fig. Ic). Uterus always
extends posteriorly at least to anterior margin

Fig. 1. Schistorchis stenosoma Itom Cantherims pardalis
(Ruppell), file fish, a, Ventral view; b, pattern of vasa
efferentia and vasa deferentia (diagrammatic); c, ter-
minal genital ducts (ventral view); <7, ootype region
(dorsal view), a, c, and d drawn with the aid of a camera
lucida. Projected scale in millimeters. Abbreviations:
a — anus, e = egg, ev ~ excretory vesicle, gp = genital
pore, gs = genital sinus, Ic — Laurer’s Canal, mg =
Mehlis’ gland, ov = ovary, sr = seminal receptacle,
sv = seminal vesicle, / = testis, ut = uterus.

of ovary, usually to posterior margin of ovary,
occasionally slightly beyond ovary although
never reaching anterior testis. Mehlis’ gland
dorsal to ovary, diffuse, extending from an-
terior edge of anterior testis to level of mid-
ovary, intercecal. Laurer’s canal well devel-
oped, opening midway between acetabulum
and ovary, dorsal to left cecum; a short, nar-
row tube directed first medianly and then
posteriorly, enlarging into receptacle-like por-

450

PACIFIC SCIENCE, Vol. VI^I, October, 1953

tion, then again narrowly tubular to ootype.
Connection between ootype and true seminal
receptacle clearly discernible (Fig. \d). Sem-
inal receptacle 0.065 to 0.190 mm. long by
0.078 to 0.218 mm. wide, as large as or larger
than ovary; slightly anterior to, more median
and more dorsal than ovary; vitellaria folli-
cular from level of anterior testis (3 or 4
follicles anterior to .anterior testis) to posterior
end of body; lateral and ventral to ceca, very
few intercecal follicles anterior to posterior
testis, then becoming confluent. Eggs rela-
tively few (7 to 15), usually collapsed, 37 to
68 by 22 to 43 }i, usual range 51 to 56 by
31 to 36 Excretory pore terminal between
anal furls; excretory vesicle I-shaped, extend-
ing to posterior edge of posterior testis, joined
there by principal pair of collecting tubules.
Tubules extend anteriorly just median to ceca,
cross ceca ventrally at acetabular level, narrow
and tend to coil slightly at pharyngeal level,
then continue forward lateral to oral sucker.

COMPARISONS: S. stemsoma, with the 11 tes-
tes in a single, median row, with the uterus
found at least as far posterior as mid-ovary
and sometimes slightly posterior to the pos-
terior border of the ovary, and with body
spines, differs from all previously described
species of Schistorchis.

S. carnetis was originally described with
cirrus sac and without ani, but Yamaguti
(1942: 362) concluded that the cirrus sac was
absent, and Odhner (1928: 5) described the
ani. S. carneus differs from 5. stenosoma in its
bulkier body proportions, the retracted oral
sucker, the smaller pharynx, the apparent ab-
sence of an esophagus, the long excretory
vesicle which extends anterior to the ovary,
and the arrangement of the testes in two
lateral groups.

5. callyodontis differs from S. stenosoma by its
lack of ani, the longer excretory vesicle, the
ovary which is at the same level as the anterior
testis, the nearly equal size of the suckers,
the testes arranged irregularly in two rows,
and the heavy musculature of the anterior
portion of the body.

5. sigani differs from 5’. stenosoma by its lack
of ani, the longer excretory vesicle, the very
short esophagus, the wide area between the
intestinal bifurcation and the acetabulum, the
nearly equal size of the suckers, the small
pharynx, the testes which are arranged irregu-
larly in two rows, and the larger eggs.

The name stenosoma is a combination of
steno- (narrow) and soma (body), referring
to the relative slenderness of the body.

Schistorchis zancli n. sp.

Fig. 2a, b

HOST: Zanclus cornutus (L.), Moorish idol or
kihikihi loulu; in 1 of 9 specimens examined.

LOCATION: Intestine.

DESCRIPTION (based on two specimens):
Body elongate, 2.409 to 2.643 mm. long,
0.606 to 0.686 mm. wide; widest at acetabular
level, tapering abruptly anteriorly at esopha-
geal level and posteriorly at level of second
testis; width of narrower posterior portion
nearly equal; anterior end blunt, posterior end
rounded. A few small, scattered spines present |
anterior to level of esophagus. Traces of eye- |
spot pigment present. Oral sucker subqua- '
drate, terminal, 0.168 to 0.196 mm. long by i
0.256 mm. wide. Acetabulum l/3 body length
from anterior end; rounded, 0.197 to 0.204
mm. long by 0.183 to 0.190 mm. wide; sucker
ratio 1:0.715 to 1:0.742. Very short prepha-
rynx observed in one specimen ; pharynx 0. 1 19
to 0.133 mm. long by 0.128 to 0.146 mm. j
wide, nearly rectangular; esophagus 0.043 to |
0.085 mm. long; ceca wide anteriorly, nar- i
rowing posterior to acetabulum, nearly paral- |j
lei to body walls, each with ventral, sub- !
terminal anus. Genital pore slightly to left j!
of median line at anterior border of acetab- |
ulum. Testes 11, largely intercecal but oc-
casionally ventral to median portion of ceca, ||
serial and contiguous, 0.080 to 0.168 mm.
long by 0.141 to 0.199 mm. wide, ellipsoidal |
to ovoid in shape. Seminal vesicle 0.256 to
0.285 mm. long by 0.183 to 0.219 mm. wide,
saccular, with slightly swollen tube extending

Genus ScMstorchis — HANSON

451

from seminal vesicle to shallow genital sinus.
Seminal vesicle ovoid^ extending posteriorly
nearly to ovary, narrowing into nonmuscular
pars prostatica surrounded by several pro-
static cells. Cirrus lacking. Ovary rounded,
0.124 to 0.153 mm. long by 0.146 to 0.153
mm, wide, lying to right of median line, con-
tiguous with anterior testis. Short uterus with-
out lateral coils, with 2 straight, longitudinal
limbs, one ventral, one dorsal; latter joining
pars prostatica near genital pore (Fig. 2^),
intercecal, to left of median line, preovarian
and well filled with eggs. Mehlis’ gland dorsal
and to left of ovary, diffuse,- extending from
level of mid-ovary anteriorly, slightly larger
than ovary, Laurer’s canal well developed,
opening dorsal to left cecum at level of sem-
inal vesicle, directed at first anteriorly than
at mid-acetabular level turning backward;
basal portion near ovary swollen. Seminal
receptacle large, claviform, lying dorsal to
and slightly anterior to ovary, Vitellaria of
small follicles often grouped 2 or 3 together,
extending from level of posterior edge of
acetabulum to very near posterior end of
body, lateral and ventral to ceca, somewhat
confluent posterior to testes. Prominent yolk
ducts extend from both sides at level of ovary,
ducts seemingly swollen into reservoirs in one
specimen. Eggs usually collapsed, 48 to 54
by 36 to 43 /i (majority 51 to 53 by 36 to
39 m)- Excretory pore ventral, subterminal,
slightly anterior to ani and on median line;
I-shaped vesicle, extending to posterior edge
of posterior testis, joined there by principal
pair of collecting tubules; tubules extend
anteriorly just median to ceca, cross ceca
ventrally beginning at level of ovary, conti-
nuing forward to pharyngeal level.

COMPARISONS: 5. zancli is most similar to
S. stemsoma of the other known species of the
genus. S. zancli is like 5. stenosoma and different
from all other species in the genus in the
possession of body spines, and in the presence
of 11 testes arranged in a single, median row.
It differs from S. stenosoma in that it has a more
rhomboidal oral sucker, a larger, more rec-

Fig. 2. Schistorchis zancli from Zanclus cornutus (L.),
Moorish idol or kihikihi loulu. a. Ventral view; b, ter-
minal genital ducts (ventral view). Drawn with the aid
of a camera lucida. Projected scale in millimeters.
Abbreviations: a = anus, gp = genital pore, gs —
genital sinus, mg = Mehlis’ gland, pp = pars prosta-
tica, sr ~ seminal receptacle, sv = seminal vesicle,
t = testis, ut — uterus.

tangular pharynx, a greater angle at the origin
of the ceca, a relatively more posterior ace-
tabulum, more anterior extent of the vitel-
laria, fewer and smaller vitelline follicles, a
more robust body, a preovarian uterus, ellip-
soidal or ovoid rather than round testes, a
relatively shorter posttesticular space, a non-
muscular pars prostatica, a non-bipartite sem-
inal vesicle, subterminal ani without cuticular
furls, and a subterminal excretory pore.

The name zancli refers to Zanclus, the genus
of the host.

452

PACIFIC SCIENCE, Vol. VII, October, 1953

REFERENCES

Cable, R. M., and A. V. FIunninen. 1942.
Studies on Deropristis inflata (Molin), its
life history and affinities to trematodes of
the family Acanthocolpidae. Biol. Bui. 82
(2): 292-312.

Johnston, S. J. 1913. On some Queensland
trematodes, with anatomical observations
and descriptions of new species and genera.
Quart. Jour. Micros. Sci. [n. s.] 59(3): 361-
400.

Luhe, Max. 1906. Report on the trematode
parasites from the marine fishes of Ceylon.
In: Report to the government of Ceylon on the
pearl oyster fisheries of the Gulf of Manaar,
by W. A. Herdman. Part 5: 97-108. Royal
Society, London.

Manter, Harold W. 1937. A new genus of
distomes (Trematoda) with lymphatic ves-
sels. Allan Hancock Pacific Exped., Rpts. 2(3):
11-23.

1940. Digenetic trematodes of fishes

from the Galapagos Islands' and the neigh-
boring Pacific. Allan Hancock Pacific Exped.,

Rpts. 2(14): 329-496.

1947. The digenetic trematodes of

marine fishes of Tortugas, Florida. Amer.
Midland Nat. 38(2): 257-416.

and Harley J. Van Cleave. 1951.

Some digenetic trematodes, including eight
new species, from marine fishes of La Jolla,
Calif. IJ. S. Natl. Mus., Proc. 101(3279):
315-340.

Odhner, T. 1928. Weitere Trematoden mit
Anus. Arkiv for Zool. 20(2): 1-5.

PoCHE, F. 1926. Das System der Platodaria.
Arch. f. Naturgesch. 91(2): 1-458.

Stunkard, Horace W. 1931. Further ob-
servations on the occurrence of anal open-
ings in digenetic trematodes. Ztschr. f
Parasitenk. 3(4): 713-725.

Yamaguti, S. 1938. Studies on the helminth
fauna of Japan. Part 21. Trematodes of fishes,
IV. 139 pp., 83 figs., 1 pi. Maruzen Com-
pany, Ltd., Kyoto.

1942. Studies on the helminth fauna

of Japan. Part 39. Trematodes of fishes
mainly from Naha. Biogeog. Soc. Japan,
Trans. 3(4): 329-397.

The Aedes Mosquitoes of the Philippine Islands
IIL Subgenera Aedimorphus^ Banksinella^ Aedes^ and Cancraedes

(Diptera^ Culiddae)^

Kenneth L. Knight and William B. Hull^

INTRODUCTION

This is the last of a series of three papers
dealing with the Aedes fauna of the Philippine
Islands. Part I (Knight and Hull, 1951) con-
tained the general introduction, the keys to
the adults and larvae, and the species treat-
ments for the subgenera Mucidus, Ochlerotatus,
and Finlaya. Part II (Knight and Hull, 1952)
dealt with the subgenera Skusea, Christopher -
siomyia, Geoskusea^ Rhimskusea, and Stegomyia.
In this part the remaining aedine subgenera
known from the Philippines are considered,
i.e., Aedimorphus, Banksinella, Aedes, and Can-
craedes.

All the necessary introductory information
is given in Part I, and only explanations need-
ed for an understanding of the abbreviations
and symbols used are repeated here.

A dagger(t) denotes types that have been
examined by us.

The following abbreviations are used in
the descriptions. Adult: apn, anterior pronotal
lobe; ppn, posterior pronotal lobe. Larva: isc,
inner or upper caudal hair of the anal segment;
osc, outer or lower caudal hair of anal segment;
Ih, lateral hair on anal plate.

Figures are not drawn to scale.

1 The opinions or conclusions contained herein are
those of the authors and are not to be construed as
official or reflecting the views of the Navy Department
or of the Naval service at large. Manuscript received
August 12, 1952.

^ Naval Medical Research Institute, Bethesda, Mary-
land. Lieutenant Commander, MSC, USN, now with
Malaria and Mosquito Control Unit #1, Naval Air Sta-
tion, Jacksonville, Florida, and Lieutenant, MSC, USN,
respectively.

The following abbreviations are used in the
species distribution sections to indicate the
present locations of specimens examined.

A. N.S.P., Academy of Natural Sciences of
Philadelphia; C.A.S., California Academy of
Sciences; U.S.N.M., U. S. National Museum;

B. M., British Museum (Natural History);

C. C., Cornell University; R.K.L., specimens
from the collection of Naval Medical Research
Unit No. 2 on which nothing has been pre-
viously published and which are now de-
posited in the U. S. National Museum, the
Johns Hopkins University, the California
Academy of Sciences, the British Museum,
and the Knight collection.

SYSTEMATICS

Subgenus Aedimorphus Theobald
1903. Aedimorphus Theobald, Mon. CuL 3:
290. Genotype: domesticus Theobald
(Africa) .

1905. Reedomyia Ludlow, Canad. Ent. 37: 94.
Genotype: pampangensis Ludlow (Phil-
ippines).

ADULT: Brown species, some with distinc-
tive areas of silvery scaling. In general related
to Stegomyia, but all the Philippine species
distinct from that subgenus in the form of the
male palpi (except the undescribed species
mentioned in the discussion of wainwrighti)
and of the dististyle of the male terminalia.
Male palpi longer than the proboscis; seg-
ments IV-V down-tilted, numerous long hairs
arising laterally and apically on the distal por-
tion of III and all along IV-V (the unde-

453

454

PACIFIC SCIENCE, Vol. VII, October, 1953

scribed species mentioned above has slender,
relatively hairless palpi as in Stegomyia). Vertex
dorsum with a variable area of narrow scales
medially (confined to nape in wainwrighti and
in some specimens of ostentatio; all narrow-
scaled in V. nocturnus), remainder broad-scaled,
Scutellum either broad- or narrow-scaled. Pa-
ratergite with or without scales. No lower
mesepimeral bristles. Fore and mid-tarsal
claws of male unequal, each unidentate; hind
claws equal, simple. Tarsal claws of female
equal, fore and mid each unidentate; hind
simple. Terminalia: Basistyle without true
apical lobes and without basal lobe, except
in V. nocturnus where a setose basal plaque
occurs. Dististyle greatly swollen apically, ex-
cept in V. nocturnus, the enlarged portion bear-
ing a complicated arrangement of hairs and
modified setae. Claspettes absent. Mesosome
divided into two lateral plates, each of which
bears several teeth laterally.

LARVA: Separable from all the other sub-
genera in the Philippines, except Ochlerotatus
and some specimens of the Finlaya species
leucopleurus and aureostriatus, by the possession
of an acus on the anal plate. Antennal hair
tuft with 5-13 branches. Head hair 4 with
2-8 branches, 5 and 6 with 1-7. Comb with
16-26 small scales in a patch, or with 8-10
large teeth in an irregular, partially double
row. Siphon with an attached acus; 12-21 pec-
ten teeth, each with one or more large ventral
denticles; distal teeth more widely spaced and
without denticles. Anal plate incomplete;
ventral brush of 12-16 tufts, all but basal
1-4 borne on a prominent, laterally connected
barred area. Habitat chiefly rain-filled tem-
porary pools, but also occasionally such
places as stream margins and permanent pools
and ditches.

DISTRIBUTION: Seven species are known
from the Philippines. This subgenus is largely
confined to the Ethiopian and Oriental re-
gions.

SYSTEMATICS: Edwards (1932: 166) divid-
ed the subgenus into eight groups. Knight
and Hurlbut (1949: 29) modified this classifi-

cation by combining Group D with Group C.

No sharp division occurs between Stegomyia
and Aedimorphus , the two subgenera being
apparently intimately connected through
Group A {apkoannulatus-gtoup) of the Ethio-
pian region and through Group B {nummatus-
group) of the Oriental region (via Group E
of Stegomyia).

In the Philippine fauna, Group C is rep-
resented by alboscutellatus, mindoroensis, pam-
pangensis, and punctifemore. In that fauna it is
separable from Group G, represented only by
vexans nocturnus, in the absence of a basal lobe
on the basistyle and in the possession of an
apically swollen and markedly modified dis-
tistyle. The positions of ostentatio and wain-
wrighti are uncertain because the males are as
yet undescribed. The other groups are not.
represented in the Philippines.

Aedes (Aedimorphus) pampangensis
(Ludlow)

Figs. 1, 2

1905. Reedomyia pampangensis Ludlow, Canad,
Ent. 37: 94 (3 females). Type locality:
Philippines. Angeles, Pampanga Prov,,
Luzon (Whitmore) . However, the spec-
imen in the U.S.N.M. is labeled "Camp
Wm. McKinley, Rizal, P. L, Nov. 3.
05.” in Ludlow’s handwriting. Type:
Female (holotype) in U.S.N.M, j
1922. Aedes (Ecculex) alhoscutellatus , Theobald.
Edwards, Indian Jour. Med. Res. 10:
467. In part.

1925. Aedes {Aedimorphus) niveoscutella Theo-
bald. Dyar and Shannon, Insecutor In- f
scitiae Menstruus 13: 76. 5

1932. A. (A) alhoscutellatus Theobald. Ed-
wards, Genera Insectorum. Ease. 194,
p. 167, Barraud, 1934, 250. In part.

1945. Aedes {Aedimorphus) pampangensis (Lud-
low). Bohart, U. S. NavMed. 580, p.

53. Different combination.

ADULT: Vertex dorsum with a median nar-
row-scaled area. Scutum uniformly brown-

Aedes of the Philippines — Knight AND Hull

scaled. Scutellum with broad silvery scales.
Tarsi all dark.

Male. Head: Proboscis pale, apical one-
fourth dusky, particularly dorsally. Palpi
slightly longer than proboscis; dark; seg-
ments IV-V down-tilted, numerous long hairs
arising apically and all along IV-V. Vertex
with a narrow area of narrow pale scales along
either side of mid-line, a line of narrow pale
scales along the dorsal portion of the eye
margins; remainder of vertex with broad
yellowish-white scales; brown upright-forked
scales over the whole dorsum. Thorax: Scutal
integument brown; clothed with small, nar-
row brown scales, may be a few narrow yel-
lowish scales along the anterior margin, over
the wing base, and on the prescutellar area.
Scutellar lobes covered with broad silvery-
white scales. Apn with a few narrow yellowish
scales; ppn sparsely covered with narrow
brownish and yellowish scales. Propleuron,
subspiracular area, and paratergite each with
some narrowed yellowish scales; upper ster-
nopleuron, lower posterior sternopleuron, and
upper mesepimeron each with small patch of
broad creamy- white scales. Legs: Fore femur
with anterior surface dark, paler basally; pos-
terior surface with dorsal half pale, remainder
dark; dorsal surface pale. Mid-femur dark an-
teriorly except for a few silvery-white scales
dorsally at apex; posterior surface pale except
for a narrow dorsal dark area on apical half.
Hind femur pale, dark dorsally, this dark scal-
ing encroaching onto the anterior and pos-
terior surfaces along apical one-half; a narrow
apical band of silvery-white scales anteriorly.
Tibiae with posterior surfaces pale-scaled.
Tarsi all dark-scaled. Fore and mid-tarsal claws
unequal, each claw unidentate; hind tarsal
claws equal, simple. Wings: Dark-scaled. Hal-
ter stem pale, knob with dusky scales. Ab-
domen: Tergites dorsally dark (may be a few
median basal yellowish scales on IV-V); a
lateral patch of creamy-white scales on I,
II-VII with a complete lateral band of brown-
ish-yellow scales, these basally encroaching
upon the dorsum on V-VIL Sternite VIII

455*

Fig. 1. A. {Aedimorphus) pampangensis. Male ter-
minalia (Luzon).

with a dense divided tuft of very long yellow
hairs. Terminalia: Basistyle broad, short; a
dense band of very long setae on tergal sur-
face; a subbasal tuft of short, curved setae on
inner tergal surface, seme of these setae being
distinctively broadened; basad of this tuft and
on the inher sternal margin is a small group
of setae, 2-3 of these being rather stout;
apically on the inner sternal surface is a group
of very long setae. Dististyle very distinctive
in form; inner surface as figured; outer surface
with numerous very short setae and with a
large tuft of long setae directed toward base
of dististyle (shown by the group of dashed
lines in the figure).

Female. Differs from the male mainly as
follows: Proboscis dusky, pale beneath from
near base to about the apical one-fourth. Palpi

456

Fig. 2. A. {Aedimorphus) pampangensis. Larva (Min-
danao). a. Head; b, terminal segments.

about one-sixth as long as the proboscis;
dusky. Torus with dusky hairs mesally. Vertex
with dorsum narrow-scaled, these narrow
scales brownish along anterior margin, re-
mainder pale; sometimes a small dark anterior
patch of broad dark scales at lateral margin
of dorsal narrow scales. Scutellar lobes cov-

PACIFIC SCIENCE, Vol. VII, October, 1953

ered with broad silvery-white scales, some
narrow ones may also be present. Postspira-
cular area with narrow yellowish scales. Fore
femur with posterior dark-scaled portion more
reduced than in male. Fore and mid tibiae
pale except for dorsal surface; hind tibia with
a pale longitudinal line on both anterior and
posterior surfaces. Tarsal claws equal, fore
and mid each unidentate, hind simple. Ter-
gites dorsally dark, IV-VII with an indefinite
median brownish-yellow area which may ex-
tend the whole width of the segment; the pale
lateral areas not encroaching dorsally. Stern-
ites with whitish-yellow scales.

LARVA: Head: Antennal shaft with heavy
spicules, shaft darkened except for small basal
area; antennal hair tuft with 7-13 branches.
Mouth brush with comb-like tips. Hair 4 with
5-8 branches, very small; 5 with 6-7; 6 with

4- 6; 7 with 7-11; 8 with 2-4; 9 with 3-5;

12 with 5-7; 13 with 2-4; 14 with 1-2; 15
with 5-9; 17 and 18 single, spine-like; 20 with
2-4. Mentum with about 16-17 small lateral
teeth. Thorax: Prothoradc hair 1 single (con-
siderably longer than 2 or 3) ; . 2 with 2-3
branches; 3 with 3-4. Mesothoracic hair 9
with 8-10 branches; 10 and 12 single; 11 with
1-2, very small. Metathoracic hair 9 with 6-8
branches; 10 single; 11 with 1-2, very small;

12 with 1-2, much smaller than on meso-
thorax. Abdomen: Dorsolateral hair on I with

5- 7 branches; on II with 6-7. Lateral hair on

1 with 3-4; on II and III with 4-7; no dis-
tinctly developed lateral hairs on remainder
of segments. Pentad hair 1 with 6-8 branches;

2 with 2-4 (on same basal tubercle as hair 1);

3 with 15-20; 4 with 2-4; 5 with 5-9- Comb '
with about 16-20 small scales in a patch, each
scale with base broadly ovoid and spine proper
very slender and short, a fringe along most
of basal portion of scale. Siphon long, slender,
acus present, index about 7.0; 12-18 pecten
teeth, the last 2-4 teeth simple and more
widely separated from one another, remainder
darker and each with 1-2 ventral denticles;
siphon hair tuft with 5-7 branches, very small
and pale, inserted at apical one-fourth. Anal

Aedes of the Philippines — Knight and Hull

457

plate incomplete, an acus present; Ih with

2- 4 branches, very small and obscure; isc with
13-17 branches, about equal to anal plate in
length; ventral brush of 12 tufts, all but basal

3- 4 arising from a connected barred area, each
tuft with 11-15 branches. Anal gills long,
lanceolate, dorsal pair slightly longer than
ventral pair and about twice as long as anal
plate.

BIONOMICS: The adult was not encountered
in nature. The larvae were collected from
scattered rain pools in a grassy area, from a
stream margin containing algae and other veg-
etation, and from a grassy pool in the bed of
a temporary stream.

DISTRIBUTION: Specimens examined. U. S.
N.M. Luzon: 4 males, 5 females, 8 sets assoc,
skins, Olongapo, Subic Bay, Zambales Prov.
(Rozeboom, Zolik). One female, 1 assoc,
larval skin, 1 larva, Dau, Pampanga Prov.
(Cowell, Ingal). One male, 6 females, 4 sets
assoc, skins, Quezon City, Manila Prov. (de
Guia, Bruce, Baisas). Six males, 26 females.
Camp Nichols, Rizal Prov. (Carraway, Callon,
Sinclair). Five females, Camp Stotsenberg,
Pampanga Prov. Mindanao: 1 larva, San Ra-
mon, City of Zamboanga Prov. (Knight,
Laffoon.) One female. Camp Gregg, Panga-
sinan. Pettit Barracks, Zamboanga, City of
Zamboanga Prov. (Visaya.) C.A.S. Mindoro:
San Jose (Ross).

Unknown outside the Philippines.

DISCUSSION: As pointed out by Bohart
(1945: 63) this species has been confused by
various authors with niveoscutellum (Theobald)
and alhoscutellatus (Theobald). However, it is
separable from the first on details of male
terminalia and from the second on the char-
acters given in the key.

Aedes (Aedimorphus) alhoscutellatus
(Theobald)

Figs. 3, 4

1905. Lepidotomyia alboscutellata Theobald,
Mus. Nat. Hungarici Ann. 3: 80 (2
females). Type locality: New Guinea.

Simbang, Huon Gulf and Friedrich-
Wilhelmshafen (Biro). Type: Females
(cotypes) in Hungarian National Mu-
seum, Budapest.

1907. Reedomyia alhoscutella Theobald. Theo-
bald, Mon. Cul. 4: 261. Different com-
bination, and lapsus.

1909. Culex argentinotus Banks, Philippine
Jour. Sci. 4: 547 (male, female). Type
locality: Philippines. Pinagsalaan well,
Taytay, Rizal Province, Luzon (Banks).
Type: Male, female (cotypes) in Bu-
reau of Science, Manila. Probably non-
existent since World War 11.

1921. Aedes ~omurensis Yamada, Annot. Zool.
Jap. 10: 73 (males, females). Type lo-
cality: Japan. Omura, Kiushu (Ya-
mada). Type: 2 males, 4 females (co-
types) in Institute for Infectious Di-
seases, Tokyo.

1922. Aedes {Ecculex) alhoscutellatus Theo.
Edwards, Bui. Ent. Res. 13: 101. Syn-
onymy of omurensis.

1922. Aedes {Ecculex) alhoscutellatus, Theobald.
Edwards, Indian Jour. Med. Res. 10:
467. Synonymy of argentinotus.

Adult described by Barraud (1934: 250)
and the larva by Penn (1949: 1).

ADULT: Male. Similar to pampangensis but
differs as follows. Head: Proboscis all dark,
or with a paler band-like area just beyond the
middle, this may be obsolete (except ventral-
ly). Palpi longer than proboscis by slightly
more than length of apical segment. Thorax:
Scutum marked on either side with two small
clusters of yellow scales, one on anterior mar-
gin and the other on a line with the first and
on a level with the scutal angle, a variable
amount of yellowish scales may be scattered
over the scutum. Scutellum with areas be-
tween the lobes free of scales. Apn with a few
broad silvery scales; ppn bare. Propleuron,
upper sternopleuron, lower posterior sterno-
pleuron, and upper mesepimeron each with
small patch of broad silvery-white scales; a
few narrow yellowish scales on the subspira-

458

Fig. 3. A. {Aedimorphus) alhoscutellatus. Male ter-
minalia (Palawan).

cular area. Legs: Femora and tibiae each with
apical patch of silvery=white scales anteriorly
(that on tibiae is largely dorsal); fore and
mid tibiae dark, some pale scaling posteriorly;
hind tibia dark posteriorly. Wing: Wing with
spot of silvery-white scales at base of costal
vein. Abdomen: Tergite I with lateral band of
silvery-white scales; III-VII with basal bands
of brownish-yellow scales, VIII largely pale-
scaled. Sternites with brownish-yellow scales
basally, brown scales apically; VIII covered
with silvery-white scales (since this sternite
is dorsal on most specimens taken, it creates
a very distinctive dorsal abdominal marking),
not possessing distinctive hair tufts. Termin-
alia: Basistyle without lobes; tergal surface
with numerous very small setae, these not
extending to lateral margin setae along distal
portion; sternal surface with a regular longi-
tudinal row of very long hairs near to mesal
margin. Dististyle distinctively shaped as fig-
ured. Ninth tergite strongly concave medially.

Female. Differs from the male mainly as
follows. Proboscis dark. Vertex with dorsum
narrow-scaled, a median anterior diamond-

PACIFIC SCIENCE, Vol. VII, October, 1953

shaped group of these dusky, the remainder
golden; the lateral broad-scaled area creamy-
yellow except for dark patch at dorsal margin
and sometimes one on a level with apn. Apn
with a few narrow yellowish scales; ppn with
a few dark narrow scales dorsally. Subspira-
cular area with a few dusky hairs. Tergites
with a basolateral patch of silvery-white scales,
the basal bands (which are variable in occur-
rence and width) either separate from these, or
only narrowly connected. Sternites largely
covered with brownish-yellow scales, some
apical dusky scales usually present.

LARVA: Similar to pampangensis, but differ-
ing chiefly as follows. Spicules on antennal
shaft sparser and not as heavily developed;
antennal hair tuft with 5-13 branches. Head
hair 5 with 3-4 branches; hair 6 with 3. Men-
tum with 16-18 lateral teeth. Comb with
20-26 scales in a patch, the scales blunt,
fringed laterally and apically and without an
enlarged central spine. Siphon index 3. 5-3. 9,
acus large; 18-21 pecten teeth, all with 1-3
ventral denticles, the distal 1-2 teeth more

Fig. 4. A. {Aedimorphus) alhoscutellatus. Larval ter-
minal segments (Mindoro).

Aedes of the Philippines — Knight AND Hull

459

widely spaced. Only 1-2 tufts of ventral brush
off the barred area basally.

BIONOMICS: The adults were collected while
attempting to bite humans and on several
occasions were netted in densely shaded
woods. Also, females were taken from a ca-
rabao-baited trap. Edwards (1928^?.' 53) re-
ported females taken at a light in Malaya.
The larvae were taken on one occasion from
road ruts through a coconut grove. Penn
(1949: 245) found a larval collection on Min-
doro in a shaded permanent ditch in which
the water was clear, stagnant, and with a
of 6.5. Leicester (1908: 132) collected the
larvae from jungle pools in Malaya.

DISTRIBUTION: Specimens examined. U.S.
N.M. Samar: 1 female, Osmena (Rozeboom,
Zolik). Males, females, San Antonio (Paul-
lus). Palawan: 14 males, 3 females, Bacungan
(Laffoon). Seven females, Puerto Princesa
(Johnson, Laffoon). Mindanao: 1 male, 2 fe-
males, Zamboanga, City of Zamboanga Prov.
(Johnson, Laffoon, Knight). One male, Mer-
cedes, Zamboanga Prov. (Laffoon, Knight).
Jinamoc Island (between Samar and Leyte)
(Medler). C.A.S. Mindoro: 2 larvae, San Jose
(Ross).

Literature records. Luzon: Los Banos, Laguna
Prov. (Bohart, 1945: 62). Mindoro: Camina-
wit Point (Penn, 1949: 1). Calapan (Edwards,

1929: 5).

Outside the Philippines, this species has
been reported from the Solomon Islands, Ad-
miralty Islands, New Guinea, northern Aus-
tralia, Duke of York Island, Sumatra, Java,
Ceram, Borneo, Malaya, India, Assam, Bur-
ma, and Japan.

Aedes (Aedimorphus) mindoroensis
Knight and Hull

1929. A. {Aed) lowisi Theo. Edwards, Notu-
lae Ent. 9: 5.

1934. Aedes {Aedimorphus) sp. Edwards, foot-
note, in Barraud, fauna Brit. India,
Dipt. 5: 250. Called attention to the
fact that material named as lowisi by

Edwards (1929: 5) in actuality repre-
sented another species.

1951. Aedes {Aedimorphus) mindoroensis Knight
and Hull, Pacific Sci. 5: 199 (1 male,
4 females). Type locality: Philippines.
Calapan, Mindoro Island. Type: Male
(holotype) in B.M.f Terminalia sepa-
rated.

DISTRIBUTION: Specimens examined. Type se-
ries. Mindoro: Calapan.

DISCUSSION: A. mindoroensis is most closely
related to lowisi, a species that is known only
from the Andaman Islands and from Morotai
in the Moluccas. Externally, the adult of lowisi
is indistinguishable from that of mindoroensis.
However, the male genitalia of lowisi differs
in having the narrow basal portion of the
dististyle equal to no more than half the total
length and in being evenly slender instead of
rather broadly tapered, in having the short
tergal setae in a rather narrow band along the
mesal margin, and in lacking any special
arrangement of the sternal setae of the basi-
style. Because of the nature of the similarities
between these species, it seems likely that a
single polytypic species is concerned. How-
ever, much more material must become avail-
able before this can be determined definitely.

This species may be distinguished from the
other members of Group C occurring in the
Philippines by its banded tarsi.

Aedes (Aedimorphus) punctifemore
(Ludlow)

Lig. 5

1921. Stegomyia punctifemore Ludlow, Military
Surgeon 49: 69O (1 female). Type lo-
cality: Philippines. Port Wm. McKin-
ley, Rizal Prov., Luzon. Type: female
(holotype) in U.S.N.M.f

Male described by Dyar (1925: 217) and
the female by Edwards, in Barraud (1934: 443) .

ADULT: Scutum with several small spots of
broad silvery scales. Tarsi all dark.

Male. A poor terminalia mount in the U.
S. National Museum is the only existing spec-

460

Fig. 5. A. (Aedimorphus) punctifemore. Dististyle of
male terminalia (Luzon).

imen at present known of the male of this
species. Figure 5 illustrates the dististyle of
this specimen.

Female. Wing length about 3.5 mm. Pro-
boscis dark, paler beneath. Palpus about one-
sixth as long as the proboscis, dark. Torus'
with broad silvery scales medially. Vertex with
some broad dark scales medially, narrow white
scales medially on nape, maybe a few medial
dark narrow scales, and in the type these latter
scales extend to the eyes; remainder of dor-
sum and all of lateral surface broad-scaled,
pale except for two dark-scaled areas; some
dark upright-forked scales on nape. Thorax:
Integument dark. Scutum with fine reddish-
black scales, 2 pairs of small silvery broad-
scaled spots on disc of scutum, a small patch
of similar scales above wing base, one on
either side of prescutellar space and one on
prescutellar space. Scutellum with broad white
scales on each lobe. Apn with some broad
white scales with narrow dark ones above,
some white broad scales below. Following
pleural areas each with a patch of small white

PACIFIC SCIENCE, Vol. VII, October, 1953

broad scales: propleural, subspiracular (2
patches), paratergite, dorsosternopleural, low-
er posterior sternopleural, and mesepimerah
Legs: Femora and tibiae anteriorly dark, with
silvery anterior knee spots and with scattered
broad translucent silvery scales present. Tarsi
dark. Wings: Dark, a small white spot basally
on costa, Flalter knob dark. Abdomen: Tergites
dark, basolateral white spots present.

LARVA: Unknown.

BIONOMICS: Unknown except that females
were taken in a carabao-baited trap on Samar.

DISTRIBUTION: Specimens examined. R.K.L.
Samar: Numerous females, Osmena. U. S.
N.M. Luzon: 1 male terminalia, 15 females,
Camp Nichols, Rizal Prov. (Carraway.) Fe-
male type, Ft. Wm. McKinley, Rizal Prov.

Outside the Philippines, known only from
Bihar, India,

Aedes (Aedimorphus) vexans
nocturnus (Theobald)

Figs, 6, 7

1903. Culex nocturnus Theobald, Mon. CuL
3: 159 (females). Type locality: Fiji
(Hall). Type: 2 females (cotypes) in
B.M.f

1913. Culex nocturnus niger Theobald, Nova
Caledonia 1: 164 (1 specimen). Type
locality: New Caledonia. Canala (Sara-
sin and Roux). Type: Sex unknown;
in Bale Museum.

1946. Aedes {Aedimorphus) vexans nocturnus
(Theobald). Bohart and Ingram, Mosq.
Okinawa and Islands in Central Pacific,
NavMed. 1055, p, 15.

ADULT: Vertex dorsum and scutellum nar-
row-scaled, Scutum with bronzy-brown, yel-
low, and whitish scales intermixed. Hind tarsi
with ba'sal bands on all segments.

Male. Wing length 3. 0-3. 4 mm. Head: Pro-
boscis dark, a poorly defined broad paler area
near the middle. Palpus longer than proboscis
by about length of last segment; II-IV with
basal white bands; numerous long hairs aris-
ing apically on III and laterally along IV-V

Aedes of the Philippines — Knight and Hull

461

(these two segments being down-tilted). To-
rus bare. Vertex dorsum covered with narrow
and narrow-curved p^le scales and dusky and
pale upright-forked scales, lateral portion cov-
ered with broad pale scales, sometimes an
anterior spot of dusky broad scales at the
margin of the narrow-scaled area. Thorax:
Scutum covered with an indefinite pattern of
narrow bronzy-brown, yellow, and whitish
scales. Scutellum with pale narrow scales. Apn
and ppn with narrow and narrow-curved scales,
pale (the dorsal scales on ppn are frequently
brownish, however). The following pleural
areas each with a patch of broad pale scales;
propleural, subspiracular (2 patches), post-
spiracular, paratergite, dorsal sternopleural,
medio-posterior sternopleural, and mesepim-
eral; usually a very few scales on the prealar
knob, particularly in female. Legs: Femora
speckled with pale scales anteriorly, knee
spots present. Tibiae may have some scattered
pale scales anteriorly, apical pale scaling pres-
ent on fore and hind, basal pale scaling pres-
ent at least ventrally on all the tibiae. Fore
and mid-tarsi with first two or three segments
with basal pale bands (may be none on I of
fore tarsus). Hind tarsi with basal pale bands
on I-V. Fore and mid-tarsal claws unequal,
each unidentate; hind claws equal, simple.
Wings: Dark-scaled, a small basal spot of pale
scales may be present on the costa. Abdomen:
Tergite I with a lateral band of pale scales,
II-VII with basal bands, those on the more
posterior segments prolonged laterally. Stern-
ites largely pale-scaled, dark scaling may be
present medially. Terminalia: Basistyle with
a basal setose plaque on mesal membranous
surface; a rather regular longitudinal row of
stout bristles on the sternal surface near the
mesal margin. Dististyle with appendage dis-
tinctly subapical. Ninth tergite concave me-
dially.

Female. Length of wing about 3.5 mm.
Torus with a patch of pale scales medially.
Differs from male chiefly as follows: Palpus
approximately one-fifth as long as the pro-
boscis, apex pale-scaled, may be a few pale

Fig. 6. A. {Aedimorphus) vexans nocturnus. Male ter-
minalia (Samar).

scales medially. Proboscis with pale area larg-
er and more pronounced. Mid-tarsus with
basal pale bands on I-IV, I with some pale
scales medially. Tarsal claws equal, fore and
mid each unidentate, the hind simple. One or
more of tergites IV-VII with apical pale scal-
ing, I-VII with lateral pale bands that are
separate from the basal bands. Wing as in
male, or sometimes possessing considerable
pale scaling along costa and subcosta and
even some pale scales on vein 1.

LARVA: Head: Antenna spiculated, tapering
slightly from base to apex; antennal hair tuft
with 6-10 branches, inserted at about basal
one-third. Mouth brushes with comb-like
tips. Hair 4 very small, with 2-6 branches;
5 single or double (rarely triple); 6 single
(sometimes double); 7 with 5-12; 8 single;
9 with 1-3; 12 with 5-10; 13 with 1-3; 14
with 1-3; 15 with 3-8; 17 and 18 single, 18
occasionally forked; 20 with 3-8. Mentum
with 11-16 lateral teeth. Thorax: Prothoracic
hairs 1 and 2 single; 3 with 2-4 branches.
Mesothoracic hair 9 with 5-9 branches; 10 and

462

PACIFIC SCIENCE, VoL VII, October, 1953

Fig. 7. A. {Aedimorphus) vexans nocturnus. Larva
(Samar), a. Head; b, terminal segments.

12 single, stout; hair 11 very small, with 1-3;
a short acutely tapered spine at base of tuber-
cle. Metathoracic hair 9 with 4-6 branches;

10 single, stout; 12 single, much reduced;

11 very small, with 1-5. Abdomen: Dorso-
lateral hair on I with 3-8 branches; on II
with 3-6. Lateral hairs on I and II with 2-4;
on III-V with 1-4; on VI with 1-3. Pentad
hair 1 with 5-11 branches; 2 and 4 with 1-3;
3 with 5-9; 5 with 6-10. Comb with 8-10
teeth, long, acutely tapered, with attached
portion oval in shape and fringed laterally.

Siphon smooth with a narrowly attached acus,
index 2. 1-3. 2; 13-19 pecten teeth in a row,
the distal two more widely separated, each
tooth except the distal two or three with 1-3
ventral denticles; siphon hair tuft with 3-7
branches. Anal plate incomplete, smooth,
with an attached acus; Ih small, single or
double; isc with 6-9 branches, osc single; ven-
tral brush of 16 (rarely 15-18) tufts with 3 or
4 off the barred area basally, elements of
barred area joined laterally by a bar, tufts of
ventral brush with 4-11 branches. Anal gills
subequal, narrow and gradually tapering to
apex, or broad at base and rapidly tapering
to apex; 1.7 to 2.4 times longer than anal
plate.

BIONOMICS: The adults were commonly
collected while trying to bite humans at night,
and from a light trap. The larvae were fre-
quently encountered in temporary rain-filled
ground- pools and depressions of all types,
but principally from those in grassy areas.

DISTRIBUTION: Specimens examined. R.K.L.
Luzon: Olongapo, Subic Bay, Zambales Prcv.
(Rozeboom, Zolik, McMillan). Samar: Os-
mena (Laffoon, Rozeboom). Palawan: Iwahig
Penal Colony, Puerto Princesa, Bacungan,
Tacburos, and Balsahen River (Johnson, Laf-
foon, Fitzgerald). Mindanao: Zamboanga and
San Ramon (Johnson, Knight, Laffoon). C.

C. Luzon: Sison, Pangasinan Prov. (Francle- |
mont). Baisas Coll. Luzon: Tunkong Manga,
Bulacan Prov. Colo. State U. Coll. Mindoro:

San Jose. (Howe.) i

Literature records. Mindoro: Caminawit '
Point (Penn, 1948: 245). Tunkulan (Edwards, |
1929: 5). Luzon: Manila (Edwards, 1929: 5).^ |

Outside the Philippines, this subspecies is
known from Samoa, Tonga, Fiji, New He- j
brides, New Caledonia, Australia, New
Guinea, Marshalls, Marianas, and Nether- i
lands Indies.

DISCUSSION: Bohart and Ingram (1946: 15) |

resurrected the name nocturnus for the vexans f
of the Australasian region and of the Philip- !|
pines. They consider the material known to j
date from these areas to be subspecifically 1

Aedes of the Philippines — Knight AND Hull

463

distinct on the basis of larval head hairs 5
and 6, which are single to triple and single,
respectively. Typical vexans larvae have these
hairs with 3-5 and 2-4 branches (the latter
occasionally single) . Philippine larvae seen by
us have hair 5 single or double (rarely triple)
and 6 single (sometimes double). Conse-
quently, the name nocturnus is employed in
this paper. Larvae from Europe (Edwards, in
Barraud, 1934: 253), Japan (La Casse and
Yamaguti, 1948: 102), and the United States
(Carpenter, Middlekauff, and Chamberlain,
1946: 232) have the hair branching given
above for typical vexans.

The two cotype females of nocturnus were
compared in the British Museum with the
types (all females) of eruthrosops Theobald
(Ceylon), minuta Theobald (India), and nip-
ponii Theobald (Japan). The only difference
noted was that in nocturnus the wings are dark
except for a small basal patch of pale scales
on the costa. In the other three types, the
wings possess a variable amount of scattered
pale scaling along the costa (as is frequently
the case in Philippine material) .

Edwards (in Barraud, 1934: 253) gives evi-
dence that indicates the probable occurrence
of additional races or subspecies, but, until
much more work is done on this subject, the
status of the Philippine material cannot be
further clarified.

Aedes (Aedimorphus) ostentatio
(Leicester)

1908. Aioretomyia Ostentatio Leicester, CuL
Malaya, p. 193 (females). Type locality:
Malaya. Pahang road (at AVi miles) and
Gombak road (at 5 miles), Kuala Lum-
pur (Leicester). Type: 2 females (co-
types) in BM.f

1910. Pseudohowardina chrysoscuta Theobald,
Mon. Cul. 5: 228 (female). Type local-
ity: Ceylon. Peradeniya (Green). Type:
Female (holotype) in B.M.f

1911. Danielsea pagei 'Ludlow , Psyche 18: 128
(7 females). Type locality: Philippines.

Fort Pikit, Mindanao (Page). Type: 7
females (cotypes) in U.S.N.M.f One
of these specimens here designated
lectotype.

1913. Ochlerotatus ostentatio (Leicester). Ed-
wards, Bui. Ent. Res. 4: 228. Synony-
my of chrysoscuta.

1922. Aedes ? ostentatio Leic. Edwards, Indian
Jour. Med. Res. 10: 468. Synonymy
of pagei.

1925. Aedes (Aedes) ostentatio Leicester. Dyar
and Shannori, Insecutor Inscitiae Men-
struus 13: 78. Different combination.
1928. Aedes ip Aedimorphus) ostentatio (Leic.).
Barraud, Indian Jour. Med. Res, 16:
374. Different combination.

Female described by Barraud (1934: 267).

ADULT: Vertex dorsum partially broad-
scaled. Scutellum narrow-scaled. Scutum with
narrow lines of yellow scaling. Tarsi all dark.

Male. Unknown.

Female. Wing length 3.6 mm. Head: Pro-
boscis dark-scaled. Palpus only about one-
sixth length of proboscis, dark-scaled. Torus
ochreous, with a few fine dark scales and hairs
medially. Vertex dorsum with broad dark
scales, a line of narrow golden scales along
eye margins and a patch of similar scales me-
dially on nape (this narrowing a variable dis-
tance anteriorly, reaching anterior margin in
some specimens), a patch of upright-forked
dusky to golden-colored scales on nape; lat-
eral portion with yellowish broad scales, a
median patch of broad dark scales. Thorax:
Scutum with narrow yellow scales along lat-
eral (broken in region over paratergite) and
anterior margins and along scutal angles, a
narrow median longitudinal band of yellow
scales that forks at the prescutellar area, re-
mainder with dark narrow scales; acrostichal
bristles absent, dorsocentral bristles present.
Scutellar lobes with narrow yellow scales,
sometimes a few median narrow dark scales
on mid-lobe, Apn with a few dark scales above
and a patch of narrow yellow scales below,
ppn bare except for a sparse dorsal line of

464

PACIFIC SCIENCE, VoL VII, October, 1953

narrow dark scaling. Each of the following
pleural areas with a patch of broad pale scales :
propleural, postspiracular (few or none), para-
tergite (few or none), dorsal sternopleural,
ventroposterior sternopleural, and mesepim-
eral. Legs: Fore and mid-femora dark ante-
riorly, hind with anterior surface pale except
for apical fourth and most of dorsal margin,
a line of pale scales anteriorly on apex of hind
and mid femur. Tibiae and tarsi dark. Tarsal
claws equal, fore and mid unidentate, hind
simple. Wings: Dark-scaled. Halter knob with
dusky scaling. Abdomen: Tergites dorsally
dark, basolateral pale scaling on I-VII, this
visible dorsally on II-VI. Sternites III-VII
with basal pale bands.

larva: Undescribed.

BIONOMICS: The single adult from Palawan
was netted in a densely shaded area near a
river. No larvae were found in the Philippines.
Leicester (1908: 193) collected larvae from
jungle pools in Malaya. Further, he reported
the female to be a vicious daytime biter.

DISTRIBUTION: Specimens examined. R.K.L.
Palawan: 1 female, Bacungan (Lalfoon).
C.A.S. Mindoro: 3 females, San Jose (Ross).

Literature records. Mindanao: Ft. Pikit (type
series of ^^gV).

Outside the Philippines, reported from In-
dia, Ceylon, Malaya, Indo-China, Sumatra,
Java, Borneo, and Ceram.

discussion: The Palawan female was com-
pared with Leicester’s two cotype females in
the British Museum. It was found to agree
sufficiently well to be considered conspecific.

Theobald’s type female of chrysoscuta was
examined and found to agree well with the
Palawan specimen except that the pleurae are
noticeably paler in color.

Aedes (Aedimorphus) wainwrighti

Baisas

1946. Aedes {Aedimorphus) wainwrighti^Lis^iS,
Philippine Mon. Bui. Bur. Health 22:
34 (1 female). Type locality: Philip-
pines. Llavac, Infanta, Tayabas Prov.

Luzon (Sunico). Type: Female (holo-
type) in Philippine Bureau of Science,
Manila.

ADULT: A dark-brown species, with a broad
median silvery line on anterior half of scutum,
broad silvery scales on the paratergite, mid-
scutellar lobe, and torus. Hind tarsi banded
with white.

Male. Unknown.

Female. Modified from the original descrip-
tion, Head: Proboscis dark brown, nearly as
long as fore femur. Palpus about one-sixth
length of proboscis, dark brown. Torus dark
brown, with a small patch of broad silvery
scales medially. Vertex broad-scaled, a tri-
angular silvery patch medially that broadens
anteriorly and extends on either side as a short
fine line along the eyes, laterad to this median
area is a dark-scaled spot, the remainder of
the dorsal surface and the entire lateral surface
pale brownish (almost whitish under different
lighting), a median area of narrow dark scales
and a few upright-forked dark scales on the
nape. Thorax: Integument brownish on scu-
tum, darker on pleurae. Scutal scales narrow
dark brown; a broad median silvery line on
the anterior half of the scutum; a pair of
dorsocentral bristles anteriorly and a stronger
pair above level of wing bases, two pairs of
prescutellars, rather few supra-alars. Scutel-
lum with broad scales, silvery on mid-lobe
except for a narrow dark line on posterior
margin, dark on lateral lobes. Apn with a small
patch of silvery flat scales; ppn without scales,
3 bristles present. Each of the following pleu-
ral areas with a patch of broad silvery scales :
propleural, subspiracular, paratergite (cover-
ing entire area), upper sternopleural, medio-
posterior sternopleural, and upper mesepim-
eral. Legs: Dark brown. Femora pale basally
and below, hind with basal half pale both
anteriorly and posteriorly, mid and hind with
a conspicuous silvery spot at apex. Fore and
mid-tarsi with a basal white band on I, re-
mainder dark. Hind tarsus with a basal white
band on I, remainder missing. Wings: Dark.

Aedes of the Philippines — Knight AND Hull

465

Abdomen: Tergites dark dorsally, silvery lat-
eral patches visible above on V-VII: lateral
margin of I entirely silvery, of II-VII broadly
white basally. Sternites II-VII basally pale,
apically dark, segment VIII retractile and hid-
den in VII.

LARVA: Unknown.

BIONOMICS: The single known specimen
was taken in a mosquito trap.

distribution: Known only from the single
type specimen. Luzon: Llavac, Infanta, Taya-
bas Prov.

DISCUSSION: As Baisas pointed out in the
type discussion, this species has a strong re-
semblance to members of Group E {alholinea-
///j-group) of Stegomyia but is separable from
them on the possession of paratergite scaling
and in being brownish rather than black in
general appearance.

In a footnote to the type description, Baisas
stated that, subsequent to the preparation of
the description, two males and several females
of a closely similar species were received from
Mt. Apo, Davao, Mindanao (Hoogstraal). In
these, the hind tarsi had I with basal one-
fifth white, II with the basal one-third white,
III all white except for a narrow ring at apex,
IV-V all dark (paler under certain lights).
The male palpi were slender throughout, with-
out hairs, and upturned apically. The male
terminalia was that of an Aedimorphus.

Until the male terminalia of these species
are available for study, it will not be definitely
possible to place them, but from the available
information it seems likely that they repre-
sent a separate group near Group A {apico-
annulatus-gtoup) . This group is unusual in
that the included species are all tree-hole
breeders.

Subgenus Banksinella Theobald

1907. Banksinella Theobald, Mon. Cul. 4:
468. Genotype: luteolateralis Theobald
(Africa).

ADULT: Male palpi longer than proboscis
by one-half or more of segment IV ; segment

V so reduced as to appear absent, segment
IV strongly upturned, numerous ventrally di-
rected long hairs arising apically on III and
all along IV. Vertex dorsum and scutellum
narrow-scaled. Dorsocentral and acrostichal
bristles present (sparse and short) . Paratergite
bare. Lower mesepimeral bristles present or
absent. Fore and mid-tarsal claws of male un-
equal, each unidentate; hind equal, uniden-
tate or simple. Tarsal claws of female equal,
each unidentate, hind simple in lineatopennis.
Terminalia: Basistyle with a distinct basal lobe.
Dististyle inserted subterminally, with or with-
out articulated appendage. Claspettes absent.
Mesosome divided into lateral plates, each of
which are crenulated or toothed laterally.

LARVA: Similar to Aedimorphus in having the
distal pecten teeth more widely spaced but
differing in lacking an acus on the anal plate.
Antennal hair tuft with 8-11 branches. Head
hair 4 with 5-9 branches; 5 with 4-7; 6 with
3-6. Comb with 6-12 large teeth in an irre-
gular row, each tooth with a sharp spine and
basal fringe. Siphon with an attached acus;
12-24 pecten teeth, each with 1 or 2 strong
basal denticles except for the distal 2-3 which
may be simple, these latter more widely spaced.
Anal plate complete, or narrowly incomplete;
ventral brush of 12-16 tufts, basal 3-7 tufts
off the barred area. Habitat chiefly rain-filled
temporary pools.

DISTRIBUTION: Two species, Edwards

and imprimens Walker, are Australasian, with
the latter also occurring in the Philippines.
One species, lineatopennis (Ludlow), extends
from the Ethiopian region into the Australa-
sian and Oriental regions. The remaining spe-
cies are Ethiopian.

SYSTEMATICS: Closely related to the sub-
genus Aedimorphus, differing most markedly
in the structure of the male palpi.

Aedes (Banksinella) imprimens

(Walker)

Figs. 8, 9

1861. Culex imprimens WdXkts, Linn. Soc. Lon-

466

PACIFIC SCIENCE, Vol. VII, October, 1953

don, Proc. 5: 144. I860 (female). Type
locality: Amboina (Wallace). Type: Fe-
male (holotype) in B.M.f Identified as
type by E. A. Waterhouse.

1901, Culex imprimiens, Walker, Theobald,
Mon. CuL 1: 422. Lapsus.

I9O8. Culex Auratus Leicester (nec Grabham,
1906), CuL Malaya, p. 153 (female).
Type locality: Malaya. Type: No in-
formation. Not in B.M.

1913. Ochlerotatus imprimens (Walker). Ed-
wards, BuL Ent. Res. 4: 228. In part.
Questioned synonymy of auratus (no
material seen). Different combination.
1946. Aedes {Banksinella) imprimens (Walker).
Hsiao and Bohart, Mosq. Japan, Nav-
Med 1095, p. 22. Different combina-
tion.

Larva is described by Bohart (1945: 63);
systematics treated by Edwards, in Barraud
(1934: 257) and Hsiao and Bohart (1946: 22).

ADULT: A large yellowish-brown species.
Hind tarsal segments all basally banded.

Male. Wing length about 5.0 mm. Head:
Proboscis dark-scaled. Palpus longer than
proboscis by over one-half length of terminal
long segment, dark-scaled. Torus bare. Vertex

Fig. 8. A. {Banksinella) imprimens (Dutch New
Guinea). Male terminalia (Dutch New Guinea). Para-
proct omitted on right.

dorsum with narrow golden scales, dark up-
right-forked scales posteriorly and pale ones
anteriorly; lateral portion of vertex with broad
pale scales. Thorax: Scutum brown, rather
thinly covered with hair-like brownish-gold
scales, paler over wing bases, and on pre-
scutellar area. All scutellar lobes with narrow
yellow scales. Apn with broadened and mar-
row yellow scales; ppn with narrow yellow
scales. Each of the following pleural areas
with a patch of broadened whitish scales:
subspiracular, dorsal sternopleural, ventro-
posterior sternopleural, and mesepimeral (pin
through postspiracular area). Legs: Fore and
mid-femora dark anteriorly, some scattered
pale scales ventrally, and pale scales at apices;
hind femur pale-scaled anteriorly except for
a dark apical area, dark scales on dorsal sur-
face, a ring of pale scales on extreme apex
anteriorly. Tibiae dark. Fore tarsi dark except
for a few pale scales basally on II; mid-tarsi
with narrow basal pale bands on I-II; hind
tarsi with narrow basal pale bands on I-V,
broadest on I and narrower on each succeed-
ing segment. Fore and mid-tarsal claws un-
equal, each unidentate; hind tarsal claws equal,
each unidentate. Wings: Dark-scaled, wing
membrane with a pronounced yellowish iri-
descence. Halter knob dark. Abdomen: Terg-
ites dark, lateral margin of I with a band of
pale scales, III-VII with narrow basal ochreous
bands. Sternites pale-scaled, an apical line of
dark scales on most of them. Terminalia:
Drawing and description from a New Guinea
specimen. Basistyle short, broad; a prominent
subbasal lobe present, this bearing numerous
very stout setae. Dististyle broad, deeply di-
vided apically into a pair of unequal arms, the
shorter of these bearing several setae. Ninth
tergite broadly divided medially.

Female. Similar to the male but differing as
follows. Palpus approximately one-fifth as
long as proboscis, dark. Torus with a few dark
hairs and scales medially. Vertex with a medio-
anterior area of dark narrow scales and with
a patch of broad dark scales at lateral margin
of dorsal narrow-scaled area. Scutum with a

Aedes of the Philippines — Knight and Hull

467

Fig. 9. A. {Banksinella) imprimens. Larva (Dutch New Guinea), a. Head; h, thorax, dorsal aspect; c, ter-
minal segments.

variegated pattern of blackish-brown and
golden scales. Ppn with some dark scales in-
terspersed dorsoanteriorly. Propleuron and
postspiracular area also each with a scale
patch. Fore and mid-tarsi with small basal
white patches on I-III, hind tarsi as in male.
Tarsal claws equal, unidentate. Base of costa
with a few pale scales. Tergites II-V with
basal ochreous bands, a dorsobasal median
pale patch on VI, nearly complete lateral
ochreous bands on I-VII. ■

LARVA: Description based on a whole larva
from Toem (E. S. Ross), and a larval skin from
Hollandia, both of Dutch New Guinea, and
a whole larva and larval skin from Bougain-
ville, Solomon Is. (A. B. Gurney), all of which
are in the U.S.N.M. No larval specimens from
P. I. available. Head: Integument with a fine
droplet-like rugosity. Antenna largest at base
and tapering slightly toward apex, with prom-
inent spicules over most of length; antennal
hair tuft with 8-11 branches. Hair 4 with 7-9
branches; 5 with 4-7; 6 with 3-5; 7 with 9-13;
8 double (once single); 9 with 2-3; 12 with
8-10; 13 and 14 with 2-3; 15 with 5-8; 17
and 18 single (rarely forked at tip); 20 with
5-9. Mentum with 20-24 lateral teeth. Thorax:
Meso- and metathoracic 9-12 hair groups
arising from large heavily sclerotized tuber-
cles, each tubercle also bearing a strong blunt

spine; mesothoracic hair 9 with 6-7 branches.
Abdomen: Dorsolateral hair on I with 3-6
branches; on II with 4-5. Lateral hair on I
with 2-3; on II with 4-6; on III with 5; on
IV-VI with 2-4. Pentad hair 1 with 7-13
branches; 2 with 1-2; 3 with 6-11; 4 with
2-3; 5 with 5-11. Comb with 10-12 teeth in
an irregular row, each tooth large with a sharp
spine and lateral fringe on the attached por-
tion. Siphon brown, acus attached, index 3.2-
3.5; 19-24 pecten teeth, each with 1-2 strong
basal denticles except the distal 2-3 which
are simple, these latter teeth more widely
spaced; siphon hair tuft moderate, with 4-7
branches. Anal plate complete, with a number
of short acutely tapered spicules along dorso-
lateral margin (one specimen also had a small
round detached acus) ; Ih single (once double),
shorter than length of anal plate; tsc with 7-11
branches; oic with 3-5 branches. Ventral brush
of 16-17 tufts, basal 4-7 tufts off barred area
and piercing anal plate, barred area not con-
nected laterally. Anal gills (one specimen)
subequal, dorsal pair 2.3 times longer than
anal plate.

BIONOMICS: No specimens were collected
by us in the Philippines. However, females
were collected in New Guinea and the Solo-
mons while attempting to bite humans. A
single male was netted in deep jungle in New

468

Guinea. The females are vicious biters and
are present in great numbers shortly after the
beginning of isolated rainy periods. Gurney
(Bohart, 1945: 63) collected larvae on Bou-
gainville Island from shaded leafy woodland
pools. Causey (1937: 413) reported larvae
from a buffalo wallow in Siam. Dr. E. S. Ross
has reported to us in personal correspondence
that he collected the larvae on Mindoro in
shaded temporary puddles.

DISTRIBUTION: Specimens examined. U.S.
N.M. Palawan: 4 females, Puerto Princesa
(McGregor). Mindanao: 1 female, Ft. Pikit
(Spry).

Literature records. Mindanao: Port Banga
(Edwards, 1929: 5).

Also, the above unpublished record from
Dr. E. S. Ross of San Jose, Mindoro.

Outside the Philippines reported from Ja-
pan, Siam, Malaya, Sumatra, Java, Borneo,
Ceram, Amboina, New Britain, New Guinea,
and the Solomons.

DISCUSSION: The type has been studied in
detail. The head and hind tarsal segments
III-V are completely missing, and the scutum
is rather badly rubbed. The remainder of the
specimen is in sufficiently good condition to
make out all the characters. A female specimen
from Cyclops Mts., Hollandia, Dutch New
Guinea (Laffoon) in the Knight collection
was compared with the type and found to
agree in all details. A male which was cap-
tured at the same time as this female, and
which agrees very well with it, is believed to
represent the male of imprimens. This pair of
specimens was used for the above adult de-
scription. Three wild-captured females from
Pavuvu, Russell Group (Downs) and one
from Guadalcanal (Downs) in the Knight
collection agree in all details with the New
Guinea female.

The larva has not previously been described
except for some brief notes by Bohart (1945:
63) from larvae taken on Bougainville Island,
Solomons (Gurney).

Apparently, no authentic material of aura-
tus Leicester exists, but based on Leicester’s

PACIFIC SCIENCE, Vol. VII, October, 1953

description (female only) this species agrees
very well with the description of imprimens
given above.

The Dutch New Guinea male specimen
referred to previously was compared to the
type (male) of A. (B.) hrugi Edwards in the
British Museum. The two were found identi-
cal in all details, including terminalia, except
that the hrugi specimen was distinctly smaller
(wing length described as 3.8 mm. in original
description) . This is also true of a single male
in the Knight collection that was reared from
larvae collected in a road rut at Lake Sentani,
Hollandia, Dutch New Guinea. This spec-
imen was also compared to the type of hrugi
and found to agree perfectly. It is believed
that hrugi is a synonym of imprimens, but,
until evidence is forthcoming that the size
difference is inconsequential,, it is thought
better to hold the two species separate.

Aedes (Banksinella) lineatopennis
(Ludlow) •

Figs. 10, 11

1901. Culex luteolateralis Hhtobdld (type male
only), Mon. Cul. 2: 71 (males, females).
Type locality: Africa. Durban, Natal
(Christophers). Salisbury, Rhodesia
(Marshall). Malaya. Straits Settlements,
Perak (Wray). Type: Male (lectotype)
in B.M.

1905. Taeniorhynchus lineatopennis Ludlow,
Canad. Ent. 37: 133 (females). Type
locality: Philippines. Camp Gregg, Ba-
yambang, Pangasinan Prov. Luzon
(Chamberlain). (However, the type la-
bel has "Camp Gregg, Angeles, Pam-
panga Province.”) Type: 3 females (co-
types) in U.S.N.M.f The specimen
bearing the locality label is here des-
ignated lectotype.

1913. Pseudohowardina line alts Taylor, Rpt.
I9II5 Austral. Inst. Trop. Med., p. 10
(female). Type locality: Australia.
Ching Do and Townsville, Queens-
land. Type: No information.

Aedes of the Philippines — Knight AND Hull

469

1915. Banksinella lineatopennis, Ludlow. Ed-
wards, Bui. Ent. Res. 5: 274. Different
combination. Restricted luteolateralis to
type female and applied lineatopennis to
type male.

Adult and larvae described by Barraud
(1934: 269).

ADULT: A small dark species readily recog-
nized by the broad longitudinal lateral area
of yellow scaling on the scutum.

Male. Wing length about 2.6 mm. Head:
Proboscis dark. Palpus longer than proboscis
by nearly the length of the terminal segment;
dark. Vertex dorsum with a large central area
of yellow narrow and upright-forked scales.
Thorax: Scutum with a broad .area of yellow
scales along the. lateral margin, central por-
tion dark-scaled, some dark scales present be-
yond the yellow scales on the lateral margin
over the spiracle; dorsocentral, acrokichal,
and prescutellar bristles present. Mid-scutellar
lobe with narrow yellow scales, sometimes a
few dusky scales apically; lateral lobes essen-
tially bare of scales. Apn bare of scales; ppn
with a few narrow dark scales. Pleuron nearly
bare of scales, a few usually being present on
the propleuron, dorsally and medioposteriorly
on the sternopleuron, and on the mesepim-
eron; 1-2 lower mesepimeral bristles present.
Legs: All dark. Fore and mid-tarsal claws un-
equal, each unidentate; hind claws equal,
simple. Wings: Dark, pale scales along vein
1 to junction with stem of vein 2 and along
basal portion of vein 5. Abdomen: Tergite I
dark-scaled dorsally; II-VII with dorsal basal
pale bands. Sternites dark. Terminalia: Basi-
style elongate; with numerous hairs along in-
ner surface, a row of short peg-like setae
interspersed among these apically; with a pro-
jecting basal lobe that bears several strong
apical spines. Dististyle inserted before apex,
prominently widened just beyond middle.
Ninth tergite deeply concave medially.

Female. Differs from male chiefly as follows.
Palpus approximately one-seventh as long as
the proboscis. Scutum with dark scales laterad

Fig. 10. a. {Banksinella) lineatopennis. Male termina-
lia (Samar).

of the yellow-scaled band from the anterior
margin to the level of the wing base. Lateral
scutellar lobes with a few narrow yellow scales.
Tarsal claws equal, unidentate on fore and
midlegs, simple on hind. Some of the sternite
scaling appearing paler. Wing with almost all
of veins Sc and 5 pale-scaled also, some pale
scales may be present basally on vein 4.

LARVA: Head: Antenna stocky, broadest at
base and tapering somewhat toward apex;
prominent spicules over most of length; an-
tennal hair tuft with 5-10 branches, inserted
before middle. Mouth brushes with comb-
like tips. Hair 4 small, with 5-9 branches; 5
with 4-7; 6 with 3-6; 7 with S-13; 8 with
2-3; 9 with 1-5, usually 3-4; 12 with 4-7;
13 and 14 with 2-5; 15 with 5-10; 17 and 18
usually single, occasionally double; 20 with
6-10. Mentum with 17-20 lateral teeth. Tho-
rax: Prothoracic hair 1 single or double; 2
stalked, with 1-3 branches; 3 stalked, with
2-5; the whole group small in size. Meso-

470

Fig. 11. a. {Banksinella) lineatopennis. Larva (Samar).
Head; terminal segments.

thoracic hair 9 with 6-10 stout branches; 10
and 12 single, stout; 11 not seen. Metatho-
racic hair 9 with 3-4; 10 single; 12 much
reduced, with 1-2; 11 not seen. Abdomen:
Dorsolateral hairs on I and II with 2-4
branches. Lateral hair on I single or double;
on II with 2-7; on III-VI single, rarely with
2-3. Pentad hair 1 with 4-8 branches; 2 and
4 with 2-3; 3 with 8-11; 5 with 4-9. Comb
with 6-8 (usually 8) teeth, long, acutely ta-
pering, attached portion oval with lateral

PACIFIC SCIENCE, Vol. VII, October, 1953

fringe. Siphon smooth with a narrowly at-
tached acus, index 2. 1-2. 5; 12-16 pecten
teeth in a straight row, the distal two or three
more widely spaced, the teeth usually with
just one ventral denticle; siphon hair tuft with
2-7 branches, the tuft no longer than the
distal pecten tooth. Anal plate incomplete,
smooth; Ih single or double; he with 8-15,
osc single; ventral brush of 12 tufts with 3-5
off the barred area basally, the tufts with 4-12
branches. Anal gills subequal, narrow, taper-
ing to a slender point, 2. 7-3.0 times longer
than the anal plate.

BIONOMICS: The adults were taken in light
traps. Nine larval collections were made, all
in temporary, rain-filled, grassy ground de-
pressions. According to Edwards (1941: 201),
this species habitually feeds upon human
blood,

DISTRIBUTION: Specimens examined. R.K.L.
Luzon: Olongapo, Subic Bay, Zambales Prov.
(Rozeboom, McMillan). Samar: Osmena
(Laffoon, Rozeboom). Mindanao: Zambo-
anga, City of Zamboanga Prov. (Johnson,
Knight, Laffoon). C. C. Luzon: Sison, Pana-
gasinan Prov. (Franclemont) . Baisas Coll.
Luzon: Tunkong Manga, Bulacan Prov.
U.S.N.M. Jinamoc Island (nr. Samar) (Med-
ler). Mindanao: Males, females, Parang (Pal-
lus). Pettit Barracks, Zamboanga (Visaya).
Luzon: Males, females. Camp Nichols, Rizal
Prov. (McDonald). Camp Stotsenberg, Pam-
panga Prov. S. Fernando, LaUnion Prov,
(Shields.) Leyte: Males, females, Abuyog
(Graham). One larva, Dulag (Starkey). C.A.S,
Mindoro: San Jose (Ross),

Literature records. Luzon: B'ayambang, Los
Banos, Camp Nichols, Camp Stotsenberg
(Bohart, 1945: 66). Mindoro: Calapan. Luzon:
Manila (Edwards, 1929: 5).

Outside the Philippines this species is known
from Africa, India,- Ceylon, Andamans, Siam,
Malaya, Sumatra, Java, Borneo, Celebes, Ka-
baena, Soemba, Timor, Amboina, and
Queensland.

Aedes of the Philippines — Knight and Hull

471

Subgenus Aedes Meigen s. str.

1818. Aedes Meigen, Syst. Beschr. 1: 13.

Genotype: cinereus Meigen (Europe,

North America).

ADULT: Dark brownish or reddish species
with little or no ornamentation, tarsi all dark.
Palpi short in both sexes. Vertex dorsum
mostly broad-scaled. Scutellum narrow-scaled.
Patches of broad white scales on propleuron,
upper and posteromedian portions of sterno-
pleuron, and on upper part of mesepimeron.
Paratergite not scaled. Lower mesepimeral
bristles absent but mesepimeron often with
fine hairs posteriorly. Legs dark. Male with
tarsal claws of fore leg unequal or equal, both
toothed, midlegs with claws unequal or equal
and with all combinations of being simple
or toothed, hind legs equal and toothed or
simple. Lemale tarsal claws equal, toothed,
those on hind tarsi simple in some species.
Tergite I with white scales along lateral bor-
der. Terminalia: Basistyle short, stout, no true
basal lobe bearing various strikingly modi-
fied apical and subapical structures. Disti-
style extremely various in form, sometimes
bearing hairs but never with a true appendage
present. Phallosome various but always di-
vided into a pair of plates or rods. Claspettes
absent. Various striking processes may be de-
veloped from the ninth or tenth segments.
Lemale terminalia specifically modified.

LARVA: Similar to Banksinella. Antennal
hair tuft single or branched (2-10). Head hair
4 with 2-7 branches; 5 single, or with 2-10;
6 single, or with 2-8. Comb with 8-14 scales
arranged in regular or irregular row. Siphon
with attached acus; distal pecten teeth usually
more widely spaced. Anal plate incomplete;
ventral brush with 10-15 tufts, all but basal
1-4 arising from a laterally connected barred
area. 'Habitat a wide variety of fresh- and
brackish-water ground pools, temporary or
permanent.

DISTRIBUTION: Approximately 68 species
are now recognized in the subgenus Aedes,
of which the majority occur in the Australa-

sian and Oriental regions. Thirteen species
are known from the Philippines.

SYSTEMATICS: To date it has not been pos-
sible to divide the known species into any
logical system of subgroups. Lor a detailed
treatment of the known Philippine species,
see Laffbon (1946).

Aedes (Aedes) robertsi Laffoon

1946. Aedes {Aedes) robertsi Laffoon, Jour.
Wash. Acad. Sci. 36: 230 (males, fe-
males, larvae). Type locality: Philip-
pines. Tacloban, Leyte (Roberts) . Type:
Male (holotype) in Academy of Natural
Sciences of Philadelphia.

DISTRIBUTION: Literature records. Known
only from type series.

Aedes (Aedes) hamistylus Laffoon

1946. Aedes {Aedes) hamistylus Laffoon, Jour.
Wash. Acad. Sci. 36: 232 (males, fe-
males, pupae, larvae). Type locality:
Philippines. Cape Melville, Balabac
(Laffoon). Type: Male (holotype) in
U.S.N.M.

DISTRIBUTION: Literature records. Known
only from type series.

DISCUSSION: This species is apparently in-
distinguishable from robertsi except for the
following characters. Male with claws of fore
leg unequal, both toothed; claws of midleg
unequal, the larger one simple, the smaller
one toothed. Apical prolongation of basi-
style more rounded than in robertsi, base of
its inner posterior projection larger, with only
a short blunt projection arising from it. Basi-
style with two bristles close to outer base of
prolongation, but with an area free of bristles
for some distance anterior to these, a total of
about 10 bristles on the basistyle. Lemale
terminalia with the regions of densest sclero-
tization in the postatrial area differing. No
means of separating the larva from that of
robertsi has been found. The larvae of robertsi
and of this species are unusual in that the

472

PACIFIC SCIENCE, VoL VII, October, 1953

anal papillae are extremely long and possess
prominent tracheation. This rather unusual
structural detail is apparently correlated with
their ability to remain nearly indefinitely at
the bottom.

Aedes (Aedes) dux Dyar and Shannon

1925. Aedes {Aides) dux Dyar and Shannon,
Insecutor Inscitiae Menstruus 13: 81
(male). Type locality: Philippines.
Type: Male (holotype) in U.S.N.M.
1928. Aedes {Aedes) sigmoides Barraud, Indian
Jour. Med. Res. 16: 373 (female). Type
locality: Andaman Islands. Port Blair
(Christophers). Type: Female (holo-
type) in B. M.j Terminalia mounted.
1937. Aedes {Aedes) dux Dyar and Shannon.
Causey, Amer. Jour. Hyg. 25: 414.
Synonymy of sigmoides.

1945. Aedes {Aedes) macrodixoa Dyar and
Shannon. Bohart, Synopsis of Philip-
pine Mosq., NavMed 580, p. 65 and
figure 52 (female only).

Adult and larva described by Laffoon (1946:
233).

DISTRIBUTION: Specimens examined. C.A.S,
Mindoro: 2 females, 1 set assoc, skins, 1 larva.
San Jose (Ross). C. C. Luzon: 19 females,
Santo Tomas, LaUnion Prov. (Franclemont).

Literature records. Luzon: Camp Nichols and
Las Pinas, Rizal Prov. Tungkong Manga, San
Jose, Bulacan Prov. Manila, Manila Prov.
Subic Bay, Zambales Prov. Camp Stotsenberg,
Pampanga Prov. Mindoro: Caminawit Point.
Mindanao: Zamboanga, City of Zamboanga
Prov. (Laffoon, 1946: 235.)

Outside the Philippines this species is
known from Java, Malaya, Siam, Andaman
Islands, and Indo-China.

DISCUSSION: Laffoon (1946: 235) ques-
tioned the synonymy of sigmoides by Causey
because Barraud’s original description stated
that the type had small, lateral, pale markings
on the abdominal tergites. However, the se-
nior author has had the opportunity of com-
paring Philippine specimens with the type of

sigmoides, and there seems to be no doubt
but that they are conspecific. Philippine spec- t
imens show paler brown areas basolaterally |
on some tergites, and in some specimens these :
areas even show a grayish color as does the
type of sigmoides.

In general appearance dux most closely re-
sembles rohertsi and hamistylus. However, it
may be readily distinguished from them be-
cause of the hind tarsal claws, which are sim-
ple in the former and toothed in the latter two.

Aedes (Aedes) johnsoni Laffoon 1

1946. Aedes {Aedes) johnsoni Laffoon, Jour. |
Wash. Acad. Sci. 36: 235 (males, fe- |
males, pupae, larvae). Type locality:
Philippines. Pasanonco, City of Zam-
boanga Prov., Mindanao (Johnson). '

Type: Male (holotype), with associated |

larval and pupal skins, in U.S.N.M.

DISTRIBUTION: Literature records. Known
only from the type series. Leyte: Tanauan.
Mindanao: Pasanonco, San Ramon, and
Zamboanga Prov. Basilan: Isabela.

DISCUSSION: Aedes johnsoni, margarsen, adus-
tus, and andamanensis (not recorded from the
Philippines) make up a complex that is prob-
ably either polytypic or superspecific in form.

Aedes (Aedes) adustus Laffoon

1946. Aedes {Aedes) adustus Laffoon, Jour.
Wash. Acad. Sci. 36: 236 (males, fe-
males). Type locality : Philippines. Cape
Melville, Balabac Island (Howell).
Type: Male (holotype) in U.S.N.M.

DISTRIBUTION: Literature records. Known
only from the type series.

DISCUSSION: In the adult stage this species
is very similar to johnsoni and margarsen, dif-
fering only in details of the male terminalia.

The larva is undescribed.

A very unusual specimen was taken as a
larva in the pool from which part of the type
series of adustus was taken (Laffoon, 1946:
236). It is inseparable from the male of adustus

Aedes of the Philippines — Knight and Hull

473

on external characters, but the terminalia are
asymmetrical (Laffoon, 1946, fig. 14), One
side is rather similar to adustus, while the other
is very odd, showing some similarity to cam-
pylostyhis and some to adustus. The larvae of
campylostylus , hamistylus, and uncus were col-
lected from this same pool. This specimen is
in the U.S.N.M.

While at the British Museum in 1946, the
senior author compared a Philippine male
specimen of adustus with the holotype male
of andamanensts Edwards and made a note to
the effect that the two were conspecific. Word
of this was sent to Laffoon, and, after reviewing
the problem, he replied that he had used the
figure given by Barraud (1934, fig. 71g) for
his concept of the male terminalia of anda-
manensis. This figure shows three apical pro-
cesses in addition to the tergal large one and
is accordingly similar to johnsoni (differing,
however, in the shape of the anterior fork
elements of the dorsal large prolongation);
and probably represents an undescribed spe-
cies. No specimen that could have been used
for this description was located in the col-
lection of the B.M., so perhaps the specimen
in question is still in the collection at Kasauli.

Barraud (1934: 290) recorded seeing spec-
imens of andamanensts from the Andamans,
Assam, E. Bengal, and the Malabar Coast.
Four of Barraud’s specimens from Assam are
in the B.M., and an examination of the term-
inalia of a male in this series showed it to be
unlike his figure and completely similar to
the type of andamanensts in lacking the third
of the three apical processes below the large
tergal prolongation. Therefore, Barraud’s fig-
ure must have been made from either E. Ben-
gal or Malabar Coast specimens. However,
the identity of his species must remain un-
known until someone has the opportunity
of re-examining his material that is still at
Kasauli.

To return to the possibility of adustus being
a synonym of andamanensts, Laffoon pointed
out (in personal communication) that Ed-
wards’ (1922, fig. 37) original figure showed

a series of short hairs along the tergal-lateral
margin of the basistyle, whereas adustus pos-
sesses a group of about 10-15 long bristles
tergal-laterally on the basistyle, and these are
all arranged apically at the base of the tergal
prolongation. No such group is shown in the
figure given by Edwards. Other differences
pointed out by Laffoon that might serve to
separate the two species are the different shape
of the anterior fork of the large apical pro-
longation of the basistyle and the different
form of the group of stout spines on the inner
basal surface of the basistyle. Unfortunately,
Lafifoon’s letter did not arrive until after the
departure of the senior author from the B.M.,
and these possible points of difference have
not been checked. So, for the present, adustus
has been maintained as a valid species.

Aedes (Aedes) margarsen Dyar and
Shannon

1925. Aedes {Aedes) margarsen Dyar and Shan-
non, Insecutor Inscitiae Menstruus 13:
80 (males, females). Type locality:
Philippines. Camp Eldridge, Laguna
Prov., Luzon (Vasquez). Type: Male
(lectotype) in U.S.N.M.

Adult and larva described by Laffoon (1946:
237).

DISTRIBUTION: Literature records. Luzon:
Camp Eldridge, Los Banos, and Calamias,
Laguna Prov. Tungkong Manga, San Jose,
Bulacan Prov. Subic Bay, Zambales Prov. San
Anastacio. Mindoro: San Jose. (Laffoon,
1946: 237.)

Unknown outside the Philippines.

DISCUSSION: The adult of this species is
distinguishable from andamanensts, johnsoni,
and adustus only in details of the male ter-
minalia. The larva is apparently inseparable
from the larva oi johnsoni.

Aedes (Aedes) nubicolus Laffoon

1945. Aedes {Aedes) uncus (Theobald). Bohart,
Syn. Philippine Mosq., NavMed 580,
p. 66 and fig. 50. Male only.

474

PACIFIC SCIENCE, Vol. VII, October, 1953

1946. Aedes {Aedes) nubicolus Laffoon, Jour.
Wash. Acad. Sci. 36: 237 (2 males).
Type locality: Philippines, Sibulan Riv-
er, at 7,000-8,000 feet on Mount Apo,
Cotabato Prov., Mindanao (Clagg).
Type: Male (holotype) in Museum of
Comparative Zoology, Cambridge,
Mass.

DISTRIBUTION: Literature records. Known
only from the type series.

DISCUSSION: This species, which is known
only from the male, is apparently indistin-
guishable externally from macrodixoa and un-
cus. However, the male terminalia is distinct
from that of all species known.

Aedes (Aedes) uncus (Theobald)

1901. Ctdex uncus LAstchAd., Mon. Cul. 2: 53
(2 females). Type locality: Malaya.
Klang, Selangor (Butler). Type: 2 fe-
males (cotypes) in B.M.j Terminalia
mounted. These two specimens are not
conspecific. Therefore, at this time the
name uncus is restricted to the specimen
bearing the following penciled label:
"Klang. Among plantains.” The other
female, labeled "Klang jungle. IX. 99”
in ink on the back of the card support-
ing the pin, is an unnamed species.
1908. Aioretomyia Varietas Leicester, Cul. Ma-
laya, p. 185 (1 male, 2 females in type
series). Type locality: Malaya. Kuala
Lumpur (Leicester). Type: 1 male, 2
females (cotypes) in B.M.f Both male
and female terminalia mounted. The
two females are conspecific with uncus
(new synonymy). The male is a dis-
tinct species, and at this time the name
varietas is restricted to this male spec-
imen.

Adult described by Laffoon (1946: 237).

DISTRIBUTION: Literature records. Culion.
Palawan: Puerto Princesa. Irahuan River Val-
ley. Balabac Island: Cape Melville. (Laffoon,
1946: 238.)

Known outside the Philippines only from
Malaya.

The Mt. Apo, Mindanao, record of Bohart
(1945: 66) was based upon the type specimens
of nubicolus.

DISCUSSION: The two specimens (Edwards,
1917: 223, stated that there was only a single
type specimen) used by Theobald in describ-
ing this species have been examined by the
senior author and found not to be conspecific.
As pointed out in the synoptic table above,
the specimen bearing the label ' 'Klang. Among
plantains.” has been selected as the type for
uncus. The other specimen apparently rep-
resents an undescribed species or at least one
in which the female stage is unknown.

The specimen selected as type has a rather
poor terminalic mount, but it was possible
to see all of the structures except the preatrial
sclerite and the preatrial plate. In all details
seen, the terminalia compared well with a
female terminalia of what has been considered
to be uncus in the Philippines by Laffoon
(1946: 237). However, externally this type
specimen differs slightly, as follows: only 7-9
hairs on the anterior part of the sternopleuron,
and about 20 fine hairs scattered all along the
posterior margin of the sternopleuron. This
difference in the number of hairs on the an-
terior portion of the sternopleuron is suffi-
cient to keep this specimen from keying to
uncus in the key to the Philippine species of
the subgenus Aedes prepared by Laffoon
(1946: 229).

The other female type specimen differs
completely from uncus as treated here in lack-
ing anterior sternopleural hairs altogether and
in having only up to 4 hairs on the mese-
pimeron below the dorsal hair tuft '(these
occur posterior to the scale patch). The ter-
minalia of this specimen resembles that of cam-
pylostylus except that there are apparently no
postatrial hairs. However, the slide is very
faded, and it was only possible to say that
it was not uncus as figured by Laffoon (1946,
hg. 21).

Edwards (1917: 223) arbitrarily associated
two Malayan male specimens from Leicester’s
collection with Theobald’s types of uncus. Ex-

Aedes of the Philippines — Knight AND Hull

475

ternally, these two specimens appear unques-
tionably to be conspecific with the specimen
selected above as the type for uncm. They
differ externally from Philippine uncus in the
same manner described above for the female.
Also, the terminalia, as figured by Edwards
(1917, fig. 8d) shows some minor differences.
Unfortunately, no notes were made of these
differences so it is not possible to evaluate
them now.

In summary, it seems likely that the Philip-
pine material may eventually be shown to be
either subspecific to uncus, or a separate, close-
ly related species. On the other hand, more
extensive collecting in Malaya and the ad-
joining portions of the Philippines may show
the variations mentioned to be normal and
intraspecific. This is the only fresh-water spe-
cies of the Philippines with a distribution
elsewhere, and it seems likely that this may
be additional evidence to show that the Phil-
ippine form is a distinct and endemic form.

In examining the types of varietas Leicester,
it was noted that externally the two female
cotypes are similar to the female selected as
type for uncus, in both general and specific
characters. In these, the number of hairs on
the anterior portion of the sternopleuron was
slightly higher (and thus more intermediate
with the Philippine material), being 10-12 in
number. Although the terminalic mount avail-
able was not too good, it appeared to resem-
ble completely that of the female type of
uncus.

The male cotype of varietas is not con-
specific with the females and is here regarded
as the type for that species. It is a smaller
species than uncus and has a general grayish-
brown appearance, very small basolateral
spots on the tergites, and delicate, thin hind
tarsal claws.

Aedes (Aedes) macrodixoa
Dyar and Shannon

1925. Aedes (Aedes) macrodixoa Dyar and
Shannon, Insecutor Inscitiae Men-
struus 13: 79 (1 male). Type locality:

Philippines. Infanta, Tayabas Prov.,
Luzon. Type: Male (holotype) in
U.S.N.M. Terminalia mounted.

Adult and systematics treated by Laffoon
(1946: 238).

DISTRIBUTION: Literature records. Luzon: In-
fanta, Tayabas Prov. Mindoro: San Jose. Ley-
te: Tacloban. Mindanao: Mercedes, City of
Zamboanga Prov. Parang. (Laffoon, 1946:

239.)

Unknown outside the Philippines.
DISCUSSION: This species is apparently in-
distinguishable externally from uncus and nu-
hicolus. Laffoon (1946: 239) has pointed out
that the female which Bohart (1945, fig. 52)
identified and figured as macrodixoa is in ac-
tuality dux.

According to King and Hoogstraal (1947:
125), the male from the upper Digoel River
in southern Dutch New Guinea that was
identified by Brug (1932: 79) as macrodixoa
was probably neomacrodixoa King and Hoog-
straal.

Aedes (Aedes) nigrotarsis (Ludlow)

I9O8. Pseudoskusea nigrotarsis Ludlow, Canad.
Ent. 40: 52 (female). Type locality:
Philippines. Infanta, Tayabas Prov.,
Luzon (Warriner). Type: Female (holo-
type) in U.S.N.M.

I9O8. Pseudoskusea medioUneata Ludlow. Ca-
nad. Ent. 40: 332. Misidentification.
Adult, larva, and systematics treated by
Laffoon (1946: 239).

DISTRIBUTION: Literature records. Luzon: In-
fanta. Subic Bay, Zambales Prov.; Camp
Nichols, Rizal Prov.; Calauan, Laguna Prov.;
Quezon City, Manila Prov.; Wackwack;
Manila, Manila Prov. Samar: Osmena. Jina-
moc Island. Leyte: Tacloban. Abuyog. Min-
danao: Zamboanga and San Ramon, City
of Zamboanga Prov. (Laffoon, 1946: 240.)
Leyte: Tolosa (Bick, 1949: 4).

Unknown outside the Philippines.
DISCUSSION: This species is closely related
to indicus (Theobald), which is known only

476

PACIFIC SCIENCE, VoL VII, October, 1953

from India. Of the Philippine fauna, campylo-
stylus most nearly resembles nigrotarsis.

Aedes (Aedes) campylostylus Laffoon
1946. Aedes {Aedes) campylostylus Laffoon,
Jour. Wash. Acad. Sci. 36: 240 (males,
females, pupae, larvae). Type locality:
Philippines. Balsahan River, Palawan
(nr. Iwahig) (Fitzjarrell). Type: Male
(holotype), with associated larval and
pupal skins, in U.S.N.M.

DISTRIBUTION: Literature records. Known
only from the type series. Leyte: Tacloban.
Mindoro: San Jose. Palawan: Balsahan and
Irahuan River Valleys. Balabac: Cape Mel-
ville, Mindanao: San Ramon, City of Zam-
boanga Prov. (Laffoon, 1946: 242.)

Aedes (Aedes) panayensis Ludlow

1914. Aedes panayensis Ludlow, Psyche 21:
159 (49 males and females). Type lo-
cality: Philippines. Iloilo, Panay (Ew-
ing). Type: Male (lectotype) in U.S.
N.M.

1929. Aedes {Aedes) panaygensis Ludl. Ed-
wards, Notulae Ent. 9: 3. Lapsus.

Adult and systematics treated by Laffoon
(1946: 242) and adult by King and Hoog-
straal (1947: 119).

DISTRIBUTION: Literature records . Panay: Ilo-
ilo. Calicoan Island (nr. Guiuan, Samar).
Culion Island. Balabac. Mindanao: Zambo-
anga, City of Zamboanga Prov. (Laffoon,
1946: 243.)

Outside the Philippines, known from Mo-
rotai, Schouten Islands, and Ceram.

DISCUSSION: The male of this species is es-
pecially distinctive in having the fore and
mid-tarsal claws equal and toothed. On the
basis of male terminalia, the closest relatives
to panayensis appear to be the Sumatran species
dermajoensis Brug and prioekanensis Brug.

The larva is undescribed.

Aedes (Aedes) butleri Theobald
1901. Aedes Butleri Theobald, Mon. CuL 2:
230 (3 females). Type locality: Malaya.

Selangor (Butler). Type: 3 females (co-
types) in B.M.f The specimen which
does not have the terminalia mounted
is probably a Skusea species. The other
two specimens, both of which have the
terminalia mounted, are conspecific and
I are here selected as representing butleri.
1903. Skusea diurna Theobald, Entomologist
36: 259 (female). Type locality: Ma-
laya. Jugra, nr. Kuala Lumpur (Dur-
ham). Type: Female (holotype) in
B.M.f Terminalia mounted.

1903. Verralina Butleri. Theobald. Theobald,
Mon. Cul. 3: 295. Different combina-
tion.

1910. Stegomyia hatiensis Carter, Entomologist
43: 275 (5 females). Type locality: Co-
chin China. Ha-tien (Broquet). Type:
2 females id lectotypes) in B.M.f
Terminalia mounted.

1924. Aedes {Skusea) urnhrosus Brug, Bui. Ent.
Res. 14: 437 (4 males, 2 females). Type
locality: Borneo. Tanah Grogot, S. E,
Borneo (Brug). Type: 2 males, 1 fe-
male (cotypes) in B.M.f Terminalia
mounted for 1 male and 1 female.
1928. Aedes {Aedes) umbrosus, Brug. Edwards
and Given, Bui. Ent. Res. 18: 345.
Description of larva.

1946. Aedes {Aedes) urnbrosus Brug. Laffoon,
Jour. Wash. Acad. Sci. 36: 243. Ten-
tative synonymy to butleri.

Barraud (1934: 296), female only, male is
probably lugubris Barraud. Laffoon (1946:
243), adult, larva, and systematics.

DISTRIBUTION: Literature records. Leyte. Ta-
cloban and Palo. Culion Island. Palawan:
Iwahig and Puerto Princesa. (Laffoon, 1946:
244.)

Outside the Philippines known from Bor-
neo, Java, Malaya, Cochin China, India, and
the Andamans.

DISCUSSION: The types of butleri have been
examined, and a female terminalic slide of a
Palawan specimen was compared with them.
There appears to be no essential differences.

Aedes of the Philippines — Knight and Hull

All

As pointed out in the synonymic table, the
female without terminalia separated is an un-
determined species. It possesses broad scales
on the scutellum and on ppn and lacks fine
hairs on the mesepimeron. It is probably a
Skusea species, but this point was not checked.

Two female terminalia labeled as this spe-
cies by Barraud in the British Museum collec-
tion from the Andamans (ix. 1911. Chris-
tophers) were examined and seemed to check
satisfactorily with the types.

The types of dturna and hatiensis were ex-
amined and found to be hutleri , as previously
indicated by Edwards and Barraud. Laffoon
(1946: 245) had questioned the synonymy of
hatiensis because Carter in his description said
that the scutellar scales were similar to those
covering the greater portion of the head, thus
implying that they were broad. However, in
examining the two Ha-tien specimens pinned
under hutleri in the British Museum, it was
found that both are pinned through the scu-
tellum, making it now impossible to know
what type of scales were present. It seems
likely that the missing three specimens (two
from Ha-tien and one from Saigon) had broad
scutellar scales and were moved by a later
worker (probably Edwards) to an Aedes spe-
cies with broad scutellar scales.

The types of umhrosus were studied, and the
female terminalia was compared with that of
a specimen from Palawan. No essential dif-
ferences were found. No Philippine male
specimen was available for direct comparison,
but the type male terminalia was compared
to the figure given by Laffoon (1946, fig. 1)
and found to agree perfectly, except that the
type specimen (only one had the terminalia
mounted) had three proximated subapical
spines instead of only two as figured by Laf-
foon, had only 13-15 verticillate bristles on
the antennal segments, and was somewhat
smaller. No differences were found between
the female type of umhrosus and the types of
hutleri, diurna, and hatiensis. The Javan spec-
imen mentioned in the original description of
umhrosus was not seen in the British Museum.

As pointed out by Laffoon (1946: 245),
the male terminalia described and figured by
Edwards (in Barraud, 1934, fig. 73e, f) as the
male of hutleri is probably the male of lugu-
hris. No Indian specimens of hutleri were seen
in the British Museum.

Subgenus Cancraedes Edwards

1929. Cancraedes Edwards, Bui. Ent. Res. 20:

342. Genotype: cancricomes Edwards

(Andaman Islands).

ADULT: Brownish species without special
ornamentation. Palpi very short in both sexes.
Vertex and scutellum all broad-scaled. Stout
dorsocentral bristles present, but no acrosti-
chals. Paratergite not scaled. Lower mese-
pimeral bristles present. Tarsal claws simple
in both sexes. Terminalia: Basistyle short, with
a complex group of apical and subapical ap-
pendages. Dististyle very short, simple, with-
out articulated appendage. Claspettes absent.
Mesosome divided, each lobe with an elon-
gate spine externally.

LARVA: The larva of only one species has
been described. As in subgenus Aedes except:
Comb with about 70 scales arranged in a
patch. Ventral brush with about 8 tufts, all
borne on a laterally connected barred area.
Habitat, crab holes and pools in mangrove
swamps.

DISTRIBUTION: The four known species (a
fifth species, kanarensis Edwards, 1934, was
questioningly placed in this subgenus by the
describer) are confined to the Oriental region.

SYSTEMATICS: Except for kanarensis, this
subgenus contains a very uniform group of
species. A. kanarensis, which has not been
included in the above subgeneric description,
differs markedly from the other known spe-
cies in the subgenus because of its scutal and
tarsal pale scaling.

Aedes (Cancraedes) miachaetessus
Dyar and Shannon

Fig. 12

1925. Aedes {Skusea) miachaetessa Dyar and

478

PACIFIC SCIENCE, Vol. VII, October, 1953

Fig. 12. A. (Cancraedes) miachaetessus. Male termina-
lia (Samar), a, Lateral aspect of basistyle, lateral surface
of basistyle omitted; b, left half of mesosome.

Shannon, Insecutor Inscidae Men-
struus 13: 78 (2 females). Type locality:
Philippines. Camp Stotsenberg, Pam-
panga Prov. Luzon (Telft). Type: 2
females (cotypes) in U.S.N.M.f One
of these specimens is here designated
lectotype.

1929. A. Cancraedes) miachaetessa, Dyar and
Shannon. Edwards, Bui. Ent. Res. 20;
342. Different combination.

ADULT: A very small dark species, without
special markings.

Male. Wing length about 2.1 mm. Head:
Proboscis dark. Palpus very short, no longer
than the clypeus, dark. Torus bare. Vertex
with broad dark scales, a few dark upright-
forked scales on nape. Thorax: Scutal inte-
gument dark brown; with narrow dark scal-
ing. Scutellar lobes with broad dark scales.
Apn and ppn with a few narrow-curved dark
scales. Pleuron yellowish brown in color, bare
of scales except for a few translucent dusky
broad scales dorsally on sternopleuron; a few
short pale hairs dorsoanteriorly and ventro-
posteriorly on sternopleuron; 1-2 stout lower
mesepimeral bristles. Legs: All dark. Tarsal
claws simple, those of fore and midlegs un-
equal. Wings: Dark-scaled. Halter knob dark-
scaled. Abdomen: Tergites and sternites dark-
scaled. Segments V-VIII noticeably enlarged,
VII-VIII much retracted, sternite VIII with

apical half densely covered with very long
hairs. Terminalia: As in figures. The one fig-
ure is a lateral view of the left basistyle. The
basistyle in this species is extremely bulbous,
and in order to show the dististyle and its
subapical appendages adequately the outer
surface of the basistyle is omitted from the
drawing.

Female. Similar to the male except that the
tergites have white basolateral patches, and
the sternites are white basally. Tarsal claws
equal, simple.

LARVA: Unknown.

BIONOMICS; Two males were collected from
the mouths of crab holes.

DISTRIBUTION: Specmens examined. R.K.L.
Samar: 1 male, Osmena (Laffoon, Knight).
Mindanao; 1 male, Zamboanga (Knight,
Laffoon). U.S.N.M. Luzon: 2 female types.
Camp Stotsenberg, Pampanga Prov.

Not known from outside the Philippines.
The two females from Koh Kut Island, Gulf
of Siam, listed by Dyar and Shannon (1925:
78) have been examined (in U.S.N.M.) and
are believed to be curtipes on the basis of the
dark pleurae.

DISCUSSION: The male of this species had
not previously been described. Of course, it
is quite possible that the association of sexes
made here is incorrect as the correlation is
made solely on general appearance.

Aedes (Cancra^des) curtipes Edwards

Fig. 13

1915. Aedes ?curtipes Edwards, Bui. Ent. Res.

5: 283 (4 females). Type locality: Bor-
neo. Kuching (.^), Sarawak (Moulton), j
Type: Female (holotype) in B.M.f

1928. Aedes {Sk.usea) curtipes, Edw. Edwards,
Bui. Ent. Res. 18: 274. Different com-
bination.

1929. A. {Cancraedes) curtipes, Edw. Edwards,
Bui. Ent. Res. 20: 342. Different com-
bination.

Male described by Edwards (1928: 274).
Larva described by Edwards and Given (1928:
346).

Aedes of the Philippines — Knight and Hull

479

ADULT: Male. Wing length about 2.0 mm.
Very similar to miachaetessus but differing
slightly as follows: Pleural integument gray-
ish brown to brown in color. Ppn with several
fine short hairs ventroposteriorly. Mesepim-
eron with several fine pale hairs between the
dorsal hair tuft and the lower mesepimeral
bristles. Hind femur with basal half partially
pale-scaled anteriorly. Terminalia: Resembling
that of miachaetessus but the subapical append-
ages differing as figured. The basal portion
of the basistyle of the single available spec-
imen was somewhat broken so that details in
that portion may not be accurately illustrated.

Female. As in male except that there are
laterobasal pale patches on most of the terg-
ites and basal paler scales on the sternites.
Torus with some dark scales and hairs me-
dially.

LARVA (from Edwards and Given, 1928:
346. No larvae taken in P.I.): Head: Antenna
almost as long as head, with numerous spic-
ules; antennal hair tuft with 5-6 branches,
inserted just before middle and reaching the
tip of the antenna. Preclypeal spines long,
bristle-like. Head hair 4 with about 6 branches,
very small; 5 with 3; 6 with 2; 7 with 7-10;
8 single; 5, 6, and 7 all large and plumose,
and arranged in a slightly curved line. Men-
tum with about 15 lateral teeth. Thorax: Lat-
eral tufts large and plumose; associated spines
very short. Abdomen: Lateral tufts double,
plumose on I and II. Comb with about 70
scales, each long, narrow, sharp, and simple.
Siphon rather strongly tapering on the outer
half, index about 2.7; acus distinct, narrow;
6-8 pecten teeth, mostly rather widely spaced
and extending a little beyond middle of si-
phon, each tooth with 2-3 very strong denti-
cles; siphon hair tuft with 3-5 branches,
inserted beyond distal pecten tooth. Anal
plate small; ventral brush of about 8 tufts,
each tuft with a rather long stalk, no tufts
off the barred area. Anal gills short, somewhat
pear-shaped, dorsal pair almost as long as
anal plate, ventral pair shorter.

BIONOMICS: Adults were collected flying at

Fig. 13. A. {Cancraedes) curtipes. Male terminalia
(Palawan), a, Mesal aspect of basistyle; b, left half of
mesosome.

the edge of a mangrove swamp and from
resting places inside the openings of crab
holes. The larvae were not taken. However,
Edwards and Given (1928: 346) reported the
larvae from mangrove pot-holes.

DISTRIBUTION: Specimens examined. R.K.L.
Palawan: 1 male, 5 females, Iwahig Penal
Colony (Laffoon). U.S.N.M. Palawan: 2 fe-
males, Puerto Princesa (19th M.G.L.).

Not previously reported from the Philip-
pines.

Outside the Philippines this species is
known from Borneo and Malaya. As pointed
out in the treatment of miachaetessus, it is be-
lieved that the two female specimens in the
U.S.N.M. from Koh Kut Island, Gulf of
Siam, are this species.

DISCUSSION: The type was studied, but un-
fortunately a male terminalia was not seen.
However, the male terminalia of the Palawan
specimen agrees with Edwards’ (1928, fig. 1)
figure of the male terminalia of curtipes, except
that he does not show the subapical append-

480

PACIFIC SCIENCE, VoL VII, October, 1953

age A that is present in the Palawan specimen

(see figure) .

AEDES OF UNKNOWN SUBGENERA
Aedes (?) platylepidus Knight and Hull

1951. Aedes (?) platylepidus Knight and Hull,
Pacific Sci. 5: 201 (3 females). Type
locality: Philippines. Puerto Princesa,
Palawan (Laffoon and Johnson). Type:
Female (holotype )in U.S.N.M.f

DISTRIBUTION: Specimens examined. Known
only from type series. Palawan: Puerto Prin-
cesa. Balabac: Cape Melville.

DISCUSSION: The male is unknown and the
larva is not described. The type series was
obtained from a larval collection but unfor-
tunately no associated larval material was re-
tained.

This species is strikingly different, on the
basis of the broad plate-like dark (with metal-
lic reflections) scutal and scutellar scales, from
all other known Pacific Aedes species. Lack of
the male and larva, together with the rather
aberrant characters of the female, prevent the
making of a definite decision at this time as
to the subgeneric position in Aedes of platy-
lepidus, The possible relationships of this spe-
cies are discussed in some detail in the original
description.

Aedes, species unknown

Three larval specimens from Palawan and
Balabac represent an unknown aedine species.
It is quite possible that this is either the larva
of Aedes platylepidus or of Heizmannia scintillans
Ludlow, but in the absence of associated mat-
terial it is impossible to say definitely at this
time.

LARVA: Head: Antenna smooth, rather
stocky; antennal hair single, inserted near
apex. Mouth brush with comb-like tips. Hair

4 single, rather long, dark, curved inward;

5 single; 6 with 2-3 branches; 7 single or
double, small; 8 single; 9 double; 12 with
2-3; 13 and 14 single; 15 with 1-3; 17 and 18
single; 20 double; 18 is inserted just mesad
and ventral to 17, rather than near the lower

corner of the mentum. Mentum with 8-9 lat-
eral teeth. Thorax: Pro thoracic hairs 1 and 2
single; 3 with 3 branches. Mesothoracic hair
9 single; thoracic hairs in general quite sparse-
ly branched. Abdomen: Dorsolateral hair on I
with 2-3 branches; on II double. Lateral hair
on I double; on II single or double; on III
and IV with 3-4; on V and VI with 5-6.
Pentad hair 1 with 2-3; 2 and 4 single; 3 with
2-6; 5 with 2-3. Comb with 6 teeth in a
curved row, each tooth long, slender, sharply
pointed, with a small amount of fringe lat*
erally at the base of the spine. Siphon smooth,
with an attached acus, index 2. 1-2. 4; 7-12
pecten teeth in an irregular row, each with
fine fringe basally on ventral side and occa-
sionally on dorsal side also; siphon hair tuft
with 4-6 branches, inserted either before, near,
or beyond the last pecten tooth. Anal plate
apparently complete, with a few small spicules
laterally on the posterior margin; Ih with 3-4
short, rather stout branches; isc single (once
double), osc single; ventral brush of 10 hairs,
each single except for basal pair with 3-5
branches; no barred area present. Anal gills
finger-like, dorsal pair slightly shorter than
ventral pair and 2. 2-2. 5 times longer than
anal plate.

BIONOMICS: This larva was taken on one
occasion from water collected in a large metal
packing container in the jungle and on an-
other from water collected in a large, thorny
palm frond lying on the ground at the edge
of a mangrove area.

DISTRIBUTION: Specimens examined, R.K.L.
Palawan: 2 larvae (#854) Irahuan River, 3
miles inland (Laffoon). Balabac: 1 larva
(#1611), Cape Melville (Johnson).

One of the Palawan larvae is deposited in
the U. S. National Museum as Aedes sp.
#46. The other two are retained in the Knight
collection.

REFERENCES

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Aedes of the Philippines — Knight AND Hull

481

BicKj G. H. 1949. Notes on mosquitoes from
Leyte^ Philippine Islands (Diptera, Culici-
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Bohart, R, M. 1945. A synopsis of the 'Philip-
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Brug, S. L. 1932. Notes on Dutch East Indian
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Carpenter, S. J., W. W. Middlekauff, and
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Causey, O. R. 1937. Some anopheline and
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Dyar, H. G. 1925. Note on the male of
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and R. C. Shannon. 1925. The types

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Edwards, F. W. 1917. Notes on Culicidae
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1922. A synopsis of adult Oriental

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1928. Mosquito notes. VII. Bui. Ent.

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1928^?:. Diptera Nematocera from the

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1929. Philippine nematocerous Dip-
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1932. Diptera: Family Culicidae. In

Genera Insectorum [Wytsman ed.], Fasc. 194.

258 pp. Brussels.

1934. Appendix, and numerous foot-
notes in Barraud 1934.

1941. Mosquitoes of the Ethiopian region.

III. Culicine adults and pupae. 499 pp.
British Museum, London.

and D. H. C. Given. 1928. The early

stages of some Singapore mosquitoes. Bui.
Ent. Res. 18: 337-357.

Hsiao, T.-Yu, and R. M. Bohart. 1946. The
mosquitoes of Japan and their medical im-
portance. NavMed 1095. 44 pp. Bureau
of Medicine and Surgery, Navy Depart-
ment, Washington, D. C.

King, W. V., and H. Hoogstraal. 1947.
New Guinea species of mosquitoes of the
genus Aedes, sub genus Aedes. Jour. Wash.
Acad. Sci. 37: 113-134.

Knight, K. L., and W. B. Hull. 1951. The
Aedes mosquitoes of the Philippine Islands.
1. Keys to species. Subgenera Mucidus,
Ochlerotatus , and Pinlaya (Diptera, Culici-
dae). Pacific Sci. 5: 211-251.

1952. The Aedes mosqui-
toes of the Philippine Islands. IL Subgenera
Skusea, Christophersiomyia, Geoskusea, Rhin-
oskusea, and Stegomyia (Diptera, Culicidae).
Pacific Sci. 6 (2): 157-189.

and H. S. Hurlbut. 1949. The mos-
quitoes of Ponape Island, eastern Carolines.
Jour. Wash. Acad. Sci. 39: 20-34.

LaCasse, W. J., and S. Yamaguti. 1948. Mos-
quito fauna of Japan and Korea. IL 273 pp.
Office of the Surgeon General, Washington,
D. C

Laffoon, j. L. 1946. The Philippine mos-
quitoes of the genus Aedes, subgenus Aedes.
Jour. Wash. Acad. Sci. 36: 228-245.

Leicester, G. F. 1908. The Culicidae of Ma-
laya. Fed. Malay States, Inst. Med. Res., Stud-
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Penn, G. H. 1948. Biological notes on ”dry
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Mindoro, P. L Ent. Soc. Wash., Proc. 50:
241-248.

1949. The larva and pupa of

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Culicidae). Nat. Hist. Miscellanea 40: 1-4.

The Response of a Tropical Fish to Direct Current and Its
Application to the Problems of Electrofishing in Sea Water ^

Morris E. Morgan^

Methods of electrofishing have been de-
veloped successfully for use in fresh-water
ponds and streams, but this equipment is not
effective in sea water because the high con-
ductivity of salt water greatly changes the
power requirements (Smolian, 1944). If this
obstacle can be overcome, it is reasonable to
assume that electrofishing can be conducted
in the sea with a success which would be at
least equivalent to that experienced in fresh
water.

Electrofishing in fresh water is now widely
practiced in Germany, the United States, and
Canada, where it is used mainly as a new tool
to supplement the facilities for study of fish
populations. The progress made in Germany
has been described by Smolian (1944) in a
comprehensive report which includes a de-
scription of the equipment and techniques
used in that country. In the United States,
the first efforts concerned with the reaction
of fish to electrical stimulation centered
around the development of electric fish screens
and began as early as 1917 (Holmes, 1948).
However, a method of actually collecting fish
in fresh-water streams was first developed in
1939 (Haskell, 1940).

Equipment used for electrofishing in fresh
water has consisted mainly of small, portable,
gas-driven generators which produced alter-
nating current. However, Smolian (1944) re-
ported that the greatest effect seemed to be
realized from using direct current interrupted
at frequent intervals. Large fish were affected

^ Contribution No. 34, Hawaii Marine Laboratory.
Manuscript received July 2, 1952.

2 Research Biologist, California Academy of Sciences.

more strongly than small ones. There was
also considerable variation in the sensitivity
of various species to electrical stimulation.

In reports from both the United States and
other countries it has been emphasized that
paralyzing, narcotizing, and attracting the fish
can be accomplished only in the immediate
vicinity of the electrodes. Outside this area
the current merely irritates the fish and fright-
ens them away.

While some reports have been made con-
cerning the injurious or fatal effects of elec-
trofishing (Hauck, 1949), it appears that fish
can be collected by using current strengths
below those which will injure or kill.

German scientists have been investigating
the possibility of adapting electrofishing tech-
niques for use in salt-water fisheries. At the
present time it has been reported (Anon.,
1951) that they are in the process of installing
electrical fishing gear on a boat of the type
used in the trawl fishery. Only general news-
paper-type reports have been published con-
cerning this work. The basic research was
financed personally by the scientists con-
cerned so it is assumed that no detailed in-
formation will be forthcoming until their
work has reached a patentable stage.

These reports indicate that their claims of
success in the development of electrofishing
equipment suitable for use in sea water are
based on the more efficient utilization of elec-
tricity rather than in merely increasing the
amount. Special mention is made of pulse
shape, pulse rate, and electrode arrangement.

The response of living organisms to elec-
trical stimulation was investigated as early as

482

Response of Fish to Direct Current — MORGAN

483

1885 by Hermann. He reported that tadpoles,
when in a field of direct current of proper
strength, would react by orienting themselves
with their heads toward the positive pole.
Since that time, experiments have been con-
ducted with numerous animals, including
crabs, fish, and frogs. Practically all types of
animals tested show an orientation to the
current. Many protozoans, however, orient
and migrate toward the negative pole rather
than the positive (Heilbrunn, 1948: 597).

Considerable research has been done in an
attempt to explain the physiological reasons
for this reaction (Breuer, 1905; Scheminsky,
1924; Van Harreveld, 1937). However, these
reports indicate that it is still imperfectly un-
derstood. As nearly as can be determined, no
satisfactory explanation of this phenomenon
has been given.

I wish to express my appreciation for the
advice and assistance given by Dr. A. L.
Tester in the preparation of this manuscript
and over-all supervision of the investigation.

METHODS

The period covered by the investigation
here reported extended from May, 1950, to
March, 1951. Work was begun at the Hawaii
Marine Laboratory on Coconut Island and
continued there throughout the summer
months. At the beginning of the academic
year, all the equipment was transferred to the
Waikiki branch of the Hawaii Marine Labor-
atory. This branch, operated in conjunction
with the Honolulu Aquarium, offered certain
advantages over the previous site, principally
its proximity to the University of Hawaii. All
the detailed experimental work reported in
the following pages took place at the latter
site.

Power Supply

The generating equipment was selected to
give the widest possible range in current and
voltage while still being portable and of
standard design.

As direct current was more likely to pro-
duce the reaction desired, an Onan two-

cylinder, gas-driven, air-cooled, portable-type,
direct-current, motor- generator set was pur-
chased. It was rated at 5,000 watts (21.8 am-
peres at 230 volts). Special equipment in-
cluded a voltmeter, ammeter, rheostat, and
circuit breaker.

Electrodes

The high conductivity of the sea water
caused serious overloading of the generator.
Plans for overcoming this difficulty centered
around the use of rheostats and suitable elec-
trodes. However, the rheostat installed to con-
trol the voltage and current was ineffective
and various other resistors devised to control
the current flow proved unsatisfactory. A spe-
cial electrode was finally developed for use
during the experiments. This consisted of a
carbon rod 6 inches long and !4 inch in dia-
meter inserted into a plastic tube (Fig. 1).
The size and number of holes in this tube
governed the flow of current. In effect the
column of sea water inside the tube became
the electrode, as the area of this column in

Copper clamp, adju.s'table
Su-pporlri ng wire

#14 Insulated wire t-o

Wat-erproof ‘i nau.lai^ing compound

Wirs and solder

Carbon electrode , 6" long -2^ diam.

■plastic tube, 12" long — l" diam.

Fig. 1. Electrode design.

Fig. 2. Experimental tank and current-interrupting equipment.

contact with the main body of water in the
tank determined how much current would
flow. As the entire electrode was covered by
water, gases formed on its surface evenly, and
their escape did not affect the current flow.
With this arrangement, it was possible to low-
er the enclosed electrode to any depth without
changing the current flow, and a more sym-
metrical field could be created at each end of
the tank.

Carbon was chosen as the most suitable
electrode material because it is a fair con-
ductor, inexpensive, and does not erode rapid-
ly with electrolysis as do most metals.

To remove the products of electrolysis as
they were formed and to prevent contamina-
tion of the water (chiefly by chlorine) in the

tank, the water within each electrode was
siphoned off continuously be means of a rub-
ber tube. The loss of water from the tank was
compensated for by a steady addition of new
water.

Tank

A tank measuring 2 X 2 X 12 feet was
constructed from redwood. The inner surface
was thoroughly impregnated with heated par-
affin to prevent leaching and to aid in water-
proofing. Net barriers were placed about 1
inch in front of each electrode to prevent the
fish from injuring themselves by contact with
the electrode. Two other net barriers were
arranged across the center of the tank to form
a small area in which the fish could be held

485

Response of Fish to Direct Current ^ — Morgan
when not actually under observation (Fig. 2).
Current Interrupter

After overloading had been controlled, pre-
liminary experiments showed that the full
load of the generator, approximately 20 am-
peres and 230 volts, would not attract and
hold fish about 4 inches long in a concrete
tank 10 feet square filled with water to a depth
of 2 feet. When the fish swam near the elec-
trodes, they were effectively held, but they
would remain at the greatest distance possible
from the electrodes or in the corners of the
tank near the bottom.

Thus it became obvious that, if a steadily
flowing current was to be used, a much greater
source of power would be needed. A similar
observation was made by Peglow (1949). To
attract the fish to the positive pole a much
more efficient use of the available power would
be necessary. The best possibility of accom-
plishing this, according to the general reports
of previous investigators, would be through
the use of interrupted current (Smolian, 1944;
Anon., 1950).

Various mechanical devices were con-
structed, mainly from war surplus material,
for the interruption of direct current. First
attempts centered around the use of relays,
but for various reasons these did not prove
practicable. It was difficult to determine the
exact duration of the power-on period com-
pared with the power-off period. When opera-
ting at higher interruption frequencies and
with the higher power values, the arc became
very difficult to control, even with condensers.
Observation of the wave form on an oscillo-
scope showed that, if the power-off period
was short, the arc often lasted over the entire
period. The relay points frequently became
heated and failed to separate. It was also dif-
ficult to determine the exact number of in-
terruptions per second.

A revolving disc-type interrupter was fi-
nally decided upon. Although not satisfactory

■Revolving

Detail of i nterrupfri ng disks ,

Ra kel ita -N ^T3ronz.a

Fig. 3. Diagram of current-interrupting equipment
used in experiments.

in every respect, it at least overcame most of
the objectionable features of previous equip-
ment (Fig. 3).

Carbon brushes bearing on either side of
the disc provided a constant connection when
the bronze portion of the disc was between
them. When the disc revolved, the connec-
tion was definitely broken when the Bakelite
portion was between the brushes. Some arc
was unavoidable, but an attempt was made
to minimize this by three condensers (10
mfd,, 600 volts, d.c.) arranged in parallel.
A variable speed motor controlled by a ”ve-
riac” was used to turn the disc.

A small generator was connected directly
to the shaft of the motor. The voltage pro-
duced by the generator was recorded on a
voltmeter. The relationship between voltage
and speed was determined so that, by reading
the voltage on the voltmeter, the speed of the
generator and, therefore, of the motor could
be calculated.

486

• PACIFIC SCIENCE, Vol. VII, October, 1953

Fig. 4. The experimental fish, Kuhlia sandvicensis (Steindachner), attracted to the positive pole and immo-
bilized there by the use of interrupted direct current.

Fish

The fish chosen for the experiments was
Kuhlia sandvicensis (Steindachner) (Fig. 4), lo-
cally called aholehole. This fish is bright sil-
very when viewed from the side but appears
dark blue when seen from above. It spawns
along the shoreline and is quite plentiful in
the brackish water near the mouths of streams
and canals where it grows to a size of approx-
imately 25 cm.

, Aholehole were recommended for the ex-
periments because of their hardiness and abil-
ity to survive considerable handling. This

turned out to be a fortunate choice, for enough
of the fish were caught in traps in Ala Wai
Canal to supply all experimental needs. The
fish were held and fed in tanks of the Hono-
lulu Aquarium when not actually being used
for experimentation. They proved to be ad-
mirably suited for the purpose intended. After
a rather high initial mortality due to injury
by trapping, practically none died, either from
handling or from the effects of electrical
shocks.

Experimenters have been unanimous in
their reports that larger fish are affected more

Response of Fish to Direct Current — MORGAN

strongly than smaller fish, so an attempt was
made to control this factor by selecting fish
of approximately the same size for the ex-
periments. The range in size was from 9-5
cm. to 15.0 cm., the mean size was 11.8 cm.,
and the standard deviation was 1.04 cm.

Procedure

The collection of quantitative data on the
reaction of fish to electrical stimuli was con-
sidered highly desirable, and an experimental
plan was formulated with this in mind. The
main object of the experimental work was to
cause the fish to move, involuntarily, to the
positive pole, where it would be held, either
by a strong attractive force or by passing into
a condition of electronarcosis. To eliminate,
as far as possible, subjective observations and
to measure this effect under different elec-
trical conditions, the experimental design dis-
cussed in the following paragraphs was used.

The tank was considered to consist of two
equal sections, positive and negative. The fish
were held in the center of the tank between
two net barriers. When the power was turned
on, the barriers were removed. The power
remained on for 60 seconds. During this pe-
riod, the time the fish spent in the positive
section was recorded. Under optimum elec-
trical influence, the fish, when released from
the center area, would move directly to the
positive pole and become immobilized, thus
spending the full period of 60 seconds in the
positive section. When no influence was in
effect, the fish (over an average) would be
expected to spend 30 seconds in the negative
section and 30 seconds in the positive section.

A two-pole, double-throw switch was placed
in the circuit so the polarity could be reversed.
This obviated the possibility that some in-
fluence other than electrical stimulation might
cause the fish to seek one end of the tank
more consistently. Each fish was given two
timed trials, the polarity being reversed in the
second trial. To determine whether the move-
ments of the fish in the tank were random,
preliminary trials were made with the power

487

off. Ten tests were made using individual fish.
They spent an average of 31.8 seconds in the
positive section and 28.2 seconds in the nega-
tive section. These average times did not de-
viate significantly from the expected times of
30 seconds in each end of the tank, so it was
concluded that movement throughout the
tank was random when the current was off.
Timed trials were also made with several fish
in the tank at the same time. However, in-
dividual variation made accurate timing of
each fish difficult, so it was decided that fish
would be used individually rather than in
groups.

During the experiments, the voltage re-
mained relatively constant (220-230 volts).
The depth of water in the tank was 12 inches,
and the distance between electrodes was 11
feet. These factors were constant for all tests.

One group of experiments was conducted
with continuously flowing (uninterrupted)
current. In it were included a series of experi-
ments at amperages of 1 to 14, inclusive. The
current in amperes may be converted to den-
sity in milliamperes per square inch by mul-
tiplying the amperage by the factor 3.62.
Thus, 14 amperes is equivalent to a current
density of 51 milliamperes per square inch.

Three other groups of experiments were
conducted with interrupted current using dif-
ferent periods of impulse duration (pulse
time). The variation in impulse duration was
obtained by using rotating discs with different
bronze-Bakelite combinations as already de-
scribed. In the first group, the disc used was
half bronze and half Bakelite. Thus, during
each revolution the power was on for one half
the time and off for one half the time (1:1
on-off ratio). In the second group of experi-
ments, the disc was three-fourths bronze and
one-fourth Bakelite. During each revolution
the power would then be on three times as
long as it would be off (3:1 on-off ratio). In
the third group of experiments one fourth of
the interrupting disc was bronze while three
fourths was Bakelite, In this case the power
was on one fourth of the time and off three

488

PACIFIC SCIENCE, Vol. VII, October, 1953

fourths of the time during each revolution
(1:3 on-off ratio).

Each group of experiments with interrupted
current included several series at successively
greater amperages. In group I, amperages of
1, 2, 3, 4, and 5 were used. In group II, am-
perages of 4, 5, 6, 7, and 8 were used. In
group III, amperages of 2, 3, and 4 were used.
At each amperage within a group, experiments
were conducted at several frequencies of in-
terruption. Frequencies used in group I were
5, 10, 15, 20, and 25 interruptions per second.
In groups II and III, frequencies of 5, 15, and
25 interruptions per second were used. In
each individual experiment, five fish were used
and two trials were made with each. The po-
larity was changed with each trial. In all
groups of experiments, successive tests were
conducted until a current value was reached
that would cause all fish to move to the posi-
tive pole and remain there for the full 60
seconds of each time period. If the frequency
of current interruption proved to be a factor
in causing the desired effect, this would be
shown by variation in the times recorded for
the different interruptions when both current
and voltage remained constant.

RESULTS

Continuous Direct Current

The results of tests using uninterrupted
(continuously flowing) current are shown in
Table 1. The fish showed no effects of the
current flow at 1, 2, and 3 amperes. At a
current value of 4 amperes, the fish swimming
near the electrodes, within about 6 inches,
would suddenly veer away as though shocked.
This effect became noticeable at greater dis-
tances from the electrode as the current was
increased. At 7 amperes some fish were more
strongly shocked in the immediate vicinity
of the poles, but reactions were entirely eva-
sive. The fish would swim toward one end of
the tank; upon nearing the pole and receiving
a shock they would turn and swim rapidly
toward the opposite end, where the experience
would be repeated.

At a current value of 9 amperes the fish
entering the area near the positive pole would
show the typical electronarcotic effect, in-
cluding cramped, slow movements and slow-
ing of opercular action, which resulted in
respiratory difficulties. If the fish were at right
angles to the lines of force between the elec-
trodes, the body would be bent with the head
and tail pointing toward the positive elec-
trode. However, the electrical field was still
not powerful enough to hold the fish and they
were able to escape this influence, avoid the
vicinity of the pole, and swim to an area in
the tank where they were affected least.

At current values above 10 amperes practi-
cally all fish started swimming toward the
positive pole when the center barriers were
removed but were repulsed rather than held
by the increasing effects as they neared the
positive pole. The result was that, they re-
mained either in a low-current density area
at the bottom of the tank near the pole bar-
riers, or near the center of the tank away from
both poles. Their swimming movements were
slow and unnatural. The dorsal fin was stiffly
elevated.

At a current value of 14 amperes all fish
swam toward the positive pole until the bar-
rier was reached. Many tried to force their
way through the protective net in their at-
tempts to get nearer the electrode. Soon after
reaching the net barrier the electronarcotic
effect became evident. The fish could not

TABLE 1

Mean Time (in Seconds for 5 Fish, 2 Trials Each)
Spent in the Positive Section of the Tank
Using Uninterrupted Direct Current

AMPERES

MEAN

TIME

AMPERES

MEAN

TIME

1

33.9

8

36.9

2

27.9

9

19.7

3

34.9

10

38.1

4

26.9

11

30.2

5

30.9

12

41.5

6

38.2

13

46.2

7

32.5

14

60.0

Response of Fish to Direct Current — Morgan

489

TABLE 2

Mean Time (in Seconds for 5 Fish, 2 Trials Each)
Spent in the Positive Section of the Tank at
Various Frequencies and Amperages of Inter-
rupted Current with 1:1 ON-dFF Ratio

amperes

FREQUENCIES

5

10

15

20

25

1

35.0 .

45.8

34.6

24.3

38.2

2

30.4

31.4

33.1

42.2

28.6

3

25.0

41.5

32.5

36.8

41.0

4

28.2

33.2

46.2

48.2

43.1

5

50.3

60.0

60.0

60.0

60.0

maintain equilibrium and would sink slowly
toward the bottom of the tank, ventral side
up. In some, all visible signs of life, including
opercular movements, ceased. However, when
the power was turned off, all the fish recov-
ered, usually within a few seconds. None died
as a result of this series of tests using a con-
stantly flowing current. Of the five fish which
were tested at various amperages, one showed
in both trials the complete electrotactic and
electronarcotic effect at 12 amperes, one at
13 amperes, and all five at 14 amperes. There-
fore, 14 amperes was considered the value
necessary to cause the fish to come to the
positive pole and become immobilized when
uninterrupted current at 225 volts was used.

A statistical analysis of the quantitative
data of Table 1 was made to supplement the
observations reported above. It may be seen
that the mean time spent in the positive half
of the tank fluctuated between 19.7 and 38.2
seconds in the experiments conducted over a
range of amperages from 1 to 11. As indicated
by a *'t” test, none of these means differed
significantly from an expected mean of 30
seconds. At 12 amperes the mean of 41.5
seconds differed significantly from the ex-
pected mean of 30 seconds (t = 2.06; P =
0.07) . At 13 amperes the mean of 46.2 seconds
was also significant (t = 4.68; P < 0.01).
Thus, with the ahoiehole, a positive electro-
tactic effect at a distance of 5.5 feet (the center
of the tank) from the electrodes was demon-

strated quantitatively at 12 amperes. However,
as already pointed out, complete electrotaxis
and electronarcosis did not result until a cur-
rent of 14 amperes was used.

Interrupted Direct Current
1:1 On-Off Ratio

The tabulated results of experiments with
various values of interrupted direct current
are shown in Tables 2, 3, and 4. In the first
group of experiments the power-off period
was equal to the power-on period at all fre-
quencies. Examination of the tables again dis-
closes marked variation in the reaction of
individual fish. The desired effect, which im-
mobilized all fish at the positive pole for the
full 60 seconds of each test period, was reached
at an average current value of 5 amperes
(Table 2). This effect occurred at each fre-
quency with the exception of five interrup-
tions per second. An effect comparable to
this was not obtained with uninterrupted pow-
er until a current of 14 amperes had been
reached. This seems to indicate that the in-
terruption in some way of direct current makes
it much more effective.

It is obvious that by interrupting the direct
current so that the power alternates on and
off for equal periods, more efficient use is
made of the available power. The question
is then raised as to whether the reaction of
the fish varies with the duration of the on-or
off-period. To obtain information on this

TABLE 3

Mean Time (in Seconds for 5 Fish, 2 Trials Each)
Spent in the Positive Section of the Tank at
Various Frequencies and Amperages of Inter-
rupted Current with 3:1 On-Off Ratio

AMPERES

FREQUENCIES

5

15

25

4

41.9

40.3

29.4

5

38.7

37.7

22.8

6

32.3

36.2

41.5

7

52.4

52.4

52.6

8

53.7

56.5

60.0

490

pointj the second and third groups of experi-
ments were conducted, the results of which
are given in the following sections.

Interrupted Direct Current
3:1 On-Off Ratio

The results of tests using a 3:1 on-off ratio
are shown in Table 3. From this table it can
be seen that the desired effect of holding the
fish at the positive pole for the full time of
each test was not achieved on all fish until
an average current value of 8 amperes was
reached, and then only in the tests having
25 interruptions per second. However, an
analysis of the results shows that a significant
deviation from the expected average of 30
seconds was reached in the tests using 7 am-
peres; for example, at five interruptions per
second t = 3.18, P == 0.01. The higher am-
perage required when using the 3:1 on-off
ratio resulted in a loss rather than a gain in
the efficient use of power. It was then decided
to decrease the duration of the on- (pulse)
period, using a 1:3 on-off ratio.

Interrupted Direct Current
1:3 On-Off Ratio

Recorded results of these tests are shown
in Table 4. A significant variation from the
expected average of 30 seconds was reached
at an amperage of 2. At a current value of 3
amperes and a frequency of 25 interruptions
per second, all fish tested were immobilized
at the positive pole. At a current value of 4
amperes all fish tested at all frequencies
showed the maximum effect possible.

The quantitative data in the preceding ta-
bles do not, of themselves, indicate the full
effect which results when interrupted current
and decreased on-period is used. Observation
of the reaction of the fish left no doubt that
the experiment using the shortest pulse pe-
riod produced the most violent effects. With
uninterrupted current the fish’s movements
were slowed and inhibited. With interrupted
current, as the individual shocks became
shorter, the effect it produced became greater.

PACIFIC SCIENCE, VoL VII, October, 1953

At the shortest pulse period, with an effective
current, the fish would often leap out of the
water and would frequently skip along the
surface, always moving directly toward the
positive pole.

Variation of Response
with Frequency of Interruption

A statistical analysis was made of the rela-
tionship between frequency of interruption
and time spent at the positive pole. From each
group using interrupted current the recorded
times at frequencies of 5, 15, and 25 were
chosen. These were taken only from the two
amperages just below the value that caused
maximum effect (60 seconds at the positive
pole). The only exception to this was the 8
ampere 3:1 on-off series; these data were add-
ed because the maximum effect was shown
only in the 25 interruptions per second tests
(Table 3). The following were included in the
analysis: 1:1 ratio, 3 and 4 amperes; 3:1 ratio,
6, 7, and 8 amperes; 1:3 ratio, 2 and 3 amperes.
The effect of different on-off ratios, as well
as the effect of the various amperages, was
separated by analysis of variance methods
from the effect due to frequencies of inter-
ruption. The data were transformed logarith-
mically to conform more closely to normal
distribution. The analysis provided the in-
formation given in Table 5.

There are significant differences among fre-
quencies (F = 6.73; P = 0.01). Inspection of
the data shows that the higher time averages
are associated with the higher frequencies.

TABLE 4

Mean Time (in Seconds for 5 Fish, 2 Trials Each)
Spent in the Positive Section of the Tank at
Various Frequencies and Amperages of Inter-
rupted Current with 1:3 On-Off Ratio

AMPERES

frequencies

5

15

25

2

41.3

■ 44.5

55.9

3

56.3

52.0

60.0

4

60.0

60.0

60.0

491

I Response of Fish to Direct Current — MORGAN

I

TABLE 5

Results of Statistical Analysis of Relationship between Frequency of Interruption

AND Time at Positive Pole

SOURCE

DEGREES OF FREEDOM

SUMS OF SQUARES

MEAN SQUARES

Total

20

0.243032

Blocks

6

0.175656

0.029276

Frequencies

2

0.035628

0.017814**

Discrepance

12

0.031748

0.002645

Observation of the effect at the various
interruptions leads to the conclusion that the
lower frequencies are not as effective as those
in the middle range. If the pulse duration is
short, this is particularly true. At low fre-
quencies there is sufficient time after the fish
receives the shock to enable it to turn par-
tially or make other movements to escape the
current. As a result, the fish pursues an erratic
rather than straight course toward the posi-
tive pole. In certain respects, the frequencies
near the value of 15 per second have some
advantages over higher frequencies used. At
higher frequencies the fish receives such a
rapid series of shocks that there seems to be
insufficient time for it to react between shocks,
thus greatly restricting its movements and
slowing its progress toward the positive pole.
At these higher frequencies, with an effective
current, the fish often becomes immobilized
at some distance from the pole.

Relationship between Average
and Peak Currents

One very important fact that enters into
the interpretation of the results of the experi-
ments is that all interrupted-current values
are average values. These values were adjusted
to the desired reading on the ammeter while
the interrupting mechanism was in operation.
The peak amperage was not indicated by the
ammeter during the tests because of the in-
dicator’s inertia. The peak amperage is the
maximum value that will be reached during
the current-on period. For example, when the
on-period equaled the off-period and the aver-

age current value, as read on the ammeter,
was 5 amperes, the peak current reached dur-
ing each pulse was 10 amperes. This value
was read on the ammeter when the interrupter
was not in operation.

From Figure 5 it can be seen that the peak
current in all cases is similar. It is quite possi-
ble that, were the exact current values neces-
sary to achieve the desired effect known, the
peak current in all cases might be equal. As
it is, the effective average current value in

Pea k

Unit- •time period

Unit" time period 1

Fig. 5. Average and peak current considerations.
Total power comparisons.

492

graph A (Fig. 5) is known only to be between
7 and 8. If the true average current values
were 7.5 amperes for graph A, 5.0 amperes
for graph B, and 2.5 amperes for graph C,
then the peak current in all cases would be
10.0 amperes.

Decreasing the on- (pulse) period results
in a more pronounced electrotactic effect at a
lower amperage and also represents a saving
in power. It is quite possible that further de-
creases in the on-time would result in equal
electrotactic effects and a still greater saving
in power.

The success of electrofishing in the open
sea will depend mainly on the ability to make
still more efficient use of available power.
This- is necessary because generators must be
able to produce sufficient power, yet be of a
size that will permit installation in a fishing
boat.

The use of large amounts of electrical pow-
er presents a constant element of danger to
personnel unless proper precautions are taken.
Safety regulations must be strictly observed
both in the laboratory and in the field.

CONCLUSIONS

1. Interrupted direct current is more desirable
for electrofishing purposes than uninter-
rupted current because of the increased
electrotactic and electronarcotic effects on
the fish.

2. The use of interrupted current in electro-
fishing represents a considerable saving in
power over the use of uninterrupted current.

3. The pulse length of interrupted current can
be decreased, at least to some extent, with-
out sacrificing any effectiveness. This re-
sults in further power savings.

PACIFIC SCIENCE, Vol. VII, October, 1953

4. The peak current value reached during a
pulse may be the important consideration
in causing the response desired in electro-
fishing.

5. With interrupted direct current, fish can
be attracted to the positive pole and im-
mobilized without being killed or injured.

REFERENCES

Anonymous. 1951. Note. Com. Pish. Rev. 13
(1): 53-54.

Breuer, Josef. 1905. Uber den Galvanotrop-
ismus (Galvanotaxis) bei Fischen. Akad.
der Wiss. Wien, Math.-Nat. Ki., Sitzber. Abt.
Ill, 114 (2): 27-56.

Haskell, David C, 1940. An electrical meth-
od of collecting fish. Amer. Fisheries Soc.,
Tram. 69: 210-215.

Heilbrunn, L. V. 1948. Outline of general phy-
siolo§i. [2nd ed.] xii + 748 pp. W. B.
Saunders Company, Philadelphia.

Hermann, L. 1885. Eine Wirkung galvan-
ische Strome auf Organismen. Pfliigers Arch.
/. die Gesam. Physiol. 37: 457-460.

Holmes, H. G. 1948. The history and devel-
opment of the electrical fish screen. U. S.
Fish and Wildlife Serv., Spec. Sci. Rpt. 53:
1-62.

ScHEMiNSKY, F. 1924. Versuche fiber Elektro-
taxis und Elektronarkose. Pfliigers Arch, f
die Gesam. Physiol. 202: 200-216.

Smolian, Kurt. 1944. Die Electrofischerei.
Samml. Fischer. Zeitfragen der Deut. Fischerei
35: 1-27.

Van Harreveld, A. 1938. On galvanatrop-
ism and oscillotaxis in fish. Jour. Expt. Biol.
15 (2): 197-208.

Biological Succession in the Aleutians^

Theodore P. Bank, IP

World War II provided "shock treatment”
to American interest in the Aleutian Islands.
Prior to 1942, the Aleutians were only vaguely
known, and thought of, if at all, as cold,
desolate, and totally inhospitable. This ig-
norance was shared by scientists, military
leaders, and, too frequently, by governmental
agents responsible for the administration of
that part of the Alaskan Territory. A sudden
awakening came when the United States and
Canada were forced by the exigency of war
to land and maintain a large force of men on
these hitherto neglected shores, and imme-
diately they assumed new importance as our
North Pacific frontier. Attesting to this new
importance is the fact that eight university-
sponsored expeditions have been sent to ex-
plore the islands in the past 4 years. Greatly
outnumbering these have been governmental
survey parties and individual investigators. In
all, more than 30 scientific projects have been
directly concerned with the Aleutian Chain
in postwar years. Moreover, this surge of
interest seems likely to continue, as several
current programs are organized on a long-
term basis.

Wartime and early postwar studies were
naturally concerned more with the strategic
potentialities of the islands for military oc-
cupation than with purely scientific questions.
However, from these investigations have come
an increasing appreciation of the peculiar eco-

^ Contribution No. 971 from the Botanical Gardens
and the Department of Botany, University of Michi-
gan. Manuscript received September 30, 1952.

2 Field Director, University of Michigan Expeditions
to the Aleutian Islands, 1948-52.

logical relationship which exists between
Aleutian environment and biota. Here a
unique set of environmental factors has mold-
ed flora, fauna, and human culture into pat-
terns quite different from those on most of
the Mainland. This paper will outline briefly
some of these patterns, which are transitional
stages of biological succession.

Field work upon which this paper is based
has been sponsored by the University of
Michigan, the Michigan Phoenix Project, and
the Office of Naval Research. In all, four
trips have been made to the Aleutians, during
which more than 20 islands were visited.
Critically important localities such as glacial
margins, hot springs, volcanic craters, and
prehistoric village sites were given special
attention by members of our expeditions, and
an attempt was made to correlate biological
and anthropological investigations with a
view toward establishing a chronology of
postglacial events in the Aleutians.

PLEISTOCENE PHYTOGEOGRAPHY

The most ambitious phytogeographical
treatment of arctic and subartic regions in
Pleistocene times is that of Dr. Eric Hulten
(1937^). For reviews of his hypotheses, the
interested reader is referred to several excel-
lent summaries which have been published
(Raup, 1941, 1947; Cain, 1944). It will be
useful, however, to outline quickly Hulten’s
major conclusions regarding the Aleutians.

Hulten undertook extensive field work in
both Kamchatka and the Aleutians, supple-
menting this with an examination of all avail-
able herbarium material from the arctic and

493

494

PACIFIC SCIENCE, Vol. VII, October, 1953

Fig. 1. The Aleutian Islands. Probable extent of ancient "Beringia” is indicated by diagonal hatching in insert.

Biological Succession in Aleutians — Bank

boreal regions. Natural ranges of hundreds
of plants were mapped. Ranges which were
more or less congruous were grouped. Each
group was thus made up of superimposed
ranges which were approximately equiformal
and of progressively larger area. Hulten termed
these "equiformal progressive areas.” He
showed that each such area has a center, i.e.,
a region where the largest number of bio-
types occurs. This is presumed to be the
center of origin, and from here the plants
dispersed to occupy their present ranges.
Those with ranges that approximate closest
the center are termed "centrants” and those
with wider ranges are termed "radiants.”
Geographically, the centers all lie outside the
maximum advance of the ice sheets and thus
probably served as the major plant refuges
during Pleistocene glaciation. Some plants
which survived in the more extensive refugia
retained their genetical ability to reproduce
and quickly occupy new ice-free areas follow-
ing the glacial periods. Others, on more iso-
lated, smaller refugia, became depauperated
of biotypes with the result that their spreading
ability was reduced. Hulten has termed them
"rigid” species. The quickly spreading plants
are termed "plastic” species.

One of the most important plant refuges
was an ice-free lowland which Hulten has
called "Beringia,” a land area thought to have
existed at various times during and imme-
diately following the Pleistocene when suf-
ficient water was withdrawn from the ocean
to expose much of the sea floor in the north-
ern and eastern Bering Sea and southern
Arctic Ocean regions (Fig. 1). From Ber-
ingia, from unglaciated portions of the eastern
Aleutians and extensive ice-free areas in Kam-
chatka, biota surviving the pleistocene are
presumed to have spread to adjacent areas as
the ice withdrew. The most highly plastic
species were the first to re-establish them-
selves on such places as the central Aleutians,
which Hulten thinks were heavily glaciated
and hence mostly devoid of plant life during
the glacial periods. Through such differential

495

migrations of plants from both ends of the
Aleutian Arc due to unequal spreading of
plastic and rigid species, the present Aleutian
flora came into being. Thus, the characteristic
middle-Aleutian gap which occurs in the dis-
tribution of many species (Hulten, 1937^.'
348-381) is the result of the inability of rigid
types to invade the islands as quickly as
plastic ones.

Hulten’s methods of arranging the equi-
formal progressive areas are open to certain
criticisms (Raup, 1947: 231): (1) additions
to his lists are undoubtedly forthcoming and
may alter some equiformal areas; (2) many
of the gaps in plant distribution can be shown
to be gaps in exploration; and (3) Hulten’s
sorting of species to illustrate equiformal
areas may represent a faulty technique. How-
ever, such criticism is comparatively minor,
and none of the recent work has seriously
altered Hulten’s general hypothesis.

An example of the second criticism of Hul-
ten’s work is found in our own studies. We
are now able to fill in many gaps of distribu-
tion of Aleutian plants. Eriophorum russeolum,
for example, was supposed by Hulten not to
occur between Umnak in the eastern Aleu-
tians and Kamchatka, but we have since
found it on Atka, Adak, Kiska, and Attu
(Fig. 1). Likewise, Prunella vulgaris var. aleu-
tica occurs on several islands of the middle
Aleutians, where it was supposedly absent.
Thus, of 105 species which theoretically have
had distributional gaps in the middle Aleu-
tians, 24 have been found on islands within
the gaps. It seems likely that more will turn
up as further exploration covers additional
ground.

Such new and important range extensions
indicate that much additional field explora-
tion is needed. Hulten’s supposition that the
middle islands were heavily glaciated during
the Pleistocene is not entirely correct, because
subsequent work has indicated that a number
of ice-free islands probably existed (U.S.G.S.,
1947): for example, west of Adak, the flat,
platform islands of Ogliugla, Ilak, Kavalga,

496

PACIFIC SCIENCE, VoL VII, October, 1953

Fig. 2. Newly exposed postglacial ash profile on Tanaga Island, August, 1950. Ash, humus, and clay strata
are clearly defined.

Unalaga, and Shemya, and parts of Kanaga,
Tanaga, etc. These undoubtedly served as
refugia for a number of hardy plants which
were able to withstand the climatic rigors and
the proximity of nearby ice caps on Adak,
the northern parts of Kanaga and Tanaga,
Amchitka, and Semisopochnoi.

POSTGLACIAL PLANT SUCCESSION

The ecological concept of succession is
hard to apply to communities which continu-
ally fluctuate in an unstable way so that a
so-called climax grouping today may be
shown later to have been only temporary.
Under such conditions, succession is difficult
indeed to define and more difficult to under-
stand, especially if brief field studies attempt
to name characteristic or dominant species
as criteria of certain stages. In the Aleutians
what one thinks of as a successional stage
or climax community is often found to con-

sist of characteristic species which occur
widely in other habitats and make up a large
part of the local flora. Moreover, the com-
munity may be replaced by another during a
relatively short period of time only to re-
appear again in slightly different form. Hulten
has called attention to this by remarking
(1937^) that the Aleutian plant associations
are very similar to those of Kamchatka, but
in the Aleutians the associations are much
less stable. There is considerable shift back
and forth in species within communities,
which results in the dropping out of various
members. There are also abrupt shifts in en-
tire plant communities which result in a rad-
ically new vegetative character of a compara-
tively large area.

In postglacial times, as Aleutian plant life
reinvaded the newly ice-free areas from va-
rious refugia, there undoubtedly occurred ma-
jor disruptive phenomena which seriously in-

Biological Succession in Aleutians — Bank

terfered with normal plant succession. During
volcanic eruptions, for instance, large areas
were completely covered by ash, and the vege-
tation was thus exterminated. Major ash falls
are clearly marked in the soil profiles (Fig. 2).
A number of such ash falls must have origi-
nated from violent eruptions which gradually
diminished to a rather continuous, compara-
tively quiet eruption of long duration. The
results of this activity appear in the profile
as layers composed of coarse pumice overlaid
by ash which becomes increasingly finer to-
ward the top. These deposits are often 0.5
meters or more thick and are almost pure
volcanic material devoid of any organic re-
mains. Frequently they separate layers of al-
most pure humus (Fig. 3). The vegetation
was destroyed by the ash fall and had to
reinvade the area, in time producing the over-
lying humus. Thus, it is somewhat possible
in the Aleutians to follow both the volcanic
history and the major accompanying disrup-
tions in plant succession.

Pollen studies of humus intercalated be-
tween ash strata offer one method of arriving
at a history of postglacial plant succession,
and for this purpose hundreds of soil samples
were secured from various ash profiles and
archaeological sites. Several of these samples
have already been examined briefly for pollen
by S. T. Anderson, and a preliminary report
of the results has been released (Anderson
and Bank, 1952). These are interesting and
worth summarizing. Two humus samples
from soil strata, one at 0.92 meters depth
and the other at 1.83 meters depth, came from
a single ash profile on Tanaga Island in the
middle Aleutians. Usually, terrestrial ash-soil
deposits should not contain much pollen, as
the grains would ordinarily be destroyed by
oxidation. It was therefore somewhat surpris-
ing to find that these samples contained suf-
ficient pollen for quantitive studies. This is
probably because of the unusual Aleutian
climate, marked by consistently high precipi-
tation and fairly low soil temperatures. Many
Aleutian soils are constantly saturated with

497

water, which undoubtedly is a major factor
in the preservation of pollen and spores.

The samples were treated according to the
procedure described by Faegri and Iversen
(1950). Pollen from the lower part of the
profile represents an entirely different plant
community from that of the upper part. The

Fig. 3. Diagrammatic profile at the edge of an Aleut
kitchen midden on Nazan Bay, Atka Island. Vertical
and diagonal cross-hatching indicates clam shell, fish
bone, and sea urchin shell deposits (archaeological).
Also shown are sea lion bones and a stone lamp in
position. The nonarchaeological strata are:' A, recent
humus; B, sand; c,E,i, fine ash, slightly stratified;
D,G,K, humms with fine ash admixture, stratified; F,
H,J, coarse pumiceous ash; L, brown clay with large
pumice (till ?); m, bedrock; 1,2,3, ferruginous bands
(from leaching from above?). Surface vegetation, es-
pecially lush on such sites, is indicated and includes:
Heracleum lanatum, Elymus arenarius subsp. mollis^ Co-
nioselinum Gmelini, Angelica lucida, Aconitum maximum,
Claytonia sibirica, Achillea borealis, and Calamagrostis
canadensis var. Langsdorffii.

498

older pollen and spores consist of high per-
centages of Erkales (55 per cent) , Gramineae
(22), and Lycopodium (17), with some Cypera-
ceae (2.9). Thus, the lower sample represents
the Empetrum heath of higher altitudes (Hul-
ten, 1937^; Bank, 1952). The younger pollen
and spores show high percentages of Grami-
neae (56), Cyperaceae (23) , and Ranunculaceae
(9.0), with some Ericaks (3.7), Dryopteris (3T),
and Umbelliferae (2.2). This sample indicates
a vegetation very similar to the Calamagrostis
meadow which exists today at the place of
the profile.

The results of this preliminary pollen study
are, of course, inconclusive. What they do
indicate is that pollen analysis of at least some
strata in Aleutian ash profiles is possible.
Certain changes in vegetation are indicated,
i.e., from an earlier Lycopodium-Empetrum
heath to a Calamagrostis-UmhtlYiitmt meadow,
and these changes can be interpreted as due
to climatic fluctuation from dry, continental-
arctic to moist, oceanic-subarctic. However,
great dependence upon pollen shifts at iso-
lated sites as an indication of widespread cli-
matic changes is extremely hazardous for two
main reasons: (1) it seems likely that heavy
ash falls not only destroyed existing vegeta-
tion but also produced such radical changes
in the local habitat that the succeeding vege-
tation may have been of totally different char-
acter for edaphic reasons; and (2) the normal
instability of Aleutian biota, to which I have
already called attention, operates to produce
major changes in natural biotic groupings for
reasons much less basic than climatic inter-
ference.

Raup (1950) has discussed this latter gen-
eral problem of physiographic ecology in
arctic and subarctic regions. He says (p. 8):

' 'Approaches to equilibrium or relative sta-
bility such as we find in the temperate regions
and are wont to call climax, are almost im-
possible to define in the Arctic and much of
the Subarctic. At best they are ephemeral in
time and space.” And again (p. 10), . . it

becomes apparent that one of the prime re-

PACIFIC SCIENCE, Vol. VII, October, 1953

quisites for development of dominance and
succession-long continued stability in the
substratum— is absent or very poorly repre-
sented over vast areas in the Arctic and Sub-
arctic.”

An example of a constantly unstable sub-
stratum in the Aleutians is found in the alpine
regions where the unceasing wind, more than
any other ecological factor, restricts plant
growth, molds vegetational structure so to
speak, and, when aided by subsurface water,
often radically changes the vegetational char-
acter of large areas in a comparatively short
period of time. Thus in high, exposed places
one frequently comes upon bare patches of
gravel or ashy soil having a slight ridge along
the lower margin. These areas are wet and are
usually on slopes. Underground seepage has
saturated the soil, which lies thinly over bed-
rock, with the result that large soil patches
have flowed downhill carrying vegetation with
them. Surface water further erodes these areas,
and wind removes all loose plant fragments.
The bare patches may be reinvaded by plants,
but irregular shallow depressions in the sur-
face of the vegetational mat identify the spot
as a former slough area. Frequently these are
continually eroded by the high winds, and
scalloped gravel pits are formed. The gravel-
blasting effect of the wind prohibits all but
the hardiest plants from gaining a foothold.
At first only gnarled, almost leafless, dwarfed
(6 inches tall) willows may grow. In time
these eroded patches are probably covered by
willows and lichens, which in turn offer some
protection from the wind to less hardy plants,
among them Campanula lasiocarpa, Primula
cuneifolia, Antennaria dioica, and Polygonum
viviparum.

Probably the best examples of vegetational
instability are found in the wet habitats. Often
these develop in the middle of an otherwise
dry Aleutian meadow or heath area. In such
places subsurface water may begin to seep
from the hillside and form bog-like drainage
slopes on which a number of bog plants,
especially Sphagnum, begin to grow. Soon

Biological Succession in Aleutians — BANK

499

Fig. 4. Temporary bog in the middle of a lowland meadow, Atka Island. Seepage from surrounding hillsides
collects here, and plants typical of wet habitats invade the area. These plants include: Carex Lyngbyaei and other
Carex spp., Flantago macrocarpa. Polygonum vhiparum, Platanthera dilatata, Juncus haltkus, Erlgeron perigrinus,
Eriophorum Chamissonis, etc.

these areas are inhabited by liverworts, Sphag-
num capillaceum, S. Girgensohnii and other
mosses, Saxifraga punctata subsp. insularis,
Leptarrhena py roll folia, Viola Langsdorffi, Pin-
guicula vulgaris, sometimes Scirpus spp., and
Carex spp. Their growth often becomes ex-
tensive and deep enough to necessitate a wide
detour when walking across a hillside. These
are usually temporary features, however, and
such wet places, with their typical vegetation,
constantly appear and disappear under the
influence of fluctuating subsurface drainage
patterns, which are in turn greatly affected
by yearly variations in snow accumulation.
True peat bogs, such as those common to
parts of mainland Alaska and Canada, are
not found in the Aleutians. Instead the bogs
are of the type just described, formed from
seepage, or are essentially low areas contain-
ing water- saturated mud overlaid by scattered

growths of sedges, Juncus spp., short grasses,
and sometimes Sphagnum (Fig. 4). Usually
such areas are shallow, but some are deep
enough to completely mire large vehicles.
Many such mud bogs are only a transitional
stage for shallow ponds and meadows. I ob-
served an amazingly quick replacement of
Carex Lynghyei and Eriophorum Chamissonis by
Calamagrostis canadensis var. Langsdorjfi and
other grasses after one such bog was drained
by military operations.

Other evidences of instability are noticeable
in stream ravines. Usually occurring here are
ferns {Athyrium Filix-femina and Dryopteris
spp.), grasses such as Deschampsia heringensis,
mosses, Streptopus amplexifolius , Epilohium an-
gusti folium var. macrophyllum, frequently Ar-
temisia unalaskensis, and various umbellifers.
The profuse vegetation frequently grows out
over the stream and in time may entirely cover

500

PACIFIC SCIENCE, VoL VII, October, 1953

Fig. 5. Diagrammatic profile of part of a typical
"slumped” stream bed and underground channel. Such
stream tunnels, A-c, are fairly common in the Aleutians
and can be dangerous should a person fall through the
ceiling B. The lower outlet A and the upper entrance
C are usually choked with ferns and other plant growth.
The tunnel ceiling usually possesses a meadow vege-
tation, consisting of grasses, umbellifers, Streptopus,
Claytonia, etc. At d a series of terraces are formed from
a former tunnel ceiling which slumped gradually and
finally caved in. The vegetation here is usually bog-
like, with Sphagnum, Carex, Juncus, liverworts, and
Saxifraga predominating.

the ditch so that the stream flows through an
underground tunnel (Fig. 5). These places
can be quite dangerous. Hulten (1937^) men-
tions falling through the ceiling of one such
tunnel. Hidden streams may persist this way
for a considerable time, but usually the over-
hanging mat is penetrated by surface drainage,
holes reappear, the overhanging mat slumps,
and the ditch becomes a series of depressions
or wet terraces composed of sphagnum and
typical bog plants, a much different vegeta-
tion than that which formerly occurred in the
ravine.

In the Aleutians, this instability of sub-
stratum and plant cover does not have precisely
the same causes as on the mainland. Perma-
nent subsurface ice (permafrost), for example,
is not the very important factor that it is over
widespread areas of Siberia, mainland Alaska,
and Canada. Rather, inst;ability is probably a
result of the interaction of the edaphic and
climatic factors which make the Aleutians
more or less unique: namely, a comparatively

thin soil mantle largely of volcanic ash, more
or less continuous ash deposition, temperate
conditions with heavy precipitation, cool
summers, and unusually high, rather constant
winds.

Kellogg and Nygard (1951) have charac-
terized the Aleutian substratum as "tundra
without permafrost” and composed of silty
or sandy loam with ash. There are, however,
small areas of permanent subsurface ice in the
Aleutians which may be fairly extensive albeit
sporadic and seemingly of little influence on
surface vegetation. I observed one such area
on Great Sitkin Island where Glacier Creek
has cut through an ancient mud flow. Ice is
exposed in the profile about 20-30 feet below
the surface of the old flow. Thick sheets and
occasional lenses of ice can be traced along
the floor of the stream bed for distances of 25
to 50 yards. Strictly speaking this is not per-
mafrost but relict glacial ice. The surface
topography and vegetation are unmodified,
and it is presumed that the depth at which
the ice occurs is too great to affect plant
growth above. Melting of the ice masses,
however, will undoubtedly cause local land
slumps leading to the formation of kettles
and, thus, different microhabitats which will
have their peculiar plant communities.

RECENT PLANT INVASIONS

The foregoing examples of fluctuating veg-
etation in the Aleutians illustrate the spread-
ing ability, i.e., plasticity, of some plants of
this area. This ability has been noted by
Hulten (1937^) who writes that on Bogoslof,
the "jack-in-the-box” island that has appeared
and disappeared a number of times in the last
150 years, a pioneering vegetation is already
well established. It includes Lathyrus mari-
timus, Elymus arenarius subsp. mollis, Honckenya
peploides major, Euccinellia pumila, and Merten-
sia maritima, all of which are widespread lit-
toral plants. Senecio psetido-arnica also occurs
on the still- warm shores. Our own field par-
ties studied briefly several pioneer plant com-
munities near a small glacier on the upper

Biological Succession in Aleutians — Bank

slopes of Great Sitkin Island (Fig. 1). We
visited the place on two occasions 12 months
apart. It was noted in 1950 that during the
preceding year a considerable extent of newly
ice-free area had been successfully invaded by
a number of plants, mostly mosses and lichens
but also including Luzula arcuata, Agrostis
exarata, Hieracium triste, Carex circinata^ and
Sibhaldia procumhens. Although the actual
creep of vegetation was in most cases only
a few feet, this was all along the margin of
the newly exposed ash, a sizeable area, and
seems remarkable for having occurred during
a single growing season.

Contrasting with this is the inability of
Picea to establish itself on Aleutian soil, de-
spite the fact that extensive spruce forests
occur on the mainlands of Alaska and Kam-
chatka at the same latitudes. On Amaknak
Island in Unalaska Bay in 1805 Russian set-
tlers planted 24 seedlings of Pkea sitchensis
which came from Sitka (Hulten, 1937^).
Other Sitka spruces were later planted on
Expedition Isle in Roose Bay and in Unalaska
Village. All these groves have been depleted
by dying trees, but parts of the original plants
are still alive, and, except for the stunted ap-
pearance and gnarled form of some of the
trees, they seem healthy. Many regularly pro-
duce cones. However, none has successfully
reproduced by seedlings. More recently, at
Adak, several dozen Sitka spruces were planted
on a hill and on a lowland. These plantings
are failures; the trees that have survived are
stunted, and there has been no reproduction.

Such lack of success on the part of the
spruce in taking hold in the Aleutians has
been attributed to various causes, among
which the acid soil conditions, high winds,
and constant fog are most often given. None
of these answers satisfactorily explains the
real reasons why the trees fail to grow in the
Aleutians, however. The so-called extreme
acidity of Aleutian soils is probably a mis-
conception (Kellogg and Nygard (1951) have
found that soil pW in the Aleutians is about
the same as that on Kodiak where there are

501

spruce forests), many of the planted groves
are fairly well sheltered from the winds, and
it is difficult to see how fog could prohibit
tree growth entirely, I can only guess at the
true reasons, as the proper studies have not
been made. It is suggested that the primary
ecological factor operating against spruce re-
generation is the absence of any prolonged
summer period of uniformly high enough
temperatures to allow maturation of seed.

CULTURAL SUCCESSION

Turning from a discussion of the effects of
Aleutian environment upon biological suc-
cession to a brief consideration of the ways
in which human culture has been conditioned
by the same environment, we find that the
Aleut was confined to a rather restricted range
of cultural adaptations, which depended upon
the sea and the shore. The Aleut seldom
visited the inland regions of his islands, partly
because topography and weather factors in
those parts made such journeys a hardship,
but mainly because the majority of his needs
were found only on the shore, in the lowland
meadows, valleys, and marshes near the sea,
or in the sea (Bank, 1952). Of more than 50
local plants which the Aleut gathered for food,
drink, medicines, and for use in the manu-
facture of such articles as grass mats, baskets,
bidarki (kyak) frames, etc., almost all occurred
along the margins of the islands. The Aleut
depended upon fish, sea mammals, shell fish,
and birds for food, and these also came from
the sea or near the shore. Thus, even if Aleut
culture had not been predominately maritime
prior to its arrival in the Aleutians, it certainly
was unable to be anything else after the Aleut
took up his habitation in the Archipelago.
Recent archaeological studies (Jochelson,
1925; Laughlin, 1951) have shown that Aleut
culture changed only slightly during 3,000
years of occupation in the islands before the
coming of the white man.

Early Russian accounts (Jochelson, 1925:
23) suggest that Aleut villages were on ex-
posed points of land and isthmuses prior to

502

PACIFIC SCIENCE, Vol. VII, October, 1953

1741. This was because survival of an attack
of a stronger warring party required access to
an escape route by sea on more than one side.
After the Russians had put an end to inter-
island warfare, the villages were supposedly
moved to sheltered bays which offered many
advantages of improved fishing and better
harbors. This view is somewhat substantiated
by vegetational patterns, for old villages lo-
cated on points of land (thus presumably pre-
Russian) usually possess a comparatively less
dense plant cover than do old villages located
in bays. The latter sites (presumably more
recently abandoned) often have a particularly
lush plant cover.

However, an outright assumption that the
oldest sites are to be found on points of land
or isthmuses along the coasts should not be
made. It is known that the islands have been
rising throughout the history of the Archi-
pelago. Jaggar (1908: 32) mentions finding
old beach terraces on Unalaska at 130 meters
altitude. Veniaminof (Petrof, 1884: 148) and
Hulten (1937^/ 24) also call attention to old
shore lines, and more recently they have been
reported by members of the U. S. Geological
Survey field parties (U.S.G.S.,1947: 102). These
places have not been explored archaeologi-
cally, although Hrdlicka (1945) mentions
several village sites in the eastern part of
Agattu at a considerable distance above the
sea, and I have also noted from the air an old
village situated on a cliff high above the sea
in the western part. It is entirely possible that
future reconnaissance will uncover the oldest
sites on such uplifted beaches, and it may be
that here is where the true beginning of Aleut
culture will have to be sought.

SUMMARY

Study of biotic patterns in the Aleutians
shows them to be extremely unstable. There
are frequent vegetational shifts which results
in a totally new plant cover over compara-
tively large areas. In postglacial times some
of these shifts may have been caused by cli-
matic changes, but it is believed that most of

them have come about through unstable soil
conditions, which in turn have been influ-
enced by fluctuations in seasonal weather.
Wind is a very important factor determining
vegetational structure and composition ar
higher altitudes. Any semblance of normat
plant succession as it is known in similal
latitudes on the mainlands of Asia and Amer-
ica is frequently altered by the combined
effects of volcanic ash eruptions, wind ero-
sion, periodic moisture surpluses (after
heavier-than-usual winter snowfalls), etc. In
general, vegetation is much less stable and
biological succession more erratic in the Aleu-
tians then on the mainlands. This is in sharp
contrast to Aleut culture, which apparently
remained unchanged in most of its major
aspects during 3,000 years of existence in the
Aleutians.

REFERENCES

Anderson, Sven T., and T. P. Bank, II.
1952. Pollen and radiocarbon studies of
Aleutian soil profiles. Science 116 (3004):
84-86.

Bank, T. P., IT 1952. Aleutian vegetation and
Aleut culture. Mich. Acad. Sci., Arts and
Letters, Papers 37: 13-30. [1951.]

Cain, Stanley A. 1944. Foundations of plant
geography, xiv -f 556 pp., 63 figs. Harper
& Bros., New York.

Faegri, K., and J. Iverson. 1950; Textbook
of modern pollen analysis. 168 pp. Munks-
gaard, Copenhagen.

Hrdlicka, Ales. 1945. The Aleutian and
Commander islands and their inhabitants.
XX + 630 pp., 239 figs. Wistar Institute
of Anatomy and Biology, Philadelphia.
Hulten, Eric. ¥)yia. Flora of the Aleutian
Islands. 397 pp., 6 figs., 16 pis., 33 maps.
Bokfoerlags Aktiebolaget Thule, Stock-
holm.

1937^. Outline of the history of arctic

and boreal biota during the Quaternary Period.
168 pp., 14 figs., 43 pis. Bokfoerlags Ak-
tiebolaget Thule, Stockholm.

Jaggar, T. A., Jr. 1908. Journal of the Tech-
nology expedition to the Aleutian Islands,

Biological Succession in Aleutians — Bank

1907. Technol. Rev. (Ellis, Boston) 10 (1):
1-37.

JOCHELSON, W. 1925. Archaeological investiga-
tions in the Aleutian Islands. 145 pp. Carnegie
Institution, Washington, D. C.

Kellogg, C. E., and 1. J. Nygard. 1951.
The principal soil groups of Alaska. U. S.
Dept. Agr., Monog. 7: 1-138.

Laughlin, W. S. 1951. The Alaska gateway
viewed from the Aleutian Islands. In: The
physical anthropology of the American Indian
(Viking Fund PubL). vii + 202 pp. Ed-
wards Bros., Inc., Ann Arbor, Michigan.
Petrof, I. 1884. Report on the populations, in-
dustries, and resources of Alaska, vi + 189

503

pp., 8 pis., 8 maps. U. S. Government
Printing Office, Washington, D. C.

Raup, H. M. 1941. Botanical problems in
boreal America. Bot. Rev. 7: 147-248.

1947. Some natural floristic areas in

boreal America. Ecol. Monog. 17: 221-234.

1950. Physiographic ecology in Alaska.

[Paper delivered before the botany session
of the First Alaska Science Conference (Na-
tional Research Council), Washington,
D. C. Distributed in mimeographed form.]
17 pp.

U. S. Geological Survey. 1947. Alaskan
volcano investigations. Rpt. 2. 105 pp., 27
figs., 8 pis. U. S. Government Printing
Office, Washington, D. C.

New Alaskan Records of Eurytemora (Crustacea, Copepoda)

I

Mildred Stratton Wilson^

The copepod genus Eurytemora is known
from all the coastal areas of North America,
though the records are not extensive. The
species occur in salt, brackish, and fresh water.
As the genus is widely spread throughout the
northern regions of the world, it is to be
expected that the coastal areas of Alaska
would have an abundance of all the habitat
forms. Unfortunately, in the Alaskan collec-
tions made during the Canadian Arctic Ex-
peditions no mature specimens were found
(Marsh, 1920: 4j). The present report is there-
fore of interest in recording two species from
western Alaska. These include a fresh-water
form new to science and an Asiatic brackish-
water species not before known in North
America.

I am indebted for these collections to Dr.
Clifford O. Berg of the Arctic Health Re'
search Center and to Mr. Frank P. Pauls of
the Alaska Department of Health. Specimens
of each species have been deposited in the
United States National Museum.

Eurytemora yukonensis n. sp.

Figs. 1-5

SPECIMENS EXAMINED. Four lots, fresh wa-
ter, lower Yukon River area, western Alaska.
C. O. Berg, collector.

Type lot: 25 9 (many ovigerous); 13 cf ;
many copepodids. Wilson Creek, about a mile
southeast of Fortuna Fedge (formerly and
popularly known as Marshall), July 27, 1951.

1 Arctic Health Research Center, United States Pub-
lic Health Service, Federal Security Agency, Anchorage,
Alaska. Manuscript received October 10, 1952.

Holotype 9, U.S.N.M. 93268; allotype ci^,
U.S.N.M. 93269.

Lot 2: A 9,5 cf, many copepodids. Near
outlet of stream into Yukon River, about 14
mile north of Fortuna Fedge, July 28, 1951.

Lot 3: A 9 , copepodid stage V. Small lake
southwest of Holy Cross, June 15, 1951.

Lot 4: 1 9 , 8 d^, 4 copepodids. Widened
portion of stream emptying into Andreafsky
River, about 1 mile east of Andreafsky, Au-
gust 16, 1951.

Female: Total length, dorsal mid-line,
1.645 mm. (metasome 1.01 mm., urosome
0.635 mm.). Head rounded, body tapering,
the greatest width in segment 2 (0.415 mm.);
width of segment 5, 0.34 mm. First segment
showing in lateral view (Fig. 1) a pronounced
cephalic depression and a prominent medial
distal protuberance. "'Wings” of last segment
very wide, slanting outward, without pro-
nounced inner lobes, entire expanse greater
than width of second segment (0.48 mm.),

Urosome equaling 38 per cent of total
length. Relative lengths of segments and
caudal rami, exclusive of caudal setae:

1 2 3 CR

50 33 45 61

Genital segment constricted in middle, a
little asymmetrical, both proximal and distal
lobes prominent, proximal wider than distal,
both armed with a few lateral sensory hairs.
Genital operculum small, nearly hemispheri-
cal, distal edge quite rounded. Segments 2
and 3 without ornamentation.

Caudal rami shorter than segments 2 and
3 combined, their length about 5.5 X their
width; unarmed except for inner marginal

504

Alaskan Records of Burytemora — -Wilson

hairs. Longest of caudal setae equaling length
of rami.

Antennules 24-segmented, reaching to near
end of metasome segment 5. Division be-
tween segments 8 and 9 distinct on inner side,
but not always on outer. Aesthetes with stalk-
like bases (Fig. 2). Setae not long, none
reaching farther than to near end of succeed-
ing two segments. Numerical setation as fol-

lows (s—

■seta, sp—spine.

a— aesthete) :

Segment

1—35, a

13~2s

2— 3s, a

14— 2s, a

3-2s, a

15— 2s

4— Is

16— 2s, a

5— 2s, a

17— 2s

6— Is

18— 2s

7— 2s, a

19-2S, a

8— sp

20— Is

9— 2s, a

21— Is

10— sp

22 — Is 4“ Is

11— 2s, a

23— Is+ls, a

12— Is, a, sp

24— 6s, a

Mandible blade (Fig. 5) with anterior side
rounded, produced to moderately sized den-
ticle with small transparent cap; symmetrical,
moderately deep gap present between it and
following denticle.

Leg 5 (Fig. 2) having basipod and exopod
subequal in length. Lateral spines of exopod
1 short, not as long as width of segment, tips
sinuous. Inner process nearly straight, form-
ing right angle with segment, its width greater
than length of segment (22:15, measured
from notch at inner lower base); margins
serrate. Terminal inner spine of exopod 2 as
long as total exopod, proportion to outer
lateral spine 25:7; segment produced to min-
ute point between the two spines ; inner mar-
gin of segment with two extremely fine hairs.

Male: Total length dorsal mid-line 1.38
mm. (metasome 0.81, urosome 0.57 mm.).
First segment with cephalic depression and
prominent medial dorsal protuberance as in
female.

Urosome 41 per cent of total length of
body. All segments lacking ornamentation.
Caudal rami a little shorter than segments 3-5

505

Fig. 1. Eurytemora yukonensis n. sp. Female. Meta-
somal segment 6 and urosome, dorsal view; b, genital
segment with operculum, ventral view; c, habitus, lat-
eral view of cephalic segment; <7, habitus, dorsal view.

combined, about 35 per cent of total length
of urosome; their length about 7 X their
width. Rami unornamented, except for long
hairs on inner margins. Longest caudal setae
longer than rami.

Antennules reaching a little beyond meta-
some. The left differing from that of female
in having only 1 seta on segment 11, and in
having aesthetes on all of segments 1-19,
except on 4. Aesthetes longer than in female,
those of segments 12 and 14 particularly long,
reaching to about end of following two seg-
ments. Right antennule with spines on seg-
ments 8-12, length of all less than width of

506

PACIFIC SCIENCE, Vol VII, October, 1953

Fig. 2. Eury femora yukonensis n. sp. Female, a. Leg
5; b, leg 5, detail of armature of exopod segments 1-2;
c, antennule, aesthete of segment 11.

segments, those of 9 and 12 longest, subequal
to each other (Fig. 3). Segments 14-16 very
swollen. Segments 17 and 18 with marginal,
hyaline, depressed processes ornamented with
double row of spinous serrations. Penultimate
segment with proximal curving group of long,
slender spinules, with striated lamina follow-
ing; distal portion without ornamentation.
Terminal segment somewhat longer than pre-
ceding segment (50:40).

Leg 5 (Fig. 3) having both rami 4-seg-
mented. Basal portion narrow, first segment
of left leg longer than that of right. Right
leg characterized by an extremely long ter-
minal segment. Relative lengths of basipod
2 and exopod segments 1 and 2: 40:50:73.
Inner distal portion of second basipod seg-
ment swollen a little. Terminal segment not
only extremely long but very slender, without

any inner basal swelling, and with distal inner
margin very irregular.

Second basipod segment of left leg short,
swollen inside. Terminal segment a little long-
er than exopod 1 (45:40), apex broadened
but hardly produced on either side; inner por-
tion and most of anterior side set with short
hairs.

COPEPODID STAGES (Fig. 4). Copepodid
stages 3, 4, and 5 were present in the collec-
tions. All had a prominent dorsal protuber-
ance of the cephalic segment as in the adult.
Stage V is briefly described.

Female, stage V: Total length 1.29 mm.
(metasome 0.8, urosome 0.49 mm.). Uro-
some 3-segmented, genital segment without
lateral protrusions. Proportion of caudal rami
to rest of urosome 12:20. Antennule 23-seg-
mented. Leg 5 having inner process of exopod
short, lateral spines all short as in adult, ter-
minal spine also about as long as exopod,
spinous process between terminal spines more
pronounced than in adult.

Male, stage V: Total length 1.175 mm.
(metasome 0.725, urosome 0.45 mm.). Uro-
some 4-segmented. Leg 5 with 4-segmented
rami, left a little shorter than right, apical
segments very similar, others showing a slight
asymmetry, most pronounced difference be-
ing rounded protrusion of inner margin of
right basipod 2.

DIAGNOSTIC CHARACTERS. The diagnostic

characters of this species may be summarized

as follows:

Pronounced dorsal protuberance of cephalic
segment in both sexes in adult and copepodid
stages 3-5.

Genital segment of female strongly con-
stricted, proximal lateral lobes wider than dis-
tal; genital operculum hemispherical.

Urosome and caudal rami in both sexes
lacking dorsal ornamentation.

Caudal rami of female shorter than seg-
ments 2 and 3 combined; those of male short-
er than segments 3-5, their length about 7 X
their width.

Alaskan Records of Eurytemora — Wilson

507

Fig, 3. Eurytemora yukonensis n. sp. Male, a, Leg 5, posterior view; b, leg 5, detail of apex of left exopod, pos-
terior view; c, left antennule, segments 11-13, with detail of aesthetes; d, right antennule, apical segments;
e, right antennule, segments 8-12, with detail of spines;/, urosome, dorsal view.

Mandible blade with symmetrical gap be-
tween anterior edge and succeeding denticle.

Spines on segments 8-12 of right antennule
of male shorter than width of segments, those
of 9 and 12 longest.

Leg 5 of female with 2 spines on exopod
1; inner process straight, wider than length
of segment; lateral spines short. Inner ter-
minal spine of exopod 2 as long as whole
exopod; this segment with inner hairs.

Leg 5 of male with both rami 4-segmented,
claw of right side very much longer than pre-
ceding segment (73:50). Terminal portion of
left leg expanded but without digitiform'
protrusion.

COMPARISON WITH OTHER SPECIES

The fifth leg of the female most closely re-
sembles that of Eurytemora lacustris Poppe (as

described by Sars, 1903) . It further agrees with
this species in lacking dorsal ornamentation
on the urosome and caudal rami. It differs in
having the last metasomal segment produced
into conspicuous wings and the genital seg-
ment swollen and constricted laterally. The
left fifth leg of the male of yukonensis differs
from lacustris in not having the terminal por-
tion produced into a prominent digit.

Eurytemora composita Keiser
Figs. 5-7

Eurytemora composita Keiser, 1929: 301, figs.
1-6.

Eurytemora composita Smirnov, 1929: 318, figs.
1-2.

Eurytemora composita Rylov, 1930: 234, fig.
77 (after Keiser).

508

Fig. 4. Eurytemora yukonensh n. sp. Copepodid stage
V. a. Female, leg 5 with detail of apex of exopod;
Z’j male, leg 5, posterior view.

lEurytemora spec. Kiefer, 1938: 78, figs.

1-3 (d^).

SPECIMENS EXAMINED. One 9 . Tundra pond
(brackish.^) near Unalakleet, on Norton
Sound, western Alaska, August 7, 1951, C.
O. Berg.

Five 9 (4 ovigerous),8 cf , 21 copepodids.
Pond (fresh water which might become brack-
ish at times). Boxer Bay, southwest cape of
St. Lawrence Island, Bering Sea, Alaska, July
6, 1952, F. P. Pauls.

DESCRIPTION OF ALASKAN SPECIMENS.

Female: Total length dorsal mid-line: St. Law-
rence Island specimens 1.21-1.46 mm.; Un-
alakleet specimen 1.41 mm. (metasome 0.68,
urosome 0.55 mm.). Body form in dorsal view

PACIFIC SCIENCE, Vol. VII, October, 1953

similar to that of yukonensis. Relative lengths
of the segments:

1 2 3 4 5 6

110 50 40 32 15 9

Greatest dorsal width of metasome in seg-
ments 2 and 3 (Unalakleet specimen 0.37
mm.; reduced to 0.295 mm. in segment 5).
First segment in lateral view (Fig. 6) showing
no cephalic depression or medial dorsal pro-
tuberance. Wings of sixth segment a little
asymmetrical, each with well-developed,
rounded inner lobe; outer portion produced
into outwardly and backwardly directed pro-
trusions reaching about to middle of genital
segment and armed with scattered hairs.

Urosome equaling 39 per cent of total
length. Relative lengths of urosome segments
(mid-line) and caudal rami, exclusive of setae:

1 2 3 CR

46 27 31 60

Genital segment constricted in the middle,
upper part with nearly symmetrical, rounded
lobes produced laterally beyond those of dis-
tal part, armed marginally with hairs. Genital
operculum conspicuous in lateral profile (Fig.
6d), occupying a comparatively large part of
ventral area which is without cuticular folds
or tumid protrusions. A heavy, sclerotized,
semicircular line encloses area of receptacula,
portion of operculum distad to this consists
of a large, rounded flap.

Fig. 5. Apex of mandible blade, female, a, Eury-
temora yukonensis n. sp.; b, Eurytemora composita Keiser.

Alaskan Records of Eurytemora — Wilson

Dorsal surface of segments 2 and 3 spinul-
ose except in medial portion.

Caudal rami subequal in length to segments
2 and 3 combined; length about 6 X their
width (measured without pressure) ; their dor-
sal surface entirely covered with short, stiff
hairs extending over outer margin to base oj
lateral seta; inner margin with closely set long
hairs. Inner terminal seta longest, equaling
length of ramus.

Antennules shorter than metasome, reach-
ing to about middle of fifth somite; 24-
segmented, segments 8 and 9 imperfectly
separated, Setation typical as in female of
yukonensis, aesthetes on narrow basal stalks.
Setae comparatively long, particularly on seg-
ments 7-20; those of 15 and 20 longest, those
of 15 of nearly uniform length, reaching to
near end of or beyond segment 18.

Mandible blade (Fig. 5) having anterior
edge produced to a large triangular tooth,
usual gap between it and succeeding denticle
filled by a rounded protuberance.

Leg 5 (Fig. id) having exopod (exclusive
of spines) only a little longer than basipod,
25:20. Spines of exopod 1 longer than width
of proximal part of segment. Lateral spine of
exopod 2 nearly as long as terminal inner
spine. Relative lengths of spines to one an-
other:

Segment 1 Segment 2

9 : 11 13 : 18

Exopod 1 not quite twice length of exopod
2 (16:9)- Inner process, measured from inner
base of segment 2, a little longer than total
length of its segment (19:16); tips markedly
upcurved, margins with a few exceedingly
fine serrations. Lateral spines appearing un-
armed except under high power which shows
that their margins are also minutely serrate.
Terminal inner spine of exopod 2 relatively
short, about twice length of segment which
is not produced to a spinous point between
it and lateral spine. Two very fine hair-like
setae on inner margin of segment.

Male: Total length dorsal mid-line about
1.07 mm. (metasome 0.605, urosome 0.465

509

Fig. 6. Eurytemora composita Keiser. Female, a, Meta-
somal segment 6 and urosome, dorsal view; b, oper-
culum and lateral outline of genital segment, ventral
view; c, habitus, lateral view; d, outline of metasomal
wings and genital operculum, lateral view.

mm.). Cephalic segment in lateral view show-
ing no depression or distal protuberance. Last
segment not produced laterally, armed with
a few hairs.

Urosome (Fig. 7) 43 per cent of total length
of body. Segments 1-4 armed laterally with
scattered hairs, segment 5 with short spinu-
lose hairs on outer dorsal surface and lateral
margins.

Caudal rami (exclusive of setae) a little
longer than segments 3-5 combined; equaling
about 39 per cent of total length of urosome;
their length nearly 10 X their greatest width.
Outer margin with closely set coarse hairs
which may or may not extend onto dorsal

510

surface in area of lateral seta; in an individual,
this dorsal ornamentation may be present on
one ramus but not on the other. Inner margin
with long hairs. Terminal inner setae longer
than caudal rami (about 70:58).

Left antennule with setation typical, differ-
ing from that of female in having 1 seta on
segment 11, and in having aesthetes on all of
segments 1-19, except on segment 4; aesthetes
also considerably longer, as in yukonensis.
Right antennule with segments 12-16 very
swollen. Segments 8-12 with spines, all of
which are shorter than width of segments;
those of 9 and 12 only a little longer than the
others. Terminal segments armed as m yukon-
ensis. Apical segment only a little longer than
the preceding.

Mandible as in female.

Leg 5 (Fig. la) having left basal portion
wider than right and with an inner protrusion.
Right second basal segment enlarged, its
greatest width to length 20:30, without
marked inner protrusion. Right exopod 2-
segmented, first segment only a little longer
than basipod 2, but much narrower. Terminal
segment longer than exopod 1, 47:35; inner
basal portion without a conspicuous swollen
portion; terminal part with cuticle of both
margins irregular. Left leg having second basal
segment shorter than that of right side, with-
out marked inner protrusion. Exopod 2-seg-
mented, both segments with a stout outer
spine; terminal segment longer than the first,
45:30. Apical portion of segment 2 broad-
ened, outer part extended as a swollen, round-
ed digit, tipped with small seta; inner portion
having cuticle crenate, anterior side with pad
of thickly set fine hairs which protrude be-
yond margin.

COMPARISON OF ASIATIC AND ALASKAN
FORMS

Eurytemora composita has been known
only from the type locality. Lake Issyk-Kul,
a large, brackish lake in western Turkestan.
Some of Reiser’s specimens were examined
by Smirnov, who supplemented the descrip-

PACIFIC SCIENCE, Vol. VII, October, 1953

tion of the type material, giving a more exact
illustration of the male fifth leg. Since the
two descriptions otherwise agree very well,
it may be assumed that the Asiatic form is
well enough known to compare it with the
Alaskan form with some degree of certainty.
The details of the antennules and mandible
are not known for the typical form.

The Asiatic and Alaskan females exhibit
the following similarities, all of which are
diagnostic characters of the species:

Size— 1.2-1. 4 mm.

Metasomal wings produced, with inner
lobes.

Genital segment with lateral hairs, con-
stricted medially, with prominent, symme-
trical, proximal lobes.

Genital operculum rounded distally, recep-
tacula located above medial sclerotization.

Anal segment and caudal rami with dorsal
ornamentation.

Caudal rami subequal to segments 2 and 3
(or a little longer), length 6 X their width.

Shape and relative proportions of segments
and spines of leg 5.

Presence of inner setae on exopod 2 of leg 5 .

The Alaskan female differs from the typical
form in:

Presence of dorsal ornamentation on second
segment of urosome.

Possible greater development of genital
operculum.

Leg 5 — Upcurved tip of inner process of
exopod 1 and presence of two slender hair-
like setae rather than a single stout seta on
inner margin of exopod 2.

No illustration of the male urosome has
been given for typical composita, but Keiser
states that the length of the rami is 7 X their
width, and that hairs are present only on the
inner margin. The Alaskan specimens differ
in having the length of the rami about 10 X
their width, and in the presence of hairs on the
anal segment and on both margins of the
rami. The fifth leg appears to be very similar
in the proportions of the segments and the
structure of the apical part of the left leg.

Alaskan Records of Eurytemora — Wilson

511

Kiefer’s brief description of two males from
Paramushir Island in the Kuriles of northern
Japan agrees in most points with Eurytemora
composita. It is like the Alaskan form in having
hairs on the anal segment. There are some
slight differences in the fifth leg, though the
proportions are similar. Without knowing the
characters of the female, it would not be en-
tirely correct to ascribe Kiefer’s specimens to
composita. However, the widely separated Tur-
kestan and Alaskan localities suggest the

Fig. 7. Eurytemora composita Keiser. a, Male, leg 5,
posterior view; b, male, leg 5, detail of apex of left
exopod, posterior view; f, male, urosomal segments
4-5 and caudal ramus; d, female, leg 5.

probable occurrence of the species in inter-
mediate places.

The occurrence of Eurytemora composita in
an isolated lake of western Asia led Gurney
(1931: 186) to suggest its origin from one of
the species of the Caspian Sea (£. affinis, velox,
grimmi). Such a supposition is now difficult
to maintain. The wider distribution of the
species, as evidenced by its presence on the
Bering coast of Alaska and its possible oc-
currence on the northeast coast of Asia, sug-
gests instead a northern origin. This is sup-
ported by the fact that composita is closely
allied to E. raboti Richard, which occurs in
fresh and brackish ponds in the Arctic regions
(Spitzbergen, Novaya Zemlya, the White Sea
coast). The relationship of these two species
has been pointed out by Smirnov (1929: 320)
and Gurney (1931: 186). The presence of E.
composita in an isolated brackish lake in eastern
Turkestan may be fortuitous or may indicate
the western edge of its distributional range.

SUMMARY

A new fresh-water species, Eurytemora yu-
konensis, is described from several localities
on the lower Yukon River, Alaska.

Eurytemora composita Keiser is recorded and
described from ponds at Unalakleet, Norton
Sound, and St. Lawrence Island, Bering Sea,
Alaska. It has been known only from eastern
Turkestan. Two unidentified males described
by Kiefer from the Kuriles, Japan, are similar
to the Alaskan material and may be the same
species. Since E. composita is alliod to the Arc-
tic species, E. raboti, its occurrence in Alaska
suggests a northern rather than a Caspian
origin.

REFERENCES

Gurney, Robert. 1931. British fresh- water
Copepoda. Vol. I. 238 pp., 344 figs. Ray
Society, London.

Keiser, N. 1929. Uber eine neue Art der
Gattung Eurytemora— Eurytemora composita
sp. nova. Zool. Anz. 80: 301-305, 6 figs.

512

PACIFIC SCIENCE, Vol. VII, October, 1953

Kiefer, Friedrich. 1938. Freilebende Siiss-
wassercopepoden von den Nordkurilen.
Biogeog. Soc. Japan, Bui. 8 (4): 75-94, 30

figs.

Marsh, C. Dwight. 1920. The fresh water
Copepoda of the Canadian Arctic Expedi-
tion 1913-18. Rpt. Canad. Arctic Exped.
1913-18, 7 (Pt. J): 1-25, pis. 1-5.

Rylov, W. M. 1930. The fresh-water cala-
noids of the U. S. S. R. In: Keys to determina-

tion of fresh -water organisms of the U. S. S. R.
A. Fresh-water fauna. 288 pp., 88 figs.
Leningrad.

Sars, G. O. I9OI-I903. An account of the
Crustacea of Norway. 4. Copepoda Cala-
noida. 171 pp., 102 pis. Bergen Museum,
Bergen.

Smirnov, S. S. 1929. Notiz iiber Eurytemora
composita Keiser. Zool. Anz. 85: 317-320,

2 figs.

The Bibionidae of New Zealand (Diptera)

D. Elmo Hardy^

The family Bibionidae is poorly represented
in the New Zealand fauna; only two genera
have been recorded to date. These are repre-
sented by six species of Philia and one species
of Bibio, The Bibio is an Australian species,
but the Philia species are endemic and known
only from New Zealand. It is interesting to
note that two of the New Zealand species
represent very nearly the minimum and maxi-
mum ranges in size for the Philia of the world.
Specimens of P. tuthilli n. sp. range down to
about 2.2 mm. in length. I know of only one
species in the entire genus which is possibly
smaller than this— P. gratiosa (Bigot) of India
is reputed to be 2.0 mm. in length. Specimens
of P. nigrostigma (Walker) measure up to 10.0
mm. in length. The only other species which
I know that is this large is P. gigantea (Mac-
quart) from the Neotropical Region.

This study is based largely upon material
which was collected by Dr. L. D. Tuthill of
the University of Hawaii and Mr. R. A. Har-
rison, Entomologist, Department of Scientific
and Industrial Research, Auckland, New Zea-
land. I am indebted to these workers for the
privilege of studying this interesting material.

Bibio imitator Walker

Bibio imitator 1834, Ent. Mag. 2: 470.

B. substitutis Walker, 1848, List. Dipt. Brit.

Mus. 1: 121 Q synonym).

B. fulvipennis Macquart, 1850, Dipt. Exot.

Suppl. 4: 17.

B. ruficoxisyi2^cogdd.n, 1850, Dipt. Exot. Suppl.

4: 17.

. 1 Department of Zoology and Entomology, Uni-
versity of Hawaii. Manuscript received May 5, 1952.

B. elegans Jaennicke, 1867, Abhandl. Senck-

end. Ges. 6: 317.

B. helioscops Schiner, 1868, Reise Novara, Dipt.

2: 20.

This is the only species of Bibio which has
been recorded from New Zealand. It is dis-
tinguished from other Pacific Bibio by the
all-black coloring, brown fumose wings, and
dense yellow pile on the -body of the males
and by the all-rufous head, thorax, and ab-
domen, and the dark reddish-brown to black
legs (excluding the coxae) of the females.

length: Body, 6. 5-7. 5 mm.; wings, 6.5-
8.0 mm.

TYPE locality: "New Holland.”

Type in the British Museum (Natural His-
tory). The species occurs in Australia, Tas-
mania, and New Zealand. I have not seen it
from the latter country but have seen a large
series of specimens from many localities in
Australia.

Key to Philia known from New Zealand

1. Wing venation normal, no incomplete

veins and crossveins 2

Venation incomplete, the m crossvein and
the bases of veins M1+2 and Mi are lacking;

very tiny species, body 2.2-2.63 mm

tuthilli n. sp.

2. Eront tibiae each with 3 spines, arranged

on a strong prominence at the upper third
of the segment, and a single spine situated
at the apical two thirds of the tibia (Eig.
^a). Wings brownish fumose. Large spe-
cies, body 6.5-10.0 mm. in length

nigrostigma (Walker)

Spines on front tibiae not arranged as

513

514

PACIFIC SCIENCE, VoL VII, October, 1953

Fig. 1. P. crinata Hardy, a. Front tibia, median portion;

tergum of male; d, male genitalia (ventral aspect).

above. Wings hyaline or but faintly yellow-
ish fumose. Smaller species, body length
under 6.0 mm 3

3. Only 3 spines on each front tibia, above

the apical set (Fig. 2a) . . . .harrisoni n. sp.
Five spines present on front tibia, above
the apical set; these are arranged in two
groups (Fig. la) 4

4. The two sets of spines located rather close
together near the middle of the tibia (Fig.
la). Apical spurs on the front tibiae much
stronger than the spines (Fig. \a). Hind
metatarsi of males not broader than apices

of tibiae 5

The two sets rather widely spaced, the first
is at the basal third of the tibia and the
second is near the apical two thirds (Fig.
Gh). The apical spur about equal in size to
the spines (Fig. Ga). The hind metatarsi of
the males broader than the apices of the
tibiae segnis (Hutton)

5. Pile of body very dense and long, the hairs
on the scutellum and in the dorsocentral
areas of the mesonotum are about equal
to one half the length of the halteres.

, apical portion of front tibia (lateral aspect) ; c, ninth

Wings slightly yellow fumose, posterior
veins brownish. Radial vein setulose
throughout its length. Tibiae of male
yellowish colored, lighter than the femora.

The spur at the apex of each front tibia
about 1.5 times as long as the apical spines

(Fig. ih) crinata Hardy

Pile rather sparse and short, scarcely ex-
tending higher than the spines of the an-
terior row on the dorsum of the thorax.
Wings hyaline, posterior veins colorless.
Radial vein nearly bare. Legs of male
entirely dark reddish brown to black. The |
spur on the front tibia twice as long as |
the apical spines (Fig. Aa) insolita (Hutton) Jf

Philia crinata Hardy
Fig. la- d

Philia crinata Hardy, 1951, Hawaii. Ent. Soc.,
Proc. 14: 262.

This species is related to P. insolita (Hutton)
but is distinguished by the characters given
in the above key and by genital characters
(cf. Figs. Ic, id, Ah, and Ac). The species has

Bibionidae of New Zealand — HARDY

been adequately described except for the gen-
ital characters. The antennal segments are
more distinctly separated in the specimens
at hand than they were in the type series, and
the antennae appear to possess 13 segments.

MALE GENITALIA: The ninth sternum is
rounded on the sides and is slightly broader
than long. The cleft on the hind margin ex-
tends about 0.25 the length of the segment
and a tumescence is present in the middle of
the hind margin. The claspers are rather blunt-
ly pointed on their inner apices (Fig. \d).
The ninth ter gum is about 1.3 times broader
than long, its hind margin is straight, and its
posterior margin has a flat-topped projection
in the median portion (Fig. Ic).

length: Body and wings, 4.5-5. 5 mm.

TYPE locality: Mt. Wellington, New Zea-
land.

Type in the United States National Mu-
seum.

I have seen additional specimens of this
from the following localities in New Zealand:
National Park, 4,000 feet, Nov. 6, 1949 (K.
P. Lamb), and Waiouru, Oct. 19, 1943, labeled
"X galls? on Olearia virgatd' (D. Hatch).
[These would not have come from the galls;
they were probably attracted to the plant for
feeding.]

Philia harrisoni n. sp.

Fig. 2a-d

This species is related to Philia varipes
(Skuse) from Australia. It is distinguished
by the short spurs on the front tibiae (Fig.
2a) (equal in size to the apical spines, not
twice as long); by the more slender basitarsi
of the hind legs (Fig. 2b) ; by the black pilose
thorax and abdomen; and by the differences
in the male genitalia as shown in Figures 2c,
d and 3^, h. It is also related to P. acutidens
(Edwards) but is separated by having only 3
spines in the middle of each front tibia and
by the black pilose body.

MALE: Entirely polished black except for a
slight rufous tinge in the ground color of the

515

Fig. 2. P. harrisoni n. sp. a, Front tibia; b, hind leg
of male; c, ninth tergum of male; d, male genitalia
(ventral aspect).

pleura, humeral ridges, tibiae, and tarsi. All
pile is dark brown to black. Head: The rostrum
is about 0.5 as long as the lower portion of
the compound eyes and is not produced be-
yond the bases of the antennae. The antennae

516

PACIFIC SCIENCE, Vol. VII, October, 1953

= .16 mm.

Fig. 3. P. varipes (Skuse). a. Ninth tergum; h, male
genitalia (ventral aspect).

are 13 segmented, the apical 4 segments are
very closely joined. Thorax: With abundant
long pile down the dorsocentral areas, around
the margins, and on the scutellum. There are
10 teeth in the anterior comb and 12 in the
posterior. The stems of the halteres are yellow-
ish, the knobs are dark brown to black. Legs:
The front tibiae have 3 spines arranged in a
transverse row near the middle of the segment.
Seven spines are in the apical set and the apical
spur is nearly equal in size to the spines (Fig.
2a). The hind tibiae are clavate, at their apices
they are about equal in width to the widest
portion of the femora. The tarsal subsegments
are strongly swollen. The basitarsus is not as
wide as the apex of the tibia and is about
twice as long as wide (Fig. 2h). Wings: Nearly
hyaline, very faintly fumose. The stigma is
brown in color and oval in shape. The costa
extends to about halfway between the tips of
veins Rs and Mi. The anterior veins are brown.

and the posteriors are colorless or but slightly
yellowed . The venation is typical for the genus .
Abdomen: About equal in length to the re-
mainder of the body. Compared to varipes and
related species, the abdomen is rather sparsely
pilose. Genitalia: The ninth tergum is over
twice as broad as long and is straight or very
slightly convex on the hind margin (Fig. 2c).
The ninth sternum is cleft 0.3 to 0.25 of its
length on the hind margin and has a tume-
scence in the middle of this concavity. The
claspers are simple and are slightly pointed
at their apices (Fig. 2d).

length: Body, 3.6 mm.; wings, 3.8 mm.
female: Unknown.

Holotype male, Chateau Track, National
Park, Feb. 27, 1949 (R. A. Harrison). Eight
paratypes, all males: one same data as type;
one from Mangatepopo, National Park, Feb,
26, 1949 (R. A. Harrison) and six from ” Above
Mangatepopo Hut,” National Park, Feb. 26,
1949 (R. A. Harrison).

The type and four paratypes have been re-
turned to Mr. Harrison, Department of Scien-
tific and Industrial Research, Auckland, New
Zealand, One paratype has been deposited in
the British Museum (Natural History), one
is at the U. S. National Museum, one in the
Bishop Museum, Honolulu, Hawaii, and one
is at the University of Hawaii.

Philia insolita (Hutton)

Fig. Aa-c

Dilophus insolitus Hutton, 1901, New Zealand
Inst., Trans. 34: 193.

This species has been adequately described,
except for genital characters, in my paper on
the Pacific Philia (Hardy, 1951, Hawaii. Ent.
Soc., Proc. 14: 264-265). It is related to P.
segnis (Hutton) and P. crinata Hardy and is
distinguished by the characteristics given in
the above key and by the male genitalia as in
Figures. Ab, c, 6c, \d, and as described below.

MALE GENITALIA: The ninth tergum is about
1.3 times as wide as long, and the hind margin

Bibionidae of New Zealand — HARDY

517

is straight or nearly so (Fig. Ab). The ninth
sternum is about as broad as long, and the
cleft on the hind margin extends about 0.25
the length of the segment. The claspers are
sharp pointed on their inner apices (Fig. 4c).

length: Body and wings, 4. 3-5. 5 mm.

TYPE LOCALITY: Christchurch.

The type is in the Canterbury Museum,
Christchurch.

I have previously recorded this species from
several localities in New Zealand.

Fig. 4. P. insolita (Hutton), a, Apex of front tibia
(lateral aspect); b, ninth tergum of male; c, male geni-
talia (ventral aspect).

Philia nigrostigma (Walker)

Fig. ^a-c

Bibio nigrostigma Walker, 1848, Cat. Dipt.

Brit. Mus., p. 121.

B. zealandicus^2L\]s.tt, 1858, Ent. Soc. London,

Trans, (n.s.) 4: 235. New synonymy.
Dilophus spectabilis Nowicki, 1875, Krakauer

K. K. Akad. d. Wissen., Mem. 2: 10.

Bibio zealandicus Walker has been pre-
viously treated as a variety of nigrostigma,
distinguished from the typical form by having
the abdomen of the female ferruginous below
instead of black. The coloration of the fe-
males has been found to be so variable that
it does not appear practical to recognize this
variety. This species has been adequately de-
scribed, except for the genital characters, in
my paper on the Pacific Bhilia (Hardy, 1951,
Hawaii. Ent. Soc., Proc. 14: 268-270). It can
be readily distinguished from the other New
Zealand species by its large size, the brown
fumose wings, and the arrangement of the
spines on the front tibiae (Fig. 5^).

MALE GENITALIA: The ninth tergum is near-
ly twice as wide as long and is gently concave
on its hind margin (Fig. 5^). The posterior
lateral margins of the ninth sternum are pro-
duced into a pair of short lobes which extend
about halfway to the apices of the claspers.
The claspers are large and conspicuous and
are enlarged at their apices and shaped like
a hammer head (Fig. 5c). The ninth sternum
is slightly broader than long.

length: Body and wings, 6.5-10.0 mm.

TYPE LOCALITY: "New Zealand.”

The type is in the British Museum (Natural
History) .

I have previously recorded this species from
several localities in New Zealand. Specimens
have since been seen from the following:
Southeast of Taupo, 3,000 feet, Dec. 7, 1949
(L. D. Tuthill); Mt. Wellington, Lava Fields,
Nov. 12, 1948 (S. A. Rumsey); Owairaka,
Nov. 16, 1940 (D. Spiller) and Avondale,
Auckland, Dec., 1949 (R. A. Harrison).

518

PACIFIC SCIENCE, Vol. VII, October, 1953

Fig. 5. P. nigrostigma (Walker), a. Front tibia, median
portion; b, ninth tergum of male; r, male genitalia
(ventral aspect).

Philia segnis (Hutton)

Fig. 6a-c

Dilophus segnis Hutton, 1901, New Zealand
Inst., Trans. 34: 194.

This species is related to P. insolita (Hutton)
but is distinguished by the arrangement of

the spines on the front tibiae (Fig. 6b) ; by the
small, poorly developed apical spurs of the
front tibiae (Fig. 6a) ; and by having the hind
metatarsi of the males more strongly swollen,
broader than the apices of the tibiae. The
ninth tergum is twice as wide as long, and
the hind margin is very slightly concave (Fig.
6c). The ninth sternum and the claspers are
shaped as in P. insolita (Fig. 4c). For further
descriptions and figures of this species see
Hardy, 1951 (Hawaii. Ent. Soc., Proc. 14:
272-273).

LENGTH: Body, 4. 5-5.0 mm.; wings, 4.0-
4.2 mm.

TYPE LOCALITY: Christchurch, New Zea-
land.

The type is in the Canterbury Museum,
Christchurch.

I have previously recorded this species from
several localities in New Zealand. Two female
specimens have recently been seen which ap-
pear to be this species; they were from Blen-
heim, Nov. 15, 1942 (D. Spiller) and Titi-
rangi, Auckland, Oct. 2, 1949 (E. Bray).

Philia tuthilli n. sp.

Fig. la-f

This remarkable species is readily differen-
tiated from any known member of the genus
Philia by its incomplete wing venation. This
is the only species which I have seen from
the entire world which lacks the m crossvein
(posterior crossvein) and which has the bases
of veins Mi-j-2 and Mi evanescent (Fig. id).
This is also one of the smallest species of
Philia which has been recorded, and the shape
of the head (Fig. la) is unique for this genus.

MALE: Tiny, chiefly shining black species
with a faint, reddish tinge in the ground color
of the pleura, humeral ridges, scutellum, and
legs. Pile all black. Head: The upper portion
of each compound eye is brownish red and
is densely covered with short pile; the lower
part of each eye is black. The front margins
of the compound eyes are produced beyond
the bases of the antennae and completely ob-

Bibionidae of New Zealand — HARDY

= .I6 mm.

Fig. 6. P. segnis (Hutton), a. Apex of front tibia
(lateral aspect); b, front tibia, median portion; c, ninth
tergum of male.

scare the basal 3 or more segments in most
specimens (Fig. la). The face is not at all
produced, its margin is sunken below the eye
margins and is not visible from a lateral view.
The antennae are apparently about 9 seg-
mented. The flagellum is clavate in shape, and
the segments are so closely joined that they
cannot be accurately counted. The antennae
project downward (Fig. la) rather than up-
ward as is the usual condition in Philia. The
labella and palpi are largely hidden beneath
the margins of the compound eyes. Thorax:
With rather long black dorsocentral and mar-
ginal hairs. The scutellum has long hairs
around its hind margin. The halteres are black

519

except for a yellowish tinge on their bases.
Legs: With all segments slender. The front
tibiae have two sets of spines, composed of
two spines each, above the apical set. The
first two spines are close together and are
situated at about the basal one third of the
segment. The second set has the spines more
widely spaced and situated near the apical two
thirds of the segment (Fig. lb). The apical
spur is small, approximately equal in size to
the spines. The segments of the hind tarsi
are slender, not at all swollen. Wings: Very
faintly tinged, almost hyaline. The stigma is
light brown and is oblong in shape. The an-
terior veins are dark brown, the posteriors
are hyaline to faintly yellowish. The costa
extends about 0.4 the distance between veins
Rs and Mi. The m crossvein and the bases of
veins Mi-1-2 and Mi are completely lacking
(Fig. id). Except for the basal portion, the
anal vein is represented only by a slight fold
in the wing. The shape of cell R is very dif-
ferent from that of other Philia. The r-m
crossvein is more nearly longitudinal in posi-
tion, and there is no sharp angle at the point
of junction of the r-m and vein Mi-1-2. Veins
Mi-f-2 and Rs give the appearance of being a
single continuous vein (Fig. id). Abdomen:
About twice as long as the thorax and sparsely
pilose. Genitalia: The ninth tergum is about
twice as wide as long. The anterior margin is
straight, or nearly so, and the posterior mar-
gin is convex (Fig. le) . The sternum is about
as wide as long and has a small tumescence
in the middle of the hind margin. The con-
cavity on the hind margin extends about 0.25
the length of the segment. The claspers are
rather sharply pointed at their apices (Fig. if) .
length: Body, 2.63 mm.; wings, 2.26 mm.
female: Head: Of usual shape for genus,
the sclerotized portion in front of the eyes
is about 0.3 as long as the eyes, and the por-
tion behind the eyes is nearly as long as the
eyes. The antennae appear to be 10 or 11
segmented. The segments are very closely
joined and are difficult to discern as in the
male. The ocellar tubercle is not well devel-

520

PACIFIC SCIENCE, Vol. VII, October, 1953

j

\

\s

V

/r ■

*> /f[Y>

= .16 mm.

Fig. 7. P. tuthilll n. sp. a. Head of male (lateral aspect); b, front tibia of male; c, front tibia of female; d, wing;
e, ninth tergum of male;/, male genitalia (ventral aspect).

Oped and extends only the height of the ocelli
above the level of the vertex. The head is
entirely polished black and is sparsely brown
to black pilose. Thorax: Chiefly polished
black, tinged with red. The propleura and the
area of the dorsum between the combs are
rufous tinged with brown. The anterior comb
contains 12 blunt teeth, the posterior comb
10 sharp teeth. Legs: The front coxae and the
bases of all the femora are rufous. The re-
mainder of the legs are brown to black, tinged
with red. The spines on the front tibiae are
much stronger than in the males and 3 spines
are present in the middle set; 2 spines placed
one above the other on the outside dorsal

surface and 1 spine placed on the inner dorsal
surface (Fig. 7c). The extra spine on the fe-
males is smaller than the others and it possibly
may occur, at least as a rudiment, in the males.
I have been unable to detect it on the spec-
imens at hand but the smaller spines of the
males are sometimes difficult to differentiate;
in small species a tiny spine could easily be
obscured by the bristles of the legs. Wings:
Slightly more fumose than in the males. Ab-
domen: Brown, tinged with red. The abdomen
is sparsely covered with short black pile.

LENGTH: Body, 2.2-2.45 mm.; wings, 2.5-
2.65 mm.

HOLOTYPE male, allotype female, and twen-

Bibionidae of New Zealand — HARDY

521

ty paratypes (ten males and ten females) : Lake
Taupo, New Zealand, Feb. 20-22, 1950 (L.
D. Tuthill).

The type, allotype, and a series of paratypes
are being deposited in the Cawthron Institute,

Nelson, New Zealand (at Dr. Tuthill’s re-
quest). The paratypes are in the following
collections: U. S. National Museum, British
Museum (Natural History), Bishop Museum,
Honolulu, and the University of Hawaii.

NOTES

Hawaii as a Cloud Physics Laboratory

The major islands of the Hawaiian chain
lie across the Pacific northeast trade winds
near north latitude 21°, about 2,400 miles
southwest of San Francisco. It is the purpose
of this note to discuss very briefly the great
advantages of one of these islands (Hawaii)
as a place to study the relationships between
the precipitation elements in clear marine air
and these elements present in clouds formed
in this air.

Recent developments in the study of rain
formation have led to the hypothesis that
large sea-salt particles are the basic nuclei
upon which many raindrops initially form
(Ludlam, 1951, Quart. Jour. Roy. Met. Soc.
77: 402—417; Woodcock, 1952, Jour. Met.
9: 200—212; Squires and Woodcock, MS.).
These developments have re-emphasized the
need in cloud physics for further detailed
physical-chemical studies of rain which will
enable us to ". . . know how nature operates
to produce precipitation” (Byers, 1953, In-
dian Acad. Sci., Proc. 37(A): 237—247).

In order to make preliminary tests of the
salt particle-raindrop hypothesis by making
measurements of natural rains, it was neces-
sary to find a geographic location where the
contents of the lower atmosphere could be
determined before, during, and after the for-
mation of cloud and rain. Recent work has
indicated that the windward side of the island
of Hawaii, in the region of Hilo, is almost
ideal for this study (see Fig, 1). Here, on
most of the days throughout the year, rain
falls near area A from orographic clouds which
have formed over the island and within the

trade-wind stream. If one follows the stream-
lines of this trade-wind air upwind of area A
and over the sea, only scattered cumulus
clouds are found upon arrival near area B. If
the air is followed downstream from A, the
clouds are found to dissipate, leaving clear
air at c. Hence it is possible to determine, by
appropriate measurement, the contents of a
parcel of clear air at B, the nature of the
precipitation within clouds formed in this air
when it arrives at A, and the final modified
parcel after the clouds have dissipated at c,
a distance of about 30 miles.

Much of the mountain slope near Hilo is
readily accessible on paved roadways. Rain
and cloud drop sizes may be sampled within
the orographic clouds from the 2,000 foot
level to the 6,500 foot level on these road-
ways, thus avoiding the problems of evapora-
tion, accretion, etc., which can greatly modify
rains falling through clear air from cloud base
to ground.

The presence of the trade-wind inversion at
an average height of about 6,000 feet over the
sea concentrates the clouds and water vapor
in the lower air (Riehl et aL, 1951, Quart.
Jour. Roy. Met. Soc. 77: 598—626; U.S.
Weather Bureau, Staff Members, Honolulu,
MS.; Leopold, 1949, Jour, Met. 6: 312—320).
The inversion also limits the vertical extent
of the salt nuclei, as shown in Figure 2. The
distribution curves representing salt particles
at the 1,550 and 2,740 meter altitudes show
a great reduction in the number and weight
of particles at the base of the inversion and
above it (see temperature on insert diagram.

522

NOTES

523

Fig. 1. Streamlines and mean annual rainfall on NE part of Hawaii. Isohyetal lines are in inches. The arrows
represent the hypothetical mean streamlines of the [flow of the trade winds in this part of the island, as given
by Leopold (1949). (Scale in statute miles.)

Fig. 2). This concentration of the precipita-
tion elements (i.e., clouds, water vapor, and
nuclei) in the lower air is found during most
of the year in this area and is one of the
major advantages of Hawaii for precipitation
studies such as those discussed here. Measure-
ments of water vapor, temperature, salt nuclei,
etc., can therefore be limited to the lower
atmosphere, and there is usually no need to
be concerned with the higher levels.

A further result of the presence of the trade-
wind inversion is that the great majority of
the clouds never ascend to heights at which
the temperature is below freezing. Thus, most
of the time there is no confusion as to whether
or not the Bergeron rain-forming process may
have occurred. It seems clear that accretional
processes must account for practically all the
rains in this region, and hence it is an ex-
cellent area in which to attempt to relate the

524

PACIFIC SCIENCE, Vol. VII, October, 1953

2.4 5.1 1 1.0 23.7 61.1

PARTICLE RADIUS AT 99 % RH ( W )

Fig. 2, Vertical distribution of sea-salt particles over
the sea in Hawaii (21°30'N, 157°40'W), May 30, 1952,
1300 local time. Surface wind 80°, force 4 to 5. Height
of scattered cumulus cloud bases varied from 690 to
820 meters. Temperature at 2,740 meters 12.8°C. The
short transverse lines on the distribution curves mark
the first quartile, median, and third quartile weight
distribution points, reading left to right. The distribu-
tion curves are read as follows: at a height of 518
meters there are about 10,000 particles larger than 100
/x/i grams.

detailed physical changes in the air associated
with these frequently recurring processes.

The presence of the great mountain Mauna
Kea (see Fig. 1), extending up to almost
14,000 feet, makes it possible to obtain direct
visual estimates of maximum cloud heights
among the orographic clouds over the high-
rainfall region during extended time periods
which make air craft observation impractic-
able. These estimates can be made at the same

time that rain and cloud drops are being sam-
pled within the cloud along the roadway be-
low. One can drive a car up to a point at the
9,500 foot level, which is near a site over-
looking the high-rainfall region A.

In this area of Hawaii, nature is producing
rain very regularly and under simple condi-
tions usually uncomplicated vertically by mul-
tiple layering of air masses of different origin
and physical history. Geophysical events of
the past and present seem to have fortuitously
conspired to produce a setting in which sim-
ple natural experiments in rain-making are an
almost daily occurrence. Through the use of
local roadways and aircraft, the whole ex-
perimental area is readily accessible for study,
and the observations of one day can com-
monly be tested or extended on the following
day under the nearly constant natural condi-
tions. The author knows of no other geo-
graphical setting so uniquely advantageous
for cloud physics studies.

The importance of rainfall to the economy
of the Hawaiian Islands has led to extensive
activity by local organizations in the study of
various weather factors and in the accumula-
tion of data from numerous observing sta-
tions. These organizations— the U. S. Weather
Bureau, the Pineapple Research Institute, the
Experiment Station of the Hawaiian Sugar
Planters’ Association, and the Territorial Cat-
tlemen’s Council— located at Honolulu, Ha-
waii, are interested in fostering cloud physics
research and are able to supply valuable
climatological data from long-period records
as well as current radiosonde and other data
from numerous local stations. — A. H.
Woodcock^ Woods Hole Oceanographic Institu-
tion. (Contribution No. 660 of the Woods
Hole Oceanographic Institution.)

Index

Acacia confusa, 181, 271
Achyranthes aspera, 293
canescens, 168

Adult and Larval Stomatopod Crustaceans Occur-
ring in Hawaiian Waters, 399-437
Aedes adustus. All

alboscutellatus, 454, 457
andamanensis, All
aureostriatus, 454
brugi, 465
butler i, 476

campy lostylus, Al^, AlA, Al6
cancricomes. All
cinereus, All
curtipes, 478
dux. All

hamistylus. All, All)
imprimens, 465
johnsoni. All
kanarensis. All
kochi, 373
leucopleurus, 454
lineatopennis, 465, 468
lowisi, 459
luteolateralh, 465
macrodixoa. All, AlA, Al'b
margarsen. All, All
miachaetessus. All
mindoroensis, 459
neomacrodixoa. All
nigrotarsis. All
niveoscutellatus, AlA
nubicolus, All
omurensis. All
ostentatio, 463
pampangensis, AlA
panayensis, AlG
platylepidus, 480
punctifemore, 459
robertsi. All
sigmoides. All
umbrosus, AlG
uncus. All, AlA
vexans nocturnus, 460
wainwrighti, 464

Aedes Mosquitoes of the Philippine Islands III.
Subgenera Aedimorphus, Banksinella, Aedes, and
Cancraedes (Diptera, Culicidae), 453-481
Aedimorphus, 453
Ageratum cony zo ides, 301
Aglaia elliptici folia, 181, 182
formosana, 182
Agrostis exarata, 501
Aioretomyia ostentatio, 463
varietas, AlA
Alepisaurus, 244
Allophylus timorensis, 295, 304
Alocasia, 319

macrorrhiza,191, 301, 305
sp., 380

Amerigoniscus, 111
Anapagurus, 111
hyndmanni, 265

Anisocheleo?nyia, 315
nivipes, 315
Annona muricata, 293
Anopheles farauti. 111
lungae, 340
nataliae, 340
solomonis, 340
Ant, 302

Antennaria dioica, 498
Anuraea cruciformis, 156
eichwaldi, 157
serrulata. 111
Anuraeopsis sp., 155
Apateodus, 244
Apocreadium, 448
Arctozenus, 219, 235
Armigeres, 381
Artemisia unalaskensis, 499
Artocarpus, 111
altilis, 295, 304
incisus, 167

Asclepias curassavica, 295
Asplanchna priodonta, 154, 157, 158, 160
Astronia cumin giana, 184
ferruginea, 184
formosana, 184
pulchra, 184

Athyrium Filix-femina, 499

Bank, Theodore P., II:

Biological Succession in the Aleutians, 493-503
Banksinella, 453
lineatopennis, 469
Banner, Albert H.:

Crangonidae of Hawaii {see index, p. 147)
Barringtonia asiatica, 299, 303
racemosa, 299
Bathysudis, 219
Bauhinia monandra, 294
Belkin, John N.:

Mosquitoes of the Genus Uranotaenia in the Solo-
mon Islands, 312-391
Bibio elegans, 513
fulvipennis, 513
helioscops, 513
imitator, 513
nigrostigma, 517
ruficoxis, 513
substitute, 513
zealandicus. 111

Bibionidae of New Zealand (Diptera), 513-521
Biological Succession in the Aleutians, 493-503
Birgus, 265
Bironella, 340
hollandi, 340

Boerlagiodendron pectinatum, 181
Brachionus angularis, 11 A

angularis var. bidens, 154, 162
budapestinensis, 11 A, 162
calyciflorus, 11 A, 160, 162
calyciflorus N2X. dorcas, 154, 160
forficula, 11 A, 158, 162
quadridentatus, 154

[ 525 }

526

PACIFIC SCIENCE, Vol. VII, October, 1953

quadridentatus V2ii. hrevispinus, 154, 162
urceolaris, 154, 162
Brackenridgia, 175

Braconidae, index to genera and species, 215
Bruguiera conjugata, 300
Bryophyllum pinnatum, 293

Calamagrostis canadensis var. Langsdorffii, 499
Calophyllum inophyllum, 271, 294, 303
Campanula lasiocarpa, 498
Canavalia microcarpa, 300, 304
Cancer arenarius, 416
ciliatus, 412
Cancraedes, 453
Cantherines pardalis, 448
Capsicum frutescens, 296, 305
Carex circinata, 501
Lyngbyei, 499
spp., 499

Carica papaya, 294, 303
Cassia occidentalis, 294
Cassytha fliformis, 167, 293, 300, 303
Casuarina, 271
equisetifolia, 295
Caucasonethes, 175
borutzkii, 175
henroti, 175
rothi, 175

Cento theca lappacea, 302
Chloridella affinis, 404
Chlorophthalmus , 220
Choanodera, 448
Citrus aurantifolia, 304
erythrocarpa, 183
Cladophora membranacea, 300
Clerodendrum inerme, 305
Clibanarius, 255
misanthropus, 261
Cocos nucifera, 166, 298, 301, 305
Codiaeum variegatum, 300
Coenobita, 265
Collinsiella tuberculata, 292
Colocasia, 319
esculenta, 297, 305
sp., 380

Colurella adriatica, 155
lepta, 155
obtusa, 155

Conochiloides dossuarius, 153
Conochilus hippocrepis, 153, 157, 158
unicornis, 153
Cordia cumingiana, 184
kanehirai, 183

subcordata, 166, 296, 301, 305
Corethrella, 373
Coronida, 402, 403, 425, 426
multituber culata , 420
sinuosa, 399, 400, 404, 419, 434
sp?, 433, 434

Crangon (see index, p. 147)

Crangonidae, or Snapping Shrimp, of Hawaii,
3-147

Crataeva speciosa, 303
Crinum sp., 298, 305
Croton cumingia, 181
Culex, 187, 319, 320, 340
annulirostris, 358
archegus, 187
argentinotus, 457
auratus, 466

imprimens, 465
luteolateralis, 468
nocturnus, 460
nocturnus niger, 460
quitensis, 190
sitiens, 332
uncus, 474
Curcuma, 319
Cycas circinalis, 293
Cynoglossum alpestre, 185
lanceolatum, 185
Cyperus javanicus, 298, 305
sp., 305

Cyrtosperma chamissonis, 168, 297, 305

Danielsea pagei, 463
Dardanus asper, 255-266
Derris trifoliata, 304
Deschampsia beringensis, 499

Development of the Pedunculate Spermatophore of
a Hermit Crab Dardanus asper (De Haan),
255-266

Diaschizza gibba, 153
Dictyosphaeria favidosa, 292
Digitaria pruriens, 298, 302, 306
Dilophus insolitus, 516
spectabilis, 517
Diogenes, 255
pugilator, 261, 264, 265

Discussion of the Trematode Genus Schistorchis
(Family Lepocreadiidae) with Descriptions of
Two New Species from Hawaii, 447-452
Diurella brachyura, 154
porcellus, 154
sty I at a, 154
tigris, 154, 157, 160
weberi, 154
Dodonea viscosa, 181
Drimys, 244
defensor, 245
Dryopteris spp., 499
Dysoxylum cumingiana, 182

Ectropothecium monumentorum, 300, 303
Ehretia formosana, 185
navesii, 185
resinosa, 185
electrofishing, 482-492
Eleusine indica, 166, 298, 302, 306
Ellisiophyllum pinnatum, 180
Elymus arenarius subsp. mollis, 500
Enteromorpha proUfera, 303
Epilobium angustifolium var. macrophyllum, 499
Eragrostis amabilis, 298, 306
Eriophorum chamissonis, 499
russeolum, 495
Escualdoniscus, 175
Euchlanis alata, 155, 156
dilatata, 155
triquetra, 155, 156
Eupagurus, 255

bernhardus, 260, 261, 265
cuanensis, 261, 265
hyndmanni, 261
prideauxi, 261, 265
Euphorbia Atoto, 167
heterophylla, 294
Eurytemora, 504
affinis, 511
composita, 507

Index

527

grimmi, 511
lacustris, 507
raboti, 511
velox, 511
yukonensis, 504
Evodia confusa, 183
merrillii, 183
roxburghiana, 183

Faunal Speciation in New Georgia, Solomon Is-
lands, 250-251
Ficus carolinensis, 304
tinctoria, 295, 301, 304
Filinia longiseta, 155, 157
Fimbristylis atollensis, 298, 306
Finaloniscus, 175

Fleurya ruderalis, 166, 295, 301, 304
Floristic Interchanges between Formosa and the
Philij?pines, 179-186
Freycinetia sp., 373

Gastroschiza triacanthum, 156
Glassman, S. F.:

Plant Records from Caroline Islands and Study of
Native Plant Names, 291-311
Glycosmis cochinchinensis, 183
citrifolia, 183
erythrocarpa, 183
pentaphylla, i83
Gomphrena globosa, 293
Goniothalamus amuyon^ 181
Gonocaryum calleryanum, 184
diospyrosi folium, 184
larlacense, 184

Gonodactylus, 402, 403, 425, 426
guerini, 399, 404, 423
Guettarda speciosa, 167, 181, 295, 304

Flalimeda opuntia, 292
Hanson, Mary Louise:

The Trematode Genus Schistorchis with Descriptions
of Two Hawaiian Species, 447-452
Hardy, D. Elmo:

Bibionidae of New Zealand, 513-521
Harry, Robert R.:

The Bathypelagic Fishes of the Family Paralepididae.
1. Survey of the Genera, 219-249
Hawaii as a Cloud Physics Laboratory, 522-524
Hedyotis biflora, 296, 301
Heizmannia scintillans, 480
Hemiphragma heterophyllum, 180
Hernandia ovigera, 167
sonora, 293, 300, 303
Hibiscus tiliaceus, 166, 294, 300, 303
Hieracium triste, 501
Hippobro7na longiflora, 296
Hodgesia cairnsensis, 340
Holosteus, 244
esocinus, 245
Homalometron, 448
Honckenya peploides major, 500
Hull, William B., and Kenneth L. Knight:

Aedes Mosquitoes of the Philippines III, 453-481

Ichthyotringa, 243
Ilex arisanensis, 184
asprella, 184
llligera luzonensis, 181
Inocarpus fagiferus, 300
Intsia bijuga, 271, 299, 304

Ipomoea. gracilis, 297, 304
polymorpha, 181
tuba, 166

Ixora casei, 296, 301

J uncus spp., 499
Jussiaea suffruticosa, 294, 303

Kanehiro, Yoshinori, et al.:

The Role of Dehydration in Development of Laterite,
438-446

Katsuwonis pelamis, 422, 431
Keratella cochlearis, 154, 157, 158, 160
cochlearis x2lx:. macracantha, 154, 157, 160
cochlearis tecta, 154, 160
cruciformis eichwaldi, 155, 156, 160
quadrata, 155, 157, 158, 160
serrulata, 155, 157, 160
valga, 155, 158, 160
Knema glome rata, 183

Knight, Kenneth L., and William B. Hull:

Aedes Mosquitoes of the Philippines III, 453-481
Kuhlia sandvicensis, 486

Landa, Antonio:

Relationship between Body Length and Scale Length
in the Pacific Pilchard, 169-174
Laportea batanensis, 182
kotoensis, 182

laterite, development of, 438
Lathyrus maritimus, 500
Laurencia mariannensis, 303
Lecane brachydactyla, 155
ludwigii, 155
luna, 155

Lepadella acuminata, 155
borealis, 155
ovalis, 155, 157

Lepidotomyia alboscutellata, 457
Leptarrhena pyrolifoliq, 499
Lepturus repens, 298, 306
Lestichthys, 244
porteousi, 245
Lestidiops, 219, 239
sphyraenopsis, 239
Lestidium, 219, 233, 239
afline, 230, 240
atlanticum, 230, 240
atrox, 231, 240
bathyopteryx, 230
bellottii, 230, 240
danae, 230

intermedium, 230, 240
japonicum, 230, 240
jayakari, 230, 240
luetkeni, 230, 240
mirabile, 230, 240
neks, 227, 231, 240
nudum, 221, 230, 239, 240
paciflcum, 221 , 230, 240
parri, 230
philippinum, 230
pofl, 231, 240
prolixum, 231, 240
proximum, 230, 240
pseudosphyraenoides, 230, 240
r in gens, 230
simile, 230, 240
sphyraenoides, 230, 240
sphyraenopsis, 230, 240

528

PACIFIC SCIENCE, Vol. VII, October, 1953

thori, 230

thunnorum, 231, 240
vanderhilti, 231, 240
Leucaena glauca, 271
Leucophanes glauculum, 300
Lever, R. J. A. W.:

Faunal Spedation in New Georgia, 250-251
Levi-Castillo, Roberto:

The Subgenus Phalangomyia of the Genus Culex in
Ecuador, 187-192
Li, Hui-Lin:

Floristic Interchanges between Formosa and the
Philippines, 179-186
Litsea cubeba, 181
Lophoceraomyia, 340
Lucio sudis, 220
Luzula arcuata, 501
Lysiosquilla, 402, 403, 425, 426
maculata, 399, 404, 416, 432, 433
sp?, 431, 433

Mob a buxifolia, 181, 182
Macaranga dipterocarpifolia, 182
McCracken, Ralph J.:

Soils of Saipan, 267-277
Macroparalepis, 219, 233, 240
affine, 231, 240, 241
brevis, 231, 241
danae, 231, 241
egei, 231, 241
Magnisudis, 225, 233, 234
baryosoma, 225, 230, 235
M allot us catervarius, 230
Mansonia, 319

Mason, Leonard E., and Harold St.John:

Vernacular Names of the Plants of Bikini, 165-168
Matsusaka, Yoshito, et al.:

The Role of Dehydration in Development of Lat-
erite, 438-446
Matthews, Donald C.:

Development of the Spermatophore of Dardanus
asper, 255-266
Mertensia maritima, 500
Messerschmidia argentea, 167, 296, 305
Microdictyon okamurai, 292
Microlejeunea bullata, 292
Microsorium scolopendria, 293, 303
Mirabilis jalap a, 294
M is cant bus flo ridula?, 271
Mokil, 292

Monommata grandis, 153
Monostyla arcuata, 155
crenata, 155
hamata, 155
lunaris, 155
pyriformis, 155
Morgan, Morris E.:

Response of a Tropical Fish to Direct Current, 482-

492

Morinda citrifolia, 167, 181, 182, 296, 304
Mosquitoes of the Genus Uranotaenia in the Solo-
mon Islands (Diptera: Culicidae), 312-391
Murry a paniculata, 181
Musa paradisiaca, 297, 301
Myristica cagayanensis, 181, 182
glomerata, 182
heterophylla, 183
Mytilina trigona, 155
ventralis, 155
ventralis brevispina, 155

Neosudis, 220

Neothunnus macropterus, 422, 428, 431
Nepenthes, 319
Nephrolepis exaltata, 303
sp., 293

Nerium oleander, 295

New Alaskan Records of Burytemora (Crustacea,
Copepoda), 504-512

New Plant Records from the Eastern Caroline
Islands, with a Comparative Study of the Native
Plant Names, 291-311

New Terrestrial Isopod from Oregon, Caucasonethes
rothi n. sp., 175-178
Nicotiana tabacum, 305
Notholca acuminata, 155, 160
labis, 155, 160
Nothopanax fruticosum, 295
Notolepis, 219, 233, 237
coatsi, 226, 230, 237, 239
coruscans, 226, 230, 239
rissoi, 226, 230, 238
Notommata sp., 153
Notosudis, 220

Oahu Research Center of the University of Cali-
fornia [Note on establishment}, 392-394
Observations on the Subgenus Phalangomyia of the
Genus Culex in Ecuador with Description of a
New Species (Diptera: Culicidae), 187-192
Ochlerotatus imprimens, 466
ostentatio, 463

Ochrosia oppositifolia, 299, 304
Ocimum sanctum, 301
Odontodactylus, 402, 403, 425, 426
hanseni, 399, 404, 421
latirostris, 421
sp?, 434

Olearia virgata, 515
Omosudis, 221
Oplismenus compositus, 299
Oregoniscus nearcticus. 111
Oreocnide trinervis, 181

Pagurus, 255
arrosor, 265

Palmer, Clarence E.:

Note on Establishment of Oahu Research Center of
University of California, 392-394
P and anus, 271

sp., 166, 298, 301, 305, 319, 373
Paralepididae, list of genera and species, 230
Paralepis, 219, 233, 23_5
albyi, 245
borealis, 235

brevirostris, 221, 230, 237
brevis, 230, 237
bronsoni, 230, 237
coregonoides, 230, 235, 237
danae, 230, 237
elongata, 226, 230, 237
philippinus, 240
speciosa, 230, 235, 237
sphekodes, 245
Parascopelus, 244
Parrott, Arthur W.:

A Systematic Catalogue of Australian Braconidae,
193-218

Paspalum conjugatum, 306
vaginatum, 299, 302
Pedalia mira, 155

Index

529

Peltophorum sp., 300
Pemphis acidula, 166, 181, 303
Philia acutidens, 515
crinata, 514, 516
harrisoni, 514, 515
insolita, 514, 516
nigrostigma, 513, 517
segnis, 514, 516, 518
tuthilli, 513, 518 ^

varipes, 515

Phlehocalymna calleryanum, 184
Phyllanthus niruri, 294
Phy salts minima, 305
Picea sitchensis, 501
Pile a microphylla, 295, 301
Pingelap, 299
Pinguicula vulgaris, 499
Piper ponapense, 303
Pipturus argenteus, 295, 304
Pisonia sp., 294, 303
grandis, 166
Platyias patulus, 154
Pleorchis, 448
oligorchis, 448
Ploesoma hudsoni, 154
triacanthum, 154, 156
truncatum, 154
Plumeria rubra, 295
Poinciana pulcherrima, 295
Polyarthra euryptera, 154
trigla, 154, 157
trigla var. major, 154
trigla var. remata, 154
Polygonum viviparum, 498
Portulaca oleracea, 293
samoensis, 167, 293

Preliminary Report on the Soils of Saipan, Mariana
Islands, 267-277

Preliminary Studies on the Rotatorian Fauna of
Korea, 151-164
Premna integrifolia, 185
gaudichaudii, 297, 301, 305
nauseosa, 185
ohtusifolia, 185
odor at a, 185
vest it a, 185

Primula cuneifolia, 498
Prinos asprella, 184
Procris pedunculatus , 304
Profundisudis, 233, 238 ■

Protosquilla, 424
guerinii, 423
Protrichoniscus, 175
Prunella vulgaris var. aleutica, 495
Prymnothonoides, 237
regani, lyi
Prymnothonus, lyi
hookeri, 237

Pseuderichthus communis, 400
Pseudoficalbia, 315
pandani, 315

Pseudohowardina chrysoscuta, 463
linealis, 468

Pseudoskusea mediolineata, 475
nigrotarsis, 475

Pseudosquilla, 402, 403, 425, 426
ciliata, 399, 400, 403, 412, 428
monodactyla, 415
oculata, 399, 403, 415, 429
sty lifer a, 412

Pseudouranotaenia, 315
parangensis, 349
rowlandi, 315
Pteristripartita, 293, 300
Pterodina bident at a, 156
Puccinellia pumila, 500

Reedomyia, 453
alboscutella, 457
pampangensis, 453, 454

Relationship betw^een Body Length and Scale
Length in Five Year-Classes of the Pacific Pil-
chard or Sardine, Sardinops caerulea (Girard,
1854), 169-174 _

Response of a Tropical Fish to Direct Current and
Its Application to the Problems of Electrofishing
in Sea Water, 482-492
Rhinellus, 243

Role of Dehydration in the Development of Laterite,
438-446
Rotaria sp., 155

Saccharum offcinarum, 299

St.John, Harold, and Leonard E. Mason:

Vernacular Names of the Plants of Bikini, 165-168
Sardinops caerulea, 169
Saxifraga punctata insularis, 499
Scaevola frutescens, 167, 296, 304
Scaridium eudactylotum, 153
Schistorchis, 447, 448
callyodontis, 448
carneus, 447
sigani, 448
stenosoma, 447, 448
zancli, 447, 450
Schizostachyum diffusum, 181
Scirpus spp., 499
Scopeloides, 244
Scytonema figuratum, 292
hofmannii, 303

Seasonal Variations in the Physical Environment of
the Ponds at the Hawaii Marine Laboratory and
the Adjacent Waters of Kaneohe Bay, Oahu,
278-290

Semecarpus gigantifolia, 183
vernicifera, 183
Senecio pseudo- arnica, 500
Sherman, G. Donald, et al.:

The Role of Dehydration in Development of Lat-
erite, 438-446
Sibbaldia procumbens, 501
Sidafallax, 167, 294
Skusea diurna, 476
Sophora tomentosa, 167
Sphagnum capillaceum, 499
Girgensohnii, 499
Sphyraena, 224
Splachnobryum luzonense, 292
Squilla, 402, 403, 425, 426
affinis, 404
alba, 399, 403, 408
arenaria, 4l6
boops, 403, 406
calumnia, 400, 403, 410
leptosquilla, 399
maculata, 4l6
monodactyla, 415
nepa, 404
oculata, 415
oratoria, 399, 403, 404

530

PACIFIC SCIENCE, Vol. VII, October, 1953

quadraticauda, 406
quadrispinosa, 412
sp?, 426, 428
sty lifer a, 412
Stegomyia, 453
hatiensis, Al6
punctifemore, 459
Stemonosudis, 229, 231, 233, 241
elegans, 231, 242
elongata, 231, 241, 242
gracile, 231, 241, 242
intermedia, 231, 241, 242
macnira, 231, 242
miscella, 231
Sterculia glomerata, 183
Streptopus ample xifolius, 499

Studies on the Bathypelagic Fishes of the Family
Paralepididae. 1. A Survey of the Genera,

219-249

Sty rax kotoensis, 182
Sudis, 219, 232, 242
hyalina, 221, 222, 231, 242
squamosa, 230
Suriana maritima, 167
Symproptopterus, 235
Synchaeta longipes, 153
ohlonga, 153, 157
tremula, 153
spp., 153, 157

Systematic Catalogue of Australian Braconidae,
193—218 [Index to genera and species, 215}

Tacca leontopetaloides, 166, 298, 305
Taeniorhynchus lineatopennis, 468
Terminalia catappa, 303
litoralis, 166, 294, 303
Testudinella bidentata, 155, 156
patina, 155, 158
Thalassia hemprichii, 297
Thaleichthys pacificus, 230
Thespesia populnea, 294, 304
Thuarea involuta, 299, 306
Toluifera cochinchinensis, 183
Tonna perdix, 255
Townsley, Sidney Joseph:

Adult and Larval Hawaiian Stomatopods, 399-437
Trichocerca capucina, 154
cristata, 154
cylindrica, 154
elongata, 154
longiseta, 154, 157
pusilla, 154
sty I at a, 154

Trichoniscus nearcticus. 111
Trichotria tetractis, 155
truncata, 155
Tripteroides, 312

{Rachisoura) mathesoni, 380
(R.) stonei, 380

Triumfetta procumbens ,166, 294, 303
Trossulus, 244
exoletus, 245

Tseu, Winifred Shui Lin:

Seasonal Variations in Physical Environment of
Ponds at Hawaii Marine Laboratory and Adja-
cent Waters, 278-290
Turpinia lucida, 182

Uranotaenia, 312-391
albescens, 339, 350, 357, 358

alboannulata, Yll
albofasciata, 365
annandalei, 331
antennalis, 339
arguellesi, 358
argyrotarsis, 341, 349
atra, 317, 322-324, 340, 366
barnesi, 317, 322-324, 332
bimaculata, 320
cairnsensis, 324, 331
campestris, 358
cancer, 324, 331
ceylonica, 324, 331

civinskii, 317, 322-324, 340, 341, 349, 358, 366
clarae, 340

coeruleocephala lateralis, 324, 331

colocasiae, 380

delae, 340

diagonals, 380

faurati, 358

fimbriata, 339

hebes, 366

ludlowae, 340

macfarlanei, 358

nataliae, 315

neotibialis, 339

nigerrima, 317, 373, 380

nivea, 365

nivipes, 359, 365

nivipleura, 365, 372

obscura, 380

orientals, 365

painei, 380

pallidocephala, 340, 349
papua, 380
propria, 324, 331
pulcherima, 314
punctulatus, 358

quadrimaculata, 316, 317, '319-324, 357, 373
reyi, 340
rutherfordi, Yll
setosa, 339

sexaueri, 317, 321-324, 340, 358, 359, 366

solomonis, 317, 322-324, 340, 350, 358, 365, 366

tibialis, 332

tibioclada, 339

triangulata, 365

triline at a, 1)12

unguiculata, 319, 366

wysockii, 316, 319-323, 366 .

Vandel, a.:

A New Terrestrial Isopod from Oregon, 175-178
Vernacular Names of the Plants of Bikini, Marshall
Islands. Pacific Plant Studies 12, 165-168
Vernonia cinerea, 296, 301, 304
Verralina butleri, Al6
Vigna marina, 295, 300, 304
Viola Langsdorffii, 499

Wedelia biflora, 296, 304
Wilson, Mildred Stratton:

New Alaskan Records of Eurytemora, 504-512
Woodcock, A. H.:

Hawaii as a Cloud Physics Laboratory, 522-524
Yamamoto, Kokichi:

Preliminary Studies on the Rotatorian Fauna of
Korea, 151-164

Zanclus cor nut us, 450
Zephyranthes rosea, 298

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