INTRODUCTION
Several of the common ocean fishes off the California coast
are heavily infested with the isopod Livoneca vulgaris. Little is
known about Livoneca except that it is a protandrous hermaphrodite;
while young, it is a male with functional testes and as it matures.
certain tissue in the body developes into ovaries, while the testes
degenerate. It is distinquished morphologically from others of
the same genus, by the following characteristics: its eyes are sep-
arated by three times their width, the head terminates in a broad,
subtruncate lobe about twice the width of the eye, the telson is
twice as broad as it is long, the total body length is up to 32mm.
(Hatch,1947).
Most of the species of flatfish in Monterey Bay were infested
with Livoneca. This study is based on two of the flatfishes most
commonly caught, the Pacific sanddab, Citharichthys sordidus and
the starry flounder, Platichthys stellatus. These are both bottom
dwelling fish, which are born with one eye on each side of the head,
one of which migrates to the other side by the time they are  inch
in length. They are found from southern California to Arctic Alaska
(Miller, 1972). Their underside is white and smooth, their upper
side is rough and a brownish-green color.
The objectives of this investigation were to examine the diet
and the nature of attachment of Livoneca to its hosts.
METHODS AND MATERIALS
Flatfish were caught in Monterey Bay by pole-fishing and otter
trawling and kept alive in sea water tanks with constant running
sea water ataapproimately ,1%.
Isopods from dead
fish were kept in a separate aerated tank with constant flowing
sea water at 1400.
To study the position of the iso-
pod on the gill, infested fish were quick-frozen in liquid nitrogen.
The gill flap was then cut away, exposing the isopod in place in
the gill chamber.
The gut and diverticula were removed from livoneca vulgaris
by dissection after they had been rendered motionless by immersion
in magnesium chloride. This prevented rapid emptying of gut and
diverticula. Examination of gut and diverticula contents took
place under 10X and 100X power.
The presence of blood in the diverticula was determined by the
Wilkinson and Peters Benzidine Peroxidase Reaction. Benzidine di
Hydrochloride was first added to the diverticula, then a few frops
of glacial acetic acid, and then a few drops of hydrogen peroxide.
The immediate appearance of a dark blue color indicated the presence
of hemoglobin.
Blood pigments were extracted from the diverticula by adding
sodium carbonate to a sample, heating to boiling, and centrifuging
for fifteen minutes at 200 rpm. The supernatant was poured off
and a few drops of concentrated sulfuric acid was added to the
diverticula to further extract the pigments. Two mls. of water
was added and the contents centrifuged for another fifteen minutes
at 200 rpm. The supernatant was run on a Beckman DB-G spectro-
photometer to obtain wavelengths at which the hemochromogen absorbed.
The same extraction procedure was performed on samples of flat-
fish blood and spectra was taken to compare with the spectra of
the blood in the diverticula.
RESULT
Eighty two fish were examined and all but one were infested
with livoneca in both gills. Generally the size of the isopod in
the gill was proportional to the size of the fish, but there were
too many exceptions for this data to be statistically significant.
Usually one Livoneca per gill was found, but up to four isopods
were found in a gill. Since the two sides of the flatfish markedly
differ, isopods in the lower and upper gill were compared. The
dorsal side of the isopod matched the color of the side of the fish
it was on. No relationship was found between the number of isopods
or their sex and the side of the fish they occupied.
Female Livoneca were found with their heads in the anterior
part of the gill chamber, facing the anterior end of the host,
with the dorsal surface lying next to the outer side of the gill
chamber. Males were generally found with their heads facing the
anterior end of the fish, but their dorsal side could be facing
either the inner or outer side of the chamber. Young isopods, two
to five mm. in length were often found crawling on the outer gill
filaments.
Externally, parasitized fish couldn't be distinquished from
non-infected fish. Ovigerous females with large brood pouches
leave an indentation in the gill chamber where the brood pouch
rests. Both males and females have hooked pereiopods which they
use to attach to the host. These leave holes of about one mm.
diameter in the inner side of the gill. Sometimes the distal parts
of the gill filaments are destroyed by the isopod.
Females in the date-brooding stages contained no food in their
guts or diverticula. Diverticulas of males and females in the
early brooding stages contained blood, which was determined to be
hemoglobin by comparing adsorption peaks of blood pigments ex-
tracted from the blood of Citharichthys sordidus and the blood
in livoneca's diverticula. Mouthparts of Livoneca are very similar to those
in the family Aegidae, which are very frequently found with their
guts and thoracic segments full of blood. (Bowman,1960). Examina¬
tion of the gut revealed no other food preference; all guts were
empty except for minute fragments of what looked like bacteria.
Young Livoneca, up to three to four mm. would consume young dead
flatfish.
DISCUSSION
The gill chamber is a favorable place for Livoneca to live;
it is protected from predation unless its host is consumed, it is
provided with a good supply of oxygenated water, food particles,
and a convenient blood supply is available. It's characteristic
orientation could be keyed to catching small food particles that
pass through the gill. It would be advantageous for Livoneca not
to destroy this chamber by consuming it. Possibly the gill fila-
ments are destroyed to make more room for the growing isopod or as
a result of Livoneca raking them for food. Other damage to the fish
could be caused by the fact that large, ovigerous females occupy
most of the gill chamber, possibly obstructing the flow of water
through the gill, thus reducing oxygen uptake.
Results show that young isopods just hatched consume dead fish
until they are at least three to four mm. in length. However, males
larger than this and females will not eat the flesh, although they
may cling to it. Eventually, isopods deprived of a blood supply,
die. This implies that during their free-living stage, young
Livoneca can scavange for food while searching for a host. After
a certain life stage Livoneca must find a host or perish. Blood
sucking continues until the female enters the late brooding stage.
It then ceases, although there is no change or degeneration of mouth
parts. Since t he hosts blood provides a steady diet, it would
not be necessary for Livoneca to gorge itself. This might explain
the empty gut.
It was not determined why blood is stored in the diverticula
and never in the gut. Possibly there are enzymes present in the
diverticula to digest the blood. It is also a convenient place
to store blood and use it a little at a time.
Livoneca has biting and sucking mouthparts which enable it to
draw blood. It bores through the tissue until it reaches a blood
supply; this could account for the small holes in the fish found
under the mouthparts when the isopod is removed.
SUMMARY
A study of the attachment of Livoneca vulgaris to its hosts,
Citharichthys sordidus and Platichthys stellatus, showed that
their characteristic orientation is with the head in the anterior
part of the gill chamber, pointed toward the anterior of the host
and its dorsal surface lies against the outer flap of the chamber.
A parasitized gill chamber may be missing gill filaments or have
an indentation on the inner side as a result of the female's
swollen brood pouch. Livoneca's diet is fish blood which it stores
in its diverticula.
370

A

255
3900

—
D
aou
40mu
700
A


409
100.
100
KEY TO THE FIGURES
Figure 1. Livoneca vulgaris: A, maxilliped, ovigerous female; B,
second maxillae, ovigerous female; C, first maxillae.
ovigerous female; D, mandibles, ovigerous female
Figure 2. Spectra of hemochromogen: A, Hemochromogen extract from
blood of Citharichthys sordidus, peak at 409 mu.; B,
Hemochromogen extract from blood in diverticula of
Livoneca vulgaris, peak at 409 mu.
LITERATURE CITED
1. Bowman, Thomas E.. 1960. Description and notes on the Biology
of Lironeca Puhi, N. Sp. (Isopoda: Cymothoidae). Crustaceana.
1:83-91
2. Hatch, Melville, 1947. Chelifera and Isopoda of Washington
and Adjacent Regions. U. of W. Publications. 10:207,211
3. Light, Smith, Pitelka, Abbott, Weesner, 1964. Intertidal Inver-
tebrates of the Central California Coast. University of Calif-
fornia Press. Berkeley and Los Angleles. 387p.
4. Menzies R., Thomas Bowman and Franklin Alverson. 1955. Studiese
on the Biology of the Fish Parasite Livoneca convexa Richardson
(Crustacea, Isopoda, Cymothoidae). Wasman J. Biol.. 13. 277-294
5. Miller, Daniel J. and Robert N. Lead, 1972. Guide to the Coastal
Marine Fishes of California. Fish Bulletin 156. State of Calif-
ornia Department of Fish and Game. 80p.
6. Norman, J. R. 1934. A Systematic Monograph of the Flatfishes.
Oxford University Press. London. 435p.