LOAN COPY ONLY TAMU-H-95-001 C3

Handbook of Shrimp Diseases

Aquaculture

S.K. Johnson
Department of Wildlife and Fisheries Sciences
Texas A&M University

90-601 (rev)
Introduction 2
Shrimp Species 2
Shrimp Anatomy 2
Obvious Manifestations of Shrimp Disease 3
Damaged Shells , 3
Inflammation and Melanization 3
Emaciation and Nutritional Deficiency 4
Muscle Necrosis 5
Tumors and Other Tissue Problems 5
Surface Fouling 6
Cramped Shrimp 6
Unusual Behavior 6
Developmental Problems 6
Growth Problems 7
Color Anomalies 7
Microbes 8
Viruses 8
Baceteria and Rickettsia 10
Fungus 12
Protozoa 12
Haplospora 13
Gregarina 15
Body Invaders 16
Surface Infestations 16
Worms 18
Trematodes 18
Cestodes 18
Nematodes 18
Environment 20

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Handbook of Shrimp Diseases
S.K. Johnson
Extension Fish Disease Specialist

This handbook is designed as an information source and tail end (abdomen). The parts listed below are apparent upon
field guide for shrimp culturists, commercial fishermen, and outside examination (Fig. 1).
others interested in diseases or abnormal conditions of shrimp.
1. Cephalothorax
It describes and illustrates common maladies, parasites and
2. Abdomen
commensals of commercially important marine shrimp. De
3. Antennules
scriptions include information on the life cycles and general
4. Antenna
biological characteristics of disease-producing organisms that
5. Antennal scale
spend all or part of their life cycles with shrimp.
6. Rostrum (horn)
Disease is one of the several causes of mortality in shrimp
7. Eye
stocks. Death from old age is the potential fate of all shrimp,
8. Mouthparts (several
but the toll taken by predation (man being one of the major appendages for holding
predators), starvation, infestation, infection and adverse envi and tearing food)
ronmental conditions is much more important. 9. Carapace (covering of
Although estimates of the importance of disease in natural cephalothorax)
populations are generally unreliable, the influence of disease, 10. Walking legs (pereio-
like predation and starvation, is accepted as important in lower pods)
ing numbers of natural stocks whenever they grow to excess. 11. Abdominal segment
Disease problems are considered very important to success 12. Swimmerets (pleopods)
ful production in shrimp aquaculture. Because high-density, 13. Sixth abdominal seg
confined rearing is unnatural and may produce stress, some ment
shrimp-associated organisms occasionally become prominent 14. Telson
factors in disease. Special measures are required to offset their 15. Uropod Fig. 1. External anatomy of shrimp.
detrimental effects. 16. Gills (Numbers conform to list.)
Disease may be caused by living agents or other influences
Inside structures include (Fig. 2)
of the general environment. Examples of influences in the
general environment that cause disease are lack of oxygen, 1. Esophagus
poisons, low temperatures and salinity extremes. This guide 2. Stomach

concentrates on the living agents and on visual presentation of 3. Hemocoel (blood space)
the structure and effects of such agents. 4. Digestive gland (hepato-
pancreas)
5. Heart
Shrimp Species
6. Intestine
There are many shrimp species distributed world-wide. 7. Abdominal muscles
Important shrimp of the Gulf of Mexico catch are the brown The "skin" or hypodermis
shrimp, Penaeus aztecus\ the white shrimp, Penaeus setiferus; of a shrimp lies just beneath
and the pink shrimp; Penaeus duorarum. the cuticle. It is functional in
Two exotic shrimp have gained importance in Gulf Coast secreting the new exoskeleton
aquaculture operations. These are the Pacific white (white leg) that develops to replace the
shrimp, Penaeus vannamei, and the Pacific blue shrimp, old at shedding. Shedding of
Penaeus stylirostris. These two species are used likewise Fig. 2. Internal anatomy of
the cuticle (also known as shrimp. (Numbers conform to
throughout the Americas on both east and west coasts. molting or ecdysis) occurs at list.) Jagged line represents
In Asia, the Pacific, and to some extent the Mediterranean, intervals during a shrimp's cutaway of cuticle to expose
the following species are used: Penaeus monodon, Penaeus life and allows for change in internal organs.
merguiensis, Penaeus chinensis, Penaeus japonicus, Penaeus developmental stage and ex
semisulcatus, Penaeus indicus, Penaeus penicillatus and pansion in size.
Metapenaeusensis. Penaeus monodon, the giant tiger (or black The reproductive organs of adults are particularly notice
tiger) shrimp is the world leader in aquaculture. able. When ripe, the ovaries of females may be seen through
the cuticle to begin in the cephalothorax and extend dorsally
Shrimp Anatomy into the abdomen. Spermatophores, a pair of oval structures
A shrimp is covered with a protective cuticle (exoskeleton, containing the sperm in adult males, are also visible through
shell) and has jointed appendages. Most organs are located in the cuticle when viewed from the underside near the juncture
the head end (cephalothorax) with muscles concentrated in the of cephalothorax and abdomen. The principle nervous struc-
ture, the ventral nerve cord, is visible along the underside of
the body between the swimmerets.

Obvious Manifestations of Shrimp Disease

Damaged Shells
Shrimp cuticle is easily damaged in aquaculture situations
when hard structures are impacted or rubbed. (Fig. 3). Blood
runs openly (outside of vessels) under the shell of shrimps, out
through appendages and into tiny fringe parts. When injury
occurs to the shell, the blood quickly clots and protectsdeeper
parts (Fig. 4).
Shell damage may also be inflicted by the pinching or biting
of other shrimpin crowded conditions. Parts of appendages
such as antennae may be missing. Cannibalism has an impor
tant influence on survival in some phases of shrimp culture
where stronger individuals devour weak ones (Fig. 5).
Shells may also be damaged because they become infected. Fig. 3. Eyes of shrimp are normally black, but rubbing of a tank wall
has caused this eye to appear whitish because of a prominent lesion.
A protective outer layer is part of the cuticle. If underlying
portions arc exposed opportunistic microbes will invade the
shell and use it as a food base or portal for entry into deeper
tissue. Larger marks darken and become obvious (Fig. 6).

Inflammation and Melanization

Darkening of shell and deeper tissues is a frequent occur
rence with shrimp and other crustaceans. In the usual case,
blood cells gradually congregate in particular tissue areas (in
flammation) where damage has occurred and this is followed
by pigment (melanin) deposition. An infective agent, injury or
a toxin may cause damage and stimulate the process (Fig. 7).
Gills arc particularly prone to darkening due to their fragile
nature and their function as a collecting site for elimination of
the body's waste products (Fig. 8). Gills readily darken upon
exposure to toxic metals or chemicals and as a result of infec
tion by certain fungi (Fusarium sp.).
Less common but important are dark blotches that sometime
occur within the tails of pond shrimp. This manifestation of Fig. 4. Microscopic view of a lesion on a uropod (tail part). Note crease
necrosis (breakdown and death) of muscle portions followed by from bend in part and loss of fringe setae.
melanization degrades the product's market potential. It is
possible that this condition results from deep microbial inva
sions that run through spaces between muscle bundles but its
actual causes remain unknown (Fig. 9).

Fig. 6. Tail ends of two shrimp. The lower shrimp shows typical darken
ing of cuticle that involves microbial action. The darkening itself is
considered a host response. The telsons of the upper shrimp are
Fig. 5. Cannibalism usually begins as other shrimp devour the append opaque because of dead inner tissue. Successful entry and tissue
ages. destruction by bacteria was accomplished only in those parts.
 Emaciation and Nutritional Deficiency
Unfed shrimp lose their normal full and robust appearance
and exhibit emaciation. The shell becomes thin and flexible as
it covers underlying tissue such as tail meat that becomes
greatly resorbed for lack of nutrients. Molting is curtailed and
shell and gills may darken in time (Fig. 10). Emaciation may
also follow limited feeding behavior during chronic disease
conditions or an exposure to unfavorable environmental condi
tions. Empty intestines arc easily observed through transparent
cuticle and flesh.
Prepared diets deficient in necessary constituents may pre
dispose or cause disease. Vitamin C deficiency, for example,
will initiate darkening of gills or certain tissues associated with
the cuticle and eventually result in deaths.
Fig. 7. A shrimp photographed (above) near time of back injury and
(below) hours later. Injury by a toxin or disease agent will usually trig
ger a similar response of inflammation and melanization.

k ,

J
f .

Fig. 9. Areas of melanized necrotic tissue in tail musculature.

Fig. 10. Emaciated shrimp. Gills and body fringes have become obviously
darkened and the soft tail is covered with a thin and fragile cuticle.

Fig. 8. Microscopic view of damaged and melanized gill tips.

Fig. 11. Lipoid (fat) spheres in microscopic view of digestive gland tubule.
 Digestive glands sometimes will become reduced in size.
Among other things, this is an indication of poor nutrition.
Well-fed shrimp will have an abundance of fat globules within
storage cells of the digestive gland tubules that provide bulk to
the gland (Figs. 11 and 12).

Muscle Necrosis
Opaque muscles are characteristic of this condition. When
shrimp areexposed to stressful conditions, such as low oxygen
or crowding, the muscles lose their normal transparency and
become blotched with whitish areas throughout. This may
progress until the entire tail area takes on a whitish appearance
(Fig. 13).
If shrimp arc withdrawn from the adverse environment
before prolonged exposure, they may return to normal. Ex
tremely affected shrimp do not recover, however, and die Fig. 12. Pond-raised shrimp with full, normal and reduced, abnormal
within a few minutes (Fig. 14). In moderately affected shrimp, digestive gland. Arrow points to abnormal gland.
only parts of the body return to normal; other parts, typically
the last segments of the tail are unable to recover and are prone
to bacterial infection (Fig. 15). These shrimp die within one or
two days (Fig. 5). Shrimp muscles with this condition are
known to undergo necrosis (death or decay of tissue).
Tumors and Other Tissue Problems
Conspicuous body swellings or enlargements of tissues have
been reported in shrimp. In most cases, affected individuals
werecaptured from polluted waters. Occurrenceof shrimp with
evident tumors is rare in commercial catches. Miscellaneous
irritations experienced by captive shrimp in tanksystems will
sometimes result in focal areas of tissue overgrowth (Fig. 16).
A particularly vulnerable tissue of captive juvenile and adult
shrimp is found on the inner surface of the portion of carapace Fig. 13. Shrimp with necrotic muscle tissue following exposure to
stressful environment. Affected tissue at arrow.
that covers the gills. When microbes invade this tissue, it and
the adjacent outer shell may completely disintegrate exposing
the gills. In other cases a partial loss of the tissue distally may
result in an outward flaring of the exposed cuticle. A hemo-
lymphoma or fluid-filled blister also forms sometime in this

Fig. 14. Shrimp with advanced muscle necrosis (arrow) shown beside
normal shrimp.

Fig. 15. Damage to abdomen of a shrimp as a result of Vibrio infection. Fig. 16. Tumorous growth on an adult shrimp from a tank system.
Fig. 17. Blister condition. Insetshows blister removed. The blister will
darken upon death of shrimp degrading marketability of heads-on
product.

portion of the carapace in pond shrimp (Fig. 17). Primary

Fig. 18. Darkening of male reproductive tract of Penaeus stylirostris. A.
causes of these manifestations are not understood. Normal tract. B. Initial darkening. Darkening will advance until sper-
A degeneration of male reproductive tracts occasionally matophores and testes become affected. (Photos courtesy of George
occurs in captive adults of certain penaeid species. A swelling Chamberlain.)
and darkening of the tubule leading from the testes to the spcr-
matophorc is readily apparent when viewed through the trans
lucent body (Fig. 18).

Surface Fouling
The surfacesof shrimps arc prone to an accumulation of vari
ous fouling organisms. Heavy infestations can interfere with mo
bility or breathing and influence marketability (Fig. 19).

Cramped Shrimp
This is a condition described for shrimp kept in a variety of
culture situations. The tail is drawn under the body and be
comes rigid to the point that it cannot be straightened (Fig. 20).
The cause of cramping is unknown, but some research points to Fig. 19. Algal overgrowth on shrimp exposed to abundant light. (Photo
courtesy of Steve Robertson.)
mineral imbalance.

Unusual Behavior Developmental Problems

Diseased shrimps often display listless behavior and cease Deformities are quite prevalent in some populations. They
to feed. In the case of water quality extremes such as low oxy arise from complex interactions that involve environment, diet
gen, shrimp may surface and congregate along shores where and gene expression. Bodies may be twisted or appendages
they become vulnerable to bird predation. Cold water may misshaped or missing. Deformities arc less prevalent in wild-
cause shrimp to burrow and an environmental stimulation such caught larvae than hatchery populations probably because wild
as low oxygen, thermal change or sudden exposures to unusual shrimp have more opportunity for natural selection and expo
chemicals may initiate widespread molting. sure to normal developmental conditions (Fig. 21).

Fig.20. Cramped shrimp condition. Full flexure (A). Flexure maintained when pressure applied (B).
 Molt arrest occurs in affected animals of some populations.
Animals begin, but are unable to complete the molting process.
In some cases, there is abnormal adherence to underlying skin,
but most animals appear to lack the necessary stamina. Nutri
tional inadequacies and water quality factors have been identi
fied as causes.

Growth Problems
Growth problems become obvious inaquaculture stocks. A
harvested population may show a larger percentage ofranting
than expected. Some research has connected viral disease with
ranting in pond stocks and it is generally held that variable
growth may result from disease agents, genetic makeup and
environmental influences.
For unknown reasons, the shell orcuticle may become frag
ile in members of captive shrimp stocks.
Shells are normally soft for a couple of days after molting, Fig. 21. Deformed larval shrimp. Arrow points to deformed appendage.
(Photo courtesy of George Chamberlain.)
but shells of those suffering from soft-shell condition remain
both soft and thin and have a tendency to crack under the
slightest pressure. Some evidence of cause suggests pesticide
toxicity, starvation (mentioned above) or mineral imbalance. to the cuticle or underlying skin. A genetic cause is suspected.
Transformation to blue coloration from a natural brown is
Color Anomalies
known for some captive crustaceans and has been linked to
Shrimp of unusual color arc occasionally found among wild nutrition. Pond-cultured, giant tiger shrimp sometime develop a
and farm stocks. Thestriking coloration, which may begold, condition wheredigestive gland degeneration contributes to a
blue or pink, appears throughout the tissue and is notconfined reddish coloration.

Microbes

Microbes are minute, living organisms, especially vi that are capable of developing into a new individual.
ruses, bacteria, rickcttsia and fungi. Sometimes protozoa arc Bacteria are one-celled organisms that can be seen only
considered microbes. with a microscope. Compared to protozoans, they arc of less
Protozoa are microscopic, usually one-celled,animals complex organization and normally less than 1/5,000 inch
that belong to the lowest division of the animal kingdom. (1/2000 cm) in size.
Normally, they are many times larger than bacteria. The Rickettsia are microbes with similarity to both viruses
typical protozoa reproduce by simple or multiple division or and bacteria and have a size that is normally somewhat in-
by budding. The more complex protozoa alternate between between. Most think of them as small bacteria.
hosts and produce cells with multiple division stages called Viruses arc ultramicroscopic, infective agents capable of
spores. multiplying in connection with living cells. Normally, vi
Fungi associated with shrimp are microscopic plants that ruses are many times smaller than bacteria but may be made
develop interconnecting tubular structures. They reproduce clearly visible at high magnification provided by an electron
by forming small cells known as spores or fruiting bodies microscope.
 Microbes

Viruses
Our knowledge of the diversity of shrimp viruses continues to
grow. Viruses of shrimp have been assigned explicitly or tenta
tively to six or seven categories. Several shrimp viruses are recog
nized to have special economic consequence in aquaculture:

Baculoviruses
Baculovirus penaei a virus common to Gulf of Mexico
shrimp. It damages tissue by entering a cell nucleus and subse
quently destroys the cell as it develops (Fig. 23). An occlusion
is formed (Fig. 24). This virus has become a constant problem
for many shrimp hatcheries where it damages the young larval
animals. Occlusions of the same or closely related viruses are
seen in Pacific and Atlantic Oceans of the Americas. At least
ten shrimp species arc known to show disease manifestations in
aquaculture settings.
Monodon-typc baculovirus one that forms spherical
Fig. 24. Occlusion bodies of Baculovirus penaei. These bodies, visible
occlusions (Fig. 25) and whose effects arc seen mostly in the to low power of a light microscope, are characteristic of this virus. The
culture of the giant tiger prawn, Penaeus monodon. Damage of occlusions and those of other baculoviruses are found mainly in the
less importance has been seen in Penaeus japonicus, Penaeus digestive gland and digestive tract.
merguiensis and Penaeus plebejus.
Midgut gland necrosis virus a naked baculovirus harmful
to the Kuruma prawn, Penaeus japonicus, in Japan.

Fig. 25. Monodon baculovirus in a tissue squash showing groups of

spherical occlusions. Light microscopy.

Solubility in Gut
Parvoviruses
Infectious hypodcrmal and hematopoietic necrosis virus
Ingestion ot Contaminated Food a virus affecting several commercially important shrimp and,
particularly, the Pacific blue shrimp, Penaeus stylirostris.
Infection of Host
Hepatopancreatic parvo-Iikc virus a virus causing disease
Fig. 23. Baculovirus life cycle. Transmission of the virus is thought to in several Asian shrimp. Transmission to Penaeus vannamei
be initiated as a susceptible shrimp ingests a viral occlusion. Virus
initially enters cell cytoplasm either by viroplexis (cell engulfs particle did not result in disease to that species.
with surrounding fluid) or by fusion where viral and cell membranes
Nodavirus
fuse and viral core passes into cell. Secondary infection occurs as
extracellular virus continues to infect. (Redrawn by Summers and Taura virus a virus causing obvious damage to various
Smith, 1987. Used with permission of author and Texas Agricultural tissues and in the acute phase, to the hypodermis and subse
Experiment Station, The Texas A&M University System.)
quently the cuticle of Penaeus vannamei (Fig. 26). It is an
important problem for both production and marketing. During
the 1995 growing season, this virus caused large losses to
aquaculture stocks in Texas. Damage was great in Central and
South America beginning in 1992.
Other viruses
Yellow head virus a virus causing serious disease of the
giant tiger prawn, Penaeus monodon. Large losses have been
experienced in Asian aquaculture units. Gills and digestive
glands of infected shrimp arc pale yellow.
White spot diseases viruses of similar size and structure
have been shown to cause a similar manifestation and heavy
losses to Penaeus japonicus, Penaeus monodon and Penaeus
penicillatus in Taiwan and Japan. Advanced infections show
development of obvious white spots on the inside of the cuticle
(Fig. 27).
Several other viruses with relatively little known importance
arc considered as members of the rcoviruses, rhabdoviruscs,
togaviruscs.

Fig. 26. Advanced stage of infection with Taura virus showing damage
to cuticle. Smaller shrimp with acute infection do not show such dam
age but do show reddish telson and uropods.
Fig. 27. Asian shrimp showing signs of white spot disease. (Photo
courtesy of R. Rama Krishna.)

Viruses

Viruses cause disease as they replicate within a host cell and within. Other "naked" baculoviruses do not show formation of
thereby cause destruction or improper cell function. A virus is occlusions.
essentially a particle containing a core of nucleic acids, DNA
or RNA. Once inside a proper host cell, the viral nucleic acid
interacts with that of a normal cell to cause reproduction of the
virus. The ability to parasitize and cause damage may be lim
ited to a single species or closely related group of hosts, a host
tissue and usually the place within a cell in which damage
takes place.
The cause and effect for all shrimp virus disease needs care
ful attention. Some viruses cause disease only after exposure to
unusual environmental conditions. Also, impressions about
virus identity arc often based on results of routine examinations
that give presumptive results. Certainly viruses cause important A
disease in particular circumstances but key understandings of
most shrimp viruses are largely unknown: longevity within
systems, source of infection, method of transmission, normal B
and unusual carriers, and potential to cause damage.
Our ability to detect shrimp viruses is ahead of our ability to
evaluate their importance or to implement controls. For viral
identification, scientists have employed the recent technology
that detects characteristic nucleic acids. This is augmented by
careful microscopical study of tissues to detect characteristic
damage to cells. Use of electron microscopy to determine size Fig. 22. Structure of viruses reported from shrimps. A. Baculoviridae.
and shape of virus particles has also been helpful (Fig. 22). Size range is about 250 to 400 nanometers in length. B. Basic struc
A peculiar feature of some baculoviruses of shrimp and ture of most of the other shrimp viruses: Parvo-like viruses20 to 24
nm in diameter containing DNA; Reo-like viruses55 to 70 nm diam
other invertebrate animals is to occurrence of the occlusion
eter, RNA; nodavirus30 nm diameter, RNA; toga-like virus 30 diam
bodies within infected cells. These are relatively large masses eter, RNA, enveloped. Rhabdoviruses are elongated like baculoviruses
of consistent shape that contain virus particles embedded but a blunt end provides bullet-shapes150 to 250 nm, RNA.
 m

Fig. 28. View with light microscope of a tissue squash of infected di Fig. 29. Transverse section of digestive gland tubules showing pro
gestive gland. Note dark necrotized tissue of tubules (arrows). gression of granuloma formation. Normal tubules are to the left (N) and
affected tubules are to the right (G.).

Bacteria and Rickettsia larger animals, infection becomes obvious in the digestive
Bacterial infections of shrimp have been observed for many gland after harmful bacteria gain entry to it, presumably via
years. Scientists have noticed that bacterial infection usually connections to the gut.
occurs when shrimp arc weakened. Otherwise normal shrimp Digestive gland tissues are organized as numerous tubular
also may become infected if conditions favor presence and structures that ultimately feed into the digestive tract. Pond-
abundance of a particularly harmful bacterium. reared shrimp occasionally die in large numbers because of
Shrimp body fluids are most often infected by the bacterial diseased digestive glands. The specialized cells that line the
group named Vibrio. Infected shrimp show discoloration of the inside of the tubules arc particularly fragile and arc easily in
body tissues in some instances, but not in others. The clotting fected. Tubules progressively die and darken (Figs. 28 and 29).
function of the blood, critical in wound repair, is slowed or lost This kind of disease manifestation is seen in recent reports of
during some infections. Members of one group of Vibrio have rickettsial infection. Cells of the digestive gland tubules arc
the characteristic of luminescence giving heavily infected ani severely damaged as rickcttsiac invade and develop therein
mals a "glow-in-thc-dark" appearance. (Figs. 30 and 31).
Bacteria also invade the digestive tract. A typical infection If infected by bacteria capable of using shell for nutrition,
in larval animals is seen throughout the digestive system. In the exoskeleton will demonstrate erosive and blackened areas

**

...-V --<:

i. .

>'*'* 4l5
-

Fig. 30. Histological cross section of a digestive gland tubule. Rickett

sial microcolonies are shown at arrows. Rickettsiae will exhibit constant Fig. 31. Electron microscope view of tissue infected with rickettsia
brownian motion and color red with Giemsa stain, but electron micros organism (arrow). (Photo circa 1987 from Penaeus vannamei on Texas
copy is needed for definite diagnosis. (Specimen courtesy of J. Brock) coast.)
 (Fig. 6). These bacteria typically attack edges or tips of exosk
eleton parts, but if break occurs in the exoskeleton the bacteria
are quick to enter and cause damage.
Filamentous bacteria are commonly found attached to the
cuticle, particularly fringe areas beset with setae (Fig. 32).
When infestation is heavy, filamentous bacteria may also be
present in large quantity on the gill filaments. Smaller, less
obvious bacteria also settle on cuticular surfaces but arc not
considered as threatening as the filamentous type.

Fig. 32. Microscopic view of filamentous bacteria on a shrimp pleopod.

Microbial Disease and Digestive Glands

Digestive glands are routinely searched by pathologists

for signs of disease. This is done after chemical preservation,
microsection, slide-mounting and staining the tissue. Trans
verse sections of the tubules are then examined with a light
microscope. General damage is seen when bacteria such as
Vibrio species invade tubules. Rickettsiae, viruses,
microsporans and haplosporans are more selective. They
invade cells and progressively cause damage from within.
For comparative purposes, a drawing of a normal tubule is
compared with a tubule showing a variety of typical manifes
tations (Fig. 33).

Fig. 33. A. Drawing of transverse section of digestive gland tubule

with bold lines that separate several types of disease conditions. 1.
Haplosporan parasite (microsporans similar but may show fully devel
oped spores, see Figs. 42 and 43). 2. Rickettsiae. 3. Virus infection
with manifestation of inclusion in cytoplasm. 4. Virus infection with
inclusion in nucleus. 5. Virus with occlusions in swollen nucleus. As
cells are destroyed, more general lesions are formed from viruses.
Inclusions of viruses normally show distinctive shape and staining
features. Particular viruses will infect particular tissue types (not
always hepatopancreas tissue) and cell locations (nucleus or cyto
plasm) within preferred hosts. Cells enlarged by haplosporans and
rickettsiae may be initially distinguished by comparing larger internal
components of the pre-spore units of haplosporans with almost sub-
microscopic particles of microcolonies of rickettsiae. H, = early
haplosporan stage, H? - later stage; R - rickettsial microcolony; CI =
cytoplasmic inclusion of virus; Nl = nuclear inclusion of virus; SN =
swollen nucleus with occlusions within.
B. The normal tubule. Toward the digestive tract, secretory cells (1)
predominate and fibrous cells (3) become more numerous. Absorp
tive cells (2) contain varying amounts of vacuoles according to nutri
tional status.

10
Fungus
Several fungi are known as shrimp pathogens. Two groups
commonly infect larval shrimp, whereas another attacks the
juvenile or larger shrimp. The most common genera affecting
larval shrimp are Lagcnidium and Sirolpidium. The method of jSKba
infection requires a thin cuticle such as that characteristic of
larval shrimp (Figs. 34 and 35).
The most common genus affecting larger shrimp is
Fusarium. It is thought that entry into the shrimp is gained via
cracks or eroded areas of the cuticle. Fusarium may be identi
fied by the presence of canoe-shaped macroconidia that the
fungus produces. Macroconidia and examples of fungal infec
tions arc shown in Figures 36, 37 and 38. D
-Search for Host

Fig. 34. Transmission of Lagenidium. A. Fungus sends out discharge

tube from within shrimp body. B. Vesicle forms. C. Vesicle produces
motile spores that are released. D. Motile spores contact shrimp and
undergo encystment. E. Germ tube is sent into the body of the shrip
and fungus then spreads throughout.

Fig. 36. Canoe-shaped macroconidia of Fusarium. These structures

Fig. 35. Lagenidium infection in larval shrimp. Note extensive develop
bud off branches of the fungus and serve to transmit fungus to shrimp.
ment of branchings of fungus throughout the body. (Photo courtesy of
Dr. Don Lightner, University of Arizona.)

OX

Fig. 37. Shrimp photographed immediately after mole: Old appendage

(arrow) is not shed due to destruction of hypodermis by active fungal
infection. Fig. 38. Microscopic view of fungus at tip of antenna.

n
Protozoa
Protozoan parasites and commensals of shrimp will occur
on the inside or outside of the body. Those on the outside are
considered harmless unless present in massive or burdensome
numbers. Those on the inside can cause disease and arc repre
sentativeof several groups of protozoan parasites: Microspora,
Haplospora and Grcgarina. Members of these groups are
known or believed to require that some animal besides shrimp
be present in order to facilitate completion of their life cycles.
A few protozoa are known to invade weakened larval animals
directly and contribute to disease.
Microspora parasitize most major animal groups, notably
insects, fish and crustaceans. In shrimp, microsporan infections
are best known locally as cause of a condition known as "milk"
or "cotton" shrimp (Figs. 39, 40 and 41). Microsporans become
remarkably abundant in the infected shrimp and cause the
white appearance of muscle tissues. A typical catch of wild
shrimp will contain a few individuals with this condition.
These shrimp are usually discarded before processing. Depend Fig. 40. Two brown shrimp cut across tail.Shrimp with whitish flesh has
microsporan infections throughout muscle tissue.
ing on the type of microsporan, the site of infection will be
throughout the musculature of the shrimp or, in particular or
gans and tissues.
Microsporans are present in the affected shrimp in the form
of spores. Spores arc small cells that can develop into a new
individual. They are very minute and detection requires exami
nation with a microscope (Figs. 42 and 43).
Infected shrimp are noted to be agile and apparently feed as
normal shrimp. However, tissue damage occurs and no doubt
affects many life functions. No eggs have been found in "milk"
shrimp and it is suspected that all types of microsporan infec
tions can render shrimp incapable of reproduction (Fig. 44).
The life cycles of shrimp microsporans have not been satis
factorily worked out. However, examination of the cycles of
related species and miscellaneous facts contained in literature
Fig. 41. Grass shrimp with "milk" shrimp condition. The normal shrimp
indicate that the cycle presented in Figure 45 is representative in the figure is transparent.
of microsporans.

Fig. 42. Microscopic view of many spores of Ameson (-Nosema) sp.

The spores are free or unenveloped. Parasitic microsporans of com
mercially important shrimp with enclosing envelopes are assigned to
genera Pleistophora, Thelohania and Agmasoma. The latter two differ
from Pleistophora in that their membranes retain a constant spore
Fig. 39. Infected or "milk" shrimp (upper) in comparison to normal number of eight per envelope. Pleistophora sp. have more than eight
shrimp (lower). (Photo courtesy of Dr. R. Nickelson.) spores per envelope.

12
Haplospora
A member of the Haplospora, another spore forming proto
zoan group, was recently recognized in as important to shrimp
health when researchers found infected animals in an experi
mental population that had been imported into Cuba from the
Pacific Coast of Central America. The parasites invaded and
destroyed tissues of the digestive gland (Fig. 32). Such infec
tions arc not common in aquaculture.

Fig. 43. Microscopic view of many spores of Thelohania sp. Note

envelope (arrow).

Fig. 44. Agmasoma penaei in white shrimp. This parasite is always

located along the dorsal midline (arrows). Advanced infections can be
seen through the cuticle with the unaided eye.

Fig. 45. Life cycle of microsporan of

shrimp. A. Ingestion of spores by shrimp.
B. In gut of shrimp, the spore extrudes a
filament that penetrates gut wall and
deposits an infective unit. A cell engulfs
this unit. C. Infective unit enters the
nucleus of the cell, undergoes develop
ment and then divides to form schizonts.
D. Schizonts then divide and develop into
spores. E. By the time spores are
formed, they are located in a specific
tissue (muscle, tissues around intestine,
etc.). The spores are either discharged
from the shrimp while living or after
death, but the method of release and the
pathway taken is not known. F. Experi
ments designed to transmit infection by
feeding infected shrimp to uninfected
have been unsuccessful. It is assumed
particular events such as involvement of
another host may be required to com
plete passage from one shrimp to the
next. Successful transmission has been
reported when infected shrimp were fed
to fish (speckled trout) and fish fecal
material was then fed to shrimp.

13
 53 '

* - *
Fig. 46. Microscopic views of gregarines. A. and B. Nematopsis sp. tropohzoites. C. Nematopsis sp. gametocyst.
D. Trophozoites of Cephalolobus sp., a gregarine that attaches to the base of the terminal lappets of the shrimp
stomach rather than the intestinal wall (photo courtesy of Dr. C. Corkern). E. Trophozites of Paraophioidina sp., a
gregarine found recently in Pacific white shrimp larvae.
S:M'&i*. 1
Gregarina
Gregarine are protozoa that occur within the digestive tract
and tissues of various invertebrate animals. They occur in the
digestive tract of shrimp and arc observed most often in the
form of a trophozoite (Fig. 46) or occasionally a gametocyst
(Fig. 47). The life cycle involves other invertebrates such as
snails, clams or marine worms as diagrammed in Figure 48.
Minor damage to the host shrimp results from attachment of
the trophozoites to the lining of the intestine. Earlier study sug
gested that absorption of food or intestinal blockage by the proto
zoa is perhaps detrimental but that pathological damage was rela
tively unimportant. Recent study indicates that when trophozoites
Fig. 47. Microscopic view of ' Nematopsis species arc present in large numbers, damage to the
gametocyst of Nematopsis sp. gut lining occurs that may facilitate infection by bacteria.

Fig. 48. Life cycle of a gregarine

of shrimp. A. Shrimp ingests
spores with bottom debris. B.
Sporozoite emerges in the gut of
the shrimp. C. Sporozoite at
taches to the intestinal wall and
grows into a delicate trophozoite;
other trophozoites do not attach
to the wall but onto each other
and form unusual shapes (See
Fig. 47). D. The unusual forms
develop and attach to the end of
the intestine (rectum) to form
gametocysts. E. Gametocyst
undergoes multiple divisions to
produce "gymnospores" that are
set free with rupture of the
gametocyst. F. Gymnospores
are engulfed by cells at the
surface of the flesh of clams. G.
They develop to form spores in
the clam. H. Then the spores
(with sporozoite inside) are
liberated from the clam in mu
cous strings (slime).

14
 Zoothamnium is a frequent inhabitant of the gill surfaces of
shrimp, and in ponds with low oxygen content, heavily infested

*# shrimp can suffocate. Surface-settling protozoa occasionally

cause problems in shrimp hatcheries when larval shrimp be
come overburdened and arc unable to swim normally. As pro
tozoa continuously multiply in numbers, shrimp acquire an
increasing burden until shedding of the cuticle provides relief.
Members of one unique group of protozoa, the apostomc
* *
ciliatcs, have a resting stage that will settle on shrimp surfaces.
When the crustacean molts, the protozoan releases and com
pletes the life cycle within the shed cuticle before entering a
resting stage on a new crustacean (Fig. 51).

Other Surface Infestations

A variety of other organisms attach to shrimp surfaces.
Fig. 49. Weakened larval shrimp invaded by ciliated protozoans (arrows).
Their abundant presence on gills and limbs can interfere with
breathing and mobility. Small, single-cell plants called diatoms
Body Invaders are often found attached to larval shrimp in hatcheries. (Fig.
Several protozoa have been noted to invade a shrimp body 52). Shrimp from aquaculture facilities that are exposed to an
and feed on tissues as they wander throughout. Tentative iden unusual amount of sunlight often will have over-growths of
tifications name Piwauronema, Leptomonas, Paranophrys and algae of mixed variety (Fig. 19).
an amoeba. Adverse effects of these protozoa are not fully Occasionally, one will find barnacles, leeches and the colo
understood, but they are usually found associated with shrimp nial hydroid Obelia bicuspidata affixed to body surfaces (Fig.
that have become weakened or diseased (Fig. 49). 53). These organisms arc probably quite common in the vicin
ity of the shrimp and select surfaces of infrequently molting,
older shrimp as spots to take up residence. Insects will some
Surface Infestations
times lay eggs on shrimp (Fig. 54).
Ectocommensal Protozoa
Some members of the crustacean group called isopods are
parasitic on shrimp of commercial importance. Commercially
Several kinds of protozoa are regularly found on surfaces,
important shrimp of the Gulf of Mexico are apparently not
including gills, of shrimp. Apparently, shrimp surfaces are a parasitized. However, smaller shrimp of the family
favored place to live within the water environment. Common
Palaemonidae arc often seen infested along our coastline. Com
on the surfaces of shrimp are species of Zoothamnium,
mercial shrimp of the family Pcnaeidae are parasitized in Pa
Epistylis, Acineta, Eplwlota, and Lagenophry.s (Fig. 50A-E). cific areas (Fig. 55A and B).

A. Zoothamnium B. Epistylis C. Lagenophrys

Fig. 50. Microscopic views of common surface

dwelling protozoa.
.

i-rr
D. Acineta E. Ephelota

15
 Fig. 54. Shrimp with insect eggs attached to cuticular surface.

Fig. 51. Microscopic view of several apostome ciliates inside grass

shrimp molt. Proper identifiation of genus cannot be determined from
living animal. Staining with a technique called silver impregnation is
required.

Fig. 55A. Asian shrimp infested with parasitic isopods (gill cover flared
open to expose parasites).

Fig. 52. Microscopic view of diatoms (arrow) attached to gill surfaces of

larval shrimp.

Fig. 53. Grass shrimp with leech attached. Fig. 55B. Parasitic isopods removed from shrimp.

If,
 Worms
Worm parasites of shrimp are categorized as trematodes
(flukes), cestodes (tapeworms) and nematodes (roundworms).
Some species are more common than others and, as yet, none
have been known to cause widespread shrimp mortality.
Worms may be found in various parts of the body (Fig. 56).

Trematodes
Trematodes (flukes) are present in shrimp as immature
forms (metacercariae) encysted in various body tissues. Meta-
cercariae of trematodes of the families Opecoelidae,
Microphallidae and Echinostomatidae have been reported from
Fig. 56. Common sites of infesta commercial species of penaeid shrimp (Fig. 57). One species,
tion by worms. 1. Tapeworms; Opecoeloides fimbriatus, has been noted to be more common
usually associated with tissues
covering digestive gland. 2.
than others along our coast, and the hypothetical life cycle of
Roundworms; in and outside this species is illustrated in Figure 58.
organs in cephalothorax, but
also along outside of intestine. 3. Cestodes
Flukes; commonly encysted in
tissues adjacent to organ in
Tapeworms in shrimp are associated typically with the di
cephalothorax, but also in ab gestive gland. They are usually found imbedded in the gland,
dominal musculature and under or next to it, in the covering tissue. In shrimp, tapeworms are
cuticle.
present as immature forms (Fig. 59), while adult forms are
found in rays. Species of the genera Prochristianella,
Parachristianella and Renibulbus are common. Other tape
worms from wild shrimp include a relatively common pear-
shaped worm of the intestine and a less common worm of the
cyclophyllidean group. Tapeworms are most often encountered
in wild shrimp.
Differentiation between the tapeworm groups is made in
general body form and tentacular armature. A hypothetical life
cycle for Prochristianella penaei is presented in Figure 60.
Nematodes
Nematodes occur more commonly in wild shrimp than in
culturedshrimp. The degree of infection is probably related to
Fig. 57. Drawing of microscopic view of common flukes of shrimp the absenceof appropriatealternate hosts in culture systems.
(excysted). A. Microphallus sp. B. Opeocoeloides fimbriatus. Nematodes will occur within and around most body organs, as

Fig. 58. Hypothetical life cycle of a shrimpfluke, Opecoeloides fimbriatus. adult. E. Eggs laid by adult fluke pass out of fishwith wastes. Egg hatches
A. Infective stage orcercaria penetrates shrimp. B. Cercaria migrates to and an infective stage known as a miracidium is released. The miracidium
the appropriate tissue and encysts forming a stage calledmetacercaria. C. penetrates a snail and multiplies in number within sporocysts. F. Cercariae
Shrimp infected with metacercaria is eaten byfish (silver perch, red drum, develop within sporocysts. When fully developed, cercaria leaves the
sheepshead, several others). D.Shrimp is digested. This releases meta sporocyst and snail and swims in search of a shrimp. Ifcontact is made
cercaria. Metacercaria stage undergoes development until itformsan with a shimpwithin a short period, the cycleis completed.

17
 well as in the musculature. Nematodes of shrimp include
Spirocamallanus pereirai (Fig. 61), Leptolaimus sp. and
Ascaropsis sp. The most common nematode in Gulf shrimp is
Hysterothylacium reliquens (Fig. 61).
'i
?>J
It is the juvenile state of nematodes that infects shrimp with
the adult occurring in fish. An illustrated life cycle thought to
represent Hysterothylacium reliquens is depicted in Figure 62.

o "

* * 'JSfT fc
F^
V
i

B
Fig. 59. A. A drawing of the shrimp tapeworm, Prochristianella penaei Fig. 61. Drawing of microscopic view of head end of (A)
as it would appear in a microscopic view after removal from its cyst. B. Spirocamallanus pereirai and (B) Hysterothylacium sp. common round
Unnamed pear-shaped tapeworm larvae in gut. (Photo courtesy of Dr. worms found in penaeid shrimp.
C. Corkern.)

jjsyw"!
B
"f*C c i
rw/y T\
twl \Ev
^ N^^^X^\N=-:
jS^y|

Fig. 60. Hypothetical life cycle of the tapeworm, Prochristianella penaei Tapeworm develops into adult in gut (spiral valve) of ray and begins to
Kruse. A. Shrimp eats a copepod or other small crustacean infested release eggs. E. Eggs pass out of the fish with feces and are eaten by
with larval tapeworm. B. Tapeworm develops into advanced larval copepod. Eggs hatch and larval worm develops inside copepod.
stage in tissues of shrimp. C. Stingray ingests infested shrimp. D.

Fig. 62. Hypothetical life cycle of Hysterothylacium relinquens, a round larval stage in tissues ot snrimp. C. Toadfish ingests infested shrimp.
worm of shrimp. A. Shrimp eats a copepod or other small crustacean D. Roundworm devleops into adult in gut of fish with feces and are
infested with larval roundworm. B. Roundworm develops into advanced eaten by copepod.

18
 Environment

Environmental Extremes
Temperature, irradiation, gas saturation, hydrogen ion con
tent (pH), oxygen content and salinity all have appropriate
tolerable ranges for sustaining life of various shrimp species
and life stages. If these ranges arc exceeded or extremes com
bine for an interactive effect, shrimp will become diseased.
Besides the direct effect from these noninfectious agents, expo
sure may result in prcdisposal to effects of opportunistic infec
tive agents.
Gas bubbles will form in the blood of shrimp if exposed to
waters with large differences in gas saturations. If a large Fig. 63. Precipitant of iron salt on a shrimp's fringe hair (setule)
amount of bubbling occurs in the blood, death will result.
In the presence of acidic water, minerals will often precipi
tate on cuticular surfaces. Usually the precipitant is iron salt
(Fig. 63).

Toxicity
Poisoning can result from toxic substances absorbed from
the water or consumed food. Water may accumulate excessive
concentrations of ammonia, nitrite, hydrogen sulfide or carbon
dioxide, all of which can have a toxic effect on shrimp. Some
metals also may cause a toxic effect when present in excess.
Both presence and toxicity of these chemicals arc influenced
by the changeable environmental conditions. They may act
singularly or have combined effects.
Certain microbes and algae will excrete poisonous materi
als. Examples of algal release arc the occasional red tides that
occur along our coast. Aside from survival loss, affected ani
mals behave in a disoriented manner. Microbes such as bacteria
become concentrated in high density rearing systems. When
microbial species with potential for toxic release greatly in
crease therein, stocks may be damaged.
Pesticides can be harmful if they occur seasonally in surface
water supplies affected by agricultural practice. Because of
migrations into estuaries or near effluent disposal sites, wild
shrimp populations are more susceptible than cultured stocks to
the variety of pollutants released.
There arc reports of toxicity caused by the food shrimp
consume. Toxins from microbes are known to build up in feeds
stored in unfavorable conditions. Some food stuffs and live
larval food, such as brine shrimp, can contain pesticides. Per
haps more common arc undesirable effects of feeds that have
aged and become rancid.
Breakdown of lining tissues (necrosis) of the intestine have
been associated with consumption of certain algae. Because
cultured shrimp feed both on prepared feeds and bottom mate
rials, it is suspected that the occasional occurrence of detrimen
tal irritants and toxins contained within bottom surfaces could
cause tissue breakdown when such sediments are consumed.
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Bielsa, L.M., W.H. Murdich and R.F. Young,J.H. 1959.Morphology of the white bookofMariculture, Vol.I,Crustacean Aquac
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fishes andinvertebrates (south Florida) - pink Overstreet, R.M. 1973. Parasites of some
shrimp. U.S. Fish Wildl. Serv. FWS/OBS-82/ General Diseases penaeid shrimps with emphasis on reared hosts.
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27
Questions or comments should be addressed to the author
Dr. S.K. Johnson
Extension Fish Disease Specialist
Department of Wildlife and Fisheries Sciences
Nagle Hall
Texas A&M University
College Station, Texas 77843-2258