1869
THE LIBRARY
.BULLETIN OF THE
^Southern California
Academy of Sciences
LOS ANGELES, CALIFORNIA
Vol. 68 January-March 1969 Part 1
CONTENTS
Biogeographic Relationships of the Salton Sea Amphipod, Gam-
marus mucronatus Say. J. Laurens Barnard and W. Scott Gray 1
Evidence of Celestial Orientation by California Toads ( Bufo boreas)
during Breeding Migration. C. Richard Tracy and Jim W. Dole 10
Hymenodora glacialis (Decapoda: Natantia) from the Arctic Basin.
Alan D. Hävens and Wesley L. Rork 19
Uscia mexicana, new genus, new species, a Watersiporid Bryozoan
with Dimorphie Autozoids. William C. Banta 30
A New Species of Speleocola (Acarina: Trombiculidae), off a Bat,
Pizonyx vivesi, from Baja California, Mexico. Richard B.
Loomis and James P. Webb, Jr 36
A comparison of the Free Amino Acids in Two Populations of the
Polychaetous Annelid Neanthes succinea. Alan J. Mearns and
Donald J. Reish 43
Research Notes:
Notes on the Life History of Fishia evelina hanhami (Lepidop-
tera). John Adams Comstock and Christopher Henne 54
The Repository of the T. W. Cook Ant Types (Hymenoptera:
Formicidae). Roy R. Snelling . . . . 57
March 17, 1969
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t Vol. 68
OF THE SOUTHERN CALIFORNIA
ACADEMY OF SCIENCES
January-March, 1969 No. 1
Bull. So. Calif. Acad. Sei. 68(1): 1-9, 1969
BIOGEOGRAPHIC RELATIONSHIPS OF THE SALTON
SEA AMPHIPOD, GAMMARUS MUCRONA TUS SAY
J. Laurens Barnard
Smithsonian Institution
Washington, D.C. 20560
and
W. Scott Gray
Pacific Marine Station
Dillon Beach, California 94929
Introduction
The biogeographic relationships of the Texan lagoonal amphipod,
Gammarus mucronatus Say, recently introduced into the Salton Sea
of California (Barnard and Gray, 1968),‘possibly extend to European
freshwaters. Barnard and Gray created a new subgenus, Mucrogam-
marus, to receive this amphipod after comparing it with various Ameri-
can and European members of the Gammarus complex that includes
subgenera such as Gammarus (5.5.) (marine and freshwater), Rivulo-
^gammarus (freshwater), Pectenogammarus (marine), Marinogam-
narus (marine) and with unnamed subgenera such as that indicated
rby Karaman (1959).
Since 1906, when Stebbing placed G. mucronatus in the genus Cari-
L* nogammarus , it has represented the only American species in a genus
that in recent years has been restricted more and more to species occur-
_ring in Lake Baikal (Siberia).
Carinogammarus has been the repository, in some cases temporarily,
— or a number of species with diverse geographical and ecological affini-
ties. European freshwater, Northern Pacific marine, Atlantic brackish-
r marine, and Asian freshwater species all at some time have been united
in this genus. The type-species of Carinogammarus is Gammarus cin-
namomeus Dybowsky (1874); this species and several other original
members of the genus are endemic to Lake Baikal.
To consider then, the origin and relationships of Carinogammarus
—mucronatus implied by this association stimulates several questions.
I
w>
1
2
Bulletin So. Calif. Academy of Sciences
Some of these questions, although perhaps rhetorical, are, nonetheless,
basic to the systematics of a sizeable complex of gammarid amphipods.
First, why is Carinogammarus mucronatus not congeneric with the
Lake Baikal carinogammaruses? Are the dorsal Ornaments of gam-
marids, so lavishly developed in Lake Baikal, valid taxonomic charac-
ters, and if so, at what level? Why is Carinogammarus mucronatus not
congeneric with the common European freshwater genus Rivulogam-
marus, as has been supposed by some workers? Does C. mucronatus
represent an example of (parallel?) convergent evolution with respect
to other carinate species? Ecologically speaking, can C. mucronatus
be considered the northwestern Atlantic counterpart of the north Pacific
Anisogammarus1. Finally, of what origin or origins are the Gammaridea?
These questions will now be amplified and examined.
Schellenberg (1937a, b) believed Carinogammarus shouldbe limited
to species in Lake Baikal; his argument for this limitation was not
strong but the action did appear sound biogeographically. He then
transferred C. mucronatus to a carinate section in Rivulogammarus ,
together with various other keeled “carinogammaruses” of European
freshwaters. Thus, to Schellenberg and the present workers (Bamard
and Gray), possession of dorsal carinae does not present a sufficiently
strong relationship to offset the geographical and ecological gap be-
tween Rivulogammarus mucronatus and the carinate gammarids of
Lake Baikal. An interesting feature of Schellenberg’s work is that he
combined in Rivulogammarus various species irrespective of their
dorsal ornamentation or lack of it, although he did use this criterion
to establish sections within the then subgenus. By this act the traditional
importance given to dorsal carinae, teeth, or spine groups was dis-
counted.
Karaman (1959) gave further recognition to the distinctness of the
Baikalian Carinogammarus from the non-Baikalian, carinate section
of Rivulogammarus. However, he placed more importance on dorsal
ornamentation than did Schellenberg. The European carinate species
were removed from Schellenberg’s diverse genus Rivulogammarus and
united in the new subgenus Fluviogammarus (a junior homonyn of
Fluviogammarus Dorogastajskii [1917]). Karaman (1959) based his
new subgenus not only on the keeled (carinate) condition but on other
characters as well. R. mucronatus was not studied by Karaman.
What of the carinate condition in R. mucronatus ? As previously
mentioned and detailed by Barnard and Gray (1968) the carinae (teeth)
on pleon Segments 1 , 2, and 3 tend to be reduced in number, and this
tendency varies in degree geographically. Other characters such as
gnathopods remain grossly stable throughout the ränge. It seems pos-
Biogeographic Relationships in Amphipods
3
sible that convergent evolution has produced four distinct carinate
groups: the Baikalian group, the European freshwater group, the North-
western Atlantic marine group, which is represented by the single,
abundant species, R. mucronatus, and the north Pacific marine-fresh-
water Anisogammarus group. But why has not there been further radi-
ation in the northwestern Atlantic to produce several carinate species?
To explore this question let us now consider the association of species
with freshwater and marine affinities. Gammarus has species extending
in salinity preference from full marine through brackish conditions to
freshwater (Spooner, 1947); also, Rivulogammarus, sensu Schellen-
berg (1937a, b) incorporated a brackish water species within a pre-
dominantly freshwater genus. Is Rivulogammarus mucronatus more
probably of a freshwater or marine origin? Morphologically, R. mucro-
natus, while distinct in some details, shows similarities to other Rivulo-
gammarus species. It is particularly close to those species of the cari-
nate section as contrasted to those of the smooth section. Perhaps this
indicates that R. mucronatus has invaded its brackish water niche from
freshwater. Invasions of estuarine biotopes are thought to be principally
from the ocean but occur in some instances from freshwater (Hedgpeth,
1957). Perhaps the relatively impoverished estuarine fauna allowed
R. mucronatus to exploit a niche existing along the Atlantic and Gulf
coasts. The wide geographical ränge might indicate an expansion
rapidly under conditions of little or no competition. This would be
comparable to the almost explosive development in the Salton Sea by
this species. The tolerance of R. mucronatus to temperature Variation,
not only diurnal but over its ränge, could be interpreted as an adapta-
tion more typical of a freshwater than a marine animal.
An interesting Situation exists in the northern Pacific Anisogam-
marus, which has parallels to the Atlantic Situation. This genus con-
tains species with dorsal ornamentation as diverse if not more diverse
than that of Rivulogammarus sensu Schellenberg. Ecologically, the
species extend from marine to freshwater. If G. mucronatus represents
a biotopic counterpart of the brackish water complex of Anisogam-
marus, then the greater diversity of the Pacific forms could indicate a
greater diversity of niches or the absence of one species with sufficient
eurytopicity to spread throughout all Pacific biotopes. Should the
variously ornamented kinds of Anisogammarus be considered distinct
genera in analogy to freshwater genera? Has the diversity seen in Ani-
sogammarus been influenced by the proximity of Lake Baikal, which
might have served as a supply point for the various genotypes?
Speculative expansion of these questions raises a point concerning
the origin or early diversification of the Gammaridea; did it take place
4
Bulletin So. Calif. Academy of Sciences
in a freshwater complex comparable to Lake Baikal? Amphipod fossils
of pre-Miocene age do not preclude such a thesis because they are
generally not extant, and similarity between fossils of Miocene amber
deposits and recent species indicates a conservatism and great age for
the Amphipoda. Baikalian radiation also reflects evolutionary trends
seen in marine Gammaridea. Additionally, the genera of Gammaridea
show a decided preference for cold water; one may eite, for example,
the large number of genera in the North Pacific Basin as compared to
many fewer genera in the tropical areas. The penetration of springs
and wells by freshwater amphipods might also be pointed out. How-
ever, it is not certain that Lake Baikal has always been a cool-water lake.
Tzvetkova’s (1965) treatment of Anisogammarus climaxes a slowly
evolving amalgamation of numerous species from once distinct genera
under one generic appellation. As mentioned previously, Anisogam-
marus has as great an ornamental diversity as once seen in Rivulogam-
marus. Indeed, Anisogammarus now rightly includes north Pacific
marine species which formerly belonged to Echinogammarus , the
freshwater equivalents of which are still segregated from Rivulogam-
marus or Gammarus.
Anisogammarus is now composed of nearly a score of species in 4
groups, divided according to the presence or absence of dorsal orna-
mentation on pleonites 1 through 6. Formulas for these groups are
given, in Symbols, where (T) = a single dorsal tooth, (S) = bundles of
dorsal spines, and (O) = no ornamentation; thus, Group 1 [Aniso-
gammarus , sensu stricto] = O-O-O-S-T-S, Group 2 [Anisogammarus
(Eogammarus)] = O-O-O-S-S-S, Group 3 [no subgeneric name] =
T-T-T-S-S-S, and Group 4 [no subgeneric name] = (S or 0)-S-S-S-S-
S. There is little question that these 4 groups are sufficiently alike to be
associated under one generic heading, for they presumably all have
accessory branchial lobes and gnathopods with dense rows of peg-like
spine-teeth on the palms. Gnathopod 1 is slightly larger than gnathopod
2. The accessory branchial lobes are unique to these marine-brackish-
freshwater amphipods, and apparently are not homologous to the ster-
nal branchiae of African Paramelita. Apart from the branchiae, the
members of Anisogammarus clearly show affinities to their freshwater
equivalents of the Palearctic, but they are today restricted to the sea-
ward fringes of the boreal-subarctic, North Pacific Basin. The broad
concept of Anisogammarus presents a nomenclatural inconsistancy
when compared to the more narrow viewpoint of the limnologically
oriented systematist. For in Anisogammarus, in addition to dorsal
ornamentation, there are included various kinds of third uropods and
lateral cephalic lobes that mark either full genera or subgenera in the
freshwater “gammarus” group.
Biogeographic Relationships in Amphipods
5
The problem is partially semantic as to whether Gammarus, Rivulo-
gammarus, and Marinogammarus are subgenera of a generic complex
or are distinct genera. Schellenberg (1937a,b) ranked these as sub-
genera of Gammarus, and also included several other subgenera which
today are treated as genera. If it were not for the dozens of other “gam-
marus” genera in Lake Baikal, the problem would be fully semantic.
The relationships of morphology, i.e., the lack of extreme divergence,
would suggest, probably, that the various non-Baikalian gammaruses
are clearly related genetically. The intricate polyphyletism involved is
nowhere better expressed than in our subject, G. mucronatus, and in
the general ecology of the group. Marinogammarus is primarily con-
fined to brackish and saline waters, while Gammarus (Gammarus) has
both fresh and saltwater species. Rivulogammarus is traditionally a
freshwater genus. The salinity ränge of G. mucronatus is from brackish
to fully saline, with morphological characters of Gammarus, Marino-
gammarus, and Rivulogammarus being shown. Its lateral cephalic lobe
is intermediate between Gammarus and Rivulogammarus , while the
second gnathopod, with excessively sloping palm, is reminiscent of
Marinogammarus. A clear relationship to “fluviogammarus” of Europe
is shown by the truncate tooth of the gnathopodal palms and the dorsal
carinae. Uropod 3 is of the Gammarus -Rivulogammarus kind in con-
trast to that of Marinogammarus. Of the 3 main groups, Rivulogam-
marus seems to be the most primitive, and may have given rise to: (1 )
Gammarus, with sharply angled cephalic lobes, whose members have
extensively reinvaded the sea; (2) Marinogammarus , with softly and
vertically truncate head lobes; (3) European carinate members confined
to freshwaters; and, (4) the marine G. mucronatus with slightly modified
cephalic lobes and extremely modified gnathopod 2. The axial gradi-
ents of gnathopod 2 have been so altered that gnathopod 2 of G. mucro-
natus has a sloping palm similar to that of gnathopod 1 .
This above sequence is probably oversimplified, as far as the Eura-
sian freshwaters are concerned. Other subgenera and genera undoubtely
fall into the scheme, if it is assumed that many genera are the result of
the diversification of the Rivulogammarus stock stimulated by numer-
ous egresses and ingresses from the sea during glacial cycles. Neither
the gammarus complex in the Caspian Sea nor the adaptive radiation
of Baikal have been considered. Both are beyond the scope of the
present paper, but are undoubtedly an important part of Eurasian
amphipod diversity. The diversification in Eurasia is strangely not
apparent in N. American gammaruses (Bousfield, 1958); the explana-
tion involved is probably complex. One factor which might be suggested
is the lack of contiguity between the margin of the N. American ice-
sheet and a “mediterranean-like” sea which on the positive side might
6
Bulletin So. Calif. Academy of Sciences
have favored repeated migrations from sea to freshwater in circum-
stances permitting isolation. The point which can be made is that the
co-distribution of Rivulogammarus between N. America and Eurasia
is indicative of its role as an evolutionary stock in non-Baikalian waters.
The authors lack the knowledge to trace further the possible origin of
Rivulogammarus , specifically, whether it is basic to the Baikal fauna or
has evolved from Baikalian precursors. Similarly, the relationships of
such genera as Echinogammarus and Sarothogammarus to Rivulo-
gammarus can not be further developed. But Schellenberg (1937a,b)
considered Echinogammarus to be a subgenus of Gammarus, making
it equivalent to Rivulogammarus. We have examined the type-species
of Echinogammarus , E. berilloni (Catta), type locality: Pyrenee Mtns.,
and consider its gnathopodal structure and general aspect to indicate
its generic equivalence to Gammarus, sensu lato. Nothing is seen,
however, to detract from the possibility that it has evolved from the
Rivulogammarus stock. Echinogammarus has been distinguished from
Gammarus by the presence of pleonal spine bundles anterior to pleonite
4, and on this basis, various North Pacific marine species formerly in
Echinogammarus are now placed in Anisogammarus. Schellenberg
(1937a,b), again recognizing the distinct Baikal fauna, pointed out the
need to remove the forms of this area from Echinogammarus. Bazika-
lova (1945) did so, creating Eulimnogammarus for them and establish-
ing several subgenera to divide them further. One of the subgenera,
Heterogammarus, Stebbing (1899), has generic priority; thus, the
name Eulimnogammarus should be reduced to subgeneric Status under
Heterogammarus .
Schellenberg (1937a,b) also pointed out four differences between
Baikalian Carinogammarus and the carinate members of Rivulogam-
marus formerly assigned to Carinogammarus: (1) the unshortened
peduncle in antenna 1 of Rivulogammarus ; (2) the presence of calceoli,
(not universal, however) in Rivulogammarus; (3) the lack of setae on
the lower margins of the coxae; and, (4) the unshortened uropod 3.
Baikalian gammarids are still an enigma to us, however, because they
are classified with primary reference to dorsal omamentation (Bazika-
lova, 1 945). This practical Classification may be convenient for Baika-
lian students, but it may disguise the presence of true members of
Rivulogammarus or it may fail to pinpoint close relationships that
would elucidate origins of both Baikalian and non-Baikalian species.
Minute details of some of the critical species groups have not been
clarified.
Karaman (1959) has segregated Rivulogammarus argaeus (Vavra),
R. triacanthus (Schafema) and R. roeselii (Gervais) into the afore-
Biogeographic Relationships in Amphipods 7
mentioned “fluviogammarus” which he makes a subgenus of Rivulo-
gammarus. He thus continues the tradition of some European specialists
in elevating various subgenera of Gammarus. If one were to return to
Schellenberg’s treatment (followed by the American expert Bousfield,
1958), “fluviogammarus” would be reduced to the level of a super-
species. It could be elevated to subgeneric Status and would thus be
equivalent to the Rivulogammarus concept and contrary to Karaman’s
thesis. The interplay of nomenclature is again semantic for we are
really interested more in the true morphological reslationships of the
species and species groups than in their names or whether they deserve
any. But names do have a use, for they signal to the reader close rela-
tionships, or diversification, and indicate to which categorical level a
specialist considers a taxon belongs. Karaman has made the point that
the carinate rivulogammaruses of Europe have, besides carinae, charac-
ters in common not fully shared with non-carinate rivulogammaruses.
These characters include those of the pleonal epimera, pereopods,
telson, general body form, antennae and gnathopods. He thus implies
that a non-carinate (“glatt” of Schellenberg) Rivulogammarus , upon
developing carinae and additional minor divergences, has radiated
into 3 species and several subspecies, thereby forming a species group
worthy of generic recognition. Karaman is not certain that these species
do not have clear-cut connection, by the other characters mentioned,
to non-carinate species occupying territory east of middle Europe. This
would not be true to the west where exploration and morphological
elucidation are in a better state. If this relationship could be proved it
would suggest a return to the earlier importance of dorsal carinae as a
mark of strong differentiation. It would then become evident to students
of Anisogammarus that they must look further into numerous characters
of that species flock, aside from dorsal ornamentation, that might
signify generic division of the 4 kinds of Anisogammarus.
Dorsal metasomal carinae have therefore appeared several times in
gammaruses: in the “fluviogammarus” stock, in the Baikalian Carino-
gammarus, and in G. mucronatus. This may signify that carinae are not
positive evidence of a major genetic innovation but are a recurrent
mark of convergence. However, one cannot fully dismiss the possi-
bility that carinae are evidence of a relationship among Baikalian,
European and American carinate species although such a relationship
would exist at the level of a generic flock concept rather than at the
superspecies level. The problem of immediate interest is whether G.
mucronatus is a product of the American Rivulogammarus stock in
which independent carinal replication has occurred. If this thesis were
true, one could eliminate the need for an Atlantic land bridge to let
8
Bulletin So. Calif. Academy of Sciences
one European carinate species enter American marine waters. A
Crossing of that kind could be eliminated from consideration if the
mucronatus precursor were preadapted to saline waters. If so, the
connection has been broken in marine waters of Europe, possibly by
evolution of the more successful lines of Marinogammarus and the
marine species of Gammarus.
CONC LUSIONS
Mucrogammarus mucronatus seems to have a stronger relationship
to European “fluviogammaruses” than it does to the Gammarus-
Rivulogammarus stock of either hemisphere. This is reflected not only
in carinae but (1) in the occurrence of numerous bent spines on the
gnathopod palmar face and proximo-posterior margin of article 6;
and (2) in the blunt midpalmar spine of mucronatus palms that resem-
bles the truncate spine of European R. roeselii (“fluviogammarus”)
more than it does the relatively symmetrical and subacute spine of
non-carinate gammaruses. The slight tendency toward Gammarus, s.s.,
ocular lobes and the obliquity of the palm of gnathopod 2, would then be
considered to be coincidental in G. (Mucrogammarus) mcronatus, but
the latter character is one of the major distinctions of Mucrogammarus.
Literature Cited
Barnard, J. L. and W. S. Gray, 1968. Introduction of an amphipod crustacean
into the Salton Sea, California. Bull. So. Calif. Acad. Sei., 67: 219-232.
Bazikalova, A., 1945. Les amphipodes du Baikal. Trav. Station Limnologique
du lac Baical, Akad. Nauk Souza S. S. R., 11: 1-440.
Bousfield, E. L., 1958. Fresh-Water amphipod crustaceans of glaciated North
America. Canadian Field-Naturalist, 72: 55-1 13.
Dorogostajskii, V., 1917. Contributions ä la faune des Crustaces du fleuve An-
gara. Ann. Mus. Zool. Acad. Sei. Petrograd. 21: 302-322.
Hedgpeth, J. W., 1957. II. Biological Aspects in Chapter 23, Estuaries and La-
goons. Geol. Soc. Amer., Mem. 67, vol. 1, p. 693-729.
Karaman, S. and G., 1959. Gammarus ( Fluviogammarus ) riacanthus Schaferna,
argeaus Vavra und roeselii Gervais am Balkan. Institut de Pisciculture de
la RP De Macedoine, 2: 183-211.
Schellenberg, A., 1937a. Schlüssel und Diagnosen der dem Süsswasser-Gamma-
rus nahestehenden Einheiten ausschliesslich der Arten des Baikalsees und
Australiens. Zoologischer Anzeiger, 1 17: 267-280.
— 1937b. Kritische Bemerkungen zur Systematik der Süsswasser-gammariden.
Zoöl. Jahrh. Syst., 69: 469-516.
9
Biogeographic Relationships in Amphipods
Spooner, G. M., 1947. The Distribution of Gammarus species in estuaries. Part
I. J. Mar. Biol. Ass. U. K., 27: 1-52.
Stebbing, T. R. R., 1899. Amphipoda from the Copenhagen Museum and other
sources. Trans. Linnean Soc. London, ser. 2, Zool., 7: 395-432.
— 1906. Amphipoda I. Gammaridea. Das Tierreich, 21: 1-806.
Tzvetkova, N. L., 1965. Novyje rod gammaridy (Amphipoda, Gammaridea) iz
pribrezhnyx uchastkov Japonskoga morja. Zoologicheskie Zhurnal, Akad.
Nauk SSSR, 44: 1631-1636.
Accepted for publication July 29, 1 968.
Bull. So. Calif. Acad. Sei. 68(1): 10-18. 1969
EVIDENCE OF CELESTIAL ORIENTATION
BY CALIFORNIA TOADS (BUFO BOREAS)
DÜRING BREEDING MIGRATION
C. Richard Tracy1 and Jim W. Dole
Department of Biology,
San Fernando Valley State College
Northridge, California 91324
Abstract Adult male California toads collected in spring
from their breeding site or while migrating towards the
breeding area were tested for their ability to Orient using
celestial cues. Animais of both groups oriented in the direc-
tion of their migratory movement when the sun was visible;
under the night sky the choice of directions was apparently
random. When provided with a choice of moving towards
an artificial “chorus” or in the migratory direction the
majority of the toads chose the chorus at night but the
migratory direction in daytime.
Introduction
On the night of March 24, 1968, we witnessed a spectacular breeding
migration of California toads, Bufo boreas Baird and Girard, as they
moved from their wintering quarters to a breeding area along one
shore of Seminole Lake, a small man-made lake 8 km Southwest of
Agoura, Los Angeles County, California. Thousands of animals were
involved; an estimated 6000-7000 toads were seen at the breeding
site in the following days. Several two minute censuses during the
migration of toads gave counts as high as 70 animals entering the lake
along a 1 50 m section of shoreline, indicating a very large population
movement. Presumably all the toads seen came from the same general
region, for all were moving in a southwesterly direction. The animals
were observed to leave a chaparral covered hillside to the northeast
of the lake, move a distance of up to 300 m overland, enter the
lake and swim rapidly the 80 m or more to the breeding congress on
the opposite shore. So far as could be determined, all were males;
females were not seen at the breeding site until two days later.
Because of the apparent abilities of these animals to move directly
to their goal, we took the opportunity to conduct a few preliminary
experiments to determine whether or not the animals were being
guided to the breeding site by celestial cues. Recently, several species
Present address: Department of Zoology, University of Wisconsin, Madison,
Wisconsin 53706
10
Celestial Orientation
1 1
of anurans have been shown to possess the ability to use the sun, stars
or moon, in conjunction with a “biological clock,” to Orient their
movements on a compass bearing at right angles to a familiär shoreline
(Ferguson et al., 1968). Such Orientation, termed “Y-axis orientation,”
involves movement on a compass course which, had the animals been
displaced directly inland or directly offshore from their capture sites,
would result in their retum to the shoreline. However, the role of
celestial cues in directing orientation to other than a Y-axis has not
previously been investigated. We also made an attempt to evaluate
the relative importance of auditory and celestial cues in influencing
the directional choice of adult toads.
Because of insufficient time and inadequate preparation we were
unable to perform all the experiments desired. However, since the
area around the lake is now being developed by commercial interests
and has been closed to our use, it appears unlikely that we shall be
able to continue this work in the near future. Consequently, our Und-
ings, although incomplete and tentative, are reported herein.
Methods
All toads used in the experiments were collected on the night of March
24. One sample of 20 maies (group 1) was taken from the breeding
congress on the Southwest shore of the lake. Since the first toads were
known to have arrived in this area the previous night, the animals had
been at the site at most about 24 hours. Probably the majority had just
arrived as a part of that night’s migratory wave. Ten other maies (group
2) were captured the same night as they moved across the crest of an
earthen dam toward the lake from the northeast.
All experimental toads were placed in light-tight jars and taken by
car over winding mountain roads to the campus of San Fernando
Valley State College, 25 km away, where all experiments were per-
formed. The animals were released here, one group at a time, in the
center of an arena built on the roof of the Science building. The arena,
made of opaque black plastic sheeting mounted on a wooden frame
and shaped as a decagon, 8.1 m between opposite corners, was so
constructed that only the floor, walls and sky were visible at ground-
level from within. The bottom was of gravel. The investigators entered
and left the arena by climbing over the wall.
To determine if the toads could use celestial cues to guide their
movements, each group was tested several times in the arena under
both day and night sky conditions. For each test the toads were placed
together under a light-tight rectangular pan inverted in the center of
the arena to which was attached a string hung loosely from one side of
12
Bulletin So. Calif. Academy of Sciences
the enclosure to another. The investigator then left the arena, posi-
tioned himself out of sight behind its wall, and pulled the string taut,
thus lifting the pan and releasing the toads. The animals were left
undisturbed for five minutes during which time they were free to move
in any direction. At the end of this period the investigator re-entered
the arena and noted the position of each animal. For all which had
moved to within 1 m of the wall a directional choice was determined;
those which did not leave the center were recorded as not moving.
After each test the toads were collected and kept in a laboratory in
aquaria containing wet Sphagnum moss until used again.
To assess the relative importance of celestial and auditory cues in
guiding the toads to their breeding site, the 10 males of group 2 were
released in the arena under both noctumal and diurnal conditions,
but with an artificial “chorus” at various locations just outside the arena
wall. The “chorus” was created by placing several male toads together
in an aquarium; the interactions of these animals as they attempted
amplexus with each other produced nearly constant vocalization.
All data were analyzed statistically to determine the probability that
the choice of directions among those toads which reached the wall in
each test was due to chance. In most instances the Rayleigh test (Bat-
schelet, 1965) was used, probability values for “Z” being obtained
from a chart provided by Durand and Greenwood (1958). In those
tests where the response appeared to be bimodal, the modified Smimov
test was employed as suggested by Batschelet (1965).
Results
Responses to celestial cues — On the night of their capture the animals
of group 1 were taken directly to the arena and released together under
a hazy moonless night sky in which only two or three of the brightest
stars were visible. The resulting dispersion of the directional choices
of the toads is shown in Fig. la; the distribution did not differ signifi-
cantly from that expected by chance when tested with the Rayleigh test
(Z = 0. 1 0; P > 0. 1 0). Because of the possibility of a bimodal distribu-
tion the Smirnov test was also employed; the computed U2 of 0.064,
however, also indicated a random distribution (P>0.10).
The moming following their capture the same animals were again
released in the arena with the sun clearly visible in the east. The result-
ing choice of directions of those which moved under these conditions
(figure lb) was clearly non-random, (Z = 15.11; P<.0001). Signifi-
cantly, the mean direction of movement was to the Southwest, the
direction in which the bulk of the breeding toads had moved when
approaching the breeding site. Retested later the same day with the
Celestial Orientation
13
sun visible in the west (Fig. lc), the animals again showed a strong
tendency to move southwestward (Z = 1 1 .5 ; P < .000 1 ).
Düring the following night, this time under a clear, starry sky, the
same animals were released together again in the arena. As on the
previous night the choice of directions appeared to be bidirectional;
however, when tested with the Smimov test the probability of achieving
such a distribution by chance was greater than 0.10 (U2 = 0.107)
indicating no significant difference from a random dispersion.
Animals from group 2, captured as they moved to the lake, were
tested in the same manner and under the same conditions immediately
following each test of group 1 . The response of group 2 on the night
of their capture (Fig. le) did not differ significantly from random
(Z = 1.75; P>0. 10) although the majority (65 per cent) of those
moving chose the general direction of their previous migration. Both
day time releases of this group (Fig. lf, g), as with the previous animals,
resulted in non-random dispersions (Z = 4.12 and 6.84; P<0.05 and
0.001, respectively). As before, the bulk of the movement was to the
Southwest, the direction in which the animals had been traveling when
captured. When released the night following their capture under a
clear starry sky the choice of directions again did not differ significantly
from that expected by chance alone (U2 = 0.068; P>0. 10).
• TOAD ► Y-AXIS OR MIGRATIONAL COURSE PRIOR TO CAPTURE
Figure 1. Top row — Directional responses of 20 adult male toads (group 1)
captured in water at the breeding site, plotted relative to their Y-axis (compass
bearing at right angles to home shore which would result in return to the shore-
line if displacement were directly inland). Bottom row — Responses of 10 adult
males (group 2) captured while moving towards the breeding site, plotted relative
to their direction of travel when captured. Test conditions were: a, e - hazy night
sky, only 2 or 3 stars visible; b, f - following morning, sun visible; c, g - that after-
noon, sun visible; d, h - clear, starry, moonless sky.
14
Bulletin So. Calif. Academy of Sciences
Response to auditory cues. When released under the hazy sky on
the night of their capture, but with an artificial chorus placed at various
locations, the animals in group 2 showed a tendency to move in the
direction of the sound (Fig. 2 a-c). In the two tests in which the direction
of the sound could be clearly distinguished from their migratory
direction, 67 per cent of the animals went towards the chorus. The
remainder moved to the Southwest, the direction of their travel when
collected, perhaps indicating an ability to Orient using the night sky
which was not apparent in the earlier tests. The dispersions in two
tests (Fig. 2 a,b) were significantly different from random (U2 = 0.221 ;
P<.025 and U2 = 0.482; P<.005, respectively); the distribution in
the third test (Fig. 2c) however, did not differ significantly from that
expected by chance (U2 = .137; P> 0.10).
When tested the following moming with the sun visible (Fig. 2d)
the responses were again divided between the “chorus” and the migra-
tory direction, but with the majority choosing the latter. Tested that
aftemoon under clear skies all toads moved to the Southwest, ignoring
the “chorus” (Fig. 2e). Both daytime dispersions differed significantly
from that expected by chance (U2 = 1.38; P< 0.005 and Z = 7.52;
P< 0.0001, respectively). Of those moving in these two tests 88 per
cent chose the general migratory direction in preference to the “chorus.”
Discussion
Although the data are meager it nevertheless appears that adult male
Bufo boreas have the ability to use celestial cues in guiding their move-
ments, for when removed from their normal place of residence and
released in a totally unfamiliar setting the animals clearly showed a
preference for moving in the direction of their initial migration to the
breeding site when they could view the sun. Since the toads had been
transferred 25 km to the arena, it is unlikely that olfactory and auditory
cues could have been guiding them. Presumably the position of the sun,
together with a “biological clock,” permitted orientation as has been
found to be the case in Bufo fowleri (Ferguson and Landreth, 1966),
Rana catesbeiana (Ferguson et al., 1968), Ascaphus truei (Landreth
and Ferguson, 1967), Pseudacris triseriata (Landreth and Ferguson,
1966), Acris gryllus (Ferguson et al., 1965) and Acris crepitans (Fer-
guson et al., 1967). Animals of these species when placed in a terres-
trial arena generally respond as if they had been displaced inland from
the shore, moving in the direction which would return them to the
shoreline from such a position. However, the toads in our experiments,
Celestial Orientation
15
both those collected at the breeding site and those taken while on the
move, generally oriented towards the Southwest, approximately oppo-
site the direction expected on the basis of the response reported for the
aforementioned species, but corresponding very closely to the direction
of their migration to the breeding site. Perhaps the animals captured
at the breeding area had not been there sufficiently long to readjust
their Orientation to the shoreline. The length of time required by
California toads to Orient to a new Situation is not known, but in Bufo
fowleri reorientation to a new shoreline appears to begin within a
matter of hours and is nearly complete after two days (Ferguson and
Landreth, 1966).
Figure 2. Directional responses of 10 adult male toads (group 2) to an artificial
“chorus” in various positions when tested under a hazy night sky. only 2 or 3
stars visible (a-c), and the next morning (d) and afternoon (e) with the sun visible.
Ferguson and his coworkers have reported that at least some of the
anurans which they have studied (Acris gryllus, Acris crepitans, Bufo
fowleri) are capable of orienting to a home shore when released under
night skies, apparently using stellar cues; in some instances an appar-
ent bidirectional response, some animals moving in the expected
Y-axis direction and some moving 1 80° opposite, has also been seen
when tested under stars, but in no case have Statistical tests been
employed to determine the probability that the dispersions differed
from random. In our nocturnal tests with Bufo boreas a tendency to
bidirectionality similar to that reported for other species was noted,
16
Bulletin So. Calif. Academy of Sciences
but statistically the dispersion did not differ from that expected by
chance alone. Hence we were unable to obtain unequivocal evidence
for stellar Orientation capabilities in these toads. This failure of the
toads to show clear ability to Orient under a night sky is somewhat
surprising in view of the fact that the migration to the lake occurred
exclusively after dark.
If the toads are unable to use stellar cues in orienting (a hypothesis
which needs to be more carefully tested) it is still possible that their
nocturnal migrations are at least in part guided by celestial cues. Per-
haps the animals establish their direction of travel during daytime
using the sun as a guide, obtain a fix on local landmarks and then at
night move in relation to these objects. Or possibly in their usual
habitat the animals first determine the direction of travel by windbome
odors from the lake, a source of Information unavailable to them in the
arena, then maintain a more or less straight line of travel to the odor’s
source by guiding on the stars. It is likely that odors play a part in
navigation in this species, for blinded toads when released within their
familiär area are able to Orient to the breeding ponds whereas anosmic
animals appear to be incapable (Tracy and Dole, submitted manu-
script). Thus it is apparent that vision is not the only source of infor-
mation used by these animals.
Several investigators have found evidence that anurans are attracted
to conspecific choruses and it has been widely suggested that auditory
cues play a role in guiding their movements to the breeding site. Old-
ham (1966 and 1 967) found that both American toads and green frogs
tend to be attracted to tape recorded choruses when released in unfamil-
iar territory but generally Orient in the direction of the breeding site,
irrespective of the direction of the chorus, when released in a familiär
region. The few tests reported here also suggest that audition plays a
secondary role in guiding the California toad, for under a hazy night
sky when celestial Orientation apparently was not possible the animals
tended to be attracted to the “chorus,” while in daylight the majority
moved in the direction of their previous migration rather than to the
sound. Presumably visual cues, if they provide adequate information,
take precedence over auditory cues. This is not surprising since Bufo
boreas produced a very weak chorus, males vocalizing only when
clasped by other males, which is often inaudible to the human ear at a
distances of less than a hundred meters. It should be noted here that
the possibility that odors from the toads producing the “chorus” rather
than the chorus itself provided guiding cues cannot be ruled out in the
present tests. Tape recorded choruses should be used in any future
experiments.
Celestial Orientation
17
From the above data it is apparent that much more work is needed
before we can begin to understand the role which the various environ-
mental cues play in guiding the migratory movements of the California
toads. Although these data suggest that the sun plays a part, possibly
an important one, in Orientation a much more thorough investigation
is needed to analyze its role, as well as that of the stars and moon, in
direction finding. Much better control of the experimental conditions
than was possible in this study, including the use of each animal only
once or the rotation of the arena to prevent recognition of and Orienta-
tion to particular parts, the use of a tape recorded chorus, and the use
of a larger number of animals under a wider variety of sky conditions,
are certainly called for. We can only hope that this preliminary work
will encourage someone with access to a sizeable population of these
animals to continue the investigation.
Acknowledgments
We are sincerely grateful to the senior author's wife, Barbara, and brother,
William, who participated in all aspects of this study. Thanks also go to the owners
and managers of Seminole Hot Springs Trailer Park for allowing us to collect on
their property.
Literature Cited
Batschelet, E. 1965. Statistical methods for the analysis of problems in animal
Orientation and certain biological rhythms. AIBS Monogr., Washington, D.C.
Durand, D. and J. A. Greenwood. 1958. Modifications of the Rayleigh test for
uniformity in analysis of two-dimensional Orientation data. J. Geol. 66: 229-
238.
Ferguson, D. E. and H. F. Landreth. I 966. Celestial orientation of Fowler’s toad,
Bnfo fowleri. Beliavionr. 26: 105-123.
Ferguson, D. E., H. F. Landreth, and J . P. McKeown. 1 967. Sun compass orien-
tation of the northern cricket frog, Acris crepitans. Anim. Belwv. 15: 45-53.
Ferguson, D. E., H. F. Landreth, and M. R. Turnipseed. 1965. Astronomical
orientation of the Southern cricket frog, Acris gryllus. Copeia. 1965: 58-66.
Ferguson, D. E., J. P. McKeown, O. S. Bosarge, and H. F. Landreth. 1 968. sun-
compass orientation of bullfrogs. Copeia. 1 968:230-235.
Landreth, H. F. and D. E. Ferguson. 1966. Evidence of sun-compass orientation
in the chorus frog, Pseudacris triseriata. Herpetologica. 22: 106-1 12.
Landreth, H. F. and D. E. Ferguson. 1967. Movements and orientation of the
tailed frog, Ascaphns truei. Herpetologica. 23: 81-93.
Bulletin So. Calif. Academy of Sciences
Oldham, R. S. 1 966. Spring movements in the American toad, Bufo americanus.
Can. J. Zool. 44: 63-100.
Oldham, R. S. 1967. Orienting mechanisms of the green frog, Rana clamitans.
Ecology. 48: 477-491.
Tracy, C. R. and J. W. Dole. Orientation of displaced California toads, Bufo
boreas, to their breeding sites. (Submitted manuscript).
Accepted for publication December 11,1 968.
Bull. So. Calif. Acad. Sei. 68(1): 19-29, 1969
HYMENODORA G LA CI ALIS
(DECAPODA: NATANTIA) FROM THE ARCTIC BASIN
Alan D. Hävens and Wesley L. Rork
Department of Biological Sciences
University of Southern California
Los Angeles, California 90007
Abstract: A study was made of the shrimp Hymenodora
glacialis (Buchholz) collected from Fletcher’s Ice Island
from June 1965 to January 1967, when the island drifted
over the deep water of the Canada Basin. A vertical distri-
bution was determined by means of horizontal and vertical
plankton net hauls, which accounted for most of the catch;
the animal is most abundant from 350-1000 m, less abundant
from 1000-3800 m, and least abundant from 0-350 m. In
addition, some specimens were taken from near the bottom
using a small biological dredge at depths of about 2000 and
3800 m. No seasonal or geographical variations in abun-
dance, or size variations with depth, etc. were indicated. An
analysis of sampling gear used indicated that more shrimp
were caught when a higher tow speed was employed or a
larger sized mesh used: there was no correlation between
the size of the animals caught and tow speed.
Most of the animals captured were from 10 to 30 mm
long; a study of secondary sexual characteristics indicated
that few specimens were mature, and that sex determination
is difficult with individuals under 40 mm, small males re-
sembling females. Only a few females were ovigerous. A
study of gut contents suggests that the species has a diverse
diet, but that the most important food is copepods, with
chaetognaths and radiolarians also being fairly important.
Gut contents were often nearly intact in large specimens.
Two larger animals had ellobiopsid parasites.
Introduction
In the biological collections made from Fletcher’s Ice Island T-3 from
June 1965 to January 1967, the natant decapod crustacean Hymeno-
dora glacialis occurred in sufficient numbers to attract our attention.
This fact was particularly interesting, since specimens were caught
using a wide variety of sampling techniques, none of which would be
considered especially good for catching shrimp; and also because this
organism is among the largest of the Arctic pelagic invertebrates, and
may therefore be an important factor in the ecology of the region.
Previous records indicate that Hymenodora glacialis is abundant
19
20
Bulletin So. Calif. Academy of Sciences
in the Greenland and Norwegian Seas and Baffin Bay, although it has
been recorded from as far south as 30° N latitude in the Atlantic Ocean
(Stephensen, 1935; Sivertsen and Holthuis, 1956). It appears to be
primarily a meso- and bathypelagic species in the northern North
Atlantic, but has been taken at many depths. A few records exist of
the occurrence of this form in the Arctic Ocean: Sars (1900) recorded
it from north of the New Siberian Islands, Bogorou (1946) from north-
west of Sevemaya Zemlya, Dunbar and Harding (in press) from the
Beaufort Sea, and Bamard (see Mohr and Geiger, in press) from
north west of Ellesmere Island; and J. C. Yaldwyn, of the Australian
Museum, has some specimens from the Beaufort Sea under study.
Methods and Materials
Studies by biologists from the University of Southern California were
renewed on June 13, 1965, on Fletcher’s Ice Island T-3. Between then
and January 14, 1967, the island drifted within an area bounded by
74° and 79°N lat. and 139° and 177°W long. primarily over deeper
water of the Canada Basin but with several transects over the Chukchi
Rise (Fig. 1).
Hymenodora glacialis was collected primarily with Vi meter plank-
ton nets with no. 6, no. 20, and no. 24 mesh sizes, and a few specimens
were taken with a 1 meter net, mesh size no. 0 and no. 6; a wire mesh
sieve was employed in the capture of specimens at the surface of the
hydrohole, and dredging was done near the bottom with a Menzies
Trawl. Several variations of the latter were used, the modified Menzies
I, with an opening 1 by 0. 1 m; the modified Menzies II, with dimensions
of 0.9 by 0. 1 5 m, having the edges bent out into flanges, and the modi-
fied Menzies T-3, which used the MMII frame, but parachute material
for the bag, instead of the no. 0 mesh used in the other types.
A list of stations at which this species was taken, as well as a detailed
description of the collecting gear, will be deposited with the American
Documentation Institute, Auxiliary Publication Service, which is
administered by the Library of Congress, Washington, D.C. and can
also be made available upon request to the Arctic project of this depart-
ment. The collections were made by J. A. Pierce III, A. J. Mearns,
G. P. Owen, J. K. Dawson, and W. L. Rork. More than 700 samples
were examined, without the aid of magnification, and decapods were
removed for further study. Specimens were preserved in 7 per cent
formalin buffered with hexamethylenamine, or in Bouin’s and 70 per
cent ethyl alcohol, and total length (from the most anterior part of the
rostrum to the tip of the telson) was measured.
Decapods from the Arctic Basin
21
Data
All of the decapods collected from ice island T-3 were identified as
Hymenodora glacialis (Buchholz) a shrimp of the family Oplophoridae
on the basis of descriptions by Sars (1885), Kemp (1910), and Sivertsen
and Holthuis (1956); the last include a discussion of the differences
between H. glacialis and H. gracilis (Smith), which are very similar
and which at least in the North Atlantic have overlapping distributions
(Sivertsen and Holthuis, 1956). The criteria used to distinguish the
two species are as follows: in H. glacialis the lobe over the second
segment of the antennal peduncle is broadly rounded, while in H.
gracilis it is produced to a blunt point; H. glacialis has a groove on the
carapace which is lacking in H. gracilis (see Sivertsen and Holthuis,
1956, Fig. 12), and H. gracilis has a podobranch on the second max-
illiped which is lacking in H. glacialis. Finally, H. glacialis has a shorter
rostrum with a more convex lower margin and swollen upper lateral
surfaces, while in H. gracilis the rostrum is longer and the upper
lateral surfaces concave.
Adult specimens examined all lacked the long and pointed distal
end of the rostrum illustrated for H. gracilis in Sivertsen and Halthuis
(Fig. 13); the end of the rostrum was blunt and up-turned, and there
were 4 to 7 rostral teeth above. Larval shrimp in the collection are
generally similar to the description of H. glacialis larvae given by
Stephensen (1935).
22
Bulletin So. Calif. Academy of Sciences
Efficiency of Sampling Gear. The effects of variations in sampling
technique and sampling conditions were examined prior to any attempt
at analysis of possible spatial or temporal fluctuations in the shrimp
population. Vertical net tows were made by rapidly hauling the net
through the water column; for the most hauls a closing device was used,
for greater accuracy in determining depth distribution. Horizontal
net tows were accomplished by stationing the net at various depths,
and relying upon the difference in speed between the ice island drifting
on the surface and the current below to produce an effective tow speed.
A convention was adopted to give a rough indication of relative tow
speed: 0, if the cable was hanging vertically into the water, 1 , if there
was a slight cable angle, 2, if there was a marked angle, and 3, if the
cable was touching the edge of the hydrohole cut in the ice, indicating
a high tow speed. Vertical net hauls, of short duration, captured one
shrimp per six hauls, and horizontal net tows, which often lasted 10
or 20 hours, caught one per four hauls.
Table 1
Efficiency of capture, Ratio of
as related to Mesh size shrimp captured
Variation in: per haul
a. mesh size, horizontal
No. 24
1/20
and vertical hauls
No. 20
1/6
combined.
No. 6
1/3
Speed, mlmin.
b. tow speed, vertical
10-14
1/36
net hauls.
15-19
1/17
20-39
1/5
40-80
1/5
Wire angle
c. Wire angle, hori-
0-1
1/8
zontal net hauls.
2
1/5
3
1/1
Variant
No. shrimp
No. hauls
d. Design of Menzies
M MI
2
9
trawl
MMII
15
16
MM T-3
0
6
It can be seen from Table la-c, that more shrimp were captured when
a larger mesh size was used, and when tow speed for vertical hauls or
wire angle for horizontal tows was greater. The Menzies Trawl also
Decapods from the Arctic Bas in
23
captured a number of shrimp (Table ld); here again the catch was
improved when the MM II variant, with its larger mouth, was employed.
Vertical Distribution. The only precise determination of the depth
at which Hymenodora glacialis occurs was made with the vertical
closing net (Table 2a). One shrimp was caught on a tow between 1 900
and 2500 m, the record depth for H. glacialis captured with this device,
during the study. Shrimp were not caught in horizontal tows just under
the ice surface, nor were they found in traps lowered to depths of 0-4
m, though one specimen was captured at the surface of the hydrohole
with a wire mesh scoop. Vertical, non-closing net hauls between depths
over 1 00 m and the surface appear to confirm the above indicated dis-
tribution pattem (Table 2b).
Table II
Vertical Distribution
Dcptli Range
( Meters)
% Water
Filtered
Shrimp Per
Meters Filtered
a. Vertical closing
0-500
27
1/7500
net
500-1000
19
1/2300
1 000-2000
32
1/8900
2000-3780
22
0
Depth Range of Hanl,
Shrimp Per
( Meters )
Hanl
b. Vertical
0-250, 300
0
non-closing net
0-500
1/7
0-1000, 2000, 3000, 3785
1/1
Depth Range
% of
Shrimp Per
(Meters)
Tows
Tow
c. Horizontal net
0-350
34
1/11
400-800
22
4/7
900-1300
23
2/7
1400-1900
9
1/3
2000-2900
9
1/7
3000-3780
3
1/4*
*Unreliable because of small number of tows.
Horizontal net tows must be regarded as less reliable than vertical
closing net tows, because of the possibility of sample contamination
during retrieval, which will increase with depth, but the distribution
of the catch is somewhat similar (Table 2c). Though less reliable than
closing net hauls, the number of specimens (76) is enough for a Statisti-
cal analysis; a chi-square test was employed, the expected numbers
24
Bulletin So. Calif. Academy of Sciences
of shrimp being estimated on the basis of the proportions of the total
hours fished within each depth ränge. The expected and observed
numbers of shrimp were found to be significantly different at a P .01
level with 4 degrees of freedom, indicating a non-uniform distribution
with respect to depth; fewer shrimp were observed in the 0-350 m
ränge, and more in the 400-800 m ränge, than expected, while for the
remaining categories of 900-1300, 1400-1900, and 2000-3800 m,
the differences were not great.
Some 33 hauls were made with the modified Menzies Trawl, an
average of one shrimp per two hauls being taken; most of the shrimp
were caught in hauls which sampled the bottom at about 2000 and
3800 m. While a closing device was not used, contamination during
retrieval was unlikely, considering the shape of the frame; the MM II
Version proved to be much more efficient than the vertical nets at cap-
turing shrimp.
Seasonal Variations and Geographical Distribution. As the ice
island was generally drifting in a westerly direction, from 141° to
176°W, when shrimp were captured (Fig. 1), geographical variations
in the catch were looked for. It was found that fluctuations in horizon-
tal and vertical net catches were related to variations in sampling
conditions and technique, and geographical or seasonal variations are
not apparent; it can only be said that the animal is present throughout
the year.
Size Variation. Data on the number of specimens of various size
ranges is presented in Table 3. An examination of the sizes of the
shrimp caught by nets run at different tow speeds revealed no correla-
tions; there is no evidence that larger shrimp will evade the nets more
than smaller ones. The data likewise do not support any strong correla-
tion between size and vertical or geographical distribution, or month
of capture.
Table III
Size Variation
Length Number
(mm)
< 10
2
10-19.5
73
20-29.5
31
30-39.5
17
40-49.5
13
50 <
1
Decapods from the Arctic Basin
25
Sexual Identification. Determination of the sex of the animals was
made on the basis of Sars’ (1885) description of the morphology of
the first and second pleopods of the male. In this species, small males
closely resemble females; many individuals under 40mm long could
not definitely be assigned to one sex of the other. There is a great deal
of Variation in the morphology of the endopod of the first pleopod in
immature specimens; the smallest male in the collection displaying
the adult condition of the first pleopods was 33 mm long. The appendix
masculina, which is found only in males, appears considerably later
than the appendix interna, which both sexes possess on the second
pleopods; the smallest specimen having this organ was 35 mm long.
On the basis of the characters of pleopods 1 and 2, 1 1 specimens
were identified as males, and 15 as females; ambiguous specimens, all
under 40 mm long, were not sexually identified. Two adult females
were ovigerous, and the one measurable specimen was 45 mm long.
Two additional ovigerous females collected on 1 5 August 1 967 were
52 and 56 mm long. A tentative Separation of immature specimens
into males and females indicated that the proportions of the sexes in
the population are approximately equal.
Table IV
Stomach Contents
Number of shrimp
Item contained in
Copepods
59
Chaetognaths
1 1
Radiolarians
10
Polychaetes
3
Amphipods
2
Ostracods
1
Hymenodora (jaw)
Foraminifera
1
Bryozoan *
1
Filamentous Algae
Muscle Fibres or
Tubule Bundles
1
Stomach Contents. Ninety-seven shrimp were dissected for stomach
contents: 12 from vertical and 70 from horizontal tows, 15 from
Menzies Trawl catches, and 1 from the surface. Eight specimens were
still in a larval stage and lacked well defined stomachs. The stomachs
were removed from the rest of the individuals, which were 1 2 mm and
26
Bulletin So. Calif. Academy of Sciences
above, and of these, 1 7 were empty. Material in the stomachs of the
remainder was identified, and the number of specimens containing
various food items in their stomachs is listed on Table 4. Copepods
apparently constitute the most significant food item, many shrimp had
their stomachs full of them; in a few cases chaetognaths or polychaetes
filled the stomach. Radiolarians were commonly present in small num-
bers. None of the other food items appears to be a very important
element in the diet. A great number of various sizes and shapes of
unidentifiable spines and tubules was found, which may have belong
to crustaceans, polychaetes, or radiolarians, but no identification of
such fragments was attempted.
In a great many cases, especially in larger specimens, the stomach
contents were in fairly good condition, which simplified the job of
identification of food items; many of the copepods were practically
unaltered. Radiolarians were usually fragmented, but were on occasion
intact; chaetognath remains took the form of intact heads or disas-
sociated hooks. The smallest specimens with any food material in the
stomach were 12 and 13 mm long; these had copepod remains, which
were generally fragmented. No correlations were found between
stomach contents, quantitative and qualitative, and seasonal, geo-
graphical, and vertical distribution, or the size of the animals.
Parasites. Two specimens had ellobiopsid parasites under the abdo-
men, a male 47 mm long, and an individual of undetermined sex, 37
mm long. The male was somewhat retarded in the development of its
secondary sexual characteristics, having a very narrow endopod on the
first pleopod, and the appendix interna only on the second pleopod,
but this may be simply coincidence.
Discussion
Hymenodora glacialis is regarded as meso- and bathypelagic. Kemp
(1910) noted its occurrence from 250 to 539 m off the west coast of
Ireland. According to Heegard (1941) it is often found on the surface
in northern waters as well as in the stomachs of birds that have ap-
parently been feeding at the surface, and Squires (1957) States that it
occurs from 250 to 2000 m off west Greenland. Our findings tend to
Support this pattern of vertical distribution: the species seems to be
commonest in the mesopelagic ränge, and somewhat less common in
the bathypelagic; and we have only a few records from the epipelagic
ränge — 3 specimens, including one from the surface.
It is interesting to note that the vertical distribution of this shrimp
in the Arctic Supports Coachman’s (1963) conclusions concerning
Arctic water masses; that there are three layers: a surface layer of cold,
Decapods from the Arctic Basin
27
Arctic water of low salinity, from the surface to 200 m; an intermediate
layer of warmer Atlantic water, of higher salinity, from 200 to 900 m,
and a layer of bottom water below 900 m, of intermediate temperature
but salinity equal to that of the Atlantic water. Hymenodora glacialis
is commonest in what would be considered the lower Atlantic water,
below 500 m, and less common in the bottom water; it is least common
in the upper layers, especially above 350 m. This would not be sur-
prising for an Atlantic form; its distribution might simply result from
the temperature and salinity characteristics of the Arctic water masses:
decrease in abundance in the bottom water could be the result of lower
temperature, and the even greater scarcity in the Arctic water, the
result of still lower temperatures and/or low salinity.
Gut content analysis of our arctic specimens confirm the findings
of Tchindonova (1959) that this species feeds on a wide variety of
things, including copepods, ostracods, chaetognaths, radiolarians,
and polychaetes, and that large specimens frequently contain whole
organisms in the stomach. Interesting though this may be, we must
avoid over-interpreting data on stomach contents of shrimp brought
in with plankton nets, as feeding on other organisms in the net is pos-
sible, while regurgitation of stomach contents could also take place.
Summary
Some 1 37 specimens of the shrimp Hymenodora glacialis (Buchholz)
were collected by five USC marine biologists from the Arctic ice island
T-3, from June 1965 to January 1967, by means of horizontal nets,
vertical closing nets, and bottom trawls, and from the surface. An
analysis of the sampling gear showed that more shrimp were caught
when a larger mesh size was used, when towing speed was greater in
the case of vertical tows, and when the current was faster in the case
of horizontal tows.
Vertical distribution of the species was found to be similar to that
indicated by previous records, the greatest number of shrimp being
found from 350 to 1000 m, fewer from 1000 to 2000 m, records of
shrimp from greater depths being somewhat uncertain, although there
is good evidence that they were taken from the bottom; and occur-
rences of shrimp from depths under 350 m being sparse. Application
of a Chi-square test indicated that the probability of accidentally
arriving at this distribution pattern is extremely low. No seasonal or
geographical variations were found within the limits of the study, or
size Variation with depth. The fact that so many shrimp were caught
over a wide geographic ränge seems to indicate that Hymenodora
glacialis is an important factor in Arctic ecology.
28
Bulletin So. Calif. Academy of Sciences
A study was made of changes in the secondary sexual characteristics
with the increase in size of the animal, determining that specimens
under 40 mm in length may not be classifiable as males or females. A
study of gut contents, the results of which may be taken with some
reservation, indicated that the most important food consists of cope-
pods, with chaetognaths and radiolarians also being fairly important,
although the species will take a wide variety of things. In many cases,
the gut contents were nearly intact, especially those of larger animals,
confirming a previous finding that the food is not chewed very much
before ingestion. Two of the larger specimens had ellobiopsid parasites.
Acknowledgements
Much thanks is due to Mr. Stephen R. Geiger for his many helpful
suggestions; and to the previously mentioned biologists who were
responsible for collection of the specimens. Thanks is also due to Dr.
Lowell Wayne, for his advice on the application of statistics and to
Dr. John Yaldwyn, for his helpful criticism of the manuscript, and the
use of the laboratory facilities of the Allan Hancock Foundation of the
University of Southern California is gratefully acknowledged. This
research was supported by contract Nonr 228 (19), NR 307-270
between the Office of Naval Research, Department of the Navy and
the University of Southern California; Professor John L. Mohr, Princi-
pal Investigator.
Literature Cited
Bogorov, V., 1946. Zooplankton collected by the “Sedov” expedition 1937-39.
(In Russian, English summary.) In V. Buinitski (ed.), Trudy dreifujusjtsjei
ekspeditri glawseiomorputu na ledokoljnom parocliode “T. Sedov”, 3: 336-
370.
Coachman, L. K., 1963. Water masses of the Arctic, Proceedings of the Arctic
Basin Symposium October 1962, Arctic Institute of North America, Wash-
ington, D.C. pp. 143-167.
Dunbar, M. J. and G. Harding. Arctic Ocean water masses and plankton: A re-
appraisal. In A. Jespersen and J. E. Sater, (ed.), Proc. Arctic Drifting Sta-
tion Symposium. Arctic Institute of North America. Washington, D.C.
(in press).
Heegard, P. E., 1941. Decapod crustaceans. Zoology of East Greenland. Med-
delelser om Gr0nland, 121: 1-12.
Kemp, S., 1910. The Decapoda Natantia of the coasts of Ireland. Fisheries, lre-
land, Sei. Invest., 1908, I.
Mohr, J. L. and S. R. Geiger. Arctic Basin faunal precis — Animals taken mainly
from Arctic drifting stations and their significance for biogeography and
Decapods from the Arctic Basin
29
water-mass recognition. In A. Jespersen and J. E. Sater, (ed.), Proc. Arc-
tic Drifting Station Symposium. Arctic Inst. N. Amer., Washington, D.C.
(in press).
Sars, G. O., 1885. Crustacea, 1. The Norwegian North Atlantic Expedition,
1876-1878, No. 6: 1-280.
Sars, G. O., 1900. Crustacea. The Norwegian North Polar Expedition. 1893-
1896 . 1 (5): 1-141.
Sivertsen, E. AND L. B. Holthuis, 1956.
Sivertsen, E. and L. B. Holthuis, 1956. Crustacea Decapoda (the Penaeidea
and Stenopidea excepted). Michael Sars North Atlantic Expedition. 1910,
5 (12): 1-54.
Squires, H. J., 1957. Decapod. Crustacea of the Calanus Expedition in Ungava
Bay, 1947 to 1950. J. Fish. Res. Bd. Canada 35: 463-494.
Stephensen, K., 1935. Crustacea Decapoda. The Godthaab Expedition. 1928.
Meddelelser om Grpnland, 80: 31-33, 66-74.
Tchindonova, J. G., 1959. Feeding of some groups of macroplankton in the
northwestern Pacific. Akademia nauk SSSR, Trudy Institut Okeanologii,
30: 166-189 [pp. 1 76-1 77 trans. from Russian].
Accepted for publication December 11,1 968.
Bull. So. Calif. Acad. Sei. 68(1): 30-35, 1969
USCIA MEXICANA, NEW GENUS, NEW SPECIES,
A WATERSIPORID BRYOZOAN WITH
DIMORPHIC AUTOZOIDS
William C. Banta
Department of Biological Sciences
University of Southern California
Los Angeles, California, 90007.
Abstract: Uscia mexicana is described as the monotypic
species of a new genus of the family Watersiporidae (Bryozoa,
Eurystomata, Cheilostomata). It is similar to Watersipora
in the structure of the frontal wall and epitheca, the shape of
the operculum, the presence of lucidae, and in the absence of
spines, avicularia and ovicells. It differs from Watersipora
in having erect, bilaminar colonies, larger zoecia, and in
possessing dimorphic autozoids. “Normal” A zoids pos-
sess skull-shaped opercula; B zoids, which make up less
than 1 per cent of the autozoids, possess enlarged, reinforced
opercula, augmented occlusor muscles, and distal, tooth-
like denticles. The significance and distribution of dimor-
phism of autozoids in the Cheilostomata are discussed.
Introduction
Düring March, 1949, fragments of what appears to be a single colony
of an unusually large cheilostome bryozoan were collected by the staff
of the R/V Velero 7K in a dredge sample taken at 24 m in the San
Lorenzo Channel, near La Paz, Baja California, Mexico. Examination
of the specimen reveals that it belongs to a new genus of the family
Watersiporidae.
Uscia, new genus
Diagnosis. A watersiporid ascophoran cheilostome, without spines,
avicularia or ovicells, possessing a single-layered tremocystal frontal
wall overlain by a darkened epitheca. Normal autozoids (“A zoids”)
predominate, but occasional zoids (“B zoids”) possess more heavily
reinforced opercula and enlarged opercular muscles. Genotype, Uscia
mexicana, new species.
Uscia mexicana, new species
Figures 1-4
Type locality. San Lorenzo Channel, 2 miles south of Espirito Santo
Island, Gulfof California; 24° 22' 13" N; 110° 19' 16"W;24m; 15
March, 1949; Velero Station no. 1738-49. Sample taken with a bio-
logical dredge; bottom “coral”.
30
New Species of Bryozoan
31
Holotype. Fragments of what appears to be a single colony, probably
fixed in 10 per cent formalin in sea water. The specimen was found
nearly dry in September, 1965, and placed in 70 per cent ethanol.
Deposited in the Allan Hancock Foundation, University of Southern
California, Los Angeles. AHF bryozoan type no. 1 54.
Paratype. Colony fragments at the British Museum (Natural History).
Description. The colony is erect, foliaceous and bilaminar (Fig. 1).
The Velero specimen appears to have been broken into several pieces;
fragments of other colonies may also be present. The largest piece is a
spectacular coralline growth approximately 4 cm by 7 cm (Fig. 1).
Its color is dark brown, but it is likely that the polypides and growing
edges were red in life (Banta, 1 968).
Zoecia are elongate, rectangular and unusually large, measuring
approximately 1.5 mm (1.2-1. 8 mm) long by 0.5 mm (0.4-0. 6 mm)
wide. The aperture (orifice) is terminal, occupying about a quarter of
the frontal surface. Both opercula and epithecae are dark brown,
nearly black. Chemical treatment in potassium hypochlorite solution
exposes the underlying frontal wall, a thick (approximately 90// )
single-layered lamina evenly perforated by a hundred or so evenly-
distributed pseudopores about 30 // in diameter (Fig. 2, psp ).
The colony is composed of two types of zoids. The dimorphism is
reflected in the morphology of the apertures and opercula in a way
similar to that described by Harmer (1900) in the genus Steginoporella
Smitt. Harmer named the zoids with smaller, more “normal” opercula
“A zoids”, and those with larger, more modified opercula “B zoids”.
Harmers terminology is followed here, although I do not imply that
the two types of dimorphism are necessarily related.
A zoids. A zoids make up the vast majority of autozoids. The aper-
ture is skull-shaped in outline (Fig. 2). The anter is shaped like a horse’s
hoof, and measures approximately 300 // in either dimension (270-
330 ti long by 280-350 // wide). The proximal border of the anter is
marked by a prominent pair of cardelles sunken slightly below the rim
of the peristome (Fig. 2, car ). The poster (po) is roughly hemispherical
and measures approximately 1 60 m (150-180//) wide by 80 // (70-90 // )
long. The entire aperture measures roughly 380 // long by 300 // wide.
The proximal and lateral parts of the aperture are bordered by a
low, smooth, imperforate portion of the frontal wall, the peristome
(Fig. 2 per). The distal rim of the aperture is formed by the frontal
part of the transverse wall.
The operculum of an A zoid is approximately the same size and
shape as the aperture (Fig. 3); its color is dark reddish brown. There
are three types of sclerites: (1) a thin marginal sclerite at the border of
New Species of Bryozoan
33
the porta; (2) a somewhat thicker sclerite bordering the vanna and
the most proximal parts of the porta; and (3) paired longitudinal con-
necting sclerites extending from about the middle of the vanna to the
distal third of the porta (Fig. 3, sei). At the junction of the porta and
the vanna, each connecting sclerite is extended laterally as the articu-
lation zone of the cardelles (art). A pair of tiny pits (“lucidae”; see
Banta, 1968) are present on the basal side of the vanna near the
proximal ends of connecting sclerites (lu). Viewed from the frontal side,
lucidae are represented by a pair of shining tubercles. Opercular occlu-
sor muscle fibers measure about 250 p (230-350/* ) from their origins
to their insertions on the tendon.
The operculum is surrounded by a ring of darkened epitheca 20-50 p
wide, which is herein named the “periopercular ring”. The periopercu-
lar ring covers most, but not all of the peristomial part of the frontal
wall, and overlaps parts of the distal zoid.
Avicularia are absent; there are no spines or ovicells.
Each zoid is provided with about 10 (8-12) lateral communication
organs with multiporous pore plates arranged along the basal border
of the lateral wall. Each plate is approximately 60 n in diameter and
bears about 10 communication pores. Lateral walls are three-layered,
consisting of two calcareous laminae and a central, dark brown inter-
calary cuticle (Banta, 1968). Transverse walls are unpaired and are
provided with 9-15 transverse multiporous pore plates arranged along
the sides and bottom of the septum.
B zoids. B zoids are much less common than A zoids, making up
perhaps 1 per cent of the autozoids in the colony. B zoids are similar to
A zoids in every observed respect except the morphology of the aper-
ture and operculum.
The aperture of a B zoid is very slightly larger than that of an A zoid,
measuring approximately 430 p (400-450 p ) long by 300 p (280-300 p )
wide. Anters of B zoids are proportionaltely longer than those of A
zoids, measuring about 330 p (300-350 p) long by 300 p (290-300/* )
wide. The lateral borders of the anter are decorated by a pair of longi-
Figures 1-4. Uscia mexicana, new genus, new species. 1. Holotype colony; scale
A. 2. KOCL-treated zoecia, six A zoids surrounding a B zoid; paratype; scale B.
3. Operculum and occlusor muscle of an A zoid seen from the basal side; s_cale C.
4. Operculum and ocolusor muscle of a B zoid seen from the basal side; scale C.
Abbreviations: an, anter of aperture; art, articulation region of cardelles; car,
cardelle; den, denticle; In, lucida; mus, occlusor muscle; per, peristome; po, poster
of aperture; psp, pseudopore; pt, porta of operculum; sei, sclerite; ten, tendon of
occlusor muscle; vn, vanna of operculum.
34
Bulletin So. Calif. Academy of Sciences
tudinal lappets. The distal border is overhung by a prominent bifid
denticle continuous with the skeleton of the transverse wall (Fig.
2, den).
The poster of the aperture in B zoids is significantly shallower and
broader than that of A zoids, measuring about 70 'ß (60-90 ß ) long by
200 ß ( 1 80-2 1 0 ß ) wide.
Opercula of B zoids are more heavily chitinized than those of A
zoids. The porta is roughly quadrangular; lateral borders are concave
because of the lateral apertural lappets (Fig. 4). The vanna is likewise
rectangular, corresponding to the shape of the poster. There are two
main types of sclerites: (1 ) a thick distal sclerite reinforcing the margin
of the porta; and (2) paired longitudinal connecting sclerites extending
from near the proximal edge of the vanna to the middle of the porta,
where tendons of occlusor muscles insert. The operculum is especially
thickened here (Fig. 4). Opercular occlusor muscles are much longer
in B zoids than in A zoids; they measure about 350 ß (340-450/i ) (Fig.
4, mus). A lucida occurs at the at the base of each longitudinal sclerite.
Discussion
Uscia mexicana appears to be closely related to the genus Watersipora,
which it resembles, in the following respects: (1) the frontal wall is an
evenly perforated tremocyst (see Canu and Bassler, 1920; 1930; (2)
the frontal wall is overlain by a darkly pigmented epitheca; (3) spines,
ovicells and avicularia are absent; and (4) it possesses a skull-shaped
aperture with proximal lucidae, a characteristic feature of Watersipora
(Osbum, 1 952). It differs from known species of Watersipora, however,
in three respects: (1) the colony is erect and bilaminar; (2) the zoecia
are much larger than those of any known species of Watersipora; and
(3) autozoids are dimorphic.
Although polymorphism is common (probably universal) in the
Cheilostomata, dimorphism in autozoids with functional polypides is
rare. The cases in which it occurrs can be divided into two categories:
sexual and non-sexual dimorphism.
A number of cheilostomes possess dioecious autozoids (Vigelius
1 884; Stach, 1 938), but sexual dimorphism of autozoids appears to be
known in only two cases. In Thalamoporella evelinae Marcus, female
zoids possess only 14 short (150/i) tentacles, compared with 17 long
(250 ß ) tentacles in sterile and male autozoids (Marcus, 1 949). Gordon
( 1 968) reports that in Hippopodinella adpressa (Busk), females possess
15-16 tentacles, but male zoids bear only eight (“four short and four
long”). Since males are apparently unable to feed, their Status as auto-
zoids is open to question.
New Species of Bryozoan
35
According to Hyman (1959: 327), non-sexual dimorphism of auto-
zoids is known in seven genera. Since the genera do not appear to be
closely related, it is likely that dimorphism has evolved independently
several times. In each case “normal” A zoids, with relatively unmodified
opercula, considerably outnumber B zoids, in which the opercula are
enlarged, reinforced, and provided with augmented occlusor muscles.
B zoids apparently represent incipient avicularia (Harmer, 1900;
Hyman, 1959). It is likely that dimorphism in JJscia mexicana is non-
sexual, but inasmuch as the polypides have not been adequately ex-
amined, the possibility cannot be excluded that the dimorphism is
sexual.
Acknowledgements
I thank Dr. John D. Soule, Mrs. Dorothy F. Soule, and Dr. Rüssel L. Zimmer
for their critical reviews of the manuscript. Contribution no. 327 of the Allan
Hancock Foundation.
Literature Cited
Banta, W. C., The body wall of cheilostome Bryozoa, I. The ectocyst of Watersi-
pora nigra (Canu and Bassler) J. Morph., 125: 497-506.
Canu, F. and R. S. Bassler, 1 920. North American early Tertiary Bryozoa. Bull.
U. S. Nat. Mus., 106: 1-879.
1930. The Bryozoan fauna of the Galapagos Islands. Proc. U. S. Nat.
Mus., 76: 1-78.
Gordon, D., 1968. Zooidal dimorphism in the Polyzoon Hippopodinella adpressa
(Busk). Nature, 219: 633-634.
Harmer, S. F. 1900. A revision of the genus Steganoporella. Quart. J. Micr. Sei.,
43: 225-297.
1902. On the morphology of the Cheilostomata. Quart. J. Micr. Sei.,
46: 263-350.
Marcus, E., 1941. Söbre Bryozoa do Brasil. Bol. Fac. Filos. Cienc. Letr. Univ.
Säo Paulo, Zool. 5: 3-169.
Osburn, R. C., 1952. Bryozoa of the Pacific coast of America. Part II, Cheilosto-
mata-Ascophora. Allan Hancock Pac. Exped., 14: 271-61 1.
Stach, L. W., 1938. Observations on Carhasea indivisa Busk (Bryozoa). Proc.
Zool. Soc. London, ser. B, 108: 389-399.
Vigelius, W. J., 1884. Morphologische Untersuchungen über Flustra membran-
aceo-truncata. Biol. Zentralbl., 3: 705-721.
Accepted for publication November 14, 1968.
Bull. So. Calif. Acad. Sei. 68(1): 36-42, 1969
A NEW SPECIES OF SPELEOCOLA
(ACARINA: TROMBICULIDAE), OFF A BAT,
PIZONYX VIVESI, FROM BAJA CALIFORNIA, MEXICO
Richard B. Loomis and James P. Webb, Jr.
Department of Biology,
California State College,
Long Beach
Astract: Speleocola cortezi, n. sp. is described from larvae
taken off Pizonyx vivesi (fish-eating bat) from Puertecitos,
Baja California Norte, Mexico. The genus Speleocola is
defined and new locality and host records are provided for
the other two species: Speleocola tadaridae Lipovsky from
Mexican bats, and Speleocola secunda Brennan and Jones
off bats and rodents from Costa Rica, Nicaragua and South-
ern Mexico.
Introduction
The genus Speleocola Lipovsky was proposed for a single species,
Speleocola tadaridae Lipovsky, found on the free-tailed bat, Tadarida
brasiliensis, from Oklahoma. The second species, Speleocola secunda
Brennan and Jones, was from another bat, Micronycteris hirsuta, of
Trinidad, B. W. I., and recently it was reported from a Panamanian
porcupine, Coendou rothschildi (Brennan and Yunker, 1966).
A third species is described below, based upon larvae found in the
ears of the fish-eating bat, Pizonyx vivesi , from Baja California del
Norte, Mexico.
In addition, the genus is defined and additional records are provided
for S. tadaridae from Mexico and S. secunda from Costa Rica, Nica-
ragua and Southern Mexico.
Genus Speleocola Lipovsky
Speleocola Lipovsky, 1952, type species Speleocola tadaridae Lipov-
sky, 1952.
Included species: Speleocola secunda Brennan and Jones, 1960 and
S. cortezi n. sp.
Diagnosis. — Larva. Member of subfamily Trombiculinae, tribe
Trombiculini, with scutum bell-shaped, constricted around bases of
posterolateral setae, anterolateral setae set back from anterior margin
and posterior to anteromedian seta; sensilla with expanded shaft and
expanded setules; posterior eye obscure and ocular plate indistinct;
palpal setal formula B/B/^NB; palpotarsus with six branched and
36
New Species of Chigger
37
nude setae (no subterminala); palpotibial claw trifurcate with a promi-
nent axial and two small lateral prongs; galeala nude; legs with two
claws and clawlike empodium without onychotriches; three genualae
I; and tarsi I, II and III each with several long nude setae on distal half.
Remarks. — Larvae of the genera Speleocöla and Microtrombicula
Ewing are closely similar. Crossley (1960) examined nymphs of Speleo-
cola tadaridae and recognized numerous similarities to nymphs of
various species of Trombicula, including four species currently re-
garded as members of Microtrombicula.
Speleocola cortezi , new species
Figure 1
Types. — Holotype and 1 5 paratopotypes from Puertecitos, Baja
California del Norte, Mexico, from 6 Pizonyx vivesi, fish-eating bat,
obtained 25 May 1963 by Ross Hardy and H. E. Childs: holotype and
4 paratopotypes, original number WJW630529-1; and 11 paratopo-
types, under original numbers WJW630529-5 (2), WJW630529-7
(2), WJW630529-8 (5), WJW630529-12 (1), and WJW630529-13
(1). The holotype and one paratopotype will be deposited in the Collec-
tion of the Rocky Mountain Laboratory, Hamilton, Montana and
other paratopotypes now in the chigger research collection at California
State College, Long Beach, California, will be deposited in appro-
priate institutions.
Diagnosis. — Larva of S. cortezi differing from S. tadaridae and S.
secunda in having the following characteristics: sensilla flagelliform
with shaft only slightly expanded (shaft greatly expanded with ex-
panded setules in other species), AW 31-36/i (less than 30 /t in other
species), dorsopalpotibial seta nude (branched in S. secunda ), coxa II
seta branched (nude in S. secunda ), second pair of stemal setae branched
(nude in S. secunda), pretarsala II present (absent in S. tadaridae) and
one pair of humeral setae (two pairs in S. tadaridae).
Description of holotype (all measurements in microns, with Varia-
tion of paratopotypes in parentheses). — Body engorged, 568 by 255,
eyes 2/2, with plate and posterior lens indistinct.
Dorsal setal formula 2-7-4-6-4-8-4-4 + 18, total 57; measure-
ments of humeral seta 37, seta of first posthumeral row 27, posterior
dorsal seta 22.
Ventral setal formula 2-2-4-4-4-4-6 + 28, total 54; measurements
of first sternal seta 27, posterior ventral seta 23.
Scutum: bell-shaped with constriction around bases of PL setae;
sensilla flagelliform with slightly expanded shaft, with numerous small
setules (having slightly expanded bases) along entire length.
38
Bulletin So. Calif. Academy of Sciences
Figure I . Speleocola cortezi n. sp. A. Scutum and eyes. B. Dorsal view of gnatho-
soma. C. Ventral aspect of palpal tibia and tarsus. D. Representative body setae,
1 St, first sternal, 2 St, second sternal and PD, posterior dorsal. E. Leg I; genu,
tibia and tarsus with nude and nearly nude setae and bases of branched setae,
with measurements of specialized setae in microns. F. Leg II. G. Leg III.
New Species of Chigger
39
Scutal measurements of holotype (with mean and extremes of 16
types, unless otherwise noted): AW 35 (34, 3 1 -36); PW 49 (49, 48-53);
SB 13 (13, 10-15); ASB 23 (25, 22-27); PSB 20 (20, 17-22); AP 31
(30, 26-32); AM 29 (29, 28-31); AL 17(18, 17-21); PL 35 (34, 32-38);
S 44 (1).
Gnathosoma: cheliceral blade with small tricuspid cap; cheliceral
base and capitular stemum lightly punctate. Galeala nude. Palpal
setal formula B/B/NNB; palpotarsus with 1 nude and 5 branched
setae, and tarsala, 5; palpotibial claw with 3 prongs.
Legs: branched setae of leg Segments with few branches. Specialized
setae: leg I, with 3 genualae and microgenuala, 2 tibialae and microti-
biala, and tarsus with tarsala, 13 (13, 12-15), distal microtarsala,
subterminala, parasubterminala, pretarsala, and 3 long, nude, distal
setae; leg II, with genuala, 2 tibialae, tarsala 14 (14, 13-15), micro-
tarsala and pretarsala; leg III, coxa with 1 branched seta, genuala,
tibiala, and 5 mastitarsalae of two types, three stout, two slender (see
Figure 1). All legs with Segments moderately punctate and terminating
in 2 stout claws and clawlike empodium without onychotriches. Leg
measurements, holotype and (in parentheses) mean and extremes of
16 types: 1,203 (210, 1 82-227); II, 176(184, 1 76-203); III, 190(206,
190-217); total, 569 (615, 561-628).
Taxonomie remarks. — The specific name S. cortezi refers to the
Sea of Cortez, another name for the Gulf of California, which is adja-
cent to the type locality and borders most of the known ränge of the
type host.
Ecological notes. — The larvae were found in the ears of 6 out of
17 examined fish-eating bats, Pizonyx vivesi, obtained from a cliff
crevice above the high tide zone at the eastern edge of Puertecitos.
Pizonyx vivesi occurs along the shores and on islands of the Gulf of
California and the Pacific coast of central Baja California (Hall and
Kelson, 1959). This bat has been found under rocks and roosting in
rock crevices, and it regularly forages over nearby waters.
Specimens examined. — Total, 16 larvae of type series.
Speleocola secunda Brennan and Jones
Speleocola secunda Brennan and Jones, 1960: 509-510, type from
St. Patrick, Trinidad, B. W. I., host Micronycteris hirsuta (hairy big-
eared bat), 23 June 1956; Goodwin and Greenhall, 1961 ; Brennan
and Yunker, 1966.
Specimens examined. — Total of 172 larvae: COSTA RICA:
LIMON PROVINCE, Finca La Lola, Rio Madre de Dios, 23 July
1963, 5 Saccopteryx bilineata (5); PUNTARENAS PROVINCE,
40
Bulletin So. Calif. Academy of Sciences
Finca Don Nicholas, 3 km N Tambor, 14 Nov. 1964, 2 Phyllostomus
discolor (6); SAN JOSE PROVINCE, 1 1 .3 km S La Georgina, 23 July
1963, 2 Molossus bondae (14). MEXICO: CAMPECHE, 7 km N,
51 km E Escärcega, 19 Dec. 1962. Peromyscus yucatanicus (85);
YUCATAN, 6 km S Merida, 18 Aug. 1962, Peromyscus yucatanicus
(8); 3 km N Piste, 26 July 1962, Peromyscus yucatanicus (2). NICA-
RAGUA: BOACA, 1 4 km S Boaca (220 m), 1 8 July 1 964, 2 Molossus
sinaloae (22); CHINANDEGA, San Antonio (35 m), 5 July 1966,
Nyctomys sumichrasti (4); GRANADA, 6.5 km SE Guanacaste
(660 m), 14 June 1966, Glossophaga soricina (1); RIVAS, 3 km N,
4 km W Sapoa (40 m), 26 June 1965, 5 Saccopteryx bilineata (18);
TRINIDAD: Bush Bush Forest, NarivaSwamp, 27 Aug. 1961, Micro-
nycteris megalotis (1); St. Andrew Co., Matura, 24 April 1959, Des-
modus rotundus (1); St. Patrick Co., Guapo, 23 June 1956, 2 Micro-
nycteris hirsuta (5 paratypes).
Additional record. — PANAMA: CANAL ZONE, along Pedro
Miguel River, 20 March 1962, Coendou rothschildi, (Brennan and
Yunker, 1966).
Remarks. — Larvae of this species have been taken from five in-
dividual rodents of three genera in addition to 1 9 bats of eight species,
suggesting that this species is not closely associated with one particular
kind of host or habitat. The chiropteran hosts probably have been
responsible for the widespread distribution of S. secunda from Trinidad
and Panama northward through Mesoamerica.
Measurements of tarsalae I and II and the legs of 95 specimens from
Campeche and Yucatan, Mexico revealed two size groups, the group
of 26 larger larvae and the group of 69 smaller chiggers. The larger
chiggers had longer tarsala 1(12, 1 2-1 3) and tarsala II (1 3, 1 2-1 5) in
6 larvae and longer legs (I, 154, 151-157; II, 129, 126-130; III, 151,
150-152; total, 434, 433-435) in 3 specimens. The smaller larvae had
tarsala I 9 (8-9), and tarsala II 10 (9-12), based on measurements of
15 specimens, and shorter legs (I, 141, 137-145; II, 114, 108-116;
III, 130, 124-136; total, 384, 377-395), based on 3 specimens.
Except for different sizes of certain structures no other differences
were detected. Larvae of both sizes were present in the single sample
of 85 larvae (26 large and 59 small) from the same individual host of
Peromyscus yucatanicus from Campeche. Therefore we believe that
these two size groups belong to a single taxon.
Speleocola tadaridae Lipovsky
Speleocola tadaridae Lipovsky, 1952, type from Merrihew Cave,
Woods Co., Oklahoma, 6 mi. S, 2 mi. W Aetna, Kansas, host Tada-
New Species of Chigger
41
rida mexicana( = T. brasiliensis) , Mexican free-tailed bat, 24 Aug.
1949; Loomis, 1956; Crossley, 1960; Loomis and Crossley, 1963.
Specimens examined. — Total of 42 larvae: USA: OKLAHOMA
Woods Co., Merrihew Cave, 6 mi. S, 2 mi. W Aetna, Kansas 15 Sept.
1948, Tadarida brasiliensis (paratype); TEXAS, Bexar Co., Fort Sam
Houston, San Antonio, 4 May 1954, Tadarida brasiliensis (11).
MEXICO: SINALOA, 4.3 km NW Topolobampo, 1 Aug. 1964, 5
Tadarida femorosacca (29), 6 Dec. 1964, T. femorosacca (1).
Additional records. — KANSAS, Barber Co., 3 mi. N, 2 mi. E
Sharon, 26 July 1952, Tadarida brasiliensis, (Loomis, 1956).
Remarks. — All of these larvae were taken from two species of cave
dwelling free-tailed bats, Tadarida brasiliensis and T. femorosacca.
Measurements of tarsalae I and II and of the legs of 14 larvae dis-
closed two size groups. The larger larvae (10 specimens) had longer
tarsala I (14, 13-16), and tarsala II (14, 13-15), as well as longer legs
(I, 195, 174-206; II, 173, 161-181; III, 191, 171-203; total, 559, 514-
582). The group of smaller larvae (4 specimens) had shorter tarsala I
(9, 8-9), tarsala II (12, 10-13) and shorter legs (I, 155, 148-159; II,
141, 128-150; III, 166, 158-180; total, 454, 434-466).
The presence of two size groups at the same locality (Topolobampo,
Sinaloa, Mexico) and from the same individual host ( Tadarida femoro-
sacca) seems to confirm that these two size groups belong to the same
taxon. Two size groups, seemingly of one taxon, also were found in the
samples of S. secunda.
Larvae of two sizes also have been reported in samples of another
bat chigger Whartonia glenni (Vercammen-Grandjean, Watkins and
Beck, 1965). These two size groups, presumably of the same taxon,
probably represent the predestined sexes.
Acknowledgments
Hosts and their parasites were provided by numerous individuals. We are grate-
ful to Dr. Ross Hardy for the opportunity to examine specimens of Pizonyx vivesi
and to W. J. Wrenn for the recovery of chiggers and their preparation on slides.
For the loan of specimens we would 1 ike to thank Dr. James M. Brennan of the
Rocky Mountain Laboratory, Hamilton, Montana. Appreciation is also extended
to Sr. Dr. Rodolfo Hernändez Corzo, el Director General, Direccion General de
Caza, Departamento de Conservacion de la Fauna Silvestre, Secretaria de Agri-
cultura y Ganaderia for permits to obtain mammals in Mexico. We are indebted
to Dr. J. Knox Jones, Jr., of The University of Kansas, for specimens from Nica-
ragua taken under United States Army Medical Research and Development
Command Grant DA-49-1 93-MD-221 5, and thank Dr. Fred S. Truxal of the
Los Angeles County Museum of Natural History and Dr. Charles A. McLaughlin
of the University of Wyoming for larvae from Costa Rica collected under the
United States Army Medical Research and Development Command Grant DA-
42
Bulletin So. Calif. Academy of Sciences
MD-49-1 93-63-G94. Studies upon which this paper was based were supported
by the U. S. Public Health Service Research Grant AI-03407 from the National
Institute of Allergy and Infectious Diseases. Finally, we wish to express our grati-
tude to Lee C. Spath and Elaine Katzer for the illustrations.
Literature Cited
Brennan, J. M. and E. K. Jones, 1960. Chiggers of Trinidad, B. W. I. (Acarina:
Trombiculidae). Acarologia, 2: 493-540.
Brennan. J. M. and C. E. Yunker, 1966. Ectoparasites of Panama. The chiggers
of Panama (Acarina: Trombiculidae). Field Mus. Nat. Hist., Chicago, lll.,
p. 221-266.
Crossley, D. A., Jr., 1960. Comparative external morphology and taxonomy of
nymphs of the Trombiculidae (Acarina). Univ. Kansas Sei. Bull., 40: 135-
321.
Goodwin, G. G. and A. M. Greenhall, 1961. A review of the bats of Trinidad
and Tobago. Bull. Amer. Mus. Nat. Hist., 122: 191-301.
Hall, E. R. and K. R. Kelson, 1959. The mammals of North America. The
Ronald Press Co., New York, 2 vols., 1083 p.
Lipovsky, L. J., 1 952. A new genus and species of chigger mite (Acarina, Trombi-
culidae). J. Kansas Entomol. Soc., 25: 132-137.
Loomis, R. B., 1956. The chigger mites of Kansas (Acarina: Trombiculidae).
Univ. Kansas Sei. Bull., 37: 1 195-1443.
Loomis, R. B. and D. A. Crossley, Jr., 1963. New species and new records of
chiggers (Acarina: Trombiculidae) from Texas. Acarologia, 5: 371-383.
Vercammen-Grandjean, P. H., S. G. Watkins and A. J. Beck, 1965. Revision
of Whartonia glenni Brennan, 1962 an American bat parasite (Acarina:
Leeuwenhoekiidae). Acarologia, 7: 492-509.
Accepted for publication November 27, 1 968
Bull. So. Calif. Acad. Sei. 68(1): 43-53, 1969
A COMPARISON OF THE FREE AMINO ACIDS IN TWO
POPULATIONS OF THE POLYCHAETOUS ANNELID
NEANTHES SUCCINEA
Alan J. Mearns1 and Donald J. Reish
Department of Biology,
California State College at Long Beach,
Long Beach, California 90801
Abstract: Free amino acids in ethanol extracts from whole
specimens of Neantlies succineci were measured with two-
dimensional paper chromatography. Specimens from the
Salton Sea, California, an inland saline lake, were compared
with samples from Alamitos Bay, California. Twenty-three
ninhydrin positive spots were identified in polychaetes from
both localities. Glutamine occurred in significantly higher con-
centrations in the Alamitos Bay samples while L-alanine
appeared significantly elevated in the Salton Sea specimens.
Introduction
The use of paper chromatography to detect biochemical differences in
concentrations of amino acids between related genera, species and
within species groups has been demonstrated in such invertebrates as
insects (Ball and Clark, 1953; Buzzati-Traverso and Rechnizter, 1953;
Micks, 1954 and 1956; Micks and Gibson, 1957 and Lewallen, 1957)
and gastropods (Kirk et al., 1954). Kirk et al. (1954) found species
specific differences in land snails; however, Stephen and Steinhauer
(1959) could not show statistically significant differences in amino acid
levels between laboratory-reared specimens of related species of cock-
roaches. Free amino acid surveys have been conducted at higher taxo-
nomic levels of marine, freshwater and terrestrial invertebrates where
distinct differences in composition were noted (Camien et al., 1951;
Duchäteau et al., 1 952; Simpson et al., 1 959).
Recent studies of the occurrence and role of free amino acids in the
body fluids of invertebrates have contributed new implications con-
cerning the physiology and ecology of at least the aquatic and marine
forms. For example, it has been suggested that the great differences in
amino acid levels between freshwater and marine invertebrates are
'Present address: Fisheries Research Institute, University of Washington, Seattle,
Washington 98105
43
44
Bulletin So. Calif. Academy of Sciences
indicative of their role in osmoregulation (Awapara, 1962; Camien
et al., 1951). Duchäteau et al. (1952) suggested that the abundance
of glycine in marine crustaceans, as opposed to freshwater forms, is
indicative of its participation in some unknown osmotic process.
Furthermore, it is now apparent that a number of “soft-bodied” marine
invertebrates are capable of removing free amino acids from dilute
solution in their environment through a process that does not necessarily
involve the digestive tract (Stephens, 1963, 1964). Active uptake of
free amino acids from an ambient medium may play a role in nutrition
in the polychaete Clymenella torquata (Stephens, 1 963) and the brittle-
star Ophiactis arenosa (Stephens and Virkar, 1966) or osmoregulation
in the polychaetes Nereis limnicola [ = Neanthes limnicola ] and
Nereis succinea [ = Neanthes succinea ] (Stephens, 1 964).
These observations suggest that, beyond possible genetic control,
natural environmental conditions may play a role in determining the
composition and quantity of free amino acids, at least in marine and
euryhaline invetebrates. Most of the available information has so far
been obtained through controlled laboratory studies; only a few com-
parative observations have been made on natural populations (Hillman,
1965 and Schäfer, 1961) where qualitative differences in amino acid
patterns have been reported. Insofar as is known, no one has quanti-
tatively compared the free amino acid composition of two isolated
populations of marine invertebrates. The polychaetous annelid Nean-
thes succinea (Frey and Leukart) was chosen for this study because of
previous knowledge of its biology (Banse, 1954) osmoregulatory abili-
ties (Smith, 1 959), its capacity to vary uptake of free amino acids under
various environmental conditions (Stephens, 1964), and of the geo-
graphically isolated population in Salton Sea, California.
Salton Sea is a saline lake which was formed during the period of
1 904-1907 when flood waters from the Colorado and Gila Rivers filled
a below-sea level basin which contained salts from an ancient sea. The
present-day salinity is similar to that of the ocean but the inorganic
elements are in different proportions and are changing (Carpelan, 1 961
and Pomeroy, 1 965). The polychaete Neanthes succinea was probably
introduced accidentally into Salton Sea in 1930 from Mission Bay, San
Diego, California (Carpelan and Linsley, 1961). It is possible that
some degree of differentiation, genetic or otherwise, may have or may
be occurring in these worms. No morphological differences of syste-
matic importance were noted by Hartman (1945) nor by us in 1967.
The only differences noted by us in living material in 1967 were the
smaller size and redder color of specimens from the Salton Sea.
The purposes, therefore, of the present study were threefold: (1) to
Amino Acids in Polychaetes
45
semi-quantitatively characterize the free or easily extracted amino
acids of N. succinea with a reproducable Chromatographie method,
(2) to estimate the individual Variation for each amino acid, and (3) to
determine whether or not significant quantitative differences could be
detected between two isolated populations of this species.
Materials and Methods
Free amino acids were analyzed from ethanol extracts of whole
specimens using two-dimensional descending paper chromatography
as described below.
Collection and Preparation of Material
Specimens of Neanthes succinea were obtained from the Colorado
Lagoon area of Alamitos Bay, Long Beach, California and from the
Desert Shores Marina, Salton Sea, Imperial County, California during
July 1 967. Specimens were collected from among the fouling organisms
attached to floating boat docks. An extraction similar to the procedure
described in Stephens and Virkar (1966) was employed. Immature,
undamaged specimens were placed in petri dishes containing sea water;
they were then placed briefly on absorbent tissue to remove excess
water and then transferred to individual glass vials containing 80 per
cent ethanol. The vials were transported to the laboratory in an ice
ehest at 4 to IOC.
Vials were stored in a refrigerator at 4 C for a total elapsed period of
48 to 50 hours from the time of collection. Extraction was terminated
by removing specimens from their respective vials and weighed. Extracts
representing undamaged, immature specimens weighing between 25
and 75 mg alcohol weight were utilized for the final analysis.
Desalting Extracts
The ethanol extracts were individually desalted following a procedure
modified from Allen and Awapara (1960). Dowex 50W x 8, 100-200
mesh ion exchange resin was prepared in 4N HCl a few hours prior to
use. Polyethylene columns measuring 200 x 1 5 mm were plugged with
glass wool and their flow rates equalized. The Dowex resin was rinsed
with distilled water to neutral pH and a 6 ml slurry was pipetted into
each column with continued rinsing. After the distilled water had drained
from the columns, 2.5 ml of each extract was pipetted over the resin
bed. Salts were eluted with ten 20 ml aliquots of glass distilled water.
All effluent was discarded. Each column was then eluted with two 1 2.5
46
Bulletin So. Calif. Academy of Sciences
ml aliquots of 4N NH4OH and the effluent collected in two test tubes.
The total effluent of 25 ml was concentrated to 0.5 ml with an Evapo-
Mix Rotary Evaporator at 60 C.
Chromatography
Each sample was spotted with a 2 ml automatic micropipette approxi-
mately 10.5 cm from one comer of a 46 x 57 cm sheet of Whatman
Number One Chromatographie paper. Drying was accelerated by using
a specially designed table that allowed hot air of about 60 C to come in
contact with the spot; the spot diameter did not exceed 10 mm. After
each spot was dried, the chromatograms were placed in a chromato-cab
descending chromatography chamber previously saturated with N-
butanol (reagent grade), glacial acetic acid (reagent grade) and glass
distilled water in the proportions of 240/60/200 by volume. The papers
were removed after 24 hours and dried under a hood for 1 2 hours. The
papers were chromatographed in the second dimension using redistilled
Figure I . Diagram of a typical chromatogram of the 23 ninhydrinpositive spots
from alcohol extracts of Neanthes succinea. 3, cystine and glutathione; 4, lysine
and phospho-ethanolamine; 5, aspartic acid; 6, glutamic acid; 7, L-amino adipic
acid; 8, serine; 9, glycine; 12, arginine; 13, L-threonine; 14, glutamine; 21, L-
alanine; 22, beta-alanine; 24, tyrosine; 32, valine, norvaline and L-methionine;
35, phenylalanine, leucine, isoleucine and norleucine; 36, asparagine; 38, proline;
A, cysteic acid, ornithine and hydroxylysine; B, histidine and 1-methyl histidine;
C, citrulline and hydroxy-proline; D, sariosine (possibly); E and F, unidentified.
Amino Acids in Polychaetes
47
phenol saturated with water for 1 8 hours. The papers were dried, the
chromatograms were treated with 0.2 per cent ninhydrin in 95 per cent
ethanol using a roller apparatus (Hrubant, 1961) and developed in a
CO2 saturated oven at 60 C for 30 minutes.
Ninhydrin-positive spots were identified from a previously pre-
pared master chromatogram of N. succinea (Fig. 1) which was modified
from Hrubant (1965). Individual spots were cut and placed in vials
containing 10 ml of 50 per cent ethanol. The papers were eluted for two
hours. Spots which overlapped (Fig. 1 ) were treated as a unit. A total of
23 different spots were obtained (Fig. 1). Optical densities of the nin-
hydrin eluates were measured with a Beckman Model B Spectrophoto-
meter at 570m/zfor blue and purple spots and 330m/i for orange, yellow
and brown spots.
Results
The data compare 12 individual samples from each population of N.
succinea for the 23 ninhydrin-positive spots by locality, as ranges,
means, and variances in Table 1 . The optical densities were determined
and these measurements were converted to relative percentages of the
total optical density; these calculations constitute the ranges and means
included in Table 1. The relative position and identification of each
spot is given in Fig. 1 . Some spots represented more than one substance,
and at least two spots contained unidentified material. Taurine and
cysteic sulfanilic acid were not present on the chromatograms; if they
had been present, they were lost during the desalting procedure with
Dowex 50 (Hrubant, 1965).
Ten of the 23 ninhydrin-positive spots represent at least 90 per cent
of the total ninhydrin-positive material; these data, together with their
Standard deviations, are presented in Table 2. As seen in Table 2,
glycine accounted for one-third of the ninhydrin-positive material on
the chromatograms with L-alanine, glutamic acid, serine, glutamine,
and proline accounting for about 40 per cent.
Differences in Mean Amino Acid levels between population
Each of the 23 ninhydrin positive spots was present on at least 67
per cent of the chromatograms. The Student t-test was applied to deter-
mine whether or not any significant differences between the sample
means for each amino acid occurred. The results of this analysis are
presented in Table 1. Six spots showed significant differences; three at
the 5 per cent level namely, L-amino adipic acid, beta-alanine, and
phenylalanine and the leucines, and three at the 1 per cent level, namely,
glutamine, L-alanine, and L-methionine and the valines. The first,
Table 1
Ranges, Means, Variance, Student T-Test, and Variance F-Tests for Ninhydrin-Positive Spots of Neanthes succinea
from Salton Sea and Alamitos Bay
48
Bulletin So. Calif. Academy of Sciences
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Amino Acids in Polychaetes
49
second, and fifth amino acids listed were elevated in the Salton Sea
material; whereas, the amino acid concentrations were higher for the
remaining three of these amino acids in the Alamitos Bay samples.
Variation in Amino Acid Levels
Variances for each spot were calculated separately to give an indi-
cation of intrapopulation variability (Table 1). The variances for each
amino acid from the two populations were compared using the F-test.
Six ninhydrin-positive spots exhibited significant differences at the 5
per cent level, these were: tyrosine, asparagine, L-amino adipic acid,
arginine, and possible sarcosine. Three spots showed significant differ-
ences at the 1 per cent level; these were, L-threonine, L-methionine
and the valines, and phenyalanine and the leucines.
Discussion
The free amino acid extracts of Neanthes succinea appear to contain a
relatively small group of compounds in high concentrations and a more
numerous group in lower concentrations. Glycine, L-alanine, glutamic
acid, serine, glutamine and proline accounted for more than 75 per cent
of total concentration of free amino acids detected by paper chroma-
tography. The means and Standard deviations for the most abundant
amino acids are included in Table 2. Generally, these observations
agree with those on other marine invertebrates. High levels of proline
and glutamine were reported by Clark (1964) for the polychaete
Nephtys hombergi, high levels of glycine were reported from crusta-
ceans and mollusks (Awapara, 1962; Kittredge, et al., 1962). Simpson,
et al., (1959) found alanine, aspartic acid, arginine, glycine and taurine
in 17 species from coelenterates, arthropods, mollusks, and echino-
derms. Apparently glutamic acid, serine, and lysine are more concen-
trated in N. succinea than in the above organisms.
Six ninhydrin positive spots were present in significantly different
levels between samples of Neanthes succinea (Table 1). Glutamine,
L-alamine and beta-alanine exhibited these differences without sig-
nificant variant differences within the population. Glutamine was pres-
ent in higher concentrations from Alamitos Bay specimens than from
the Salton Sea ones (9.6 ± 5.6% to 3.7 ± 1.3%). L-alanine and beta-
alamine were higher in the Salton Sea norms. The remaining three nin-
hydrin-positive spots, L-amino adipic, and L-methionine and the val-
ines, and phenylalanine and the leucines, differed not only between the
two populations but also within the population (Table 1 ).
Amino acid differences have been found among populations of other
50
Bulletin So. Calif. Academy of Sciences
invertebrates. Schäfer (1961) reported that specimens of the abalone
Haliotis cracherodi and of the shore crab Pachygrapsus crassipes from
polluted areas lacked asparagine which was present in specimens col-
lected from non-polluted areas. Chromatographie differences were
detected between two Stocks of the eastern oyster Crassostrea virginica
maintained under identical laboratory conditions (Hillman, 1964).
The Chromatographie patterns varied with changes in salinity and avail-
able food. Stephen and Steinhauer (1959) were unable to detect signifi-
cant difference of amino acid levels in several species of cockroaches.
Table 2
Means and Standard Deviations (S.D.) in Relative Per cent for the Ten
Most Abundant Amino Acids in Neanthes succinea.
Amino Acid Salton Sea Alamitos Bay
Me cm
S.D.
Me an
S.D.
Glycine
35.8
±4.9
31.8
±2.1
L-Alanine
15.9
± 3.1
8.2
± 3.8
Glutamic Acid
8.7
±3.3
9.2
±4.8
Serine
8.0
± 1.3
7.7
±2.1
Glutamine
3.7
± 1.3
9.6
±5.6
Proline
6.0
±3.3
5.8
±2.3
Lysine, phospho-ethanolamine
3.8
± 1.0
3.5
± 1.7
Unidentified (F)
3.1
± 1.2
2.9
± 1.0
phenylalanine, leucines
1.6
±0.7
4.3
±3.6
Asparagine
2.0
±0.6
2.3
± 1.2
The two samples of N. succinea from different populations can be
distinguished on the basis of the concentration of glutamine and L-ala-
nine (Table 2). In addition, the beta-alanine level is significantly higher
in the Salton Sea population, but the relative per cent concentration is
less than 1 .0. Although differences occurred among the other ninhydrin-
positive materials present on the chromatograms, their quantitative
significance could not be shown to be statistically significant. The dif-
ferences between these two populations may be the result of any of the
following, either individually or collectively: (1) genetic difference in
the ensuing 37 years of isolation, (2) environmental difference as a
result of the different proportions of inorganic elements, (3) original
differences between the population introduced from Mission Bay in
1930 and the Alamitos Bay population in 1967, (4) higher water tem-
peratures at Salton Sea, (5) food differences, (6) some factor or factors
yet unknown.
Amino Acids in Polychaetes
Acknowledgments
51
The authors thank Dr. H. Everett Hrubant for the use of his Chromatographie
apparatus employed in this study and to Mr. James Asher for his technical assis-
tance in preparing the Chromatographie procedure.
SUMMARY
1 . Free or easily extractable amino acids were quantitatively analyzed
J?y two dimensional descending paper chromatography of two geo-
graphically isolated populations of Neanthes succinea. The popula-
tion from Salton Sea which, was introduced in 1930, was compared
to one from Alamitos Bay, California.
2. A total of 23 ninhydrin-positive Spots were identified from both
localities. Glycine, L-alanine, glutamic acid, serine, glutamine and
proline accounted for about 75 per cent of the amino acids.
3. Student t-tests indicated that six amino acids differed significantly in
mean concentrations between populations. Glutamine and L-alanine
maintained homogenous variances between the two populations.
4. Variance F-tests indicated each population differed in degree of vari-
ability for 9 ninhydrin-positive Spots.
5. The reasons for these observed differences were discussed; they may
be the result of genetic, ecologic or other factors yet unknown.
Literature Cited
Allen, K. and J. Awapara, 1960. Metabolism of sulfur amino acids in Mytilus
edulis and Rangici cuneata. Biol. Bull. 1 18: 173-182.
Awapara, J., 1962. Free amino acids in invertebrates: a comparative study of
their distribution and metabolism. ln Holden, J. T., ed., Amino acid pools.
Distribution, formation and function of free amino acids. Elsivier, Amster-
dam. p. 158-175.
Ball, G. H. and E. W. Clark, 1953. Species differences in amino acids of Culex
mosquitos. Systematic Zool. 2: 138-141.
Banse, K.., 1954. Über Morphologie und Larvalentwicklung von Nereis (Nean-
thes) succinea (Leuckart) 1847. (Polychaeta errantia). Zool. Jalirb., Abt. f.
Anat. u. Ontog. Tiere. 74: 160-171.
Buzzati-Tra verso, A. A. and A. B. Rechnizter, 1953. Paper partition Chroma-
tograph in taxonomic studies. Science. 1 17: 58-59.
Camien, M. N., H. Sarlet, G. Duchateau, and M. Florkin, 1951. Non-protein
amino acids in muscle and blood of marine and freshwater crustaceans.
J. Biol. Chem. 193: 881-885.
Carpelan, L. H., 1961. Physical and Chemical characteristics. In Walker, B. W.,
ed., The ecology of the Salton Sea, California, in relation to the sport fishery.
Calif. Fisli and Game, Fish Bull. 1 13: 17-32.
52
Bulletin So. Calif. Academy of Sciences
Carpelan, L. H. and R. H. Linsley, 1961. The Pile Worm, Neanthes succinea
(Frey and Leukart). In Walker, B. W., ed., The ecology of the Salton Sea,
California, in relation to the sport fishery. Calif. Fish and Game, Fish Bull.
113: 63-76.
Clark, M., 1964. Biochemical studies on the coelomic fluid of Nephtys hom-
bergi (Polychaeta: Nephtyidae), with observations on changes during dif-
ferent physiological States. Biol. Bull. 127: 63-84.
Duchateau, G., H. Sarlet, M. N. Camien and M. Florkin, 1952. Acides
amines non proteiniques des tissue chez les mollusques lamellibranches et
chez les vers. Comparison des formes marines et des formes dulciocoles.
Arch. Intern. Pliysiol. 60: 124-125.
Hartman, O., 1945. The marine annelids of North Carolina. Duke Univ. Marine
Station., Bull. 2: 1-54.
Hillman, R. E., 1964. Chromatographie evidence of intraspecific genetic differ-
ences in the Eastern Oyster, Crassostrea virginica. Systematic Zool. 13: 12-18.
— 1 965. Chromatographie studies of allopatric populations of the Eastern Oyster,
Crassostrea virginica. Chesapeake Sei. 6: 1 15-1 16.
Hrubant, H. E., 1961. An Apparatus for applying reagent to paper chromato-
grams. J. Chromatog. 6: 94.
— 1965. Urinary amino acid differences in C57BL/6 and C3HeB/Fe inbred
mice, and their Fi hybrid. Can. J. Gen. and Cyto. 7: 530-535.
Kirk, R. L., A. R. Main and F. G. Beyer. 1954. The use of paper partition chro-
matography for taxonomic studies of land snails. Biochem. J. 57: 440-442.
Kittridge, J. S., D. G. Simonsen, E. Roberts and B. Jelinek, 1962. Free amino
acids of marine invertebrates. Invited discussion In Holden, J. T., ed.,
Amino acid pools. Distribution, formation and function of free amino acids.
Elsivier, Amsterdam, p. 176-186.
Lewallen, L. L., 1957. Paper chromatography studies of the Anopheles maculi-
pennis complex in California (Diptera: Culicidae). Ann. Entomol. Soc.
Amer. 50: 602.
Micks, D. W. 1954. Paper chromatography as a tool for mosquito taxonomy: the
Culex pipens complex. Nature. 174: 217-221.
— - 1956. Paper chromatography in insect taxonomy. Ann. Entomol. Soc. Amer.
49: 576-581.
Micks, D. W. and F. J. Gibson, 1957. The characterization of insects and ticks
by their free amino acid patterns. Ann. Entomol. Soc. Amer. 50: 500-505.
Pomeroy, R. D., 1965. A reconnaissance study and preliminary report on a water
quality control plan for the Salton Sea. California State Water Quality Con-
trol Board. 193 p. [mimeographed report].
Reish, D. J. and M. C. Alosi, 1 968. Aggressive behavior in the polychaetous anne-
lid family Nereidae. Bull. So. Calif. Acad. Sei. 67: 21-28.
Schäfer, R., 1961. Effects of pollution of the free amino acid content of two
marine invertebrates. Pacific Sei. 15: 49-55.
Amino Acids in Polychaetes
53
Simpson, J. W., K. Allen and J. Awapara, 1959. Free amino acids in some
aquatic invertebrates. Biol. Bull. 1 17: 371-381.
Smith, R. I., 1959. Physiological and ecological Problems of brackish waters.
Mar. Biol., Proc. Biology Colloquim, Oregon State College, p. 59-69.
Stephen, W. P. and A. L. Steinhauer, 1959. Intrapopulation variability as a
restrictive factor in the use of free amino acids in comparative taxonomy.
J. Kan. Entomol. Soc. 32: 123-127.
Stephens, G. C., 1963. Uptake of organic material by aquatic invertebrates. II.
Accumulation of amino acids by the bamboo worm. Clymenella torquata.
Comp. Biochem. Physiol. 10: 191-202.
— 1964. Uptake of organic material by aquatic invertebrates. III. Uptake of
glycine by brackish-water annelids. Biol. Bull. 126: 150-162.
Stephens, G. C. and R. A. Schinske, 1961. Uptake of amino acids by marine
invertebrates. Limnol. and Oceanog. 6: 175-181.
Stephens, G. C. and R. A. Virkar, 1966. Uptake of organic material by aquatic
invertebrates. IV. The influence of salinity on the brittlestar, Ophiactis are-
nosa. Biol. Bull. 131: 172-185.
Accepted for publication November 20, 1 968
Bull. So. Calif. Acad. Sei. 68(1): 54-56, 1969
RESEARCH NOTES
NOTES ON THE LIFE HISTORY OF FISHIA
EVE LI NA HANHAM1 (Lepidoptera)
This Information on Fishia evelina hanhami Smith resulted from the
capture of a gravid female taken in the Juniper Hills area of the Mojave
Desert, Los Angeles County, California, elevation 3500 ft., on October
18, 1967.
The female began ovipositing the following night after confinement.
The eggs were laid in masses on paper toweling placed within a screen-
topped jar, and remained in hibernation in this stage until the following
April. When the young larvae hatched they were fed on Linanthus
breviculus Greene. Later they were transferred to Phacelia tanacetifolia
Benth, which was more easily available, and which they readily
accepted.
Prior to the present study an adult male of this species was reared by
one of us (CH) from a mature larva collected in the field at Smokey
Valley, XYZ Creek, Tulare County, California, elevation 6200 ft.,
June 4, 1953, on an undetermined species of Linanthus. This record
simplified the choice of plants as food for rearing this species in the
laboratory.
Ovum
Figure 1A
Ovoid; width 0.8 mm; height 0.6 mm.
Surface covered with numerous ridges, approximately 60 in number,
running from base toward micropyle, but many terminate short thereof,
or fuse with others. Each ridge is topped along its length by a line of
round nodules placed close together, and the ridges themselves are so
closely approximate that it is difficult to see the character of the shell
surface between them. The micropyle is relatively very small and deep,
and many of the ridges seem to carry into it. However, some end abrupt-
ly at the micropylar edge where they form a slightly elevated circlet.
The ovum is strawcolor when freshly laid, turning to pinkish-brown
within a few days and remaining this shade throughout the winter hiber-
nation period.
Larva of 5 mm length
Head width 1.5 mm; color yellow-green; ocelli black; mandibles
dark yellow.
54
Life History of Fishia
55
Figure 1. A. Ovum. B. Larva of 15 mm length. C. Mature larva. D. Pupa, lateral
aspect. E. Pupa, ventral aspect. F. Cauda of pupa. All figures are enlarged. Exact
dimensions are recorded in the text.
56
Bulletin So. Calif. Academy of Sciences
Body, green on anterior two-thirds, shading to yellow on anal third.
A narrow longitudinal white stripe runs mid-dorsally the length of the
body. Dorso-laterally another longitudinal white line occurs. Between
it and the mid-dorsal line there are two white dots on each segment.
Latero-interior to this area is another broad longitudinal band bordered
inferiorly by a longitudinal yellow stripe, immediately superior to
which are the yellow spiracles. Below the yellow stripe the body is pale
yellow-green, as are the ventrum, legs and prolegs.
Larva of 1 5 mm length
Figure 1B
Similar in most respects to the 5 mm larva, except for size, and the
yellow shading which is absent on the caudal area. The head is a paler
yellow-green than the body. The ocelli are black on a white base. Head
width 1 .4 mm; head setae discernible without magnification; body setae
short and inconspicuous.
Mature Larva
Figure IC
Length 27 to 30 mm; width through center, 5 mm. Head, glistening
pale yellow. Width 3.5 mm; ocelli, a few black, the others concolorous
with head; mandibles tipped with black; antennae yellow; setae yellow
to colorless.
Body, first cervical segment narrow, with a black anterior margin
and dull yellow color with a slight Suggestion of a whitish mid-dorsal
stripe. Remaining body Segments are dull yellow with a slight tinge of
green. A narrow whitish mid-dorsal stripe, previously mentioned, runs
from the first thoracic segment to about the eighth or ninth Segments.
The entire dorsal half of the body is heavily speckled with irregulär
black dots as far down as the upper edge of the spiracles. The latter
have pale dull yellow centers and narrow black rims. Inferior to the
spiracles, the venter is a uniform pale yellow. Legs, concolorous with
body, with brown tips; prolegs, concolorous with venter. Crochets,
uniordinal, red-brown.
Pupa
Figures ID and E
Average length, 17 mm; width through center 6 mm. It pupates
under soil, incased in a fragile cocoon measuring 23 by 1 3 mm. Surface
texture, finely granulär. Color, brown, except for black eyes, dark brown
head and nearly black cremaster. Spiracles, glistening, also nearly black.
Cremaster terminates with a row of almost microscopic spinules.
John Adams Comstock and Christopher Henne, 1373 Crest Road,
Del Mar, California.
Accepted for publication December 10, 1968.
Bull. So. Calif. Acad. Sei. 68(1): 57-58, 1969
THE REPOSITORY OF THE T. W. COOK ANT TYPES
(Hymenoptera: Formicidae)
In his book, The Ants of California, T. W. Cook (1953) described four
new ant taxa, retaining the type specimens of all four forms in his per-
sonal collection. Following his death in 1962, his collection, including
the type material, was tumed over to The Oakland Museum, Oakland,
California by his widow. I examined the collection in 1966 and segre-
gated the types since they were not clearly marked.
Through the courtesy of the officials of The Oakland Museum, and
particularly of Dr. C. D. MacNeill, these types have now been placed
on permanent deposit in the Los Angeles County Museum of Natural
History. For the benefit of future workers, the following commentary
is offered regarding each of Cook’s forms.
The type data given below are taken from the labels on each pin
and are here transcribed as they appear on the labels. The data from
individual labels are separated by a slash mark and my comments are
enclosed in brackets. Inasmuch as holotype labels were not attached
by Cook, I have affixed to each pin such a label, except for Lasius
helveolus.
Proceratium californicum Cook, 1953: 45-46. Figure, p. 46. Type
data: “Glenwood, Cal., 27 May 1908.” This species has been discussed
in detail by Snelling (1967).
Pogonomyrmex barbatus spadix Cook, 1953: 98-99. Figure, p. 98
(labeled “ Pogonomyrmex spadix T. W. Cook”). Type data: “25 mi. E.
of Deming, N. M., June 17, 1942, H. A. Scullen, coli ./Pogonomyrmex
barbatus (F. Smith) var., det. M. R. Smith/P. spadix [pink]/DRAWN
spadix [white].”
The specimen is a worker, pinned through the anterior part of the
promesonotum, with the head secured in position by a large drop of
glue. Despite Cook’s Statement that the ant is unusually large and dark,
there is nothing at all to distinguish it from P. barbatus , and it must be
considered a synonym of P. barbatus as indicated by Cole (1968).
Pogonomyrmex californicus nitratus Cook, 1953: 99-100. Figure,
p. 99 (labeled Pogonomyrmex nitratus T. W. Cook). Type data: “Doug-
las, Ariz., 12-7-32, L. C. Murphree [large, in pencil] /Pogonomyrmex
californicus subsp. nov.? [in pencil] , NITRATUS T. W. Cook [in ink]
/DRAWN [in pencil].”
This worker specimen is point-mounted, in fair preservation, lacking
only the right foreleg beyond the coxa, the left foreleg beyond the
femur, and the left hind leg beyond the femur. This name may be safely
57
58 Bulletin So. Calif. Academy of Sciences
considered a synonym of P. californicus Emery, as stated by Cole
(1968).
Lasius (Chthonolasius) helveolus Cook, 1953: 327. Figure, p. 326
(labeled Lasius helvus, n. s.). Type data: “ Lasius helveolus T. W. Cook /
Lake Tahoe, VII. 15.50, coli. T. W. Cook/cotypes [a small red dot in
lower right comer]
Two specimens, both workers, are pointed on one pin; the upper
specimen lacks the head. A third specimen, of the original three, is in
the collection of the Museum of Comparative Zoology. This ant was
correctly assigned as a synonym of L. ( Chthonolasius) flavus (Fabricius)
by Wilson (1955).
Although Wilson stated that the Cook collection included the holo-
type and one paratype, this is not strictly true. Cook did not designate
a holotype at the time he described this ant, nor has there been any sub-
sequent selection of a lectotype. Since all three of the original specimens
are accounted for, and since the two contained in the Cook collection
are clearly marked as cotypes, Wilson’s use of “holotype” and “para-
type” is incorrect. Since Cook’s name is an obvious synonym I see
nothing to be gained by selecting a lectotype.
Literature Qted
Cole, A. C., 1968. Pogonomyrmex harvester ants. A study of the genus in North
America. Uni v. Tenn. Press, x + 222 pp.
Cook, T. W., 1953. The ants of California. Palo Alto: Pacific Books, 462 p.
Snelling. R. R., 1967. Studies on California ants. 3. The taxonomic Status of
Proceratium californicum Cook. Los Angeles Co. Mus., Contrib. Sei., No.
124, 10 p.
Wilson, E. O., 1955. A monographic revision of the ant genus Lasius. Bull. Mus.
Comp. Zool., 113: 1-202.
Roy R. Snelling, Los Angeles County Museum of Natural History,
Los Angeles, California 90007.
Accepted for publication November 1 5, 1 968
Southern California
Academy of Sciences
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