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SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM
PROCEEDINGS
OF THE
UNITED STATES NATIONAL MUSEUM
VOL ME 2.102
NUMBERS 3291-3310
UNITED STATES
GOVERNMENT PRINTING OFFICE
WASHINGTON : 1956
ADVERTISEMENT
The scientific publications of the National Museum include two
series, known, respectively, as Proceedings and Bulletin.
The Proceedings, begun in 1878, are intended primarily as a medium
for the publication of original papers, based on the collections of the
National Museum, that set forth newly acquired facts in biology,
anthropology, and geology, with descriptions of new forms and revi-
sions of limited groups. Copies of each paper, in pamphlet form, are
distributed as published to libraries and scientific organizations and to
specialists and others interested in the different subjects.
The dates at which these separate papers are published are recorded
in the tables of contents of each of the volumes.
The present volume is the hundred and second of this series.
The Bulletin, the first of which was issued in 1875, consists of a
series of separate publications comprising monographs of large zoo-
logical groups and other general systematic treatises (occasionally
in several volumes), faunal works, reports of expeditions, catalogs of
type specimens, special collections, and other material of similar
nature. The majority of the volumes are octavo in size, but a quarto
size has been adopted when large plates were regarded as indispensa-
ble. In the Bulletin series appear volumes under the heading Contri-
butions from the United States National Herbarium, in octavo form,
published by the National Museum since 1902, which contain papers
relating to the botanical collections of the Museum.
Remincton KELioaa,
Director, U. S. National Museum.
II
ZANTHSON EN.
APR16 1955
LIBRARY
CONTENTS
Pages
Axspott, R. Tucker. A study of an intermediate snail host
(Thiara granifera) of the oriental lung fluke (Paragonimus).
Figures 32-45, and plates 8, 9. Published February 26,
1952 Byes. WS DSRisf RO Aiello LAS 71-116
Cartwricut, O. L. Aphotaenius, a new genus of dung
beetle (Coleoptera: Scarabaeidae). Published March 7,
1952 “Sb PAGO BLA Ua SET LT) Ws ft. 3h 8, MOTH LSI=184
New genus: Aphotaenius.
New species: Aphotaenius colombiensis.
CHAPMAN, WILBERT M., and Scuutrz, Leonarp P. Review
of fishes of the blennioid genus EHcsenius, with descriptions
of five new species. Figures 90-96. Published April 24,
BOSD ES 6 veins es ee wee ew ine PINAL, I OURO, .th07=528
New species: Ecsenius stigmatura Fowler, E. lividanalis, E. pro-
oculis, E. opsifrontalis, E. hawatiensis.
Ciark, Austin H. Echinoderms from the Marshall Islands.
Publisitea ware 27ehOg2-0 7) ~e ek e tee ee 2G0-o05
New species: Ophiodera punctata, Ophiomastix sexradiata.
Schizostella, a new genus of brittle-star (Gor-
gonocephalidea). Plate 40. Published March 19, 1952 . 451-454
New genus: Schizostella.
New species: Schizostella bifurcata, S. bayert.
Fietp, Wint1am D. Moths of the genera Mulona Walker
and Lomuna, a new and closely related genus (Arctiidae:
Lithosimae). Plates 10, 11. Published March 19, 1952 . 221-230
New genus: Lomuna.
New species: Mulona schausi, M. manni, M. barnesi.
Moths of the genus Epeiromulona, a new genus
of Lepidoptera. Plates 41-46. Published April 25, 1952 . 455-469
New genus: Epeiromulona.
New species: Epeiromulona lephina, E. biloba, E. hamata, E.
roseata, E. thysanata, E. icterinus.
New subspecies: Epeiromulona hamata hamata, E. h. venezuelensis,
E. h. colombiensis, E. h. brasiliensis.
Boos
1V PROCEEDINGS OF THE NATIONAL MUSEUM
Fisoer, WALTER Kenrick. The sipunculid worms of Cali-
fornia and Baja California. Figure 87 and plates 18-39.
Published July 8, 1952 :
New Genus: Siphonides.
New species: Siphonides rickettsi, Golfingia laetmophila, G. elachea,
G. macginitiei, G. pugettensis, Dendrostomum dyscritum, D.
lissum, D. schmitti.
New subspecies: Golfingia margaritacea californiensis, G. eremita
californica.
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SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 Washington: 1952 No. 3291
CONTRIBUTIONS TO THE MORPHOLOGY AND TAXONOMY
OF THE BRANCHIOPODA NOTOSTRACA, WITH SPECIAL
REFERENCE TO THE NORTH AMERICAN SPECIES
By Foitxr LInvER
INTRODUCTION
Since the days when Packard’s “Monograph of the Phyllopod
Crustacea of North America’ was published (1883), the accessible
material of notostracans has grown considerably, and the principles
commonly used for the taxonomy have been discussed in several
papers. Analyses of these characters, especially by Braem (1893),
Gurney (1923, 1924), Sidorov (1927), Barnard (1929), and Gauthier
(1934), have established a considerable variation in most of the char-
acters of European, Asiatic, and African forms. This variation, of
course, does not make a survey of the Notostraca of North America
an easy task, and when I venture such a survey here, I wish to note
that in some respects it is tentative only. It is based mainly upon a
study of the collections of the U.S. National Museum, amounting te
109 lots and considerably more than 2,000 specimens. ‘This study was
performed during a 3 months’ visit to the Museum in 1946, and was
made possible by a grant from the Smithsonian Institution, to which
my thanks are due for this generous support. I am also greatly
indebted to Dr. Waldo L. Schmitt and other officials of the Museum
for having most kindly facilitated my work. Also, Prof. J. E.
Lynch, University of Washington, Seattle, kindly allowed me to
examine some important material from the northwestern United
States. For comparison I had specimens from other parts of the
world, though this material, about 3,000 specimens from 71 localities,
has not been as comprehensive as could be desired.
1
APR6 1956
2, PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 102
Gurney (1924, p. 559) says about the taxonomy of the Notostraca,
“The separation of species within the genera Apus and Lepidurus is a
matter of quite unusual difficulty, since there is much variation, and,
as a rule, the only characters on which separation can be based are
smal! and ill-defined.”” Barnard (1929, p. 230) agrees with him, and
so do I.
As for other phyllopods, the females of the Anostraca present a
similar difficulty, as, sometimes, if they occur without males, it is
not even possible to say with certainty to what family they belong.
But the males of this order give excellent specific characters in the
genital organs and the second antennae—organs which display a
striking uniformity in the Notostraca. In the Conchostraca, the
structure of the head and the carapace provides rather good spe-
cific characters, but in these respects, too, the Notostraca are fairly
consistent.
The characters actually employed—helpfully reviewed by Barnard
(1929, pp. 230-234)—for the most part represent differences of degree
only. A striking example of this is Packard’s (1883, p. 319) key to
the American species of Apus; a study of the type specimens of the
species involved immediately reveals that the number of spines on
the supra-anal plate, which seems to be the fixed point in this key,
is not constant but overlaps the boundaries between his species.
According to my experience, one seldom finds a specimen which is
in complete accordance with the description of any of these species,
and similar difficulties are met even when one goes to the type speci-
mens. These are not numerous enough to cover the range of indi-
vidual variation, nor are their descriptions extensive enough. Yet
such variation is especially important when the characters involve
differences of degree. .
Of course, one can often compare material from neighboring locali-
ties in order to learn something of this variation, or, better still, one
can try to get abundant material from the original locality. The
first course is generally the only one practicable in museum work, and
I have tried it here. It is not very satisfactory, though some results
seem to be obtainable in this way.
Each sample of Notostraca ought to contain about 50 or more spec-
imens in order to cover most of the variation. And, in my opinion,
a new species should never be described from less than 100 specimens
from the same district, preferably taken at varying times of the
season. Paratypes are, In many respects, just as important as
holotypes.
A special question arises when we try to interpret the variation.
Ghighi (1924) suggests that the variation may sometimes be due to
the presence of two species in the same pool. I think that this occurs
NORTH AMERICAN NOTOSTRACA—LINDER 3
only rarely, as in the Anostraca and in the Conchostraca, and if it
should happen to occur in a particular sample, the fact would prob-
ably be readily revealed when all characters are considered. I have
not found any indications of mixed populations in this sense in the
samples I have examined, nor can I find any definite reference to
it in the literature, with the exception that Lundblad (1920) noted
that species of different genera sometimes have been found in the
same pool. In these cases no intermediate forms are reported.
So far, I have considered only specific characters. Sidorov (1927)
and Gauthier (1934) are convinced that there are geographical sub-
species within Apus cancriformis of the Old World. I do not deny
the possibility that there may be such taxonomic units in some
North American species, most likely among the forms of Apus.
Among the anostracans, which have similar habits, I have found
some indications of geographical subspeciation in forms of the
genera Branchipus and Tanymastiz of the Old World. Smirnov (1931)
seems to be of similar opinion with regard to some conchostracans.
However, I do not find it possible to separate geographical units
out of the material to which I have had access. In my opinion the
appropriate way of doing this would be to investigate thoroughly
some limited areas, and to follow this with a comparison of samples
from interspersed districts. ‘The possible existence of clines (Huxley,
1942, and Margalef, 1948) might also be traced in this way.
During my work it appeared that a species from Asia and Europe,
Lepidurus macrurus, is a synonym of the American species L. couesit.
Further, I found that South American forms of Apus are the same
species as the North American Apus longicaudatus. This may be
significant. It is worth noting that no one since the days of Packard
(1883) and Simon (1886) has tried to compare species from all over
the world, and, with our present knowledge of the great latitude in
variation of many characters, it does not seem at all improbable
that additional species from different continents will be found
synonymous.
The present tendency to reduce the number of species in the
Notostraca (see Gurney, 1924, and Barnard, 1929) is, I think, sound,
and is well supported by our present knowledge of variation. This
paper follows the same line.
In the following pages I have tried to analyze, with the help of
material from all continents, the commonly used taxonomic charac-
ters and, also, a character which has been very much neglected, the
number of body-rings of different kinds. This character has proved
quite useful. One of the results is a new grouping of the species of
Lepidurus into what seems to be two natural groups; another, that
lower taxonomic units begin to be revealed among parthenogenetic
4 PROCEEDINGS OF THE NATIONAL. MUSEUM VOL. 102
populations, although the real status of these cannot be determined
from the accessible material. In connection with this analysis I
have also considered polypody, and the peculiar phenomenon I have
called spiral growth, both of which are problems of interest from a
more strictly morphological point of view. Finally, I have applied
to the North American material the results of this analysis of taxo-
nomic characters.
TERMINOLOGY AND MEASUREMENTS
Instead of the term ‘‘segment,”’ in referring to the postmaxillary part
of the animal, I prefer the term ‘“‘body-ring.”’ The first 11 body-rings
I call the thorax. This is followed by the abdomen, which is com-
prised of leg-bearing body-rings, legless body-rings, and the telson,
which is not a true segment, and which should not be included in the
number of body-rings. The series of legs does not always stop under-
neath a boundary between two body-rings; it may stop at any place
quite independent of these boundaries (fig. 20). I have found it
advisable to count half leg-bearing body-rings in those cases where
the series stops approximately underneath the center of a ring, and a
whole leg-bearing ring when the series covers almost the whole ring,
disregarding those cases where the series only slightly surpasses a
boundary. Incomplete rings (fig. 16) are often found at the anterior
boundary of the telson, but very rarely within the series of rings.
It is useful to have a short formula when describing the numbers
of the various kinds of body-rings in a specimen or in a species. In
this paper, I have used such an arrangement. Thus, the formula
11-+12+5—28 body-rings indicates that there are 11 thoracic, 12
abdominal leg-bearing, and 5 abdominal legless rings, forming a total
number of 28 rings (telson not included). For a species the formula
is more complicated because there is a considerable variation of
these numbers within the limits of a species (Linder, 1947). If the
abdominal leg-bearing rings number from 16 to 18.5, the legless rings
from 5 to 10, and the total from 35 to 38 rings, the formula would be,
11+ (16-18.5) + (5-10) =35-38 body-rings.. Hf there is an incomplete
ring, the abbreviation “i” is inserted, as in this example: 11--12+
5+i=28-+i body-rings. It is also possible to indicate the presence
of the interesting abnormality, spiral growth, and its place within
the series. If a spiral of 3 rounds appears after the 25th body-ring,
immediately followed by the telson, in a specimen with 12 abdominal
leg-bearing rings, the formula would run as follows: 11+12+
2+sp.3r.=25-+sp.3r.
The telson bears a rather neglected structure, viz, a pair of dorsal
sensory setae (fig. 28), surrounded by short spines, and a pair of caudal
filaments. Between the latter there is, in the genus Lepidurus, the
NORTH AMERICAN NOTOSTRACA—LINDER 5
supra-anal plate. I measure the length of this plate from the most
anterior point of the basis of the caudal filaments to its apex.
As Barnard (1929) and S@mme (1934) have pointed out, the total
length of an animal cannot be measured with proper accuracy in pre-
served material. The reason is simple enough. Anteriorly and pos-
teriorly, the integument of a ring is much softer than in the central
part. This soft integument may form a fold directed inward and for-
ward, and in such eases the rings appear short and the total length
of the animal is relatively short, too. But other specimens from the
same lot, which have a carapace of similar length, may show no deep
fold in these places, the soft integument being more or less stretched
out between the more highly chitinized parts of the rings (fig. 1). In
TA
a
ham ee Nac
b
Ficure 1.—Schematic diagram of the body-rings of a notostracan showing how the length
is influenced by varying contraction. Highly chitinized parts of the integument indi-
cated by heavy lines. a, Contracted; b, extended.
the latter case, of course, the rings appear longer and the total length
of the animal may be considerably greater than in the former cases.
Tt is quite usual to meet with both extremes and a more or less con-
tinuous series of intermediate cases in a lot containing a large number
of specimens, as they may have been fixed in varying stages of con-
traction. Rosenberg (1947) noticed a considerable shrinking of the
specimens when they were placed in preservation fluid. It is also
obvious that preservation, after an indeterminable time, weakens
the soft tissue in and between the body-rings and thus causes a length-
ening of the body to a variable degree. In very old material macera-
tion (and lengthening) may have gone rather far. It is, however,
desirable to give an idea of the length even if it cannot be accurately
stated. I measure the length on the midline from the apex of the
head to the base of the caudal filaments. Thus, in Lepidurus, the
length of the supra-anal plate is not included in the figure of the total
length. This may seem a little odd, but in this way comparable
figures for Lepidurus and Apus are obtained. The length of the supra-
anal plate in the former genus may be up to 44 percent of the length
of the carapace, and thus it would be quite misleading to include this
plate, which has no counterpart within the genus Apus.
6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Barnard (1929) also points out that we cannot give an accurate
count of the body-rings exposed behind the carapace. In order to
give some, even if not an accurate, idea of this conspicuous feature
I have mentioned the number in some specimens. But I give this
number, as well as the figures of the total length, only with a stated
reservation as to its accuracy.
As Sémme (1934) has shown for Lepidurus arcticus, the carapace
is little affected by the preservation fluid and thus can give an idea
of the size of the animal. This is only natural, because it is contin-
uously and highly chitinized. Foilowing the general custom, I
measure its length along the mid-dorsal line. Of course, we cannot
be sure that variations in this figure closely follow the variations of
the real total length of the body; the relation may vary according to
varying ages of the specimens, to say nothing of the varying contrac-
tion. To obtain the greatest width of the carapace, the usual method
seems to have been to measure it in a straight line. This is not
reasonably accurate, because the height of the carapace may vary,
sometimes to a considerable degree. It is better to measure the
greatest width from the carina to both sides (fig. 2). In cases of
Ficure 2.—Cross section of the carapace of a notostracan, showing how the width of the
carapace is measured.
distortion, I have flattened the carapace to a shape similar to normal
appearance before taking the measurement.
I count 5 endites on each leg (fig. 26). The coxal lobe is the first
endite; it is essentially similar to the other endites in the arrangement
of the setae and has on its margin similar small elevations covered
with small needlelike structures. What Packard (1883) calls the
sixth endite is quite dissimilar to the more basal, real endites; it has
only a single row of setae and spines along its margin, and no trace
of the small elevations mentioned above; I call it the endopodite,
as many other authors do. Packard’s ‘“flabellum” is the exopodite,
and his “gill” is called epipodite in this paper. Further, we some-
times find what may be the vestige of a preepipodite (fig. 22) in the
form of a small elevation with a few setae (cf. Eriksson, 1934, p. 236).
Specimens from the following museums were examined:
Muséum National d’Histoire Naturelle, Paris (Paris Mus.).
Naturhistoriska Riksmuseet, Stockholm (Stockholm Mus.).
Zoologiska Museet, Kungl. Universitetet, Uppsala (Uppsala Mus.).
United States National Museum, Washington (U.S.N.M.).
NORTH AMERICAN NOTOSTRACA—-LINDER 7
BODY-RINGS AND POLYPODY
Following the usual custom, I consider the body proper to begin with
the first postmaxillary ring, which is, however, strictly separated
from the head only on the ventral side. Packard (1883) counts the
first ten body-rings as the thorax, and the following ones as the ab-
domen. I include the eleventh body-ring in the thorax, as does Simon
(1886) in his important revision of the phyllopods and as Sars does in
his many descriptions of various notostracans. More than a hundred
years ago Zaddach (1841) showed, in an admirable work now almost
neglected, that the first 11 body-rings are essentially similar to each
other in the equipment of muscles, while the following ones differ
quite considerably from them. Further, the legs of this portion are
different, though the boundary in this respect is usually not distinct.
A boundary between different regions in this location is recognized
in crustaceans of many orders—so many, in fact, that it is not likely
to be a mere coincidence (Linder, 1941, p. 113).
I have abandoned the use of the term ‘‘segment,’’ replacing it with
the term “body-ring,” when real rings or parts of rings are concerned.
I do this for the reason that in the abdomen there are no complete,
ordinary segments but two series of parts of segments, in some respects
independent of each other—the series of rings and the series of legs.
These are united to form a composite structure unique in appearance.
This is the true meaning of the much discussed phenomenon, poly-
pody. Itis not an absolutely new view. Lankester (1904) expressed
a similar view, but did not give evidence for his opinion. However,
Zaddach (1841) had already supplied much evidence supporting that
opinion, and the following facts, most of which have not been noticed
by earlier authors, may give additional evidence:
1. The series of legs covers a varying number of body-rings in
specimens from one locality. Zaddach (1841) has given an example
of this. He also regards the number of pairs of legs as fixed; however,
I have found that this number is subject to a considerable variation.
The series of legs may stop at any place under a body-ring (fig. 20).
If we choose a certain pair of legs in the caudal part of the series
of legs, and determine its place under the series of body-rings, we shall
find that it has varying positions in various specimens from the same
locality (see table 2, p. 42). Thus, each body-ring in the abdomen
does not have a fixed number of pairs of legs, and this means that
there is no evident correlation, as is supposed by most authors, between
the series of legs and the series of abdominal body-rings. Evidence
for this fact from extra-American material is given in an earlier paper
(Linder, 1947).
2. The boundaries between the body-rings in the leg-bearing part
of the abdomen do not continue ventrally. They stop at a longitudi-
8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
nal boundary, always well marked and often formed as a ridge in the
integument (fig. 20), which delimits the leg-bearing area. This area
may be divided into “‘segments” coinciding with the legs but totally
independent of the boundaries between the body-rings. Zaddach
(1841) has already described the ventral “segmentation” and the
corresponding division of the longitudinal muscles in this area in
Apus cancriformis.
3. In two cases of abnormal growth of the body-rings that I have
found, the series of legs is not affected by the abnormal turns, or spiral
growth, of the related body-rings (fig. 17).
The first two points hold good for each of the species of Notostraca
that I have examined, though some of the details may be difficult to
observe in specimens which have just molted or are badly preserved.
It is a well-known fact that the rate of production of new
body-rings (in early postembryonic stages) is quite different from
that of the legs, with their muscles and nerve-cord ganglions. From
the caudal end of the series of legs there are produced, at an
almost frantic rate and in considerable number, the elements of the
leg-bearing area, but the body-rings, budding from the anterior margin
of the telson, increase in number relatively slowly. The body-rings
do not vary much in size, but the elements of the leg-bearing area
grow smaller and smaller caudally, as if the animal meets the frantic
activity of this part with something like starvation of its individual
elements. We might assume a kind of organizer at the end of the
series of legs (for what else could we assume?) that is, in some way
or other, less favored than the ordinary organizers at the anterior
boundary of the telson.
I have noticed in many specimens that this disproportionate
development has continued during the further growth of the animal.
Thus, we often notice that in a larger animal the series of legs covers
a smaller number of body-rings than is the case in a smaller animal
in the same lot (see table 2), indicating that the size of the body-rings,
but not necessarily their number, increases faster than does the series
of legs. Other facts also point to this conclusion: an oblique striation
occurring at the area where the body-rings and the series of legs meet,
explained as the mark of a tension between the two parts as they grow
at dissimilar rates; and the moving forward of the fiftieth pair of legs,
as observed in table 2 (we note that this moving forward does not
exactly follow the changes in size, but individual variation in these
complicated processes is to be expected).
NUMBER OF BODY-RINGS
It seems to be the rule that neither the total number of rings nor
the number of abdominal leg-bearing rings is given in descriptions
NORTH AMERICAN NOTOSTRACA—LINDER 9
of North American Notostraca. There are few exceptions to this rule.
Packard (1883) mentions the figures for one specimen of Apus lucasa-
nus and Rosenberg (1947) quotes fixed figures characteristic of his
species A. oryzaphagus and A. biggsi. The number of legless rings,
on the other hand, is almost always recorded, and a slight variation
in this respect is often noticed. Interesting information about
Lepidurus arcticus is given, though only with some hesitation, by
Chamberlin and Duncan (1924, p. 99): ‘““The number of segments
visible on the dorsal surface from the point of attachment of the
carapace to the body seems to be either 26 or 27... . Behind the
appendage bearing segments are five others, exclusive of the telson,
which are without appendages of any sort.’’ As the above-mentioned
point of attachment stretches over the first body-ring, we may con-
clude that these authors found the number of body-rings to be
11+ (11-12) +5=27-28 (telson not counted).
Even in descriptions of species from other parts of the world we
seldom get information about the number of leg-bearing abdominal
rings; at times a fixed total number is given, together with the number
of legless rings, but without any attention to a possible variation of
the former figure. Zaddach (1841) provides an exception; he says
that in A. cancriformis the number of leg-bearing abdominal rings
(in “pars abdominis posterior,’ according to Zaddach) is 17 or 18,
the number of legless rings 5 or 6, and the total number of body rings
always 34.
All authors except Chamberlin and Duncan seem to take it for
granted that the species of Notostraca are nomomeristic, but this is not
true of any of the species examined by me.
From figures 14 and 31 it can be seen that a variation in number is
found not only in the legless rings, but in the leg-bearing ones, too.
These varying numbers, together with the 11 thoracic rings, are
combined in different ways to give a total number varying within the
limits of a species. This is true of every species of which I have seen
a reasonable number of specimens.
When I now proceed to analyze what I have found concerning
the variation of these figures—a preliminary analysis in some respects
because the material is rather scanty—I shall begin, not in the tradi-
tional way, with a treatment of the legless rings, but with the total
number of rings and the number of abdominal leg-bearing rings.
These are the primary characters. The number of legless rings is not
a simple character; it is the result of the varying interplay of the two
other series.
As for the total number of body-rings, the range of variation may be
quite considerable in specimens from the same locality. In a new
species of Lepidurus from Grand Coulee, Wash. (see p. 39), the males
10 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 102
have 30-33+1 and the females 30+i-34 rings (fig. 14). The range is
also great in A. longicaudatus from Wyoming (U.S.N.M. No. 58766),
where the males have 41-44 and the females 39-++i-43 rings (fig. 31).
These are relatively extensive samples, the former containing 54 males
and 42 females, the latter 33 males and 98 females. In smaller
samples the range of variation is not so great. The extremely high and
low numbers of rings in one sample are usually represented by rather
few specimens.
No doubt the average number differs in various samples of the
same species. But we cannot reasonably compare samples with 1, 2,
or 10 specimens with samples containing a hundred or more specimens.
Small samples give very uncertain figures, and museum work, in this
respect, is much hampered by the fact that the great majority of
samples contain a very small number of specimens. Statistical
calculations, however desirable, are thus rendered very difficult.
There seems to be a general rule that species with a large number of
rings have a greater range of variation than species with few rings,
both in Lepidurus and in Apus. There are some exceptions to this
rule, but not many, in the material I have seen. The fewest rings in
Apus occur in forms that have a clear parthenogenetic tendency and
that show a very small range of variation. About these more will be
said later on (see p. 12). Similar forms belonging to a group with
relatively few body-rings are found among species of Lepidurus.
Body-rings in the species of Lepidurus range from 25 to 34, those of
Apus from 30 to 44. In the former genus, the high numbers of 30-34
are represented by two species only, L. bilobatus and a new species
described in this paper, both of which differ from the other species of
the genus in the number of leg-bearing abdominal rings, and, most
evidently, in the number of legs. Apart from this grouping, and from
what has been mentioned about parthenogenetic forms, I cannot find
clear correlations between the number of rings and other characters.
I doubt the validity of Rosenberg’s (1947) method of distinguishing taxo-
nomic units—and not lesser units at that, but species—by associating
a certain number of rings with a certain size of specimen, because
I think that further research is necessary to establish the conditions
of the variation in size, which in some lots is considerable, in others
rather small.
It is usual for the males to have a higher total number of rings than
the females of the same species, though the rule has many exceptions.
It is, of course, necessary to know whether the number of rings is
really fixed in adult specimens—non-adult stages are not considered
here—or whether it increases with an increase in size of the specimen.
I have attacked this problem, which has never before been investi-
gated, in two ways: first by comparing the number of rings in small
NORTH AMERICAN NOTOSTRACA—LINDER ll
and large specimens in the same sample, and second, by comparing
the old and the new integument of specimens in molting. All my ob-
servations agree that there is no general rule that larger specimens have
a higher total number of rings than smaller ones. On the contrary, I
have not infrequently found relatively small specimens with a high
number (see fig. 14). Also, I have thoroughly examined about a
hundred molting specimens of various species, and in none of them
have I found a new ring appearing from the telson in the underlying,
new integument. Judging from this, I think I am entitled to assume
a fixed total number of rings in an adult specimen.
As for the variation in the number of leg-bearing rings, I have
found no clear rule when comparing larger and smaller specimens
from the same lot. When examining molting specimens, however, I
have found two cases where the number of pairs of legs is mcereased
by one in the underlying, new integument. Thus we must reckon
with the possibility of an increase in the number of legs, and, conse-
quently, even a slight increase in the number of abdominal leg-bearing
rings, with an increase in the size of the animal. However, I think
we can be sure that this increase, if present, will not be great.
The usual range in the number of leg-bearing abdominal rings in
the genus Lepidurus is 9.5-13. This applies to all extra-American
species I have examined (L. apus, L. kirkii, L. viridis), and to most
American species (L. arcticus, L. packardi, L. couesii). This condi-
tion, together with the similarly small number of legless rings, at least
in some cases helps to explain the short abdomen commonly men-
tioned as a character separating Lepidurus from Apus. Packard
(1883), however, mentions that L. bilobatus has an unusually long
abdomen, with many body-rings exposed behind the carapace; and I
found, when examining a specimen of this species marked ‘‘type’’ in
the U. S. National Museum, that it has 16.5 leg-bearing abdominal
rings. Further, in material consisting of more than a hundred speci-
mens of a new species from the State of Washington, I found this
number to be 14.5—-18, while 12 specimens of a variety of this species had
16-17.5. Corresponding to the high number of leg-bearing abdominal
rings we have a higher total number of body-rings here than in other
species of the genus; there are more legs, and, generally, more rings
exposed behind the carapace. In my opinion, these figures indicate
that there are two groups of species within the genus Lepidurus,
whatever the taxonomic category we choose for them.
In the American forms of Apus, the number of leg-bearing abdom-
inal rings ranges between 16 and 21 (males 16-18.5, females 17.5-21).
In one large lot I have seen every number in this range represented (see
fig. 31). Usually, the females have more, sometimes many more,
leg-bearing abdominal rings than the males of the same species.
12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
The variation in the number of legless rings is often used as a
taxonomic character. This appears to be a useful character in South
African forms of Apus (Barnard, 1929), but in the North American
forms I have not found it to be of much aid in separating species.
In Lepidurus, the number varies within narrow limits (4—5.5 in the
great majority of forms), with no specific differences corresponding to
the differences in number; a few specimens having 3, 3.5, or 6 legless
rings show no other essential differences. There is, of course, the
possibility that Z. packardi may have a high number and that this
character may be of taxonomic importance, the type specimen having
6 legless rings. But I do not think this is probable. In two lots (5
specimens) that I consider belonging to this species I found 4.5-6
legless rings.
In Apus, the whole range in the American specimens I have seen
ig 4.5+1-13 in the females and 8-15-++-i in the males. It seems that
this range of variation ought to be great enough to include several
species (or subspecies, or forms) with varying ranges, and I will cer-
tainly not deny the possibility that lower taxonomic units than species
may be separated. However, I have seen a single lot of Apus that
covered much of this range (9+1-13 in the females and 12.5-15+1
in the males, as shown in table 6). This is an extreme example, the
range of variation being smaller in other samples, especially in those
with only a few specimens,
As arule, the males have more legless rings than the females of the
same species because they usually have fewer leg-bearing rings but
a higher total number of rings.
BODY-RINGS IN PARTHENOGENETIC POPULATIONS
It seems rather remarkable that an author like Zaddach (1841),
extremely careful in so many details, gives a fixed number of 34
body-rings in Apus caneriformis. And Rosenberg (1947) definitely
states that all specimens, several hundred in number, of his species
A. oryzaphagus have 35 body-rings (36 including the telson, according
to him). J have seen 6 of them (U.S.N.M. No. 88360) and found
5 to have 35 and 1 to have 34+1 body-rings. Rosenberg describes
another form, under the name of A. biggsi, where all specimens have
36 rings (37 including the telson), but here it seems uncertain whether
he had enough specimens for comparison. I found 3 of them having
36 and 1 having 35++i rings (U.S.N.M. No. 88361). In these cases
the statements of a fixed number may result from the counting of
incomplete rings as ordinary rings. Even so, the range of variation
is very small. =| “N Bqos0IO MW WIoIp 2288nN09 “7
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38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
European part of the U.S. S. R.; Jakutsk (Sars, 1897), several places
in the territory of Akmolinsk (Sars, 1901, 1903), territory of Koksche-
tavsk (Sars, 1903), and Kasalinsk in Turkestan (Decksbach, 1924).
Remarks.—Outside America, there is a species with a similar telson
and supra-anal plate, LZ. macrurus Lilljeborg. The general resem-
blance between this species and L. couesi was noted by Lilljeborg,
who mentioned (1877, p. 14) one difference only, that the former has
6 to 8 rings exposed behind the carapace and the latter only 5. This
character ‘is generally recognized to be somewhat unreliable” (Gur-
ney, 1923, p. 502) because of the varying state of contraction of the
specimens (p. 5), and I have seen American specimens of L. couesit
with 7 exposed rings as well as specimens of LZ. macrurus, identified by
Lilljeborg, with five exposed rings.
I have examined 25 Asiatic specimens identified as LZ. macrurus by
Lilljeborg. They, as well as some specimens from a more southern
locality, agree perfectly well with the American specimens of L. couesit
as described, and ought to bear the same name, which is 2 years older
than that given by Lilljeborg.
The relationship of this species to the European L. apus, to which
it bears a certain resemblance, is in need of clarification, but I have
not had enough material of the latter species to make a detailed
comparison.
LEPIDURUS BILOBATUS Packard
Figure 13; Puars 2 (Ficurss 3, 4)
Lepidurus bilobatus PackKarp, 1883, p. 318.
Description—A supplementary description of a female labeled
“Type, Utah, coll. Henshaw” (U.S.N.M. No. 11606) is given here.
Body-rings 11+16.5+4.5=32. Legless rings with many (19) densely
crowded spines on the ventral side, dorsally 7 spines. Supra-anal
plate slightly bilobed, with 8 dorsal spines, 4 of which are smaller than
the others, all of them placed on a slightly marked keel. Nuchal
organ oval, placed between hind parts of eye-tubercles. About 62
pairs of legs. In other respects there is reasonable agreement with
Packard’s description. Further particulars are given in table 1 (p. 36).
Remarices —The real status of this type seems to be a little uncertain,
because Packard’s type locality was not Utah, but Po Cafion, Ver-
million River, Colo. However, the specimen most probably comes
from Packard’s collection, and I feel reasonably certain that it belongs
to this species. The large number of abdominal rings agrees well
with Packard’s statements that “the abdomen is longer than usual’’
and “‘there are about sixteen segments beyond the posterior edge of
the carapace.’”’ Another species with many abdominal rings is de-
scribed later on in this paper (p. 39), but LZ. bilobatus is easily dis-
tinguished by the position of the nuchal organ, the typical arrange-
ment of the spines on the posterior emargination of the carapace, and
NORTH AMERICAN NOTOSTRACA—LINDER 39
by the varying sizes of the spines on the supra-anal plate. In the
two last mentioned respects, there is good agreement with Packard’s
description. ‘Seven more or less well-marked median spines” on the
telson, according to Packard, stands in sufficiently good accordance
with the condition in this specimen. Bilobation of the supra-anal
plate occurs here, in agreement with Packard’s description, but it is
to be remembered that this character is not an important one.
Tentatively, I have placed here another specimen (U.S.N.M. No.
82028) which bears only the label ‘“‘ Probably Arizona.”’
In a case like this, when there is a little uncertainty about the
status of the single existing type specimen, and the original description
is not very good, it may be questionable whether we ought to discard
the species or keep it on record. In my opinion, the description is
good enough to enable us to identify the species with reasonable
accuracy. However, it seems desirable to search for more material
from the type locality or its vicinity.
LEPIDURUS LYNCHI, new species
Figures 14-18; 19, c; 20-27; PLatss 3, 4, aND 7 (F1GuRE 3)
Type specimens.—Holotype (1 9), allotype (1 co), and paratypes
(Uppsala Mus.); paratypes also in U.S.N.M. (No. 82101); ponds
in the upper Grand Coulee, Grant County, Wash; collector J. E.
Lynch. Additional paratypes (699, 2o%<7, 5 juniors) from Fish Lake
Valley, Nev.; collector, La Rivers.
FEMALES (52 specimens): Holotype 11+16.5+4.5=32 body-rings;
paratypes 11+ (15-18)+ (3 +i-5.5)=30+1-34 body-rings. Surface of
all body-rings smooth, spines on dorsal and lateral quarters of caudal
border of legless rings large, smaller on ventral quarter. On penulti-
mate ring 9 large, 8 small spines; in paratypes 6-14 and 4-9, respec-
tively, the boundary between large and small spines not always
sharp, because they may diminish in size gradually. Telson (disre-
garding supra-anal plate) long, its length almost as great as its width,
smooth except for small spines around dorsal sense organs and around
bases of caudal filaments. Supra-anal plate 4 mm. in length (24
percent of carapace); in paratypes 0.6-8 mm. (7-30 percent of
length of carapace, highest percentage being in large specimens).
This plate irregularly tongue-shaped, with coarse, small spines on
its margin, and 4 rather large spines dorsally in midline; in paratypes
broadly triangular, tongue shaped or spatulate, often more or less
bilobed, often irregular, with 2—7 dorsal spines. Caudal filaments
24 mm.; in paratypes 11-39 mm. 18 body-rings exposed beyond
carapace; in paratypes 12-19. Color of preserved specimens greenish
brown. Total length disregarding supra-anal plate and caudal
filaments, 26 mm.; paratypes 12-45.5 mm. (preserved specimens!).
40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
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Ficure 14.—Diagram indicating the number of body-rings in the abdomen and the length
of the carapace in 42 female paratypes of Lepidurus lynchi, new species (males present
in the sample not considered here). Each shaded triangle at the top of the diagram
represents the telson of a specimen, below which are indicated the legless abdominal
rings (dotted) and the leg-bearing rings. Incomplete rings at the anterior boundary
of the telson are denoted by the oblique position of the triangle. Body-ring numbers
are shown in the marginal columns at the left and right of this portion of the diagram.
The carapace length of each specimen in mm. is indicated in the lower part of the figure,
according to the scale in the left and right marginal columns. Correlations of various
kinds are reserved for a later statistical report on more comprehensive material.
Carapace length 17.5 mm.; width 2 x 11.5 mm., width: length
relation 1.22; in paratypes length 8.5-26 mm., width 2 x 5-2 x 15 mm.,
width: length relation 1.14-1.30. Caudal half of lateral margin
serrated by numerous, minute, pointed spines. Dorsal carina slightly
marked, with 19 short spines evenly scattered along whole carina; in
paratypes carina often inconspicuous or wanting, with 0-20 spines.
Posterior emargination rather deep, 5 mm., with 12 short spines on
one side, 13 on the other, not quite similar in size and somewhat
irregularly placed; in paratypes 7-17 spines on each side. Supra-
antennal crest quite smooth.
NORTH AMERICAN NOTOSTRACA—LINDER 41
Ficure 15.—Dorsal view of telson and last two body-rings of a female of Lepidurus lyncht,
new species, X 18.
Ficure 16.—Three pieces of an incomplete ring at the anterior margin of the telson in a
female of Lepidurus lynchi, new species, X 11: a, Dorsal view; b, ventral view.
Eyes unusually small. Nuchal organ round, elevated to about
same height as the small eye-tubercles, and placed clearly behind
them; in paratypes more or less rounded form, rather varying height
but always similar in position.
Antennae, mandibulae, and maxillae as usual in the genus.
Pairs of legs 61; in paratypes 60-71. Endites 2, 3, 4, 5 on first
leg 1, 5, 3.4 and 8 mm., long, respectively; endopodite very short,
102
PROCEEDINGS OF THE NATIONAL MUSEUM VOL.
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NORTH AMERICAN NOTOSTRACA—LINDER 43
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Ficure 17.—a, Oblique ventral view of part of the abdomen and telson of the largest female;
of Lepidurus lynchi, new species, showing a right-handed spiral of a little more than four
rounds beginning in the leg-bearing part of the body and forming the entire legless part
of it, X 8.5. The telson is shown in ventral view because of a slight natural torsion of the
telson. Legs omitted, as well as spines on caudal filaments. Only basal parts of supra-
anal plate and caudal filaments shown. (C.f., caudal filaments; S.4.p/., supra-anal plate;
Sp, spiral; T, telson.) b, Schematic drawing of the spiral.
,
ae os
\
Ficure 18.—Dorsal view of posterior emargination of the carapace of a female paratype of
Lepidurus lynchi, new species (Uppsala Museum), X 7.
fringed with setae on its caudal margin; in paratypes (fig. 21) varying
lengths, but fifth endite always at least double the length of fourth and
reaches, when stretched backward, usually nearly to the posterior
angle of carapace. On second leg endites 3, 4, and 5 rather similar in
length, endopodite long, 2.2 mm., pointed, of about same length as
fifth endite, fringed with setae on its caudal margin. Eleventh
leg with a small epipodite, fringed with setae, and trace of a pre-
44 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 102
Ficure 19.—Eyes and nuchal organ: a, Female holotype of Lepidurus packardi Simon
(Paris Mus.), X12; 6, male paratype of Lepidurus couesii Packard from Montana (Stock-
holm Mus.), 12; c, female paratype of Lepidurus lynchi, new species (Uppsala Mus.),
8; d, male of Apus longicaudatus from Wyoming (U.S.N.M. No. 58766), X 8.
Figure 20.—Ventral view of a portion of the right side of a female of Lepidurus lyncht,
new species, from which the last 25 legs have been removed, X 21. The specimen has
11+17.5+3.5=32 body-rings and 64 pairs of legs. a, Bases of legs; b, base of last (sixty-
fourth) leg; c, endopodite of thirty-ninth leg; d, twenty-ninth body-ring.
epipodite in the form of an elevation with a few setae. Last leg very
small.
Mates (55 specimens): Allotype 11+14.5+4.5+i=30-+1 body-
rings; paratypes 11+ (14.5-17) + (8.5-5.5) =30-33-+1 body-rings. Sur-
face of body-rings smooth, spines on caudal border of legless rings
NORTH AMERICAN NOTOSTRACA—LINDER 45
about as numerous as in female. Telson, except supra-anal plate, as
in female. Supra-anal plate 6.5 mm. in length, 35 percent of length
of carapace; in paratypes 0.7-5 mm., 18-32 percent of length of cara-
pace. This plate irregularly bilobed, with 2 rather large spines dor-
sally in midline; in paratypes form varying as in female, with 2-6
spines. Caudal filaments 23 mm.; in paratypes 14-25.5 mm. Body-
rings exposed behind carapace 15+-1; in paratypes 10-19. Color as
in female.
Total length without supra-anal plate and without caudal filaments
29.5 mm.; paratypes 12—24 mm.
Carapace length 18.5 mm., width 2 x 11 mm., width: length relation
1.19; in paratypes length 11.5-16.5, width 2x 7-2x 10 mm., width:
length relation 1.08-1.33. Anterior part broader than in females.
Spines on lateral margin of carapace as in females. Carina missing;
one short spine in midline; in paratypes sometimes trace of an incon-
spicuous carina at the places where there are spines, these ranging
x
Ficure 21.—First leg of a female paratype of Lepidurus lynchi, new species (Uppsala Mus.),
X 6.
AG PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
from 0-28 in number. Posterior emargination rather deep, 5 mm.
in depth, with 13 rather conspicuous spines on one side, 14 on the
other, a little irregularly placed and not quite similar in size; in
paratypes rather deep, 7-16 spines on each side.
Eyes, nuchal organ, and supra-antennal crest as in female.
Antennae, mandibulae, and maxillae as in female.
Pairs of legs 60; in paratypes 60-67 (only 5 specimens counted).
Length of endites 2, 3, 4, and 5 on first leg 0.8, 3.4, 4 and 10.2 mm.,
respectively; endopodite very short, fringed with setae on its caudal
margin; in paratypes varying lengths, but fifth endite always more
than double the length of fourth endite, reaching, when stretched
backward, more than halfway to end of carapace. On second leg
Ficure 22. Eleventh leg of a female paratype of Lepidurus lynchi, new species (Uppsala
(Mus.), X 12: a, Epipodite; b, pre-epidodite.
Ficure 23.—Last (seventy-first) leg of a female paratype of Lepidurus lynchi, new species
(Uppsala Mus.), X 200: a-e, First to fifth endite; f, endopodite; g, exopodite; h, epipodite.
NORTH AMERICAN NOTOSTRACA—LINDER A7
Ficure 24.—Dorsal view of supra-anal plate of a male paratype of Lepidurus lynchi, new
species (Uppsala Mus.), * 10.5.
Ficure 25.—First leg of a male paratype of Lepidurus lynchit, new species (Uppsala Mus.),
Merk
endites 3 and 4 of same length, 2.1 mm., endite 5, 4.2 mm., endopodite
long and narrow, 2.2 mm., fringed with setae on its caudal margin;
VOL. 102
AS PROCEEDINGS OF THE NATIONAL MUSEUM
in paratypes varying lengths, fifth endite sometimes not much longer
Eleventh leg with trace of a pre-epipodite as in
than endopodite.
female.
Vf fi pf p
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h
Ficure 26.—Second leg of male paratype of Lepidurus lynchi, new species (Uppsala Mus.),
X 7: a-e, First to fifth endite; f, endopodite; g, exopodite;fh, epipodite.
MY fy,
HY
Figure 27.—Eleventh leg of a male paratype of Lepidurus lynchi, new species (Uppsala
Mus.), X 13.
NORTH AMERICAN NOTOSTRACA—LINDER 49
Remarks.—This new species is especially interesting, because it
belongs (see table 3) with Z. bilobatus in the group with many abdom-
inal rings. The variation of the species, as limited above, may
perhaps seem to be rather great. It includes forms with a very
spiny carapace together with forms where the carapace is quite
smooth; but intermediate forms occur in great numbers, and we find
a similar variation in several species of the genus Apus (Barnard,
1929). Furthermore, the form of the supra-anal plate is highly
variable; but, as I have shown previously (p. 26), this is true of many
species. The great range of variation must be seen in connection
with the fact that the species is described from such a great number of
specimens from the same district.
The species is easily distinguished from L. bilobatus by the size and
place of the nuchal organ and the eyes, characters to which I must
ascribe great taxonomic importance. Its relation to the following
form, however, I cannot regard as quite clear.
LEPIDURUS LYNCHI ECHINATUS, new variety
Ficure 28; Puarses 5 (Ficure 1), anp 7 (FiaureE 4)
Type specimens.—Holotype 9 (U.S.N.M. No. 82068), allotype @
(U.S.N.M. No. 82069), paratypes 6 @ and 4? (U.S.N.M. No. 82070
and Uppsala Mus.). North end of Goose Lake, about 15 miles
southwest of Lakeview, Lake County, Oreg.; altitude 4,700 feet; in
water 2 feet deep, muddy; June 14, 1940; collector, J. E. Lynch.
Ficure 28.—Dorsal view of telson and last two body-rings of a female of Lepidurus lynchi
echinatus, new variety, X 9. (S, dorsal sensory setae.)
Body-rings 11+ (16—17.5)+ (3.5-4.5) =31-33. Telson rather long.
Always many large spines on carina of carapace. In females, 2-5
dorsal spines on supra-anal plate, in males, 3. Legs 67-68 in number
(only two specimens). Total length, disregarding supra-anal plate
and caudal filaments: Female 32-41 mm., male 22-30 mm. Length
of carapace: Female 21-28 mm., male 17.5-21 mm. Width: length
relation of carapace 1.21-1.29.
956651524
50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Very similar to the typical form. All characters lie within the
range of variation mentioned for this form, except for some details
of the carapace. On the middle and caudal part of its lateral margin
there are numerous stout spines considerably larger than those in the
principal form, though not so large as the spines at the posterior
emargination. All the first-mentioned spines are directed straight
outward. This is a very conspicuous feature and the explanation of
the name echinatus. Furthermore, the carapace is more rounded and
a little larger in relation to the body than in the typical form, and its
posterior emargination is less deep, leaving a smaller number (7-10)
of rings exposed behind the carapace.
Remarks.—It is regrettable that there are not more than 12 speci-
mens of this variety, because this small lot cannot be expected to
show much of the variation. As a provisional measure I have given
it the rank of a variety within Z. lynchi, because it shows a great
similarity to this species in such important respects as the number of
abdominal rings, the characteristic appearance of the eyes and the
nuchal organ, and the array of spines on the lateral margins of the
carapace. At the same time, there are clear differences between the
two forms in the shape of the carapace.
There is a certain similarity between this variety and L. lemmoni
Holmes, especially in the array of lateral spines on the carapace and,
perhaps, also in the position of the nuchal organ.
But, as I will discuss in more detail later (p. 51), the similarities are
more or less uncertain, and I consider L. lemmoni as an insufficiently
known species. There are dissimilarities, in any event, among which
the different shape of the posterior emargination of the carapace is the
first to attract attention. I consider it better in this case to estab-
lish a new variety than to venture a highly dubious identification with
an insufficiently described species.
LEPIDURUS LEMMONI Holmes
Lepidurus lemmont Houmss, 1894, p. 585 (not Coopey, 1946, p. 338).
Description.—The principal points of the description of this insuffi-
ciently known species follow: About 4 body-rings exposed behind
carapace, 4 or less legless body-rings (the author says 5 instead of 4,
but in all probability he counts the telson as a ring). Number of
spines on last body-ring never far from 12. Supra-anal plate rather
long and may be rounded, truncated (obliquely so in some specimens),
or, in some cases, bilobed, with 2-3 spines in midline, but no median
carina except where the spines are situated. Carapace oval, carina
with 10-15 teeth, posterior emargination with 5-7 teeth, sides with
numerous small teeth. Nuchal organ round, small, considerably
elevated, and located considerably behind the posterior margin of
eyes. Length, 28 mm.
NORTH AMERICAN NOTOSTRACA—LINDER 51
Type locality.—Honey Lake, Calif.
Types.—The type specimens (4 9, 1c”) are said to have been lost
in the San Francisco fire.
Remarks.—Holmes’ description is very good for its time, but
naturally it does not mention the number of abdominal leg-bearing
rings or the total number of rings. In these respects, there are two
well demarcated groups within the genus, and these totals seem to be
essential for a proper identification. Perhaps both were low, because
he reports only 4 rings exposed behind the carapace; but we cannot
be certain of this. Equivalent totals in L. lynchi var. echinatus are
high even though as few as 7 rings are exposed. Only 12 specimens
are known of the latter form, and only 5 of LZ. lemmoni. So few speci-
mens cannot be expected to show a full range of variation; thus, it
is possible that LZ. lemmoni may fall in within the upper limits of
variation in this respect. And we must not forget that, in general,
the number of exposed rings is shown to be an unreliable character,
so its value as an indicator of the number of rings is very uncertain.
Other characters do not give clear indications, either. The nuchal
organ is “‘located considerably behind the posterior margins of the
eyes.” Perhaps we have a condition here similar to that in L. lynchi
and L. lynchi var. echinatus, but a precise comparison is impossible
because Holmes does not mention the place of the organ in relation
to the eye-tubercles, only to the eyes.
The very long caudal filaments present a character of doubtful
significance. In each large lot of Lepidurus there is considerable
variation in the length of these appendages.
The lateral spines on the carapace look quite conspicuous on Holmes’
figure. Similar spines, or larger still, are found in L. lynchi var.
echinatus, whereas in all other notostracans I have seen the correspond-
ing structures are minute and inconspicuous. But echinatus differs
clearly from ZL. lemmoni by its very shallow posterior emargination
of the carapace as well as by the structure of its legs, and so an
identification with this form seems highly improbable.
To sum up, the characters given for L. lemmoni cannot be regarded
as sufficient for a proper identification. In my opinion, we had
better not keep the species on record.
It can be abandoned without much trouble, because, as far as I
can see, it has been recorded only once since its establishment, and in
this case there is undoubtedly an error of identification. Coopey
(1946) refers some specimens from Oregon to this species, though
with the express reservation that they are only provisionally identified.
Most kindly, be has given me the opportunity of examining 3 speci-
mens from Rocky Point Pond, Klamath County. I find them to
belong to L. couesii. The telson (without supra-anal plate) is short
and broad, and on the conspicuous keel of the supra-anal plate there
52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
is a large number of spines of varying sizes (2-3 spines in L. lemmoni).
Even the place of the nuchal organ is as in L. couesw. All characters
considered, the specimens are quite typical representatives of this
species.
Genus APUS Schaeffer
Apus SCHAEFFER, 1756, p. 131.—GurNEy, 1923, p. 497.—Barnarp, 1929, p. 229.1
The principal characters of the genus are, in my opinion, the
absence of a supra-anal plate on the telson (p. 26), the relatively
large number of body-rings, 30-44 as against 25-34 in Lepidurus
(p. 42), and the reduction of the second maxillae in full-grown speci-
mens (p. 28).
Even within these limits, however, the genus presents a unit that
is not quite homogenous or well distinguished from the genus Lepi-
durus, because one species, A. cancriformis, is in several respects sug-
gestive of the latter genus.
Absence of a supra-anal plate seems to be characteristic of all
species except A. cancriformis, where we may find something rather
similar to such a plate (fig. 8). It is very short, but we can only
assume that it is essentially the same structure as the plate in Lepi-
durus, so that A. cancriformis may be regarded as an intermediate
stage, in this respect, between Lepidurus and the remaining species
of Apus.
In the reduction of the second maxillae, the same species forms an
exception. I have found it to have, in full-grown specimens, just as
well developed maxillae as in any species of Lepidurus.
It has very few body-rings as compared with almost all the other
species of the genus that I have examined. There are a minimum of
30, and, generally, a maximum of 34. In only one sample of A.
cancriformis have I found more, 35 in a female and 35 or 37 in the
males (U.S.N.M. No. 58211, from Nanking, China). I have observed
a similarly small number of rings in A. sudanicus from South Africa,
identified by Barnard (U.S.N.M. No. 75749). The male in this
sample has 32, the two females 33 and 33+ body-rings. All the other
species of Apus have a larger number of body-rings. A. longicaudatus
comes nearest with 34-++i-44 rings (see table 4); however, I am fully
aware that in many forms we know too little of the number to be able
to make definite groupings of the species according to this character.
Still, it may be considered significant that A. cancriformis, which comes
close to Lepidurus in the structure of the telson and the second
maxillae, also has a similar total number of body-rings. This is a
further indication that the species is a sort of connecting link between
the two genera.
1I fully agree with Gurney and Bamard that the name of Triops Schrank, 1803, pp. 180, 251 (sometimes
spelled Triopes or Thriops), ought to be rejected.
NORTH AMERICAN NOTOSTRACA—LINDER 53
In its general appearance, also, this species is suggestive of the genus
Lepidurus. The body, because of its small number of rings, may be
almost completely covered by the carapace, which sometimes leaves
only part of the telson exposed. Such proportions are unusual in
the species of Apus, and it is interesting to notice that they occur,
not in the species of Lepidurus that have a similar number of rings
(L. bilobatus, L. lynchi), but in those that have the fewest rings.
The number of abdominal leg-bearing rings, a figure that proved
helpful as a means of distinguishing groups within the genus Lepidurus
(p. 31), gives no clear character in Apus. In the whole genus, it
ranges from 12.5 to 20, figures that are generally higher than in Lepi-
durus. Im A. cancriformis, the range is usually 16 to 18, though I
have occasionally observed either fewer or more of such rings (the
fewest was noticed, not in this species, but in A. sudanicus, as noted
above, where a male has 12.5 and two females 13.5 and 14 such rings).
The number of pairs of legs in various species is too little known to
enable us to draw any conclusions about its taxonomic value.
On the whole, we must admit that much work remains to be done
before we will be able to make a reasonably good arrangement of the
species of Apus.
Distribution in America: Montana, Oregon, Wyoming, California,
Nevada, Utah, Colorado, Nebraska, Arizona, New Mexico, Kansas,
Oklahoma, Texas; Galapagos Islands; Hawaiian Islands; Mexico;
Haiti; St. Vincent Island; Argentina.
APUS LONGICAUDATUS LeConte
Ficures 3-6, 19, d, 29-31: Puares 5 (Ficures 2, 3), 6, and 7 (Ficures 1, 2)
Apus longicaudatus LeConve, 1846, p. 274.—PackarD, 1883, p. 324.—MackIn,
1939, p. 46.
Apus aequalis PacKARD, 1871, p. 3; 1883, p. 320.—Prarsg, 1912, p. 192; 1913, p. 2.
Apus newberryi PACKARD, 1871, p. 2; 1883, p. 321.
Apus lucasanus PacKarD, 1871, p. 2; 1883, p. 322.
Apus oryzaphagus ROSENBERG, 1947, p. 70.
Apus biggst ROSENBERG, 1947, p. 72.
As far as I can see, the various species of Apus described from the
North American continent must be united into one species, A. longi-
caudatus. A study of reasonably extensive material shows that the
three species of Packard, enumerated above, cannot be properly de-
marcated from each other and from A. longicaudatus, and the same
applies to Rosenberg’s species.
Packard’s material was probably too limited to enable him to detect
the unbroken series of variation in most of the characters used by him.
It was not so much that he had material from so few localities—
4 species from 13 localities—as that only a few specimens were avail-
able from each locality. One of his species, A. newberryi, was based
upon three females from two localities.
102
vol.
PROCEEDINGS OF THE NATIONAL MUSEUM
54
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NORTH AMERICAN NOTOSTRACA—-LINDER 57
I suspect that many carcinologists have had difficulties trying to
identify species of Apus with the aid of Packard’s descriptions.
Pearse (1912) gives a good example. He had an unusually large
number of specimens from one locality and found a remarkable
variation in the number of dorsal spines on the telson, which Packard
uses as one of the main characters: ‘The armature of the telson is
somewhat variable; on the dorsal side there is commonly one median
spine, but there are often two; there are usually two lateral spines at
the proximal edge of the telson, but there are frequently more”
(Pearse, 1912). Here is a mixture of characters from several species.
A. lucasanus and A. longicaudatus are described with one median
spine, A. aequalis with two, as is one of the specimens of A. newberryi
mentioned by Packard. ‘Lateral spines at the proximal edge of the
telson,” probably means the spines around the dorsal sensory setae, of
which one on each side is described as large in A. lucasanus. Thus
most of Pearse’s specimens ought to have agreed with A. lucasanus,
but others not. Further, Pearse found the length of the carapace to
be “about equal to the portion of the body exposed behind,” as in
A. aequalis according to Packard; it was obviously on these grounds
that he identified his specimens with this species and dismissed the
similarities with other species (especially A. lucasanus), which were
observed on the telson of most specimens.
Mackin (1939) says that Packard’s three species are synonyms of
LeConte’s species; his studies on material from Oklahoma and
neighboring States showed that the characters used by Packard are
of little value in taxonomics. However, he regards his conclusion as
only tentative.
Packard’s descriptions cover a lot of characters, but many of them
are considered in only one species, so that comparisons in some re-
spects are impossible. Those presented in table 5 are especially
stressed by him. (In this table I have, in all cases, given one less
ring than Packard, because he obviously counts the telson as a body
ring, while I do not.)
The length of carapace in relation to length of body, and the number
of body-rings exposed behind the carapace, vary according to the
contraction of the rings, a condition itself highly variable. These
characters are, of course, also influenced by the number of body
rings, though in a rather erratic way on account of the variable con-
traction of the rings. These characters of Packard’s are not simple
characters, and generally they are more or less unreliable.
As for the number of legless rings, the examples in table 6 show
how greatly this often varies in specimens from the same locality.
The sample from Wyoming (columns a, c) is significant. It contains
many more specimens than the others, and it covers the numbers
given by Packard as characteristic of three species, A. newberryt
58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 102
(10 in females), A. Jucasanus (10 in females), and A. longicaudatus (9 in
females, 13 in males). More than that, it presents an unbroken series
beginning with a number of legless rings not much greater than that
given for A. aequalis and ending with a number (15+) considerably
higher than those observed by Packard. No correlation with other
characters has been found.
TABLE 6.—Variation in the number of legless rings in populations of Apus
Females Males
U.S.N.M. | U.S.N.M. | U.S.N.M. | U.S.N.M. | Stockholm
Number of legless rings No. 58766 | No. 61750 | No. 58766 | No.61750 | Mus.
(Wyoming)} (Texas) |(Wyoming)| (Texas) (i al S
’ :
=
K
o
Ficure 40.—Map of southern Guam Island, showing collecting localities and relative mean
shell-size of colonies (see p. 103 for explanation).
SNAIL HOST OF ORIENTAL LUNG FLUKE—ABBOTT 97
LIFE HISTORY
HABITAT
As has been stated earlier, the various subspecies or geographical
races of Vhiara granifera have not been studied adequately, so that
details of its geographical distribution cannot be presented at this
time. However, the limits of its distribution as a whole are fairly well
known. This species lives in fast-flowing fresh-water streams. Its
most westerly limits include the Island of Ceylon and the eastern coast
of India. Its range extends easterly to include the coasts of Siam and
southern China, the East Indies, and the Philippine Islands. ‘To the
north it is found on Formosa (Taiwan) Island and the small islands
south of Japan. Farther east, it is common in Melanesia, especially
the Solomon Island Archipelago, the Marianas, and the Polynesian
islands of the Hawaiian and the Society Islands. Undoubtedly the
topography and nature of the areas in which this species is found
varies from place to place, but it is highly likely that certain optimum
environmental conditions are common to all its localities. Thus we
may find this species in streams very near the sea in certain small
islands, but considerably inland and at higher altitudes on such large
islands as Luzon, Leyte, and Mindanao in the Philippine Islands.
In the isolated Micronesian islands of the Marianas only three
islands are large enough to have permanent streams: Guam, Saipan,
and Rota. All three support colonies of T'hiara (Plotiopsis) scabra,
but only the first possesses streams large enough to maintain popula-
tions of Thiara (Tarebia) granifera. Guam (Guajan) Island is 82
miles long by 4 to 10 miles wide and about 225 square miles in area.
It is mountainous in the southern half, where a great number of small
streams flow down to the sea. The northern half forms a low plateau,
which is devoid of permanent streams.
The majority of the streams in the southern half of Guam are col-
onized by 7'hiara granifera. Collections were made at the localities
listed. The number following each station record indicates the number
of specimens collected (fig. 40).
A. Zadue Maagos River. 1945. D. Frey collector. 92.
B. Agafia Spring, 114 miles southeast of Agafia. April 22,1945. R.T. Abbott
collector. 198.
C. Talisai stream, under bridge, 2 miles southeast of Agat. May 1, 1945.
R. T. Abbott collector. 100.
D. Water Plant, 2 miles up Ylig River. April 26,1945. R.T. Abbott collector.
65. Flow rapid, 6 feet wide, 6 inches deep.
E. 1 mile up Pago River. April 26, 1945. R.T. Abbott collector. 50. Flow
rapid, 6 feet wide, 4 inches deep.
F. First stream 600 yards south of Piti. April 24, 1945. R. T. Abbott col-
lector. 38. Flow sluggish, 3 feet wide, 1 inch deep; shady.
G. Geus River. 1945. D. Frey collector. 97.
H
. 100 yards up Aguada Stream. April 24,1945. R.T. Abbott collector. 127.
Flow sluggish, 4 feet wide, 2 inches deep: shady.
98 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
I. 1 mile up La Sa Fua River. May 10, 1945. R. T. Abbott collector. 20.
Flow sluggish, 4 feet wide, 2 inches deep; shady.
J. % mile up Inarajan River. May 2, 1945. R. T. Abbott collector. 48.
K. Merizo River. 1945. D. Frey collector. 245.
L. Small stream near Ajino Beach. 1945. D. Frey collector. 8.
M. 1% mile up Umatac River. May 10, 1945. R. T. Abbott collector. 36.
Flow rapid. 30 feet wide, 8 inches deep; no shade.
The snails are most abundant in the shallow riffles where the flow
of water is fairly rapid and where the bottom consists of a pavement
of small stones. Occasionally the snails may be found congregated
on firm sandy bottom provided there is a healthy flow of water. Waters
of the stream that are exposed to direct sunlight during most of the
day appear to attract this species. The upper reaches of the streams
that have a very rapid flow of water are not favorable, although other
mollusks such as Veritina and Septaria may be abundantly represented.
The tiny flows of headwater tributaries, which are less than a foot
in width and 2 inches in depth, will not support 7’. granifera but are
favorable to 7’. scabra. The stream at Umatac, in the southwest end
of Guam, has the colony with the greatest number of largest individ-
uals. The maximum concentration of snails (about 10 per square
foot) is found in the fairly level section of the stream, which is about
30 feet wide and half a foot deep. This area is only a few hundred
feet from the high-tide mark of the ocean but is probably never diluted
with salt water except perhaps during typhoons, which may come from
the west.
Conditions at Agafia Spring, the most northerly located colony on
the island, are interesting in the dwarfing effect on the size of the
individuals. The spring is used as a source of water for nearby Agana
and at present has a pumping station located there. The spring
proper is about 30 feet deep, with a diameter of about 60 feet. The
upsurge of water is moderate, but the overflow is relatively fast, con-
sisting of a 100-foot sluiceway about 3 or 4 feet wide. There are
abundant algal growths in the spring. The shells and animals of this
colony are similar to those of other Guam populations except in the
reduction in size. No environmental cause for this could be found.
Even more pronounced dwarfing is evident in specimens that were
introduced to the large spring at Lithia, Fla.
On Leyte Island, Philippine Islands, where numerous collecting sta-
tions were made, 7hiara granifera was found in large rivers and small
streams. High in the central mountains, where large stream condi-
tions duplicate those on Guam, colonies of this snail are rather com-
mon. In the extensive Leyte Valley, at the northeast end of the island,
the larger rivers, with silty to sandy bottoms, meander; only rarely
is 7’. granifera found along the very edges of the rivers, where the flow
of water is moderately fast. However, smaller streams, which pass
down from the foothills bordering the valley, are excellent habitats.
SNAIL HOST OF ORIENTAL LUNG FLUKE—ABBOTT 99
The water-temperature limits of this species were not appreciated
until a survey was made of the Lithia Spring, Fla., populations. In
aquaria and the spring, where the temperature remains around 76° F.,
the snails are able to maintain themselves in adequate numbers. How-
ever, just beyond the influence of the warm waters of the spring, where
the cooler waters of the local river are about 50° F. or less, the snails
are absent. The overflow ditch outside the extensive plant aquaria
in Tampa is colonized by these snails during the warm summer months,
but in winter, when the water temperature is as low as 50° F., these
snails die off. The temperature recordings for Guam streams and
even for the rivers at 3,000 feet altitude on Leyte Island were all
above 75° F.
BIONOMICS
GRrowTH AND sizE.—The rate of growth of the shell has not been de-
termined, although, from common experience in raising these snails
in aquaria, it is assumed that adult size or maturity is reached within
6 to 12 months. Two preliminary studies were made of the character
of shell size, one on the 13 colonies collected on Guam Island, the other
on the populations of Lithia Spring. In the latter study, dissections
were made to determine the size at which individuals become sexually
mature, for there is no tell-tale flaring or thickening to the outer lip
that often accompanies maturity in many other molluscan species.
The significant differences in the size of samples taken at various
places in the spring, only a few dozen yards from each other, are
extremely important in demonstrating the wide range in size exhibited
by one species. The aquarial stock from which these snails originated
grew to a size almost four times that of its offspring in the spring.
Only in an isolated, subsidiary spring, a few yards from the main
spring, did specimens reach a size comparable to the aquarial stock.
In all likelihood the differences in size exhibited by the seven sample
populations in the spring have been brought about by ecological and
not genetic factors. Not only are the means of shell lengths and maxi-
mum sizes of shell considerably varied, but the points at which ma-
turity is reached are correspondingly shifted. This latter fact ex-
cludes the premise that these colonies represent aggregates of smaller,
younger individuals. No direct correlation between diminutive size
and type of bottom, depth of water, temperature of water, amount of
shade, or distance from source of spring could be found. The small
colony in the adjacent subsidiary spring attained a shell size nearly
three times that of colonies in the main spring. Relatively few speci-
mens per square foot were found in the smaller spring, while in the
main spring overcrowding was evident, sometimes reaching a popula-
tion density of 400 specimens per square foot. It is likely that food
availability is the most important factor, but controlled experiments
alone will support this assumption.
100 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
TaBLE 3.—Data pertaining to collections from Lithia Spring, Fla.
Number | weanshell | Point of Typeot | Depthot | , Distance
Station speniaiens length, mm. Saagatity Botton eater (ft.) paar:
1 437 7. 88 7.0 Weed 1 25
2 368 7. 50 ap) Sand 1-3 Zo
3 442 6. 28 he Sand 4 25
4 Soe oOo 625 Weed 3 100
5 487 6. 30 ? Sand 3 200
16 23 Insign 4 Sand 10 500
7 676 9.9 OAD) Sand 2 30
18 5 Insign ? Weed 1 100
19 4 Insign 1 Weed 2 250
10 105 5. 95 8. 0 Weed 1-4 Subsid-
11
1 Insufficient numbers of specimens to ascertain maturity point or significant mean of shell length.
Table 3 shows the results obtained from 11 sample collections taken
from Lithia Spring, Hillsborough County, Fla., in February 1947.
The stations are marked on the map of the spring (fig. 41). Stations
6, 8, 9, and 11 had insufficient numbers of specimens to ascertain a
reliable point at which maturity is reached, or a significant mean of
shell length.
The map has been marked with short arrows, which indicate the
main flow of the current of water coming from the spring and with
iong, broken arrows, which indicate the flow of river water that
breaks across the low land to the north of the spring during flood
periods. The river water, which is 18° C. in contrast to the 25° C.
of the spring water, is fatal to these snails and undoubtedly accounts
for their absence or low numbers at stations 8, 9, and 11. These
last-mentioned stations are well stocked with local Florida snails
(pleurocerids, amnicolids, and planorbids).
The mean shell-length data given in table 3 is of little value in
making a comparison of two populations, and, in fact, can be very
misleading. A look at the population-growth curves of shell length
(fig. 42) will reveal the fallacy of the mean shell length. Most of the
populations give a bimodal curve, as accentuated in population 4. The
first peak is simply an aggregation of immature specimens, a factor
that will vary according to the reproductive cycle of that population
and that will change in value as the young grow to an adult size. In
population 4, maturity is reached at 6.5 mm. and extends up to 12.5
mm. The mean for these adults, which are continually growing, is
about 7.5 mm., yet if the entire population, both immature and adult
specimens, is averaged, the mean must be recorded as 5.5mm. This
difficulty may be avoided in great measure by comparing only adults,
SNAIL HOST OF ORIENTAL LUNG FLUKE—ABBOTT 101
> Vo)
‘ _
7 > > = ‘\ ,
7 7 a is ti
a a ae \
ee oa \
e aie 2 \
Qan Oy Vs A
ee NV
y
|
LITHIA SPRING San ei
| /
/
if
vA
Ficure 41.—Map of Lithia Spring, Fla., showing collecting stations (1 to 11) and water
currents. Short arrows=spring water; long, broken arrows=river currents.
but, again, this is possible only if dissections can be made to determine
whether the specimens in question are adult.
It is interesting to note the distance separating the mode of the
immature curve and that of the adult curve in each population. Were
these distances proportionately the same in each population, we could
safely assume that the production of young was brought about simul-
taneously throughout the various populations of the spring and was
102 PROCEEDINGS OF THE NATIONAL MUSEUM Vou. 102
NO. SPECIMENS
190
$e
STATION
£0
7o
LENGTH OF SHELL (MM.)
Ficure 42.—Population curves based on shell length in three samples from Lithia Spring,
Fla. Circles denote point at which maturity is reached.
due to a seasonal, environmental stimulus. However, the distance
between the “wave” of young and the adults varies considerably from
6 mm. in population 7 to 0 mm. in population 3. The latter popula-
tion, which is unimodal, may represent a population in which the
“wave” of immature has grown into the mature class, and it is not
unlikely that it was bimodal a few weeks before the date of collection,
in the manner of the other populations.
The independent but pronounced production of young in the various
populations suggests a cyclical activity in the physiology of the
population itself, in which a fluctuation of sexual activity is present.
How frequently these waves of young are produced could not be
ascertained from data gathered on one visit to the spring.
McMullen (1947), in his studies on the growth rates of the fresh-
water snail Oncomelania quadrasi Moellendorff in the Philippine
Islands, found similar smaller peaks or waves in the measurements
of colonies. By revisiting his collecting areas periodically he was
able to watch these waves of young move slowly toward the main
peak of the adults. From his data he was able to ascertain that that
species reaches maturity in 4 to5 months. I have shown also (1946)
that similar waves occur in the developmental stages of the eggs of
Oncomelania, and that this was due to increased egg production
brought about by the stimulus of rains and flood periods.
SNAIL HOST OF ORIENTAL LUNG FLUKE—ABBOTT 103
Bronomics or GuAm coLonies.—A natural sequence to the bionomic
study of the Lithia Spring populations was an investigation of the
same species living on Guam Island to see if similar environmental
influences on size existed in areas natural to this species. In making a
statistical study of the 13 colonies collected on Guam Island it was
found that two serious drawbacks existed that, indeed, are common
to any material of this nature. The first was that in most of the
colonies an insufficient number of preserved animals were available
to ascertain at what point maturity was reached. Secondly, the speci-
mens were collected by hand and not in a strictly random fashion, as
would be the case had a sieve or hand net been employed. The natural
tendency of the collector is to choose the largest specimens first, then
the smaller ones, until patience or time halts collecting.
Despite these handicaps, it was felt that a reliable index of size
could be obtained by using the 10 percent of that part of the colony
that represents the largest specimens. This method would be open
to criticism were we dealing with animals which stopped growth at
maturity, and which possessed some distinctive morphological adult
character. When neither of these markers are present, as is the case
with this genus of mollusks, we seek the maximum growth to which
the snails grow and eliminate an abnormally low mean, owing to
high percentage of young. The choice of the 10 percent figure is
arbitrary, although, in population samples of 50 or more, the chances
are that this will include only adults. It is unwise to choose simply
the largest one or two specimens, for it is common knowledge that
abnormal giants or perhaps polyploids are apt to be present in any
population and will not serve as representatives. This is the reason
why, in smaller samples, of 20 or less, it is best to choose the 20 or 30
percent representing the largest specimens, for comparative purposes.
What we are essentially seeking in a study of shell length is the
largest size to which individuals will grow in any one population.
Since growth is continuous throughout the life of the individual, two
factors will delimit the length of shell—genetic and environmental.
We have seen in our study of the Lithia Spring populations that some
environmental factor is most likely responsible for length of shell,
and it is not unreasonable to assume that such is the case in any dif-
ferences found on Guam. On this island the streams are close to each
other, and accidental dispersal by birds and other creatures is likely
to keep the chances of a lengthy genetic isolation very low. To compare
the 10 percent representing the largest specimens of each population
is, in essence, to compare the ecologic conditions of each stream.
In order to test the validity of using the 10 percent referred to of
each colony, histograms were made of the 13 colonies, and the means
of shell length were calculated for the total population and for the top
10 percent. Figure 43 is a summation of this study from colonies A to
104 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
M, with the histograms in the right column and the means, to their
left, represented by straight lines. The upper line is the mean for the
entire population sample, and, the line below, the mean for the top 10
percent. These two lines could never have the same length, but the
nearer they come together, the more natural will be the curve. Any
preponderance of young would lower the mean for the entire popula-
tion. It will be noticed, from a comparison of the proximity of the
two means and the configuration of the histogram, that colonies such
as A, B, E, and K, in which the two means are closest, have more
compact histograms approaching the normal curve. (L is based on
only 8 specimens and cannot be considered significant for this purpose.)
On the other hand, samples D, F, and especially G and J, whose two
means are relatively far apart, possess histograms that are consider-
ably distorted and drawn out by a high percentage of young. A
résumé of the statistics is given in table 4.
TaBLE 4.—Shell lengths of Guam colonies
Mean of
i Number of Mean of Largest
Colony specimens colony eos specimen
A 92 13. 04 15.3 16.5
B 198 13. 44 16. 8 19.0
C 100 15. 58 20. 1 23. 0
D 65 15. 10 22. 6 27.0
E 50 19. 85 24. 0 26.0
¥ 38 18. 44 25. 1 25. 5
G 97 15. 55 25. 4 27.0
H 127 18. 33 27.3 29. 0
I 20 22. 10 29. 2 30. 0
J 43 16. 09 31. 6 33. 5
K 245 24. 46 31. 7 34. 0
L 28. 68 34. 0 35. 5
M 36 26. 41 37.1 40. 0
Although this method of comparing colonies lacks the usefulness
and accuracy of presenting the true nature of each population, i. e.,
the percentage of young, the modes of one or more growth stages, and
so forth, it seems to be the only recourse in problems confronting us
in those invertebrates that show no morphological signs of maturity,
and that grow throughout their life span. In reality, it answers only
the question “how large does this species grow in this environment?”
A map of the southern half of Guam Island has been drawn and is
presented in figure 40. The collecting localities for colonies A to M
have been spotted, and the relative size of the top 10 percent of the
individuals entered in the form of outlined shells. It will be noted
that there is no geographical cline evident in their distribution. The
only possible correlation noted is that between size of the large streams
and size of large shells, but unfortunately insufficient studies of the
SNAIL HOST OF ORIENTAL LUNG FLUKE—ABBOTT 105
COLONY MEANS HISTOGRAMS
Ficure 43.—Means and histograms of population samples from Guam Island (see p. 103
for explanation).
flow of these streams at various seasons were made. It would be
interesting to mark and to transplant snails from, say, Agana Spring
to Umatac River to see if final growth conformed with the latter
environment. A reciprocal transplant would be equally illuminating.
EFFECT OF EROSION ON SHELL LENGTH.—Whenever the water of the
habitat of fresh-water mollusks becomes acid, there is apt to be a
106 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
marked effect on the shells and, in cases where the pH is less than 6.0,
sometimes the death of the entire colony results. Many mollusks have
developed a chitinous periostracum that protects the shell from cor-
rosion. However, the abrasive action of sand particles and the attach-
ment by algae or egg cases of Neritina gastropods may break through
this outer covering and expose the calcium carbonate of the shell to
the acid waters. It is not uncommon for some colonies of Thiara
granifera to be considerably eroded at their spires. Shell-length
measurements, in these cases, can hardly serve as fair comparisons
with colonies that have suffered no loss of shell. The effect of erosion
may be avoided by comparing the width of shells or the height of
the last two or three whorls that have not been affected.
However, it is of equal interest to learn exactly what amount of
reduction in length has been brought about by stream action, for
ecologists may wish to know not only the pH values of various streams
but also what effect acidity is having on the animals. A study was
made of a population sample of 257 specimens of 7hiara granifera
(Naujan River, Mindoro Island, Philippine Islands, U.S.N.M.
258890) which showed considerable erosion. It was found that adults
were reduced in shell length by 11.6 percent. Since this type of study
has not, to our knowledge, been previously used, we have gone into
considerable detail.
]
|
‘
|
|
|
1
4mm. ig MM.
125
{ MM.
25
MM.
Ficure 44.—I. Histogram of total shell length of sample from Mindoro colony, II. Histo-
gram of same shells showing height of last whorl only.
SNAIL HOST OF ORIENTAL LUNG FLUKE—ABBOTT 107
Two histograms (fig. 44) are presented, with the upper one repre-
senting the distribution of individuals based on the measurements of
total length of shell, and with the lower histogram representing the
same individuals in which the height or length of the last whor] has
been measured. Therefore, the upper histogram carries the effect of
erosion, and the lower one is without it. Numerous small individuals
were not worn away at their spires. These were measured for width
and length to ascertain the ratio between these two figures (i. e., the
obesity of the shell). The results are shown in table 5.
TaBLE 5.— Measurements of shells unaffected by erosion
Total length Length of last whorl Total length~length of
(mm.) (mm.) last whorl
—
-
GUS STMT 90 00 9050
MDHNOWDOOMWH
iit iain ini ob ae
on
oo
9 O99 pe pe CUT OUT DD
AWORDUNN OOO
Average ratio 1. 51
In a population sample, the distribution of which approaches the
normal curve, or which consists of a uniform sample of adults, it
would be necessary merely to multiply the mean of the last whorl
measurements by 1.51 (total length/length of last whorl in perfect
specimens) to arrive at a theoretical mean of the total shell length.
Having obtained this figure, which would represent a population size
unaffected by erosion, we need only to compare it with our actual
measurements of eroded specimens to ascertain the amount of
reduction.
This direct conversion could not be applied in this case, however,
for the histogram is strikingly bimodal and in its lower peak possesses
a number of young that were not worn away at the spires. The entire
sample, therefore, was arbitrarily divided into groups A and B. As
natural a midpoint as possible was chosen between the two peaks in
the lower histogram at 6.75 mm. If the erosion-correction factor is
applied to this (1.51 x 6.75), we obtain a theoretical equivalent total
shell length of 10.19. It so happens that this figure divides the two
groups in the upper histogram in exactly the same numerical propor-
tion (i. e., 114 in group A and 142 group B) as we have in the lower
histogram.
108 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
This division into two groups has put all the few perfect specimens
in group A and has left us “adult” and all eroded specimens (except 1)
in group B for comparative studies.
Group A Group B
‘otalinumber! 2200) !) 6.029576) BY tis Gy Bane ey Sts 114 (100%) 142 (100%)
Notierod ede 27) teint Lie tae oa Bab AT 31 (27.1%) 1 (0.7%)
TOG ee it a TN TL Me ai ope LN 83 (72.9%) 141 (99.3%)
The means, their standard deviation, and the standard error of the
means were calculated for the following:
TorauL SHELL LENGTH
Mean é bn N
GroupeA 32 oe ee eee 6. 59 1. 42 0. 13 114
Group Bitte aoe eee Se 12. 46 1. 47 12 142
Lenetu or Last WHORL
GroupvAws 24. Ue ae es ee 4, 40 0. 29 0. 02 114
Group Dale oc 2-25: foe eee 9. 03 1. 04 . 09 142
The correction factor (length/width of perfect specimens) of 1.51
was then applied to the measurements of the last whorls in each
group to compare theoretical total shell length and actual total shell
length.
Reduction
Total Last whorl Last whorl X caused by
length length correction erosion
PA eres 2b NS a ee ne oes ya als Pu Te ll 6. 59 4. 40 6. 64 0. 03
1 6 pe 0 |S ce ie ame RE Type a 12. 46 9. 03 13. 63 ibe ales
It will be noted that the reduction caused by erosion in group A is
extremely small. This is due to the youth of the group, which has
been exposed to erosion for a relatively short time. In fact, 27.1 per-
cent of these specimens were without a trace of spire erosion. In
group B, representing adults that have been exposed to acid waters
for considerable time, the reduction has been 1.17 mm., or 11.6 percent.
PARASITOLOGY
TREMATODE Parasites.—This species of fresh-water snail is of par-
ticular interest to parasitologists because of its ability to serve as an
intermediate host of numerous trematodes, three of which have been
known, in their adult stages, to parasitize man. Other genera and
species of the snail family Thiaridae also have this intermediate host
ability, and their presence in the Orient is responsible for large endemic
areas of the human lung fluke (Paragonimus westermani).
In the course of dissecting specimens from many localities, it was
noted that some colonies in the Philippine Islands were infected as
high as 20 percent with unidentified heterophyidlike cercariae. Dis-
section of specimens from the introduced colonies at Lithia Spring,
Fla., failed to show any trematode infection.
SNAIL HOST OF ORIENTAL LUNG FLUKE—ABBOTT 109
A résumé of the parasites carried by this species is given here. Iden-
tification of the intermediate snail hosts as “Melania obliquegranosa
Smith” is unquestionably referable to Vhiara granifera (Lamarck). I
have examined specimens with the above identification from Formosa
Island and the illustrations published by Japanese parasitologists,
and I find that the implicated Formosan species is a minor race of the
widespread subspecies 7’. g. mauiensis Lea.
Family HETEROPHYIDAE
- HAPLORCHIS TAICHUI (Nishigori, 1924).
—
Geographical distribution—Northern and central Formosa.
Implicated by Faust, E. C. and M. Nishigori, 1926.
Intermediate snail hosts —T hiara granifera (Lamarck) (+ Melania
obliquegranosus (Smith)) and Semisulcospira libertina Gould.
Second intermediate hosts—Fishes: Cyprinus, Gambusia, Caras
stus, Zacco, Pseudorasbora, Phodeus, Ctenopharyngodon.
Definitive hosts—Birds (?), mammals, including man (experimental
infection). An intestinal parasite acquired by eating raw, infected
fish.
“The miracidium which hatches from the egg penetrates the tissues
of the snail to which it is adapted, as in the case of the miracidial
larva of M. taihokui, and metamorphoses into a sporocyst. Rediae are
produced parthenogenetically within these sporocysts, and after a pe-
riod of five to six weeks mature cercariae develop within the rediae.
These carcariae erupt from the snail tissues and are found freely swim-
ming about in the water.” (Faust and Nishigori, 1926.)
2. DIORCHITREMA FORMOSANUM Katsuta, 1932.
Geographical distribution.—Formosa.
Implicated by Katsuta, 1932.
Intermediate snail hosts—Thiara granifera (Lamarck), and
Semisulcospira libertina Gould.
Definitive hosts—Mammals. An intestinal fluke acquired by eating
raw, infected fish.
3. METAGONIMUS YOKOGAWAI Katsurada, 1912.
Geographical distribution —Japan, Korea, Formosa, Rumania, Si-
beria, Dutch East Indies, Palestine, Russia, and Spain.
Intermediate snail hosts—Thiara granifera (Lamarck), Semiscul-
cospira libertina Gould, Hua amurensis Gerstfeld. The record of
“Melania ebenina” in China is probably referable to Hua toucheana
Heude. Unknown in the Philippine Islands and Eurasia.
Second intermediate hosts —Fishes: the salmonoid Plecoglossus al-
tevelis and the cyprinoid Richardsonium.
110 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Family TROGLOTREMATIDAE
4. PARAGONIMUS WESTERMANI Kerbert, 1878.
Geographical distribution.—In the Orient, Japan, Korea, Man-
churia, Formosa, China (especially Chekiang Province), French Indo-
China, the Philippine Islands, Siam, the Malay Peninsula, Assam,
India, New Guinea, Java, and Sumatra. Also South America, prob-
ably Africa, and one record for North America.
Intermediate snail hosts —T hiara granifera (Lamarck) (Formosa),
Thiara (Melanoides) tuberculata Miller (Formosa), Semisulcospira
libertina Gould (Formosa and Japan), Hua species, and Syncera lutea
A. Adams (China). Unknown elsewhere.
Second intermediate hosts——The fresh-water crabs Potamon (eo-
thelphusa) obtusipes Stimpson, P. dehaanii White, and Eriocheir
japonicus De Haan.
Definitive hosts —Lungs of mammals, especially the felines and man.
The life cycle was first elucidated by K. Nakagawa in Formosa in
1917.
“When the snails are placed in water containing miracidia, the
latter swarm around them and become attached to the heads, Jaws and
feet, but rarely to the tentacles and mantles. They cling with their
suckers, insert proboscis into the tissue of the host and enter the body
of the snail like the cercariae of Schistosomum [sic] japonicum, as de-
scribed by Miyairi (1915). Unlike the miracidia of Schistosomum,
those of the pulmonary distoma Paragonimus shed their cilia in this
act.
“Besides the cercariae, sporocysts of various sizes are found abun-
dantly in the liver of Melania [+Zhiara and Semisulcospira]. They
are sometimes found in the heart and kidneys.” (Nakagawa, 1917,
pp. 301-302).
REMARKS ON SNAIL HOST SPECIFICITY.—Gastropod mollusks serve as
the obligatory first intermediate host of all digenetic trematodes or
flukes. While in several cases the relationship between the parasitic
fluke and the host snail is restricted to one species in certain areas, on
the whole there does not appear to be any set pattern for certain snails
to serve as hosts to any particular spe¢ies or even genus of trematode.
Of the three important blood flukes that infect man, Schistosoma
japonicum appears most restricted in its choice of only one genus
of snails, Oncomelania. In the Philippine Islands, Oncomelania
guadrasi Moellendorff is the only known carrier; in Japan, only O.
nosophora Robson. This genus of snails is a gill-breathing amni-
colid. Yet Schistosoma mansoni Sambon and S. haematobium Bil-
harz, of Africa and tropical America, are carried by the snails Bulinus,
Physopsis, and Australorbis, all of which are lung-breathing Planor-
SNAIL HOST OF ORIENTAL LUNG FLUKE—ABBOTT t11
bidae. The phylogenetic relationships of these two groups of gastro-
pods could hardly be farther apart. In all likelihood the choice of
mollusks is based on the physiological adaptations and ecological
preferences of the snails in each case.
Thiara granifera serves as host for two different superfamilies of
trematodes, the Heterophyoidea (Metagonimus, Haplorchis, and Di-
orchitrema) and the Troglotrematoidea (Paragonimus). ‘These same
trematode groups are also carried by other genera of Thiaridae (Semz-
sulcospira, Hua, Goniobasis, and the subgenus of Tiara called Mela-
noides). In addition, however, members of the snail families Amni-
colidae and Synceridae may also act as intermediate hosts, in some
cases being the only snail hosts in the local endemic area.
In all likelihood Thiara granifera may be considered a potential host
cf the American Paragonimus kellicottii on epidemiological and eco-
logical grounds.
An understanding of snail-host specificity will probably not arrive
from a study of molluscan phylogeny, but rather will have to await
an intensive study of the physiology of the many fresh-water species
that serve as hosts. The seeming tendency for certain families of
snails to serve as hosts to particular groups or species of flukes is
probably merely an expression of a common physiological condition
possessed by these snails.
INTRODUCTION INTO THE UNITED STATES
Thiara granifera has established itself some 5,000 miles east of its
normal geographical limits in the Lithia Springs of Hillsborough
County, Fla. The thriving American colonies should be considered as
potential hosts for one or more of the trematodes that they are capable
of carrying in their native habitats. (See under Parasitology, p. 109.)
It appears, however, for the several reasons discussed subsequently,
that no danger exists in the establishment of these snails in our country.
It appears from the information now at hand that there was only
cne introduction of the snail into the United States. On March 23,
1935, an aquarium dealer of San Francisco, Calif., sent four specimens
to the United States National Museum for identification. In a letter
(March 23, 1935) to Dr. G. S. Myers, then curator of the division of
fishes, this dealer states, “I do not know just where their native habitat
might be, because here and there, from different points, Australia,
China, Hawaii, etc., where I get a small shipment, plant life comes
along and some small ones [snails] may be adhering to it.”
No further notice was taken of this species, since at that time the
parasitological importance of this species was overlooked. It was not
until 1947 that it reappeared, when Dr. C. Wythe Cooke, of the U. S.
Geological Survey, collected a number of specimens in Lithia Spring,
iia bY PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Fla., and kindly forwarded them to the United States National Mu-
seum. A few weeks later I was sent to this spring to determine its
prevalence, manner of introduction, and its possible spread.
Thiara granifera is present in the main Lithia Spring in extraordi-
nary numbers, sometimes as many as 400 specimens per square foot.
A small, connecting subsidiary spring contains a few larger individ-
uals. A fuller account of the spring condition appears in the section
on bionomics (p. 100).
An aquatic-plant and fish dealer in Tampa related that he acquired
specimens in 1937 while on a visit to California. Since that time,
his plant vats have been stocked with this mollusk which, as an oddity,
had been sold over a hundred times in a year as the “Philippine horn
of plenty.” It was his opinion that the snails were probably acci-
dentally introduced to the spring around 1940, when improperly
washed tubs were used to gather native plants. I have seen specimens
in home aquaria in Silver Spring, Md., which had been acquired
from Washington, D. C., dealers.
It will be seen by the above experiences that any foreign snail
that has sales value because of its attractive appearance or peculiar
habits will soon be distributed to many parts of the country and in
many cases will be introduced purposely or accidentally to native
waters. Probably the most likely solution to the problem of con-
trolling introduced, dangerous mollusks lies in encouraging dealers,
particularly the importers, to send their mollusks for identification
to museums that have specialists in mollusks on the staff. The present
programs of the Bureau of Plant Quarantine and Insect Control and
the United States Public Health Service appear to be extensive enough
for our protection. It must be realized that even the most stringent
of laws prohibiting the import of foreign mollusks are not going to
offer perfect screening, since accidental introductions by various means
are likely to occur. In the past hundred years approximately 50
species of exotic species of land and fresh-water mollusks have estab-
lished themselves in the United States and Canada.
The geographical origin of these snails will probably remain un-
certain. From the condition and size of individuals in the Lithia
Spring colonies, it appears that an abnormally stunted ecological form
exists, which makes fair comparisons with races from endemic Pacific
areas difficult. The more closely resembling lots in the United States
National Museum are from the Hawaiian and Marianas Islands.
They are possibly members of the subspecies or geographical race
Thiara granifera mauiensis Lea, 1856.
It is highly unlikely that this snail will spread to many other sec-
tions of the United States, except in rare spring localities where water
SNAIL HOST OF ORIENTAL LUNG FLUKE—ABBOTT 113
temperatures remain above 75° F. Its public health menace is negli-
gible by virtue of the complex life cycle of the parasite that it is
capable of carrying. A second intermediate host, usually a fresh-
water crayfish or crab, is a necessary part of the life cycle of
Paragonimus. Furthermore, in order to accomplish infection of the
definitive host the crayfish must be eaten raw, a custom which is infre-
quent or rare among our people.
IDENTIFICATION OF THIARA GRANIFERA AND CLOSELY
RESEMBLING SPECIES IN THE UNITED STATES
In addition to a snail of similar appearance, which is native to
Florida springs, there are two species of thiarid snails sometimes
found in aquaria that are likely to be confused with T’hiara granifera.
Pleurocera (or Ceriphasia?) catenaria (Say) is found in the same
habitat as 7. granifera at Lithia Spring, but rarely succeeds in an
aquarium. Hemisinus cubanianus (Orbigny), from Cuba, and
Pachychilus glaphyrus (Morelet), from Central America (Honduras),
have been raised by fish fanciers with moderate success. ‘The synopses
of distinguishing characters presented herewith will aid in identify-
ing these species.
THIARA (TAREBIA) GRANIFERA (Lamarck)
Figure 45, a; PLATE 8, Ficurss 1, 2
At Lithia Spring: Shell 10 to 15 mm. (about 1% inch) in length,
amber to reddish amber, rarely with green algal attachments. Sides
of whorls in spire flat, without strong spiral cords.
In aquaria: Shell sometimes 20 mm. in length, and if in stagnant
tank, heavily varnished with black coating. Animal with young in
brood pouch under skin of back; mantle edge with tiny, fleshy fingers
or papillae. Operculum with nucleus at one end.
PLEUROCERA CATENARIA (Say)
Fiaure 45, c; PLATE 8, FIGuREs 3, 4
Shell 15 to 20 mm. (about 34 inch) in length, amber brown to black-
brown, often with green algal attachments. Sides of whorls in spire
slightly rounded and carinate near the bottom, with strong spiral
cords, which form well-developed tubercles as they cross the small
axial ribs. Often with dark brown, spiral color band at base of shell.
Mantle of animal wavy, but without papillae. No brood pouch.
Operculum with nucleus near center.
114 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Figure 45.—Animal characters of four genera of Thiaridae: a, Thiara (Tarebia) granifera
(Lamarck), from Lithia Spring, Fla., and the Orient; b, Hemisinus cubanianus (Orbigny),
from Cuba; c, Pleurocera catenaria (Say), from Florida springs; d, Pachychilus glaphyrus
(Morelet), from Honduras. (From left to right: Mantle edge, central radular tooth,
outer marginal tooth, and operculum.)
SE
SNAIL HOST OF ORIENTAL LUNG FLUKE—ABBOTT 115
HEMISINUS CUBANIANUS (Orbigny)
Figure 45, 0
Shell 20 to 25 mm. (about 1 inch) in length, smooth, greenish brown,
with many spiral rows of elongated, black-brown color spots. Animal
with a few, large young in brood pouch under skin or back. Mantle
edge with tiny, fleshy papillae. Operculum with nucleus at one end.
PACHYCHILUS GLAPHYRUS (Morelet)
Fiaure 45, d
Shell 35 to 45 mm. (about 114 to 134 inches) in length, smooth,
black-brown, and without spiral rows of color dots. Animal without
brood pouch. Mantle edge smooth. Operculum with nucleus near
center.
LITERATURE CITED
ABBOTT, R. T.
1946. The egg and breeding habits of Oncomelania quadrasi Mildff., the
schistosomiasis snail of the Philippines. Occ. Papers on Mollusks,
Harvard Univ., vol. 1, No. 6, pp. 41-48, pl. 9.
1948. Handbook of medically important mollusks of the Orient and Western
Pacific. Bull. Mus. Comp. Zool., vol. 100, pp. 245-328, 5 pls., 14 figs.
1948a. A potential snail host of Oriental schistosomiasis in North America
(Pomatiopsis lapidaria). Proc. U. 8. Nat. Mus., vol. 98, pp. 57-68,
pls. 3—4.
Asport, R. T., and HuntTER, G. W., III.
1949. Studies on potential snail hosts of Schistosoma japonicum. I. Notes
on the amnicolid snails Blanfordia, Tricula and a new genus, Fukuia
from Japan. Proc. Helminth. Soc. Washington, vol. 16, No. 2, pp.
73-86, 3 pls.
Bouvier, M. E. I.
1887. Systéme nerveux morphologie generale et classification des gastero-
pods prosobranches. Ann. Sci. Nat. Zool., vol. 3, No. 7, pp. 1-510,
pl. 1-19.
Faust, E. C. and M. NISHIGORI.
1926. The life cycles of two new species of Heterophyidae, parasitic in
mammals and birds. Journ. Parasit., vol. 18, no. 2, pp. 91-128.
McMULLEN, D. B.
1947. The control of Schistosomiasis Japonica. I. Observations on the
habits, ecology and life cycle of Oncomelania quadrasi, the mollus-
can intermediate host of Sohistosoma japonicum in the Philippine
Islands. Amer. Journ. Hygiene, vol. 45, pp. 259-273.
116 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Moorg, J. B.S.
1889. Nassopsis and Bythoceros. Quart. Journ. Micr. Sci., new ser., vol. 17,
pp. 187-201, pl. 20-21.
1899. Tanganyika rufifilosa, and the genus Spekia. Quart. Journ. Micr.
Sci., new ser., vol. 42, pp. 155-185, pls. 20-21.
1898. The anatomy of Typhobias, with a description of the new genus
Bathanalia. Quart. Journ. Micr. Sci., new ser., vol. 41, pp. 181-204,
pl. 11-14.
NAKAGAWA, K.
1917. Human pulmonary distomiasis caused by Padragonimus westermanni.
Journ. Exper. Med., vol. 26, pp. 297-823.
Riccu, BD.
1937. Systematische, Anatomische, Okologische und ‘Tiergeographische
Undersuchungen tiber die Stisswassermollusken Papuasiens und
Melanesiens. Archiv fiir Naturg., new ser., vol. 6, No. 3, pp. 35-153.
SrEsHatya, R. V.
1929. The stomach of Paludomus tanschaurica (Gmelin). Rec. Indian Mus,
vol. 31, pp. 7-12.
1934. Anatomy of Paludomus tanschaurica (Gmelin). Rec. Indian Mus., vol.
82, pt. 2, pp. 185-212, 15 figs.
1940. A free larval stage in the life-history of a fluviatile gasteropod.
Current Science, vol. 9, No. 7, pp. 331-332. Calcutta.
THIELE, J.
1929. Handbuch der systematischen Weichtierkunde, vol. 1, 376 pp. Jena
WENz, W.
1939. Handbuch der Paliozoologie, vol. 6, Gastropoda, part 3: Prosobran-
chia, pp. 481-720. Berlin.
U.S GOVERNMENT PRINTING OFFICE: 1952
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 8
, 2. Shells of Thtara (Terebia) granifera (Lamarck), from Lithia Springs, Fla. (> 5).
, 4. Shells of Pleurocera catenaria (Say), from Lithia Spring, Fla. (> 4).
U S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 9
THIARA (TEREBIA) GRANIFERA (LAMARCK).
Contents of a brood pouch, showing development from egg to young ready to emerge.
(25):
ee
ee
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
issued
SMITHSONIAN INSTITUTION
U. S. NATIONAL. MUSEUM
Vol. 102 Washington : 1952 No. 3293
SOME MARINE ASELLOTE ISOPODS FROM NORTHERN
CALIFORNIA, WITH DESCRIPTIONS OF NINE NEW
SPECIES
By Rosrerr J. Menzies
In THIS report certain of the species of the isopod families Mun-
nidae and Janiridae from northern California are described. The
recorded species were all collected from the intertidal zones, with the
exception of Janiropsis magnocula, which was taken from about 30
fathoms off the California coast. Hitherto only the genus /aniropsis,
represented by two (really one) species, had been known from the
California coast.
In order to eliminate repetition, those characteristics mentioned
in the family diagnoses are not mentioned in the generic diagnoses;
furthermore, characteristics enumerated in the generic diagnoses ap-
ply to the specific diagnoses and descriptions as well.
Tn all instances, the width of specimens is taken at the widest part
of the second peraeonal somite; the length is measured on the mid-
line from the rostrum to the tip of the telson.
Dr. Fenner A. Chace, Jr., curator of the division of marine inver-
tebrates, United States National Museum; Dr. Melville Hatch, Zo-
clogy Department, University of Washington; and Dr. Frank A.
Pitelka, Zoology Department, University of California, have all as-
sisted through the loan of certain specimens utilized in this report.
Dr. Chace and Dr. Hatch kindly lent some type specimens, which
proved very helpful in clearing up the confused concept con-
cerning Janiropsis kincaidi kincaidi. Most Monterey County speci-
mens were lent by Dr. Pitelka. They form part of a fine collection
956648—52——-1 117
118 PROCEEDINGS OF THE NATIONAL MUSEUM you. 102
of isopods, made by John Davis, which was deposited in the Univer-
sity of California collections and subsequently lent me for study.
To these gentlemen I extend my sincere thanks for their kind as-
sistance.
Family MUNNIDAE
(Cf. Nierstrasz and Schuurmans Stekhoven, Jr., 1930, p. X, e 108)
Genus MUNNA Kroyer, 1839
Munna Kr¢gyer, Naturh. Tidsskr., vol. 2, p. 612, 1889. (Genotype, Munna boeckii
Kré¢yer, 1839.)
Generic diagnosis—In general the generic diagnosis is modified
after G. O. Sars (1897-99, p. 106). Body subpyriform, with dorsal
surface somewhat vaulted; last three peraeonal somites small, with
lateral parts directed somewhat posteriorly. Cephalon usually broad.
Pleotelson consisting of two somites: A narrow, short, anterior somite
and a pyriform last somite. Eyes distinct (G. O. Sars, 1897-99) or
Jacking (H. J. Hansen, 1916, p. 34). First antenna with a 3-jointed
peduncle and a flagellum composed of only a few joints. Second
antenna with a 6-jointed peduncle and a multiarticulate flagellum.
Mandible with a well-developed molar process, palp 3-jointed. Maxil-
liped with a 5-jointed palp. Gnathopods of adult male subchelate,
often enlarged and swollen. Remaining peraeopods ambulatory, not
subchelate, increasing in length, with carpal and propodal joints much
elongated and bordered with spinelike 2-pointed setae; dactylar joint
small, biunguiculate. Uropods with one (?) or two branches.
Remarks.—Even today this genus remains inadequately understood
and contains a vast number of species. The genotype, Munna boeckii,
is still imperfectly known, especially in respect to the minute structure
of the mouth parts, uropods, and pleopods. For that reason the di-
agnosis here given is very liberal, and I believe that once the genotype
is better known it will be possible to divide the genus into at least
two, and perhaps more, very distinct genera. It has been possible to
detect on the species that I have examined a minute but clearly dis-
cernible dorsal (outer) uropodal branch, which usually has but a
single seta at its apex, and which generally is concealed partly or totally
by the edge of the pleotelson. This isa rather significant observation,
in that the genus Afunna has long been considered different from the
other genera belonging to the family Munnidae in lacking biramous
uropoda, e. g., “uropoda very small, simple” (G. O. Sars, 1897-99, p.
106); “uropoda small, simple” (Richardson, 1905, p. 480); “mit
kurzen eingliedrigen uropoden” (Vanhdéffen, 1914, p. 561) ; “uropoden
einiistig” Nierstrasz and Schuurmans Stekhoven, Jr. (1930, p. X, e
108) ; “uropoder enkla” (Hult, 1937, p. 9); et cetera.1 It is believed
1 After the preparation of this paper, Dr. Raymond Amar (1948) discovered an additional
uropodal branch in Munna petiti Amar.
MARINE ASELLOTE ISOPODS—MENZIES 119
that the omission of reference to this minute dorsal uropodal branch
has been due to its being overlooked by the several researchers who
have worked on the genus, and that it will in all probability be found
on the other species belonging to the genus. Of course this makes the
differentiation of Munna from related genera a more difficult task, but
at the same time it indicates closer affinities.
In three of the examined species a small but distinct squama was
detected on the third joint of the peduncle of the second antenna.
In a fourth species, I. ubiquita, there was only a slight indication of
such a scale, in the form of a large seta. Reference to this observation,
like that concerning the dorsal uropodal branch, appears to be lacking
in the published description of species of Munna to which I have had
access, and this character appears to have been overlooked. The
squama also indicates a closer affinity between Munna and its related
genera.
As mentioned, it is possible to divide Munna species into several
groups, some of which may become recognized genera when I. boeckii
is better known. The following attempt at a division of the genus,
while obviously incomplete, seems none the less worthwhile:
A. Species having a leaflike, flattened, ventral uropodal branch
lacking spinelike protuberances. Each male first pleopod not ex-
panded laterally at apex, apex pointed. Gnathopods of adult male
and female specimens similar. Male third pleopod with terminal
joint of exopod not equal in width to width of terminal joint of endo-
pod. Probably includes Munna nana Nordenstam, Munna acarina
Miller, Munna petiti Amar, and Munna ubiquita, new species.
B. Species having rounded (in X-section) ventral uropodal branch
bearing at least one large spinelike protuberance. Each male first
pleopod expanded laterally at apex. Gnathopods of adult male en-
larged and swollen, unlike those of adult females and subadult males.
Male third pleopod with terminal joint exceeding in width the width
of terminal joint of endopod. Probably includes Munna avatshensis
Gurjanova, M. kréyeri Goodsir, M. palmata G. O. Sars, W/. stephenseni
Gurjanova, and If. chromatocephala, new species.
C. Species similar to those in B, except ventral uropodal branch ap-
pears to lack any large spinelike protuberance. A distinct, dorsally
visible, dentate, suburopodal shelf is present, which structure is lack-
ing in species mentioned in B. Probably includes Munna minuta
Hansen and Munna halei, new species.
D. Species lacking eyes. Minute structure of mouth parts, pleopods,
and uropods not known. Equals Coecimunna Richardson, 1908, and
includes Munna truncata Richardson and M. acanthifera Hansen.
E. Species to which such assignments are impossible at the present
time, owing to their being inadequately known. Includes a vast
majority of the species.
120 PROCEEDINGS OF THE NATIONAL MUSEUM vou. 102
The four species herein recorded, three of which are new, represent
the first intertidal records for the genus from California.
KEY TO THE CALIFORNIA SPECIES OF MUNNA
a’. Uropods small, leaflike, lacking spinelike protuberances. Male first pleopods
apically. pointed 2222 2 = sas Se atent Sete re eal M. ubiquita, new species
a. Uropods round in X-section. Male first pleopods with apices laterally
expanded.
b*. Uropoda at least with one large spinelike protuberance. Dentate suburo-
podal shelf lacking.
c’. Lateral borders of pleotelson smooth, devoid of large 2-pointed setae.
M. chromatocephala, new species
c’. Each lateral border of pleotelson with 2 to 3 large 2-pointed setae.
M. stephenseni Gurjanova
b*. Uropods lacking large spinelike protuberances. Dentate suburopodal shelf
VISibles ing dorsal svie wees aa eee M. halei, new species
MUNNA UBIQUITA, new species
Ficures 46-48
Munna minuta Hansen, HarcH, 1947, p. 1738, not figs. 42-44,
Holotype-—Male, length 0.8 mm., width 0.4 mm.
Allotype—Ovigerous female, length 1.2 mm., width 0.6 mm.
Diagnosis.—Kyes on fairly long immovable stalks, preorbital lobes
well developed. Pleotelson with about six serrations on each ventro-
lateral margin. Flagellum of first antenna composed of three joints,
last joint slightly shorter than second, which is about twice length of
first, terminal joint with e single sensory filament at apex. Adult male
and female gnathopods similar. Male first pleopod apically pointed,
not laterally expanded. Second male pleopod with apex of exopod
acutely pointed. Suburopodal shelf not evident. Uropodal ventral
branch thin, leaflike, lacking spines.
Character of body.—Minute, mitelike, legs spindly. Patch of black
chromatophores, above uropods and on male pleopods, distinctive.
Antennae and body witn scattered black chromatophores.
Cephalon.—F¥ rontal margin slightly emarginate, devoid of spinelike
setae.
Peraeon.—Epimeral plates evident in dorsal view on somites 2-7.
Pleon.—Pleotelson with few setae and no 2-pointed setae.
Second antenna.—As loug as body. No squama observed, although
a large seta occupies the place usually occupied by squama on third
peduncular article. Flagellum with about 10 separate articles, first
much longer than second.
Maxilliped.—With three coupling hooks.
First mawilla.—Outer lobe with about 11 apical setae, inner lobe
with 38 apical setae.
MARINE ASELLOTE ISOPODS—MENZIES 121
Second mawilla—Each of the two outer lobe lappets with 4 apical
setae. Inner lobe with 11 apical setae and at least 4 pectinate scales
along outer surface.
Mandibles —Left mandible incisive part with 4 teeth, lacinia with
4 teeth, setal row with 3 setae, molar process expanded at denticulate
a g h
Ficure 46.—Munna ubiquita, new species: a, Dorsal view, male paratype; b, apex of
telson; c, seventh peraeopod; d, female operculum; e, uropods, left; f, first maxilla; g,
second maxilla; h, pleotelson. (Magnification as indicated by scale: f and g same as D;
c and d same as fh.)
distalend. Right mandible lacking lacinia, incisor with 5 teeth, setal
row with 4 setae. Terminal article of palp with two apical setae.
Gnathopod.—Carpus with three 2-pointed setae on inner distal
angle, merus with a large 2-pointed seta at outer distal angle. Large
black chromatophores on all articles except dactyl.
Seventh peraeopod.—Seven 2-pointed setae along inferior margin
and four on superior margin of propodal joint. Other peraeopods
similar in a general way to seventh.
122 PROCEEDINGS OF THE NATIONAL MUSEUM vou. 102
Uropod.—Dorsal branch with a single seta, ventral branch with four
setae on lateral margin and two apical setae.
Types.—Holotype, allotype, and 15 female paratypes, collected at
the type locality, Marshalls, Tomales Bay, Marin County, Calif., in
shoreline plankton haul, October 31, 1947, by R. J. Menzies. Type
specimens have been deposited in the collections of the following
institutions :
United States National Museum, holotype (No. 89520), allotype,
and 16 paratypes; Allan Hancock Foundation, 6 paratypes; Rijks-
museum van Natuurlijke Historie, Leiden, Holland, 3 paratypes;
Pacific Marine Station, 29 paratypes, accession numbers 1283-1290
Arth.
Geographical distribution.—Seattle, Wash., to Tomales Bay, Marin
County, Calif.
Material examined Specimens examined were collected from the
following localities:
Wasurnerton : Seattle, Carkeek Park, intertidal, on Polystphonia.
CaxuirorniA : Sonoma County, Bodega Harbor, off Dusty’s pier, No-
vember 29, 1947 (R. J. M.), 1 ovigerous female; Bodega Bay, whistle
buoy, 40 feet, July 14, 1948 (R. J. M.), 1 male. Marin County,
Dillon Beach, First Sled Road, January 21, 1948 (R. J. M.), 8 males,
13 females; Tomales Bluff, Tomales Point, bay side, August 18, 1947
(R. J. M.), 1 specimen; in tide-pool plankton haul, November 29, 1947
(R. J. M.), 3 specimens; Tomales Bay, Marshalls, shore-line plankton
haul, November 13, 1947, 11 specimens, mostly ovigerous females
(R. J. M.).
Hcology.—This species appears to be one of the most abundant
species of intertidal marine isopods at localities near Dillon Beach,
and because of its apparent ability to thrive in vastly different ecologi-
cal conditions I have given it the name wbiquita. Specimens were
found on the Bryozoa Bugula pacifica and Tricellaria occidentalis and
in most tide pools and shore-line plankton hauls. The species appears
intertidally abundant in both protected bay and exposed rock-coast
localities. Specimens taken from Tomales Bay demonstrated a posi-
tive phototropism in the laboratory. Ovigerous specimens were
noted during January, October, and November.
Remarks.—An examination of some paratypes of Munna acarina
Miller (1941, pp. 311-313, fig. 2) indicates that the species is closely
related to Uf. ubtquita. ‘The specimens were kindly lent by Dr. Miller
from his personal collection. Munna nana Nordenstam (1933, pp.
222-295, figs. 56-57) similarly appears closely related to U/. ubiquita;
however, in the paper cited Dr. Nordenstam makes no mention of the
species having 2-branched uropods. In a letter recently received
from Dr. Nordenstam he remarks that he has been unable to find a
dorsal uropodal branch on specimens of Munna nana. Munna ubi-
MARINE ASELLOTE ISOPODS—MENZIES 123
Ficure 47.—Munna ubiquita, new species: a, Terminal articles of seventh peracopod; },
right mandible, distal end; ¢, maxilliped; d, first antenna; e, gnathopod; f, male first
pleopods; g, distal end of left mandible. (Magnification adjacent to a applies to all
figures.)
124 PROCEEDINGS OF THE NATIONAL MUSEUM you. 102
quita differs from both acarina and nana in having minute serrations
along the ventrolateral borders of the pleotelson. Such serrations
appear to be lacking in acarina and nana. I have examined the speci-
mens recorded by Dr. Hatch (1947, p. 173) as Munna minuta Hansen,
and in my opinion they belong to this species.
a
Ficure 48.—Munna ubiquita, new species: a, Third pleopod, male; b, second pleopod, male.
MUNNA STEPHENSENI Gurjanova
Ficures 49-51
Munna stephenseni GuRJANOVA, 1938, p. 88 (in Russian), p. 91 (in English), fig.
15, a-f.
Munna kroyeri Goodsir, Frr, 1926, p. 22—Hatcu, 1947, p. 174 (not the figures
given by Hatch).
The specimens taken in Marin County exhibit sufficient variation to
make identification of them with Gurjanova’s species seem quite rea-
sonable. Those identified by Fee as Munna krgyeri Goodsir in all
MARINE ASELLOTE ISOPODS—MENZIES 125
probability belong to this species, although the description given by
Fee applies equally well to J/. krgyeri or to M. stephenseni in most
details. Fee’s remark that there is one abdominal segment in the
species is probably incorrect, because the genus is characterized in part
by having two pleon somites.
Figure 49.—Munna stephensent Gurjanova: a, Gnathopod, adult male; b, distal portion
of right mandible; c, pleotelson; d, lateral border of pleotelson; ¢, first maxilla; f, ovigerous
female; g, second maxilla; h, right uropod. (Magnification of ¢ the same as a; magnifica-
tion of d and h the same as 5; magnification of g the same as ¢.)
The following descriptive material offers a brief diagnosis of the
species as well as some supplementary notes concerning structures not
mentioned in the original description. The adult male gnathopod is
figured for the first time.
956648—52 2
126 PROCEEDINGS OF THE NATIONAL MUSEUM voL. 102
Diagnosis.—Eyes on short immovable stalks, preorbital lobes strong.
Pleotelson with two to three large 2-pointed setae on each side an-
terior to uropods. Body covered with setae, which are especially evi-
dent at posterior border of pleotelson. Flagellum of first antenna in
adult composed of three articles, last article about one-eighth length
of second and about two-thirds width of that article; second and third
articles each with a sensory filament at apex. No suburopodal shelf
Ficure 50.—Munna stephenseni Gurjanova: a, Third pleopod, male; b, second pleopod,
male. (Magnification as indicated by scale.)
evident. Ventral uropodal branch with a strong posteromedially di-
rected spinelike tooth at posteromedial angle, in addition to several
smaller teeth.
Supplementary descriptive notes:
Pleon.—Small specimens with two spinelike 2-pointed setae on
each side, large specimens with one more such seta on each side.
First antenna.—Gurjanova describes two flagellar segments,’ the
second of which is long, and figures (enlarged drawing) a flagellum
composed of three segments (plus a very minute terminal article), the
long last article of her description being figured as two subequal
joints. In one of the specimens that I examined the flagellum of
the left antenna was composed of two articles, much as Gurjanova
describes (plus the minute terminal article, which Gurjanova did not
consider a separate article), while the right antennal flagellum re-
2 Antenna 1 has a 2-jointed flagellum, with a large long last joint” (Gurjanova, 1933,
p. 91).
MARINE ASELLOTE ISOPODS—MENZIES 197
sembled that figured (enlarged drawing) by Gurjanova. All the
other specimens that were examined had a flagellum similar to that
shown in figure 51, a, that is, one consisting of three articles, including
the minute terminal article.
Ficure 51.—Munna stephenseni Gurjanova: a, First antenna; b, carpus of seventh peraeo-
pod; c, maxilliped; d, female gnathopod; e, male first pleopods; f, left mandible; g, distal
articles of seventh peraeopod; h, left mandible. (Magnification as indicated by scales:
a, d, g, and h same as b; ¢ and f same as e.)
128 PROCEEDINGS OF THE NATIONAL MUSEUM vou. 102
Second antenna.—Basal flagellar segments pink in life.
Maxilliped—With three coupling hooks.
First maxilla—Outer lobe with 12 setae, inner lobe with 4 apical
setae.
Second mawilla—Each of the two outer lobe lappets with 4 apical
setae, inner lobe with about 13 large setae.
Mandible —Left mandible incisor with five teeth, lacinia with four
teeth, setal row with four setae. Right mandible lacking lacinia,
incisor with four teeth, setal row with four setae. Terminal article of
palp with three denticulate setae at apex. Molar process expanded at
apex.
Adult male gnathopod.—With two blunt teeth on cutting edge of
propodus and two longer teeth on cutting edge of carpus.
Subadult male gnathopod—Resembles female gnathopod.
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 Washington = 1952 No. 3294
AUSTRALASIAN STILT-LEGGED FLIES (DIPTERA: TYLI-
DAE) IN THE UNITED STATES NATIONAL MUSEUM
By Georce C. SreyskaL
Cottecrions made during recent years by members of the armed
forces in the course of malaria-control work and similar activities have
added materially to our knowledge concerning the flies of the family
Tylidae in areas not covered in Hennig’s recent (1935-1936) mono-
graph. This and other material in the United States National
Museum were made available to me through the kindness of the au-
thorities of the Smithsonian Institution, and study has led to the
following results:
Eleven new forms are described, for one of which a new genus is
erected. The genus Afimomyrmecia is transferred from the sub-
family Trepidariinae to the Taeniapterinae, and descriptive notes are
given. Examination of material determined as Furybata hexapla has
revealed a compact group of species around that form and the fact
that other species placed in the genus are abundantly distinct. A new
genus, based upon the second species referred to EHurybata, E.
semilauta Osten-Sacken, has been set up for the latter group. Sufli-
cient material was examined to make possible a preliminary study of
the subspecies of Mimegralla albimana, a “Formenkreis” widespread
in the East and Oceania.
Geographical spellings are taken from maps of the National Geo-
graphic Society, especially those of the Pacific Ocean (December
1936) and southeast Asia (October 1944), except that Japanese forms
of names for islands formerly under Japanese mandate are not used.
161
956650-—52———-1
162 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Family TYLIDAE
Subfamily TAENIAPTERINAE
Genus GRAMMICOMYIA Bigot
1. GRAMMICOMYIA HALLI, new species
FIGURE 72, @
Male.—Length 8.5-9.5 mm., exclusive of ensiform hypopygial proc-
ess. Very similar to G. bergi Steyskal (1947, p. 7), differing as
follows: Anterior part of frons bright creamy yellow, somewhat wider
than in G. bergi, pale centrally back to ocellar region; third antennal
joint largely blackish, but brown basally ; anterior hump of mesonotum
rather fiatter than in G. bergi, not rugulose, brown pruinose antero-
medianly. Pruinosity of venter and pleura rather thin. Middle and
hind femora very little swollen centrally, tapering to each end, dark
brown in middle. Hind tibiae black at base, grading into brown in
apical half; middle tibiae wholly black. Copulatory fork as figured.
Wings as in G. bergi. Palpi flat, elongate-oval, with small hairs only,
dark brown, somewhat paler apically.
Types.—Holotype, male, U.S.N.M. No. 58304, and two male para-
types, New Gurvea: Finschhafen, November 9, 1944 (David G. Hall).
Remarks.—The hind femora, swollen centrally, place this form in-
termediate between @. bergi, in which the femora are swollen basally,
and the more typical forms from farther west, which have the hind
femora strongly swollen distad of the middle.
2. GRAMMICOMYIA SONDAICA Hennig
Grammicomyia sondaica HENNIG, Konowia, vol. 14, p. 80, 1935 (Siam, Sumatra,
Java).
Mauay States: Singapore, 1 specimen.
Genus MIMEGRALLA Rondani
3. MIMEGRALLA ALBIMANA (Doleschall)
Taeniaptera albimana DotescHaLt, Nat. Tijdschr. Nederl. Ind., vol. 10, p. 413,
pl. 10, fig. 4, 1856.
Mimegralla albimana (Doleschall) Hennia, Konowia, vol. 14, p. 198, 1935.
The various subspecies of M. albtmana may be rather easily dis-
tinguished from other species by the following combination of charac-
ters: First posterior cell narrowly open; postvertical bristles lacking;
front wholly black (except frequently reddish anteriorly in &. a.
contraria) ; antennae reddish yellow, brownish apically or dorsally
only; hind basitarsi (except extreme base) whitish.
AUSTRALASIAN STILT-LEGGED FLIES—STEYSKAL 163
Ficure 72.—a, Grammicomyia halli, new species, copulatory fork. b, Mimegralla perfulva,
new species, copulatory fork. c,d, Townesa spinosa, new species: c, Copulatory fork,
d, posterior aspect of left foreleg of male. ¢-h, Mimomyrmecia tessellata Frey: e¢, Dorsal
aspect of head, f, ventral aspect of half of copulatory fork, g, lateral aspect of head,
h, lateral aspect of copulatory fork.
164 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
The examination of a considerable amount of material has enabled
me to construct a key, which is presented here in the hope that it will
be found to be an improvement over Hennig’s rather sketchy one,
wherein the subspecies are brought out in several different places
among other species. A map (fig. 73), based upon Hennig’s data and
material examined by me, is also offered as a graphic résumé of the
distribution of the various races.
KEY TO THE SUBSPECIES OF MIMEGRALLA ALBIMANA ?
1. Hind femora yellow, with three or less brown or blackish rings, the third
ring always clearly subapical, the tip of the femora yellow; when the rings
are of some length: Brown with four yellow rings, the fourth covering
the |, tips. 2 ae oe ee 2
Hind femora brown or blackish, with 0 to 3 yellowish rings : Basal, distomedian,
and apical when complete (sometimes a nerrow proximomedian ring divides
the median black section in M. a. striatofasciata) _________-________-__ 4
. Mesopleura with brassy pruinose area posteriorly, which is part of a sterno-
mesopleural band; mesonotum posteriorly usually with a pair of similar but
less distinct longitudinal stripes (Philippine Islands: Taiwan; Ryukyu
Islands; Marianas Islands; Caroline Islands; Palau Islands).
3c. M. a. galbula (Osten-Sacken)
Pruinosity of mesopleura indistinct, not sharply delimited, less distinct than
to
that: of, sternopleurass.22- 2 222. 8. ee ee ee eee 3
3. A reddish interantennal spot present; palpi wholly yellow (Malay States;
IBOENEO! Davia) a2 oe ee ee ee 3a. M. a. albimana (Doleschall)
Interantennal spot lacking; tip of palpi black; femoral rings strong (Celebes,
Halmahera, Amboina, Aroe Islands; ? Ternate).
3b. M. a. sepsoides (Walker)
4. Hind femora unicolorous black (New Hebrides) __-- 3f. M. a. extrema Hennig
Hindafemorawith, yellow rings ee eee 5
5. Basal hind femoral ring about four times width of femur, the rings well-
defined; wings dilute brownish, with a single more distinct crossband in
middle; anterior margin of front frequently reddish (New Guinea).
3d. M. a. contraria (Walker)
Basal hind femoral ring twice or less width of femur___------_____---__-~- 6
6. Wings hyaline with 2 well-defined crossbands (Admiralty Islands; Bismarck
Islands ; Solomon Islands) --------~- 3e. M. a. striatofasciata (Enderlein)
Wings dilute brownish with a single more distinct crossband in middle--__ 7
. Middle and hind femora with a distinct subapical ring, also yellow at tip
below; thorax rather shining above (Samoa; ? Fiji).
3g. M. a. samoana (Czerny)
Middle and hind femora without distinct subapical ring; thoracic dorsum
grayish centrally (Tonga Islands) ----__----- 3h. M. a. tongana (Czerny)
=~]
3a. MIMEGRALLA ALBIMANA ALBIMANA (Doleschall)
Hennig adduces from a study of type material that this, the typical
subspecies, was first described from Java. It is also known from the
Malay States and Borneo. I have not seen specimens.
1M. a. keiensis Hennig is not included; see discussion under M. a. ketensig (p. 167).
165
AUSTRALASIAN STILT-LEGGED FLIES—-STEYSKAL
*(AusazQ) vunduo; *v “py (A *(Aui9z2) vuvowns ‘v “py PB :stuuezy vwayxa vy po O f(urspiepuy) vyo12s0,
~OjvLdis “DB “PW GH F(AoyTeA\) Vtsv4jU0I ‘vy *Y @ {(uayxoeg-uoisQ) vjngq7v3 -v “Py CS ‘(s1oyeAd) Saprosdas -v “pr VW {(jpeyosejoq)
pupmiq “Do “Wy W z([[eyssajoq]) vurwiqzy vipossamtpy jo satoadsqns ay Jo uonnqinsip 3uimoys dep—cy aunog
SE
or 08?) (OM: ( o«/
166 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
3b. MIMEGRALLA ALBIMANA SEPSOIDES (Walker)
Calobata sepsoides WALKER, Proc. Zool. Soc. London, vol. 3, p. 124, 1859.
Mimegralla albimana sepsoides (Walker) HennNic, Konowia, vol. 14, p. 200,
1935.
I have not seen material referable to the subspecies, but Hennig cites
records from Celebes, Amboina, Halmahera, Aroe Islands and
Ternate (2).
3c. MIMEGRALLA ALBIMANA GALBULA (Osten-Sacken)
Calobata galbula OsTEN-SACKEN, Berliner Ent. Zeitschr., vol. 26, p. 202, 1882.—
Swezey, Bernice P. Bishop Mus. Bull. No. 189, p. 198, 1946.
Mimegralla albimana galbula (Osten-Sacken) HrNnNic, Konowia, vol. 14, p. 200.
Mimegralla albimana palauensis Henntc, Konowia, vol. 14, p. 201, 1935 (new
synonym).
Ryukyu Isitanps: Okinawa, 12 specimens. Tarwan: Hokuto, 1
specimen. Puuitippinr Isuanps: Luzon, 10 specimens; Samar, 1 speci-
men; Negros, 1 specimen; Palawan, 2 specimens; Calamian group
(Culion Island), 1 specimen. Martanas Istanps: Guam, 9 speci-
mens; Saipan, 1 specimen. Carottne Istanps: Yap, 1 specimen;
Ulithi Atoll, 7 specimens; Truk, 7 specimens; Nukuoro Atoll, 2 speci-
mens; Ponape, 2 specimens. KaprncaMARANGI, 1 specimen. Panau
Istanps: Peleliu, 1 specimen; Koror, 5 specimens; Babelthuap, 3
specimens.
Hennig (1985, p. 200) cites MZ. a. galbula from the Philippine Is-
lands (Palawan, Mindanao, Luzon) and Taiwan (several localities,
including Lambe Island). Our extra-Philippine material is gener-
ally a little smaller and paler; the dark femoral bands are narrower
or reduced to indistinct spots on the dorsal side of the femora and the
wing pattern is very indistinct; but the Philippine specimens also
vary in these respects and it is deemed unwise to recognize any dis-
tinctions. The Palau Islands material varies enough to bring it well
within the limits of galbula.
3d. MIMEGRALLA ALBIMANA CONTRARIA (Walker)
Calobata contraria WALKER, Proc. Linn. Soc. London, vol. 5, p. 253, 1861.
Mimegralla albimana contraria (Walker) HENNIc, Konowia, vol. 14, p. 201, 1935.
New Guinea: Cyclops Mountains, 5 specimens; Finschhafen, 20
specimens; Hollandia, 4 specimens; Milne Bay, 2 specimens; Nadzab,
Markham River Valley, 3 specimens; Toem, 2 specimens.
The type was from Dorey, and Hennig cites the following localities,
also all in New Guinea: Ramoi; Soron; Lorentz River; Alkmaar;
Biak Island; Rivier Kamp; Simpsonhafen. This large form is one of
the more distinct ones. Most specimens have the anterior margin of
the front reddish, an exceptional character in the J/. albimana circle.
AUSTRALASIAN STILT-LEGGED FLIES—STEYSKAL 167
38e. MIMEGRALLA ALBIMANA STRIATOFASCIATA (Enderlein)
Calobata striatofasciata ENDERLEIN, Arch, fiir Naturg., vol. 88 (A5), p. 184, 1922.
Mimegralla albimana striatofasciata (Hnderlein) HenNnia, Konowia, vol. 14, p.
202, 1935; Stettiner Ent. Zeit., vol. 98, p. 49, 1987—SrrysKat, Occas. Pap.
Mus. Zool. Univ. Michigan, No. 502, p. 9, 1947.—Brre, Occas. Pap. Mus. Zool.
Univ. Michigan, No. 503, pp. 3-8, pls. 1-2, 1947.
ApMIRALTY IsLaAnps: 2 specimens; Los Negros, 1 specimen. Soxo-
MON IsLANDs: Guadalcanal, 17 specimens; Russell Island, 1 specimen;
New Georgia, 3 specimens: Florida Island, 4 specimens; Treasury
Island, 1 specimen; Bougainville Island, 7 specimens.
The type of 1. a. striatofasciata was from Ralum, in the Bismarck
Islands, and Hennig reports the subspecies from Mioko (Duke of
York Island, 1935) and New Britain (1937). Our Admiralty Islands
material is most nearly typical, with distinct wing bands and a strong
hind femoral pattern, including a distinct preapical dark band.
A wide variation exists, however, in the coloration of this form
and although some evidence of a cline toward the east is present,
some of the specimens from Bougainville, Florida, and New Georgia
approach those from the Admiralties.
3f. MIMEGRALLA ALBIMANA EXTREMA Hennig
Mimegralia ailbimana extrema HENNIa, Konowia, vol. 14, p. 203, 1935.
New Hesrwes: Espiritu Santo (Turtle Bay and Segond Channel)
29 specimens.
This form was described from Pentecost Island and Malekula Island,
also in the New Hebrides. It isa large and distinct form.
8g. Mimegralla ALBIMANA SAMOANA (Czerny)
Cyclosphen samoanus CzERNY, Stettiner Ent. Zeit., vol. 93, p. 271, 1932.
Mimegralla albimana samoana (Czerny) HENNIG, Konowia, vol. 14, p. 202, 1935.
Calobata (Neocalobata) deferens MAttocH, Insects of Samoa (British Mus.
Nat. Hist.), pt. 6, fasc. 9, p. 346, Feb. 23, 1935 (new synonym).
Samoa: Tutuila, October 26, 1946, D. G. Hall collector, 1 specimen.
3h. MIMEGRALLA ALBIMANA TONGANA (Czerny)
Cyclosphen tonganus CzERny, Stettiner Ent. Zeit., vol. 93, p. 272, 1932.
Mimegralla albimana tongana (Czerny) HENNIG, Konowia, vol. 14, p. 203, 1935.
I have not seen specimens of this form, which was described from
the Tonga Islands.
$k. MIMEGRALLA ALBIMANA KEIENSIS Hennig
Mimegralla albimana keiensis HENNIeG, Konowia, vol. 14, p. 201, 1935.
Not seen by me. It was described from three female specimens
from the Kai (Kei or Key) Islands. It is likely that it is but a variant
of M. a. sepsoides, which is known from the neighboring Aroe (Aru)
Islands.
168 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
4. MIMEGRALLA BINGHAMI (Enderlein)
Calobata binghami ENDERLEIN, Arch. fiir Naturg., vol. 88, ser. A5, p. 186, 1922.
Mimegralla binghami (Enderlein) HEeNNiIe, Konowia, vol. 14, p. 210, 1935.
Inpra: Assam (Hellgate), September 22, 1943, D. KE. Hardy col-
lector, 3 specimens.
Described from Sikkim.
5. MIMEGRALLA COERULEIFRONS (Macquart)
Calobata coeruleifrons Macquart, Dipteéres exotique nouveaux ou peu connus,
vol. 2, No. 3, p. 24, pl. 33, fig. 2, 1843.
Mimegralla coeruleifrons (Macquart) HENNIG, Konowia, vol. 14, p. 198, 1935.
Matay States: Singapore, 5 specimens. Inp1A: Bengal, Tezgaon,
1 specimen.
A widely distributed species from India and South China to the
Sunda Islands.
6. MIMEGRALLA CEDENS CHRYSOPLEURA (Osten-Sacken)
Calobata cedens WALKER, Proc. Linn. Soc. London, Vol. 1, p. 1385, 1857.
Calobata chrysopleura OSTEN-SACKEN, Berliner Ent. Zeitsechr., vol. 26, p. 201,
1882.
Mimegralla cedens chrysopleura (Osten-Sacken) Hennic, Konowia, vol. 14,
p. 205, 1935.
Pumirrint Isutanps: Luzon, Mount Maquiling, 3 specimens.
Known only from the Philippine Islands; the typical subspecies
is from Borneo.
6a. MIMEGRALLA CEDENS THAIENSIS Cresson
Mimegralia thaiensis Cresson, Trans. Amer. Ent. Soc., vol. 52, p. 269, 1926.
Mimegralla niveimana Cresson, Trans. Amer. Ent. Soc., vol. 52, p. 269, 1926.
Mimegralla cedens thaiensis Cresson, HENNia, Konowia, vol. 14, p. 204, 1935.
Hennig lists the above synonymy of J/. niveimana with a query.
I have compared male paratypes of J/. thaiensis with the male type
of I. niveimana, and I feel that I can confirm the synonymy. All the
specimens are from the same locality in lower Siam. The copulatory
fork of this form differs somewhat from that of A/. cedens chryso-
pleura.
7. MIMEGRALLA CONFINIS (Walker)
Calobata confinis WALKER, Proc. Linn. Soc. London, vol. 1, p. 37, 1857.
Mimegraila conjinis (Walker) HENNIa, Konowia, vol. 14, p. 211, 1935.
PuiiierIne Isuanps: Luzon (Mount Maquiling) 3 specimens; Min-
danao (Davao), 1 specimen.
A widely distributed species already known in the Philippines from
Luzon and Basilan Island.
AUSTRALASIAN STILT-LEGGED FLIES—STEYSKAL 169
8. MIMEGRALLA CONTINGENS CONTINGENS (Walker)
Calobata contingens WALKER, Proc. Linn. Soe. London, vol. 7, p. 221, 1864.
Mimegralla contingens (Walker) EnprerteIn, Arch. fiir Naturg., vol. 88 (A5),
p. 195, 1922.—Hernnie, Konowia, vol. 14, p. 90, 1935.
New Guinea: Nadzab, 2 specimens.
Already known from several localities in New Guinea, as well as
from Misoél and Halmahera. Other subspecies are known from Ter-
nate (M. c. lunaria), northern Australia (1. c. australica), and the
Solomon Islands (J. c. solomonis).
8a. MIMEGRALLA CONTINGENS NOVAEHEBRIDEANA, new subspecies
Female.—Length 10 mm. Preapical white fascia of wing very
broad, in middle a little wider than first posterior cell; wing veins
R, and R, ending slightly apicad of tp; mesofrons with distinct
tomentum; base and apex of middle and hind femora yellowish, apex
of hind femora for about one-fifth length of femora with traces of a
preapical dark ring, but yellowish color is rather dark and not sharply
delimited; middle and hind tibiae and tarsi wholly black.
Remarks.—This race has the tomentose mesofrons of J. ¢. luwnaria
and If. c. salomonis. 'The former, however, has a very narrow pre-
apical wing fascia and the tips of the middle and hind femora but
little paler, while from the latter it may be distinguished by the
wholly black middle and hind tibae and strong dark wing fascia.
Types.—Holotype and one paratype, U.S.N.M. No. 58305, New
Hesrwwes: Espiritu Santo Island, Nav. Med. School Ser. No. 18/34,
sp. 20; two paratypes, same locality, Segond Channel, August 29,
19438, Laffoon collector.
9. MIMEGRALLA PERFULVA, new species
FIgukeE 72, b
Male and female.—Length of body 8-10 mm.; length of wings 6.6-
“7mm. Color brownish yellow, except as further noted. Head of
general body color, including mesofrons; ocellar triangle blackish;
third antennal joint, except at extreme base, black; antennal grooves
shining black with whitish pruinose patch below; palpi black in api-
cal third. Posterior frontal bristle anterior to anterior ocellus by
length of ocellar triangle; two well-developed anterior frontals;
arista blackish, brown at base, toward base with a few short hairs.
Postvertical bristles lacking.
Thorax with presutural lateral border of mesonotum rather broadly
dark brown, pruinose, and with traces of median longitudinal dark
956650—52——-2
{|
170 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
margins postsuturally; only one notopleural bristle, situated in pos-
terior corner of notopleural area; scutellum dark brown laterally.
Abdomen very dark brown, but somewhat paler basally and api-
cally. Copulatory fork figured is yellowish with black mesal spinules.
Legs of general body color; tips of femora black, on fore femora
for distance equal to twice the width, on middle and hind femora for
distance equal to width, an indistinct darker ring at apical third of
femora. Fore tibiae black except at base, middle and hind tibiae
dark brown. Fore tarsi black, except apical half of basitarsi and
basal two-thirds of second joint, which parts are pale yellow; middle
and hind tarsi brown, blackish apically, the hairs black, except part
of the plantar brush.
Wings uniformly tinged with hight brown; the basal crossvein its
own length apicad of vein closing the anal cell; first posterior cell
narrowly open; second vein ending 0.36 to nearly 0.5 the distance
from tp to apex of wing. Knobs of halteres gray-brown.
Ty pes.—Holotype, male, U.S.N.M. No. 58981, Carortinr IsLanps:
Ponape (Mount Dolennankap), 1,700 to 2,000 feet, August 10, 1946
(H. K. Townes No. 1648) ; allotype, female, same locality, 1,800 feet,
August 11, 1946 (H. K. Townes No. 1651) ; paratypes, four males, five
females, same data as allotype; one female, same data as holotype.
Remarks.—Since this species runs in Hennig’s key (1935, p. 83) to
Mimegralla leucopeza, it may be a subspecies of that form. However,
the general yellowish color is more extensive than in any race of J.
leucopeza, and the wide separation from the nearest station of that
species (M. 1. albitarsis, Manila, Philippine Islands) also makes
doubtful its identity with leucopeza.
10. MIMEGRALLA PONAPENSIS PONAPENSIS Hennig
Mimegralla ponapensis HENNIG, Konowia, vol. 14, p. 207, 1935.
CaROLINE Istanps: Ponape Island, Colonia, 29 specimens.
10a. MIMEGRALLA PONAPENSIS KUSAIEANA, new subspecies
Male and female—Middle femora yellow at base for distance equal
to twice the diameter of the part, then black to 0.6 the whole length,
then yellow for about 1.5 times the diameter and finally brown, but
yellow again apicoventrally. The hind femora are yellow for 3.5
diameters, then black to 0.6 the whole length, the yellow portion with
a preapical brown section.
The middle and hind tibiae, which in Mf. p. ponapensis are practi-
cally wholly black or sometimes a little brownish distomedially, in
M. p. kusaieana are yellowish except at each end.
AUSTRALASIAN STILT-LEGGED FLIES—STEYSKAL 171
The whole insect is otherwise as in the typical race, the thoracic
dorsum of both races largely brownish pruinose, the antennae and
palpi blackish apically, and the frons reddish anteriorly.
Types.—Holotype male, allotype female, and three each male and
female paratypes, U.S.N.M. No. 58982, Carotinr Istanps: Kusaie
Island (Lele), August 21, 1946 (1. K. Townes No. 1815) ; paratypes,
four each males and females, same locality, August 19, 1946 (Oakley
No. 1726) ; four females, Kusaie Island (Mount Tafeyat), 500 to 800
feet, August 2, 1946 (H. K. Townes No. 1812) ; five males, three fe-
males, Majuro Atoll (Majuro Village), August 28, 1946 (H. K.
Townes Nos. 1989, 1992).
Genus MIMOMYRMECIA Frey
11. MIMOMYRMECIA TESSELLATA Frey
FIGurRE 72, e-h
Mimomyrmecia tessellata Frey, Not. Ent., vol. 7, p. 75, 1927—Bryan, Proc.
Hawaiian Ent. Soc., vol. 8, p. 34, 1932.—HENNIe, Konowia, vol. 14, p. 309, 1935.
Puirrine Istanps: Luzon (Mount Maquiling), 7 specimens;
(Mount Banahao), 2 specimens; (Los Banos), 2 specimens; (Limay,
Bataan), 1 specimen; (Manila), 1 specimen; Negros Occidentale
(Victoria) 1 specimen.
Examination of the above material has revealed that the genus is
definitely taeniapterine, rather than trepidariine, as Hennig, in the
lack of material, presumed it tobe. It will run in my key to the genera
of African and Oriental Taeniapterinae (1947, p. 6) to couplet 3, the
first three characters of the first alternative suit. The arista, how-
ever, is plumose. The aflinities seem to lie definitely with @Grammi-
comyia, but the development of the back of the head into three pro-
tuberances distinguishes Mimomyrmecia from any other Paleotropical
genus.
TOWNESA, new genus
Genotype.—T ownesa spinosa, new species.
Very similar to Mimegralla, to which genus it will run in my key
(1947, p. 6), agreeing in all particulars. The type of Mimegralla, M.
coeruleifrons (Macquart), as well as all other species of Mimegralla
known to me, has no armature of any kind on the fore femora.
Townesa, however, has a row of 8 to 12 strong bristles nearly as long
as the diameter of the femora along the whole length of each lower
side of the fore femora of the male, and 3 to 5 similar but somewhat
smaller bristles on each side of the apical half of the fore femora of
the female. The legs are very long and uniformly slender; the first
172 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
posterior cell of the wing is very narrow apically but not closed; the
second vein exceeds the level of tp by the length of tp, postvertical
bristles are lacking; two well-developed anterior frontals are present
definitely within the tomentose mesofrontal area; the posterior
frontals are about opposite the anterior ocellus; and the bristles of the
mesonotum include 1 de, 2 sa, and 2 strong ntpl. The arista is dis-
tinctly and abundantly short plumose.
Remarks.—I take pleasure in naming this genus in honor of the
discoverer of its type species, Dr. H. K. Townes.
12. TOWNESA SPINOSA, new species
FIGURE (2, C, d
Male and female—Length of body 8 to 9 mm.; length of wings 6
to6.8mm. Color of body black.
Head with mesofrons black-tomentose; parafrontals shining black
with bluish reflection; medifacies brown with black lower margin;
clypeus, palpi, and antennae brown, the antennae with third joint black
above and in apical half and with arista black apically, brown at base,
and furnished with short plumosity.
Thorax lightly greenish-brown-pruinose on pleura and sternum,
dark-brown-pruinose on notum, except broad sublateral greenish sub-
shining stripes, which meet anteriorly across the anterior hump. Post-
alar calli shining brownish.
Abdomen black, dark-brown-pruinose with gray incisures, except
segment bearing the copulatory fork and those posterior to it, which
are shining. Copulatory fork as figured (fig. 72, c), bearing dense,
rather long dark brown hairs ventrally and mesally, also with denser
and shorter black bristles mesally.
Legs brownish yellow; fore femora of male as in figure 72, d, the
shaded parts black, in female similar but with fewer bristles (cf.
generic diagnosis above) ; fore tibiae black; fore tarsi light brown
basally, apically blackish; middle and hind femora with extreme apex
black, preapical narrow ring of black, and indistinct distomedian ring
or dorsal spot; middle and hind tibiae dark brown, their tarsi yellow
basally, brown apically.
Wings evenly light brown; ¢a interstitial with vein closing anal cell.
Halteres with yellow stem, blackish knob.
Types.—Holotype and four paratypes, males, U.S.N.M. No. 58983,
Caroutne Istanps: Kusaie Island (Mount Tafeyit), 500 to 800 feet,
August 20, 1946 (H. K. Townes No. 1810) ; allotype and one paratype,
females, same locality, August 2, 1946 (H. K. Townes No. 1812) ; para-
types, three males, Kusaie (Lele Island), August 19, 1946 (Oakley
No. 1726).
AUSTRALASIAN STILT-LEGGED FLIES
STEYSKAL 173
Subfamily TREPIDARIINAE
KEY TO THE AUSTRALASIAN GENERA OF TREPIDARIINAE ?
1. Upper part of metanotum conical and surpassing scutellum,
Nestima Osten-Sacken
Metanotumuar Ordinary-cLonnmee = 2s we She ay ae a ee Pee eat eee ese 2
2. Middle and hind femora on lower side before the tip with a series of spinules_ 3
Middle‘and hind) femora’ unarmed below 202. 222s ee ees 4
3. Wings strongly fasciate or tessellate with dark brown; mesofrons shining
medially, flat and strongly angulate apically______ Eurybata Osten-Sacken
Wings with but indistinct pattern; mesofrons tomentose, strongly convex in
frone andetrunestese se Bese Se ee se eS senses Crosa, new genus
4. Seutellum turned upward; head spheroidal; three fronto-orbital bristles.
Gongylocephala Czerny
Scutellum horizontal above; head somewhat depressed ; wing with third costal
-
division more than half as long as ultimate section of fourth vein--____- 5
5. Middle and hind femora thickened basally; arista plumose.
Crepidochaetus Enderlein
Middle and hind femora slender; arista bare to plumose.
Trepidarioides Frey
Cothornobata Czerny
Metopochetus Enderlein
Of the last three genera I have seen only one representative, 7'rep7-
darioides territa Osten-Sacken, and I am unable to find good distinc-
tive characters in the literature.
Genus EURYBATA Osten-Sacken
On examining a series of specimens included in Lurybata hexapla
it became evident to me that Osten-Sacken was correct when, in deserib-
ing the second species of Lurybata, EF’. semilauta, he stated that later
authors might see fit to place it in another genus. Hurybata may be
distinguished from Crosa, new genus (p. 176), which is based upon £/.
semilauta, as in the foregoing key, by the characters of the head rep-
resented in figure 74, a, b, and in the male by a lack of long hairs on the
basal ventral side of the fore femora. The abdomen is shining apically
and near the segmental sutures.
It was also evident that two new species were included with £.
hexapla. These species are included in the following key to the species
of the restricted genus Hurybata. All species are known only from
the Philippine Islands.
2The genus Mimomyrmecia Frey has been removed from this subfamily (see p. 171).
The genera Formicosepsis De Meijere and Cypselosoma Hendel have been referred here
by Hennig (1941), although Hennig has later (in litt. Sabrosky, 1949) indicated that
Cypselosoma is a clusiid. These two genera may be distinguished from all other Tylidae
by the possession of strong vibrissae and ocellar bristles. Calycopteryr Eaton, from the
Kerguelen and Heard Islands, a fly with rudimentary wings, has also been referred here
by Hennig (1934).
174 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
KEY TO SPECIES OF THE GENUS EURYBATA OSTEN-SACKEN
1. Dark fasciae of wing uniform and straight; fore tibiae whitish; mesonotum
wholly shining or very indistinctly pruinose; male: copulatory fork with
two long divergent prongs (fig. 74, e) _--__-------_ E. hexapla Osten-Sacken
The wing fascia distad of tp (fifth from base of wing) projecting basad be-
tween third and fourth veins or ail fasciae broken, forming a checkered
PUGS T a HS yee se Ee Ee Re EN ase Vel ce eA LO Sen 2
2. Only the fifth fascia uneven; fore tibiae blackish except at tips; mesonotum
with distinct prescutellar brown-pruinose spot; male: copulatory fork with
short arms inflexed at ends (figs. 74 f, g) 15_---__ E. nigritibia, new species
Wings with checkered pattern; fore tibiae only a little brownish distomedi-
ally; mesonotum wholly shining; male: copulatory fork with short arms,
knobbed at tip and slightly convergent (fig. 74, h) -_. E. tessellata, new species
13. EURYBATA HEXAPLA Osten-Sacken
Ficure 74, a, b, e€
Hurybata hetapla OSTEN-SACKEN, Berliner Ent. Zeitschr., vol. 26, p. 206, 1882.—
BerEzzI, Philippine Journ. Sci., vol. 2, ser. 3D, p. 155, 1917.—Frey, Not. Ent.,
vol. 7%, p. 7, 1927.
Puitiepine Istanps: Luzon (Mount Maquiling), Baker collector, 12
specimens; (Ube Laguna), May 12, 1924, R. C. McGregor collector, 2
specimens; Tayabas (Malinao), Baker collector, 1 specimen.
Bezzi lists Los Banos and Mount Maquiling as localities, probably
from part of the same material that is still in the Museum, but I saw
no specimens from Los Bajfios, a town at the foot of Mount Maquiling;
Frey lists (Mount) Banahao (Tayabas) and Surigao in Mindanao.
The latter locality is the source of the type of FZ. tessellata, new species.
Hennig did not see specimens.
The following characters will assist in distinguishing #’. hexapla
from the new species: Arista short plumose on barely the basal half;
third antennal joint 2.5 times as long as broad, blackish except at
base; fore legs yellow, except apical half of femur and apical joints
of tarsi; middle and hind femora with brownish preapical band; one
sternopleural bristle; one posterior notopleural bristle; one dorso-
central bristle; wing fasciae all even and straight; pterostigma very
small; apical wing spot as long as width of first posterior cell at base
of spot; copulatory fork as figured (fig. 74, e).
14. EURYBATA NIGRITIBIA, new species
FIGuRE 74, f. g
Male—Length, 10 mm. Differs from /. hexapla in having yellow-
ish brown antennae, the third joint only about 1.5 times as long as
broad; the shining mesofrontal wedge extending as a narrow strip
all the way to the anterior ocellus; fore femora colored similarly to
AUSTRALASIAN STILT-LEGGED FLIES—STEYSKAL 175
h
A
aw,
aman i
Ficure 74.—a, b, e, Eurybaia hexapla Osten-Sacken: a, Dorsal aspect of frons and vertex;
b, lateral aspect of head; ¢, copulatory fork. c, d, Crosa semilauta (Osten-Sacken):
¢, Dorsal aspect of frons and vertex; d, lateral aspect of head. f, g, Eurybata nigritibia, new
species: f, Copulatory fork, ventral aspect; g, copulatory fork, lateral aspect. h, Eurybata
iessellata, new species, copulatory fork. i, Crosa yapensis, new species, copulatory fork.
176 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 102
FE. hexapla, but the tibiae are blackish except for a short distance at
tip and the tarsi are brown from tip of basitarsi; all tarsi apically
darkened; mesonotum with a heavily brown-pruinose roundish patch
behind transverse suture, a band before scutellum, and one at mesal
corner of notopleural area.
Wing fascia beyond tp (fifth from base of wing) extending basad
between third and fourth veins, making the band twice as wide at
this point; two sternopleurals; one strong posterior and one small
anterior notopleural bristle; apical wing spot longer than width of
first posterior cell at base of spot; copulatory fork as in figures 74,
hig.
Type.—Holotype, male, U.S.N.M. No. 58302, Puitrerrnr Isnanps:
Luzon (Ube Laguna), May 12, 1929, R. C. McGregor collector.
15. EURYBATA TESSELLATA, new species
FIGURE 74, h
Male and female—Length, 10 mm. (exclusive of ovipositor in
female). Antennae blackish except at base, twice as long as wide;
the shining mesofrontal wedge very rapidly running out into a short
point not more than halfway to ocellus; fore femora blackish except
short distance at base; fore tibiae a little brownish apically; all tarsi
light brown apically; mesonotum without distinctly set-off pruinose
areas; wings with the transverse fasciae, especially in middle of wing,
broken into a series of alternating squarish blotches; a hyaline spot
anterior to tp lying directly posterior to a brown blotch in submarginal
cell; marginal cell wholly brown; apical wing spot indistinct; two
subequal notopleural bristles; two sternopleurals; copulatory fork of
male as in figure 74, A.
Types.—Holotype, male, U.S.N.M. No. 58303, PHILtppIng IsLanps:
Mindanao (Surigao), Baker collector; allotype, female, Island of
Basilan, Baker collector.
CROSA, new genus
Genotype.—Eurybata semilauta Osten-Sacken.
Distinguished from related genera as in the foregoing key, as well
as by the following characters: Mesofrons (fig. 74, c) wholly dull,
vertical in front, not projecting, parafrontals ending bluntly; ocelli
well foreward; one dorsocentral bristle; pterostigma very small; ab-
domen wholly pruinose; male fore femora with many long loose hairs
on basal ventral side.
It is possible that with the exception of the preceding three species
all those heretofore included in H'urybata should find their place here.
AUSTRALASIAN STILT-LEGGED FLIES—STEYSKAL Cli
Crosa is dedicated to that master dipterologist, Carl Robert von
Osten-Sacken, and the name is formed from the initial letters of his
name.
16. CROSA SEMILAUTA (Osten-Sacken), new combination
FiGurRE 74, ¢c, d
Burybata semilauta OSTEN-SAcKEN, Berliner Ent. Zeitschr., vol. 26, p. 207,
1882.—F rey, Not. Ent., vol. 7, p. 76, 1927.—HENNIe, Konowia, vol. 14, p. 302,
1935.
PHiipPPiIneE Istanns: Luzon ( Los Banos; Mount Maquiling) ; Min-
danao (Zamboanga) ; Tayabas (Lucban) ; Samar (Osmefia).
Previously recorded from localities in Luzon, Mindanao, and
Samar.
17. CROSA TETRAS (Steyskal), new combination
Hurybata tetras STEYSKAL, Occas. Pap. Mus. Zool. Univ. Michigan, No. 502, p. 2,
1947.
The type material, from the Solomon Islands, is evidently con-
generic with C. semilauta.
18. CROSA YAPENSIS, new species
FIGURE 74, 1
Male.—Length of body, 6.5 mm.; length of wings, 4.5 mm. Color
yellow, except extreme tip of femora and tibiae, dorsum of abdomen
and anterior part of parafrontal stripes, which parts are brown.
There is a faint preapical band of brown on fore femora and the
apical tarsal joints are a little darkened. Mesofrons golden brown,
becoming blackish in ocellar region.
The entire insect is shining or subshining except mesofrons, frontal
orbits anterior to parafrontal stripes, antennae, and a heavily tomen-
tose pleural stripe. Anterior frontal orbits narrowly golden tomen-
tose. Pleural stripe bright white when viewed very obliquely, but
otherwise dull lead colored; it covers the narrow lower margin of
propleura and continues across mesopleura as a stripe as wide as fore
tibiae. ‘The stripe, were it to continue across the sternopleura, would
run into the uppermost two posterior sternopleural setae. Para-
frontal stripes with a bluish reflection.
Head very similar to that of C. semélauta (fig. 74, c, d), but aristal
hairs only half as long as in that species.
Wings clear hyaline with brownish-yellow veins. Third and fourth
veins converge to within a distance equal to length of ta. Pterostigma
obsolete.
178 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 102
Fore femora furnished below with loose yellowish hairs, at base of
femora twice as long as width of femur, but rapidly becoming shorter
until they give out at mid-femur.
Copulatory fork in figure 74, 7, the right arm apparently broken
off.
Type.—Holotype, male, U. S. N. M. No. 58984, Carorinz IsLanps:
Yap Island, near Yaptown, July 14, 1946 (H. K. Townes No. 1239).
Remarks—This species is apparently near Hurybata cuneifrons
De Meijere and £. petasibarba Enderlein, as keyed in Hennig (1935,
p. 801), but the “wedge-shaped spot on the hind margin of the meso-
pleura” is here a slender stripe. The pale general color and the
wholly hyaline wings are distinctive.
Genus GONGYLOCEPHALA Czerny
19. GONGYLOCEPHALA PALLIDA LUZONICA, new subspecies
Gongylocephala pallida StrysKat, Occas. Pap. Mus. Zool. Univ. Michigan, No.
502, p. 4, 1947.
Male and female.—Subspecies agrees with G. pallida (Guadalcanal,
Solomon Islands), except that base of fore tibiae is not brownish, but
tibiae are uniformly yellow; middle tibiae are uniformly dark brown,
not grading to yellow basally; fore femora are a little darker than
tibiae in well-preserved specimens and hind tibiae are a little darkened
basally; whitish fascia of wing is quite straight; male copulatory fork
as in G. pallida.
Types.—Holotype male, allotype, female, and four male and two
female paratypes, U.S.N.M. No. 58306, Puiiprinr IsLanps: Luzon
(Mount Maquiling), Baker collector.
Genus NESTIMA Osten-Sacken
20. NESTIMA POLITA Osten-Sacken
Nestima polita OSTEN-SAcKkEN, Ann. Mus. Genova, vol. 16, p. 458, 1880.—HEnNnNIG,
Konowia, vol. 14, p. 308, 1935.
New Guinea: Nadzab (Markham River Valley), May 4, 1944
(K. V. Krombein), one female.
Although the two known species of Westima, VN. polita Osten-
Sacken, and NV. prolixa (Walker), are both inadequately described,
I believe that the above specimen agrees well enough with Osten-
Sacken’s description to make the determination. There must remain
some doubt as to the identity of the following forms, described as
new, but I feel certain that they are specifically distinct.
AUSTRALASIAN STILT-LEGGED FLIES—STEYSKAL 179
21. NESTIMA PLEURALIS, new species
Female.—Length 10 mm., exclusive of ovipositor. Similar to Osten-
Sacken’s description of WV. polita, although the thorax is not “uni-
formly dark brown, shining,” but moderately pruinose on dorsum
with a pitchy brown, largely polished band of equal width extending
all around thorax, including entire pteropleura, crossing notopleural
suture, and including neck, humeri, and anterior face of anterior
hump of mesonotum. Below this band on pleura of each side is a
heavy stripe of appressed white hair, very narrow above fore coxae
end gradually widening to include entire hypopleura from middle
coxal to pteropleural sutures. There are several whitish sternopleural
bristles and 1+2 notopleurals. Lower part of thorax is yellow.
Fore trochanters are black; fore femora yellow at base and at tip
for a distance equal to three times the width, preceded by a blackish
band of approximately same length; fore tibiae blackish except at
extreme base; and fore tarsi whitish except two brown apical joints.
Middle and hind legs have tips of tibiae and entire tarsi black.
Halteres with brown stem and yellow knob.
Type—Holotype, female, U.S.N.M. No. 58308, New Gurnea:
Nadzab (Markham River Valley), May 14, 1944 (K. V. Krombein).
22. NESTIMA VIRIDINSULA, new species
Female.—Length 10 mm., exclusive of ovipositor. Very similar
to NV. polita and N. pleuralis. Thorax dark brown, moderately prui-
nose on dorsum, blackish in a humeral stripe on each side of anterior
hump. Mesopleura with pitchy, shining stripe in upper part, taper-
ing and becoming evanescent forward. No trace of pruinosity or
hair on pleura anterior to hyposternopleural suture. Several whitish
sternopleural bristles and 1+2 notopleurals.
Fore legs yellow; tibiae light brown on most of length, yellow
basally and whitish apically; tarsi whitish, but brownish on apical
two or three joints; middle and hind legs yellow, except black tips of
tibiae and entire tarsi.
Halteres with brown stem and yellow knob.
Type.—Holotype, female, U.S.N.M. No. 58307, Bismarck Isuanps:
Green Island, 1944 (W. G. Downs).
Genus TREPIDARIOIDES Frey
23. TREPIDARIOIDES TERRITA (Osten-Sacken)
Calobata territa OsTEN-SACKEN, Berliner Ent. Zeitschr., vol. 26, p. 203, 1882.
Trepidarioides territa (Osten-Sacken) Frey, Not. Ent., vol. 7, p. 76, 1927.—
HENNIG, Konowia, vol. 14, p. 307, 1935.
180 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
PuierPine Istanps: Mindanao (Surigao); Luzon (Mount Ma-
quiling; Laguna).
The copulatory fork of the male of this species, type of the genus
Trepidarioides Frey, is a simple U-shaped organ, as narrow at base
as the arms are wide, unadorned, and quite different from the organ
figured by Hennig (1935, p. 306) for 7’. eyanea (Hendel).
LITERATURE CITED
Hennia, W.
1934. Ueber Bau und Verwandtschaft der Kerguelenfliege Calycopteryx
moseleyi Hat. Zool. Anz., vol. 108, pp. 196-201.
1935. Revision der Tyliden (Dipt., Acalypt.). IJ, Teil: Die ausserameri-
kanischen Taeniapterinae, die Trepidariinae und Tylinae. Allge-
meines tiber die Tyliden. Konowia, vol. 14, pp. 68-92, 192-216,
289-310.
1941. Formicosepsis de Meijere und Cypselosoma Hendel, zwei Gattungen
der Tyliden. Stettiner Ent. Zeit., vol. 102, pp. 129-131.
STEYSKAL, G. C.
1947. Micropezidae (Diptera) from the Solomon Islands. Occas. Pap. Mus.
Zool. Univ. Michigan, No. 502, pp. 1-9.
U3. GUVERNMENT PRINTING OFFICE: 1952
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
issued {6
” en Je" i, y)
Cres)
nih NCTOR
Shere:
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 Washington: 1952 No. 3295
APHOTAENIUS, A NEW GENUS OF DUNG BEETLE
(COLEOPTERA: SCARABAEIDAE)
By O. L. Cartwricut
A new South American beetle, which I discovered as I studied the
collection of Aphodiinae (Coleoptera: Scarabaeidae) in the United
States National Museum, has certain characters, especially those of
the hind tibiae, that are shared by Ataenius carolinus Van Dyke.
These peculiar characters, which have precluded the satisfactory in-
clusion of carolinus in any known genus of the Aphodiinae, would
seem to indicate the advisability of setting up a new genus for the
reception of both species.
APHOTAENIUS, new genus
General shape elongate oval, very convex, lateral edges of elytra
and pronotum posteriorly not visible from directly above. Head
moderately large, smooth except for a band of coarse punctures across
occiput. Clypeus shallowly emarginate, finely reflexed margin
flattened and dorsally angulately widened between two widely placed
marginal teeth or denticles, when viewed from directly above the
widened margin appearing as an angulate carina and hiding extreme
lower edge, which has a much smaller median angulation directed in
the opposite direction. Genae bent downward as in Ataenius.
Pronotum not crenate, not fimbriate, posterior angles truncate-obtuse,
slightly depressed. Pygidium as in Ataenius, the anterior basal por-
tion with a longitudinal groove receiving the ventrally dentate tips
of the elytra, the exposed apical portion with a depressed central area.
Middle and posterior tibiae with distinct oblique carinae as in Apho-
956657—52 181
182 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
dius but without the apical fringe of spinules, the apical margin
ventrally with two well-separated triangular teeth, each tooth with
a very fine hairlike seta basally on each side, the outer apical angle
spinelike as in Ataentus. Mouthparts as in Ataendus.
Genotype.—Ataenius carolinus Van Dyke=Aphotaenius carolinus
(Van Dyke).
A photaenius, though having distinct oblique carinae on the middle
and hind tibiae as in Aphodius, should be placed in the tribe Eupariina,
since the head, genae, pygidium, and other parts are much closer to
Ataenius and allied genera than to Aphodius. The middle and hind
tibiae are unique.
APHOTAENIUS CAROLINUS (Van Dyke)
Ataenius carolinus VAN Dyker, Pan-Pacific Ent., vol. 14, p. 157, 1928.
Aphodius carolinus Hinton, Ann. Mag. Nat. Hist., ser. 10, vol. 20, p. 196, 1937.
Piceous black, shining, head anteriorly and legs reddish, antennae
and palpi testaceous. Head three-fourths as wide as pronotum,
strongly convex, basally with a band of coarse punctures, front finely
punctate, anterior smooth, impunctate; clypeus with finely reflexed
margin, bidentate, teeth strongly developed, acutely pointed and
widely separated, slightly emarginate between teeth, margin flattened
and dorsally angulately widened, this appearing as an angulate carina
from directly above, the extreme ventral margin, invisible from above,
with a small median angulation directed in the opposite direction.
Pronotum convex, margined basally and laterally, surface with mixed
coarse and very fine punctures, coarse punctures uniformly distrib-
uted, separated by less than one to two diameters. Elytra convex,
coarsely striate, striae strongly crenate punctate, intervals moderately
convex, minutely punctate. Covered basal portion of pygidium
deeply grooved under tips of elytra, exposed portion with depressed
eroded area divided by a longitudinal carina. Posterior prosternal
spine strongly developed, laterally compressed. Mesosternum closely
moderately punctate. Metasternum quite closely coarsely punctate,
median longitudinal line deep. Second abdominal segment closely
shallowly setigerously punctate along anterior margin, carinate be-
tween the coxae; remaining segments smooth, coarsely deeply crenate
in front. Posterior femoral line obsolete, only faintly indicated, mid-
dle femoral line and anterior ventral line of profemur wide, deep, and
noticeably alutaceous. Anterior tibiae tridentate externally. Middle
and posterior tibiae with oblique carinae as in Aphodius, without
fringe of spicules apically but with two well-separated triangular
teeth, each tooth with a fine hairlike seta basally on each side, the
outer apical angles prolonged, spinelike as in Ataenius, long spur,
first tarsal joint, and three following tarsal joints combined equal in
APHOTAENIUS, A NEW DUNG BETTLE—CARTWRIGHT 183
length. Terminal tarsal joint about as long as two preceding com-
bined, claws minute. Length, 2.75 mm.
Holotype.—In California Academy of Sciences, No. 2549.
Type locality —Black Mountains, N. C.
Remarks.— A photaenius carolinus is a common species in deer drop-
pings in June in Pisgah Forest, near Brevard, N. C., and was taken
there as early as April 29 and as late as September 10. It has been
taken in sheep droppings on Sassafras Mountain, S. C.; in rather old
cow dung in woods near Pine Mountain, Ga.; and in the same in
V’On Swamp, Fair Lawn Plantation, 7 miles from Awendaw, near
Charleston, S. C. Mark Robinson and I collected several specimens
at the last-named place on June 4, 1948. These were the first seen from
other than mountain localities, except possibly a single specimen in
the M. A. Cazier collection bearing an old and very doubtful label,
“Key West, Fla.” Recently a single specimen was found in an early
private collection of H. S. Barber, which had been determined by
E. A. Schwarz as “Aphodius n. sp.” This specimen bears label data
as follows: “Marlboro, Md. May 13. Collection H. S. Barber.” It
was probably collected between 1898 and 1901.
Still more recently I found a specimen in the Blatchley collection
at Purdue University. This specimen of Aphotaenius carolinus was
collected in Crawford County, Ind., May 19, 1908, by W. 5. Blatchley
and was erroneously reported as Ataenius lecontet Harold in his
“Catalogue of the Coleoptera in Indiana.”
APHOTAENIUS COLOMBIENSIS, new species
Piceous black, shining, anterior margin of head and thorax and legs
reddish, antennae and palpi testaceous. Head about three-fourths
as wide as pronotum, strongly convex, basally with a band of close
moderate punctures, front with more widely spaced fine to minute
punctures; clypeus slightly depressed and emarginate at middle with
a low angulation or denticle each side, lateral margins finely reflexed,
genae bent down as in Ataenius, margin between denticles flattened
and dorsally angulately widened to twice the height of the denticles,
this appearing as a carina from directly above, lower edge of widened
margin, invisible from above, with a small median angulation in
the opposite direction. Pronotum convex, laterally and basally mar-
gined, not fimbriate, anterior angles obtusely rounded, posterior angles
truncate-obtuse, slightly depressed, base distinctly lobed medially,
surface with mixed fine and coarse punctures, the latter generally
distributed but closer laterally, separated by one to four or five diam-
eters on the disc. Elytra convex, deeply striate, striae coarsely
crenately punctate, intervals weakly convex, with scattered minute
punctures. Mesosternum closely moderately punctate. Metaster-
184 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
num with scattered very fine punctures at middle, a few coarse shallow
punctures and alutaceous sculpture at sides, median longitudinal line
moderately impressed. Second abdominal segment carinate between
coxae, closely, shallowly, setigerously punctate anteriorly, remaining
segments smooth, coarsely deeply crenate anteriorly. Covered portion
of pygidium longitudinally grooved to receive tips of elytra, which
lock together with tongue-and-groove sutural edges, disc of exposed
portion depressed but smooth, with a few scattered very small low
tubercles basally. Anterior ventral profemoral line and posterior fem-
oral line of middle femurs entire, wide, moderately deep and alutace-
ous. Posterior femurs smooth, with scattered minute punctures, with-
out femoral line. Anterior tibiae externally tridentate. Middle and
hind tibiae with well-developed oblique carinae as in Aphodius, ter-
minally without fringe of spinules but with two widely separated
triangular teeth on ventral apical edge, each tooth with a fine hairlike
seta basally on each side, outer apical angle prolonged, spinelike as
in Ataentus. Long spur slightly shorter than first tarsal joint, which
is as long as the three following joints combined, tarsal claws minute.
Sex not determined but probably a male. Length 4 mm.; width 1.75
mm.
Holotype.—UvU. 8S. N. M. No. 59444.
The unique specimen of colombiensis was collected at 2,900 meters,
above Guasca, Cundinamarca, Colombia, March 10, 1942, by Dr. E. A.
Chapin.
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
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SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 Washington: 1952 No. 32%
PRELIMINARY ANALYSIS OF THE VERTEBRATE FOSSIL
FAUNA OF THE BOYSEN RESERVOIR AREA
By Tueovors FE. Wuitre
As a part of the salvage program of the River Basin Surveys, a
cooperative project between the Smithsonian Institution, the National
Park Service, the Bureau of Reclamation, and the Corps of Engineers,
Department of the Army, in the prospective reservoir sites in the Mis-
souri Valley, the Boysen Reservoir area near Shoshoni, Wyo., has been
prospected for vertebrate fossils for parts of two seasons. During the
first period, from October 23 to November 7, 1947, I worked alone, and
considerable time was lost because of early snows. The area was again
worked, with the aid of John C. Donohoe, a student at Montana State
College, and Ernest L. Lundelius, a student at the University of Texas,
from June 4 to July 12, 1948. Although the specimens have not been
credited to individuals, I wish to state that these men have proved
themselves competent collectors, and we three found about equal
amounts of material. Although it is planned to visit this area for as
many seasons as possible before the reservoir is flooded, it seems de-
sirable to make the information gathered to date available to other
paleontologists.
185
956647—52——-1
186 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
SYSTEMATIC DESCRIPTION OF FOSSIL VERTEBRATES
Class REPTILIA
Order SQQUAMATA
Suborder SERPENTES
Family BOIDAE
Genus BOAVUS Marsh
BOAVUS cf. OCCIDENTALIS Marsh
About 30 associated thoracic vertebrae (loc. No. 48FR78);1 2
thoracic vertebrae (loc. No. 48F R80).
Although there is considerable difference in size between the two
specimens, I am inclined to be extremely cautious about differenti-
ating species of snakes on the size of the vertebrae only, since age is not
readily reflected in the surface texture of the bone. Consequently, the
principal importance of this material is the presence of this genus in
the Lost Cabin faunal zone of the Wind River formation.
Suborder SAURIA
Family VARANIDAE
Genus SANIWA Leidy
SANIWA sp.
One dorsal and five caudal vertebrae (loc. No. 48FR65) ; two dorsal
vertebrae of presumably a young individual (loc. No. 48FR78) ; one
caudal vertebra (loc. No. 48F R80).
This material is too imperfect for more than generic identification
and its value is only that it establishes this genus in these deposits.
Family ANGUIDAE
Genus GLYPTOSAURUS Marsh
GLYPTOSAURUS DONOHOETI, new species
FIGURE 75
Type.—U.S.N.M. No. 18316 (fig. 75), a badly damaged skull lack-
ing the tip of the snout, both maxillae, and the right temporal region
(loc. No. 48F R65).
Referred material—U.S.N.M. No. 18317, skull and jaw fragments
with scutes (loc. No. 48F R65).
1¥or a description of localities see pp. 203-206.
BOYSEN RESERVOIR VERTEBRATE FOSSILS—-WHITE 187
Horizon and locality Lower Eocene, Lost Cabin, NE1%{SW1,{, sec.
5, T. 4 N., R. 6 E., of Wind River meridian; White Hill, south side of
Cottonwood (Dry Muddy) Creek, 11 miles north-northwest (air line)
of Shoshoni, Fremont County, Wyo.
Diagnosis —A medium-sized species; interorbital breadth 33 per-
cent less than in @. Aéilst Gilmore; interorbital area with 5 regular
alternating rows of bony scutes, supraorbital and median rows larger
‘~
Ficure 75.—Glyptosaurus donohoet, new species, type, U.S.N.M. No. 18316; squamation of
dorsal surface of skull, X 1. ise,
than second and fourth rows; scutes raised into a boss as in rugosus
and nodosus, vertical diameter of orbit equal to interorbital breadth ;
scutes of the temporal region less regular in outline and about twice
the diameter of those of the interorbital area.
Discussion.—This specimen, in conjunction with a braincase from
Pipestone Springs (U.S.N.M. No. 18805), permits a few additions to
Gilmore’s (1928, 1938) discussions of the genus.
The various elements that make up the braincase are securely fused,
as in Peltosaurus. The former location of some of the sutures can be
made out by lines of roughened bone. The condyle is elongate-oval in
outline, twice as broad as deep. The tubera basioccipitalia project
ventrolaterally from the basicranial axis and are expanded anteriorly
and posteriorly at their bases as though reinforced by flying but-
tresses. ‘These expansions are thickened along their edges so that the
188 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
tubera are triradiate from their terminations, with the median portion
the heaviest. ‘The foramen for the twelfth cranial nerve is located
beside the condyle and below the paroccipital process, at the termina-
tion of the posterior wing of the tubera basioccipitalia. The ninth
and tenth nerves exit through a dorsoventrally elongated foramen at
the ventral side of the jugular groove a little posterior to the median
portion of the tubera basioccipitalia. The fenestra ovale lies just above
this foramen. The foramen for the exit of the venus capitis lateralis
lies just above the anterior termination of the anterior wing of the
tubera at the bottom of the jugular groove. The foramen for the
hyoid branch of the seventh nerve lies at the top of the jugular groove
slightly posterior to the foramen for the venus capitis lateralis. A
thin, fairly deep ridge of bone extends downward from the paroc-
cipital process of the prodtic so that the jugular groove is partially
enclosed laterally.
The region of the hypophyseal fontanelle is so badly damaged in
both specimens that reliable data cannot be obtained. The basiptery-
goid processes of the basisphenoid are elongate and flattened as in most
Sauria. They are separated from the tubera basioccipitalia by a deep
notch, which extends to the main body of the basisphenoid.
The anterior edge of the prodtic is damaged in both specimens, but
enough of this region is preserved in U.S.N.M. No. 18316 to indicate
that the ossification of the prefacial commissure very nearly or en-
tirely encircled the facialis branch of the seventh nerve as it left the
braincase.
A fragment of the maxilla in U.S.N.M. No. 18317 shows that the
anterior maxillary teeth are much smaller than the posterior teeth.
They increase rapidly in size to the fifth tooth, which is as large as
the remainder.
The collection of Glyptosaurus material in the United States
National Museum, which contains most of the types, was examined
in connection with this material. Many of the species were founded
on the characters of the frontal and interorbital regions only, and as
yet some of the species are known only from the type specimens.
Although the taxonomy of a genus based on such a limited portion of
an animal leaves much to be desired, it is possible to make a morpho-
logical grouping of the species of this genus by means of the characters
presented by this region of the skull. Only with the aid of better
material can the validity of this grouping be determined. The known
species are tentatively grouped as follows:
I. Interorbital region with four rows of osseus scutes with one or two odd
seutes interpolated between the median rows__--------- G. montanus group
IY. Interorbital region with five regular, alternating rows of osseus scutes.
G. hillsi group
BOYSEN RESERVOIR VERTEBRATE FOSSILS—-WHITE 189
IlI. Interorbital region with six irregular rows of scutes; odd scutes may or
may not be present between the median rows___-_____---_ G. giganteus group
Vee ronal recion UNENOWNe == ae see ee ee ee eee ee G. sphenodon
TABLE 1.—Stratigraphical distribution of the Glyptosaurus montanus, G. hillsi,
and G. giganteus groups
Periods Stages montanus group hillst group giganteus group
Whitneyian
Oligocene | Orellan + giganbens
Chadronian montanus i tuberculatus
Duchesnian
Uintan R x
Eocene Bridgerian Be cee eee | brevidens nodosus 7
princeps sylvestris
rUgosUus
Huerf@no BB’ || °° halletar we? CR SP aie ae ae
Huerfano A i hillst?
Wasatchian | obtusidens donohoet
Class MAMMALIA
Order INSECTIVORA
Family DELTATHERIDIIDAE
Genus DIDELPHODUS Cope
DIDELPHODUS VENTANUS Matthew
Fiaure 76
U.S.N.M. No. 18369 (fig. 76), a badly crushed skull with left P?
to M° and both lower jaws from which all the teeth have been broken
(loc. No. 48F R65); U.S.N.M. No. 18483, fragment of left mandible
with M, (loc. No. 48F R80).
If the skull is correctly referred to this form its characters are
sufficiently distinctive to warrant designation as a separate species,
rather than citation as a mutation of D. absarokae. The distinctive
characters presented by the teeth of this specimen are : P? two-rooted,
anterior and posterior cingula well developed and with minute
erenulations; P* submolariform and differs from P* only in being
slightly smaller; M‘ exhibits several minute tubercles on the external
cingulum between the parastyles and metastyles. Other characters
190 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
of the teeth agree with Matthew’s (1918, p. 583) figures of D.
absarokae.
Although this skull is very badly broken and crushed, it adds a
few details to our knowledge of the genus. Because /ctops is relatively
well known, comparisons will be made with it, although the two
forms are not closely related:
(1) The frontonasal suture les a short distance in front of the
orbit. (2) The zygoma is a little heavier than in Jctops. (8) Post-
orbital process is short but very well defined. A companion process
was not observed on the fragment of the zygoma preserved. (4) The
orbit appears to be as large relatively as in Zetops. (5) The sagittal
crest is single and moderately high. (6) The parietal foramen appears
to lie closer to the crest than to the squamosal. (7) Squamosoparietal
Ficure 76.—Didelphodus ventanus Matthew, U.S.N.M. No. 18369; occlusal view of left
P?-M3, X4.
foramina were not observed. (8) The union of the mastoid portion
of the petrosal and the squamosal appears to have been similar to
that in Jctops. (9) The mastoid appears to form as much of the
occiput as in Jetops. (10) The relationship of the glenoid to the
periotic suggests that the postglenoid and posttympanic processes
of the squamosal were separated by a meatal notch, although these
processes were broken away. (11) The tympanic ridge of the ali-
sphenoid is lacking, but there is a short one on the glenoid portion
of the squamosal. (12) The foramina in the alisphenoid appear to
have been much the same as in Jetops, but this region is badly crushed
and difficult of interpretation. (13) The alisphenoid appears to be
fused to the basisphenoid. (14) The periotic appears to be rather
large judged by the dimensions of the skull that can be observed.
(15) The inferior border of the massateric fossa is sharply defined
by an abrupt indentation as in Deltatheridium.
MEASUREMENTS OF TEETH OF DIDELPHODUS VENTANUS (IN MILLIMETERS)
Length Width
By AB pee mete Np pA ee ye eg LN - V3 Ou eee
AVENE BE he hn 9S DRT TO ey tok Bat oh fd Rd a SHOU Ss ha Saas
Miastenia Ae pee es les eee ey EEG ae ae ee ee Le) ite ees
PPG 0s 2 pearl SUNS Ale gop ph SSP PRUE AS 2 cals een PA 2
1 BN RR Oy eGR Aaa hy SG ON ea WIM EVAR ES or 2.8 one
NPA epee en LAs tee Me ce elie Al ed 8 ee eee 2.6 3. 4
TY a eat eet pl ly iby agra pret PM Pde A clo 3. 0 S10)
I Bea RA tes et Mae Ek tc 0 a a 2. 4 ah §5)
IIS eee Os A ae oe Nene Seat at le Rian tna eee 1.7 3. 0
>
BOYSEN RESERVOIR VERTEBRATE FOSSILS—-WHITE 19]
The dentition of this specimen is distinctly more advanced than
that of D. absarokae of the Gray Bull. Unfortunately the upper
dentition of this genus is unknown from the Alkalai Creek exposures.
Consequently it is impossible to evaluate the stratigraphic significance
of this specimen.
Family MIXODECTIDAE
Genus CYNODONTOMYS Cope
CYNODONTOMYS SCOTTIANUS COPE
U.S.N.M. No. 18436, fragment of left mandible with posterior
half of P, and M,-; (loc. No. 48F R76) ; U.S.N.M. No. 18434, frag-
ment of right mandible with M,-; (loc. No. 48F R80).
The limited material pertaining to this species does not permit any
additions to Matthew’s (1915c, pp. 470-477) discussion of the genus.
CYNODONTOMYS LUNDELIUSI, new species
FIGURE 77
Holotype—U.S.N.M. No. 18371 (fig. 77), fragment of a right
mandible with posterior half of M,, M,, posterior half of Ms, and the
roots of P;., (loc. No. 48F R65).
Figure 77.—Cynodontomys lundeliusi, new species, type, U.S.N.M. No. 18371; occlusal
view of teeth and lateral view of right mandible, X 2.
Horizon and locality—Lower Eocene, Lost Cabin. NW14SW14,
sec. 5, T. 4 N., R. 6 E. of Wind River meridian, south side of Cotton-
wood (Dry Muddy) Creek, 11 miles (air line) north-northwest of
Shoshoni, Fremont County, Wyo.
Diagnosis —Size large, 33 percent larger than the average for
C. scottianus (Matthew, 1915c, p. 471); M; relatively shorter than
in that species; heel of M, narrower than on M.; paraconid on M,
distinct ; external and posterior cingula as in @. scottianus.
192 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Discussion — Although the teeth in this specimen are broken and
badly worn, the characters presented, especially the size, are distinct
enough for the species to be easily recognized.
MEASUREMENTS OF TEETH OF CYNODONTOMYS LUNDELIUSI (IN MILLIMETERS)
Length
y= Migs 62 Bo er ee 30. 7
PEM, 2222 ee ee ee ee ee ee ee Dawe
Moy (estimated) =. ee ee ee 5. 0
My 225222 cous ee Be ee ee eee 5.5
Mie AS 00 2 RD eR A SSE ee eee 6.5
Depth of jaw at. Mauna 2) ee ee oo eee 12.8
Depth:of jaw atiMy-- 2s}. 228 a Se ee 130
Order TILLODONTIA
Family TILLOTHERIIDAE
Genus ESTHONYX Cope
ESTHONYX ACUTIDENS Cope
U.S.N.M. No. 18267, fragment of right mandible with I,-; and P;-M,
(loc. No. 48FR78) ; U.S.N.M. No. 18470, fragments of both mandibles
(loc. No. 48F R65); U.S.N.M. No. 18469, skull and jaw fragments
(loc. No. 48F R80).
This material is being studied by Dr. C. L. Gazin and will be dis-
cussed in his revision of the order.
Order PRIMATES
Family ADAPIDAE
Genus NOTHARCTUS Leidy
NOTHARCTUS VENTICOLUS Osborn
U.S.N.M. No. 18487, left mandible with M,-3, roots of P3-4, and
alveoli of P,.. (loc. No. 48F R77).
This specimen does not add anything to our knowledge of the species.
Family APATEMYIDAE
Genus TEILHARDELLA Jepson
TEILHARDELLA sp.
U.S.N.M. No. 18488, right mandible with only the incisor (loc. No.
48FR80).
This specimen is provisionally referred to this genus on the charac-
ters of the mandible, which exhibit a number of differences from the
genotype, but these differences cannot be properly appraised until the
dentition is known. The characters exhibited by this specimen are:
BOYSEN RESERVOIR VERTEBRATE FOSSILS—-WHITE 1938
P; procumbent ; P, with a single large root; M, and M, with posterior
root the larger ; posterior root of M, very long and narrow ; massateric
fossa very deep and broad.
MEASUREMENTS OF ALVEOLI OF TEILHARDELLA sp. (IN MILLIMETERS)
Length
PM, --- — = Le =e 8
1 a ae eee ae Seen a = ee Ou
Vie ae ee a a ee oe Reh = a alae
Va ee ee ae oe ee eR eS = a = ile
Mae eS Ss Dh
Family ANAPTOMORPHIDAE
Genus LOVEINA Simpson
LOVEINA ZEPHYRI Simpson
U.S.N.M. No. 18439, portion of a left mandible with part of P,, the
base of M,, and M. and M; (loc. No. 48F R76).
This specimen is provisionally referred to this species on the basis
that M, and M; agree with those of ? LZ. vespertina (Matthew) better
than with those of any other genus. It differs from that species in
the proportionally shorter M, and M,, in the broader trigonid, and in
the presence of a minute entoconid on the heel of M;. Since M, and
M; are unknown in the genotype any attempt at comparison of the two
specimens would be futile.
MEASURMENTS OF TEETH OF LOVEINA ZEPHYRI (IN MILLIMETERS)
Width (at base)
Length Trigonid Teel
IVIg ee es ees s - 7. 4 cee 0 ues
Migs Seo 2 & e 2.3 ? 1.8
1 8, See ene os ea 2.3 2.0 Dae,
Nig western fre ere See Lee tede 2.7 1.8 6
Order TAENIODONTA
Family STYLINODONTIDAE
Genus STYLINODON Marsh
STYLINODON CYLINDRIFER (Cope)
U.S.N.M. No. 18440, portion of right canine (loc. No. 48F R76).
This specimen is referred to S. cylindrifer on the basis of the dis-
tribution of the enamel, which is in two bands, one on each side of
the tooth. It is of uniform thickness and width and shows the obso-
lete vertical striation and the stronger transverse growth lines which
Cope (1884, p. 192) describes for the type of the species. The cement,
which covers the areas between the enamel bands, overlaps the enamel
for a short distance on each side, but there is no evidence that the
bands were covered.
956647—52
9
194 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
MEASUREMENTS OF CANINE OF STYLINODON CYLINDRIFER (IN MILLIMETERS)
Digmeter (transverse) Lee Ee ee ee eee 10.5
Diameter! (anteroposterion)= een ee eee 1a be 5
Width between enamel bands:
PAM CLL O Dee sees ae ee ee ee ee ee 2.8
Posterior 22 22) ee eee ee ee 4.8
Order RODENTIA
Family ISCHYROMYIDAE
Genus PARAMYS Leidy
PARAMYS MAJOR Loomis
U.S.N.M. No. 18442, right mandibular fragment with M,» and roots
of M; (loc. No. 48F R76) ; U.S.N.M. No. 18441, left mandibular frag-
ment with P,—M, (loc. No. 48F R80).
This material does not permit anything to be added to Matthew’s
(1918, p. 614) discussion of the species.
PARAMYS MURINUS Matthew
U.S.N.M. No. 18443, right mandible with M,. and roots of P, (loc.
No. 48F R80).
This specimen does not agree with the figures (Matthew, 1918,
p. 617) of the type in that the enamel is entirely smooth and not
rugose. Consequently, it is only provisionally referred to this species
pending the acquisition of better material.
Order CARNIVORA
Family HYAENODONTIDAE
Genus PROLIMNOCYON Matthew
PROLIMNOCYON ANTIQUUS Matthew
U.S.N.M. No. 18444, left mandible with P;_, and roots of M2, (loc.
No. 48F R76) ; U.S.N.M. No. 19445, both mandibles with only roots
of teeth preserved (loc. No. 48F R65).
This material does not permit anything to be added to Matthew’s
(1915a, p. 70) discussion of the species.
Genus SINOPA Leidy
SINOPA STRENUA (Cope)
U.S.N.M. No. 18446, mandibular fragments with M,-; of both sides
and associated skeletal fragments (loc. No. 48F R77).
This specimen does not permit the addition of anything to
Matthew’s (1915a, p. 74) discussion of the species.
BOYSEN RESERVOIR VERTEBRATE FOSSILS—-WHITE 195
Genus DIDYMICTIS Cope
DIDYMICTIS ALTIDENS Cope
U.S.N.M. No. 18447, skull with calvarium and occiput eroded away,
right and left P'"-M? present (loc. No. 48F R75).
This specimen differs from the one figured by Matthew (1915a, p.
23) in that P* is 2-rooted, and there is no diastema between it and the
canine. The parastyle on M? is better developed and the internal
cinglum is continuous. M? has a greater transverse diameter for its
length and is more advanced.
MEASUREMENTS OF TEETH OF DIDYMICTIS ALTIDENS (IN MILLIMETERS)
Length Width
Pe Mig ee) ee Fe eS poe eee ny OO TE) ee ae
2 SIN es Re ee eee Re a eee eY POO OE «AP eee
Mun eens ae eas eee ay She Js gees Si! 9.0 17.0
Met! ee ee mee (OLE eee te eA alt heed BONO 9. 0
Genus MIACIS Cope
MIACIS ef. LATIDENS Matthew
U.S.N.M. No. 18448, right maxillary fragment with M'*, roots of
P*, and associated skull fragments (loc. No. 48F R80).
This specimen is intermediate in size between the types of J/.
exiguus and latidens (Matthew, 1915a, p. 33-35). It agrees with the
former in the extended parastyle cn the upper molars and with the
latter in that the paracone is much larger than the metacone. ‘The
internal cingulum is interrupted medially below the protocone. Mat-
thew (1915a, p. 33) states that it is continuous in both species but the
illustrations show it to be the same as in this specimen. Although
M' shows considerable wear, there is a suggestion of a small hypocone,
and this specimen may be prophetic of Jf. parvivorus of the Lower
Bridger.
MEASUREMENT OF TEETH OF MIACIS CF. LATIDENS (IN MILLIMETERS)
Length Width
aN pee A ae EN Se eet eee SR eT Re Bea GSD we eee
| ei Sse ae a Bs, ee kee ee Si Nees ao, Vat al GS 3 egy hi i Mapied eeete
ites = Ba snk ee ee ee ee ee ee 6.0 8.4
IN gee yea ere ares ee cee eee eee eee 4.0 8.0
Genus VULPAVUS Marsh
VULPAVUS AUSTRALIS Matthew
U.S.N.M. No. 18449, left mandibular fragment with M,, (loc. No.
48F R76).
This specimen does not permit anything to be added to Matthew’s
(1915a, p. 89) discussion of the species.
196 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
MEASUREMENTS OF TEETH OF VULPAVUS AUSTRALIS (IN MILLIMETERS)
Width
Length trigonid Heel
Meat aya Sie ee ET ie Eee ee PU Cah a eet hh eee
Mia es ee ee 2 DS a eee 6.9 4.5 4.0
Manes re pe) OL eed ae ee eee 4.8 3.9 3.3
Order CONDYLARTHRA
Family MENISCOTHERIIDAE
Genus MENISCOTHERIUM Cope
MENISCOTHERIUM TERRARUBAE Cope
U.S.N.M. No. 18451, fragment of right mandible with P.-M; (loc.
No. 48F R80): U.S.M.N. No. 18450, fragment of left mandible with
P.M; (loc. No. 48F R80).
The limited material of this form is uniform in size and is larger
than the material from Alkalai Creek listed by Granger (1915, p. 359).
On the basis of size the material agrees better with terrarubae than
with chamense, but whether these should receive full specific status
or be considered as varieties will not be considered here.
Although the genus was identified with certainty from one
locality, it would be premature to attempt any discussion of the
paleoecology of the Boysen Reservoir area on the basis of such limited
data.
Family HYOPSODONTIDAE
Genus HYOPSODUS Leidy
HYOPSODUS POWELLIANUS Cope
U.S.N.M. No. 18452, right mandible with P,—M; and right maxilla
with P?* (loc. No. 48F R75).
This specimen is referred to H. powellianus on the basis of size and
the position of the mental foramina, both of them lying below Py. The
teeth are so badly worn that certain identification is impossible.
MEASUREMENTS OF TEETH OF HYOPSODUS POWELLIANUS (IN MILLIMETERS)
Length
1 PSST (een eee NE Fa em a ps A le ey Sea eens aE Fee 8 2340
VW Pee eee es ae ME tre Rese eae RR Ft os Soa ae ek nr ee ee 18.5
Se ae Mee AERO pe EAT AT Ege ee ee 5.0
Mie et ae 2 es Sone ee he 2 ae ea ee 5.5
I Cpe oe enya ee pee oe abe 2 oe ok ees ee Sd eee ea 6.0
BOYSEN RESERVOIR VERTEBRATE FOSSILS—WHITE 197
HYOPSODUS WORTMANI Osborn
U.S.N.M. Nos. 18453, 18454, two specimens, one with upper and
lower molars associated (loc. No. 48F R78); U.S.N.M. Nos. 18455-
18460, six specimens, including one maxilla with M?®* (loc. No.
48F R80).
There appears to be some confusion in the literature concerning
the size range of the lower molars of this species. Osborn (1902,
p. 185) in the original description gives the size range from 11 to 13
millimeters. Loomis (1905, p. 422) found that his material exhibited
a uniform measurement of 12 mm. Matthew (1915b, p. 317), in his
key to the species of the genus, gives the length of the lower molars
as 10mm. In the material from the Boysen Reservoir area, the two
specimens with M,; have a molar length of 13 mm. In all the speci-
mens the length of M._; varies between 8.0 and 9.0 mm. Van Houten
(1945, p. 425) pointed out that most of the Lower Eocene genera
could be revised profitably. ‘This is certainly true of Hyopsodus.
Hyopsodus wortmani has never been adequately characterized in the
literature, and if this material is correctly referred, the lower dentition
may be characterized as follows: P, submolariform, anterointernal
style well developed and joined to the protoconid by a distinct crest;
deuteroconid well developed; anterointernal style, protoconid, and
deuteroconid forming a distinct trigonid; heel well developed and
trenchant, hypoconid centrally placed and prominent, entoconid small
and indistinct from posterior cingulum; M, with metaconid distinctly
twinned, crescents on protoconid well developed, hyopoconulid and
entoconid distinct, anterior and posterior cingula present; M, similar
to M, except that metaconid is indistinctly twinned; M, long and nar-
row, narrowing rapidly from in front posteriorly, hypoconulid as
large as, or larger than, hypoconid and forming a distinct heel;
entoconid small but distinct, metaconid may or may not be indistinctly
twinned.
This material is not readily distinguishable from H. paulus Leidy, of
the Lower Bridger, by size, but the teeth of the type are so badly worn
that their characters cannot be properly evaluated.
Order PANTODONTA
Family CORYPHODONTIDAE
Genus CORYPHODON Duméril and Bibron
CORYPHODON sp.
Right P** (loc. No. 48F R65).
This material is inadequate for more than generic identification.
198 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Order PERISSODACTYLA
Family EQUIDAE
Genus HYRACOTHERIUM Owen
HYRACOTHERIUM VENTICOLUM Cope
Figure 78
U.S.N.M. No. 18368, left mandibular fragment with Dp, (fig. 78)
(loc. No. 48FR65) ; U.S.N.M. No. 18462, right mandibular fragment
with the heel of Dp; and Dp, (loc. No. 48F R76); U.S.N.M. No.
18461, right mandibular fragment with P,., and M; (loc. No.
48F R78).
Although this material is very fragmentary, it shows the character
of the lower deciduous premolars, which was not treated in Granger’s
(1908) revision. Unfortunately, only the heel of Dp, is preserved.
USNM - 1/8368
FicurE 78.—Hyracotherium venticolum Cope, U.S.N.M. No. 18368; lateral and occlusal
views of right DPy, X 144.
The characters that this limited material presents are as follows:
Dp; with posteroexternal crescent well developed, cross-crest between
hypoconid and entoconid well developed, hypoconulid small but dis-
tinct, external and posterior cingula present; Dp, with anterior and
posterior external crescents well defined, metaconid distinctly twinned
and higher than protoconid, cross-crest between hypoconid and
entoconid well developed, entoconid higher than hypoconid, hypoconu-
lid small but well defined, well developed anterior, external, and
posterior cingula.
The deciduous teeth described here are somewhat higher crowned
than the permanent teeth, and can be distinguished from the perma-
nent dentition of Orohippus only with difficulty. In fact, these
teeth were originally referred to that genus and it was only after Dr.
C. L. Gazin and I spent some time comparing them with the material
in the U.S. National Museum that their true identity was learned.
I have been told by Morris Skinner that this characteristic—the
deciduous teeth of horses being more advanced than the permanent
teeth—has been observed in the later Tertiary horses. In view of the
growth processes involved in the formation of horse teeth (White,
1942, p. 26) it is logical to correlate the above phenomenon with the
activities of the endocrine glands, which stimulate and regulate
BOYSEN RESERVOIR VERTEBRATE FOSSILS—WHITE 199
growth, during the period of postnatal development (ibid., p. 45).
That the thyroid (Goldzieher, 1939, p. 83) plays an important role in
influencing the morphogenic processes, particularly in the ossification
of the skeleton and in the growth of the teeth, has been demonstrated
by the administration of thyroid extract to hypothyroid children and
by thyroidectomy of normal laboratory animals. Its action is by no
means independent but is closely integrated with that of the pituitary,
parathyroid, and adrenals. Nor do these glands function only in com-
bination with each other but in combination with the other glands of
the system to maintain an endocrine balance and a favorable “internal
environment” or homeostasis (ibid., p. 11).
In order to maintain homeostasis the endocrine glands must respond
to external factors. The changes in external environment that are
accompanied by changes in the activity of the glands are: Altitude,
temperature, climate, quantity and type of food, and accessory food-
stuffs such as mineral salts and vitamins. The data on the responses
to these factors are limited almost entirely to the clinical observations
on man and laboratory animals. These data indicate that when less
than radical changes in the external factors prove deleterious there
is a strong probability that an endocrine imbalance already existed
(Goldzieher, 1939, pp. 11-13).
Of equal importance to the activities of the endocrine glands are
the responses of the receptor tissues to the stimuli of the hormones.
These responses may be affected by a variety of factors, such as: The
condition of the tissues, ennervation, chemicals, and the age and stage
of development of the individual. With regard to the two last-
mentioned factors, which are probaby the most fundamental, little
is known except that the responses of the tissues are characteristic
of the stage of development. Thus, in a young and growing individ-
ual, the response to hormones is growth and maturation, while in the
adult the hormones are capable only of maintaining the orderly
function of the tissues (Goldzieher, 1939, p. 6). It is a well-known
fact that in animals there is a noticeable slowing down of growth soon
after puberty, though the postpubertal growth period may be as long
as the prepubertal. However, this may be due to the interaction of
the endocrine glands.
In regard to the environment offered by this region during Lower
Eocene time, Van Houten (1945, pp. 442-444) characterizes it as a
humid lowland with a warm-temperate to subtropical climate sup-
porting a luxuriant vegetation of both woodland and savannah types.
As to the soils, the parent rocks from which they were derived were
igneous (extrusives and intrusives) and sedimentary (limestones,
dolomites, shales, and sandstones). Weathering processes would make
available phosphorus, potassium, calcium, and some sodium from
200 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
the igneous rocks and calcium and magnesium from the limestones and
dolomites. The result would be a very fertile soil containing an ample
supply of all the minerals necessary for a luxuriant and nutritious
vegetation. In view of the above considerations, it can be concluded
that this region offered a very nearly optimum environment for
herbivorous animals, and that factors which would seriously disturb
the endocrine balance were absent.
The estrogens and androgens (sex hormones) (Goldzieher, 1939,
p. 289) and the hormone of the adrenal cortex (ibid., p. 93) have an
inhibitory effect on the thyrotropic hormone of the anterior lobe of the
pituitary, which results in decreased activity of thethyroid. Although
estrogens and androgens are present in nearly all foods (ibid., p. 748)
and growing plants, and do not appear to be altered by the processes
of digestion, they are not secreted in quantity by the gonads till a
short period before puberty. It is believed that only a portion of
these hormones taken with food find their way into the blood stream.
It has been shown that these hormones are inactivated in the liver
and are rapidly destroyed by oxidation in the lings (ibid., p. 748),
which greatly reduces their effectiveness when administered by mouth.
In view of the role played by the thyroid in the formation of the
teeth by its effect on metabolism and growth, the low estrogen and
androgen content of the blood while the deciduous teeth were being
formed could result in the advanced type of teeth. That the third
permanent premolar is more advanced than the fourth in some species
of Hyracotherium (Granger, 1908) may be due to the sudden increase
in the estrogen and androgen content of the blood during the interval
between the formation of these two teeth. If the appearance of the
physiological brake on the thyroid furnished by the secretions of the
gonads and the adrenal cortex were postponed until after the deter-
mination of the form of the permanent teeth by the growth of the
tooth germ, it is conceivable that the advanced type of tooth would
result. In view of the antagonism between the gonads and the thyroid
(Goldzieher, 1939, p. 94), such a deferment could be the result of
mild hyperthyroidism.
Family BRONTOTHERIIDAE
Genus LAMBDOTHERIUM Cope
LAMBDOTHERIUM POPOAGICUM Cope
U.S.N.M. No. 18464, right P?-M? (loc. No. 48F R76); U.S.N.M.
18463, right P?-M! (Shoshoni Reservoir) ; U.S.N.M. No. 18465, loose
upper teeth (loc. No. 48FR75).
This genus is generally accepted as the index fossil for the Lost
Cabin faunal zone of the Wasatchian, Lower Eocene. Unfortunately,
this material is too fragmentary to add anything to our knowledge of
BOYSEN RESERVOIR VERTEBRATE FOSSILS—-WHITE 201
this species. Bonillas (1936) has given very good reasons for con-
cluding that only one species existed in the Wind River Basin.
Genus EOTITANOPS Osborn
EOTITANOPS sp.
U.S.N.M. No. 18466, fragment of a right mandible with M,—M, (loc.
No. 48FR65) ; U.S.N.M. No. 19104, loose teeth including P;-, and part
of M, (loc. No. 48F R79).
This material is too fragmentary to add anything to our knowledge
cf the genus.
Family ISECTILOPHIDAE
Genus HEPTODON Cope
HEPTODON BROWNORUM Seton
U.S.N.M. No. 18467, badly broken right mandible with P,-M; and
portions of the left mandible (loc. No. 48F R65) ; U.S.N.M. No. 18468,
right mandible with P;-M, (loc. No. 48F R80) ; U.S.N.M. No. 18471,
badly crushed skull and jaws with associated skeletal fragments (loc.
No. 48F R75).
This material is referred to Heptodon brownorum on the basis of
size, but it is too fragmentary to add anything to our knowledge of
the species.
Order ARTIODACTYLA
Family DICHOBUNIDAE
Genus BUNOPHORUS Sinclair
BUNOPHORUS ETSAGICUS (Cope)
FIGURE 79
U.S.N.M. No. 18370, left mandibular fragment with P.—-M; and
alveolae for P._; (loc. No. 48F R76).
This specimen is a younger individual than the type (Sinclair, 1914,
p. 273) and shows the characters of the teeth much better. While
there are some differences the material is not adequate for specific
separation. Anterior mental foramen between P, and P, and the pos-
terior below the posterior root of P;; very short diastema between P,
and P;, both double rooted; P, with a small anterior tubercle, pro-
toconid and deuteroconid well developed, deuteroconid nearly as high
as protoconid (Cope, 1884, pl. 25e, fig. 24a, and Sinclair, 1914, fig. 7,
indicate that a deuteroconid may have been present but was obliterated
by wear), posterior cingulum with tubercle just lateral to median line;
paraconid appears to be absent on all of the molars, a faint, discontinu-
ous cingulum present on anterior, external, and posterior borders;
hypoconulid on M; larger than entoconid and forming a distinct heel.
202 PROCEEDINGS OF THE NATIONAL MUSEUM Vou. 102
MEASUREMENTS OF MANDIBULAR FRAGMENT OF BUNOPHORUS ETSAGICUS (IN
MILLIMETERS)
Length Width
| 2a] ye ae ene EN RSE gla OR! nD SLOW eee
35 apa ae Nee Drege 2 3g J Nd oir a ped De siehul) \: i ae
7.5 4.4
7.0 6.0
U4 6.5
940 G0
Ficure 79.—Bunophorus etsagicus (Cope), U.S.N.M. No. 18370; occlusal view of teeth and
lateral view of left mandible, X 1.
Genus DIACODEXIS Cope
DIACODEXIS OLSENI Sinclair
U.S.N.M. No. 18472, left mandibular fragment with P,-M, (loc.
No. 48F R78).
This material does not permit anything to be added to Sinclair’s
(1914, p. 292) discussion of the species.
SUMMARY
Since Tertiary reptiles are yet too poorly known to be useful as
horizon markers, they will be omitted from the summary. From the
Boysen Reservoir area 23 species of fossil mammals have been identi-
fied; 14 of them (see Table 2) are common to the Lost Cabin faunal
zone of the Lower Eocene, 2 of them to the Gray Bull, and 2 others
are common to both the Gray Bull and the Lysite. Consequently,
there can be little doubt that these deposits must be referred to the
Lost Cabin faunal zone. However, on structural grounds these beds
may be somewhat younger than the type section on Alkalai Creek.
There are about 250 feet of the Wind River formation exposed in
the Boysen Reservoir area. For the most part the formation consists
of drab greenish-gray gypsiferous clays with yellowish, usually fine-
grained, channel sandstones forming nearly vertical cliffs. While
most of the gypsum is probably secondary there are numerous areas of
local concentration caused by seepage of ground water or by capillary
BOYSEN RESERVOIR VERTEBRATE FOSSILS-—-WHITE 203
TABLE 2.—Species of fossil mammals identified from the Boysen Reservoir area
Gray Bull] Lysite | Lost Cabin
Didelphodusiventanus4- .---2 5. . 22 4tuth. S24. acu ties lage 8 EY a ge ms
Ccnodontonnys:scOublanUs= =- 95 a5 eee Dee ee ae ee 2
Gynoucontoniys lundeliusi, new species. — _ 2] -.-----|2--22.----|_2-2---: Ade.
mennOmum AcutiGens: 22. 22s fester tA pescoraibrs eeu lees ce eed x
INobharetus venticolus: 42-52-52 5255 2 o6--)455-5-— EA) eS Pe. X
END Un owsvixo Led Me pet) os RRS a SOE ET I aN a ae a [ee neal z
Loveina zephyri !____---- eh a al epee Glee Soa fone SEE oe
DONearOn eNO el cae ee oo ee Ree ee
RATAN SUD AOL: 22 oe ken eee ee Dae Ai eal eee aes a ae
params murinus)) o_o. 22.--22 34. 1 x cee 3 eee By,
Prolimnocyon antiquus-—___-~- RISE AE A ee | ieee a
PINOMssstrenua 265-5 ele tee AUN x x
Dicivimuchisnaltidenssss. 266 ee eb eT ec Joe Ne eee
Witlnavls australisei= = 25 2 os ee ee x x
Muistcistlatidens!t22 Sef) aa Shot MELE t t2 oA SE eh as face
Meniseotheriumyterrarubaes: £5 Ses 22 nae = ee oe
bivopsodus powellianus... 2... ...2. 2.22. -le<---3-.-- x
Hyopsodus wortmani____.__._---------- AN |e eee RE APA | of ORS See ee
Feaicotherum, venticolumé ..2 5-42 22... 3/2) Sees tele. 2-5
fambaouherium popoagicum_--..-..._-2-_|. 42. =-.-=--|. =2----==-
HS TINANODSADOreRHSe Loe | SHO UE ed Oh ae B oc) Sete es
Re poo On brO yorum = an) Anat ee ee eee | eee ae See
IBunophorus etsagicus. 22.225 === ----= 4 Ny (ee ae a
WIRCOGeXISHOISEMIEs alee kes eek Sib Ses FS LE ot ele | SE ee
CRU PG GINS ie pole eh he te ele aE et Se ae ee
i
|
uo -
a
i I et metion
— Loeal
K
4%
C74
tees Say Pew Dae
PA pe = ter
sali al
ma
i eT
fosio7d yelieV nyse
O08
5
3
Re @ ert
E-OS6S ov gom AS mol beigabA
i
|
ay
-
BOYSEN RESERVOIR VERTEBRATE FOSSILS—-WHITE
205
There appears to be less gypsum in this locality than in the others.
This locality was the most productive in both quantity and variety of
fossil mammals.
Table 3 shows the species obtained at each locality.
TaBLE 3.—Distribution of forms by localities
Mammals recognized from
the Boysen Reservoir area
Didelphodus ventanus_------
Cynodontomys scottianus
Cynodontomys lundeliusi___-
Esthonyx acutidens____---_-
Notharetus venticolus__-_---_-
WMeilhardellaisp.--=-.-=2_--==
Moveina zephyrie-=--22--.--
Stylinodon cylindrifer______-
Raramysimajors2-52---— 52
Paramys murinus-_-_--.------
Prolimnocyon antiquus-__-----
Sinopa strenua-.—_—-=- 4 = —
Didymictis altidens____-_----
Miacis latidens_.__....------
Vulpavus australis____..----
Meniscotherium terrarubae_ _
Hyopsodus powellianus-_-----
Hyopsodus wortmani
Hyracotherium venticolum _-_
Lambdotherium popoagicum_
ROtiLanO PSs Sp! aise ee eee
Heptodon brownorum___--__-
Bunophorus etsagicus-- -----
Diacodexis olseni___._.-_---
Coryphodonisp-==5-=2-._ 2
Locality number
65 75 76 TiC 78 79 80
RMA |e Bete citer yes | eats ick een oe eee x
Sears pe eee Xi le soe lot a. eee ali ok
> mm (see ya PPR Me (eS g R ir La iaP
> a ee pars >t eee > ae i ee x
Se ey PD 28 Mth Soe Sal ee eee
BE oe ee eae ee mere] emer ae eee = (6) eee ee be
i Ne ee > Se (Eee ae HE Re aed eee se
a ary | a ee Eig Vee ee lee alee ee
pe pe) 3 A at eae pea aby oa Earl Hse kl eoept gt | 028
ESA eye SEE Ses i ae ce eee Lie x
>. | eee > Gat | (ebepe ype eee Tee |e Nea om Pa OP Fe
APS RY eee ee Tee Neer See el ree |e
pret x >on) hesacantimeet ean iol ee probe eS
BAPE by | UN Real ISR daha peer a apd bance Neer TO aaa x
paces (eae) ect > rs es pee napelis (Fe Reese mad eel (Sa Vig
deeietlee 5.23 rs esl de thee |e oe x
apn ee Mr ei she hn ted ae ee ee ee a ere
Re eee eee ead eee 5 a oy are x
Sef eee > EP LEN ah ee eee eee
epee. x pan ER Oe ee oe cha iy poe SP ae eee
RF, ete es Ki ulideceehihecas Sollee ee
x x Gras veer wal ener ey 22, | UI a x
Ne ec ee Boy tetas ee ee | Cao
Sipe nd bce | ee ae SC >. i Weenteeneane des [AC >
> ir Kae tec) Pe eed Pde Seal th ee a mgt Se
LITERATURE CITED
Baver, C. M.
1934. Wind River Basin.
Boniiiasg, Y.
Bull. Geol. Soc. Amer., vol. 45, pp. 665-696.
1936. The dentition of Lambdotherium. Journ. Mammal., vol. 17, No. 2,
pp. 189-142.
Corr, B. D.
1884. The Vertebrata of the Tertiary formations of the West. Report of
the U. S. Geological Survey of the Territories, vol. 8, Book 1.
Gazin, C. L.
1936. A taeniodont skull from the Lower Eocene of Wyoming. Proc. Amer.
Philos. Soc., vol. 76, pp. 597-612.
GiumoreE, C. W.
1928. Fossil lizards of
North America. Mem. Nat. Acad. Sci., vol. 22,
38d Mem., 169 pp.
1938a. Fossil snakes of North America.
No. 9.
Geol. Soc. Amer., Special Paper
1938b. Descriptions of new and little-known fossil lizards from North
America.
Proce.
U. S. Nat. Mus., vol. 86, pp. 11-26.
206 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
GOLDZIEHER, MAX A.
1939. ‘The endocrine glands. New York.
GoopricH, E. S.
1930. Studies on the structure and development of vertebrates. London.
GRANGER, WALTER.
1908. A revision of the American Eocene horses. Bull. Amer. Mus. Nat.
Hist., vol. 24, pp. 221-264.
1910. Tertiary faunal horizons in the Wind River Basin, Wyoming, with
descriptions of new Hocene mammals. Bull. Amer. Mus. Nat.
Hist.,. vol. 28, pp. 235-251.
1915. A revision of the Lower Eocene Wasatch and Wind River faunas.
Part IiI. Order Condylarthra, families Phenocodontidae and
Meniscotheriidae. Bull. Amer. Mus, Nat. Hist., vol. 34, pp. 829-361.
Grecory, W. K.
1910. The orders of mammals. Bull. Amer. Mus. Nat. Hist., vol. 27,
pp. 1-524.
Grecory, W. K., and Simpson, G. G.
1926. Cretaceous mammal skulls from Mongolia. Amer. Mus. Nov., No. 225.
JEPSEN, G. L.
1930. New vertebrate fossils from the Lower HKocene of the Bighorn Basin,
Wyoming. Proc. Amer. Philos. Soc., vol. 69, pp. 117-181.
1934. Tet fina alk alee) cgfi x Tal NY PUR AE a
Kish cor, Roth + ME >, 4 Ah eae eee
Power yet “1 a Me LS eine ‘
7 : - : j ih ate
:
- thy | H ‘ i
a 2 : ,
_ { il pean P
— )
. tt I 4 : 7
‘ | Wear} na
© ae ie > a
' 7 ; iis is oi Ue tee
, i { ¥
( j a ee ‘
Pai ‘ais
; _— ;
| | | ene rk: Ap Lae Be Sai
? ' r 1 ‘
: MrT es to Peer es Y,
Le Wd ALS 1
>
A
: q
i
‘ '
\
‘ om, :
> \ ; 7
tans ;
1 i} Ls + a
Li fn ay en le
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
issued
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 Washington: 1952 No. 3297
A NEW CRAYFISH FROM ALABAMA, WITH NOTES
ON PROCAMBARUS LECONTEI (HAGEN)
By Horron H. Hosss, JR.
THE COMBINED ranges of the members of the Blandingii section of
the genus Procambarus, excluding mountainous areas, extend from
Mexico to Massachusetts, and from Florida to Ohio and Minnesota.
The new species described herein is found in a region in which the
members of this section have shown the greatest degree of speciation
(i. e., in the southeastern part of the United States). Since the
species is represented in my collection by only six specimens, taken
from the type locality, little is known of its habits and variation,
and its range can be postulated only in terms of the known distribu-
tion of its close relatives, which presumably have similar habitat
preferences. On the basis of the latter it seems probable that it is
confined to the Tallapoosa River and its tributaries.
Procambarus lecontei (Hagen, 1870, p. 47) has never been ade-
quately diagnosed, and the published figures (first pleopods, antennal
scale, and epistome by Hagen, 1870, and dorsal aspect by Faxon,
1885a) are inadequate and not entirely accurate. Furthermore, since
its original description there has been considerable confusion as
to its range. In addition to presenting a revised diagnosis of
P. lecontei and illustrating the diagnostic features, I am including
a bibliography of the species and summarizing its known range.
I wish to express my gratitude to Dr. E. C. Raney, R. D. Suttkus,
and J. Kezer, of Cornell University, for the gift of the specimens
on which the new species is based, as well as to others, mentioned in
connection with collection data, who have assisted in collecting
specimens of P. lecontez.
967611—52 209
210 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Abbreviations used to indicate repositories of specimens are as
follows:
HHH, my personal collection at the University of Virginia. Note
that the date collected may be obtained from the catalog number—
6-1649-2a, specimens were collected on June 16, 1949.
MCZ, Museum of Comparative Zoology.
TU, Tulane University.
U.S.N.M., United States National Museum.
Genus PROCAMBARUS Ortmann
Cambarus ORTMANN, 1905b, p. 487.
PROCAMBARUS LECONTEI (Hagen)
Ficure 81
Cambarus lecontei HAcren, 1870, pp. 10, 45-47, 48,7 52, 97,’ 106,7 107 (pp. 100, 101,
and 106 not lecontei), figs. 15, 18, 145.—CrraAsrr, 1934, p. 4.—Faxon, 1884,
pp. 110, 1387; 1885a, pp. 17, 19, 22, 24, 29-30,? 31, 32, 33, 158,7 167," 1687 173,
pl. 2, fig. 2; 1885b, p. 358; 1914, p. 418.—Harris, 1903, pp. 58,7 107,’ 188,7 144,
151 (pp. 148 and 152 not lecontei).—Hay, 1899, pp. 959, 963.—OrTMANN, 1902,
p. 277; 1905a, pp. 102, 129.
Procambarus lecontei Hosss, 1942b, p. 342 (by implication) ; 1942c, pp. 94, 95, 98.
Diagnosis.—Rostrum with lateral spines; areola moderately broad
with four or five punctations in narrowest part; cephalothorax
granulate laterally, punctate dorsally ; a single lateral spine present on
each side of carapace. Male with hooks on ischiopodites of third
and fourth pereiopods; palm of chela of first-form male not bearded
but bearing a row of seven to nine tubercles on inner margin; post-
orbital ridges terminating cephalad in spines. First pleopod of first-
form male with no marked hump on cephalic surface but with a
noticeable “knob” on lateral surface opposite the caudal process;
tip terminates in four distinct parts (of which all except caudal proc-
ess are directed caudad at approximately a right angle to main shaft
of appendage): mesial process long, slender, subcylindrical, non-
corneous, lies considerably proximad of and extends much farther
caudad than the other terminal elements; cephalic process moderately
short, compressed, corneous, and somewhat hoods the more prominent
central projection; caudal process slender, somewhat flattened,
corneous, lies proximad of the central projection, and is directed at
about a 45° angle to the main shaft of the appendage; central projec-
tion compressed, corneous, beaklike, and extends slightly farther
caudad than either cephalic or caudal processes. Annulus ventralis
as in fig. 81, d, decidedly broader than long with a prominent raised
area on cephalolateral (dextral or sinistral) side of sinus; sternum
immediately cephalad of annulus with multituberculate prominences
which extend caudad to cover a portion of annulus.
1In part, excluding records from Georgia, Florida, Mississippi, and North Carolina.
A NEW CRAYFISH FROM ALABAMA—HOBBS oi
b= yy,
z Zend
> \
a
Ficure 81.—Procambarus lecontet (Hagen), from Moores Creek, 8 miles southwest of Mobile
on United States Highway 90, Mobile County, Ala.: a, Mesial view of first pleopod of
male, form I; 6, lateral view of carapace of male, form I; c, lateral view of first pleopod of
male, form I; d, annulus ventralis; ¢, mesial view of first pleopod of male, form I; f, lateral
view of first pleopod of male, form I; g, epistome of male, form I; A, basipodites and ischio-
podites of third and fourth pereiopods of male, form I; 7, dorsal view of carapace of male,
form I; 7, mesial view of first pleopod of male, form II; k, antennal scale of male, form I;
d, lateral view of first pleopod of male, form II; m, upper view of chela of male, form I.
a=Mesial process; b=cephalic process; d=caudal process; z=central projection.
Pubescence removed from all structures illustrated except m.
Bie PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Specimens examined.—AuaBaMa: Mobile County, MCZ No. 217%,
Mobile (types) (131,24 ¢11,7? 2); MCZ No. 4958, Mobile (types)
(161); HHH No. 6-1649-2a, Clear Creek, 4.7 miles east of Semmes
[R. D. Suttkus, C. F. Cole, and R. H. Gibbs] (1¢1,1?); HHH No.
6-340-1b, 5.6 miles east of Irvington [L. Berner, C. Benton, and HHH]
(1¢J,1¢I1I,1°); HHH No. 6-340-9c, 2.5 miles south of Bucks [LB,
CB, and HHH] (1 2 ); HHH No. 6-240-5b, 3.5 miles south of Irving-
ton [LB, CB, and HHH] (246 641,24 ¢11,1¢ imm): Baa:
6—240-2a, 3.8 miles west of Grand Bay [LB, CB, and HHH] (1 ¢ I);
HHH No. 6-140-9a, 15 miles southwest of Mobile [LB, CB, and
HHH] (1¢1); HHH No. 6-140-8a, 8 miles southwest of Mobile
[LB, CB, and HHH] (2¢ 31; 1341,29 9,29 9 1mm) MOZiino
number), Mobile (141 dry).
Mississtpp1: Stone County, HHH No. 6-1649-1, 16.6 miles east of
Wiggins [RDS, CFC, and RHG] (1éII, 22 2); in collection of
George H. Penn, Jr., Tulane University, TU. No. 1233, 16 miles east
of Wiggins (6/8/49) (26 éII,1¢).
Remarks.—F axon (1885a, p. 30) has discussed the locality records
cited by Hagen (1870, p. 48), and has indicated that only the speci-
mens mentioned by the latter that were collected from Mobile, Ala.,
belong to Procambarus lecontei.2, Following his discussion of Hagen’s
locality records he stated that “besides the Mobile types, I have found
but one other specimen of this species in the Museum, a young female
in a jar with @. spiculifer (Cat. No. 172), from Athens, Ga.” I have
examined this specimen, which is in a poor state of preservation,
and am uncertain as to its identity, but even if it should prove to be
lecontei there is every reason to doubt the reliability of the label.
I have collected on several occasions in the vicinity of Athens, and
have received numerous collections made by Dr. Donald C. Scott
from that region, and neither of us has found P. lecontet. In fact,
only two species belonging to the genus Procambarus have been found
in the Athens region: P. spiculifer (LeConte, 1856, p. 401) and P.
pubescens (Faxon, 1884, p. 109). Furthermore, no other specimen of
lecontet has been reported east of Mobile County, Ala. Thus P.
lecontet is known only from the southwestern part of Alabama and
the southeastern part of Mississippi, where it is found in tributaries
of the Mobile and Pascagoula Rivers.
PROCAMBARUS VERRUCOSUS, new species
FIcure 82
Diagnosis —Rostrum with small lateral spines or tubercles, which
set off the acumen from basal portion; areola moderately narrow with
2T may add that the specimens from Pensacola, Fla., which Faxon stated were not
leconteit but of which he made no specific determination, are Procambarus evermanni
(Faxon, 1890, p. 620) [MCZ No. 249].
A NEW CRAYFISH FROM ALABAMA—HOBBS 213
three punctations in narrowest part; cephalothorax markedly granu-
late laterally and in the caudal portion of the areola; a single lateral
spine present on each side of carapace. Male with hooks on ischi-
opodites of third and fourth pereiopods; palm of chela of first-form
male not bearded but bearing a row of seven to nine tubercles along
inner margin; postorbital ridges terminating cephalad in spines.
First pleopod of first-form male with a rounded hump on cephalic
margin near distal end, and terminating in four distinct parts: The
noncorneous mesial process prominent, acute, and extends caudodistad,
and laterad of the other terminal elements; the small, corneous, acutely
triangular cephalic process arises from middistal end of appendage;
caudal process corneous, and flattened cephalocaudad, corneous, sub-
triangular, and concave in caudal aspect, and closely applied to
caudolateral surface of central projection; central projection corne-
ous, slender, and terminating bluntly, directed caudodistad as are
cephalic and caudal processes. Annulus ventralis as in figure 82, d,
and sternum immediately cephalad of annulus with multituberculate
prominences, which extend caudad to cover portion of annulus.
Holotypic male, form I.—Body subovate, somewhat compressed
laterally; abdomen narrower than thorax (12.9-14.6 mm. in widest
parts, respectively) ; width of carapace slightly less than depth in
region of caudodorsal margin of cervical groove (14.6-15.0 mm.).
Areola moderately narrow, about 7.8 times longer than broad with
three punctations in narrowest part, caudal third granulate; cephalic
section of carapace about 2.1 times as long as areola (length of areola
about 31.9 percent of entire length of carapace).
Margins of rostrum gently converging cephalad, and terminating
at base of acumen in a small acute tubercle on each side; acumen not
upturned ; rostrum excavate above and without swollen margins, and
studded with many inconspicuous setae arising from very minute
punctations.
Postorbital ridges grooved laterally, and terminating cephalad in
acute spines; subrostral ridges moderately well developed, but evident
in dorsal aspect only at base; suborbital angle small and obtuse;
branchiostegal spine well defined and acute; sides of carapace with
an acute spine on each side. Surface of carapace granulate except for
cephalodorsal portion of carapace and cephalic two-thirds of areola
which are punctate.
Abdomen longer than thorax (34.7-31.9 mm.).
Cephalic section of telson with two spines in each caudolateral
corner. Outer dextral spine bidentate.
Epistome semiovate with a distinct cephalomedian projection; en-
tire margin bearing plumose setae.
Antennule with a strong spine on ventral surface of basal segment.
214 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Antenna broken, but in other specimens extends caudad to fourth
abdominal segment; antennal scale broad with a well-developed spine
on outer distal margin; lamellar portion rounded mesially, and broad-
est proximad of middle (fig. 82, %).
Right chela slender, with inflated palm; palm studded with seti-
ferous tubercles on all surfaces. Inner margin of palm with a row of
nine tubercles which are only slightly more prominent than others
flanking this row. In addition to squamous tubercles on lower surface
of palm a distinctly larger tubercle present near base of dactyl.
Fingers not gaping. Opposable margin of immovable finger with a
row of three small tubercles on basal fifth, the distal one largest; a
strong tubercle extends mesiad from lower opposable margin near
midlength of finger ; otherwise opposable margin with crowded minute
denticles. Opposable margin of dactyl with 2 proximal rows of
rounded tubercles: an upper row of 10 and a lower one of 6; between
and distad of these 2 rows are crowded minute denticles. A low
longitudinal ridge present on upper surface of both fingers; all sur-
faces of fingers with setiferous punctations except along proximo-
mesial portion of dactyl where squamous tubercles occur in basal sixth.
Carpus of first right pereiopod about 1.7 times longer than broad
with the usual oblique furrow on upper surface represented by a broad
shallow depression; all surfaces with squamous tubercles, more abun-
dant in mesial half of segment; mesial surface with an oblique row
of three spikelike tubercles. Lower mesiodistal margin with a strong
acute tubercle, and a similar one on lower laterodistal margin.
Merus of first right pereiopod with 2 irregular rows of tubercles
on upper margin and a few scattered ones between; 2 strong acute
tubercles in distal portion; lateral surface with scattered punctations ;
lower surface with two rows of tubercles: lateral row of 14 and mesial
row of 16, and in addition a few tubercles on both sides of these 2
rows.
Ischiopodites of third and fourth pereiopods with hooks; hooks
simple, that on third extending proximad of ischiopodite, and that
on fourth opposed by a tubercle on basipodite. Coxopodites of fourth
and fifth pereiopods with ventrally projecting prominences: those
on fourth swollen, and those on fifth smaller and less bulbiform.
First pleopods symmetrical and reaching coxopodite of third pereio-
pod when abdomen is flexed. (See description of pleopod under
Diagnosis.)
Morphotypic male, Form II.—Differs from the holotype in the fol-
lowing respects: Rostrum more contracted distally ; caudal portion of
areola punctate; all spines on telson simple; mesial row of tubercles on
A NEW CRAYFISH FROM ALABAMA—HOBBS 215
Ficure 82.—Procambarus verrucosus, new species: a, Mesial view of first pleopod of holo-
type; b, lateral view of carapace of holotype; c, lateral view of first pleopod of holotype;
d, annulus ventralis of allotype; ¢, mesial view of first pleopod of holotype; f, lateral view
of first pleopod of holotype; g, epistome of holotype; h, mesial view of first pleopod of
morphotype; 1, lateral view of first pleopod of morphotype; 7, dorsal view of carapace of
holotype; &, antennal scale of holotype; /, basipodites and ischiopodites of third and fourth
pereiopods of holotype; m, upper view of chela of holotype. a=Méesial process; b=
cephalic process; d=caudal process; z=central projection. Pubescence removed from
all structures illustrated except m.
216 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
carpus of cheliped consisting of 4 instead of 3; only 1 acute tubercle
(spine) present near upper distal end of merus of cheliped, and lower
surface of same podomere with 14 tubercles in mesial row. The usual
secondary sexual differences occur with smaller hooks on ischiopodites
of third and fourth pereiopods and less well-developed armature of the
coxae of the fourth and fifth pereiopods. First pleopod with all ter-
minal elements represented and disposed as illustrated (fig. 82, A, 2).
Accessory shoulder, typical of the group of which this species is a
member, lies more laterad in the second-form male. (See Afeasure-
ments. )
Allotypic female—Differs from the holotype in the following re-
spects: Caudal portion of areola punctate; sinistral outer spine of
telson broken but was bidentate; chela comparatively broader and
shorter with upper opposable margin of immovable finger bearing
a row of 5 tubercles, of which the second from base is largest, and
lower opposable margin with 1 large tubercle at base of distal two-
fifths; opposable margin of dactyl with a row of 7 tubercles, of which
third from base is largest; lower mesial row of 14 tubercles on merus
of cheliped.
Sternum immediately cephalad of annulus ventralis with paired
caudally projecting tuberculate prominences which extend over (ven-
trally) the cephalic margin of the latter. Annulus ventralis (fig.
82, d) irregularly shaped with its greatest length in the transverse
axis. Dextral wall very high and relatively thin; cephalosinistral wall
much thicker and of more irregular contour. Sinus originates near
cephalic margin of annulus and extends caudodextrad slightly caudad
of midlength; here it makes an S-curve just crossing the median line
and turns caudad and slightly dextrad almost to caudal margin of
annulus.
Measurements (in millimeters) —
Carapace: Holotype Allotype | Morphotype
ReRerg hit tyee or ele ee A Re 15. 0 15. 9 14, 2
AVAL CU IN eee ae Bey ea 14. 6 15.8 15.1
en gti 122i va Soe Oe wen eee 31.9 33. 2 30. 6
Areola
engthsss ave ieee. Re ee at 10. 1 10. 4 9.9
WALES ae ee ek ee 1.5 1.5 1.4
Rostrum:
Benet £27k ee be aS See Ee 9. 0 9. 2 8. 6
Wile hic aslo te tek eee ee ere > 6. 4 5. 6 5. 3
Right chela:
Length, inner margin of palm_-__-_ 10. 9 6.1 7.3
Width of palms iw boa BS 6. 9 6. 0 4.6
Length, outer margin of hand-_- -- 28. 2 17.0 19, 2
Length ofidactyli 22 4. . Ueitwenets 15.3 10. 1 10. 5
A NEW CRAYFISH FROM ALABAMA—HOBBS Al
Type locality— A tributary of Calebea Creek [Alabama River sys-
tem], 3.9 miles south of Tuskegee, Macon County, Ala., on United
States Highway 29. The specimens on which this description is
based were collected by Dr. E. C. Raney, of Cornell University, on
March 24, 1948, and he has kindly furnished me with the following
data: At this locality the stream is clear, about 10 feet wide and 3
feet deep, sandy-bottomed, and with a volume flow of 5 cubic feet per
second. The stream flows through an open pasture, and at the time
the collection was made the temperature of the air was 80° F., and
that of the water 65° F.
Disposition of types —The holotypic male, form I, allotypic female,
and morphotypic male are deposited in the United States National
Museum (No. 90743). The paratypes, consisting of one male, form
I, one male, form IT, and one female, are in my personal collection at
the University of Virginia (No. 3-2448-3b).
Relationships —Procambarus verrucosus, « member of the Bland-
ingii group (Hobbs, 1942b, p. 341), has its closest affinities with Pro-
cambarus blandingii acutus (Girard, 1852, p. 91). However, it may
readily be distinguished from it by the more complexly appearing
annulus ventralis of the female, and the structure of the first pleopod
of the male. The caudal knob of the first pleopod has assumed a
markedly different position in P. verrucosus from that in some of the
other species of this group, in which this knob more closely resembles
that of the hypothetical generalized pleopod (Hobbs, 1942a, p. 58).
In P. hayi (Faxon, 1884, p. 108) the caudal knob (which has been
prolonged proximally into a long irregular fold) lies on the caudo-
lateral face of the appendage at the base of the caudal process; in
P. blandingii acutus, while the caudal knob is distinctly knoblike, it
has shifted caudolaterad and lies at the cephalolateral base of the
cephalic process; in P. verrucosus there has been a still stronger degree
of shifting in a cephalic direction so that it forms a rounded shoulder
on the cephalic border of the appendage. Were it not for the in-
termediate stages of shifting of the “caudal knob” which have been
observed in specimens tentatively identified as P. blandingii acutus
(a subspecies that has never been clearly defined) one would hardly
suspect that the “shoulder” on the pleopod of P. verrucosus had any
relationship to that of the more “typical” caudal knob as occurs in
P. pictus (Hobbs, 1940, p. 419) and the “less typical” one in P. havi.
Remarks. —The annulus ventralis of the allotype contains a sperm
plug, which indicates that this species breeds in the early spring;
however, it should be pointed out that this does not mean that this
species does not breed during summer, fall, or winter.
218 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Collected with P. verrucosus in the type locality were five specimens
of P. versutus (Hagen, 1870, p. 51).
The name verrucosus refers to the granulate condition of the
carapace.
LITERATURE CITED
CREASER, FE. P.
1934. A new crayfish from North Carolina. Oce. Pap. Mus. Zool. Univ.
Michigan, No. 285, pp. 1-4, 1 fig.
FAxoN, WALTER. :
1884. Descriptions of new species of Cambarus ; to which is added a synonym-
ical list of the known species of Cambarus and Astacus. Proc.
Amer. Acad. Arts and Sci., vol. 20, pp. 107-158.
1885a. A revision of the Astacidae. Mem. Mus. Comp. Zool., vol. 10, No. 4,
pp. 1-179, 10 pls.
1885b. A list of the Astacidae in the United States National Museum. Proce.
U. 8. Nat. Mus., vol. 8, pp. 356-861.
1890. Notes on North American crayfishes—family Astacidae. Proc. U. 8.
Nat. Mus., vol. 12, pp. 619-634.
1914. Notes on the crayfishes in the United States National Museum and
the Museum of Comparative Zo6dlogy with descriptions of new species
and subspecies to which is appended a catalogue of the known species
and subspecies. Mem. Mus. Comp. Zool., vol. 40, No. 8, pp. 351-427,
pls. 1-13.
GIRARD, CHARLES.
1852. A revision of the North American Astaci with observations on their
habits and geographical distribution. Proc. Acad. Nat. Sci. Phila-
delphia, vol. 6, pp. 87-91.
Hacen, H. A.
1870. Monograph of the North American Astacidae. Illus. Cat. Mus. Comp.
Zool., pt. 3, pp. 1-109, pls. 1-11.
Harris, J. ARTHUR.
1903. An ecological catalogue of the crayfishes belonging to the genus Cam-
barus. Kansas Univ. Sci. Bull., vol. 2, No. 3, pp. 51-187, 5 pls.
Hay, W. P.
1899. Synopses of North American invertebrates, VI: The Astacidae of
North America. Amer. Nat., vol. 33, pp. 957-966, 1 fig.
Hosgss, Horton H., Jr.
1940. Seven new crayfishes of the genus Cambarus from Florida, with notes
on other species. Proc. U. S. Nat. Mus., vol. 89, pp. 387-423, figs. 14—
22,
1942a. On the first pleopod of the male Cambari (Decapoda, Astacidae).
Proce. Florida Acad. Sci., vol. 5, pp. 55-61, 2 pls.
1942b. A generie revision of the crayfishes of the subfamily Cambarinae
(Decapoda, Astacidae) with the description of a new genus and
species. Amer. Midl. Nat., vol. 28, No. 2, pp. 334-357, 3 pls.
1942c. The crayfishes of Florida. Univ. Florida Publ., biol. sci. ser., vol. 3,
No. 2, pp. 1-179, 3 figs., 24 pls., 11 maps.
LECONTE, JOHN.
1856. Description of new species of Astacus from Georgia. Proc. Acad. Nat.
Sci. Philadelphia, vol. 7, pp. 400-402.
A NEW CRAYFISH FROM ALABAMA—HOBBS 219
ORTMANN, A. E.
1902. The geographical distribution of fresh-water decapods and its bearing
upon ancient geography. Proc. Amer. Philos. Soc., vol. 41, pp. 267-
400.
1905a. The mutual affinities of the species of the genus Cambarus, and their
dispersal over the United States. Proc. Amer. Philos. Soc., vol.
44, pp. 91-136, 1 map.
1905b. Procambarus, a new subgenus of the genus Cambarus. Ann. Car-
negie Mus., vol. 3, No. 3, pp. 485-442, 1 fig.
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U. S. NATIONAL MUSEUM
Vol. 102 Washington : 1952 No. 3298
MOTHS OF THE GENERA MULONA WALKER AND LOMUNA,
A NEW AND CLOSELY RELATED GENUS (ARCTIIDAE:
LITHOSIINAE)
By Wruuiam D. Frevp
THE TWO genera of moths! treated in this paper are found only in
the Greater Antilles and the Bahama Islands. Until the present time
they have been treated as a single genus with three species, Mulona
lapidaria Walker, Mulona nigripuncta Hampson, and Mulona grisea
Hampson.' A careful study of the 47 known specimens from the
collections of the United States National Museum, the British
Museum (Natural History), the American Museum of Natural
History, the Carnegie Museum, and Cornell University, disclosed
two genera and six species involved in the complex.
Palpal, antennal, and venational characters are identical in the
two genera treated in this paper and are given here to avoid repetition
in the generic descriptions. Labial palpus upturned, reaching middle
or slightly above middle of frons. Antenna of male and female fili-
form and gradually more slender to the tip; each subsegment with
two pairs of bristles, one from near or above middle of ventrolateral
margin of subsegment, the second much smaller and just behind the
first; subsegments pubescent. Venation of forewing with vein 2 from
middle of cell or from just before or after middle, downward curved
at base; vein 3 from just below lower angle of cell; veins 4 and 5 sepa-
rate, 4 from lower angle, 5 from slightly above lower angle (in one of
the 5 specimens of Lomuna nigripuncta Hampson, 4 and 5 are con-
nate or extremely short stalked); vein 6 from below upper angle of
cell; vein 9 from stalk of 7 and 8 or rarely 7 from stalk of 8 and 9;
1 The species Autoceras phelina Druce was placed in Mulona by Hampson (Catalogue of the Lepidoptera
Phalaenae in the British Museum, vol. 2, p. 387, 1900). It does not belong in Mz/ona and was transferred
to the genus Gaudeator Dyar by Forbes (Bull. Mus. Comp. Zool., vol. 85, No. 4, p. 183, 1939).
967606—52 221
222 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 102
veins 10 and 11 free; vein 10 from middle or nearer stem 7, 8, and 9
than to 11; vein 11 from beyond middle of cell and curved toward 12,
distally coincident with 12. Venation of hindwing with vein 2 from
outer third of cell or beyond; vein 3 stalked with 4 from lower angle
of cell; vein 5 absent; vein 6 separate from 7 or sometimes connate
or stalked with 7 (in the single female specimen of Mulona manni),
6 from upper angle, 7 from before upper angle of cell; vein 8 from
before middle of cell. —
KEY TO THE INCLUDED GENERA, BASED UPON GENITALIA
1. Male with uncus short and broadly bilobed; gnathos absent; anellus bifur-
cate with arms weakly sclerotized and short, one-eighth or less the length
of harpes; female with ductus bursa cylindrical, not at all flattened; pos-
terior genital plates ADSent= 222.2 82325 a ee Lomuna, new genus
Male with uncus long, not bilobed, either slender, curved and hooklike, or
broad and hoodlike; gnathos present; anellus bifurcate with arms strongly
sclerotized and with arms long, nearly as long or longer than harpes;
female with ductus bursa flat, not sclerotized or only partially sclerotized;
posterior genital plate present.o- 2-22. 5. - =. -22=-=2-2 Mulona Walker
Genus MULONA Walker
Mulona Waker, List of specimens of lepidopterous insects in the collection of
the British Museum, vol. 35, p. 1896, 1866.—Hampson, Catalogue of the
Lepidoptera Phalaenae in the British Museum, vol. 2, pp. 386-388, 1900.—
Draupt, in Seitz, Gross-schmetterlinge der Erde, vol. 6, p. 252, 1918.—
Forses, Scientific survey of Porto Rico and the Virgin Islands, vol. 12,
pt. 1, p. 33, 1930.—Srranp, in Wagner, Lepidopterorum catalogus, pars 26,
p. 732, 1922.
Genotype.—Mulona lapidaria Walker. Monobasic.
Labial palpus, antenna, and venation as described on page 221.
Male genitalia (pl. 10, figs. 2-6) with uncus slender, curved and
hooklike in some species and broad and hoodlike in others, terminating
in a short constricted point in all species, this point usually blunt;
enathos weakly sclerotized and divided, forming two broad short
lobes; anellus bifurcate with stongly sclerotized arms and these arms
nearly as long or longer than harpes; transtilla present and wholly
or partially fused; vinculum triangular to nearly round; aedeagus
thick throughout, distally broad and blunt and armed with various
broad, sometimes dentate lobes; ductus ejaculatorius entering anterior
end of aedeagus; vesica with numerous scobinations; harpe with an
apical lobe or arm (the cuiller) and with a clavus and with a third
arm from base of costa (except in M. schausz where apex of harpe is
a broad rounded lobe and clavus is greatly reduced).
Female genitalia (pl. 11, figs. 8-11) with ductus bursa nearly flat
and only partially sclerotized, with two separate sclerotized latero-
ventral plates near ostium bursae or with ductus bursae flat and
almost entirely sclerotized, triangular or subtriangular; bursa copu-
MOTHS OF GENERA MULONA AND LOMUNA—FIELD 223
latrix obpyriform or subspherical, longer than broad and heavily and
completely serratulate through middle two-thirds or more; seventh
sternum not modified, weakly sclerotized; eighth sternum with sclero-
tized bilobed lip at posterior margin (the posterior genital plate),
each lobe with sclerotized arm extending anteriorly to ductus bursa;
with bilobed invaginated sac on dorsal surface immediately before
ovipositor.
Remarks.—As defined above Mulona contains five species and is
confined to the Greater Antilles and the Bahama Islands.
KEY TO THE SPECIES OF MULONA
aa MvMialeg sis oso a4 J. ee Deut tO ees Wait 1.6 UO isp $i 2
Bemales! = 250 foes se ah ee ee a bp th By Adie Eade eb epaye 6
Zewuncus broad and hoodlike (pl. 10, figs. 5, 6)..-.2-~.--~==-. ==-.4: =. =-4- 3
Uncus slender, curved and hooklike (pl. 10, figs. 2, 3, 4)
3. Apex of harpe pointed and sharply bent back; clavus as broad as long; harpe
without long sharp arm from base of costa (pl. 10, fig. 5)__.manni, new species
Apex of harpe rounded and recurved but not sharply bent back; clavus
longer than broad; harpe with a long sharp arm from base of costa (pl. 10,
1045 (6) pee SR es ee 2 eee Serer ee anne Ceres et ee barnesi, new species
4. Anellus with arms of equal length; harpe with costal arm absent (pl. 10,
TYAS Gis CB SS le EE fae 5 tah eet a AE Ip en a, Ay Nl 5
Anellus with arms subequal in length; harpe with costal arm nearly equal in
length and subparallel to apical arm (pl. 10, fig. 2)___~__ grisea Hampson
5. Harpe with apical arm present, about four times as long as broad, recurved;
arms of anellus divergent at free end (pl. 10, fig. 3)--lapidaria Walker
Harpe without apical arm, apex broadly rounded; arms of anellus directed
toward each other at free end (pl. 10, fig. 4)__-_-_- schausi, new species
6. Ductus bursa partially selerotized with two separate, sclerotized lateroventral
bilobed plates near ostium bursae (pl. 11, figs. 10, 11)_-___.---_---- i
Ductus bursa almost entirely sclerotized and without separate sclerotized
plstess(olalal sige ASO) se oak ae ee DS ee ee 8
7. These plates nearly flat and only slightly upturned anteriorly (pl. 11, fig. 11).
barnesi, new species
These plates distinctly curved and greatly upturned anteriorly (pl. 11,
fiery AO) sins Ary st gyi beet epistles e eS Lay a Oecd _ manni, new species
8. Ductus bursae with sclerotized plate nearly triangular (pl. 11, fig. 9).
schausi, new species
Ductus bursae with sclerotized plate bilobed near entrance to bursa copu-
Peutrxer (Oe a lel hi ons) ee ae eee ee ae rc fy nee grisea Hampson
1. MULONA GRISEA Hampson
Puate 10, Figures 2, 2A; PLATE 11, Fiaure 8
Mulona grisea Hampson, Catalogue of the Lepidoptera Phalaenae in the British
Museum, vol. 2, p. 388, pl. 29, fig. 30, 1900.—Dravupt, in Seitz, Gross-
schmetterlinge der Erde, vol. 6, p. 252, pl. 32, fig. L 8, 1918.—Srranp, in
Wagner, Lepidopterorum catalogus, pars 26, p. 732, 1922.
Male.—Palpus light gray or white below, dark fuscous above.
Antenna and head white or light gray tinged with yellow or brown,
sometimes darker. Mesoscutum, patagium, tegula, and meso-
scutellum white or light gray, sprinkled with brownish scales. A light
224 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 102
brown spot at center of hind margin of mesoscutellum. Legs dirty
white to pale yellow or brown, fuscous along inner sides of foreleg
and with midlegs and hindlegs sometimes faintly banded. Abdomen
dirty white. Forewing above gray or grayish white, heavily irrorated
with light brown and with the following small dark brown or black
spots: one at costa near base of wing; one just below cell and at about
the basal one-fourth of cell; a row of four, crossing wing just before
middle, the first of these on costal margin, the third just below middle
of cell, and the fourth immediately below the third and on the anal
vein, the second out of line with the other three and located below
front margin of cell near origin of subcosta. There are three of these
spots just beyond end of cell, one near lower angle of cell at the base
of veins 3, 4, 5, one near upper angle of cell at the base of veins 6 and
7+8-+-9 and the third just above these on costal margin; three or four
submarginal spots cross the wing from costa to hind margin, these
usually very indistinct. In the middle of cell near the outer end is a
minute orange or orange-yellow spot. Forewing below pale brown,
whitish along middle of costa, fringe dirty white above and below.
Hindwing above and below pale brown or light fuscous, base much
lighter, sometimes dirty white. A large patch of white or grayish
white specialized scales along middle of costa covered by forewing.
Length of forewing, 8—9.5 mm.
Male genitalia as illustrated (pl. 10, fig. 2) and with characters as
given in the key.
Female.—Habitus entirely like the male.
Length of forewing, 10 mm.
Female genitalia as illustrated (pl. 11, fig. 8) and with characters
as given in the key.
Type locality —Jamaica.
Additional type data.—Described from the holotype (as type), female
(Jamaica; J. J. Bowry; 2 genitalia preparation, British Museum No.
1947-272) and from a male (same locality as female) not designated
type in original description.
Location of type.—In the British Museum (Natural History.)
Distribution. —Jamaica: St. Andrew Parish, Kingston, Mount
Mansfield House, Gordon Town (July); Clarendon Parish, Milk River
(August); Manchester Parish, Mandeville (June, July, November),
Newport (February); St. Elizabeth Parish, Balaclava (June). Locality
in Jamaica for which the parish is uncertain: May Hill (July).
Twenty-six specimens examined.
Remarks.—The genitalia of the type of M. grisea was compared with
the figure of the female of this species (pl. 11, fig. 8) by D. S. Fletcher,
of the Department of Entomology, British Museum (Natural History).
He found the type of grisea to be the species I have illustrated.
MOTHS OF GENERA MULONA AND LOMUNA—FIELD 225
2. MULONA SCHAUSI, new species
PuateE 10, Fiaures 4, 4A; Puatre 11, Figure 9
Male.—Habitus very much like that of Mulona grisea Hampson.
All dark spots on forewing above slightly more indistinct, orange spot
as in grisea. Hindwing above paler, nearly white, slightly darker
along outer angle and with a large dark brown patch of specialized
scales along middle of costal margin, extending to below front margin
of cell, this area covered by the forewing. Abdomen white above,
darker, more brownish below.
Length of forewing, 10-11 mm.
Male genitalia as illustrated (pl. 10, fig. 4) and with characters as
given in the key.
Female.—Habitus similar to that of male, forewing above suffused
with a dirty yellowish color, particularly along outer margin. Hind-
wing above and below entirely light fuscous with white fringes.
Length of forewing 10 mm.
Female genitalia as illustrated (pl. 11, fig. 9) and with characters
as given in the key.
Type locality—Matanzas, Province of Matanzas, Cuba.
Additional type data.—Described from the holotype, male (locality
as listed above, type U. 8S. N. M. No. 34830; collection William
Schaus; o genitalia slide W. D. F. No. 1655, 1941); allotype, female
(Santiago de las Vegas, Province of Habana, Cuba; January 24, 1933;
A. Otero; 2 genitalia slide W. D. F. No. 1661, 1941) and one para-
type, male (Province of Habana, Cuba; Father Roberto; @ genitalia
preparation, British Museum No. 88-1949).
Location of types.—Holotype and allotype in the United States
National Museum. Paratype in the British Museum of Natural
History.
Distribution —Cusa: Province of Habana, Habana, Santiago de las
Vegas (January); Province of Matanzas, Matanzas.
Three specimens (all known) examined.
Remarks.—Species named for the late William Schaus.
3. MULONA MANNI, new species
Puiate 10, Figures 5, 5A; Puare 11, Ficure 10
Male.—Very similar to Mulona grisea but smaller, usually with a
dark spot along middle of outer margin of patagium. Forewing
above with a marginal row of small dark spots not found in M. grisea
or MM. schausi, orange spot absent. Hindwing above and below
somewhat paler than in grisea and about as in schausi. Specialized
scales near base at costa and pale in color. Abdomen white to pale
brown.
Length of forewing, 6.5-8 mm.
226 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Male genitalia as illustrated (pl. 10, fig. 5) and with characters as
given in the key.
Female.—Habitus very much like the male, hindwing above entirely
white.
Length of forewing, 7 mm.
Female genitalia as illustrated (pl 11, fig. 10) and with characters
as given in the key.
Type locality Mangrove Cay, Andros Island, Bahamas.
Additional type data.—Described from the holotype, male (locality
as listed above; type U. S. N. M. No. 34831; William M. Mann;
o& genitalia slide W. D. F. No. 1663, 1941); allotype, female (same
locality and data as holotype; 2 genitalia slide W. D. F. No. 1664,
1941); paratypes Nos. 1-4, males (same locality as holotype; January
11, 1902; J. L. Bonhote), paratype No. 5, male (Bahamas; Carter;
Walsingham collection), and paratype No. 6, male (Little Abaco,
Bahamas; March 1902; J. L. Bonhote; @ genitalia preparation,
British Museum No. 274-1947).
Location of types——Holotype and allotype in the United States
National Museum. Paratypes Nos. 1-6 in the British Museum
(Natural History).
Distribution —Bauamas: Andros Island, Mangrove Cay (January),
Little Abaco (March).
Hight specimens (all known) studied.
Remarks.—Species named for Dr. Wiliam M. Mann, who collected
the holotype and allotype.
4. MULONA LAPIDARIA Walker
PLATE 10, Ficures 3, 3A
Mulona lapidaria WALKER, List of specimens of lepidopterous insects in the
British Museum, vol. 35 (supplement part 5), p. 1,896, 1866.—KrrBy, Syn-
onymic catalogue of Lepidoptera Heterocera, vol. 1, p. 366, 1892.—Hampson,
Catalogue of the Lepidoptera Phalaenae in the British Museum, vol. 2, p.
287, 1900.—Draupt, in Seitz, Gross-schmetterlinge der Erde, vol. 6, p. 252,
pl. 34, figure D 9, 1918.—Srranp, in Wagner, Lepidopterorum catalogus,
pars 26, p. 732, 1922.
Male.—This species is closely related to Mulona manni. Head and
thorax white tinged with pale brown. Forewing above white irro-
rated with brown, with marginal row of small dark spots as in manni.
The row of four small dark-brown spots crossing wing just before
middle are faintly connected, forming a zigzag line. There is a faint
black submarginal streak in lower half of wing nearly parallel to
outer margin and a similar streak starting at costa from just before
middle of wing, bending down from costa and continuing parallel to
costa to near apex, where it is bent upward to costa. With a minute
pale yellow spot in middle of cell near outer end. Hindwing above
white, outer margin and apex tinged with brown.
MOTHS OF GENERA MULONA AND LOMUNA—FIELD 297
Length of forewing, 9 mm.
Male genitalia as illustrated (pl. 10, fig. 3) and with characters as
given in the key.
Female.—Unknown.
Type locality.— Santo Domingo (“St. Domingo’).
Additional type data.—Described from a single male specimen, the
holotype (St. Domingo; Tweedie; genitalia preparation, British Mu-
seum No. 1947-275).
Location of type-—In the British Museum (Natural History).
Distribution.—Santo Dominco.
Remarks.—D. 58. Fletcher, of the Department of Entomology, Brit-
ish Museum (Natural History), was kind enough to dissect this holo-
type and send me a photograph of the genitalia. A drawing was
made from this photograph and this drawing was checked by Mr.
Fletcher for accuracy.
5. MULONA BARNESI, new species
Puate 10, Figures 6, 6A; Puate 11, Fiaure 11
Male.—Similar to the other species of Mulona. Forewing above
white, the small dark spots usually found in species of this genus are
greatly reduced and are missing in the lower half of the wing. Small
orange spot in middle of cell near outer end as in M. grisea and M.
schausi. Forewing below white suffused with pale brown. Hind-
wing above and below slightly brownish along outer margin.
Length of forewing, §.5-10 mm.
Male genitalia as illustrated (pl. 10, fig. 6) and with characters as
given in the key.
Female.—Habitus similar to that of male. Dark spots on forewing
above more abundant than in male and hindwing above light brown,
paler and whitish at base.
Length of forewing, 9 mm.
Female genitalia as illustrated (pl. 11, fig. 11) and with characters
as given in the key.
Type locality —Santiago de Cuba, Province of Oriente, Cuba.
Additional type data.—Described from the holotype, male (locality
as given above; type U.S. N. M. No. 34832; William Schaus; @ geni-
talia slide W. D. F. No. 1657, 1941); allotype, female (same locality
as holotype; collection Schaus and Barnes; September; 2 genitalia
slide W. D. F. No. 1660, 1941); paratype No. 1, male (same data as
holotype, o@ genitalia slide W. D. F. No. 1656, 1941) and paratype No.
2, male (Holguin, Province of Oriente, Cuba; H. S. Parish; @ genitalia
preparation, British Museum No. 1949-89).
Location of types ——Holotype, allotype, and paratype No. 1 in the
United States National Museum. Paratype No. 2 in the British
Museum (Natural History).
228 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Distribution.—Cusa: Province of Oriente, Santiago de Cuba (Sep-
tember), Holguin.
Four specimens (all known) studied.
Remarks.—Species named for the late John T. Barnes, old friend,
constant companion, and assistant to the late William Schaus.
LOMUNA, new genus
Genotype.—Mulona nigripuncta Hampson.
Labial palpus, antenna, and venation as described in the introduc-
tion.
Male genitalia (pl. 10, fig. 1) with uncus very short and broadly
bilobed; gnathos absent; anellus with ventrobasal plate broad and
rectangular, entirely fused to harpes and bifurcate at posterior end
and with arms very weakly sclerotized and very short, one-eighth or
less the length of harpes; transtilla present and partially fused across
middle; vinculum with saccus expanded, almost as long as broad and
rounded below; aedeagus thick throughout, distally broad and bilobed;
ductus ejaculatorius entering anterior end of aedeagus; vesica with
numerous scobinations; harpe deeply divided apically into three long
and slender arms.
Female genitalia (pl. 11, fig. 7) with ductus bursa sclerotized, cylin-
drical, not at all flattened; bursa copulatrix subspherical, slightly
longer than broad and heavily and completely serratulate through
middle two-thirds; seventh and eighth sternae not modified and weakly
sclerotized (without anterior and posterior genital plates) ; with bilobed
invaginated sac on dorsal surface immediately before ovipositor.
Remarks.—As defined above Lomuna contains a single species known
only from Puerto Rico (see remarks on type locality under L. nigri-
puncta Hampson).
LOMUNA NIGRIPUNCTA (Hampson)
Puate 10, Figure 1, 1A; Puatse 11, Ficure 7
Mulona nigripuncta Hampson, Catalogue of the Lepidoptera Phalaenae in the
British Museum, vol. 2, pp. 387, 388, pl. 29, fig. 6, 1900.—Dravupt, in Seitz,
Gross-schmetterlinge der Erde, vol. 6, p. 252, pl. 29, fig. L 9, 1918.—Srranp,
in Wagner, Lepidopterorum catalogus, pars 26, p. 732, 1922.—Forpss, Scien-
tific survey of Porto Rico and the Virgin Islands, vol. 12, pt. 1, p. 38, 1930.—
Wotcort, Journ. Agr., University of Puerto Rico, vol. 20, No. 1, p. 415, 1936.
Male.—Habitus somewhat like that of Mulona grisea Hampson but
with black spots on forewing more distinct and more nearly round.
Palpi dark fuscous except along ventral surface, where color is white.
Head, antenna, and thorax white, sometimes pale yellow or brown on
thorax. Forewing above white with black spots arranged as in grisea
but with an additional black spot in base of wing on the base of anal
vein. The orange or yellow spot found in the middle of the cell near
MOTHS OF GENERA MULONA AND LOMUNA—FIELD 229
the outer end in M. grisea, M. schausi, M. barnesi, and M. lapidaria
is present in M. nigripuncta and is orange or yellowish orange. Fore-
wing below nearly white along hind and outer margins, pale brown
over rest of wing. Hindwing above and below entirely white.
Specialized scales found in grisea near costal margin of hindwing are
absent in nigripuncta. Legs as in grisea.
Length of forewing, 8-8.5 mm.
Male genitalia as illustrated (pl. 10, fig. 1).
Female.—Habitus similar to that of the male. Forewings above
irrorated with black scales. This irroration sometimes heavy and
therefore habitus sometimes much darker than male. Forewing
below entirely light fuscous, sometimes paler along hind margin.
Hindwing above and below pale fuscous, nearly white or pale gray in
base of wing.
Length of forewing, 8-9 mm.
Female genitalia as illustrated (pl. 11, fig. 7).
Type locality—Probably Puerto Rico. Norz: Hampson cites th
type locality as “Colombia” and his type is labeled “nigripuncta, Z.,
Columb., M. Ber.” in Zeller’s own handwriting. It is a specimen that
Zeller intended to describe and the specimen came originally from the
Berlin Museum. This species is known only from Puerto Rico and
is not known from Colombia. The species of this group and other
closely related groups are very limited in their distribution and hence
it is extremely doubtful that the type of nigripuncta came from
Colombia.
Additional type data.—Described from a single specimen, the holo-
type, female (locality cited as Colombia but probably is Puerto Rico;
collection Zeller; Berlin Museum; 92 genitalia preparation, British
Museum No. 1947-273). Hampson also cites Puerto Rico as a habitat
for nigripuncta but he does not list number of specimens or sexes for
this locality. He had evidently seen specimens other than the type
from Puerto Rico; however these were not before him at the time he
wrote his original description, hence the single specimen mentioned
above is indeed the holotype.
Distribution.—Purrto Rico: District of Guayama, Palmas Abajas,
near Guayama (June); District of San Juan, Manati (June), Bayamon
(July); District of Aguadilla, Lares (July); District of Mayagiiez, San
German (April).
Five specimens examined.
Remarks.—The genitalia of the type of nigripuncta was compared
with the figure of the female of this species (pl. 11, fig. 7) by D. S.
Fletcher of the Department of Entomology, British Museum (Natural
History). He found the type of nigripuncta to be the species there
illustrated.
‘230 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 102
PLATES
The drawings for figures 3 and 3A were made by the author from a
photograph of the genitalia of the type. These two drawings were
corrected and modified by D. S. Fletcher, of the Department of
Entomology, British Museum (Natural History), by comparison with
the actual type preparation. Arthur D. Cushman, of the U. S.
Bureau of Entomology and Plant Quarantine, made the drawing of
the bursa copulatrix and ductus bursae in figure 8 (from an imperfect
specimen in the Carnegie Museum) and the drawings of the remaining
figures. The rest of figure 8 was drawn from the type by Mr. Fletcher.
The following figures were based upon the preparations of the holo-
types of the respective species: 3, 3A, 4, 4A,5,5A,6and6A. Figures
9, 10, and 11 were drawn from the allotypes of the respective species.
Figures 1 and 1A were drawn from slide W. D. F. No. 1651, 1941.
Figures 2 and 2A were drawn from slide W. D. F. No. 1658, 1941.
Figure 7 was drawn from slide W. D. F. No. 1662, 1941. These three
slides were all compared with the genitalia preparations of the types
in the British Museum (Natural History).
The size of the genitalia on these plates is indicated by the straight
lines placed near the figures. These lines indicate 1 mm.
U.S. GOVERNMENT PRINTING OFFICE: 1952
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 10
1, 1A, Lomuna nigripuncta (Hampson): 1, Male genitalia; 1A, aedeagus. 2, 2A,
Mulona grisea Hampson: 2, Male genitalia: 2A, aedeagus. 3, 3A, M.
lapidaria Walker: 3, Viale genitalia; 3A, aedeagus. 4, 4A, M. schausi, new
species: 4, Male genitalia; 4A, aedeagus. 5, 5A, M. manni, new species:
5, Male genitalia: 5A, aedeagus. 6, 6A, M. barnesi. new species: 6, Male
genitalia; 6A, aedeagus.
NO Jeyvesais: ih!
PROCEEDINGS, VOL.
U.S. NATIONAL MUSEUM
lulona grisea
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 Washington : 1952 No. 3299
A NEW SPECIES OF COMMENSAL AMPHIPOD FROM A
SPINY LOBSTER
By Ciarence R. SHOEMAKER
In January 1942 the late E. F. Ricketts, in connection with the work
of his Pacific Biological Laboratory, at Pacific Grove, Calif., examined
a living spiny lobster Panulirus interruptus (Randall) at the local
fish market. The lobster, which was presumably sent from Santa
Barbara, had some amphipods adhering to its pleopods, and six of
these were sent to me for identification. I find that these specimens
represent a new species. The structure of the peraeopods indicates
that they were modified and developed for the purpose of grasping,
and the animals appear to have been living commensally upon the
pleopods of the lobster. The mouthparts are of the normal type and
are not modified in any way, thus indicating that the animal is not
parasitic. The specimens are all females possessing partially de-
veloped marsupial plates.
PARAPLEUSTES COMMENSALIS, new species
Figure 83
Female—Head with rostrum rather short and blunt; lateral lobes
rounding; eye rather large, black, and reniform. Antenna 1: Pedun-
cular joints short; first joint not twice as long as second; second joint
not twice as long as third; flagellum about twice as long as peduncle
and containing 14 joints (an unknown number of terminal joints are
missing). Antenna 2: Peduncle short; fourth joint about twice as
long as third and equal in length to the fifth; flagellum a little longer
than peduncle and consisting of 13 joints.
231
967610—52
232 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Right mandible with 10 or 11 teeth in cutting plate; no accessory
plate; molar conical with very small triturating surface; 12 spines
in spine-row; palp with third joint slightly longer than second and
not apically produced. Maxilla 1: Inner plate small with one plumose
apical seta; outer plate with nine spine-teeth; palp with eight apical
spines, and three submarginal setae on the outer surface. Maxilla 2:
Inner plate with one long plumose seta on inner margin. Maxilliped:
Inner plate reaching to the base of the first joint of palp, armed distally
with four or five very short spine-teeth and two slender spines, and on
the inner margin with one long spine; outer plate reaching a little
beyond the base of the second joint of palp, armed on inner edge with
eight slender submarginal spine-teeth and on the outer surface with
six small submarginal spinules; palp with first three joints subequal
in length.
First four coxal plates a little deeper than their respective body seg-
ments. Gnathopod 1 as shown in fig. 83, 7; sixth jomt with palm
oblique, convex, armed with a row of submarginal spinules, and defined
by two spines, below which on the hind margin of joint is another pair
of spines; seventh joint fitting palm. Gnathopod 2, like gnathopod 1,
but a little longer. Peraeopods 1 and 2 alike; sixth joint strongly
developed, hind margin armed with seven pairs of stout spines against
the distal four of which the seventh joint closes, forming a grasping
organ. Peraeopods 3 to 5 are alike, but the fourth is somewhat the
longest; second joint well expanded; sixth joint strongly developed
and armed on the front margin with groups of stout spines, which to-
gether with the seventh joint form an effective grasping organ.
The lower posterior angle of the metasome segments minutely and
sharply produced. Uropods as shown by Sars for Stenopleustes
malmgreni (Crustacea of Norway, 1893, vol. 1, pl. 125, fig. 1) except
that the outer ramus of uropod 3 is proportionately shorter in Sars’
figure. Uropod 1 reaching a little farther back than uropod 2.
Uropod 3 reaching back about as far as uropod 2. Telson reaching to
the distal end of peduncle of uropod 3, three-fourths as wide as long
with the convex sides converging to the evenly rounding extremity.
Length from front of head to end of uropods 5.5 mm.
Type.—A. female, U.S.N.M. No. 85260, taken from the pleopods of a
spiny lobster Panulirus interruptus, from Santa Barbara, Calif., Janu-
ary 22, 1948, by E. F. Ricketts.
A NEW COMMENSAL AMPHIPOD—SHOEMAKER 933
Ficure 83.—Parapleustes commensalis, new species, female: A, Head; B, man-
dible; OC, maxilla 1; D, maxilliped; E, lower lip; F, gnathopod 1; G, gnathopod
2; H, peraeopod 2; J, peraeopod 3; J, metasome; K, telson.
U_ S. GOVERNMENT PRINTING OFFICE: 1952
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
issued
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 Washington; 1951 No. 3300
A NEW GENUS OF CENTRAL AMERICAN MILLIPED
(FAMILY EURYURIDAE), WITH NOTES ON THE AMER-
ICAN GENERA.
By Ricuarp L. Horrman
WHILE sortiINne out unidentified millipeds in the collection of the
United States National Museum, I encountered a single male speci-
men that represents an unnamed genus (here described) of the family
Euryuridae. Investigation of pertinent literature has revealed sev-
eral taxonomic errors that may be corrected at this time, and it appears
that at least one genus (Polylepiscus) is readily separable into two
more natural groups.
Together with these various additions and emendations, I include
new distributional records for two species of Amplinus, and a consid-
eration of the known American euryurid genera with reference to
some diagnostic characters whose value seems questionable. A tenta-
tive key to the genera is provided.
The Euryuridae is a small family (12 genera and about 50 species)
with many of the characters of the Platyrrhacidae and with nearly the
same distribution. Both families are represented in the Indo-Aus-
tralian region and in tropical America; species of Huryurus and
Auturus also invade the North Temperate Zone as far as Ohio and
Minnesota.
The most recent, and only complete, account of the Euryuridae is
to be found in Attems’ monumental Polydesmoidea monograph (Das
Tierreich, Lief. 68-70, 1937-1940). This manual lists all known spe-
cies and because of its conservatism affords an excellent starting point
for taxonomic studies. Attems combines the Euryuridae and
235
967609—52
236 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Platyrrhacidae (Lief. 69, p. 202), the latter being represented in his
arrangement by the single genus Platyrhacus. Seven other genera of
Attems’ “Platyrrhacidae” have the characters of the Euryuridae as
here understood. Since the appearance of Attems’ work, Chamberlin
has described a number of species and genera from northeastern Peru
(1941, Bull. Amer. Mus. Nat. Hist., pp. 498 et seq.).
I wish to express my indebtedness to Dr. E. A. Chapin, curator,
division of insects, United States National Museum, who has greatly
facilitated my work at that institution; to R. L. Wenzel, division of
insects, Chicago Natural History Museum, for the loan of many
specimens from collections under his charge; and to Dr. Clarence J.
Goodnight, department of biological sciences, Purdue University, who
has kindly given me valuable material from his Central American
collecting. |
REMARKS ON TAXONOMIC CHARACTERS
Attems’ key to the five American genera recognized by him is based
to a considerable extent upon sculpture of the dorsum. H'wryurus and
Pycnotropis are separated off in couplet 4 by the statement “Metazoni-
ten glatt oder nur mit Spuren einer polygonalen Felder,” as opposed
to “Metazoniten mit Querreihen grosser Tuberkeln,” which leads to
Polylepiscus and Amplinus.
This reliance upon tergite modification leads to considerable difii-
culty. Even in the four genera mentioned, one finds smooth species
in “tuberculate” genera, and vice versa. The issue is complicated by
the addition of Chamberlin’s four Peruvian genera and one from
Panama to be described herein. There seems to be such variability in
the development of polygonal areas on the dorsum that its practical
value is largely negated. The new Panamanian form keys out to
Pycnotropis, and the gonopods are of the type found in that genus, but
other characters are those of Phinotropis (Polylepiseus, in part, of
Attems). Because of the difficulty attendant upon the use of sculp-
ture, I believe it advisable to appeal to other characters, such as sub-
antennal swellings, anal sternite, and male gonopods, for generic
diagnosis.
The following key appears to be satisfactory at present for the rec-
ognition of American genera:
1The genus Aphelidesmus has been placed in the Euryuridae by recent American workers
at one time or another, since the time of Cook (1895) and Pocock (1909). Despite its
somewhat truncate terminal segment, Aphelidesmus properly belongs in the family
Strongylosomidae, a group with somewhat distant affinities.
EURYURID MILLIPEDS—HOFFMAN 237
TENTATIVE KEY TO THE AMERICAN GENERA OF EURYURIDAE
1. Anal sternite distally truncate, with two large setiferous tubercules.
Amplinus
Anal sternite semicircular, with very small tubereules or none______--____ 2
2. Male gonopod without solenomerite ; tibiotarsus, if present, not set off from
Ef CEVA TD Tsp eee ee a ra | a Re og te na ate yar aie eee ee ee 3
Male gonopod with a definite solenomerite branech______________--_____-__ 4
3. Gonopod without perceptible tibiotarsus_— ~__~ Net Neches on ee Oe AL ED Auturus
Gonopod with a long slender tibiotarsal portion___________-_____- Euryurus
4. Gonopod with two tibiotarsal branches____-__.______________ Polylepiscus
Gonopod* with, 2.singlertibiotarsal’ processs2s = - 2) Biles Se se eee 5
5. Tibiotarsal portion of gonopod broad, short, laminate, solenomerite very
SNODG ee == eae ee a es SR ie eT a ae i ae eee 6
Tibiotarsus long, slender, usually terete in cross-section, solenomerite
GTO S79] 1a ee a ce er er eee fe NAS ee Fa {5
6. Tergites with three transverse rows of large tubercules___ Protaphelidesmus
Meroiteseompletely Smooth. =e fe ee Ptyxogon
T Aclarge ovoid. swelling below each antenna_-2@_~._____L__4_ ee 8
No swellings understhe Antennae. [22 he ee ee Pycnotropis
8. Distal end of tibiotarsus of gonopod pointed, or at least unbranched.
Phinotropis
Mistaleend.of tibiotarsus variously modified *2—=——~_2-=*_-s—t see 9
9. Tibiotarsus straight or simply curved, parallel sided, distally bifid.
Thrinoxethus
Tibiotarsus long and slender, bent into a sigmoid curve, with the terminal
CNG et ril ewe eee eke 2d ea ye a) ete EE a Sigmogonotropis
Family EURYURIDAE Pocock
Buryurinae Pocock, Biologia Centrali-Americana, Diplopoda, p. 147, 1909.
Genus AMPLINUS Attems
Pachyurus (Amplinus) AtteMs, Denkschr. Akad. Wiss. Wien, vol. 68, p. 281,
1899. (Genotype: Polydesmus klugi Brandt.)
Range—Mexico (Veracruz and Hidalgo) south to Costa Rica.
Species—Amplinus abtrusus (Karsch), areatus Pocock, armatus
Pocock, erenus Chamberlin, convexus (Carl), erichsoni (Brandt),
flavicornis Pocock, klugi (Brandt), manni Chamberlin, niteuws Cham-
berlin, nitédus (Brélemann), orphinus Chamberlin, palicaudatus
(Attems) , tapachulae Chamberlin, triramus Pocock, vergelanus Cham-
berlin, and velitdws Chamberlin.
AMPLINUS FLAVICORNIS Pocock
Amplinus flavicornis Pocock, Biologia Centrali-Americana, Diplopoda, p. 151,
pl. 11, fig. 2, 1909.
Cutapas: Palenque Ruins, near Palenque, July 8, 1949, C. J. Good-
night collector.
This is the first definite locality for the species, which was described
from “Central America.” The specimens at hand match the original
238 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
description closely. The prozonites are very dark brown, the meta-
zonites light brown or tan; keels, antennae, legs, and last tergite yellow.
In the male the apical prongs of the gonopod are not bent to the extent
shown in Pocock’s figure, but the difference in this respect is hardly
significant.
AMPLINUS ORPHINUS Chamberlin
Amplinus orphinus CHAMBERIN, Proce. U. S. Nat. Mus., vol. 60, art. 8, p. 42, pl. 16,
fig. 8, 1922.
Britisn Honpuras: Middlesex, February 1, 1923, K. P. Schmidt and
L, Walters collectors. A single female tentatively referred to this
species (Chicago Natural History Museum).
GuatTeMALa: Chimaltenango, La Jolla Grande (Finca Monserrat),
northwest slope of Volcan de Fuego, 5,700 feet, May 3, 1948, R. L. Wen-
zel and R. D. Mitchell collectors (Chicago Natural History Museum).
Honvvuras: Rio Santa Ana, 2,500 feet, March 21, 1923, Schmidt and
Walters collectors (Chicago Natural History Museum).
This species has previously been recorded from La Ceiba, Honduras
(the type locality) and Livingston, Guatemala. The following locali-
ties represent considerable range extensions for orphinus, which
appears to be one of the most abundant and widespread members of
the genus. The record from Chimaltenango, Guatemala, is of especial
interest in carrying the range up into the high mountains.
Genus AUTURUS Chamberlin
Auturus CHAMBERLIN, Bull. Univ. Utah, biol. ser., vol. 6, No. 8, p. 7, 1942. (Geno-
type: Auturus phanus Chamberlin.)
Range.—Eastern United States. Mississippi Valley from Minne-
sota to Louisiana, east to Georgia.
Species.—Auturus dixianus Chamberlin, evides (Bollman), georgi-
anus Chamberlin, louistanus (Chamberlin), mimetes Chamberlin.
phanus Chamberlin, scotius Chamberlin.
Genus EURYURUS Koch
Furyurus Kocu, Krit. Rev. Insectf. Deutschl., vol. 3, p. 59, 1847. (Genotype:
Polydesmus erythropygus Brandt.)
Range.—Southeastern United States, north as far as Ohio.
Species.—Euryurus australis Bollman, erythropygus (Brandt).
EURYURUS AUSTRALIS Bollmen
Euryurus erythropygus australis BottMAN, Proc. U. S. Nat. Mus., vol. 11, p. 346,
1889.
Buryurus faicines Loomis, Bull. Mus. Comp. Zool., vol. 92, p. 403. fig. 15, 1943
(type locality: Torreya State Park, Liberty County, Fla.).
EURYURID MILLIPEDS—-HOFFMAN 239
Range.—The species is known to occur from west Florida north
through Georgia (Indian Springs), Alabama (Auburn), and Ten-
nessee to extreme northern Kentucky (Crittenden).
Loomis was misled into describing falcipes by a misunderstanding
of Bollman’s statement that the upper branch of the male gonopod
of australis is five times as long as the lower. This refers to the
branches in their position on the living animal. In Loomis’ descrip-
tion and figure, the gonopod is reoriented so that the actual position
is reversed. Bollman’s type of australis has been discovered at the
National Museum and its gonopods verify this idea, matching per-
fectly with the figure of falcipes.
Genus PHINOTROPIS Chamberlin
Phinotropis CHAMBERLIN, Bull. Amer. Mus. Nat. Hist., vol. 78, p. 499, 1941.
(Genotype: P. tidws Chamberlin.)
Range.—Northwestern Brazil, northeastern Peru, Ecuador.
Species—Phinotropis acuticollis (Attems), braueri (Carl), kalo-
notus (Attems), mammatus (Attems), ?rorert (Chamberlin), t¢dus
Chamberlin.
The original descr iption of Phinotropis is as follows: “This genus is
erected for the species described below in which the male gonopods
differ from those of 7Arinoxethus in having the major distal branch
entire and distally acute like the minor branch.”
Thrinowethus was characterized as follows: “Agreeing in general
structure with Polylepiscus but differing in the form of the gonopods
of the male. In these there are two distal branches of which one is
distally acute and the other, larger one, furcate at its distal end .. .”
Insofar as these two diagnoses alone go, it is obvious that Phino-
tropis is identical with Polylepiscus as used by Attems and others.
However, with the restriction of the latter name to Guatemalan species
having three terminal processes on the male gonopod, Chamberlin’s
name becomes available for the South American forms and must be
used, although originally a synonym.
Polylepiscus rorert Chamberlin, from Trinidad, is known only from
the female type specimen, and probably pertains to another, possibly
new, genus.
Genus POLYLEPISCUS Pecock
Polylepiscus Pocock, Biologia Centrali-Americana, Diplopoda, p. 154, 1909.
(Genotype: Polylepiscus stolli Pocock. )
Range.—Guatemala.
Species —Polylepiscus actaeon Pocock, alah Pocock, heteros-
culptus (Carl), stolli. Pocock.
The considerable distance separating the ranges of the Guatemalan
and upper Amazonian species hitherto cored to Polylepiscus fur-
240 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
ther serves to emphasize the lack of any close affinity between the two
groups.
Genus PROTAPHELIDESMUS Brélemann
Protaphelidesmus BROLEMANN, Ann. Soc. Ent. France, vol. 84, p. 559, 1916.
(Genotype: Platyrrhacus ligula Brélemann. )
Range.—Venezuela.
Species —Protaphelidesmus ligula Brélemann, levigatus Attems.
Genus PTYXOGON Chamberlin
Ptyxogon CHAMBERLIN, Bull. Amer. Mus. Nat. Hist., vol. 78, p. 500, 1941. (Geno-
type: Ptyzogon incus Chamberlin. )
Range.—Northeastern Peru.
Species.—Ptyxogon incus Chamberlin.
Genus PYCNOTROPIS Carl
Pycnotropis CARL, Mem. Soc. Sci. Nat. Neuchatel, vol. 5, p. 932, 1914. (Genotype:
Polydesmus (Euryurus) taenia Peters.)
Amydrinus CHAMBERLIN, Bull. Amer. Mus. Nat. Hist., vol. 78, p. 500, 1941.
(Genotype: Amydrinus pongus Chamberlin. )
Range.—Northeastern Peru, Ecuador, Colombia, Panama.
Species—Pycnotropis devillei (Silvestri), flavocarinata (Silves-
tri), haenschi Carl, latzeli Attems, polygonata (Gervais), pongus
(Chamberlin), taenia (Peters).
SIGMOGONOTROPIS, new genus
Genotype.—Sigmogonotropis serratus, new species.
Diagnosis—Head smooth, clypeal furrow distinct, a prominent
ovoid swelling under each antennal socket. Collum and other tergites
smooth, underparts without tubercules or bristles. Anal sternite semi-
circular, without enlarged tubercules. Gonopod of male as follows:
Coxa small, without hairs; prefemur and femur coalesced, straight,
setose. Distally, set off by a definite constriction, are a long tubular
solenomerite and a long, slender, sigmoidally curved tibiotarsus, ter-
minally trifid and with a broad subterminal toothed Jamia.
This genus seems to be intermediate between Pycnotropis and Phi-
notroms.
SIGMOGONOTROPIS SERRATUS, new species
FIGURE 84
Type specimen.—Male holotype, U.S.N.M. No. 1900, collected at
Cana, east Panama, by E. A. Goldman, June 1912.
Diagnosis —Characterized by the configuration of the male gonopod
as represented in figure 84.
241
EURYURID MILLIPEDS—-HOFFMAN
‘(uMoYs InuTayzaid UO aves []e JOU) yWeadse oIeYdad ‘sngvssas stdosjouos0Ms1g Jo podouod ajeul yoT—'Fg IANSIY
i)
/;
242 PROCEEDINGS OF THE NATIONAL’ MUSEUM VoL. 102
Description of holotype—Specimen much broken. Approximate
length, 51 mm.; greatest width, 6.9 mm.
Head entirely smooth, glabrous except for a few fine hairs near
labral margin. Clypeal furrow very distinct. A prominent trans-
verse ovoid swelling directly below each antennal socket. Antennae
rather stout, short, reaching back to anterior margin of third tergite;
articles 2-6 of nearly equal size.
Collum perfectly smooth, with a fine but distinct margining ridge
except on caudal edge. Anterior edge nearly straight, caudal edge
bent forward laterally. Lateral ends slightly truncate and marginal
ridges more prominent than across back. Second tergite wider than
collum and third tergite, its keels pronouncedly swept forward.
Tergites completely smooth, except for extremely faint suggestions
of polygonal areas on the keels, to be seen when the dorsum is dry.
Keels of moderate width and length, never overlapping, set above
middle of body and nearly horizontal, thus interrupting slope of
the rather convex dorsum; anterior corners rounded off, posterior
corners produced caudad into a point on all keels back of third seg-
ment. Both anterior and posterior edges are set off by a marginal
ridge. Lateral edges inflated and smooth but for a slight notch on
the anterior shoulder. On poriferous keels the pores are lateral in a
large swelling. On the 19th segment pores are definitely inferior on
the small, distally rounded keel. Last tergite broadly truncate,
nearly tongue shaped, lateral edges almost parallel. A small median
terminal lobe.
Anal valves smooth, with well-defined median ridges. Preanal
sternite nearly semicircular without lateral setiferous tubercules.
Last pair of legs separated by a distance equal to diameter of their
coxae; penultimate pair slightly farther apart. Bases of all legs fused
into a raised area very distinct from and higher than the prosternites.
Legs long and slender, end of third joint visible past keels in dorsal
aspect. Lengths of joints of legs, in order of decreasing length, 3,
6, 5, 2,4, 1. Terminal claw slender and nearly straight. No special
processes associated with any of the legs.
Sides and underparts all smooth and glabrous. A deep constric-
tion between prozonite and metazonite. In going toward the head,
the sternites tend to develop a small conical tubercle at base of each
leg.
Gonopod aperture of medium size, oval, with caudal and lateral
raised edges. Pregenital legs unmodified. Seminal apertures small,
in end of coxae of second pair of legs, no special processes for them.
Gonopods: Coxa short with a long slender trachial process and a
small hook just above exsertion of solenite. Prefemur and femur fused
into a straight, heavy, upright trunk, somewhat concave on one side
and densely setose; the opposite side is traversed by the seminal
EURYURID MILLIPEDS—-HOFFMAN 243
groove. Terminal elements of gonopod (solenomerite and tibiotar-
sus) set off by a distinct joint. Solenomerite long, gently curved
across tibiotarsus, with a small distal expansion. ‘Tibiotarsus sig-
moidally curved, with a subterminal lateral lamination, the outer edge
of which bears three teeth; distally trifurcate with the median tooth
the longest.
Original color completely lost through preservation.
Genus THRINOXETHUS Chamberlin
Thrinowvethus CHAMBERLIN, Bull. Amer. Mus. Nat. Hist., vol. 78, p. 498, 1941.
(Genotype: Thrinozvethus hermosus Chamberlin.)
Range.—Northeastern Peru (Department of Loreto).
Species.—Thrinoxethus bombonus Chamberlin, cainarachus Cham-
berlin, hermosus Chamberlin, iguitus Chamberlin, lamprus Chamber-
lin, nétens Chamberlin, phanotypus Chamberlin, ucayalus Chamberlin.
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SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 Washington: 1952 No. 3301
AN EMENDED DIAGNOSIS OF THE COPEPOD GENUS
PUPULINA (CALIGOIDA), WITH DESCRIPTIONS OF
NEW SPECIES AND A REDESCRIPTION OF THE GENO-
TYPE
By Mivprep Srratron WILson
THE CALIGOID copepod parasite Pupulina has been an imperfectly
known genus since its discovery in 1892. The original description by
van Beneden was incomplete, and attached to it, as the description
of the male, was the diagnosis of an immature female of another genus.
Not until 1935 was its validity definitely established, by Dr. Charles
Branch Wilson, through the description of both sexes of the genotype,
Pupulina flores. One subsequent record of the species has been made,
that of Bere (1936).
This seeming rarity and the incompleteness of knowledge that
always results when but a single member of a genus is known add
interest to the recent discovery of two new species. These occurred
together on a specimen of the giant ray M/obula lucasana Beebe and
TeeVan, taken off the coast of California. The copepods were re-
moved from its surface by Prof. G. E. MacGinitie, who referred them
to the United States National Museum for identification. The host,
the second of its kind to be drawn to the attention of science, was
also of particular interest, and salient facts concerning it have already
been noted in a publication by MacGinitie (1947).
The collections of both Wilson and Bere are in the United States
National Museum and thus, fortunately, available for study. A
direct comparison of the new species with Pupulina flores has made
possible a new, nearly more complete diagnosis of the genus; ; especially
has it mmaeated the importance of certain characters fatherts not
stressed or noted. In addition, it has been found that some of the
245
246 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
specimens identified by Bere as Pupulina flores are referable to the
smaller of the two new species.
A new description, based on the Wilson and Bere specimens, has
been prepared for the genotype, as much detail having specific value
has been previously omitted. Development forms, present in his ma-
terial and not mentioned by Wilson, are also described.
For critical readings of the manuscript I am indebted to Dr. Charles
H. Martin, of Oregon State College, and to Paul L. Illg, of the United
States National Museum.
HOST RECORDS
According to each of the three published instances of the occurrence
of Pupulina, it has been found on a species of large ray belonging
to the family Mobulidae. From this fact and MacGinitie’s discovery
it would appear that the genus may be a specific parasite of this
group of rays. It seems very probable that records of its occurrence
have been infrequent because the hosts themselves are seldom caught.
In this connection, it is necessary to qualify Wilson’s statement
(1935a, p. 593) that his specimens, taken near the Galapagos Islands,
were from “Manta birostris, the same species of fish as that from
which Beneden’s specimens were collected.” Van Beneden did not
so name the species of the host, merely alluding to it as Ceraptopterus.
The latter is a genus of Coleoptera, and so far as is known to me,
the name has not been used in the fishes. From van Beneden’s de-
scription of the size of the host, it is probably safe to assume that he
had confused the name with Ceratoptera Miller and Henle, a
synonym of Manta. Van Beneden’s specimens occurred off the Azores,
a locality considerably removed from the Galapagos, and whether the
host was the identical species of Manta is certainly open to question.
Genus PUPULINA P. J. van Beneden
Pupulina VAN BENEDEN, 1892, p. 254.—C. B. Witson, 1935a, p. 593.
Diagnosis (emended).—Body form of more or less specific varia-
bility in females; strikingly similar in males.
Frontal plates well defined, without lunules. First three thoracic
segments fused with head; fourth segment free, without dorsal plates.
Lateral, cephalic, and thoracic areas of carapace demarcated dorsally
by well-defined grooves; ventral supporting ribs of lateral areas stout,
the posterior of double rods, their outlines usually conspicuous
dorsally.
Genital segment of female moderately enlarged, its width greater
than that of free segment, about one-half to three-fifths that of
carapace; the posterior corners produced into backwardly directed
COPEPOD GENUS PUPULINA (CALIGOIDA)—WILSON 247
lobes; dorsal surface with a large medial clear area in which the
integument is considerably thinner than that surrounding it, creating
in the mature forms a characteristic design for each species. Abdomen
narrow and elongate in both sexes, 3-segmented in the female, the divi-
sion between the first two segments sometimes indistinct ; 2-segmented
in the male. Caudal rami linear in male; either linear or short and
broad in female; terminal setae very short, sparsely plumose or naked.
Second antenna of female with either two or three segments well
defined, apical claw stout, basal segment with a strong, posteriorly
directed spinous process; that of male 3-segmented, tipped with a
stout claw and armed variously with laminae and spines. Prehensile
lateral hooks present, the basal portion bearing posteriorly two papil-
lae armed with branched or simple setae.
Oral appendages similar to those of the Caliginae. Mouth tube in
female constricted near the middle, its greatest width a little more
than half the length of the entire tube; the whole more slender and
hardly constricted in the male; opening subterminal. Mandible 4-
segmented, segments 2 and 3 imperfectly defined, claw heavily chitin-
ized, with teeth on the inside margin only.
First maxilla consisting of two heavily chitinized pieces and a very
small, membranous papilla, the outer chitin piece having its outside
edge more or less produced distally, being extended in the genotype
into an elongated process; the inner piece (usually referred to as
endopod by recent writers) extended posteriorly into a slender spine-
like process, in the male either branched distally or bearing an articu-
lated spine. The papilla (the exopod of recent writers) arising from
the ventral face between the basal portions of the two chitin structures ;
bearing three comparatively short setae which exhibit sexual as well
as pronounced specific differences. Second maxilla elongated; first
segment imperfectly defined, represented by a slightly enlarged basal
portion; third segment very slender, with two thin, curving, terminal
claws, the posterior of which is much shorter than the other. Maxil-
liped of female with short terminal claw, not more than half the length
of basal segment; male having claw a little longer and the basal seg-
ment variously armed.
Two pairs of accessory processes arising from the ventral face,
sexually similar. The anterior pair spinelike, situated near distal
end of the inner chitin piece of the first maxilla. The posterior pair
heavily chitinized or membranous, shape specifically variable, located
behind the base of the maxilliped, directly opposite the posterior
supporting lateral ribs of the carapace, but in nowise connected with
them or with the central area usually occupied by the furca, which is
lacking.
Legs 1-3 biramose. Leg 1 with both rami 2-segmented, the endopod
well developed ; the inside spine of the second exopod segment charac-
QAS PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
teristically branched in each species. Curved chitin rods, sometimes
armed with spines or spinous processes, between bases of legs 1 and
2. Leg 2 with trimerous rami; outer spines of exopod 1 and 2 only
moderately developed; endopod 1 and 2 with outer marginal laminae.
Leg 3 having both rami 3-segmented, their bases set closely together.
Exopod well developed, the first segment elongate as in the second leg,
outer spines weaker. Endopod 1 and 2 with outer laminae, that of
segment 1 greatly enlarged and overlying the exopod, that of seg-
ment 2 much smaller; the second segment the longest, with two inner
setae arranged like those of the second leg; third segment approxi-
mately the same size as the first segment, with four terminal setae.
Leg 4 uniramose and stout; 4-segmented, the basal segment greatly
broadened; somewhat sexually dimorphic in armature; some species
having short spines on the inner margins of segments 2-4; the sutures
between these segments transverse. No rudiments of legs 5 and 6
visible on genital segment of adult, but present on the ventral side in
female chalimi; condition in immature male not observed.
Length of known females, 6-15 mm. ; of males, 4.8-12 mm.
Parasites of giant rays.
Genotype —Pupulina flores P. J. van Beneden.
PUPULINA FLORES van Beneden, new description
PLATE 12; PuLatTe 14, Ficures 8-11
Pupulina flores VAN BENEDEN, 1892, p. 254, pl. 3, figs. 6-8, female.
Lepeophtheirus flores, BASSETT-SMITH, 1899, p. 455.
Lepeophtheirus flores, C. B. WILSON, 1905, pp. 617, 618, in key.
Pupulina flores, C. B. Witson, 1935a, p. 594, figs. 1-13, female and male.
Pupulina flores, C. B. WiLson, 1935b, p. 331, refers van Beneden’s male to Para-
lebion elongatus female.
Pupulina flores, Bere, 1936, p. 590, the “large female.”
Specimens examined.—Six mature, two immature females; two ma-
ture males, U.S.N.M. No. 60489. Host: Manta birostris (Walbaum).
Locality: near the Galapagos Islands. Identified by Charles Branch
Wilson.
One female, U.S.N.M. No. 79150. Host: “probably Mobula hypos-
toma (Bancroft).” Locality: Lemon Bay, Fla., Gulf of Mexico.
Identified by Ruby Bere.
Diagnosis.—Anterior (cephalothorax) and posterior parts of body
approximately equal in length in both sexes. Processes of female
genital segment reaching a little beyond the middle of the abdomen.
Caudal rami attached terminally, linear; those of female only a little
shorter than the abdomen; those of male longer. Outer chitin piece
of first maxilla extended distally almost to tip of the process of the
inner piece; the latter bifurcate in male. Maxilliped of male with
a short papillalike process on the basal segment. Posterior ventral
COPEPOD GENUS PUPULINA (CALIGOIDA)—-WILSON 249
accessory processes large, heavily chitinized, triangular spines. Leg
4 without inside spines on segments 2-4, but with a cuticular process
on 4.
Description—FEMate (pl. 12, fig. 1) : Galapagos specimens: Total
length 13.9-15.0 mm. Carapace about 6 mm. long, 5.8-6.0 mm. wide.
Gulf of Mexico specimen: The single, nonovigerous female only 10
mm. in length, but otherwise like the Galapagos specimens.
Anterior margin of carapace curved, median sinus slitlike; posterior
margin nearly straight. Grooving of carapace as in pl. 12, fig. 1;
lateral lobes with long slender hairs dorsally; each side of distal edge
of thoracic area set with four or five spines. Free segment about three
times wider than long, with short dorsal spines posteriorly.
Genital segment joined by short neck to the free segment, rounded
anteriorly; posterior processes reaching slightly beyond midpoint of
second segment of abdomen, ending in slender, rounded tips, diverg-
ing inside directly from the base of the abdomen. Sides of segment
and parts of dorsal surface set with very short spinules. Design on
dorsal surface sloping in a wavy line from the rather straight top to
the widened central area; the posterior part narrowed with a marked
central incision of the entire bottom margin.
Abdomen narrow and elongate, a little longer than the genital seg-
ment in midline; the first segment very short and indistinctly defined ;
the second and anal segments plainly demarcated from each other, the
second twice the length of the anal; both distal segments with a midline
of long, coarse hairs and scattered surface spinules; anal segment with
a pair of dorsal longitudinal ridges distally.
Caudal rami a little shorter than the abdomen, flattened laterally,
proximally with a raised longitudinal ridge; dorsal margins set
with coarse hairs; terminal setae very short and nonplumose, three
in number. Ovisacs slender, longer than the caudal rami.
The first antenna (pl. 14, fig. 9) with a double row of stout plumose
setae on the upper side of the first segment and two short spines distad
on the lower side; the narrowed terminal segment having a slender
seta midway on the lower margin, and the apex set with numerous
short spines and varied setae (pl. 14, fig. 8).
Second antenna (pl. 12, fig. 4) 2-segmented, basal spinous process
very slender, terminal claw curved inwardly, only moderately
elongated. Lateral prehensile hooks (pl. 14, fig. 5) comparatively
long and strongly curved, setae of basal papillae branched. Mouth
tube as described for the genus; mandible with 12 teeth.
The anterior accessory process of the ventral face (pl. 14, fig. 7)
located near the distal inner side of the inner piece of the first maxilla ;
consisting of a very stout spine about one-fourth as long as this portion
of the maxilla. The outer piece of the first maxilla (pl. 12, fig. 8) well
developed, its outer edge elongated and spinelike distally, reaching
250 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
beyond the middle of the inner piece; the papilla seemingly closely
associated with the tissue of its inner basal portion. The three papil-
lary setae unequal and unlike (pl. 12, fig. 15), the shortest having an
accessory spine near its midline. The inner piece of the maxilla longer
than the extension of the outer piece, ending in a similar stout, slightly
curved spine.
The second maxilla (pl. 12, fig. 9) of the long slender form charac-
teristic of the genus, tipped with two slender, unequal claws with
marginal hyaline flanges, the posterior about half the length of the
other. The claw of the maxilliped (pl. 12, fig. 10) slender, about half
the length of the basal segment, bearing a stalked seta on its posterior
side. The accessory process just posterior to the base of the maxilliped
(pl. 12, fig. 11) a very broad, stout, chitinous spine, irregularly
triangular, with the distal point somewhat attenuated. (The basal
part of this spine is probably a bar corresponding to that found in
the other species, but the main body is so strongly chitinized that any
division between the two is obscured.)
Chitin rod between legs 1 and 2 narrow, curved backward, each
side armed with a stout upwardly directed spine. Form of leg 1 as
shown in plate 15, figure 14; endopod reaching to distal third of basal
exopod segment; terminal exopod segment with the upper two spines
stout and dentate on both margins, the modified inner spine and ac-
companying seta as shown in plate 12, figure 16.
Basal segment of leg 2 armed inside with a plumose seta and a
triangular lamina. Exopod (pl. 12, fig. 18) with outer spine of first
segment reaching to distal end of third; spine of third segment with
smooth margins, reaching considerably beyond distal edge of seg-
ment; second spine about one-third the length of the first, smooth
and slightly curved. Laminae on outer margins of endopod segments
1 and 2 similar to those shown in plate 15, figure 12, except that the
edges are more rounded. (There is no lamina on the inner edge of
the second segment as stated by C. B. Wilson, 1935a. As is usual,
the margin of the segment is extended beyond the point of attachment
of the succeeding segment, and bears two long setae at the distal
terminus of the extension.) The third segment rounded and very short,
bearing six setae.
The exopod of leg 3 having the basal segment as long as the two
succeeding segments, the outer spines of segments 1 and 2 smooth,
slender, and scarcely longer than the following segment. The inner
setae not so long as those of leg 2. Segment 3 (pl. 12, fig. 14) with
three outer unequal spines. The laminae of the outer margins of the
endopod attached as shown in plate 12, figure 17; that of segment 1
very large and overlying the first two segments of the exopod. The
inner Margin of segment 2 expanded as in leg 2.
COPEPOD GENUS PUPULINA (CALIGOIDA)—WILSON 951
As my findings disagree with some of those of C. B. Wilson (1935a),
the following summary of the armature of legs 1-3 is given (sp=
spine; se=seta; Jam=lamina) :
Exopod Endopod
Leg 1 sp 0
3sp 4se 3se
Leg 2 sp se lam se
sp se lam 2se
2sp 6se 6se
Leg 3 sp se lam 0
sp se lam 2se
3sp 4se 4se
Basal segment of leg 4 (pl. 12, fig. 20) with small spines on anterior
and posterior margins; second segment with similar spines along outer
margin; none on margins of segments 3 and 4. ‘Terminal outer spines
of segments 2 and 8 about as long as the succeeding segment, inner
spines lacking. Distal end of fourth segment with three stout spines
and one slender spine, the outermost shorter than the segment, the next
as long as, and the third a little longer than the segment; the inner
spine attached laterally, slender, curved, reaching beyond the end of
the segment by one-half its own length; a thin, cuticular process,
spinelike distally, present at about the middle of the inner margin.
All large major spines of the leg coarsely toothed, the longest of the
terminal segment having some of the teeth considerably enlarged on
the inner margin, with three stout additional teeth on the outer side.
Mate (pl. 12, fig. 3) : Total length 11.4-12.1 mm. Carapace about
equal in length and greatest width, 5.1-5.4 mm.; cephalothorax about
5.8-6.0 mm. long. Posterior part of body averaging 5.8 mm. in length;
of this the approximate divisions are: genital segment, 1.3 mm.; ab-
domen, 1.9 mm.; caudal rami (exclusive of setae), 2.6 mm. The
caudal rami are therefore longer than the abdomen, but are shorter
than the genital segment and abdomen combined. None of the speci-
mens, including the allotype (kindly measured for me by Paul Illg, of
the United States National Museum), exhibit the condition described
and figured by C. B. Wilson (1935a, p. 597) in which “the caudal
rami are as long as the entire body behind the carapace.”
Carapace similar to that of female except that the medial posterior
edge is somewhat curved. Free thoracic segment little more than
twice as wide as long; posterior part considerably widened at midline,
with long, stiff marginal hairs.
Genital segment with sides slightly rounded, set with conspicuous
spinules, both marginally and on dorsal and ventral surfaces; four-
fifths as wide as long; posterior corners produced dorsally, well
rounded ventrally. Basal segment of abdomen distinctly set off;
252 PROCEEDINGS OF THE. NATIONAL MUSEUM VOL. 102
anal segment not demarcated by discernible transverse grooving either
dorsally or ventrally in specimens examined, but with a pair of raised
longitudinal dorsal ridges as in female, particularly noticeable in
stained specimens. Both ventral and dorsal surfaces covered with
scattered spinules. Caudal rami slightly less than one-third longer
than the abdomen, flattened laterally as in female, with proximal
dorsal ridges, inside margins with long hairs, outside with spinules.
Second antenna (pl. 14, fig. 10) reduced in size but very stout; basal
segment unarmed, strongly united to ventral face of carapace (this
segment is apparently omitted in C. B. Wilson’s illustration, 1935a) ;
second segment stout and well rounded, projecting upward, armed
below with a broad but rather short spine, and a comparatively large,
2-layered, fringed lamina placed distad to the spine. Third seg-
ment with a stalked spine, segmented near its tip, and a stalked seta
at the base of the claw. Claw short, but very strongly curved, with
a rounded projection at its inner base. (The illustration of this ap-
pendage, pl. 14, fig. 10, is a flattened, semidiagrammatic view, turned
so as to include detail of all the armature.)
Lateral prehensile hooks scarcely larger than those of female, but
much more strongly curved, papillary setae unbranched. Anterior
accessory process like that of female, but comparatively smaller. In-
ner piece of first maxilla (pl. 12, fig. 12) bifurcate at tip, inner branch
the longer; basal portion of the outer piece broad, extended distally
into a long spine as in female; setae of papilla unequal (pl. 12, fig. 15).
Maxilla 2 as in female. Maxilliped (pl. 14, fig. 11) about twice the
length of the second antenna, with a stout claw bearing a stalked seta;
basal segment with a short, rounded process whose tip is more or
less demarcated by a roughened area. Posterior accessory process
behind maxilliped a stout triangular spine like that of female. Chitin
rod between legs 1 and 2 also resembling that of female.
Legs 1-3 with setal formula and laminae like female. Basal seg-
ment of leg 4 (pl. 12, fig. 19) armed marginally with spinules; more
spinules scattered over the upper part of the dorsal surface, and the
lower part with a patch of long, stiff hairs running crosswise. The
large spines of segments 2-4 more slender and comparatively longer
than those of female; all having toothed margins, that of the longest
terminal spine with the proximal teeth enlarged on the inner margin.
Length of the shorter inner spine not known, as it is unfortunately
broken off in all the specimens, including the allotype.
DEVELOPMENT Sraces: Two female chalimi, not mentioned by C. B.
Wilson in his report, are present in the Galapagos collection. In the
following descriptions, they are referred to as Stages a and 6.
Stace @ (pl. 12, fig. 2): Total length, 4.2 mm. All the first three
thoracic segments incorporated in the carapace, which is as wide as
long. Frontal plates indistinct. Free thoracic segment only slightly
COPEPOD GENUS PUPULINA (CALIGOIDA)—WILSON 253
wider than long, with small distal lobes; fourth legs attached near
the middle of the segment. Genital segment separate from abdomen,
widened anteriorly, narrowed posteriorly, with cuticular lobes on the
sides at the midpoint; behind these, another pair of lobes that repre-
sent the beginnings of the posterior processes. Ventrally, these latter
lobes carry the rudiments of leg 5, consisting of a small, rounded flap
hardly differentiated from the surface and bearing terminally two
short plumose setae; leg 6 not discernible in this specimen. Abdo-
men and caudal rami together only a little longer than the genital
segment. Abdomen showing no segmentation. Caudal rami flat-
tened dorsoventrally, with the plumose terminal setae longer than in
the adult.
Ventrally, the fore part of the body showing the distinguishing
characters of the species already developed. The outer piece of the
first maxilla well extended distally; both anterior and posterior ven-
tral accessory processes large and of distinctive shape, the posterior
process more membranous than in the adult, with the distal attenuated
point sharper. A pair of spinous processes between legs 1 and 2,
the connecting rod between them scarcely discernible.
The second antenna segmented like that of the adult, but more
slender, the terminal claw very strongly bent as in adult; prehensile
lateral hook comparatively shorter. The second maxilla not at all
strongly developed, the basal segment being shortened and not ex-
tending outward beyond the maxilliped as in the adult; the terminal
claws (pl. 12, fig. 2, a@) not demarcated basally from the apex, the
posterior claw very short, broad and dentate.
Leg 1 with both rami 2-segmented and all setae present, the branch-
ing of the modified spine of the exopod incomplete. Rami of legs
2 and 3 only 2-segmented, the laminae not developed. Leg 4 rela-
tively small, consisting of the basal and two other smaller segments.
Srace 6 (pl. 12, fig. 13): Total length about 9.3 mm. Metasome a
little longer than urosome. Carapace slightly longer than wide.
Fourth legs attached near the distal end of free segment, the widened
area behind having disappeared; distal edges of segment somewhat
angular.
Genital segment with processes reaching to about the proximal
third of abdomen; rudiments of leg 5 on the ventral side having three
setae, one lateral, two terminal, all shorter than the two setae of the
preceding stage. Rudiments of leg 6, consisting of a very much ab-
breviated lobe and a single seta, located outside and slightly anterior
to leg 5.
Abdomen 2-segmented dorsally, the short basal segment of the
adult indistinctly demarcated ventrally; the anal segment distally
with a pair of dorsal longitudinal lines, undoubtedly the beginnings of
967612522
254 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
the prominent ridges of the adult. Caudal rami a little shorter than
the abdomen, terminal setae shorter than in Stage a.
All appendages considerably larger and more chitinized than in
Stage a. Outer piece of maxilla 1 a more slender spine than in
adult; claws of second maxilla longer, but of same general shape as in
Stage a. Rod between legs 1 and 2 well developed. All legs with
rami completely segmented; leg 4, however, not large, and its spines
weak and flattened.
This chalimus is almost as long as Bere’s specimen from the Gulf of
Mexico which, however, is as completely developed as the adult.
Hosts and distribution —Manta sp., Atlantic Ocean, near the Azores
(van Beneden) ; Manta birostris (Walbaum), Pacific Ocean, near the
Galapagos Islands (C. B. Wilson) ; “probably” Mobula hypostoma
(Bancroft), Gulf of Mexico, in Lemon Bay, Fla. (Bere).
Remarks.—The identification of these Pacific and Gulf of Mexico
specimens with the female of the type species from the Atlantic
would seem from a comparison with van Beneden’s original descrip-
tion and illustrations to be correct. His drawing of the female shows
the same general shape of the various parts of the body, and the dorsal
design on the genital segment is closely similar, the midinvagination
of the posterior margin being characteristic of the species. His illus-
tration differs in that the anterior lobe of this design is considerably
narrower and the sides longer. In his figure, also, the posterior inside
margins of the genital segment are more rounded with the corners
less produced, and the abdomen is unsegmented and longer in pro-
portion to the genital segment.
The only illustrations van Beneden gave of the appendages are
some very incomplete figures of those of the cephalic area. These,
however, show two important characteristics of the species: The ex-
tension of the outer portion of the first maxilla into a slender spine,
and the triangular shape of the posterior ventral accessory processes.
As these are distinctive features of the Pacific and Gulf of Mexico
specimens it seems reasonable to assume that they are identical with
those of van Beneden.
The form described by van Beneden as the male of Pupulina flores
has been stated by C. B. Wilson (1985a, 1935b) to be an immature
female of Paralebion elongatus C. B. Wilson.
PUPULINA MINOR, new species
PratE 13
Pupulina flores, in part, BERE, 1986, p. 590, the ‘‘smaller’” specimens.
Specimens examined.—Two females, one ovigerous; two males; col-
lected by G. E. MacGinitie, from the giant ray Mobula lucasana Beebe
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 12
PUPULINA FLORES VAN BENEDEN
2, Female, dorsal view of chalimus, stage a.
3, Male, dorsal view. 4-11, Female, cephalic appendages in
situ: 4, Antenna 2; 5, lateral hook; 6, outline of mouth tube; 7, anterior ventral accessory
process; 8, maxilla 1; 9, apex of maxilla 2; 10, maxilliped; 11, posterior ventral accessory
1, Female, dorsal view of adult. 2, a, Claws
of maxilla 2, stage a.
12, Male, maxilla 1. 13, Female, dorsal view of chalimus, stage b. 14, Female,
leg 3, exopod segment 3. 15, Male and female, papillary setae of maxilla 1. 16, Female,
leg 1, modified spine of exopod. 17, Female, leg 3, endopod. 18, Female, leg 2, exopod.
19, Male, leg 4. 20, Female, leg 4. (Scale applies to figures 1-3 and 13.)
process.
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 13
PUPULINA MINOR, NEW SPECIES
1, Female, dorsal view. 2, Male, dorsal view. 3-9, Female, cephalic appendages in
situ: 3, Antenna 2; 4, lateral hook; 4, a, papillary seta of hook; 5, maxilla 1; 6, anterior
ventral accessory process; 7, maxilla 2; 8, maxilliped; 9, posterior ventral accessory proc-
ess. 10, Male, maxilla 1. 11, Female, papillary setae of maxilla 1. 12, Male, papil-
lary setae of maxilla 1. 13, Female, leg 2, exopod. 14, Male, maxilliped, with detail
of papilla. 15, Male, antenna 2, greatly enlarged. 16, Male, leg 4. 17, Female, leg 1,
modified spine of exopod. 18, Female, leg 4. (Scale applies to figures 1 and 2.)
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 14
PUPULINA BREVICAUDA, NEW SPECIES, AND P. FLORES VAN BENEDEN
Pupulina brevicauda: 1, Female, dorsal view of adult; 2, young female, dorsal view; 3,
male, dorsal view; 4, female, dorsal view of chalimus, stage b; 4, a, ventral view of genital
segment, stage b, with legs 5 and 6; 5, female, stage c, posterior part of body, dorsal view;
6, male, antenna 2, greatly enlarged; 7, female, terminal portion of antenna 1. P. flores:
8, Female, enlarged terminal portion of antenna 1; 9, female, antenna 1; 10, male, antenna
2, greatly enlarged; 11, male, maxilliped, with detail of papilla. (Scale between figures
1 and 2 applies to figures 1, 2, and 4; scales at figures 3 and 5 apply to those figures only.)
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 15
10,
PUPULINA BREVICAUDA, NEW SPECIES
1-7, Female, cephalic appendages in situ: 1, Antenna 2; 2, lateral hook; 3, anterior ventral
accessory process; 4, maxilla 1; 5, maxilla 2; 6, maxilliped; 7, posterior ventral accessory
process. 8, Female, papillary setae of maxilla 1. 9, Male, maxilla 1. 10, Male, papil-
lary setae of maxilla 1. 11, Female, mandible. 12, Female, leg 2. 13, Male, maxilliped, |
greatly enlarged. 14, Female, leg 1. 15, Female, leg 1, modified spine of exopod. |
16, Female, leg 3, exopod segment 3. 17, Female, leg 4. 18, Male, leg 4.
COPEPOD GENUS PUPULINA (CALIGOIDA)—WILSON 255
and TeeVan; off Santa Catalina Island, Calif.; October 3, 1946.
Three females, two ovigerous; three males; collected by Ruby Bere,
“around the mouth of a devilfish (probably Afobula hypostoma) ,”
Lemon Bay, Fla., Gulf of Mexico. Identified and reported as Pupu-
lina flores by Bere (1936).
Types.—Holotype female, U.S.N.M. No. 85973; allotype male,
U.S.N.M. No. 85972; Gulf of Mexico specimens.
Diagnosis —Length of anterior and posterior parts of body ap-
proximately equal in both sexes. Processes of genital segment of
female reaching to near the middle of the abdomen. Caudal rami
linear and a little longer than abdomen in both sexes, attached later-
ally, divergent in female. Outer piece of first maxilla not produced
distally in female, only slightly so in male; inner piece in male with
small spine near distal outer margin. Male maxilliped with process
of basal segment elongated. Posterior ventral accessory processes
with small lobed membranous extension distally. Leg 4 with inside
spine on segment 3, cuticular process on segment 4.
Description —FrMatp (pl. 18, fig. 1): Gulf of Mexico specimens:
Total length, 6.8—7.0 mm.; average length and width of carapace, 3.2
mm. California specimens: Total length 5.9 mm.; length and width
of carapace, 2.5 mm.
Anterior and posterior margins of carapace curved; sides slightly
rounded. Median sinus a small rounded pit; posterior sinuses rather
shallow. Distal edges of lateral area set thickly with long hairs, edge
of thoracic area with a few spinules. Main lines of dorsal grooving
well defined, that of central cephalic area not distinct in available
specimens.
Free thoracic segment almost three times as wide as long; posterior
margin set with very minute spinules. Genital segment two-thirds
wider than long at middorsal line; sides tapering gradually, armed
marginally itl short spinules. orcas with rounded points, ex-
tending nearly to the middle of the abdomen; inside, the processes
not arising on either side of the base of the abdomen as in P. flores,
the posterior margin of the genital segment being extended beyond
this point. Design on dorsal surface of genital segment with rounded
peak at top, and single lobes at side and bottom; this pattern identical
in the two lots of specimens. Ovisacs reaching slightly beyond end
of caudal rami in all specimens.
Abdomen slender, indistinctly 3-segmented; a dorsal inflation of
the anterior portion of the proximal part the only indication of its
division into two joints; this basal portion almost twice the length
of the anal segment; both set with marginal spinules, dorsal surface
with long slender atest Caudal rami slender, a little longer than
bbdomen: attached to the anal segment laterally, and divergent from
256 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
one another; inner and outer margins set with very short, scattered
spinules. End of rami bearing terminally a short, broad, spiniform
seta and three longer, nonplumose setae, one lateral and one short
ventral seta.
Second antenna (pl. 13, fig. 3) with spinous process of basal seg-
ment well developed; third segment more or less distinct; terminal
claw comparatively weak and little curved, with a slender seta at
its base. Prehensile lateral hook (pl. 13, fig. 4) short, with broadened
base, not strongly curved; setae of papillae branched (pl. 18, fig. 4, a).
Mouth tube as described for the genus, well developed and seeming
large in proportion to the comparatively small size of the copepod.
First maxilla (pl. 18, fig. 5) having the inner piece well extended
distally and the outer portion with a small posterior extension not
reaching beyond the curved basal part of the inner piece; the longest
of the three papillary setae (pl. 18, fig. 11) very broad and partially
divided near its base, the other two very slender, the longer consisting
of two unequal segments. The anterior ventral accessory process
(pl. 18, fig. 6) located near the distal inside end of the inner piece
of the first maxilla, a small spine terminating a membranous fold
that runs backward about half the length of the maxilla (this
membrane not shown in pl. 13, fig. 6).
Posterior claw of second maxilla (pl. 18, fig. 7) only one-third the
length of the anterior. Claw of maxilliped (pl. 18, fig. 8) very short,
with an accessory seta. Posterior ventral accessory process (pl. 18,
fie. 9) consisting of a stout curved chitin rod and a membranous ex-
tension with a small distal lobe. Rod between the bases of legs 1
and 2 broadened, unarmed, curved backwardly.
Legs 1-3 of the same form as in other species of the genus; modified
spine of leg 1 as illustrated in plate 13, figure 17. Exopod of leg 2
(pl. 13, fig. 13) with outer spine of first segment very stout, reaching
only a little beyond the second segment; that of segment 2 reaching
to about the middle of the third segment. First spine of third seg-
ment curved downward, reaching only a little beyond distal margin
of the segment; second spine straight, about half the length of the
first. Leg 3 similar to that of P. flores.
Leg 4 (pl. 13, fig. 18) with the basal segment broadened and armed
with scattered spines; segment 2 having marginal outer spinelets
and a thin ventral Jamina distally. AJl the major spines with toothed
margins. The terminal spines of segment 4 considerably elongated,
the longest being about twice the length of the segment; the innermost
spine attached laterally and unusually short. Segment 8 with a
short lateral spine inside; the fourth segment with a curved cuticular
process (or imperfectly separated spine), covered ventrally by the
lamina which extends outward from the distal edge of the segment.
COPEPOD GENUS PUPULINA (CALIGOIDA)—-WILSON 25%
Mater: Gulf of Mexico specimens: Total length, 5.7-6.1 mm.; width
of carapace, 2.2-2.5 mm.; length of carapace, 2.3-2.8 mm. California
specimens: Total length, 4.8 mm.; width of carapace, 2.0 mm.; length
of carapace, 2.1 mm.
Carapace suborbicular, longer than wide; posterior sinuses shallow;
thoracic area somewhat elongated and narrowed behind the sinuses.
Pattern of cephalic area not distinct in specimens available, except
for two anterior ridges, between which are a pair of rather stout
spines; another pair of spines located a short distance posterior to
the eyes; these spines apparently easily broken off, as in no single
specimen are they all intact.
Free segment about two and one-half times wider than long, with
long dorsal hairs posteriorly. Distal corners of genital segment
hardly produced. Abdomen about as long as genital segment,
2-segmented, set marginally and dorsally with fine spinules. Caudal
rami one-fourth longer than abdomen, attached laterally as in female,
but not divergent; not twisted and flattened laterally to such an
extent as those of P. flores, but with similar long hairs inside. Caudal
setae as in female.
Prehensile lateral hook hardly stouter than that of female, setae
of papillae unbranched. Second antenna (pl. 13, fig. 15) compara-
tively stout; basal segment well differentiated from ventral surface,
second segment armed with a large, ridged lamina and a stout spine;
the third with a 2-layered lamina, a spine and a seta, both of which
arise from large stalks; claw short but strongly curved.
First maxilla (pl. 13, fig. 10) with inner piece elongated distally
as in female, but bearing at outer distal third a stout recurved spine;
basal outer piece with distal outer lobe larger than in female and
extended slightly beyond the curved basal part of the inner piece;
papilla (pl. 13, fig. 12) with the three usual setae, the medial being
very slender.
Maxilliped (pl. 13, fig. 14) with stout terminal claw, armed with
a seta on a long stalk; basal segment having a prominent process
similar to that of P. flores, but more elongated. Anterior ventral
accessory process with smaller spine than in the female; posterior
process like that of female.
Rod between legs 1 and 2 with posterior, upwardly directed lobes,
apparently unarmed. Legs 1-3 like female; leg 4 (pl. 18, fig. 16) like
female except that in addition to scattered spines, the basal segment
is set dorsally with very long slender hairs.
Color.—MacGinitie (1947) has recorded the following notes on the
color of the California specimens: “The smaller of the two new species
of copepods had a wine-colored eyespot. The body and tail were trans-
parent, with wine and blue lines and patches over the body. The
258 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
eggs were salmon colored.” Bere (1986) referred to the “smaller
females” (Gulf of Mexico specimens) which are here designated as
belonging to this species, as having the carapace “creamy with much
branched reddish purple pigment spots; genital segment and egg
strings rose colored; eyes black.”
Remarks.—The only noticeable differences between the Gulf of
Mexico specimens and those from the Pacific are in the larger size
of the former and in the thicker integument of the latter.
It is interesting to note that this species occurred in both collections
with another species of the genus, in the Gulf of Mexico with Pupulina
flores and in the Pacific with the other new species herein described.
The name minor refers to the notably small size of this species, com-
pared with that of the other known species.
PUPULINA BREVICAUDA, new species
PLATE 14, Figures 1-7; PLATE 15
Specimens examined.—Kight females, 2 adult, nonovigerous, others
in various development stages; 16 males, 2 attached to immature
females; collected by G. E. MacGinitie, from around the mouth and
anterior end of the giant ray Mobula lucasana Beebe and TeeVan;
off Santa Catalina Island, Calif.; October 3, 1946; occurring with
Pupulina minor.
Types.—Holotype female, U.S.N.M. No. 85977; allotype male,
U.S.N.M. No. 85976.
Diagnosis.—Posterior part of body in female longer than the an-
terior; a little shorter in the male. Both abdomen and processes of
the genital segment of female greatly elongated, the latter reaching
a little beyond distal end of caudal rami. Caudal rami of female
short and broad, only about one-fifth of the total length of the ab-
domen; linear in male, almost as long as abdomen. Basal portion
of outer piece of first maxilla not produced distally in female, hardly
so in male; inner piece of male with minute spine near distal outer
margin. Male maxilliped with large curved, clawlike process at its
base. Posterior ventral accessory processes large membranous lobes
in female, not so well developed in male. Leg 4 with inside spines
on segments 2-4.
Description—FrMate (pl. 14, fig. 1): Measurements of holotype:
Total length about 14.8 mm.; length of cephalothorax, 6 mm.; pos-
terior part of body, 8.8 mm.; length of carapace, 5.2 mm.; width of
carapace, 6.0 mm. ‘Total length of paratype, 14.5 mm.; width of
carapace, 6.0 mm.
Carapace a little wider than long; anterior and posterior margins
only slightly curved, sides well rounded. Anterior sinus hardly dis-
tinguishable; posterior sinuses deep. Dorsal pattern of carapace
COPEPOD GENUS PUPULINA (CALIGOIDA)——-WILSON 259
uniquely distinguished by the pair of transverse lines of the mid-
cephalic area, these carried backward by curved lines ending in small
loops; midanterior portion with a design similar to that of the male
(pl. 14, fig. 3), but not entirely clear in available specimens. Ventral
supporting ribs of the lateral area not visible dorsally in holotype,
partially so in paratype.
Free segment about one-fourth wider than long, posterior edge
straight. Genital segment with a narrow neck, anterior portion (ex-
clusive of neck and processes) a little longer than wide; sides round-
ing out from neck and extending straight backward in long, narrow
processes, almost twice as long as the main portion of the segment
and reaching a little beyond the tip of the caudal rami; in the para-
type female, the processes slightly shorter than the rami. Processes
well rounded apically, partially covering the abdomen in the proximal
inner area; sides curved under and more or less grooved ventrally ;
the surface integument relatively thin and lacking hairs or spinules.
Top of dorsal pattern of segment with well-rounded lobe in middle,
slanting sides, and five posterior lobes, of which the central is the
longest.
Abdomen slender, elongate, with three distinct segments; the mid-
dle the longest, the anal the shortest; the first two segments together
six times the length of the anal; margin set with very fine short hairs.
Caudal rami about one-fourth as broad as long; differing from those
of other females of the genus in being somewhat flattened, very short
and broad ; equaling only a little more than a fourth of the total length
of the abdominal segments; slightly less than twice the length of the
anal segment ; tipped with three very short setae.
First antenna with fewer plumose setae on the basal segment than in
P. flores, armed terminally as in plate 14, figure 7. Second antenna
(pl. 15, fig. 1) with a stout basal spinous process, terminal claw ex-
ceptionally long, armed proximally with a stalked spine and a long
seta. Lateral hook (pl. 15, fig. 2) very short, hardly curved, setae of
basal papillae little developed. Mouth tube as for the genus; claw of
mandible (pl. 15, fig. 11) with 12 teeth. Anterior ventral accessory
processes short, narrow, membranous spines (pl. 15, fig. 3), larger than
those of minor, smaller than those of flores.
First maxilla (pl. 15, fig. 4) with posterior lobe of outer portion not
extended beyond base of inner piece; setae of papilla subequal (pl.
15, fig. 8), the shortest divided near the tip, the middle one with the
basal part rounded and swollen; inner piece with distal extension very
slender. Posterior claw of second maxilla (pl. 15, fig.5) about half the
length of the anterior. Maxilliped (pl. 15, fig. 6) with moderately
strong claw, set with a slender seta. Posterior ventral accessory proc-
ess (pl. 15, fig. 7) a somewhat large, membranous, rounded flap at-
tached to a narrow chitinous bar.
260 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Rod between legs 1 and 2 narrow, unarmed. Legs 1-3 of the usual
form for the genus. Leg 1 (pl. 15, fig. 14) with endopod reaching to
distal end of first exopod seoment; etd spine and seta of exopod
2 as in plate 15, figure 15.
Leg 2 (pl. 15, fig. 12) with the outer spine of first exopod segment
reaching to the svisannl third of segment 3; that of segment 2 reach-
ing a little beyond the middle of the third segment. First spine of
third segment curved downward, shorter than the second, which is
setiform and sparsely plumose. Laminae of endopod with somewhat
straight outer edges.
Exopod of leg 3 with outer spine of segment 2 reaching to about
middle of third segment; outer spines of segment 3 (pl. 15, fig. 16)
all very short, subequal; differing from flores and mznor in the pres-
ence of a fifth plumose seta inside, this seta shorter and more slender
than the others. Endopod lke that of flores except that the lower
part of the lamellar expansion of segment 2 is much larger, extending
distally almost to the end of the third segment.
Leg 4 (pl. 15, fig. 17) with the basal segment comparatively narrow,
set with short spines posteriorly. Segments 2-4 bearing short spin-
ules inside, that of segment 4 the longest, placed just below the middle
of the segment; this latter spinule apparently homologous to the im-
perfectly separated cuticular process of the other two species. Second
segment with an irregular double row of outer marginal spinelets;
segments 5 and 4 with narrow marginal laminae. Major spines of all
segments stout and comparatively short, the longest of the terminal
spines being only a little longer than its segment. The distal portion
of segment 4 almost squarely truncated, with the inner spine thus
placed in a terminal position; this spine straight and stout, a little
less than one-third the length of the longest terminal spine.
Mate (pl. 14, fig. 3): Average total length, 8.7 mm.; length of
cephalothorax, 4.7 mm.; width of carapace, 3.7 mm. Carapace longer
than wide, anterior and posterior margins rounded, sides nearly
straight. Cephalic area with well-defined dorsal patterns, transverse
lines more or less distinct as in female. Spinules set thickly on dorsal
surface of thoracic area, thinly scattered over the rest of the carapace.
Free thoracic segment almost three times wider than long, set dor-
sally with spinelets and posteriorly with long hairs.
Genital segment and abdomen approximately equal in length.
Genital segment about twice as long as broad, with small distal proc-
esses; surface covered dorsally and ventrally with minute spines.
Anal segment a little longer than the basal segment of the abdomen;
caudal rami slightly shorter than the abdomen, linear as in other males
of the genus. Abdomen with center line of long coarse hairs; caudal
rami with similar coarse hairs on inside margins. |
COPEPOD GENUS PUPULINA (CALIGOIDA)—-WILSON 261
Lateral prehensile hook stouter than that of female, about as long
as terminal claw of second antenna; the setae of basal papillae long,
slender, and unbranched. Antenna 2 (pl. 14, fig. 6) with a cuticular
process on basal segment; a stout, curved spine and small laminate
process on segment 2; a 4-jointed, broadly stalked seta on segment 3;
terminal claw stout and strongly curved, though relatively short, the
seta at its base long and 2-jointed. First maxilla (pl. 15, fig. 9) having
the basal outside portion of the outer piece widened transversely with
a slight distal extension inside; papilla (pl. 15, fig. 10) of two parts, the
upper with a very short spine, the lower with two subequal setae, the
outer seta very broad and set on a stout basal stalk; inner piece slender
throughout, having at its distal outer third a very small curved spine.
Maxilliped (pl. 15, fig. 18) with a small lamina at the middle of the
basal segment, and an unusual clawlike process with a divided tip,
attached basally.
Leg 4 (pl. 15, fig. 18) with broad basal segment set with stout spin-
ules and long coarse hairs. Segments 2-4 with inside spines and outer
marginal laminae as in female; apex of segment 4 not so squarely
truncated, terminal inside spine slender and curved, comparatively
longer than that of the female.
DEVELOPMENT SraceEs: Three development stages of the female are
represented in the collection. None is so immature as the earliest
stage (a) described for Pupulina flores. The youngest, however, cor-
responds very closely to stage b of P. flores, and so is likewise des-
ignated as stage 6. Stage c is probably the one immediately follow-
ing this in natural development. Other forms designated as young
females are considered to represent early growth stages of the adult
copepod.
No filament was ascertained to have been present in any of the speci-
mens, but there is a small ventral structure that is interpreted as being
a filament gland.
Stace 6 (pl. 14, fig. 4) : Two specimens, to one of which a male is
attached to the dorsal neck of the genital segment, clinging by means
of the claws of its second antennae.
Total length, 7.8 mm.; cephalothorax, 5.5 mm.; posterior part of
body, including caudal rami, 2.3 mm. All the first three thoracic
segments incorporated in the carapace, which is a little longer than
wide; frontal plates distinct.
Processes of genital segment reaching almost to the end of abdo-
men. Abdomen 2-segmented, the anal segment not being differen-
tiated. Caudal rami having the short, broad form characteristic of
the species; in this stage they are almost as long as the abdomen.
All of the appendages completely segmented, but not so strongly
developed as those of the adults. The inner piece of the first maxilla
262 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
not reaching beyond the tip of the anterior ventral process; maxillae
and maxilliped crowded together as in adult. Claws of second maxilla
short and weak, as noted for immature forms of P. flores. Anterior
ventral accessory processes very small spines; the membranous flap
of the posterior process hardly separated from the tissue of the ventral
face, but the outline discernible.
Legs 1-8 completely segmented, with the laminae also well devel-
oped. Leg 4 with all the segments indicated, but very short, the whole
structure appearing thin; all the spines, including the inside marginal
ones, present but very short and weak. Rudiment of leg 5, a lobe
hardly differentiated from the surface and bearing three short setae,
present ventrally on the proximal surface of the genital process near
the outer edge (pl. 14, fig. 4, a) ; leg 6, represented by a single seta at-
tached to a minute lobe, located anterior to the fifth pair, the seta
projecting beyond the margin of the genital segment.
This form is closely similar to that described as stage } for P. flores,
and undoubtedly represents the same period in development. ‘That it
can properly be interpreted as a chalimus stage, though a very late
one, can be seen not only from the shape and size of the genital segment
and abdomen, but by the incompleteness of segmentation of the latter.
The appendages, though completely segmented, are thin and weak, and
the claws of the second maxillae have not assumed the adult form.
In addition, legs 5 and 6 are at the height of their development, re-
quiring passage through another stage before disappearing.
Srace e (pl. 14, fig. 5) : One specimen, male attached. Total length,
i1.0 mm.; cephalothorax, 6.5 mm.; posterior part of body, including
caudal rami, 4.5 mm.; width of carapace, 6.5 mm.
Genital processes only slightly longer than in stage 6, but abdomen
considerably longer, showing division into three segments.
All appendages more strongly developed than in preceding stage.
Distal extension of inner piece of first maxilla reaching well beyond
the tip of the anterior ventral process. Claws of maxilla 2 as in the
adult. Spines of leg 4 considerably longer and thicker. Ventral ac-
cessory processes as in stage 6. Legs 5 and 6 still present, but the
setae much shorter than in preceding stage.
Whether this stage is to be interpreted as a chalimus or a growth
form of the adult is not clear. Although the appendages are more
strongly developed than in Stage d, they are still thin and weak com-
pared to the young or adult female. In addition to the comparatively
short abdomen and the small genital segment, there are still present the
rudiments of legs 5 and 6.
Youne FEMALE (pl. 14, fig. 2): Three females, measuring between
13.1 and 13.5 mm., show differences in the length of the processes of
the genital segment: Slightly shorter than the anal segment (as fig-
COPEPOD GENUS PUPULINA (CALIGOIDA)—WILSON 263
ured) ; slightly shorter than the caudal rami; as long as the caudal
rami. These must represent different growth stages of the mature
copepod; they differ from the adult only in their lesser length, in hav-
ing the anterior and posterior parts of the body approximately equal
in length, the carapace not so wide, and in very slight differences in the
dorsal pattern of the genital segment.
Color—MacGinitie (1947) has recevded this species as being trans-
parent and without colored markings; eyespot black.
Remarks.—MacGinitie has further noted that this species was para-
sitized by a trematode, the egg cases being attached mainly to the ab-
dominal area. It may or may not be of significance that the right
caudal ramus of the holotype female, which is a little shorter than the
left, and the left genital process of the paratype female, which is a
little shorter than the right, are each much more heavily infested by
the trematode egg cases than any other area.
The name brevicauda refers to the very shortened caudal rami,
particularly conspicuous in the female, though it is to be noted that
those of the male are comparatively much shorter than those of other
species.
LITERATURE CITED
BASSETT-SMITH, P. W.
1899. A systematic description of parasitic Copepoda found on fishes, with
an enumeration of the known species. Proc. Zool. Soc. London, 1899,
pp. 488-507, pl. 26.
BENEDEN, P. J. VAN.
1892. Quelques nouveaux Caligidés de la céte d’Afrique et de l’archipel des
Acores. Bull. Acad. Roy. Sci. Belgique, ser. 3, vol. 24, pp. 241-262,
4 pls.
Bere, RuBY.
1936. Parasitic copepods from Gulf of Mexico fish. Amer. Midl. Nat., vol.
17, pp. 577-625, 12 pls.
MAcGINITIE, G. E.
1947. Notes on the devilfish, Jl/obula lucasana, and its parasites. Copeia,
1947, No. 4, pp. 276-278, 2 figs.
WILSON, CHARLES BRANCH.
1905. North American parasitic copepods belonging to the family Caligidae.
Part I.—The Caliginae. Proc. U. S. Nat. Mus., vol. 28, pp. 479-672,
pls. 5-29.
1935a. A parasitic copepod, Pupulina flores, redescribed after forty years.
Parasitology, vol. 27, pp. 593-597, 13 figs.
1935b. Parasitic copepods from the Dry Tortugas. Papers Tortugas Lab.,
vol, 29, Carnegie Inst. Washington Publ. 452, pp. 327-347, 6 pls.
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SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 Washington: 1952 No. 3302
2 ==
ECHINODERMS FROM THE MARSHALL ISLANDS
By Austin H. CiarKk
THE ECHINODERMS from the Marshall Islands recorded in this re-
port were collected during Operation Crossroads by the Oceano-
graphic Section of Joint Task Force One under the direction of
Commander Roger Revelle in 1946, and by the Bikini Scientific Re-
survey under the direction of Capt. Christian L. Engleman in 1947.
The number of species of echinoderms, exclusive of holothurians, in
these two collections is 80, represented by 2,674 specimens. Although
many of these have not previously been recorded from these islands,
a number known from the group were not found, while others that
certainly occur there still remain undiscovered.
Of the 80 species collected, 22 were found only in 1946 and 24 only
in 1947; only 34, about 40 percent, were found in both years. It is
therefore impossible to appraise the effects, if any, of the explosion
of the atomic bombs. But the specimens of the 54 species collected in
1947 are all quite normal. On the basis of the scanty and inadequate
data available it would seem that the bombs had no appreciable effect
on the echinoderms.
Some of the species are represented by young individuals only.
This is always the case in any survey of the echinoderm fauna of any
tropical region. A few localities are found to yield nothing but
young individuals of certain species at a given time, or possibly unless
collections are made over a series of years.
A few of the records are from depths greater than any heretofore
recorded for the species. These records are based mostly on dead
material, which may have washed down from the shallower water of
the reefs, though there is no reason to believe that the animals could
not have lived at the depths given.
265
967608—52 1
266 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
The echinoderms represented in the collections, with the year or
years in which they were found, are listed on page 299. Subsequent to
the completion of this report some specimens collected in 1948 have
come to hand. They have been noted in the text under Localities, and
for convenience they have been included with the entries for 1947 in
the list just mentioned.
This collection, unusually extensive for any region in the Poly-
nesian area, was assembled by 16 members of the expeditions, none
of whom had a special interest in the echinoderms. These were:
Capt. R. H. Draeger, M. C., U. S. Navy; Lt. Comdr. Fred C. Ziesen-
henne, U. S. N. R.; Capt. Earl S. Herald, U. S. Army; and Frederick
M. Bayer; Vernon E. Brock; Frank Cali; Martin W. Johnson;
Thomas F. Kohler; Harry S. Ladd; Joseph P. E. Morrison; R. Dana
Russell; Leonard P. Schultz; William Randolph Taylor; Joshua R.
Tracey; Douglas M. Whitaker; and Fred C. Zimmerman. It is a
pleasure to thank these gentlemen and to congratulate them on the
notable contribution they have made to our knowledge of the echino-
derms of Polynesia.
Class CRINOIDEA
Family COMASTERIDAE
Genus COMASTER L. Agassiz
COMASTER GRACILIS (Hartlaub)
Actinometra gracilis HartLaun, Nachr. Ges. Gottingen, May 1890, pp. 170, 187
(description ; Pulo Edam, near Batavia, Java).
Locality.—Bikini Atoll; 150-265 feet; August 7, 1947. One speci-
men, U.S.N.M. No. E. 7236.
Notes.—This specimen has 8 arms on each of the 5 rays, making 40
arms in all. In recording 11 specimens of Comaster multifida (J.
Miiller) from the Gilbert Islands Prof. Torsten Gislén expresses doubt
in regard to the specific distinctness of C. gracilis. It is quite possible
that gracilis will prove to be a synonym of multifida.
Genus COMANTHUS A. H. Clark
COMANTHUS BENNETTI (J. Miiller)
Alecto bennetti J. Mitier, Monatsb. preuss. Akad. Wiss., 1841, p. 187 (description ;
locality unknown).
Locality—Rongelap Atoll; Kieshiechi Island, lagoon side; Mor-
rison, July 24, 1946. Two large specimens, U.S.N.M. Nos. E. 6995,
E. 6996.
MARSHALL ISLANDS ECHINODERMS—CLARK 267
Family MARIAMETRIDAE
Genus STEPHANOMETRA A. H. Clark
STEPHANOMETRA INDICA PROTECTUS (Liitken)
Antedon protectus LUTKEN, Mus. Godeffroy Cat., vol. 5, p. 190, 1874 (Tonga
Islands ; nomen nudum) ; iz P, H. CARPENTER, Trans. Linn. Soc. (Zool.), ser.
2, vol. 2, p. 19, 1897 (character of the oral pinnules).
Localities —Rongerik Atoll; Latoback Island; Bayer, Zimmerman,
and Morrison, August 18, 20, 21, 1947; shallow water, the only depth
given being 5 feet. Seven specimens, U.S.N.M. Nos. E. 7232-E. 7235,
K. 7240, E. 7241.
Rongelap Atoll; on a coral head on the sandy bottom of the lagoon
off the center of Tufa Island; 10 feet; Morrison, July 18, 1946. One
large specimen, U.S.N.M. No. E. 7559.
Class ECHINOIDEA
Family CIDARIDAE
Genus EUCIDARIS Pomel
EUCIDARIS METULARIA (Lamarck)
Cidarites metularia LAMARCK, Histoire naturelle des animaux sans vertébres,
vol, 3, p. 56, 1816 (Ocean of the Great Indies, Mauritius, and Santo Domingo).
Localities —Bikini Atoll; 200-290 feet; August 6, 1947. One small
specimen, U.S.N.M. No. E. 7341.
Bikini Atoll; seaward side of Bikini Island; 150-275 feet; August
7, 1947. Two worn spines, slightly fusiform, the longest 31 mm. long,
U.S.N.M. No. E. 7375.
Bikini Atoll; 800-900 feet; August 7, 1947. One small spine,
slightly fusiform, 15 mm. long, U.S.N.M. No. E. 7376.
Bikini Atoll; Namu Island; outside reef; Johnson, April 4, 1946.
One specimen, U.S.N.M. No. E. 7609.
Eniwetok Atoll; lagoon; Bogon Island, intertidal; Johnson, June 1,
1946. One specimen, U.S.N.M. No. E. 7614.
Rongerik Atoll; Rongerik Island; under rock in shallow water;
Whitaker, August 20, 1947. One specimen, U.S.N.M. No. E. 7242.
Genus CHONDROCIDARIS A. Agassiz
CHONDROCIDARIS GIGANTEA A. Agassiz
Chondrocidaris gigantea A. AGAssiIz, Bull. Mus. Comp. Zool., vol. 1, No. 2, p. 18,
1863 (Hawaiian Islands).
Locality —Bikini Atoll; 800-900 feet; August 7, 1947. A perfectly
clean subambital spine, U.S.N.M. No. E. 7372.
Notes.—This spine agrees fairly well with some of the subambital
268 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
spines in a specimen of Chondrocidaris gigantea at hand from the
Hawaiian Islands, and I have little hesitation in referring it to that
species. It is 87 mm. long and 4 mm. thick without the winglike
processes. The wings are in seven rows along the lateral and aboral
side of the spine, those of one row alternating with those of the rows
on either side. The adoral side of the spine is flattened and is with-
out processes. The processes begin 3 mm. from the base. There are
three or four in each row. Those nearest the base are high conical
tubercles. From the base outward they become progressively elon-
gated and flattened, the outermost being high thin ridges 4-6 mm. long
with a slightly convex crest. The terminal 7-12 mm. on the spine
carries five high thin ridges about 2 mm. high, highest at the proximal
end, thence curving downward to the apex. The color is white with
streaks and spots of ight pink.
The spine resembles in a general way that from Lau, Fiji, figured by
Dr. H. L. Clark under the name of Chondrocidaris problepteryx
(Bernice P. Bishop Mus. Bull. 181, p. 314, pl. 41, fig. EK, 1945), but the
wings are more numerous, are not truncated distally, and are continued
downward to the base.
Genus PHYLLACANTHUS Brandt
PHYLLACANTHUS IMPERIALIS (Lamarck)
Cidarites imperialis LAMARCK, Histoire naturelle des animaux sans vertébres,
vol. 3, p. 54, 1816 (Red Sea).
Locality —Bikini Atoll; west end of the target area, 3 miles south-
west by south of Bikini Island, in the lagoon; 150 feet; broken coral
bottom; Morrison, August 26, 1947. Four spines, U.S.N.M. No. E.
(ee
Note.—These spines are of the typical form with three narrow
evenly spaced white bands; the largest (broken) is 9 mm. in diameter.
The fact that the detached spines of this species and of Chondro-
cidaris gigantea and E'ucidaris metularia were dredged in deep water
does not necessarily indicate that the individuals from which these
spines came lived at these depths. Although this condition is pos-
sible, it is more likely that they lived in shallower water and that
after their death the spines were washed into the localities in which
they were found.
Family ARBACIIDAE
Genus COELOPLEURUS L. Agassiz
COELOPLEURUS, sp.
Locality —Bikini Atoll; 800-900 feet; August 7, 1947. Portion of
a spine, U.S.N.M. No. E. 7370.
MARSHALL ISLANDS ECHINODERMS—CLARK 269
Note.—The spine fragment is 31 mm. long, 1.8 mm. in diameter at
the larger end, and 1 mm. at the smaller. It is rounded triangular in
section, smooth and polished, bright scarlet above, white below. The
lower side has five equally spaced fine longitudinal ridges.
Family DIADEMATIDAE
Genus ECHINOTHRIX Peters
ECHINOTHRIX DIADEMA (Linné)
Echinus diadema LiNn&, Systema naturae, ed. 10, vol. 1, p. 664, No. 7, 1758
(Indian Sea).
Localities —Bikini Atoll; Namu Island, flats a little west of the
northern point, under loose flat coral heads; Morrison, April 4, 1946.
Two specimens, U.S.N.M. No. E. 7148.
Bikini Atoll; Bokonfuaaku Island, on ocean side of reef; Schultz,
April 15, 1946. Four specimens, U.S.N.M. Nos. E. 7222, KE. 7223,
K. 7969, E. 7970.
Bikini Atoll; southeast corner of Bikini Island, lower intertidal
zone, near outer reef, under rocks; Morrison, March 7, 1946. One
specimen, U.S.N.M. No. E. 7006.
Eniwetok Atoll; Ziesenhenne, February 1, 2, 1946. One specimen,
U.S.N.M. No. E. 7285.
Eniwetok Atoll; Arambiru Island, ocean reef; Schultz, June 3,
1946. Two specimens, U.S.N.M. Nos. E. 7224, E. 7225.
Eniwetok Atoll; outer reef at east end of Bogen Island; Morrison,
May 21, 1946. Three specimens, U.S.N.M. No. E. 7536.
Eniwetok Atoll; outer reef flats, Pujiyoru Island; Morrison, June 2,
1946. One small specimen, U.S.N.M. No. E. 7546.
Johnston Island, outer reef; Bayer, August 28, 1947. Three speci-
mens U.S.N.M. Nos. E. 7387, E. 7388.
ECHINOTHRIX CALAMARIS (Pallas)
Hehinus calamaris PALLAS, Spicilegia zoologica, ..., vol. 1, fase. 10, p. 31,
pl. 2, figs. 4-8, 1774.
Localities —Eniwetok Atoll; East Rigili Island; Morrison, May
30, 1946. One specimen, U.S.N.M. No. E. 7583.
Rongelap Atoll; intertidal; Johnson, July 1946. One small speci-
men, U.S.N.M. No. E. 7611.
Johnston Island, outer reef; Bayer, August 28, 1947. Two speci-
mens, U.S.N.M. Nos. E. 7220, E. 7389.
270 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 102
Family TEMNOPLEURIDAE
Genus TEMNOPLEURUS L. Agassiz
TEMNOPLEURUS TOREUMATICUS (Leske)
Cidaris toreumatica LeskE, Additamenta ad Jacobi Theodori Kiein . . ., p.
155, pl. 10, fig. H, 1778.
Locality—Rongelap lagoon; 23+ fathoms; Johnson, June 16,
1946. One small specimen, U.S.N.M. No. E. 7607.
Genus MESPILIA Desor
MESPILIA GLOBULUS (Linné)
Echinus globulus LINNE, Systema naturae, ed. 10, vol. 1, p. 664, No. 2, 1758
(Indian Ocean).
Localities —Bikini Atoll; lagoon; Schultz, March 29, 1946. Two
specimens, U.S.N.M., No. E. 7276.
Eniwetok Atoll; Lidilbut Island, lagoon reef; Schultz, June 1, 1946.
Two fragments, U.S.N.M. No. E. 7187. :
Bikini Atoll: 3 miles off Bikini Island, in lagoon; 150 feet; Hadi-
meda bottom; Morrison, July 11, 1946. One specimen, U.S.N.M.
No. E. 7299.
Bikini Atoll; about one-third of a mile southwest of the southeast
point of Bikini Island; 27 feet; April 25, 1946. Two specimens,
U.S.N.M. No. E. 7532.
Bikini Atoll; Namu Island, outer reef; Johnson, April 4, 1946. One
specimen, U.S.N.M. No. E. 7604.
Bikini Atoll; Enirik Island; intertidal; Johnson, 1946. One speci-
men, U.S.N.M. No. EK. 7623.
Eniwetok Atoll; south end of Rigili Island; Morrison; May 26,
1946. Two dead tests, U.S.N.M. No. E. 7558.
Eniwetok Atoll; East Rigili Island; Morrison, May 30, 1946. Three
specimens, U.S.N.M. No. E. 7530.
Rongelap Atoll; lagoon, 2 miles west of Busch Island; 120 feet;
Morrison, June 21, 1946. One specimen, U.S.N.M. No. E. 7531.
Rongelap Atoll; lagoon; 23+ fathoms; Johnson, June 16, 1946.
One small specimen U.S.N.M. No. E. 7605.
Genus DESMECHINUS H. L. Clark
DESMECHINUS RUFUS (Bell)
Salmacis rufa Brix, Proe. Zool. Soe. London, 1894, p. 411, pl. 26, figs. 2, 3,
(Macclesfield Bank 30-44 fathoms).
Locality—Bikini Atoll; 800-900 feet; August 7, 1947. One speci-
men, U.S.N.M. No. E. 7371.
MARSHALL ISLANDS ECHINODERMS—CLARK Zeet
Notes—The specimen is a bare test 9 mm. in diameter and 4 mm. high.
The sculpture is strongly marked. The color is bright orange red
fading to white on the oral surface. The region around the periproct
is white, the white color running in a broad wedge down the
interambulacral areas about halfway to the ambitus, and in narrower
wedges about two-thirds as far down the midambulacral areas. There
is a yellow-green spot in the middle of each genital. The interior of
the test is white with a tapering red stripe running along each outer
border of the interambulacral areas as far as the coronal ring. The
lower half of the test is lighter and mottled red.
Family TOXOPNEUSTIDAE
Genus CYRTECHINUS Mortensen
CYRTECHINUS VERRUCULATUS (Liitken)
Psammechinus verruculatus LUTKEN, Vid. Medd. Naturh. Foren. Kgbenhavn,
1864, p. 166, 1865.
Localities —Bikini Atoll; lagoon; 180-200 feet; Schultz, March
29, 1946. One specimen, U.S.N.M. No. E. 7286.
Bikini Atoll; in the lagoon, dredged about 3 miles off Bikini Island;
150 feet; Halimeda bottom; Morrison, July 11, 1946. Six specimens,
U.S.N.M. No. E. 7300.
Bikini Atoll; lagoon; Morrison, August 26, 1947. Seven specimens,
U.S.N.M. No. E. 7275.
Rongelap Atoll; lagoon; 23+ fathoms; Johnson, June 16, 1946.
One specimen, U.S.N.M. No. E. 7606.
Genus TRIPNEUSTES L. Agassiz
TRIPNEUSTES GRATILLA (Linné)
Echinus gratilla Linnt, Systema naturae, ed. 10, vol. 1, p. 664, No. 4, 1758
(Indian Ocean).
Localities —Eniwetok Atoll; Ziesenhenne, February 1, 2, 1946. One
specimen, U.S.N.M. No. E. 7282.
Eniwetok Atoll; Arambiru Island, ocean reef; Schultz, June 3, 1946.
Fifteen specimens, U.S.N.M. Nos. E. 7971, E. 7972.
Eniwetok Atoll; East Rigili Island; Morrison, May 30, 1946. One
specimen, U.S.N.M. No. E. 6998.
Eniwetok Atoll; Pujiyoru Island, ocean reef; Morrison and Cali,
June 2, 1946. Eleven specimens, U.S.N.M. No. E. 7308.
Johnston Island; through Dr. David Starr Jordan. One specimen,
U.S.N.M. No. 5948.
Johnston Island, outer reef; Bayer, August 28, 1947. Three speci-
mens, U.S.N.M. No. E. 7390.
262 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Family PARASALENIIDAE
Genus PARASALENIA A. Agassiz
PARASALENIA GRATIOSA A. Agassiz
Parasalenia gratiosa A. AGAssiz, Bull. Mus. Comp. Zool., vol. 1, p. 22, 1863
Kingsmill [Gilbert] and Society Islands).
Localities —Bikini Atoll; lagoon; 180-200 feet; Schultz, March 29,
1946. ‘Two specimens, U.S.N.M. No. E. 7263.
Bikini Atoll; lagoon, from staghorn coral at anchorage about 1 mile
inside Bikini Island; July 16, 1947. One specimen, U.S.N.M. No.
E. 7218.
Bikini Atoll; lagoon; 40-80 feet; Bayer, August 8, 1947. Fourteen
specimens, U.S.N.M. No. E. 7219.
Bikini Atoll; lagoon; 30-75 feet; Bayer, August 15, 1947. Two
specimens, U.S.N.M. No. E. 7216.
Bikini Atoll; lagoon; 50-140 feet; Bayer, August 15, 1947. One
specimen, U.S.N.M. No. E. 7215.
Bikini Atoll; Bikini Island, lagoon reef; Bayer, August 24, 1947.
One specimen, U.S.N.M. No. E. 7214.
Bikini Atoll; Bikini Island; Whitaker, August 1947. Nine speci-
mens, U.S.N.M. No. E. 7163.
Eniwetok Atoll; lagoon; approximately 5 miles north of anchorage,
about 3 miles west of Bogen Island; 90-120 feet; Taylor. One speci-
men, U.S.N.M. No. E. 7246.
Family ECHINOMETRIDAE
Genus ECHINOMETRA Gray
ECHINOMETRA MATHAEI (de Blainville)
Echinus mathaei DE BLAINVILLE, Dict. sci. nat., vol. 37, Oursin, p. 94, 1825.
Localities —Bikini Atoll; Bikini Island, lower intertidal zone near
outer reefs, under rocks; Morrison, March 7, 1946. Three specimens,
U.S.N.M. No. E. 7005.
Bikini Atoll; Bikini Island, under loose flat coral heads on flat
inside of outer reefs near north point, at about low tide level ; Morrison,
April 4, 1948. Three specimens, U.S.N.M. No. E. 7003.
Bikini Atoll; outer reef at the north end of Bikini Island; Whitaker,
July 1947. Seven specimens, U.S.N.M. No. E. 7406.
Bikini Atoll; drift at the south end of Bikini Island; Morrison,
April 9, 1946. One dead test, U.S.N.M. No. E. 7297.
Bikini Atoll; flats inside outer reef, southeast side of Bikini Island;
Morrison, April 5, 1946. One small specimen, U.S.N.M. No. E. 7572.
Bikini Atoll; outer reefs, south end of Bikini Island; Morrison, July
3, 1946. Two specimens, U.S.N.M. No. E. 7553.
MARSHALL ISLANDS ECHINODERMS—CLARK 273
Bikini Atoll; southeast corner of Enyu Island, flats inside outer
reef above low tide line; Morrison, March 16, 1946. Two specimens,
U.S.N.M. No. E. 7541.
Bikini Atoll; southeast corner of Enyu Island, rocky flats inside
outer reef; Morrison, March 16, 1946. One specimen, U.S.N.M. No.
K. 7579.
Bikini Atoll; Ourukaen Island, narrow reef on northeast side; Ladd
and Tracey, May 3, 1946. Three specimens, U.S.N.M. Nos. E. 7580,
E. 7581.
Bikini Atoll; outer reef, south (ocean) side of Bokororyuru Island;
Morrison, April 30, 1946. One specimen, U.S.N.M. No. E. 7582.
Bikini Atoll; outer reef at south end of Bokororyuru Island; Ladd
and Tracey, April 29, 1946. Sixteen specimens, U.S.N.M. Nos. E. 7548,
BE. 7549, E. 7550.
Bikini Atoll; east end of Namu Island, ocean drift; Morrison, April
14, 1946. Fragments, U.S.N.M. No. E. 7298.
Bikini Atoll; Eninman Island, outer reef flats; Morrison, July 17,
1947. Three specimens, U.S.N.M. No. E. 7274.
Eniwetok Atoll; Ziesenhenne, February 1, 2, 1946. Ten specimens,
U.S.N.M. No. E. 7284.
Eniwetok Atoll; Arambiru Island; Schultz, June 3, 1946. Five
specimens, U.S.N.M. No. E. 7260.
Eniwetok Atoll; outer reef flats, northeast side of Arambiru Island;
Morrison, June 3, 1946. Nine specimens, U.S.N.M. Nos. E. 7560,
E. 7561.
Eniwetok Island; south end of Rigili Island; Morrison, May 26,
1946. One dead test, U.S.N.M. No. E. 7555.
Eniwetok Atoll; East Rigili Island; Morrison, May 30, 1946.
Twelve specimens, U.S.N.M. No. E. 7565.
Eniwetok Atoll; outer reef, east end of Bogen Island; Morrison,
May 21,1946. One specimen, U.S.N.M. No. E. 7554.
Eniwetok Atoll; outer reef flats, Pujiyoru Island; Morrison, June
2, 1946. Six specimens, U.S.N.M. No. E. 7547.
Eniwetok Atoll; outer reef, southwest side of Grinem Island; Mor-
rison, May 29, 1946. Four specimens, U.S.N.M. No. E. 7551.
Eniwetok Atoll; flats on outer reef, southwest side of Buganegan
Island; Morrison, May 28, 1946. Two specimens, U.S.N.M. No. E.
7552.
Kwajalein Atoll; Draeger, July 1946. Nine bare tests, U.S.N.M.
No. E. 7244.
Rongerik Atoll; Eniwetak Island, under rocks on reef flat ; Morri-
son, June 29, 1946. Two specimens, U.S.N.M. No. E. 7569.
Rongelap Atoll; Rongelap Island, outer reef flats; Morrison, June
17, 1946. One specimen, U.S.N.M. No. E. 7004.
967608—52 2
274 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Rongelap Atoll; Rongelap Island, intertidal; Johnson, July 1946.
One specimen, U.S.N.M. No. E. 7610.
Johnston Island, outer reef; Bayer, August 28, 1947. Four speci-
mens, U.S.N.M. No. E. 7217.
Note.—These specimens are all small or medium sized. The largest,
from the outer reef flats on the northeast side of Arambiru Island,
Eniwetok Atoll, is 51 mm. in length, 40 mm. broad, and 32 mm. high
(E. 7561).
ECHINOMETRA MATHAEI var. OBLONGA (de Blainville)
Echinus oblongus pE BLAINVILLE, Dict. sci. nat., vol. 37, Oursin, p. 95, 1825.
Localities —Bikini Atoll; Bikini Island, from borings in coralline
rock at and just behind the edge of the outer reef on the southeast
side of the island; Morrison, April 5, 1948. ‘Twenty-eight specimens,
U.S.N.M. No. E. 7001.
Bikini Atoll; Bikini Island, outer reef opposite the center of the
island; Ladd, April 17, 1946. Three specimens, U.S.N.M. No. E. 6700.
Bikini Atoll; Bikini Island, outer edge of reef opposite the east
central part; Morrison, April 18, 1946. One specimen, U.S.N.M.
No. E. 7002.
Bikini Atoll; Bikini Island; Herald, July 15, 1946. Two speci-
mens, U.S.N.M. No. E. 7259.
Eniwetok Atoll; Lidilbut Island, near outer reef edge; Morrison,
May 1,1946. Five specimens, U.S.N.M. No. E. 7568.
Rongelap Atoll; Rongelap Island, outer reef flats; Johnson, July
1946. One specimen, U.S.N.M. No. E.7281.
Johnston Island, outer reef; Bayer, August 28, 1947. One speci-
men, U.S.N.M. No. E.7218.
Genus ECHINOSTREPHUS A. Agassiz
ECHINOSTREPHUS ACICULATUS A. Agassiz
Echinostrephus aciculatus A. AGassiz, Bull. Mus. Comp. Zool., vol. 1, p. 20, 1863
(Kingsmill [Gilbert] and Hawaiian Islands).
Localities —Bikini Atoll; Eniairo Island; Bayer, July 21, 1947.
One specimen, U.S.N.M. No. E. 7228.
Bikini Atoll; Namu Island, outer reef; Bayer, August 25, 1947.
Four specimens, two in their excavations in coral rock, U.S.N.M. Nos.
E. 7226, E. 7368, E. 7369.
Eniwetok Atoll; Arambiru Island, ocean reef; Schultz, June 3,
1946. Two specimens, U.S.N.M. Nos. E. 7227, E. 7266.
Eniwetok Atoll; south end of Rigili Island; Morrison, May 26, 1946.
One dead test, U.S.N.M. No. E. 7556.
Eniwetok Atoll; East Rigili Island; Morrison, May 30, 1946. One
specimen, U.S.N.M. No. E. 7529.
MARSHALL ISLANDS ECHINODERMS—CLARK 275
Eniwetok Atoll; Pujiyoru Island, outer reef flats; Morrison, June
92,1946. One specimen, U.S.N.M. No. E. 7544.
Rongerik Atoll; Bock Island, outer reef; Bayer and Zimmerman,
August 19, 1947. One specimen.
Note.—One of the specimens from Namu Island is at the bottom of
a cylindrical hole with perfectly smooth sides, 80 mm. deep and 380 mm.
in diameter (EF. 7368).
Genus HETEROCENTROTUS Brandt
HETEROCENTROTUS TRIGONARIUS (Lamarck)
Echinus trigonarivus LAMARCK, Histoire naturelle des animaux sans vertébres,
vol. 8, p. 51, 1816 (?Mediterranean).
Localities —Bikini Atoll; reef off the southeast corner of Bikini
Island; Morrison, March 7, 1946. Four specimens, U.S.N.M. Nos.
E. 7007, E. 7008, E. 7533.
Bikini Atoll; outer reef, southern end of Bikini Island; Morrison,
July 3, 1946. One specimen, U.S.N.M. No. E. 75384.
Bikini Atoll; Enyu Island, southeast corner, outer reef; Morrison,
March 16, 1946. Five specimens, U.S.N.M. No. E. 6988.
Bikini Atoll; Namu Island, edge of outer reef; Morrison, April 4,
1946. Three specimens, U.S.N.M. Nos. E. 6992, E. 6993.
Bikini Atoll; Bikini Island, outer reef opposite the center of the
island; Ladd, April 17, 1946. One specimen, U.S.N.M. No. E. 6991.
Bikini Atoll; Bikini Island, edge of reef opposite the east central
part of the island; Morrison, April 18, 1946. Two specimens,
U.S.N.M., No. E. 6989.
Bikini Atoll; Enyu Island, outer reef, south point of the island;
Bayer, August 1, 1947. Two specimens, U.S.N.M., Nos. E. 7183,
E. 7184.
Bikini Atoll; Namu Island, outer reef; Morrison, August 7, 1947.
One specimen, U.S.N.M., No. E. 7207.
Bikini Atoll; Namu Island, outer reef; Bayer, August 25, 1947.
Two specimens, U.S.N.M., Nos. E. 7179, E. 7180.
Eniwetok Atoll; Ziesenhenne, February 1, 2, 1946. One specimen,
U.S.N.M., No. E. 7283.
Eniwetok Atoll; Arambiru Island, ocean reef; Schultz, June 3,
1946. Two specimens, U.S.N.M., Nos. E. 7967, E. 7968.
Eniwetok Atoll; Pujiyoru Island, ocean reef; Morrison, June 2,
1946. Two specimens, U.S.N.M., Nos. E. 7965, EK. 7966.
Eniwetok Atoll; flats on east side near south end of Eniwetok Island ;
Morrison, May 20, 1946. One specimen, U.S.N.M., No. E. 7535.
Eniwetok Island; Grinem Island, outer reef on southwest side;
Morrison, May 29, 1946. Seven specimens, U.S.N.M., Nos. E. 6990,
E. 6994.
276 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Eniwetok Atoll; Lidilbut Island, near outer reef edge; Morrison,
May 1, 1946. Two small specimens, U.S.N.M., No. E. 7557.
Rongelap Atoll; 2 miles west of Busch Island; 120 feet; Taylor,
June 21, 1946. One small dead test, U.S.N.M., No. E. 7564.
Rongelap Atoll; Kabelle Island; lithothamnion ridge, intertidal;
Johnson, July 28, 1946. One small specimen, U.S.N.M., No. E. 7603.
Johnston Island; outer reef edge, northwest side; Bayer, August
28, 29, 1947. Two specimens, U.S.N.M., Nos. E. 7181, E. 7182.
HETEROCENTROTUS MAMMILLATUS (Linné)
Echinus mamillatus LINNE, Systema naturae, ed. 10, vol. 1, p. 664, No. 9, 1758 (no
locality).
Localities —Johnston Island; through David Starr Jordan. One
specimen, U.S.N.M., No. E. 5949.
Johnston Island, outer reef edge on the northwest side of the island;
Bayer, August 28, 1947. Twenty-four specimens, U.S.N.M., Nos.
E. 7185-E. 7206, E. 7237, E. 7316.
Family CLYPEASTRIDAE
Genus CLYPEASTER Lamarck
CLYPEASTER RETICULATUS (Linné)
Echinus reticulatus LiInNNE, Systema naturae, ed. 10, vol. 1, p. 666, No. 15, 1758
(American Ocean).
Locality —Bikini Atoll; Bikini Island; 150-265 feet; August 7,
1947. One specimen, U.S.N.M. No. E. 7262.
Family LAGANIDAE
Genus LAGANUM Linck
LAGANUM DEPRESSUM L. Agassiz
Laganum depressum L. AGAssiz, Monographie des Scutellidae, p. 110, pl. 23, figs.
1-7, 1841 (Moluceas).
Localities.—Bikini Atoll; about one-fourth mile southwest of the
southeast point of Bikini Island; 21 feet; sandy bottom with Forami-
nifera; Morrison, April 25, 1946. Thirty-two specimens, U.S.N.M.,
Nos. E. 7505, E. 7508, E. 7294.
Bikini Atoll; one-third of a mile southwest of the southeast point
of Bikini Island; 21 feet; Morrison, April 25, 1946. Three speci-
mens, U.S.N.M., No. E. 7291.
Bikini Atoll; one-third of a mile west of the southeast point of
Bikini Island; 24 feet ; sandy bottom; Morrison, April 23, 1946. Two
specimens, U.S.N.M., No. E. 7507.
Bikini Atoll; one-third of a mile southwest of the southeast point
of Bikini Island; 27 feet; sandy bottom; Morrison, April 25, 1946.
Five specimens, U.S.N.M., Nos. E. 7510, E. 7577.
MARSHALL ISLANDS ECHINODERMS—CLARK 277
Bikini Atoll; one-third of a mile west of the southeast point of
Bikini Island; 30 feet; coral and foraminiferal sand; Morrison, April
23, 1946. Seventeen specimens, U.S.N.M., Nos. E. 6999, E. 7511.
Bikini Atoll; one-third of a mile west of the southeast point of
Bikini Island; 36-42 feet; Morrison, April 23, 1946. Four specimens.
Bikini Atoll; one-half mile south of the west end of Bikini Island,
in the lagoon; 150 feet; sandy bottom with Halimeda; Morrison,
August 7, 1946. One specimen, U.S.N.M., No. E. 7293.
Bikini Atoll; three-fourths of a mile south of the west end of Bikini
Island; 90 feet; sandy bottom; Morrison, August 26, 1947. Two
specimens, U.S.N.M., No. E. 7238.
Bikini Atoll; south of the west end of Bikini Island; coral-algae
bottom; Morrison, April 28, 1946. Two specimens, U.S.N.M., No.
E. 7292.
Bikini Atoll; in the lagoon 3 miles off Bikini Island; 150 feet; Hali-
meda bottom; Morrison, July 11, 1946. Four specimens, U.S.N.M.
Nos. E. 7295, E. 7296.
Bikini Atoll; 4 miles south of the west end of Bikini Island; 168
feet; Morrison, April 25, 1946. One specimen, U.S.N.M. No. E. 7265.
Bikini Atoll; in the lagoon one-eighth to one-fourth mile off-
shore inside Bokonfuaaku Island; 18-380 feet; foraminiferal sand cov-
ering rock bottom; Morrison, July 11,1946. Eleven small specimens,
U.S.N.M. No. E. 7509.
Eniwetok Atoll; Pujiyoru Island, lagoon reef; Schultz, June 2,
1946. Fragments, U.S.N.M. No. E. 7512.
Eniwetok Atoll; in the lagoon approximately 5 miles north of the
anchorage, about 8 miles west of Bogen Island; 90-120 feet; Taylor.
Six specimens, U.S.N.M. No. E. 7247.
Rongerik Atoll; ocean side of Rongerik Island; August 13, 1947.
Two specimens, U.S.N.M. No. E. 7239.
Rongelap Atoll; lagoon; 23+ fathoms; Johnson, June 16, 1946.
One specimen, U.S.N.M. No. E. 7608.
Note.—One of the specimens from about one-fourth mile southwest
of the southeast point of Bikini Island has the petals broadly outlined
in purple.
Family FIBULARIIDAE
Genus FIBULARIA Lamarck
FIBULARIA AUSTRALIS Desmoulins
Fibularia australis DesMovLins, Etudes sur les Echinides, p. 240, 1837 (southern
seas).
Localities—Bikini Atoll; 3 miles off Bikini Island, in the lagoon;
150 feet; Halimeda bottom; Morrison, July 11, 1946. Four speci-
mens, U.S.N.M. No. E. 7308.
278 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Eniwetok Atoll; north end of Bogon Island; Draeger, 1948. Eight-
een specimens, U.S.N.M. No. E. 7814.
Eniwetok Atoll; south end of Rigili Island; Morrison, May 25,
1946. Eleven specimens, U.S.N.M. No. E. 7495.
Rongerik Atoll; in drift on the sandy beach at the north end of
Bock Island; Morrison, August 19, 1947. Twenty specimens,
U.S.N.M. Nos. E. 7587, 7816.
Rongelap Atoll; sandy flats on the lagoon side of Kabelle Island;
Morrison, June 20, 1946. One specimen, U.S.N.M. No. E. 7496.
FIBULARIA OVULUM (Linné)
Echinus ovulum Linnk, Systema naturae, ed. 13, vol. 1, pt. 6, p. 3194, No. 83,
1788 (no locality).
Localities —Bikini Atoll; in the lagoon, one-half mile south of the
west end of Bikini island; 150 feet; sandy bottom with Halimeda;
Morrison, August 7, 1946. One specimen, U.S.N.M. No. E. 7304.
Bikini Atoll; a mile and a half south of the west end of Bikini Is-
land; 72 feet; coral and sand bottom; Morrison, April 4, 1946. Two
specimens, U.S.N.M. No. E. 7305.
Bikini Atoll; west side of Arriikan Island; Morrison, May 14, 1946.
One specimen, U.S.N.M. No. E. 7503.
Eniwetok Atoll; north side of Pujiyoru Island, at high tide line;
Morrison, June 2, 1946. One specimen, U.S.N.M. No. E. 7306.
Eniwetok Atoll; south end of Rigili Island; Morrison, May 25,
1946. ‘Thirty specimens, U.S.N.M. Nos. E. 7497, E. 7500, E. 7829.
Eniwetok Atoll; in the lagoon approximately 5 miles north of the
anchorage, about 3 miles west of Bogen Island; 90-120 feet; Taylor.
Three specimens, U.S.N.M. No. E. 7249.
Eniwetok Atoll; north end of Bogen Island; Draeger, 1948. One
specimen, U.S.N.M. No. E. 7815.
Eniwetok Atoll; islet west of Elugelab Island; Draeger, 1948.
Eleven specimens, U.S.N.M. No. E. 7826.
Rongerik Atoll; lagoon side of Bock Island; Morrison, June 27,
1946. Two specimens, U.S.N.M. No. E. 7303.
Rongerik Atoll; in drift on the sandy beach at the north end of
Bock Island; Morrison, August 19, 1947. One thousand six hundred
and two specimens, U.S.N.M. Nos. E. 7586, E. 7813, E. 7824.
FIBULARIA VOLVA L. Agassiz and Desor
Fibularia volva L. AGassiz and DeEsor, Ann. Sci. Nat., ser. 3, vol. 7, p. 142, 1847
(Red Sea).
Localities —Bikini Atoll; in the lagoon; Morrison, August 26, 1947.
Two specimens, U.S.N.M. No. E. 7277.
Eniwetok Atoll; islet west of Elugelab Island; Draeger, 1948.
One specimen, U.S.N.M. No. E. 7830.
MARSHALL ISLANDS ECHINODERMS—CLARK 279
Rongelap Atoll; sandy flats on the lagoon side of Kabelle Island;
Morrison, June 20, 1946. Seven specimens, U.S.N.M. No. E. 7498.
Rongerik Atoll; in drift on the sandy beach at the north end of
Bock Island; Morrison, August 19, 1946. Sixteen specimens,
U.S.N.M. Nos. E. 7589, E. 7817, E. 7821.
FIBULARIA ACUTA Yoshiwara
Fibularia acuta YoSHIWARA, Annot. Zool. Japon., vol. 2, p. 60, 1898.
Rongerik Atoll; north end of Bock Island; Morrison, August 19,
1948. ‘Two specimens, U.S.N.M. No. E. 7818.
Genus ECHINOCYAMUS van Phelsum
ECHINOCYAMUS INCERTUS H. L. Clark
Hehinocyamus incertus H. L. CLarxK, Mem. Mus. Comp. Zool., vol. 46, No. 1, p. 64,
pl. 128, figs. 1-3, 1914 (Albatross station 4045, off Kawaihae Light, Hawaii,
147-198 fathoms).
Locality.— Bikini Atoll; 800-900 feet; August 7, 1947. One speci-
men, U.S.N.M. No. E. 7374.
Note.—This specimen is 7 mm. long, 5 mm. in maximum width, and
2.5mm. high, a little larger than the type specimen.
ECHINOCYAMUS MEGAPETALUS H. L. Clark
Echinocyamus megapetalus H. L. CLark, Mem. Mus. Comp. Zool., vol. 46, No. 1,
p. 60, pl. 128, figs. 5-8, 1914 (Mauritius).
Localities—Eniwetok Atoll; south end of Rigili Island; Morrison,
May 25, 1946. One specimen, U.S.N.M. No. E. 7501.
Eniwetok Atoll; Chinieero Island; Draeger, 1948. Six specimens,
U.S.N.M. No. E. 7827.
Bikini Atoll; Namu Island; Morrison, March 1, 1946. One speci-
men, U.S.N.M. No. E. 7828.
Rongerik Atoll; in drift on the sandy beach at the north end of
Bock Island; Morrison, August 19, 1947. One hundred thirty-four
specimens, U.S.N.M. Nos. E. 7590, E. 7819, E. 7825.
Rongerik Atoll; Latoback Island, northeast side, just above the
high-tide line; Morrison, June 28, 1946. One specimen, U.S.N.M. No.
E. 7499.
Note.—The specimen from Latoback Island, Rongerik Atoll, is 6
mm. long, 4 mm. wide, and 2.5 mm. high.
ECHINOCYAMUS ELONGATUS H. L. Clark
Echinocyamus elongatus H. L. CLarK, Mem. Mus. Comp. Zool., vol. 46, No. 1, p. 61,
pl. 126, figs. 9-11, 1914 (Albatross station 3846, south coast of Molokai,
Hawaiian Islands, 117-110 meters).
Locality.—Bikini Atoll; west side of Arriikan Island; Morrison,
May 14,1946. Eight specimens, U.S.N.M. No. E. 7502.
280 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 102
Family ECHINONEIDAE
Genus ECHINONEUS van Phelsum
ECHINONEUS ABNORMALIS de Loriol
Echinoneus abnormalis Dr LorioL, Mém. soc. phys. nat. hist. Genéve, vol. 28, No. 8,
p. 41, 1888 (Mauritius).
Locality—Eniwetok Atoll; east side of Rigili Island; Morrison,
May 30, 1946. One specimen, U.S.N.M. No. E. 7488.
Note.—This specimen is large, 50 mm. long, 42 mm. wide, and 28
mm. high.
ECHINONEUS CYCLOSTOMUS Leske
Echinoneus cyclostomus LesKr, Additamenta ad Jacobi Theodori Klein, p. 178,
pl. 37, figs. 4, 5, 1778.
Localities —Bikini Atoll; Eniairo Island; Bayer, July 21, 1947.
One specimen, U.S.N.M. No. E. 7267.
Bikini Atoll; ocean drift on the west part of the north side of
Namu Island; Morrison, March 380, 1946. One specimen, U.S.N.M.
No. E. 7302.
Bikini Atoll; drift on the ocean side of Namu Island; Morrison,
April 14, 1946. One specimen, U.S.N.M. No. E. 7301.
Bikini Atoll; rocky flats inside the outer reef, southeast corner
of Enyu Island, near low tide line; Morrison, March 16, 1946. One
specimen, U.S.N.M. No. E. 7493.
Eniwetok Atoll; east side of Rigili Island; Morrison, May 30, 1946.
One specimen, U.S.N.M. No. E. 7489.
Eniwetok Atoll; south end of Rigili Island; Morrison, May 25,
1946. One specimen, U.S.N.M. No. E. 7494.
Rongelap Atoll; outer reef flat of Rongelap Island; Johnson, July
1946. Two large specimens, U.S.N.M. No. E. 7280.
Rongerik Atoll; in drift on sandy beach at the north end of Bock
Island; Morrison, August 19, 1947. Eighteen specimens, U.S.N.M.
Nos. E. 7589, E. 7820.
Family NUCLEOLITIDAE
Genus ECHINOLAMPAS Gray
ECHINOLAMPAS ALEXANDRI de Loriol
Echinolampas alexandri pe Lori0oL, Mém. soc. phys. nat. hist. Genéve, vol. 24,
p. 660, 1876 (Mauritius).
Localities —Bikini Atoll; in the lagoon; 180-200 feet; Schultz,
March 29, 1946. One specimen, U.S.N.M. No. E. 7279.
Bikini Atoll; 4 miles south of the west end of Bikini Island; 180
feet; Morrison, April 4, 1946. One specimen, U.S.N.M. No. E. 7278.
Bikini Atoll; one-third of a mile west of the southeast point of
MARSHALL ISLANDS ECHINODERMS—CLARK 281
Bikini Island; 80 feet; coral and foraminiferal sand bottom; Mor-
rison, April 28, 1947. One very young specimen, U.S.N.M. No. E.
7506.
Notes.—The very small specimen (E. 7506) bears little resemblance
to the adult, but instead suggests a small slender spined “’chinometra.
It is slightly oval, two-thirds as broad as long, and twice as long
as high, with the aboral surface gently convex. It measures 7 mm.
in length, 5.5 mm. in width, and 3.5 mm. in height.
The genitals are fused into a somewhat irregular five-sided plate
1.5 mm. long and 1 mm. broad. The posterior genital appears to be
much larger than the others and bears a large perforated tubercle
surrounded by a broad areole. Numerous other tubercles about half
as large are scattered over the surface of the combined genitals.
There are four large madreporic pores, each on the summit of a short
tube, the four tubes crowded into a square with rounded angles and
slight notches on the sides. No pores are visible in the genital plates.
The triangular oculars are very small, each situated on one of the
sides of the combined genitals.
There are from 10 to 12 large interambulacrals in each column.
These are at first hexagonal, toward the ambitus gradually becoming
transversely elongated, and at the ambitus about twice as broad as
long; below the ambitus they gradually become shorter again.
Aborally each interambulacral plate bears a large central perforated
tubercle surrounded by a broad areole margined by a circlet of small
and glassy, usually contiguous, tubercles. On the broad interambu-
lacral plates at and below the ambitus there are commonly two tu-
bercles to a plate.
The ambulacrals are at first minute. After a series varying from
12 to 19, much larger ones, each with a large central tubercle, appear,
and soon all are large with usually three pores in a slightly curved
line along the outer border. At the ambitus the ambulacrals are
hexagonal and about as high as the much broader interambulacrals,
with central tubercles about as broad as those of the latter. On the
aboral surface the very small ambulacrals always extend farther
down on one side of the ambulacral areas than on the other.
There is a single glassy spheridium situated in a deep and capacious
pit at the adoral end of each ambulacrum.
The peristome is circular, 2 mm. in diameter, densely and evenly
covered with very minute spinous plates. The mouth is closed and
no teeth are visible.
The periproct is large, situated just below the ambitus and sloping
slightly inward, diamond shaped with rounded angles, transversely
elongated, 2.3 mm. wide and 1.4 mm. high. Most of its surface is
covered by three large plates of which the two outer reach halfway
967608—52—-3
282 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
from the lateral apices to the anterior angle. The remaining area,
from the anterior angle about halfway to the posterior, and halfway
to the lateral apices, is covered by several much smaller plates.
The primary spines are 2 mm. long, cylindrical or slightly tapering,
with eight high and rounded longitudinal ridges. In section they
resemble the spine of Lchinolampas alexandri var. sibogae figured
by Mortensen (Monograph of the Echinoidea, vol. 4, part 1, p. 281,
fig. 272, c. 1948). The secondary spines are very fine, 0.7 mm. long
and cylindrical.
The ophicephalous pedicellariae resemble the one from £’. a. sibogae
figured by Mortensen (pl. 14, figs. 5, 6).
The color is light olive-green. The test is yellowish white with
some olive-green patches and a purple patch on the genitals.
This little specimen bears a close resemblance to the young E'chino-
lampas depressa figured by Agassiz (Revision of the Echini, pl. 16,
figs. 1, 2, 8, 1872), but it is much lower than is shown in figure 3, in
lateral view being more like the larger specimen shown in figure 10.
The periproctal structure is as shown in figure 14, but the three large
plates are smaller.
Dr. Morrison’s specimen (E. 7278) is 77 mm. long, 69 mm. in maxi-
mum width, 35 mm. high at the apical system, and 86 mm. high at
the posterior end. It agrees wel] with a specimen from Tavoy figured
and described by Koehler, but is slightly larger and more depressed.
Dr. Schultz’s specimen (E. 7279) agrees with the preceding, but is
somewhat smaller.
This species has heretofore been reported only from Mauritius,
Ceylon, and the Bay of Bengal.
Family SPATANGIDAE
Genus METALIA Gray
METALIA DICRANA H. L. Clark
Metalia dicrana H. L. CrarK, Mem. Mus. Comp. Zool., vol. 46, No. 2, p. 211, pl.
146, fig. 16, pl. 160, figs. 1-4, 1917 (Panglao, Bohol Province, Philippines).
Localities —Bikini Atoll; in the lagoon, about one-eighth to one-
fourth mile offshore inside Bokonfuaaku Island; 18-30 feet; Morri-
son, July 11, 1946. One specimen, U.S.N.M. No. E. 7487.
Eniwetok Atoll; in the lagoon, approximately 5 miles north of the
south anchorage, about 3 miles west of Bogen Island; 90-120 feet;
Taylor. Seven specimens, U.S.N.M. No. E. 7245.
Notes—The specimen from Bikini is 26 mm. long, 23 mm. wide,
MARSHALL ISLANDS ECHINODERMS—CLARK 283
and 17 mm. high. The seven specimens from Eniwetok are small,
about 17 mm. long, and are very variable in shape.
Genus BRISSUS Leske
BRISSUS LATECARINATUS (Leske)
Spatangus brissus var. latecarinatus Leskre, Additamenta ad Jacobi Theodori
Klein, pp. X_X, 185, 1778.
Localities —Bikini Atoll; Namu Island; drift on the north side of
the east end; Morrison, April 14, 1946. Two bare tests, U.S.N.M.
No. E. 7258.
Bikini Atoll; blasted out of a coral head in making a channel across
the flats just west of the beach on the west side of Bokororyuru
Island; Morrison, August 12, 1946. Fragments, U.S.N.M. No. E.
7290.
Rongelap Atoll; lagoon side of Eniaetok Island; 12 feet; Brock
and Herald, July 20, 1946. One specimen, U.S.N.M. No. E. 7492.
Note.—The two specimens from Namu Island (KE. 7258) are 110
and 115 mm. in length.
Genus MARETIA Gray
MARETIA OVATA (Leske)
Spatangus ovatus Leske, Additamenta ad Jacobi Theodori Klein, p. 188, pl. 49,
figs. 12, 13, 1778.
Localities —Bikini Atoll; 50-80 feet; August 15, 1947. One speci-
men, U.S.N.M. No. E. 7256.
Bikini Atoll; in the lagoon; 100-140 feet; August 6, 1947. Three
small broken specimens, U.S.N.M. No. E. 7373.
Eniwetok Atoll; Lidilbut Island, north side, outer reef flats, under
sand; Morrison, June 1, 1946. One specimen with parasitic gastro-
pods, U.S.N.M. No. E. 7504.
Class ASTEROIDEA
Family ASTROPECTINIDAE
Genus ASTROPECTEN Schultze
ASTROPECTEN POLYACANTHUS Miiller and Troschel
Astropecten polyacanthus MU.tier and TroscHer, System der Asteriden, p. 69,
1842 (Red Sea).
Locality —Eniwetok Atoll; in the lagoon approximately 5 miles
north of the anchorage and about 3 miles west of Bogen Island;
Taylor, 1947. One small specimen, U.S.N.M. No. E. 7248.
284 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Family LUIDIIDAE
Genus LUIDIA Forbes
LUIDIA MASCARENA Déderlein
Luidia mascarena DODERLIEN, Die Asteriden der Siboga-Expedition, pt. 2, p. 261,
fig. 5, 1920 (Mauritius).
Locality.—Bikini Atoll; in the lagoon; Morrison, August 26, 1947.
One specimen, U.S.N.M. No. E. 7255.
Note.—This specimen was kindly identified by Prof. Walter K.
Fisher.
Family GONIASTERIDAE
Genus CALLIASTER Gray
CALLIASTER ELEGANS Doderlein
Calliaster elegans DOpERLIEN, Bijdragen tot de Dierkunde, Amsterdam, Afi. 22,
p. 49, pl. 1, figs. 1-1b, 1922 (Solor Strait, 118 meters).
Locality —Bikini Atoll; off Bikini Island; 400-450 feet; Russell,
August 14, 1947. One specimen, U.S.N.M. No. E. 7365.
Notes —R=45 mm., r=18 mm.; R=2.4 r. This specimen agrees
fairly well with Déderlein’s type specimen, which was dredged by
the Siboga in the Solor Strait near Flores in 113 meters, though it is
rather more than twice as large. The plates of the disk agree with
those of the type. The innermost spines of the carinal row are 6 mm.
high, those following decreasing rapidly in height and disappearing
at the level of the third superomarginal. The plates of the disk, ex-
cept for those of the two rows adjoining the superomarginals, each
have a prominent central tubercle. There are no pedicellariae.
There are eight superomarginals of which the outer five are in con-
tact in the midradial line. The two interradial superomarginals
each bear a prominent spine 2 mm. high on the inner edge, a shorter
spine or tubercle in the middle, and usually one or two tubercles near
the outer edge. The next two superomarginals on each side bear
similar but smaller spines. The following superomarginals bear
usually two or three tubercles near the inner edge, one in the middle,
and from one to three near the lower edge.
The inferomarginals bear a group of from two to four tubercles
near the upper edge, or sometimes a group of three long spines.
Each actinal plate bears a short stout spine.
The adambulacral plates bear two large stout spines on the outer
surface and a comb of usually seven subequal furrow spines.
The slight differences between this specimen and the type, the only
other specimen known, are probably due chiefly to size and age.
MARSHALL ISLANDS ECHINODERMS—CLARK 285
Family OREASTERIDAE
Genus CULCITA L. Agassiz
CULCITA NOVAE-GUINEAE Miller and Troschel
Culcita novae-guineae MULLER and TRoscHEL, System der Asteriden, p. 38, 1842.
Localities —Bikini Atoll; north side of the lagoon, by diving to a
depth of 15 feet; March 11, 1946. Three specimens, U.S.N.M. Nos.
FE. 7960, E. 7961, E. 7962. The diameter of the largest (in alcohol) is
225 mm.
Bikini Atoll; Bokororyuru Island, west side; Morrison, August 12,
1948. One specimen, U.S.N.M. No. E. 6997.
Eniwetok Atoll; Arambiru Island, ocean reef; Schultz, June 3, 1946.
One specimen, U.S.N.M. No. E. 7964.
Eniwetok Atoll; Pujiyoru Island, ocean reef; Schultz, June 2, 1946.
One specimen, U.S.N.M. No. E. 7963.
Family LINCKIIDAE
Genus NEOFERDINA Livingstone
NEOFERDINA OCELLATA (H. L. Clark)
Ferdina ocellata H. L. CLarkK, The Echinoderm fauna of Torres Strait, p. 60, pl.
6, fig. 5 (colored), pl. 31, figs. 1, 2, 1921 (Mer, Murray Islands, Torres Strait).
Locality.—Bikini Atoll; Namu Island, ocean drift on the north side:
Morrison, March 30, 1946. One beach-worn specimen, U.S.N.M. No.
KE. 7364.
Notes—R=33 mm., r=10 mm.; width of rays at base 12 mm. This
specimen agrees well with Dr. H. L. Clark’s original description and
figures, based upon a single specimen from Mer, Murray Islands,
Torres Strait. Since the species was originally described two addi-
tional specimens have been recorded, another from Mer and one from
Northwest Islet.
Neoferdina ocellata is probably conspecific with MN. cancellata
(Grube), described from a specimen without locality and later recorded
from Fiji.
Genus FROMIA Gray
FROMIA BALANSAE E. Perrier
Fromia balansae FE. Perrier, Arch. zool. experim., vol. 4, p. 178, 1875 (New
Caledonia).
Locality —Eniwetok Atoll; East Rigili Island; Morrison, May 30,
1946. One specimen, U.S.N.M. No. E. 7524.
Notes —R=25 mm.,r=7 mm. This specimen agrees well with the
figure of the type specimen published by Koehler (Echinoderma of
the Indian Museum, Part vi, Asteroidea IT, pl. 18, figs. 7, 8, 1910).
286 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
FROMIA HEMIOPLA Fisher
Fromia henmiopla F1sHer, Proc. U. S. Nat. Mus., vol. 46, p. 218, 1913 (Tonquil
Island, Gumila Reef, south of Mindanao, Philippine Islands).
Localities —Bikini Atoll; in the lagoon; Schultz, March 29, 1946.
One specimen, U.S.N.M. No. E. 7264.
Bikini Atoll; 150-180 feet; Schultz, April 18, 1946. One small
specimen, U.S.N.M. No. E. 7288.
Notes.—The specimen from the Bikini lagoon has R=16 mm., r=4
mm. Inthe specimen from 150-180 feet R=5 mm.
FROMIA EUSTICHA Fisher
Fromia eusticha FisHer, Proc. U. 8. Nat. Mus., vol. 46, p. 213, 1918 (Albatross
station 5146, in the vicinity of Siasi, Tapul group, Jolo Archipelago; 24
fathoms).
Locality.—Bikini Atoll (lat. 11°34’30’’ N., long. 165°30’30’’ E.) ;
180 feet; August 22, 1947. One specimen, U.S.N.M. No. E. 7287.
Genus DACTYLOSASTER Gray
DACTYLOSASTER CYLINDRICUS PACIFICUS Fisher
Dactylosaster cylindricus pacificus FisHer, Bernice P. Bishop Mus. Bull. No. 27,
p. 75, pl. 8, 6. 1925 (Laysan Island).
Locality—Bikini Atoll; Schultz, April 13, 1946. One specimen,
U.S.N.MINo. E. 7271.
Notes—R=11 mm. The animal is completely covered with smal]
granules, with one large pointed granule on the summit of each plate.
Genus OPHIDIASTER L. Agassiz
OPHIDIASTER GRANIFER Liitken
Ophidiaster granifer LUTKEN, Vid. Med., vol. 23, p. 276, 1871 (Tonga Islands).
Localities —Bikini Atoll; Eninman Island, outer reef flats; Bayer,
July 17,1947. One specimen, U.S.N.M. No. E. 7289.
Pian Atoll; Bikini Island, lagoon reef; Bayer, eo 24, 1947.
‘Two specimens, U. S.N.M. No. E. 7270.
Bikini Atoll. One specimen, U.S.N.M. No. E. 7269.
Bikini Atoll; Draeger, 1946. Five specimens, U.S.N.M. No. E. 7822.
Bikini Atoll; Johnson, May 1946. One specimen, U.S.N.M. No. E.
7621.
Eniwetok Atoll; outer reef, southwest side of Grinem Island ; Morri-
son, May 29, 1946. Two specimens, U.S.N.M. No. E. 7513.
Eniwetok Atoll; East Rigili Island; Morrison, May 30, 1946. Two
specimens, U.S.N.M. No. E. 7514.
Rongelap Atoll; outer reef flats of Mellu Island; Morrison, June
19, 1946. One specimen, U.S.N.M. No. E. 7515.
Notes.—In the largest specimen R=33 mm. One of the specimens
from Grinem Island is four-rayed.
MARSHALL ISLANDS ECHINODERMS——CLARK 287
OPHIDIASTER SQUAMEUS Fisher
Ophidiaster squameus Fisurr, U. 8. Fish Comm. Bull. for 1903, pt. 3, p. 1,079,
pl. 81, figs. 6, 6a, 6b, pl. 37, fig. 4, 1906 (Albatross station 4100, Pailolo Chan-
nel, between Maui and Molokai, Hawaiian Islands, 150-151 fathoms).
Locality —Rongelap Atoll; outer reef flats on the ocean side of
Enybarbar Island; Morrison, June 18, 1946. One specimen, U.S.N.M.
No. E. 7516.
Notes —R=40-45 mm. Pedicellariae with straight alveolae are
rather numerous on the abactinal surface, some of them raised on
smooth elongate elevations. The color is bright red and yellow in
broad irregular bands.
OPHIDIASTER LORIOLI Fisher
Ophidiaster lorioli F1isuer, U. 8S. Fish Comm. Bull. for 1903, pt. 8, p, 1,077, pl. 31,
figs. 4, 4a—d, pl. 39, fig. 3, 1906 (Albatross station 3834, south coast of Molokai,
Hawaiian Islands, on reefs).
Locality —Eniwetok Atoll; outer reef, east end of Bogen Island;
Morrison, May 21, 1946. One specimen, U.S.N.M. No. E. 7517.
Notes.—All the rays are of different lengths, up ts 25 mm. long.
Only a single pedicellaria is present, in the center of an interbrachial
area on the oral side. There are two small madreporites.
OPHIDIASTER PUSILLUS Miiller and Troschel
Ophidiaster pusillus MULLER and TRoscHEL, Arch. fiir Naturg., Jahrg. 10, vol. 1,
p. 180, 1844 (Philippines).
Localitiy—Bikini Atoll; Bokororyuru Island; Morrison, April 30,
1946. One specimen, U.S.N.M. No. E. 7518.
Notes —R=20mm. There are seven pedicellariae, each situated ona
second superomarginal, two in two of the interbrachial areas, and one
in each of the other three.
Genus LINCKIA Forbes
LINCKIA MULTIFORA (Lamarck)
Asterias multifora LamMaArcK, Histoire naturelle des animaux sans vertébres,
vol. 2, p. 565, 1816 ( ?Seas of Europe).
Localities.—Bikini Atoll; in the lagoon; 40-80 feet; August 6, 1947.
One specimen, U.S.N.M. No. E. 7272.
Bikini Atoll; Draeger, 1946. Two specimens, U.S.N.M. No. E. 7833.
Bikini Atoll; sandy reef flats behind the outer reef, opposite the
center of Bikini Island; Ladd, April 17, 1946. One comet, U.S.N.M.
No. E. 7539.
Bikini Atoll; Romurikku Island, reef; Schultz, May 18, 1946. One
specimen, U.S.N.M., No. E. 7273.
ISS PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Bikini Atoll; rocky flats inside outer reef, southeast corner of Enyu
Island; Morrison, March 16, 1946. One small specimen, U.S.N.M.,
No. E. 7566.
Bikini Atoll; lagoon side of Yurochi Island, under rock in tide
pool; Morrison, March 22, 1946. One comet, U.S.N.M., No. E. 7542.
Bikini Atoll; outer reef, south (ocean) end of Bokororyuru Island;
Morrison, April 30, 1946. One specimen, U.S.N.M., No. E. 7541.
Bikini Atoll; Enirik Island, lagoon tide pool; Schultz and Brock,
March 20, 1946. Three specimens, U.S.N.M., No. E. 7486.
Bikini Atoll; Enirik Island; Johnson, 1946. One specimen,
U.S.N.M., No. E. 7619.
Bikini Atoll; Johnson, 1946. Two specimens, U.S.N.M., No.
E. 7615.
Eniwetok Atoll; Bogombogo Island; Johnson, May 31, 1946. One
specimen, U.S.N.M., No. E. 7616.
Eniwetok Atoll; outer reef flats, Pujiyoru Island; Morrison, June
2, 1946. One six-rayed comet, U.S.N.M., No. E. 7545.
Eniwetok Atoll; East Rigili Island; Morrison, May 30, 1946.
Thirteen small specimens, of which nine are comets; one comet has six
rays and three madreporites, U. S. N. M., No. E. 75387.
Rongelap Atoll; inner reef flats, sandy and rocky, of Burok Island;
1-2 feet; Morrison, July 24, 1946. One specimen, U.S.N.M., No.
Ee. 7540.
Rongelap Atoll; 2 miles west of Busch Island; 120 feet; Taylor,
June 20, 1946. One comet, U.S.N.M., No. E. 7538.
Rongelap Atoll; Kabelle Island, lithothamnion ridge, intertidal;
Johnson, July 28, 1946. One specimen, U.S.N.M., No. E. 7617.
Rongerik Atoll; Latoback Island, lagoon reef; Bayer and Zim-
merman, August 20, 1947. One specimen, U.S.N.M., No. E. 7338.
Johnston Island, outer reef on the northwest side; Bayer, August
28, 1947. One specimen, U.S.N.M., No. E. 7339.
LINCKIA GUILDINGII Gray
Linckia guildingii Gray, Ann. Mag. Nat. Hist., vol. 6, p. 285, December 1840 (St.
Vincent, West Indies).
Localities —Bikini Atoll; Johnson, May 16, 1946. One specimen,
U.S.N.M., No. E. 7612.
Rongelap Atoll; lagoon; Johnson, June 16, 1946. ‘Two specimens,
U.S.N.M., Nos. E. 7618, EK. 7620.
Notes.—In the specimen from Bikini Atoll one ray is about 35 mm.
long, and the other four are 9-10 mm. The color is purple blotched
with yellowish white.
MARSHALL ISLANDS ECHINODERMS—CLARK 289
LINCKIA LAEVIGATA (Linné)
Asterias laevigata Linn&, Systema naturae, ed. 10, vol. 1, p. 662, No. 8, 1758
(Mediterranean and Indian Seas).
Localities —Eniwetok Atoll; Chinieero Island, reef on outer side
of island, close to island but in water on bottom not likely to be
exposed at low tide; Taylor and Schultz, May 25, 1946. One speci-
men, U.S.N.M., No. E. 7775.
Rongelap Atoll; west side of Naen Island; Herald, July 30, 1946.
One specimen, U.S.N.M., No. E. 7776.
Notes.—Dr. Schultz recorded the color of the specimen from Chi-
nierro Island as a “bright grayish blue or delft blue.” In this speci-
men R=132 mm., r=23 mm. In the one from Naen Island R=177
mm., r=25 mm.
Family ASTERINIDAE
Genus ASTERINA Nardo
ASTERINA CEPHEA (Miiller and Troschel)
Asteriscus cephus (Valenciennes, MS.) MU tier and TroscHEL, System der
Asteriden, p. 41, 1842 (Batavia, Java).
Locality —Eniwetok Atoll; East Rigili Island; Morrison, May 30,
1946. One specimen, U.S.N.M. No. E. 7528.
ASTERINA CORONATA CRISTATA Fisher
Asterina cristata Fisurr, Proc. Biol. Soc. Washington, vol. 29, p. 27, 1916
(Ponapé, Caroline Islands).
Localities —Bikini Atoll; lagoon; 180-200 feet; Schultz, March 29,
1946. One specimen, U.S.N.M. No. E. 7361.
Bikini Atoll; Johnson, May 1946. One specimen, U.S.N.M. No.
E. 7622.
Rongerik Atoll; Latoback Island; lagoon reef, from a clump of
Stylophora mardaz; Bayer and Zimmerman, August 20, 1947. One
specimen, U.S.N.M. No. E. 7362.
Notes—The specimen collected by Dr. Schultz at Bikini is 9 mm.
in diameter and has two regenerating rays. Dr. Johnson’s specimen
from Bikini has seven rays, the two longest 12 mm. long, the other five,
9 mm. The specimen from Latoback Island, Rongerik Atoll, has
seven rays and is 6.5 mm. in diameter.
ASTERINA ANOMALA H. L. Clark
Asterina anomala H. L. CrarK, The Echinoderm fauna of Torres Strait, p. 95,
pl. 7, fig. 8 (colored), pl. 23, fig. 5, pl. 26, figs. 2, 3, 1921 (reef flat, Mer,
Murray Islands, Torres Strait).
Locality. Bikini Atoll; lagoon, about 1 mile inside Bikini Island;
290 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 102
picked off staghorn coral; Bayer, July 16, 1947. One specimen,
U.S.N.M. No. E. 7368.
Notes —R=11 mm., r=5 mm. There are seven rays of nearly
equal length. This specimen agrees well with Dr. H. L. Clark’s orig-
inal description. The form, size, and proportions are the same as
those of the specimen shown in his colored figure. This species is
known from Mer and Lord Howe Islands, from Cape Leveque, West-
ern Australia, and from the Hawaiian Islands. It is abundant at
Lord Howe Island.
Family ECHINASTERIDAE
Genus OTHILIA Gray
OTHILIA LUZONICA Gray
Othilia luzonica Gray, Ann. Mag. Nat. Hist., vol. 6, p. 282, December 1840 (Isle
of Luzon, Philippine Islands).
Localities —Bikini Atoll; lagoon, off western end of Bikini Island;
Cali, April 29,1946. Onespecimen, U.S.N.M. No. E. 7521.
Bikini Atoll; Bikini Island, drift; Morrison, April 5, 1946. One
specimen, U.S.N.M. No. E. 7520.
Bikini Atoll; drift from south side of Uorikku Island; Morrison,
March 28, 1946. One small specimen.
Bikini Atoll; Airukiraru Island, lagoon side; Schultz, April 17,
1946. Three specimens, U.S.N.M. No. E. 7483.
Bikini Atoll; narrow outer reef on southwest (ocean) side of
Ourukaen Island; Ladd, May 2, 1946. One specimen, U.S.N.M. No.
K. 7523.
Bikini Atoll; lagoon reef, a poor narrow reef on the northeast side
of Ourukaen Island; Ladd and Tracey, May 3, 1946. Two small
specimens, U.S.N.M. No. E. 7567.
Bikini Atoll; east side of Chieerete Island, under rocks in tide pools;
March 20, 1946. Two specimens, U.S.N.M. No. E. 7139.
Bikini Atoll; Bokororyuru Island, flats inside outer reef on south
(ocean) end; Morrison, April 30, 1946. Two specimens, U.S.N.M.
No: Ee 519:
Eniwetok Atoll; Rigili Island; Morrison, May 25, 1946. One speci-
men, U.S.N.M. No. E. 7522.
TOU Atoll; east side of Rigili Island; Morrison, May 30, 1946.
Seven specimens, U. S.N.M. No. E. 7490.
Eniwetok Atoll; Lidilbut Island, lagoon reef; Schultz, June 1, 1946.
One specimen, US.NM. No. E. 7484,
Eniwetok Atoll; outer reef, east end of Bogen Island; Morrison,
May 21, 1946. One specimen, U.S.N.M. No. E. 7491.
MARSHALL ISLANDS ECHINODERMS—CLARK 291
Family ACANTHASTERIDAE
Genus ACANTHASTER Gervais
ACANTHASTER PLANCI (Linné)
Asterias planci Linnf, Systema naturae, ed. 10, p. 823, Appendix, 1758 (Goa
[Portuguese India]).
Locality —Bikini Atoll; Eninman Island; Bayer, July 17, 1947.
One specimen, U.S.N.M. No. E. 7391.
Class OPHIUROIDEA
Family OPHIOMYXIDAE
Genus OPHIOMYXA Miiller and Troschel
OPHIOMYXA AUSTRALIS Litken
Ophiomyza australis LUTKEN, Additamenta ad historiam Ophiuridarum, part 35,
p. 99, 1869 (Bass Strait, between Australia and Tasmania).
Localities —Bikini Atoll; one-half mile from Aomoen Island, in
the lagoon, in coral at 42 feet; Brock, Kohler, and Herald, July 10,
1946. Four specimens, U.S.N.M. No. E. 7350.
Rongerik Atoll; Latoback Island, lagoon reef; Bayer and Zimmer-
man, August 18, 20, 21,1947. Four specimens, U.S.N.M. Nos. E. 7351-
E. 7354.
Genus OPHIODERA Verrill
OPHIODERA PUNCTATA, new species
Description.—The disk is stellate with evenly rounded interbrachial
borders (dried), and is 27 mm. in diameter; the arms are 160 mm.
long.
The disk is thickly studded with minute oval concretions uniformly
distributed over the surface but becoming smaller toward the inter-
radial borders; these do not extend out onto the arms, though there
may be a few just beyond the ends of the radial shields. The disk thus
resembles that of Ophiomywxa bengalensis Koehler, but the concretions
are smaller and end abruptly at the arm bases. The radial shields
are conspicuous, 5 mm. long and about 1.5 mm. broad. Their distal
ends are connected across the arm bases by a row of six small rounded
contiguous plates.
There are no upper arm plates, but there are a few scattered rounded
plates on the aboral side of the first two arm segments.
The interbrachial areas below are naked. The genital slits are long,
reaching nearly to the sixth spine-bearing side arm plates. Parallel
to the genital slits and near them is a row of eight or nine short black
lines at right angles to the slits, the outermost about 2 mm. long and
292 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
the following regularly decreasing in length to that just beyond the
oral shield, which is merely a dot.
The oral shields are reniform, nearly twice as broad as long. The
adoral shields are broad, outwardly adjoining the first under arm
plates; they do not quite meet beneath the oral shields. The jaw plates
are narrow with parallel sides, so that there is a long triangle of
membrane between them. There are four mouth papillae. The outer-
most is narrow, conical, and sharply pointed. The next is conical, but
stouter. The third is broader with almost or quite parallel sides and
a broad chisellike tip. The innermost is broadly fan shaped with a
curved and finely serrate outer edge and resembles the teeth.
The under arm plates are large and well developed with a sharp
angle proximally and a deep distal notch; at the arm bases they are
almost in contact; distally they become narrower and somewhat more
separated, and the distal notch becomes deeper. At the arm tips they
are small and widely separated, the side arm plates almost meeting
between them.
The side arm plates beyond the disk bear four, rarely five, subequal
spines of which the uppermost is slightly longer and considerably
stouter than the others, and the two lowest are the most slender.
Toward the arm tips the number of spines becomes reduced to three,
and the upper transforms into a stout hook with three or four teeth.
Later the middle spine also becomes hooklike. At the end of the arms
there are only two spines, both in the form of hooks.
The disk is rather dark brown with numerous white specks and
conspicuous dull yellow radial shields. The arms and the oral surface
are brownish yellow, with a conspicuous row of short black lines along
the genital slits.
Locality —Bikini Atoll; off Enyu Pass, 700-725 feet; August 22,
1947. One specimen, the type, U.S.N.M. No. E. 7367.
Family OPHIACTIDAE
Genus OPHIACTIS Liitken
OPHIACTIS SAVIGNYI (Miiller and Troschel)
Ophiolepis savignyi MULLER and TrROoscHEL, System der Asteriden, p. 95, Species
12, 1842 (Egypt).
Localities —Eniwetok Atoll; in the lagoon, approximately 5 miles
north of the anchorage, about 3 miles west of Bogen Island; 90-120
feet; Taylor. Five specimens, U.S.N.M. No. E. 7250.
Rongerik Atoll; Bock Island; Bayer and Zimmerman, August 19,
1947. One specimen, U.S.N.M. No. E. 7252.
Rongerik Atoll; Latoback Island, lagoon reef, from a sponge;
Bayer and Zimmerman, August 21, 1947. Twenty-one specimens,
U.S.N.M. No. E. 7251.
MARSHALL ISLANDS ECHINODERMS—CLARK 293
Family OPHIOTRICHIDAE
Genus OPHIOTHRIX Miiller and Troschel
OPHIOTHRIX LONGIPEDA (Lamarck)
Ophiura longipeda LAMARCK, Histoire naturelle des animaux sans vertébres, vol.
2, p. 544, 1816 (Mauritius).
Locality —Bikini Atoll; Bikini Island, under rocks on the outer
reef flat; Bayer, August 8, 1947. One specimen, U.S.N.M. No. E. 7342.
OPHIOTHRIX ELEGANS Liitken
Ophiothria elegans LiitKen, Additamenta ad historiam Ophiuridarum, part 3,
pp. 57, 99, 1869 (China Sea).
Locality—Rongelap Atoll; lagoon, 2 miles west of Busch Island;
120 feet; Taylor, June 21, 1946. Three specimens, U.S.N.M. No.
EK. 7584.
OPHIOTHRIX VIRGATA Lyman
Ophiothriz virgata LYMAN, Proc. Boston Soc. Nat. Hist., vol. 8, p. 82, 1861 (Kings-
mill [Gilbert] Islands).
Localities —Bikini Atoll; Bikini; Schultz, April 13, 1948. Three
specimens, U.S.N.M. No. E. 7345.
Bikini Atoll; in the lagoon; Morrison, August 26, 1947. One speci-
men, U.S.N.M. No. E. 7344.
Bikini Atoll; Johnson, May 1946. Six specimens, U.S.N.M. No.
E. 7625.
OPHIOTHRIX TRILINEATA Liitken
Ophiothriz trilineata LirKen, Additamenta ad historiam Ophiuridarum, part
3, pp. 58, 100, 1869 (Samoan Islands).
Locality —Bikini Atoll; Bikini Island, lagoon reef; Bayer, August
24,1947. One specimen.
OPHIOTHRIX EXIGUA Lyman
Ophiothria erigua LyMAN, Bull. Mus. Comp. Zool., vol. 3, part 10, p. 236, pl. 4, figs.
14-26, 1874 (Philippine Islands).
Locality —Eniwetok Atoll; in the lagoon approximately 5 miles
north of the anchorage, about 3 miles west of Bogen Island; 90-120
feet; Taylor, 1947. One specimen, U.S.N.M. No. E. 7329.
OPHIOTHRIX LEPIDA de Loriol
Ophiothria lepida pe LoRIoL, Mém. soe. phys. nat. hist. Genéve, vol. 32, part 1,
No. 3, p. 45, pl. 25, figs. 1-1f, 1893 (Mauritius).
Locality —Bikini Atoll; off Bikini Island; 400-550 feet; August 6,
1947. One specimen, U.S.N.M. No. E. 7340.
Note.——This specimen is of the typical form, not the Hawaiian
subspecies (hawatiensis A. H. Clark). It was taken from a muriciid
alcyonarian.
294 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 102
Family OPHIOCHITONIDAE
Genus OPHIODESMUS Ziesenhenne
OPHIODESMUS DEGENERI A. H. Clark
Ophiodesmus degeneri A. H. Ciark, Bernice P. Bishop Mus. Bull. 195, p. 46,
fig. 17, a, b, p. 47, 1949 (Albatross station 4160, in the vicinity of Moku Manu
[Bird Island], Hawaiian Islands, 57-71 meters).
Localities —Bikini Atoll; in the lagoon, 180-200 feet; Schultz,
March 29, 1946. Two small specimens, U.S.N.M. No. E. 7357.
Bikini Atoll; in the lagoon, 150-180 feet; Schultz, April 18, 1948.
Two specimens, U.S.N.M. No. E, 7358.
Family OPHIOCOMIDAE
Genus OPHIOCOMA L. Agassiz
OPHIOCOMA ANAGLYPTICA Ely
Ophiocoma anaglyptica E1y, Journ. Washington Acad. Sci., vol. 34, No. 11, p. 378,
fig. 1, p. 374, 1944 (Canton Island, reef).
Localities—Bikini Atoll; Namu Island, west of the north point,
under loose flat coral heads; Morrison, April 4, 1946. One specimen,
U.S.N.M. No. E. 7141.
Eniwetok Atoll; reef on the ocean side of Pujiyoru Island; Mor-
rison and Cali, June 2, 1946. Six specimens, U.S.N.M. No. E. 7309.
Eniwetok Atoll; under rocks on rocky shore, south end of Rigili
Island; Morrison, May 25, 1946. Two specimens, U.S.N.M. No. E.
T57A.,
Notes——The apparently quite distinct and easily recognized
Ophiocoma anaglyptica was described in 1944 from Canton Island
in the Phoenix group and has not since been reported. The granules
on the disk are lower and broader than those of O. scolopendrina,
a point not mentioned in the original description.
Mr. Ely has kindly presented his type specimen to the National
Museum (No. E. 6847). The National Museum also possesses speci-
mens from Asor Island, Ulithi Atoll, Caroline Islands, collected and
presented by Lt. Comdr. F. C. Ziesenhenne, U. S. N. R.
OPHIOCOMA SCOLOPENDRINA (Lamarck)
Ophiura scolopendrina Lamarck, Histoire naturelle des animaux sans
vertébres, vol. 2, p. 544, 1816 (Mauritius).
Localities —Bikini Atoll; Bikini Island, under rocks in the lower
intertidal zone near the outer reef; Morrison, March 7, 1946. Three
specimens, U.S.N.M. No. E. 7188.
Bikini Atoll; Namu Island, under loose flat coral heads a little west
of the north point; Morrison, April 4, 1946. One specimen, U.S.N.M.
No. E. 7142.
MARSHALL ISLANDS ECHINODERMS—CLARK 295
Bikini Atoll; Enyu Island, ocean reef; Schultz, May 8, 1946. One
specimen, U.S.N.M. No. E. 7348.
Bikini Atoll; rocky flats inside outer reef, southeast corner of Enyu
Island; Morrison, March 16, 1946. Five specimens, U.S.N.M. No. E.
1573.
Bikini Atoll; Eninman Island, outer reef flats; Bayer, July 17,
1947. One specimen, U.S.N.M. No. E. 7312.
Bikini Atoll; east side of Yomyaran Island; Morrison, April 10,
1946. Two specimens, U.S.N.M. No. E. 7576.
Eniwetok Atoll; Eniwetok Island, reef at low tide; Ziesenhenne,
February 1, 2, 1946. Twenty-two specimens, U.S.N.M. Nos. E. 7333,
E. 7334, E. 7335.
Eniwetok Atoll; Arambiru Island, ocean reef; Schultz, June 3,
1946. Three specimens, U.S.N.M. Nos. E. 7328, E. 7329.
Rongerik Atoll; outer reef flats of Latoback Island; Morrison, June
28, 1946. One specimen, U.S.N.M. No. E. 7575.
Rongelap Atoll; Enybarbar Island, northeast end; Schultz, June
18, 1946. Thirteen specimens, U.S.N.M. Nos. E. 7310, E. 7311.
Kwajalein Atoll; Gugegwe Island; Ziesenhenne, February 9, 1946.
Thirteen specimens, U.S.N.M. No. E. 73387.
OPHIOCOMA ERINACEUS Miiller and Troschel
Ophiocoma erinaceus MULLER and TROSCHEL, System der Asteriden, p. 98, 1842
(Red Sea; Indian Ocean).
Localities —Bikini Atoll; outer reef flats of Bokonfuaaku Island;
Morrison, July 3, 1946. One specimen, U.S.N.M. No. E. 7570.
Bikini Atoll; outer reef, south (ocean) end of Bokororyuru Island;
Morrison, April 30, 1946. One specimen, U.S.N.M. No. E. 7571.
Eniwetok Atoll; Eniwetok Island; Morrison, June 17, 1946. One
specimen, U.S.N.M. No. E. 7140.
Johnston Island, outer reef on northwest side; Bayer, August 28.
1947. Three specimens, U.S.N.M. Nos. E. 7318, E. 7314.
OPHIOCOMA PICA Miiller and Troschel
Ophiocoma pica MULirr and TroscHet, System der Asteriden, p. 101, 1842
(locality unknown).
Localities —Bikini Atoll; Bikini Island; Bayer, 1947. One speci-
men, U.S.N.M. No. E. 7563.
Bikini Atoll; Eniairo Island; Bayer, July 21, 1947. One specimen.
Eniwetok Atoll; outer reef flats, northeast side of Arambiru Island;
Morrison, June 3, 1946. One specimen, U.S.N.M. No. E. 7562.
Rongerik Atoll; Latoback Island, lagoon reef, from a clump of
Stylophora mardax; Bayer and Zimmerman, August 18, 1947. One
specimen, U.S.N.M. No. E. 7356.
Johnston Island; outer reef on northwest side; Bayer, August 29,
1947. One specimen, U.S.N.M. No. E. 7315.
296 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
OPHIOCOMA BREVIPES Peters
Ophiocoma brevipes PETERS, Monatsb, preuss. Akad. Wiss., 1851, p. 465.
Localities —Eniwetok Atoll; East Rigili Island; Morrison, May 30,
1946. One specimen, U.S.N.M. No. E. 7578.
Rongerik Atoll; outer reef flats on the north side of Bigonattam
Island; Morrison, August 21, 1947. One specimen, U.S.N.M., No.
E. 7330.
OPHIOCOMA, sp.
Locality.—Bikini Atoll; lagoon; Morrison, August 26, 1947. Six
very small specimens, U.S.N.M. No. E. 7355.
Genus OPHIOCOMELLA A. H. Clark
OPHIOCOMELLA CLIPPERTONI A. H. Clark
Ophiocomella clippertoni A. H. CLARK, Smithsonian Misc. Coll., vol. 98, No. 11,
p. 7, pl. 1, figs. 1, 2 (as O. parva), June 2, 1939 (Clipperton Island).
Localities —Bikini Atoll; Bikini Island, outer reef; Bayer, July 25,
1947. One specimen, U.S.N.M. No. E. 7347.
Eniwetok Atoll; East Rigili Island; Morrison, May 30, 1946. Two
specimens, U.S.N.M. No. E. 7585.
Rongerik Atoll; Bock Island, in algae on the outer reef flats; Tay-
lor, June 27, 1946. One specimen, U.S.N.M. No. E. 7346.
Genus OPHIOMASTIX Miiller and Troschel
OPHIOMASTIX MIXTA Litken
Ophiomastir miata LUTKEN, Additamenta ad historiam Ophiuridarum, pt. 8,
pp. 42, 99, 1869 (Samoa and Fiji).
Locality —Eniwetok Atoll; Pujiyoru Island, ocean reef; Morrison
and Cali, June 2, 1946. Two specimens, U.S.N.M. No. E. 7821.
OPHIOMASTIX BISPINOSA H. L. Clark
Ophiomastiv bispinosa H. L. CiarKx, Bull. Mus. Comp. Zool., vol. 61, No. 12,
p. 442, pl. 2, figs. 1, 2, 1917 (Paumotu [Tuamotus] Islands, Makemo).
Localities—Bikini Atoll; Bikini Island; Schultz, April 13, 1946.
One specimen, U.S.N.M. No. E. 7324.
Bikini Atoll; in the lagoon; Schultz, March 29, 1946. One speci-
men, U.S.N.M. No. E. 7326.
Bikini Atoll; Johnson, May 1946. One specimen, U.S.N.M. No. E.
7624.
Bikini Atoll; Bikini Island; Schultz, April 13, 1946. Two speci-
mens, U.S.N.M. No. E. 7327.
Bikini Atoll; one-half mile north of Aomoen Island, in the lagoon;
42 feet, in coral; Brock, Kohler, and Herald, July 10, 1946. Two
specimens, U.S.N.M. No. E. 7323.
MARSHALL ISLANDS ECHINODERMS—CLARK 297
Bikini Atoll; 30-75 feet; August 15,1947. One specimen, U.S.N.M.
No. E. 7322.
Notes.—In the last-mentioned specimen (E. 7322) the stellate disk
is 13 mm. in diameter and the arms are 70 mm. long. The disk is
thickly covered with short sharp spinules. There are two tentacle
scales on the first four or six pores. The arm spines are usually three,
sometimes four; the enlarged uppermost arm spines are 5 mm.
long, usually slender, occasionally thickened. The color (as in E.
7324) is olive-brown with the arms obscurely banded.
Specimen No. E. 7323 has the disk 9 mm. in diameter and the
arms 50 mm. long. ‘The disk is thickly beset with very small sharp
spinules. There are two tentacle scales on the first three, sometimes
four, pores. The color is uniform olive-brown, the spines lighter.
Specimen No. E. 7326 has the disk 5 mm. in diameter, and beset with
scattered sharp spinelets. There are two tentacle scales on the first two
or three pores. There are three arm spines. This specimen agrees
well with the type as described and figured by Dr. H. L. Clark.
Dr. Johnson’s specimen (No. E. 7624) has the disk 8 mm. in diameter
and the arms 50 mm. long. The disk is rather thickly beset with
short spines. The arm spines are usually three, occasionally two.
OPHIOMASTIX SEXRADIATA, new species
Description.—The disk is 3 mm. in diameter and the six arms are
14 mm. long. The disk is covered with small imbricating scales and
studded with numerous short stout conical spinelets resembling those
on the disk of Ophiocomella clippertoni.
The general structure is similar to that of O. clippertoni, but there
are only three arm spines—four on the first three or four side arm
plates beyond the disk—and the uppermost arm spine is from half
again to twice as long as the others, slender and tapering gradually
to a sharp tip; near the arm bases it may be considerably longer than
an arm segment. The middle spine is shorter, but usually stouter.
The lowest spine is about as long as the middle spine, and usually
somewhat more slender.
The disk is light brown, the spinules white. The upper arm plates
are light brown with a narrow white border and a narrow white curved
band extending across the plates from the spines on one side to those
on the other. The under arm plates are light brown with a narrow
distal white border. The oral and adoral shields have a dark center
and broad white borders.
In two of the specimens all six arms are of the same size. In the
remainder, three of the arms are much smaller than the others, in-
dicating autotomous division.
Locality Bikini Atoll; in the lagoon; Morrison, August 26, 1947.
Four specimens, U.S.N.M. Nos. E. 7359 (type), E. 7360.
298 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 102
Notes.—In spite of the close agreement in the form of the arm plates
and the plates of the oral surface, and in the number of arms, with the
species of Ophiocomella, this little species should be referred to
Ophiomastixz, with which it agrees in the relations of its arm spines
and in the general scheme of its color pattern. It does not agree with
any of the known species of Ophiomastia, none of which are six-rayed
in any stage. The color pattern also is unique.
Family OPHIODERMATIDAE
Genus OPHIOPEZELLA Ljungman
OPHIOPEZELLA SPINOSA (Ljungman)
Ophiaracna spinosa LIUNGMAN, Ofv. K. Vet.-Akad. Firh., vol. 23, pt. 6, p. 305,
1867 (Foua).
Localities —Bikini Atoll; Eninman Island, outer reef flat; Bayer,
July 17,1947. One specimen, U.S.N.M. No. E. 7210.
Rongerik Atoll; Bock Island; Bayer and Zimmerman, August 19,
1947. One specimen, U.S.N.M. No. E. 7212.
Rongerik Atoll; Latoback Island, lagoon reef; Bayer and Zimmer-
man, August 20, 1947. One specimen, U.S.N.M. No. E, 7211.
Family OPHIOLEPIDIDAE
Genus OPHIURA Lamarck
OPHIURA KINBERGI Ljungman
Ophiura (vel Ophioglypha) kinbergi LaunamaN, Ofv. K. Vet.-Akad. Forh., vol. 23,
p. 166, 1866 (Sydney, New South Wales, Australia).
Locality—Rongelap Atoll; lagoon; 23+ fathoms; Johnson, June
16,1946. One specimen, U.S.N.M. No. E. 7613.
Genus OPHIOLEPIS Miiller and Troschel
OPHIOLEPIS CINCTA Miiller and Troschel
Ophiolepis cincta MULLER and TROSCHEL, System der Asteriden, p. 90, 1842 (Red
Sea).
Locality —Rongerik Atoll; Latoback Island, lagoon reef; Bayer and
Zimmerman, August 21,1947. Twospecimens, U.S.N.M. Nos. E. 7253,
KE. 7254.
Genus OPHIOPLOCUS Lyman
OPHIOPLOCUS IMBRICATUS (Miiller and Troschel)
Ophiolepis imbricata MULLER and TroscHer, System der Asteriden, p. 93, 1842
(Mauritius; Timor).
Localities—Bikini Atoll; Eninman Island, outer reef flats; Bayer,
July 17,1947. One specimen, U.S.N.M. No. E. 7209.
Bikini Atoll; Namu Island, outer reef; Morrison, August 7, 1947.
One specimen, U.S.N.M. No. E. 7208.
MARSHALL ISLANDS ECHINODERMS—CLARK
299
ECHINODERMS FROM OPERATION CROSSROADS, 1946, AND THE
BIKINI SCIENTIFIC RESURVEY, 1947
Species 1946 1947 | Spieces 1946
Crinoidea Asteroidea—Continued
Womaster GraCtts2=—--- oon Rah hae seri sar RATA ARO c OMe ee es 2
Comanthus bennetti__------ oP Se eenmiemenines 2 ten “
Stephanometra indica protec- RES HA eee
tug —— -—-—----------~-~ x X | Dactylosaster cylindricus pa-
Echinoidea cificug _..--------------- x
Ophidiaster granifer.__----- x
Eucidaris metularia___----- x X | Ophidiaster squameus___---- x
Chondrocidaris gigantea__-__ ---- X | Ophidiaster lorioli_...---_-- x
Phyllacanthus imperialis__.. ~--- X | Ophidiaster pusillus_.------ x
Cocopleurus sp--i5------- --_- x |. Linekia multifora_...-.-.- me
Echinothriz diadema_ ------ x X | Linckia guildingii___------ x
Echinothriz calamarts - - - - -- x 'X | Linckia laevigata____----- x
Temnopleurus toreumaticus-. X ----| Asterina cephea__-.-------- x
Mespilia globulus____.----- x ----| Asterina coronata cristata... xX
Desmechinus rufus--------- ---- x | Asterina anomala___.-..-.-. ~-=-
Cyrtechinus verruculatus.-.-. xX X | Othilia luzonica_._....-._- x
Tripneustes gratilla____---- x 1X | Acanthaster planci-...----- ----
Parasalenia gratiosa__-_.___- x x
Echinometra mathaei__----- > lla ib. Ophiuroidea
Echinometra mathaei oblonga. x ?x | Qphiomyzxa australis_.---- x
Echinostrephus aciculatus_._. x X |Ophiodera punctata, new
Heterocentrotus trigonarius_. xX x BDECIEs a eum NOLO re Tiamat
Heterocentrotus mammuillatus. ---- !X | Ophiactis savignyi--.------ ----
Clypeaster reticulatus_._..... ---- X | Ophiothriz longipeda__----- ----
Laganum depressum__------ x X | Ophiothriz elegans_-------- x
Fibularia australis_______-- X 7X | Ophiothrizx virgata__..__---- x
Fibularia ovulum_--------- > Saar. * Ophiothriz trilineata_-_-__=— =
Fibularia volva___--------- x ?x OphiotlinicieLug ia a ee
Pic se Dee hee 7X | Ophiothriz lepida___------- ----
Echinocyamus incertus_..-._ ---- X | Ophiodesmus degeneri_----- x
Echinocyamus megapetalus_. x ?x Ophiocoma anaglyptica.- --- x
Echinocyamus elongatus---.- xX ---- Ophiocoma scolopendrina_.-- xX
Echinoneus abnormalis____~ X ----| Ophiocoma erinaceus_-__---- x
Echinoneus cyclosiomus___-- x X | Ophiocoma pica__--------- x
Echinolampas alexandri__-_-- xX X | Ophiocoma brevipes_------- x
Metalia dicrana__.-__-__-_- en Seal pitoconian apt. |: ie aumento)
Brissus latecarinatus - -- ~~ -- Xx ----| Ophiocomella clippertoni___. x
Maretia ovata____.-------- x X | Ophiomastiz mizta___-_---- Ds
Met aeoiden Ophiomastix bispinosa Chis bande >
Ophiomastix sexradiata, new
Astropecten polyacanthus___- ___~ x BC CIOS eee ne ee eee nee
Luidia mascarena___.--..--- ---- x | Ophiopezella spinosa__-..-- ----
Calliaster elegans____.._..-_ -_-- x | 'Ophrura kinbergr_ 2-2 x
Culcita novae-guineae______- x) * x i Onnhroleprs Cincla= ees eee fos
Neoferdina ocellata_______-- x 2-2] | Onhzoplocisanbricaius oo ==
1 Johnston Island only.
2 Collected in 1947 and 1948 (See text, p. 266).
3 Collected in 1948 only (See text, p. 266).
1947
300 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
ISLANDS ON WHICH COLLECTIONS WERE MADE, WITH THE SPECIES
COLLECTED ON EACH
BIKINI ATOLL
AIRUKIRARU ISLAND: Othilia luzonica.
AOMOEN ISLAND: Ophiomyza australis; Ophiomastiz bispinosa, 42 feet.
ARRIIKAN IstAnD: Fibularia ovulum; Echinocyamus elongatus.
BIKINI ISLAND and vIcINITy: Comaster gracilis, 150-265 feet; Eucidaris me-
tularia, 150-275, 200-290, 800-900 feet; Chondrocidaris gigantea, 800-900
feet; Phyllacanthus imperialis, 150 feet; Coelopleurus, sp., 800-900 feet;
Echinothria diadema; Mespilia globulus, Desmechinus rufus, 800-900 feet ;
Cyrtechinus verruculatus, littoral, 188+, 150, 180-200 feet; Parasalenia
gratiosa, littoral, 30-75, 40-80, 50-140, 180-200 feet; Echinometra mathaei;
Echinometra mathaei var. oblonga; Heterocentrotus trigonarius; Clypeaster
reticulatus, 150-265 feet; Laganum depressum, littoral, 21, 24, 27, 30, 36-42,
90, 150, 168 feet; Fibularia australis; Fibularia ovulum; Fibularia volwa;
Echinocyamus incertus, 800-900 feet; Hehinolampas alexandri, 180, 180-200
feet; Maretia ovata, 50-80, 100-140 feet; Luidia mascarena; Calliaster ele-
gans, 400-450 feet ; Culcita novae-guineae; Fromia hemiopla, littoral, 150-180
feet; Fromia eusticha, 180 feet; Dactylosaster cylindricus pacificus ; Ophidi-
aster granifer; Linckia multifora, littoral, 40-80 feet; Linckia guildingii;
Asterina coronata cristata, littoral, 180-200 feet; Asterina anomala; Othilia
luzonica; Ophiothrix longipeda; Ophiothria virgata, Ophiothria trilineata,
Ophiothrix lepida, 400-550 feet; Ophiodesmus degeneri, 150-180, 180-200
feet; Ophiocoma scolopendrina; Ophiocoma pica; Ophiocoma, sp.: Ophioco-
mella clippertoni; Ophiomastix bispinosa, littoral, 30-75 feet; Ophiomastir
sexradiata.
BOKONFUAAKU ISLAND: EHchinothria diadema; Laganum depressum; Metalia
dicrana ; Ophiocoma erinaceus.
Boxkororyuru ISLAND: Echinometra mathaei; Brissus latecarinatus; Culcita
novae-guineae; Ophidiaster pusillus; Linckia multifora; Othilia luzonica;
Ophiocoma erinaceus.
CHIEERETE ISLAND: Othilia luzonica.
ENrArrO ISLAND: Hchinostrephus aciculatus; Hchinoneus cyclostomus; Ophio-
coma pica.
ENiriK Istanp: Mespilia globulus ; Linckia multifora.
ENINMAN ISLAND: Echinometra mathaei; Ophidiaster granifer; Acanthaster
planci; Ophiocoma scolopendrina; Ophiopezella spinosa; Ophioplocus im-
bricatus.
Enyvu ISLAND: EHchinometra mathaei; Heterocentrotus trigonarius ; Echinoneus
cyclostomus; Linckia multifora; Ophiodera punctata; Ophiocoma scolopen-
drina,
NaMu Istanp: Eucidaris metularia; Echinothrix diadema; Mespilia globulus;
Echinometra mathaei; Echinostrephus aciculatus, Heterocentrotus trigo-
narius; Echinocyamus megapetalus; Echinoneus cyclostomus; Brissus la-
tecarinatus; Neoferdina ocellata; Ophiocoma anaglyptica; Ophiocoma
scolopendrina ; Ophioplocus imbricatus.
OURUKAEN ISLAND: EHchinometra mathaei; Othilia luzonica.
ROMURIKKU ISLAND: Linckia multifora.
UorikkKu ISLAND: Othilia luzgonica.
YoMYARAN ISLAND: Ophiocoma scolopendrina.
YuROCHI IsLAnD: Linckia multifora,
MARSHALL ISLANDS ECHINODERMS—CLARK 301
ENIWETOK ATOLL
ARAMBIRU ISLAND: Echinothrix diadema; Tripneustes gratilla; HEchinometra
mathaei; Echinostrephus aciculatus; Heterocentrotus trigonarius; Culcita
novae-guineae; Ophiocoma scolopendrina; Ophiocoma pica.
Bocen (Jrerorv) IstaAnp: Echinothriv diadema; Parasalenia gratiosa; Echino-
metra mathaei; Laganum depressum; Fibularia ovulum; Metalia dicrana;
Astropecten polyacanthus ; Ophidiaster lorioli; Othilia luzonica; Ophiothria
exvigua; Ophiactis savignyi.
Bocomsoco Istanp: Linckia multifora.
Bocon Istanp: Hudicaris metularia; Fibularia australis.
BuGANEGAN (Mut) IsLtanp: Hehinometra mathaei.
CHINIEERO IstAND: Hehinocyamus megapetalus; Linckia laevigata.
Bast Rieti Istanp (a rock just east of Rigili Island) : Hchinothriz calamaris ;
Mespilia globulus; Tripneustes gratilla; Echinometra mathaei; Echino-
strephus aciculatus ; Fromia balansae; Ophidiaster granifer ; Linckia multi-
fora; Asterina cephea; Ophiocoma brevipes; Ophiocomella clippertoni.
HBLuGeELAB ISLAND: Fibularia ovulum; Fibularia volwa.
ENIWETOK ISLAND: Echinothriz diadema; Tripneustes gratilla; HEchinometra
mathaei ; Heterocentrotus trigonarius ; Ophiocoma scolopendrina; Ophiocoma
erinaceus.
GRINEM ISLAND: Echinometra mathaei; Heterocentrotus trigonarius; Ophidi-
aster granifer.
LipInBput (TEITEIRIPUCCHI) ISLAND: Mespilia globulus; Hchinometra mathaet
var. oblonga; Heterocentrotus trigonarius ; Maretia ovata; Othilia luzonica.
Puutlyoru (Rugoru) Istann: EHchinothriz diadema; Tripneustes gratilla ; Hchino-
metra mathaei; Echinostrephus aciculatus; Heterocentrotus trigonarius;
Culcita novae-guineae; Laganum depressum; Fibularia ovulum; Linckia
multifora; Ophiocoma anaglyptica; Ophiomastia micta.
Ricirr ISLAND (southwestern or seaward end): JZespilia globulus ; Echinometra
mathaei; Echinostrephus aciculatus; Fibularia australis; Fibularia ovulum;
Echinocyamus megapetalus; Hchinoneus abnormalis; Echinoneus cyclo-
stomus; Othilia luzonica; Ophiocoma anaglyptica.
JOHNSTON ATOLL
JOHNSTON ISLAND (F. M. Bayer) : Echinothriz diadema; Echinothria calamaris ;
Tripneustus gratilla; EHchinometra mathaei; Echinometra mathaei var.
oblonga; Heterocentrotus trigonarius; Heterocentrotus mammillatus;
Linckia multifora ; Ophiocoma erinaceus ; Ophiocoma pica.
KWAJALEIN ATOLL
GUGEGWE (BERLIN) ISLAND: Ophiocoma scolopendrina.
KWAJALEIN ISLAND: Hcehinometra mathaei.
RONGELAP ATOLL
Burok IsLAnp: Linckia multifora.
Busow IsLtanp: Mespilia globulus; Heterocentrotus trigonarius; Linckia multi-
fora; Ophiothriz elegans.
ENIAETOK ISLAND: Brissus latecarinatus.
ENYBARBAR ISLAND: Ophidiaster squameus; Ophiocoma scolopendrina.
KABELLE ISLAND: Heterocentrotus trigonarius; Fibularia australis; Fibularia
volwa; Linckia multifora.
KIESHIECHI IsLAND: Comanthus bennetti.
302 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 102
MELLU ISLAND: Ophidiaster granifer.
NAEN ISLAND: Linckia laevigata.
RONGELAP ISLAND: Hechinothriz calamaris; Temnopleurus toreumaticus; Mespilia
globulus, 1388+ feet; Cyrtechinus verruculatus, 138+ feet; Hchinometra
mathaei; Echinometra mathaei var. oblonga; Laganum depressum, 138+
feet; Hchinoneus cyclostomus; Linckia guildingii; Ophiura kinbergi,
138+ feet.
Tura IsLtanp: Stephanometra indica protectus.
RONGERIK ATOLL
BIGONATTAM ISLAND: Ophiocoma brevipes.
Bock IsLanpb: Echinostrephus aciculatus; Fibularia australis; Fibularia ovulum;
Fibularia volwa; Fibularia acuta; Echinocyamus megapetalus; Echinoneus
cyclostomus; Ophiactis savignyi; Ophiocomella clippertoni; Ophiopezella
spinosa.
ENIWETAK (HNYVERTOK) ISLAND: Echinometra mathaei.
LATOBACK ISLAND: Stephanometra indica protectus; Echinocyamus megapetalus;
Linckia multifora; Asterina coronata cristata; Ophiomyza australis;
Ophiactis savignyi; Ophiocoma scolopendrina; Ophiocoma pica; Ophiopezella
spinosa; Ophiolepis cincta.
RONGERIK ISLAND: Hucidaris metularia; Laganum depressum.
ADDITIONAL RECORDS OF ECHINODERMS FROM THE MARSHALL
ISLANDS
From time to time scattered references to specimens from the Mar-
shall Islands have appeared in the literature, based upon material
that has found its way into museums from various sources, much of it
from the Reverend B. G. Snow, a resident missionary. Most of the
specimens are from Ebon (Boston Island), with some from Jaluit
(Bonham Island), both in the extreme south of the Marshall group.
The only systematic collecting that has been done in the Marshalls
was by the Swedish expedition to the South Sea Islands under the
leadership of Dr. Sixten Bock in 1917. A report on the Ophiuroidea
of this collection was published by Prof. René Koehler in 1927
(Géteborgs Kungl. Vet. Vitt. sam. Handl., vol. 33, No. 3), which lists
16 species, mostly from Jaluit and Ebon, of which the following are
not represented in the present collection.
Ophiomyxa brevispina von Martens (Jaluit)
Amphilimna serradiata (Duncan) (Jaluit)
Ophionereis porrecta Lyman (Ebon)
Ophiothrix demessa Lyman (Jaluit)
Ophiothrix galatheae Ltitken (Jaluit)
Ophiothrix triloba von Martens (Jaluit)
Ophiopezella dubiosa de Loriol (Jaluit)
It is probable that Professor Koehler’s Ophiothrix galatheae is the
same as O. longipeda of the present list.
The Crinoidea of this expedition, all collected by Dr. Chr. Hessle,
were described by Professor Torsten Gislén in 1940 (Kungl. Svenska
Vetenskapsakademiens Handlingar, ser. 3, vol. 18, No. 10). He re-
MARSHALL ISLANDS ECHINODERMS—CLARK 303
corded five species of which four are not represented in the present
collection. These are:
Eudiocrinus tenuissimus Gislén (Jaluit)
Lamprometra palmata (J. Miiller) (Ebon; Jaluit)
Cenometra bella var. magnifica Gislén (Jaluit)
Dorometra nana (Hartlaub) (Ebon; Jaluit)
Earlier records of crinoids from the Marshall Islands are Stephano-
metra protectus from Ebon, collected by the Reverend B. G. Snow
in 1877 (A. H. Clark, Bull. Mus. Comp. Zool., vol. 51, p. 242, 1908,
as Himerometra heliaster), recorded also by Gislén and represented
in the present collection; and Lamprometra palmata, also collected
by Mr. Snow at the same time (Hartlaub, Mem. Mus. Comp. Zool.,
vol. 27, No. 4, p. 409, 1912, as Antedon brevicuneata).
Dr. Hubert Lyman Clark in 1915 (Mem. Mus. Comp. Zool., vol. 25,
No. 4, p. 343, No. 13868) recorded Ophiolepis superba from Ebon, where
it had been collected by Mr. Snow in 1877. It has not since been
reported from the Marshall Islands.
Additional species known from Johnston Island (A. H. Clark,
Bernice P. Bishop Mus. Bull. 195, p. 119, 1949) are: Ychinostrephus
aciculatus, EF’. molaris, Brissus latecarinatus, Culcita novae-guineae
forma novae-guineae, Acanthaster planci, Mithrodia fisheri, and
Ophiactis savignyit.
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
Issued
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 : Washington : 1952 art ie ; No. 3304
NEW AMERICAN CYNIPID WASPS FROM GALLS
By Lewis H. Wetp
TWENTY-FOUR new species of cynipids and their galls and two
inquilines are described herein, together with some notes on biology
and synonymy. One new generic name is proposed for three described
gall-making species from the Pacific coast.
Types of the new species are in the United States National Museum
(U.S.N.M.). Paratypes are deposited in institutions that are indi-
cated by the following abbreviations: the American Museum of
Natural History (A.M.N.H.), the Chicago Museum of Natural His-
tory (C.M.N.H.), the Academy of Natural Sciences of Philadelphia
(A.N.S.P.), the California Academy of Sciences (C.A.S.), the
Museum of Comparative Zoology (M.C.Z), Cornell University (C.U.),
the Hlinois Natural History Survey (1I.N.H.S.), and the University
of Utah Museum (U.U.M.).
Genus SAPHONECRUS Dalla Torre and Kieffer
SAPHONECRUS BREVICORNIS (Ashmead), new combination
Synergus brevicornis ASHMEAD, Trans. Amer, Ent. Soc., vol. 23, p. 189, 1896.
The types have the radial cell open; hence the species belongs in
Saphonecrus.
Genus SYNOPHRUS Hartig
SYNOPHRUS MEXICANUS (Gillette)
Synergus mexicana GILLETTE, Trans. Amer. Ent. Soc., vol. 23, pp. 90, 96, 1896.
The type, in the U. S. National Museum, has the petiole smooth and
315
967604—52——1
316 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
the radial cell open; hence the species belongs in Synophrus and was
transferred in “Hymenoptera of America North of Mexico,” p. 613,
1951.
Genus SYNOPHROMORPHA Ashmead
Synophromorpha ASHMEAD, Psyche, vol. 10, p. 45, 1903.
Ashmead, in his key to genera, designated Synophromorpha salicis
as the type of this genus, but no further description of it was ever
published, and no specimen so labeled was found in the Ashmead
collection. It was supposed to have been reared from a dipterous
gall on willow and considered to be a guest fly. He characterized
the genus as having an open radial cell, the petiole not longitudinally
striate, and separable from Synophrus Hartig by having a coriaceous
instead of transversely sculptured mesoscutum with more or less
distinct parapsidal grooves, smaller foveae, incomplete areolet, and
the third segment of the antenna longer than the fourth in the male.
SYNOPHROMORPHA SYLVESTRIS (Osten Sacken), new combination
Synophrus (?) sylvestris OSTEN SACKEN, Ent. Zeit. Stettin, vol. 22, p. 415, 1861.
This species agrees with the diagnosis of Synophromorpha and is
here transferred to that genus, and as the original spelling is permissi-
ble Latin it is retained. It was reared from the gall of Diastrophus
nebulosus Osten Sacken on blackberry in the Washington, D. C., area.
Adults agreeing with paratypes of this species have often been reared
from this gall in this, the type, area and in many other localities,
Range in length of females 2.1-3.25 mm. Average of 31 specimens
2.86 mm. Males 1.8-2.65 mm. Average 2.25 mm. Similar adults
have been reared from galls of Diplolepis tgnota (Osten Sacken) in
the Washington area.
Two new species are described in the present paper. Like sylvestris
they have the face striate, antennae and legs straw yellow, antennae
12-segmented in the female, mesoscutum broader than long, parapsidal
grooves percurrent and broadened posteriorly, foveae smooth, carinae
on propodeum straight, diverging above, abdomen with a hairy ring
at base and a faint indication of a suture between tergites IT and III
as in Ceroptres.
SYNOPHROMORPHA TERRICOLA, new species
Differs from sylvestris in having the mesopleuron smooth instead of
finely aciculate and a fainter coriaceous sculpture on mesoscutum. The
parapsidal grooves closely approach each other at scutellum where
they are separated only by a distinct median groove. Length of fe-
males 2.2-2.9mm. Average of 32 specimens 2.57 mm. Males have 14-
AMERICAN CYNIPID GALL WASPS—WELD 317
segmented antennae. Length 1.8-2.7 mm. Average of 20 specimens
2.42 mm.
Types —U.S.N.M. No. 60105: Type female, allotype, and six para-
types. Paratypes also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., and
M.C.Z.
Habitat—Emerged early in May from galls of Diastrophus radicum
Bassett at East Falls Church, Va. Part of the paratypes are from
galls of D. bassettii Beutenmueller, from Washington, D. C.
SYNOPHROMORPHA RUBI, new species
Differs from S. terricola in having the parapsidal grooves widely
separated behind with a median groove wanting or a mere notch.
Wemales measure 2.2-2.9 mm. Average of 15 specimens 2.46 mm.
Males 1.6-2.2 mm. Average of six specimens 1.77 mm.
Types.—U.S.N.M. No. 60106: Type female, allotype, and four para-
types. Paratypes also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., and
M.C.Z.
Habitat—The types emerged in July from galls of Diastrophus
euscutaeformis Osten Sacken collected at Winnetka, III.
Genus CEROPTRES Hartig
CEROPTRES OBTUSILOBENSIS Weld
Diplolepis q.-obtusilobae KarscH, Zeitschr. fiir Ges. Naturw., vol. 58, pp. 292, 298,
pl. 6, figs. 3, 8a, 3b, 1880.
Cynips obtusilobae DALLA Torre, Catalogus hymenopterorum, vol. 2, p. 47, 1893
(cites Karsch).
The Karsch gall in Berlin is an abrupt terminal stem swelling on
Quercus stellata, from Texas, labeled Cynips quercus obtusilobae.
The three type adults reared from it, which Karsch considered to be
the makers of the gall and related to Veroterus, are all guest flies of the
genus Ceroptres. Dalla Torre was the first to apply a binomial desig-
nation to the species. As obtusilobae was preoccupied in Ceroptres in
1885 by Ashmead (Trans. Amer. Ent. Soc., vol. 12, p. 300) the name
obtusilobensis was proposed in the “Hymenoptera of America North of
Mexico,” p. 614, 1951, for the Karsch species.
Genus LIPOSTHENES Foerster
LIPOSTHENES GLECHOMAE (Linnaeus)
Cynips glechomae LINNAEUS, Systema naturae, ed. 10, p. 553, 1758.
Aulaz latreillei Kierrer, Ent. Zeit. Wien., vol. 17, p. 257, 1898.
Linnaeus did not describe his species, but he cited seven references ;
six of these refer to galls only. However, in one, Reaumur (Memoirs,
1 See p. 315 for explanation of abbreviations used throughout this paper.
318 PROCEEDINGS OF THE NATIONAL MUSEUM Vou. 102
vol. 8, pp. 460-462, figs. 1-5, 1737) says he reared a brown fly with
four wings, similar to those he had reared from oak but with slight
and sufficient differences to indicate it is a distinct species. This
brief characterization of the maker of the gall validates the name that
Linnaeus gave to the species in 1758. He himself evidently had not
seen the maker, for in Fauna Suecica, p. 386, No. 1520, 1761, he says
of the gall “insectum tamen non dum vidi” (however, I have not yet
seen the insect).
Kieffer makes two misstatements when he writes (loc. cit.) : “Linné
hat zuerst aus den Gallipfeln von Glechoma hederacea den Erzeuger
erhalten, und denselben Cynips glechomae genannt; die Beschreibung
lautet: ‘fusca thorace villoso.” Fauna Suecica 1761, pag. 386, Nr.
1520.” Reaumur, in France, was the first to rear the maker. The
phrase “fusca thorace villoso” does not occur in the reference that
Kieffer cites. There is in Geoffroy (Histoire abregée des insectes qui
se trouvent aux environs de Paris . . ., vol. 2, p. 203, No. 20, 1762),
the statement, “Cynips totus fuscus, thorace subvillosus. Sa coleur
est brun et noiratre; son corcelet est un peu velu.” Then, in 1787,
Fabricius (Mantissa insectorum . . . , vol. 1, p. 252), in what is merely
a key to species in each genus, shortens this to “fusca thorace villoso,”
to distinguish glechomae from the preceding species in his key.
The sides of the pronotum are pubescent and the mesoscutum is
bare. In 1841 Hartig described adults he had from Foerster as having
a shining mesonotum and makes the comment that “Cynips glechomae
Lin. (fusca thorace villoso) muss notwandig ein andere Insect gewesen
sein, da die Brust nichtsweniger als behaart ist.” He does not name
it, however. He takes the three Latin words to be a condensed
description instead of a mere phrase to separate it from another species.
This Hartig-Foerster material is what Foerster in 1869 made the type
of his genus Liposthenes.
Latreille also (Histoire naturelle générale et particuliére, des
crustacés et des insectes . . . , vol. 13, p. 207, 1803) reared the maker
and described it as very black, smooth, shining—the same insect
that Hartig and Foerster examined. In 1898 Kieffer gave a new
name to this species, calling it Awlax latreillei, and in 1910 (Das
Tierreich, Lief. 24, p. 668) it appeared as a species distinct from
glechomae Linnaeus from Sweden (sic), its gall not distinguishable
from that of glechomae. Aulawx latreillei is listed as from Britain,
Germany, France, Austria, Italy, and North America.
If the above reasoning about the Latin phrase is correct, then Kieffer
was not justified in giving a new name to the Hartig-Foerster-
Latreille material, and the name Jatrezlec should go into synonymy.
There is only one species making the characteristic gall on Nepeta in
AMERICAN CYNIPID GALL WASPS—WELD 319
Europe and this species was introduced, together with its host plant,
into the eastern United States. It should be known by the specific
name that Linnaeus gave toit. Taylor (Amer. Journ. Botany, vol. 36,
pp. 222-230, February 1949) published an excellent study of the early
stages of this gall.
Genus DIPLOLEPIS Geoffroy
Rhodites Hartig, 1840, was founded on three species and Foerster
in 1869 designated Cynips rosae Linnaeus as the type. The name is
transliterated from a Greek word meaning “pertaining to a rose.”
Thus the name when applied to an insect implies an association with
rose. In both the American and European literature up to 1917 the
maker of a rose gall has been placed consistently in Rhodites.
Neither Hartig nor Foerster seems to have consulted the old litera-
ture. Geoffroy in 1762 (Histoire abregée des insectes, vol. 2, p. 308)
proposed the genus Diplolepis for six species of gall makers, which
he designated simply by number. Under No. 1 he cites Rosel, ins.
vol. 3, suppl., plates 35, 36, and 53, figs. 10, 11, where he describes
and figures an insect and gall on the leaf of oak without name.
(Fourcroy in 1785 gave the name Diplolepis quercus to Geoffroy’s
No. 1.) In the bibliography of his No. 2 he cites among others
Systema Naturae, ed. 10, p. 553, No. 1, 1758, which is Cynips rosae
Linnaeus. The other numbered species lack a bibliography and
have no standing. Thus by a reference the genus is established on
one named species, and the definite designation of Cynips rosae as the
type by Rohwer and Fagan in 1917 (Proc. U. S. Nat. Mus., vol. 53,
p. 365) was unnecessary. They seem to feel that there is some slight
doubt about the supposed designation of C. rosae as type by Geoffroy.
The genus is monobasic. It is next-to-the-oldest name in the Cyni-
pidae and was plainly intended to apply to a gall maker. And yet
it does not appear in the key to genera in Foerster (1869), Mayr
(1851), or Ashmead (1903), either as a valid name or a synonym. It
was used erroneously in 1910 by Dalla Torre and Kieffer in Das
Tierreich, Lief. 24, Cynipidae.
As Fhodites and Diplolepis are isogenotypic, Rhodites disap-
pears in synonymy, although there is sentiment for having it
placed on the conservanda list. Kinsey and Ayres (Indiana Uni-
versity Studies, vol. 9, Study 53, 1922) were the first to adopt Diplo-
lepis in place of Rhodites, and Felt in 1940 (Plant galls and gall
makers) followed their example. The name is coming into increased
use among American authors. Europeans still use Rhodites. Itisa
case of an appropriate name made familiar by a century of usage
against the plain intent of an older author. The establishment of
a name by a reference is not an unusual or doubtful procedure. Lin-
320 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
naeus thus established his Cynips glechomae. Whether plenary
power should be exercised in a case where the obvious intent of an
author is so evident and his procedure so little open to question is a
matter for discussion.
DIPLOLEPIS LENS, new species
Female.—Head and thorax black, abdomen and legs red. Vertex
coriaceous. Antennae 14-segmented. Mesoscutum almost shining but
not smooth, slightly rugose behind; parapsidal grooves percurrent
with a median groove posteriorly. Scutellum rugose, longer than
broad. Mesopleuron with a smooth and shining spot above and a
larger one below. Wing pubescent and ciliate, clouded on radial and
part of cubital cells as in resae. Abdomen not so long as head plus
thorax. Length 1.9-2.6 mm. Average of 23 specimens 2.24 mm.
Smaller than D. bicolor (Harris), whose mesoscutum is more coarsely
rugose behind, without a median groove.
Types.—U.S.N.M. No. 60107: Type and 4 paratypes. Paratypes
also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., and M.C.Z.
Host.—Rosa nutkana.
Gall.—aA lentil-shaped thickening, up to 3.2 mm. in diameter by
1.7 mm. thick, in the parenchyma of the leaf in the fall. Single or
several on a leaf. Similar to the gall of D. rosaefolit Cockerell in
the eastern United States.
Habitat.—The types are from galls collected at Corvallis, Oreg., on
October 16, 1939, along a road northwest of the university campus.
Adults emerged June 7-14, 1940. Similar galls were seen at Holland,
Siskiyou, and Klamath Falls, Oreg.; Chelan and Waitsburg, Wash.,
and Walnut Creek, Calif.
DIPLOLEPIS TERRIGENA, new species
Female.—Head and thorax variegated, red and black, abdomen and
legs red. Antennae 14-segmented, three basal segments slightly red-
dish. Mesoscutum shining, smooth, with scattered punctures; parap-
sidal grooves broad, rugose, percurrent, ending in a rugose area at
pronotum, median groove represented posteriorly by confluent punc-
tures and continued forward by a dark median line. Scutellum ru-
gose, less coarsely so in center. Wing not clouded, areolet reaching
one-fifth way to basal. Propodeum with a smooth median groove.
Abdomen nearly as long as head plus thorax, exposed parts of tergites
back of II finely coriaceous, tergites IV-VII knife-edged dorsally.
Using width of the head as a base the length of mesonotum ratio is
1.27; antenna 2.4; wing 3.0; ovipositor 3.8. Length 3.45-5.0 mm.
Average of 65 specimens 3.99 mm.
Male—Head and thorax black, abdomen red under black, legs be-
yond coxae red. Antennae 15-segmented, third bent. Abdomen
AMERICAN CYNIPID GALL WASPS—WELD 321
shorter than thorax, twice as long as high. Length 2.25-3.25 mm.
Average of 43 specimens 2.92 mm.
Differs from D. fulgens (Gillette), the median groove of which is
as distinct as a parapsidal in posterior third and whose propodeum is
rugose medially.
Types.—U.S.N.M. No. 60108: Type female, allotype, and 20 para-
types. Paratypes also in A.M.N.H., C.M.N.H., A.N.S.P., C.AS.,
M.C.Z., and C.U.
Host—Rosa sp.
Gall (pl. 17, fig. 17) —A large abrupt “root” gall at the crown, up
to 50 mm. in diameter, surface somewhat botryoidal.
Habitat—The types are from galls collected at Ojai, Calif., on
April 6, 1922. They contained pupae at that time. Adults emerged
April 26 to May 20. Galls were seen also at St. Helena and Los Gatos,
Calif.
DIPLOLEPIS POLITA (Ashmead)
Rhodites polita ASHMBAD, Colorado Biol. Assoc. Bull. 1, pp. 14, 38, 1890.
Rhodites occidentalis BEUTENMUELLER, Brooklyn Ent. Soe., vol. 17, p. 45, 1922
(new synonymy).
Ashmead described polita from a gall similar to the eastern Diplo-
lepsis bicolor (Harris), found on Rosa californica, but adds the state-
ment that he has seen galls from Dakota and Colorado also. As the
types in the U. S. National Museum are from Marin County, Calif:,
and bear Koebele’s label with No. 70 in red ink, the type locality is
California, and the Dakota and Colorado material was misdetermined.
There are males of this species in collections from Los Angeles but not
marked as types.
Fullaway reared adults, which he determined to be Rhodites bicolor
(Harris), from Cupertino (Stevens Creek), Calif., and described them
in 1911 (Ann. Ent. Soc. Amer., vol. 4, p. 377). In 1922 Beutenmueller
gave this bicolor Fullaway the name of Rhoditis occidentalis. The
U.S. National Museum has a female specimen from Cupertino (Stev-
ens Creek) which must be a type of the Fullaway description. It
agrees with types of 2. polita Ashmead. It measures 3.1mm. The
5 mm. measurement in the description must be a misprint for 3 mm.
Genus NEUROTERUS Hartig
NEUROTERUS TANTULUS, new species
Female.—Black. Head from above about twice as wide as long,
broader than thorax, cheeks slightly broadened behind eyes; from in
front broader than high, malar space one-third eye with a faint groove,
interocular space broader than high. Antennae 12-segmented, lengths
as (scape) 25(12):18(14) :25(6):17:16: 16: 16:16:15: 14:12:22
322 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
(9). Mesoscutum without trace of parapsidal grooves. Wing pubes-
cent and ciliate, second cross-vein at angle of 55 degrees with basal,
areolet reaching one-fourth way to basal. Claw with a weak tooth.
Abdomen higher than long, ovipositor often extruded. Using width
of the head as a base the length of mesonotum ratio is 1.17; antenna
1.8; wing 4.3; ovipositor 3.8. Range in length of 100 measured speci-
mens 1.75-2.05 mm. Average 1.91 mm.
Types.—U.S.N.M. No. 60109: Type and 20 paratypes. Paratypes
also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., M.C.Z., I.N.HLS., and
C.U. A gall is on each pin.
Host.—Quercus alba.
Gall (pl. 16, fig. 1) —Small saucer-shaped galls occurring in large
numbers attached by a broad base on under side of leaf in early sum-
mer. Individual galls measure .9-1.6 mm. in diameter by .6-.7 mm.
high, with an elevation in center of the concave free surface, brown,
almost bare, dropping off when mature and leaving a scar .7 mm. in
diameter on the gall and a brown spot on the leaf.
Habitat—Type locality, College Park, Md. On May 24, 1948, W. H.
Anderson swept up a tablespoonful of the galls, which had fallen on
a table under a heavily infested white oak tree. These were placed
in a breeding cage on the ground outdoors and when examined on
April 1, 1949, scores of adults had emerged and many had already
died.
Genus TRICHOTERAS Ashmead
TRICHOTERAS PERFULVUM, new species
Female.—Piceous; mandibles, parts of scutellum and legs, brown,
Head coriaceous, pubescent; from above transverse, as broad as
thorax, cheeks not broadened behind eyes; from in front almost cir-
cular, malar space one-third eye, without groove, antennae filiform,
12-segmented, lengths as (scape) 16:11: 20:19:17:17:15:11:10:9:
8:18. Pronotum and mesonotum somewhat shining with uniformly
distributed appressed hairs from prominent punctures. Mesoscutum
broader than long, parapsidal grooves percurrent. Scutellum rugose
posteriorly, with two widely separated smooth pits at base. Wing
pubescent and ciliate, veins brown, third abscissa of subcosta almost
parallel with second abscissa of radius, radial cell five times as long
as broad, areolet present. Mesopleuron largely bare, smooth, polished.
Tarsal claws with a strong tooth. Carinae on propodeum slightly
bent, area on either side pubescent. Abdomen bare except for usual
pubescent patches at base of tergite II, tergite II foliiform. Ventral
spine slender, about five times as long as broad in side view, shorter
than hind metatarsus. Using width of the head as a base the length
AMERICAN CYNIPID GALL WASPS—-WELD 323
of mesonotum ratio is 1.3; antenna 2.6; wing 4.7. Length 1.75-2.15
mm. Average of five specimens 1.99mm. Differs from 7’. coquilletti
Ashmead in having fully developed wings, from amber-colored 7’.
vaccinitfoliae (Ashmead) in having cheeks not broadened behind the
eyes.
Types —uU.S.N.M. No. 60110: Type and paratype. Paratypes also
in A.M.N.H. and C.AS.
Host.— Quercus chrysolepis.
Gall (pl. 17, fig. 20).—Tan colored, circular, 6-8 mm. in diameter
by i mm. high, single or in two’s or three’s, attached to midrib on
upper or lower side of leaf in the fall. Sparingly hairy, thin walled,
with an eccentric ellipsoidal cell inside held in place by radiating fibers.
Habitat.——The types are from galls collected at Idyllwild, Calif.,
November 9, 1948. Both living and dead adults were found in the
galls when they were cut open, February 9, 1949. Galls have been
seen in the San Bernardino, San Gabriel, and Santa Lucia Mountains,
in Giant Forest, Big Basin, on Mount St. Helena, and at Ukiah, Calif.
Fresh galls are fully grown by mid-June.
TRICHOTERAS ROTUNDULA, new species
Female.—Particolored, black and brown. Head coriaceous; from
above transverse, cheeks not broadened behind eyes; from in front
malar space one-half eye, without groove, faintly striate; antennae
12-segmented, first four segments as 10:8:15:13-last 11. Meso-
scutum broader than long, not quite smooth, with uniformly dis-
tributed pubescence, parapsidal grooves percurrent, a median groove
present posteriorly. Scutellum punctate, foveae smooth. Meso-
pleuron bare and smooth above, pubescent below. Wing pubescent
and ciliate, veins slender, brown, first abscissa of radius arcuate, radial
cell elongated, areolet small. Claws toothed. Carinae on propodeum
diverging below. Abdomen as long as head plus thorax, lengths of
tergites on dorsal margin as 65:138:4, tergite II foliiform. Ventral
spine slender, as long as hind metatarsus. Using width of head as a
base the length of mesonotum ratio is 1.3; antenna 2.4; wing 4.6.
Length 2.0-2.05 mm. Described from three specimens. Differs from
7. perfulvum in having a longer striate malar space and a longer
ventral spine.
Types. —U.S.N.M. No. 60111: Type. Paratype in C.A.S.
Host.—Quercus chrysolepis.
Gall (pl. 16, fig. 2)—A midrib cluster of a few globular galis,
usually on the under side of a leaf in the fall. Covered with short
straight hairs when young and green, later bare, smooth, tan, up to
2.7 mm. in diameter with a central cell 1.5 by 1.2 mm. supported by
stout radiating fibers.
967604522
324 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Habitat.—The type emerged in November from a gall collected at
Camp Baldy, Calif., on November 7, 1939. One paratype was cut out
September 5, 1918, from a gall collected at Los Gatos. Another was
cut out November 10, 1949, from a gall collected at Idyllwild on No-
vember 6, 1948. One (not in the type series and all amber) was cut
out dead in 1946 from a gall collected on Mount St. Helena in August
1922.
PARACRASPIS, new genus
Genoty pe-—Callirhytis quadaloupensis Fullaway.
Agamic female.—Head massive, broader than thorax, cheeks broad-
ened behind eyes, malar space without groove, antennae 14-segmented,
filiform, third longer than fourth, longer than first two. Thorax
normally arched; pronotum narrowed to one-ninth in middle. Meso-
scutum broader than long, parapsidal grooves complete or incomplete,
median wanting. Scutellum rounded behind, with a transverse groove
at base, lateral impressions prominent. Mesonotum covered with uni-
formly distributed appressed hairs which almost hide the sculpture.
Mesopleuron pubescent. Tarsal claws with a tooth. Wings reduced
but with evident and nearly normal venation, reaching about to the
tip of abdomen. Abdomen longer than head plus thorax, second ter-
gite longest but not reaching half the length of the abdomen, remain-
ing five more or less subequal, sides of III-VI not pubescent. Ventral
spine broad, rounded at end, bristly as in Aéraspis Mayr. Ovipositor
almost straight at end, rather short and stout.
Differs from Acraspis Mayr in having less reduced wings, a more
robust thorax with a normally rounded scutellum. Genus erected to
include three agamic species, all from leaf galls not of the “hedgehog”
type on Quercus chrysolepis in California. Sexual generation un-
known.
Included species—The following species are included in the new
genus:
PARACRASPIS GUADALOUPENSIS (Fullaway)
Callirhytis guadaloupensis FULLAWAY, Ann, Ent. Soc. Amer., vol. 4, p. 868, 1911.
PARACRASPIS INSOLENS (Weld)
Acraspis insolens WELD, Proc. U. S. Nat. Mus., vol. 68, art. 10, p. 59, 1926.
PARACRASPIS PATELLOIDES (Weld)
Acraspis patelloides WELD, Proc. U. S. Nat. Mus., vol. 68, art. 10, p. 60, 1926.
AMERICAN CYNIPID GALL WASPS—WELD 325
Genus LIODORA Foerster
LIODORA COMATA, new species
Female.—Black ; mandibles, palpi, base of antennae and legs yellow-
ish white. Head from above transverse, as broad as thorax, cheeks not
broadened behind eyes; from in front wider than high, malar space .3
eye, without groove, antennae 14-segmented, lengths of first five as 9:
6:11:10:9. Sides of pronotum, mesoscutum, and mesopleuron
smooth, bare, polished. Parapsidal grooves narrow, percurrent. A
distinct suture between mesoscutum and scutellum. Two smooth, shal-
low pits at base of scutellum, disk smooth, punctate and pubescent
behind and overhanging propodeum. Carinae on propodeum slightly
bent. Wing pubescent and ciliate, radial cell five times as long as
broad, veins brown, areolet small, first abscissa of radius arcuate.
Claws with a strong tooth. Abdomen as high as long, tergites smooth,
all visible on dorsal margin. Ventral spine stout, twice as long as
broad, not bristly. Ovipositor curved at tip. Length 1.45-2.15 mm.
Average of 30 specimens 1.7 mm.
Male.—Head broader than thorax. Antennae 15-segmented, lengths
of first five as 9:6: 16:11:10, third bent and lighter in color. Abdo-
men with a short pedicel. Length 1.45-1.8 mm. Average of six
1.64 mm.
Types—U.S.N.M. No. 60112: Type female, allotype, and six para-
types. Paratypes also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., and
M.C.Z.
Host.—Quercus alba.
Gail (pl. 16, fig. 11).—Conical, 3.0 by 1.4 mm., tan, thin walled, cov-
ered with crinkly white matted hairs. Produced singly at edge of
leaf in early spring when leaves are only an inch or so long.
Habitat—Type locality East Falls Church, Va. Adults emerged
April 26 and May 4-15.
Genus AMPHIBOLIPS Reinhard
AMPHIBOLIPS GLOBUS, new species
SEXUAL GENERATION
Female.—Head, thorax, antennae, legs except fore and middle tibiae
and tarsi, black; abdomen red to almost black. Head rugose, from
above narrower than thorax, cheeks not broadened behind eyes; from
in front broader than high, malar space .4 eye, with radiating ridges,
antennae filiform, 13-segmented. Sides of pronotum coarsely reticu-
late. Mesoscutum coarsely rugose, the percurrent parapsidal and
median grooves also rugose. Scutellum more coarsely rugose behind,
pits shallow, a median depression on disk hardly apparent. Meso-
326 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
pleuron rugose, more coarsely so anteriorly. Wing pubescent and
ciliate, the blotch at base of radial cell covering the areolet also. Tar-
sal claws toothed. Carinae on propodeum strongly bent, enclosed area
rugose with a short median. Abdomen about as long as thorax, lengths
of tergites on dorsal margin as 38: 8, tergite IIT smooth on hind margin,
III punctate. Ventral spine pubescent, as long as hind metatarsus.
Using width of head as a base the length of mesonotum ratio is 1.5;
antenna 2.1; wing 3.2; ovipositor 2.0. Length 4.15-4.8 mm. Average
of 10 specimens 4.58 mm.
Male.—Similar but abdomen black. Antennae 15-segmented, the
third stoutest, the flagellum tapering to tip. Abdomen shorter than
thorax, all tergites visible on dorsal curvature. Antenna ratio 3.3.
Length 3.5-4.0 mm. Average of 11 specimens, 3.74 mm.
Differs from A. spongifica (Osten Sacken), whose tergite IT is punc-
tate posteriorly.
Types —U.S.N.M. No. 601138: Type female, allotype, and three para-
types. Faratypes also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., and
M.C.Z.
Host.— Quercus palustris.
Gall (pl. 16, fig. 4, a2) —A bud gall in June produced from weak
lateral buds on previous year’s growth of saplings 1 to 3 feet high,
on sprouts from stumps and on lower limbs of small trees, occasionally
from dormant buds on main trunk of trees. Jn good light, deep green
with white spots at attachment of radiating fibers inside; in deep
shade, more or less etiolated. Globular, firm, fleshy, bare, up to 18
mm. in diameter, not deciduous. Wallimm. thick. Central cell rela-
tively large, supported by many coarse, branching, radiating fibers.
Larval cell 6 mm. by 3.2 mm. and its wall 1.5 mm. thick with big cells
at base of the radiating fibers. Exit hole 2.2 mm. in diameter made
while gall is still green. After the emergence of the maker the gall
turns brown and becomes wrinkled. Many undersized galls are para-
sitized. Out of more than 200 galls collected only 20 makers emerged.
Habitat—The type series of galls was collected at East Falls
Church, Va. June 26, 1948. Adults emerged June 28, 30, July 2, 4,
1948. None of these galls was seen at this place in 1949, in 1950, or
in 1951. Galls had been seen at Rosslyn and Lucketts, Va., Washing-
ton, D. C., and Lanham, Md.
AGAMIC GENERATION ?
In early September 1948 approximately 65 bud galls were collected
on the ground under a large pin oak tree only a few feet from where
galls of A. globus herein described were taken the previous June.
Similar galls had not been seen here before—circumstantial evidence
that it is the alternating form. It occurs in August and September at
AMERICAN CYNIPID GALL WASPS—-WELD Bye
the end of the season’s growth. It is smooth, bare, green with small
purple spots marking the attachment of numerous radiating fibers
within, up to 14 mm. in diameter, rounded at distal end, slightly taper-
ing at attached base, the wall relatively thin (.8 mm.) (pl. 16, fig. 4, b).
Larval cell ellipsoidal, 6 by 4mm. Galls drop in late August or early
September and soon turn brown and become slightly wrinkled. At
this time the nutritive layer is all used up and full-grown larvae and
pupae are found inside. Some may emerge in the fall but the adults
described below were cut out of the galls on October 18 and November
5, 1949. As they are thought to be the alternating agamic generation
of globus they are not given a different specific name.
Agamie female.—Head and thorax black, abdomen red, tibiae and
tarsi brown. Cheeks broadened behind eyes. Antennae 13-seg-
mented. Mesoscutum rugose with parallel longitudinal ridges back
of anterior lines and short transverse ridges on either side of them;
parapsidal grooves shallow, rugose, percurrent, a median streak per-
current in the sculpture. Scutellum rugose, without distinct median
depression or posterior emargination, foveae with oblique ridges in
bottom. Large spot on base of radial cell covers areolet also. A short
median carina on propodeum. Abdomen as long as thorax, all tergites
normally showing on dorsal curvature, hind margin of IT smooth, rest
punctate, VII pubescent. Ventral spine longer than hind metatarsus.
Mesonotum ratio 1.5; antenna 2.0; wing 4.0. Length 3.9-4.55 mm.
Average of three 4.3 mm.
Genus ANDRICUS Hartig
ANDRICUS BRUNNEUS Fullaway
Andricus brunneus FULLAWAY, Ann. Ent. Soc. Amer., vol. 4, p. 353, 1911.
This species was described from a “thin-walled subglobular gall,
pointed at both ends and about the size of a pea” on the leaves of
Quercus douglasii. It seems to have been associated with the wrong
gall. The type gall in the Stanford collection, agreeing with the above
description but with a central cell supported by radiating fibers,
seems to be a gall of what was described under the name of Andricus
atrimentus Kinsey. I have reared adults agreeing with the types from
a midrib cluster on this host at Stanford University, the type locality,
and also from similar galls on Quercus lobata, Q. dumosa, and Q. garry-
ana. These galls are from 3-5 mm. in diameter, in a cluster of 4 to
6 on the under side of the leaf (pl. 17, fig. 16), tan, slightly mottled,
slightly pubescent, thick walled without any radiating fibers or free
central cell, dropping in September. From galls collected on Quercus
douglastvi at Stanford University on October 27, 1948, adults emerged
October 18 to November 5, 1949.
328 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
ANDRICUS ALBICOMUS, new species
Female.—Red, often with some black on occiput, foveae, propodeum,
and parts of abdomen. Head coriaceous; from above vertex bare,
cheeks slightly broadened behind eyes, occiput not concave; from in
front malar space 0.4 eye, without groove, antennae filiform, 13-seg-
mented. Mesoscutum microcoriaceous, shining, longer than broad,
with scattered hairs, parapsidal grooves percurrent. Scutellum rugose,
pubescent, margined on sides, foveae smooth. Mesopleuron bare,
smooth. Wing hyaline, pubescent and ciliate, veins pale and slender,
radial cell four times as long as broad, areolet and cubitus almost
obsolete. Claws toothed. Abdomen jonger than head plus thorax,
almost as high as long, lengths of tergites as 45:18:8:4:4, slightly
gibbous below petiole, the curved tip of ovipositor projecting beyond
sheaths. Ventral spine longer than hind metatarsus. With width
of head as a base, the length of mesonotum ratio is 1.4; antenna 2.9;
wing 4.9; ovipositor 5.8. Range in length 1.5-2.4 mm. Average of
53 specimens 2.0 mm.
Close to Andricus kingi Bassett, whose disk is smooth back of septum
and antennae are 14-segmented.
Types.—U.S.N.M. No. 60114: Type and 10 paratypes. Paratypes
also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., and M.C.Z.
Host.—Quercus garryand.
Gall (pl. 16, fig. 5)—A thin-walled conical gall about 5 mm. in
diameter on the under side of a leaf in the fall, densely covered with
pedicelled, stellate grayish-white hairs. Inside is a transverse basal
larval cell with a distal cavity opening at apex.
Habitat.—The types were cut out October 18, 1949, from galls col-
lected at Cave Junction, Oreg., on October 14, 1948. Others emerged
in outdoor breeding cage on April 1, 1950. Galls were seen also at
Corvallis and at Siskiyou summit, Oregon, on United States High-
way 99.
ANDRICUS CHICO, new species
Female—Light brown; eyes, ocelli and sheaths black. Head coria-
ceous; from above transverse, cheeks broadened behind eyes, occiput
concave; from in front interocular area wider than high, malar space
one-third eye, without groove, fan-striae about mouth, antennae fill-
form, 13-segmented. Mesoscutum coriaceous with short uniformly
distributed pubescence from distinct punctures, parapsidal grooves
percurrent. Septum between the foveae broad, disk rugose. Meso-
pleuron smooth under hind wing, striate below, almost bare. Wing
pubescent and ciliate, veins beyond second cross-vein pale, areolet dis-
tinct. Claws toothed. Carinae on propodeum straight and parallel.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102, PLATE 16
tantulus
multicostatus subcostata
1, Neuroterus tantulus on Quercus alba; 2, Trichoteras rotundula on Q. chrysolepis; 3,
Dryocosmus grumatus on Q. wislizent; 4, Amphibolips globus on Q. palustris; 5, Andricus
albicomus on Q. garryana; 6, Andricus chico on Q. lobata; 7, Callirhytis oblata on Q.
falcata; 8, Andricus projectus on Q. chrysolepis; 9, Andricus multicostatus on Q.
subturbinella; 10, Callirhytis subcostata on Q. stellata; 11, Liodora comata on Q. alba; 12,
Zopheroteras guttatum on Q. palustris.
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102, PLATE 17
prescotti
minusculus
cistella
13, Dryocosmus minusculus on Quercus agrifolia; 14, Callirhytis hopkinst on Q. imbricaria;
15, Andricus prescotti on Q. subturbinella; 16, Andricus brunneus on Q. dumosa; 17,
Diplolepis terrigena on Rosa sp.; 18, Andricus crenatus on Q. dumosa and Q. douglasti;
19, Antron pileus on Q. subturbinella; 20, Trichoteras perfuloum on Q. chrysolepis; 21,
Callirhytis furva on Q. palustris; 22, Callirhytis cistella on Q. emoryt.
AMERICAN CYNIPID GALL WASPS—WELD 329
Abdomen as long as head plus thorax, as high as long, all tergites
showing on dorsal margin. Ventral spine slender, longer than hind
metatarsus. Using width of the head as a base the length of mesono-
tum ratio is 1.3; antenna 2.3; wing 3.8. Length 2.3-2.6 mm. Aver-
age of three specimens 2.48 mm.
Differs from A. foliatus (Ashmead) in the shorter and more sparse
pubescence on thorax and in having a distinct areolet.
Type.—U.S.N.M. No. 60115: Type. Paratype in C.A.S.
Host—Quercus lobata.
Gall (pl. 16, fig. 6) —A smooth, bare, ovoid bud gall found in fall
in the axils of leaves, greenish and mottled with purple, projecting
beyond the bud scales, 2-3 mm. in diameter.
Habitat.—The types were cut out November 10, 1949, from galls
collected at Chico, Calif., on October 18, 1948. Galls have been seen
at Stanford University, Los Gatos, Santa Margarita, Lakeport, Ukiah,
and Willits, Calif.
ANDRICUS PROJECTUS, new species
Female—Body reddish brown, infuscated dorsally; antennae and
legs lighter. Head coriaceous; from above transervse, occiput not
concave, cheeks broadened behind eyes; from in front broader than
high, interocular space broader than high, malar space one-half eye,
a few striae from corners of clypeus, antennae filiform, 13-segmented.
Mesoscutum pebbled, almost bare, parapsidal grooves percurrent, no
median. Disk of scutellum pubescent, rugose except back of septum,
foveae smooth, shallow. Mesopleuron and sides of propleura bare,
finely striate. Wing hyaline, hairs on surface short, not ciliate, veins
almost clear, areolet small, radial cell five times as long as broad.
Claws with a strong tooth. Carinae on propodeum straight, parallel.
Abdomen longer than head plus thorax, lengths of tergites along
dorsal curvature as 44:11:2:2:9; hind margin of II smooth, ex-
posed parts of rest punctate. Ventral spine slender, seven or eight
times as long as broad. Using width of head as a base, the length of
mesonotum ratio is 1.3; antenna 2.5; wing, 4.5; ovipositor 3.4. Range
in length 1.15-3.0 mm. Average of nine specimens 2.12 mm. Differs
from A. niger Tavares in having tergite IIT punctate.
Types U.S.N.M. No. 60116: Type and one paratype. Paratypes
also in C.A.S. and A.N.S.P.
Host — Quercus chrysolepis.
Gall (pl. 16, fig. 8).—Bursting out far beyond the bud scales in the
fall, cylindrical, bare, smooth, greenish with a light tan apex. When
detached the base is darker, with a depressed scar at the truncated
end, 7.0 mm. long by 2.1 mm. in diameter. Monothalamous. Usually
occurs at tip of strong sprouts from stumps.
330 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Habitat—The type emerged April 16, 1943, from galls collected
at Canyonville, Oreg., October 18, 1939. One was cut out February
17, 1941. Two paratypes were cut out October 18, 1949, from galls
collected at Canyonville on October 12, 1948. Three emerged April
15, 1941 and April 20, 1942, from galls collected on the Mt. Wilson
trail above Sierra Madre, Calif., on November 4, 1939. One is from
Colfax, Calif. Galls were seen also at the California-Oregon State
line on United States Highway 199 and at San Rafael and Idyllwild,
Calif.
ANDRICUS CRENATUS, new species
Female-——Red or amber, slightly infuscated. Head coriaceous;
from above transverse, vertex bare, occiput not concave, cheeks
broadened behind eyes; from in front as high as wide, malar space
one-third eye, without groove, antenna 14-segmented, infuscated dis-
tally. Sides of pronotum faintly striate. Mesoscutum microcoriace-
ous, shining, longer than wide, high arched in profile, with scattered
hairs, parapsidal grooves percurrent. Scutellum longer than wide,
sides margined, finely rugose, pubescent, the two deep smooth foveae
at base often infuscated. Mesopleuron largely bare, smooth. Wing
pubescent and ciliate, veins slender, radial cell 4.7 times as long as
broad, areolet almost obsolete. Claws toothed. Carinae on propo-
deum bent, enclosed area smooth, narrowed above. Abdomen longer
than head plus thorax, longer than high, all tergites usually showing
on dorsal margin, ventral spine slender, longer than hind metatarsus.
Using width of the head as a base the length of mesonotum ratio is
1.4; antenna 2.7; wing 5.1. Length 1.5-2.45 mm. Average of 24
specimens 1.9 mm.
Differs from Andricus pattersonae Fullaway in its smaller size.
Types—U.S.N.M. No. 60117: Type and three paratypes. Para-
types also in the C.M.N.H., A.N.S.P., C.A.S., and the M.C.Z.
Hosts —Quercus dumosa and Q. douglasiz.
Gall (pl. 17, fig. 18). —A spangle about 4 mm. in diameter, usually
on the upper side of the leaf, saucer shaped, with a thin crenate mar-
gin when young in August and with a prominent hump in center.
When mature in fall there is a lens-shaped larval cavity inside, on the
floor of which is a thin, white, circular disk from which prominent lines
radiate. Gallson Quercus douglasii are less crenate.
Habitat—The type is from a series of dead adults cut out of galls
collected on Quercus dumosa at Los Gatos, Calif., on December 13,
1935. Others, all from California, are from the San Bernardino
Mountains, San Jacinto Mountains, and Banning; other paratypes
emerged November 19, 1935, from galls collected at Colfax, on Quercus
douglasii a few days previously ; others are from Stanford University
AMERICAN CYNIPID GALL WASPS—-WELD 331
and Angels Camp. Similar galls have been seen on Quercus engel-
manné at Santa Anita, Calif. Adults not included in the type series
have been reared from galls on Quercus garryana at Siskiyou summit
north of Holbrook, Calif.
ANDRICUS MULTICOSTATUS, new species
Female—Straw yellow, head and thorax slightly darker, only
ventral valves and tips of mandibles infuscated. Head from above
transverse, as broad as thorax, cheeks short, not broadened behind
eyes, occiput not concave; from in front broader than high with a
median ridge above clypeus, malar space striate, half length of eye,
antennae 13-segmented, filiform. Mesoscutum coriaceous, with
scattered hairs, without evident punctures, parapsidal grooves per-
current, no median. Scutellum rugose with two deep, smooth, shin-
ing pits, septum narrow. Carinae on propodeum straight and par-
allel. Mesopleuron finely striate. Wing hyaline, pubescent and cil-
iate, first abscissa of radius arcuate, second pale, only distal vein of
areolet evident. Tarsal claws toothed. Abdomen longer than head
plus thorax, higher than long, only tergites II and III showing on
dorsal margin, both smooth; ventral spine slender, almost as long
as hind metatarsus. With the width of the head used as a base the
length of mesonotum ratio is 1.8; antenna 2.2; wing 4.0. Length
2.2 mm.
Type.—U.S.N.M. No. 60118: Holotype.
FHost.—Quercus subturbinella.
Gall (pl. 16, fig. 9)—An axillary or terminal bud gall in fall.
Cylindrical with 10 to 20 distinct longitudinal ribs, up to 5 mm. long,
protruding far beyond the bud scales. Greenish with white spots
when fresh, the bulbous distal third smooth and reddish.
Habitat—The type was cut out alive on October 14, 1944, from a
gall collected at Prescott, Ariz., September 13, 1943, by Mrs. N. W.
Capron. She collected galls at Cherry and Prescott, Ariz., on Oc-
tober 4, 1935, and September 11, 1947, respectively.
ANDRICUS PRESCOTTI, new species
Female.—Black; antennae, palpi, legs distally and abdomen ven-
trally brown. Head from above transverse, as broad as thorax, ver-
tex bare, ocellar area punctate, cheeks broadened behind eyes, occiput
slightly concave; from in front broader than high, malar space over
half eye, without groove, front finely coriaceous. Antennae filiform,
13-segmented. Pronotum pubescent except medially. Mesocutum
finely coriaceous with scattered hairs from punctures, parapsidal
grooves percurrent, a median notch or short groove behind, lateral and
332 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
anterior lines depressed. Scutellum finely rugose, pubescent, basal
pits smooth, shallow. Carinae on propodeum short, straight, parallel.
Wings hyaline, cilia and pubescence short, veins brown, first abscissa
of radius angled, areolet reaching one-fourth way to basal. Meso-
pleuron largely bare, smooth, polished. Claws toothed. Abdomen
almost globose, tergite II with usual pubescent patches at base, its hind
margin and exposed parts of rest punctate. Ventral spine bare, five
times as long as broad in side view, shorter than hind metatarsus. Us-
ing width of head as a base the length of mesonctum ratio is 1.4; an-
tenna 1.8; wing 3.7; ovipositor 2.4. Length 2.5-3.15 mm. Average
of 30 specimens 2.79 mm.
Types.—U.S.N.M. No. 60119: Type and six paratypes. Paratypes
also in the A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., and M.C.Z.
Host— Quercus subturbinella.
Gall (p. 17, fig. 15).—A bare, smooth, ellipsoidal gall, 5.0 mm. long
by 2.9 mm. in diameter with a nipple at the apex, protruding from a
shallow depression in the side of the acorn cup and dropping to the
ground when mature in late summer, and then showing a girdle of
hairs at the base. From one to eight on an acorn. Monothalmous.
Habitat.—The type locality is Prescott, Ariz. From galls collected
in October 1935 by Mrs. N. W. Capron, adults were cut out on Octo-
ber 10, 1936, and one emerged April 23, 1937. In July and August
1947 she tied bits of cloth over a lot of affected acorns, so that the
galls would not drop to the ground and be lost. Adults were cut out
of these galls on March 20, 1948 and January 7, 1949. More were
bagged in the summer of 1948, when the galls began to drop by the
middle of August, a month earlier than usual, perhaps owing to a dry
season. Many of these were blanks. Adults emerged April 15, 23,
26, 30, 1949.
ANDRICUS PILULA Bassett
This species was described from two females cut from galls from
southern Utah. The types have the claws toothed, not simple as
described. Similar galls were collected by Dr. A. W. Grundmann
on Quercus gambeli in the mountains near Salt Lake City, Utah,
and both females (agreeing with the types) and males were reared
June 1-5, 1949. A description of the male is given below.
Male.—Black, flagellum and legs in part yellowish. Antenna
nearly three times as long as width of the head, 15-segmented, third
longer than fourth and bent, flagellum tapering to tip. Mesoscutum
coriaceous, median groove shorter than in female. Disk of scutellum
finely rugose, foveae smooth. Mesopleuron bare, smooth except for
a few striae in middle. Wing pubescent and ciliate, veins brown,
areolet wanting, cubitus traceable from margin nearly to basal. Claws
toothed. Carinae on propodeum slightly bent. Abdomen shorter
AMERICAN CYNIPID GALL WASPS—-WELD 333
than thorax, lengths of tergites as 15:4. Length 1.7-2.15 mm.
Average of 15 specimens 1.83 mm.
Types.—Specimens of the above description of male are in the
U.S.N.M., A.N.S.P., C.A.F., and the U.U.M.
ANDRICUS PILULARIS, new species
Female.—Related to Andricus pilula Bassett but entirely different
in color, being uniform amber. Antennae 13- instead of 14-segmented.
Mesoscutum microcoriaceous instead of pebbled, anterior lines scarcely
evident, median groove shorter but a dark median line is percurrent,
with uniformly distributed, short, appressed pubescence instead of
almost bare. Disk of scutellum more finely rugose, foveae smooth
instead of with parallel longitudinal ridges. Mesopleuron smooth,
without striae in center. Wing with a small but distinct areolet.
Hind femur stouter. Carinae on propodeum slightly bent and en-
closed area smooth instead of rugese. Abdomen with all the tergites
visible on dorsal curvature, only the tip of sheaths projecting. Ventral
spine longer. Length 2.05-2.85 mm. Average of 12 specimens
2.45 mm.
Male.—Difters from male of piluda in having greatly bulging
eyes, the interocular space higher than broad and cheeks narrowed
behind eyes. Length 1.9-2.75 mm. Average of 8 specimens 2.28 mm.
Type.—vU.S.N.M. No. 60120: Type female, allotype, and two para-
types. Paratypes also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S.,
and M.C.Z.
Host—Quercus subturbinella.
Gall—Similar to galls of Andvicus utriculus Ashmead and A.
pilula Bassett. Globular, about 4 mm. in diameter, projecting more
on upper side of leaf.
Habitat—The types are from galls collected at Wolf Creek, near
Prescott, Ariz., by Mrs. N. W. Capron, in June and July 1935. Living
adults were cut out of the galls on July 20 and August 15. Others
were sent August 1, 1947, and adults cut out August 7. One paratype
is from a gall on Quercus oblongifolia, from Nogales. Galls have
been noted on Q. towmeyi, Q. submollis, Q. gambelii, and Q. diversi-
color.
Genus ANTRON Kinsey
ANTRON PILEUS, new species
Female—Amber. Head coriaceous; from above transverse, nar-
rower than thorax, occiput not concave, cheeks not broadened behind
eyes; from in front broader than high, malar space .28 eye, without
groove, antennae filiform, 15-segmented. Sides of pronotum pubes-
cent. Mesoscutum shining, smooth and almost bare, parapsidal
334 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 102
grooves deep, percurrent. Scutellum longer than broad, an arcuate
groove and two shallow pits at base, disk finely rugose, pubescent, over-
hanging propodeum behind. Mesopleuron smooth, shining. Wing
pubescent and ciliate, without clouds, veins distinct, brown, abscissa
II of radius enlarged at wing margin, areolet reaching one-fifth way
to basal. Claws toothed. Carinae on propodeum strongly bent, en-
closed area smooth, much broader than long medially. Abdomen as
high as long, only 2 or 3 tergites on dorsal margin. Ventral spine
stout, bristly, tapering to tip in side and ventral view. Using width
of the head as a base the length of mesonotum ratio is 1.3; antenna
9.3; wing 3.8; ovipositor 2.5. Length 1.5-2.35 mm. Average of 24
specimens 2.02 mm.
Male—Differs from female in having head and abdomen black and
16-segmented antennae. Length 1.5-1.9 mm. Average of 13 speci-
mens 1.58 mm.
Differs from some other species of Antron in having the wing clear.
Types.—U.S.N.M. No. 60121, type female, allotype, and six para-
types. Paratypes also in the A.M.N.H., C.M.N.H., A.N.S.P., C.A.S.,
and the M.C.Z.
Host.— Quercus subturbinella.
Gall (pl. 17, fig. 19).—A broadly conical bud gall, usually solitary,
at end of new growth in summer. Straw yellow above with a purple
margin, 4-7 mm. in diameter, broader than high, strongly concave
below like the pileus of the fungus Coprinus. Larval cavity at very
apex and exit hole just below the short nipple.
Habitat—The type material was sent by Mrs. N. W. Capron, from
Prescott, Ariz., July 12, 1947, and one adult (the type female) emerged
en route. More galls were sent August 1, 1947, and dead adults were
cut out of the gails. More were sent in early July 1949, and adults
emerged July 10-19. In previous years she had sent galls from Camp
Creek, Cherry, Pine, and Young, and I had collected them at Ash
Fork and Williams, Ariz.
Genus ZOPHEROTERAS Ashmead
ZOPHEROTERAS GUTTATUM, new species
Female.—Living specimens black ; head and thorax fading to brown.
Head coriaceous; from above transverse, occiput concave, cheeks not
broadened behind the eyes; from in front broader than high, inter-
ocular area 1.4 times as broad as high, malar space .5 eye, with a groove,
antennae 14-segmented, filiform, lengths of first four and last seg-
ments as 15 :5 :16:15 and 10. Mesoscutum coriaceous, bare, longer
than broad, not humped in profile, parapsidal grooves distinct, per-
current, widely separated in front, almost meeting behind. Knob
on scutellum coriaceous, almost as wide as deep groove at base, in
AMERICAN CYNIPID GALL WASPS—WELD 335
which are two indistinct smooth pits. Carinae on propodeum almost
straight, widely diverging below. Mesopleuron striate below, coria-
ceous above. Hind tibia longer than tarsus. Claws simple. Abdo-
men as long as head plus thorax, gibbous below petiole, all tergites
visible on dorsal margin, ventral spine 6.5 times as long as broad in
side view, ovipositor straight at tip. Using width of head as a base
the length of mesonotum ratio is .67, antenna 2.7, ovipositor 5.9.
Length 1.5-2.65 mm. Average of 42 specimens 2.2 mm.
Differs from Z. sphaerula Weld in its smaller size, in having mesoscu-
tum uniformly convex in profile, the mesopleuron striate in part and
ovipositor straight at tip.
Types —U.S.N.M. No. 60122: Type and nine paratypes. Paratypes
also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., and the M.C.Z.
Host.— Quercus palustris.
Gall (pl. 16, fig. 12)—Almost spherical, smooth, bare, with purple
spots on a cream background, attached singly on under side of leaf
on a secondary vein, dropping in early October. From 1.4 to 2.5 mm.
in diameter. Pure white inside with the minute larval cavity central.
Habitat.—Type locality East Falls Church, Va. Galls were col-
lected in October in eight different years; adults emerged the second
spring in each case on the following dates: March 6, 20, 25, 29, April
1,3, 7. Buds on pin oak at that time show no signs of swelling. Ovi-
position not observed. Alternating generation unknown.
Genus EUMAYRIA Ashmead
EUMAYRIA INVISA, new species
Female.—Light brown to amber. Head massive, coriaceous, dull;
from above broader than thorax, occiput not concave; from in front
broader than high, malar space one-third eye without groove, antennae
14-segmented, first four as 13:8:20:14. Pronotum coriaceous, dull.
Mesoscutum lighter in color, somewhat shining, not quite smooth,
rather flat, parapsidal grooves percurrent, straight and converging be-
hind but well separated at scutellum. Disk of scutellum margined at
sides, narrowed in front, rugose behind. Wing reduced, not reaching
tip of abdomen, pubescent and ciliate, venation almost normal, areolet
wanting. Mesopleuron with a smooth and polished spot under hind
wing. Claws simple, hind femur short and stout. Carinae on pro-
podeum almost straight, slightly converging above. Abdomen longer
than head plus thorax, tergite II with usual patches of pubescence at
base, rest subequal. Ventral spine longer than hind metatarsus, eight
times as long as broad in side view. Using width of head as a base the
length of mesonotum ratio is 1.0; antenna 2.3; wing 2.0. Length 2.3-
3.2mm. Average of 12 specimens 2.56 mm. No other species known
with reduced wings.
336 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Types.—U.S.N.M. 60123: Type and six paratypes. Paratypes in
A.M.N.H. and C.M.N.H.
Host.—Quercus myrtifolia.
Gall.—Cells under the bark producing, where numerous, a more or
less hypertrophied twig. Similar to gall of Callirhytis crypta
(Ashmead).
Habitat—Type locality Carrabelle, Fla. Living adults were cut
out of the galls on November 17, 1929. Galls collected 2 weeks
previously.
Genus BASSETTIA Ashmead
BASSETTIA FLORIDANA Ashmead
Bassettia floridana ASHMEAD, Trans. Amer. Ent. Soc., vol. 14, p. 147, 1887.
Dryophanta corrugis Bassett, Trans. Amer. Ent. Soc., vol. 17, p. 71, 1890 (new
synonymy).
The types of Dryophanta corrugis in the Bassett collection in the
Academy of Natural Sciences of Philadelphia agree with Ashmead’s
floridana.
Genus DRYOCOSMUS Giraud
DRYOCOSMUS GRUMATUS, new species
Female.—Red, often with black on occiput, anterior and lateral
lines, mesopleura, and dorsal abdomen. Head granulate; from above
transverse, cheeks slightly broadened behind eyes, antennae filiform,
14-segmented. Mesoscutum smooth and polished, parapsidal grooves
deep, smooth, percurrent, median wanting. Scutellum longer than
broad, coriaceous behind the shallow pits, rugose peripherally. Meso-
pleuron almost bare, faintly striate in center. Wing pubescent and
ciliate, veins brown, areolet small, cubitus reaching basal, radial cell
four times as long as broad. Tarsal claws simple. Abdomen in side
view as high as long, not gibbous below petiole, all tergites showing
on dorsal margin. Ventral spine nine times as long as broad, longer
than hind metatarsus. Length 2.15-2.75 mm. Average 2.3mm. De-
scribed from eight specimens, all imperfect.
Types.—U.S.N.M. No. 60124: Type and two paratypes. Another
paratype is in C.A.S.
Host.— Quercus wislizeni.
Gall (pl. 16, fig. 8) —A smooth, bare, ellipsoidal gall about 3 mm.
in diameter attached to midrib on under side of the leaf in the fall
and bearing a little fleshy knob on its summit exactly like the gall of
vileyi (Ashmead) in the eastern United States.
Habitat—Rex Hunt collected a lot of galls in the fall of 1949 at
Felton, Calif., and placed them in a wire breeding cage outdoors on
AMERICAN CYNIPID GALL WASPS—WELD 337
the ground for the winter. When examined on March 15, 1950, many
adults had emerged and died while other galls still contained larvae.
These galls occur also on Quercus agrifolia and Q. kelloggi.
DRYOCOSMUS MINUSCULUS, new species
Female.—Black; mandibles, antennae and legs mostly brownish.
Head coriaceous; from above transverse, cheeks slightly broadened
behind eyes, occiput concave; from in front broader than high, malar
space .4 eye, without groove, antennae 14-segmented, last 8 with rhi-
naria. Sides of pronotum striate in part. Mesoscutum bare, polished,
parapsidal grooves percurrent. Scutellum with two smooth circular
pits at base; disk coriaceous behind septum, rugose behind. Meso-
pleuron bare, smooth. Wing pubescent, ciliate, veins pale, areolet
small, radial cell four times as long as broad. Clawssimple. Carinae
on propodeum slightly bent, enclosed area broader than high. Abdo-
men plump, in side view gibbous below petiole, tergites along dorsal
curvature as 43: 10:10:10:18:1, III-VI sparsely punctate, VII cori-
aceous, tip of ovipositor projecting beyond sheath curved. Ventral
spine eight times as long as broad, slightly longer than hind metatar-
sus. Using width of the head as a base the length of mesonotum ratio
is 1.3; antenna 2.4; wing 5.0. Length 1.75-2.05 mm. Average of five
specimens 1.91 mm.
Differs from Dryocosmus bicornis (McCracken and Egbert), which
is straw yellow and has the ovipositor straight at the tip. D. bicornis
forms galls on the same hosts as does ). minusculus.
Types.—U.S.N.M. No. 60125: Type and one paratype. Paratypes
also in C.A.S., and A.M.N.H.
Host — Quercus agrifolia.
Gall (pl. 17, fig. 18) —Small brown galls up to 2.7 mm. in diameter,
shaped like a depressed sphere with a minute pit in the center above.
Produced on upper side of leaves of Quercus agrifolia, Q. wislizent,
and rarely on Q. kelloggii in the fall and dropping when mature. Said
to be so numerous sometimes as to defoliate the tree. Some years these
galls are not common.
Habitat—The types were reared from galls collected October 22,
1939, on Quercus agrifolia on Mount Diablo, Calif. Adults emerged
April 17, 1941. Galls have been noted on this host at some 40 local-
ities between Alpine and Ukiah and Red Bluff, Calif. They are also
common on Quercus wislizeni and have been seen on Quercus helloggi
afew times. The guest fly, Synergus agrifoliae Ashmead, often reared
from it, varies much in color pattern and has been redescribed as
Synergus maculatus Fullaway (1911) new synonymy, and as Synergus
obscurus McCracken and Egbert (1922), new synonymy.
338 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Genus CALLIRHYTIS Foerster
CALLIRHYTIS CORRUGIS (Bassett)
When I previously reported (Proc. Ent. Soc. Washington, vol. 33,
p. 225, 1931) that this species had been taken in numbers ovipositing
in buds of Quercus maxima, Q. velutina, and Q. palustris at various
dates in April at East Falls Church, Va., the gall from which they had
emerged was not known. It can now be reported that the species has
now been reared from “stone” galls in acorns of all the above oaks—
galls not distinguishable from those of Callirhytis fructuosa Weld.
Acorns of red oak were collected in Loudoun County, Va., on Septem-
ber 22, 1940, and from them fructwosa emerged April 19, 1942, and
corrugis May 4, 1943, and April 20, 1947. Acorns of pin oak were col-
lected at East Falls Church, Va., on October 20, 1940. C. corrugis
(Bassett) emerged April 12, 18, 26, 1942; April 18, 1944; April 14,
1945, and April 6, 1946. Two species thus sometimes make similar
galls in acorns of the same host oak. The alternating gall produced
by these adults which oviposit in buds has not yet been discovered.
CALLIRHYTIS MODESTA (Osten Sacken)
Tn the box of Bassett cynipid types in the American Entomological
Society the type of Cynips papula Bassett, a synonym of modesta, is a
female of Ceroptres sp., a guest fly. A pinned specimen in another
case marked “cotype” is the maker of the gall and has now been made
the type. This is in spite of the fact that the original description of
papula was drawn from specimens of two genera and combined the
head and thorax of the guest with the abdomen and wing of the
maker.
CALLIRHYTIS CISTELLA, new species
Female—Head and thorax amber, dull; abdomen red, shining;
propodeum and tips of antennae infuscated. Head from above trans-
verse, narrower than thorax, cheeks not distinctly broadened behind
eyes; from in front broader than high, malar space .4 eye, slightly
striate; antennae filiform, 13-segmented. Mesoscutum finely and
uniformly rugose, appressed pubescence short, parapsidal grooves ob-
solete anteriorly, median short. Disk of scutellum slightly coarser
posteriorly, not humped back of the narrow septum between two
smooth shining pits. Mesopleuron striate on lower half. Wing
hyaline, surface dotted, nonciliate, veins brown, not clouded, abscissa
I of radius arcuate, areolet wanting. Claws simple. Abdomen as
high as long, gibbous below petiole, only two tergites on dorsal mar-
gin; ventral spine 7 times as long as broad, longer than hind meta-
AMERICAN CYNIPID GALL WASPS-——WELD 339
tarsus. Using width of the head as a base, the length of mesonotum
ratio is 1.4; antenna 2.2; wing 3.7. Length 2.2 mm.
Type.—U.8.N.M. No. 60126: Type.
Host.—Quercus emoryi.
Gall (pl. 17, fig. 22).—A stalked, fusiform, pip gall beside young
acorns July to October. Spindle shaped, brown, 3 by 5 mm. tapering
at both ends, cavity large, wall thin.
Habitat.—The type was reared from a gall collected October 4, 1935,
at Cherry, Ariz., by Mrs. N. W. Capron. A paratype emerged April
14, 1951, from a gall she collected at Prescott, Ariz., on September
20,1950. Galls have been seen at Oracle, Ariz., and in Burro Moun-
tains in New Mexico; seen also on Quercus hypoleuca at Patagonia,
Bisbee, and in Huachuca and Chiricahua Mountains in Arizona.
CALLIRHYTIS HOPKINSI, new species
Agamic female—Amber; foveae, propodeum, middle and hind
tibiae and tarsi infuscated. Head granulate, dull; from above as
broad as thorax, cheeks broadened behind eyes, occiput concave; from
in front broader than high, malar space .6 eye with a groove, antennae
14-segmented, first four as 23(9) :9(7) :28(6) :21; last 17(6). Meso-
scutum granulate, covered uniformly with very short hairs, parapsi-
dal grooves percurrent, a median evident posteriorly. Sculpture of
disk coarser and somewhat transverse, in profile strongly humped back
of septum. Mesopleuron dull, granulate, short-pubescent. Wing
hyaline, dotted, nonciliate, veins beyond second cross-vein clear, first
abscissa of radius angulate, heavy, areolet small. Hind femur broad-
est back of middle. Claws simple. Carinae on propodeum but
slightly bent, enclosed area twice as broad as high. Abdomen length
to height to width as 37: 81: 31: lengths of tergites along dorsal curv-
ature as 95: 26:24:10:12:18. Ventral spine 8 times as long as broad,
shorter than hind metatarsus. Using width of the head as a base the
length of the mesonotum ratio is 1.3; antenna 2.5; wing 3.5; ovipositor
2.4. Length 3.14.2 mm. Average of 154 specimens 3.89 mm.
Differs from agamic Callirhytis operator (Osten Sacken) in having
a shorter ventral spine, 14-segmented antennae and in its lighter
color.
Types.—U:S.N.M. No. 60127: Type and 40 paratypes. Paratypes
also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., and M.C.Z.
Host—Quercus imbricaria.
Gall (pl. 17, fig. 14). —A “pip” gall found in fall on young acorns of
current season, green when fresh and secreting honey dew, becoming
brown later and dropping to the ground. Short-cylindrical, 4-7 mm.
broad by 2-4 mm. high, the larval cell central, the wall thick. Differs
in shape and structure from all other known “pip” galls.
340 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Habitat—The type material was collected at Mineral Wells, W.
Va., by Dr. A. D. Hopkins on September 4 and October 8, 1948. By
September 12 most of the specimens had fallen to the ground, and he
estimated that there were over a bushel of the galls (from 50 to 100
to a square foot) under this one large tree. Adults emerged from an
outdoor breeding cage at Falls Church, Va., on March 28 and April
1, 6, 14, 1945, March 29 and April 5, 1946, and April 12, 18, 1947.
Galls on this host have been seen at Washington, D. C., and Rosslyn,
Va. Alternating generation unknown.
CALLIRHYTIS OBLATA, new species
Female—Bicolored; red with black on anterior and parallel lines,
on foveae, sternum, middle and hind tibiae and ali last tarsal seg-
ments. Head from above transverse, as broad as thorax, occiput
concave; from in front broader than high, cheeks broadened behind
eyes, malar space half eye without groove. Antennae 13- or 14-seg-
mented, first six as 16:10:19:18:15:138. Mesoscutum coriaceous with
scattered punctures and short appressed pubescence. Parapsidal
grooves deep, narrow, smooth, not percurrent, median almost as long
as parapsidal. Scutellum rugose, the two pits at base separated by a
septum. Mesopleuron largely aciculate and bare. Wing hyaline,
pubescence short, nonciliate, veins beyond second cross-vein scarcely
evident. Claws simple. Carinae on propodeum straight, parallel,
enclosed area broader than high. Abdomen with an interrupted ring
of white hairs at base. Lengths of tergites along dorsal curvature
as 87:24:14:11:11. Hind margin of tergite II and exposed parts of
rest punctate. Ventral spine slender, longer than hind tibia. Using
width of the head as a base the length of mesonotum ratio is 1.2;
antenna 2.1; wing 4.2; ovipositor 4.8. Range in length 2.85-3.8 mm.
Average of 18 specimens 3.29 mm. It seems to be related to those
species of the genus that are reared from root galls. Has a much
longer ventral spine than @. fulva Weld.
Types —U.S.N.M. No. 60128: Type and four paratypes. Paratypes
also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., and M.C.Z.
Hosts.—Quercus coccinea and Quercus falcata, Spanish oak.
Gall (pl. 16, fig. 7).—A green, smooth, bare bud gall at apex of new
growth in May, dropping to ground when mature. An oblate sphae-
roid, 4-5 mm. in diameter by 2.5 mm. high, red in center above but
without a nipple, not at all hidden by bud scales; when detached with-
out a girdle of hairs at base.
Habitat—The type is selected from a series that emerged March
21, 1946, from galls collected on Spanish oak at Dyke, 3 miles south
of Alexandria, Va., on April 19, 1945. Paratypes are from galls on
scarlet oak collected near Vienna, Va., and at East Falls Church, Va.
AMERICAN CYNIPID GALL WASPS—WELD 341
CALLIRHYTIS FURVA, new species
Female.—Reddish brown with clypeus, antennae, occiput, anterior
and lateral line areas, propodeum, tibiae, tarsi, and abdomen in part
infuscated ; foveae and mesopleuron black. Head from above trans-
verse, cheeks broadened behind eyes; from in front broader than high,
malar space one-third eye, not striate, antennae filiform, 14-segmented.
Mesoscutum coriaceous, with scattered hairs, parapsidal grooves per-
current. Scutellum disk coriaceous anteriorly, rugose behind, the two
basal foveae sharply outlined. Mesopleuron bare and smooth under
hind wing, striate anteriorly, pubescent below. Wing pubescent and
ciliate, veins brown, areolet small. Claws simple. Carinae on pro-
podeum parallel above, bowed out below, enclosed area broader than
high. Abdomen compressed, longer than head plus thorax, gibbous
below petiole in side view, knifelike back of tergite II, lengths of
tergites along dorsal curvature as 73:20:17:15:10:4; tergite IT
smooth, exposed parts of rest punctate. Ventral spine shorter than
hind metatarsus, tip of ovipositor hooked. Using width of the head as
a base, the length of mesonotum ratio is 1.3; antennae 2.3; wing 2.9;
ovipositor 4.8. Length 2.15-3.65 mm. Average of 46 specimens 2.91
mm.
Differs from Callirhytis infuscata (Ashmead) in being particolored,
larger, in having setigerous punctures between parapsidal grooves and
a longer ovipositor.
Types.—U.S.N.M. No. 60129: Type and seven paratypes. Para-
types also in A.M.N.H., C.M.N.H., A.N.S.P., C.A.S., and M.C.Z.
Host.—Quercus palustris.
Gall (pl. 17, fig. 21).—Small, somewhat globular galls, 3-4 mm. in
diameter covered with short, straight, stiff brown hairs, scattered along
midrib or main veins on upper side of leaf in the fall and dropping off
singly when mature. The hairs do not weather away during winter.
Similar to galls of C. infuscata (Ashmead) on Quercus laevis in
Florida.
Habitat—The types were reared from galls collected on ground
at East Falls Church, Va., on October 12, 1943. Adults emerged
March 20, 24, 1945, and one was cut out alive on January 2, 1946.
Others emerged March 20, 29, 1948, from galls collected September 15,
1946. One was cut out December 1, 1919, from a gall collected at
Ironton, Mo., on October 15,1917. Similar galls have been noted on
nine other species of red oaks.
CALLIRHYTIS SUBCOSTATA, new species
Female —Black; sides of pronotum and legs in part amber. Head
coriaceous; from above transverse, cheeks broadened behind eyes,
342 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
occiput concave; from in front broader than high, malar space one-
third eye, striate, antennae filiform, 14-segmented. Mesoscutum
broader than long, coriaceous, almost bare, parapsidal grooves smooth,
percurrent, median short, anterior lines sunken. Scutellum pubescent,
coriaceous back of the smooth narrow pits, rugose peripherally.
Mesopleuron bare and polished. Wing pubescent and ciliate, veins
brown, areolet reaching one-fifth way to basal, second abscissa of
radius arcuate, radial cell 4 times as long as broad. Tarsal claws
simple. Abdomen in side view as high as long, lengths of tergites as
50:15:6:5:4:3, tergites smooth. Ventral spine slender, 8 times as
long as broad, shorter than hind metatarsus. Using width of the
head as a base, the length of mesonotum ratio is 1.3; antenna 2.1; wing
4.0. Length of four specimens 2.45, 2.25, 2.1 and 2.0 mm.
Types —U.S.N.M. No. 60130: Type and one paratype. Paratype
also in A.M.N.H.
Host.— Quercus stellata.
Gall (pl. 16, fig. 10).—Produced on the side of the acorn cup when
the latter is about 4 mm. in diameter and dropping off when mature
in late May. Green, 2.5-3.0 mm. in diameter, ribbed like a melon
with about 16 grooves, pinkish in the grooves.
Habitat—The types emerged April 1, 1950, from galls collected at
East Falls Church, Va., on May 29, 1949. The nutritive layer was
then about used up and the galls about ready to drop. This isolated
tree had been visited almost every spring for many years but these
galls had never been seen there before.
US, GOVERNMENT PRINTING OFFICE: 1952
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 Washington : 1952 No. 3305
NOTES ON MAMMALS FROM THE NILE DELTA REGION
OF EGYPT
By Henry W. Serzer
From Jury 12, 1946, to April 9, 1947, Ens. P. Quentin Tomich col-
lected mammals from the Nile Delta while serving with Naval Medical
Research Unit No. 3. In addition to the large collection made by
Ensign Tomich, Lt. S. M. Wheeler, of the United States Typhus Com-
mission, obtained some specimens in the Cairo area. Earlier, in 1924,
Miss Annie M. Alexander obtained a small collection from the Cairo
area and from the Fayum. The former two collections are on deposit
in the United States National Museum while the latter is on deposit at
the Museum of Vertebrate Zoology of the University of California
(MVZ) and has been made available for study at this time by Dr.
Alden H. Miller.
Since impetus is being given study of the role wild mammals
play in diseases of man, a brief diagnosis of the external characters
plus comments on taxonomic problems as relating to the commoner
mammals from Egypt, as based on the above-mentioned material,
will be useful to epidemiologists working in the Nile Delta region.
It will be noted that in certain cases names have been validated from
Etienne Geoffroy St.-Hilaire, 1803. The “Catalogue des Mammi-
féres du Muséum National d’Histoire Naturelle” meets all require-
ments for Linnaean names as established by the International Commis-
sion on Zoological Nomenclature. In all instances the descriptions
are clearly recognizable. It is believed that the statement of Isidore
Geoffroy St.-Hilaire, to the effect that his father never intended the
above work for a scientific treatise, should not be accepted, inasmuch as
the work is clear, concise, and was published and circulated.
343
967605—52——1
344 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
The Tomich collection is perhaps the largest and most complete
single collection known from the Nile Delta region. Even so, several
species that have occurred and presumably still occur in that region
were not taken by the collectors on whose work this paper is based.
For the sake of completeness, references to these have been extracted
from the literature.
In addition there are four species which have been taken from the
Delta region within historic time but have since been extirpated.
They are: Wild sheep, Ammotragus lervia; dorcas gazelle, Gazella
dorcas; saberhorned oryx, Orya algazel; and addax, Addaaw naso-
maculata.
Capitalized color terms are from Ridgway, “Color Standards and
Color Nomenclature” (1912). All measurements are given in milli-
meters and all weights in grams.
No comments have been made in regard to habitats or habitat pref-
erence, since Tomich hopes to publish an account of the natural
history of these animals and their role in epidemiology.
Order INSECTIVORA
Family ERINACEIDAE
HEMIECHINUS AEGYPTIUS (E. Geoffroy St.-Hilaire)
Hrinaceus aegyptius EH. Greorrroy St.-Hrtarre, Catalogue des mammiféres du
Muséum National d’Histoire Naturelle, p. 69, 1803 (Egypt).
Erinaceus libycus HEMPRICH and HHRENBERG, Symbolae physicae ..., dec. 2,
footnote, folio k, 1832 (near Alexandria, Egypt).
Specimen examined —One. Giza Province: Abu Rauwash, Cairo
area (MVZ).
Description—Upper parts, except head and shoulders, covered with
short (15 to 20 mm.) spines; base of spines (38 to 5 mm.) brownish,
then a buffy band, a brown and a terminal white band of about equal
width; hairs of entire head, ears, belly, fore and hind legs Warm Buff.
Tail short; ears large; eyes small; fore feet broad and short; hind feet
relatively long and narrow.
Measurements—An adult male from Abu Rauwash, Cairo area,
measured : Total length 200; length of tail 20; length of hind foot 35;
condylobasal length 45.8; length of palate 24.2; width of rostrum
at lacrimal canal 10.4; length of nasals 17.0; least interorbital width
11.0; width across zygomatic arches 26.4.
Remarks.—No ear measurements are available for the one specimen
examined, but from the appearance of the dry skin the ears appear
as though they would, when laid forward in a fresh specimen, reach
to the end of the nose.
MAMMALS FROM THE NILE DELTA REGION, EGYPT—SETZER 345
Family SORICIDAE
CROCIDURA OLIVIERI (Lesson)
Sorex olivieri Lesson, Manuel de mammalogie . . ., p. 121, 1827 (Saqqara Pyra-
mids, Giza Province, Egypt, as a mummy).
Specimen examined.—One. Giza Province: Talbia.
Description—Entire upper parts Prout’s Brown, shading rather
rapidly on the lower sides to dark gray, which covers the entire
ventral surface with the exception of a small patch of brownish hairs
on the throat; tail of uniform color around and finely covered with
long whitish hairs; dorsal surfaces of hands and feet grayish, shad-
ing to white on the digits; palms and soles naked. Lars large, nearly
naked and with a pronounced fold on the ventral surface; tail more
than half the length of head and body. Scent gland, lying immedi-
ately anterior to flank and between the dark dorsal and light ventral
colors, conspicuous and covered with short whitish hairs.
Measurements.—No external measurements were available but the
skull of an adult male from Talbia, Giza Province, measured : Greatest
length 29.0; occipitonasal length 26.4; greatest width of braincase
12.5; least interorbital width 5.5; length of palate 12.0; width across
M? 9.3; length of tooth row (incisor to M*) 12.8; width of rostrum 3.6.
CROCIDURA FLOWERI Dollman
This species is known from Giza and Beltim. It is apparently rare
and hard to obtain. It differs from Crocidura oliviert in smaller
size (head and body 57-71 mm. as opposed to 93-110 mm. in @,
olivier). No specimens have been examined.
CROCIDURA RELIGIOSA I. Geoffroy St.-Hilaire
This is the smallest Crocidura to be found in the Nile Delta. It
apparently is fairly common in the Giza Gardens and in the country
near Abu Rauwash. The head and body range from 45 to 55 mm.
in length, thus separating this species from the other, larger, species
of Crocidura known to occur in the same general area. No specimens
have been examined.
SUNCUS CRASSICAUDUS Lichtenstein
Despite efforts of several collectors to retake this species in Suez
and Suakin, Sudan, it has not been found. It is thought that it is
probably the same animal as the Swneus found along the coastal re-
gion of India and that the individuals known from North Africa
are merely fortuitous travelers come ashore from some trading ves-
sel. It may be distinguished, if ever retaken, from specimens of the
346 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
larger species of Crocidura by the presence of two more teeth in the
jaws (total of 28 in Crocidura and 30 in Suncus). No specimens
have been examined.
Order CHIROPTERA
Family PTEROPIDAE
ROUSETTUS EGYPTIACUS (E. Geoffroy St.-Hilaire)
Pteropus egyptiacus EB. Grorrroy ST.-HiLarrE, Ann. Mus. Nat. Hist. Natur. Paris,
vol. 15, p. 96, 1810; spelling of specific name corrected or emended to
aegyptiacus in Description de l’Egypte. Histoire naturelle, .. . mammiféres
. « Vol. 2, p. 134, pl. 3, fig. 2, 1818 (Great Pyramid of Giza).
Specimens examined—Seven. Giza Province: Gezira Island,
Cairo.
Description.—Upper parts between Drab and Hair Brown, shading
to between Light Drab and Drab on the underparts; all hairs uni-
form in color to base; membranes of wings and uropatagium, in dry
specimens, blackish; feet, uropatagium and forearm for half its
length, sparsely furred; uropatagium reduced to a semicircular mem-
brane, extending but slightly beyond the rump; tail short, barely ex-
ceeding the narrow rim of the uropatagium; hind feet large, strongly
clawed; thumb long; index finger with small claw; toes of hind feet
with short bristlelike hairs; ears large, somewhat pointed; tragus
absent.
Measurements —Averages and extremes of two males and five fe-
males from Gezira Island, Cairo, were, respectively: Total length 152
(148-154), 145.4 (134-162) ; length of tail 14 (18-15), 13.4 (12-14) ;
length of forearm 91.5 (88-95), 87.8 (85-95); length of hind foot
27.5 (26-29), 27.5 (27-28); length of ear from notch 23 (23), 22.2
(22-23) ; greatest length of skull 42.7 (41.8-43.6), 40.96 (39.2-42.8) ;
basal length 41.1 (40.2-41.9), 39.2 (37.2-41.5) ; occipitonasal length
41.0 (40.0-42.0), 39.2 (37.8-41.2) ; greatest zygomatic breadth 25.85
(24.6-27.1), 24.4 (22.8-26.2); breadth of braincase 16.75 (16.5-17.0),
16.7 (16.3-17.1); canine to M® 16.8 (16.4-17.2), 16.1 (15.4-16.9) ;
weight of two females 125.0 (109-141).
Remarks.—Anderson (1902) regards Vespertilio aegyptiacus Lin-
naeus, in Hasselquist, 1757, as being the first name to be applied to
these bats. This is erroneous since the description in 1757 is not ten-
able but does state that V. aegyptiacus has a tragus, that it is of small
domestic mouse size and that the tail is the same length as the body.
The name, if it were tenable, would have to be used for Rhinopoma
microphyllum.
MAMMALS FROM THE NILE DELTA REGION, EGYPT—SETZER 347
Family RHINOPOMATIDAE
RHINOPOMA MICROPHYLLUM (Brunnich)
Vespertilio microphyllus Brunnicu, Dyrenes historie og dyre-samlingen udi
universitetets natur-theater, vol. 1, p. 50, pl. 6, figs. 14, 1782 (Arabia and
Egypt).
Specimens examined.—Twelve. Giza Province: Cairo area, Saqqara
Pyramids, 11; Pyramids of Giza, 1.
Description—Upper parts Mouse Gray shading to whitish below;
membranes of wing brownish; ears light brownish, nearly naked;
face but sparsely covered with fine grayish hairs; tragus free of pinna
and spatulate. Tail as long as or longer than head and body, three-
fourths of which is completely free of the uropatagium; ears joined
by a ridge across the forehead; hind legs long.
Measurements.—Averages and extremes of seven males and four
females from Saqqara Pyramids, Cairo area, were, respectively : Total
length 139 (127-145), 141 (139-143); length of tail, 59.8 (54-67),
60.5 (58-63) ; length of forearm 68.4 (66-71), 68 (67-69) ; length of
hind foot 16.5 (16-17), 16.2 (16-17) ; length of ear from notch 20.2
(19-22), 20.2 (19-21) ; greatest length of skull 20.6 (20.2-21.3), 20.2
(19.8-20.3) ; condylobasal length 18.6 (17.7-19.5), 18.1 (17.8-18.4) ;
occipitonasal length 17.6 (16.9-18.4), 17.2 (16.7-17.3) ; greatest zygo-
matic width 12.1 (11.5-12.6), 11.95 (11.8-12.1); width of braincase
8.5 (8.2-8.8), 8.5 (8.4-8.6) ; canine to M® 7.7 (7.3-8.0), 7.5 (7.4-7.6) ;
weight 19 (14-25), 17.5 (17-19).
Remarks.—As may be noted from the measurements, there is a
considerable amount of variation in size of these bats. On the basis
of size, Thomas described Rhinopoma cystops from Luxor, Egypt.
This species may be distinguished from the smaller 2. cystops by the
longer forearm, darker coloration, and larger skull in all measurements
taken.
RHINOPOMA CYSTOPS Thomas
Rhinopoma cystops THomMas, Ann. Mag. Nat. Hist., ser. 7, vol. 11, p. 496, May
1903 (Luxor, Egypt. Type: Adult female, B. M. No. 2.1.17.2).
Specimens examined.—Sixteen. Giza Province: 8 miles southwest
of Cairo, at the Pyramids, 12; Pyramids of Giza, 4.
Description—Upper parts Light Mouse Gray shading to whitish
below; membranes of wing brownish; ears light brownish, nearly
naked; face but sparsely covered with fine grayish hairs; tragus free
of pinna and spatulate. Tail as long as or longer than head and body,
three-fourths of which is completely free of the uropatagium; ears
joined by a narrow ridge across the forehead; hind legs long.
348 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Measurements —Averages and extremes of nine males and three fe-
males from 8 miles southwest of Cairo, at the Pyramids, were, re-
spectively : Total length 135 (128-143), 131 (125-188) ; length of tail
68.0 (64.0-73.0), 65.0 (61.0-70.0) ; length of forearm 57.5 (55.0- 60.0),
55.3 (53.0-57.0) ; length of hind foot 13.3 (18.0-14.0), 13.8 (13.0-14.0) ;
length of ear from notch 19.0 (18.0-20.0), 19.8 (18.0-21.0) ; greatest
length of skull 18.1 (17.9-18.4), 17.6 (17.2-17.9) ; condylobasal length
16.1 (15.9-16.2), 15.6 (15.5-15.7) ; occipitonasal length 14.9 (14.7-
15.1), 14.4 (14.2-14.6) ; greatest zygomatic width 10.4 (10.2-10.7),
10.86 (10.8-10.4) ; width of braincase 7.6 (7.4-7.7), 7.3 (7.8) 5 canine
to M* 6.3 (6.2-6.4), 6.0 (5.9-6.1).
Remarks.—The specimens listed above vary in some detail from the
measurements given by Thomas in his description of the species.
These measurements of the Cairo specimens are all somewhat larger
but are definitely smaller than those given for the larger species
Rhinopomamicrophyllum. Thanks to C. C. Sanborn, curator of mam-
mals, Chicago Natural History Museum, measurements of specimens
of &. cystops from various localities throughout the Middle East have
been made available to me. A topotypical series of 17 specimens of
FR. cystops are only slightly smaller than the measurements given for
the Cairo series above. It is apparent that Flower (1932, p. 385) was
in actuality comparing specimens of P. cystops from the type locality
and from the Cairo area. He thus arrived at the conclusion that #.
cystops was conspecific with R. microphyllum. That this conspecific-
ity is not real is shown by a comparison of the measurements listed
for the two species in the present paper.
The two species were found together, by Tomich, at the Pyramids
of Giza. Among these specimens the larger, darker 2. microphyllum
is easily differentiated.
Family EMBALLONURIDAE
TAPHOZOUS PERFORATUS E. Geoffroy St.-Hilaire
Taphozous perforatus ©. Grorrroy St.-Hilaire, Description de ’Figypte. Histoire
naturelle, ...mammiféres .. ., vol. 2, p. 126, 1818 (Egypt).
Specimens examined.—Fifteen. Giza Province: north base of
Mount Abu Rauwash.
Description—Upper parts Hair Brown shading to Drab on the
belly and throat; all hairs white at base; face and chin but sparsely
haired; wing membranes and uropatagium brownish; radio-ulna
lightly furred for one-fourth its length with whitish hairs; tail, for
about half its length, free of uropatagium and tipped with a few
grayish hairs; ears small and haired at base, rest naked; tragus free
MAMMALS FROM THE NILE DELTA REGION, EGYPT—SETZER 349
from pinna, small and papillate in shape; calear about three-fourths
the length of the tibio-fibula.
Measurements—Averages and extremes of six males and nine fe-
males from the north base of Mt. Abu Rauwash were, respectively :
Total length 101 (99-102), 102.3 (101-106); length of tail 22.6
(22-23), 23.4 (22-26); length of forearm 64 (63-65), 64 (63-66) ;
length of hind foot 14 (14), 13.6 (18-15) ; length of ear from notch
17.3 (17-18), 17.5 (17-18) ; greatest length of skull 20.3 (20.1-20.6),
20.1 (19.8-20.3) ; basal length 19.4 (19.0-19.6), 19.2 (18.8-19.6) ; oc-
cipitonasal length 17.3 (17.2-17.3), 17.1 (16.9-17.4) ; greatest zygo-
matic breadth 11.7 (11.6-11.8), 11.6 (11.4-11.7) ; breadth of braincase
9.5 (9.3-9.7), 9.85 (9.0-9.5) ; canine to M® 8.5 (8.5), 8.5 (8.48.6) ;
weight 24.0 (20-27), 24.2 (16-28).
Remarks —¥F rom the other bats available at this time from the
Cairo area, this species may be distinguished by its rather long,
pointed ears, the peculiar tuft of hair on the rather short, free tail,
by the white basal portion of the hairs, and by the long calcar which
supports the posterior edge of the uropatagium.
LIPONYCTERIS NUDIVENTRIS (Cretzschmar)
Taphozous nudiventris CRETZSCHMAR, in Riippell, Atlas zu der Reise im nérdlichen
Afrika, p. 70, pl. 27b, 1828 (Egypt).
Specimens examined.—Five. Giza Province: Abu Rauwash
(MVZ).
Description.—Upper parts Hair Brown shading into Cinnamon-
Drab on the lower parts; wings, uropatagium, feet and ears, blackish ;
muzzle, chin, rump, under surfaces of femora, uropatagium, ears and
wing membranes, naked, with the exception of a narrow strip of
whitish hairs running the entire length of the membrane on the under-
side of the forearm. Tragus large and truncate; tail about one-fourth
of total length, free from uropatagium for about one-half its length,
sparsely covered with long whitish hairs.
Measurements.—Averages and extremes of three males from Abu
Rauwash were: Total length 124 (121-126) ; length of tail 80 (28-32) ;
length of hind foot 18 (18) ; greatest length of skull 28.2 (26.8-29.6) ;
basal length 21.6 (21.4-21.8) ; occipitonasal length 24.4 (24.0-24.8) ;
greatest zygomatic breadth 16.2 (15.8-16.7) ; breadth of braincase 11.2
(11.0-11.3) ; length of tooth row from M® to canine 11.3 (11.0-11.5).
Remarks.—This bat may be told from a close relative living in the
same area, Taphozous perforatus, by the bare rump, the bare under-
side of the femora and the much larger skull.
It is interesting that in 1924 Miss Alexander obtained only Lipony-
cteris nudiventris at Abu Rauwash and in 1946 Tomich obtained only
the closely allied Taphozous perforatus.
350 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Family RHINOLOPHIDAE
RHINOLOPHUS ACROTIS Heuglin
Rhinolophus acrotis HreuGcuin, Noy. Act. Acad. Caesareae Leopoldino Carolinae,
Halle, vol. 29, pp. 4, 10, December 1861 (Keren, Eritrea, about lat. 15°45’
N., long. 88°30’ E. Type: Adult male, labeled No. 986, Stuttgart Museum).
Specimens ewamined—Four. Giza Province: Pyramids of Giza.
Description—Upper parts Drab, underparts and base of hairs of
upper parts Pale Drab-Gray; membranes of wing and uropatagium
blackish brown; ears moderately large, rather acutely pointed, naked
except at base and basal half of medial margin, tragus but a small
rounded projection; noseleaf large, horseshoe shaped, dorsal projec-
tion free and rather small; face and muzzle fully haired; wings and
uropatagium naked; tail, enclosed in uropatagium for its entire
length, about half the length of the head and body.
Measurements.—Average and extremes of three males from the
Pyramids of Giza were: Total length 85.0 (84-87) ; length of tail 28
(24-33) ; length of forearm 45.6 (45-47) ; length of hind foot 10.3
(9-11) ; length of ear from notch 19.3 (19-20); greatest length of
skull 19.6 (19.5-19.7) ; basal length 15.6 (15.4-15.8) ; occipitonasal
length 14.23 (14.2-14.3) ; zygomatic breadth 9.7 (9.6-9.8) ; breadth of
braincase 8.7 (8.5-8.9) ; canine to M? 7.0 (7.0); weight of two indi-
viduals 8.0 (8.0).
Remarks.—The two species of Rhinolophus supposedly occurring in
the Cairo area are represented here by only Rhinolophus acrotis. This
species is apparently much the same in size and dorsal coloration as
Rhinolophus euryale. The two species can be distinguished by the
light belly and rounded dorsal projection of the basioccipital in &.
acrotis and the dark belly and pointed basioccipital in F. euryale.
Family HIPPOSIDERIDAE
ASELLIA TRIDENS (E. Geoffroy St.-Hilaire)
Rhinolophus tridens ®. Grorrroy St. Hi~arre, Ann. Mus. Nat. Hist. Natur., Paris, ‘
vol. 20, p. 265, 1813 (Egypt).
Specimens examined—Fifty. Lisht, Upper Egypt, 32 (alcoholic).
El Maabdeh Cave, 1 (alcoholic). Oasis of Kharga, 12 (alcoholic).
Henneh, 1. Nile, 4 (alcoholic).
Description.—Since only one conventional skin and skull combina-
tion is available, and that old and faded, an exact color diagnosis is
not possible. Based on the one specimen, the color appears to
have been light brown above and paler below; wing membranes and
uropatagium dark brown. Nose with prominent “horseshoe” with
trilobed leaf on dorsal surface; ears large, connected by narrow band
MAMMALS FROM THE NILE DELTA REGION, EGYPT—-SETZER 35]
across top of head; no tragus appears on either dry or alcoholic speci-
mens; tail for three-fourths of its length enclosed in uropatagium,
other one-fourth free.
Measurements.—No skin measurements were available, but average
and extreme skull measurements from five females from Lisht, Upper
Egypt, were: Greatest length 18.0 (17.8-18.2); basal length 16.1
(15.6-16.6) ; occipitonasal length 14.3 (13.6-14.5) ; zygomatic breadth
10.0 (9.9-10.1) ; width of braincase 8.35 (8.38-8.5) ; canine to M?* 6.5
6.5).
Pee pee this bat is not to be found in the lower Nile
region, since all records thus far have come from no farther north than
Saqqara, Giza Province.
Family VESPERTILIONIDAE
PIPISTRELLUS KUHLII (Natterer)
Vespertilio kuhlii NATTERER, in Kuhl, Die Deutschen Fledermiiuse. Ann. Wetterau.
Gesel. Hanau, vol. 4, p. 199, 1817 (Trieste).
Specimens examined.—Seven. Giza Province: 5 miles west of
Simbillawein, 6; Maadi, 1.
Description.—Hairs of upper parts tipped with Buffy Brown, black
at bases; underparts lighter than back; wing membranes, in dry speci-
mens, dark brown, uropatagium lighter and strongly veined; posterior
margins of wings and uropatagium bordered with white; ears small,
brownish and naked; tragus large; base of uropatagium furred;
calcar keeled.
Measurements.—Averages and extremes of five females from 5 miles
west of Simbillawein and measurements of one male from the same
locality are respectively: Total length, 87.2 (84-92), 89; length of tail
35.8 (382-87), 87; length of forearm 35.5 (35-86), 35; length of hind
foot 8.5 (8-9), 8; length of ear from notch 12 (12), 12; greatest length
of skull, 13.4 (13.2-13.6), 13.4; basal length 12.9 (12.7-18.2), 13.0;
occipitonasal length 11.2 (11.1-11.8), 11.2; zygomatic breadth 8.4 (8.3-
8.6), 8.5; breadth of braincase 6.6 (6.5-6.7) , 6.6; canine to M® 4.9 (4.9),
4.9; weight 4.25 (4.0-5.0), 5.0.
Remarks.—The series of bats available for study are remarkably
uniform in color, in the whitish margins of the wings and uropatagia,
as well as in the measurements of the skull. In one character of the
skull, however, there is some variation that is rather peculiar. The
first premolar on one side of the upper jaw is missing in two specimens.
The tooth is extremely reduced in size in any event and even under high
magnification no alveolus can be ascertained where the tooth is sup-
posed to be in those specimens where the tooth is missing.
These two specimens present a condition that appears to tend toward
the reduced dentition of Scotozous.
967605—52——_2
302 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 109
EPTESICUS INNESI Lataste
To date, there are two specimens of this bat known. They are the
pair, on which the original description was based, taken by Dr. Walter
Innes from a house in Cairo about 1885. Since that time no additional
specimens have been recorded anywhere else in Egypt, although sought
after. This bat may be distinguished from Pipistrellus kuhli by the
nearly naked face and the lack of white edging on the wings and uro-
patagium. No specimens have been examined.
PLECOTUS AURITUS Linnaeus
This bat may be distinguished from other species of bats occurring
in the Nile Delta region by means of its large ears. The ears are
almost as long as the head and body (head and body 45 mm., ear 35
mm.). It has been recorded from Mena House, Second Pyramid, and
Minia. No specimens have been examined.
Family MOLOSSIDAE
NYCTINOMUS TENIOTIS (Rafinesque)
Cephalotes teniotis RAFINESQUE, Precis des découvertes . . . Somiologiques. .
p. 12, 1814 (Sicily).
Specimen examined—One. Giza Province: near Cairo (alco-
holic).
Remarks —There was available to me only one specimen, preserved
in alcohol. Any color diagnosis would, therefore, be worthless. The
ears are large and narrowly joined across the forehead. Approxi-
mately half the tail is free of the uropatagium. Nyctinomus teniotis
is considerably larger than Vyctinomus aegyptiacus, at least as far as
the skull is concerned, from the specimens available. In addition,
N. aegyptiacus has but two lower incisors on each side of the lower jaw,
while J. tenzotis has three.
It appears that there has been a great deal of doubt as to the names ap-
plied and as to what the respective authors were describing. Geoffroy
St.-Hilaire (1818, p. 129), in deseribing WV. aegyptiacus, says in regard
to the teeth, “Les dents deviennent un excellent indicateur de cette
organisation ; les incisives sont au nombre de deux en haut, et de quatre
en bas: celles-la sont fortes, coniques et contigues, quand les secondes
sont tres-petites et comme entassees au devant des canines.” It thus
appears that the plate and the published description are at variance,
since he states in the latter that there are four incisors below and the
plate appears to show six. Is it possible that the artist interpreted
the cingulum of the canine as an incisor?’ In De Winton’s review of
Nyctinomus he states there are four incisors in V. aegyptiacus and six
in J. tentotis, yet he raises a question as to what JV. aegyptiacus actually
is in the published work of Anderson on the mammals of Egypt.
MAMMALS FROM THE NILE DELTA REGION, EGYPT—SETZER 353
From specimens of identified V. aegyptiacus from Basutoland, X.
teniotis is at once distinguished by its larger size and the occurrence
of three instead of two incisors in each mandible.
Order LAGOMORPHA
Family LEPORIDAE
LEPUS AEGYPTIUS Desmarest
This is the largest of the hares to be found in Egypt. It has long,
thin hind legs; the hind feet are long and narrow; the ears are about
134 times the length of the head; and the tail is black bordered with
brown. It is known to occur in Giza and Fayum Provinces. No speci-
mens have been examined.
LEPUS ROTHSCHILDI De Winton
A medium-sized hare, characterized by short, stout hind legs; broad,
short hind feet; ears about 114 times the length of the head; and the
tail black, edged with white. On these characters it can be distin-
guished from L. aegyptius. It also is known to occur in Giza and
Fayum Provinces.
Order RODENTIA
Family CRICETIDAE
GERBILLUS GERBILLUS ANDERSONI De Winton
Gerbillus andersoni Dp WINTON, Ann. Mag. Nat. Hist., ser. 7, vol. 9, p. 45-46
[p. 45], January 1902 (Mandara, east of Alexandria, B.M. No. 92.7.1.5).
Specimens examined.—Kight. Behera Province: one-half mile
west of Dikheila Airfield, 4; one-half mile east of Muntazah, 4.
Description.—Upper parts between Ochraceous-Buff and Ochrace-
ous Orange; hairs plumbeous at base with a subterminal band of the
above color and a terminal band, very minute, of black. Black hairs
freely intermixed with banded hairs, thus presenting a somewhat
subdued hue. Color purest on sides, flanks, cheeks, rump, and nose.
Entire underparts, muzzle, most of the vibrissae, hind and forelegs,
dorsal and ventral surfaces of hands and feet, supraorbital and
postauricular spots, white. Tail penicillate, ochraceous above shad-
ing to yellowish white beneath; tip on dorsal surface with longish
brown hairs. Ears blackish, sparsely covered with fine white hairs
inside and ochraceous on the outside. Eye bordered by narrow black
ring.
354 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Measurements.—Averages and extremes of two males and two fe-
males from one-half mile east of Muntazah were, respectively: Total
length 222 (219-224), 215 (207-223); length of tail 122 (120-125),
119 (113-125) ; length of hind foot 30 (30), 28 (27-29) ; length of ear
from notch 15.5 (15-16), 15.5 (15-16); occipitonasal length 29.1
(28.9-29.3), 28.2 (28.2); length of palate 12.95 (12.8-13.1), 12.5
(12.3-12.7) ; length of palatal foramina 5.5 (5.4-5.6), 5.2 (5.1-5.3) ;
length of auditory bulla 8.85 (8.8-8.9), 8.75 (8.7-8.8) ; crown length of
upper molar series 3.85 (3.8-8.9), 3.85 (3.8-3.9); least interorbital
width 5.9 (5.8-6.0), 5.75 (5.7-5.8).
Remarks.—Gerbillus gerbillus andersoni may be distinguished from
what is here considered the nominate race from 5 miles northwest of
the Pyramids of Giza by darker dorsal color and longer more pig-
mented ear. Cranially it may be distinguished by longer rostrum,
larger teeth, more inflated auditory bullae, longer, narrower braincase,
longer incisive foramina, and more robust zygomata.
GERBILLUS GERBILLUS GERBILLUS (Olivier)
Dipus gerbiilus OxrvieR, Bull. Sci. Soc. Philom. Paris, vol. 2, p. 121, 1801 (Giza
Province, Egypt).
Specimens examined.—Thirty-seven. Giza Province: Abu Rau-
wash, 1 (MVZ) ; 5 miles east of Helipolis, 4; 8 miles east of Helipolis,
8; 5 miles northwest of Pyramids of Giza, 10; Saqqara,2. Suez Canal
Zone: 3 miles northwest of Fayid, 12.
Description—Upper parts Pinkish Cinnamon, hairs plumbeous at
base, subterminal band of above color, minute terminal band of some
hairs blackish. Color purest on sides, flanks, cheeks, and nose; entire
underparts, muzzle, most of the vibrissae, hind and fore legs, dorsal
and ventral surfaces of hind and fore feet, supraorbital and postauric-
ular spots, white. Ears flesh-colored, nearly naked. Eye bordered
by narrow black ring.
Measurements—Averages and extremes of four males and six fe-
males from five miles northwest of the Pyramids of Giza were: Total
length 222 (216-227), 211 (203-217) ; length of tail 127 (123-181),
120 (113-128) ; length of hind foot 30 (29-81), 28.5 (28-29) ; length
of ear from notch 12 (12), 12 (12); occipitonasal length 27.3
(27.2-27.4), 26.6 (26.2-26.8) ; length of palate 12.2 (12.1-12.5), 12.1
(11.7-12.5) ; length of palatal foramina 4.5 (4.3-4.6), 4.4 (4.8-4.6) ;
length of auditory bulla 8.7 (8.5-8.8), 8.4 (8.3-8.8) ; crown length of
upper molar series 3.45 (3.4-3.5), 3.5 (3.4-8.6) ; least interorbital width
5.85 (5.8-5.9), 5.7 (5.4-6.0) ; weight 21 (19-22), 19 (18-21).
Remarks.—This small gerbil, represented by 37 specimens, is re-
markably constant in measurement and color. In external measure-
ments it is about the same as Gerbillus gerbillus andersoni. The two
MAMMALS FROM THE NILE DELTA REGION, EGYPT—SETZER 355
subspecies can, however, be readily distinguished by their coloration
and the longer darker ear and by the noticeably larger skull of
G. g. gerbillus.
GERBILLUS HENLEYI MARIAE (Bonhote)
Dipodillus mariae Bonnore, Proce. Zool. Soc. London, 1909, p. 792, April 8, 1910
(Mokkattam Hills, near Cairo, Egypt).
~
Specimen examined.—One. Giza Province: 5 miles east of
Heliopolis.
Description—Upper parts Pinkish Cinnamon, but little pure color
on sides, all pigmented areas strongly suffused with black; hairs of
upper parts plumbeous at base; entire underparts, muzzle, fore and
hind legs, upper surface of feet, supraorbital and postauricular
spots, and narrow ring at base of tail, white; eye and pinnae of ears
bordered by narrow black line; ears light brownish, finely covered with
whitish hairs; tail long, bicolored (same as back on dorsal surface
proximally, grading to blackish distally), penicillate; soles of hands
and feet naked.
Measurements.—A male from 5 miles east of Heliopolis measured:
Total length 172; length of tail 98; length of hind foot 21; length of
ear from notch 9; occipitonasal length 21.3 ; length of palate 9.0; length
of palatal foramina 3.8; length of auditory bulla 7.5; crown length
of upper molar series 2.7; least interorbital width 3.8; weight 10.
GERBILLUS NANUS GARAMANTIS Lataste
Gerbillus garamantis LATASTE, Le Naturaliste, 1881, p. 507 (Sidi-Roueld, Ouargla,
Algeria. Female, B. M. No. 19.7.7.1596, and its skull, B. M. No. 19.7.7.1597).
Specimens examined.—Three. Giza Province: 1 mile northeast of
Kom Aushim.
Description—Upper parts Clay Color, purest on sides and cheeks;
with a strong admixture of black on the back; hairs plumbeous at base
in all pigmented hairs; underparts, forelimbs, dorsal surfaces of fore
and hind feet, ventral half of tail, supraorbital and postauricular spots
and muzzle, white. Ears brownish, nearly naked, bordered by thin
blackish line; dorsal surface of nose, blackish ; eye bordered by narrow
black line; dorsal surface of tail as back color but with more admixture
of black, which increases toward the black penicillate tip. Soles of
hands and feet naked.
Measurements.—Averages and extremes of three males from 1 mile
northeast of Kom Aushim, were: Total length 195.3 (191-202) ; length
of tail 110 (106-115) ; length of hind foot 23 (22-24); length of ear
from notch 12.3 (12-13) ; occipitonasal length 25.0 (24.2-25.8) ; length
of palate 11.2 (10.6-11.7) ; length of palatal foramina 4.5 (4.34.7) ;
length of auditory bulla 7.9 (7.7—-8.2) ; crown length of upper molar
356 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
series 3.43 (8.4-3.5) ; least interorbital width 4.6 (4.5-4.9); weight
19 (16-22).
Remarks.—It seems hardly logical that this gerbil, in view of the
apparent plasticity of the genus, could be the same subspecies in the
Nile Valley and in Algeria. However, owing to the lack of compara-
tive material, it is felt best to place the above specimens in this
subspecies.
GERBILLUS PYRAMIDUM PYRAMIDUM (E. Geoffroy St.-Hilaire)
Dipus pyramidum BH. GEOFFROY ST.-HILAIRE, Catalogue des mammiféres du
Muséum National d’Histoire Naturelle, p. 202, 1803 (near Pyramids of Giza,
Giza Province, Egypt).
Specimens examined.—Twenty-three. Fayum Province: 1 mile
northeast of Kom Aushim, 4; Sennuris, 3 (MVZ); The Fayum, 2
(MVZ). Giza Province: Abassia Fever Hospital, Cairo, 2; 3 miles
northeast of Mount Abu Rauwash, 5; 1 mile northeast of Mount Abu
Rauwash, 3. Nubia: Abu Simbel, 1 (MVZ) ; Amanda, near El Derr,
3 (MVZ).
Description—Upper parts Pinkish-Cinnamon, purest on sides,
flanks, cheeks and muzzle; hairs of the dorsum plumbeous at base,
finely tipped with black, remainder of hairs of body white at base;
entire underparts, dorsal and ventral surfaces of fore and hind feet,
supraorbital and postauricular spots, lower half of cheeks and muzzle,
and posterior ventral half of tail, pure white; tail above, same pure
color as on sides, shading in the posterior one-fourth to a penicillate
black tip; ears finely covered with hair, margins of pinnae blackish;
narrow black ring around eye. Tail not quite so long as head and
body; hind feet relatively short.
Measurements —Two males from 1 mile northeast of Abu Rauwash
and two females from 3 miles northeast of Mt. Abu Rauwash meas-
ured, respectively: Total length 232, 255; 229, 252; length of tail 128,
135; 129, 189; length of hind foot 34, 35; 33, 83; length of ear from
notch 15, 15; 15, 15; occipitonasal length 31.5, 33.0; 29.9, 82.1; length
ef palate 14.5, 15.0; 13.8, 14.5; length of palatal foramina 5.9, 6.0;
5.8, 6.1; length of auditory bulla 9.0, 9.7; 9.1, 10.0; crown length of
upper molar series 4.3, 4.3; 4.2, 4.2; least interorbital width 6.6, 7.2;
6.5, 6.8; weight 37, 56; 32, 42.
Remarks.—In the series of specimens available at this time are
what appear to be two color phases. However, when these animals
are examined critically, these “phases” are seen to be juvenile and adult
pelages. The juvenile pelage is darker than that of the adult and
appears to be retained longer than in most species of rodents. With
respect to pelage these darker animals appear to fit the description
of Gerbillus pyramidum tarabuli, and it is possible that this subspecies
is based on immature specimens of @. p. pyramidum. However, until
MAMMALS FROM THE NILE DELTA REGION, EGYPT—SETZER 357
such time as a concerted effort is made to obtain specimens and to
study this genus throughout its range, all of the animals cited above
are here referred to Gerbillus pyramidum pyramidum.
PACHYUROMYS DUPRASI NATRONENSIS De Winton
Pachyuromys dupresi (sic) natronensis DE WintToN, Noy. Zool., vol. 10, p. 285,
August 25, 1903 (Bir Victoria, on the way to Wadi Natron from the Nile
Valley, Egypt).
Specimens examined.—Two. Giza Province: West of Abu Rau-
wash, Cairo area, 1 (MVZ). Egypt: Presumably near Cairo, 1.
Description—Upper parts Pinkish-Cinnamon, individual hairs
plumbeous at base and finely tipped with black; color purest on sides,
flanks, cheeks and top of muzzle; hairs of cheeks and muzzle, where
pigmented, white at base; entire underparts, lower half of cheeks,
thighs, dorsal surfaces of hands and feet, tips of ears, and supra-
orbital spots, white; all unpigmented hairs white to base; tail finely
covered with white hairs. Tail short and heavy, becoming expanded
more or less club-shaped distally and in dry specimens flattened dorso-
ventrally; hind feet small; ears rather large.
Measurements.—An adult female, labeled only Egypt but presum-
ably from near Cairo, measured: Total length 165; length of tail
55; length of hind foot 24; occipitonasal length 35.3; length of palate
15.8; length of palatal foramina 6.4; length of bulla 16.5; crown
length of upper molar series 4.4; least interorbital width 6.3.
Remarks.—This animal superficially resembles Meriones libycus but
may be readily distinguished from that species by the short, club-
shaped, fatty tail and by a greater extension of the nasals and much
larger, more inflated bullae.
The immature animal lacks the bright color of the adult and appears
instead much darker, owing to the greater amount of basal plumbeous
and black tipping of the hairs. In the immature skull, the general
conformation is the same but minor changes occur as adulthood is
reached. ‘The space between the mastoid portions of the auditory
bullae is greater in the immature, no supraorbital ridge is developed,
the nasals appear to encroach more into the frontals, the interparietal
becomes slightly smaller and the wings of the supraoccipital become
more defined as the animal ages.
MERIONES LIBYCUS LIBYCUS Lichtenstein
Meriones libycus LICHTENSTEIN, Verzeich. Doubl. Zool. Mus. Berlin, 1823, p. 5,
No. 9. (Type locality given by Ellerman (1947, p. 485) as near Alexandria,
Egypt.)
Specimens examined.—Thirteen. Giza Province: Suez Road,
8 miles east of Heliopolis, 1; Suez Road, 5 miles east of Heliopolis, 10.
Suez Canal Zone: 3 miles northwest of Fayid, 2.
358 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 102
Description.—Animals from near Heliopolis have the upper parts
Warm Buff with a moderate admixture of black; color purest on sides,
flanks and cheeks; entire ventral surface, dorsal and ventral surfaces
of hands and feet, muzzle, postorbital and postauricular spots, white;
tail above as color of back, shading to a black penicillate tip; purer
color on ventral half of tail. AJIl pigmented hairs plumbeous at
base. Eye bordered by narrow black ring. Ears light brownish,
finely covered with white hairs inside and out. Part of vibrissae
white, part black.
Measurements.—Averages and extremes of three males and seven
females from five miles east of Heliopolis, were, respectively: Total
length 265.3 (256-274), 242 (226-261) ; length of tail 126.3 (123-182),
120.0 (108-132) ; length of hind foot 33.3 (83-34), 31.5 (380-33) 5 ear
from notch 18 (17-19), 16 (11-18) ; occipitonasal length 39.2 (38.6-
39.7), 35.9 (34.8-37.6); length of palate 16.7 (16.4-17.1), 16.1
(15.5-16.9) ; length of palatal foramina 6.6 (6.5-6.7), 6.3 (5.6-6.6) ;
length of auditory bulla 14.8 (14.5-15.6), 13.9 (18.1-14.7); crown
length of upper molar series 4.7 (4.6-5.1), 4.5 (4.4-4.7) ; least inter-
orbital width 6.0 (5.5-6.4), 5.9 (5.4-6.3); weight 85.6 (77-100),
59.0 (50-71).
Remarks.—Ellerman, in March 1941 (p. 533), places the species
Meriones shawii as a pure synonym of Meriones libycus libycus, and
on the same page (533) places Meriones crassus as a subspecies of
M. libycus. In May 1947 he elevated Mf. crassus to full specific rank
and removed M. shawii from synonomy, making it a subspecies of
M. Uibycus. In raising M. crassus to full specific rank, he considered
that nine subspecies properly belonged to this former subspecies.
The subspecies he placed under M. crassus belonged to the species
M. libycus, M. schouesboei, M. erythrourus, M. ismahelis and M.
arimalius. In October 1947 Ellerman and Chaworth-Musters listed
eight subspecies of M. libycus, nine of M. crassus, and placed M. shawit
as a full species with three subspecies. In none of the papers cited
above do Ellerman, or his coauthor, Chaworth-Musters, give any
reason for suppressing or resurrecting species or subspecies.
In “A Revision of the Genus Meriones,” Chaworth-Musters and
Ellerman (1947) cite as key characters: Hind claws dark, M. libycus,
and hind claws pale, M. crassus. In addition to the claw character he
gives the joined or closely aproximated processes around the supra-
meatal triangle as a character distinguishing WM. libycus. In M.
crassus the processes are not supposed to be closed or closely approxi-
mating. In specimen U.S.N.M. No. 282509 one hind foot has dark-
colored and the other hind foot has pale-colored claws. Several other
specimens in the series show a similar condition but not quite so pro-
MAMMALS FROM THE NILE DELTA REGION, EGYPT—SETZER 359
nounced. The processes around the suprameatal triangle are also
variable. In U.S.N.M. Nos. 282509 and 282512, as examples, one
side shows the processes closed or nearly approximating, while the
other side is open and not closely approximated.
From this and from the indecision in placing species, it thus ap-
pears that the degree of variation being dealt with is certainly not on
a specific but rather on a subspecific level.
PSAMMOMYS OBESUS NICOLLI Thomas
Psammomys obesus nicolli Tuomas, Ann. Mag. Nat. Hist., ser. 8, vol. 2, p. 92,
July 1908 (Damietta, northern Egypt).
Specimens examined.—Two. Damietta.
Description.—Upper parts Mikado Brown, strongly suffused with
black; color purest on top of head, rump and on the shoulders; dark
dorsal color shading over sides into the lighter Clay Color tipped hairs
of the underparts; all hairs of the body plumbeous at base; dorsal sur-
faces of hands and feet yellowish white; narrow dorsal stripe of tail
brownish black, expanding distally to form a brush; underside of tail
as pure color of the dorsum; vibrissae mixed black and white; cheeks
grayish. Tail shorter than head and body; ears short and rounded,
sparsely covered with whitish hairs inside and out; forefeet small,
four-toed but vestige of thumb remaining as stout claw. Skull re-
sembling that of Meriones, except upper incisors are not grooved;
supraorbital ridges pronounced.
Measurements——No external measurements were available; a sub-
adult male skull from Damietta measured: Length of palate 17.3;
length of palatal foramina 5.6; crown length of upper molar series
5.53 least interorbital width 6.5; length of nasals 13.7.
Remarks.—In the absence of comparative material and since these
specimens are topotypical they are here referred to Psammomys obe-
sus nicolli.
PSAMMOMYS OBESUS OBESUS Cretzschmar
This subspecies of sand rat is known from the Alexandria area and
may be distinguished from P. 0. nicolli, from Damietta, solely on the
basis of its lighter color. No specimens have been examined.
Family SPALACIDAE
SPALAX AEGYPTIACUS Nehring
The mole rat is apparently only locally distributed in the region
around Mariut and Ramleh near Alexandria. Its presence may be
ascertained by small mounds of earth thrown out of its burrow system
in much the same manner as the pocket gopher of North America.
360 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Even though these animals show no extenal eye, there is supposed to
be a minute, completely organized eye present under the skin. No
specimens have been examined.
Family MURIDAE
ARVICANTHIS NILOTICUS (Desmarest)
Arvicola niloticus DESMAREST, Mammalogie . . ., pt. 2, p. 281, 1822 (Egypt).
Specimens examined.—Two. Behera Province: Alexandria, 1
Fayum Province: Lake Fayum, 1.
Description.—Dorsal hairs tipped with Ochraceous-Tawny, bases
and other hairs of the dorsum throughout their length, blackish ;
strong admixture of blackish hairs on dorsum, becoming fewer on
sides and rump; nose, ring around eyes and hairs of ramp Buckthorn
Brown; tail bicolored, blackish on dorsal surface, but sparsely haired ;
ears rather large, rounded and but finely covered with hair; hairs of
belly, ventral surfaces of fore and hind legs white-tipped, these hairs
with blackish bases; immature pelage darker. Tail about equal to
head and body in length; hind feet rather long; thumb expressed as a
rudimentary claw only.
Measurements.—One adult male from Lake Fayum, Fayum Prov-
ince, measured: Total length 321; length of tail 141; length of hind
foot 38; length of ear from notch 20; length of palate 16.9; length of
palatal foramina 8.5; crown length of upper molar series 6.5; length
of nasals 14.6; least interorbital width 5.4; weight 56.0.
Remarks—On superficial examination these animals are surpris-
ingly like the North American Cricetine genus Sigmodon, both ex-
ternally and cranially. From recorded habitat observations they
both occupy much the same ecological niche.
RATTUS NORVEGICUS (Berkenhout)
Mus norvegicus BERKENHOUT, Outlines of the natural history of Great Britain,
vol. 1, p. 5, 1769.
Specimens examined.—Twenty-eight. Behera Province: Alexan-
dria, 7. Suez Canal Zone: Port Said, 6; Ismaila, 8; Suez, 7.
Description.—Upper parts near Verona Brown, no pure color, all
hairs on back black tipped and plumbeous based, with the intermediate
area of the above color; sides, flanks and shoulders somewhat lighter
than dorsal color; throat, underside of forelimbs, entire belly and
dorsal surfaces of hands and feet, white; hairs of belly and throat,
plumbeous based ; tail blackish, but sparsely covered with short whitish
hairs, annulations distinct. Ears small; tail equal to or somewhat
less than head and body; eyes small. Skull: Temporal ridges paral-
lel or nearly so posteriorly; anterior molar with small accessory cusp
on anterior edge.
MAMMALS FROM THE NILE DELTA REGION, EGYPT—SETZER 361
Measurements.—Averages and extremes of five males and two fe-
males from Alexandria, Behera Province, were, respectively: Total
length 398 (370-428), 415 (415) ; length of tail 195.4 (182-207), 196.5
(194-199) ; length of hind foot 42 (40-46), 40 (89-41) ; length of ear
from notch 19.6 (19-20), 19 (19) ; greatest length of skull 46.4 (44.5-
49.5), 27.2 (47.0-47.4) ; condylobasal length 41.1 (39.3-48.9), 41.8
(41.5-42.1) ; length of palate 22.3 (21.5-24.2) , 22.2 (21.9-22.6) ; length
of palatal foramina 7.9 (7.4-8.4), 7.85 (7.7-8.0) ; alveolar length of
upper molar series 7.5 (7.47.6), 7.4 (7.3-7.5) ; least interorbital width
6.6 (6.4-7.0), 7.0 (6.9-7.1) ; length of nasals 17.2 (15.8-19.3), 17.75
(17.5-18.0) ; greatest zygomatic width 22.7 (21.0-25.4), 22.85 (22.8-
22.9).
Remarks.—As may be noted from the measurements, these rats show
a great amount of age variation. No effort was made to segregate
senile from subadult individuals for the purposes of measurement.
It is interesting to note that the averages of individuals from the four
areas represented are quite close to one another. There are no observa-
ble peculiarities in these introduced rats.
RATTUS RATTUS (Linnaeus)
Mus rattus LINNAEUS, Systema naturae, ed. 10, p. 61, 1758 (Sweden).
Specimens examined.—Thirty-four. Behera Province: Alexandria,
11. Daqahlia Province: 5 miles west of Simbillawein, 4. Giza
Province: Cairo, 11; Kafr Taharmes, 2; Talbia, 4. Suez Canal Zone:
Fayid, 2.
Description.—Three basic color types of this species occur in the
Delta region. They are: (1) Blackish brown upper parts, with plum-
beous belly and hands and feet; (2) grizzled brown upper parts with
buff-tipped plumbeous-based hairs on the belly and brownish hands
and feet, the toes of both hands and feet whitish; (3) dorsal color as
in (2) but somewhat lighter, with underparts white, washed with
yellowish, the hands and feet whitish. Tail longer than head and
body, but finely covered with hair; annulations pronounced; ears
large and rounded. Temporal ridges of skull more or less in the form
of a lyre, never parallel as in R. norvegicus; anterior upper molar
without accessory cusp.
Measurements.—Averages and extremes of eight males and nine
females, all of the third color type, from several localities were, re-
spectively: Total length 412 (378-432), 406 (355-460) ; length of tail
227.5 (208-249), 294.2 (194-262) ; length of hind foot 36.5 (34-38),
35.6 (33-87) ; length of ear from notch 23.5 (21-25), 24.8 (24-96) ;
greatest length of skull 42.3 (40.6-44.3) , 42.7 (39.6-46.6) ; condylobasal
length 36.9 (35.0-37.8), 36.7 (33.4-40.7) ; length of palate 19.6 (18.6-
362 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
20.0), 19.6 (18.1-21.4) ; length of anterior palatal foramina 7.4 (6.8-
8.3), 7.6 (7.0-8.3) ; alveolar length of upper molar series 6.8 (6.3-7.2),
6.9 (6.5-7.8) ; least interorbital width 6.1 (5.8-6.3), 6.1 (5.6-6.6) ;
length of nasals 14.9 (18.5-15.8), 15.2 (13.3-17.0).
Remarks.—lf one were to recognize subspecies of the commensal rat
then the three color types mentioned under description would become
(1) Rattus rattus rattus, (2) Rattus rattus alecandrinus (which may
be considered topotypical), and (8) Rattus rattus frugivorous. How-
ever, since the three basic color types do not appear to segregate out in
any given series from any given locality as just these three color types,
but show in six of thirty-four specimens intermediate conditions be-
tween any two of the three, it is felt that here, as in Acomys, 1s an ex-
pression of some genetic factor, called by some a single gene character
for melanism. Certainly, one of the criteria for subspecies is a free
interbreeding between two or more contiguous kinds which would be
expected to produce intermediate types. It is extremely hard, though,
to visualize three distinct subspecies occupying the same environmenta!
niche. Flower (1932, p. 409) and many others mention the fact that
no habitat differences could be ascertained between the so-called three
subspecies. It may be that if these animals were to be found ferally,
one or the other of the three types would segregate and be com-
pletely recognizable. It is worthy of mention, though, that the four
wild-caught animals from 5 miles west of Simbillawein show all
three color types plus one that is intermediate between types two and
three.
It is felt that, since there appears to be no segregation, so far as is
known, it is best to ignore the subspecific names for the three above-
mentioned rats until such time as these names can be shown to be valid.
MUS MUSCULUS Linnaeus
Mus musculus LINNAEUS, Systema naturae, ed. 10, p. 62, 1758 (Upsala, Sweden).
Specimens exvamined.—Eighteen. Behera Province: Alexandria, 6;
1% mile west of Dikheila Airfield, 1; Heliopolis, 1; Maadi, 1. Dagahlia
Province: 5 miles west of Simbillawein, 3. Giza Province: Kafr
Taharmes, 1; Kuniessa, 1; 'Talbia, 1. Suez Canal Zone: Port Said, 1;
Suez, 2.
Description—Three color phases are present in the small series
from Egypt. They are: (1) Pallid Neutral Gray rather strongly in-
termixed with black; (2) Cinnamon-Brown without so much black
intermixed; and (3) Mummy Brown, strongly suffused with black.
In all color phases the hands and feet are white; the bellies in the first
two phases are white; in the first the hairs are white to the bases, in
the second the hairs are plumbeous at the base; in the third phase
the belly is buffy, with the hairs plumbeous at the base. In all three
MAMMALS FROM THE NILE DELTA REGION, EGYPT—SETZER 363
phases the tail is more or less bicolored and the dark dorsal stripe cor-
responds to the back color. In the first phase, the ears are pale, re-
sembling the dorsal color; in the second and third phases the ears are
dark, as isthe dorsal color. The skulls show no one feature that might
tend to segregate any of the above color phases as subspecies. They
are all regarded as Mus musculus.
Measurements—Averages and extremes of two males and four
females from Alexandria, Behera Province, were respectively : Total
length 157 (156-158), 166.2 (157-173); length of tail 78 (78), 78.7
(74-82) ; length of hind foot 17 (17), 17.3 (17-18) ; length of ear from
notch 18 (13), 13.3 (18-14) ; greatest length of skull 21.7 (21.2-22.2),
21.7 (21.0-22.4); occipitonasal length 21.2 (20.8-21.6), 21.0 (20.5-
21.7); length of palate 10.05 (9.8-10.3), 9.9 (9.6-10.4); length of
palatal foramina 5.2 (5.2), 4.7 (4.5-5.0) ; crown length of upper molar
series 3.15 (3.1-3.2), 3.2 (3.0-3.4) ; length of nasals 7.9 (7.8-8.0), 7.8
(7.5-8.3) ; least interorbital width 3.55 (38.5-3.6), 3.5 (8.5); greatest
zygomatic width 11.05 (11.0-11.1), 10.9 (10.6-11.1).
Remarks.—The names applied to the house mice of the Delta region
appear to be in as much confusion as are the names applied to Acomys
and to Rattus. It is not thought wise to try to solve the nomen-
clatural mix-up on so few specimens. The material available is ex-
tremely heterogeneous and shows all the types that have been rec-
ognized from the Nile Delta and southward.
Mus musculus gentilis, which occurs in a feral condition farther
south along the Nile, appears to be the name most applicable to the
largest numbers of animals from the Alexandria-Cairo-Suez area.
There are, however, individuals that would appear to belong to the
north European stock of Mus musculus musculus and Mus musculus
domesticus. In addition, there are specimens of a light color phase
from Alexandria that do not correspond to any of the named kinds of
Mus musculus supposed to be in this area. Cranially, all the speci-
mens examined appear to be of one subspecies.
It is thought best, at this time, merely to refer the house mice of
northern Egypt to the species Mus musculus.
ACOMYS CAHIRINUS (E. Geoffroy St.-Hilaire)
Mus cahirinus BE. Grorrroy St.-HImarre, Catalogue des mammiféres du Muséum
National d’Histoire Naturelle, p. 195, 1803 (Cairo, Egypt).
Specimens examined.—Thirty. Behera Province: Alexandria, 8.
Daqahlia Province: 5 miles west of Simbillawein, 1. Giza Province:
Cairo, Abassia Fever Hospital, 5; Abu Rauwash, 2 (MVZ); Cairo,
4; Kafr Taharmes, 1; Saket Meki, 2. Suez Canal Zone: Suez, 6.
Nubia: Abu Simbel, 1 (MVZ).
364 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 102
Description—Upper parts from between Ochraceous-Orange and
Ochraceous-Tawny to deep slate gray; belly yellowish to light slate
gray. Color on body nowhere pure; in light phase strongly suffused
with black-tipped hairs; in dark phase uniformly slaty. Hands, feet
and tail of both color phases as belly color. Tail equal to head and
body in length, strongly annulated and with small bristlelike hairs.
Pelage coarse but becoming spinose between the ears and spines increas-
ing in size and number over the rump. Ears large, rounded, and but
sparsely covered with fine yellowish hairs.
Measurements.—Averages and extremes of two males and six fe-
males from Alexandria, Behera Province, Egypt, were, respectively :
Total length 209.5 (198-221), 218.6 (208-225) ; length of tail 113.6
(108-121), 106.6 (97-112) ; length of hind foot 19 (18-20), 18 (18);
length of ear from notch 18.5 (17-20), 18 (17-20); greatest length
of skull 28.05 (27.0-29.1), 28.28 (27.9-28.9); condylobasal length
23.85 (22.7-25.0), 24.0 (23.4-24.8) ; length of palate 14.55 (13.5-15.6),
14.32 (18.7-15.0) ; alveolar length of upper molar series 4.3 (4.2-4.4),
4.4 (4.4); length of palatal foramina 6.3 (6.2-6.4), 62 (5.8-6.6) ;
least interorbital width 4.8 (4.8), 4.7 (4.64.8) ; length of nasals 10.3
(9.9-10.7), 10.6 (10.2-11.1).
Remarks.—These small spiny mice have, at various times, been
designated by various specific or subspecific names. Most of these
names have, apparently, been based on differences in color. From
the specimens now at hand, it appears that the only name strictly
applicable is Acomys cahirinus. KE. Geoffroy St.-Hilaire evidently
had before him a specimen of the light-colored phase when he de-
scribed this animal, and subsequent authors must have had repre-
sentatives of the other more melanistic types. In all, the two basic
types of pelage color are light above and light below and dark above
and dark below. From these two phases all possible gradations can
be demonstrated by specimens from Abassia Fever Hospital, in Cario,
or from Alexandria.
From ail appearances this peculiarity of color is nothing more than
the expression of melanism. In many ways these small mice recall
the variation of pelage found in Rattus rattus and considered by many
to be subspecies.
Family DIPODIDAE
JACULUS JACULUS JACULUS (Linnaeus)
Mus jaculus LINNAEUS, Systema naturae, ed. 10, p. 68, 1758 (type locality given
by Linnaeus as “In Arabia, Calmukia,’” emended by Allen (1939, p. 423) as
Pyramids of Giza).
Specimens examined.—Twenty-two. Giza Province: Abu Rau-
wash, 5 (MVZ) ; Suez Road, 5 miles east of Heliopolis, 4; 1 mile north-
MAMMALS FROM THE NILE DELTA REGION, EGYPT—-SETZER 365
east of Kom Aushim, 1; Alexandria Road, 5 miles northwest of Pyra-
mids of Giza, 6. Suez Canal Zone: 3 miles northwest of Fayid, 6.
Description—Upper parts between Vinaceous-Cinnamon and Pink-
ish-Cinnamon; hairs plumbeous at base, grading into above color and
finely tipped with black; sides and flanks with hairs white to base
with a slight admixture of the above colors; entire ventral surface,
anterior and ventral surface of thighs, forelegs, hip stripe, postauricu-
lar and postorbital spots, muzzle, cheeks, tip and ventral half of
tail, white. Ears about as dorsal coloration, sparsely covered with
fine white hairs. Eye bordered by narrow black ring. Tail above,
for proximal two-thirds, as dorsal color; other one-third divided sub-
equally by proximal black ring, incomplete ventrally, and a distal
white tip.
Measurements.—Averages and extremes of two males and four
females from 5 miles northwest of the Pyramids of Giza were,
respectively: Total length 293 (292-294), 294.5 (284-801) ; length of
tail 179 (176-182), 182 (176-188) ; length of hind foot 62 (60-64),
61.7 (60-65) ; length of ear from notch 19 (19), 19.2 (19-20) ; occipi-
tonasal length 30.0 (30.0), 30.2 (29.6-30.7) ; greatest length of skull
33.1 (83.1), 33.1 (32.6-33.9) ; length of nasals 10.65 (10.4-10.9), 10.9
(10.4-11.4) ; least interorbital width 11.9 (11.8-12.0), 12.1 (11.5-12.7) ;
length of palate 15.6 (15.6), 15.6 (15.2-15.9) ; length of palatal foram-
ina 4.35 (4.3-4.4), 4.25 (4.24.3); length of bulla 14.0 (14.0), 13.8
(13.5-14.2) ; width of skull across bullae 22.6 (22.6), 21.3 (21.1-21.5) ;
width across maxillary arches 22.5 (22.3-22.7) , 22.0 (21.3-22.5) ; crown
length of upper molar series 4.5 (4.44.6), 4.75 (4.7-4.8) ; weight 50.0
(48.0-52.0), 53.0 (51.0-56.0).
Remarks.—If the animals from 5 miles northwest of the Pyramids
of Giza are considered topotypical, a slight difference is noted in the
size of the crania and the length of the hind foot from those animals
from 8 miles northwest of Fayid and 5 miles east of Heliopolis. The
pelage in the animals from the latter two localities has less reddish
and more black admixture, thus presenting a somewhat darker tone.
Order CARNIVORA
Family CANIDAE
CANIS LUPASTER Hemprich and Ehrenberg
Apparently the jackal is not uncommon. The status of the name
applied to the jackal of this region is in some doubt. It can be dis-
tinguished from the domestic dog by its shorter muzzle, wider face,
and brighter coloration. No specimens have been examined.
366 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
VULPES VULPES NILOTICA (KE. Geoffroy St.-Hilaire)
Canis niloticus B. GEOFFROY StT.-HILAIRE, Catalogue des mammiféres du Muséum
National d’Histoire Naturelle, p. 134, 1803 (Egypt).
Vulpes vulpes aegyptiaca, ANDERSON and DE WINTON, Zoology of Egypt, Mam-
malia .. ., p. 227, pl. 82, 1902 (Egypt).
Specimen examined —One. Dagqahlia Province: 5 miles west of
Simbillawein.
Description—Dorsal stripe, beginning between shoulders and con-
tinuing onto dorsal one-fourth of tail, Sanford’s Brown, strongly in-
termixed with black hairs; all except black hairs with a subapical band
of yellowish white and wide basal band of plumbeous; Sanford’s
Brown purest on cheeks, around eyes, inside and back of fore limbs and
back of hind limbs; throat, chest, and belly to peneal region smoky
gray with some admixture of whitish hairs; inside of ears, lower part
of cheeks and continuing toward and beyond angle of jaws, lateral
and ventral surfaces of base and distal half of tail, white; white hairs
of distal half of tail sparsely tipped with black; sides, shoulders, and
flanks, grayish; areas on side of neck, behind shoulder and groin, yel-
lowish white; anterior face of foreleg with blackish brown, white and
black longitudinal stripes; anterior face of thigh and continuing onto
dorsal surface of hind foot with yellowish-white stripe; posterior
dorsal one-half of ears black.
Measurements.—An adult male from 5 miles west of Simbillawein
measured: Total length 1,052; length of tail 416; length of hind foot
157; length of ear from notch 101; greatest length of skull 149.6; oc-
cipitonasal length 130.3; condylobasal length 136.2; length of palate
73.1; width across M' 40.7; greatest width of braincase 46.3; least
interorbital width (behind supraorbital processes) 20.0; length of
nasals 40.5; width of rostrum 25.1.
Family MUSTELIDAE
MUSTELA NIVALIS SUBPALMATA Hemprich and Ehrenberg
Mustelae subpalmatae HeMpPRicH and EHRENBERG, Symbolae physicae ..., dec.
2, folio k, p. 2, September 1852 (in houses of Cairo and Alexandria, Egypt).
Specimens examined.—Six. Daqahlia Province: 5 miles west of
Simbillawein, 2. Behera Province: Alexandria, 1. Giza Province:
Kafr Taharmes, 1; Kuniessa, 1; Saket Meki, 1.
Description.—All dorsal coloration, including hands, feet, ears, and
tail all around, between Brussels Brown and Raw Umber; chin, throat,
chest, and most of belly, white or a yellowish white; brown of dorsal
coloration extending onto belly, sometimes continuous across belly but
MAMMALS FROM THE NILE DELTA REGION, EGYPT—SETZER 367
more frequently merely spotted; tail with faint suggestion of black
tip. Ears and eyes small; tail about half as long as head and body.
Measurements.—An adult male from Kuniessa, Giza Province, and
an adult female from 5 miles west of Simbillawein measured, respec-
tively: Total length 381, 330; length of tail 120, 92; length of hind
foot 47, 39; length of ear from notch —, 17; basilar length 46.7, 42.1;
canine to M® 13.0, 12.0; breadth of rostrum across lacrimals 13.7, 11.2;
least interorbital width 8.5, 7.4; mastoid breadth 25.0, 20.9; zygomatic
breadth 27.7, 22.7.
Remarks.—Apparently, in Egypt, this animal is more an inhabitant
of houses than it is of the countryside. All the specimens studied
were caught in buildings during rat-control programs.
POECILICTIS LIBYCA (Hemprich and Ehrenberg)
Mustela libyca HEMPRICH and HHRENBERG, Symbolae physicae . . ., dec. 2, folio
k, p. 6, 1832 (Egypt).
Specimen examined.—One. Giza Province: Cairo Area, Abu
Rauwash (MVZ).
Description.—Colors of this animal are white and blackish brown.
White distributed on the body as small mystacial patches; a narrow
white ring occurs between the eyes and ears, then turning ventrad
around the point of the jaw and meeting on the bottom of the throat;
tip of ears white; base of a more or less lyre-shaped band of white
begins in back of the ears and behind the forehead stripe, the band
running posteriorly and somewhat lateroventrad to cross the flanks
and continue to the base of the tail; inside this large lyre is a smaller
lyre-shaped band of white, which becomes closed at the base of the
tail; inside the loop of the median lyre is a more or less diamond-
shaped patch completely encircled by black and with a small black spot
in the center. The outermost lyre-shaped band continues onto the
dorsal surface of the tail to the tip and for approximately one-half the
ventralsurface. The posterior ventral half of the tail has black-tipped
white hairs. Entire ventral surface covered with blackish brown hairs.
Ears small and rounded; soles of hands and feet, with the exception of
the pads, haired; eyes small.
Measurements—An adult male from Abu Rauwash measured:
Total length 388; length of tail 158; length of hind foot 43; condyloba-
sal length 53.6; length of palate 25.1; M' to canine 18.1; breadth across
P* 18.5; width of zygomatic arches 33.5; width of rostrum 11.4; width
of cranial constriction behind postorbital processes 11.5; width across
bullae 31.1.
Remarks.—This small mustelid is remarkably like the North Amer-
ican genus Spilogale in its markings.
368 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Family VIVERRIDAE
HERPESTES ICHNEUMON (Linnaeus)
Viverra ichneumon LinnaEvs, Systema naturae, ed. 10, p. 48, 1758 (Egypt).
Specimen examined.—One. Daqahlia Province: 5 miles west of
Simbillawein.
Description—Color : Under fur shading from Isabella Color on neck
and shoulders to Cinnamon-Rufous on rump, thighs, and base of tail.
Guard hairs long (about 65 mm.) and eight-banded, with alternating
white and black bands; tips of guard hairs brownish instead of black
as rest of bands; face, chin, muzzle, and dorsal surfaces of hands and
feet, blackish; palms and soles naked; belly sparsely haired. Entire
animal, owing to banded hairs, presents a grizzled appearance. Ears
small and rounded; tail not quite so long as head and body, and owing
to the banded hairs appears to be ringed; tail more or less tufted at tip.
Measurements—One male from 5 miles west of Simbillawein meas-
ured: Total length 990; length of tail 453; length of hind foot 104;
length of ear from notch 37; condylobasal length 104.6; zygomatic
width 53.7; postorbital width 17.5; interorbital width 18.5; maxillary
width 32.4.
Family FELIDAE
FELIS CHAUS Schreber
This cat is apparently to be found throughout the Delta region,
usually in low marshy places. It may be distinguished from the feral
Felis catus by its shorter tail (less than one-half the head and body
length) ; by its larger, black-tipped ears; and by its reddish feet. It is
apparently not uncommon. No specimens have been examined.
MAMMALS FROM THE NILE DELTA REGION, EGYPT—SETZER 369
LITERATURE CITED
ALLEN, G. M.
1939. A checklist of African mammals. Bull. Mus. Comp. Zool., vol. 83,
pp. 763. (February.)
ANDERSON, JOHN
1902. Zoology of Egypt: Mammalia (revised and completed by W. HB.
De Winton), pp. xvii + 374, pls. 638, figs. 7.
CHAWoRTH-MustTers, J. L., and ELLerRMAN, J. R.
1947.
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 Washington: 1952 No. 3306
THE SIPUNCULID WORMS OF CALIFORNIA AND BAJA
CALIFORNIA
By Water Kenrick FIsHer
SIPUNCULID is the name informally bestowed on members of a small
phylum of marine wormlike animals of world-wide distribution—the
Sipunculoidea.!. These creatures live from the intertidal zone to
oceanic depths, in tubular excavations in mud and sand, in clefts of
rocks, in masses of coral, in borrowed shells, under rocks, in holdfasts
of kelp, or in almost any protected situation that affords access to
reasonably clear water and food. They are not found in shifting
sand or mud, since in such habitats they are likely to be buried or
else deprived of protection. Their food is finely divided detritus
selected or trapped by the ciliated tentacles, or bottom material
actively swallowed. This latter material is not necessarily finely
divided. It is sometimes coarse, including the miscellaneous small
fragments of the hard parts of Foraminifera, corals, bryozoans, an-
nelids, mollusks, echinoderms, and crustaceans, often very abundant,
especially in calcareous sands.
1 Adam Sedgwick, in his ‘Students’ Textbook of Zoology”’ (vol. 1, p. 594, 1898), first accorded this group
phyletic rank. It had long occupied in the Annelida the position of a poor relation, as one of the orders of
the class Gephyrea, in which Quatrefages also placed the Echiuroidea and Priapuloidea. ‘‘Gephyrea’’
either as a class or a phylum is an unnatural and unsatisfactory association, but one which has been singu-
larly viable. It was used by Shipley (1910) in the ‘‘Cambridge Natural History” and by Borradaile and
Potts (1932) in ‘““I'he Invertebrata’’—to mention two instances. For a good general account, consult Grace
E. Pickford, “Sipunculida,’’ Encyclopedia Brittanica, 1947; J. W. Spengel, Handworterbuch der Natur-
wissenschaften, vol. 9, pp. 97-106, 1913; Yves Delage and Edgard Hérouard, Traité de Zoologie Concréte, vol.
5, pp. 12-26, 1897; F. Baltzer, ‘“Sipunculida’”’; Kiikenthal and Krumbach’s “Handbuch der Zoologie,”’ vol.
2, pt. 9, pp. 15-61, 1931.
371
372 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
The region covered by this report centers in the coastal waters of
California and Baja California but includes what little is known of
the fauna north to the Alaskan Peninsula.
The material has been accumulating over a considerable period of
years and includes specimens dredged by the United States Fisheries
steamer Albatross in 1904. Unfortunately only about a third of the
Albatross collection was in a condition suitable for study when it
became available in 1946. The Harriman Alaska Expedition (1899),
through Dr. Wesley R. Coe and the late Dr. William E. Ritter, made
a small but important contribution. By far the largest collections
were contributed by Prof. G. E. MacGinitie, Dr. W. F. Thompson,
and the late Edward F. Ricketts. I am also indebted to the United
States National Museum and the Museum of Comparative Zoology
for the loan of material, to Dr. J. H. Gerould for the loan of reprints,
and to Dr. Elisabeth Deichmann and Dr. Elise Wesenberg-Lund for
extracts from the important works of Leuckart and Diesing.
For contributions of material thanks are due to the following
persons: Dr. M. W. de Laubenfels, Dr. D. F. Hoffmeister, Dr. C. L.
Hubbs, Prof. Trevor Kincaid, Dr. S. F. Light, W. H. MacGinitie,
E. B. and T. T. McCabe, R. J. Menzies, Dr. E. H. Myers, Ida S.
Oldroyd, D. L. Reish, Dr. N. W. Riser, Dr. Richard Synder, Dr.
Emery F. Swan, Dr. H. B. Torrey, W. J. Waidzonas, Patrick H.
Wells, Dr. D. L. Ray, and Dr. D. M. Wootton.
As compared with Japan the sipunculid fauna of the west coast of
North America is meager. Phascolosoma agassizii is the dominant
intertidal species from Kodiak, Alaska, to Point Conception, Calif.
Its range extends to northern Baja California. Ph. puntarenae is its
equivalent from Cape San Lucas to Panama. Dendrostomum pyroides
occurs from Coos Bay, Oreg., to northern Baja California, while D.
dyscritum has been taken between Point Conception and the northern
boundary of California. D. zostericolum has been found from Point
Conception to Ensenada, Baja California. D. perimeces, a consistent
inhabitant of sandy mud, probably colonizes estuaries and sloughs from
an off-shore, shallow-water population. It has been taken between
Bodega Head and Venice, Calif. D. hexadactylum, which resembles
pyroides, has been found only in 10 to 20 fathoms, in Monterey Bay.
It was described from Japanese waters. The conspicuously large
Siphonosoma ingens is a southern California species, extending north
to Monterey Bay. Sipunculus nudus is an off-shore cosmopolite,
which finds its way into estuaries of southern California and into the
intertidal zone of Baja California and the Mexican coast. The genus
Golfingia (formerly Phascolosoma) is a negligible element in the shore
SIPUNCULID WORMS OF CALIFORNIA—FISHER 373
fauna of California, only three inconspicuous species having been
found: @. margaritacea californiensis, G. hespera, and G. macginitiet,
all rare or very local in occurrence.
Only two sipunculids are known from Alaska: Golfingia margaritacea
(Sitka, Kilisnoo, Dutch Harbor, Point Barrow) and Phascolosoma
agassizii (Cape Fox, Prince William Sound, Kodiak). Undoubtedly
G. vulgaris occurs in shallow water.
Phascolosoma agassizii has been taken from intertidal stations
on Vancouver Island and the Queen Charlotte Islands, British Colum-
bia, and G. margaritacea has been taken at Vancouver Island.
Washington has four species, all from Puget Sound and vicinity:
Golfingia vulgaris, G. margaritacea, G. pugettensis, and Phascolosoma
agassizi.
Two species are known from Oregon (Coos Bay): Dendrostomum
pyroides and Phascolosoma agassizv.
With this unimpressive showing, I need hardly emphasize that this
paper is not a definitive treatment of the fauna. While it is improb-
able that many new intertidal forms will come to light, it is equally
probable that the list of shallow-water and deep-sea species is very
far from complete.
In this paper I have used Golfingia Lankester in place of Phas-
colosoma authors and have replaced Physcosoma Selenka with Phas-
colosoma Leuckart in order to conform to accepted rules of nomen-
clature. For the same reason I have employed Dendrostomum Grube,
the original spelling, instead of Dendrostoma, a later emendation by
Keferstein.
KEY TO THE GENERA OF SIPUNCULOIDEA
a’. No horny or calcareous shield or cone at anterior end of trunk, although
enlarged horny papillae and tubercles sometimes present in anal region.
b'. Tentacles surround mouth; tentacles few or many, simple or branched;
nuchal organ, when present, dorsal to tentacles.
c!, Longitudinal muscle layer of wall of trunk divided into separate bundles.
d@', Skin of usually thick body wall provided with longitudinal coelomic
canals or with independent coelomic pouches longitudinally ar-
ranged.
e!, Integumental coelomic spaces are longitudinal canals under cuticle;
4 retractors; intestinal coil with a long, anteriorly anchored loop;
introvert with subtriangular scalelike papillae directed posteriorly.
Sipunculus Linnaeus, 1766 (p. 375)
e?, Integumental coelomic spaces are in form of independent, usually
irregular pockets arranged in longiseries; no accessory anterior
loop in intestinal coil.
f}. Scalelike papillae on introvert; anterior to the 4 retractors are 2
small protractors arising in front of anus; irregular integumental
coelomic spaces independent in each quadrilateral division of
skin and sometimes with several long papillae; spindle muscle
arises on rectum ..-....--.- Xenosiphon Fisher, 1947 (p. 377)
374 PROCEEDINGS OF THE NATIONAL MUSEUM yOL. 102
f?. No scalelike papillae on introvert but rings of simple tiny spines;
2 retractors; skin strongly marked in rectangles; spindle muscle
arises by 2 roots from body wall behind anus, and a third root is
attached to esophagus; nuchal organ prominent; tentacular
divisions of oral disk 8, the 2 ventral bifid; no Polian villi.
Siphonomecus Fisher, 1947
f*. No-scalelike papillae on introvert; skin not divided into rectangles
by furrows; hooks or spinelets present or absent on introvert;
4 retractors; spindle muscle arises by 3 roots, a median in front
of anus and 2 laterals behind anus; some species with incomplete
transverse coelomic dissepiments; contractile vessel usually
with Polian villi__..._-- Siphonosoma ? Spengel, 1912 (p. 380)
ad. Body wall thin without coelomic canals or pockets.
e!. Nephridia with prominent anterior lobe; spindle muscle. with one
root attached to body wall in front of anus and not anchored
posteriorly; no Polian villi; tentacles very small, numerous,
filiform; type with tiny complex hooks on introvert.
Siphonides, new genus (p. 386)
e. Nephridia not bilobed; contractile vessel with lateral pouches but
no villi; no introvert hooks in adults.
Golfingia (subgenus Phascolopsis Fisher, 1950) (p. 398)
c. Longitudinal muscle layer of body wall forming a continuous layer,
without separate bands (except Phascolopsis in Golfingia).
d', Anus in anterior part of trunk behind introvert.
e!. Two nephridia; intestine forms a-close spiral surrounding spindle
muscle; 2 or 4 retractor muscles.
jf}. Tentacles relatively small, digitiform to filiform, few to many,
in which case they are arranged in longitudinal double series
radiating from circumoral disk.
Golfingia ? Lankester, 1885 (p. 388)
f. Tentacles large, bushy, 4 to 8; retractors 2, rarely 4; contractile
vessel with accessory tubules, usually conspicuously long,
Miformaas=)28 he see Dendrostomum ‘ Grube, 1859 (p. 404)
e. Only 1 nephridium, fixed by muscular threads; intestine not in
close spiral but in loops extending forward and backward and held
to body by several strands of muscle fiber; 1 to 3 retractor muscles;
body often spirally twisted in adaptation to life in empty gastropod
and scaphopod shells; single circle of tentacles.
Phascolion > Théel, 1875
d?, Anus on introvert more or less in neighborhood of mouth; one retractor
muscle, attached to posterior end of trunk; 1 nephridium; esophagus
forms a few coils and passes into a short somewhat irregular spiral.
Onchnesoma Koren and Danielssen, 1875
2 The coelomic puuches have not been verified for several small species and may be rudimentary or absent
if body wall is thin, as in Siphonides. Small species with head retracted sometimes resemble Golfingia.
’ Phascolosoma auct., nec Leuckart, 1828.
* Dendrostoma Keferstein, 1865.
5 Cryptosomum Quatrefages, 1866.
SIPUNCULID WORMS OF CALIFORNIA—FISHER 375
v2. Tentacles few to very numerous, arranged in a circle (interrupted dorsally),
enclosing nuchal organ and situated dorsal to mouth; longitudinal
muscle layer of trunk divided into separate anastomosing bundles;
retractors 4, rarely 2; skin covered with papilliform glands which some-
times become tubercular in anal region and at posterior extremity.
Phascolosoma ® Leuckart, 1828 (p. 422)
a?, A specialized shield or cone present at anterior end of trunk.
b!, A horny or calcareous shieldlike structure present at both ends of trunk;
introvert arises on ventral side of anterior shield; longitudinal muscle
layer continuous, or divided into bundles__-Aspidosiphon Diesing, 1851
b?, A hard, caleareous, cone-shaped appendage present at anterior end of
trunk ventral to which is introvert; no posterior shield.
Lithacrosiphon Shipley, 1902
b§, A round caplike structure made up of calcareous plates present at anterior
end of trunk, from center of which the introvert is extruded; no posterior
SS TNA] Cle tee oe fees maranatha ee Cel Cloeosiphon Grube, 1868
Genus SIPUNCULUS Linnaeus
Sipunculus LiInNAbBUus, 1766, p. 1,078. (Type, S. nudus Linnaeus.)
Diagnosis.—Usually large species with long cylindrical body and
short, sharply differentiated introvert covered with squamiform
papillae. Trunk generally thick walled, and longitudinal and circular
muscle layers divided into regular fascicles. Skin divided into ree-
tangular areas by longitudinal and circular furrows. Posterior end
of body rounded or bluntly pointed and sometimes marked off from
main trunk by a limiting ring fold of skin. A flat tentacular fold
surrounds oral disk and from its margin tentacles of varying com-
plexity develop. No hooks on introvert or papillae on the trunk.
Description.—The longitudinal muscle bundles rarely anastomose.
Characteristic of the genus are longitudinal integumentary canals
corresponding to the intervals between the muscle bundles and com-
municating with the coelom by slits between the regular circular
fascicles of muscles that are external to the longitudinal. The canals
contain coelomic fluid and anything floating in it. Retractor muscles
four, separated to head; nephridia two; spindle muscle present or
absent; a dorsal and a ventral contractile vessel without appendages;
especially characteristic is an accessory intestinal spiral (pl. 18, fig.
1, A) between the end of esophagus and the beginning of the true
spire, and coiled in the latter. Esophagus and intestine anchored by
very numerous fixing muscles. ‘‘A median-dorsal unpaired epithelial
tube opens upon the surface of the head immediately behind the
tentacular fold, and leads backward to a cerebral sense organ anterior
¢ The genus erroneously called Physcosoma Selenka, 1897.
376 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
and ventral to the brain. The esophageal connectives do not surround
the attachments of the retractor muscles to the head, as in most
sipunculids, but lie behind and beneath these attachments” (Gerould,
1913, p. 427). The brain is conspicuous. On its front is a con-
spicuous cerebral organ or frons, sometimes elaborately lobed (Fisher,
1947, pl. 10, fig. 2; pl. 11, fig. 3). The ventral nerve cord is not
attached anteriorly by a mesentery to the wall of the introvert, as in
Siphonosoma. It is supported on each side by a strong paraneural
muscle originating from the first, or first and second, muscle bundles
just posterior to beginning of introvert.
Remarks.—Plate 18, figure 1, shows the accessory intestinal spiral
A dissociated from the regular spiral B. This is easily accomplished
by clipping the fixing muscles that hold it in place. I have figured
(Fisher, 1947, pls. 10, 11) this extra spiral in situ in Sipunculus
polymyotus and S. galapagensis, where the bends X and Y are indicated.
The details of the fixing muscles are so different in these two species
as to suggest that they may be of value in classification.
The accessory spiral is not present in Xenostphon or in Siphonosoma.
SIPUNCULUS NUDUS Linnaeus
PuaTE 18
Sipunculus nudus LinNaEvs, 1766, p. 1,078.
Remarks.—This is perhaps the best known and certainly one of the
most widely distributed species of sipunculids—a truly eurythermal
type. On the west coast of America it occurs from Monterey Bay,
Calif., to Panama. It is found on the coast of Brazil, in the West
Indies, and north to Beaufort, N. C. In Europe it inhabits the
North Sea, English Channel, and the Mediterranean and Adriatic
Seas. It is reported from the Red Sea, Indian Ocean, East Indies,
China, Japan, along the western part of the Pacific (Yap, Loyalty
Islands), and South Australia.’
The specimens recorded below have 30 to 33 muscle bundles. The
ventral retractor muscles arise from longitudinal muscles 1-7, 1-6,
or 2-5 (Ensenada) ; the dorsal retractors arise from 9-15, 10-14, 10-15,
11-16. A large specimen from southern California is 190 mm. long,
but others are considerably smaller, circa 130 mm. The longest
specimen is 290 mm. and was washed ashore at Pacific Grove, Calif.,
during a heavy surf.
The species has not been found at Monterey Bay in intertidal
situations, even at Elkhorn Slough, which has been rather thoroughly
explored by Prof. G. E. MacGinitie and others; but it occurs in sub-
tidal situations, since many were washed ashore during a period of
spring tides and heavy surf. It seems probable that Newport Bay
7 A specimen sent me by S. J. Edmonds from Port Willunga, south of Adelaide, South Australia, is super-
ficially like S. nudus but belongs to another, probably new, species.
SIPUNCULID WORMS OF CALIFORNIA—FISHER 377
and Anaheim Slough, in southern California, were colonized by an
off-shore population, as was the Estero de Punta Banda, south of
Ensenada. Professor MacGinitie took it also at a lagoon, Miramar
Beach, Guaymas, Mexico, while Ricketts established a record at
La Paz.
Specimens examined.—Thirty-five, as follows:
Guaymas, Mexico, lagoon at Miramar Beach, Feb. 9, 1948, G. E. MacGinitie,
1 specimen.
El Mogote, Baja California, near La Paz, March 22, 1940, EK. F. Ricketts, 1
specimen.
Estero de Punta Banda, Baja California, 6 miles south of Ensenada, Dec. 19, 1930,
sand, G. E. MacGinitie, 5 specimens.
Anaheim Landing, Calif., March 1918, gravel, 2 specimens.
Newport Bay, Calif., January to February 1930, G. E. MacGinitie, 20 specimens.
Pacific Grove, Calif., Jan. 6, 1939, washed ashore by high seas, 6 specimens.
Genus XENOSIPHON Fisher
Xenosiphon Fisuer, 1947, p. 360. (Type, Sipunculus branchiatus W. Fischer.)
Diagnosis.—Differing from Sipunculus sensu stricto in the following
particulars: Integumental coelomic spaces in the form of independent
sacs of irregular outline; an extra pair of muscles functioning as
retractors and protractors arising from posterior border of introvert
and inserted in front of brain; rectum unusually long, the anus being
in front of nephridiopores; postesophageal intestine without a long
forward loop; nephridia long, slender, attached to body wall for
nearly their entire length; squamiform papillae of very short introvert
increasing in size toward the tentacles, which have very many leaflets
arranged in subtriangular pads surrounding the mouth; type species
with papilliform dermal outgrowths.
XENOSIPHON BRANCHIATUM (Fischer)
PuaTE 19
Sipunculus mundanus var. branchiatus W. Fiscuer, 1895, p. 3, pl. 1, figs. 1, la, 2.
Sipunculus branchiatus SPENGEL, 1913, p. 74.
Xenosiphon branchiatum FisuER, 1947, p. 360, pl. 12.
Description.—The following description is based chiefly on three
specimens from Panama. Length 310 mm.; introvert and tentacle
crown 20-25 mm.; thickness of cylindrical body 8-12 mm., this
varying according to constriction of ring muscles. The specimen from
La Paz, 420 mm. long, is constricted in the middle of body to 8 mm.
diameter. Longitudinal muscle bands 29-34, only rarely anasto-
mosing. When the body is fully inflated the longitudinal and circular
muscles divide the surface into flat rectangular areas separated by
rather inconspicuous grooves, but when constriction takes place there
is apparent a series of more or less convex annuli. The middle third
of the body, except for a ventral zone, about six muscle bands in width,
378 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
is closely covered with slender pointed papilliform outgrowths of the
cuticle, 1-1.5 mm. long, which give a furry appearance to skin.
These papillae communicate with irregularly zigzag subcutaneous
canals, above which the cuticle usually forms slight welts, which have
a direction oblique to the longitudinal axis of the dermal rectangles.
Each rectangle has its own canal, independent of the others (pl. 19,
figs. 4, 5). Beyond the papuliferous area these canals—or more
properly spaces, as they are usually branched—can be traced forward
half the distance to the head, and also posteriorly, as they are often
self-injected with yellowish material from the coelom. On the
periphery of the papuliferous area a papilla usually appears, first at
the anterior end of the canal, next at the posterior end, then in between,
until there are four or five to each rectangle. Brown or yellow, finely
divided material, which is sometimes loose in the canals, is also found
in the bottom of the papillae. If the top of the canal is stripped off, a
pore at either end is seen to lead deeper into the tissue (arrows in
pl. 19, fig. 4). If ordinary ink is forced from the coelomic side into the
pores that exist at intersection of longitudinal and transverse muscle
bands, it appears in these pores at the ends of the subcutaneous
canals but is usually blocked by material already in the canal. The
papillae are highly iridescent in sunlight. The area strongly reminds
one of the papularium of a sea star, and the function is probably the
same, i. e., respiratory.
The terminal knob of the body is very short, broadly rounded to
subtruncate, and the slight margin is capable of disappearing under
distension. ‘There is a conspicuous terminal pore, and the skin, either
smooth or longitudinally ridged, is closely beset with microscopic pores
of at least two sizes.
The short introvert is covered with squamiform papillae, which
increase in size toward the front, near which they decrease over a
narrow zone to the bare zone behind the tentacles. The largest
papillae are 0.75-1 mm. in length and breadth.
The tentacles are composed of very numerous small grooved foliate
elements in subtriangular mats or groups, radiating from the mouth,
which is ventral to the center. There are seven of these from which
ridges of tissue converge to the mouth, two dorsolateral, two lateral,
and two ventral, the odd one being the middorsal and much the
largest. On the periphery of the crown the space between the major
groups is filled in with one to three small groups of tentacles, which
probably increase in number as growth proceeds.
The anus is equivalent to about five muscle rings (not clearly
differentiated) behind the posterior papillae of the introvert.
Interiorly the longitudinal muscles form flat bands becoming
angular in section only when the body is much constricted. The
SIPUNCULID WORMS OF CALIFORNIA—FISHER 379
introvert and the four retractors occupy about one-seventh of the
body length. The retractors are free from one another and arise at
approximately the same level. Both ventrals arise from muscles 1-4,
while both dorsals arise obliquely from muscles 7-11. The two
protractors arise from muscles 12-15 at the posterior border of the
introvert. Before insertion, 4 mm. in front of the brain, they pass
over the dorsal retractors. Their form and position, with the introvert
out and in, are shown in plate 19, figures 1 and 2. The rectum passes
far forward and opens close behind the (dorsal) origin of the pro-
tractors. (Muscles 17 and 18 are the two middorsal in figure 1; 18
and 19 are really 17 and 16 of the left side.) The rectum lacks thin,
fan-shaped wing muscles.
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U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 22
SIPHONIDES RICKETTSI, NEW SPECIES.
SEE PAGE 443 FOR EXPLANATION.
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FORMS OF GOLFINGIA MARGARITACEA (SARS).
SEE PAGE 443 FOR EXPLANATION.
U.S. NATIONAL MUSEUM
PROCEEDINGS, VOL. 102 PLATE 24
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GOLFINGIA HESPERA (CHAMBERLIN) AND G. MARGARITACEA CALIFORNIENSIS, NEW
SUBSPECIES.
SEE PAGE 444 FOR EXPLANATION,
PROCEEDINGS, VOL. 102 PLATE 25
U. S. NATIONAL MUSEUM
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GOLFINGIA ELACHEA, NEW SPECIES, AND G. LAETMOPHILA, NEW SPECIES.
SEE PAGE 444 FOR EXPLANATION
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 26
AR NS EE RR TP ES a aR I ah w mae tC anamnad
GOLFINGIA MACGINITIEI, NEW SPECIES.
SEE PAGE 445 FOR EXPLANATION.
PROCEEDINGS, VOL. 102 PLATE 27
U.S. NATIONAL MUSEUM
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PERIMECES FISHER.
DENDROSTOMUM PYROIDES CHAMBERLIN AND D
SEE PAGE 445 FOR EXPLANATION.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 28
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DENDROSTOMUM PERIMECES FISHER AND D. PYROIDES CHAMBERLIN.
SEE PAGE 446 FOR EXPLANATION.
PROCEEDINGS, VOL. 102 PLATE 29
U. S. NATIONAL MUSEUM
DENDROSTOMUM PYROIDES CHAMBERLIN.
SEE PAGE 446 FOR EXPLANATION.
U.S. NATIONAL MUSEUM
PROCEEDINGS, VOL. 102 PLATE 30
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SPECIES OF DENDROSTOMUM.
SEE PAGE 447 FOR EXPLANATION.
PROCEEDINGS, VOL. 102 PLATE 31
U.S. NATIONAL MUSEUM
DENDROSTOMUM ZOSTERICOLUM CHAMBERLIN.
SEE PAGE 447 FOR EXPLANATION,
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 32
DENDROSTOMUM ZOSTERICOLUM CHAMBERLIN.
SEE PAGE 448 FOR EXPLANATION,
PROCEEDINGS, VOL. 102 PLATE 33
U.S. NATIONAL MUSEUM
DENDROSTOMUM PERIMECES FISHER.
SEE PAGE 448 FOR EXPLANATION,
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 34
DENDROSTOMUM DYSCRITUM, NEW SPECIES.
SEE PAGE 448 FOR EXPLANATION.
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 35
DENDROSTOMUM LISSUM, NEW SPECIES
SEE PAGE 448 FOR EXPLANATION,
PROCEEDINGS, VOL. 102 PLATE 36
U.S. NATIONAL MUSEUM
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PHASCOLOSOMA PUNTARENAE GRUBE AND PH. AGASSIZII KEFERSTEIN.
SEE PAGE 449 FOR EXPLANATION.
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 37
»
PHASCOLOSOMA PUNTARENAE GRUBE AND PH. AGASSIZII KEFERSTEIN.
SEE PAGE 449 FOR EXPLANATION.
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 38
PHASCOLOSOMA AGASSIZII KEFERSTEIN.
SEE PAGE 450 FOR EXPLANATION.
PROCEEDINGS, VOL. 102 PLATE 39
U. S. NATIONAL MUSEUM
SPECIES OF PHASCOLOSOMA.
SEE PAGE 450 FOR EXPLANATION.
>]
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
issued
SS —
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
No. 3307
Vol. 102
SCHIZOSTELLA, A NEW GENUS OF BRITTLE-STAR
(GORGONOCEPHALIDAE)
Washington ; 1952
By Austin H. Ciark
THanks 10 the enthusiastic interest of Count Pourtalés, Alexander
Agassiz, Prof. René Koehler, Dr. H. L. Clark, Dr. Th. Mortensen,
and others more extensive and intensive work has been done on the
echinoderm fauna of the Caribbean area than of any other part of
the tropical seas. Yet within the past few years a very considerable
number of new and surprising forms have come to light. During re-
cent dredging operations Frederick M. Bayer added another to this
growing list of unexpected new types, a new genus of Gorgonocephali-
dae, which may be known as—
SCHIZOSTELLA, new genus
Diagnosis —A genus of Gorgonocephalidae, subfamily Astrocheli-
nae, resembling Asteroporpa and Astrocnida but with seven rays, the
arms bifurcate at about the middle, the double rows of hook-bearing
granules with the bordering larger granules forming bands as broad
as the distance between them, and reproducing by fission.
Genotype.—Schizostella bifurcata, new species.
Notes —The subfamily Astrochelinae of the family Gorgonocephali-
dae, distinguished from the subfamily Gorgonocephalinae by the ab-
sence of interradial accessory plates, includes seven genera of which
five, Astrochele, Astrogomphus, Asteroporpa, Astrothorax, and Astro-
thrombus, have undivided arms, while in two, Astrocnida and Astro-
clon, the arms branch at the tip. The genera Astrogomphus, As-
teroporpa, and Astrocnida are represented in the Caribbean region,
the two first also in the west Pacific.
451
969587—52
452 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Schizostella seems to be most closely related to Asteroporpa, espe-
cially to the group of species including lindneri of the Caribbean,
australiensis of Australia, and hadracantha of Japan in which the
double bands of hook-bearing tubercles on the arms are bordered on
each side by a regular row of contiguous enlarged granules, the quad-
ruple bands composed of enlarged and hook-bearing granules are only
slightly elevated above the general surface of the arms, and the hooks
are small, usually without an accessory tooth. In spite of the forked
arms it is less closely related to Astrocnida, and much less closely to
Astroclon.
In several ophiurans in different families the young have six or
sometimes seven arms and reproduce by fission, giving rise to adults
with usually five arms. But Schizostella cannot be the young of any
known West Indian species as it differs from all related species in
details of its structure and ornamentation.
Although a number of species in several genera of the Trichasteridae
are known to reproduce by fission when young, this is the first case
to be reported in the Gorgonocephalidae.
SCHIZOSTELLA BIFURCATA, new species
PLATE 40
Description.—The disk is 4 mm. in diameter and is composed of
seven triangular segments elevated in the middle and with rounded
outer angles which are separated from each other by rather deep
grooves, becoming deeper outwardly and ending at a deep notch in
the interradial border. The seven segments each contain two radial
ribs, but appear as single units, seldom giving any indication of a
double structure.
The disk is densely covered with hemispherical granules, the smaller
brownish and the larger white, which are irregularly arranged except
at the distal ends of the segments, where there are two regular rows
of white tubercles separated by two rows of hook-bearing tubercles.
The seven arms are at the base very nearly as broad as the distal
width of the disk segments. They are roughly twice as long as the
diameter of the disk and bifurcate at about the middle, on or about
the thirteenth brachial.
The ornamentation of the arms resembles that of the outer part
of the disk segments. There are evenly spaced transverse rows of
white hemispherical granules between which are double rows of hook-
bearing granules alternating with, at the base of the arms, irregular
double rows of larger, lower, polygonal granules which farther out
become single irregular rows and disappear almost completely at the
arm tips. The hooks are minute, very stout, thick-crescentic, without
an accessory tooth, and become very prominent on the slender arm
tips.
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 40
SCHIZOSTELLA BIFURCATA, NEW GENUS, NEW SPECIES.
The topmost figure (both sides) is the type, U.S.N.M. No. E. 7875; the four lower figures
are two cotypes. XX 3.
A NEW GENUS OF BRITTLE-STAR—-CLARK 453
The oral surface is studded with large white granules resembling
those on the aboral surface, but widely scattered.
The second tentacle pore has a double arm spine. Farther out there
seem to be three arm spines arising from the same base. Distally these
become transformed into hooks.
The genital openings, two in each interradius, are very large.
The color is pinkish brown, the under surface and the large tubercles
yellowish white.
Type—U.S.N.M. No. E. 7875, from off Triumph Reef, Elliott Key,
Fla.; 20-25 fathoms; collected by Frederick M. Bayer, November
28, 1949.
Notes.—Three additional specimens were collected at the same sta-
tion. ‘The type and two others were on Swiftia exserta (Ellis and
Solander) and one was on Filigorgia, sp.
The type and the specimen from Filigorgia are symmetrical with
seven rays each. The two others have two full-sized rays and two
much smaller, evidently the result of fission.
For comparison with these specimens there are at hand four exam-
ples of the ophiocrenoid stage of one of the species of Gorgonocepha-
linae. These have five arms which divide once, in some cases with
the rudiments of a second forking at the tips of the secondary arms,
and the interradial areas of the disk are deeply concave. They were
dredged at Albatross station 3729, off Omai Zaki Light, Japan, in
34 fathoms and were clinging to Afelitodes, sp. (?nodosa). They re-
semble in general the specimens described by Matsumoto (Journ.
Coll. Sci., Tokyo Imper. Univ., vol. 38, art. 2, pp. 68, 69, fig. 18, p. 69,
March 31, 1917) from Sagami Bay in 300 fathoms.
With the four specimens from Albatross station 3729 were dredged
two specimens of Astrocladus dofleini Déderlein (recorded by H. L.
Clark, U.S. Nat. Mus. Bull. 75, p. 298, 1911, as Astrophyton pardalis)
of which they are presumably the young.
Matsumoto said that his ophiocrenoids may be young either of
Astroboa or Astrochalcis. They are from deeper water than the
maximum recorded for any of the five species of Astrocladus, all of
which occur in the sublittoral zone.
While studying the specimens of Muricea collected by the Albatross
in the West Indies Mr. Bayer found a second species of Schizostella
which may be known as
SCHIZOSTELLA BAYERI, new species
Description.—Resembling S. bifurcata and, like it, with seven
bifurcate arms; but the white granules on the disk, instead of being
scattered, are contiguous and arranged in regular lines to form a con-
spicuous, white, close reticulation, while those on the arms are con-
454 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
tiguous instead of slightly separated, forming regular narrow trans-
verse lines across the arms.
Type—U.S.N.M. No. E. 7997, from Albatross station 2188, southeast
of Jamaica (lat. 17°44’05”’ N., long. 75°39’00’" W.) ; 23 fathoms; coral
and broken shells; February 29, 1884.
Notes.—The type and two additional specimens (KE. 7998) were
found on Muricea pendula Verrill.
U.S, GOVERNMENT PRINTING OFFICE; 1952
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
Issued by the
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 102 | Washington: 1952 No. 3308
MOTHS OF THE GENUS EPEIROMULONA, A NEW GENUS
OF LEPIDOPTERA
By Wiuu1am D. Frevp
THe GENUs treated in this paper (Arctiidae, Lithosiinae) has for-
merly been represented by a single species originally described as
phelina by Druce and doubtfully referred by him to the genus Avto-
ceras Felder in the text of his description? and placed in the genus
Melania Wallengren on the plate accompanying this text.2 Subse-
quently Hampson ® placed it in /ulona Walker and Forbes,‘ recog-
nizing that it did not belong with Afudona, transferred it tentatively to
the genus Gaudeator Dyar. In reality it belongs to quite a distinct
and new genus.
A study of the 75 specimens from the collections of the United States
National Museum, the British Museum (Natural History), and Cor-
nell University discloses the fact that the genus contains 7 species (one
of these with 3 subspecies), all but 1 being new.
EPEIROMULONA, new genus
Type.—E peiromulona lephina, new species.
Labial palpus upturned, short, barely reaching above lower margin
of frons.
Antennae of male and female filiform and gradually more slender
to the tip; each subsegment heavily setose underneath. Each sub-
segment with a pair of bristles arising from middle of ventrolateral
1 Biologia Centrali-Americana, Insecta, Lepidoptera-Heterocera, vol. 1, p. 139, 1885.
2 Ibid., vol. 3, pl. 13, fig. 13, 1885
3 Catalogue of the Lepidoptera Phalaenae in the British Museum, yol. 2, pp. 386-387,
1900.
4 Bull. Mus. Comp. Zool., vol. 85, p. 1838, 1939.
455
969585-—52
456 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
margin and with a second pair of bristles arising from just above
middle of dorsolateral margin of subsegment and with a single sen-
sory cone arising from near the middle of front margin.
Venation of forewing with vein 2 from outer fourth of cell, nearly
straight, not downward curved at base; vein 3 separate from just
before lower angle of cell (3 very rarely stalked with 4); 4 from
lower angle and 4 and 5 separate; 6 from below upper angle of cell;
9 from stalk of 7 and 8; 10 and 11 free; 10 from cell about halfway
between stem of 7, 8, and 9 and base of 11; 11 from outer third of
cell and curved distally toward 12.
Venation of hindwing with 2 from outer fourth of cell; 3 stalked
with 4 from lower angle of cell; 5 separate and from just below middle
of cell; 6 and 7 stalked from upper angle of cell; 8 from about middle
or slightly after middle of cell.
Male genitalia (see all figures on pls. 41-48 and figs. 13-17 on pl. 45)
with uncus hooklike, broad at base and sharply constricted and curved
distally; gnathos absent; anellus present (dorsal plates absent) and
represented by a strongly sclerotized, nearly H-shaped juxta; vinculum
broadly U-shaped; inner surface of harpe with downward-projecting
lobe from near middle (this lobe sometimes fingerlike or hooklike)
except in icterinus,; saccus very small; aedeagus broader posteriorly
than anteriorly and bent or at least curved near middle; vesica armed
with numerous spinelike cornuti; bulbus ejaculatorius from about
anterior fourth or fifth of aedeagus.
Female genitalia (see all figures on pl. 44 and figs. 11 and 12 on
pl. 45) with pouch between posterior margin of seventh abdominal
segment and anterior margin of eighth abdominal segment; seventh
abdominal segment sometimes strongly sclerotized above this pouch,
sometimes so much so that the pouch is more like a lip than a pouch;
strongly sclerotized species with a pair of lateral lateroventral cup-
like shallow pouches on seventh segment, lying on each side of raised
central area; eighth segment not at all or only weakly sclerotized ven-
trally, never forming a solid ventral plate, but with a plate that is
broken through the middle; ostium bursae before middle of eighth
segment; ductus bursae heavily sclerotized, broad at ostium bursae and
somewhat twisted and narrowed at bursa copulatrix; entrance to bursa
copulatrix armed with a furcate plate, this plate inwardly covered
with short spines; bursa copulatrix with ductus seminalis from right
side or slightly dorsal and near ductus bursae; bursa copulatrix sub-
spherical, slightly bent and extended to the left and with a ribbonlike
scobinate plate at anterior end.
Comparatiwe remarks.—This genus is somewhat related to Mulona
Walker and Gaudeator Dyar. Vein 2 of forewing is nearly straight,
not at all downward curved at base in Gaudeator and E’peiromulona,
MOTHS OF GENUS EPEIROMULONA—FIELD 457
whereas in Mu/lona vein 2 is downward curved at base. Vein 5 of hind-
wing is present in Gaudeator and E'peiromulona, absent in Mulona.
Veins 3 and 4 of hindwing are usually connate or separate in Gaudea-
tor, stalked in Mulona and E'peiromulona. Epeiromulona is easily
distinguished by the male and female genitalia from the other two
genera. In males of /u/ona the juxta is bifurcate and U- or V-shaped.
In Epetromulona the juxta is H-shaped and in Gudeator a broad,
slightly curved plate. The transtilla is absent in H’peiromulona and
present in Mulona and Gaudeator. The aedeagus is short and broad in
Mulona, short and not so broad in Gaudeator, and is rather long in
Epeiromulona. Numerous cornuti are present on the vesica in Gaude-
ator and E’peiromulona and absent in Mulona. In females of Mulona
the middle two-thirds or more of bursa copulatrix is heavily serra-
tulate while Gaudeator and E'petromulona females are not serratulate
but contain instead two areas of armature, one each at anterior and
posterior end of bursa copulatrix. The anterior armature in E’petro-
mulona consists of a ribbonlike scobinate plate and in Gaudeator this
armature consists of a small, nearly round scobinate plate. The
posterior armature in E'petromulona consists of a bifurcate plate
covered with short spines and in Gaudeator this armature consists of
a collar covered with long spines, this collar almost half encircling
bursa copulatrix near entrance of ductus bursae.
KEY TO THE SPECIES OF EPEIROMULONA
1::, ‘Vertex, of head onange.l Sse) es icterinus, new species (p. 467)
Wertexcofrheagn whit eis a2 has P= 5 ee. ee ee Poe ee Pe
>. Hindwings, salmon pink ______-___— roseata, new species (p. 465)
Ena win eSSyellOwAOLKOLaANS Cea ose ee tee ee cr ee ee 2?
a Legs: yellow with black bands22))2_ 80 Olt uke UN Si ee ae 4
Legs white with black bands__~______2) 42 biloba, new species (p. 460)
4. Fringe of forewing entirely orange or pale orange-yellow; apex of first
LEMETEY CllG Wee es ee ee eee es ee oe ne ee sue 5
Fringe opposite middle of outer margin of forewing black; apex of first
ETI TaD 8 C Keer pee ce yea 2 oe eat er thysanata, new species (p. 466)
Co EU OS ee er a TS ene es Ss ad AEE a aA 5 6
HCTIATE Sit. = x LATTE aR FAs tT) CAEL ED oer ee Fe gyre! 9
_ 6. Upper elements of H-shaped juxta distinctly shorter than bottom ele-
mentsy (pl. 41g. lh) Seo ree lephina, new species (p. 459)
Upper elements of H-shaped juxta as long as or slightly longer than
bottomselements™ (pl 42) tigi) 3) ee ee ee en eee {s
7. Distal end of harpe narrow, much narrower than base_______________ 8
Distal end of harpe as broad or nearly as broad as base (pl. 45,
1 -ees W@ eee ee See Ae ee ee 2 hamata colombiensis, new subspecies (p. 464)
8. Harpe with costa not at all or only slightly expanded at apex (pl. 42,
fig. 3a; pl. 45, figs. 18, 14).
hamata hamata, new species and subspecies (p. 462)
Harpe with costa well expanded at apex (pl. 45, figs. 15, 16).
hamata venezuelensis, new subspecies (p. 463)
458 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
9. Posterior margin of seventh abdominal segment below, with three dis-
tinet lobes: .(pliv44. Wigs sO edi O)e oe he et ae ee eee a ee ee 11
Posterior margin of seventh abdominal segment below without these
1c) =) pmena RE pa on te pee TUIN rie a ENA elon nae Raph ey see yn ng oe L 10
10. Shallow cuplike pouches on seventh abdominal segment extending well
into ventral surface (pl. 44; fig. 7)---------- phelina (Druce) (p. 458)
Shallow cuplike pouches on seventh abdominal segment lateral in posi-
tions (plec445 hie! 8) ee es lephina, new species (p. 459)
11. Lobes on each side of mesial lobe of posterior margin of seventh
abdominal segment small (pl. 44, fig. 9).
hamata brasiliensis, new subspecies (p. 465)
These lobes large, nearly as large as mesial lobe (pl. 44, fig. 10).
hamata hamata, new species and subspecies (p. 462)
EPEIROMULONA PHELINA (Druce)
PLate 44, Ficure 7; PLate 46, Fraure 22
Autoceras (?) phelina Druce, Biologia Centrali-Americana, Insecta, Lepidop-
tera-Heterocera, vol. 1, p. 139, 1885.
Melania phelina (Druce), Biologia Centrali-Americana, Insecta, Lepidoptera-
Heterocera, vol. 3, pl. 13, fig. 18, 1885.
Cincia phelina (Druce), Kirsy, A synonymic catalogue of Lepidoptera Hetero-
cera, p. 366, 1892.
Mulona phelina (Druce), Hampson, Catalogue of the Lepidoptera Phalaenae
in the British Museum, vol. 2, pp. 386, 3887 (2), 1900.—Drauvpt, in Seitz,
Gross-schmetterlinge der Erde, vol. 6, p. 252, pl. 34, fig. ec 9, 1918.—Srranp,
in Wagner, Lepidopterorum catalogus, pars 26, p. 732, 1922.
Male——Unknown.
Female.—(pl. 46, fig. 22) —Palpus, head, base of antenna, patagium,
tegula, pronotum, and mesonotum pale dirty orange-yellow. Except
for palpi this color may be due to a stain and may have originally
been white (in the single known specimen of this species). Metano-
tum and thorax underneath pale orange-yellow. Antenna beyond
base black with a long white streak just before apex. There is a black
spot in middle of thorax and another on base of tegula. Foreleg pale
orange-yellow with most of tibia and last four tarsal subsegments
black. First tarsal subsegment nearly white. Hindleg pale orange-
yellow (midlegs missing in the single known specimen). Wings white,
spotted with 15 black spots and bars, these arranged as in £. bdloba.
Fringe of forewing pale yellowish orange.
Length of forewing 9 mm.
Female genitalia as illustrated (pl. 44, fig. 7) and with characters
as given in the key.
Type locality and distribution —V olean de Chiriqui,” Panama.
Additional type data——Described from a single specimen, the holo-
type female (elevation 2,000 to 3,000 feet; Champion; ¢ genitalia
preparation, British Museau No. 1951-22).
Location of type.—In the British Museum (Natural History).
MOTHS OF GENUS EPEIROMULONA—FIELD 459
Remarks.—The photographs of the type (pl. 46, fig. 22) and of the
genitalia of the type (pl. 44, fig. 7) were furnished through the cour-
tesy of D. S. Fletcher, of the British Museum (Natural History),
Department of Entomology.
EPEIROMULONA LEPHINA, new species
PLate 41, Fiacures 1, la, 1b; PLare 44, Ficure 8; Puate 46, Ficures 18-20
Mulona phelina (Druce), Dyar nee Druce, in part., Proc. U. 8. Nat. Mus., vol.
47, p. 167, 1914.
Gaudeator (?) phelina (Druce), Forses nee Druce, in part., Bull. Mus. Comp.
Zool., vol. 84, No. 4, p. 163 (on chart), p. 188, August 1939.
Male.—(pl. 46, figs. 18, 20).—Palpus pale orange-yellow. Head,
base of antenna, patagium, tegula, pronotum, and mesonotum white.
Metanotum and thorax underneath pale orange-yellow. Antenna be-
yond base black with a long white streak just before apex. A black
spot in middle of thorax and on base of tegula (this last spot usually
hidden by patagium). Legs pale orange-yellow with foretibia, apex
of midtibia and hindtibia and last four tarsal subsegments of all legs
black; apex of forefemur yellow. Abdomen orange-yellow, pale yel-
lowish white at base above. Forewings and hindwings above and
below very similar to béloba with fringe of forewing slightly paler
orange-yellow near the margin.
Length of forewing 6.5-7 mm.
Male genitalia as illustrated (pl. 41, figs. 1, la, 1b) and with char-
acters as given in the key.
Female (pl. 46, fig. 19) —Habitus like that of the male.
Length of forewing 7-8 mm.
Female genitalia as illustrated (pl. 44, fig. 8) and with characters
as given in the key.
Type locality—Porto Bello, Col6n Province, Panama.
Additional type data.—Described from the holotype, male (locality
as given above; U.S.N.M. type No. 34839; August Busck; March 1912;
g genitalia preparation W. D. F. No. 3191, 1949) ; allotype, female
(same data as holotype; 2 genitalia preparation W. D. F. No. 3218,
1949) ; paratypes Nos. 1-5, two males and three females (Rio Trini-
dad, Panama; March, May, June 1911, 1912; August Busck. W. D. F.
genitalia preparations as follows: Paratype No. 1, ¢, No. 3190, 1949;
paratype No. 2, 3, No. 3192, 1949; paratype No. 3, 9, No. 3212, 1949;
paratype No.4, 2, No. 1666, 1941; paratype No. 5, 9, No. 3211, 1949) ;
paratype No.6, ¢ (La Chorrera, Panama; May 1912; August Busck;
3 genitalia preparation W. D. F. No. 1675, 1941) ; paratypes Nos.
7-9, males (Cayuga, Guatemala; Schaus and Barnes; March, May,
August. W.D.F., ¢ genitalia preparations as follows: Paratype No.
7, No. 3187, 1949; paratype No. 8, No. 1672, 1941; Paratype No. 9, No.
460 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
3239, 1949) and paratypes Nos. 10-18, females (Barro Colorado
Island, Panama Canal Zone; March, November 1934, 1940, 1941;
James Zetek and Marston Bates. W.D.F., ¢ genitalia preparations
as follows: Paratype No. 10, No. 3213, 1949; paratype No. 11, No.
3214, 1949; paratype No. 12, No. 3226, 1949; paratype No. 13, No.
3244, 1949).
Locations of types.—Holotype, allotype, and paratypes Nos. 1, 3, 4,
5, 6, 7, 18 in the United States National Museum. Paratypes Nos. 8,
12 in the collection of Cornell University. Paratypes Nos. 9, 11 in the
British Museum (Natural History). Paratypes Nos. 2, 10 in the
American Museum of Natural History.
Distribution—Panama: Province of Colén, Porto Bello (May) ;
Rio Trinidad (March, May, and June); Province of Panama, La
Chorrera (May). Canat Zone: Barro Colorado Island (March,
April, November). Guatemata: Department of Izabal, Cayuga
(May).
Remarks.—Fifteen specimens (including the genitala preparations
of all these) were examined.
Comparative remarks.—This species differs in habitus from F.
biloba in having the underside of thorax and the legs colored pale
orange-yellow with black bands on the legs, and in having the femur of
foreleg pale orange-yellow. In d¢loba the underside of thorax and
the legs are white with black bands on the legs and with the apical
half of the femur of foreleg black.
EPEIROMULONA BILOBA, new species
PLATE 41, FIGURE 2, 2a, 2b; PLATE 46, FIGURE 21
Mulona phelina (Druce), Dyar nee Druce, in part., Proc. U. S. Nat. Mus., vol. 47,
p. 167, 1914.
Male (pl. 46, fig. 21).—Palpus, head, base of antenna, patagium,
tegula, and thorax (above and below) white. A black spot in middle
of thorax and another at the base of tegula (this spot usually hidden
by patagium). Antenna beyond base black with a long white streak
just before apex. Forelegs white or very pale yellowish white with
black on apical half of femur, basal half of tibia, and the last four sub-
segments of tarsus. Midlegs and hindlegs white with apex of tibia
and last four subsegments of tarsus black. Abdomen orange-yellow
except at base above, where it is yellowish white. Forewing above
white with 15 small black spots and bars. Five black bars along
costal margin, the first near base, the second opposite the beginning
of the last one-third of cell, the fourth just beyond end of cell and
C-shaped and the fifth subapical and nearly touching the lower ele-
MOTHS OF GENUS EPEIROMULONA—FIELD 461
ment of the C-shaped bar. Four spots lie on lower margin of cell and
on vein 2, the first on lower margin of cell just before middle, the sec-
ond at point of origin of vein 2, the third in the middle of vein 2, and
the fourth just beyond this. There is a series of four spots along
vein 1, the first near base of vein 1, the second near middle of this
vein, the fourth at the margin, and the third between the second and
fourth. There are two spots opposite end of cell on vein 4, one in
the middle of this vein and the other beyond this near outer margin
of wing. Fringe of outer margin of forewing bicolored, orange near
margin and pale orange-yellow beyond this. Hindwing above and
below, including fringe, pale orange-yellow with an apical black spot
on costal margin at apex. Forewing below, including fringe, orange-
yellow, usually with a pink tinge on the cell and sometimes also on
the surrounding interspaces in middle of wing, and with a black spot
on middle of costal margin, a black streak extending from base of
wing outward, sometimes to the black spot, a U-shaped black sub-
apical marking on costal margin and a small black spot at end of
vein 4 on the margin (this black spot not extending into the fringe
opposite).
Length of forewing 6.5-7 mm.
Male genitalia as illustrated (pl. 41, figs. 2, 2a, 2b).
Female.—Unknown.
Type locality—Rio Trinidad (labeled Trinidad River), Colon,
Province, Panama,
Additional type data.—Described from the holotype, male (locality
as given above; June 1912; August Buseck; U.S.N.M. type No. 34808;
é genitalia preparation W. D. F. No. 3189, 1949) ; paratypes Nos. 1-3,
males (same locality and data as holotype except collected in March
and June; ¢ genitalia preparation W. D. F. No. 3188, 1949 for para-
type No. 1; ¢ genitalia preparation W. D. F. No. 1665, 1941 for
paratype No. 2; ¢ genitalia preparation W. D. F. No. 3186, 1949 for
paratype No. 3) and paratypes Nos. 4 and 5, males (Muzo, Colombia;
400-800 meters; Fassl; ¢ genitalia preparation W. D. F. No. 3235,
1949 for paratype No. 4; ¢ genitalia preparation W. D. F. No. 3284,
1949 for paratype No. 5).
Location of types—Holotype and paratypes Nos. 1 and 2 in the
United States National Museum. Paratype No. 3 in the entomological
collection of Cornell University. Paratypes Nos. 4 and 5 in the British
Museum (Natural History).
Distribution.—Panama: Province of Colén, Rio Trinidad (March,
June). Corompr1a: Department of Boyacd, Muzo.
Remarks.—Six specimens (including their genitalia preparations)
studied.
462 PROCEEDINGS OF THE NATIONAL MUSEUM Von. 102
EPEIROMULONA HAMATA, new species
Male (pl. 46, figs. 23, 25, 26).—Palpus pale orange-yellow. Head,
base of antenna, patagium, tegula, pronotum, and mesonotum white.
A black spot in middle of thorax and another on base of tegula (usu-
ally covered by patagium). Metanotum and thorax underneath pale
orange-yellow (or sometimes whitish yellow). Legs pale orange-
yellow (sometimes whitish yellow) and marked with black as in EF,
biloba, except that apex of first femur is yellow, not black. Abdomen
pale orange-yellow, slightly whitish at base above. Antenna beyond
base black with a long white streak just before apex. Forewings and
hindwings above and below very similar to #. phelina and hardly
separable from £. biloba. Apical spot on hindwing variable, some-
times greatly reduced or absent and sometimes enlarged.
Length of forewing 6.5—8 mm.
Male genitalia as illustrated (pl. 42, figs. 3, 3a, 3b; pl. 45, figs.
13-17). This species divides into three subspecies on the basis of ap-
parently constant differences in the harpes (see descriptions, pp.
462-464).
Female (pl. 46, figs. 24, 27) —Habitus like that of the male.
Length of forewing 7.2-8.2 mm.
Female genitalia as illustrated (pl. 44, figs. 9,10). On the basis of
the female genitalia there is a fourth subspecies (see description, p.
465).
Distribution.—Trinidad, northern South America and Brazil.
Comparative remarks.—In addition to the great differences in the
genitalia this species differs from ZF. biloba in lacking the black on
the femur of foreleg and in having the legs yellow with black bands
instead of white with black bands. It is hardly distinguishable from
EB. lephina and E. phelina except in the genitalia. However, the
black bars in middle and base of costa on forewing above are usually
thicker and more round or subquadrate than in phelina or biloba.
Subspeciation—As stated above this species divides into four sub-
species on the basis of the male and female genitalia. These sub-
species appear to be geographically isolated. They do not appear to
differ in color and habitus. When more material becomes available
for study so that the extent of individual variation can be worked
out it is possible that these subspecies will be elevated to specific rank.
EPEIROMULONA HAMATA HAMATA, new subspecies
Puate 42, Ficures 3, 3a, 3b; PLATE 44, FIGURE 10; Puate 45, Ficures 138, 14;
PLATE 46, FIGURES 23, 24
Male (pl. 46, fig. 23).—Palpus, head, thorax, abdomen, and wings
as described above.
Length of forewing 7.5-8 mm.
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102, PLATE 41
Pr
ate
1-lb, Epeiromulona lephina, new genus and species: 1, Male genitalia; la, harpe; 1b,
aedeagus; male genitalia preparation No. W.D.F. 3191, 1949; holotype. 2-2b,
E. biloba, new species: 2, Male genitalia; 2a, harpe; 2b, aedeagus; male genitalia
preparation No. W.D.F. 3189, 1949; holotype.
US. NATIONAL MUSEUM PROCEEDINGS, VOL. 102, PLATE 42
3-3b, Epeiromulona hamata hamata, new species and subspecies: 3, Male genitalia; 3a,
harpe; 3b, aedeagus; male genitalia preparation No. W.D.F. 1667, 1941; holotype.
4-4b, E. thysanata, new species: 4, Male genitalia; 4a, harpe; 4b, aedeagus; male
genitalia preparation No. W.D.F. 1670, 1941; holotype.
U.S NATIONAL MUSEUM PROCEEDINGS, VOL. 102, PLATE 43
5-5b, Epeiromulona roseata, new species: 5, Male genitalia; 5a, harpe; 5b, aedeagus; male
genitalia preparation No. W.D.F. 1674, 1941; holotype. 6-6b, E£. tcterinus, new
species: 6, Male genitalia; 6a, harpe; 6b, aedeagus; male genitalia preparation
No. W.D.F. 1673, 1941; holotype.
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102, PLATE 44
4
oe
7, Epeiromulona phelina (Druce): Female genitalia; British Museum genitalia preparation
No. 1951-22; holotype. 8, E. lephina, new species: Female genitalia; genitalia prepara-
tion No. W.D.F. 3218, 1949; allotype. 9, E. hamata brasiliensis, new subspecies:
Female genitalia; W.D.F. genitalia preparation No. 3222, 1949; holotype. 10, £.
hamata hamata, new subspecies: Female genitalia; W.D.F. genitalia preparation No.
1677, 1949; allotype.
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102 PLATE 45
11, Epetromulona thysanata, new species: Female genitalia; genitalia preparation No.
W.D.F. 1678, 1941; allotype. 12, E. icterinus, new species: Female genitalia; geni-
talia preparation No. W.D.F. 3217, 1949; allotype. 13, FE. hamata hamata, new
subspecies: Left harpe; male genitalia preparation No. W.D.F. 3208, 1949; paratype
No. 1. 14, £. A. hamata, new subspecies: Left harpe; male genitalia preparation No.
W.D.F. 3227, 1949; paratype No. 5. 15, E. hamata venezuelensis, new subspecies:
Left harpe; male genitalia preparation No. W.D.F. 3231, 1949; holotype. 16, E.
hamata venezuelensis, new subspecies: Left harpe; male genitalia preparation No.
W.D.F. 3233, 1949; paratype No. 2. 17, E. hamata colombiensis, new subspecies:
Left harpe; male genitalia preparation No. W.D.F. 3236, 1949; holotype.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 102, PLATE 46
-
18, Epeiromulona lephina, new species, holotype, male. 19, E. lephina, new species, allo-
type, female. 20, E. lephina, new species, underside of paratype No. 7, male. 21. E.
biloba, new species, holotype, male. 22. E. phelina (Druce), holotype, female. 23,
E. hamata hamata, new species and subspecies, holotype, male. 24, E. h. hamata,
new species and subspecies, allotype, female. 25, E. hamata venexuelensis, new
subspecies, holotype, male. 26, E. hamata colombiensis, new subspecies, holotype,
male. 27, E. hamata brasiliensis, new subspecies, holotype, female. 28, E£. roseata,
new species; holotype, male. 29, E£. icterinus, new species, holotype, male. 30, £.
thysanata, new species, holotype, male. 31, E&. thysanata, new species, allotype,
female. (All figures twice natural size.)
MOTHS OF GENUS EPEIROMULONA—FIELD 463
Male genitalia as illustrated (pl. 42, figs. 3, 3a, 3b; pl. 45, figs. 13,
14). This subspecies differs from the others in having the distal end
of harpe narrower and more elongate, with costa not greatly expanded
at apex. In this subspecies the downward-projecting, hooklike struc-
ture from middle of costa is usually greatly recurved and elongated
at tip and the ampullalike spiny knob is located much closer to the
ventral margin than in the other subspecies.
Female (pl. 46, fig. 24). —Habitus like that of the male.
Length of forewing 7.2-8 mm.
Female genitalia as illustrated (pl. 44, fig. 10), differing from /.
hamata brasiliensis in having the apex of ductus bursae slightly
broader and with broad lobes along posterior edge of seventh sternum
above shallow cuplike depressions that lie on either side of the central
raised transverse ridge.
Type locality——Cayenne, French Guiana.
Additional type data.—Described from the holotype, male (locality
as given above; William Schaus; U.S.N.M. type No. 34818; ¢ genitalia
preparation W. D. F. No. 1667, 1941); allotype, female (Trinidad,
West Indies; August Busck; 9 genitalia preparation W. D. F. No.
1677, 1941) and from two male and three female paratypes (various
localities: Cayenne, French Guiana; Trinidad, West Indies and
Caparo, west-central Trinidad, West Indies. Genitalia preparations
as follows: Paratype No. 1, ¢, W. D. F. No. 3208, 1949; paratype No.
2, 29, W. D. F. No. 3215, 1949; paratype No. 3, 9, W. D. F. No. 3216,
1949; paratype No. 4, 2, W. D. F. No. 3221, 1949 and paratype No. 5,
3, W. D. F. No. 3227, 1949).
Location of types.—Holotype, allotype, and paratypes Nos. 1-3 in
the United States National Museum. Paratypes Nos. 4 and 5 in the
British Museum (Natural History).
Distribution —Frencw Guiana: Cayenne. Trrnmap: Capara in
west-central Trinidad; several specimens are labeled simply “Trini-
dad.”
Remarks.—Six specimens (including their genitalia preparations)
were studied.
EPEIROMULONA HAMATA VENEZUELENSIS, new subspecies
PuaTE 45, Fiacures 15, 16; PLate 46, Ficure 25
Male (pl. 46, fig. 25).—Palpus, head, thorax, abdomen, and wings
not different from typical Z’. hamata.
Length of forewing 6.5-7.2 mm.
Male genitalia as illustrated (pl. 45, figs. 15, 16). This subspecies
differs from typical hamata in having the distal end of harpe elongate
at ventral margin and with costa expanded at apex. The downward-
projecting hooklike structure originates from just before middle of
costa and is recurved and elongate at the tip but not as recurved and
464 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102:
elongate as in F. hamata hamata. The ampullalike spiny knob is
located almost midway between the costal and ventral margins.
Female.—Unknown.
Type locality and distribution.—Las Quiguas, Esteban Valley, Ven-
ezuela.
Additional type data.—Described from the holotype, male (locality
as given above; é genitalia preparation W. D. F. No. 3231, 1949);
paratype No. 1, male (same locality; ¢ genitalia preparation W. D. F.
No. 3232, 1949) and paratype No. 2, male (same locality; ¢ genitalia
preparation W. D. F. No. 32383, 1949).
Location of types.—Holotype and paratype No. 1 in the British
Museum (Natural History). Paratype No. 2 in the United States
National Museum.
Remarks.—Three specimens (including their genitalia prepara-
tions) were studied.
EPEIROMULONA HAMATA COLOMBIENSIS, new subspecies
PLATE 45, FIGURE 17; PLATE 46, FIGURE 26
Male (pl. 46, fig. 26).—Palpus, head, thorax, abdomen, and wings
not different from the other subspecies.
Length of forewing 7.5-8 mm.
Male genitalia with harpe as illustrated (pl. 45, fig. 17). #.h. colom-
biensts differs from the other subspecies in having the distal end of
harpe very much broader, only slightly elongated at ventral margin.
Costa is more greatly expanded at apex than in £. h. venezuelensis.
The downward-projecting hooklike structure arises from just before
middle of costa and is broad and short, only slightly recurved and
elongated at tip. The ampullalike spiny knob is located below middle
of harpe but is not nearly so close to ventral margin as it is in
FE. hamata hamata. 'The ventral margin of harpe is greatly expanded
just below this ampullalike structure.
Female.—Unknown.
Type locality and distribution—Upper Rio Negro, Colombia.
Additional type data.—Described from the holotype, male (locality
as given above; 800 meters; collection Fassl; ¢ genitalia preparation
W. D. F. No. 3236, 1949) and paratype No. 1, male (same locality
and data; ¢ genitalia preparation W. D. F. No. 3237, 1949) and
paratype No. 2, male (same locality and data; ¢ genitalia preparation
W. D. F. No. 3288, 1949).
Location of types.—Holotype and paratype No. 1 in the British
Museum (Natural History). Paratype No. 2 in the United States
National Museum.
Remarks.—Three specimens (including their genitalia prepara-
tions) were studied.
MOTHS OF GENUS EPEIROMULONA—FIELD 465
EPEIROMULONA HAMATA BRASILIENSIS, new subspecies
Puate 44, Figure 9; PLATE 46, FIGURE 27
Male——Unknown.
Female (pl. 46, fig. 27).—Palpus, head, thorax, abdomen, and
wings not different from typical hamata.
Length of forewing 7 mm.
Female genitalia as illustrated (pl. 44, fig. 9) with apex of ductus
bursae slightly narrower than in 2. hamata hamata and with narrow
lobes projecting from posterior edge of seventh sternum above shallow
cuplike depressions that lie on each side of the central raised trans-
verse ridge.
Type locality and distribution —Melguina, 10 miles south of Dia-
mantino, Mato Grosso, Brazil.
Additional type data.—Described from a single specimen, the holo-
type, female (locality as given above; 2,000 feet; C. L. Collenette; ¢
genitalia preparation W. D. F. No. 3222, 1949).
Location of type.—In the British Museum of Natural History.
EPEIROMULONA ROSEATA, new species
PLATE 43, Fiaures 5, 5a, 5b; PLATE 46, Faure 28
Male (pl. 46, fig. 28).—Palpus, head, base of antenna, patagium,
tegula, pronotum and mesonotum white.
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
f SAF ions NN
issued
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Washington : 1952 No. 3310
Vol. 102
REVIEW OF THE FISHES OF THE BLENNIOID GENUS
ECSENIUS, WITH DESCRIPTIONS OF FIVE NEW
SPECIES
By Witzert M. Cuapman! and Lronarp P. Scuurrz
McCottocu described the fish genus #’csenius on the basis of a new
species, #. mandibularis, from Australian waters. Norman further
defined the genus and indicated that several other species of blennies,
formerly referred to Salarias Cuvier, should properly be included
in Eesenius. The present study is a review of the genus and the
species included in it.
Ecsenius is a closely knit genus of 11 known marine species occurring
only in the tropical Indo-Pacific faunal area. Within that area
species have been found from the Red Sea on the west through the
Marshall and the New Hebrides Islands to the Hawaiian Islands on
the east and from Formosa on the north to Queensland, Australia, on
the south. While this genus occurs in the coral area, in relation to
coral heads, it typically occurs not on the shallow reefs but apparently
in moderate depths, and this perhaps accounts for the relative scarcity
of specimens in ichthyological collections.
The dentition clearly sets off Zesenius from all other genera of blen-
nies. In the upper jaw are more than 100 tiny, independently movable
teeth, a condition typical of the genera Jstiblennius Norman and
Salarias, whereas in the lower jaw, the labial teeth are fewer than 50
in number, they are much stouter than those above (usually with the
one or two teeth on each side farthest from the symphysis somewhat
1The senior author was aided in these studies by grants in aid from the John Simon
Guggenheim, Jr., Memorial Foundation and by the California Academy of Sciences,
507
969588—52——-1
508 PROCEEDINGS OF THE NATIONAL MUSEUM yOu. 102
enlarged or stronger than the others), and they are only slightly flex-
ible. This condition suggests Rhabdoblennius, and is distinctly in
contrast with that of Salarias and Istiblennius. The canine far back
on, the lower jaw, typical of salariin blennies, is tiny but stout, and
may be one or two in the same species. Furthermore, as Norman has
reported, traces of more teeth in this same dentary series can be de-
tected. Indeed, in the genotype #. mandibularis there are six or
seven plainly apparent teeth on each dentary in this series, quite dis-
tinct from the series of labial teeth on the anterior part of the dentaries,
a situation quite unknown in other salariin blennies. Norman has
suggested, on this account, that . mandibularis might be properly
placed in a separate subgenus. A study of the other species of the
genus has, however, inclined us to reject this suggestion. In all other
particulars, #. mandibularis agrees closely with the other species of
the genus. The types of that species are considerably larger than
specimens available in the other species. It is possible that these
latter are mostly juvenile and that the greater number of dentary
teeth in #. mandibularis is simply a factor of age, representing an
adult condition detectable as traces of teeth noted by Norman in other
species.
While the condition of the dentition might indicate a relationship
somewhere between /stiblennius, Salarias, and Rhabdoblennius, there
is little else that does, and no lineal phyletic connection is apparent
between H’csentus and those three genera. The lack of orbital or
nuchal cirri, the low number of dorsal spines, the unbranched caudal
rays, the full membranous connection of the last anal ray to the
caudal peduncle, the slender genital tube of the male, the elongation
of one or more caudal rays well beyond the fin membrane, which is
normal in most species, the lack of crenulation on the lips, the pelvics
1,3, the short lateral line with its double, rather than single, line of
pores, are characters that are mostly confined to H'csenius among the
salariin blennies, and, when put together, set the genus off sharply
from its relatives. There are other more subtle but just as definite
characteristics—the slender fin rays, which give an air of diaphanous
fragility to the vertical fins, and the anterior profile, which is either
vertical or even slopes backward from the forehead to the margin of
the upper lip.
The distribution of the known specimens and species of the genus
is suggestive of the division of the vast tropical Indo-Pacific fish fauna
into subareas, illustrated by the other genera of Salariinae and other
families of fishes studied by the authors. Two species have been found
FISHES, GENUS ECSENIUS—-CHAPMAN AND SCHULTZ 509
only in the Red Sea and at Djibouti; two have been found only
in the Solomon Islands; one each in the Marshall Islands, northeast
Australia, the Philippines, Formosa, the Hawaiian Islands, the west-
ern Indian Ocean, and the Persian Gulf. Only one species has been
determined to have a broad range, occurring in Ceylon, Bengal, Christ-
mas Island, Timor, and the New Hebrides Islands. In discussions
under each species some notes have been made of the possible relation-
ships of the several species, but clarification of these relationships must
await further collections throughout the range of the genus.
Genus ECSENIUS McCulloch
Besenius McCuttocu, Ree. Australian Mus., vol. 14, p. 121, 1923. (Genotype,
#. mandibularis McCulloch.)—Norman, Ann. Mag. Nat. Hist., ser. 11, vol. 10,
p. 810, 1943.
Description—Dorsal rays XII, 12-20; anal II, 13-21; pectoral
13-15 ; pelvic 1,3.
No crest on head in either sex (except low ridge on large male of
mandibularis) ; nasal cirrus either simple, bifid, or trifid; no supra-
orbital or nuchal cirrus; both lips smooth; one or more tiny, hidden
canines posteriorly on each dentary; 45 to 50 firmly placed teeth in
lower jaw (typical of Blennius); more than 100 independently
movable, slender teeth in upper jaw (typical of Jstéblennius and
Salarias) ; no teeth on vomer; depth 3.6 to 6.0, head 3.3 to 4.8, both in
standard length; lateral line a double series of pores anteriorly, end-
ing under tenth or eleventh dorsal spine, not turning downward on
side, continued on as a faint line with no visible pores to end of dorsal;
dorsal notched or not, reaching to or over first small rays of caudal;
last ray bound to caudal peduncle by membrane; caudal usually with
12, oceasionally 13, unbranched principal rays, with 6 or 7 small rays
above and below, mostly hidden under skin ; upper and lower principal
caudal rays typically exserted beyond margin of fin, by as much as
length of shortest caudal rays in large individuals of certain species;
2 spines in anal, always plainly visible in male and usually so in
female, but very minute in latter; no crenulated pads on first anal
rays of male, instead these rays on adult males are normally a little
swollen or bulbous at distal tip; no anal ray extends beyond margin of
fin in males; last anal ray fully bound to caudal peduncle by mem-
brane; males with a slender, tubular genital pore midway between
anus and insertion of anal spine; pelvics always with I, 3 rays, the
third soft ray normally bound to second and not visible without dis-
section ; pelvic spine always present but not visible without dissection.
VOL. 102
PROCEEDINGS OF THE NATIONAL MUSEUM
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FISHES, GENUS ECSENIUS—-CHAPMAN AND SCHULTZ 511
KEY TO THE SPECIES OF ECSENIUS
la. (See also 1b and 1c.) Pectoral with 15 rays and no dark spot around anus
(Red Sea and Djibouti).
2a. Color of head, body, and vertical fins a rich, uniform brown, with no
markings; dorsal XIJ,19; anal I1,20 (Red Sea).
Ecsenius frontalis (Cuvier and Valenciennes)
2b. Color of head and body light tan with no markings other than a few scat-
tered black specks on upper part of caudal peduncle; a jet black band
as wide as eye diameter running along base of both spinous and soft
dorsal, outer part of fin hyaline; top and bottom of caudal fin jet black
to end of long rays, rest of fin hyaline; anal fin colorless; dorsal XII, 18;
ans EE ZO CD NDouUtl ee eee Ecsenius gravieri (Pellegrin)
1b. Pectoral rays 13 to 15, usually 14; anal rays II, 18 or 19; a distinct dark spot
around anus; a large dark brown spot on base of caudal fin; dark brown
streak from lower edge of eye backward across gill cover; dorsal rays
XII, 16 or 17 (Philippines)__-Ecsenius stigmatura Fowler, new species
1c. Pectoral with 13 or 14 rays, rarely 12; if area around anus is black anal
rays are II,13 or 14 (tropical Indo-Pacific oceans).
8a. Nasal cirrus simple and single.
4a. Dorsal with 19 or 20 soft rays; anal with 21 or 22; a dark band along
base of dorsal (western Indian Ocean and Persian Gulf).
Ecsenius pulcher (Murray)
5a. Body evenly brown, without markings._._(Ecsenius pulcher, female)
5b. Head, throat, and body to level of sixth dorsal ray rich, light brown
(paler below), without markings; rest of body to caudal abruptly
pale, crossed by 5 or 6 equally spaced narrow, sharply defined
vertical bars, reaching from dorsal to ventral edge of body.
(Ecsenius pulcher, male)
4b. Dorsal with 12 to 14 soft rays; anal with 13 to 17 soft rays; no dark
streak along base of spinous dorsal and never sharply defined if
present on soft dorsal.
6a. Spinous and soft dorsal separated by shallow notch in young but no
notch at all in adults; sharply defined black spot, half the area of
eye, around anus, set off from rich brown body by light peripheral
band; no other markings on head, body, or fins; dorsal with 12 or 18
soft rays; anal with 13 or 14 soft rays (Solomon Islands).
Ecsenius lividanalis, new species
6b. Spinous and soft dorsal separated by a deep notch reaching nearly
to base of fin; last dorsal spine minute, nearly hidden in basal fin
membrane; no dark spot around anus; well-defined dark streaks,
bars or spots on body; dorsal with 13 to 16 soft rays; anal with
15 to 18 soft rays.
7a. A row of 6 or 7 small teeth laterally on ridge of dentary; anal
with 16 to 18 soft rays; double row of small, dark, roundish
spots along side of body, but no bars or bands (Queensland,
AUS EGS a) pee es Ecsenius mandibularis McCulloch
7b. One or 2 tiny teeth posterolaterally on the dentary, but never
more; anal with 15 soft rays; no dark spots on body, but dark
bars or bands present on sides; 2 pale horizontal bands, each as
wide as pupil, running full length of body, one from upper corner
of opercle, the other from under pectoral.
512 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 102
8a. Dorsal with 14 soft rays; a dark band as wide as pupil extending
back from eye to opercular edge; no vertical dark bars on body;
nasal cirrus about a third the diameter of eye in length; mem-
brane from last dorsal ray covering first 8 or 4 small caudal
shorter than postorbital length of head by about one-third
the diameter of eye (Solomon Islands).
Ecsenius pro-oculis, new species
8b. Dorsal with 13 soft rays; dark band from eye extending beyond
opercular edge onto body to a vertical from middle of spinous
dorsal; 8 or 9 narrow, dark, vertical bars on body extend-
ing to base of dorsal, where 2 or 3 may be divided; nasal
cirrus about half the diameter of eye in length; membrane of
last dorsal ray not covering first few caudal rays; pectoral
shorter than head by less than half the diameter of eye; pelvics
longer than postorbital length of head by nearly half the
diameter of eye (Marshall Islands).
Ecsenius opsifrontalis, new species
35. Nasal cirrus always double, ventral one may be bifid or double; dorsal
with 16 to 20 soft rays; anal with 19 to 21 soft rays.
9a. Head and anterior part of body evenly dark with no markings; posterior
part of body and adjacent vertical fins often abruptly pale (bright
golden in life) with no markings.
10a. Dorsal with 16 of 17 soft rays; anal with 18 to 20 soft rays (eastern
Indian Ocean; Christmas Island; Timor; New Herbrides Islands).
Ecsenius bicolor (Day)
10b. Dorsal wth 20 soft rays; anal with 21 soft rays (Formosa).
Ecsenius namiyei (Jordan and Evermann)
9b. Middle of side of body with 5 to 7 short but distinct white bars evenly
spaced; a dark spot behind eye; middle caudal fin rays dusky to
blackish; body plain olive to light greenish brown, undersides pale
OLIV ORES eae oe ee Ecsenius hawaiiensis, new species
ECSENIUS FRONTALIS (Cuvier and Valenciennes)
Salarias frontalis Cuvier and VALENCIENNES, Histoire naturelle des poissons,
vol. 11, p. 828, 1836 (type locality, Massuah, Red Sea).—Gunruer, Catalogue
of the fishes in the British Museum, vol. 3, p. 245, 1861 (in synonymy of
?8. fuscus).—Bamber, Journ. Linn. Soc., vol. 31, p. 484, pl. 46, fig. 1, 1915
(Suakim, Red Sea).
Specimens.—Three specimens, 37 to 49 mm. long, from Baie de
Djibouti, Gravier (in a jar with specimen of Enchelyurus), labeled as
Salarias fuscus), P. M. 04.318; 1 specimen, 52 mm. long, collected by
Crossland in Red Sea, B. M. 1915.10.25.9.
Description —Dorsal rays XIJ,19; anal IT,20; pectoral 15; pelvic
1,3.
Nasal cirrus single and simple, on level with upper level of pupil,
lanceolate, nearly as long as eye diameter; snout vertical or somewhat
projecting; one or two tiny canines below on either side; depth 4.8
to 5.4, head 3.6 to 4.4, both in standard length; dorsal spines and rays
FISHES, GENUS ECSENIUS—-CHAPMAN AND SCHULTZ 513
of same height, with no trace of notch, the fin ascending gradually
to greatest height in middle of soft dorsal; last ray with its tip free
from membrane binding it to caudal peduncle, the membrane not
covering first small caudal rays; caudal truncate, upper and lower
two rays extended out from level of fin; anal with tip of last ray free
from membrane; pectoral extending back to anal insertion; pelvics
are equal to two-thirds the postorbital length of head.
Coloration.—Matix: Color of head, body and vertical fins a rich
uniform brown with no markings and the abdomen only a little more
pale than the rest of body; peritoneum black; anterior top edge of
spinous dorsal white; caudal somewhat dusky in the two larger males;
pectorals pale and clear. Frain: As male, except caudal clear.
Remarks.—The four specimens listed are without doubt the Salarias
frontalis of Cuvier and Valenciennes. Except for the lack of mark-
ings, the higher pectoral count, and the single nasal cirrus this species
resembles #. bicolor, and it should be considered the Red Sea cognate
of that species.
ECSENIUS GRAVIERI (Pellegrin)
Salarias gravieri PELLEGRIN, Bull. Mus. Hist. Nat. Paris, vol. 12, p. 93, 1906
(type locality, Djibouti).
Specimen.—One male, 53 mm. long, collected by Gravier in Baie de
Djibouti, the type of the species. P.M. 04-319.
Description.—Dorsal rays XII,18; anal 11,20; pectoral 15; pelvic
I,3.
Nasal cirrus single and simple, no longer than diameter of pupil;
snout vertical or forehead somewhat projecting; one or two tiny
canines on either side below; depth 4.8, head 3.5, both in standard
length; all dorsal spines except first excised, the first few to a third
their depth; shallow notch between the spinous and soft dorsal, the
last spine short and hidden in basal membranes; only tips of soft
dorsal free; all caudal rays excised at tips, but upper and lower two
principal rays extended by a length equal to two-thirds the length of
the middle rays; dorsal and anal attached to caudal peduncle, the
membranes reaching to, but not over, first small rays of caudal.
Coloration.—MA.x: Except for a few scattered black specks on the
upper part of the caudal peduncle the body and head are pale tan
with no markings; peritoneum black; a jet black band running along
base of both dorsals, as wide as eye diameter; outer part of fin per-
fectly clear and unmarked; a few scattered black specks on base of
caudal; top and bottom of caudal jet black to end of long rays; rest of
fin hyaline; anal and paired fins colorless. Frmare: None available.
Remarks.—This species is closely related to #. frontalis (from the
same locality), but the character of the dorsal and the striking color
pattern seem to set it off clearly. This can scarcely be an older male
514 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
of E’. frontalis, for specimens of the latter, examined in the Paris and
British Museums, were of practically the same size as those of £.
gravieri.
ECSENIUS STIGMATURA, Fowler, new species
Figure 90
Holotype.—U.S.N.M. No. 99379, Philippine Islands, Dammi Island
between Jolo and Tawi Tawi Straits, lat. 5°52’12’’ N., long.
120°31’00’" E., depth 244 fathoms, September 21, 1909, Albatross,
standard length 46.5 mm., female.
Paratypes—U.S.N.M. No. 111878, taken with the holotype and
bearing same data, standard length 37 mm., female; U.S.N.M. No.
122444, Philippine Islands, Cataingan Bay, east of Masbate Island,
April 18, 1908, Albatross, standard length 36.2 mm., female.
~ Ficure 90.—Ecsenius stigmatura F owler, new species. (From photograph of color
drawing from the Philippine Albatross collection.)
Description——The following counts were made on the holotype
and paratypes, respectively: Dorsal rays XII,17; XII,17; XII,16;
anal I1,19; II,19; II,18; pectoral 13-14; 14-14; 15-15; pelvic 1,3;
1,3; 1,3; no branched caudal rays; nasal cirri 1-1; 1-1; 1-1.
Certain measurements were made on the types and these data are
recorded in thousandths of the standard length in table 2.
Head (tip of upper lip to end of gill cover) 3.6 to 4.0; greatest
depth 3.8 to 4.7, longest pectoral ray 5.2 to 5.8, snout tip (upper lip) to
anus 1.8 to 1.9, anal fin base 2.5 to 2.7, all in the standard length.
Kye 2.7 to 3.3, snout 3.5 to 3.7, postorbital length of head 1.5, greatest
depth of body 1.1, least depth of body 2.2 to 2.3, longest pectoral ray
1.4 to 1.5, longest dorsal spine 1.8 to 1.9, all in the length of head
from tip of upper lip (snout) to rear of gill cover. Fleshy inter-
orbital space 2.5 to 3.0 in eye.
No orbital cirrus; no nuchal cirrus; a single, simple, nasal cirrus
arising on upper edge of nasal opening; snout profile with a notable
backward slant of 25 to 30 degrees from the vertical; edges of both lips
smooth; lateral line running a dorsal course over pectoral fin and
ending about opposite tip of pectoral fin and base of eighth or ninth
dorsal spine, not descending to middle of side; vertical line through
FISHES, GENUS ECSENIUS—-CHAPMAN AND SCHULTZ 515
dorsal origin passes through rear edge of opercle, base of pectoral fin,
and notably far behind base of pelvics; last dorsal ray attached by
membrane to dorsal edge of caudal peduncle but not to base of caudal
fin; anal origin opposite base of last dorsal spine; last anal ray at-
tached by membrane to lower edge of caudal peduncle; pectoral fins
reaching about two-thirds the way to the anus; anal spines small, first
embedded; posterior canine of lower jaw short but strong; teeth in
upper jaw about 110, movable, those in lower jaw larger, movable,
about 40 to 45 in number, the teeth on each side farthest from the
symphysis notably enlarged; no vomerine teeth; posterior margin of
caudal fin probably truncate; no cephalic crest; a vertical line through
front edge of upper lip passes through front edge of eye; forehead a
little in front of tip of upper lip.
Coloration—In aucouou: Background coloration plain reddish
brown; a large area in front of anus blackish brown; on base of caudal
fin and on caudal peduncle a dark brown spot, somewhat angular
posteriorly ; pectoral, pelvics, dorsal, and caudal fins hyaline or pale;
anal fin dusky brown; a narrow dark brown streak extends from lower
edge of eye backward to rear angle of opercle, below this dark streak
on opercle is a narrow pale streak that continues on body behind
head only, just dorsal to base of pectoral fin.
Autve (based on color sketch made on A/batross): Background
coloration reddish brown; anal spot purplish black; caudal! spot black;
anal reddish brown; dorsal spines orange; an orange streak along
base of dorsal fin; iris orange; dark streak behind eye is dark blue,
and pale streak below it is orange; upper lip slightly yellowish brown.
Ecology.—This species probably came from deep water, as the
Albatross dredge record indicates 244 fathoms.
Remarks.—£, stigmatura is one of two known species with a black
area (purplish black when alive) in front of the anus; £. lividanalis,
new species, lacks the caudal spot and streaks behind the eye. Addi-
tional characters that distinguish 2. stigmatura from the other species
referred to the genus may be found in the accompanying key.
ECSENIUS PULCHER (Murray)
Salarias pulcher Murray, Journ. Bombay Nat. Hist. Soc., vol. 2, p. 47, 1887
(type locality, Kurrachee Manora Rocks, India) ; Indian Ann. Mag. Nat. Sci.,
vol. 1, p. 23, 1887 (Kurrachee, India).
Salarias phantasticus BouLENGER, Ann. Mag. Nat. Hist., ser. 6, vol. 20, p. 422,
1897 (type locality, Mekran Coast, Persia).
Salarias anomalus Recan, Journ. Bombay Nat. Hist. Soc., vol. 16, pp. 327, 331,
pl. B, fig. 4, 1905 (type locality, Persian Guif; Mekran Coast, Karachi) ;
Journ. Zool. Soe. London, p. 406, 1909 (on type material).
Specemens.—T wo males, each 45 mm. long, collected by Murray at
Kurrachee (the types of Salarias pulcher), B. M. 87.9.22.59—60; 2
969588—52——_2
516 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
males, 56 and 58 mm. long, collected by Townsend on Mekran Coast,
Persia (types of Salarias phantasticus) , B. M. 97.9.22.20-21 (these are
like pulcher types except that the posterior dorsal spines and caudal
rays are even longer and more extended; black vertical lines, respec-
tively, 7 and 8; two tiny black dots on posterodorsal quarter of
orbit) ; 16 female, 82 to 58 mm. long, collected by Townsend in Persian
Gulf (the type series of Salarias anomalus), B. M. 1900.5.9.47-56 ;
1 female, 41 mm. long, collected by Stephens at Henjam Islands, Per-
sian Gulf, B. M. 1932.2.18.43; 1 female, 58 mm. long, collected by
Townsend at Hinderabi Islands, Persian Gulf, B. M. 98.6.29.163; 2
females, 29 and 34 mm. long, collected by Townsend at Jask, Mekran
Coast, Persia (these specimens have a short horizontal white bar on
the cheek under posterior part of eye—caudal and spinous dorsal
only barely lengthened) ; female, 51 mm. long, collected by Knox at
Muscat, B. M. 1912.11.26.1.
Description.—Dorsal rays XII,19-20; anal I7,20-21; pectoral 14;
pelvic I,3.
Nasal cirrus slender, single, simple, pointed, about equal to eye in
length; forehead projecting, a concavity between it and upper lip;
eye high and large, reaching into anterior profile and almost into
dorsal profile; a single tiny canine on either side below, hidden in fold
of membrane at corner of jaw; depth 5.2 to 5.7; head 4.3 to 4.5, both in
standard length; dorsal deeply notched, the last spine about one-third
length of the next preceding; posterior dorsal spines elevated, their
tips free from the fin membrane, the fourth from the last as long as
head, its distal one-fourth free from membrane; membrane from last
ray reaching to, but not over, first small ray of caudal; caudal with
12 principal rays, none branched, the second and eleventh rays long-
est, the fin forked, almost filamentous, longer than head in specimens
as much as 45 mm. long; anal spines of male both small and hidden
at base of first ray; no pads or swellings at tips of anal rays; last
ray bound fully to caudal peduncle but not reaching first caudal
rays; longest pectoral ray equal to postorbital length of head; long-
est pelvic ray less than two-thirds postorbital length of head.
Coloration.—Mats: Head, throat, and body rich, light brown (paler
below) and with no markings back to level of sixth dorsal ray; on
this pale posterior part of body five or six equally spaced, narrow,
sharply defined dark vertical bars reaching from dorsal to ventral
edge of body; the anterior line a little wider than pupil, the posterior
one a little narrower; in some specimens there may be a shorter
fine line between some of the main lines; basal half of spinous dorsal
and all of soft dorsal evenly dusky; this area, while pigmented,
transparent and with no markings; distal half of spinous dorsal
pale, but bearing markings—an oval dark spot the size of pupil
FISHES, GENUS ECSENIUS—CHAPMAN AND SCHULTZ 517
between spines 1 and 2, another between 2 and 3, another between 5
and 6, another near end of 7, and another near ends of spines 8 and
9, last two spots thickened and involving the spines (on one speci-
men only the third and fifth of these dorsal spots are present) ;
anal clear, narrowly edged with black; caudal dusky distally but
with no actual markings and transparent; paired fins clear. f'rmaun:
Like male except that the body is plain brown without the markings
so typical of the male.
Remarks.—As in most other species of the genus, the males and
females of this species cannot normally be differentiated by the ex-
ternal sex organs. Examination of the gonads of the types of both
pulcher and anomalus demonstrates that the former is the male, the
latter is the female of the species. It is quite possible that this is the
cognate in the western Indian Ocean of opsifrontalis from the Marshall
Islands and pro-oculis from the Solomon Islands.
ECSENIUS LIVIDANALIS, new species
Wieurr 91
Holotype.—U.S.N.M. No. 144723, male, 34 mm. in standard length,
collected by Chapman and Cheyne at Munda, New Georgia, Solomon
Islands, June 15, 1944.
Paratypes.—_U.S.N.M. No. 144291, 3 specimens, 24 to 33 mm., col-
lected with holotype and bearing same data; U.S.N.M. No. 144292,
6 specimens, 24 to 30 mm., collected by Chapman and Cheyne in Munda
Lagoon, New Georgia, Solomon Islands.
Description.—Dorsal rays X11J,12-13; anal IT,13-14; pectoral 13;
pelvic I,3.
Nasal cirrus simple, slender, pointed, equal to about half diameter
of pupil in length; forehead projecting slightly, but profile rounded ;
a single, tiny canine on each side below, rather far forward and
no more than one-third the size of labial teeth; lateral tooth occasion-
ally set off a little from the labial teeth; depth 3.6 to 4.3, head 3.3 to
3.5, both in standard length; spinous dorsal slightly rounded ante-
riorly, a little lower than soft dorsal; younger specimens show slight
notch between spines and rays but older specimens have no notch,
only a difference in level between spines and rays; last spine does not
come up to edge of fin membrane and is only two-thirds height of
next spine; membrane attaching last dorsal ray to caudal peduncle
reaching over first two or three small caudal rays; last ray of anal
similarly attached to caudal peduncle; genital papilla of female simi-
lar to that of Salarias and hides first tiny anal spine; second anal spine
half length of first ray; anal highest in middle where it is two-thirds
height of soft dorsal; male genital tube slender, nearly as long as
first anal spine and situated midway between it and anus; caudal
518 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
evenly rounded, shorter than head by length of snout and one-third
the diameter of eye; none of rays forked or exserted; longest pectoral
ray shorter than head by snout and half the eye; longest pelvic ray
equals two-thirds postorbital length of head, spine completely hidden,
slender third ray scarcely visible without dissection.
Coloration--Matrz: Over-all color a rich, reddish brown, only a
little highter on throat than elsewhere; brown extends onto dorsals
Ficurr 91.—E£csenius lividanalis, new species. Holotype. (Drawn by Louise Horne.)
but fades distally so that outer half of fin more or less transparent;
anal rather evenly dusky with edge lighter; caudal with only a little
pigment along rays and on membrane basally; pectorals clear; eye
black; a sharply defined black spot (brilliantly blue in life) half the
area of eye surrounding anus, and set off from brown of body by a
light peripheral band; no other markings on head, body, or fins.
Fruate: Same as male.
ECSENIUS MANDIBULARIS McCulloch
Eesenius mandidularis McCuritccn, Ree. Australian Mus., vol. 14, No. 2, p. 122,
pl. 15, figs. 1 and 2, 1923 (type locality, Masthead Island, off Port Curtis,
Queensland, Australia).
Specimens.—None examined. Description after McCulloch.
Pescripiion.—Dorsal rays X11,18-16; anal I1,16-18; pectoral 18;
pelvic 1,3.
Nasal cirrus single, simple, borne on posterior nasal pore and about
half diameter of eye in length; forehead slightly projecting but evenly
rounded; six or seven small conical teeth on each side on ridge of
dentary and behind labial teeth (in position occupied by the one or
two tiny canines found in other species of the genus) ; depth 4.7, head
4.1, both in standard length; spinous dorsal rounded posteriorly, the
ast spine tiny, buried in membranes at bottom of deep notch between
spinous and soft dorsals; longest dorsal spine a trifle shorter than
longest ray; last dorsal ray bound to caudal peduncle by membrane
FISHES, GENUS ECSENIUS—CHAPMAN AND SCHULTZ 519
extending to first small rays of caudal; first anal spine buried in geni-
tal pad of female, but visible in male; anal lower than soft dorsal with
a small fleshy pad at tip of each ray in male; caudal with 12 or 13
principal rays, those above and below irregularly exserted beyond
fin margin in males; pectoral as long as postorbital length of head and
half diameter of eye; longest pelvic ray about two-thirds postorbital
length of head.
Coloration —Ma.x: Head and body uniformly light brown with a
bluish tinge on sides; two rows of well-defined, evenly spaced, dark
spots each smaller than pupil along the sides, the upper, of about nine
spots, above level of lateral line, the lower, along middle of side, be-
ginning behind pectoral, and containing about seven spots; no other
markings on head or body; fins without markings except pads on
tips of anal rays distinctly lighter than rest of fin. Fraarm: Like
male.
Remarks.—Set off from other species in the genus chiefly by the
number of teeth in the canine series on the dentary.
Ficure 92.—Ecsenius pro-oculis, new species. Holotype. (Drawn by Louise Horne.)
ECSENIUS PRO-OCULIS, new species
FIGURE 92
Holotype—U.S.N.M. No. 144722, a male, 40mm. in standard length,
‘collected by Chapman and Cheyne, May 20, 1944, at a little island in
Munda Lagoon near Sessavelle, New Georgia, Solomon Islands, among
coral heads in about 10 feet of water.
Description—Dorsal rays XII,14; anal IJ,15; pectoral 13; pelvic
1,3.
Nasal cirrus single and simple, rather bluntly pointed, as broad
as nasal pore on whose upper edge is it borne, about one-third diameter
of eye in length; forehead strongly projecting so that anterior edge
of orbit is a little beyond a vertical from margin of upper lip, giving
520 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 162
the eyes the effect of protruding; a single canine on either side below,
rather far forward and no larger than corner labial teeth of lower
jaw; the two corner teeth on each side of the lower jaw somewhat
detached from rest of series and caninelike; depth 4.3, head 4.2, both
in standard length; third spine of dorsal highest, with fin shortening
rapidly posteriorly, next-to-last spine only half length of third spine
or first ray, last spine a tiny nubbin almost buried in basal membrane
of fin; third ray of soft dorsal longest; this part of fin also shortening
rapidly posteriorly; membrane binding last ray to caudal peduncle
covering first three or four small rays of caudal; soft dorsal a little
higher than spinous dorsal, with deep notch between two parts of
fin; anal lower than spinous dorsal; both anal spines plainly visible,
the second being two-thirds the length of first ray, the first two-thirds
the length of second; all rays except last with a small fleshy pad at
the tip; last fully bound to caudal peduncle by membrane; genital
papilla or tube of male a little longer than wide; caudal with 13 prin-
cipal rays and 7 small rays both above and below; middle caudal rays
equal head in length; third and fourth dorsal principal rays greatly
elongated, extending beyond level of fin by two-thirds the length of
middle rays; third principal ray from ventral edge of fin only slightly
extended; none of caudal rays forked ; longest pectoral and pelvic rays
approximately equal in length and shorter than postorbital length
of head by one-third the diameter of eye; third ray of pelvic, while
entirely bound to, and much more slender than, second, plainly visible;
pelvic spine not visible without dissection.
Coloration —Matx: Peritoneum black; throat noticeably paler than
sides of head and without markings; rest of head and the body a very
dark reddish brown; a darker band as wide as pupil extending back
from eye to edge of opercle; two horizontal light bands, each as wide
as pupil, run full length of body, one from upper corner of opercle
to upper part of caudal base, the other from under the pectoral to
lower part of caudal base; no spots or other markings on head or body:
spinous dorsal clear and transparent; soft dorsal with a clear band
at base, then a narrow dusky band which shades off above to leave
most of fin clear and transparent; middle third of caudal dusky, upper
and lower thirds mostly clear; anal uniformly dusky with pads at
tips of rays a little paler; pectorals clear; pelvics uniformly dusky.
Femase: None available.
Remarks.—This species, and #. opsifrontalis from the Marshall
Islands, are distinguished from the species by the marked projection
forward of the orbits and the resultant backward slope from the
forehead to the upper lip.
FISHES, GENUS ECSENIUS—-CHAPMAN AND SCHULTZ 521
ECSENIUS OPSIFRONTALIS, new species
Fiqure 93
Holotype.—U.S.N.M. No. 142065, Rongelap Atoll, Rongelap Island,
Marshall Group, collected by Brock, Herald and Kohler, July 25, 1946,
in lagoon at depth of 18 feet, standard length 31 mm.
Paratype.—U.S.N.M. No. 142066, Bikini Atoll, collected by Brock
and Schultz in lagoon at depth 20 to 25 feet, March 26, 1946, 1 specimen,
26.3 mm.
Description—Dorsal rays XII,18; anal I1,15; pectoral 18;
pelvic 1,3.
Certain measurements were made on the types, and these data are
recorded in thousandths of the standard length in table 2.
Nasal cirrus single and simple, slender, rising on dorsal side of
nasal pore, about half diameter of eye in length; forehead projecting
so that anterior slopes from level of eye back to margin of upper lip
at an angle of 30 to 45 degrees from the vertical ; a single, small canine
close behind other teeth on dentary, hidden by fold of membrane at
Ficure 93.—Ecsenius opstfrontalts, new species. Holotype. (Drawn by Dorothea B. Schultz.)
corner of mouth; depth 4.8 to 5.1, head 3.8 to 3.9, both in standard
length, dorsal spines all about same length, except that last is no
more than two-thirds length of next-to-last; membrane definitely
notched behind spinous dorsal; soft dorsal highest in middle, the
longest ray a little longer than longest spine; last ray bound to caudal
peduncle by a membrane which does not reach first small caudal rays;
both anal spines visible in both sexes; females with a rounded, rather
large genital pad with a tiny nubbin on posterior edge; males with a
slender tube, half length of first spine, between anus and first anal
spine; tips of first anal rays not swollen; longest anal rays shorter
than longest dorsal spine or ray; caudal approximately truncate, the
upper and lower rays not exserted in specimens of the sizes available;
13 principal rays, with about 6 or 7 small rays above and below;
none of rays branched; pectoral shorter than head by less than half
the diameter of eye; pelvic fin longer than postorbital length of head
by more than half the diameter of eye, spine completely hidden, third
O22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
ray slender, shorter than second and so-closely bound to it as to be
clearly visible only by dissection.
Coloration—Matzs: Background of head and body light brown;
a dark band as wide as eye extending straight back from posterior
edge of eye to a vertical from about twiddle of spinous dorsal; two
faint light bands about as wide as pupil running horizontally on ‘body
to base of caudal, one from upper edge of operele, the other from be-
low pectoral; eight narrow, dark, transverse bars spaced on body, run-
ning from hes of dorsal com to, but not across, lower pale longt-
tudinal band; two or three of these bars may be nit dorsally ; the last,
on caudal peduncles is C-shaped with the opening posterior; no spots
or other markings on head or body; both lips dusky; anal fin with a
dusky submarginal band, tips of rays white; other fins with no mark-
ings. HrmAa.e: essentially the same coloration as male.
Remarks—This species is probably the Marshall Island cognate of
E. pro-oculis, with which it shares many characters, but from which it
differs as indicated in the key. The color is generally less dark than
in #. pro-ocults and the markings are therefore less sharply defined.
ECSENIUS BICOLOR (Day)
Figure 94
Salarias bicolor Day, Supplement to the fishes of India, p. 798. (on Tickell ms.)
1888 (type locality, Saddle Island, Kyoukphyoo Aracan); The fauna of
British India .. ., vol. 2, Fishes, p. 323, 1889 (on type material)..WEBER,
Die Fische der Siboga-Wxpedition, vol. 57, p. 533, 1913 (eastern tip of Timor).
Salarais furcatus (non De Vis 1884) JomnstTonr, Report of Ceylon Pearl Oyster
Fisheries, vol. 2, No. 15, p. 2138, pl. 1, fig. 4, 1904 (type locality, Chilam Paar,
Ceylon, at 15 to 20 meters depth).—WHrttey, Rec. Australian Mus., vol. 17,
No. 3, p. 136, 1929 (notes that this species is not the same as that of De Vis).
Salarias burmanicus Hora and Muxexsi, Rec. Indian Mus. Calcutta, vol. 38, D.
34, 1936 (type locality, Maung. Magan, Tavoy District, lower Burma).
Specitmens.—11 specimens, 31 to 49 mm. long, collected by Chapman
and Cheyne at Espiritu Santos, New Hebrides Island, U.S.N.M. Nos.
144716, 144293 and 144294; 6 specimens, 30 to 49 mm. long, collected
by Andrews at Christmas Island (the type series of Salarias melano-
soma Regan), B. M. 1909.3.4.52-57; 1 specimen, 44 mm. long, from
Godeffroy Museum, from Ponape, B.M. 81.10.20.189; 2 specimens, 42
and 49 mm. long, from Godeffroy Museum, from Ponape, B.M. 81.10.-
20.187 ; 1 specimen, 38 mm. long, collected by Weber at Siboga station
282, Oued of Timor.
Btscription._Dorel rays X1J,16-17 ; anal IT,18-20; pectoral 13-14;
pelvic, 1,3.
Nasal cirrus double; cirrus on dorsal side of nasal pore simple,
slender, pointed, and equal to or a little greater than diameter
of eye in length; cirrus on ventral edge of pore usually simple, but
may be forked or even nearly double, half the length of the dorsal
FISHES, GENUS ECSENIUS—CHAPMAN AND SCHULTZ 520
cirrus and of same shape; anterior profile either vertical or with fore-
head slightly projecting; a single small canine close behind labial
teeth, hidden by fold of membrane in corner of mouth on each side
of lower jaw in males; not found in females; depth 5.0 to 5.2, head
4.5 to 4.8, both in standard length; dorsal spines all approximately
same height except last, which are half height of first ray and two-
thirds height of next-to-last spine; soft dorsal highest in its middle,
and longest spine a little shorter than longest ray; spines not extend-
ing beyond fin membrane; in small specimens fin membrane comes
down to height of last spine, making a distinct, if shallow, notch; in
larger specimens of both sexes the membrane extends straight back
with no notch between fins, only difference in level; membrane bind-
ing last ray of dorsal and anal to caudal peduncle reaching to, but
Figure 94.—Ecsenius bicolor (Day). Specimen from New Hebrides Islands. (Drawn by
M. Nickerson).
not over, first small ray of caudal; caudal with 13 principal rays and
6 smaller ones both above and below; second and third principal rays
extending beyond edge of middle rays, third to a distance equaling
half length of middle rays; third principal ray from bottom extending
even farther; in females of the same size the elongated caudal rays
project by no more than one-fourth the length of the middle rays, and
in smaller specimens of both sexes they scarcely extend beyond normal
edge of fin; both anal spines plainly visible in both sexes, the second
three-fourths the length of the first ray, the first one-half the length
of the first ray; in male, but not female, each anal ray bears a small
bulbous enlargement on anterior side of its tip; in male a slender
genital tube half the length of first spine between anus and first spine;
in females a rounded, rather large genital pad with a tiny nubbin on
its posterior edge; longest pectoral ray shorter than head by length
of snout and two-thirds eye; longest pelvic ray equal to about two-
thirds postorbital length of head; pelvic spine and third ray com-
pletely hidden and visible only by dissection.
Coloration—Mate: Head and anterior two-thirds of body almost
uniformly dark brown (bluish in life) ; blackness of peritoneum makes
abdomen darker than above; in posterior third of body this dark area
524 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
shades off into a very pale area, which extends over the caudal; this
pale area bright golden in life; pale area extends anteriorly farther
along base of anal, and a little way along base of dorsal; on smaller
specimens the dark pigment extends farther posteriorly until there
may even be a little on the base of the middle caudal rays, but always
the fish is lighter posteriorly than anteriorly ; no spots or markings on
the head or body, except for an indistinct light band from the eye down
across the upper lip and an indistinct dark streak directly behind the
orbit; a continuous dusky band along basal portion of dorsal back
to where light area of body begins, covering basal two-thirds of
spinous dorsal, where it contains, over the first three spines, a horizon-
tal, elongate, black streak; anteriorly the distal part of spinous dorsal
clear except for margin; posteriorly this dark margin becomes wider
until it melts into lower dusky band; soft dorsal with a dusky marginal
band, but remainder of fin clear except for basal pigment anteriorly
and a little pigment along each ray; in smaller specimens the dusky
basal band extends to end of soft dorsal; anal fin with a fine clear band
mesially, on anterior rays, which broadens posteriorly to cover base
and more than half of fin; rest of fin, including whole distal edge,
dusky except for fleshy pads on tips of rays, which are lighter ; caudal
entirely pale; pectoral rays dark, especially lower ones, but membranes
clear and light; pelvics dusky, but lighter than body next to them;
eyes almost black, as are the nasal cirri. FrmaLte: Same as male ex-
cept light posterior part of body continued to part of caudal peduncle,
more of anal dusky, and there is some pigment on the distal two-
thirds of middle of caudal.
Remarks. —While the description of bécolor by Day is brief and the
fin counts contain either none of the spines, or not all of them,
there is no question but that he was dealing with this well-marked
species, as is true of the specimens described by Johnstone as S.
furcatus and Hora and Mukerji as S. burmanicus.
The series of specimens described as S. melanosoma from Christmas
Island might be considered as being subspecifically distinct from the
above, for all these specimens have 18 soft rays in the anal (all the
New Hebrides specimens have 19 or 20 soft anal rays), the upper
and lower caudal rays either do not project, or project less than one-
fourth the length of middle caudal rays, and the color is a uniform
rich brown, scarcely lighter below than above. However, of the three
specimens from Ponape one had 18 soft anal rays and the other two
had 19, and there is no clearly distinguished color break posteriorly.
One has the caudal rays extended by the length of the middle caudal
rays, the other two by less than one-third the length of the middle
caudal rays. There is no dependable evidence in this group of sub-
speciation.
FISHES, GENUS ECSENIUS—CHAPMAN AND SCHULTZ 525
The specimen from Timor is identical with the New Hebrides speci-
mens except that it has only 18 soft anal rays. Coloration well marked.
Ecsenius namiyei is doubtless the Formosan cognate of this species,
marked particularly by the greater number of dorsal and anal rays,
and the uniform coloration of the body.
Escenius frontalis is very likely the Red Sea cognate of this species,
well marked, however, by the single nasal cirrus and the 15 pectoral
rays.
ECSENIUS NAMIYEI (Jordan and Evermann)
FievureE 95
Salarias namiyei JORDAN and EvERMANN, Proc, U. 8S. Nat. Mus., vol. 25, p. 362,
fig. 25, 1902 (type locality, Pescadores Islands).
Specimens.—None seen; description from Jordan and Evermann.
Description.—Dorsal rays XII,20(%); anal I1,21; pectoral 13;
pelvic 1,3 (?).
A pair of tentacles on nasal pore, the longest no longer than diameter
of pupil; forehead vertical, the orbits reaching the dorsal but not
Figure 95.—Ecsenius namtyei (Jordan and Evermann). Holotype. (Drawn by A. H.
Baldwin.)
anterior profile of head; depth 4.5, head 4.5, both in standard length;
dorsal rays all about the same height except for last three rays, no
trace of notch; membrane from last dorsal and anal rays free from
first caudal rays; caudal with 12 principal rays, none branched, trun-
eate behind; pectoral shorter than head by snout and half the eye dia-
meter; pelvics two-thirds postorbital length of head.
Coloration——Body purplish red, paler posteriorly; dorsal fin with
narrow oblique darker markings; anal darker along margin, with a
similar median band.
Remarks.—This is undoubtedly the Formosan cognate of F. bicolor.
The three more rays in dorsal, one more in anal, and the oblique mark-
ings of the dorsal prevent synonymizing it with bicolor in the absence
of a series of specimens for comparison.
526 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 102
ECSENIUS HAWAIIENSIS, new species
Figure 96
Holotype.—U.S.N.M. No. 112293, Oahu Island, Pearl Harbor, from
a drydock, “hauled from Guam about a year ago and left standing,”
April 1950, collected by Spencer Tinker, standard length 66 mm.
Paratypes.—U.S.N.M. No. 112294, taken with holotype and bearing
same data, 12 specimens, 48 to 70 mm.
Description.—In addition to the counts recorded in table 1, the
pelvics were always 1,3; none of the caudal fin rays is branched.
Certain measurements were made on the holotype and two paratypes
and these data are recorded in table 2.
TABLE 2.—Measurements on certain species of Ecsenius, recorded in thousandths
of the standard length
|
| opsifrontalis stigmatura hawaiiensis
Characters ae “ oo =
type reid ie Paratypes sie Paratypes
Standard length in mm_-_.___..._---- 31.0 26.3 46.5 37 36. 2 66 66.7 48
Head, tip of snout to front of upper \
Vipeeee fe oni tcccsenccsc an hneeee eee 255 266 249 251 263 227 220 227
Total length of head_-._____- eee ee. 281 278 262 273 279 242 234 240
Greatest: depth... 2-2 2 NOs 197 209 215 224 248 207 202 217
Least depth of body_...-.--..._--_-- 97 95 105 116 111 112 111 104
Postorbital length of head_________-_- 181 182 178 184 180 165 157 161
HG YG xe rin ei eh reo) ORs ek ae 77 80 69 81 83 59 68 71
Snot oS 74 76 67 73 72 77 73 67
Interorbital space (fleshy) - -.--- BUSA 31 3 32 30 33 38 30 38
Length of posterior nasal cirrus_____- 39 38 49 65 83 52 60 69
Longest fin ray:
Horsalispine-=) == 2252-2 a ee ee 158 141 114 135 | 146 121 | 135 150
Morsalisoltrayossses oe ae 177 145 142 119 163 189 165 133
Amal iene lee soa acol. ae eae 126 126 118 122 105 118 112 125
Pectoral. tes3 2 ase eter ks 2 242 236 183 184 191 151 145 133
Pelvieuasc-2=- == eS 27-1} 198 110 132 122 103 84 110
Oand als! es. ee eee ee 232 240 221 249 265 313 3sl 202
Snoutitipito anus. soonest e558 559 542 552 530 508 487 516
Snout tip to dorsal origin._.._.______] 258 285 236 257 | 248 212 | 210 219
Length of anal fin base.._____._____- 355 327 | 380 386 | 378 484 | 408 406
|
Head 4.0 to 4.2; greatest depth 4.5 to 4.8; longest pectoral ray 6.1
to 6.3; snout tip (upper lip) to anus 2.1; anal fin base 2.3 to 2.4; all
in the standard length. Eye 3.5 to 4.0; snout 3.1 to 3.4; postorbital
length of head 1.4 to 1.5; greatest depth of body 1.2 to 1.3; least depth
of body 2.2 to 2.5; longest pectoral ray 1.5 to 1.6; longest dorsal spine
1.7 to 1.8; all in the length of head from tip of upper lip to rear
of gill cover. Fleshy interorbital space 1.8 to 2.0 in eye.
No orbital cirrus; no nuchal cirrus; a single, simple nasal cirrus
arises on upper edge of anterior nasal opening, and a shorter cirrus,
sometimes bifid to base, arises on lower edge of nasal opening; snout
FISHES, GENUS ECSENIUS—CHAPMAN AND SCHULTZ 527
profile with a notable backward slant from the vertical of about 15
degrees ; edges of both lips smooth, lateral line running a dorsal course
and ending opposite bases of last or next-to-last dorsal spines, about
an eye diameter behind tip of pectoral fin; lateral line does not descend
to middle of side; vertical line through dorsal origin passes a trifle in
front of upper edge of gill opening; base of pectoral fin notably behind
base of pelvics; last dorsal ray attached by membrane to dorsal edge
of caudal peduncle, but not to base of caudal fin; anal origin opposite
base of last dorsal spine; last anal ray attached by membrane to lower
edge of caudal peduncle; pectoral fins reaching about two-thirds the
way to anus; anal spines small, first embedded on females; posterior
canine of lower jaw short but strong; teeth in upper jaw about 110,
movable; those in lower jaw movable but firmer, in one specimen 38,
the tooth farthest from the symphysis on each side enlarged; no
vomerine teeth; posterior margin of caudal fin truncate on smallest
Ficure 96.—Ecsenius hawattensis, new species. Holotype. (Drawn by Aime M. Awl.)
specimen, with an outer ray in each lobe slightly elongate, whereas in
largest specimens the inner sixth or seventh caudal rays are shortest
and of nearly equal length, the fourth and fifth from center of fin in
upper caudal lobe are greatly elongate, and the third and fourth from
center in lower lobe are greatly elongate, sometimes twice the length
of shortest caudal fin ray; no cephalic crest; a vertical line through
front edge of upper lip passes through front edge of eye; the forehead
is a little in front of front edge of upper lip.
Coloration.—In alcohol, olive-brown to light greenish brown;
darker dorsally, paler ventrally ; the posterior part of body not notably
paler than anterior part of body as is #. bicolor; middle of sides with
five to seven short narrow vertical white bars not wider than pupil
and about two to four times longer than wide, the lower edge of these
bars a little below midlengthwise axis of body, then they continue
dorsally, fading out gradually before reaching base of dorsal fin;
pectoral, dorsal, and anal fins dusky; pelvics slightly dusky; caudal
fin with outer rays white or nearly so, the middle rays dusky; behind
eye is an obliquely oblong blackish blotch, margined with pale; a pale
streak extends posteriorly from behind middle of eye a short distance;
corner of mouth white or pale.
528 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 102
Remarks.—This new species belongs to the group of species center-
ing around £. bicolor but differs from that species chiefly in coloration ;
hawaiiensis has short white bars on the midsides, whereas bicolor is
plain brown without markings.
Doubt might be cast on the source of this material, “a dry dock
hauled from Guam about a year ago,” if with this new species (which
did not occur in the extensive collections made in the Marshall and
Marianas Islands by a group of collectors during and after the war)
other species had not been taken as follows: Scorpaenopsis cacopsis
(known only from the Hawaiian Islands), Cirrhitichthys aprinus,
a specimen of Apogon, several specimens of gobies not yet identified,
2 specimens of an unidentified species of Acanthurus in the settling
stage of late metamorphosis, specimens of Pseudochromis tapeinosoma
and Tripterygion hemimelas (common to the Marshall, Marianas, and
Hawaiian Islands) having about one more scale on the average than
those from the Marshall and Marianas Islands, and Gymnothorax
undulatus (common to all three named localities). We conclude that
the fishes in this collection are endemic to the Hawaiian Islands and
were not transported in the drydock brought from Guam.
U.S. GOVERNMENT PRINTING OFFICE: t952
INDEX
(New genera, species, etc., are printed in italics)
Abbott, R. Tucker; A study of an inter- | albitarsis, Mimegralla leucopeza, 170
mediate snail host (Thiara grani- | Alecto bennetti, 266
fera) of the Oriental lung fluke | alexandri, Echinolampus, 280, 299, 300
(Paragonimus), 71
abnormalis, Echinoneus, 280, 299, 301
absarokae, Didelphodus, 189, 190, 191
abtrusus, Amplinus, 237
abyssorum, Golfingia, 395, 397, 399
Acanthaster, 291
planci, 291, 299, 300, 303
Acanthasteridae, 291
acanthifera, Munna, 119
Acanthurus sp., 528
acarina, Munna, 119, 122, 124
Acerentomidae, 305
Acerentulus travassosi, 305
aciculatus, Echinostrephus, 274, 299,
300, 301, 302, 303
Acomys, 362, 363
cahirinus, 363
Acraspis, 324
insolens, 324
patelloides, 324
acrotus, Rhinolophus, 350
actaeon, Polylepiscus, 239
Actinometra gracilis, 266
acuta, Fibularia, 279, 299, 302
acuticollis, Phinotropis, 239
acutidens, Esthonyx, 192, 203, 205
acutus, Procambarus blandingii, 217
Adapidae, 192
Addax nasomaculata, 344
Aedematosomum, 423
aegyptiaca, Vulpes vulpes, 366
aegyptiacus, Nyctinomus, 352, 353
Pteropus, 346
Spalax, 359
Vespertilio, 346
aegyptius, Erinaceus, 344
Hemiechinus, 344
Lepus, 353
aequalis, Apus, 53, 56 (table), 57, 60
(table), 61, 63, 64, 65, 66
agassizii, Phascolosoma, 372, 373, 423,
424, 429, 430, 431, 432, 433, 435
Phymosoma, 424
Physcosoma, 424, 430
agrifolia, Quercus, 337
agrifoliae, Synergus, 337
alba, Quercus, 322, 325
albicomus, Andricus, 328
albimana, Mimegralla, 161, 162, 164
(key), 165 (map), 166
Mimegralla albimana, 164, 165
(map)
Taeniaptera, 162
364736—56——_2
alexandrinus, Rattus rattus, 362
pieezcl Oryx, 344
Allopora californica, 426, 429
altidens, Didymictis, 195, 203, 205
altivelis, Plecoglossus, 109
alutaceum, Dendrostomum, 405
amamiense, Siphonosoma, 381, 382
Amaroucium californica, 158
amarula, Thiara (Thiara), 88
americanus, Leptoconops kerteszi, 476
Amitermes, 475
Ammotragus lervia, 344
Amnicolidae, 81, 111
Amphibolips, 325
globus, 325
spongifica, 326
Amphipod, new commensal, from
spiny lobster, 231
Amphilimna sexradiata, 302
Amplinus, 235, 237
abtrusus, 237
areatus, 237
armatus, 237
convexus, 237
crenus, 237
erichsoni, 237
flavicornis, 237
klugi, 237
manni, 237
niteus, 237
nitidus, 237
orphinus, 237, 238
palicaudatus, 237
tapachulae, 237
triramus, 237
vergelanus, 237
xelithus, 237
amurensis, Hua, 109
Amydrinus, 240
pongus, 240
suse Pua, Ophiocoma, 294, 299, 300,
analoga, Ianiropsis, 135, 136, 141, 142
(fig.), 143 (fig.), 144 (fig.)
Anaptomorphidae, 193
anceps, Golfingia, 396
andersoni, Gerbillus, 353
Gerbillus gerbillus, 353, 354
anderssonia, Golfingia, 390
Andricus, 327
albicomus, 328
brunneus, 327
chico, 328
529
530
Andricus—continued
crenatus, 330
foliatus, 329
kingi, 328
multicostatus, 331
niger, 329
pattersonae, 330
pilula, 333
pilularis, 333
pipula, 332
prescotti, 331
projectus, 329
utriculus, 333
Anguidae, 186
anguinea, Golfingia, 390
anomala, Asternia, 289, 299, 300
anomalus, Salarias, 515, 516, 517
Anomalobittacus, 475
Anostraca, 2, 3
antarctica, Golfingia, 390
Golfingia margaritacea, 391
Antedon brevicuneata, 303
protectus, 267
antillarum, Phascolosoma,
424, 434
Phymosoma, 434
Physcosoma, 434
antiquus, Prolimnocyon, 194, 203, 205
Antron, 333
pileus, 333
Apatemyidae, 192
Aphelidesmus, 236
Aphodiinae, 181
Aphodius, 182, 184
carolinus, 182
Aphotaenius, 181
carolinus, 182
colombiensis, 183
Aphrodite sp., 426, 427
Apiocera, 475
Apioceridae, 475
Apionsoma, 388
trichocephalus, 388
Apodis cancriformis, 69
Apogon sp., 528
appendiculata, Golfingia, 390
aprinus, Cirrhitichthys, 528
Apterobittacus, 475
apus, Lepidurus, 11, 13, 14 (table), 26,
31, 42, (table)
Apus, 2, 3, 5, 10, 11, 12, 16, 17, 18, 19,
26, 27, 28, 29, 31, 42 (table), 52,
387, 422,
54-55 (table), 56 (table), 58
(table), 63
aequalis, 53, 56 (table), 57, 60
(table), 61, 63, 64, 65, 66
australiensis, 27, 28, 42 (table),
54-55 (table), 61, 65
Ding, 12, 13, 16, 53, 61 (fig.),
cancriformis, 3, 8, 9, 12, 13, 15
(table), 16, 17, 23 (fig.), 29, 42,
perl) 52, 58, 54-55 (table), 64,
domingensis, 64
granarius, 22 (fig.), 27, 28, 42
(table), 54-55 (table), 61, 65
guildingi, 64
PROCEEDINGS OF THE NATIONAL MUSEUM
VOL. 102
Apus—continued
longicaudatus, 3, 10, 13, 17, 18, 19
(fig.), 20 (fig.), 21 (fig.), 28, 30,
42 (table), 52, 53, 54-55 (table),
56 (table), 58, 59 (fig.), 61 (fig.),
62 (table), 63 (fig.), 64, 65, 66
lucasanus, 9, 19 (fig.), 58, 56 (table),
57, 58, 59 (fig.), 60 (table), 61,
63, 64, 65, 66
namaquensis, 42 (table), 54-55
(table)
newberryi, 53, 56 (table), 57, 60
(table), 64, 65, 66
numidicus, 27, 28, 42 (table), 54-55
(table)
obtusus, 66
oryzaphagus, 9, 12, 13, 16, 53, 61
(fig.), 64, 65
sudanicus, 42 (table), 52, 53, 54-55
(table)
Arbaciidae, 268
arcassonense, Siphonosoma, 381
arcticus, Lepidurus, 6, 9, 11, 13 (table),
14 (table), 16, 17, 18, 19, 26, 30,
31, 32, 33, 34, 42 (table), 69
Monoculus, 33
arimalius, Meriones, 358
Arctiidae, 455
areatus, Amplinus, 237
armatum, Eosentomon, 306, 309, 310,
311, 312, 314
armatus, Amplinus, 237
Arthropoda, 16
Artiodactyla, 201
Arvicanthis niloticus, 360
Arvicola niloticus, 360
Asellia tridens, 350
Aspidosiphon, 375
rutilofuscus, 395
aspidosiphonoides, Phascolosoma, 395
asser, Phascolosoma, 435
Asterias laevigata, 289
multifora, 287
planci, 291
Asterina, 289
anomala, 289, 299, 300
cephea, 289, 299, 301
coronata cristata, 289, 299, 300
cristata, 289
Asterinidae, 289
Asteriscus cephus, 289
Asteroidea, 283, 299
Asteroporpa, 451, 452
australiensis, 452
hadracantha, 452
lindneri, 452
Astroboa, 453
Astrochalcis, 453
Astrochele, 451
Astrochelinae, 451
Astrocladus, 453
dofleini, 453
Astroclon, 451, 452
Astrocnida, 451, 452
Astrogamphus, 451
Astropecten, 283
polyacanthus, 283, 299, 301
Astropectinidae, 283
INDEX
Astrophyton pardalis, 453
Astrothorax, 451
Astrothrombus, 451
Ataenius, 181, 182, 183, 184
carolinus, 181, 182
lecontei, 183
Aulax latreillei, 317, 318
auritus, Plecotus, 352
Australasian stilt-legged flies sd Noa
Tylidae) in the United States
National Museum, 161
australe, Siphonosoma, 381
australica, Mimegralla contingens, 169
australiensis, Apus, 27, 28, 42 (table),
54-55 (table), 61, 65
Asteroporpa, 452
australis, Euryurus, 238, 239
Euryurus erythropygus, 238
Fibularia, 277, 299, 300, 301, 302
Golfingia eremita. 397
Ophiomyxa, 291, 299, 300
Sipunculus, 380
Vulpavus, 195, 203, 205
Australorbis, 110
Autoceras, 455
phelina, 221, 455, 458
Auturus, 235, 237, 238
dixianus, 238
evides, 238
georgianus, 238
louisianus, 238
mimetes, 238
phanus, 238
scotius, 238
avatshensis, Munna, 119
balansae, Fromia, 285, 299, 301
barnesi, Mulona, 223, 227, 229
Bassettia, 336
floridana, 336
bassettii, Diastrophus, 317
bayeri, Schizostella, 453
Beetle, dung, Aphotaenius, a new genus
of, 181
bengalensis, Ophiomyxa, 291
benhami, Golfingia, 396
bennetti, Alecto, 266
Comanthus, 266, 299, 301
bergi, Grammicomyia, 162
bicolor, Diplolepis, 320, 321
Eesenius, 510, 512, 518, 522, 523
(fig.), 525, 527, 528
Rhodites, 321
Salarias, 522
bicornis, Dryocosmus, 337
bifurcata, Schizostella, 451, 452, 453
biggsi, rare 9, 12, 13, 16, 53, 61 (fig.),
?
Bikini Atoll, list of Echinoderms, 300
billitonense, Dasmosiphon, 386
biloba, Epeiromulona, 457, 459, 460, 462
bilobatus, Lepidurus, 10, 11, 18, 25
(fig.), 26, 28, 30, 31, 32, 36
(table), 38, 42 (table), 53
Bilobus, 30, 31
binghami, Calobata, 168
Mimegralla, 168
birostris, Manta, 246, 248, 254
531
bispinosa, Ophiomastix, 296, 299, 300
Bittacidae, 475
blandum, Dendrostoma, 406, 409
Dendrostomum, 404, 405
Blennius, 509
Boavus, 186
occidentallis, 186
boeckii, Munna, 118, 119, 134
boholense, Siphonosoma, 381, 382
bombonus, Thrinoxethus, 243
bonhouri, Siphonosoma, 381, 382
borealis, Eotitanops, 203
Bouteloua curtipendula, 490, 493
Boysen Reservoir area, 204 (map)
Preliminary analysis of the verte-
brate fossil fauna of the, 185
branchiatum, Xenosiphon, 377
branchiatus, Sipunculus, 377
Sipunculus mundanus, 377
Branchiopoda Notostraca, Contributions
to the morphology and taxonomy of,
with special reference to the North
American species, 1
Branchipus, 3
braasten st Epeiromulona hamata, 458,
465
braueri, Phinotropis, 239
brevicauda, Pupulina, 258
brevicornis, Saphonecrus, 315
Synergus, 315
brevicuneata, Antedon, 303
brevidens, Glyptosaurus, 189 (table)
brevipes, Ophiocoma, 296, 299, 301, 302
breviremis, Ianiropsis, 134, 145
brevispina, Ophiomyxa, 302
Brissus, 283
latecarinatus, 283, 299, 300, 301,
303
Brittle-star, Schizostella, a new genus of,
451
Brontotheriidae, 200
Brotia, 94
brownorum, Heptodon, 201, 203, 205
brunneus, Andricus, 327
Bugula pacifica, 122
Bulinus, 110
Bunophorus, 201
etsagicus, 201, 202 (fig.), 203, 205
burmanicus, Salarias, 522, 524
Bythoceros, 88
cacopsis, Scorpaenopsis, 528
cahirinus, Acomys, 363
Mus, 363
eainarachus, Thrinoxethus, 243
calamaris, Echinothrix, 269, 299, 301,
302
Kehinus, 269
California, and Baja California, Sipun-
culid worms of, 371
Some marine asellote isopods from
northern, 117
californianus, Mytilus, 426
californica, Allopora, 426, 429
aroucium, 158
Golfingia eremita, 389, 396
Janiropsis, 135
Rosa, 321
532
californiense, Pleurogonium, 149
californiensis, Golfingia margaritacea,
373, 389, 392
Calliaster, 284
elegans, 284, 299, 300
Callirhytis, 338
cistella, 338
corrugis, 338
crypta, 336
fructuosa, 338
fulva, 340
furva, 341
guadaloupensis, 324
hopkinsi, 339
infuscata, 341
modesta, 338
oblata, 340
operator, 339
subcostata, 341
Calobata binghami, 168
cedens, 168
chrysopleura, 168
coeruleifrons, 168
confinis, 168
contingens, 169
contraria, 166
(Neocalobata) deferens, 167
galbula, 166
sepsoides, 166
striatofasciata, 167
territa, 179
Calycopteryx, 173
Cambarus, 210
lecontei, 210
cancellata, Neoferdina, 285
cancriformis, Apodis, 69
Apus, 3, 8, 9, 12, 13, 15 (table), 16,
17, 23 (fig.), 29, 42 (table), 52,
538, 54-55 (table), 64, 67
Triops, 67
Triops (Apus), 67
Canidae, 365
Canis lupaster, 365
niloticus, 366
capensis, Golfingia, 390
capsiforme, Golfingia, 390
Carasius, 109
carditoides, Petricola, 419
Carnivora, 194, 365
carolinense, Dasmosiphon, 386
carolinus, Aphodius, 182
Aphotaentius, 182
Ataenius, 181, 182
Cartwright, 0. L.; Aphotaenius, a new
genus of dung beetle (Coleop-
tera: Scarabaeidae), 181
catenaria, Ceriphasia, 113
Pleurocera, 113, 114 (fig.)
catherinae, Golfingia, 400, 401
catus, Felis, 368
cedens, Calobata, 168
Cenometra bella magnifica, 303
Central American milliped, a new genus
of, 235
Cephalotes teniotis, 352
cephea, Asterina, 289, 299, 301
cephus, Asteriscus, 289
Ceraptopterus, 246
PROCEEDINGS OF THE NATIONAL MUSEUM
VOL. 102
Cerataulax, 472, 480, 484
per ee at 472, 480, 484, 498,
vittatus, 472
Ceratoptera, 246
Cerianthus sp., 389, 395
Ceriphasia catenaria, 113
Ceroptres, 316, 317
obtusilobensis, 317
sp., 338
chamense, Meniscotherium, 196
Chapman, Wilbert M., and Leonard P.
Schultz; Review of the fishes of
the Blennioid genus Ecsenius,
with descriptions of five new
species, 507
charcoti, Golfingia, 390
chaus, Felis, 368
Chenopodium oblongi, 492
pratericola desiccatum, 493
chico, Andricus, 328
Chiroptera, 346
Chloris radiata, 485
Chondrocidaris, 267
gigantea, 267, 299, 300
problepteryx, 268
chromatocephala, Munna, 119, 120, 128,
129 (fig.), 131 (fig.)
chrysolepis, Quercus, 323, 324, 329
chrysopleura, Calobata, 168
Mimegralla cedens, 168
chuni, Golfingia, 396
Cidaridae, 267
Cidaris toreumatica, 270
Cidarites imperialis, 268
metularia, 267
Cincia phelina, 458
cincta, Golfingia, 396
Ophiolepis, 298, 299, 302
cinerea, Golfingia, 396
Circulifer, 476
tenellus, 476
Cirrhitichthys aprinus, 528
cistella, Callirhytis, 338
Clark, Austin H.; Echinoderms from
the Marshall Islands, 265
Schizostella, a new genus of brittle-
star (Gorgonocephalidae), 451
clippertoni, Ophiocomella, 296, 297,
299, 300, 301, 302
Cloeosiphon, 375
cluthensis, Golfingia, 390
Clypeaster, 276
reticulatus, 276, 299, 300
Clypeastridae, 276
coccinea, Quercus, 340
Coecimunna, 119
Coelopleurus, 268
sp 268, 299, 300
coeruleifrons, Calobata, 168
Mimegralla, 168, 171
Coleoptera, 181, 246
colombiensis, Aphotaenius, 183
Epeiromulona hamata, 457, 464
Comanthus, 266
bennetti, 266, 299, 301
Comaster, 266
gracilis, 266, 299, 300
INDEX
Comaster—continued
multifida, 266
Comasteridae, 266
comata, Liodora, 325
commensalis, Parapleustes,
fig.)
Conchostraca, 2, 3, 16
Condylarthra, 196
confinis, Calobata, 168
Mimegralla, 168
confusa, Golfingia, 396
contingens, Calobata, 169
Mimegralla, 169
Mimegralla contingens, 169
contraria, Calobata, 166
Mimegralla albimana, 164,
RAD) 166
convexus, Amplinus, 237
Copepod genus Pupulina (Caligoida),
an amended diagnosis of, 245
Coprinus, 334
coquilletti, Trichoteras, 323
coriacea, Golfingia, 396
corrugis, Dryophanta, 336
Callirhytis, 338
Coryphodon, 197
sp., 197, 203, 205
Coryphodontidae, 197
costata, Thiara (Melanoides), 88
Cothornobata, 173
couesii, Lepidurus, 3, 11, 17, 18, 23 (fig.),
24, (fig.,, 25 (fig.), 26, 27, 30, 31,
32, 34, 36 (table), 38, 42 (table),
51, 52
231, 233
165
crassicaudus, Suncus, 345
crassum, Hesperosiphon, 386
crassus, Meriones, 358
Crayfish, new from Alabama, with notes
on Procambarus lecontei
(Hagen), 209
crenatus, Andricus, 330
crenulata, Stenomelania, 94
crenus, Amplinus, 237
Crepidochaetus, 173
Cricetidae, 353
Crinoidea, 266, 299, 302
cristata, Asterina, 289
Asterina coronata, 289, 299, 300
Crocidura, 345, 346
floweri, 345
olivieri, 345
religiosa, 345
Crosa, 173, 176
semilauta, 175 (fig.), 177
tetras, 177
yapensis, 175 (fig.), 177
crypta, Callirhytis, 336
Cryptosomum, 374, 423
Ctenopharyngodon, 109
cubanianus, Hemisinus,
114 (fig.), 115
Culcita, 285
ge aN ea aoa 285, 299, 300, 301,
08,) 794,113,
cumanense, Dasmosiphon, 386
cuneifrons, Eurybata, 178
curtipendula, Bouteloua, 490, 493
cuscutaeformis, Diastophus, 317
533
cyanea, Trepidarioides, 180
yelestheria hislopi, 16
Cyclosphen samoanus, 167
tonganus, 167
cyclostomus, Echinoneus, 280, 299, 300,
301, 302
cylindrata, Golfingia, 390
cylindrifer, Stylinodon, 193, 203, 205
Cynipidae, 319
Cynips glechomae, 317, 320
obtusilobae, 317
papula, 338
quercus obtusilobae, 317
rosae, 319
Cynodontomys, 191
lundeliusi, 191 (fig.), 203, 205
scottianus, 191, 203, 205
Cyprinus, 109
Cypselosoma, 173
Cyrtechinus, 271
verruculatus, 271, 299, 300, 302
cystops, Rhinopoma, 347
Dactylosaster, 286
cylindricus pacificus, 286, 299, 300
Dasmosiphon, 386
billitonense, 386
carolinense, 386
cumanense, 386
edule, 386
formosa, 386
hataii, 386
koreae, 386
dayi, Siphonosoma, 381
deferens, Calobata (Neocalobata), 167
degeneri, Ophiodesmus, 294, 299, 300
dehaanii, Potamon, 110
dehamata, Dendrostomum, 406
delagei, Golfingia, 396
delicatum, Eosentomon armatum, 311,
312
Deltatheridiidae, 189
Deltatheridium, 190
demessa, Ophiothrix, 302
Dendrostoma, 373, 374, 404, 405
blandum, 406, 409
hexadactylum, 410
mytheca, 411, 415
perimeces, 415
petraeum, 406
pyroides, 406, 409
zostericola, 411
Dendrostomum, 373, 374, 404, 405
(rey), 406 (key), 413 (fig), 435,
alutaceum, 405
blandum, 404, 405
dehamata, 406
dyscritum, 372, 404, 405, 406, 413
(fig.), 414, 417, 420, 421
ellipticum, 405
hexadactylum, 372, 404, 405, 406,
410
lissum, 406, 419, 422
minor, 405
perimeces, 372, 404, 405, 406, 415
peruvianum, 405, 411, 414, 415,
422
534
PROCEEDINGS
Dendrestomum—continued
pinnifolium, 405
pyroides, 372, 373, 404, 405, 406,
410, 419, 426
schmitti, 406, 413 (fig.), 422
signifier, 405
spinifier, 405
stephensoni, 406
tropicum, 405
zostericolum, 372, 404, 405, 410,
411, 413 (fig.), 415, 416, 417,
418, 422
dentigerum, Phascolosoma, 423, 432
Phymosoma, 432
depressa, Golfingia, 395
depressum, Laganum,
301, 302
derjugini, Inniropsis kincaidi, 135, 136,
137 (fig.), 138 (fig.), 139 ‘(fig.),
140 (fig.)
Janitopers 135, 139
desiccatum, Chenopodium pratericola,
276, 299, 300,
Desmechinus, 270
rufus, 270, 299, 300
devillei, Pycnotropis, 240
Diacodexis, 202
olseni, 202, 203, 205
diadema, Echinus, 269
Echionthrix, 269, 299, 300, 301
Diadematidae, 269
Diastrophus bassettii, 317
cuscutaeformis, 317
nebulosus, 316
radicum, 317
Dichobunidae, 201
dicrana, Metalia, 282, 299, 300, 301
Dictypharidae, 475
Didelphodus, 189
absarokae, 189, 190, 191
ventanus, 189, 190 (fig.), 203, 205
Didymictis, 195
altidens, 195, 203, 205
Diorchitrema, 111
formosanum, 109
Diplolepis, 319
bicolor, 320, 321
fulgens, 321
ignoto, 316
lens, 320
polita, 321
quercus, 319
q.-obtusilobae, 317
rosae, 320
rosaefolii, 320
terrigena, 320
Diplopoda, 16
Dipodidae, 364
Dipodillus mariae, 355
Dipus gerbillus, 354
pyramidum, 356
diversicolor, Quercus, 333
dixianus, Austurusm 238
dofleini, ’Astrocladus, 453
domesticus, Mus musculus, 363
domingensis, Apus, 64
donohoet, Glyptosaurus, 186, 187 (fig.),
189 (table)
OF THE NATIONAL MUSEUM
VOL. 102
doreas, Gazella, 344
Dorometra nana, 303
douglasii, Quercus, 327, 330
Dryocosmus, 336
bicornis, 337
Grumatus, 336
minusculus, 337
Dryophanta corrugis, 336
dubia, Golfingia, 390
dubiosa, Ophiopezella, 302
dumosa, Quercus, 327, 330
dyscritum, Dendrostomum, 372, 404,
405, 406, 413 (fig.), 414, 417,
420, 421
ebenina, Melania, 109
Echinasteridae, 290
Echinocyamus, 279
elongatus, 279, 299, 300
incertus, 279, 299, 300
megapetalus, 279, 299, 300, 301, 302
aia ha from the Marshall Islands,
monmoiies 267, 299
Echinolampus, 280
alexandri, 280, 299, 300
alexandri sibogae, 282
Echinometra, 272
mathaei, 272, 299, 300, 301, 302
mathaei. oblonga, 274, 299, 300,
301, 302
Echinometridae, 272
Echinoneidae, 280
Echinoneus, 280
abnormalis, 280, 299, 301
cyclostomus, 280, 299, 300, 301, 302
Echinostrephus, 274
aciculatus, 274, 299, 300, 301, 302,
303
molaris, 303
Echinothrix, 269
calamaris, 269, 299, 301, 302
diadema, "269, 299, 300, 301
Echinus calamaris, 269
diadema, 269
globulus, 270
gratilla, 271
mamillatus, 276
mathaei, 272
oblongus, 274
ovulum, 278
reticulatus, 276
trigonarius, 275
Eesenius, 507, 510 (table), 511 (key)
bicolor, 510, 512, 513, 522, 523
(fig.), 525, 527, 528
frontalis, 511, 512, 5138, 514, 525
gravieri, 511, 513
hawatiensis, 510, 512, 526, 527
(fig.)
Hetienele. 510, 511, 515, 517, 518
(fig.)
mandibularis, 507, 508, 509, 511,
518
namiyei, 525 (fig.)
opsifrontalis, 510, 512, 517,
520,
521 (fig.), 526 (table)
INDEX
Ecsenius—continued
PED cet 510, 512, 517, 519 (fig.).
5
pulcher, 511, 515
Review of the fishes of the Blen-
nioid genus, with descriptions of
5 new species, 507
stigmatura, 510, 514 (fig.), 526
(table)
edule, Dasmosiphon, 386
Egregia sp., 139, 145
Egypt, Mammals from the Nile Delta
region of, 343
egyptiacus, Pteropus, 346
Rousettus, 346
elachea, Golfingia, 390, 395, 399
elegans, Calliaster, 284, 299, 300
Ophiothrix, 293, 299, 301
ellipticum, Dendrostomum, 405
elongata, Golfingia, 390
elongatus, Echinocyamus, 279, 399, 300
aralebion, 248, 254
Emballonuridae, 348
emoryi, Quercus, 339
Enchelyurus sp., 512
engelmanni, Quercus, 331
Eniwetok Atoll, list of Echinoderms, 301
eniwetoki, Siphonosoma, 381
Eosentomidae, 305
Eosentomon armatum, 306, 309, 310,
311, 312, 314
armatum delicatum, 311, 312
germanicum, 306, 309
pallidum, 306, 309, 311
ribagai, 306, 309
rostratum, 306
transitorium, 309, 311
venezuelense, 305, 307 (fig.), 308
fig.
yosemitense, 311
Kotitanops, 201
borealis, 203
sp., 201, 205
Epeiromulona, 455, 457 (key), 469
biloba, 457, 459, 460, 462
hamata, 462, 463, 466
hamata brasiliensis, 458, 465
hamata colombiensis, 457, 464
bes hamata, 457, 458, 462, 464,
hamata venezuelensis, 457, 463, 464
icterinus, 456, 457, 466, 467
eprne, 455, 457, 458, 459, 466,
phelina, 458, 459, 462, 465, 466,
468, 469
roseata, 457
thysanata, 457, 466
epilittoralis, Taniropsis, 136, 149, 150
(fig.), 151 (fig.), 156
Eptesicus innesi, 352
Equidae, 198
pemnite eearngls, 389, 395, 396, 398,
Phascolosoma, 396
Phascolosomum, 396
Sipunculus, 395, 396
erichsoni, Amplinus, 237
535
Erinaceidae, 344
ae Ophiocoma, 295, 299, 300,
i!
Hrinaceus aegyptius, 344
libyeus, 344
Eriocheir japonicus, 110
erythropygus, Euryurus, 238
Polydesmus, 238
erythrourus, Meriones, 358
Esthonyx, 192
acutidens, 192, 203, 205
etsagicus Bunophorus, 201 202 (fig.),
203, 20
Eucidaris, 267
saa laa 267, 268, 299, 300, 301,
3
Eudiocrinus tenuissimus, 303
Kumayria, 335
invisa, 335
Eupariina, 182 *
Eurybata, 161,°173, 174 (key), 176
cuneifrons, 178
eee 161, 173, 174; 175 (fig:);
176
Eurybata nigritibia, 174, 175 (fig.)
petasibarba, 178
semilauta, 161, 173, 176, 177
tessellata, 174, 175 (fig.), 176
tetras, 177
eurydale Rhinolophus, 350
Euryuridae, 235, 236, 237 (key)
Euryurinae, 237
Euryurus, 235, 236, 237, 238
australis, 238, 239
erythropygus, 238
erythropygus australis, 238
falcipes, 238, 239
eusticha, Fromia, 286, 299, 300
evides, Auturus, 238
excerta, Swiftia, 453
exigua, Ophiothrix, 293, 299, 301
exiguus, Miacis, 195
extrema, Mimegralla albimana, 164, 165
(map), 167
falcata, Quercus, 340
falcipes, Euryurus, 238, 239
Felidae, 368
Felis catus, 368
chaus, 368
Ferdina ocellata, 285
Fibularia, 277
acuta, 279, 299, 302
australis, 277, 299, 300, 301, 302
ovulum, 278, 299, 300, 301, 302
volva, 278, 299, 300, 301, 302
Fibulariidae, 277
Field, William D.; Moths of the genera
Mulona Walker and Lomuna, a
new and closely related genus
(Arctiidae: Lithosiinae), 221
Moths of the genus Epeiromulona,
a new genus of Lepidoptera, 455
filiformis, Golfingia, 395
Filigorgia sp., 453
fimbriata, Golfingia, 395
finmarckica, Golfingia margaritacea, 392
536
Fisher, Walter Kenrick; The sipunculid
worms of California and Baja
California, 371
fisheri, Mithrodia, 303
Fishes, description of the Blennoid
genus Ecsenius, 507
flagrifera, Golfingia, 395
flavicornis, Amplinus, 237
flavocarinata, Pycnotropis, 240
Flies, Australasian stilt-legged (Diptera:
Tylidae) in the United States
National Museum, 161
flores, Lepeophtheirus, 248
Pupulina, 245, 246, 248, 254, 255,
256, 257, 258, 259, 260, 261, 262
floridana, Bassettia, 336
floweri, Crocidura, 345
foliolata, Luidia, 149
foliatus, Andricus, 329
Formicosepsis, 173
formosa, Dasmosiphon, 386
formosanum, Diorchitrema, 109
Fromia, 285
balansae, 285, 299, 301
eusticha, 286, 299, 300
hemiopla, 286, 299, 300
frontalis, Ecsenius, 511, 512, 513, 514, 525
Salarias, 512
fructuosa, Callirhytis, 338
frugivorous, Rattus rattus, 362
fulgens, Diplolepis, 321
fulva, Callirhytis, 340
funafuti, Siphonosoma, 381, 382
furcatus, Salarias, 522, 524
furcifer, Polylepiscus, 239
urva, Callirhytis, 341
fusca, Golfingia, 390
galapagensis, Sipunculus, 376
galatheae, Ophiothrix, 302
galbula, Calobata, 166
Mimegralla albimana, 164, 165
(map), 166
Galls, New American cynipid wasps
from, 315
gambelii, Quercus, 332, 333
Gambusia, 109
garamantis, Gerbillus, 355
Gerbillus nanus, 355
arryana, Quercus, 327, 328, 331
astropoda, 72
Gaudeator, 221, 455, 456, 457
phelina, 459, 466, 467
Gazella doreas, 344
gentilis,s Mus musculus, 363
georgiana, Golfingia, 390
georgianus, Auturus, 238
gerbillus, Dipus, 354
Gerbillus, 354
Gerbillus andersoni, 353
garamantis, 355
gerbillus, 354
gerbillus andersoni, 353, 354
gerbillus gerbillus, 354
henleyi mariae, 355
nanus garamantis, 355
pyramidum pyramidum, 356
pyramidum tarabuli, 356
PROCEEDINGS OF THE NATIONAL MUSEUM
VOL, 102
germanicum, EKosentomon, 306, 309
gigantea, Chondrocidaris, 267, 299, 300
giganteus, Glyptosaurus, 189 (table)
glacialis, Golfingia, 395
Lepidurus, 33, 66
Glance, Grace; A new species of insect
of the order Protura, 305
Glaphytus} Pachychilus, 113, 114 (fig.),
1
glauca, Golfingia, 395
Glechoma hederacea, 318
glechomae, Cynips, 317, 320
Liposthenes, 317
globulus, Echinus, 270
Mespilia, 270, 299, 300, 301, 302
globus, Amphibolips, 325
glossipapillosa, Golfingia, 390
Glyptosaurus, 186, 188 (key),
(table)
brevidens, 189 (table)
donohoet, 186, 187 (fig.), 189 (table)
eteaetous, 189 (table)
illsi, 187, 188, 189 (table)
montanus, 188, 189 (table)
nodosus, 189 (table)
obtusidens, 189 (table)
princeps, 189 (table)
sphenodon, 189
sylvestris, 189 (table)
tuberculatus, 189 (table)
ones, 372, 373, 374, 380, 388, 389
ey
abyssorum, 395, 397, 399
anceps, 396
anderssoni, 390
anguinea, 390
antarctica, 390
appendiculata, 390
benhami, 396
capensis, 390
capsiforme, 390
catherinae, 400, 401
charcoti, 390
chuni, 396
cincta, 396
cinerea, 396
cluthensis, 390
confusa, 396
coriacea, 396
cylindrata, 390
delagei, 396
depressa, 395
dubia, 390
elachea, 390, 395, 399
elongata, 390
eremita, 389, 395, 396, 398, 399
eremita australis, 397
erelmita californica, 389, 396
filiformis, 395
fimbriata, 395
flagrifera, 395
fusca, 390
georgiana, 390
glacialis, 395
glauca, 395
glossipapillosa, 390
gouldii, 383, 388, 393, 403
hanseni, 390, 391
189
Golfingia—continued
harveii, 390
hespera, 373, 389, 393
hozawai, 400, 401
hudsoniana, 390
hyugensis, 400, 401
tkedai, 390
intermedia, 396
improvisa, 396
johnstoni, 396
laetmophila, 389, 395, 397
liljeborgi, 395
lutea, 390
macginitiet, 373, 390, 400, 401,
436
macintoshii, 388, 389, 390
macra, 395
margaritacea, 373, 390, 391,
398
margaritacea antarctica, 391
margaritacea californiensis,
389, 392
margaritacea finmarckica, 392
margaritacea meridionalis, 391
margaritacea siberica, 391, 392
martensi, 400, 401
minuta, 396
misakiana, 393, 395
mucida, 395
muricaudata, 390
nigra, 400, 401
nordenskjéldi, 390
noto, 390
novaezealandiae, 395
obscura, 390
ohlini, 390
okinoseana, 390
onagawa, 400, 401
ownstoni, 390
papillifera, 396
papillosa, 390
pavlenkoi, 400
pellucida, 396
prioki, 395
procera, 390, 400, 401, 402
pugettensis, 373, 390, 400, 401
pyriformis, 400, 401
recondita, 396
rutilofusca, 395
sabellariae, 396
sanderi, 390
sarsii, 396
semperi, 400, 401
signa, 390
sluiteri, 396
socia, 390
soyo, 390
subhamata, 396
trybomi, 390, 391
valida, 390
verrilli, 395
vulgaris, 373, 389, 390
zenzibakensis, 400, 401
Gongylocephala, 173, 178
pallida, 178
pallida luzonica, 178
Goniasteridae, 284
Goniobasis, 111
364736—56——3
INDEX 537
402,
392
373,
Gorgonocephalidae, 451, 452
Schizostella, a new genus of, 451
Gorgonocephalinae, 451
gouldii, Golfingia, 383, 388, 393, 403
Sipunculus, 393
gracilis, Actinometra, 266
Comaster, 266, 299, 300
Grammicomyia, 162, 171
hergi, 162
halla, 162, 163 (fig.)
sondaica, 162
granarius, Apus, 22 (fig.), 27, 28, 42
(table), 54-55 (table), 61, 65
granifer, Saat saad 286, 299, 300, 301,
30
Tarebia, 72
Thiara, 71, 72, 74 (table), 75 (fig.),
77 (fig.) 79, 82 (fig.), 83 (fig.), 87
(fig.), 89 (fig.), 90, 91 (fig.), 93
(table), 95 (fig.), 96 (fig.), 97, 98,
100 (table), 101 (fig.), 102 (fig.),
104 (table), 105 (fig.), 109, 110,
elds
Thiara (Tarebia), 97, 113, 114 (fig.)
granulatum, Phascolosoma, 388, 422,
423, 429
Phymosoma, 388, 423
Physcosoma, 423
gratilla, Echinus, 271
Tripneustes, 271, 299, 301
gratiosa, Parasalenia, 272, 299, 300, 301
gravieri, Ecsenius, 511, 513
Salarias, 513
grisea, Mulona, 221, 223, 228, 229
grumatus, Dryocosmus, 336
guadaloupensis, Callirhytis, 324
Paracraspis, 324
guildingi, Apus, 64
guildingii, Linckia, 288, 299, 300, 302
guttatum, Zopheroteras, 334
Gymnothorax undulatus, 528
hadracantha, Asteroporpa, 452
haematobium, Schistosoma, 110
haenschi, Pyenotropis, 240
halapense, Sorghum, 490
haleyi, Munna, 119, 120, 132 (fig.), 133
(fig.)
halli, Grammicomyia, 162, 163 (fig.)
hamata, Epeiromulona, 462, 463, 465
Epeiromulona hamata, 457, 458,
462, 464, 465
hanseni, Golfingia, 390, 391
hansenii, Stephanostoma, 388
Haplorchis, 111
harveii, Golfingia, 390
hataii, Dasmosiphon, 386
hawaiiensis, Eecsenius, 510, 512, 526,
527 (fig.)
hayi, Procambarus, 217
hederacea, Glechoma, 318
Heleidae, 476
heliaster, Himerometra, 303
Hemiechinus aegyptius, 344
hemimelas, Triptweygion, 528
hemiopla, Fromia, 286, 299, 300
Hemiptera, 471
538
Hemisinus, 79
cubanianus, 78, 79, 113, 114 (fig.),
5
11
Heptodon, 201
brownorum, 201, 203, 205
hermosus, Thrinoxethus, 243
Herpestes ichneumon, 368
hespera, Golfingia, 373, 389, 393
Phascolosoma, 393
Hesperapis, 475
Hesperosiphon, 386
crassum, 386
parvum, 386
vastum, 386
Heterocentrotus, 275
mammillatus, 276, 299, 301
trigonarius, 275, 299, 300, 301
Heterophyidae, 109, 111
Heteroptera, 486
heterosculptus, Polylepiscus, 239
hexadactylum, Dendrostoma, 410
ee, 372, 404, 405, 406,
10
hexapla, Eurybata, 161, 173, 174, 175
(fig.), 176
hillsi, Glyptosaurus, 187, 188, 189 (table)
Himerometra heliaster, 303
Hipposideridae, 350
hislopi, Cyclestheria, 16
Hobbs, Horton H., Jr.; A new crayfish
from Alabama, with notes on
a ie lecontei (Hagen),
9
Hoffman, Richard L.; A new genus of
Central American milliped (fam-
ily Euryuridae), with notes on
the American genera, 235
Homalosoma, 388, 389
hopkinsi, Callirhytis, 339
hozawai, Golfingia, 400, 401
Hua, 111
amurensis, 109
sp., 110
toucheana, 109
hudsoniana, Golfingia, 390
Hyaenodontidae, 194
Hyopsodontidae, 196
Hyopsodus, 196, 197
paulus, 197
powellianus, 196, 203, 205
wortmani, 197, 203, 205
hypoleuca, Quercus, 339
hypostoma, Mobula, 248, 254, 255
Hyracotherium, 198, 200
venticolum, 198 (fig.), 208, 205
hyugensis, Golfingia, 400, 401
Tais, 135
Ianiropsis, 117, 134, 136 (key)
analoga, 135, 136, 141, 142 (fig.),
148 (fig.), 144 (fig.)
breviremis, 134, 145
epilittoralis, 136, 149, 150 (fig.), 151
(fig.), 155
kineaidi derjugini, 135, 136, 137
tae’ 188 (fig.), 189 (fig.), 140
g.
PROCEEDINGS
OF THE NATIONAL MUSEUM
VOL. 102
Janiropsis—continued
kincaidi kincaidi, 117, 135, 136, 138
(fig.), 141, 151, 156
magnocula, 117, 136, 145, 146 (fig.),
147 (fig.), 148 (fig.)
minuta, 136, 155 (fig.), 157
montereyenis, 136, 152, 153 (fig.),
154 (fig.), 158
pugettensis, 135, 136, 139
tridens, 136, 156, 157 (fig.)
ichneumon, Herpestes, 368
Viverra, 368
irae eenoneane, 456, 457, 466,
Ictops, 190
ignoto, Diplolepis, 316
ikedai, Golfingia, 390
imbricaria, Quercus, 339
imbricata, Ophiolepis, 298
imbricatus, Ophioplocus, 298, 299, 300
imperialis, Cidarites, 268
Phyllacanthus, 268, 299, 300
improvisa, Golfingia, 396
incertus, Echinocyamus, 279, 299, 300
incus, Ptyxogon, 240
indica, Mecidea, 473 (map), 474, 480,
481, 493, 500
infuscata, Callirhytis, 341
ingens, Siphonomecus, 382
Siphonosoma, 372, 381, 382
ingramsi, Mecidea, 494, 495
innesi, Eptesicus, 352
Insectivora, 189, 344
insolens, Acraspis, 324
Paracraspis, 324
intermedia, Golfingia, 396
interruptus, Panulirus, 231, 232
invisa, Eumayria, 335
iquitus, Thrinoxethus, 243
Ischyromyidae, 194
Isectilophidae, 201
ismahelis, Meriones, 358
Isopods, Some marine asellote isopods
from northern California, with
descriptions of nine new species,
117
Istiblennius, 507, 508, 509
jaculus, Jaculus jaculus, 364
Mus, 364
Jaculus jaculus jaculus, 364
Janira maculosa, 141, 145
Janiridae, 117, 134
Janiropsis, 134
californica, 135
derjugini, 135, 139
kincaidi, 135, 136, 139
japonicum, Phascolosoma, 390, 429
Physcosoma, 390, 424
Schistosomum, 110
japonicus, Eriocheir, 110
Johnston Atoll, list of Echinoderms, 301
johnstoni, Golfingia, 396
kalonotus, Phinotropis, 239
keiensis, Mimegralla albimana, 164, 167
kellicottii, Paragonimus, 111
kelloggii, Quercus, 337
INDEX
kerteszi, Leptoconops, 476
kinbergi, Ophiura, 298, 299, 302
kincaidi, Ianiropsis kincaidi, 117, 135,
136, 138 (fig.), 141, 151, 156
Janiropsis, 135, 136, 139
kingi, Andricus, 328
kirkii, Lepidurus, 11, 31, 42 (table)
klugi, Amplinus, 237
Polydesmus, 237
koreae, Dasmosiphon, 396
kristenseni, Mecidea, 473 (map), 474,
ih 480, 482, 491, 494, 495, 497,
50
krgyeri, Munna, 119, 124, 125
kuhlii, Pipistrellus, 351, 352
Vespertilio, 351
kusaieana, Mimegralla ponapensis, 170
ee Atoll, list of Echinoderms,
1
laetmophila, Golfingia, 389, 395, 397
laevigata, Asterias, 289
inckia, 289, 299, 301, 302
laevis, Quercus, 341
Laganidae, 276
Laganum, 276
depressum, 276, 299, 300, 301, 302
Lagomorpha, 353
Lambdotherium, 200
popoagicum, 200, 203, 205
Laminaria sp., 139, 145, 158
Lamprometra palmata, 303
lamprus, Thrinoxethus, 243
lapidens: Mulona, 221, 222, 223, 226,
9
pee ee, Brissus, 283, 299, 300, 301,
303
Spatangus brissus, 283
latidens, Miacis, 195, 203, 205
latreillei, Aulax, 317, 318
latzeli, Pycnotropis, 240
lecontei, Ataenius, 183
Cambaris, 210
Procambarus, 209, 210, 211 (fig.)
lemmoni, Lepidurus, 26, 28, 30, 33, 34,
35, 50, 51
lens, Diplolepis, 320
lenticularis, Limnadia, 68
Lepeophtheirus florea, 248
lephina, Epeiromulona, 455, 457, 458,
459, 466, 468
lepida, Ophiothrix, 293, 299, 300
Lepidoptera, Moths of the genus Epei-
romulona, a new genus of, 455
Lepidurus, 2, 3, 4, 5, 9, 10, 11, 12, 16,
17, 18, 19, 21, 22, 23, 26, 27, 28,
29, 30, 31, 32 (key), 35, 42 (table);
51, 52, 53
apus, 11, 13, 14 (table), 26, 31,
38, 42 (table)
arcticus, 6, 9, 11, 13 (table), 14
(table), 16, 17, 18, 19, 26, 30, 31,
32, 33, 34, 42 (table), 69
bilobatus, 10, 11, 18, 25 (fig.), 26,
28, 30, 31, 32, 36 (table), 38, 42
(table), 53
539
Lepidurus—continued
couesii, 3, 11, 17, 18, 23 (fig.), 24
fig.), 25 (fig.), 26, 27, 30, 31, 32,
Be 36 (table), 38, 42 (table), 51,
glacialis, 33, 66
kirkii, 11, 31, 42 (table)
lemmoni, 26, 28, 30, 33, 34, 35, 50,
51
lyncht, 39, 40 (fig.), 41 (fig.), 42
(table), 43 (fig.), 44 (fig.), 45
(e.), 46 (fig.), 47 (fig.), 48 (fig.),
lynchi echinatus, 32, 42 (table), 49
(fig.), 51
macrurus, 3, 17, 34, 35, 38
packardi, 11, 12, 17, 23, 24 (fig.),
26, 28, 31, 32, 33, 36 (table), 42
(table)
patagonicus, 17, 31, 42 (table)
viridis, 11, 31, 42 (table)
lepineyi, Mecidea, 473, 479, 480, 483
499, 501, 502
Leporidae, 353
Leptoconops, 476
kerteszi, 476
kerteszi americanus, 476
Lepus aegyptius, 353
rothschildi, 353
lervia, Ammotragus, 344
leucopeza, Mimegralla, 170
levigatus, Protaphelidesmus, 240
libertina, Semisulcospira, 109, 110
libyca, Mustela, 367
Poecilictis, 367
libycus, Erinaceus, 344
Meriones, 357, 358
Meriones libycus, 357
ligula, Platyrrhacus, 240
Protaphelidesmus, 240
liljeborgi, Golfingia, 395
Limnadia, 16
lenticularis, 68
Linckia, 287
guildingii, 288, 299, 300, 302
laevigata, 289, 299, 301, 302
multifora, 287, 299, 300, 301
Linckiidae, 285
Linder, Folke; Contributions to the
morphology and taxonomy of the
Brachipoda Notostraca, with spe-
cial reference to the North Ameri-
can species, 1
lindneri, Asteroporpa, 452
linearis, Mecidea, 472, 479, 480, 483,
497, 498, 499, 501
Liodora, 325
comata, 325
Liposthenes, 317
glechomae, 317
lissum, Dendrostomum, 406, 419, 422
Lithacrosiphon, 375
Lithosiinae, 455
lividanalis, Eesenius, 510, 511, 515, 517,
518 (fig.)
lobata, Quercus, 327, 329
540
Lobster, spiny, a new species of com-
mensal Amphipod from, 231
Lomuna, 222 (key), 228
longicaudatus, Apus, 3, 10, 13, 17, 18,
19 (fig.), 20 (fig.), 21 (fig.), 28,
30, 42 (table), 52, 53, 54-5
(table), 56 (table), 58, 59 (fig.),
61 (fig.), 62 (table), 63 (fig.), 64,
65, 66
lenge Ophiothrix, 2938, 299, 300,
302
Ophiura, 293
longula, Mecidea, 473 (map), 476, 478,
479, 480, 484, 486, 490, 491, 495,
497, 500
lordi, Phascolosoma, 424
lorioli, Ophidiaster, 287, 299, 301
nigripuncta, 221, 228
louisianus, Auturus, 238
Loveina, 193
vespertina, 193
zephyri, 193, 203, 205
lucasana, Mobula, 245, 258
lucasanus, Apus, 9, 19 (fig.), 53, 56
(table), 57, 58, 59 (fig. ye 60
(table), 61, 63, 64, 65, 66
Luidia, 284
foliolata, 149
mascarena, 284, 299, 300
Luidiidae, 284
lunaria, Mimegralla contingens, 169
lundeliusi, Cynodontomys, 191 (fig.),
203, 205
Lung fluke, A study of an intermediate
snail host (Thiara granifera) of
the (Paragonimus), 71
lupaster, Canis, 365
lutea, Golfingia, 390
Syncera, 110
lutzt, Mecidea, 474, 479, 480, 485, 493
luzonica, Gongylocephala pallida, 178
luzonica, Othilia, 290, 299, 300, 301
lynchi, Lepidurus, 39, 40 (fig.), "41 (fig.),
42 (table), 43 (fig.), 44 (fig.), 45
(Se.), 46 (fig.), 47 (fig.), 48 (fig.),
Lyponycteris nudiventris, 349
macginitiet, Golfingia, 373, 390, 400,
401, 402, 436
Machilinus, 476
macintoshii, Golfingia, 388, 389, 390
macra, Golfingia, 395
Macrocystis sp., 134, 145, 152
macrurus, Lepidurus, 3, 17, 34, 35, 38
maculatus, Synergus, 337
maculosa, Janira, 141, 143
magnifica, Cenometrra bella, 303
magnocula, Ianiropsis, 117, 136, 145,
146 (fig.), 147 (fig.), 148 (fig.)
major, Mecidea, 473 (map), 474, 476,
478, 479, 480, 481, 484, 485, 486,
489 (fig.), 491, 493, 494, 500
major, Paramys, 194, 203, 205
malmgreni, Stenopleustes, 232
mamillatus, Echinus, 276
Mammalia, 189
PROCEEDINGS OF THE NATIONAL MUSEUM
VOL. 102
Mammals from the Nile Delta region of
Egypt, notes on, 343
mammatus, Phinotropis, 239
mammillatus, Heterocentrotus,
276,
299, 301
5 mandibularis, Ecsenius, 507, 508, 509,
511, 518
mannt, Mulona, 223, 225, 226
manni, Amplinus, 237
mansoni, Schistosoma, 110
Manta, 246
birostris, 246, 248, 254
sp., 254
mardax, Stylophora, 289, 295
Maretia, 283
Maretia ovata, 283, 299, 300, 301
margaritacea, Golfingia, 373, 390, 391,
392, 398
margaritaceum, Phascolosoma, 388, 391
margaritaceus, Sipunculus, 391
mariae, Dipodillus, 355
Gerbillus henleyi, 355
Mariametridae, 267
Mariel Islands, Echinoderms from,
65
martensi, Golfingia, 400, 401
mascarena, Luidia, 284, 299, 300
mathaei, Echinometra, 272, 299, 300,
301, 302
Echinus, 272
mauiensis, Thiara, 72
Thiara granifera, 109, 112
maxima, Quercus, 338
Mecidea, 471, 472, 473 (map), 476, 477,
478-480 (key), 480, 485, 494,
496, 497, 499
indica, 473 (map), 474, 480, 481,
493, 500
ingramsi, 494, 495
kristenseni, 473 (map), 474, 479,
480, 482, 491, 494, 495, 497, 502
Mecidea lepineyi, 473, 479, °480, 483,
499, 501, 502
linearis, 472, 479, 480, 483, 497,
498, 499, 501
longula, 473 (map), 476, 478, 479,
480, 484, 486, 490, 491, 495, 497,
500
lutzi, 474, 479, 480, 485, 493
major, 473 (map), 474, 476, 478,
479, 480, 481, 484, 485, 486, 489
(fig.). 491, 493, 494, 500
minor, 473 (map), 474, 478, 479,
480, 482, 488, 490, 495
pallida, 473 (map), 474, 476, 479,
480, 481, 483, 484, 485, 487, 493,
499, 500, 501
pallida virens, 493, 4
pallidissima, 473 aie 474, 479,
480, 494
pampeana, 473 (map), 474, 478,
479, 480, 484, 491, 495
prolixa, 473 (map), 474, 479, 482,
484, 497, 499, 501
quadrivittata, 479, 480, 498
rungsi, 479, 480, 499
sahariana, 479, 480, 500
INDEX
Mecidea lepineyi—continued
ierr bugs of the genus, review of,
4
straminea, 479, 480, 499, 501
tellinii, 474, 479, 480, 484, 501
vidali, 479, 480, 501
vittata, 472, 498
Mecideini, 475
megapetalus, Echinocyamus, 279, 299,
300, 301, 302
Melania, 72, 110, 455
ebenina, 109
obliquegranosa, 109
phelina, 458
taihokui, 109
Melaniidae, 72
Melanoides, 72, 111
melanosoma, Salarias, 524
Melitodes nodosa, 453
sp., 453
Meniscotheriidae, 196
Meniscotherium, 196
chamense, 196
terrarubae, 196, 203, 205
Mendora, 474
Menzies, Robert J.; Some marine
Asellote isopods from northern
California, with descriptions of
nine new species, 117
meridionalis, Golfingia margaritacea,
9
3
Meriones, 358, 359
arimalius, 358
crassus, 358
erythrourus, 358
ismahelis, 358
libycus, 357, 358
libycus libyeus, 357
schouesboei, 358
shawii, 358
Mesochaetopterus sp., 389, 395
Mespilia, 270
globulus, 270, 299, 300, 301, 302
Metagonimus, 111
yokogawai, 109
Metalia, 282
dicrana, 282, 299, 300, 301
Metopochetus, 173
metularia, Cidarites, 267
Eucidaris, 267, 268, 299, 300, 301,
302
mexicana, Synergus, 315
mexicanus, Synophrus, 315
Miacis, 195
exiguus, 195
latidens, 195, 203, 205
parvivorus, 195
microdentigerum, Phascolosoma, 434
microphyllum, Rhinopoma, 346, 347,
348
microphyllus, Vespertilio, 347
Milliped, Central American, a new genus
of, 235
Mimegralla, 162, 171
albimana, 161, 162, 164 (key), 165
(map), 166
albimana albimana, 164, 165 (map)
541
Mimegralla—continued
albimana contraria, 164, 165 (map),
166
albimana extrema, 164, 165 (map),
167
albimana galbula, 164, 165 (map),
166
albimana keiensis, 164, 167
albimana palauensis, 166
albimana samoana, 164, 165 (map),
167
albimana sepsoides, 164, 165 (map).
166, 167
albimana striatofasciata, 164, 165
(map), 167
albimana tongana, 164, 165 (map),
167
binghami, 168
cedens chrysopleura, 168
cedens thaiensis, 168
coeruleifrons, 168, 171
confinis, 168
contingens, 169
contingens australica, 169
contingens contingens, 169
contingens lunaria, 169
contingens novaehebrideana, 169
contingens solomonis, 169
leucopeza, 170
leucopeza albitarsis, 170
niveimana, 168
perfulva, 163 (fig.), 169
ponapensis, 170
ponapensis kusaieana, 170
ponapensis ponapensis, 170
thaiensis, 168
mimetes, Auturus, 238
Mimomyrmecia, 161, 171, 173
tessellata, 163 (fig.), 171
minor, Pupulina, 254, 258, 259, 260
minor, Dendrostomum, 405
Mecidea, 473 (map), 474, 478, 479,
480, 482, 488, 490, 495
minusculus, Dryocosmus, 337
minuta, Ianiropsis, 136, 155 (fig.), 157
minuta, Golfingia, 396
Munna, 119, 120, 124
minutum, Phascolosoma, 388
misakiana, Golfingia, 393, 395
Mithrodia fisheri, 303
Mitosiphon, 393
Mixodectidae, 191
mixta, Ophiomastix, 296, 299, 301
Mobula hypostoma, 248, 254, 255
lucasana, 245, 258
Mobulidae, 246
modesta, Callirhytis, 338
molaris, Echinostrephus, 303
Mollusea, 72
Molossidae, 352
Monoculus arcticus, 33
montanus, Glyptosaurus,
(table)
montereyensis, Ianiropsis, 136, 152, 153
fig.), 154 (fig.), 158
188, 189
542
PROCEEDINGS OF THE NATIONAL MUSEUM
VOL. 102
Moths of the genera Mulona Walker | namiyei, Ecsenius, 525 (fig.)
and Lomuna, a new and closely
related genus (Arctiidae: Litho-
siinae), 221
mourense Siphonosoma, 381, 384
mucida, Golfing ia,
Mulona, 221, 399 eee 223 (key), 455,
456, 457
barnesi, 223, 227, 229
grisea, 221, 223, 228, 229
lapidaria, 221, 222, 223, 226, 229
manni, 223, 225, 226
nigripuncta, 221, 228, 229
phelina, 458, 459, 460, 467
schaust, 223, 225, 229
multicinctus, Siphonomecus, 385
multicostatus, Andricus, 331
multifida, Comaster, 266
multifora, Asterias, 287
Linckia, 287, 299, 300, 301
mundanus, Sipunculus, 399
Munna, 118, 120 (key)
acanthifera, 119
acarina, 119, 122, 124
avatshensis, 119
boeckii, 118, 119, 134
chromatocephal, 119, 120, 128, 129
(fig.), 131 (fig.)
halet, Hoe 120, P39 (fig.), 133 (fig.)
kr¢gyeri, 119, 124, 125
minuta, 119, 120, 124
nana, 119, 122, 124
palmata, 119
petiti, 118, 119
stephenseni, 119, 120, 124, 125
(fig.), 126 (fig.) 127 (fig.), 134
subneglecta, 131
truncata, 119
ubiquita, 119, 120 (fig.), 123 (fig.),
124 (fig.)
Munnidae, 117, 118
muricaudata, Golfingia, 390
Muricea, 453
pendula, 454
Muridae, 360
murinus, Paramys, 194, 203, 205
Mus cahirinus, 363
jaculus, 364
musculus, 362
musculus’ domesticus, 363
musculus gentilis, 363
musculus musculus, 363
norvegicus, 360
rattus, 361
musculus, Mus, 362
Mus musculus, 363
Mustela libyca, 367
nivalis subpalmata, 366
Mustelae subpalmatae, 366
Mustelidae, 366
myrtifolia, Quercus, 336
mytheca, Dendrostoma, 411, 415
Mytilus californianus, 426
Mytilus-Mitella biotype, 131
namaquensis, Apus, 42 (table), 54-55
(table)
Salarias, 525
nana, Dorometra, 303
Munna, 119, 122, 124
nasomaculata, Addax, 344
Nassopsis, 88, 94
natronensis, Pachyuromys duprasi, 357
Pachyuromys dupresi, 357
nebulosus, Diastrophus, 316
Neoferdina, 285
cancellata, 285
ocellata, 285, 299, 300
Nepeta sp., 318
Neritina, 98
Nestima, 173, 178
pleuralis, 179
polita, 178, 179
prolixa, 178
viridinsula, 179
Neuroterus, 317, 321
tantulus, 321
newberryi, Apus, 53, 56 (table), 57, 60
(table), 64, 65, 66
nicolli, Psammomys obesus, 359
niger, Andricus, 329
nigra, Golfingia, 400, 401
ipa oo Phascolosoma, 429, 431, 432,
nigripuncta, Lomuna, 221, 228
Mulona, 221, 228, 229
nigritibia, Eurybata, 174, 175 (fig.)
Nile Delta region, Egypt, Notes on
mammals from, 343
nilotica, Vulpes vulpes, 366
niloticus, Arvicanthis, 360
Arvicola, 360
Canis, 366
nitens, Thrinoxethus, 243
niteus, Amplinus, 237
nitidus, Amplinus, 237
niveimana, Mimegralla, 168
nodosa, Melitodes, 453
nodosus, Glyptosaurus, 189 (table)
nordenskjéldi, Golfingia, 390
norvegicus, Mus, 360
Rattus, 360, 361
nosophora, Oncomelania, 110
Northarctus, 192
venticolus, 192, 203, 205
noto, Golfingia, 390
Notostraca, 2 a 5 (fig.), 6 (fig.), 16
novae-guineae, Culcita, 285, 299, 300,
301, 303
novaehebrideana, Mimegralla contingens,
novaezealandiae, Golfingia, 395
Nucleolitidae, 280
nudiventris, Lyponycteris, 349
Taphozous, 349
nudus, Sipunculus, 372, 375, 376
numidicus, Apus, 27, 28, "42 (table),
54-55 table)
nutkana, Rosa, 320
Nyctinomus aegyptiacus, 352, 353
teniotis, 352
obesus, Psammomys obesus, 359
oblata, ’Callirhytis, 340
INDEX
obliquegranosa, Melania, 109
oblonga, Echinometra mathaei,
299, 300, 301, 302
oblongi, Chenopodium, 492
oblongifolia, Quercus, 333
oblongus, Echinus, 274
obscura, Golfingia, 390
obscurus, Synergus, 337
obtusidens, Glyptosaurus, 189 (table)
obtusilobae, Cynips, 317
Cynips quercus, 317
obtusilobensis, Ceroptres, 317
obtusipes, Potamon (Geothelphusa),
11
274,
obtusus, Apus, 66
ocellata, Ferdina, 285
Neoferdina, 285, 299, 300
occidentalis, Boavus, 186
Rhodites, 321
Tricellaria, 122
ohlini, Golfingia, 390
okinoseana, Golfingia, 390
oleracea, Spinacia, 493
olivieri, Crocidura, 345
Sorex, 345
olseni, Diacodexis, 202, 203, 205
onagawa, Golfingia, 400, 401
Onchnesoma, 374
Oncomelania, 102, 110
nosophora, 110
quadrasi, 102, 110
operator, Callirhytis, 339
Ophiactidae, 292
Ophiactis, 292
savignyi, 292, 299, 301, 302, 303
Ophiaracna spinosa, 298
Ophidiaster, 286
granifer, 286, 299, 300, 301, 302
lorioli, 287, 299, 301
pusillus, 287, 299, 300
squameus, 287, 299, 301
Ophiochitonidae, 294
Ophiocoma, 294
anaglyptica, 294, 299, 300, 301
brevipes, 296, 299, 301, 302
erinaceus, 295, 299, 300, 301
pica, 295, 299, 300, 301
scolopendrina, 294, 299, 300, 301
sp., 296, 299, 300
Ophiocomella, 296, 298
2 296, 297, 299, 300, 301,
Ophiocomidae, 294
Ophiodera, 291
punctata, 291, 299, 300
Ophiodermatidae, 298
Ophiodesmus, 294
degeneri, 294, 299, 300
Ophioglypha, 298
Ophiolepididae, 298
Ophiolepis, 298
cincta, 298, 299, 302
imbricata, 298
savignyi, 292
superba, 303
Ophiomastix, 296, 298
bispinosa, 296, 299, 300
mixtra, 296, 299, 301
543
Ophiomastix—continued
sexradiata, 297, 299, 300
Ophiomyxa, 291
australis, 291, 299, 300
bengalensis, 291
brevispina, 302
Ophiomyxidae, 291
Ophionereis porrecta, 302
Ophiopezella, 298
dubiosa, 302
spinosa,{298, 299, 300, 302
Ophioplocus, 298
imbricatus, 298, 299, 300
Ophiothrix, 293
demessa, 302
elegans, 293, 299, 301
exigua, 293, 299, 301
galatheae, 302
lepida, 293, 299, 300
lepida hawaiiensis, 293
longipeda, 293, 299, 300, 302
trilineata, 293, 299, 300
triloba, 302
virgata, 293, 299, 300
Ophiotrichidae, 293
Ophiura, 298
kinbergi, 298, 299, 302
longipeda, 293
scolopendrina, 294
Ophiuroidea, 291, 299
opsifrontalis, Ecsenius, 510, 512, 517,
520, 521 (fig.) 526 (table)
Oreasteridae, 285
Orgeriini, 475
Orgerius, 475
Orohippus, 198
orphinus, Amplinus, 237, 238
Oryx algazel, 344
Othilia, 290
luzonica, 290, 299, 300, 301
oryzaphagus, Apus, 9, 12, 13, 16, 53, 61
(fig.), 64, 65
ovata, Maretia, 283, 299, 300, 301
ovatus, Spatangus, 283
ovulum, Echinus, 278
Fibularia, 278, 299, 300, 301, 302
ownstoni, Golfingia, 390
Pachychilus glaphyrus, 113, 114 (fig.),
115
Pachyuromys duprasi natronensis, 357
dupresi natronensis, 357
Pachyurus (Amplinus), 237
pacifica, Bugula, 122
pacificus, Dactylosaster cylindricus, 286,
2
packardi, Lepidurus, 11, 12, 17, 23, 24
fig.), 26, 28, 31, 32, 33, 36
(table), 42 (table)
palauensis, Mimegralla albimana, 166
palicaudatus, Amplinus, 237
pallida, Gongylocephala, 178
Mecidea, 473 (map), 474, 476, 479,
480, 481, 483, 484, 485, 487, 493,
499, 500, 501
pallidissima, Mecidea, 473 (map), 474,
479, 480, 494
pallidum, osentomon, 306, 309, 311
544 PROCEEDINGS OF
palmaeformis, Postelsia, 130
palmata, Lamprometra, 303
Munna, 119
Paludomus, 81, 88, 90, 94
tanschaurica, 78
palustris, Quercus, 326, 335, 338, 341
pampeana, Mecidea, 473 (map), 474,
478, 479, 480, 484, 491, 495
Pantodonta, 197
Panulirus interruptus, 231, 232
papillifera, Golfingia, 396
papillosa, Golfingia, 390
papula, Cynips, 338
Paracraspis, 324
insolens, 324
guadaloupensis, 324
patelloides, 324
Paragonimus, 110, 111, 113
kellicottii, 111
westermani, 71, 108, 110
Paralebion elongatus, 248, 254
Paramys, 194
major, 194, 203, 205
murinus, 194, 203, 205
Parapleustes commensalis, 231, 233 (fig.)
Parasalenia, 272
gratiosa, 272, 299, 300, 301
Parasaleniidae, 272
pardalis, Astrophyton, 453
parvivorus, Miacis, 195
parvum, Hesperosiphon, 386
patagonicus, Lepidurus, 17,
(table)
patelloides, Acraspis, 324
Paracraspis, 324
pattersonae, Andricus, 330
paulus, Hyopsodus, 197
pavlenkoi, Golfingia, 400
pellucida, Golfingia, 396
Peltosaurus, 187
pendula, Muricea, 454
Pentatomidae, 471
perforatus, Taphozous, 348, 349
perfulva, Mimegralla, 163 (fig.), 169
perfuluum, Trichoteras, 322, 323
perimeces, Dandrostoma, 415
Dendrostomum, 372, 404, 405, 406,
31, 42
415
Perissodactyla, 198
peruvianum, Dendrostomum, 405, 411,
414, 415, 422
escadolense, Siphonosoma, 381
etalosoma, 388, 389
petasibarba, Eurybata, 178
petiti, Munna, 118, 119
petraeum, Dendrostoma, 406
Petricola carditoides, 419
phanotypus, Thrinoxethus, 243
phantasticus, Salarias, 515, 516
phanus, Auturus, 238
Phascolion, 374, 389, 423
strombi, 389
Phascoloides, 389, 395
Phascolopsis, 374, 393
Phascolosoma, 372, 373, 374, 375, 388,
389, 422, 423 (key), 429, 436
agassizii, 372, 373, 423, 424, 429,
430, 431, 432, 433, 435
THE NATIONAL MUSEUM
VOL, 102
Phascolosoma—continued
antillarum, 387, 422, 424, 434
aspidosiphonoides, 395
asser, 4385
dentigerum, 423, 432
eremita, 396
granulatum, 388, 422, 423, 429
hespera, 393
japonicum, 390, 429
lordi, 424
margaritaceum, 388, 391
microdentigerum, 434
minutum, 388
nigrescens, 429, 431, 432, 433
procerum, 400, 402
puntarenae, 372, 387, 423, 429, 430,
431, 433
scolops, 429
vulgare, 388, 423
vulgaris, 388
Phascolosomum, 388, 389, 422, 423
eremita, 396
puntarenae, 430
phelina, Autoceras, 221, 455, 458
Cincia, 458
Epeiromulona, 458, 459, 462, 465,
466, 468, 469
Gaudeator, 459, 466, 467
Melania, 458
Mulona, 458, 459, 460, 467
Phinotropis, 236, 237, 239
acuticollis, 239
braueri, 239
kalonotus, 239
mammatus, 239
roreri, 239
tidus, 239
Phodeus, 109
Pholadidea sp., 419
Phyllacanthus, 268
imperialis, 268, 299, 300
Phyllospadix sp., 426
Phymosoma, 388, 422, 423
agassizii, 424
agassixii puntarenae, 430
antillarum, 434
dentigerum, 432
granulatum, 388, 423
Phymosomum, 388, 422, 423
Physconosoma, 422, 423
ERY RAGRON 373, 375, 388, 389, 422,
42
agassizii, 424, 430
antillarum, 434
granalatum, 423
japonicum, 390, 424
Physopsis, 110
pica, Ophiocoma, 295, 299, 300, 301
pictus, Procambarus, 217
pileus, Antron, 333
pilula, Andricus, 333
pilularis, Andricus, 333
pinnifolium, Dendrostomum, 405
Pipistrellus, kuhlii, 351, 352
pipula, Andricus, 332
planci, Acanthaster, 291, 299, 300, 303
Asterias, 291
Planorbidae, 110
INDEX
Platyrhacus, 236
Platyrrhacidae, 235, 236
Platyrrhacus, ligula, 240
Plecoglossus altivelis, 109
Plecotus auritus, 352
pleuralis, Nestima, 179
Pleurocera catenaria, 113, 114 (fig.)
Pleurogonium californiense, 149
Plotiopsis, 72
Pocillopora sp., 434
Poecilictis libyca, 367
polita, Diplolepis, 321
Nestima, 178, 179
Rhodites, 321
polyacanthus, Astropecten, 283, 299,
01
Polydesmus erythropygus, 238
klugi, 2
(Euryurus) taenia, 240
polygonata, Pycnotropis, 240
Polylepiseus, 135, 236, 237, 239
actaeon, 239
furcifer, 239
heterosculptus, 239
stolli, 239
polymyotus, Sipunculus, 376
Polysiphonia, 122
pommeraniae, Siphonosoma, 381
ponapensis, Mimegralla, 170
Mimegralla ponapensis, 170
Amydrinus, 240
pongus, Pyenotropis, 240
popoagicum, Lambdotherium, 200, 203,
205
porrecta, Ophionereis, 302
Postelsia palmaeformis, 130
Potamon dehaanii, 110
(Geothelphusa) obtusipes, 110
powellianus, Hyopsodus, 196, 203, 205
prescotti, Andricus, 331
Primates, 192
princeps, Glyptosaurus, 189 (table)
prioki, Golfingia, 395
problepteryx, Chondrocidaris, 268
Procambarus, 209, 210, 212
blandingii acutus, 217
hayi, 217
lecontei, 209, 210, 211 (fig.)
pictus, 217
pubescens, 212
spiculifer, 212
verrucosus, 212, 215 (fig.)
versutus, 218
procera, Golfingia, 390, 400, 401, 402
procerum, Phascolosoma, 400, 402
projectus, Andricus, 329
Prolimnocyon, 194
antiquus, 194, 203, 205
prolixia, Mecidea, 473 (map), 474, 479,
482, 484, 497, 499, 501
Nestima, 178
pro-oculis, Ecsenius, 510, 512, 517, 519
(fig.), 522
Prophymosoma, 422, 423
Prosobranchiata, 72
Protaphelidesmus, 237, 240
levigatus, 240
ligula, 240
545
protectus, Antedon, 267
Stephanometra, 303
Stephanometra indica, 267, 299, 302
Protura, 305
New insect of the order, 305
Psammechinus verruculatus, 271
Psammomys obesus nicolli, 359
obesus obesus, 359
Pseudochromis tapeinosoma, 528
Pseudorasbora, 109
Pteropidae, 346
Pteropus aegyptiacus, 346
egyptiacus, 346
Ptyxogon, 237, 240
incus, 240
pubescens, Procambarus, 212
pugettensis, Golfingia, 373, 390, 400, 401
pugettensis, Ianiropsis, 135, 136, 139
pulcher, Ecsenius, 511, 515
Salarias, 515, 517
punctata, Ophiodera, 291, 299, 300
puntarenae, Phascolosoma, 372, 387,
423, 429, 430, 431, 433
Phascolosomum, 430
Phymosoma agassizii, 430
Sipunculus (Phymosomum), 430
Pupulina, 245, 246
brevicauda, 258
Copepod genus, with descriptions of
new species, 245
flores, 245, 246, 248, 254, 255, 256,
257, 258, 259, 260, 261, 262
minor, 254, 258, 259, 260
pusillus, Ophidiaster, 287, 299, 300
Pyenotropis, 236, 237, 240
devillei, 240
flavocarinata, 240
haenschi, 240
latzeli, 240
polygonata, 240
pongus, 240
taenia, 240
pyramidum, Dipus, 356
Gerbillus pyramidum, 356
pyriformis, Golfingia, 400, 401
pyroides, Dendrostoma, 406, 409
Dendrostomum, 372, 373, 404, 405,
406, 410, 419, 426
q.-obtusilobae, Diplolepis, 317
quadrasi, Oncomelania, 102, 110
quadrivittata, Mecidea, 479, 480, 498
quadrivittatus, Cerataulax, 472, 480,
484, 498, 499
quercus, Diplolepis, 319
Quercus agrifolia, 337
alba, 322, 325
chrysolepis, 323, 324, 329
coccinea, 340
diversicolor, 333
douglasii, 327, 330
dumosa, 327, 330
emoryi, 339
engelmanni, 331
falcata, 340
gambelii, 332, 333
garryana, 327, 328, 331
hypoleuca, 339
546
Quercus agrifolia—continued
imbricaria, 339
kelloggii, 337
laevis, 341
lobata, 327, 329
maxima, 338
myrtifolia, 336
oblongifolia, 333
palustris, 326, 335, 338, 341
stellata, 317, 342
submollis, 333
subturbinella, 331, 332, 333, 334
toumeyi, 333
velutina, 338
wislizeni, 336, 337
radiata, Chloris, 485
radicum, Diastrophus, 317
rattus, Mus, 361
Rattus, 361, 364
Rattus rattus, 362
Rattus norvegicus, 360, 361
rattus, 361, 364
rattus alexandrinus, 362
rattus frugivorous, 362
rattus rattus, 362
recondita, Golfingia, 396
religiosa, Crocidura, 345
Reptilia, 186
reticulatus, Clypeaster, 276, 299, 300
Echinus, 276
Rhabdoblennius, 508
Rhinolophidae, 350
Rhinolophus, 350
acrotis, 350
eurydale, 350
tridens, 350
Rhinopoma cystops, 347
microphyllum, 346, 347, 348
Rhinopomatidae, 347
Rhodites, 319
bicolor, 321
occidentalis, 321
polita, 321
ribagai, Eosentomon, 306, 309
Richardsonium, 109
rickettst, Siphonides, 386
requetii, Sermyla, 79
Rodentia, 194, 353
Rongelap Atoll, list of Echinoderms, 301
Rongerik Atoll, list of Echinoderms, 302
roreri, Phinotropis, 239
Rosa californica, 321
nutkana, 320
rosae, Cynips, 319
Diplolepis, 320
rosaefolii, Diplolepis, 320
roseata, Epeiromulona, 457
rostratum, Eosentomon, 306
rothschildi, Lepus, 353
rotumanum, Siphonosoma, 381
rotundula, Trichoteras, 323
Rousettus egyptiacus, 346
rubt, Synophromorpha, 317
rudis, Tarebia, 72
rufa, Salmacis, 270
rufus, Desmechinus,*270,'299, 300
rungsi, Mecidea, 479,480, 499
PROCEEDINGS OF THE NATIONAL MUSEUM
VOL. 102
rutilofusca, Golfingia, 395
rutilofuscus, Aspidosiphon, 395
sabellariae, Golfingia, 396
sahariana, Mecidea, 479, 480, 500
Sailer, R. I.; A review of the stink bugs
of the genus Mecidea, 471
Salarias, 507, 508, 509
anomalus, 515, 516, 517
bicolor, 522
burmanicus, 522, 524
frontalis, 512
furcatus, 522, 524
gravieri, 513
melanosoma, 524
namiyei, 525
phantasticus, 515, 516
pulcher, 515, 517
salicis, Synophromorpha, 316
Salmacis rufa, 270
samoana, Mimegralla albimana,
165 (map, , 167
samoanus, Cyclosphen, 167
sanderi, Golfingia, 390
Saniwa, 186
sp., 186
Saphonecrus, 315
brevicornis, 315
sarsii, Golfingia, 396
Sauria, 186
savignyi, Ophiactis, 292, 299, 301, 302,
303
Ophiolepis, 292
scabra, Thiara, 81
Thiara (Plotiopsis), 79, 97
Scarabaeidae, 181
schaust, Mulona, 223, 225, 229
Schistosoma haematobium, 110
mansoni, 110
Schistosomum, 110
japonicum, 110
Schizostella, 451
bayer, 453
bifurcata, 451, 452, 453
New genus of brittle-star (Gorgono-
cephalidae), 451
schouesboei, Meriones, 358
schmitti, Dendrostomum, 406, 413 (fig. ),
422
Schultz, Leonard P., see Chapman
Wilbert M., and, 507
scolopendrina, Ophiocoma, 294, 299
, 301
Ophiura, 294
scolops, Phascolosoma, 429
Scorpaenopsis cacopsis, 528
scotius, Auturus, 238
Scotozous, 351
scottianus, Cynodontomys,
205
164
191, 203,
Scrupocellaria sp., 158
semilauta, Crosa, 175 (fig.), 177
EKurybata, 161, 173, 176, 177
Semisulcospira, 94, 110
libertina, 109, 110, 111
semperi, Golfingia, 400, 401
Senecio‘sp., 490
INDEX
sepsoides, Calobata, 166
Mimegralla albimana,
(map), 166, 167
Septaria, 98
Sermyla riquetii, 79
Serpentes, 186
serratus, Sigmogonotropis, 240, 241 (fig.)
Setzer, Henry W.; Notes on mammals
from the Nile Delta region of
Egypt, 343
sexradiata, Amphilimna, 302
sexradiata, Ophiomastix, 297, 299, 300
shawii, Meriones, 358
Shoemaker, Clarence R.; A new species
of commensal amphipod from a
spiny lobster, 231
siberica, Golfingia margaritacea, 391,
392
164, 165
sibogae, Echinolampus alexandri, 282
Sigmoden, 360
Sigmogonotropis, 237, 240, 241 (fig.)
serratus, 240, 241 (fig.)
signa, Golfingia, 390
signifer, Dendrostomum, 405
Sinopa, 194
strenua, 194, 203, 205
Siphonides, 374, 386
rickettsi, 386
Siphonomecus, 374, 385
ingens, 382
multicinetus, 385
te ele 376, 380, 381 (key),
, 38
amamiense, 381, 382
arcassonense, 381
australe, 381
boholense, 381, 382
bonhouri, 381, 382
dayi, 381
eniwetoki, 381
funafuti, 381, 382
ingens, 372, 381, 382
mourense, 381, 384
novae-pommeraniae, 381
pescadolense, 381
rotumanum, 381
takasukii, 381
Sipunculoidea, 371, 373 (key)
Sipunculus, 373, 375, 379, 380, 423
australis, 380
branchiatus, 377
eremita, 395, 396
galapagensis, 376
gouldii, 393
margaritaceus, 391
mundanus, 379
mundanus branchiatus, 377
nudus, 372, 375, 376
Sipunculus polymyotus, 376
(Phymosomum) puntarenae, 430
sluiteri, Golfingia, 396
socia, Golfingia, 390
solomonis, Mimegralla contingens, 169
sondaica, Grammicomyia, 162
Sorex olivieri, 345
Sorghum halapense, 490
Soricidae, 345
soyo, Golfingia, 390
547
Spalacidae, 359
Spalax aegyptiacus, 359
Spatangidae, 282
Spatangus brissus latecarinatus, 283
ovatus, 283
sphaerula, 5 lpr 335
sphenodon, Glyptosaurus, 189
Spiculida, 371
spiculifer, Procambarus, 212
Spilogale, 367
Spinacia oleracea, 493
spinifer, Dendrostomum, 405
spinosa, Townesa, 163 (fig.), 171, 172
spinosa, Ophiaracna, 298
Ophiopezella, 298, 299, 300, 302
spongifica, Amphibolips, 326
Squamata, 186
squameus, Ophidiaster, 287, 299, 301
stellata, Quercus, 317, 342
Stenomelania, 79
crenulata, 94
Stenopleustes malmgreni, 232
Stephanometra, 267
indica protectus, 267, 299, 302
protectus, 303
Stephanostoma, 388, 389
hansenii, 388
stephenseni, Munna, 119, 120, 124, 125
(fig.), 126 (fig.), 127 (fig.), 134
stephensoni, Dendrostomum, 406
Steyskal, George C.; Australasian stilt-
legged flies (Dyptera: Tylidae)
in the United States National
Museum, 161
stigmatura, Ecsenius, 510, 514 (fig.),
526 (table)
Stink bugs, a review of the genus
Mecidea, 471
stolli, Polylepiscus, 239
straminea, Mecidea, 479, 480, 499, 501
strenua, Sinopa, 194, 203, 205
striatofasciata, Calobata, 167
Mimegralla albimana,
(map), 167
strombi, Phascolion, 389
Strongylosomidae, 236
Stylinodon, 193
cylindrifer, 193, 203, 205
Stylinodontidae, 193
Stylophora mardax, 289, 295
subcostata, Callirhytis, 341
subhamata, Golfingia, 396
submollis, Quercus, 333
subneglecta, Munna, 131
subpalmata, Mustela nivalis, 366
subpalmatae, Mustelae, 366
subturbinella, Quercus, 331, 332, 333,
334
sudanicus, Apus, 42 (table), 52, 53,
54-55 (table)
Suncus crassicaudus, 345
sp., 345
superba, Ophiolepis, 303
Swiftia excerta, 453
sylvestris, Glyptosaurus, 189 (table)
Synophromorpha, 316
Synophrus, 316
Syncera lutea, 110
164, 165
548
Synceridae, 111
Synergus agrifoliae, 337
brevicornis, 315
maculatus, 337
mexicana, 315
obscurus, "337
Sy nophromorpha, 316
rubi, 317
salicis, 316
sylvestris, 316
terricola, 316, 317
Synophrus, 315, 316
mexicanus, 315
sylvestris, 316
PROCEEDINGS
taenia, Polydesmus (Euryurys), 240
Pycnotropis, 240
Taeniaptera albimana, 162
Taeniapterinae, 161, 162
Taeniodonta, 193
Taenioglossa, 72
taihokui, Melania, 109
takasukii, Siphonosoma, 381
Tanganyicia, 94
tanschaurica, Paludomus, 78
tantalus, Neuroterus, 321
Tanymastix, 3
tapachulae, Amplinus, 237
tapeinosoma, Pseudochromis, 528
Taphozous nudiventris, 349
perforatus, 348, 349
tarabuli, Gerbillus pyramidum, 356
Tarebia, 72
granifera, 72
rudis, 72
Teilhardella, 192
sp.; 192, 203, 205
tellinii, Mecidea, ‘474, 479, 480, 484, 501
Temnopleuridae, 270
Temnopleurus, 270
toreumaticus, 270, 299, 302
tenellus, Circulifer, 476
teniotis, Cephalotes, 352
Nyctinomus, 352
tenuissimus, Kudiocrinus, 303
terrarubae, Meniscotherium, 196, 203,
terricola, Synophromorpha, 316, 317
terrigena Diplolepis, 320
territa, Calobata, 179
Trepidarioides, 173, 179
tessellata, Eurybata, 174, 175 (fig.)
tessellata, Mimomyrmecia, 163
171
tetras, Crosa, 177
Eurybata, 177
thaiensis, Mimegralla, 168
Mimegralla cedens, 168
Thiara, 71, 72, 78,%79, 88, 90, 94, 110)
111
176
(fig.),
(Thiara) amarula, 88
(Melanoides) costata, 88
granifera, 71, 72, 74 (table), 75
77 (fig.) 79, 82 (fig.), 83
, 87 (fig.), 89 (fig.), 90, 91
, 93 (table), 95 (fig.), 96
(fig.), 97, 98, 100 (table), 101
(fig.), 102 (fig.), 104. (table),
OF THE NATIONAL MUSEUM
VOL. 102
Thiara—continued
granifera—continued
105 (fig.), 109, 110, 111, 112, 113
Saou, eranifera, 97, 113, 114
g
granifera mauiensis, 109, 112
mauiensis, 72
scabra, 81
(Plotiopsis) scabra, 79, 97
(Melanoides) tuberculata, 88, 110
Thiaridae, 71, 72, 79, 94, 111, 114 ‘(figs.)
Thiarinae, 72
Thrinoxethus, 237, 239, 243
bombonus, 243
cainarachus, 243
hermosus, 243
iquitus, 243
lamprus, 243
nitens, 243
phanotypus, 243
ucayalus, 243
Thriops, 52
thysanata, Epeitromulona, 457, 466
Thysanocardia, 388, 400, 401 (key)
Tia ropsis, 72
tidus, Phinotropis, 239
Tillodontia, 192
Tillotheriidae, 192
Tiphobia, 94
tongana, Mimeogralla albimana,
165 (map), 167
tonganus, Cyclosphen, 167
toreumatica, Cidaris, 270
boroumanicus, Temnopleurus, 270, 299,
30
toucheana, Hua, 109
toumeyi, Quercus, 333
Townesa, 171
spinosa, 163 (fig.), 171, 172
Toxopneustidae, 271
transitorium, Eosentomon, 309, 311
travassosi, Acerentulus, 305
Trepidariinae, 161, 173 (key)
Trepidarioides, 173, 179, 180
cyanea, 180
territa, 173, 179
Tricellaria occidentalis, 122
Trichasteridae, 452
trichocephalus, Apionsoma, 388
Trichoteras, 322
coquilletti, 323
perfuluum, 322, 323
rotundula, 323
tridens, Ianiropsis, 136, 156, 157 (fig.)
tridens, Asellia, 350
Rhinolophus, 350
trigonarius, Echinus, 275
Heterocentrotus, 275, 299, 300, 301
trilineata, Ophiothrix, 293, 299, 300
triloba, Ophiothrix, 302
Triopes, 52
Triops, 52
(Apus) cancriformis, 67
cancriformis, 67
Tripneustes, 271
gratilla, 271, 299, 301
Tripterygion hemimelas, 528
tritamus, Amplinus, 237
164,
INDEX
Troglotrematidae, 110
Troglotrematoidea, 111
tropicum, Dendrostomum, 405
truncata, Munna, 119
trybomi, Goltingia, 390, 391
Tylidae, 161, 162, 173
oer Thiara (Melanoides), 88,
tuberculatus, Glyptosaurus, 189 (table)
ubiquita, Munna, 119, 120, 121 (fig.), 123
(fig.), 124 (fig.)
ucayalus, Thrinoxethus, 243
undulatus, Gymnothorax, 528
utriculus, Andricus, 333
valida, Golfingia, 390
Varanidae, 186
vastum, Hepserosiphon, 386
velutina, Quercus, 338
venezuelense, Eosentomon, 305,
(fig.), 308 (fig.)
Epeiromulona hamata, 457, 463, 464
ventanus, Didelphodus, 189, 190 (fig.),
03, 205
venticolum, Hyracotherium, 198 (fig.),
203, 205
venticolus, Notharctus, 192, 203, 205
vergelanus, Amplinus, 237
verrilli, Golfingia, 395
verrucosus, Procambarus, 212, 215 (fig.)
verruculatus, Cyrtechinus, 271, 299,
307
Psammechinus, 271
versutus, Procambarus, 218
Vespertilio aegyptiacus, 346
kuhlii, 351
microphyllus, 347
Vespertilionidae, 351
vespertina, Loveina, 193
vidali, Mecidea, 479, 480, 501
virens, Mecidea pallida, 493, 494
virgata, Ophiothrix, 293, 299, 300
viridinsula, Nestima, 179
viridis, Lepidurus, 11, 31, 42 (table)
vittata, Mecidea, 472, 498
vittatus, Cerataulax, 472
Viverra ichneumon, 368
549
Viverridae, 368
volva, Fibularia, 278, 299, 300, 301, 302
vulgare, Phascolosoma, 388, 423
vulgaris, Golfingia, 373, 389, 390
Phascolosoma, 388
Vulpavus, 195
australis, 195, 203, 205
Vulpes vulpes aegyptiaca, 366
vulpes nilotica, 366
Wasps, New American cynipid, from
galls, 315
Weld, Lewis H. New American
cynipid wasps from galls, 315
westermani, Paragonimus, 71, 108, 110
White, Theodore E.; Preliminary analy-
sis of the vertebrate fossil fauna
of the Boysen Reservoir area, 185
Wilson, Mildred Stratton; An emended
diagnosis of the copepod genus
Pupulina (Caligoida), with de-
scriptions of new species and a
redescription of the genotype, 245
wislizeni, Quercus, 336, 337
Worms, The sipunculid, of California
and Baja California, 371
wortmani, Hyopsodus, 197, 203, 205
xelithus, Amplinus, 237
Xenosiphon, 373, 376, 377
branchiatum, 377
177
109
311
yapensis, Crosa, 175 (fig.),
yokogawai, Metagonimus,
yosemitense, Kosentomon,
Zacco, 109
zenzibakensis, Golfingia, 400, 401
zephyri, Loveina, 193, 203, 205
Zopheroteras, 334
guttatum, 334
sphaerula, 335
Zostera sp., 385, 404, 417
zostericola, Dendrocola, 411
zostericolum, Dendrostomum, 372, 404,
405, 406, 410, 411, 413 (fig.),
415, 416, 417, 418, 422
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