TANV -8-89-001 C3

Handbook of Shrimp Diseases

.'"Etllhkl,'II; I',gI yBa Ir~ol 0l[rri~ory

Aquaculture

S.K. JohnsonDepartment of Wildlife and Fisheries Sciences

Texas A8 M University NATlONAL SEA CRANT DEPOSITORYPELL LiBHARY BUILUiNG

URi, NARP�'..";.~NSETT BAY CAMPUSNARRAGANSETT, R> 0288'

Q,es', ons or comme."'.s shou',d be addressed to the a.thor-Dr. S K. JohnsonExtens,ori Fish Disease Specia'. stDepartment of W IC'ife and Fisheries SciencesNagle ka!ITexas AB,M Univers,tyCOIIege StatiOr, TexaS 778'3-2258

TAMU-SG-904012 M September 1989

NA89AA-0-SG 139Afl-1

$2.00ArtdfSonaf copies available frorrr:Saa Grant Cotfaga ProgramTaxaa ASM University at GalvestonP.O. Box $675Galveston, Texas 77S53-1675

National Sea Grant DepositoryPell Library Building - GSOUniversity of Rhode Island

Narragansett, RI 02882-1197USA

LOAN GOOPY ONLy

Handbook ofShrimp Diseases

Contents

Introduction

Shrimp Species

Shrimp Anatomy .

Microbes ............VirusesRickettsiaBacteriaFungiProtozoa

Other Infestations

WorrnS .TrematodesCestodesNematodes

Miscellaneous Conditions

InjuryDevelopmental ProblemsColor AnomaliesDigestive Gland ManifestationsInflammation and MelanizationEmaciation and Nutritional DeficiencyCramped ShrimpMuscle NecrosisTumors and Other Tissue ProblemsEnvironmental Extremes

Toxicity

.... 1 5

Bibliography . 20

Publication of this handbook is a cooperative effort of the Texas A& M University Sea Grant College Program, the Texas A8 MDepartment of Wildlife and Fisheries Sciences and Texas Agricultural Extension Service, Publication is partially supponed byInstitutional Grant No. NA89AA-D-SGt 39 to the Texas A&M University Sea Grant College Program by the National Sea GrantProgram, National Ocearsc and Atmospheric Administration, Department of Commerce.

Handbook Of Shrimp Diseases

S. K JohnsonExtension Fish Disease Specialist

Shrimp Artatorny

6- 4

-- � 56 7

Shrimp Species

Fig. t . External anatomy at shrirrip 'Numh.rs ~nfc;m te li t,!

This handbook is designed as an informationsource and field guide for shrimp culturists, commer-cial fishermen, and others interested in paras:tes orabnormal conditions of shrimp. In addition to descrip-tions and illustrations of the common parasites andcomrnensals of commercial penaeid shrimp, thepublication includes information on the life cycles andgeneral biological characteristics of these disease-producing organisms that spend all or part of their lifecycles with shrimp, Several conditions of unknowncause are also described.

Disease is an important factor in reducing shrimpnumbers in natural populations. Natura! mortality ordeath from old age is the potential fate of all shrimp,but the toll taken by predation man being one of themajor predators!, starvation, infestation, infection andadverse environmental conditions is highly significant.Therefore, conditions described in the followingpages exert considerable influence on shrimp nurn-bers in natural stocks.

Disease problems are considered important tosuccessful production in shrimp aquaculture. Be-cause high-density, confined rearing is unnatural andproduces stress, some shrimp-associated organismswill become prominent. Special measures are re-quired to offset their detrimental effects.

Disease may be caused by living or non-livingagents, as well as by physical or chemical factors.Non-living causes include lack of oxygen, poisons,low temperatures, salinity extremes, etc, This guideconcentrates on the living agents and on visualpresentation of the structure and effects of suchagents,

There are many shrimp species distributed world-wide. ImpOrtant Shrimp Of the Gulf of Mexico catchare the brown shrimp, Penaeus azfecus: the whiteshrimp, Perraeus setiferus; and the pink shrimp,Penaerrs duorarum.

Two exotic shrimp have gained importance in GulfCoast aquaculture operations, These are the Pacificwhite shrimp, Pertaeus vannamet, and the Pacific

blue shrimp, Penaeus stytirostns. These two speciesare used likewise throughout the Americas on botheast and west coasts.

ln Asia, the Pacific, and to some exten'. the Medi-terranean, the following species are used: Penaeusmonodon, Penaeus merguiensis, Penaeus chinensts,Penaeusjaponicus, Penaeus semisuicatus andMetapenaeus ensis.

A shrimp is covered with a protective cutie'.e exoskeleton, shell! and has lointed appendages.Mast organs are located in the head end cephalotho-rax! with muscles concentrated in the tail end abdo-rnen!. The parts listed below are apparer t uponou tside e x am i nation Fig.1!:

Cephalathorax2. Abdomen3. Ante nnui es4. Antenna5, Antennal Scale6. Rostrum harn!7. Eye6. Mouthparts

severa! append-ages for holdingand tearing food!

9. Carapace covering of cephalothor-ax!

10. Walking Legs pere iopods!

11. Abdominal Seg-rnent

12. Swimrneretes

pleo pods!13. Sixth Abdominal

Segment14. Telson15. Uropod16, GiIs under

carapace!

Mlcrobes

Micrabes

Inside struc turesinclude Fig 2!:

Esophagus2. Stomach3. H emocoe! blood

space!4 Digestive Gland

hepatopancreas!5 HeartI . Intestine7, Abdominal

Muscles

The 'skin' Or hypo- Fig. 2 Internal anatomy ot shrimpdermis Of a Shrimp lieS Numhors conform to kst ! Jagged

line represenLs cut away ot culidelust beneath the io expose internal organs,cuticle. It is functional

in secreting the new exoskeleton that develops toreplace the old at shedding Shedding of the cuticle also known as molting or ecdysis! occurs at intervalsduring a shrimp's life and allows for expansion ofsize.

The reproductive organs ol adults are particularlynOtiCeable. When ripe, the OvarieS Of femaleS may beseen through the cuticle to begin in the cephalothoraxand extend dorsally into the abdomen.

VIRUSES

Viruses cause disease as they replicate within asuitable host cell type and thereby cause destructionor improper cell function. The damage varies accord-ing to host species and is usually confined to a singlespecies or closely related group of hosts.

Much knowledge is needed about shrimpviruses. Exposure to particular environmental condi-tions may predispose shrimp to disease by viruses

Microbes are minute, living organisms, especiallyviruses, bacteria, rickettsia and fungi. Sornetirnesprotozoa are considered rnicrobes,

Protozoa are microscopic, usually one-celled,animals that belong to the lowest division of theanimal kingdom. Normally, they are many timeslarger than bacteria.

Bacteria are one-celled organisms that can beseen only with a microscope. Compared to protozo-ans, they are of less complex organization andnormally less than 1 S,000 inch in size.

Viruses are ultramicroscopic, infective agentscapable of multiplying in connection with living cells.Normally, viruses are many times smaller thanbacteria but may be made clearly visibfe at high

that act only as opportunists. Cause and effect for allshrimp virus disease needs careful attention Anotherproblem worth considering is that known viruses maybe more diverse than currently recognized. This isbecause presuinptive identifications are often mademerely on the basis of size and shape. In the case ofsome detected viruses, are they rept cating in theshr rnp, or are they merely accumulated transientIOdgerso Little is known alSO Of how shrimp viruseSare transmitted from one shrimp to another.

Viruses of shrimp have been assigned explicitly ortentatively to four viral groups Fig. 3!.

Fig. 3 Structure ot viral groups reported fram shrimp. A. Bacutovtri-dae tour repiesentative in penaid shrimp ! Size range is about 300to 400 nanometers in ienglh B Basic structure of other knownshnrnp viruses tour! A Prcornavirudae-like virus is 20 nm indiamefer and contains DNA A Patvovtrtdae-tike virus is 22 to 24nm in diameter and contains RNA. Two Reovtridae like viruses are55 tO 61 and 60 to 70 nm in diameter and COntain ONA

magnification provided by an electroRickettsia are rnicrobes with sim

viruses and bacteria and have a sizesomewhat in-between,

Fungi associated with shrimp aplants that develop interconnectingtures. They reproduce by forming srnas spores or fruiting bodies that are capaoping into a new individual.

The microbial world is vast in number and kind andhas both beneficial and harmful rnernbers. Those thatlive in association with shrimp are normally eitherparasitic and capable of causing disease whenpresent in sufficient numbers, or commensal andcapable of living in association without causing injury.

Shrimp Viruses

~ ~grn' -g I

!~P us l Oil

i,r1'Idgui Cell

i euodir r;r vl'rpeKis

reSe pncar, ir feclipn a

cells end Te~,je~

Pr mary.nlect;rynji. ~

spiuo r1y ig ciil

Ingestia oi Contaminated epoc

Ireec.tiop ryt Hoyu

Bacutovirus

BaCtrtavfrVS penael � a vi'uS COmmOn tO Gulf OfMexico shrimp, This virus damages t ssue by enterir ga cell nucleus and subseq ently dos:roy ng tl e ce! asthe virus develops Pig d! I'. has becon;e a constantprOblern fOr many shrimp halcherieS where it darn-ages the young larval anirr als 8acutovirus penaerhas a characteristic that d stinguisnes it from some o'the other shrimp viruses, some of the small nfec',iveviruS partiCleS fOrm w:thin a Oy. amid-Shaped rnaSScalled an occlusion T! e occlusion is large enough tobe seen easily with a light microscope f ig. 5!.

Fig 4 BacutcrviruS ate Cyote TranSmiesion of the virus is thoughtto be initiated as a susceptible shrimp ingests a v ral occlusionVirus initially enters cell cyioplaSm erther by v:roplexiS cell engulfsparticle with surrounding tluid! or by fusion where viral and cellmembranes tuse and viral core passes into cell Secondaryinlectiori occurs as extracellular virus continues to intact tRedrawntrom Summers and Smith, 1987 Used with permission ot authorand Texas Agncultural Expenment Station, Texas A8 M UniversitySystem !

Fig 6. Monodon baculovirus in a tissue squash showing groups ofspherical occlusions.

MOnOdOn-tyPe bacuiov "US o"'e "a. '"-': S

sphe ca, occfus ons iF g 6; and w: ose e".ec.s areSeen mOS! y in the C' Iture of;ne AS ar g 'l': t ge'prawn, PertaeuS n. n dort A recent eo- l Ol tranS-mission to Penaeus;,!rina ie. did t~o'. ";c ve causa

tior of d,seasePlebelus oacuiovirus � A spi e ca occ'�s -n-

forrn ng virtis of tile Australian shr mo Perv<ir i.sptebetus

Midgu'. glano necrosis virus � - a vu us haritffu Eothe Kuyuma prawrl, PertaeUS iapeniCUS. ii Japan

Picornavirus

Irtfectious hypode~mal and her, atopoietic necros svirus � a virus affec'.ing several con.mercially tntpor-tar t shrimp and. particularly, lhe Pac:fic hft.e s rimp,Penaeus styttrostris

Parvovirus

Hepatopancreatic parvo-like virus a virus-causing disease in several Asian sh imp A recentlynoted transmission to Penaeus vanrtamei did no'.result in disease of that species.

Reovirus

Reo-like virus of Ku uma prawn � a newly recognized virus of Penaeus JapontCus.

Heo-like virus of Tiger prawn � a newly recog-nized virus of Penaeus monocton.

Fig 5 Ocdusion bodies of Racurotrrrus penaei These bodiesvisible to low power Of light rhicroscope, are CharaCteriStiC ot thiSvirus. The occlusrons and those of other baculoviruses are toundmainly in the digeslive gland and digesbve tract

R ICK ETTS! A!f cki''' a nlpc'. Ons i-aie ~e,-;n' v i;.-n~ i<~

Ir'ei'leo as a' se o sn' 'ilp d,'. i'ase Cp is .i! ''ii'nshvuga! can' i 'r 'w q'so'p a t ed',q

! I NO ir iect o .- i ave been OOC . "poled I On s- i ino Nor[I R,r e 'c<t i si'I '!n!!.

8 ACT E RIR

Bacterial ir fpct ons or sf'nntp l.ave beer Observedfo' many yea s Scierilists liave no'need tna: bacte;alinfec:;on in!he b ood or d gies: ve g 'and u,.i;iiiyOCCurS wheri Snr rnp are weakeried Otfierw se,r Or>a' Shrimp a',So may beCOme ii-'eCted i cond.' o" sfavOr preSenCe Ot a part,Cula ly I artTtul baCte'iun.

Shriinp body fluids are n.OSt Of.en ir feet'ed by 'thebacterial group named Vibrio S'rimp infected withbacteria show disco'oration of t te body tissues inSame inStanCeS. bu'. nOt in Others Ttie CIOtting f,.nCtten of '.he blood C.itiCal in wOund repair, iS S Owec Orlost during some nfections. Occasionally parts of thebody are iost Fig. 6!

If infected by bacteria capab'e of us,ng snell fornut ition, Ihe exoskeleton will demonstrate erosiveand olackened areas Figs 9 and 10!. These oac;eriatyOically attack edges o tips of exOSke eton parts butif breakS Occur m the exOSkeletOn the baCteria arequick to eriter and cause damage

f-ilarnentous bacteria are COmr.",Only fOund attaChedtO the Cu'. Cle. partiCularly frir ge areaS beSet withSetae Fig 11!. When infeStatiOn iS heavy, filamen-tauS baCteria may alSO be preSent in large quantity Onthe gill filaments Smaller, less obvious bacteria alsosettle on cuticular surfaces but are not considered asthreatening as the filamentous type.

FUNGI

Several fungi are known as shrimp pathogensThree groups commonly infect larval shrimp, whereasanother attacks the luveniie or larger shrirrp. The

Fig 7 Histological cross section ot a digestive gland tubule.Rickettsial microcolonies are shown at arrow Rickettsia will exhibitconstant brownian motion and color red with Giemsa stain, butelectron microscopy is needed tor definte diagnosis. Specimencourtesy ot J Brook !

Fig B Oamagi lo at%onion of a slirimp as a insult ol V.brioIrl'iecti on

Fig 9 Shrimp with numerous black erosive areas considered aresult ot schon by bacteria The darkening itselt is considered ahost response

Fig. to Microscopic view of a lesion on a uiopod tail part I

S ,lrCh lur HOSI

erg 1'I MiCrOScopic view Of iilamentcuS basteria On a Shnmppioopod

Frg 1 2 TranernieeiOn Ot Lagenrdium A FunguS SendS Out diS-Charge tube Out Oi Shiimp bOdy B Veaicle terre C VeerCIe prO-durxrs motile spores that are released D Motile spores contactshrimp and urxlergo encystment e Germ tube is sent inlo thebody ol the shrimp and fungus then spreads throughout

Frg t 3 Lagoirrtfium rnleCten in larVal Shrimp. Note ertensivedevelOPrnent Of blancfringS Of luriguS lhrcughcut Ihe bbdy. Phptctsturleey Ot Dr OOn Ughlner, Univareily Ot AriZOna.!

moSt COmmcn geflus af leo! irg Iarv,i. S.";r imp isf agenfCkvm. Less common b t important a eS,rotpig'ium and HahphtfforOS. The me:t'Od cf in',eCtlorlrequires a thin cuticle such as tnat charac'.e is; c oflarval shnrtlp Ffg. t 2!.

The most common genus a'fectir g '.arqer shrimp isFvSarium. It iS thOught '.hat In S furquS gair s eniryintc the bOdy thrOugh cracks cr erOdec areaS on thecuticle Fvsarivm may be identified by t ie oresenceOf CanOe-Shaped maCrOCOnidia that the funguSpraduCeS. MaCrOCOnidia and exampleS Ot lur galinfeCticnS are Shcwn in FigureS 13-16.

Fig 14. Canoe shaped macroconxfia ot Fusarrurn Thesestructures bud otf ot branches ot the fungus artd serve io transmitfungus to shrimp

Fig 1 5. Microscopic view of fungus in gill hlament.

Fig. 18. h4crasmpic view of fungus at tip ot antenna

PROTOZOA

Protozoan parasites and commensals of shrimpwill occur on the inside or outside of the body. Thoseon the outside are considered harmless unlesspresent in massive or burdensome numbers. Thoseon the inside can cause disease and are representa-tive of several groups of protozoan parasites. Micro-sporidia, Haplosporidia and Gregarina. Members ofthese groups are known or believed to require thatsome animal besides shrimp be present in order tofacilitate completion of their life cycles, A few proto-zoa are known to invade weakened larval animalsdirectly and contribute to disease.

Micrasporidians

Microsporidians parasitize most major animalgroups, notably insects, fish and crustaceans, Inshrimp, microsporidian infections are best knownlocally as cause of a condition known as "milk or"cotton shrimp Figs. 17-19!. Microsporidians be-come remarkably abundant in the infected shrimp andcause the white appearance of muscle tissues. A

Fig, 17. Infected or "milk" shrimp upper! in comparison to normalshrimp lower.! Photo courtesty of Dr. R. Nickeison.!

Fig. 18. Two brown shrimp cut across tail. Note infected area arrow!.

Fig tg. Grass shrimp with "milk" shnrnp condition. The normalshrimp in the figure is transparent.

Fig. 20. Microscopic view of many spores of Ameson =Nosema!sp. The spores are free or unenveloped. Parasitic microsporidiansof commercially important shrimp with enclosing envelopes areassigned to genera P/eisfophora, Tftefohania and Agmasoma. Thelatter two differ from Pfeisfophora in that their members retain aconstant spore number of eight per envelope. Pfeisfophora sp.have more than eight spores per envelope.

Fig. 2t. Microscopic view of many spores of Thelohania sp. Noteenvelope arrow!.

Fig 23 Life cyde of microsporidian of shrimp A Ing esson ofspores by shrimp. B In gut ot shrimp, the spore extrudes afilament that penetrates gut Wall and depositS an infeciivC unit Acell engulfs this unit C Intoctive unit enters the nucleus of the cell,undergooS development and lhen d.'videS to form schiZonts 0Schizonts then divide and develop into spores E By the timespores are formed, they are located in a specific tissue lrnuscle,TTSSueS arOund TnteStfne, etc ! The Spores a ro either d schargod

from the shrimp while living or affer death, but the method ofrelease and the pathway taken is nol knov n F ExperimentsdeSigned IO transmit infOCtidn by feeding infeoted Shrin p tOuninfected have been unsuccessful I I is assumed particular eventssuch as involvement of another host may bu required to completepassage from one shrimp to the next Successful transmission hasbeen reported when infected shrimp were fed ro fish speckledtrouT! and fish fecal material was then fed to shrimp

Fig. 22 Agmasoma penaef in white shnrnp This parasite is alwaysIOcated alOng the dOrsal midline arrowS! Aduanced infectiOnS canbe Seen thraugh the cutfofe vnth the unaided eye

typical catch of wild shrimp will contain a few individu-als with this condition. These shrimp are usuallydiscarded before processing. Depending on the typeof microsporidian, the site of infection will be through-out the mueculafure of the st rimp Or,',n parti ular,organs and tissues.

fvticrosporidians are present in the affected shrimpin the form of spores, Spores are small cells that candevelop into a new individual. They are very minuteand detection requires examination with a microscope Figs. 20-21!.

Infected shrimp are noted to be agile and appar-ently feed as normal shrimp However, tissue damageoccurs and no doubt affects many life functions. Noeggs have been found in "milk shrimp and it isSuSpeCted that all typeS Of miCrOSpOridian infeCIIOnScan render shrimp incapable of reproduct,on Fig. 22!.

The life cycles of shrimp rnicrosporidians have notbeen satisfactorily worked out. However, examinationof the cycles of related species and the rniscellane-ous facts contained in literature indicate that the cyclepresented in Figure 23 is representative of rnicrospo-ridians.

HapfosporidiaA member of the Haplosporidia, another spore-

forming protozoan group, was recently recognized asimportant to shrimp health when researchers foundinfected animals in an experimental population thathad been imported into Cuba from the Pacific Coastof Central America. The pares.es invaded ar.destroyed tissues of the digestive gland Fig 24!.Gregarirtes

Gregarines are protozoa that occur within thedigestive tract and tissues of various invertebrateanimals. They occur in the digestive tract of shrimpand are observed most often in the form of a tropho-

0

Fio 24 A Orating cl transverse sect, on of digestive glandI inule con'.a nin; haplospor dian parasites Straight lineseparates apoe-rance of a tubule into early treater's le'!! andlate s'.ages r:I r.,'ecticn Drawn !rom Oyhova et al,; QRB ! Arent -r ropcrt from I.tslaysian Penac~s mcnodon tigures a cuitesimilar man,;Cat",ion by an aSSurned mi rOf Feridian infeCtianEf. For compo ison a seel,on of a normal tubule Toward thedigest.~r. tract secretory ci 'Is r, 1! prerlom.note ano '�braes cellsf3; become more r un.erous Rhsorp',ve ce'Is ',2! conte,nvarying amounts oi va uole; accord. r 9:o nutritional status

� H

Fig. 29. Microscopic view of gametocyst ofhf orna iopsfs sp

Fig 25 Microscopic vi ew ottrophozoi te of Afematopsfs sp

' ~P.J-'@ ~

Ftg. 26. Microscopic view ot trophozoite oftttematapsis sp.

Fig 27 Left.! Microscopic view oftrophozoites of Nematopsis sp.Fig 28 fAbove.t Troohozoit~ nrCephafabus, a gregar inc thatattaches to tho base of the terminallappets of the shrimp stomach ratherthan the intestinal wall Photocourtesty of Dr C. Carkem.!

10

Fig 30 Life cycle of a gregarine of shrimp A Shrvnp ingestsspareS with bOtlam debris e. SpOroZOde emerges in the gut of theShnrnp C SpOroZoite attaches to the rntest;nai walI and grows intoa delicate trophozoife; other trophozoites do not atach to the sallbut onta each other and form unusual shapes See Figs 26, 27 !D The unusual forms develop and attach to the end af tho

zoite Fig. 25 -28! or a garnetocyst Fig. 29!, The lifecycle involves marine snails or clams as diagrammedin Figure 30.

Minor damage to the host shrimp results fromattachment af the trophozoites to the lining of theintestine. Absorption of food or intestinal blockage bythe protozoa is perhaps detrimental. lt is the consen-sus of most parasitologists that these effects arerelalively unimportant and, consequently, theyconsider gregarines of ffttfe pathological importance.Body Invaders

Severaf protozoa have been noted to invade ashrimp body and feecf on tissues as they wanderthroughout. Tentative identifications nameParauriorfema, Lepfomonas, Parartophrys and anamoeba. Adverse effects of these protozoa are notfully understood, but they are usually found associ-ated with shrimp that have become weakened ordiseased Fig. 3t !.

Ectocommensal ProtozoaSeveral kinds of protozoa are regularly found on

surfaces, including giffs, of shrimp. Apparently, shrimpsurfaces are a favored place to live within the waterenvironment, Common on the surfaces of shrimp are

intestine irectum! tO form gametooysfs E Garnetocyst und'eigoosmultiple divisions to produce gymnospores" that are sei free wiihrupture ol the gainetocyst F Gymnospares aie engulfed by cedsat the surface of the flesh of clams G They develop to formspores in the clam. H Then the spores lwith sporozoife inside! arehberaied from the dam in mucous stniigs lslime!

Fig. 31 Weakened larval shrimp invaded by ciliated protozoansla rrows!

species of Zootharrfrfium, Episfylis, Acirfeta, Ephelota,and Lagertophrys Fig 32!.

Zoothamryium is a frequent inhabitant of the gillsurfaces of shrimp, and in ponds with fow oxygencontent, heavily infested shrimp can suffocate.Surface-settfing protozoa occasionally cause prob-lems in shrimp hatcheries when larval shrfrn p become

B F pis~rraA Zoo fharrimumL. igurihPhit ~

ig ~Z CorhmOn Quite i Qwf Ji igpf Dto2OR

E EphercraD. Acinera

Other Infestations

overburdened and are unable to swim normally. Asprotozoa continuously multiply in numbers, shrimpacquire an increasing burden until shedding of thecuticle provides relief.

Members of one unique group of protozoa, theapostome ciliates, have a resting stage that will settleon shrimp surfaces. When the crustacean rnolts, theprotozoan releases and completes the life cyclewithin the shed cuticle before entering a resting stageon a new crustacean Fig. 33!.

A variety of other organisms attach to shrimPSurfaces. Their abundant preSenCe on gillS artd urnbscan interfere with breathing and mobility. Small,single-ceil plants called diatoms are often foundattached to larval shrimp in hatcheries. Fig. 34!-Shrimp from aquaculture facilities that are exposed toan unusual amount of sunlight often will have o>+rgrowths of algae of mixed variety Fig. 35!.

Fig 34. Diatoms attached to gill surfaces of larval shrettP-

Frg 33 Microscopic view oi several apostome criates inside grassshrimp molt Proper ldenbfication pl genus Gaflhof be deterntlfledfrcm Irving animal Straining witri a teChnique called SilVer impreg-nation Is requried

Fig 35. Afgat overgrowth c,n shrimp errposed to ahundant light.tphote courtesy of stews Rotrerteon l

32

Occasionally, one will find barnacles, leeches andthe colonial hydroid Obejia brcuspfdata affixed tobody surfaces. These organisms are probably quitecommon in the vicinity of the shrimp and selectsurfaces of infrequently molting, older shrimp asspots to take up residence. Insects will sometimes layeggs on shrimp and a strange group of animals calledellobiopsids seltle on the carapace and prov.de tl-eshr rnp with a feathered-cap appearanCe FigS 36-38!

Some members ot the crustacean group calledisopods are paras t c on shrirr.p of commercialimportance. Commercially important shrimp of theQulf of Mexico are appa. en;ly not parasitized. How-ever, smaller shrimp of the fam y Palaernonidae areoften seen infested along our coastline. Cornrnercialshrimp of the family Penaeidae are parasitized inPacific areas Fig. 39!.

Fig 36 Grass shnrnp with leech attached Photo txttirtesy ofKenneth A H Read!

Fig. 37. Shrimp with insect eggs attacherf ttt crt ttctrtar surface.

ig, 38. Etlobfopsrds attached to shrimp cuticle

Frg 39A. Asian shnmp infested with parasitic isopods lgill coverflared open to expose parasites!

Fig, 39B. Paraaiaaa rernOved fram shrimp.

Worms

L

A

13

Worms!ha; have been!cond ir sl rivp a e lrer a-todes '.,kes!, cestodes taoeworms! anc nematodesIroL!ndworrns! Soi iespecies are r, ore cc mo< t'ariOthifrs arid, aS yet. nOrie have been ~now i lo CauSewideSpread Snrin p riOr',ality Wor+S rr ay be fdi!nd inv'irious partS of tfie body I-tg 40i

Fig 40 Ccr imon sites oiinlestation by wormsTapeworr s usually assoc.-atod with tissues coveringdirgeSSVe g'and 2 Raund-worms, in and outside oforgans in cepha'othorax butalso in and along outside ofintestine 3 iFlukCS, can-rnonly encysted in tissuesadiacent to cigars in ccpha-lothorax, but also inabdaniina' musculature andunder cu'.i ale

Fig. 42. Hypothebcal life cycle of a shrimp fluke, Opecoe!aidesfi!rrtrnatus. A. Infective stage or oercaria penetrates shrimp. B.Cercana migrates to the appropriate tissue and encysts forming astage called metacercaria. C. Shrimp infected with metacercaria iseaten by fish silver perch, red drum, sheeps head, several others JD Shrimp is digested. This releases metacercaria. Metacercariastage undergoes development until it forms an adult. E. Eggs laid

TREMATODES

Trern;tlodes ",ukes! a e presenti in Shr rnp as!mrna!art. forn s metacercariae! ercysteo in variousbocy t.ssues bletacerca tae o' . 'enia',odes of tnefar'iii es OpecOel cae MiCrapha'lidae and ECh nOStO-na', dae hav. Oeen;epo.ted f"Orr. COrnmerCial SpecieSof peiiaeid Sn '. o IF.g 4T!. One speC,es, Opecoe/ot-deS !rn!br!aftts.haS de en nc ted tO be mare COmrnOnthari oil-e s. ancf '.ne hypo.het,cat I fe cycle of thiSSpeC eS iS . IuStra'.ed in Fig~re 42.

Fig 4' Drawing at miCrOscopic view Of common flukeS Of Shrimptexcysted ! A Mrc!opha !us sq B Opecoe!aides trnttrrratus

by adult fluke pass out of fish with wastes Egg hatches and anirlfeCtiVe Stage knOWn aS a miraatdtum is releaaed, The miracrdtumpenetrates a snail and multiplies in number within sporocysts. F.Cercariae develop withrn sporocysts When fully developed,cercaria leaves the sporocyst and snail and swr ms in search of ashrimp. If coritact is made with a shrimp within a short period, Ihecycfe is completed.

14

CESTODES

Tapeworrns in shrimp are assoc,ated typ ca'Ily withthe digeStive gland. They are uSually fOund imbeddedin the gland, or next to it, in the covering tissue. Inshrimp, tapeworrns are present as irnrnature forms Fig, 43!, while adult forms a'e found in rays Speciesof the genera Prochnstfarteifa. Parachrisfianefla andRenfbu!bvS are COrnmOn Other tapeworm s from witdshrimp include a relatively common pear-shapedworm of the intestine and a iess common worin of thecyclophyllidean group. Tapewo.ms are rr,ost of;enencountered in wild shrimp.

Differentiation between the tapeworm groups isrrtade in general body form and tentacular arma ureA hypothetical life cycle for Prochrisffane!ia penaef ispresented in Figure 44

NEMATODES

NematOdes OCCur mOre COmrnOnly in wild shrimpthan in cultured shrimp The degree of infection isprobably related to the absence of appropriatealternate hosts in culture systems. Nernatodes willoccur within and around most body organs, as well asin the musculature. Nematodes of shrimp includeSpirocamalfanvs pereiraf Fig, 45A!, Leptolaimus sp.and Ascaropsfs sp. The inost common nematode inGulf shrimp is Hysferothylacfvm reiiq'uens Fig. 458!.

It is the juvenile state of nematodes that infectsshrimp with the adult occurring in fish. An illustratedlife cycle thought to represent Hystercftttylaciumretiqvens is depicted in Figure 46.

Rg, 44, HypCNWlicat rife cycle of the tapeworm, procftristlaneffapenaei Kruse. A. Shrimp eats a copepod or other smait crustaceaninfested with larval tapeworm. B, Tapeworm develops intoadvanced larval stage iln tissues of shrimp. C. Stfngray ingests

Fig 43 A A drawing of the shrimp tapeworm, Prochn'st.'aneiiapenaei as it would appear in a microscopic view after removal fromits cyst B Unnamed pear-shaped tapeworm larvae in gut Photocourtesy of C Corkern !

Fig. 45 Drawing of intctosoopic view of head end of A! S piro-Camalianus pefeirai and B! Hysterprhylaoium Sp COmmpn rOundworms found in penaid shrimp.

infested shrimp. D. Tapeworm develops into adult in gut spiralvalve! of ray and begins to release eggs, E. Eggs pass out of thefish with feces and are eaten by copepcd. Eggs hatch and larvalworm develops inside copepod.

Fig. 46 Hypothetical life cycle of Hysferorhyfaciurrt reirquens, aroundworm of shrimp A Shnmp eats a copepod or other smallcrustacean infeSted with larval roundwOrm 8 Roundwormdeve'lops into advanced larval stage in tissues of shrimp C

Fig 47. Eyes of shnmp are normally black, but rubbing ol a tankwall has cauSed thiS eye tO appear WhitiSh beCauee Of a preminentlesion

Fig. 48. Cannibalism usually begins as other shrimp devour ~appendages.

TcadfiSh ingOSts infested shrimp O Roundworm dOVOiOpS intOadult in gut of hsii and begins to release eggs E Eggs pass oui oftho hsh with feces and are. eatun by copopod

Miscellaneous Conditions

INJURY

Shrimp CutiCle iS eaSily damaged in aquaCulturesituations when hard structures are impacted orrubbed. Fig. 47! Shell damage may also be inflictedby the pinching or biting of ottter shrimp in crowdedCOnditiOnS. CannibatiSm haS an irnpdrtant influenoeon survival in some phases of shrimp culture wherestronger individuals devour weak ones Ftg. 48!

DEVELOPMENTAL PROBLEMS

Deformaties are quite prevalent in some popula-tions. They arise from complex interactions thatinvOlve enviranrnent, diet and gene expreSSiOrl.Bodies may be twisted or appendages misshaped ormissing Deforrnaties are less prevalent in wild caughtlarvae than hatchery populations probably becausewild shrimp have more opportunity for natural selec-tion and exposure to normal developmenta! condi-tio ns F fg. 49!.

F. 49. Deformed ktlval shrimp Arrow points to deformedFigappendage

I

16

!vttolt arrest occurs i«affected a ima s of somepOpulatiOns AnirnalS begin Oui ire i,-.able to cOI",-pleio '.I.e n'citing process !n so-e CaSes,!here isabnO ma adherenCe;O under'ying Sk r. bu' mOStanimals appear to lack the r ecessa y s!aminaNutrilional inaceauacies and wa',er qua .y fa tarshave been icent.f,ed as causes

Tl.e Shell Or Cut C e r, ay beco;ite Iragile dur. igdevelOpment Tl-e rr.asonS IOr SOfl SI'iel'.S rel, ain amystery. but many th nk ',I e conc!it on res '.s or rarl yfrOrn inadeOuate nut itiOn StarvatiOn! Shet'IS arenOrmally sott for a coup e of cayS after molt g, butShe', S Of thOSe suffering frOTt SOft-Shell conC lionrema,n bo'.h soft and thin and ave a tenclency 'ioC'ao~ under the sltgnteSI preSSu e

COLOR ANOMALIES

Shrimp Of unusual COIOr are OCCaSiOna ly loundarnOng wild and '.arm SIOCkS Inc Str king cc oratiOn,whloh may be g Old. blue Or p»t'I. appea S throi.ghoiitthe t,SSue and iS nOt COnf.n.d:O the CutiCle Or unde-ly-ing skit . A genet,c cause is suspecteo I ransfo malion;o blue co'.orat. on from a na'. 'ral brown is know Ifor some capt ve crustaceans and has oeen linked tonu'.ritiOn. Pond-Cultured PaCil C tiger Shr,mp SOme-timeS dove!Op a COnditiOn where d gestive glar d de-generatiOn COn',ributeS tO a reOOiSh COIOratiOn.

DIGESTIVE GL.AND MANIFESTATIONS

POnd-reared Shr mp occasionally die in largenumbers because of diseased digestive glandsShrimp digestive gland tissues are organized asnumerOuS tubular StructureS that u.:imately feed intOthe digestive tract The Speo alized Cells that line theinSide Of the tubuleS are par',icu',arly fragile and maybecome damaged by a var ety of causes. Oncedamage is initiated, the shrimp will beg,n to respondin a predctable manner that progresses toward thefarmat.On Of granu!OrnaS FigS. 50, 51! Many Shrimpthat exib,ted this manifesta',ion were lost duri" g 1985-1988 on Texas shr,mp farms.

Fig. 52. A Shrimp photographed IA-left! near time Of baok iniury and B-nght! hours later Injury by a toxin or disease agent will usuallytrigger a similar teSpOnSe Ot inflammatiun and melanizatiOn

Fig ha T.~ui~.u;:a. si. r~«i I d Si St,c Clune '. !,ii:sprogiession oi granuior,ia format,or No~nial i.oules aie io I' u ii Iifhl! and aiiucted iubulos are to tiii; iigi I f' !

Fig St Tissue squaSh O'I hepatepaneieaS with advancedgian Ionia formation Note dark necrotizeo tissue ot tubulesIarrows!

17

Fig 53 State of gills during initial stages of darKening condition.

Fig 54 Emaciated shrimp. Gills and body fringes have becomeobviously darKened and the SOft Lail iS COvered with a thin andfragile cuticle

Fig 55. Lipoid fat! spheres In microscopic view of digestive glandtubule

INFLAMMATION AND MELANIZATIONConditions of tissue darkeninening is a frequent

occurrence with shrimp and otheer cr usta c cans. In th eUSual ~se, blood CellS gradually con re ate lnpartlou ar tlSSue areaS Inflammatlan! where damagehaS OCCurred and thiS iS fOllOwed by pigment mela-nin! depos'tion. An infective agent. inlury or a toxinmay cause damage and stimulate the process Fig.52!. G~lls are particularly prone to darkening due totheir frag~le nature and their function as a collectingSite IOr eliminatlen Of the bOdy'S waSte prOduCtS Fig.53! GillS readily darken upOn eXpOSure tO taxlcmetals or chemicals and as a result of infection bycertain fungi Fusanurrf sp !

EMACIATION AND NUTRITIONAL PEFiCIENCYUnfed shrimP lose their normal full and robust

appearance. The Shell becOmeS thin and flexible as ltcovers underlying tissue such as tail meat thatbecomes greatly resorbed for lack of nutrients.Molting is curtailed and shell and gills may darken intime. Emaciation may result from unavailable nutri-ents or loss of feeding behavior after exposure tounfavorable environmental conditions Fig. 54!.

Prepared diets deficient in necessary constituentsmay predispose or cause disease. Vitamin C defi-CienCy, fOr example, Will initiate darkening Of gillS Orcertain tiSSueS aSSOCiated with the cuticle and eventu-ally result in deaths.

DigeStive glandS SOrnetirneS wil! beCOme ~educedin size, Among other things, this is an indication ofpoor nutrition. Well-fed shrimp will have an abun-dance of fat globules within storage cells of thedigestive gland tubules that provide bulk to the gland Fig. 55!.

CRAMPED SHRIMPThis is a condition described for shr~mP kept in a

variety of culture situations, The tail is drawn underthe body and becomes rigid to the point that it cannotbe straightened Fig. 56!. The cause of cramping lsunknown,

Fuff flexure IA-felt.! FtaxureFfg se, Cramped Shrimp Cenmaintained when piessu~ ~

MUSCLE NECROSIS

Opaque muScles are CharaCter S'..C O' thiS COndi-tiOn Witch Shrimp are expOSed IO SlreSS'ul COndl-tiOnS, SOCh aS IOw Oxygen Or Crawdir g. the mu ClesI Se their rOrmal transparenCy and beCOime b',O',CheCwith whitiSh areaS thrOughCu'. Tiiis may progreSS untilthe en;,re tail area '.akeS On a wmuSn appea anCe

Fig 57!,If shrimp are w!nd awn from Ihe adverse eri. iron-

ment before proloriged stress exoos."e they mayreturn tO nOrma Extremely af lee'cd Shrimp CO nOIrecover, however, arid die with n a few m nu'.es Irir iodera'.ely affected shrimp, only oar'.s of;he bcdyreturn tO narma, Other par,S typiCa',ly Ihe laSt Seg-mertS Of the '.ail are unable:O reCover F ig. 5FI!These shrimp die w ttiin One Or twO dayS Srrirtpmuscles with this concitior are kr own lo unde gonPCi'OSiS death Or decay of '. Ssuel. Trit! wh .ishCOnaitiOn Can be COnfuSed with milk Shrimp miclo-sporidian infectionl, out exarr ir ation of the 'iguresshould ad in distinguishing the two diseases

TUMORS AND OTHER TISSUE PROBLEMS

COnSpioucuS bOdy SweiingS Or er argerhentS oftiSSueS have been repOrted ir. Shrirhp. In rrOSt CaSeS,affec',ed ir,divid a's were captured from po lu',edwa'.ers Occurrence of shrimp with evident tumors srare:n commercial catches Miscellaneous irritationsexperienced by captive shrirr p ir tank sys;errs w IIsometimes reeult in fOCal a eaS Of OSSue OvergrOwth Fig. 59!

A hemolymphoma or fluid-fi'led bl.Ster may fOrm onthe inner surface of the portior of carapace tfiatcovers the gills Fig 60! The cause Of this rare con-dition is not known Another rare condition is necrosisand subsequent melanization of tissue within themuSCulature t.ike hemOlyrnphcma, '.he CauSe S un-known and marketability of the procuct is greatlyinfluenced Fig 61!

A degeneration of male reproductive trac'.s occa-sionally occurs in captive adults of certair penaeidspecies. A swelling and darkenir.g of the tubule lead-ing from the testes to the spermatophore is readiiyapparent when viewed through the translucer.t body Fig 62!.

A condition where the the basement layer of tissuelining the intestine thickens and results in loss of foodabsorption and some nerve dysfunction is knownfrom Penaeus japaniCuS and PenaeIJS merguienSiS. Acause has not been established.

ENVIRONMENTAL EXTREMES

Temperature, irradiation, gas saturation, hydrogenion content pH!, oxygen content and salinity all haveappropriate tolerable ranges for sustaining life ofvarious shrimp species and life stages. If theseranges are exceeded or extremes combine for an

Fig 5? S'iii.'-ip W li ne.re; c -iu C i i;,;ii, IO. i.vir9 .,iAffected ossui: a; snow

Fig 58 Shrimp with adxahced rnuSCle neomsis larrowl shownbeside normal shnmp

Fig 59 Tuinoious growths on an adult shnrnp hom a tarh systoin

Fig, SO. BliSter COnditiOn Inset shows biteter removed The bliSterwill darken upon death of shrimp

18

Fig 61 Areas ot rnelanizeci necrotic tissue in tail musculature

Fig 62 Darkening of male reproductive tract of Penaeus sryiirosins. A Normal tract 8 initial darkening Darkening will advanceuntil spermatophores and testes become affected. Photoscourtesy of George Chamberlain !

Fig, 63. Precipitant of iron salt an a shrimp's fringe hair setule!.

interactive effect shnrnp wiil becon e seasedestdes,fte d.rect effect Irottt these noninf

a ents,exnoninfectious

g, expOSure may reSult in predtSpOSal t ffof opporturt stic ttlfecttve agents. o e ects

Gas bubbles will form in the blood of h0 s rimp if ex-in gas satur-poseo to waters with large differences in g

at OnS. If a large aincunt of bubbiing OCCurS n thebloOd. death will reSult

In the preSenCe of aCidic wa',er, rnineralS will Oftenprecipitate on cutioufar surtaCes USualf th p p-e precipi-tant is iron salt if g 63!

TOXICITV

Poisoning car result from toxic substances ab-sorbed f;om the water or consumed food. Water mayaCCumulate exCeSSive COnCentratiOnS Ol amrnOna,nitrite, hydrOgen sulfide Or CarbOn dioxide, all ol whiohCan have a tOx iC effeCt On Shrimp. SOme meta'S alSOmay cause a toxic effect when present in excessBOth preSenCe and IOxictty of theSe CherniCalS areinfluenCed by the changeable environmental cOndi-liOnS rnenl Oned abOve They may aCt Singularly Orhave combined effects.

Certain micrObeS and algae will exCrete pOisOnOuSmaterialS. FxampleS Of algal releaSe are the OCCa-sional red tides that occur along our coast Aside fromsurvival loss, affected animals behave in a disorien-tated manner. MiCrObeS beCOme COnCentrated in highdenSity rearing systemS and SuCh SysternS are knOwnto develop species that can cause damage by toxicrelease.

PeStiCideS Can be harmful ff they OCcur SeaSOnallyin surface water supplies affected by agricultura!practice. Because of migrations into estuaries or neareffluent disposal sites, wild shrimp populations aremore susceptible than cultured stocks to the variety ofpollutants released.

There are reports of toxicity caused by the foodshrimp consume. Toxins from microbes are known tobuild up in feeds stored in unfavorable conditionsSome foodstuffs and live larval food, such as bnneshrimp, can contain pesticides. Perhaps more com-rnon are undesirable effects of feeds that have agedand become rancid.

Breakdown of lining tissues necrosis! of theintestine have been associated with consumption ofcertain algae. Because cultured shrimp feed both onprepared feeds and bottom materials, it is suspectedthat the occasional occurrence of detrimental irritantsand toxins contained within bottom surfaces couldcause tissue breakcfown when such sediments areconsumed. 8

Selected Bibliography

SHRIMP SPECIESBielsa, LM, W H Murdich and R F

tabrsky 1983. Species proviles Ide historiosand environmental requirements of coastalkshes and invertebrates south Florida!�pink shrimp IJ.S. Fish Wildl. Berv. FWSIOBS4t2/1 1, I 7 IJ.S. Army Corps af f ngl-neere, TR ELd82-4I 21 p

Hoithuis, L 8 1980 FAD species cata-logue. Vot I Shnmps and prawns ot theworld FAO Rsherles Synopsis No. 125,Volume 1. Food and Agriculture Orgentza-llon ol the United featlone, Rome. 271 p

Lassuy, D R 1983 Brown shrimp SpacleS prOtilea LdO hiSIOriaS and environmentalrequiremonts of coastal tishes and invorlebraleS Gulf Of MexiCO! VSFWS Publ, No.FWS7OBSJt2I I t.t

Muncy, R J 1984 Whiteshrimp Speciesprofiles 4fe histones and environmentalrequirements ol coastal fishes and inverte-brates Gull of Mexico! UBFWS Publ. No.FWSIOBS-$2I11.20 also South Atlanbc.FWSIOBS-82I I 1.21!

Perez. Faitante, I 1969.Western Atlanticrhrlrnps of the genus Penaeus L.B, FishWildl. Berv. Fish. Bull., 67 461 591

� � - 1988 illustrated key to periaeoidshrimps ofcornmerce in the Americas NOAATechnical Report t4MFS 84, 32 p

Yu. H-P. and T- Y Chan. 1986 The lllua-traled Penaeoid prawns ol Taiwan. Sauth-em Materials Center, inc, Taipei, 183 p.

SHRIMP ANATOMY

Belt, TA and D V Lighlner. 1988. Ahandbook ol normal PensHd shrimp his-tology. Workl Aquaculture Sordety, BatonRouge, Louisiana, 114 p.

Johnson, P T 1980 Histology ol thebtue crab, Cefffrrecree sett!due, A modelfor the Deca pads. Praeger Publ, N, Y, 440

P- Foster, C A end H.D Howse 1978 Amorphological study on gills of the brownshrimp, Peitaeus azfecus Tissue ~ Celt,10:77-92.

Young, J.H 1959. Morphologyofthe whiteshrimp Penaeue se Nerve Unnaeus 1758!.Fishery Bulletin, 59 �45!:1-168.

GENKRAL DSEASES

Couch, J.A. 1978. Diseases, parasites,and toxic responses of commercial shrimpsof the Gulf of Mexico and South AtfantfcCoasts ot North America. Fishery Bugetin,78:1~.

Davidson, E.W., ed., 1981. Pethayene-ele rrl lrlvertebrete IIRIvrbtet dteeeaea, Al-

lanheid. Osmun Publishers, Totova, NewJersey

Foigenbaum, D L. 1975. Parasites of thecommercial shrimp Penaevs vannamei Booneand Penaevs brasdiensis Latreiile Bull. Mar.ScL, 25'491-514

Fontaine, C T 1985. A survey ol potentialdisease causirig organisms in bait shrimptram West Galveston Bay, Texas NOAATechnlcalMemorandurn NMFS-SEFC-169,25 p + 16 figs

Johnson. S K 1977 Crawhsh and freshwater shrimp diseases Texas AB M Univer-sity Sea Grant College Program Publ, No.TAMU-SG-77-605

Kruse, D N. 1959 Parasites of the cornmercial shrimps, Penaeu azfecus Ives, Pduorarum Burke nioad, and P. sefiferus�nnaeus! Tulane Stud, Zoal., 7 123-144.

Lewis, D H and J K. Leong, eds 1979Proceedings of the Second Biennial Crusta.cean Health Workshop. Texas ABM SeaGrant College Program Publ. Na. TAMU-SG-78 114, 400 p

4ghtner, D.V. 1915. Some potentiallyserious disease problems in the culture ofpenaeid shrimp in Norlh Amenca. Pages 75-97 In Proceedings af the Third U,S.-JapanMeeting on Aquaculture at Tokya, Japan,OCtOber, 1$-18, 1974. Special Publicauan ofFishery Agency, Japanese Government andJapan Sea Regional Fisheries ResearchLaboralory, Niigata, Japan

1983 Diseases of cultured pe-naeid shrimp, Pages 289-320 in McVey,J.P., ed. CRC Handbook of Mariculture.Volume 1. Crustacean Aquacutture CRCPress, lnc., Boca Raton, FL

Lighlner. D.V., R.M. Redrnan, D A Danald,R.R. Williams, and L A Perer 1984 malardiseases encountered in controlled environ.ment culture of penaeid shrimp at PuertoPenasco, Senora, Mexico Pages 25-33 irrC.J Sinderrnann editor!, Proc. 9th and 18thL,B. Japan lteetlnge on Aquaculture,NOAA Tech. Rept NMFS 16.

Qverstreet, R.M. 1973 Parasites of somepenaeid Shrimps With emphasis an rearedhosts. Aquaculture, 2:105-140.

1978. Manna maladies? Worms,germs and other symbionts from the North-ern Gultof Mexico. Mississippi-Alabama BeaGrant Corieortlum Publ No. MASGP-76-021, 140 p.

1986. Solving parasite-relatedprobkams in cultured crustaceans. Pages 309-318, trt M.J. Howtdf ed., Parasttotogy-QuoVedi& Proc. 8th fntt.Congress Pareailol.

Provenzano, A.J. 1983. Pathobkdogy.The Biology ot Cruetacee. Volume B.Ar~: Press, New York, NY, 290 p.

Sindermann, C J and D V I.ightner1968 Developments in aquaculture and fisheries science. '17 Disease diagnosis sndcontrol ln Norlh American marine aquac-ulture 2nd edition Elsevier, Amslei dam,431 p

Tareen, I LI 1982 Control of diseases inthe cultured populakan of penaiiid shrimpPenaevs semisvlcalvs de Haan! J. WorldMariculture Sac., 13 157 16 i

Viliella, J B, E S Iversen and C J Sindermann 1970 Comparison of the parasitesof pond reared and wild pink shrimp Penaevs dvorarvm Bur kenr cad! in South FloridaTrans. Am. Fish. Boc., 99 169 794

VIRUSBell, T A and D V Lightner 1963 The

penaeid shrimp species affe~ted and knowngeagraphic distribution of IHHN virus inFirstBiennial Conference on Warm WaterAquaculture Crustacea. Brig ham YoungIJnrv. Hawaii Campus!, Lare, Hl AVI Pross,Westport, CT.

1984 IHHN virus Infectivity andpathogenicity studies in Penaeus stylirostiisand Penaevs vannamei Aquaculture,38 185-194

1987 IHHN disease of Penaeusstylirostre: Effects of shrimp sire on diseaseexpression. Journal al Fish Diseases,10 165-170

Bonarni, J.R. 1976. Viruses from crusta-ceanS and annelidS Our Slate Of knOwledgeProc. Inl Cagoq. lnverlabr Pslhol� 1 20-23

Banarni, J R, M. Brehelin and M Weppe1986 Observations sur la pathogenfcr te Latransmission et la resistance du MBV mono-don bacutovlrus!. Proc, Ind. InternationalColloquium on Pathology ln Marine Aq-Iieculture, Porta, Portugal. 7-11 Sept. 1986,p. 119.

Bove, G. Ceshcra, G Giorgetti and MVanefli. 1984 Isalalion of an IPN-like virusfrom adult kur uma shnm p Penaeusj aponicvsBull. Eur. Assn. Fish Pathal., 4 21

Brock, J.A., D.V. Lightner and T.A, Bell1983. A review of four virus BP, MBV, BMNand IHHNV! diseases of pe nacrd shrimp withparbcular reference tO CliniCal Significance,and conhol in shrimp aquaculture. In'tama-tlonel Council for the Exploration of theSee C.M. 983/Gen:IOIMrni-Symposium.

Chen, S.N. and G H, Kou. 1989 infectionof cultured cells from the lymphoid organ ofPenaeua monodon Fabricius by rnonodon-type baculovirus IMBV! Journaf ol FishDleeeeee, 12;73-76.

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1981 Viral diseases of invurleorated Other than ineectS PaaeS 127 160 inPathogeneeis of invertebrate microbialdiseases E W Davidson, ed AllanheldIosrnum Publ ! Totowa, Naw Jersey

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Couch, J A and L Courlnoy 1977 Inter-acban ol chemical pollutants and virus in acrustacean A navel bioassay system AnnN Y. Aced. Sci . 298 497-504

Diamant, A, A Colorni 1987 Viral infec-tions in penaeid shnmp in Israel and use otacndine orange fluoresce nse as a rapid diag-nosbc method E.A.F.P. 3rd InlernatlonalConference, Bergen Norway!. Aug 31-Sept3, 1967 Abstract!

Foster C A, C A Farley and P TJohnson 1981. Virus-like parbcJes in cardiaccells of the brown shrimp, Penaeus azfecusIves J. Submlcroac. Cytol�13 723-726

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1983 Hrstopatholagy and diag-nosdc methods for IHHN and M 1 V diseaseSin cultured penaeid shrimp. Proc, Sympo-~ turn on Warm Waler Aquaculture Cru sf a-c»a. Brigham Young Univ. Laic, Hawaii, Feb.9-1'I, 1983.

1983 Qbltervalfane an the geo-graphic distnbution, pathagenesis and mor-phology of the tuscufovtrus from Penaaua~ Fabrfckat. Aquaculture. 32:209.233.

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Sano, T Ntshtmwa, H Fukuda and T.Hayaahida. 1985 Baculavtraf tnfeobvity vialsOn Kuruma Shrimp larvae, Penaeus !apani-cus, of dillerent ages, Pages 397-403, in A EElle, ed, Rah and Shellfish Pathology,Academic Press, Landon

Sana, T. and H. Fukuda. 1987 Principalmfcrobfaf rJiseases of mariculture in Japan.Aquapu flute, 67 59-69.

SWnmera, M.D. 1977. Charaateriaabon Ofshrimp baculovirus Ectdogfcal ResearchRerkse, U,B. Envbonmental ProtectionAgency KPA-0000-77-130, November,1977. Iifatfonaf Technical fnformatkrn Serv-ke, Sprktglefd, VA. 22161, USA, 36 p.

Summers, M 0 and G IE Smith 1987 Amanual ot methadS tar baaufavtrue veatpraand insect cell CultWe prapedurfte TeaseAgricultural Experiment StaSen guf latin,No 'f555. 56 p

Tsrng A, D V Lightner, J R Bahama andA M Rodman 1985 Is gul and nerve syn.drome IGNS! in the tiger shrimp Panacea!aper ircus Bate! ot vkaf origin> 2ild lrl lerrie-tional Conf. pl the European Ae socle lionaf Fish PalhOIOglala. MOntpellier. Francelabs tract!

Tsing, A and J R Banamr 'l987 A newviral daeaSe Of the tiger Shrimp PanaceaAfporecus Bate Journal of Fish Ofseaaee,10 139 'I41

R I CK E TTSI ABiock, J A . L K Nakagawa, T Hayashi.

S Teruya and H Van Campen1986 Hepa-tapancieabc rickettsial infection al the pe.naaid shrimp, Penaeus margrnarus Ran-dalll, trom Hawaii Joume!of Fish Diseases9 353-355

Brack, J A . 1988 Rickettsial inlecbon olponaekt shrimp Pages 38 41 in C Sinder-inann and D Lightner, eds Otaease d4gno-~ ls and control In North American aquac-ulture Efsevter, Amslnrdam

Colomi,A, T Sainocha and B Caiorni1987 Pathogenic viruses intraduced intoIsraeli martcufture Syatema by impOrted pe.naeid shrimp Bamfdgeh, 39 21 28

BACTERIABoemare, N, F Cousserans and J R.

Bonarni 1978 Epizootieauibrionaosaedansles elevages de crevetles Penaeidae Ann.Zoal. Ecol. Anlm., 10 227-238

Boyle, P J and R Mitchell 1978 Ab-sence of microorganisms in crustacean dl-gesbve tracts Science, 20 t157-1159

Cipnani, G R . R S Wheeler and R K.Sizemore 1980 Characterization of brownspot disease af Gull Coast shrimp. J. In-vertebr. Pathol., 36.255.

Cook. D W. and S R. Loflon 1973. ChNno.clasbc bacteria associated with shell diseasein Penaeus shrrmp and the blue crab Calfinecfee sap>Ac! J, Wffdflfe Diseases,9 154-158.

Hood, M A and S. P. Meyers. 1977 aA'dr�-otv�olagrca and chitinodastfc activities assoa-ated with ~a see ferua. J, Oceanogra-phical Society, Japan, 33:235.

Leong, J.K. and C.T. Fonlaine. 1979.Expenmental assessment of the virulence oftour species pf Vibrfa bacteria in ptmtaridshrimp Pages 109.132. irr. Lewis, D E andJ.K Leong, eda. PrCroeerdkrtge Of lhe BepcndBiennlaf Crustapemt Hetdth WolkahtO. EeÃaeAkll IJnlv, Sea Orassi Cogege ProgramPubl. iso. TAIIU4MI.7$-f 18,

Lanate. D.H, 1973. Atatponfm af brownshrimp lo infection with Vfbnb sp�ploc.World Maffoultuse 8oc., 4.333.

Lewis, D.H., J.K. Leong and C. Mock.1982. Aggregation ol penal ehdmp larvae

21

due la mrcrabiaf epibtonts Aquaculture,27 149-155

Lewis, D H and A I. Lawrence 1985Immunoprophytaxis to rirrbno sp in pondroared ponaoid shrimp Proceedings Sym-posium on Warm Water Crustacean Aq-uaculture Bngham Young Unrv Laic, Hi,pp 304-307

Ughtner, D V and D H Lewis 1975 ASeptlaemio bactenal disease syndrome olpenaerd shrimp Marine Fisheries Review,37 25 28

Lightner, D V and R M Redmen 1986 Aprobable Mycobacfonum sp infection at themarine shrimp Penaous vannamei ICrusta-cea Decapodai Journal af Fish Diseases,9 357 359

Sparks, A K, 1981 Bacterial disoasos ofinvoitobratos othor than insocts, Pagos 323363 in Davidson E W, Ed, Pathogenesi ~of invertebrate microbial diseases, Allanheld, Qsmum. Totawa, Now Jersey, pp323

Vanderzani C. R Nrckelson and J Cparker 1970 Isolaeon af Vibrio parahaemofyficus from Gulf coast shrimp J. Milk FoodTechnol, 33 161 162

Vanderzant, C., E Mroz and R Nicks!.son 1970 Microbial tiara ot Gulf of Moxicoand pond shrimp J. Food Milk Tschnol,33 346.

Vanderaant, C., R Nickelaon and P WJudkins 1971 Microbial tlora ol pond rearedbrown shrimp Penaaus azfecus! App.Microbial . 21916.

Vanderzant,C and R Nickolson1973 Vibno parahaemoiyficus A problem inmaricufture'7 J. Milk Food Technol, 36 135-139

Yaauda, K and T Kitao 1980 Bacterialflora in the digestive tract ot prawns, PenaausJapan/cus Bate Aquaculture, 19 229

FUNGUSAquacop. 1977 Observabons on diseases

of crustacean cultures in Palynesia Proc.8th. Ann. Mtg, World Mericulture Boc.,8.685-703.

Bland, C E 1974 A survey of fungal die.casse Of marine Organlsrna With empha SIS Oncurrerit research concerning Lagenidiumcafffne ates. Proc. Guff Cecal RegionalBymposlurn on Diseases o< Aquatic ani-mals, Baton Rouge, LA, pp. 47-53.

� 1075 Fungal diseases of marinecrustacee, Proceedings of the Third US-Jepen Meeting on Aquaculture et Tokyo,October 1974, pp 41-48.

Sand, C.E., D.G. Ruch, B.R. Salser andD.V. Lightner. 1976 Chemical control ofLl janidi' ma to/sral pctg iogrrA ofiiiark rirraus-taoea. University of feorth Carolina BeeOrant Pubf lection, tJfrfC-BG-7842, 38 pp.

Chu, S.H. 1976. llole on the fungus dis-~ ase of the Pond cuhured grass prawn Pe-nsees mtatofyon. Jour. Rsh. Boc. Taiwan,4:31-35.

Cook, H.L 1171. Fungi ~on shrimp,FAO Atytaebtft tee Buff., 3:1$.

Equsa, S and T Veda 1972 A Fusar umsp associated with black gill disease ot thekuruma prawn, Penaeus iaponicus BateBulls lin af ihe Japanese Sociely af Scien-tific Fisheries 38! f I I 125'3-1260

Guary, J C .M Geary and S Egusa 1974Infections bacteriennes et for giques de crustaces per aides Penaeus fapoiricus Bate enelevage Colloquesur L'Aquaculture. Aciesde Colloque a. No. 1 1975 CNEXO ed, pp'125-135

Hatai, K and S Fgusa 1978b Studies onthe pathOgenic lungue aSsacialed with blaCkgill disease of Kuruina prawn. Perraeus iaporricus -II Some of the note on the BG-tungus Fish Pathology, 12 225-231

Hatai. K Furuya and S Egusa 1978Studios on the pathogenic fungus as sea a iudwilh black gill disease a! Kuruma prawn,Ponaous laponicus I isolation and ido nri ficebon ot the BG tusarium Fish Palhofagy,12 219 224

Halai,K. BZ Bien, CA Batrcados, andS Egusa 1980 Studier an the fungal dis-eases in C r uS taCe an s I I Hali ph thor os phdip-pensis sp nov isolated from cultivated lar-vae Ot the Iumba bger praWn Penaeusmoriadon Transactions ol the Mycologi-cal Society of Japan, 21:47 55.

Hose, J E, D V Lfghtner. R M Redmanand D A Danald 1984 Observations on!hepathogenesis of the imperfect fungus, Fusar-ium sofani, in the Cali amia brown shnmp,Penaeus cafrfornensis J. Inveriebr. Rethal.,44 292-303

Johnson, S K 1974 Fusanumsp.infaboratory held pink shrimp Texas A6M Univ.Agr. Ext, Serv. Fish Disease DiagnosticLaboratory Publication FDDL-BI.

Lararnore, C R, J A Barkate and H.0Persyn 1977 Fusanum infection in eyes ofmature shrimp Penaeus vannamei. TexasAS M University Agr. Ext. Serv. Fish Die-~ aee Diagnostic Laboratory PublicationFDDL-Bg

Ughtner, D.V and C T. Fontaine 1973 Anew fungus disease of the white shrimpPenaeus serrferus J. Inverlebr. Pethol.,22 94-98

tightner, D V, D. Moore and D A. Danald.1979 A rnycotic disease of cultured penaeidshrirnpcuased bylhelungus Fusan'umsolaniin Proc. 2nd. Biennial Crustacean HealthWotftshop, D.H. Lewis and J.K. Leon g, edsTexas ARM Univ. See Gtettt Publ. No.TAMU-BG-yg-t 1 4, pp. 137-'158.

Solangi, M.A. and D.V. Ughtner. 1976.Cellular Inflammatory response of PenaeusaaleCue and P. Setffarus tO the athOgeniCfungue, Fuaan'um Sp. iSOlated frOm the Cali-fomla brown shrimp. P, ca yoiniensis. J,Invertebs. Rethol., 27 77-86

PROTOZOAAnderson, I.G., M. Sheriff and G,

hktsh. tggg. A hepatopanoeasc microspo-ridian in pond-reared tiger shrimp, ~ntonodcn, from Malaysia. J. fnvertefsr. Pa-lhoL, 53:278. 280.

Baxter I'. h' R H Rigi!on anc C ifanna1970 P mslopharasp iMi rospoiir.iii Ni,semabnae! a new Pnr,i;iie ol snrinio J, inver'lebr. Pathof 16 289 331

Bradbury P C 1966 Tire lie cycle aiidn>orpiiulegy of thi' apo�loni,i!niis rili,iiiiHyafophysa ch,ii!on n g n sp J Proto-zool. 13 209 225

Bradbury p C,J C Cl,ii" p,ind J T Lyon,III 1974 Tercbro: prr.r err,rftonisp n } par }sno Ot thc endOCuticle ol liic,liiimpPalaomonr.frspugio Holthur.. J. Protozoal.21 678 686

Constransitch, M J 197n Descriptionpathology ariCI incrdenoe ol Pssiopi.ora peani.i n sp Microsporidia, Nosi.mal dae! apm as ite of cornme real s lir imp Master sThesis, Northwestern St 'nrv, 35 p

Corkern, C C Jr 1977 Pralozcan andmetazoan syn.bionts of natural and niancuitured pop ula lions ot cami herc ial s fir imp fromthe northwestern coast of the Gulf of MexicoDissertation Texas A6M University p 167

Couch, J A 1983 Diseases caused byprotozoa Pages 79 1'll. in Provenzano, Jr .A J,ed Pathobiology. TheblologyofCrus-tacae,

Dykova, I and J Loin 1988 A new haplospOrean Infeoting the hepatapancreas inthe penaeid shrimp, Penaeus vanriameiJournal ol Fish Diseases, 11 15 22

Foster, C A, TG Sarphie and W EHawkins 1978 Fine structure af theeCtOCOmmensal Zoolhamnium sp With emphasis on iis mode of attachment to penaerdshrimp. Journal of Fish Diseases, 1 321335.

Gacutan, R Q, A T Llobrera, C B Santiago, P J Guberroz an G L Po 1979 Asuclorean parasite oi Panacea monodonlarvae Pagos 202-213, in D H. Lewis andJ K. Leang, eds . Proceedings ol the Sec-ond Biennial Crustacean Health WorkshopTAMU-SG-79-114. Texas Aa M UnwersitySea Grant College Prograrn, College Station.Texas

GrimeS, B H. 1976 Notee on the diSeibutron of Hyaiophysa and Gymnodrnroryres oncrustacean hosts in coastal North Carolinaand a desert piton Of Hyaiophyaa frageri 5 p riJ. Protozoal.. 23 246-251

Hutton, R F, F S Bemal, B Eklred, R.MIngle and K D. Woodburn. 1959. Invesbga-bons on lhe pa rasites and diseases af saltwa-ter shrimps Penaeidae! of sports and corn.mortal importance to Rorida Tech. Ser. No.28, Fla. St Bd. Conserv, 36 p

Iversen, E.S. and R B Manning 1959. Anew micsosparidian parasite from the pinkShrimp R aeue due an.mi Trene. AmFish. Soc., 88:130-132

Iversen, E.S. and N.N. Van Melar 1964-A reOOrd Of the mioraaperfdlan, Theaducriarw parasnizing the shrimp. Panaeuabrasih'ensis. Bull. Mer. ScL Gulf Cerlb.,'I 4:549-553.

Iversen, E,S. and J.F. Kelly. 1976. fxlcro.

«por id as» «ucc issiully transmitted 'xpenment.iily in pink sii imp J Invertebr. Pa.thol., 27 4tf. 408

Jofinson. C A 1976 Observatiori an th iOCCurun 'e ai ilia paraaitie Ciliate 6! naphiy;tin du 'ipods in const.il watois otf the so ilfii .isti!r» Ui! fad Slalii«J. Protozoal. 23 '52"50

Jaiin an 5 K 1976 Cfi in! col coiiliol ulpi!illiiCt!O IS Cilia !San ya ing panaeld Si!Il�'pTexas ASM Univ Fish Diseaso Diagno«!icLaboratory Leaflet No FDDL $7

;976 Twenty foui hour toxic tytest«ot six chemicals to rnyss larvae olpenaeuS SoriferuS TeXas AB M Univ FiahDisease Diagnostic Laboratory Leaflet NoFDDL SB

Johnson, S K, J C Parker andHolcomb 1974 Control ol Zoofhamnrumspan penna d shnmp Proceedings of theFourth Annual Workshop ot the WorldMeric elt ere Soc le I y, Monterrey, Mexioo'l973 4 321 331

Kelly, J F 1975 A description of the his-tologtcat structure of normal and rnicrospori-dan infected pink shiimp. Per!acus duorarum Burkenroad Ph D Thesis, University olM arch Miami. Fla

1979 Tissue spec ffc ties of The-lohania duorara, Agmasoma penaei, andPleisfophora sp, microsporidian parasites ofpink shrimp, Penaeus duorarum J. tnverlebr.Pathol., 33 331-339.

Kruse, D N 1959 Parasites of the com-rnerctal shrimps, Penaeus aztocus Ives, Pduora rum Bur kenroad and P. so rife rusfLinnaeus! Tulane Stud loof�7 123 144

1966 Life cycle studies on Nema-tapsis duorarr n sp Gregarina Porospo-rrd ea!, a paras i te of the pink shnrnp lPenaeusduorarurn! and pelecy pod molluscs Disser-tation Flonda Stale University, p. 174 Forabstract see diss abstr. Ph D 278, 2919-8,l f 966!

Laramore, C R and J A Barkate f979.MOrtalrlies produCed in the protoZOae Stageaof penaeid shrimp by an unspeciated amoebaTexas ABM Univ. Fish Disease DiagnosticLaboratory Leaflet No. FDOL-S12, 7 p.

Mtglarese, J V. and M.H. Shealy. 1974.Incidence of rnicrospondian and trypan-orhynCh OeatOdes in white Shrimp, PenaeuSseri ferus Linnaeus in Sou th Carolinaestuaries S.C, Aced Scl, Buff., 36 93.

Overstreet, R.M. and S Safford. 1980.Diatome in the gillS Of the carnmercfaf whiteshrimp. Gutf Research Reports, 6 421-422.

Sawyer. T K. and S.A MacLewt. 1978.Some protozoan diseases of decapodctustaceans Marine Fieherlee Review pa-per, 1342 32 n5

Sprague, V. 195ct. Notes on three rnfcro-sporidian parasites of decapod Crustacea ofLouisiana coastal waters. Occ. Pap. Mar,Latt. St. Univ., 8:1-8.

� 1970, Some protozoan parasitesand hyperparasites in marine decapacl crus-tacea. IPagsa 4'18-430, fit: Snieszko, S.F, ed.

A syn pe«!urn on di«eas is ol fi«lies and«'! !its'ias Spec. Publ. No. 5, Amer. Fish.Soc. W,i«hington D C

Spr,igua. V and J Couch 1971 Anani,oiatud li«t ol protozoan para«i os hyperp,iiasitu«and commensal«of dec, podci, sl,icea J. Protozool., 18 526 537

GENERAL SURFACEINF ESTATION

Abu Hahiinii, R 1984 Piet senary obserV alia r>S Of t',i i etfcietS Of EP PO ureert e/i,g JnaCliopra fsopoda Bopyi decl on the reproduClien Ol PenaeuS S rrn SufC,tiuS d i HaanlDecopoda Penaeidael lnt. J. tnvertebr.Repro. 7 51 62

Baurdon. R 1981 lsopod Crust,icea IBopyridae parasites on penaeids Mem. Qr.slrarn. 91 237-260

Cheng, W W and W Y Tseng 1982 Aparasite, Eprpenaeon ingens Nobik ot thecommencal shrimp Penaeus semisufcaiusde Harm in Hong Kong Pages 393-399. in 8Morton, ed, Marine flora and fauna at HongKong and southern China. introductionand Taxonomy.

Dawson, C E 1957 Baianus fouhng ol«hnrnp Science, 'l26 1068

1958 Observations an lhe infecbon of the shrimp Penaeus semisufcarus byEpipenaeon efegans in the Persian Gult J.Psrasltol�44 240-24 1

Eidred, 8 1962. The attachment ot thebarnacle Balanus amph fete nivens Darwinand other touling organism «of lbe rock shrimp.Sfcyonra dorsafrs Kingsloy Crustaceans,3 233-206

El- Mesa, M, M Al Hossaimi and A RAbdal-Ghaffar 1981 infestation of the shrimpPanacea seiniscuicarus fde Haan! by thebopyrid Epipenaeon eiegans fChopra! inKuwait Kuwsff inst. Sci, Ree. Tech. Rep.,369 1-22

Hellman, E G and R M Yancey 1966Ellobiopsidae of Alaskan coastal watersPeclgc Science, 20 70-78.

Johnson, S K 1974 Ectocommensals andparasites of shrimp from Texas rearing pondsTexas ABM University Sea Grant CollegePubllcatlan No. TAMU-SG-74-207, 19 p

Markham, J.C. 1982. Bopyr d isopodsparasibc on decapod crustaceans in HongKong and southern China Pages 325-391,/n 8 Morton, ed, Marine tfors snd fauna ofHong Kong snd southern Chlns. lntroduc-lion and taxonomy.

McCauley, J.E 1962 Ellobopsidae fromthe Pacihc. Science, 137:867-868.

Malhewa, C P., M El-Muaa. M. Al-HOS-saini, M. Samuel and A.R. Abdul Ghaffar.1966. Inftxstatiuns of Ep pevaae�&i e'cigaron Penaeus sernrsufcalue and their use asbfofogtcaf tags J. Crated Stot., 8:53-62.

Nearhos, S.p. 1980 Aspects ol parasia.zation cl Ctu~ penaefd prawns by twomembers of the family Bopyridae I fsopca4:Eprcaridea!. M. Sc Qual. Tttesis University olCf ueensland.

hloarho«, S P and A J G Laster 1984Ncnx records ot Bapy dao iCrustacea isopoda E p c,fr iduni i!om Queen sf and watt!raMem. Qveenef. Mus. 21 257 259

Owens. L 1983 Bopyrid para«ite Eprpe- xt!wn viqiin«Nobili as a b otog caf markerior tMnan,t pawn Piin,tous mergurensrs deM,in Aust J. Mar. Fresh. Ree�34 477-481

Thontas, MM 1977 A new reoord atEprprfrtaoon ingiins Nob k Bopyr dae, Iso.poda I pai.tsitic on Penaeus semisulcarus deHarm from Palk Bay and Gulf of Mannerindian J. Ffeh. 24 258 261

vad !r w 1973 A bibhography ot IheE'lobiopsidau. 1959 1971 v nth a list of Tha-i,issomyces speoes and their hosts Sars4,Na 52, pagus 175 179

WORMSAldrich. D V 1965 Observaaons on Ihe

ecology and life cycle al Prochnshanelra pe.neer Kruse lCustoda Trypanorhynchal J.Paresltol., 51 370 376

Corhern, C C 1970 Invest gabons of thehofminthes lrom the hepatopancreas of thebrawn shnmp Penaeus azfecus Ives, tramGalveston Bay, Texas M Sc Thesis, TexasAft M University, 51 p

- 1977 Protozoan and metazoansymbionts of naluia I and inar cultured popufat ons af commercial shrimp from the north-western coast ot the Gulf af Mexico 0 sser-talion Texas AftM University, 167 p

1978 A larval cyciophyadeanICeatoda TryPanarhynohaf J. Paraeltal45 490

Deardartf T L and R M. Overstreet 1981,Larval Hyaierorhyfa arum - ThynnaSCar!S!iNornatoda Anfsak daei trom hshes andinvertebraleS in the Gulf Ot Mexfac. Prac.Helminthok Soc, Wash., 48'113-126

1981. Review ot Hysferotffytaciumancf lhenngascans both previouslyThynnascaris! Nematode. Arusakrdae! fromthe northern Gull of Mexico Prac. Blot. Scc.Weeh., 931035-1079.

Fergenbaum, D L 1975 Parasites ot thecommercial shrimp Panacea vannamefBoone and Penaeus braau'ernie Latreile Btdf.ScL 25.491-514

Fe genbaum, 0 and J Carnuccao. 1978.Companson between the trypanorhynchidces tode infections of Penaeus duorarum andPenaeus brasrfrensrsin Biscayne Bay. Rorida.J. Inwrtebr. Pathal,. 28:127-130

Fusco, A C and R M Overstreet. 1978.Sprrocamaflanus encores sp n. and $. haf-ifrophus sp n lNematoda: Camaftwtidea!from hah as in the northern Guff ot Mexico. J.Paraaitol., 64.239-244.

Hutton, R. F.. T. Balf and B. Efdred. 1962.Immature nematodes of the genus Cantra-caecum Ra!liat and Henry, 1912, fromshrimps. J, ParasltoL, 48 327-332.

HuttOn, R.F., F Scgantkees-Barnet astd8 Eldred, 1959. Another species of igcfcpfHaffus Ward, tggt from the pink shrimp, Pe-naeus ducran'tref Burkenroad. J. ParaafaeL,45 490-490.

Hutton R F., F Sogandares-Bernal, BEldred, R M. Ingle and K D. Woodbum 1959Invesb gati ons on the parasites and diseasesof the saltwater shrimps of sports and com-mercial imporancee ot Florida. Tech. Ber. Fle.Bd. Conserv., 26:1-38

Kruse, D N 1959 Parasites ot lhe com-rnerical shrimps, Penaeus azfecus Ives, Pduerarum BurkOnr cad and P Serif'erus Unnaeus!. Tulane Stud. Zoot., 7:123-144,

Natarafan, P 1979. On the occurrence ottrypanorhynchan pleracercoid in lhe marineprawn Penaeusindicvs. Curr. Res. Ifndfa!,8 63-64

Norris, D.E. and R.M OverstreeL 1976The public health implicabons of larvalThynnascaris nernatodes from shellfish J.Milk Food Technol., 39.47-54.

Overstreet, R M 1983 Metazoan syrnbi-onts of crustaceans Pages 155-250, in A.J.Pravenzano, ed., Pathoblotogy. The biol-ogy of Crustecea, Academic Press hl. Y

Owens, L. 1985 Poiypocephaius sp Cestocla Lecanicephafidae! as a biologicalmarker fot banana prawns, Penaevsrnerguiensfs de Man, in the Gulf of Carpen-taria. Aust. J. Mar. Freshw. Res., 36 291-299.

Owens, L. and J.S. Gfazebraok 1985The biology of bopyrid isopads parasitic oncornmerciaf penaeid prawns in northernAusoafia. Pages 105-1 13, in P.C. Rothlis-berg, B,J, Hill and D.J. Staples, eds. SecondAust. Nat. Prawn Bem. NPS2, Cleveland,Australia.

Owens, L. 1987. A checklist of metazoanparasiles from hlatantia ex duding the crust-acean parasitess of the Cartdea! J. ShellfishResearch, 6:117-124.

Ragan, J.G and D V. Aldrich. 1972 Infect-ionn of brown shrknp. Penaeus azfecus ivesby Prochrr'sflaneffa penaei Kruse Cestoda:Try panorhyncha! in southeastern LouisianaBays. Trans. Am. Fish. Soc., 10 1:226-238.

Sogandares-Bernal, F. and R.F. Hutton.1959. The identity of Metacercte ia B reportedfrom the pink shrimp, Penaeus duorarvmBurkenroad. by Woo&urn, et al in 1957. J,Paraeftol, 45:362 and 378.

INJURYFontaine, C.T. and R.C. Dyjak. 1973. The

development ol scar tissue in the brownshrimp, Penaeus azfecvs, after woundingwith the Pelersen disk tag. J. fnvertebr.Pathof.. 22:476-477

Fontaira, C.T. and D.V. Lightner. 1073.Observations on lhe process of wound repairin pen~d shrimp J. Invertabr. Pafhof.,2223-33.

1975 Cot tufar response to injury inaenasd shrimp. Mar. RsfL Rev., 37:4-10.

Hafcfotv, K 1988. Absence of epicutfcfelrom lhe repair caMe produced by ttour ma-acosfrat~ ciustacaaits. J. ~staceahMoL, 8:346-364.

Nyhfan, L and T, Unastam, t 98tf. Woundeetctfcns and Aftaftnt rttytass asafci growth in

crayhsh cuticle J. Invertebr. Pathat. 36 187-197

DEVELOPMENTAL PROBLEMSAbaraz. M and F Sanda 1981 Oxygen

consumpbon by Nephrops norvegicvs L ! Crustacea, Decapoda! in relationship to itsmoulting cyde J. Exp. Mar. Biol, EcoL,54 113 118

Chen, HC. 1981. Same abnormal as-pects of the prawn Paiaemon eiegans exposed to heavy metals. Proc. R.O.C.-U.B.Coop. Bcl. Seminar an Fish Disease Na-bonal Sci Council, R O.C, 3.75-83.

Clark, J, V, 1986. Inhibition of moul ting inPenaeus semisuicalus de Harm! by long-term hypoxia Aquaculture, 52.253-254.

Menthe, D., R Malone and H Perry1984 Factors affecting molting success ofthe blue crab Caifinectes sapicrus Rathbunheld in dosed, recirculating seawater systemsAbstracts, 1984 Ann. Meeting, NagonalShe tlfisheriee Assn., June 25-28, Tampa,Florida, page 40

Stern. S and O Cohen. 1982, Oxygenconsumption and ammonia excretion duringthe moult cyde of the freshwater prawn,tufacrobraohiumraaenbergii de Man!. COmp.8 ice hem. Physi of. A., 73:417-419

Wickins, J F 1976 Prawn biology andculture. Oceancgr. Mar. Blot. Annu. Rev.,14 435-5D7

COI OR ANOMALIESLiao, I.C, F, R. Yang and S W. Liu. 1977.

Preliminary report on some diseases of cultured prawn and their control method. Re-ports on Fish Disease Research f!, JCRRFisheries Settee No. 29.28-33.

Johnson, S.K. 1978. Handbook ot shrimpdiseases. Texas ABM Univ. Bea GrantPublication SG-75-603, 23 p.

DIGESTIVE GLANDMANIFESTATION

Baubsta, M.N. 1986. The response ofPenaeus monodon juveniles to varying pro-tein/energy ratios in test diets Aquaculture,53 229- 242.

Doughtie, D.G. and K.R. Rao. 1983. Ul-trastructural an histological study of degen-erative changes in the antennal glands,hepatopancreas, and midgut of grass shrimpexposed lo bvo dithiocarbamate biocides. J.fnverfebr. PathoL, 41:281-300

Egusa S., Y. Takahashi, T. llami and K.Momoyama. 1988. Histopathofogy of vfbrf-osis in the Kunana prawn, Penaeus iaponi-cus Bate. Rsh Pathology, 23:59-65.

Ughtner, D.V., R.M. Rodman, R.P, Priceand M.g, Wiseman. 1982. Htstopathology otaffatoxfcosis in the marine shrimp Penaeussfyfimsfrfs and P. vannamei. J. Inverfabr.Pathoi., 40:279-29 1.

Lightner, O.V, and R,fiif. Redman. 1g85.Necrosh of lhe hep dopancraas in ~rncvtcrdon tatd P. sffrffrosfifswith red disease.Jotwnaf of Bah Bfseeeea, 8:181-168.

Sparke. A.K 1988 Mull pie g:an.ilon asin the midgut of the dungei,ess crab, Caixcermagister. J. Invertebr. Pathol., 35 3 3-3?4

Rosernark, R, P R Bowser and N Baurn1980 Histological observalians ol lhe hepatopancreas in juvenile lobsters subiected todietary stress. Proc. World Mariculture Soc,11 471 478

Vogt,G 1987. Moni:oringofenvironn entalpollutants such as pest,eicos in prawn aquaculture by histological diagnosis Aquacul-lure, 67 157-164

INFLAMMATION ANDMELANIZATION

Couch, J A, 1971 Ultrastiuctural study oflesions in gills of a marine shrimp exposed tocadrniurn J. Invertebr. Pathol.. 29 26 -288

Doughbe, D G, P J Conklin and K RA ac. 1983 Cuticular lesions induced in areasshrimp exposed to hexavalent chromium. J.lnvsrtebr. Psthol., 42 249-258

Daughtie DG and K.R, Aao 1983 Ul-trastructural and histological study ol degenerabve changes leading to black gills in grassshrimp exposed to a dothiocar ham ate biocide.J, lnvertebr, PathoL 41.33-50

Ughtnet, D.V. and R.M. Redrnan 1977Histochemical demonstration of melanin incellular inflammatory processes of penaeidshnmp. J. Invertebr, Pathol., 30.298-302

Nimrno, D.R,, Lightner, D.V. and L HBahner 1977 Effects of cadmium on theshrimps, Penaeus duorarvm, Paiaemonetespugio, and P vvigaris..Pages 131-183, inF.J. Vernberg, A Calabrese, F.P Thurbergand W.B. Vernberg, eds Physiologicalresportaea of marine blots to poffulants.Academic Press, N.Y.

Soderhall, K, L. Hall, T Unestam and LNyhlen. 1979 Attachment of phenoloxidaselo fungal cell walls in arthropod immunity J.tnvertebr. Pslhof., 34 285-294

Solangi, M.A. and D V. Lightner 1976Cellular inflammatory response ot Penaeusazfecus and p. setirerus to the pathogenicfungus, Fusarium sp, isolated trom the Cali-fornia brown shrimp, P. cafrforniensis. J.lnvertebr Pathoi., 27 77-86

EMACIATION AND NUTRITIONALDEFICIENCY

Baticados, C.L., A.M Coloso and R.C.Duremidez. 1986. Studies on the chronicsoR shelf syndrome in the tiger prawn, Pe-naeus rnonodon Fabncius. from brackishwater ponds. Aquaculture, 56:271-285.

Central Marine Fisheries Research Insti-tute, Cochin, india. 1982-83, 1983-84, 1984-85. Physiology, Nutrition and PathologyDivision, Annual Reports.

Dali, W and D M. Smith. 1986. Oxygenconsumption and ammonia-N excretion infsd and starved tiger prawns, Periaeusescufert fee Haswell. Aquaculture, 55:23-33,

Lightner, D.V., L.B. Calvin, C Brand andD.h. Danafd. 1977. Black dealh, a diseasesyndrome related lo a dietary deficiency of

iii�,i. id Proc. World IylaricullureSoc,8 rii ' 8,"3

i i D V, B hunter, P C Magarelli.;ind L B Colvin 1979 Ascarbic acid

iiuiiiiional iequiriirnent and role in woundii i,iii in pnnarid Shiimp PrOC. WOrld MariC-ult. Soc, 6 34 7 3F5

M,ig,iriiili I' C, Jr, B Hunter, D V Lightner and I 8 Colvin 1979 Black doalh anaSCOrbic .icid duhoienCy diSOaSe in penaeidshrimp Comp. Biochem. Physiol., 63A 103108

Vogi, G, V Starch, E T Quinitio and F PPasrual 1985 Midgul gland as monitor or-gan ior Iho nutriuonal value of diets in Pe-naeus maiiodon Docapoda! Aquaculture.48 i 12

CRAMPED SHRIMPJolinson, S K 1975 Cramped Conditian

in pond reared shrimp Texas AAM UnivFish Disease Diagnostic Laboratory Leaf le No FDDL S6

MUSCLE NECROSISAki yam a D M . J A Brack and S R Hel ay

1982 Idiopathic muscle necrosis in the cul-tured freshwater prawn, Macrobrachiumrosenbergii VMiSAC, 11 '19-1121

Delves-Broughton. J and C W Poupard,1976 DiSeaSe prOblemSOI prawnsin recircu-lation systems in the U K, Aquaculture 7 201-217

Lakshmi. G J, A Venkataramiah and H.D.House, 1978 Effect at salinity and temperature on spontaneous muscle necrasis irlPenaeus aztecus Ives. Aquaculture, 13.35-43

McDonakt, D G.,B.R. McMahon and C M.Wood 1979 Ananalysisofacid-base distur-bances in the haemolymph tollawing stre-nous activity in the Dungeness crab, Cancermagisler J. Exp. Biol�79:49 58

McMahon 1 R .. D G McDonald and C.M.Wood 1979 Ventilation, oxygen uptake andhaemOlymph oxygen tranSpOrt, follOWrngenfOrCed exhausting activity in the DungeneSScrab, Cancermagisfer J. Exp, Biol.,80.271-285.

Momoyarna. K and T. Matsuzata 1987,Muscle necrosis of cultured kururna shrimp Penaeusiaponicus!. Fish Palhology, 22.69-75

Nash, G, 5 Chlnabut and c. Urnsuwan.1987 Idiopathic muscle necrosis in the fresh-water prawn, Macro brachium rosenbergil deMan, cultured in Thailand, Journal ol FfshDiseases, 10 109- 120

Phillips, J.W, R.J.W. McKinney, F.J,RHird and D.L. MacMillan. 1977. Lactic acid

tu malice in cruslaceans and the liver tunc-tian Oi thi midgiil gland quCStiOned Cam-parative Biochemistryandphystology56B,427 433

Rigdon, R H and K N Baxter 1970Sponlaneous necroses in rnusclos of brownshi mp. Periadiis 4= locus IvOS Tran S. Am.Fish Soc. 99 583 587

Spotts, D G and P L Lutz 1981 L-lacticarid aCCumulatian during aChVity StruSS inAtrcrabrachrurn rosengergii and Penaeusduorarurn J. World Mar. Soc., 12 244 249

TUMORS AND OTHER TISSUEPROBLEMS

Lig'itner, D V, R M Redman, T A Belland J A Brock 1984 An idiopathic proliferative disease syndiome ot the midgut andvenlral nerve in the Kuruma prawn, Penaeuslaponicus Bate, cuilured in Hawaii Journalof Fish Diseases, 7 183-191

Overstreot, R M and T Van Devender1978 Implication of an onvironmonlally in-duced hamartoma in commercial shrimps J,Invertebr. Pathol., 31 234-238

Sparks, A K 1972 Inverlebrale palhol-ogy: Noncommunicable diseases. Aca.demic Press, New York

Sparks, A K. and D V Lightner 1973. Atumorlike papillilorm growth in the brownshrimp I Penaeus az focus!. J. Invertebr. Pa-thol 22 203 212

ENVIRONMENTAL EXTREMESBrissen, S, 1985 Gas-bubble disease

observed in pink shrimps, Penaeus brasrfren-sis and Penaeus paulensis. Aquaculture,47.97-99.

Lightner, D V., B.R. Saiser and RSWheeler. 1974. Gaa-bubble diSeaae in thebrown shrimp Penaeus azfecus!. Aquacul-ture, 4 81-84

Nash, G, I G Anderson and M. Shantf1988. PathOIOgiCal Changee in the bger prawn,Penaeus monoobn Fabricius. associated withculture in brackishwaler ponds developedfrOm pateniralty aCid Sulfate SOile. Jour tisl ofFish Diseases, 11;113-123.

Simpson, H.J., H.W. Ducklow, B Deckand H L Cook. 1983. BraCikiSh-Water aquaC-ulture in pyrite-bearing trOpiCal SOilS. A quac-ullura, 34 333-350

Supplee, V.C, and D V. Lightner. 1976.Gas-bubble disease due to oxygen super-sa turation inraoeway-reared Ca4f amia brawnshrimp. Prog. Fish-Cult., 38:158-159.

Wicluns, J.F. 1984 The effect of hyper-capnic sea water on growth and mineraliza-tion in penaeid prawns Aquaculture, 41:37-48.

TOXIC ITYChen J C Y Y Ting H Lin and T C

Lian 1 85 Heavy metal conoentrations insea water from giass prawn hatche e andtlie coast ot Taiwan J. World Marlcuh Boc.,16 316 332

Couch. J A 1978 Diseases, parasiteS,and taxic responses oi commeroal penaeidshrimps ol the Gulf of Mexico and SouthAtlanhc coasts of North America Fish. Bull,76 1-44

Couch, J A and I . Courtney. 1977. Inter-action of chemical pollutants and virus in acrustacx.an a novel bioassay system. Ann.N.Y, Aced. Scl., 228 497 504

Liener, I E, 1980 Toxic constlluents ofplanl foodstuffs. Academic Press, N Y

Lightner, D V, D A Danald, R M Red-man, C Brand, B.R Salser and J. Reprieta1978 Suspected blue-green algal prxsoningin the blue shrimp. Proc. World Ibxricutt.Soc. 9:447-458

Lightner, D.V 1978. Possible toxiceffectsof the marine blue-green alga. Spirulrnasub-salsa, on the blue shrimp, Pen acus styli res tris.J. Inverlebr. Palhal., 32 139-150

1988 Bunted head syndrome ofpenaeid shnmp. Pagos 93-95, inC.J Sinder-rnann and D V Lightner, eds, Diseasediag-nosla and coinlrol tn North American aq-uaculture. Elsevier, Amsterdam.

1988 Alfatoxicosis of penaeidshrimp. Pages 96-99, in C J. Sinderrnannand D.V IJghtner. eds . Disease diagnosis~ nd control in North American aquacul-ture. Elsevier, Amsterdam.

LOVell, R.T. 1984 Micrabial tckinS in fiahfeeds Aquacult. Nag., 10 34.

Medi yn, R A. 1980. Susceptibility of fourgeographical strains ol' adult Artemis toPfychodtSCuSbreviStOXin S!. Pages 225-231,in G Persaone, P. Sorgeloos,0 Roeis, aridE. Jaspers. eds. The brine shrimp Artemis,Volume I, ikorphology, 13enetfcs. Radlobl-OIOgy, Toxi colOgy. UniVerea PreSs, Wettern,Belgium. 345 p

Crverstreet, R M. 1988. Aquatic pollutionproblems, Southeastern U.S. coasts: histo-pathological indicators. Aquatic Totdcotogy,11:213 239

Sindermann, C J. 1979 Pollution-associ-ated diseases and abnormakties of tish andshellhsih: a review Fish. Bull.. 76 717-749

Vogt, G 1987 Monitoringofenvironmentalpollutants such as pesticides in prawn aquac-ulture by histological diagnosis. Aquacul-ture, 67 157 164,%

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